Novon a journal of botanical nomenclature from the Missouri Botanical Garden.

Volume 5
Number 1
1995

GARDEN LIBRARY.

Notes on the Asclepiadaceae of China

Michael G. Gilbert

Flora of China Project, Missouri Botanical Garden, c/o Department of Botany,
Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom

W. D. Stevens

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Li Ping-tao

College of Forestry, South China Agricultural University, Wushan, Guangzhou 510642,
Guangdong Province, China

Abstract. This paper is a precursor to the account
of the Asclepiadaceae in the Flora of China. A new
genus and species, Sichuania alterniloba, is de¬
scribed. Twenty-four new species are described in
Biondia (B. crassipes, B. laxa, B. parviurnula,

B. revoluta, and B. tsiukowensis ), Ceropegia (C.
sinoerecta), Cynanchum (C. bicampanulatum, C.
brevicoronatum, C. duclouxii, C. kingdonwardii,

C. longipedunculatum, C. megalanthum, C. ping-
shanicum, C. rock 'd, and C. sinoracemosum ), Hoya
(H. commutata and II. mekongensis ), Marsdenia
( M. brachyloba, M. tend, and M. yuei), and Ty-
lophora (T. forrestd, T. rockii, T. tuberculata, and
T. uncinata). Jasminanthes is resurrected. Eleven
new combinations are proposed in Ceropegia (C.
exigua), Cynanchum ( C. boudieri subsp. cauda-
tum), Heterostemma (//. menghaiense), Hoya (H.
chinghungensis), Jasminanthes ( J. chunii, J. mu-
cronata, J. pilosa, and J. saxatilis), and Tylophora
(T. costantiniana, T. oligophylla, and 7. tsiangii).
The new name Cynanchum triangulare is proposed
to replace C. deltoideum Hooker, not Hance. New
synonymy is proposed in Cynanchum, Lygisma,
Marsdenia, and Tylophora. The status of Merril-
lanthus is discussed.

The following new taxa and new combinations are
required for the forthcoming account of the Ascle¬

piadaceae for the Flora of China. Some new syn¬
onymy is also presented. The taxa are arranged
alphabetically.

Biondia Schlechter.

Biondia, a genus of 13 species endemic to China,
is characterized by narrow leaves with minute pale
dots on the adaxial surfaces, small flowers mostly
with well-developed corolla tubes, and reduced, often
annular, coronas. Such leaf dots are also seen in
some species of Tylophora, but they have restricted
occurrence in genera related to Biondia. The most
important exception is Cynanchum thesioides K.
Schumann, which also has narrow leaves. This spe¬
cies is somewhat isolated in Cynanchum and has
been placed in its own genus, Rhodostegiella C. Y.
Wu & D. Z. Li. There is perhaps a case for regarding
it as a Biondia with a more normally developed
corona than usual.

Most species of Biondia are too poorly repre¬
sented in herbaria to establish patterns of variation,
and it is difficult to have much confidence in species
delimitation. A number of collections cannot be placed
within the known range of variation of existing taxa.
These are herein described as distinct species. It is
likely that more information on variation within spe-

Novon 5: 1-16.1995.

2

Novon

cies will require a reassessment of the number of
species recognized.

Biondia crassipes M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Sichuan: Yiajiang Xian,
2700 m, 10 Sep. 1977, B. Z. Guo & W. Y.
Wang 23624 (holotype, HNWP 88274).

Species nova Biondiae tsiukowensi M. G. Gilbert &
P. T. Li similis sed ab ea foliis angustatis nervatione laterali
non prominenti, rachidibus inflorescentiarum plus minusve
pedicellos aequantibus, rachillis cymularum elongatis plus
minusve incrassatis differt.

Twining herbs. Stems puberulent along 2 sides
and at nodes. Petioles 2-4 mm, adaxially densely
puberulent; leaf blades linear-oblong, to 5 x 0.3
cm, paler abaxially, base cuneate, apex acute; midrib
raised adaxially, lateral veins obscure, glabrous. In¬
florescences 2—3.5 cm, mostly with two cymules
separated by a rachis ca. as long as pedicels; rachillas
of cymules slightly elongated and ± fleshy, very
sparsely puberulent. Pedicels 6-10 mm. Sepals el¬
liptic, ca. 1.5 x 0.6 mm, sparsely puberulent. Co¬
rollas apparently green, bell-shaped, 2.5 mm, ex¬
terior glabrous, interior minutely hairy, hairs short,
stiff, white; tube ca. 1.5 mm; lobes triangular, rev¬
olute, twisted. Coronas reduced to inconspicuous
collar concealing bases of anthers. Pollinia oblong-
reniform. Follicles solitary, narrowly fusiform, ca.
5.5 x 0.4 mm.

The narrow leaves and small flowers of Biondia
crassipes suggest a relationship to B. longipes P.
T. Li and B. revoluta. The corolla form is most
similar to those of B. microcentra (Tsiang) P. T. Li
and B. tsiukowensis, while the lax inflorescence is
similar to that in B. laxa. Biondia crassipes differs
from all other species of Biondia by the slightly
elongated and distinctly thickened rachillas of the
cymules; all other species have contracted umbel-
liform cymules. Biondia longipes differs further
from B. crassipes by the glabrous corollas, while
B. revoluta differs by the longer and more distinctly
tubular corollas.

Biondia laxa M. G. Gilbert & P. T. Li, sp. nov.
TYPE: China. Yunnan: Ouei cha (Yunpe), 13
May 1920, Ten 404 (holotype, E).

Species nova Biondiae insigni Tsiang similis sed ab
ea foliorum nervis lateralibus utroque costae latere 4-6,
omnibus similibus, rachidi inflorescentiae longa cymulas
plerumque 3 gerente differt.

Twining herbs. Stems densely pubescent along 2
sides. Petioles to 7 mm; leaf blades lanceolate, 4.5-
6 x 1.2-2 cm, glabrous, base rounded, apex acute;

lateral veins 4-6 pairs, at ca. 45° to midrib, all
similar, adaxially slightly raised and minutely pu¬
berulent. Cymes extra-axillary; peduncles to 2.5 cm,
glabrous; cymules umbel-like, up to 3 along zigzag
rachis to 2 cm. Pedicels 1-1.3 cm, very slender.
Sepals ovate, ca. 1 x 0.5 mm, glabrous or sparsely
pilose on margins. Corolla tubes bowl-shaped, ca. 1
mm, much shorter than lobes; lobes triangular-acu¬
minate, ca. 2.5 mm, revolute, apex twisted, very
minutely puberulent. Coronas ± annular with fleshy,
square to ± rounded lobes to base of anthers. Gy-
nostegium slightly exserted from corolla tube; pol¬
linia cylindric, as long as corpusculum. Fruits not
seen.

Biondia laxa is immediately separable from all
other members of the genus by the inflorescence,
which has a distinct rachis to 2 cm long and up to
three separate umbel-like cymules. Older leaves have
thick, pale veins very similar to those of the closely
related B. insignis. However, B. insignis has a
distinctive leaf venation with two major lateral veins
at an acute angle to the midrib, as well as up to
eight pairs of minor veins at a wider angle. Biondia
insignis also differs by the ringlike corona.

Paratypes. CHINA. Xizang (Tibet): Rong To Val¬
ley, Layul, 1800 m, 20 Apr. 1933, Kingdon-Ward 10352
(BM). Yunnan: Lao Kouy Chan, near My le, 1906, P.
Ngueou in Ducloux 4171 (P); Shweli-Salwin Divide at
25°30'N, 2700 m, Aug. 1917, Forrest 15718 (BM, E).

Biondia parviurnula M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Anhui: Jinzhai, Guan
Cai Gou, Bai Ma Zhai, 800 m, 4 Mar. 1984,
K. Yao 8956 (holotype. A; isotype, K).

Species nova Biondiae hemsleyanae (Warburg) Tsiang
similis sed ab ea nervatione foliorum prominentiore, in-
florescentiis minimis, corollis minoribus urceolatis differt.

Twining herbs. Stems densely minutely puberu¬
lent along 2 sides. Petioles to 7 mm, adaxially pu¬
berulent; leaf blades lanceolate, to 6.5 x 2.2 cm,
base rounded, apex acute-apiculate, glabrous except
for adaxially densely puberulent midrib; lateral veins
3 or 4 pairs, slightly raised on both sides; upper
leaves relatively narrower, ± elliptic to elliptic-ob¬
long. Inflorescences umbel-like, ca. 4-flowered; pe¬
duncles ca. 2 mm, puberulent. Pedicels 1.5-2 mm,
glabrous. Sepals lanceolate, ca. 1.5 x 0.7 mm, apex
acute, pubescent, somewhat glabrescent. Corollas
urn-shaped, 3-3.5 mm, interior pale purple, exterior
glabrous; tube ca. 2.5 x 2 mm, throat ca. 1.2 mm
wide, interior sparsely hairy; lobes ovate, ca. 1.2
mm, apparently not revolute. Corona a collar cov¬
ering bases of anthers, thin when dried. Stigma head
conical. Fruits not seen.

Volume 5, Number 1
1995

Gilbert et al.

Asclepiadaceae from China

3

The type of Biondia parviurnula was identified
as B. hemsleyana, but B. parviurnula is readily
distinguished by having leaves with more prominent
venation, very small inflorescences (less than 1 cm
long), and smaller, more distinctly urn-shaped co¬
rollas. The corolla of B. parviurnula is most similar
to those of B. microcentra and B. yunnanensis (H.
Leveille) Tsiang but is glabrous inside. The type
material also differs from those species by having
internodes with two lines of pubescence.

Biondia revoluta M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. NW Yunnan/E Xizang (Ti¬
bet): Tung-chu-ling, 3000 m, May 1913, King-
don-Ward 310 (holotype, E).

Species nova Biondiae pilosae P. T. Li similis sed ab
ea internodiis bilateraliter pubescentibus, foliis manifeste
revolutis nervatione laterali prominentiore, pedicellis lon-
gioribus differt; a speciebus aliis lobis corollarum tubum
aequantibus differt.

Twining herbs. Stems hairy along 2 sides. Petioles
to 3.5 mm, puberulent; leaf blades linear-oblong, to
2.8-6 x 0.4-0.55 cm, base ± rounded, margins
strongly revolute, apex subacute, apiculate, glabrous
except for adaxial midrib, lateral veins not raised,
underside distinctly paler, wrinkled when dried,
probably fleshy when fresh. Peduncles 1-1.5 cm,
hairy along 1 side; cymules umbel-like, 2-4-flow-
ered, solitary or 2 separated by rachis to 5 mm,
puberulent. Pedicels 7-9 mm, glabrous. Sepals ovate,
0.7-1 x 0.5-0.7 mm, sparsely puberulent, apex
acute. Corollas bell-shaped, inside minutely puber¬
ulent, 2.5-3.5 mm, lobes ovate, ca. 1.5 x 1.5 mm,
apex rounded, slightly revolute. Coronas annular,
obscurely 5-toothed. Fruits not seen.

Biondia revoluta is most closely related to B.
pilosa, which differs by having internodes with only
one line of hairs, leaves with acuminate apices and
more conspicuous lateral veins, and shorter pedun¬
cles. The collection from Gansu ( Wang 19120) has
smaller leaves and flowers than the type collection,
and the lateral leaf veins are distinctly darker than
the lamina and are not raised. In all other respects,
it is a good match and is included here as a paratype
despite the disjunct distribution.

Paratype. CHINA. Gansu: Wen Xian, 1500 m, 30
Apr. 1966, Z. B. Wang 19120 (HNWP).

Biondia tsiukowensis M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Yunnan: Salween-Me-
kong Divide, Tsiu kow, 2400 m, May 1911,
Kingdon-Ward 110 (holotype, E).

Species nova Biondiae microcentrae similis sed ab ea
internodiis bilateraliter pubescentibus, pedunculis longior-
ibus, lobis corollinis ca. % partes longitudinis tubi ae¬
quantibus differt.

Twining herbs. Stems densely pubescent along 2
sides. Petioles ca. 6 mm, adaxially puberulent; leaf
blades lanceolate, ca. 6x1.2 cm, glabrous except
for veins, base rounded, apex ± acute, slightly apic¬
ulate; lateral veins 5-7 pairs, adaxially slightly raised
and minutely puberulent, at an acute angle to midrib.
Cymes umbel-like; peduncles 2-18 mm, almost gla¬
brous. Pedicels to 11 mm. Sepals ovate, ca. 1.4 x
0.7 mm, glabrous. Corolla tube bell-shaped, ca. 2.3
mm long, longer than lobes; lobes triangular-acu¬
minate, 2x1.4 mm, revolute toward apex, mi¬
nutely puberulent inside. Corona lobes ± square,
reaching base of anthers. Gynostegium ca. 1.4 mm
high; pollinia cylindrical, as long as corpusculum.
Fruits not seen.

Biondia tsiukowensis is distinguished from other
species of Biondia by having longer peduncles and
corolla lobes only slightly shorter than the tube.
Other species of Biondia have corolla lobes either
less than half as long as or distinctly longer than
the corolla tube. It also differs by its more or less
square corona lobes that reach the bases of anthers.
Biondia microcentra is in some ways most similar
because it also has the lateral veins of the leaves
raised adaxially and a hairy interior to the corolla,
but it differs markedly by having much shorter (to
4 mm) peduncles and pedicels.

Ceropegia L.

Ceropegia , which includes about 170 species
mostly in Africa, is represented in China by 17
species.

Ceropegia exigua (Huber) M. G. Gilbert & P. T.
Li, comb, et stat. nov. Basionym: Ceropegia
longifolia Wallich subsp. exigua Huber, Mem.
Soc. Brot. 12: 42. 1957. TYPE: China. Si¬
chuan: Tung Valley, Wilson 4112 (holotype,
BM; isotypes, K, P).

Ceropegia exigua is perhaps the most distinctive
member of the C. longifolia Wallich complex. It
can be recognized instantly by the very short corolla
lobes, which are abruptly contracted from the prom¬
inently spreading top of the corolla tube to form a
short narrow beak, even when the flower is fully
open. The corolla lobes in the other members of this
group are much longer (usually at least three-fourths
as long as the corolla tube, sometimes longer), and
the upper part of the tube does not spread and

4

Novon

passes much more gradually into the lobes, which
are not abruptly contracted into a beak. Further¬
more, in C. exigua the corolla lobes are only slightly
darker than the rest of the corolla and are quite
coarsely pilose, while in the rest of the complex, as
in many other species in the genus, the corolla lobes
are deeply colored and finely and densely pubescent,
contrasting greatly with the rest of the corolla.

Ceropegia sinoerecta M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Yunnan: au pied du
Hee Chan men (Ho Kin), 2000 m, 30 May
1887, Delavay 2625 (holotype, P).

Species nova Ceropegiae mairei (H. Leveille) Huber
similis sed ab ea corollae angustioris basi quam ostio latiore
lobis linearibus fere ad basim atratis manifeste differt; a
C. wallichii White partibus omnibus gracilioribus differt.

Erect herbs. Rootstock a cluster of fusiform roots.
Stems erect, to 20 cm, uniformly finely puberulent.
Petioles 4-6 mm, narrowly winged; leaf blades el¬
liptic, 2-5 x 0.6-1.6 cm, base cuneate, apex acute,
lowermost leaves sometimes cuspidate, abaxially pale
and sparsely hairy on veins only, adaxially densely
puberulent. Peduncles 4-17 mm. Cymules umbel¬
like, 2-4-flowered, sometimes producing a second
cymule separated by a short rachis. Pedicels 5-17
mm. Sepals linear-lanceolate, ca. 3 x 0.6 mm,
glabrous. Corollas 3.6-4.3 cm, glabrous except for
lobes, tube dark for basal half, pale for apical half,
basal swelling ovoid, 4-6 mm wide, tube 1.3-1.6
mm wide, increasing gradually to 3.5-5.5 mm at
base of lobes; lobes darkly colored almost to base,
minutely puberulent, 14-15 mm, linear, slightly
curved inward at base, otherwise parallel except for
sharply incurved tip. Outer corona lobes each with
2 linear teeth ca. 1.5 mm, ciliate; inner corona lobes
linear, ca. 2.5 mm, erect. Fruits not seen.

Ceropegia sinoerecta is perhaps most closely re¬
lated to C. mairei, from which it differs by the more
slender flowers widest at the basal inflation and by
the differentiated corolla tips extending almost to
the base of the lobes instead of less than halfway
down. Ceropegia wallichii White (Nepal and west¬
ern Himalayas), which also has erect stems and
similarly shaped corollas, is a robust plant with un¬
usually thick stems and much larger leaves and
flowers, whereas C. sinoerecta is delicate in all fea¬
tures.

The holotype has simple umbel-like inflorescences
and parallel corolla lobes, while the paratype has
inflorescences with pairs of umbels separated by a
distinct rachis and corolla lobes bowing slightly out¬
ward. More collections are needed to properly assess
the significance of these differences. There are sev¬

eral other collections by Delavay numbered as 2625,
but these differ in details of date and locality, and
all belong to C. mairei.

Paratype. CHINA. Yunnan: Kou ty pres Pin tchouay,
Apr. 1907, Jean Py in Ducloux 5305 (P).

Cynanchum L.

With 57 species, Cynanchum is the largest and
most difficult genus of Chinese asclepiads. It is being
treated here in the wider sense, i.e., including Vin-
cetoxicum N. M. Wolf, Cyathella Decaisne, and
Rhodostegiella. When the group as a whole is con¬
sidered, it is currently not possible to define adequate
discontinuities among the potential segregate gen¬
era.

Cynanchum acutum L. subsp. sibiricum (Willd-
enow) F. Rechinger, FI. Iranica, Asclepiada-
ceae 9. 1970. Cynanchum sibiricum Willd-
enow, Ges. Naturf. Freunde Berlin Neue Schrif-
ten 2: 124. 1799. TYPE: B-WILLD 5233
(microfiche).

Cynanchum cathayense Tsiang & Zhang, Acta Phytotax.
Sin. 12: 110. 1974. Syn. nov. Cyathella cathay-
ensis (Tsiang & Zhang) C. Y. Wu & D. Z. Li, Acta
Phytotax. Sin. 28: 465. 1990. TYPE: China. Gansu:
Chiu-Chuan Hsien, 1350 m, 2 Dec. 1956, R. C.
Ching 30 (PE).

Cynanchum cathayense was not compared with
C. acutum in the protologue. Numerous collections
from Xinjiang match the description of C. cathay¬
ense and are not separable from C. acutum subsp.
sibiricum. There seems to be no alternative but to
include C. cathayense in that taxon.

Cynanchum acutum L. subsp. acutum is wide¬
spread in the Mediterranean region from Spain east
to Turkey, where there is an area of overlap with
C. acutum subsp. sibiricum. It differs by the ovate
leaves with convex rather than concave sides, but
individuals with hastate leaves are found from
throughout the range, and it does not seem possible
to treat the two as species.

Cynanchum ascyrifolium (Franchet & Savatier)
Matsumura, Ind. PI. Jap. 2, 2: 509. 1912.
Vincetoxicum ascyrifolium Franchet & Sa¬
vatier, Enum. PI. Jap. 2: 441. 1879. TYPE:
“prob. ins. Nippon, cum V. multinerve mixtum,
ex Tanaka accepit Dr. Savatier” (holotype, P).

Cynanchum ascyrifolium has been used widely
recently in both Japan and China for a very dis¬
tinctive erect plant with large membranous acumi¬
nate leaves and small inflorescences. Unfortunately,

Volume 5, Number 1
1995

Gilbert et al.

Asclepiadaceae from China

5

the type of C. ascyrifolium is a plant with oblong-
elliptic leaves with a thick texture and rounded-
apiculate tips. It is clearly different from any other
species known to us. The correct name for most
material commonly included under C. ascyrifolium
is C. acuminatifolium Hemsley.

Cynanchum auriculatum Royle ex Wight, Contr.
Bot. India 58. 1834. SYNTYPES: India. Ka-
moun, Wallich 8228 (“asclep. 137”) (K-
WALL); Kimour, Royle s.n. (LIV not seen).

Cynanchum saccatum W. T. Wang in Tsiang & P. T.
Li, Acta Phytotax. Sin. 12: 91. 1974. Syn. nov.
TYPE: China. Xizang (Tibet): Chang-Tu, Chi-na-
tung, Tsa-wa-rung, 3000 m, Aug. 1935, C. W.
Wang 65215 (holotype, IBSC).

Cynanchum saccatum was described without the
examination of authentic material of C. auricula-
turn, from which it is indistinguishable. Specimens
of C. boudieri were often misidentified as C. auri-
culatum, but C. boudieri has strongly reflexed co¬
rollas, whereas C. auriculatum has rotate corolla
lobes.

Cynanchum bicampanulatum M. G. Gilbert &
P. T. Li, sp. nov. TYPE: China. Sichuan:
Kangting (Tachienlu), 2700 m, 11 July 1934,
Harry Smith 10443 (holotype, UPS; isotypes,
MO, S).

Species Cynancho vincetoxico sensu lato affinis sed ab
eo corollis coronisque campanulatis, lobulis coronae ad
antheras corollasque adnatis facile distinguenda.

Herbs. Roots many, little branched, fleshy. Stems
forming fairly dense cluster, little branched, erect
or ascending, to 30 cm, minutely puberulent mainly
along 2 sides. Petioles 3-9 mm; leaf blades oblong-
ovate, 5.5-7 x 4-4.5 cm, base shallowly cordate,
truncate or rounded, apex acute to slightly acumi¬
nate; basal veins 5-7, lateral veins 4 or 5 pairs,
glabrous or minutely puberulent on some. Inflores¬
cences ± umbel-like, cymules several flowered, sep¬
arated by a very short rachis; peduncles (5-)8-18
mm, puberulent; bracts linear, 0.9—2 mm. Pedicels
5—8 mm (to 12 mm in fruit), puberulent. Sepals
oblong, ca. 1.5 x 0.5 mm, apex rounded, almost
glabrous. Corollas bell-shaped, ca. 4 mm; glabrous
except for a few stiff hairs within tube, purple (fide
Guo & Wang 21046); tube ca. 2 x 3 mm; lobes
oblong-ovate, ca. 1.7 x 1.5 mm, apex rounded,
usually revolute. Corona tube ± as high as anthers,
membranous with indexed, rounded, fleshy lobes,
adnate to corolla tube and to anthers opposite lobes.
Anthers square, slightly gibbous; anther appendage
ovate, indexed over stigma head; pollinia ovoid, ca.

0.2 mm. Stigma head dark, rounded. Follicles paired
or solitary, spreading, glabrous, beaked-fusiform, at
least 3.5 x 1.5 cm.

Distribution and ecology. China, Gansu and Si¬
chuan Provinces. In open fields (“prato aprico”);
2570-2700 m.

At first glance Cynanchum bicampanulatum
looks very similar to some of the forms of C. forrestii
Schlechter, a member of the C. vincetoxicum (L.)
Persoon complex. However, in C. bicampanulatum
the corolla and corona are campanulate, in contrast
to the shallow-tubed, rotate corolla and bowl-shaped
corona of members of that complex. Cynanchum
stenophyllum Hemsley resembles C. bicampanu¬
latum in having the corona adnate to both the an¬
thers and corolla tube but differs in having narrow
leaves and almost no indumentum.

Paratypes. CHINA. Gansu: Erh-lau-shan ad Min-
chow, 2570 m, 30 June 1930 (fl), Hummel 3894 (S);
Minchow, 2410 m, 8 Sep. 1930 (fr), Hummel 5097 (S).
Sichuan: Daofu Xian, 5 July 1977 (fl), Guo & Wang
20861 & 21046 (HNWP).

Cynanchum boudieri H. Leveille & Vaniot, Bull.
Soc. Bot. France 51: CXLIV. 1904. TYPE:
China. Sichuan: Kouy-Tcheou, Pin-fa, ruisseau
du Tu-chang, 5 Oct. 1902, Cavalerie 620
(holotype, E).

Cynanchum amphibolum C. K. Schneider in Sargent,
PI. Wils. 3: 346. 1916. Syn. nov. TYPE: China.
Hubei Province: Patung Hsien, Aug. 1907, Wilson
2247 (holotype, A; isotype, K).

Cynanchum boudieri subsp. caudatum (Miquel)
P. T. Li, M. G. Gilbert & W. D. Stevens, comb,
et stat. nov. Basionym: Endotropis caudata
Miquel, Ann. Mus. Bot. Lugduno-Batavum 2:
128. 1866. TYPE: Japan, Keiske (?) (not seen).

The Chinese material of Cynanchum boudieri,
which has often been misidentified as C. auricula¬
tum, has elongated inflorescences, whereas the Jap¬
anese collections have umbelliform inflorescences.
Plants of the two countries are otherwise very sim¬
ilar, and they are best treated as subspecies.

Cynanchum brevicoronatum M. G. Gilbert &
P. T. Li, sp. nov. TYPE: China. Hubei (Hupeh):
Ichang, A. Henry 6418 (holotype, BM; iso¬
types, K, P).

Species Cynancho uiallichii simiiis sed ab eo calyce
reflexo corona brevissima cupuliformi membranacea in-
tegra appendicibus carente facile distinguenda.

6

Novon

Twining herbs. Steins puberulent along 1 line.
Petioles to 3 cm; leaf blades abaxially slightly glau¬
cous, ovate or ovate-triangular, 6-7.5 x 3-4.2 cm,
base cordate, apex acute to slightly acuminate, basal
lobes ± incurved, adaxially very sparsely and mi¬
nutely hairy, abaxially puberulent on veins, some¬
times pale gray-green; basal veins 3(-5), lateral
veins 2 or 3 pairs; axils often with reduced stipule¬
like shoots. Inflorescences simple, shortly and dense¬
ly raceme-like, to 1.5 cm; peduncles 1-5 mm, pu¬
berulent; cymules 2-flowered. Pedicels to 5.5 mm,
pubescent. Sepals lanceolate, ca. 1.5 x 0.7 mm,
ciliate, minutely puberulent outside, strongly re¬
flexed at anthesis. Corollas ± erect, lobes oblong-
ovate, ca. 3.5 x 1.7 mm, slightly fleshy, glabrous.
Coronas a shallow ± membranous cup ca. half as
high as gynostegium, internal appendages absent.
Gynostegium ca. 2.5 mm high; anther appendages
ovate, acute. Follicles often solitary, lanceolate in
outline, ca. 6 x 1.5 cm.

Distribution. China in the provinces of Hubei
and Sichuan. There is no information on ecology.

The corona in Cynanchum brevicoronatum is so
short that at one point it was suspected to be a
growth abnormality, but the flowers seem uniform
and fully functional. Initially C. brevicoronatum was
confused with C. decipiens, which also has a short
corona, but closer examination shows that the co¬
rona is much shorter, membranous rather than fleshy,
and lacks the inner appendages seen in C. decipiens.
Another distinctive feature of C. brevicoronatum is
the reflexed sepals. It is probably most closely related
to species such as C. wallichii Wight and C. cal-
lialatum Hamilton ex Wight. Similar very short
coronas are seen in Biondia, which can easily be
distinguished by the much narrower leaves with pale
dots on the adaxial surface and the distinctly fleshy
corona.

Paratype. CHINA, t. Sichuan: Tchen-keou-tin,
Farges 92 bis (P).

Cynanchum duclouxii M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Yunnan: Yunnansen, 3
Aug. 1904, Ducloux 176 (holotype, E).

Species Cynancho kintungensi similis sed ab eo corolla
parva lobis erectis, corona brevioribus differt, ab speciebus
aliis appendicibus coronae parvis vel reductis plus minusve
minute longitudinaliter bicarinatis pilis caulium unifariis
distinguenda.

Twining herbs. Stems puberulent along 1 line.
Petioles 2.5—3.2 cm; leaf blades triangular-ovate,
4.5-8 x 1.7-5.1 cm, adaxially uniformly minutely
puberulent, abaxially slightly paler and glabrous ex¬
cept for puberulent veins, base deeply cordate, basal

lobes slightly enlarged, sometimes overlapping, apex
acute to slightly acuminate, basal veins 3-5, pedate,
lateral veins 2 or 3 pairs. Inflorescences raceme¬
like, sometimes terminal on short axillary branches;
peduncles 0.7-5.5 cm; rachis to 1.5 cm; cymules
2-flowered, in a lax spiral. Pedicels to 6 mm, pu¬
berulent. Sepals ovate, ca. 1 x 0.8 mm, sparsely
puberulent, apex acute. Corollas white, ± erect,
deeply divided, lobes 2.5-3 x 1-1.4 mm, glabrous
outside, minutely puberulent inside. Coronas mem¬
branous, prominently 5-lobed, tube ca. as high as
middle of anthers, shorter than lobes; lobes longer
than anther appendages, often inflexed, with 2 prom¬
inent adaxial gibbosities at base, rarely with a small
adaxial appendage. Anther appendages ovate, acute,
prominent. Follicles solitary, lanceolate in outline,
ca. 6.5 x 0.9 cm, base tapered, apex slightly acu¬
minate.

Distribution. China, Yunnan and Sichuan Prov¬
inces. Bushland in valleys; no indication of altitude.

Cynanchum duclouxii is related to C. wallichii
Wight, C. sinoracemosum M. G. Gilbert & P. T.
Li, and C. kintungense Tsiang. From these, C. du¬
clouxii is separated by its smaller flowers with erect
corolla lobes and its coronas, which are shorter than
the anthers. In C. wallichii the corolla lobes are
reflexed, whereas in C. kintungense they are spread¬
ing, and in both species the corona tube is longer
than that of C. duclouxii. The characters separating
C. duclouxii from C. sinoracemosum are listed un¬
der the latter.

Paratypes. CHINA. Sichuan: Tianquan Xian, Erlong
Mountain, 14 Aug. 1953, X. Y. Jiang 35246 (HNWP).
Yunnan: Yunnansen, E. E. Maire 1141 (E).

Cynanchum kingdonwardii M. G. Gilbert & P.
T. Li, sp. nov. TYPE: China. Yunnan: Yung¬
ning, 29 June 1922, Kingdon-Ward 5266 (ho¬
lotype, E).

Species Cynancho callialato similis sed ab eo laminis
foliaribus linearilanceolatis basibus cuneatis manifeste dis¬
tinguenda.

1 wining herbs. Internodes with poorly defined line
of pubescence. Petioles to 11 mm; leaf blades linear-
lanceolate, to at least 11 x 1.2 cm, adaxially thinly
puberulent, abaxially glabrous, base cuneate, apex
acute; lateral veins ca. 10 pairs. Inflorescences um¬
bel-like; peduncles to 3 mm, pubescent along 1 side.
Pedicels to 7 mm, puberulent. Sepals lanceolate, ca.
2 x 0.8 mm, thinly hairy. Corollas very pale green,
deeply divided, glabrous; lobes oblong-lanceolate, ca.
5 x 2.4 mm, blunt. Coronas white, cup-shaped, ca.
2.5 mm high, membranous, adnate to anthers, mar-

Volume 5, Number 1
1995

Gilbert et al.

Asclepiadaceae from China

7

gin with 5 low, acute teeth, smooth adaxially. Gy-
nostegium slightly higher than corona; anther ap¬
pendages narrowly ovate. Fruits not seen.

Distribution and ecology. China, Yunnan Prov¬
ince. On scrub-clad limestone slopes, in shade, at
3000-3300 m.

Cynanchum kingdonwardii resembles C. calli-
alatum and C. wallichii in having cup-shaped cor¬
onas with reduced or no internal appendages. It
differs from those and all related species by the
linear-lanceolate leaf blades with cuneate bases, in¬
stead of more or less ovate blades with deeply cor¬
date or sometimes rounded bases.

Cynanchum longipedunculatum M. G. Gilbert
& P. T. Li, sp. nov. TYPE: “Western China,”
12,000 ft. [3600 m], June 1904, Wilson 4103
(holotype, BM; isotype, P).

Species Cynancho decipienti affinis sed ab eo inflo-
rescentiis subcapitatis pedunculis longioribus corona long-
iore cupuliformi non carnosa diifert.

Twining herbs. Stems puberulent along 1 side.
Petioles ca. 2.5 cm, slender; leaf blades lanceolate,
ca. 5.6 X 2.3 cm, adaxially uniformly puberulent,
abaxially slightly glaucous, hairy on veins only, base
deeply cordate with incurved, often overlapping,
basal lobes, apex long-acuminate; basal veins usually
7, lateral veins 3 or 4 pairs. Inflorescences long
pedunculate, umbel-like, very congested; peduncles
6-9 cm, longer than subtending leaf, densely hairy
on 1 side; cymules usually all aggregated, sometimes
an isolated flower below main inflorescence. Pedicels
3.5-6 mm, puberulent on 1 side. Sepals oblong-
lanceolate, ca. 2 x 0.9 mm, puberulent, sometimes
only on margins. Corollas white (fide Wilson), ex¬
terior glabrous, interior densely pilose, tube ca. 0.5
mm; lobes ca. 5.5 x 1.7 mm, ± erect. Coronas
cup-shaped, longer than stigma head, not very fleshy,
tube longer than triangular marginal lobes, interior
with 5 internal appendages opposite to and nearly
as long as marginal lobes. Anther appendages ovate,
nearly acute, scarious except for narrow midrib,
higher than corona. Stigma head rounded. Fruits
not seen.

Distribution. Known only from the type collec¬
tion without precise locality or notes on ecology apart
from the unusually high altitude, 3600 m; most
likely collected in Sichuan Province.

Cynanchum longipedunculatum differs from the
related C. decipiens C. K. Schneider by having
subcapitate inflorescences up to 2 cm in diameter
and coronas longer than the gynostegium. Cynan¬

chum decipiens has umbel-like inflorescences 3-15
cm long and corona shorter than the gynostegium.

Cynanchum lysimachioides Tsiang & P. T. Li,
Acta Phytotax. Sin. 12: 89. 1974. TYPE: Chi¬
na. Yunnan: between Likiang, Youngnung &
Yungpei, en route to Muli, Rock 5317 (holo¬
type, NY; isotype, E).

Cynanchum likiangense W. T. Wang in Tsiang & P.
T. Li, Acta Phytotax. Sin. 12: 89. 1974. Syn. nov.
TYPE: China. Yunnan: Li-Kiang, 6 Aug. 1939, Y.
C. Chao 21216 (holotype, KUN).

Comparison of the types of Cynanchum lysi¬
machioides and C. likiangense shows that the sup¬
posed difference in habit, the one erect and the
other twining, is not real because the habit is basi¬
cally erect but with a distinct tendency toward twin¬
ing. Other supposed differences in density and length
of indumentum and flower size are well within the
range found in many other species and seem unlikely
to be of real significance.

Cynanchum megalanthum M. G. Gilbert & P.
T. Li, sp. nov. TYPE: China. Yunnan: 20 mi.
S of Yung-ning, 2700 m, 27 May 1921, King-
don-Ward 4103 (holotype, E).

Species Cynancho corymboso similis sed ab eo corolla
magna parce pubescenti, coronae lobulis marginalibus del-
tatis tubo semper longioribus distinguenda.

Robust twining herbs. Stems puberulent along 1
(or 2) lines. Petioles ca. 2.5 cm; leaf blades trian¬
gular-ovate, ca. 8.5 x 4.8 cm, thinly papery, abax¬
ially gray-green, minutely puberulent along veins,
adaxially minutely puberulent, base deeply cordate,
basal lobes rounded, not incurved, apex acute to
acuminate; basal veins 5, lateral veins ca. 3 pairs.
Inflorescences umbel-like or sometimes raceme-like
with many-flowered cymules; peduncles robust, ca.
5 cm, thinly puberulent. Pedicels 10-14 mm, pu¬
berulent along 1 side. Sepals lanceolate, ca. 5 x
1.4 mm, sparsely and minutely puberulent. Corollas
cream or pale greenish pink, deeply divided, rotate;
lobes linear-lanceolate, ca. 10 x 2.2 mm, very
sparsely hairy. Coronas cup-shaped, ca. 1 mm high,
with 5 triangular marginal lobes to 3 mm high and
small internal appendages. Anther appendages oval,
incurved over stigma head. Fruits and seeds not
seen.

Distribution and ecology. China (Yunnan Prov¬
ince), Myanmar. Along thicket margins; 2700 m.

Cynanchum megalanthum is superficially similar
to C. corymbosum but with the corolla much larger
and more sparsely hairy and with the corona tube

8

Novon

shorter than the well-defined triangular marginal
lobes. The internal corona appendages are always
small and sometimes more or less obsolete, sug¬
gesting a relationship also to C. callialatum and C.
duclouxii, which show comparable variation.

Paratypes. CHINA. Yunnan: To ngay pres Lou Pou
prefecture de Cong tibouray, June 1906, S. Ten in Du-
cloux 4143 (P). MYANMAR. Forrest 10327 (E).

Cynanchum pingshanicum M. G. Gilbert & P.
T. Li, sp. nov. TYPE: China. P’ing Chan, ca.
500 km S of Beijing, mountains SW of Nan
Yeli, Mei-huei-t’ouo, 5 June 1923, L. Chanet
& J. H. Serre A178 (holotype, P).

Species Cynancho acuminatifolio Hemsley affinis sed
ab eo paginis adaxialibus foliorum uniformiter puberulis,
lobis corollae angustioribus, inflorescentiis longioribus dif-
fert.

Erect herbs. Stems 30-60 cm high, ± tending
to twine at tip in larger plants; internodes at first
pubescent, soon glabrescent. Lower and uppermost
leaves reduced, other leaves larger; petioles 1-2
cm; leaf blades ovate to elliptic, 7.5-15 x 3.7-8.5
cm, adaxially uniformly puberulent, abaxially pu-
berulent along veins, base cuneate to ± truncate
and decurrent along petiole; veins 5-8 pairs. Inflo¬
rescences at upper nodes, up to 4 per node, up¬
permost usually longer than subtending leaf, irreg¬
ularly forked to ± umbel-like, many flowered; pe¬
duncles 1-6.5 cm, glabrescent. Pedicels 6-9 mm,
glabrous or sparsely puberulent. Sepals triangular-
lanceolate, ca. 2 x 0.6-0.8 mm, glabrous except
for ciliate margin. Corollas probably white, ca. 12
mm diam., glabrous; lobes oblong-triangular, ca. 6
x 2 mm, apex ± obtuse. Coronas fleshy, 5-lobed
to near base, slightly shorter than anthers; lobes
bluntly triangular, ± free from gynostegium, without
internal appendages. Fruits not seen.

Cynanchum pingshanicum , which is closely re¬
lated to C. acuminatifolium Hemsley and C. ja-
ponicum Morren & Decaisne, is known only from
five collections all made by Chanet and Serre be¬
tween 1923 and 1926. It differs from both of those
species by the uniformly puberulent adaxial leaf
surface, from C. acuminatifolium by the width of
corolla lobes (2 vs. 3 mm wide), and from C. ja-
ponicum by the shorter inflorescences.

Paratypes. CHINA. P’ing Chan, Nan ye li, 1 June
1926, L. Chanet & J. H. Serre A724 (P); P’ing Chan,
Pai lin t’ong, 4 June 1926, L. Chanet & J. H. Serre
A722 (P); no data, L. Chanet & J. H. Serre 15 (P); no
data, L. Chanet & J. H. Serre 1223 (P).

Cynanchum rockii M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Sichuan: Muli, Mutirong,
Muti Konka, snow range E of the Yalung, 3000
m, May-June 1932, Rock 23718 (holotype,
K; isotypes, BM, E, HUH).

Species Cynancho vincetoxico sensu lato affinis sed ab
eo corolla atrorubenti lobis angustatis erectis glabris tubo
interne lanato manifeste distinguenda.

Erect herbs. Stems 2, to 30 cm, branched, pu¬
bescent. Petioles 5-11 mm; leaf blades ovate, ca.
5x3 cm, adaxially sparsely pubescent, abaxially
densely puberulent along veins, base rounded to
broadly cuneate, apex acuminate; lateral veins 3 or
4 pairs. Inflorescences up to 7-flowered, slender,
often forked with umbel-like clusters of flowers, to
3.6 cm; peduncles ca. 1.8 cm. Pedicels ca. 6 mm.
Sepals lanceolate, ca. 2.2 x 0.5 mm. Corollas dark
red, tube ca. 2 mm, lobes erect, ca. 4 x 1.5 mm,
triangular-acuminate, apex blunt, glabrous except
the throat with woolly hairs. Corona lobes almost
free, ovate, higher than gynostegium, apex rounded,
adnate to anther, without internal appendage. Fruits
not seen.

Distribution and ecology. China, Sichuan. Al¬
pine meadow; 3000 m.

The type of Cynanchum rockii was cited by
Tsiang (1939) as C. muliense Tsiang. However, the
type of C. muliense is clearly a member of the
complex including C. vincetoxicum, C. inamoenum
(Maximowicz) Loesener, and C. forrestii Schlechter.
Cynanchum rockii is easily distinguished by the
dark red, narrowly lobed corollas, which are glabrous
except for the prominently woolly throat. It also
grows at a higher altitude than members of the C.
vincetoxicum complex.

Cynanchum sinoracemosum M. G. Gilbert &
P. T. Li, sp. nov. TYPE: China. Yunnan: Kou
ly Region of Pin tchouan, 1910, J. Py in F.
Ducloux 7653 (holotype, P).

Species nova Cynancho corymboso, C. wallichii , et
C. kintungensi similis sed ab eis internodiis uniformiter
puberulis, foliis angustioribus, lobis corollarum erectis gla¬
bris differt; etiam C. duclouxii similis sed ab eo floribus
majoribus foliis angustioribus basibus non dilatatis differt.

Twining herbs. Stems puberulent all round when
young, later along 1 or 2 sides only. Petioles 1.4-
3.5 cm, slender, puberulent along adaxial groove;
leaf blades lanceolate, 4.3-6(-8) x 1.5-2.5(-4.5)
cm, adaxially sparsely pubescent, abaxially paler and
more sparsely hairy, sometimes ± glabrous except
for veins, base deeply cordate, basal lobes incurved
and sometimes overlapping, apex long acute to

Volume 5, Number 1
1995

Gilbert et al.

Asclepiadaceae from China

9

slightly acuminate; basal veins 5-7, lateral veins 3
or 4 pairs; minute stipule-like axillary leaves present.
Inflorescences raceme-like, 2-4 cm, cymules 2-
flowered, in regular spiral; peduncles 0.4-2.5 cm,
puberulent. Pedicels 3-7 mm. Sepals ca. 1.3 x 0.7
mm, puberulent. Corollas white, erect, glabrous; lobes
oblong-lanceolate, 3.3-4.5 x 1.3-1.5 mm. Coronas
white, deeply cup-shaped, 2-3 mm high, concealing
gynostegium, plicate, marginal lobes 5, ± bluntly
triangular, to 0.4 mm, alternating with well-defined,
± appendage-like plicae. Anther appendages lan¬
ceolate, erect, white. Fruits not seen.

Distribution. China, Sichuan and Yunnan Prov¬
inces. Growing in hedges, at least at 1900 m.

Cynanchum sinoracemosum resembles C. du-
clouxii in having erect glabrous corollas and reduced
corona appendages. Cynanchum duclouxii has dis¬
tinctly smaller flowers with corona appendages re¬
duced to paired gibbosities, whereas C. sinorace¬
mosum has prominent longitudinal plicae and broad¬
er leaves often with enlarged basal lobes. Other
species with similar coronas and inflorescences (e.g.,
C. callialatum, C. corymbosum Wight, C. kintun-
gense, and C. wallichii ) differ by their spreading
to reflexed, mostly hairy, corollas and broader leaves.
In all these closely related species the internodes
have the hairs restricted to one or two narrow lines,
but in C. sinoracemosum the internodes are uni¬
formly hairy when young and do not develop the
well-defined lines of hairs when glabrescent.

Paratypes. CHINA. Sichuan: Kouy Tcheou, June
1914, J. Esquirol 5037 (P); Kouy Tcheou, near Gan-
pin, 29 Aug. 1897, E. Bodinier & L. Martin 1958 (P).
Yunnan: Yo lin chan, near Tong miy tcheou, 1911, F.
Ducloux (P); near Ou se tchong, 1 Aug. 1904, J. Ten
in F. Ducloux 2904 (P); Yunnan Fu, 1900 m, 30 July
1916, Schoch 255 (US).

Cynanchum triangulare M. G. Gilbert, W. D.
Stevens & P. T. Li, nom. nov. Replaced name:
Cynanchum deltoideum J. D. Hooker, FI. Brit.
India 4: 24. 1883, not Cynanchum deltoideum
Hance, Ann. Sci. Nat., ser. 5. 5: 228. 1866.
Vincetoxicum deltoideum [“deltodeum”] O.
Kuntze, Rev. Gen. 2: 424. 1891.1 YPE: India.
Khasia Mts.: Kalapanee and Moflong, 5000—
6000 ft.. Hooker & Thomson s.n. (holotype,
K; isotype, P).

western India. This leaves C. otophyllum as a Chi¬
nese endemic.

Heterostemma Wight & Arnott.

Heterostemma has about 30 species distributed
from India east through the tropics to northern
Australia and Samoa. It is represented in China by
nine species, of which four are endemic.

Heterostemma menghaiense (H. Zhu & H.
Wang) M. G. Gilbert & P. T. Li, comb, et slat,
nov. Basionym: Heterostemma villosum Cos-
tantin var. menghaiense H. Zhu & H. Wang,
Acta Bot. Yunnan. 16: 27, fig. 2. 1994. TYPE:
China. Yunnan: Xishuangbanna, Menghai,
Mengsong, 1000 m, 14 May 1989, Zhu H.
& Wang H. 2443 (holotype, HITBC not seen).

Heterostemma menghaiense was first collected
in 1938 by T. T. Yii (Yu 16307), and it is strange
that it remained undescribed for so many years. It
has possibly been confused with the Indochinese H.
villosum Costantin, which has a much less distinctive
indumentum. The uniformly densely long pilose in¬
dumentum is not matched by any other member of
Heterostemma. Several species have lines of similar
long hairs along the internodes, but the hairs on the
leaves and flowers of these other species are always
much finer and sparser than in H. menghaiense
and are often appressed. The large flowers of H.
menghaiense are also distinctive, matched best by
those of H. grandijlorum Costantin.

Second collection. CHINA. Yunnan: Shunning, T. T.
Yii 16307 (A, E).

Hoya R. Brown.

Hoy a is a genus of perhaps 100 species distrib¬
uted primarily in southeastern Asia into Oceania.
Seventeen of the 32 Chinese species are endemic.

Hoya chinghungensis (Tsiang & P. T. Li) M. G.
Gilbert, P. T. Li & W. D. Stevens, comb. nov.
Basionym: Dischidia chinghungensis Tsiang
& P. T. Li, Acta Phytotax. Sin. 12: 130. 1974.
TYPE: China. Yunnan: Che-li Hsien, Meng-
soong, Dah-Meng-Lung, 1900 m, Sep. 1936,
C. W. Wang 78311 (holotype, IBSC; isotype,
A).

Cynanchum triangulare has usually been dis¬
missed (e.g., Tsiang & Li, 1977) as a synonym of
C. otophyllum C. K. Schneider, but it differs from
that by the much better developed corona tube and
appears to be a good species restricted to north¬

The collections cited in the protologue of Dis¬
chidia chinghungensis are all in fruit, and flowers
were not known to the original authors. Flowering
collections are now available, and it is obvious that
this is a species of Hoya closely allied to H. Ian-

10

Novon

ceolata D. Don. Material of H. chinghungensis
collected in Thailand (at Tee-Lao-Su waterfall, Mae
Sod District, Tak Province, near the border with
Myanmar), which was not seen by us, along with a
good color photo on the cover, were discussed by
Thorut (1993) as “ Hoya bella subsp. nov.” How¬
ever, Thorut failed to make the connection with D.
chinghungensis, even though he cited all the Chi¬
nese collections in the protologue of that species.
There may well be a case for treating this and H.
bella W. J. Hooker as subspecies of H. lanceolata,
but this should only be done after a detailed study
of the complex as a whole, which is otherwise entirely
extra-Chinese.

Material seen. CHINA. Yunnan: Jinghong Xian, 1700
m, 5 June 1992, Tsi Zhanhuo 92-362 (MO). MYAN¬
MAR. Northern Triangle, Kingdon-Ward 21152 (E).

Hoya commutata M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Guangxi: Shap Man Taai
Shan, near Iu Shan village, SE of Shang-sze,
near border with Guangdong, W. T. Tsang
22375 (holotype, P, mixed with H. fusca).

Species nova a congeneris ramis fioriferis dense pu-
bescentibus foliis caducis, floribus purpureis, lobulis co-
ronae acutis gracilibus sinus corollae excedentibus distin-
guenda.

Habit not known, possibly a hanging epiphyte.
Stems pubescent. Leaves not known. Inflorescences
pubescent, ca. 30-flowered; peduncles ca. 2 cm.
Pedicels to 4 cm, slender, flexuous in dried speci¬
mens. Sepals triangular, ca. 2.5 x 0.9-1.3 mm,
densely pubescent. Corollas rotate with revolute
margins, “purple,” ca. 1.5 cm diam., densely pa¬
pillate-pubescent, limb ca. 0.8 cm wide. Corona
lobes acute, stellately spreading, extending beyond
sinus between corolla lobes; inner margins acute, ±
meeting in center. Fruits not seen.

The only piece of Hoya commutata seen from
China is a bare twig with a single inflorescence
included with a collection of H. fusca. Other sheets
of Tsang 22375 have only young fruiting material
of H. fusca. The two taxa can easily be separated
by the fact that H. commutata is more or less
uniformly pubescent, while H. fusca is glabrous.

The paratype is very similar to the holotype, again
a bare stem with leaves from another plant, this
time belonging to Alstonia (Apocynaceae). It differs
from the holotype only by the sparser indumentum.
The inflorescence and the long corona lobes, which
extend beyond the sinuses between corolla lobes, are
very similar indeed, though there is no note on the
color. It would seem that Hoya commutata flowers
on rather specialized hanging branches with cadu¬

cous leaves, most unusual in a genus normally noted
for its long-lived leaves.

Paratype. MYANMAR. Northern Triangle, Kingdon-
Ward 21276 (BM).

Hoya mekongensis M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Xizang/Yunnan; Tsekou
(Haut Mekong), 10 Oct. 1895, Biet in Soulie
1598 (holotype, P; isotypes, P 2 sheets).

Species nova Hoyae lii Burton similis sed ab ea ramulis
petiolisque non setosis inflorescentiis lateralibus differt.

Lianas climbing by adventitious roots. Stems mi¬
nutely puberulent, glabrescent. Petioles 1.2-1.6 cm,
minutely puberulent; leaf blades oblong-oblanceo-
late, 10-13 x 4-5 cm, minutely hairy, base round¬
ed, apex acuminate to caudate-acuminate; lateral
veins ca. 7, poorly defined. Pseudumbels extra-ax¬
illary, apparently flat topped, at least 8-flowered;
peduncles (0.3—)1 —2.5 cm, pubescent. Pedicels 2-
2.7 cm, sparsely hairy on 1 side. Sepals ovate, ca.
1.5 x 1.5 mm, sparsely hairy. Corollas white, sweet
scented, 1.5-1.7 cm diam., reflexed, lobes ca. 6 x
5 mm, minutely papillate with short-ciliate margins.
Corona lobes ± ovoid, erect, ca. 2 times as high as
gynostegium, dorsal grooves very narrow, inner tooth
incumbent on gynostegium, just meeting in center.
Anther margins short; appendages inconspicuous.
Fruits not seen.

Hoya mekongensis appears to be most closely
related to H. lii Burton, which has a similar reflexed
corolla and erect, rounded corona lobes. It differs
by having minutely puberulent stems and petioles
and extra-axillary inflorescences. In contrast, H. lii
has setose stems and petioles and terminal inflores¬
cences.

Jasminanthes Blume.

We believe that the genus Stephanotis is best
treated as a Malagasy endemic. Therefore, the fol¬
lowing new combinations are needed for the Flora
of China.

Jasminanthes chunii (Tsiang) W. D. Stevens &
P. T. Li, comb. nov. Basionym: Stephanotis
chunii Tsiang, Sunyatsenia 3: 165. 1936.
TYPE: China. Guangdong: Tung-Wu Shan, 6
May 1928, W. T. Chun 6417 (holotype, IBSC,
originally in SYS).

Jasminanthes mucronata (Blanco) W. D. Ste¬
vens & P. T. Li, comb. nov. Basionym: Apo-
cynum mucronatum Blanco, FI. Filip. 852.
1837. TYPE: from seed sent from China (ho¬
lotype, not preserved; see Merrill, 1918).

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1995

Gilbert et al.

Asclepiadaceae from China

11

Jasminanthes pilosa (Kerr) W. D. Stevens & P.
T. Li, comb. nov. Basionym: Stephanotis pi¬
losa Kerr, Bull. Misc. Inform. 1938: 453.
1938. TYPE: Thailand. Loi, Dan Sai, Kao Keo
Kang, 1300 m, Kerr 5769 (holotype, K).

Jasminanthes saxatilis (Tsiang & P. T. Li) W.
D. Stevens & P. T. Li, comb. nov. Basionym:
Stephanotis saxatilis Tsiang & P. T. Li, Acta
Phytotax. Sin. 12: 118. 1974. TYPE: China.
Yunnan: Foo-Ning, 1100 m, 3 May 1940, C.
W. Wang 89139 (holotype, KUN).

Lygisma J. D. Hooker.

A genus of 3(-6) species known from Malaysia,
Myanmar, Thailand, Vietnam, and China. One spe¬
cies in China.

Lygisma inflexum (Costantin) Kerr, Bull. Misc.
Inform. Kew 1939: 457. 1939. Basionym: Pi-
lostigma inflexum Costantin in Lecomte, FI.
Indo-Chine 4: 73. 1912. TYPE: Vietnam.
Cochinchine, Thorel s.n. (holotype, P).

Tylophora hainanensis Tsiang & P. T. Li, Acta Phytotax.
Sin. 10: 35. 1965. Syn. nov. TYPE: China. Hainan:
Po-Ting, 120 m, 1 Oct. 1957, H. D. Zhang 948
(holotype, HC-HSNU).

Lygisma is easily recognized by the small flowers
with very characteristic inflexed corolla lobes. The
correct generic position of the Chinese material had
been questioned since it was first collected. It was
placed in Tylophora, but with reservation. The type
of T. hainanensis is a very good match with the
type of Lygisma inflexum.

Marsdenia R. Brown.

Marsdenia includes about 100 species distributed
in America, Asia, and tropical Africa. Of the 25
native species in China, 14 are endemic.

Marsdenia braehyloba M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Yunnan: Feng Chen Lin
Mountain, A. Henry 11196 (holotype, K).

Species nova a speciebus aliis inflorescentiis umbelli-
formibus, corolla purpurea, lobulis coronarum brevibus
lateraliter compressis basibus profunde sulcatis distin-
guenda.

Lianas, glabrous except for sepal and corolla mar¬
gins. Nodes with distinct interpetiolar line. Petioles
2.5-4.5 cm; leaf blades elliptic, 10-14.5 x 5-7.5
cm, abaxially very pale, adaxially dark with paler

zone along midrib, base cuneate, apex acuminate;
lateral veins 5 or 6 pairs, raised on both sides.
Inflorescences umbel-like, up to 20-flowered; pe¬
duncles to 3 cm. Pedicels 1-1.7 cm. Sepals ±
circular, ca. 1.5 x 1.5 mm, margin ciliolate. Co¬
rollas bell-shaped with spreading lobes, purple; tube
ca. 2 mm, broader than long; lobes ca. 4x2 mm,
apex rounded. Corona lobes ca. as high as anthers,
laterally compressed with almost hooked tip, basal
half deeply grooved. Anther appendages oblong,
rather fleshy toward base, half as high as stigma
head. Stigma head ovoid with acuminate, divided
beak, exserted from corolla tube. Follicles solitary,
lanceolate in outline, ca. 16x4 cm, base narrowed
into a stalk 1 cm long. Seeds ovate, ca. 17 x 7
mm; coma off-white, to 3 cm.

Because of its somewhat reduced corona lobes,
material of Marsdenia braehyloba has usually been
filed under Gongronema. Each corona lobe is closely
appressed to the anther in the form of an inverted
“V” with the apex laterally compressed and some¬
what recurved and the lower part deeply grooved,
with the two legs of the “V” slightly spreading and
almost winglike. This contrasts very strongly with
the short, spreading, dorsiventrally compressed scales
inserted at the bases of the anthers that characterize
Gongronema. The conspicuously exserted stigma
head of M. braehyloba is also similar to that of
many other species of Marsdenia in the narrow
sense.

Paratype. CHINA. Yunnan: Feng Chen Lin Moun¬
tain, A. Henry 11073 (K).

Marsdenia tenii M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Yunnan: Siu tchoang, Sim¬
eon Ten 395 (holotype, E).

Species nova Marsdeniae hainanensi Tsiang similis
sed ab ea lamina foliari basi rotundata vel truncata, in¬
florescentiis longis simplicibus cymulos umbelliformes ger-
entibus differt.

Lianas, yellow-brown tomentose except for flow¬
ers. Petioles to 4 cm; leaf blades oblong-ovate, to
12.5 x 7.5 cm, sparsely hairy, base rounded to
truncate, apex acute; lateral veins ca. 5 pairs. In¬
florescences with several umbel-like cymules along
an unbranched rachis; peduncles to 3 cm; rachis to
at least 2 cm. Pedicels ca. 5 mm. Sepals elliptic,
ca. 3x2 cm. Corollas white, rotate, ca. 6 mm,
glabrous except for lobes and retrorsely pilose throat;
lobes ca. 3.5 x 2.5 mm, densely appressed-tomen-
tose in center. Corona lobes to base of anthers,
almost flat. Stigma head bifid-conical, equaling an¬
ther appendages. Fruits not seen.

12

Novon

Marsdenia tenii is similar to M. hainanensis,
M. tomentosa Morren & Decaisne, and M. sinensis
Hemsley, with which it shares ovate tomentose leaves.
It is easily separated from these by the inflorescence,
which has a long, unbranched rachis. The other
species have much-branched inflorescences that are
often wider than long.

Marsdenia tinctoria R. Brown, Mem. Wern. Soc.
1: 28. 1810. TYPE: Indonesia. Sumatra: “Tar-
am akkar Mars. Sumat. 78” (holotype, BM).

Marsdenia glubifera Tsiang, Sunyatsenia 3: 199. 1936.
Syn. nov. TYPE: China. Kwangtung: Yung Yuen,
S. K. Lau 24527 (isotype, P).

Marsdenia tinctoria var. brevis Costantin in Lecomte,
FI. Indo-Chine 4: 94. 1912. Syn. nov. SYNTYPES:
Laos, Phon-thane, Spire s.n. (P); Pon-pissay, Thorel
s.n. (P).

At first glance, the flowers of Marsdenia globi-
fera are very distinct, and it is difficult to compare
them with those of any other species. However, the
stems and leaves are so similar to those of the
common and widespread M. tinctoria that the two
plants have to be closely related. A closer exami¬
nation revealed that the flowers of M. globifera are
diseased, and their lower part is an undifferentiated
solid tissue. This is hinted at by the drawing of a
longitudinal section of a flower in the protologue.
Therefore, we have no hesitation in including this
diseased material in M. tinctoria.

Marsdenia yuei M. G. Gilbert & P. T. Li, sp. nov.
TYPE: China. Yunnan: Mienning, Taheching,
2300 m, T. T. Yil 17801 (holotype, E).

Species nova Marsdeniae koi Tsiang similis, sed ab ea
pedunculis brevioribus, corollis glabris parvilobis, lobulis
coronae tubo corollae longioribus differt.

Lianas, glabrous except for flowers. Stems stout.
Petioles 4 cm; leaf blades ovate, ca. 9.5 x 5.8 cm,
base shallowly cordate, apex bluntly cuspidate to
shortly acuminate, lateral veins 4 or 5 pairs, flat¬
tened. Inflorescences umbel-like, up to 9-flowered;
peduncles ca. 15 mm. Pedicels to 9 mm. Sepals
rounded, ca. 3 x 2.5 mm, finely appressed puber-
ulent. Corollas white, ± campanulate, glabrous ex¬
cept for sparsely ciliate margins; tube ca. 4.5-5 x
4 mm; lobes oblong-obovate, ca. 5.5 x 2.5-3.2
mm, apex rounded. Gynostegium ca. 6 mm high.
Corona lobes narrowly triangular, as long as anther
appendages, apex rounded, exserted from corolla
tube. Anther appendages ovate, membranous mar¬
gin relatively narrow. Stigma head hemispherical,
conspicuously exserted from anther appendages and
corolla tube. Fruits not seen.

We take pleasure in dedicating Marsdenia yuei
to the collector, one of the most highly regarded of
all Chinese plant taxonomists. The type collection
was cited as M. medogensis P. T. Li in the proto¬
logue of that species, which has much larger flowers
with corolla lobes ca. 9x9 mm, discoid stigma
head only slightly exserted from the anther ap¬
pendages, and oblong leaf blades more than four
times as long as broad with rounded base and acute
apex. Vegetatively M. yuei seems more closely re¬
lated to M. koi Merrill than to M. medogensis.
Marsdenia koi has much longer pedicels, larger
flowers, and corona lobes shorter than the anthers
and corolla tube.

Merrillanthus Chun & Tsiang.

The original account of this monotypic genus
described the pollinia as pendulous, which would
make it a member of the Asclepiadeae. The general
morphology, especially the form of the corona lobes,
suggested that this might not be correct, but most
material available had flowers too poorly preserved
to check the pollinia. An isoparatype sheet, Lau
3550 (P), has flowers well enough preserved to
examine the corona in more detail. This proved to
be extremely Tylophora-like in all respects, includ¬
ing the horizontal pollinia. Tylophora augustiniana
(Hemsley) Craib is particularly similar, differing only
by the much smaller flowers. Merrillanthus can be
easily separated from known species of Tylophora
by the large fruit with a thick fibrous mesocarp, in
contrast to the very thin, often almost papery follicle
walls of all species of Tylophora familiar to us. Thus
Merrillanthus is kept, at least until Tylophora as
a whole is revised. However, the fruits of I. au¬
gustiniana have not been described, and Marsden¬
ia, even in the narrow sense adopted for the Flora
of China, shows a comparable range of fruit types.
A collection from Cambodia, M. A. Martin 310 ( P),
is almost certainly M. hainanensis, which would
mean that Merrillanthus is no longer endemic to
China.

Sichuania alterniloba M. G. Gilbert & P. T. Li,
gen. et sp. nov. TYPE: China. Sichuan: S
Wushan, A. Henry 7262 (holotype, E).

Genus et species nova Holostemmate R. Brown similis
sed ab eo inflorescencentiis racemiformibus, floribus mi-
noribus, lobulis coronae liberis et antheris alternantibus,
antheris pallidis lucentibus marginibus longis dorsaliter
sulcatis appendicibus prominentibus scariis manifeste dif¬
fert.

Robust twining herbs. Stems minutely puberulent
along 1 side (sometimes along 2 at base of internode).

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Gilbert et al.

Asclepiadaceae from China

13

Petioles 2—3 cm; leaf blades ovate-triangular, 5-8
x 3.3-7 cm, abaxially pale, adaxially much darker
and apparently glossy, base cordate, thick, glabrous
throughout; lateral veins 4-6 pairs. Inflorescences
raceme-like, with few-flowered cymes arranged in
lax spiral along rachis; peduncles 1.5-5.5 cm, some¬
times sharply reflexed; rachis up to 2 cm; bracteoles
few, ca. 1 mm. Pedicels up to 1 cm, puberulent
along 1 side. Sepals lanceolate, ca. 2.7 x 1.3 mm,
apex acute, almost glabrous. Corollas very shallowly
bowl-shaped, almost rotate, white, glabrous; lobes
ovate, ca. 5.5 x 3.5 mm. Corona lobes separate,
alternating with anthers, ovate to ± semicircular,
short and inconspicuous. Gynostegium ca. 2.5 mm
high; anthers ca. 2 mm, uniformly pale and glossy
with long margins and dorsal groove; anther ap¬
pendages ovate, scarious except for narrow midrib.
Retinaculum ca. 0.5 mm; translator arms short and
rigid; pollinia cylindrical-ovoid, slightly compressed,
ca. 0.6 mm. Stigma head apiculate, concealed by
anther appendages. Fruits not seen.

Distribution. Endemic to China, Sichuan Prov¬
ince (South Wushan and Tianquan Xian).

The generic position of Sichuania alterniloba is
so questionable that it seems best to place it in a
genus of its own. The original collection had been
named as Holostemma, a view supported by the
presence of an interpetiolar line, fairly large, hard-
textured anthers more or less decurrent at the base
to the corolla, pendent pollinia, and fairly thin-tex-
tured corona. However, the distinctive corona with
separate, spreading, ovate scalelike lobes alternating
with the anthers (i.e., opposite the corolla lobes) is
quite different from the annular corona of Holos¬
temma and from that of any other genus known to
us. Other genera such as Gongronema have free,
scalelike corona lobes inserted at the base of the
gynostegium. In all taxa with similar scalelike lobes
known to us, these are opposite the anthers. Rou-
linia parviflora Decaisne (Cynanchum contrape-
talum E. Sundell; see Sundell, 1981) is described
as having corona lobes opposite the corolla lobes,
but these are erect and grooved. Cynanchum con-
trapetalum is apparently in all other ways typical
of Cynanchum subg. Mellichampia and thus is un¬
related to Sichuania. The anther form of Sichuania
is also different from that of Holostemma in that
the anther tips are more or less retuse with very
sharply delineated scarious appendages, not acu¬
minate with poorly differentiated inconspicuous ap¬
pendages.

Paratype. CHINA. Sichuan: Tianquan Xian, 1000
m, 11 Sep. 1963, West of Sichuan Expedition Group
3334 (HNWP).

Tylophora R. Brown.

Tylophora is in need of an overall revision. Most
African material has never been named. The dis¬
tinction from allied genera such as Belostemma
Wallich ex Wight, Merrillanthus, and Pentastelma
Tsiang & P. T. Li needs closer investigation. Some
of the species from tropical Asia look very different
from typical members of the genus and equally
deserve investigation.

The separation between Cynanchum and Tylo¬
phora on floral morphology is sometimes difficult.
It is often hard to interpret the orientation of the
pollinia in the smaller-flowered asclepiads with only
slightly elongated pollinia, especially in dried ma¬
terial. In most cases the characteristic erect corona
lobes clearly inserted on the anthers of most species
of Tylophora is diagnostic. However, there are some
species, such as T. glabra Costantin, otherwise typ¬
ical of the genus, in which the corona lobes are
almost free from the anthers, spreading and laterally
joined at the base to form a very Cynanchum -like
structure. Several non-floral characters often enable
the confident recognition of virtually all species of
Tylophora, at least in mainland Asia. These include
often elongated, zigzag, inflorescence rachises, wide¬
ly spaced cymules frequently elongated with age,
very narrow bracts, threadlike pedicels, and often
paired, widely divaricate fruits. None of these char¬
acters is of itself diagnostic, but in combination they
form a distinctive facies that leaves little doubt in
the recognition of Tylophora.

Tylophora costantiniana (Tsiang) M. G. Gilbert,
W. D. Stevens & P. T. Li, comb. nov. Bas-
ionym: Cynanchum costantinianum Tsiang,
Sunyatsenia 4: 119. 1939, new name for Cy¬
nanchum hirsutum Costantin, in Lecomte, FI.
Gen. Indo-Chine 4: 66, fig. 10 (12-13). 1912,
not C. hirsutum Vahl, Eclog. Amer. 2: 24.
1798, nor Tylophora hirsuta (Wallich) Wight,
Contr. Bot. India 49. 1834. TYPE: Vietnam.
Tonkin, Mekong, Thorel s.n. (holotype, P).

The type material of Cynanchum hirsutum Cos¬
tantin is mostly in fruit. However, one stem has an
inflorescence with buds developed sufficiently to con¬
firm that this is a Tylophora closely related to 7!
tsiangii.

Tylophora forrestii M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Yunnan: hills NW of
Tiugyueh(?) (25°30'N, 98°25'E), 2100 m. May
1931, Forrest 29601 (holotype, E).

14

Novon

Species Tylophorae ovatae (Lindley) Hooker ex Steu-
del similis sed ab ea internodiis glabrescentibus, foliis mem-
branaceis, inflorescentiis longioribus cymulas umbelli-
formes gerentibus, floribus majoribus differt; a T. brownii
foliis membranaceis basi cuneatis vel rotundatis differt.

Lianas. Stems uniformly pubescent when young
but glabrescent and soon glabrous or pubescent along
1 side only. Petioles to 2 cm; leaf blades ovate-
elliptic, to 9.4 x 4.2 cm, membranous, base broadly
cuneate to ± rounded, apex acuminate; lateral veins
4 or 5 pairs, prominently pilose when young, gla¬
brescent. Cymes to 8 cm; peduncles to 4.5 cm,
pubescent, cymules ± umbel-like, ca. 6-flowered;
bracts linear, ca. 1.5 mm. Pedicels to 15 mm. Sepals
linear-lanceolate, ca. 3 x 0.4 mm, apex acute,
margins prominently ciliate. Corollas yellow-green,
tube ca. 0.7 mm, lobes ca. 6 x 1.6 m wide, glabrous.
Corona lobes narrowly ovoid, appressed to gynos-
tegium and reaching base of anthers, ± acute. Gy-
nostegium ca. 2 mm high; anther appendages ovate,
erect. Corpusculum longer than ellipsoid pollinia.
Fruits not seen.

The indumentum suggests an affinity to Tylo-
phora ovata, but T. forrestii is a more vigorous
plant with softer leaves, larger flowers, more strictly
umbel-like cymules, and readily glabrescent stems.

Tylophora glabra Costantin in Lecomte, FI. Indo-
Chine 4: 109. 1912. TYPES: Vietnam. Tonkin
vers Ninh-Binh, Bon s.n. (syntype, P); vers
Ouonbi, au nord de Guang-yen, Balansa 609
(syntypes, P, K); Sept Pagodes, Mouret s.n.
(syntype, P).

Tylophora longipedicellata Tsiang & P. T. Li, Act.
Phytotax. Sin. 12: 136. 1974. Syn. nov. TYPE:
China. Hainan: Lung-Kiang, 15 Sep. 1933, C. H.
Tsoong 545 (holotype, IBSC).

Tylophora renchangii Tsiang, Sunyatsenia 3: 232. 1936.
Syn. nov. TYPE: China. Guangxi (“Kwangsi”): Tien-
chen, N of Sup-man-ta Shan, 900 m, 13 Oct. 1928,
R. C. Ching 7821 (holotype, IBSC; isotype, A).

Tsiang (1936) first drew attention to the unusual
corona of this species in his protologue for Tylo¬
phora renchangii. The corona lobes are almost
Heterostemma- like and distinct from any other Ty¬
lophora seen from mainland Asia. Typical Tylo¬
phora corona lobes are closely appressed to the
stamens, dorsally gibbous, and have an acuminate
tip. In T. glabra the corona lobes are more or less
circular, inserted at the base of the anthers, and
spread at an angle of about 45° to them, while the
acuminate tip is often strongly reflexed. Superficially
these are so different that it is difficult to accept

this species in Tylophora, but all other features of
T. glabra are typical of that genus.

The status of Tylophora longipedicellata de¬
serves more investigation. Most material from Hai¬
nan has the broad spreading corona lobes with a
blunt inflexed tip characteristic of T. glabra. Other
collections have short erect corona lobes with an
erect acuminate tip, as illustrated in the protologue
of T. longipedicellata. In other features, the two
species are nearly identical. Henry 8275 (K) has
corona lobes somewhat intermediate between those
of the two species. Therefore, we tentatively reduce
T. longipedicellata to synonymy of T. glabra.

Tylophora oligophylla (Tsiang) M. G. Gilbert,
W. D. Stevens & P. T. Li, comb. nov. Bas-
ionym: Absolmsia oligophylla Tsiang, Suny¬
atsenia 6: 121. 1941. TYPE: China. Yunnan:
Che-li Hsien, Aug. 1936, C. If . Wang 75455
(holotype not seen; isotype, PE).

The generic placement of Absolmsia oligophylla
is uncertain. It differs profoundly from the type and
only other species of Absolmsia, A. spartioides
(Bentham) Kuntze, which is a very specialized rel¬
ative of Hoya, known only from rot holes in large
trees in Borneo. Absolmsia spartioides has fleshy
leaves when young but then depends on persistent,
long, stiff, photosynthetic peduncles, which seem to
take over the function of leaves. A sheet labeled in
Tsiang’s handwriting as the holotype of A. oligo¬
phylla, deposited in the herbarium of the Fan Me¬
morial Institute of Biology and now housed in the
Institute of Systematic Botany, Beijing (PE), consists
of the lower part of a single stem and completely
lacks any trace of inflorescences and flowers. A
photograph of a different sheet showing the upper
parts was reproduced in the protologue. A search
for an isotype at the South China Institute of Botany
was not successful. The material seen appears to
come from a prostrate plant with no sign of the
stem’s twining and with soil adhering to the under¬
sides of the leaves, very like Tylophora rotundifolia
Hamilton ex Wight, which also has similar almost
circular leaves. The illustration of the flowers ac¬
companying the protologue, particularly the small
gynostegium, is also suggestive of a Tylophora. The
spreading corona lobes would be unusual for Tylo¬
phora, but T. glabra Costantin has even larger
spreading lobes that are at least superficially similar.
The description of pollinia as pendulous, if correct,
would rule out a placement in Tylophora, but, as
discussed above, Merrillanthus was described as
having pendulous pollinia whereas examination of
material cited in the protologue showed the pollinia

Volume 5, Number 1
1995

Gilbert et al.

Asclepiadaceae from China

15

to be horizontal. The large, stiffly papery leaves and
indumentum of short stiff hairs of 71 oligophylla
are distinctive, and it should be easy to make a
positive match with new material, should this become
available.

Tylophora rockii M. G. Gilbert & P. T. Li, sp.
nov. TYPE: China. Sichuan: Muti Konka, 3300
m. Rock 23717 (holotype, E).

Species nova Tylophorae yunnanensi Schlechter sim-
ilis sed ab ea inflorescentiis sessilibus, floribus majoribus,
capite stigmatico elevato differt.

Erect herbs to 70 cm. Stems little branched,
tending to twine at tips, uniformly puberulent. Pet¬
ioles ca. 15 mm; leaf blades oblong-oblanceolate, to
9.5 x 4 cm, sparsely puberulent, uppermost and
lowermost reduced, base cuneate to ± rounded,
apex rounded-apiculate, margins ± undulate, veins
4 or 5 pairs. Inflorescences extra-axillary from up¬
per nodes, to 8 cm, mostly ± sessile with long first
internode, cymules shortly and densely raceme-like
to umbel-like. Pedicels to 1(—2) cm. Sepals linear-
triangular, ca. 3 mm. Corollas purplish, stiffly rotate,
8-10 mm diam.; lobes oblong, 5(-7) x 2.5-3 mm,
apex rounded; interior densely puberulent. Corona
lobes reaching base of anther appendage, ± acute.
Gynostegium ca. 1.5 mm high; anther appendages
broader than long. Stigma head broadly dome-shaped
with light-colored center. Fruits not seen.

Tylophora rockii is most closely related to T.
yunnanensis, which has distinctly smaller flowers,
sepals ca. 1.5 mm long, and corolla lobes 3-4 x
1.5-2 mm. Tylophora rockii has mainly sessile
inflorescences with long first internodes, whereas T.
yunnanensis has peduncles to 6 cm long. Tylophora
rockii has a broadly dome-shaped stigma head in
contrast to the depressed stigma head of T. yun¬
nanensis.

Tylophora tsiangii (P. T. Li) M. G. Gilbert, W.
D. Stevens & P. T. Li, comb. nov. Basionym:
Cynanchum tsiangii P. T. Li, Bull. Bot. Lab.
North-E. Forest. Inst. 3(1): 103. 1983. TYPE:
China. Guizhou: Xing-yi, 1300 m, 7 Oct. 1960,
Y. T. Chang et al. 6880 (holotype, SCBI).

The hispid indumentum, brownish coloring of the
dried leaves, and inflorescence architecture (a long
zigzag rachis with elongated raceme-like cymules
and linear bracts) of Tylophora tsiangii are all very
similar to those of Tylophora ovata (Lindley) Hook¬
er ex Steudel. The short, elliptic corona lobes adnate
to and clearly shorter than the anthers are typical
for Tylophora. Though placed in different parts of

the family, the distinction in floral morphology be¬
tween these two genera can be very subtle. Tylo¬
phora tsiangii is clearly related to the Indochinese
T. costantiniana, which is very similar in habit and
was also misplaced within Cynanchum.

Tylophora tuberculata M. G. Gilbert & P. T.
Li, sp. nov. TYPE: China. Yunnan: Kouy-
Tcheou, Echang loing, 800 m. May 1913,
Esquirol 4377A (holotype, P, mixed with Cy¬
nanchum corymbosum Wight).

Species nova Tylophorae tenui Blume similis sed ab
ea et speciebus similibus caulium indumento ex trichom-
atibus erectis crassisque constante, foliis adaxaliter tub-
erculatis, lobulis coronae basaliter connatis, dorsaliter di-
latatis umbonem subpatentem truncatum vel leviter emar-
ginatum formantibus manifeste distinguenda.

Lianas. Stems slender, short pilose along 2 sides.
Petioles 5-10 mm, adaxially hairy, glabrescent; leaf
blades lanceolate, 2.5-4 x 1.1-1.2 cm, thick pa¬
pery, base rounded-cordate, apex acute, glabrous,
adaxially with prominent scattered tubercles. Inflo¬
rescences to 4 cm, much branched; peduncles to 6
mm, intemodes to 14 mm. Pedicels 4-6 mm, gla¬
brous. Sepals lanceolate, ca. 0.7 mm, tip recurved,
acute, glabrous. Corollas rotate to shallowly bowl¬
shaped, ca. 5 mm diam., greenish white, glabrous
outside, densely short pilose inside; lobes ovate, ca.
2 x 1.5 mm. Corona lobes joined at base, reaching
base of anther, with very prominent spreading, trun¬
cate or slightly emarginate, dorsal projection. Anther
appendages short, rounded; pollinia ellipsoid, almost
vertical. Stigma head rounded. Old follicles linear
lanceolate in outline, ca. 6.5 x 0.5 cm.

Tylophora tuberculata resembles 7. tenuis Blume
in having small lanceolate leaves and delicate much-
branched inflorescences. It differs from that and all
similar species of Tylophora by the thick stiffly erect
hairs forming two lines down the internodes of young
stems and along the margins of the petioles, the
well-defined small tubercles scattered adaxially on
leaf blades, the inflorescence with the peduncle clearly
shorter than the intemodes of the branched rachis,
and the distinctive corona lobes, which are enlarged
dorsally into a more or less spreading, fleshy, trun¬
cate to slightly emarginate projection and joined
laterally at the base by a thin flange. The corona
can be interpreted as forming a link between the
corona lobes of typical species of Tylophora, in
which the lobes are only slightly swollen dorsally,
and the corona lobes of 71 glabra, in which the
dorsal swelling seems to be grossly expanded into a
flat disc. The only sheet seen also has a fruit of
Cynanchum corymbosum Wight mounted on it.

16

Novon

Tylophora uncinata M. G. Gilbert & P. T. Li,
sp. nov. TYPE: China. Hainan: Po-ting, How
73159 (holotype, IBSC; isotypes. A, IBSC).

Species nova Tylophorae ovatae (Lindley) Hooker ex
Steudel affinis sed ab ea lamina foliari adaxialiter glabra
abaxialiter trichomatibus apice uncinatis basi papillosis
manifeste differt.

Lianas. Stems glabrescent. Petioles ca. 1.5 cm;
leaf blades ovate, 3.5-9 x 1.3-5.5 cm, stiffly pa¬
pery, adaxially glabrous, abaxially uniformly pu¬
bescent, hairs with papillate bases and strongly
hooked tips, base cordate, sinus usually deep and
narrow, apex ± acute, apiculate. Cyme extra-ax¬
illary; peduncles to 2 cm, rachis zigzag, sometimes
± absent, cymules densely raceme-like; bracts lin¬
ear-lanceolate, to 1.5 mm. Pedicels 6-8 mm. Sepals
ca. 1.7 x 0.5-0.7 mm, oblong-lanceolate, apex
acute, with few to many hooked hairs. Corolla pale
green at apex, dull green at base, 3.5-5 mm; tube
ca. 0.5 mm; lobes ovate to ovate-oblong, 2.5-3.5
x 1.2-1.3 mm. Corona lobes ovate, ca. 0.5 x 0.2
mm, pouched, apex obtuse, covering base of anthers.
Anther appendage rounded; pollinia globose, hori¬
zontal. Stigma head ± flat, 5-angled, depressed in
center. Follicles linear-lanceolate in outline, ca. 5
x 0.6 cm, apex acuminate. Flowering August-Sep-
tember.

Tylophora uncinata had been named mostly as
T. ovata (Lindley) Hooker ex Steudel var. brownii
(Hayata) Tsiang & P. T. Li, but closer examination
has shown that it differs from all species of Tylo¬
phora known to us by the very distinctive hooked

hairs on the underside of the leaves. Tylophora
uncinata differs further from otherwise similar spe¬
cies such as T. ovata by the more papery, adaxially
glabrous leaves.

Paratypes. CHINA. Guangdong: Gao-zhou, 4 Apr.
1929, K Tsiang 2088B (IBSC). Guangxi: Shang-si, 2
Aug. 1951, W. C. Ko 901 (CANT); without locality, Nov.
1889, A. Henry 8105 (K, P).

Acknowledgments. Thanks are due to Roy Ger-
eau for correcting the Latin diagnoses, to Marshall
R. Crosby, Ihsan Al-Shehbaz, Amy McPherson, and
Diana Gunter for facilitating the production of this
paper in many ways, and to the directors of the
following institutions for help during visits and/or
loan of material: E, HNWP, IBSC, K, P, PE, S,
SYS, UPS, US. The Stanley Smith Horticultural
Trust is thanked for the financial support of Gilbert.

Literature Cited

Merrill, E. D. 1918. Species Blancoanae. Publ. Dept.

Agric. Nat. Resour. Manila 12: 1-423.

Sundell, E. 1981. The New World species of Cynan-
chum , subgen. Mellichampia (Asclepiadaceae). Evol.
Monogr. 5: 1-63.

Thorut, C. 1993. A very special Hoya. The Hoyan 15:
29-30.

Tsiang, Y. 1936. Notes on the Asiatic Apocynales III.
Sunyatsenia 3: 121-239.

-. 1939. An enumeration of the Asclepiadaceae

plants collected by Tsai and Wang from Yunnan in
1933-1936. Sunyatsenia 4: 95-130.

- & P.-T. Li. 1977. Asclepiadaceae. In: FI.

Reipubl. Popularis Sin. 63: 249-575.

Nomenclatural Notes on the Boraginaceae of China

Chu Ge-lin

Institute of Botany, Northwest Normal University, Lanzhou, Gansu 730070,
People’s Republic of China

Rudolf Kamelin

Herbarium, V. L. Komarov Botanical Institute, Russian Academy of Sciences,
Prof. Popov Street 2, 197376 St. Petersburg, Russia

Robert R. Mill

Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, U.K.
Michael G. Gilbert

Flora of China Project, Missouri Botanical Garden, c/o Department of Botany,
Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ABSTRACT. Three new combinations, Tournefortia
sibirica L. var. angustior (A. DC.) G. L. Chu &
M. G. Gilbert, Lappula ferganensis (Popov) Ka¬
melin & G. L. Chu, and L. lasiocarpa (W. T. Wang)
Kamelin & G. L. Chu, plus a new name, Trigonotis
harrysmithii R. R. Mill, are validated.

Two precursory papers on the Boraginaceae for
the Flora of China have already been published
(Kamelin, 1993; Riedl, 1994). The following ad¬
ditional nomenclatural adjustments are herein pro¬
posed to make the names available for the forth¬
coming Volume 16 of the Flora of China.

Tournefortia sibirica L. var. angustior (A. DC.)
G. L. Chu & M. G. Gilbert, comb. nov. Bas-
ionym: Tournefortia arguzia Roemer &
Schultes p angustior A. DC., Prodr. 9: 514.
J845. SYNTYPES: China. Nei Mongol (“Chi¬
nese Mongolia”), Turczaninow s.n., Bunge s.n.;
Kazakhstan. E shore of Caspian Sea, Karelin
s.n. and Hohenacker s.n.; Russia. Astrakan,
Bieberstein s.n.; Turkey (“Asia Minor”),
Aucher-Eloy s.n.; Ukraine. S Podolia, Tur¬
czaninow s.n. (All syntypes in G, not seen.)

The new combination is needed because the va¬
riety does not have a valid combination in Tour¬
nefortia sibirica. A lectotype should be designated
from de Candolle’s original material at G. Such
material has not been examined by us.

Trigonotis harrysmithii R. R. Mill, nom. nov.
Replaced name: Trigonotis smithii W. T.
Wang, Bull. Bot. Res., Harbin 13(1): 3. 1993,
not Trigonotis smithii Banerjee, Bull. Bot. Surv.
India 8; 325. 1966 (1967). TYPE: China.
Sichuan: Sungpan, 3200 m, 14 July 1922,
Harry Smith 2775 (holotype, UPS; isotype,
PE).'

The earlier name Trigonotis smithii Banerjee,
which applies to a species from Bhutan and the east
Himalaya so far not recorded from China, was named
after William Wright Smith, the collector of the
type specimen and a former Regius Keeper of the
Royal Botanic Garden, Edinburgh. The later hom¬
onym was dedicated to the Swedish botanist Harry
Smith, an expert on the Gentianaceae who collected
in China.

There has been a reappraisal of the characters
dividing Lepechiniella from Lappula. Lappula is
readily distinguished from Lepechiniella in having
styles exserted from the nutlets instead of concealed
by them. No Chinese material belongs to Lepechi¬
niella as thus defined, which makes necessary the
following generic transfers.

Lappula ferganensis (Popov) Kamelin & G. L.
Chu, comb. nov. Basionym: Lepechiniella fer¬
ganensis Popov, FI. USSR 19: 715. 1953.
TYPE: [Tajikistan]. Alai Valley, Mount Kultz-
cha, 26 June 1931, Lipschitz 248 (holotype,
MW).

Novon 5: 17-18. 1995.

18

Novon

Lappula platyptera C. J. Wang, Bull. Bot. Res., Harbin
1(4): 91. 1981. Syn. nov. TYPE: China. Xinjiang:
Wuqia Xian, 3300 m, 4 Aug. 1978, Xinjiang Ex-
pediton 1911 (holotype, WUK).

Lappula lasiocarpa (W. T. Wang) Kamelin &
G. L. Chu, comb. nov. Basionym: Lepechi-
niella lasiocarpa W. T. Wang, Bull. Bot. Res.,
Harbin 4(2): 7. 1984. TYPE: China. Xinjiang:
Fuhai, 21 May 1959, Y. F. Wang s.n. (holo¬
type, PE).

Lepechiniella balchaschensis M. Popov, Spisok. Rast.
Gerb. FI. SSSR 12: 50. 1953. Syn. nov. Lappula
betpakdalensis Nabiev, Opred. Rast. Sred. Azii 8:
138. 1986, not Lappula balchaschensis M. Popov
ex N. Pavl., Zhurn. Bot. U.R.S.S. 30(4): 190. 1945.
TYPE: Kazakhstan. Lake Balchasch, southern shore
at Bertys Bay, 15 June 1934, M. Popov s.n. (ho¬
lotype, LE).

Lepechiniella balchaschensis is the oldest name
for this species, but the epithet is not available in
Lappula because Popov had already described Lap¬
pula balchaschensis based on a different type. The
transfer to Lappula was made in 1986 by Nabiev,
who introduced the replacement epithet betpak¬
dalensis. However, Lepechiniella lasiocarpa W.
T. Wang is the older name that should be used in
Lappula.

Literature Cited

Kamelin, R. 1993. Nomenclatural adjustments in Chi¬
nese Boraginaceae. Novon 3: 263.

Riedl, H. 1994. A new species and new combinations
of Boraginaceae from China. Novon 4: 46-47.

Onosma apiculatum (Boraginaceae), a New Species from China

Harald Riedl

Botanische Abteilung, Naturhistorisches Museum, Postfach 417, Burgring 7, A-1014 Wien,

Austria

ABSTRACT. Onosma apiculatum from Xinjiang,
China, is described. Its relationship to the three
species of series Tianschanica is discussed, and its
distinguishing characters from these are given.

Onosma apiculatum H. Riedl, sp. nov. TYPE:
China. Xinjiang: Zhao Moa Hong Qi, 2nd wood¬
cutting grounds, 2100 m, 9 June 1965, Cheo
Tai-yien et al. 650713 (holotype, SZ; isotype,
SZ; photos, W).

Herba perennis, foliis rosulae linearibus vel lineari-
subspathulatis, basin versus petiolatim attenuatis, 4-13
cm longis, 4-8 mm latis; foliis caulinis inferioribus bas-
alibus similibus basin versus paulo tantum attenuatis, su-
perioribus prope basin latissimis manifeste brevioribus,
omnibus viridibus, supra setis patentibus densis tuberculis
albis vix elevatis e circulis 2 vel 3 cellularum compositis
insidentibus et papillis nonnullis minutis, infra setis im¬
primis in nervo mediano densis, ceterum laxis pilisque
brevibus numerosis densissimis vestitis, cymis singulis vel
geminatis, calyce ca. 1.5 cm longo, laciniis linearibus,
subobtusis, usque ad basin liberis, corolla 2-2.2 cm longa,
compressa 8 mm fere lata, extra minutissime papillosa,
intus glabra, antheris tota longitudine cohaerentibus apice
sterili vix 1 mm longe exsertis; filamentis antheris paulo
brevioribus, in parte tertia inferiore corollae insertis, stylo
corollam 2.5-3 mm superante stigmate bilobo insignis.

Herbs perennial, with a single flowering stem and
sterile rosette of leaves arising from either a stout
vertical rhizome or from its ± horizontal branches.
Stem lateral, ascending, 20-35 cm high, un¬
branched, straw-colored when dry, densely covered
by a mixture of white patent bristles ca. 2 mm long
and very short patent hairs. Leaves of the sterile
rosette 4-13 cm long, 4-8 mm wide, linear-spat-
ulate to linear oblanceolate, tapering into a petiole¬
like base, revolute at margin, acute to subobtuse at
apex, green, densely covered adaxially with patent
to subappressed white bristles arising from a white,
nearly flat tubercle composed of 2 or 3 circles of
roundish to radially elongated, strongly convex cells
and numerous minute papillae, abaxially covered
with similar bristles mainly along the midrib and
only sparsely on other parts, with dense short hairs
as those on stem; lower stem leaves similar to rosette
leaves in size and shape but only a little narrowed
toward base; uppermost leaves much shorter, 2.5-
4 cm long, widest near base, acute, bristles and hairs
as in rosette leaves. Cymes single or paired, involute,
4-6 cm long in flower; lower bracts similar to upper

stem leaves, to 2 cm long, broad at base, decreasing
in size upward. Lower pedicels to 5 mm long, upper
ones to 2 mm. Calyx ca. 1.5 cm long in flower,
divided to base into 5 linear, subobtuse lobes densely
covered with fairly soft, appressed or subappressed
bristles and short hairs, elongated soon after flow¬
ering. Corolla pale yellow, turning brown after flow¬
ering and when dry at least in apical part, club-
shaped, 2-2.2 cm long, ca. 8 mm wide just below
lobes, ca. 3 mm wide at base in the compressed
state, very minutely papillate outside, glabrous inside
and on nectary; lobes ca. 1.5 mm long, recurved,
ca. 4.5 mm wide at base, abruptly tapering into an
obtuse apex ca. 1.5 mm wide. Anthers coherent for
their entire length, ca. 9.5 mm long, fertile part ca.
3.5 mm; sterile tips horizontally truncate at apex,
exserted from corolla for less than 1 mm; filaments
ca. 7 mm long, ca. 0.8 mm wide, tapering toward
anther. Style surpassing corolla by 2.5-3 mm; stig¬
ma 2-lobed. Nutlets not seen.

Onosma apiculatum belongs to series Tian¬
schanica Popov, the three other species of which
are 0. gmelinii Ledebour, O. irritans Popov ex
Pavlov, and O. baldshuanicum Lipsky. From O.
gmelinii, the new species is distinguished by its
green instead of grayish leaves, bracts longer than
calyx, club-shaped corolla, and slightly longer an¬
thers; from 0 . irritans by its markedly wider leaves
without a distinct petiole, presence of short hairs
between the bristles, bracts longer than calyx, short¬
er calyx, and club-shaped corolla; and from O. bald¬
shuanicum by its narrower leaves without a distinct
petiole, much denser cover of bristles and short
hairs, bracts longer than calyx, and shorter anthers.
Onosma apiculatum also differs from these three
species by its shorter corolla and by the shape of
its corolla lobes, which are very broad at base and
suddenly contracted into a narrow tip, hence the
specific epithet apiculatum. The type collection was
misidentified as O. echioides L., but this species is
native to Europe and adjacent parts of Turkey and
differs by several characters, the most notable of
which are the shorter calyx scarcely half as long as
corolla and the asterotrichous indumentum.

Acknowledgment. I thank Ihsan Al-Shehbaz for
his help in the preparation of this manuscript and
in obtaining the loan of specimens.

New Combinations in Neotropical Grammitidaceae (Pteridophyta)

Alan R. Smith

University Herbarium, University of California, Berkeley, California 94720, U.S.A.

Abstract. Continuing study on neotropical Gram¬
mitidaceae for various floras in press or in progress
necessitates the following new combinations: Cera-
denia oidiophora, C. phalacron, Lellingeria cili-
olepis, L. dissimulans, L. hombersleyi, Melpomene
gracilis, M. zempoaltepetlensis, Terpsichore bi-
pinnata, T. elastica, and 7. Jlexuosa.

Examination of additional types of several rare
and geographically restricted species of Grammiti¬
daceae, as well as further study of problematic groups,
has resulted in the need for several new combina¬
tions. This work is a continuation of observations
previously published on the genera Ceradenia, Lel¬
lingeria, Melpomene, and Terpsichore (Bishop,
1988; Smith et ah, 1991; Smith & Moran, 1992;
Smith, 1993).

Ceradenia oidiophora (Mickel & Beitel) A. R.
Smith, comb. nov. Basionym: Grammitis oi¬
diophora Mickel & Beitel, Mem. New York
Bot. Gard. 46: 202. 1988. TYPE: Mexico.
Oaxaca: Dto. Ixtlan, trail from San Pedro No-
lasco N to the Llano Verde, Mickel 3822 (ho-
lotype, NY; isotype, UC).

Known only from four collections from Oaxaca
(Mickel & Beitel, 1988).

Ceradenia phalacron (Stolze) A. R. Smith, comb,
nov. Basionym: Grammitis phalacron Stolze,
Fieldiana, Bot. 32: 84. 1993. TYPE: Peru.
Cuzco: Valley of Rio Urubamba, Bites A29
(holotype, US).

Ceradenia phalacron is known only from the
type. It is somewhat aberrant in Ceradenia because
of its relatively simple or only shallowly lobed blades,
lack of laminar setae, and prominulous veins, but it
fits more comfortably there than in any other de¬
scribed genus. With other ceradenias, it agrees in
the absence of hydathodes, castaneous rhizome scales
with setulose margins, lack of spongy parenchyma,
the prominulous venation, the lack of laminar or
circumsoral setae, and the glandular paraphyses
within the sorus.

Lellingeria ciliolepis (C. Christensen) A. R. Smith,
comb. nov. Basionym: Polypodium ciliolepis
C. Christensen in Asplund, Ark. Bot. 20A: 21.
1926. Ctenopteris ciliolepis (C. Christensen)
Copeland, Philipp. J. Sci. 84: 395. 1956
[1955]. Grammitis ciliolepis (C. Christensen)
Lellinger, Amer. Fern J. 74: 58. 1984. TYPE:
Bolivia. Sur Yungas, El Chaco valley, Asplund
1501 (isotype, US).

Lellingeria dissimulans (Maxon) A. R. Smith,
comb. nov. Basionym: Polypodium dissimu¬
lans Maxon, Contr. U.S. Natl. Herb. 10: 502.
1908. Ctenopteris dissimulans (Maxon) Cope¬
land, Philipp. J. Sci. 84: 415. 1956 [1955].
Grammitis dissimulans (Maxon) F. Seymour,
Phytologia 31: 179. 1975. Melpomene dis¬
simulans (Maxon) A. R. Smith & R. C. Moran,
Novon 2: 429. 1992. TYPE: Guatemala. Alta
Verapaz: near Coban, von Tiirckheim (ed. Don¬
nell Smith 884) (holotype, US).

On the basis of descriptions, this rare and poorly
known species was mistakenly transferred to Mel¬
pomene (Smith & Moran, 1992), but examination
of the type and several additional specimens from
Guatemala (Johnson 678, NY, US; Steyermark
48781, US; Standley 71160, US) indicates that it
belongs in Lellingeria (Smith et al., 1991). Lellin¬
geria dissimulans is similar to L. randallii (Maxon)
A. R. Smith & R. C. Moran, from Panama and
Jamaica, except that the rhizome scales are larger
and lack marginal setae, and the blades completely
lack hairs and setae on the lamina and rachis abax-
ially; the sori are somewhat impressed. The species
is so far known only from Guatemala.

Lellingeria hombersleyi (Maxon) A. R. Smith,
comb. nov. Basionym: Polypodium homber¬
sleyi Maxon, Amer. Fern J. 20: 1. 1930. Cten¬
opteris hombersleyi (Maxon) Copeland, Phi¬
lipp. J. Sci. 84: 430. 1956 [1955]. Grammitis
hombersleyi (Maxon) Lellinger, Proc. Biol. Soc.
Wash. 89: 714. 1977. TYPE: Trinidad. Hom-
bersley 331 (holotype, US).

This is known only from Trinidad and, like /
dissimulans, is also related to L. randallii.

Novon 5: 20-21. 1995.

Volume 5, Number 1
1995

Smith

Neotropical Grammitidaceae

21

Melpomene gracilis (Hooker) A. R. Smith, comb,
nov. Basionym: Polypodium gracile Hooker,
Bot. Misc. 2: 239. 1831. Grammitis gracilis
(Hooker) Stolze, Fieldiana, Bot. 32: 96. 1993.
TYPE: Peru. Pasco: Huayllay, Cruckshanks
s.n. (holotype, K not seen; isotype, GH not
seen).

Melpomene zempoaltepetlensis (Mickel & Bei-
tel) A. R. Smith, comb. nov. Basionym: Gram¬
mitis zempoaltepetlensis Mickel & Beitel,
Mem. New York Bot. Card. 46: 205. 1988.
TYPE: Mexico. Oaxaca: Dto. Mixe, NW slope
of Cerro Zempoaltepetl, trail from Yacoche to
Totontepec, Mickel 4638 (holotype, NY).

I now believe this is sufficiently distinct from M.
pilosissima to warrant species status. Melpomene
zempoaltepetlensis differs in lacking setae at the
margins of segments and appears to be confined to
southern Mexico.

Terpsichore bipinnata (Stolze) A. R. Smith, comb,
nov. Basionym: Grammitis bipinnata Stolze,
Fieldiana, Bot. 32: 92. 1993. TYPE: Peru.
Cuzco: Prov. La Convencion, Loma Grande,
Biles 2167 (holotype, US not seen; isotype, F
not seen).

Terpsichore elastica (Bory ex Willdenow) A. R.
Smith, comb. nov. Basionym: Polypodium
elasticum Bory ex Willdenow, Sp. PL, ed. 4.
5: 183. 1810. Ctenopteris elastica (Bory ex
Willdenow) Copeland, Philipp. J. Sci. 84: 426.
1956 [1955]. TYPE: Bourbon [Reunion]. Bory
s.n. (holotype, B-Herb. Willd. 19675 not seen;
microfiche, UC).

I have now seen sufficient material from Mada¬
gascar and the Seychelles to be convinced that this
species is not synonymous with T. cultrata (Bory
ex Willdenow) A. R. Smith, as previously thought
(Smith, 1993). It differs by the narrower blades and

by the absence of rhizome scales, thus agreeing with
T. mollissima (Fee) A. R. Smith. Terpsichore elas¬
tica is the only known species in the genus from
Africa and islands of the Indian Ocean. Like many
of its neotropical congeners allied to 7. cultrata
(Group 3 of Smith, 1993), the spores are ellipsoid,
monolete, and binucleate when shed (van der Werff
12890 , UC, from Madagascar).

Terpsichore flexuosa (Maxon) A. R. Smith, comb,
nov. Basionym: Polypodium Jlexuosum Max¬
on, Contr. U.S. Natl. Herb. 17: 597, t. 42.
1916. Grammitis maxoniana Lellinger, Amer.
Fern J. 74: 58. 1984 (nom. nov., not G. flex¬
uosa Kunth = Eriosorus flexuosus). Lellin-
geria flexuosa (Maxon) A. R. Smith & R. C.
Moran, Amer. Fern J. 81: 84. 1991. TYPE:
Cuba. Oriente: Camp La Gloria, S of Sierra
Moa, Shafer 8037 (holotype, US).

On the basis of descriptions, this species was
mistakenly transferred to Lellingeria by Smith and
Moran (in Smith et al., 1991). Examination of the
type and other specimens from Cuba shows that
Terpsichore flexuosa is closely related to T. sta-
heliana (Posthumus) A. R. Smith, also with setose
sporangia, and not Lellingeria pendula (Swartz) A.
R. Smith & R. C. Moran, as Maxon thought.

Literature Cited

Bishop, L. E. 1988. Ceradenia, a new genus of Gram-
mitidaceae. Amer. Fern J. 78: 1-5.

Mickel, J. T. & J. M. Beitel. 1988. Pteridophyte flora
of Oaxaca, Mexico. Mem. New York Bot. Card. 46:
1-568.

Smith, A. R. 1993. Terpsichore, a new genus of Gram¬
mitidaceae (Pteridophyta). Novon 3: 478-489.

- & 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.

Arracacia ravenii (Apiaceae), a New Species from Southern Mexico

Lincoln Constance

Herbarium and Department of Integrative Biology, University of California,
Berkeley, California 94720, U.S.A.

James Affolter

The State Botanical Garden of Georgia, The University of Georgia,
2450 South Milledge Avenue, Athens, Georgia 30605-1624, U.S.A.

Abstract. A new species of Arracacia (Apiaceae)
is described from the high mountains of Chiapas and
Oaxaca, Mexico. Although sharply distinct in many
respects, it is contrasted with A. pringlei J. Coulter
& Rose, the Oaxacan taxon it most closely resem¬
bles.

Arracacia Bancroft is one of the largest and most
important apioid genera of Umbelliferae in the New
World, not least because some of its members served
as important food plants for the Incas. The genus
is central to a number of other Mesoamerican gen¬
era, such as Tauschia , Coaxana, Coulterophytum,
Myrrhidendron, and Neonelsonia, all of which have
been confused with it in one way or another. It
continues to yield well-marked but previously un¬
described forms from throughout its extensive range.
Attention is called to the following example to fa¬
cilitate its inclusion in Flora Mesoamericana.

Arracacia ravenii Constance & Affolter, sp. nov.
TYPE: Mexico. Chiapas: Municipio de Tene-
japa, steep heavily wooded slope of Cerro Zon-
tehuitz, with Quercus, Drimys & Magnolia on
NE side of hill, paraje of Matsab, alt. 9200 ft.
(2804 m), 25 Aug. 1966, D. E. Breedlove
15311 (holotype, DH; isotype, UC). Figure
la-f.

Species Arracaciae pringlei J. Coulter & Rose affinis
sed habitu procumbenti, non glauco-caulibus, divisionibus
foliorum confertis acutis non acuminatis, pedunculo so-
litario terminali, involucello evidenti, floribus flavo-viren-
tibus, fructibus late ovoideis differt.

Slender, caulescent, branching, woody-based pe¬
rennial herbs forming trailing mats to 60 cm long,
the foliage glabrous, the inflorescence scaberulous;
leaves triangular-ovate, 5-15 cm diam., temate-
bipinnate or 2-3-pinnately dissected, the ultimate
divisions oblong-ovate to lanceolate, 5-15 mm long,
1-10 mm broad, crowded, sessile, acute, spinulose-

serrate and pinnately incised toward base, the up¬
permost confluent, conspicuously reticulate, squa-
mosely tufted on the rachis above, a well-defined
nodal ring lacking; petiole 5-10 cm long, oblong¬
sheathing; cauline leaves numerous, like the basal;
inflorescence of a solitary rather stout terminal pe¬
duncle 10-20 cm long; umbels 8-12 cm diam.;
involucre lacking, or of inconspicuous filiform bracts;
rays 12-18, mostly fertile, slender, 3.5-5 cm long,
subequal, slightly webbed; umbellets 30-40-flow-
ered, the 2-8 fertile pedicels slender, 5-10 mm
long; involucel of 5-8 linear-lanceolate, scarious-
margined, acuminate bractlets to 10 mm long, equal¬
ing flowers but shorter than fruit; flowers greenish
yellow, the petals oval; stylopodium conical, the
styles slender; carpophore apparently bipartite; fruit
broadly ovoid, 4-5 mm long, 3-4 mm broad, gla¬
brous, tapering at apex, the ribs very prominent;
vittae large, usually solitary in intervals, 2 on com¬
missure; seed channeled under intervals, the face
sulcate; chromosome number unknown.

Distribution. Known only from mountains of cen¬
tral Chiapas (Cerro Zontehuitz) and southern Oaxaca
(Cerro Zempoaltepetl).

Habitat. Steep slopes with Quercus, Drimys,
Magnolia, Ilex, and Podocarpus, in wet montane
or cloud forest, at 2400-3000 m.

Arracacia ravenii keys readily to the group of
woody-based species comprising A. fruticosa Rose,
A. ebracteata (Rose) Mathias & Constance, A. ae-
gopodioides (Kunth) J. Coulter & Rose, and A.
pringlei J. Coulter & Rose, more particularly the
last. It differs from A. pringlei in its sprawling habit,
green rather than glaucous stems and rachises,
crowded leaf divisions, solitary terminal peduncle
rather than terminally clustered peduncles, evident
involucel, yellowish green rather than purple flowers,
and broadly ovoid rather than oblong-fusiform fruit.

We are pleased to name the plant for Peter
Raven, who, among so many other accomplish¬
ments, played a very significant role in initiating the
Flora of Chiapas project.

Volume 5, Number 1
1995

Constance & Affolter
Arracacia ravenii

23

Figure 1. Arracacia ravenii Constance & Affolter. —a. Habit. —b. Basal leaf. —c. Fruiting umbellet. —d. Fruit,
lateral view. —e. Mericarp transection. —f. Petal, (a, Breedlove & Raven 8105; b-f, Breedlove 15311.)

Paratypes. MEXICO. Chiapas: Mun. of San Cristobal
las Casas, SW slope of Cerro Zontehuitz, 19 Jan. 1965,
Breedlove & Raven 8105 (DH, UC), 24 May 1965,
Breedlove 10020 (DH, MICH, UC); Santa Cruz en San
Felipe, 15 Nov. 1986, Ton & M. C. Mtz. de Lopez 9529
(MO); Mun of Tenejapa, slope at Paraje Matsab, 5 Jan.
1966, Ton 487 (DH, UC); slope in Paraje Pahal Ton,
20 Jan. 1966, Ton 668 { DH, UC), Cerro Sontehuits, Las
Casas-Tenejapa, 28 June 1942, M. C. Carlson 2411

(MEXU); summit of Cerro Tzonohuitz, 92°35'W, 16°49'N,
26 Mar. 1986, Snow & Whittemore 212 (MO). Oaxaca:
steep N slope of Cerro Zempoaltepetl, 35 km N of Ayutla,
Mitla-Choapam, 17 Apr. 1988, Breedlove & Barthol¬
omew 66871 (CAS).

Acknowledgment. The figure was prepared by
Charlotte Mentges Hannan.

Two New Species of Araceae for the Guianas

Thomas B. Croat

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Abstract. Two species of Araceae from the
Guianas are described. Philodendron billietiae,
known from Guayana, French Guiana, and Brazil,
is most closely related to P. acutatum. Xanthosoma
granvillei, known only from French Guiana, is dis¬
tinguished mainly by its large, blackish-drying blades.

In order to complete the Araceae treatment for
the Flora of Saul and the Flora of the Guianas it
has become necessary to describe a number of new
species. Descriptions of other new species of Araceae
will follow.

Philodendron billietiae Croat, sp. nov. TYPE:
French Guiana. National Hwy. 2, Cayenne-
Regina, vie. km 93, 100-150 m, 4°18'N,
52°10'W, Croat 74321 (holotype, MO
4343643-5; isotypes, B, BBS, BR, BRG, CAS,
CAY, CM, F, G, GH, INPA, K, M, MG, NY,
P, US, VEN). Figures 1-4.

Planta hemiepiphytica; internodia brevia, usque 2.5-
5 cm diam.; petiolus subteres, 25-57 cm longus, pallide
aurantiacus-flavus vel viridi-flavus; lamina plus minusve
anguste triangularis et sagittata, 30-85 cm longa, 8.1-
21 cm lata; nervis lateralibus primariis 3-5 utroque;
pedunculus 9.4-27 cm longus; spatha 17.5-18.5 cm
longa, extus viridis; lamina intus alba; tubus intus pur-
purascens; spadix 16.5-18 cm longus; ovaria 6-11 lo-
cularia cum ovulis plus quam 10 in quoque loculo; affinis
P. acutato Schott sed internodis proportione brevioribus,
petiolo in vivo fiavido vel ochraceo versus constanter
viridi, lamina proportione angustiore basique lobis 35-
40° versus 60-75° divergentibus differt.

Appressed-climbing hemiepiphyte, sometimes on
rocks; stems about 1 m long, sometimes longer;
internodes shorter than broad, 2.5-5 cm diam., light
brown with irregular, weak, reticulate to mostly
transverse fissures, turning yellow-brown; petiole
scars to 4.5 cm diam.; cataphylls sharply D-shaped
near apex, otherwise terete and unribbed or some¬
times bluntly 2-ribbed, green to pale orange (some¬
times brown, orange at base) or reddish, promptly
caducous; roots moderately few, stout, ca. 5 mm
diam., purplish brown, drying smooth, semiglossy,
the epidermis often fissured; leaves usually clustered
near the end of the stem, with erect spreading pet¬
ioles and pendent blades; petioles broadly spreading,
25-57 cm long, averaging 40 cm long, broadly

Novon 5: 24-29. 1995.

sulcate at base, obtusely flattened toward the apex,
weakly angular near the apex, sheathed 4-8 cm at
the base, firm, olive green to golden-yellow, pale
yellow-orange or greenish yellow, when fresh, some¬
what asperous and rather densely purplish-lineate,
drying light yellow-brown to dark brown, typically
closely fissured; blades pendent, narrowly triangular
to narrowly ovate-triangular to oblong-triangular,
30-85 cm long, 8.1-21 cm wide (averaging 52 x
15 cm), 2.7-4 times longer than wide, averaging
3.4 times longer than wide, broadest at the base or
near the petiole attachment, sometimes weakly con¬
stricted above the petiole attachment, 0.85-1.4 times
as long as the petioles (averaging 1.07 times longer
than petioles), moderately coriaceous, dark green
and semiglossy above, much paler and matte to
weakly glossy below; the margins often weakly un¬
dulate, hyaline or reddish; posterior lobes directed
at about 170° from the midrib (the two lobes sep¬
arated by an angle of 35-40°), sometimes directed
up slightly at an angle to the midrib, 8-22 cm long,
3—10 cm wide, about twice as long as wide; sinus
much longer than wide, acute at apex, usually barely
naked along the margin, but sometimes weakly na¬
ked up to 2.5 cm (the leaf tissue merging almost
imperceptibly with the posterior rib); midrib flat to
very broadly raised and paler (sometimes orange-
yellow) with purplish dots above, narrowly raised
and convex, yellow-brown to yellow-red or yellow-
green below; primary lateral veins 3-5, widely spaced
and usually not prominent, weakly sunken to flat
and yellow-green above, convex to weakly raised
and sometimes reddish to purplish below, arising at
70-105° angle, mostly at nearly 90° angle, some¬
times prominently down-turned at the midrib, some¬
times splayed out at the midrib; minor veins nu¬
merous, moderately distinct when fresh, weakly raised
above, flat and distinctly visible below, drying prom-
inulous on both surfaces, with alternate veins fre¬
quently turned to join another minor vein just before
the margin; cross veins inconspicuous; young leaves
reddish. Inflorescences 1 3 per axil, erect; peduncle
9.4-27 cm long, ca. 1 cm diam., somewhat flat¬
tened, ribbed on one side, medium yellow-green,
green-lineate to purplish-lineate; spathe 17.5-18.5
cm long, yellow-green to dark green outside, slightly
paler toward apex, densely pale purplish-lineate on

Volume 5, Number 1
1995

Croat

Araceae from the Guianas

25

Figures 1-4. Philodendron billietiae Croat. —1. Habit, plant creeping over rocks in full sun; Croat 74230. —
2. Close-up of stem showing old intact cataphyll and leaf scars, roots, and petioles; Moonen s.n., cultivated in Cayenne.
— 3. Inflorescence at anthesis showing male portion of spadix barely emergent; Moonen s.n. —4. Open inflorescence;
Moonen s.n.

26

Novon

tube, dark purplish-lineate on blade, the outer mar¬
gin of the tube slightly paler, the blade white to
greenish white within, the margins weakly turned
back at anthesis; tube sometimes reddish-spotted
outside, mauve to maroon within, becoming dark
maroon-purple within after anthesis with conspicu¬
ous, closely spaced resin canals restricted to the
area of the constriction, blending in with the red of
the tube and the white of the blade; spadix 16.5-
18 cm long; staminate portion white, to 9 cm long,
to 11 mm diam. at base, weakly constricted to ca.
9 mm somewhat above the sterile portion, tapered
to the narrowly rounded apex; pistillate portion pale
green, to 7.2 cm long on front side, 5.3 cm long
on back side; pistils ca. 3 mm long, pale yellow-
green, turning pale yellow post-anthesis; ovaries 6-
11-locular; ovules more than 10 per locule; sterile
male flowers broader than the pistillate portion, scal¬
loped and undercut at base, 2.2 cm diam., 10-11
mm long; fertile portion constricted 8.5 cm above
base, creamy white, bluntly pointed at apex.

Philodendron billietiae is known from Guyana,
French Guiana, and Brazil (Para) in tropical rain¬
forest at 140 to 340 m elevation; certainly it is to
be expected in Suriname as well. In Brazil it has
been collected only near Belem, and only a live
collection cultivated at the Moscow Main Botanical
Garden has been seen.

The species is a member of Philodendron subg.
Philodendron (formerly Polyspermium) and is
characterized by its long, mostly triangular leaf blades
with a very narrow sinus, elongated posterior lobes,
relatively few, widely spaced primary lateral veins,
and especially by its greenish yellow or pale orange
petioles, which are about as long as the blade.

Philodendron billietiae was first collected by H.
A. Gleason of the New York Botanical Garden in
1921 in Guyana, where it has not been collected
subsequently. It is named in honor of Frieda Billiet
of the Jardin Botanique National de Belgique, Meise,
who studied the species at several localities in French
Guiana, brought it into cultivation, and first brought
it to my attention. The species is in cultivation in
the botanical garden at Meise, as well as at the
Missouri Botanical Garden and the Moscow Main
Botanical Garden.

Philodendron billietiae is most easily confused
with and probably most closely related to P. acu-
tatum Schott, a similar species with which it may
occur. Philodendron acutatum differs in having
proportionately longer intemodes, consistently me¬
dium green petioles, and proportionately wider,
mostly narrowly ovate blades that often have the
posterior lobes divergent from one another at an

angle of 60° or more. In addition, P. acutatum is
ecologically different from P. billietiae in French
Guiana, occurring principally in the coastal swamp
forests with Avicennia.

Paratypes. FRENCH GUIANA. Pic Coudreau-Mont
Bakra, Region des Emerillons, 670 m, 3°18’N, 52°57'W,
de Granville & Cremers 11833 (CAY, MO, P, U, US);
Savane Roche de Virginie-Approuague River basin, 140
m, 4°1 l'N, 52°9'W, Cremers & Petronelli 11683 (CAY,
K, MO, P, U); Montagnes de Kaw, 4°33'N, 52°09'W,
Cremers 12718 (CAY); Montagnes de Kaw, along road
to Montagne Favard, vie. jet. to Fourgrassie, 200 m,
4°38'N, 52°17'W, Croat 74335 (CAY, K, MO, US);
Route Nationale, RN 2, 4°35'N, 52°20'W, Cremers 12723
(CAY); km 95.5, Prevost 1739 (CAY); km 67, Crique
Tibourou, along old route for RN 2, 5 m, 4°29'N, 52°19'W,
Billiet & Jadin 5740 (B, BR, CAY, K, MO, NY); km
93, 100-150 m, 4°18'N, 52°10'W, Croat 74321 (B,
BR, CAY, K, NY, MG, P, US, VEN); road to Montagnes
de Kaw, km 34, 4°33'N, 52°09'W, 340 m, Billiet &
Jadin 4600 (BR); La Compte River, vie. km 40 on RN
2, 4°N, 52°21'W, Billiet & Jadin 4714 (BR, CAY, MO);
between La Compte & Cacao, S of Cayenne, 1981, Billiet
81-0879 (BR), Croat 76441 (MO); Saul, vie. Eaux Claires,
Sentier Botanique near headwaters of St. Eloi River, 350
m, Croat 74230 (CAS, CAY, CM, M, MO, W). GUY¬
ANA. Rockstone, Gleason 846 (GH).

In addition to the localities listed above, the spe¬
cies has been seen in French Guiana on the road to
Petit Saut, km 20 (F. Billiet, pers. comm.).

Xanthosoma granviliei Croat & Thompson, sp.
nov. TYPE: French Guiana. Saul, vie. Eaux
Claires, along road between Eaux Claires and
Belizon, 230 m, 3°37'N, 53°12'W, Croat
74131 (holotype, MO-4342303; isotypes, B,
CAY, CM, K, NY, P, US). Figures 5-8.

Planta terrestris; caulibus 15-30(-50) cm longis; in-
ternodia usque 6(—15) cm diam.; petiolus teres, 39-
75(-100) cm longus; lamina ovato-hastata, 19-70 cm
longa, (16-)21-50 cm lata, 2.1-2.3 plo longior quam
lata, in sicco subdenigrata; cum costa postica nuda 1.0-
5.5 cm; nervis primariis lateralibus 3-6 utroque; inflo-
rescentiae 3-6 per axila; pedunculus 8-30 cm longus;
spatha 11.5-20 cm longa; tubus 4.5-6.5 cm longus,
extus viridus, intus atropurpureus; lamina 8-8.5 cm lon¬
ga, extus et intus albida; spadix usque 18 cm longus,
parte pistillata plus minusve aurantiaca; ovaria 2-3 lo-
cularia, loculi 2-3 ovulati; fructus aurantiacus.

Terrestrial; stems rhizomatous, the older portion
creeping over ground, the younger portion erect,
15—30(—50) cm long; internodes short, to 6(— 15)
cm diam.; cataphylls to 25 cm long, persistent,
turning dark brown, the uppermost persisting semi¬
intact; petioles 39-75(-100) cm long, terete but
somewhat flattened near apex, succulent and juicy,
tinged purplish at base, sheathed usually less than
1 4 its length (up to 20 cm when subtending an in¬
florescence); blades ovate-hastate, 19-70 cm long.

Volume 5, Number 1
1995

Croat

Araceae from the Guianas

27

Figures 5-8. Xanthosoma granvillei Croat & Thompson; Croat 74131. —5. Habit showing an inflorescence and
an emerging new leaf. —6. Stem with old persistent leaf sheaths. —7. Leaf blade. —8. Close-up of inflorescence
showing an unopened spathe.

28

Novon

(16-)21-50 cm wide, 2.1-2.3 times longer than
broad, prominently lobed at base, acuminate at apex,
dark green and semiglossy above, moderately paler
and weakly glossy below, drying somewhat black¬
ened, dark olive-green above, dark brownish green
below; the lateral margins usually convex, rarely
concave; posterior lobes 1.2-1.8 times longer than
wide, subacute at apex, sometimes turned weakly
outward, often held somewhat upward at an angle
to the midrib; sinus hippocrepiform or sometimes
obovate, 9-19 cm deep; posterior rib straight and
extending outward at ca. 60° angle to the midrib,
naked 1-5.5 cm along sinus; basal veins 6-9 pairs,
the uppermost free, the remainder pinnately ar¬
ranged along the posterior rib; primary lateral veins
3-6 pairs, arising at 40-60° angle; minor veins
weakly etched above, flat and darker than surface
below. Inflorescences 3-6 per axil; peduncle green¬
ish, 8-30 cm long, longer than or shorter than the
spathe; spathe 11.5-20 cm long, constricted at low¬
er !4, the tube 4.5-6.5 cm long, the blade white,
sometimes lined with pink, the tube green outside,
sometimes tinged purplish, dark purple within, some¬
times becoming deep red outside; spathe blade 8-
8.5 cm long, whitish on both surfaces; spadix to 18
cm long, several centimeters shorter than spathe,
female portion bright orange to yellow-orange; ova¬
ries 2-3-locular; locules usually 2-3-ovular. In-
fructescences with peduncles to 22 cm long, fruiting
spathe to 7 cm long, fruiting spadix 3.5-4.5 cm
long, 2-2.5 cm diam. Fruits orange; immature seeds
1.6-1.8 mm long, 1 mm diam., obtuse at apex,
cordulate at base, ca. 10-ribbed longitudinally. Very
common and conspicuous in wet season but surviving
as an underground stem during the dry season.

Xanthosoma granvillei is known only from
French Guiana, mostly between 200 and 500 m,
but rarely to nearly sea level; it is to be expected
in adjacent Suriname. The species was first collected
in 1966 by Oldeman at Cacao, south of Cayenne.
It is named for Jean Jacques de Granville from
ORSTOM, in Cayenne, who collected more well-
prepared specimens of the species than any other
collector.

Xanthosoma granvillei is recognized by its short
stems emerging usually only a short distance above
the ground level, its ovate-cordate, blackish-drying
blades with triangular posterior lobes, sunken minor
veins, and a partly naked, usually hippocrepiform
sinus. Additionally, the cluster of 3—6 inflorescences
with peduncles about as long as the spadix, with
orange female spadices, distinguish it from other
species of Xanthosoma.

Xanthosoma granvillei is most easily confused

with another blackish-drying species that also ap¬
pears to be new to science. That species differs by
having tuberous growth, smaller, more hastate leaf
blades, and an inflorescence with slender peduncles
that are 3-4 times longer than the spathe. It is
represented by collections from the Montagnes de
Kaw (Cremers 2719, 8702) and from the nearby
region at Crique Armontabo, a tributary of the Oya-
pock River ( Cremers 7055). It will be described
later.

Xanthosoma granvillei might also be confused
with X. undipes K. Koch & Bouche, which differs
in having a longer stem, foul-smelling sap, and blades
that dry greenish rather than blackened. Superfi¬
cially, X. granvillei is very similar to X. holivar-
anum Bunting from the El Dorado-Gran Sabana
area of Venezuela. That species, described from
sterile material, has similar stems and blades but
differs in having obtusely sulcate petioles sheathed
to about the middle (versus merely obtusely flattened
and usually less than ‘4 its length for X. granvillei),
and leaf blades that dry gray-green above and light
yellow-green below (not at all blackened). In addi¬
tion, X. bolivaranum lacks any obvious lateral veins
directed off the posterior rib toward the base, where¬
as X. granvillei has 2-3 prominent lateral veins
branched off the posterior rib and directed toward
the base.

Paratypes. FRENCH GUIANA. Cacao, 60 km S of
Cayenne, Oldeman B-450 (P); Montagne Cacao, SE of
Cayenne, Cremers 13031 (CAY, MO), Feuillet 548 (CAY);
Basin de la Comte River basin, 4°34'N, 52°27'W, Cre¬
mers 12728 (CAY); Region de Paul Isnard, Citron Petite
piste Boeuf Mort, Feuillet 653 (CAY, K); Mont Atachi
Bacca-Inini River region, 9 km SE of Gobaya Soula, 420
m, 3°33'N, 53°55'W, de Granville et al. 10649 (CAY,
K); E of Gobaya Soula, 440 m, 3°33'N, 53°55'W, de
Granville et al. 10482 (B, CAY, FTC, K, US); Montagnes
de Kaw, road to Montagne Tresor, side trail to Placer
Tresor, 4°35'N, 52°18’W, 150 m, Croat 74283 (CAY,
MO); Montagnes de Kaw, Cascade de Fourgassie, 4°37'N,
53°18'W, Cremers & Feuillet 12968 (CAY, MO); Mon¬
tagnes de Kaw, road to Degrad Lalanne, 300 m, 4°34'
N, 52‘TO’W, Hoff et al. 6308 (CAY); Station des No-
uragues, Prevost 2503 (CAY); Saul, Mont Galbao, 550
m, 3°36'N, 53°17'W, de Granville et al. 8608 ( B, CAY,
K, NY, P, U, US), 500 m, de Granville et al 8837
(CAY, K, US); Saiil, in village, jardin de Richard, Capus
2 (CAY); 10 km SSW of village, de. Granville 5420 (BR,
CAY, K, P); 18 km S of village, de Granville 5531 (BR,
CAY, K, P); Saiil, Pied de Mont Galboa, 200 m, 3°37'N,
53°16’W, de Granville et al. 9042 (CAY); Mont La
Fumee, 3°37'N, 53°12'W, Mori & Pipoly 15575 ( NY);
Saiil, Fumee, Prevost 1798 (MO); Saul, La Fumee Moun¬
tain trail, 250 m, Mori & Grade 18865 (NY); Mont
Galboa, 10 km SW of Saiil, 650 m, de Granville 2401
(CAY, P); Saul, road to Eaux Claires, 250 m, Mori &
Grade 21083 (NY); Saiil, vie. Eaux Claires, along road
between Eaux Claires and Belizon, 220-240 m, 3°37'N,
53°12'W, Croat 74131 (B, CAY, CM, K, MO, NY, P,

Volume 5, Number 1
1995

Croat

Araceae from the Guianas

29

US), de Granville 4480 (K); Saul, road to Belizon, 0-2
km S of Eaux Claires, Mori et al. 22194 (MO, NY);
Saiil, Grand Boeuf Mort trail, 300 m, Mori & Grade
18935 (CAY, NY); vie. Saiil, trail to Mont Galboa, de
Granville 4486 (CAY, K); Rte. de Belizon, km 7.7,
4°20'N, 52°23'W, Billiet & Jadin 4424 (BR, CAY);
Saiil, ENE of village, Pic Matecho, Cremers 6184 (CAY,

K); Saiil Region, Saut Quaimicouare, Yaroupi River (trib¬
utary of the Oyapock), de Granville 345 (P); Bassin du
Sinnamary, Crique Kourcibo, Saut Kawene, 12 m, 4°53'N,
53°3'W, Hoff et al. 6475 (CAY, MO); Chemin de fer
Gare-Tigre-St. Elie, km 0-5, 20 m, 4°57'N, 53°8'W,
Hoff et al. 6585 (CAY, MO).

Paspalum niquelandiae (Poaceae: Paniceae), a New Species from the
Serpentine Outcrops of Central Brazil

Tarciso S. Filgueiras

Reserva Ecologica do IBGE, Caixa Postal 08770, 70312-970 Brasflia, DF, Brazil

ABSTRACT. Paspalum niquelandiae Filgueiras, a
new species of Poaceae from the serpentine outcrops
of Niquelandia, a municipality in the Brazilian state
of Goias, is described, illustrated, and compared with
Paspalum soboliferum Chase, P. humboldtianum
Flugge, and P. inconstans Chase. The conserva-
tional importance of a serpentine endemic grass flora
in central Brazil is briefly discussed.

Recent fieldwork carried out in the municipality
of Niquelandia, in the Brazilian state of Goias, where
serpentine soils have been reported (Brooks et al.,
1990), resulted in the description of several new
taxa in the Poaceae (Davidse & Filgueiras, 1993;
Filgueiras et al., 1993; Filgueiras & Davidse, 1994).
A new species of Paspalum L. from the same area
is herein described, illustrated, and compared to
similar species.

Paspalum niquelandiae Filgueiras, sp. nov.
TYPE: Brazil. Goias: Municipio de Niquelandia,
ca. 14'T8'S-48°23'W, morro pedregoso, ca.
2 km Leste da localidade de Macedo, planta
crescendo entre pedras, proximo a pequena
floresta de galeria, ocasional na area, 19 maio
1993, T. S. Filgueiras & F. C. de A. Oliveira
2461 (holotype, IBGE; isotypes, F, G, ICN, K,
MEXU, MG, MO, R, SI, SP, US). Figure 1.

Paspalo sobolifero Chase spiculis ellipticis subtiliter
pilosis, gluma superiore marginibus ciliatis simile, sed cul-
mis non ramosis, ligula 0.2-0.3 mm longa, laminis 20-
45 cm longis et 8-22 mm latis, communi rhachidi 9-15
cm longa, gluma superiore ciliis densis, numerosis, co-
roniformibus abunde differt.

Coarse, caespitose perennial. Rhizomes well de¬
veloped, knotty, covered with cataphylls. Culms 120-
185 cm tall, erect, unbranched in the vegetative
portion, with 5-8 elongated internodes; internodes
hollow, stramineous, glabrous; nodes dark, glabrous.
Prophyll not seen. Leaves mostly cauline; sheaths
strongly striate, glabrescent to pilose in the lower
half, becoming papillose-hispid upward, the margins
free, both ciliate or one glabrous toward the base;

Novon 5: 30-33. 1995.

auricles absent; ligule a minute, brown, glabrous
membrane, v-shaped, 0.2-0.3 mm long; cilia at the
ligular area 5-8 mm long, colorless; blades 20-45
cm long, 8-22 mm wide, flat, lanceolate, glabrescent
to papillose-hispid, acuminate apically, the midrib
projecting abaxially, the margins serrate-denticulate
to ciliate; uppermost blade reduced, minute. Inflo¬
rescence of unilateral, ascending racemes, divergent
in maturity; peduncle well exserted, 12-22 cm long,
glabrous; common rachis 9-15 cm long, glabrous,
cylindrical at the base, becoming 3-4-angulate up¬
ward, the margins denticulate, the internode at the
base of each raceme with a tuft of colorless hairs
2-2.5 mm long; racemes 8-30, 2.5-8.5 cm long;
rachis 1-1.5 mm wide, zig-zag, flat abaxially, with
a central ridge adaxially, arcuate in maturity, bear¬
ing spikelets to the tip, the margins ciliate or den¬
ticulate, or both. Pedicels of unequal length, 0.1-
0.3 mm long, scabrid, the apex discoid. Spikelets
paired, each pair alternate on each side of the rachis,
pedicelled, 2.3-3.3 mm long, 1-1.2 mm wide,
broadly elliptic-lanceolate in outline, pale to purplish,
dorsally compressed, abaxial, awnless, with 2 florets,
disarticulating below the glume and falling as a unit;
lower glume absent; upper glume slightly to strongly
convex on the back, 2.3-3.3 mm long, 3-nerved,
glabrous to finely appressed pubescent, the margins
densely ciliate-pectinate; the cilia colorless, 0.3-0.7
mm long, with bulbous bases, the apex apiculate;
lower floret sterile, consisting only of a lower lemma;
lower lemma 2.2-2.9 mm long, narrower than the
upper glume, 3-nerved, navicular, hyaline, with a
depressed area at the base, the surface glabrous to
short-pubescent, the apex glabrous to slightly ciliate;
lower palea absent; upper floret 2.2-2.7 mm long,
bisexual, slightly chartaceous, pale; upper lemma
obscurely 5-nerved, navicular, glabrous, the surface
finely papillose-striate, the apex acute; germination
flap evident; upper palea 2.1-2.2 mm long, acu¬
minate, glabrous, slightly chartaceous, the surface
finely papillose-striate; lodicuies 2, fleshy, condu-
plicate, 0.1-0.2 mm long; stamens 3; anthers 1.2-
2 mm long, laterally exserted; ovary 0.5—0.8 mm
long, glabrous; styles 2, separate, 1.5-1.8 mm long;
stigmas 2, plumose, yellow or purplish. Caryopsis
1.8-1.9 mm long, 0.9-1 mm wide, light-colored.

Volume 5, Number 1
1995

Filgueiras

Paspalum niquelandiae

31

Figure 1. Paspalum niquelandiae Filgueiras, illustrated from the type collection (Filgueiras & Oliveira 2461 ).
— A. Habit. — B. Ligular area of leaf. —C. Portion of a raceme with paired spikelets. — D. Dorsal view of spikelet.
—E. Upper floret, lemma side. —F. Upper floret, palea side.

32

Novon

Table 1. Main distinguishing characteristics of Paspalum

niquelandiae , P. humboldtianum , and P. soboliferum.

Character

P. niquelandiae

P. humboldtianum

P. soboliferum

Habit

Erect

Reptant

Erect

Height (cm)

120-185

40-120

ca. 60

Culm

Unbranched

Branched

Branched

Nodes

Glabrous

Pilose

Glabrous

Ligule (mm)

0.2-0.3

1.8-3.8

2-2.5

Blades (cm x mm)

20-45 x 8-22

1-8 x 6-17

5-15 x 6-9

Common rachis (cm)

9-15

1.5-6

4-6

Number of racemes

8-30

1-8

3-7

Raceme length (cm)

2.5-8.5

4-10

2-4.5

Rachis width (mm)

1-1.5

1-3

1-1.2

Spikelet size (mm)

2.3-3.3

3-4

2.8-2.9

Cilia on glume (mm)

0.3-0.7

1.9-2.1

0.4-0.5

with persistent stylar bases; embryo %-% as long as
the caryopsis; hilum punctate, basal, Z l0 -'/ s as long
as the caryopsis.

Paspalum niquelandiae is morphologically most
closely related to Paspalum soboliferum Chase be¬
cause of the shape of the spikelets and especially
because of the cilia on the margins of the upper
glume; it differs in the unbranched culms, smaller
ligule, longer blades, longer common rachis, greater
number of racemes, and size and density of the cilia
on the margins of the upper glume. Paspalum so¬
boliferum is known only from a single plant (the
holotype) from Ecuador (Hitchcock, 1927). The new
species is more distantly related to Paspalum hum-
boldtianum Flugge; the latter is readily distinguished
by its sprawling habit, the pilose nodes, the size of
the ligule, blades, and common rachis, and the num¬
ber of racemes. Paspalum humboldtianum occurs
from Mexico to Argentina (Chase, 1929). Table 1
summarizes the main differences among P. nique¬
landiae , P. humboldtianum , and P. soboliferum.
It is evident from Table 1 that these species form
a trio of closely related species with some overlap¬
ping characteristics, but whose distinctiveness is also
readily apparent.

The spikelet shape and the ciliate lower lemma
remotely link Paspalum niquelandiae to P. incon-
stans Chase, a species from Bolivia, Ecuador, and
Peru. The latter is of smaller stature (culms 50 to
60 cm high), has both terminal and axillary racemes,
and possesses a lower number of racemes (1 to 8,
but commonly 2 to 5). A further distinguishing char¬
acteristic is the presence of a lower glume in the
lower spikelets of P. inconstans (occasionally lack¬
ing in some spikelets). In addition, the cilia on the
lower lemma of P. inconstans are smaller and much
less dense than in P. niquelandiae.

The classification of Paspalum niquelandiae is
not yet clear. It is obviously related to a group of
species included by Chase in her unpublished mono¬
graph of the South American species of Paspalum
[accessible from the Hitchcock & Chase Grass Li¬
brary of the Department of Botany, Smithsonian
Institution, Washington, D.C., U.S.A] in subgenus
Ceresia (Persoon) Reichenbach. This group of spe¬
cies (which includes Paspalum bicilium Mez, P-
carinatum Kunth, P. polyphyllum Nees, among
others) has a characteristic upper glume with con¬
spicuous marginal cilia. The inclusion of P. nique¬
landiae in subgenus Ceresia is not acceptable be¬
cause this would make the subgenus exceedingly
heterogeneous. Although related to some species in
subgenus Ceresia, P. niquelandiae is best left un¬
grouped until more convincing evidence of relation¬
ships becomes available.

Paspalum niquelandiae is known from two pop¬
ulations in the municipality of Niquelandia in the
Brazilian state of Goias. It is apparently restricted
to the serpentine outcrops in the area. It grows in
scattered clumps among rocks and boulders, near
the edge of some gallery forests, but was never found
inside the gallery forest itself. Although the clumps
seen were rather robust and, therefore, easy to spot,
large populations of the species were never located.

This is the third species of Paspalum described
from the locality of Macedo in Niquelandia. The
other two are P. longiaristalum Davidse & f “•
gueiras (Davidse & Filgueiras, 1993) and P■ biar-
istatum Filgueiras & Davidse (Filgueiras & Davidse,
1994). Ophiochloa Filgueiras, Davidse & Zuloaga,
a panicoid grass genus, has also been recently de¬
scribed from the same area (Filgueiras et al., 1993).
The discovery of so many new taxa in such a small
area (ca. 3 km 2 ) in central Brazil suggests that Ni¬
quelandia is a center of high plant diversity and

Volume 5, Number 1
1995

Filgueiras

Paspalum niquelandiae

33

endemism, primarily due to the presence of serpen¬
tine soils, with their high concentrations of nickel,
chromium, cobalt, and other minerals (Brooks, 1987).

The presence of a highly specialized flora in Ni-
quelandia is of utmost concern because the area is
the center of the largest nickel mining and smelting
operation in Brazil (Brooks et al., 1990). There is
not a single conservation unit in the area to grant
legal protection to this unique flora. A detailed bo¬
tanical survey should be undertaken in the region
to assess the diversity of the entire flora and its level
of endemism. The conservational implications of the
presence of such a peculiar flora in Niquelandia will
be addressed in a forthcoming publication.

The specific epithet refers to the town of Nique¬
landia, in whose municipality the new species was
collected.

Paratypes . BRAZIL. Goias: municipio de Niquelan¬
dia, ca. 1 km Leste da localidade de Macedo, morro
pedregoso, 19 maio 1993, T. 5. Filgueiras & F. C. de
A. Oliveira 2456 (IBGE, RB, UB), 4 ago. 1992, T. S.
Filgueiras & R. D. Lopes 2427 (B, IBGE).

Acknowledgments. I am grateful to CAPES (Bra¬
zilian Post-Graduation Federal Agency) and IBGE
(Instituto Brasileiro de Geografia e Estatistica) for a
post-doctoral fellowship that enabled me to spend
18 months at the Missouri Botanical Garden. I also

thank the Missouri Botanical Garden staff for mak¬
ing their facilities available for my research. I es¬
pecially thank my friend Clarence J. Lovejoy, Jr.,
and his family for their friendship and support during
my stay in St. Louis. Vladimiro S. Dudas, San Isidro,
Argentina, provided the excellent illustration.

Literature Cited

Brooks, R. R. 1987. Serpentine and Its Vegetation.
Dioscorides Press, Portland.

Brooks, R. R., R. D. Reeves, A. J. M. Baker, J. A. Rizzo
& H. D. Ferreira. 1990. The Brazilian serpentine
expedition (Braspex). 1988. Natl. Geogr. Res. 6:
205-219.

Chase, A. 1929. The North American species of Pas¬
palum. Contr. U.S. Natl. Herb. 28: 1-310.
Davidse, G. & T. S. Filgueiras. 1993. Paspalum lon-
giaristatum (Poaceae: Paniceae), a new serpentine
endemic from Goias, Brazil, and the first awned spe¬
cies in the genus. Novon 3: 129-132.

Filgueiras, T. S. & G. Davidse. 1994. Paspalum biar-
istatum (Poaceae: Paniceae), a new serpentine en¬
demic from Goias, Brazil, and the second awned
species in the genus. Novon 4: 18-22.

-,- & F. O. Zuloaga. 1993. Ophioch-

loa , a new endemic serpentine grass genus (Poaceae:
Paniceae) from the Brazilian cerrado vegetation. No¬
von 3: 360-366.

Hitchcock, A. S. 1927. The grasses of Ecuador, Peru,
and Bolivia. Contr. U.S. Natl. Herb. 24: 291-556
[Paspalum, contributed by Agnes Chase, pp. 434-
455].

A New Combination in Peltostigma (Rutaceae)

Roy E. Gereau

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. Peltostigma parviflorum Q. Jimenez
& Gereau is reduced to synonymy under P. gua-
temalense (Standley & Steyermark) Gereau, comb,
nov., based on Galipea guatemalensis Standley &
Steyermark.

Peltostigma Walpers (Rutaceae: Rutoideae) be¬
longs to subtribe Choisyinae of the tribe Zanthox-
yleae, while Galipea Aublet (Rutaceae: Rutoideae)
belongs to subtribe Cuspariinae of the tribe Cus-
parieae. The two subtribes may be contrasted as
follows (Engler, 1931):

Choisyinae: flowers actinomorphic; corolla poly-
petalous; stamens all antheriferous; seeds
with endosperm.

Cuspariinae: flowers usually zygomorphic; corolla
usually gamopetalous; some stamens often
reduced to staminodes; seeds with little or
no endosperm.

Peltostigma parviflorum Q. Jimenez & Gereau
is the only species of the genus with unifoliolate
leaves and small flowers (petals 8.2-10.1 mm long)
(Jimenez & Gereau, 1991); its congeners (P. pte-
leoides (Hooker) Walpers and P. eximium C. V.
Morton) possess 3—5-foliolate leaves and large flow¬
ers (petals 15-25 mm long) (Morton, 1933; Stan¬
dley & Steyermark, 1946). Galipea guatemalensis
Standley & Steyermark was named from a fruiting
specimen lacking the floral characters necessary for
correct generic (or even subtribal) placement in the
Rutaceae. Examination of its holotype has shown
that G. guatemalensis, an earlier name, is the same
species as P. parviflorum, necessitating the follow¬
ing new combination.

Peltostigma guatemalense (Standley & Stey¬
ermark) Gereau, comb. nov. Basionym: Gali¬
pea guatemalensis Standley & Steyermark,
Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 165.
1944. TYPE: Guatemala. Izabal: Rio Dulce,
between Livingston and 6 mi. upriver, on N
side, 1—25 m, 14 Apr. 1940 (fr), Steyermark
39448 (holotype, F).

Peltostigma parviflorum Q. Jimenez & Gereau, Arm.
Missouri Bot. Gard. 78: 527. 1991. Syn. nov. TYPE:
Costa Rica. San Jose: 9°43T0"N, 84°23T0"W, Zona
Protectora La Cangreja, Santa Rosa de Puriscal,
400-500 m, 4 sep. 1987 (fl), Q. Jimenez M. & L.
J. Poveda A. 494 (holotype, CR; isotypes, K, MO).

Jimenez & Gereau (1991) discussed geographical
trends in the morphology of Peltostigma guate¬
malense (as P. parviflorum) in terms of a “northern
race” consisting of the collections from Guanacaste
and Puntarenas, Costa Rica, and a “southern race
comprising the type and four collections from An-
tioquia, Colombia. The “races” may be contrasted
as follows: “Northern race”: petioles 0.2-2.1 cm
long, not extremely variable, the longest in a given
specimen exceeding the shortest by 0.5-1.5 cm;
leaflet blade 3.2-16.0 x 1.2-7.3 cm, the apex
obtuse or obtusely acuminate, the base acute to
rounded; petals 8.2-9.3 x 8.0-8.5 mm. “Southern
race”: petioles 0.3-5.6 cm long, extremely variable,
the longest in a given specimen exceeding the short¬
est by 2.0-4.5 cm; leaflet blade 4.2-20.6 x 2.2-
10.2 cm, the apex acutely acuminate, the base
usually attenuate; petals 9.8-10.1 x 8.5-9.7 mm.

The type of Galipea guatemalensis has larger
leaflets (to 19.5 x 8.7 cm) and longer petioles (to
2.6 cm) than is characteristic for the “northern
race,” but is otherwise more similar in foliar char¬
acters to the northern than to the southern popu¬
lations. A previously unreported collection of this
species from Peru has foliar characters entirely with¬
in the parameters of the “southern race,” but in¬
sufficiently mature flowers to assess the correlation
between geography, floral measurements, and foliar
characters. With few collections and little available
data on flower size, it is preferable to continue to
consider Peltostigma guatemalense a single species
with some observable correlation between geography
and morphological characters. Further investiga¬
tions based on more ample collections could reveal
the desirability of recognizing the two informal “rac¬
es” as subspecies.

Specimen examined. PERU. Madre de Dios: Manu,
Parque Nacional Manu, Rio Manu, Pakitsa station, Tach-
igali Trail, 4 km N of camp, 11°56'S, 71'16'W, 350 m,
11 Sep. 1989 (fl), R. B Foster & H. Beltran 11371
(F, MO).

Novon 5: 34-35. 1995.

Volume 5, Number 1
1995

Gereau

Peltostigma

35

Acknowledgments. I thank J. A. Kallunki and
A. Pool for bringing the type of Galipea guate-
malensis to my attention; J. S. Miller and G. E.
Schatz for useful discussions of geographic vari¬
ability; and C. M. Taylor for a careful reading of
the manuscript.

Literature Cited

Engler, A. 1931. Rutaceae. Nat. Pflanzenfam. ed. 2,
19a: 187-359.

Jimenez M., Q. & R. E. Gereau. 1991. Peltostigma
parviflorum (Rutaceae), nueva especie de Costa Rica
y Colombia. Ann. Missouri Bot. Gard. 78: 527-530.

Morton, C. V. 1933. A new species of Peltostigma
from Mexico. Proc. Biol. Soc. Wash. 46: 83-84.

Standley, P. C. & J. A. Steyermark. 1946. Rutaceae.
In: P. C. Standley & J. A. Steyermark, Flora of
Guatemala, Fieldiana, Bot. 24(5): 389-425.

Triniochloa talpensis (Poaceae: Pooideae), a New Species with Deciduous
Leaves from Mexico

Manuel Gonzalez-Ledesma, Maricela Gomez-Sanchez, Stephen D. Koch
Centro de Botanica, Colegio de Postgraduados, 56230 Chapingo, Edo. de Mexico, Mexico

Teresa Mejia-Saules

Instituto de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico

ABSTRACT. Triniochloa talpensis from Mexico is
described and illustrated. Its morphology and leaf
anatomy are compared with those of T. laxa Hitch¬
cock.

RESUMEN. Se describe e ilustra Triniochloa tal¬
pensis de Mexico. Se comparan caracteristicas de
morfologia y de anatomia foliar con aquellas de 7.
laxa Hitchcock.

Triniochloa is a small genus of grasses that was
described by Hitchcock in 1913, the members of
which grow in temperate forests from Mexico to
Bolivia. It is distinguished from other genera of the
Pooideae by its spikelets 1-flowered without a rach-
illa extension, floret callus obtuse and bearded, and
lemma 5- or 7-nerved with a bifid apex and genic¬
ulate awn inserted above the middle. This genus was
included in the tribe Meliceae of the subfamily Pooi¬
deae by Reeder (1968) on the basis of its connate
leaf sheaths, membranous ligules, truncate lodicules
that lack vascular traces, and small chromosomes.
He suggested that it is closely related to Schizachne.
This placement is supported by Clayton & Renvoize
(1986) and Watson & Dallwitz (1992).

Triniochloa, as presently circumscribed, includes
four species: T. laxa Hitchcock and T. micrantha
(Scribner) Hitchcock, which are restricted to Mex¬
ico; T. stipoides (H.B.K.) Hitchcock, distributed
from central Mexico to Colombia, Venezuela, and
Bolivia; and T. andina Luces, known only from
Venezuela. The new species from western Mexico
increases the number to five.

Mature blades of basal leaves from herbarium
specimens were used for anatomical studies. A seg¬
ment about 10 mm long was removed from the
middle of the blade, rehydrated by boiling in water,
sectioned free-hand, and mounted in glycerin jelly.
Epidermis preparations were made following Met¬
calfe’s (1960) method. Drawings were made using
an X-100-1 Ken-A-Vision Microprojector.

Novon 5: 36-39. 1995.

Triniochloa talpensis M. Gonzalez-Ledesma &
M. Gomez-Sanchez, sp. nov. TYPE: Mexico.
Jalisco: Municipio de Talpa de Allende, alrede-
dores de la mina Zimapan en la Sierra de Cuale,
ca. 2150 m, 10 Feb. 1992, M. Gonzalez-
Ledesma & ]. A. Perez de la Rosa 496 (ho-
lotype, CHAPA; isotypes, IBUG, MEXU,
MICH, MO, US, XAL). Figure 1.

Plantae perennes, rhizomatosae. Culmi 60-110 cm
alti. Vaginae foliorum pubescentes prope nodos. Ligulae

1.5- 2.5(-5) mm longae. Laminae foliorum planae 22-
27 cm longae et 3.0-5.0 mm latae inferioribus caducis.
Panicula erecta, angusta, 16-34 cm longa, ex vagina
exserta. Spiculae 11.0-16.8 mm longae. Glumae inae-
quales, prima spiculis breviore, 7.5-13.6 mm longa; se-
cunda quam lemma aequanti vel longiore, 11.0-16.5 mm
longa. Lemma 11.0-14.5 mm longum, 7-nerve, arista
dorsali geniculata, 13.0-23.0 mm longa. Antherae (4.8-)

5.5- 7.0 mm longae.

Perennial herbs with rhizomes to 20 cm long, 2-
3 mm diam., rhizome sheaths strigose near the nodes.
Culms 60-110 cm tall, 1.5-2.0 mm diam. near the
base, robust, strongly decumbent, pubescent below
the upper nodes. Leaves, at least the lower, ciliate
near the collar. Leaf sheath margins connate, the
upper sheaths shorter than the internodes and pu¬
bescent below. Ligules 1.5-2.5(-5) mm long, mem¬
branous. Blades 22 27 cm long, 3.0-5.0 mm wide,
flat, deciduous at level of the collar or beneath,
adaxial surface scabrous to pubescent, with promi¬
nent ribs, abaxial surface glabrous. Panicle 16—34
cm long, 1.0—2.0(—2.5) cm wide, erect, narrow,
completely exserted, with 30-65(-90) spikelets;
panicle axis glabrous below, scabrous on the angles
toward the apex; lowest node 1-2-branched; branch¬
es appressed or ascending, scabrous at least toward
the apex. Spikelets 11.0-16.8 mm long, 1-flowered.
Glumes unequal, hyaline except near the nerves.
First glume 7.5—13.6 mm long, shorter than the
spikelet, 1-nerved. Second glume 11.0-16.5 mm
long, as long as the spikelet, 3-nerved. Lemma 11.0—

38

Novon

Figure 2. Leaf blade anatomy. A-D. Triniochloa talpensis (Gonzalez-Ledesma & Perez de la Rosa 496 , CHAPA).
—A. Outline of half of the blade. —B. Detail of transverse section, including the midrib. —C. Abaxial epidermis as
seen in surface view. —D. Prickle. E-G. Triniochloa laxa (T. Mejia-Saules 1820 , CHAPA). —E. Outline of the
blade. —F. Detail of transverse section, including the midrib. — G. Abaxial epidermis as seen in surface view. Stippling
indicates the costal region in both C and G.

14.5 mm long, with a basal stipe 0.5 mm long,
rounded dorsally, glabrous or scabrous at the apex,
7-nerved, the central 3 converging at or above the
middle to form the awn; apex bifid; awn 13.0-23.0
mm long, geniculate; callus with hairs up to 5 mm
long. Palea shorter than lemma. Rachilla not pro¬
longed beyond floret. Stamens 3, anthers (4.8-)5.7-
7.0 mm long. Lodicules 2, rounded, connate, short,
and turgid. Caryopsis not seen.

Leaf Blade Anatomy (Fig. 2A-D)

Transverse section. Outline of the blade flat to
involute. Adaxial surface with squared, flat-topped
ribs associated with first-order vascular bundles and
rounded ribs associated with second-order bundles;
furrows narrow, moderately deep. Abaxial surface
smooth, midrib only slightly projecting. Outer vas¬
cular bundle sheaths complete or interrupted by
abaxial or adaxial sclerenchyma, round in outline.
Sclerenchyma girders present, the adaxial T-shaped.
Chlorenchyma nonradiate, its cells of various sizes

and shapes, continuous between adjacent vascular
bundles. Colorless parenchyma cells absent. Bulli-
form cells in fan-shaped, adaxial groups not pro¬
jecting above the adjacent epidermis.

Abaxial epidermis. Long cells longer than wide
with undulating anticlinal horizontal walls. Stomata
absent. Intercostal short cells solitary, square or tall
and narrow. Papillae absent. Prickles absent, or a
few in the costal zone near the margin of the blade.
Silica cells in costal zone with dumb-bell-shaped silica
bodies, sometimes associated with cork cells. Costal
short cells solitary or in pairs, seldom in short rows.

Triniochloa talpensis is clearly a member of the
genus Triniochloa in its anatomical features (Wat¬
son & Dallwitz, 1992) and morphology, but it is
unique in the genus because of its deciduous leaves.
The basal and lower culm leaves simply die, and if
the blade becomes detached it does so at the apex
of the sheath, as in most grasses. The upper culm
leaves have a special way of separating; when they
are mature and dry, the blade and the portion of
the sheath above the point of closure separate in

Volume 5, Number 1
1995

Gonzalez-Ledesma et al.
Triniochloa talpensis

39

Table 1. Salient characters distinguishing Triniochloa talpensis and T. laxa.

T. talpensis

T. laxa

2.0-3.0 0.8-1.0

60-110 40-80

deciduous

completely exserted

Rhizome diameter (mm)

Plant height (cm)

Leaf blade
Panicle

Panicle length (cm)

Spikelets per panicle

Adaxial ribs of first-order bundles

Adaxial furrows

Abaxial projection of the midrib
Adaxial sclerenchyma girders
Bulliform cells

Anticlinal horizontal walls of long
cells

Prickles (abaxial epidermis)

16.0-34.0

30-65(-90)

squared

narrow

inconspicuous

T-shaped

fan-shaped groups not projecting
above the epidermis
undulating

absent or a few in the costal zone
near the margin of the blade

persistent
partially exserted
7.0-13.0
6-18
rounded

wide and shallow
conspicuous
I-shaped

rows projecting above the level of
the epidermis
straight

abundant in all costal zones

one piece from the remainder of the sheath (Fig.
1C). This is especially noticeable in pressed, dried
material when the upper blades are moved in order
to expose the ligule or when they are slightly pulled.
Detachment is along a well-defined line, but there
is no articulation nor evidence of an abscission layer.
Detachment was not seen in fresh material when
the plants were collected and pressed. It appears
that an abscission layer develops when the plants
are drying. This suggests that the phenomenon is
possibly a response to drought conditions.

Triniochloa talpensis grows in open pine-oak
forests and is only known from the Sierra de Cuale
in the municipio of Talpa (whence the specific ep¬
ithet), state of Jalisco, in the western part of the Eje
Neovolcanico. It is closely related to T. laxa , a
species that grows in forests of spruce and pine
mixed with poplar, and is restricted to the northern
Sierra Madre Occidental in the state of Chihuahua.

Although both species have rhizomes and similar
spikelets, Triniochloa talpensis is more robust than
T. laxa, and there are various other characters that
distinguish them (Table 1). In addition, important
anatomical differences were found in the arrange¬
ment of the adaxial sclerenchyma, the protrusion of

the bulliform cells, the shape of adaxial ribs, the
abundance and distribution of the prickles, and other
features (Fig. 2, Table 1). Deciduous leaves, narrow
adaxial leaf furrows, and more abundant adaxial
prickles (Fig. 2B) suggest that T. talpensis is adapt¬
ed to drier habitats than T. laxa.

Acknowledgments . We thank Fernando Chiang
Cabrera for reviewing the Latin description. We also
thank the curators of COL, ENCB, MEXU, MO,
and US for lending specimens under their care, and
the anonymous reviewers for their valuable obser¬
vations.

Literature Cited

Clayton, W. D. & S. A. Renvoize. 1986. Genera Gra-
minum. Kew Bull., Addit. Ser. 13: 1-389.
Hitchcock, A. S. 1913. Mexican grasses in the United
States National Herbarium. Contr. U.S. Natl. Herb.
17(3): 181-389.

Metcalfe, C. R. 1960. Anatomy of Monocotyledons. I.

Gramineae. Oxford Univ. Press, Oxford.

Reeder, J. R. 1968. Systematic position of the genus
Triniochloa (Gramineae). Amer. J. Bot. 55: 735.
Watson, L. & M. J. Dallwitz. 1992. The Grass Genera
of the World. C. A. B. International, Wallingford,
U.K.

Didymocarpus miniatus (Gesneriaceae), a New Species from
Peninsular Malaysia

Ruth Kiew

Department of Biology, Universiti Pertanian Malaysia, 43400 UPM Serdang,
Selangor, Malaysia

Abstract. Didymocarpus miniatus (Gesneri¬
aceae), a new species in Didymocarpus sect. Het¬
eroboea, is described from Bukit Bauk in Treng-
ganu, Peninsular Malaysia.

Didymocarpus miniatus Kiew, sp. nov. TYPE:
Peninsular Malaysia. Trengganu: Bukit Bauk,
R. Kiew RK 3792 (holotype, UPM; isotypes,
L, SING).

Differt a Didymocarpo atrosanguineo Ridley foliis
petiolatis, pedunculis brevioribus et floribus omnino min-
iatis.

Herb. Stem woody, up to 1 m tall and 0.7 cm
thick. Indumentum of young stem, petioles, and
lower surface of veins silky ferrugineous with uni-
seriate 3-4-celled trichomes ca. 1.5 mm long. Leaves
petiolate, spirally arranged on the upper 4-12 cm
of stem, separated by internodes up to 2 cm long.
Lamina oblanceolate, (15-)17-24 x (5.5-)6-10
cm, apex acuminate, acumen 1-2 cm long, strongly
narrowed to base, drying chartaceous, margin cre-
nate, indumentum of 2-celled uniseriate hairs ca.
0.5 mm long, dense and roughly velvety above, less
dense beneath, in life dark green above and pale
grayish green beneath. Midrib prominent above and
beneath, secondary veins 20-21 pairs, plane above
and prominent beneath, tertiary veins obscure above,
prominent beneath, areoles near the midrib irregular
in shape, those toward the margin rectangular. Pet¬
iole (1.5—)2(—3) cm long. Flowers solitary. Peduncle
slender, 6-12 mm long. Bract pairs ligulate, 3 mm
long, finely hairy. Calyx divided to base, lobes linear,
4 mm long, densely hairy. Corolla completely ver¬
milion with two yellow longitudinal nectar guides in
the base of the throat, finely pubescent outside, hairs
glandular uniseriate, 2-3-celled stalk up to 0.5 mm
long, nectar guides with scattered minute ellipsoid
vesicles, tube 2.5-3.7 cm long, narrow, 3-4 mm
wide in the lower half, dilating distaliy to 10 mm
wide, lobes 5, broadly rounded, upper two 4x7
mm, lower three 8x6 mm and projecting beyond
the upper two. Stamens 2, filament slender, 10 mm
long, anthers white, oblong, 2x1 m m, connivent,
positioned within corolla tube. Ovary 5 x 0.5 mm,

Novon 5: 40-41. 1995.

style slender, 30 mm long, densely pubescent with
glandular hairs, stigma discoid, peltate, 1 mm across.
Nectary encircling base of ovary, 0.75 mm high,
distaliy lobed for ca. one-third of its length. Capsule
slender, 7.5-8 cm long, finely pubescent when young.

Among the 85-odd Didymocarpus species in
Peninsular Malaysia, only D. miniatus has com¬
pletely vermilion flowers. Only one other species,
D. atrosanguineus, has flowers with red lobes, but
its corolla tube is usually cream-colored. (One or
two plants of this latter species in Sekayu Forest
Reserve, Trengganu, produce flowers that are com¬
pletely red, but they are deep crimson not vermilion.)
In lamina indumentum, D. miniatus most closely
resembles the broad-leaved form of D. atrosangui¬
neus (Kiew, 1989), but it is readily distinguished
from this form by its petiolate leaves.

Didymocarpus miniatus belongs to section Het-
eroboea, which includes species with large, trumpet¬
shaped, solitary flowers and large oblanceolate leaves
(Ridley, 1923). However, it is atypical of this sec¬
tion, in which the species have leaves with a winged
petiole, by its petiolate leaves. In addition, D. min¬
iatus is distinct in its short peduncles, 0.6-1.2 cm
long, as opposed to peduncles 2.5-10 cm long in
other species in this section.

In common with other large tubular Didymo¬
carpus flowers (Kiew, 1992), the stigma in D. min¬
iatus is discoid and peltate, and the nectary is large
and cylindrical (although it is the first species in
section Heteroboea to be described with a lobed
nectary). The glandular indumentum on the style
resembles that of D. quinquevulnerus Ridley, which
is also a member of section Heteroboea.

Didymocarpus miniatus has an extremely local
distribution, at the foot of Bukit Bauk, where it grows
on slopes above small swampy areas or seasonal
streams, and at about 20 km south on Bukit Cha-
bang, 5 km inland of Kerteh (G. W. H. Davison,
pers. comm.). Both of these hills belong to the same
coastal sandstone formation.

Paratypes. PENINSULAR MALAYSIA. Bukit Bauk,
S. Anthony SA 602 (UPM), G. IP. H. Davison GD5
(UPM), K. M. Kochumen KEP 9491 (KEP).

Volume 5, Number 1
1995

Kiew

Didymocarpus miniatus

41

Acknowledgments. This work was supported by
IRPA grant number 4-07-05-813. I thank Geoffrey
Davison, who first drew my attention to this new
species.

Literature Cited

Kiew, R. 1989. Didymocarpus (Gesneriaceae) on Gun-
ung Tahan, Malaysia. Gard. Bull. Singapore 42: 47-
64.

-. 1992. Five new species of Didymocarpus

(Gesneriaceae) from Peninsular Malaysia. Gard. Bull.
Singapore 44: 23-42.

Ridley, H. N. 1923. Didymocarpus. Flora Malay Pen¬
insula. 2: 506-523. Reeve, London.

A New Species of Campyloneurum (Polypodiaceae)
from Northwestern Ecuador

Blanca Leon 1

Department of Botany, Field Museum, Chicago, Illinois 60605, U.S.A. and Museo de Historia
Natural, Av. Arenales 1256, Apartado 14-0434, Lima-14, Peru

ABSTRACT. Campyloneurum oellgaardii is newly
described from a humid western premontane Andean
forest of Ecuador. It appears to belong to the sphen-
odes group, which is characterized by undivided
primary areoles and long petiolate leaves, and which
includes C. coarctatum, C. inflatum, C. sphenodes,
and C. sublucidum.

During a study of the fern genus Campyloneurum
C. Presl, a distinctive new species was recognized
and is described below.

Campyloneurum oellgaardii B. Leon, sp. nov.
TYPE: Ecuador. Carchi: drainage of Cerro Go-
londrinas mountains, 0°52'N, 78°07'W, 21 Dec.
1987, Hoover 2211 (holotype, MO; isotype,
QCA). Figure 1.

Species C. inflato proxima, a qua rhizomate longe
repenti, 6 mm crasso, atrofusco, dense paleaceo, squamis
adpressis, brunneolis, foliis ampliori lanceolatis, 147 cm
longis, 22 cm latis differt.

Epiphyte; stem long-creeping, black, not prui-
nose, 6 mm wide. Stem scales lax, light brown in
mass, ovate, 3-4 mm long, 2-2.5 mm wide, bases
auriculate, apices obtuse, margins entire with scat¬
tered hairs, slightly clathrate, the cells oblong or
broadly oblong, cell walls 6—9 pm wide, several
central cells with dark brown walls, walls of marginal
cells yellowish or brownish, cell lumina transparent.
Phyllopodia 4-5 mm long, 7-10 mm wide, 1-15
cm distant. Leaves pendulous (?), 147 cm long,
petiole 55 cm long, shiny dark stramineous; laminas
lanceolate, 22 cm wide, herbaceous-chartaceous,
base cuneate, apex long-acuminate, margins carti¬
laginous, sinuate, leaves with inconspicuous bicel-
lular glandular hairs scattered abaxially; stomata

1 Present addresses: Department of Geography, Uni¬
versity of Maryland Baltimore County, Baltimore, Mary¬
land 21228, U.S.A. and Department of Botany, NHB-
166, Smithsonian Institution, Washington, D.C. 20560,
U.S.A.

polocytic or rarely copolocytic; costa prominent,
slightly angular abaxially; primary veins prominent,
75° divergent from the costa, straight, lighter in
color than the adjacent tissue, 7-9 mm distant;
secondary veins slightly prominulous on both sides
of the lamina, transverse secondary veins forming
19—21 primary areoles between the costa and mar¬
gin; primary areoles undivided, with 2(—3) excurrent
free veinlets, the marginal ones sometimes divided
with (0)— 1 veinlet. Sori subapical on the excurrent
veinlet; paraphyses and spores not seen.

Campyloneurum oellgaardii is known only from
the type material, collected in northwestern Ecua¬
dor, at 1200 m elevation in a perhumid premontane
forest. This locality borders on what Dodson & Gen¬
try (1991) considered an extension of the Choco
pluvial forest. The specimen was found growing as
an epiphyte several meters above the ground. Based
on the curvature of the petiole (Fig. la), the leaves
are probably pendently arched.

Campyloneurum oellgaardii is characterized by
well-spaced and extraordinarily large leaves, more
than 1 m long, in addition to its undivided primary
non-costal areoles. Because of its habit, leaf mor¬
phology, and pattern of venation this species fits
within the sphenodes group of Leon (1992). Besides
this new species, the sphenodes group consists of
C. chrysopodum (Klotzsch) Fee, C. coarctatum
(Kunze) Fee, C.falcoideum (Kuhn ex Hieronymus)
M. Meyer ex Lellinger, C. inflatum M. Meyer ex
Lellinger, C. sphenodes (Kunze ex Klotzsch) Fee,
and C. sublucidum (Christ) Ching. Campyloneurum
oellgaardii differs from the other species of the
group in having a larger stem diameter (6 mm vs.
2-3 mm), obtuse stem scales, and longer (more than
100 cm vs. (17-)30-70(-85) cm) and broader (20
cm vs. (2-)3-6(-10) cm) leaves.

Most species of Campyloneurum have leaves less
than 1 m long. Leaves longer than this are otherwise
found only in: (1) those species with entire leaves,
short-creeping stems and phyllitidis-, brevifolium-
venation ( phyllitidis -, latum -venation of Lellinger,
1988) such as C. abruptum (Lindman) B. Leon, 6.

Novon 5: 42-44. 1995.

Volume 5, Number 1
1995

Leon

Campyloneurum oellgaardii

43

Figure 1. Campyloneurum oellgaardii B. Leon (from Hoover 2211). a. Stem scale. b. Cellular detail of stem
scale. —c. Habit. —d. Pattern of venation. (Drawn by the author.)

brevifolium (Link) Link, C. pascoense R. M. Tryon
& A. F. Tryon, C. phyllitidis (L.) C. Presl, and C.
tucumanense (Hieronymus) Ching; and in (2) those
pinnate-leaved species with undivided primary ar-
eoles, such as C. decurrens (Raddi) C. Presl and C.

magnificum T. Moore. All these species are low
epiphytes, epipetrics, or terrestrials, and they usu¬
ally grow in partially closed forests. In contrast, C.
oellgaardii is a high-canopy epiphyte, according to
the collector, and its gigantism may be an adaptation

44

Novon

to a dense canopy and/or low risk of leaf damage
due to breakage.

This species is dedicated to Benjamin 011gaard in
honor of his knowledge of the Ecuadorean flora,
especially the pteridophytes.

Acknowledgments. I thank David B. Lellinger
and Kenneth R. Young for comments on the manu¬
script. Thanks are also due to Altec Tangerini and
David Brunner for assistance.

Literature Cited

Dodson, C. & A. H. Gentry. 1991. Biological extinction
in western Ecuador. Ann. Missouri Bot. Card. 78:
273-295.

Lellinger, D. B. 1988. Some new species of Campy-
loneurum and a provisional key to the genus. Amer.
Fern J. 78: 14-35.

Leon, B. 1992. A Taxonomic Revision of the Fern
Genus Campyloneurum (Polypodiaceae). Ph.D. The¬
sis, University of Aarhus, Denmark.

A New Species of Beilschmiedia (Lauraceae) from Guerrero, Mexico

Francisco G. Lorea Hernandez

Departamento de Biologla, Facultad de Ciencias, UNAM, Ciudad Universitaria, Circuito
Exterior, 04510 Mexico, D.F., Mexico. Current address: Missouri Botanical Garden,
P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. Beilschmiedia angustielliptica is de¬
scribed and illustrated. Morphologically this species
is easy to recognize because of its long, reticulate
(but not foveolate) lanceolate leaves, its glabrous
anthers, and its glabrate ovary. So far, this species
is known only from the central mountains of Guer¬
rero, Mexico.

Resumen. Se describe e ilustra Beilschmiedia an¬
gustielliptica. Entre las otras especies del genero
esta nueva especie es facil de reconocer por sus
hojas grandes, lanceoladas, reticuladas (pero no fo-
veoladas), sus anteras glabras, y su ovario glabrado.
Hasta ahora esta especie solo se conoce de las mon-
tanas centrales de Guerrero, Mexico.

Extensive collections carried out during the last
ten years in Guerrero, Mexico, under the auspices
of the “Flora del Estado de Guerrero” project, or¬
ganized by the Laboratorio de Plantas Vasculares
(Facultad de Ciencias, UNAM), have resulted in
more complete collections for several formerly poor¬
ly known families in that state. For example, about
a dozen species of Lauraceae were previously re¬
ported from Guerrero (Standley, 1922; Allen, 1945).
Today, the assembled material represents more than
35 species. Several collections belong to undescribed
species in different genera. A new species of Beilsch¬
miedia is here described.

The Latin American species of Beilschmiedia
were last revised more than 50 years ago by Kos-
termans (1938), who recognized 15 neotropical spe¬
cies in the genus. Currently, the increase in collec¬
tions suggests the merger of some names, but also
the recognition of new species, so the actual number
of taxa is around 10-20 species (van der Werff,
1991).

In Mexico there exist about eight species of
Beilschmiedia. For a time, only B. mexicana (Mez)
Kostermans was recognized as occurring in the
country (Kostermans, 1938; Allen, 1945). Several

Mexican specimens of Beilschmiedia have been an¬
notated in herbaria as B. pendula (Swartz) W. B.
Hemsley, implying the presence of this species in
the region. But B. mexicana is suspected to be a
synonym of B. pendula (Burger & van der Werff,
1990). Early in the second half of this century B.
riparia F. Miranda was described from Chiapas
(Miranda, 1953) as a distinct species with pilose and
subcoriaceous leaves; this species is now known from
the tropical semideciduous forests in Guerrero, Oa¬
xaca, and Chiapas. Besides B. pendula, B. riparia,
and the newly described species, several collections
from different places in Mexico indicate the presence
of at least five other taxa, some of them conspecific
with or related to species known in Mesoamerica,
namely: B. anay (S. F. Blake) Kostermans, B. hon-
durensis Kostermans, and B. ovalis (S. F. Blake)
C. K. Allen.

Until a thorough revision of the American species
of Beilschmiedia is done, I present the following
provisional key to separate the Mexican species.

Key to the Mexican Species of Beilschmiedia

la. Leaf undersurface glabrous, rarely some straight
appressed hairs on main veins.

2a. Leaf undersurface finely reticulate (can-
cellate), minor veins evenly thickened
forming areolae 0.1-0.3 mm diam., no or
almost no free veinlets.

3a. Leaves stiff coriaceous, rather thick,

obtuse to acute . B. aff. mexicana

3b. Leaves chartaceous, acute to long

acuminate. B. mexicana

2b. Leaf undersurface roughly reticulate, mi¬
nor veins unevenly thickened, then areolae
usually 1.0 mm diam. or larger and ap¬
pearing with several free veinlets.

4a. Secondary veins in 14-18 pairs, leaves

narrowly elliptic.

. B. angustielliptica

4b. Secondary veins in 8-10 pairs, leaves

elliptic. B. hondurensis

lb. Leaf undersurface pubescent, if hairs only on
main veins, these not straight appressed.

Novon 5: 45-47. 1995.

46

Novon

^'7 ; B p U ^'"‘ l ? d i a angushelliptica Lorea-Hernandez. -A. Branchlet with inflorescence. -B. Branchlet

Tiew/of a Za k i ? ' ‘ aSt d ‘ visi ° ns - “ D ' °P en flower (°™y removed). -E. Lateral and abaxial

vtews of a thtrd whorl stamen. -F. Stammode. -G. Ovary. -H. Adaxial and lateral views of an external stamen.

5a. Hairs on undersurface straight and

appressed. fi. aif. pendula

5b. Hairs on undersurface straight to curly,
erect or loosely appressed.

6a. Hairs curly, erect to loosely

appressed. B. ovalis

6b. Hairs straight and erect.

7a. Leaf blades widely elliptic to ob-
lanceolate; widest leaves 11-14

cm. B. aff. anay

7b. Leaf blades elliptic, widest leaves

5-8.5 cm. B. riparia

Volume 5, Number 1
1995

Lorea Hernandez
Beilschmiedia angustielliptica

47

Beilschmiedia angustielliptica Lorea-Hernan-
dez, sp. nov. TYPE: Mexico. Guerrero: Mpo.
Atoyac de Alvarez, aprox. 2 km S to El Molote,
on the trail to El Eden, 1580 m, 19 May 1993
(fl, fr), F. Lorea & L. Lozada 5540 (holotype,
FCME; isotype, MO). Figure 1.

Arbores parvae (7-8 m altae). Folia opposita vel su-
bopposita, lamina (7-)ll-18(-24.5) x (1.5-)2.5-4.5(-
5.5) cm, anguste elliptica, aliquantum oblanceolata, basi
apiceque acuta vel acuminata, utrinque glabra, margine
interdum Ieviter undulata, nervis lateralibus subtus prom-
inentibus, costa supra complanata, subtus valde promi-
nenti ac in sicco manifeste rubiginosa. Inflorescentiae 2.5-
5.5(-9.5) x 1.5-3 cm, paniculatae, axillares in ramulos
aphyllos breves crebiter positae, vel terminales; axibus
puberulis vel tomentosis. Flores hermaphroditi, 2.2-2.7
mm longi, tepalis sex, ovatis vel late ovatis, erectis, abax-
ialiter pro parte maxima puberulis, adaxialiter praecipue
basi puberulis, margine glabris; staminibus novem, 1.1-
1.4 mm longis, antheris glabris, eis verticillorum primi et
secundi complanatis, late ovatis, eis verticilli tertii pyr-
amidalibus, filamentis omnino pilosis vel tomentosis; stam-
inodiis fere sessilibus, apice triangularibus, basi truncatis
vel subcordatis, ovario 0.7-0.85 mm longo, pyriformi,
glabrato; hypanthio tomentoso. Fructus 3-3.5 x ca. 1.7
cm, ellipticus, anthracinus.

Low trees (7-8 m), young branches compressed
to flattened, yellow-golden puberulent to tomentose,
bark reddish brown, glabrous and light brown with
rather smooth bark when older, buds densely to¬
mentose. Leaves opposite, a few subopposite, peti¬
oles (0.5-)0.7-1.3(-1.8) cm, slightly canaliculate,
puberulent, soon glabrous, blades (7—)11 — 18(—24.5)
x (1.5-)2.5-4.5(-5.5) cm, narrowly elliptic, some
oblanceolate, base and apex acute or acuminate,
chartaceous, glabrous on both surfaces, sparsely pu-
berulous when young, margin sometimes slightly
undulate, veins on upper and lower surface slighty
elevated, secondaries in 14-18 pairs, ascending,
prominent on lower surface, midvein flattened on
upper surface, very prominent and distinctly reddish
brown when dry on lower surface. Inflorescences
2.5-5.5(-9.5) x 1.5-3 cm, axillary, close to each
other on short aphyllous branchlets, or terminal,
paniculate, pedunculate, peduncle 0.6-2.5(—4.5) cm,
major axes 2.5-5.5 mm, puberulent to tomentose,
flower pedicels (2-)2.6-3.6(-4.2) mm, puberulent,
bibracteolate close to the base or not, bracteoles
minute and soon falling. Flowers hermaphroditic,
2.2-2.7 mm long, tepals six, 1.5-1.7 x 1-1.3 mm,
ovate to widely ovate, erect, outside puberulent for
most of the central area, inside mainly on the base,
margins glabrous, stamens nine, 1.1-1.4 mm long,
anthers of first and second whorls 0.7-0.8 mm long,
flattened, widely ovate, thecae introrse, anthers of
third whorl 0.5-0.6 mm long, pyramidal, thecae
extrorse-lateral, all filaments pilose to tomentose,
anthers glabrous, connectives protruding beyond the
anther cells, apparently glandular, glabrous, glands

irregularly round, 0.4-0.5 mm, staminodes 0.4-
0.6 mm long, almost sessile, apex triangular, acute,
base truncate or subcordate, adaxial face plane,
abaxial face convex and tomentose in the middle,
ovary 0.7-0.85 mm long, pyriform, glabrate, style
0.4-0.5 mm long, stigma apical, oblique, hypan-
thium 0.5-0.8 mm, tomentose. Fruit 3-3.5 x 1.7
cm, when mature, elliptic, black, smooth, and shiny.

Beilschmiedia angustielliptica is well distin¬
guished morphologically from other species in Amer¬
ica by its narrow, elliptic leaves conspicuously re¬
ticulate (but not foveolate), and glabrous anthers.
The general appearance of leaf reticulation in B.
hondurensis suggests some similarity with B. an¬
gustielliptica, but the former has more or less
abruptly acuminate, short elliptic leaves and the
ovary is glabrous, whereas in B. angustielliptica
the leaves are long lanceolate and the ovary is sparsely
pilose. So far, B. angustielliptica is known only
from mesophyllous mountain forests in Guerrero,
where the canopy is dominated by Quercus spp..
Magnolia schiedeana D. F. L. v. Schlechtendal,
Pinus strobus L. var. chiapensis M. Martinez, Per-
sea americana P. Miller, Licaria aff. triandra
(Swartz) Kostermans, Alchornea latifolia Swartz,
Cinnamomum sp., Prunus sp., and Pouteria sp.

Paratypes. MEXICO. Guerrero: Mpo. Atoyac, El
Molote, 1630 m, (fr), A. Nunez 1159 (FCME); Mpo.
Tecpan, km 16 on the road from El Porvenir to San
Antonio de Las Tejas, 1360 m, 7 m tree, (fl), G. Lozano
227 (FCME).

Acknowledgments. This paper was carried out
with support of a scholarship from DGAPA (Univ-
ersidad Nacional Autonoma de Mexico), and partial
support for fieldwork from the Missouri Botanical
Garden. I thank Henk van der Werff for critical
comments on the manuscript, and Roy Gereau for
having called my attention to some recommenda¬
tions of the Code of Botanical Nomenclature, and
for checking the Latin description. The illustration
was drawn by John Myers.

Literature Cited

Allen, C. K. 1945. Studies in the Lauraceae. VI. Pre¬
liminary survey of the Mexican and Central American
species. J. Arnold Arbor. 26: 280-434.

Burger, W. C. & H. van der Werff. 1990. Lauraceae.
In: W. Burger (editor). Flora Costaricencis. Fieldiana,
Bot. n.s. 23: 1-138.

Kostermans, A. J. G. H. 1938. Revision of the Lau¬
raceae V. Recueil Trav. Bot. Neerl. 35: 834-931.
Miranda, F. 1953. Plantas nuevas o notables de la flora
de Chiapas. Anales Inst. Biol. Univ. Nac. Mexico 24:
69-96.

Standley, P. C. 1922. Lauraceae. In: Trees and shrubs
of Mexico. Contr. U.S. Natl. Herb. 23(2): 286-298.
Werff, H. van der. 1991. A key to the genera of
Lauraceae in the New World. Ann. Missouri Bot.
Gard. 78: 377-387.

Passiflora sanctae-mariae in Series Luteae (Passifloraceae), a New
Species and Series in Subgenus Decaloba

John M. MacDougal

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT. Passiflora sanctae-mariae, restricted
to Chiapas, Mexico, and adjacent Guatemala, is new¬
ly described and assigned to a species group that
includes P. lutea L. of the United States. The five
species of this group lack extrafloral nectaries and
have very reduced or no bracts. The group is rec¬
ognized as Passiflora series Luteae (Small) J.
MacDougal, comb, et stat. nov.

Passiflora sanctae-mariae is a new species of
small vining passionflower, known from nine dried
collections and one living collection. The living col¬
lection, field collected by the author, was grown and
observed in the greenhouse for several years, and
it furnished the type collection. The species is re¬
stricted to the volcanic mountains of northwestern
Guatemala and adjacent Chiapas, Mexico. It is close¬
ly related to P. lutea L. of the United States, the
Mexican and Guatemalan P. pavonis Masters, an
as yet undescribed species in east-central Mexico
(Passiflora dictamo sensu Killip, not DC.), and P
filipes Bentham, which ranges from Texas to Ec¬
uador. These five species lack extrafloral nectaries
and have very reduced or no bracts, and form a
related species alliance here recognized as a series.

Passiflora series Luteae (Small) J. MacDougal,
comb, et stat. nov. Passiflora [no rank] Luteae
Small, Man. S. E. FI. 895. 1933. TYPE: Pas¬
siflora lutea L.

Small (1933), in his flora of the southeastern
United States, subdivided the seven species of Pas¬
siflora listed there into four categories, one of which
he named Luteae. It contained P. lutea and P.
sexflora A. L. Jussieu. This infrageneric unit was
published without rank, among the many such key¬
ing categories used throughout that work. Small’s
name is valid under Article 35.2 of the Code and
serves as the basionym for the name of the series.
Killip, in his taxonomic monograph of 1938, placed
P. lutea and P. sexflora in subgenus Plectostemma
Masters but separated the species into their own
separate series under section Decaloba (DC.) Mas¬
ters. He correctly associated P. pavonis and P.
filipes with P. lutea. Killip made no reference to

Novon 5: 48-51. 1995.

Small’s work, even in the bibliography of the mono¬
graph, and appears to have independently used the
name Luteae. Killip’s “series” are invalidly pub¬
lished, however, as they lack Latin diagnoses, and
the name is now validated as a series here. The
series is most easily recognized within subgenus De¬
caloba by the absence of extrafloral nectaries, ex¬
treme reduction or absence of floral bracts, and the
small purple berry. Further characteristics include
small flowers with the coronal filaments in two series,
the outer series filiform, greenish white to greenish
yellow or pale yellow, and sometimes dull purplish
or dull violet at base. The ovaries are glabrous or
with few short hairs. The Passiflora bilobata A. L.
Jussieu complex of the West Indies appears to be
the sister group to series Luteae based on floral
morphology and lack of extrafloral nectaries.

The delineation and lectotypification of subgenus
Plectostemma and the earlier and generally over¬
looked subgenus Decaloba (DC.) Reichenbach is
nearly the same (see discussion in MacDougal, 1994),
so the series and following new species are here
placed in subgenus Decaloba section Decaloba.

Passiflora sanctae-mariae J. MacDougal, sp. nov.
TYPE: Cultivated at Duke University, June
1982, from stems of MacDougal & Miley 602
collected Jan. 1980 in Guatemala, MacDougal
& Miley 602GR (holotype, DUKE; isotypes,
BM, CAS, CR, EAP, HUA, MEXU, MO, USCG,
US). Figures 1, 2b.

Passiflora scandens; petioli eglandulosi; folia eglan-
dulosa bilobata vel trilobata, lobis lateralibus acutis, mar-
ginibus integris, lobo centrali obtuso vel obsolete, vel trun-
cata; bracteae nullae; coronae filamenta biseriata, fila-
mentis exterioribus filiformis, 2.1-3.0 mm longis; oper¬
culum plicatum; ovarium glabrum; bacca atropurpurea;
semina 10-13 sulcata.

Slender perennial vine 2-5 m long, minutely pu-
berulent throughout at the shoot tip with trichomes
(0.05-)0.08-0.2(-0.3) mm long, except the adaxial
surface of the lamina glabrous (or a few trichomes
at the base of the primary veins), older growth
sparsely puberulent to glabrescent. Stems terete,
tardily glabrescent, drying subangulate and striate,

2.7cm

Volume 5, Number 1
1995

MacDougal

Passiflora sanctae-mariae

49

Figure 1. Passifiora sanctae-mariae J. MacDougal (from MacDougal & Miley 602GR, except as noted). A.
Habit. —B. Stipule. —C, D. Seed. —E-G. Leaves at prereproductive nodes; F (Steyermark 37098), G (Steyermark
37215). —H, I. Leaves at flowering nodes; H (Breedlove 50768), I (Purpus 7292).

3mm

50

Novon

with little secondary growth; phyllotaxy 2/5; nodes
sometimes purplish (especially prophylls and junc¬
tion of stipules); posture of shoot tip cernuous. Stip¬
ules (1.1 —)2—3.5 x 0.3-0.5(-1.0) mm, linear-lan¬
ceolate to broadly lanceolate, attenuate to acumi¬
nate, falcate. Petioles 1.1-2.6 cm long, eglandular,
minutely puberulent. Laminas at fertile nodes (2.5-)
4-7.3 X (4.7-)6-l 1.2 cm, entire, not variegated,
the general outline depressed obovate, truncate-ob¬
late, semi-circular, or transversely truncate-elliptic,
subtruncate to shallowly 2(-3)-lobed less than 0.15
the distance to the very shallowly cordate to broadly
rounded base, the lateral lobes acute (rarely slightly
obtuse or somewhat rounded), the central lobe obtuse
to obsolete, lobes not mucronulate or the mucro 0.5
mm long or less, the angle between the primary
lateral veins (62-)67-90(-11 2f , the ratio of central
to lateral vein lengths 1.1 — 1,5(—1.7), the ratio of
laminar width to length 1.2-2.0(-2.3); leaves of
juvenile plants often variegated along lateral veins,
narrowly depressed obovate to transversely trun¬
cate-elliptic in outline, 2(-3)-lobed to !4 the distance
to the base, the angle between the lateral veins 60-
130°, the ratio of central to lateral vein lengths 1.3-
2.1, the ratio of laminar width to length 1.2-3.6;
laminar nectaries absent. Tendrils straight during
development at shoot apex. Prophylls of the vege¬
tative ramifying bud 2, broadly lanceolate to ovate,
usually purplish, long acuminate or 3-toothed with
central tooth caudate. Peduncles 1.1-2.6 cm long,
geminate, uniflorous. Bracts absent. Hypanthium
4.5-5 mm diam., flowers ca. 1.5 cm diam., sub¬
erect, pale yellowish green with a dark center, with
faint sweet odor; stipe 3-6 mm long (to 8 mm in
fruit); sepals 7.5-10.5 x 2.8-4.2 mm, elliptic-
oblong to ovate-oblong, rounded to subacute, ecor-
niculate; petals 5-8 x 2.4—2.6 mm, oblong to ovate-
oblong, rounded, whitish; filamentous corona in 2
series, the outer filaments ca. 37-40, 2.1-3.0 mm
long, ca. 0.2-0.3 mm diam., filiform, curved-
spreading, slightly thicker in the distal half but at¬
tenuate at the very apex, cream to whitish, basally
light greenish and often with a small purplish mark,
the inner filaments 1.5 mm long, clavate, erect,
purplish with pale apex; operculum 1.5 mm long,
membranous, plicate, dark purple; nectary not con¬
cave; limen (disk) ca. 2.0 mm diam., uniformly dark
purple, the raised edge only ca. 0.2 mm high; sta-
minal filaments connate 3.1-4.1 mm along andro-
gynophore, unmarked except dark purple at junction
with limen, the free portions ca. 3-3.5 mm long,
unmarked; anthers 2.8-3.1 mm long, not marked
with purple; ovary 1.5-1.9 x 1.2-1.5 mm, widely
ellipsoid, glabrous, light green; styles ca. 3-3.5 mm
long; stigmas ca. 0.8 mm diam., cream. Fruit 1.1-

Figure 2. —A. Silhouette of flower of P. filipes

(MacDougal & Miley 579GR ). — B. Silhouette of flower
of P. sanctae-mariae (MacDougal & Miley 602GR ).

1.4 x 0.9-1.1 cm, widely ovoid-ellipsoid, purplish
black, estipitate; seeds 3.0-3.3 mm long, 2.0-2.3
mm wide, 1.2-1.3 mm thick, obovate to widely
obovate, nearly symmetric to slightly campylotro-
pous, transversely sulcate with 10-13 sulci, the
ridges somewhat verrucose, the chalazal beak broad¬
ly conical and slightly inclined toward the raphe.
Germination epigeal.

Passiflora sanctae-mariae is restricted to the
western slopes of the Serrania Transistmica, or Cor¬
dillera Madre, of Chiapas, Mexico, and adjacent
western Guatemala. It may be distinguished from
the other members of series Luteae in Mesoamerica
by the following key.

la. Plant completely glabrous; peduncles (1.5—)3—

5(-6) cm long; petals 3-4.5(-5) mm long; seeds
usually 3-4(-5)-sulcate; elevation 1100 m or
less. P. filipes

lb. Plant minutely puberulent to pubescent (except
adaxial surface of lamina) at least on young
growth and petioles; peduncles 1.1-2.6 cm long;
petals 5-8 mm long; seeds usually 6-13-sul-
cate; elevation 1300-2800 m.

2a. Stems pubescent, or sparsely so below;
leaves 1.5-5.5(-7) cm wide; corona 3-5
mm long; operculum pale greenish; andro-
gynophore 4.8-5.5 mm long; seeds 6-7-
sulcate (rarely irregularly rugose); eleva¬
tion (1350-)l 900- 2800 m. P. pavonis

2b. Stems minutely or sparsely puberulent, or
glabrescent below; leaves (4.7-)6-11.2 cm
wide; corona 2-3 mm long; operculum dark
purple; androgynophore 3.0-4.1 mm long;
seeds 10-13-sulcate; elevation 1300-1500

m. P. sanctae-mariae

Passiflora sanctae-mariae was collected in the
field in flower and fruit, then cultivated as a single

Volume 5, Number 1
1995

MacDougal

Passiflora sanctae-mariae

51

individual 1980-1986 at Duke University, with the
type specimens collected in June 1982. Observations
made on living material allowed the detailed de¬
scriptions of the flowers provided above. The flowers
opened in the greenhouse around midday and closed
in the late afternoon (the other species in the series
open in early to mid-morning and close in the early
to late afternoon). No autogamy was observed in six
years of cultivation, and fifteen self pollinations failed
to set fruit, good evidence that the species is self¬
incompatible.

Fruits and seeds are known only from my field
collection; the number of seeds per fruit was 26,
29, 30, and 38 + 2 aborted, i.e., seeds 26-40 (N
= 4). The number of seeds per fruit is significantly
higher than its relatives in the series, which range
from 2 to 12 (MacDougal, unpublished). A few of
the seeds were germinated in the greenhouse to
determine germination type, but the seedlings were
not grown. The germination type is epigeal with
elliptic foliaceous cotyledons, the norm for the ge¬
nus, but is worth noting because the close relative
Passiflora lutea has hypogeal germination, ex¬
tremely rare in the genus (MacDougal, 1994). Ger¬
mination type for the other species in the section is
unknown.

As is the case in many of the passifloras in section
Decaloba, the leaves of the juvenile plants of this
species are often variegated (cf. Fig. IF), but this
condition is not retained at reproduction. The leaves
tend to be larger than the other species of the section
and are characteristically apically truncate or sub¬
truncate, often giving the leaves a hemispherical or
bat-wing shape. The common name ala murcielago
is attached to some of the specimens, a name gen¬
erally associated with many of the bilobed passifloras
in northern Mesoamerica. One collector noted its
use as a diuretic, a use also shared by several other
bilobed passifloras in this area.

Passiflora sanctae-mariae is found only in pre-
montane and lower montane wet forest. In Quezal-
tenango it is associated with P. prolata Masters, P.
helleri Peyritsch, P. biflora Lamarck, and P. mem-
branacea Bentham.

This species is dedicated to the Blessed Mother
who is the namesake of the volcano where the clone
of the type was collected, and who bore Jesus Christ
whose sufferings are commemorated in the generic
name.

Paratypes. GUATEMALA. Quezaltenango: 100 m
up dirt road at Km post 197 on Hwy. 9S ca. 8 km below
tunnel near Santa Maria de Jesus, old finca Pirineos on
slopes near Volcan Santa Maria, ca. 1340 m, 20 Jan.
1980 (fl, fr), MacDougal & Miley 602 (BM, DUKE, F,
MEXU, MO); rich forested slopes along Quebrada San
Geronimo, finca Pirineos, lower S-facing slopes of Volcan
Santa Maria between Santa Maria de Jesus and Cala-
huache, 1-2 Jan. 1940, Steyermark 33331 (F); top of
ridge between finca Pirineos and finca Soledad, lower S-
facing slopes of Volcan Santa Maria, 1300-1400 m, 5
Jan. 1940, Steyermark 33525 (F). San Marcos: half¬
way up slopes above finca El Porvenir on “Todos Santos
Chiquitos,” lower S-facing slopes of Volcan Tajumulco,
1300-1500 m, 7 Mar. 1940, Steyermark 37093 (F),
Steyermark 37215 (F). MEXICO. Chiapas: Mpio. de
Angel Albino Corzo, above finca Tuxtepec, 1380 m, 21
Oct. 1980 (fl), Breedlove & Strother 46719 (CAS), 7
Apr. 1981 (fl), Breedlove 50768 (CASJ, 24 June 1981
(fl), Breedlove 51184 (CAS); finca Irlanda, June 1914,
Purpus 7292 (UC).

Acknowledgments. The type material was col¬
lected during fieldwork supported by NSF grant
DEB-7912607 while I was a graduate student on
an NSF Fellowship. I am grateful to Donald E. Stone
and the other members of my graduate committee
at Duke University, as well as the staff of the green¬
houses there, for their guidance and assistance. John
Myers prepared the drawing.

Literature Cited

Killip, E. P. 1938. The American species of Passiflor-
aceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1 —
613.

MacDougal, J. M. 1994. Revision of Passiflora sub¬
genus Decaloba section Pseudodysosmia (Passiflor-
aceae). Syst. Bot. Monogr. 41: 1-147.

Small, J. K. 1933. Manual of the Southeastern Flora.
New York. [Published by the author.]

New Combinations in Asian Krascheninnikovia Gueldenstaedt
(Chenopodiaceae)

Sergei L. Mosyakin

N. G. Kholodny Institute of Botany, 2 Tereshchenkivska Str., Kiev, 252601 Ukraine

ABSTRACT. New combinations in the genus Kras¬
cheninnikovia are proposed: K. arborescens (Los-
ina-Losinskaja) Mosyakin, K. compacta (Losina-
Losinskaja) Grubov var. longipilosa (Tsien Cho-po
& Ma Cheng-gung) Mosyakin, and Krascheninni¬
kovia sect. Caudalae (Aellen) Mosyakin.

Krascheninnikovia Gueldenstaedt (1772: 551)
is the correct name for the genus previously com¬
monly known as Eurotia Adanson and Ceratoides
Gagnebin. This nomenclatural problem has been dis¬
cussed by several authors over the years (Lozina-
Lozinskaya, 1930; Reveal & Holmgren, 1972; Gru¬
bov, 1976; Guterman, 1975; Tomsovic, 1990;
Tsvelev, 1993).

Unfortunately, when establishing the genus Cer¬
atoides, Gagnebin (1755: 59) did not write a di¬
agnosis, but rather only mentioned a pre-Linnaean
work by Tournefort (1703). In the latter work the
genus Ceratoides included both an annual species
(later named by Linnaeus as Ceratocarpus aren-
arius L.), and a perennial one, which is conspecific
with Krascheninnikovia ceratoides (L.) Guelden¬
staedt. At the time of its establishment (Gagnebin,
1755) the genus Ceratoides included Ceratocarpus
arenarius, the type of Ceratocarpus L. (1753); thus
Ceratoides is a synonym of Ceratocarpus. The next
available name for the genus is Krascheninnikovia.
[More detailed discussion is provided by Guterman
(1975) and Tsvelev (1993).] In the Flora Reipub-
licae Popularis Sinicae (Tsien Cho-po & Ma Cheng-
gung, 1979) the name Ceratoides was accepted for
this genus. Thus, the following new nomenclatural
combinations are necessary for a forthcoming edition
of the Flora of China.

Krascheninnikovia arborescens (Losina-Los-
inskaja) Mosyakin, comb. nov. Basionym: Eu¬
rotia arborescens Losina-Losinskaja, Izv. Akad.
Nauk SSSR, Ser. 7, Otd. Fiz.-Mat. Nauk 1930,
No. 9: 999. 1930. Ceratoides arborescens
(Losina-Losinskaja) Tsien Cho-po & Ma Cheng-
gung, Flora Reipublicae Popularis Sinicae 25(2):
27. 1979.

Krascheninnikovia compacta (Losina-Losinska¬
ja) Grubov var. longipilosa (Tsien Cho-po &
Ma Cheng-gung) Mosyakin, comb. nov. Bas¬
ionym: Ceratoides compacta (Losina-Losin¬
skaja) Tsien Cho-po & Ma Cheng-gung var.
longipilosa Tsien Cho-po & Ma Cheng-gung,
Flora Reipublicae Popularis Sinicae 25(2): 28.
1979.

When first describing the new variety Ceratoides
compacta (Losina-Losinskaja) Tsien Cho-po & Ma
Cheng-gung var. longipilosa Tsieng Cho-po & Ma
Cheng-gung, the authors did not cite the basionym
for their new combination at the species level (see
Kung Hsien-wu et al., 1978). Thus, the complete
combination (species plus variety) was validated later
in the Flora Reipublicae Popularis Sinicae (Tsien
Cho-po & Ma Cheng-gung, 1979).

Two sections, based mostly on the differences in
the structure of trichomes, were established by Ael¬
len (1952) for the genus Eurotia. The correct names
for these sections, as well as a new combination for
one of them, are given below.

Krascheninnikovia sect. Krascheninnikovia.

TYPE: Krascheninnikovia ceratoides (L.)
Gueldenstaedt (type of the genus).

Eurotia sect. Ecaudatae Aellen, Verh. Naturf. Ges. Basel
63: 267. 1952. TYPE: not designated.

Because in Aellen’s (1952) treatment the section
Ecaudatae included the type of the genus, this name
should be replaced by the appropriate autonym.

Krascheninnikovia sect. Caudatae (Aellen) Mo¬
syakin, comb. nov. Basionym: Eurotia sect.
Caudalae Aellen, Verh. Naturf. Ges. Basel 63:
267. 1952. TYPE: Eurotia compacta Losina-
Losinskaja = Krascheninnikovia compacta
(Losina-Losinskaja) Grubov.

The native North American species previously
commonly known as Eurotia lanala (Pursh) Mo-
quin-Tandon was transferred to Krascheninnikovia
by Meeuse & Smith (1974). If accepted as a distinct

Novon 5: 52-53. 1995.

Volume 5, Number 1
1995

Mosyakin

Asian Krascheninnikovia

53

entity, another North American taxon described by
Rydberg (1912) as Eurotia subspinosa is in need
of the same transfer or a new infraspecific combi¬
nation. However, in view of the current work on
the Flora of North America project, I believe this
nomenclatural decision should be left to American
botanists.

Literature Cited

Aellen, P. 1952. Ergebnisse einer botanisch-zoologisch-
en Sammelreise durch den Iran. Botanische Ergeb¬
nisse II. Chenopodiaceae: Agriophyllum, Esfandi-
aria, Eurotia. Verh. Naturf. Ges. Basel 63: 253-
272.

Gagnebin, A. 1755. Observations faites sur le systeme
des autheurs de botanique et sur l’Ophris minima
C.B. Acta Helv. Phys. Math. 2: 56-75.

Grubov, V. I. 1976. Proposal for conservation of the
genus name Eurotia Adans. against Axyris Linn.
Taxon 25: 362.

Gueldenstaedt, A. I. 1772. Krascheninnikovia , novum
plantarum genus. Novi. Comment. Acad. Sci. Imp.
Petrop. 16: 548-560.

Guterman, W. 1975. Notulae nomenclaturales. Phyton
(Horn) 17 (1-2): 31-50.

Kung Hsien-wu, Chu Ge-ling, Tsien Cho-po, Li An-jen &

Ma Cheng-gung. 1978. The Chenopodiaceae in Chi¬
na. Acta Phytotax. Sin. 16(1): 117.

Lozina-Lozinskaya (Losina-Losinskaja), A. S. 1930. Ma-
terialy po izucheniyu roda Eurotia (Materials for the
study of the genus Eurotia). Izv. Akad. Nauk SSSR,
Ser. 7, Otd. Fiz.-Mat. Nauk 1930, No. 9: 977-
1007.

Meeuse, A. D. J. & A. Smith. 1974. A new combination
in Krascheninnikovia (Chenopod.). Taxon 20: 644.

Reveal, J. L. & N. M. Holmgren. 1972. Ceratoides ,
an older generic name for Krascheninnikovia and
Eurotia. Taxon 21: 209.

Rydberg, P. A. 1912. Studies of the Rocky Mt. Flora —
XXVII. Bull. Torrey Bot. Club 39: 310-328.

Tomsovic, P. 1990. Patri druh Ceratoides latens (Eu¬
rotia ceratoides ) do ceskoslovenske kveteny? (Does
Ceratoides latens (Eurotia ceratoides ) belong to the
flora of Czechoslovakia?). Preslia (Praha) 62: 33-
39.

Tournefort, J. P. 1703 (repr. 1719). Corollarium in-
stitutionum rei herbariae. Lugdunii.

Tsien Cho-po & Ma Cheng-gung. 1979. Ceratoides
(Tourn.) Gagnebin. Pp. 24-28 in Flora Reipublicae
Popularis Sinicae. Vol. 25(2). Beijing.

Tsvelev, N. N. 1993. Zametki o marevykh (Cheno¬
podiaceae) Vostochnoy Evropy (Notes on Chenopo¬
diaceae of Eastern Europe). Ukrayins’k. Bot. Zhurn.
50(1): 78-85.

A New Species of Oserya (Podostemaceae) from Jalisco, Mexico

Alejandro Novelo R.

Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico,
Mexico, D.F. 04510, Mexico

C. Thomas Philbrick

Department of Biological and Environmental Sciences, Western Connecticut State University,
Danbury, Connecticut 06810, U.S.A.

ABSTRACT. A new species of Podostemaceae, Os¬
erya longifolia, from Jalisco, Mexico, is described
and illustrated. Leaf length is the most prominent
feature that distinguishes this species.

RESUMEN. Se describe e ilustra una especie nueva
de la familia Podostemaceae, Oserya longifolia, del
estado de Jalisco, Mexico. La longitud de la hoja es
la caracteristica mas prominente que distingue a
esta especie.

Oserya Tulasne & W eddell is an American genus
of six species, most recently monographed by van
Royen (1954). Like all Podostemaceae, species of
Oserya grow attached to rocks in swift-moving river
currents. Five of the six species ( O. biceps Tulasne
& Weddell, 0. flabellifera Tulasne & Weddell, 0.
minima van Royen, O. perpusilla (Went) van Roy¬
en, O. sphaerocarpa Tulasne) occur in northern
Brazil and northeastern South America, while 0.
coulteriana Tulasne occurs in Mexico.

Little is known of the geographic distributions of
species of Oserya. In fact, four of the South Amer¬
ican species are known only from the type collec¬
tions. At the time of van Royen’s monograph, 0.
coulteriana was known from few collections in the
Mexican states of Jalisco and Michoacan. During
our recent studies of Mexican Podostemaceae we
have found this species to be relatively widespread
in Colima, Nayarit, and Guerrero, in addition to
Jalisco and Michoacan. A collection from southern
Sinaloa is also now known (Novelo & Philbrick,
unpublished).

While conducting field studies of Mexican Podo¬
stemaceae, we made collections that did not cor¬
respond to any described species. These collections
serve as the basis for the description of a new species,
Oserya longifolia Novelo & Philbrick.

Oserya longifolia Novelo & Philbrick, sp. nov.
TYPE: Mexico. Estado de Jalisco: mumcipio
de La Huerta, Rio Purificacion, 4 km al oeste
de La Huerta, 19°30'N, 104°40'W, 300 m,
26 Mar. 1993, Novelo & Philbrick 1166 (ho-
lotype, MEXU; isotypes, MO, NY, WCSU).
Figure 1.

Herbae aquaticae plerumque caulibus stoloniferis ap-
planatis. Folia circinata usque ad 40 cm longa; petiolus
laevis; lamina repetite 2-4(-5)-divisa, divisionibus ultunis
0.3-0.6 mm latis, applanatis, apice acutis. Flores her-
maphroditi, zygomorphi, pedicellati, axillares, solitarii. Te-
pala 3, filamentis alternata, uno in furca duorum stam-
inum affixo. Stamina 2, andropodio portata; antherae el-
lipticae, dorsifixae, per 2 rimas laterales longitudinaliter
dehiscentes. Ovarium 2 carpellis, 2 stylis 0.7-1.2 mm
longi. Fructus 2 valvis, quaque valva 3-costata.

Aquatic herbs usually with stoloniferous and flat¬
tened stems 0.8-1.0 mm diam., strongly adhering
to rocks. Leaves alternate, circinate, up to 40 cm
long, petiole 2-17 cm long, cylindrical, smooth,
0.7-1.6 mm diam., with a broadened base; blade
2—4(—5) divided, the ultimate divisions 0.3-0.6 mm
wide, flattened, apex acute. Flowers hermaphroditic,
zygomorphic, pedicellate, borne singly, axillary, pro¬
tected by a spathella; spathella up to 7 mm long,
thin clavate. Pedicels 3.5-8.0 mm long, elongating
during anthesis, not expanded at capsule base, re-
pals 3, alternate with the filaments, free, membra¬
nous, subulate, one of them attached in the fork
between the two stamens; lateral tepals 0.9-1-2 mm
long, medial tepal 0.7-0.9 mm long. Stamens 2,
borne by an andropodium, andropodium 0.8-1-3
mm long, elongating during anthesis; filaments 1 -0"
1.8 mm long, subulate, flattened, elongating during
anthesis, deciduous; anthers 0.4-0.6(—0.8) mm long,
elliptic, dorsifixed, 2-celled, dehiscing longitudinally
by 2 lateral slits. Ovary superior, 1.7—2.1 mm long,

Volume 5, Number 1
1995

Novelo & Philbrick
Oserya longifolia

55

Figure 1. Oserya longifolia Novelo & Philbrick. Drawings based on the holotype. —A. General habit of plant with
several parts of the leaves interrupted (longest leaf 40.1 cm). — B. Top of the stem with several circulate new leaves.
—C. Detail of a single leaf (26.7 cm long). —D. Several stages of flower development from between leaf bases,
before (left) and after (right) the spathella is ruptured. — E. Lateral view of the mature flower showing orientation of
the styles, anthers, and tepals at the time of dehiscence. —F. Abaxial view of the mature flower showing the
andropodium and tepals. —G. Mature styles with receptive stigmas. —H. Cross section of ovary showing placenta
and numerous ovules. —I. Abaxial view of a single anther. —J. Adaxial view of a single anther. —K. Cross section
of anther. —L. Mature fruit. —M. Cross section of a capsule showing ribs and dehiscence points of the capsule
valves. —N. Seed.

56

Novon

ellipsoid, 2-locular; styles 2, 0.7-1.2 mm long, free,
conical, ovules numerous, placenta axile. Fruit 1.7—
2.6 mm long and 0.8-1.2 mm diam., 2-locular
capsule, suture margins thickened; valves 2, each
3-ribbed. Seeds 0.24-0.26 mm long, 0.16-0.20
mm wide, obovoid. Pollen 14.88 gm (SD 0.9, N =
30) diam., tricolpate.

Oserya longifolia grows submerged in the swift
currents of river rapids, attached directly to rocks
of various sizes. No other species of Podostemaceae
were observed growing with Oserya longifolia, al¬
though a large population of Eichhornia crassipes
(Martius) Solms-Laubach occurs in areas of slow
current in association with 0. longifolia.

Thus far, Oserya longifolia has only been col¬
lected in the Purificacion River, in areas of full sun.
The largest leaves (up to 40 cm long) were found
on plants that are submerged in areas of very fast
current. In areas of slow current, or when the plants
are exposed as the water level drops, the terminal
portions of the leaves fall away, leaving the short,
coarse bases.

Two species of Oserya are now known from Mex¬
ico: O. coulteriana and O. longifolia. These species
are readily distinguished. Oserya longifolia is the
larger, more robust of the two (Table 1). The longer
leaf of O. longifolia (15-40 cm vs. 2-10 cm for
O. coulteriana) is particularly notable. However,
the tepal that occurs between the anthers at the
apex of the andropodium is shorter in O. longifolia
than in O. coulteriana (Table 1). Two qualitative
features also distinguish the species: the petioles of
O. longifolia are cylindrical in cross section, where¬
as those of O. coulteriana are markedly flattened,
and distinct petiolar spines are common in O. coul¬
teriana whereas they are lacking in O. longifolia.

Leaf length also distinguishes Oserya longifolia
from the five South American species of the genus.
Leaf length of all South American species ranges

Table 1. Vegetative and floral characteristics that
distinguish Oserya longifolia from O. coulteriana.

Characteristic

O. longifolia

0. coulteriana

Petiole shape (cross

cylindrical

flattened

section)

Petiolar spines

absent

usually present

Leaf length

15-40 cm

2-10 cm

Width of ultimate leaf

0.3-0.6 mm

0.1-0.3 mm

division

Spathella length

5-7 mm

3-6 mm

Pedicel length

3.5-8.0 mm

2.0-6.0 mm

Length of medial tepal

0.7-0.9 mm

1-1.3 mm

Andropodium length

0.8-1.3 mm

0.2-0.5 mm

Anther length

0.4-0.6

0.75-1.10 mm

Fruit length

(-0.8) mm
1.7-2.6 mm

1.8-2 mm

from 2 to 3 cm, while leaves of O. longifolia range
from 15 to 40 cm. In addition, all five South Amer¬
ican species have a single stamen; in contrast, both
O. longifolia and O. coulteriana possess two sta¬
mens (occasional plants of the latter species have a
single stamen).

Acknowledgments. We thank Fernando Chiang
for providing the Latin description, Fernando Chiang,
Ronald Aajkar, and Donald H. Les for helpful com¬
ments on the manuscript, and Albino Luna for draw¬
ing Figure 1. Support for this study was provided
by The Intercambio Academico Office at the Na¬
tional University of Mexico, a grant to CTP by the
National Science Foundation, and a grant to ANR
by the Consejo Nacional de Ciencia y Tecnologia.

Literature Cited

Royen, P. van. 1954. The Podostemaceae of the New
World. III. Acta Bot. Neerl. 3: 215-263.

Impatiens kulamavuensis , a New Species of Balsaminaceae from India

A. G. Pandurangan and V. J. Nair

Botanical Survey of India, TNAU—Campus, Coimbatore—641 003, India. Present address
of A. G. Pandurangan: Tropical Botanical Garden and Research Institute, Thiruvananthapuram-
695 562, India. Present address of V. J. Nair: The Herbarium, Royal Botanic Gardens, Kew,
Richmond, Surrey, TW9 3AB, U.K.

ABSTRACT. Impatiens kulamavuensis , a distinct
new species from India allied to I. parasitica Bed-
dome, is described.

Impatiens kulamavuensis A. G. Pandurangan
& V. J. Nair, sp. nov. TYPE: India. Kerala:
Idukki Dist., Meenmutty-Kulainavu, ± 700 m,
6 Oct. 1983, A. G. Pandurangan 82037 (ho-
lotype, CAL; isotype, MH). Figure 1.

Ab Impatiente parasitica Beddome caulibus stoloni-
feris, foliis glandulosis, nervis lateralibus non manifestis;
sepalis lateralibus et petalis pilosis; stigmatibus curvatis,
sericeo-pubescentibus et seminibus pyriformibus differt.

Epiphytic stoloniferous herbs; stems short, stout,
fleshy, less than 1 cm long. Leaves alternate; petioles
gland-dotted, 2-7 mm long, with 2 prominent stip-
itate glands near the base of the blade; lamina ovate-
lanceolate, 2-5.5 x 1.5-2.5 cm, membranous,
gland-dotted above, glabrous below, deep green, the
base cuneate, the apex acuminate, margins entire
to obscurely crenulate-serrate, crenules 2-4 mm
apart; lateral veins 4—6 pairs, indistinct. Inflores¬
cence a 1-3-flowered raceme; peduncles 2.5-5 cm
long; pedicels 3.5-5 cm long. Flowers greenish yel¬
low. Lateral sepals linear, 8-10 x 2-3 mm, hairy
outside, glabrous inside. Lower sepal saccate, com¬
pressed, 12-15 x 5-7 mm, glabrous; spurs strongly
curved, 7-8 x 3-5 mm, tip obtuse. Dorsal petal
erect, orbicular to ovate, 6-7 x 8-9 mm, hairy at
base. Lateral petals united close to the base, 3-lobed,
upper lobes smaller in size, 7-8 x 9-10 mm. An-
droecium 3-3.5 mm long. Ovary ellipsoid, glabrous,
5-6 x 3-3.5 mm; ovules many, 1-seriate; style
short; stigma curved, with an obtuse tip, silky pu¬
bescent. Capsules ellipsoid, 10-12 x 4-5 mm, gla¬
brous. Seeds pear-shaped, smooth. Flowering and
fruiting October-December.

Distribution. Endemic to Meenmutty-Kulamavu,
Idukki Dist., Kerala, India.

Table 1. Comparison of major differences between
Impatiens parasitica Beddome and Impatiens kulama¬
vuensis A. G. Pandurangan & V. J. Nair.

I. parasitica

I. kulamavuensis

Stems 10-12 cm long,
not stoloniferous, mo-
niliform

Stems less than 1 cm
long, stoloniferous, not
moniliform

Leaves glabrous on both
surfaces or glabrescent
above; lateral nerves
prominent

Lateral sepals glabrous
outside, glandular in¬
side at base

Leaves glanduliferous
above, glabrous be¬
neath; lateral nerves
indistinct

Lateral sepals hairy out¬
side, glabrous inside

Petals glanduliferous in¬
side at base

Petals hairy inside at base

Stigmas straight; stigmatic
lobes 5-toothed, not
pubescent

Seeds rounded to ellipsoid

Stigmas curved; lobes not
5-toothed, obtuse, silky
pubescent

Seeds pear-shaped

Habitat and ecology. Epiphytic herbs, growing
on moss-clad tree trunks at an altitude of 600-700
m. The host trees are Elaeocarpus tuberculatus
Roxburgh and Pygium wightiana Blume.

Impatiens kulamavuensis was collected in the
Idukki hydroelectric project area, Kerala. It is allied
to I. parasitica Beddome, but differs in the char¬
acters listed in Table 1.

Paratype. INDIA. Kerala: Idukki dist., Meenmutty,
± 600 m, 30 Dec. 1983, A. G. Pandurangan 62585
(MH).

Acknowledgments. We thank N. P. Balakrish-
nan. Joint Director, and N. C. Nair, former Joint
Director, Botanical Survey of India, for help and
encouragement.

Novon 5: 57-58. 1995.

58

Novon

7 mm

1

J

iomrr\

7 mm

Figure 1. lmpatiens hulamavuensis A. G. Pandurangan & V. J. Nair. —A. Habit. —B. Flower. —C. Lateral
sepals. D. Lip. E, G. Lateral petals, outer and inner views. —F. Dorsal petal. —H. Stamens. —I. Pistil. —J-
Transection of ovary. —K. Fruit. —L. Seeds.

Two New Taxa for the Flora of Thailand

John A. N. Parnell

School of Botany, Trinity College, Dublin 2, Ireland

ABSTRACT. The new subspecies Mitrasacme ero-
phila subsp. grandijlora (Loganiaceae) and Pedi-
cularis lhailandica subsp. parvula (Scrophularia-
ceae) are described.

Mitrasacme is a widespread Australasian genus,
represented in Thailand by three undercollected spe¬
cies. In order to make a name available for the
forthcoming Flora of Thailand account, the follow¬
ing new subspecies is described.

Mitrasacme erophila Leenhouts subsp. gran-
diflora J. Parnell & 0. Griffin, subsp. nov.
TYPE: Thailand. Chiang Mai, Omkoi, ca. 35
km from pine improvement centre to Omkoi,
29 Dec. 1978, C. Niyomdham, M. Suangtho
& B. Sangkhachand 168 (holotype, L; iso¬
type, AAU).

A subsp. erophila foliis glandulis minutis instructis,
corolla grandiore, stamenibusque basim corollae versus
affixis differt.

Herb to 20 cm. Stem < 1 cm high, unbranched,
minutely papillose. Leaves 2—4 pairs at the base of
the plant, ovate, < 4 mm long, with a single distinct
mid-vein; apex acute to blunt. Inflorescence ter¬
minal, lax, branched and umbellate, many-flowered;
peduncles 1-3 arising from the stem with 1-5 min¬
ute, triangular bracteoles either sterile or subtending
the inflorescence branches; pedicels to 4.2 cm, gla¬
brous. Flowers white. Calyx 1.5-2 mm long, cam-
panulate; lobes 0.7-1 mm. Corolla 5-6.7 mm long,
campanulate; lobes 1.2-1.7 mm, oblong, mucro-
nate. Fruit 1.5-2 mm high; styles connate at the
tips; becoming freer with maturity.

Mitrasacme erophila subsp. grandijlora differs
from subspecies erophila in its leaves, which bear
minute peltate scales on their upper surface, its
longer corolla (the corolla in subspecies erophila is
3-3.7 mm long) with larger, rounded corolla lobes
(the corolla lobes in subspecies erophila are oblong
and 1.2-1.7 mm long) and in the position of the
stamens’ insertion in the corolla tube (the stamens
in subspecies erophila are attached at or above the
center). Subspecies grandijlora is known only from
the type collection.

Pedicularis in Thailand has been recently revised
for the Flora of Thailand, where four species are
recognized (Yamazaki, 1990). This account does
not fully document the great variability seen in Thai
material, and some herbarium material does not
match that of any of the described species. Material
collected by the first joint expedition of the Royal
Botanic Gardens, Kew, Trinity College, Dublin, and
Khon Kaen University, although conspecific with
Pedicularis thailandica, is sufficiently distinct to
require circumscription.

Pedicularis thailandica Yamazaki subsp. par¬
vula J. Parnell, subsp. nov. TYPE: Thailand.
Chiang Mai, Doi Inthanon National Park, forest
trail at km 42 along summit road, hill evergreen
forest, 7 Oct. 1990, P. Chantaranothai, J.
Parnell, D. Simpson & R. Pooma 90/613
(holotype, TCD).

A subsp. thailandica statura minore; caulibus vix lan-
atis; foliis minutis una cum floribus ternis verticillatis et
corolla alba differt.

Herb to 17 cm. Stems erect, simple, subglabrous.
Leaves 3-verticillate; petioles 2 mm, sparsely lanate;
blades ovate, deeply pinnate, the pinnules crenate.
Flowers solitary, 3-verticillate; pedicels 0.2-0.5 mm.
Calyx tubular-campanulate, 5.5 x 4 mm, sparsely
lanate, 5-toothed; teeth rounded, ca. 2 mm long.
Corolla white, 18-20 mm long, tube straight, 12
x 2 mm, densely glandular-pilose especially on the
lobes and hood; hood (galea) ca. 3.5 mm; straight
near the mouth, abruptly curved near the tube, two¬
toothed; lower lip three-lobed, all lobes broadly ob¬
long. Stamens with glabrous filaments ca. 10 mm
long; anthers deltoid, ca. 1.5 mm long. Mature cap¬
sule unknown. Subspecies parvula is known only
from the type collection.

Pedicularis thailandica subsp. parvula differs
from subspecies thailandica in being only half its
height, with the main stem almost glabrous (vs.
spreadingly lanate); the leaves being much smaller
and not duplicate-serrate; and the flowers being in
whorls of three (vs. four) with a white (vs. yellow)
corolla.

Novon 5: 59-60. 1995.

60

Novon

Acknowledgments. I thank P. Chantaranothai
and R. Pooma, who were jointly responsible for
arranging the field excursion to Doi Inthanon, D.
A. Webb for helping with the Latin, and D. Mid¬
dleton for commenting on a draft of this paper.

Literature Cited

Yamazaki, T. 1990. Scrophulariaceae. In: T. Smitinand
& K. Larsen (editors), Flora of Thailand, 5: 139-
238.

New Species of Siparuna (Monimiaceae) I.
Four New Species from Ecuador and Colombia

Susanne S. Renner and Gerlinde Hausner
Institute of Systematic Botany, University of Mainz, Bentzel-Weg 2,
D-55099 Mainz, Germany

Abstract. Siparuna croatii and S. palenquensis
from central and Pacific Ecuador, respectively, S.
gigantotepala from Pacific Colombia and Ecuador,
and S. harlingii from the eastern Andean slopes of
Colombia and Ecuador are described, illustrated, and
discussed as to their relationships with morpholog¬
ically similar species.

Comprising an estimated 150 species, Siparuna
is the largest and least understood genus of the
Monimiaceae. The genus was last revised by Perkins
(1901, 1911), whose typological species concept
overstates diversity in certain areas, such as plan-
altan and southeastern Brazil. In Colombia, Ecuador,
and Peru, however, increased collecting activity over
the past 20 years has turned up numerous new
species, and it now appears that most species of
Siparuna are Andean in distribution. Material in
the major relevant herbaria (acronyms following
Holmgren et a]., 1990), such as AAU, BM, COL,
F, G, GB, GH, GOET, K, M, MA, MJG, MO, NY,
OXF, P, QCA, QCNE, S, SEL, TCD, UC, US, W,
WU, and Z, currently amounts to over 2500 col¬
lections representing approximately 100 species from
the Andes. Continued discovery of new taxa in upper
Amazonia strongly suggests the need for additional
collecting. As a result of ongoing work toward a
monograph of Siparuna, we herein describe four
new species in order to make their names available
for the forthcoming treatment of the Monimiaceae
for the Flora of Ecuador. Work on the Peruvian
species is still in progress, rendering description of
several apparently new species extending south be¬
yond Ecuador inadvisable.

Flowers of Siparuna are small and unisexual, and
plants are monoecious or dioecious (Perkins, 1901;
Feil, 1992). The flowers have an obconical or ur-
ceolate floral cup in which the free carpels or sta¬
mens are more or less completely enclosed (e.g..
Figs. 1A, and 4B and F) and usually minute (1-4
mm) tepals. A feature characteristic of the genus is
that the floral apex forms a roof, called a velum,
with a central pore through which the upper parts
of the stamens or styles protrude (for details of floral

ontogeny and a morphological interpretation of the
velum, see Endress, 1980). In the male flowers, the
velum is more or less raised at anthesis and the pore
is relatively large (Fig. 1A); in the female flowers,
the inner margin of the velum is always distinctly
raised and the pore is narrower (Fig. 1C).

The stamens are sessile and initiated centripetally,
the innermost ones often remaining small or even
staminodial. The anthers are two-locular and open
around the perimeter of both pollen sacs except at
the distal end (Figs. IB, G, N; 4G; compare also
Endress & Hufford, 1989, figs. 97-100). Dehis¬
cence results in a flaplike structure that is hinged
distally and bent upward. In fully anthetic male
flowers, these flaps usually can be seen with the
unaided eye.

Anthesis typically lasts 9-16 days in male flowers
and 16-24 days in females (Feil, 1992), and the
fertilized flowers then expand. Measurements of flo¬
ral parts and descriptions of their shape are therefore
of limited value. More diagnostically useful are the
shape, color, and pubescence of the leaves and the
number of stamens and styles.

Siparuna croatii Renner & Hausner, sp. nov.
TYPE: Ecuador. Cotopaxi: 63.4 km SE of
Quevedo, 6 km NW of El Corazon, primary
forest on steep slopes near a waterfall above
Rio Angamarca, 1030 m, 4 Apr. 1983 (female,
fl), T. B. Croat 55752 (holotype, QCNE; iso¬
types, AAU, MO, QCA). Figures 1A-D, 2.

Species ramulis quadrangularis valde sulcatis et foliis
late ellipticis (18-42 x 10-20 cm) a congeneribus div-
ersa.

Dioecious shrub or tree, 4-5 m tall, partly with
a liana-like habit, the twigs quadrangular, deeply
sulcate, and khaki-colored due to a dense indumen¬
tum of minute stellate hairs. Leaves opposite, the
petioles 4-5 cm long; the lamina drying brownish
to olive-green, chartaceous, elliptic to broadly ellip¬
tic, 18-42 x 10-20 cm, the base obtuse or sub¬
acute, the apex shortly apiculate, the lower surface
with minute appressed stellate hairs, the upper sur-

Novon 5: 61-70. 1995.

62

Novon

Volume 5, Number 1
1995

Renner & Hausner
Siparuna

63

face scantily stellate-pubescent to glabrescent and
rather smooth, with 22-24 pairs of secondary veins,
the secondary and tertiary veins distinctly promi-
nulous and yellow below, the margin inconspicuously
denticulate. Cymes of both sexes axillary in groups
of 2-4, ample and multiflorous, 3-5 cm long, with
minute yellowish khaki stellate hairs. Male flower at
anthesis 1.9-2.1 x 1.8-2.1 mm, the floral cup
urceolate to globose with minute stellate hairs, the
floral roof moderately raised, glabrous and drying
dark brown, the 4-5 tepals obtusely triangular and
0.4-0.6 mm long, when fresh greenish; stamens 6,
fleshy and containing white globules (large oil cells),
the 4 outer ones slightly exserted at anthesis. Female
flowers at anthesis of the same size and shape as
the males but the floral roof raised to a cylindrical
bulge separated by a distinct groove from a second
innermost tube sheathing the style bases; the styles
5-7, basally united. Fruit fleshy and red when fresh;
mature fruit and seeds unknown.

Distribution, habitat, and phenology. Endemic
in central Ecuador; growing in primary montane
forest on steep slopes and in secondary scrub at
elevations of 1000-1400 m; collected flowering and
with young fruits in April and May.

Siparuna croatii differs from all other species in
the genus in its khaki-colored, deeply sulcate, qua¬
drangular branchlets, olive-green leaves with sec¬
ondary and tertiary veins distinctly raised below,
and, for the genus, ample, khaki-colored inflores¬
cences with small flowers.

Paratype. ECUADOR. Manabi: trail from El Corazon
to Facundo Velo, 1-3 km S of El Corazon, 1400 m, 17
May 1980 (male, fl), Harling & Andersson 19216( AAU,
GB, MJG, QCA).

Siparuna palenquensis Renner & Hausner, sp.
nov. TYPE: Ecuador. Pichincha: Hcda. Cov-
adonga on Rio Pilaton, 1000 m, 2 July 1955
(male, fl), E. Asplund 16764 (holotype, S; is¬
otypes, MJG, QCA). Figures 1K-N, 3.

A Siparuna eggersii Hieronymus foliis obovatis (lb-
27 x 10-17 cm) dilfert.

Dioecious tree, 4-10 m tall, young branchlets
densely pubescent with minute stellate hairs, older

branches dark brown and subglabrous, terete. Leaves
opposite, the petioles 1.5-4.5 cm long; the lamina
drying olive-green or brown, chartaceous, obovate,
16-27 x 10-17 cm, the base cordate, truncate,
or rounded, the apex shortly apiculate, both surfaces
rather rough and scantily to moderately stellate-
pubescent, with 10-13(-16) pairs of secondary veins,
veins to the third order impressed above, distinctly
visible below, the margin minutely denticulate. Cymes
of both sexes axillary in groups of 2-4 and multi¬
florous (less so in the females), 1.5-4 cm long, and
almost glabrous. Male flower at anthesis ca. 5.5-
6.3 x 5-5.5 mm, the floral cup obconical with
scant stellate hairs, the floral roof strongly raised,
glabrous and drying dark brown or black, the 4-5
tepals nearly completely fused, fresh greenish yel¬
low; stamens 6, often with 4 outer and 2 central
ones, fleshy and containing white globules (oil cells),
the outer stamens distinctly exserted at anthesis.
Female flowers at anthesis of the same size and shape
as the males; the styles ca. 15-30, basally united.
Fruit fleshy, ca. 1.3-2 x 1-2 cm, fresh immature
light green with sparse blackish scales, mature yel¬
lowish or reddish orange and smelling strongly of
lemon; the 15-30 seeds distinctly visible in dried
fruits, verrucose, gray, and with a red aril when
fresh.

Distribution, habitat, and phenology. Restrict¬
ed to Pacific Ecuador and described by collectors
as infrequent or rare; growing in disturbed premon-
tane forests on steep slopes or on cliffs; sea level to
1200 m; collected flowering and fruiting mainly
from April to September.

The suggested relative, Siparuna eggersii Hi¬
eronymus, has smaller, obovate leaves (usually 12-
15 x 8-10 cm) with cuneate bases. The two species
have previously been confused, as, for example, in
the florula of the Rio Palenque Biological Station by
Dodson & Gentry (1978), which describes and il¬
lustrates specimens of S. palenquensis as S. eg¬
gersii.

Paratypes. ECUADOR. Esmeraldas: NE across Rio
Blanco from Quininde, Little 6230 (F, K, US). Los Rios:
Rio Palenque Biological Station, Dodson 5134 (AAU, F,
MO, QCA, SEL, US), Dodson 5723 (MO, QCA, SEL,

Figure 1. A-D. Siparuna croatii Renner & Hausner (A, B, Harling A Andersson 19216; C, D, Croat 55752).
—A. Part of male inflorescence. —B. Stamen. —C. Female flowers. —D. Young fruit. E-I. Siparuna gigantotepala
Renner & Hausner (E, Barfod et al. 48903; F, G, Cuatrecasas 15746; H, I, Rubio et al. 999). — E. Young male
flower. —F. Mature male flower. —G. Stamens. — H. Female flower. —I. Young fruit. K-N. Siparuna palenquensis
Renner & Hausner (K, 0llgaard 98054; L, Jaramillo 44; M, Dodson 5134; N, Asplund 16764). —K. Female
flowers. —L. Mature fruit. —M. Male flower. —N. Stamens. The following parts share the same magnification: A,
C, E, F, H, I, K, M; B, G, N; and D, L.

64

Novon

Figure 2. Female specimens of Siparuna croatii Renner & Hausner (Croat 55752 ). —A. AAU isotype. — B-
MO isotype.

Volume 5, Number 1
1995

Renner & Hausner
Siparuna

65

Figure 3. Female specimens of Siparuna palenquensis Renner & Hausner. —A. Croat 55702. B. 0llgaard
98054, AAU, paratype.

66

Novon

US), Dodson 5930 (MO, NY, SEL, US), Gentry & Dod¬
son 17970 (MO, QCA). Pichincha: Pichincha, ca. 8
km SE of La Aurora, km 7 on Sto. Domingo-Quevedo
rd., at bridge over Rio Baba, 0llgaard 98054 (AAU,
QCA); Tinalandia, 9.6 km E of Santo Domingo de los
Colorados, above Rio Toachi, Croat 55702 (AAU, MO);
km 3 of Toachi-Las Pampas rd., Dodson & Gentry
13708 (F, MO, QCNE, SEL); old rd. Quito-Santo Do¬
mingo, 2-9 km NE of turn-off to old rd. of AUuriquin,
Luteyn et al. 8736 (AAU, NY, QCA, QCNE); Aloag-
Santo Domingo, Toachi, at the confluence between Rio
Pilaton and Rio Toachi, Sparre 13829 (MJG, S). Co¬
topaxi: island in Rio San Pablo, near La Mana, Webster
22727 (UC).

Siparuna harlingii Renner & Hausner, sp. nov.
TYPE: Ecuador. Morona-Santiago: 7-8 km N
of Gualaquiza on rd. to Indanza, 1500 m, 16
Apr. 1985 (male, fl), G. Harling & L. An-
dersson 24175 (holotype, QCA; isotypes, AAU,
GB). Figure 4.

Species Siparunae asperae (Ruiz & Pavon) A. DC.
proxima, cujus flores foliorumque texturam et colorem
habet. Differt floribus fructibusque minus pubescentibus
et pedicellis paullo incrassatis.

Dioecious tree or shrub, sometimes scandent, 2-
6(-12) m tall, the young and older branchlets dense¬
ly pubescent with stellate hairs, subangular. Leaves
opposite and those of a pair slightly unequal in size,
the petioles (2-)2.5-4(-7) cm long; the lamina dry¬
ing dark green to dark brown, papery, brittle, and
subbullate to rather smooth, elliptic to broadly el¬
liptic, sometimes narrowly elliptic, 18-35 x 9-22
cm, the base truncate to cordate, occasionally round¬
ed or acute, the apex acuminate, both surfaces
densely stellate-pubescent, glabrescent on the upper
surface, with (8—)12— 13(—15) pairs of secondary
veins, these distinctly visible on the upper surface
and slightly raised and yellowish brown pubescent
on the lower surface, the tertiary venation dense
and distinctly visible, the margin finely denticulate.
Cymes of both sexes axillary in groups of 2-4, much-
branched (less so in the females), 2-5 cm long, with
minute stellate hairs. Male flower at anthesis ca. 4-
4.5 x 4-5 mm, the floral cup obconical with minute,
pale yellowish stellate hairs, the floral roof distinctly
raised, glabrous and drying black, the 4-5 tepals
obtuse or triangular and 1-1.5 mm long, glabrous
or occasionally with a few stellate hairs, when fresh
greenish yellow to creamy white, turning red; sta¬
mens 6, often with 4 outer and 2 central ones, fleshy
and containing whitish globules (oil cells), the outer

stamens distinctly exserted at anthesis. Female flow¬
ers at anthesis of the same size and shape as the
males but the floral roof centrally conspicuously
raised; the styles 20-30, free. Fruit fleshy, 1.5 x
1.5 cm (when fresh to 2 cm diam.), when immature
green with pink spots, when mature light red and
with a strong lemon smell when crushed, glabrescent;
the 20-30 seeds distinctly visible in dried fruits,
verrucose, gray and with a red aril when fresh.

Distribution, habitat, and phenology. On the
eastern Andean slopes from Putumayo and Caqueta
in Colombia to Napo, Pastaza, Morona-Santiago,
and Zamora-Chinchipe in Ecuador (and expected
in Loja); usually collected in disturbed terra firme
forest from 300 to 2000 m altitude; in Ecuador
flowering and fruiting year-round.

Siparuna harlingii can only be confused with S.
aspera from which it differs especially in the fruit
and pedicels, which in fruit remain slender in S.
harlingii and become fleshy and almost part of the
fruit in S. aspera. The two also differ in pubescence,
S. harlingii having generally much shorter hairs,
and in the leaf venation, S. harlingii having (8-)
12— 13(— 15) lateral nerves, S. aspera (15-)16-20(-
26). Finally, the two may be distinguished by the
number of styles (20-30 in S. harlingii vs. (10-)
12— 15(— 18) in S. aspera).

As is the case with several species of Siparuna,
the crushed leaves of this plant are rubbed on the
body by the Quichua Indians in Ecuador to cure
“mal aire,” a general term used for various diseases
of the nervous system and stomach. The Quichua
name of the species is “Malaire panga,” panga
meaning leaf ( Neill & Palacios 6993).

Paratypes. ECUADOR. Zamora-Chinchipe: horse-
trail Guadalupe-San Jose de Yacuambi, along Rio Ya-
cuambi, potreros and riverside woods, Harling & An-
dersson 13937 (AAU, GB). Tungurahua: between Ban¬
os and Rio Verde, Acosta-SoKs 10267 (F); Rio Topo,
Harling et al. 10073 (AAU, GB); 30 km on rd. from
Mera towards Banos, Lawesson et al. 43293 (AAU,
QCA); Rio Margaritas, Penland & Summers 141 (F, GH,
NY). Napo: Tena, Asplund 8938 (QCA, S); Tena, rd.
to Archidona, Harling 3660 (S); Mission Shandia, Jatun
Yaku River, Barclay 4932 (COL); Puerto Napo, Benoist
4757 (P); Puerto Napo, path to Latas, Harling 3537
(S), Ceron & lguago 5518 (AAU, QCNE); 8 km down¬
river from Misahualli, Neill & Palacios 6993 (AAU, F,
MO, QCNE); km 32-35 Puyo-Tena rd., Jorgensen et
al. 61248 (AAU, QCA); Zatzayacu, Mexla 7097 (NY,
UC, US). Pastaza: Mera, Asplund 18357 (S), Harling
et al. 7807 (AAU, GB), 7840 (AAU, GB). Morona-

Figure 4. Siparuna harlingii Renner & Hausner (A, Harling 928; B, Cuatrecasas 11167; C, Harling & Andersson
13937; D, Neill & Palacios 6993; E-G, Harling & Andersson 13917). —A. Habit. —B. Female flower. C.

Volume 5, Number 1
1995

Renner & Hausner
Siparuna

67

Young fruits. —D. Mature fruit. —E. Male inflorescence. —F. Male flower. - G. Stamen. The following parts share
the same magnification: A, E; B, F; and C, D.

68

Novon

Santiago: rd. Limon-Macas, 96 km NE of Limon, Boh-
lin et al. 1477 (GB, QCA); near Mendez, Camp E-851
(NY, S), Harling 928 (S); Mendez-Limon rd., ca. 3.2
km S of turnoff to Mendez, Dorr & Valdespino 6345
(AAU, NY, QCA, QCNE); Rio Tutanangosa, rd. Sucua-
Huarani, Holm-Nielsen 20508 (AAU). Zamora—Chin-
chipe: rd. Zamora-Zumba, km 5-12, Harling & An-
dersson 13917 (AAU, GB); 10 km S of Zamora on rd.
along left shore of Rio Jamboe, Harling & Andersson
24014 (AAU, GB, QCA); rd. La Saquea-Yacuambi, 1
km N of Chapintza, Harling & Andersson 23893 (AAU,
GB, QCA); Zamora, Harling 5930 (NY, S), Knight 713
(S); Cumbaratza, Jaramillo & Winnerskjold 5922 (NY,
QCA). COLOMBIA. Putumayo: Rio Guamues, San An¬
tonio del Guamues, Cuatrecasas 11167 (F, US); Mocoa,
Cuatrecasas 11278 (F, US), Schultes 2031 (F, K, US),
Schultes & Cabrera 19064 (US); Rio Mocoa drainage
at San Antonio, Ewan 16704 (US); Puerto Umbria, King
1764 (BM, F, K, MO, NY, S, US); near Rio Putumayo,
Soejarto et al. 1277 (US). Caqueta: 62 km SE of
Guadalupe, along rd. to Florencia, Davidse et al. 5633
(COL, NY).

Siparuna gigantotepala Renner & Hausner, sp.
nov. TYPE: Ecuador. Carchi: San Marcos val¬
ley, 600 m, 20 Nov. 1983 (male, fl), A. Barfod,
L. P. Kvist & D. Nissen 48903 (holotype,
QCA; isotype, AAU). Figures 1E—I, 5.

Siparuna subscandens A. C. Smith tepalis longe acu-
tatis (2-3 mm) conspicuis distinguenda.

Dioecious shrub or treelet, 2-3(-5) m tall, bran-
chlets densely covered with minute sessile stellate
hairs, quadrangular and slightly sulcate. Leaves op¬
posite, the petioles 2.5-4.5 cm long; the lamina
drying umber, chartaceous, obovate, 18.5-33 x
8.5-14 cm, the base cuneate to acute, the apex
cuspidate with the tip (l-)1.5-2(-3) cm long, both
surfaces densely covered with minute stellate hairs,
with 9— 10(—13) pairs of secondary veins, veins to
the third order slightly raised above and distinctly
raised below, the margin doubly serrate or subentire.
Cymes of both sexes axillary in pairs and multiflorous
(less so in the females), 1.5-5(-9) cm long, densely
covered with minute stellate hairs. Male flower at
anthesis ca. 2.8—3.2 x 2.5-3 mm, the floral cup
obconic with stellate sessile hairs, a few of these also
on the inside of the tepals and the floral roof, the
floral roof distinctly raised, the (4-)5-6 tepals ob¬
long and 2-3 mm long, when fresh greenish cream
or white; stamens 4-5(-6), the outer ones somewhat
exserted at anthesis. Female flowers at anthesis of
the same size and shape as the males but the velum
distinctly raised to a central tube surrounding the
style bases; the styles 5-8, free. Fruit fleshy, ca.
1.3-2 x 1-2 cm, subglobose and crowned by the
persistent tepals, when mature red or purple with
whitish spots and strongly lemon-scented; the ca.
5-8 seeds verrucose, gray and with a red aril when
fresh.

Distribution, habitat, and phenology. Collected
in very humid primary pluvial forests in Pacific
Colombia and Ecuador at elevations of 20-1500 m;
in Esmeraldas and Carchi collected flowering and
fruiting from November to April.

Siparuna gigantotepala differs from all other
Andean species in the elongate, oblong tepals that
persist in fruit. A few specimens of S. gigantotepala
(e.g., Forero et al. 6740) possess small domatia at
the leaf bases. In this they resemble the western
Ecuadorean S. eggersii Hieronymus, the western
Colombian S. subscandens A. C. Smith, and the
Panamanian S. domatiata A. H. Gentry. All three
differ from S. gigantotepala in lacking elongate
tepals. Siparuna eggersii also has smaller, paler
green-drying leaves that lack the pronounced drip
tip of S. gigantotepala, S. subscandens has nar¬
rower leaves, and S. domatiata has much larger
domatia (as far as known) than S. gigantotepala.
There is one Amazonian species, S. macrotepala
Perkins, which has similarly conspicuous tepals. It
differs in pubescence (simple hairs in S. macrotepala
vs. stellate hairs in S. gigantotepala), number of
secondary veins in the leaves (5-6(-8) in the first
vs. 9— 10(— 13) in the second), and number of styles
(9-12 in the first vs. 5-8 in the second). Moreover,
S. gigantotepala has free styles, while those of S.
macrotepala are united into a tube.

The Coaiquer (Awa) Indians inhale the penetrat¬
ing smell of the fruits and leaves of S. gigantotepala
to clear the nasal passages (A. Barfod et al. 48903).
The local names “Limon de monte” or “Diablo de
monte limon” also refer to the strong smell of the
fruits.

Paratypes. ECUADOR. Esmeraldas: Eloy Alfaro,
Reserva Cotacachi-Cayapas, Parroquia Luis Vargas Tor¬
res, Rio Santiago, Tirado et al. 804 (QCNE); trail to Rio
Mataje Awa encampment from Rio Palavi, Hoover et al.
4007 (AAU, MO, QCA); Reserva Awa, 25 km NW of
El Chical, Municipio Maldonado, Rubio et al. 999 (AAU,
MO, QCNE), Rubio et al. 1036 (AAU, MJG, MO, QCNE);
Reserva Awa, Mataje, Jorgensen et al. 65316 (AAU,
QCA), Rubio & Quelal 1356 (AAU, MJG, MO, QCNE).
Carchi: Reserva Forestal Awa, trail Gualpi bajo Iara-
bita, Jorgensen et al. 65232 (AAU, QCA); Gaulpi Chico,
Awa encampment, Hoover et al. 2602 (AAU, MO, QCA);
Reserva Awa, Centro San Marcos, Mendez et al. 162
(QCNE); wet plateau above San Marcos de los Coaiqueres,
0llgaard et al. 57274 (AAU, QCA), 0llgaard et al.
57584 (AAU, QCA). COLOMBIA. Choco: Quibdo-Tu-
tunendo rd., 14 km E of Quibdo, Gentry & Renteria
24139 (MO); San Jose de Palmar, hoya del Rio Torito,
finca Los Guaduales, quebrada Santa Fe, Forero et al.
6740 (MO). Valle: Rio Naya, Puerto Merizaldo, Costa
Pacifica, Cuatrecasas 13964 (F, US); Rio Yurumangui.
Cuatrecasas 15746 (F, US); Rio Calima, ca. 10 km N
Buenaventura, Gentry 35263 (COL, MO). Narino: Mun¬
icipio de Tumaco, Mora 4212 (COL).

Volume 5, Number 1
1995

Renner & Hausner
Siparuna

69

Figure 5. Female specimens of Siparuna gigantotepala Renner & Hausner. —A. 0llgaard et al. 57274. B.
Barfod et al. 48903, AAU isotype.

70

Novon

Acknowledgments. We thank A. Berg and A.
Horn for help with the illustrations, D. Neill for
information on the whereabouts of duplicates in Qui¬
to, and D. Lorence and P. M. Jorgensen for insightful
reviews. Dan Nicolson went over the Latin diag¬
noses; his input is gratefully acknowledged.

Literature Cited

Dodson, C. H. & A. H. Gentry. 1978. Flora of the Rio
Palenque Science Center. Selbyana 4: 1-628 [Mon-
imiaceae, pp. 434-436].

Endress, P. K. 1980. Ontogeny, function and evolution
of extreme floral construction in Monimiaceae. PI.
Syst. Evol. 134: 79-120.

- & L. D. Hufford. 1989. The diversity of sta¬
men structures and dehiscence patterns among Mag-
noliidae. Bot. J. Linn. Soc. 100: 45-85.

Feil, J. P. 1992. Reproductive ecology of dioecious
Siparuna (Monimiaceae) in Ecuador—A case of gall
midge pollination. Bot. J. Linn. Soc. 110: 171-203.

Holmgren, P. K., N. H. Holmgren & L. C. Barnett.
1990. Index Herbariorum 1. The Herbaria of the
World, Ed. 8. New York Botanical Garden, New
York.

Perkins, J. R. 1901. Beitrage zur Kenntnis der Mon¬
imiaceae. III. Monographic der Gattung Siparuna.
Bot. Jahrb. 28: 660-705, t. 12-14.

-. 1911. Monimiaceae. Pp. 1-67 in A. Engler

(editor). Das Pflanzenreich IV, 101 (Nachtrage). W.
Engelmann, Leipzig.

Two Lesquerellas (Cruciferae) of South Central and Western Montana

Reed C. Rollins

Gray Herbarium of Harvard University, 22 Divinity Avenue, Cambridge,
Massachusetts 02138, U.S.A.

ABSTRACT. Two heretofore unknown species of
Lesquerella are described. These occur in limited
areas in the mountains of central and western Mon¬
tana. Both species, here named L. lesicii and L.
pulchella, occur in patches of limestone-derived
soils and rubble that are conspicuously barren of
trees and shrubs and other types of plant cover, or
on the borders of such areas where the limestone
detritus meets soils and rubble of crystalline rocks.
Also, they are often found in open areas in sparse
stands of low-growing trees or shrubs.

Recent samplings of populations of Lesquerella
in Montana show that species diversity in this genus
goes somewhat beyond that previously recognized
in monographic treatments (Payson, 1922; Rollins
& Shaw, 1973; Rollins, 1993). What has been
known for the area is that species of Lesquerella
are often abundant and are an important component
of the flora of specialized habitats, especially where
limestone or derivatives of limestone are involved.
The nature of some of these habitats is well shown
in Figures 1 and 2, where the highly reflective
limestone-derived soil and rubble contrast with the
more vegetated surrounding areas. These windswept
whitish areas are the places where L. pulchella
grows. Also known is that a high degree of com¬
plexity exists within some species, such as L. alpina
(Nuttall ex Torrey & A. Gray) S. Watson, which is
frequently encountered in the general region. But
now we see that there are existing taxa not previ¬
ously recognized, and these have complicated re¬
lationships with species south of Montana in Idaho
and Wyoming. The main purpose of this paper is
to describe two new species of Lesquerella and ex¬
plain their relationships to previously known taxa.
It is somewhat ironic that I had only recently (Rol¬
lins, 1993) completed a review of Lesquerella as it
occurs in North America when these two species
showed up too late to be included in my treatment.

Lesquerella lesicii Rollins, sp. nov. TYPE: U.S.A.
Montana: abundant in gravelly limestone-de¬
rived soil at edge of limber pine woodland on
the ridge W of Layout Creek, % mi. S of Mys¬
tery Cave, Pryor Mountains, Carbon County,
7500 ft., T85, R28 E, S21, SW '/„ with Shosh -
onea pulvinata, Astragalus aretoides, and A.
miser, 20 June 1992, Peter Lesica 5707 and
Rob DeVelica (holotype, GH).

Herba perennis, caudicibus simplicibus, folia basibus
erectis dense stellatis argenteis 0.5-1 cm longis, caulis
erectis vel decumbentibus gracilibus 1-1.5 dm longis,
petalis aureis spathulatis vel lingulatis 6-7 mm longis,
pedicellis fructiferis recurvatis vel divaricatis 5-10 mm
longis, siliquis globosis vel subglobosis 3-4 mm diametro
pilis stellatis adspersus, loculis 3-5 ovulatis.

Delicate perennial; caudex simple, sometimes
elongated and with old leaf bases; basal leaves erect,
usually fewer than 10, entire, 1.5-3 cm long, pet¬
ioles slender, abruptly expanded to blade, 1-2.5 cm
long, blades broadly ovate to elliptical, 0.5-1 cm
long, silvery from a dense cover of stellate trichomes;
leaf trichomes small, ca. 0.02 mm diam., ray tips
15-25, each primary ray forked near its base; flow¬
ering stems very slender, mostly filiform, simple,
erect to decumbent, 1-1.5 dm long; cauline leaves
few, remote, ± spatulate, lower somewhat petiolate,
upper cuneate at base; inflorescences lax, rarely
nodding, usually with fewer than 10 flowers; sepals
erect, densely pubescent, oblong, nonsaccate and
without scarious margins, 3.5-4 mm long; petals
yellow, often fading to light purple toward their tips,
spatulate to nearly Ungulate, 6-7 mm long; stamens
strongly tetradynymous; infructescences lax, greatly
elongated; fruiting pedicels filiform, recurved in a
single arch to widely spreading, 5-10 mm long;
siliques globose or subglobose, spreading at right
angles to rachis to pendent, 3-4 mm diam., ±
densely pubescent on exterior, valves glabrous on
the interior; styles ca. 1.5 mm long, glabrous or

Novon 5: 71-75. 1995.

72

No von

with a few trichomes near their bases; ovules 3-5
per locule; mature seeds not seen.

Lesquerella lesicii is most closely related to L.
fremontli Rollins & E. Shaw. Both species are ap¬
parently endemic to very limited areas and appear
to be restricted to Madison limestone derivatives.
Lesquerella lesicii occurs in the Pryor Mountains
of Montana, which are northwest of the Big Horn
Mountains and separated from that range by the
canyon of the Big Horn River. Lesquerella fremontii
is found at the southeastern end of the Wind River
range in Wyoming. These areas are about 200 miles
apart. The two species are singular in having small
unbranched caudices (although definitely perennial),
erect basal leaves, simple flowering stems, more or
less recurved fruiting pedicels, and globose siliques.
Both species have small, stellate, many-rayed tri¬
chomes and weak stems, which are particularly frag¬
ile in L. lesicii.

Lesquerella lesicii differs from L. fremontii in
having basal leaves with very thin petioles that
abruptly expand to a broadly ovate to orbicular
blade, whereas in L. fremontii the petioles are stouter
and expand gradually to a spatulate or nearly linear
blade. The flowering stems of L. fremontii are hor¬
izontal along the ground and the infructescences are
secund. In L. lesicii these stems are erect and the
infructescences have the usual racemose pattern.
The fruiting pedicels are definitely recurved in a
single arch and the siliques are more or less pendent
in L. fremontii, but in L. lesicii the pedicels, al¬
though tending to be recurved, are more often just
slightly arched and are widely spreading, bearing
siliques that are horizontal. The pedicels are so finely
filiform that they are extremely fragile, while those
of L. fremontii are thicker and much less fragile.

According to Peter Lesica (in litt.), Lesquerella
lesicii is not the only endemic species that occurs
in the Pryor Mountains. He points out that Penste-
mon caryi Pennell and Erigeron allocotus Blake
are endemic there and that the recently described
monotypic Shoshonea pulvinata Evert & Con¬
stance is restricted to Madison limestone in the
Pryors, and along the front of the Beartooth, Ab-
saroka, and Owl Creek mountains in Wyoming and
Montana.

Paratypes. U.S.A. Montana: Carbon County, very
common in gravelly limestone derived soil on a NE-facing
slope at the head of Big Coulee, Pryor Mtns., 6400 ft.,
T9S, R28E, NE /„ with Cercocarpus ledifolius, Musi-
neon vaginatus, and Agropyron spicatum, 19 June 1992,
Peter Lesica 5700 and Rob DeVelice (GH, MONTU);
locally abundant in stony limestone derived soil in a wind¬
swept Douglas Fir forest on a ridge ca. % mi. S of Mystery
Cave, Pryor Mtns., 7500 ft., T8S, R28E, Sec. 21, SW

/„ with Hymenoxys torreyana and Eritrichium howar-
dii, 27 June 1991, Peter Lesica 5449 (GH, MONTU).

Lesquerella pulchella Rollins, sp. nov. TYPE:
U.S.A. Montana: Beaverhead County, common
in barren calcareous soil in open spruce wood¬
land on S-facing slope along Trapper Creek just
S of Hecha Mines, Pioneer Mtns., T3S R11W,
Sec. 32, 8600 ft., with Potentilla fruticosa
and Zigadenus elegans, 25 July 1991, Peter
Lesica 5525 and Steve Cooper (holotype, GH;
isotype, MONTU).

Herba perennis, caudicibus simplicibus, folia radicalibus
simplicibus argenteis petiolata 1-2 cm longis laminis late
ovatis vel ellipticis, caulis tenuibus prostratis 2-7 cm
longis, folia caulina petiolata vel ad basi cuneatis, sepalis
oblongis 3-4(-5) mm longis dense pubescentibus, petalis
aureis spathulatis vel lingulatis (6—)7—8(—9) mm longis,
pedicellis fructiferis plerumque patuleis ± sigmoideis 5-
7 mm longis, siliquis orbicularibus vel ellipticis pubescen¬
tibus (3-)4-6(-7) mm longis, 3-4 mm latis, stylis 1-1.5
mm longis, loculis 3-5 ovulatis, seminibus oblongis crassis
± 2 mm longis.

Perennial; caudex mostly simple; caudex leaves
several to numerous, simple, silvery from a dense
cover of appressed radiately branched trichomes,
blades broadly ovate or elliptical to wider than long,
obtuse, 1-2 cm long including the slender petioles;
leaf trichomes in several layers, branches usually
forked near their bases, primary branches 6-7; stems
several to many, slender, simple or occasionally
branched, mostly prostrate, 2-7 cm long; cauline
leaves petiolate below to cuneate above, spatulate
to narrowly elliptical or broadly ovate; inflorescences
congested, flowering pedicels divaricately ascending
to nearly erect, straight or nearly so; sepals narrowly
oblong, 3-4(-5) mm long, nonsaccate, densely pu¬
bescent; petals yellow, spatulate to Ungulate, (6-)7-
8(-9) mm long; stamens erect, filaments slender,
anthers oval; fruiting pedicels divaricately ascending
to widely spreading, sigmoid to slightly so or nearly
straight, slender, 5-7 mm long, not swollen toward
the apex; siliques nearly orbicular to efliptical or
broadly oblong, rounded above and below, strongly
compressed contrary to the septum but with a def¬
inite ridge down the middle formed over the replum
edges, densely pubescent but not silvery, (3-)4-6(-
7) mm long, 3-4 mm wide; septum entire or with
a narrow perforation; styles 1-1.5 mm long, not
expanded at summit; seeds plump, oblong, brownish,
ca. 2 mm long, wingless, cotyledons accumbent.

Lesquerella pulchella (Figs. 3, 4) is most closely
related to L. paysonii Rollins. Both species have
sfliques compressed contrary (perpendicular) to the
plane of the septum. Their fohage and sfliques are

Volume 5, Number 1
1995

Rollins

Lesquerella from Montana

73

Figures 1 and 2. Typical habitats of Lesquerella pulchella Rollins. —1. Upper, East Pioneer range, viewed from
subsummit of Black Lion Mountain. —2. Lower, north of Black Lion Mountain, south of Vipond Park. Photos by
Peter Lesica.

74

Novon

Figures 3 and 4. Lesquerella pulchella Rollins (ca. natural size). —3. Upper, flowering plant (Lesica 5528).
4. Lower, fruiting plants (Lesica 5525). Photos by Peter Lesica.

covered with relatively small, radiately branched
trichomes, and in each species the foliage is silvery
from the multiple layers of trichomes present. A
single layer of trichomes on the silique surfaces does
not produce the same silvery effect. The silique
compression is more severe in L. paysonii than in

L. pulchella. The result is that the replum margin
does not protrude and the replum itself is narrower
in L. paysonii than in L. pulchella. In the latter,
the replum margin does protrude to form a riblike
ridge where the valves come together.

Generally in Lesquerella, the presence or absence

Volume 5, Number 1
1995

Rollins

Lesquerella from Montana

75

of trichomes on the interior of the valves is not an
infallible mark of distinction between species, but
most of the specimens of Lesquerella pulchella I
have examined do have trichomes on the valve in¬
terior. On the other hand, none of the specimens of
L. paysonii I have seen have trichomes there. This
is worth noting because it may turn out to be a
significant difference once enough populations have
been sampled to establish the surety of the feature
difference. Some variation in the length of the styles
is to be expected and does occur in both species.
However, this variation ranges from 2 mm in length
downward in L. pulchella , but from 2 mm upward
to 4 mm in L. paysonii.

There is a basic difference in the outline shape
of the siliques of Lesquerella paysonii and L. pul¬
chella. In the latter, the siliques are nearly orbicular
in outline or only slightly longer than broad, while
in L. paysonii the siliques are definitely longer than
broad, usually nearly twice as long as wide. The
effect of this pattern, combined with the degree of
compression on the shape and length of the replum,
is seen in the differences of the replum shape in the
two species. The replum of L. paysonii is longer,
narrower, and tapers more gradually toward the
apex than in that of L. pulchella, where the replum
tends toward a more oblong shape. In L. pulchella
the valve margins are tightly pressed together so
that the replum margin does not show and there is
a continuous cover of trichomes over the ridge formed
between the valves. But in L. paysonii the valve
margins are rarely sufficiently together to touch each
other. Here the replum margins show a break be¬
tween the valves, and the trichome cover is not
continuous from one valve to the other.

Lesquerella pulchella ranges in elevation from
around 6300 to about 9600 feet. As with other
plant species with such a wide elevational occur¬
rence, there is considerable variation from popula¬
tion to population. Some of this variation appears
to be correlated with elevation, some not. I note
that the flowers on the high elevation variants are

both larger and more numerous per plant than in
the populations of lower elevations. This was called
to my attention by Bonnie Heidel.

Paratypes. U.S.A. Montana: Beaverhead County,
common on gentle E-facing slope at the top of Keokirk
Mt., near contact between crystalline and calcareous par¬
ent materials, T3S, R11W, Sec. 10, 9600 ft., 25 July

1991, Peter Lesica 5528 and Steve Cooper (GH, MON-
TU); common in portion of upper S-facing slope on ridge
point, S of Bridger Pass, T7S, R11W, Sec. 27, 7140 ft.,
5 Aug. 1992, B. Heidel 938 (GH); common in barren
calcareous soil on a moderate E-facing slope of the ridge
N of Black Lion Mt., T2S, R11W, Sec. 29, 9200 ft.,
24 July 1991, Peter Lesica 5522 and Steve Cooper
(GH, MONTU); steep slopes of upper montane zone,
Nemesis Mt., T14S, R2E, Sec. 31, 9050 ft., 27 June

1992, B. Heidel 722 (GH); near same location, 27 June
1992, B. Heidel 721 (GH); above Scudder Creek on
ridges at the edge of Pioneer Mtns., T6S, R12W, Sec.
21, 6720 ft., 24 June 1992, B. Heidel 700 (GH); open
scree of S-facing slope, ridge E of Nemesis Mt., T14S,
R2E, Sec. 21, 9000 ft., 26 July 1992, Lisa Schassberger
Roe 493 and Jonathan Stewart Roe (GH).

Acknowledgments. I am much indebted to, and
thank, Peter Lesica of the University of Montana,
and Bonnie L. Heidel and Lisa Schassberger Roe of
the Montana Natural Heritage Program, for pro¬
viding me with excellent study material. Peter Les¬
ica, in addition to excellent collections, also provided
significant notes on the field populations of Les¬
querella lesicii and additionally the photographs of
Figures 1-4, for which I am grateful. My sister,
Aileen G. Roads Carter, has generously provided
funds to offset the cost of the colored plates, for
which I thank her sincerely.

Literature Cited

Payson, E. B. 1922. A monograph of the genus Les¬
querella. Ann. Missouri Bot Gard. 8: 103-236.
Rollins, R. C. 1993. The Cruciferae of Continental
North America. Stanford Univ. Press, Stanford, Cal¬
ifornia.

-& E. A. Shaw. 1973. The Genus Lesquerella

(Cruciferae) in North America. Harvard Univ. Press,
Cambridge, Massachusetts.

Merostachys multiramea (Poaceae: Bambusoideae: Bambuseae) and
Similar Species from Brazil

Tatiana Sendulsky

Herbario, Instituto de Botanica, Caixa Postal 4005, 01061-970 Sao Paulo, Brazil

ABSTRACT. The true identity of Merostachys mul¬
tiramea Hackel is discussed. As a result of a review
of the group of the species similar to M. multira¬
mea, the following are described as new: Meros¬
tachys abadiana, M. caucaiana, M. filgueirasii,
M. kleinii, M. magellanica, M. pilifera, M. scan-
dens, and M. skvortzovii . The description of floral
morphology is added to M. glauca McClure & L.
B. Smith, which was originally described from sterile
material. A key to the species is provided. All the
species are fully illustrated.

In the past the name Merostachys multiramea
Hackel has been applied erroneously to a number
of different species, both described and undescribed.
Because true M. multiramea is a distinct and rather
rare species, represented in herbaria by only a few
collections, a considerable amount of misidentified
material has been placed under the specific epithet
multiramea and thus left unstudied. Among these
are species that present no difficulties in identifi¬
cation: either they are clearly new or they have
already been described. However, there is a group
of about 10 species that are difficult to distinguish
one from another, but which cannot be aggregated
under one name as has so far been the case.

As botanists working with herbarium collections
of Brazilian bamboos have tended, over the years,
to assign almost any specimen of Merostachys with
medium-sized, 1- or occasionally 2-flowered spike-
lets to M. multiramea Hackel, the misidentification
has had fourfold results. A number of validly pub¬
lished species, including M. argyronema Lind man,
M. claussenii Munro, and M. clausenii Munro var.
mollior Doell, were for a long time disregarded in
the belief that they were all M. multiramea. Two
names, M. anomala Dutra and M. burchellii Mun¬
ro emend. Dutra (not M. burchelii Munro), crept
into the group. Collections that would later be con¬
sidered as distinct though unpublished taxa (M. ma¬
gellanica , for instance) went unrecognized. And the
identity of M. multiramea itself, the “umbrella”
for so many other taxa, was largely obscured.

The name Merostachys multiramea was published
in 1909 by Hackel, and M. claussenii Munro var.

Novon 5: 76-96. 1995.

mollior by Doell in 1880 (see von Martius, 1840-
1906). The two species were described again, under
new names, by Dutra in 1938: the former as M.
anomala Dutra and the latter as M. burchellii Munro
emend. Dutra. Dutra’s emendation was based on the
assumption that unidentified collections of flowering
material in his possession were a good match for the
vegetative and sterile material on which Munro (1868)
based his original description. This “graft” was pub¬
lished by him (Dutra, 1938) as a distinct species.

The situation was further complicated by the in¬
version of the captions to Dutra’s figures: his figure
2 (Dutra, 1938: 149), purporting to show leaves
and inflorescences of M. burchellii Munro emend.
Dutra, is, in fact, of M. anomala (= M. multira¬
mea), while his figure 3 (Dutra, 1938: 149), which
claims to be of M. anomala, is of M. burchellii
Munro emend. Dutra (= M. claussenii Munro var.
mollior Doell).

Return of the captions to their rightful figures
eliminates a misleading interpretation of inflorescence
and spikelet morphology in the two taxa concerned,
and permits their proper identification as synonyms
of previously published species. In this context, M.
multiramea also assumes its true identity, as a rare
bamboo from the southern part of Brazil; Dutra s
figure 2 makes it clear that it is not one of the group
of rather common species with only one floret in the
spikelet. Merostachys multiramea Hackel is char¬
acterized by a 2-flowered spikelet and by its dark
brown (nearly black), conical caryopses. Both florets
are fertile—the rudimentary apical floret and the
upper and lower glumes, called “florets” by authors
of the last century, obviously not included. Thus, far
from being widely distributed, M. multiramea has
been collected only infrequently, and mostly by Dutra,
under the name M. anomala.

Key to the Species of Merostachys Similar to

M. MULTIRAMEA

1. Culm internodes densely pilose, hairs sericeous,
translucent, ca. 1 mm long, in some areas sharp,

globose prickle hairs also present. M. pilifera

1. Culm internodes glabrous, smooth or scabrous,
with prickle hairs or mottled with dark spots
and stripes. ^

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1995

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Merostachys from Brazil

77

2( 1). Spikelets always 2-flowered or occasionally 2-
flowered (both florets fertile), otherwise bearing
different stages of development of the upper

floret . 3

2. Spikelets always 1-flowered. 5

3(2). Spikelets always 2-flowered. 4

3. Spikelets occasionally 2-flowered (both florets

fertile), otherwise bearing different stages of
development of the upper floret; florets ca. 9
mm long, caryopsis ellipsoid, 6 mm long, 1.5
mm wide. M. caucaiana

4(3). Florets 7-9 mm long, 3 mm wide, laterally
strongly compressed, imbricate; caryopsis con¬
ical, attenuate at the apex, dark brown (nearly

black), 6-7 mm long, 3 mm wide.

. M. multiramea Hackel

4. Florets ca. 10-12 mm long; caryopsis broadly

ovoid, basally and apically rounded, dark yel¬
low, 6-7 mm long, 4-4.5 mm wide.

. M. kleinii

5(2). Spikelets always with two parallel, longitudinal,
white lines, formed by the keels of the palea.
. M. magellanica

5. Spikelets without parallel, white lines. 6

6(5). Caryopsis ovoid . 7

6. Caryopsis ellipsoid, 7.5 mm long, 3 mm wide,
grayish brown, culm leaf blades narrow, 4-5
cm long, 4 mm wide, branch leaf blades lan¬
ceolate, 4-9 cm long, 8-12 mm wide, L:W

= 5-7.5. M. jilgueirasii

7(6). Branch leaf blades lanceolate, 3-11 cm long,

6-16 mm wide, L: W = 5-6.9 or 3.5-10 cm
long, 8-14 mm wide, L: W = 4.4-7.1; branch
complements with 40-200 branches. 8

7. Branch leaf blades broadly lanceolate, 4-12

cm long, 15-20 mm wide, L:W = 2.6-6 or
narrowly linear-lanceolate, 5-13 cm long, 6-
12(—17) mm wide, L: W = 8.3-10.8; branch
complements with 20-60 branches. 9

8(7). Branch complement with 150-200 branches;
branch leaf blades lanceolate, 3-11 cm long,

6-16 mm wide, L: W = 5-6.8; culm leaf blades
lanceolate, 6-10 cm long, 10 mm wide; inflo¬
rescences pectinate; caryopsis 5 mm long, ca.

3 mm wide, apically rounded. M. skvortzovii

8. Branch complement with 40-60(-100)

branches; branch leaf blades lanceolate, 3.5-
10 cm long, 8-14 mm wide, L: W = 4.4-7.1,
culm leaf blades lanceolate, 8-12 cm long, 4-
5 mm wide; inflorescence distichously arranged,
caryopsis 5.5-6.5 mm long, ca. 3 mm wide,
slightly attenuate apically. M. scandens

9(7). Branch leaf blades broadly lanceolate, 4-12
cm long, 1.5-2 cm wide, L: W = 2.6-6; lemma
finely puberulent . M. abadiana

9. Branch leaf blades narrowly linear-lanceolate,

5-13 cm long, 6-12(-17) mm wide, L: W =
8.3-10.8, lemma pilose, the hairs ca. 0.5 mm

long, silver-grayish.

. M. glauca McClure & L. B. Smith

Merostachys abadiana Sendulsky, sp. nov. TYPE:
Brazil. Estado de Sao Paulo: Municipio Itatinga,
Abadia, 26 sep. 1978 (fl), Campos Neto s.n.
(holotype, SP 154498). Figure 1.

Culmi ca. 2 m longi. Fragmenta ad extremitates cul-
morum internodium 2-5 mm diametro, laevia. Nodi prom-
inentes, crista supranodali angusta, tubiformi. Folia cul-
morum ignota. Rami in complemento 20-30. Foliorum
ramorum laminae late lanceolatae, 4-12 cm longae, 1.5-
2 cm latae. Inflorescentia speciformis, pectinata, 2-4 cm
longa. Spiculae 11-13 mm longae, late ellipticae, 1-flos-
culatae; gluma infera 3 mm longa; gluma superior 8.5
mm longa, 11-13-nervia, acuta; lemma 9 mm longum,

15- 16-nervium, late lanceolatum, puberulum; palea fu-
siformis, 9 mm longa, 10-nervia, anguste sulcata. Cary¬
opsis ovoidea, 4.5 mm longa, 3.4 mm lata, cinereo-lutea.

Woody bamboo ca. 2 m high, according to col¬
lector’s note. Fragments of internodes from tip of
the culm fistulose, 2-5 mm diam., yellowish, smooth.
Nodes salient, supranodal ridge narrow, tubiform.
Culm leaves not seen. Branch complement with 20-
30 branches, these 8-40 cm long, 1 mm thick,
glabrous, 1 -2-noded before the leaves. Branch leaves
of the flowering branches 4-5 per complement:
sheaths tight, finely puberulent, the overlapping
margin glabrous; adaxial surface dark-spotted; oral
setae 2-3 mm long, distally crisped; pseudopetiole
grayish yellow, 2 mm long, finely pubescent; inner
ligule a thick membrane, pubescent on the back;
outer ligule a slightly pubescent rim; blades 4-12
cm long, 1.5-2 cm wide, L:W = 2.6-6, broadly
lanceolate, glabrous adaxially except for a scabrous
base, with rows of fine prickle hairs and 2-3 mar¬
ginal nerves toward the apex with fine prickle hairs,
slightly pubescent abaxially, more densely so at the
base, the hairs whitish, margins glabrous or finely
scabrous toward the apex. Inflorescences terminal
on leafy branches, spikelike, pectinate, 2-4 cm long,
with 4-15 spikelets. Rachis densely pubescent. Ped¬
icels pubescent, short. Spikelets solitary, 1 -flowered,
bisexual, broadly ellipsoid, somewhat inflated, 11-
13 mm long, 3-4 mm wide. Glumes 2, unequal.
Lower glume 1-nerved, carinate, attenuate toward
the apex, finely puberulent, 3 mm long, 1 mm wide
at the base. Upper glume 8.5 mm long, 3.5 mm
wide, broadly lanceolate, acute, awnless, finely pu¬
berulent, 11-13-nerved, adaxially shiny, dark-spot¬
ted. Lemma 9 mm long, 6 mm wide, very broadly
lanceolate, deeply concave, enclosing a palea, 15-

16- nerved, finely puberulent abaxially, the adaxial
surface dark brown, shiny, densely spotted, the mar¬
gins densely ciliate, the hairs dark, reddish. Palea
fusiform, 9 mm long, 5 mm wide, 2-keeled, narrowly
sulcate abaxially, 10-nerved, the nerves densely
anastomosing, the veinlets uneven; the keels finely
ciliate; abaxial surface densely puberulent at the
apex and along the sulcus, glabrous in the lower
part; adaxial surface shiny, dark. Rachilla extension
bristle-like, with a small rudiment on the tip. Car¬
yopsis 4.5 mm long, 3.4 mm wide, ovoid, rostrate.

78

Novon

Figure 1. Merostachys abadiana Sendulsky. —a. Branch complements from the upper part of the culm- b.
Apical part of the inflorescence. c. Spikelet. —d. Caryopsis, lateral view. —e. Caryopsis, embryo view. —f. Lower
glume. g. Upper glume. h. Lemma. i. Palea. —j. Caryopsis, hilum view. —k. Upper portion of the branch
leaf sheath and base of the blade (abaxial surface). Based on Campos Neto s.n. (SP 154498).

grayish yellow, the embryo and the hilum scarcely
manifest externally.

Phenology. Known only from the type collection.
Distribution. Known only from the type locality,
in the State of Sao Paulo, Brazil.

Merostachys abadiana was collected on the
grounds of a Benedictine abbey in Sao Paulo in
1978 and sent to the Institute de Botanica, Sao
Paulo, for identification. No further information about
the plant was obtained, except that some clumps of

Volume 5, Number 1
1995

Sendulsky

Merostachys from Brazil

79

Figure 2. Merostachys caucaiana Sendulsky. —a. Flowering branch complement. —b. Spikelet, lower floret with
mature caryopsis, upper floret not fully developed. —c. Spikelets with two fertile florets. —d, e. Spikelets in different
stages of development (upper florets). —f. Lodicules. —g. Caryopsis, embryo view. —h. Inflorescence. —i. Caryopsis,
lateral view. —j. Caryopsis, hilum view. —k. Upper portion of the branch leaf sheath and base of the blade (abaxial
surface). —1. Lower glume. —m. Upper glume. —n. Lemma. —o. Palea. —p. Palea and rachilla extension with
underdeveloped upper floret. Based on O. Handro & Menezes s.n. (SP 262720).

the bamboos are still growing in that area. It is details of the lemma but differs in the glabrous
somewhat similar to M. pilifera (described later in internodes and in the wider branched leaf blades,
this paper) in the morphology of the spikelet and which are hairy on the abaxial surface.

80

Novon

Merostachys caucaiana Sendulsky, sp. nov.
TYPE: Brazil. Estado de Sao Paulo: Municipio
de Cotia, Caucaia, 1978 (fl), 0. Handro &
Menezes s.n. (holotype, SP 262720). Figure 2.

Habitus ignotus. Fragments ad extremitates culmorum
internodium fistulosa, laevia, opaca, maculis atro-viridibus
rotundatis. Nodi pilis albis, retrorsis ad basin. Folia cul-
morum ignotae. Rami in complemento 40-50. Foliorum
ramorum laminae lanceolatae, 2-7 cm longae, 5-12 mm
latae. Inflorescentia spiciformis, pectinata, 2-3 cm longa,
ramum foliorum terminans. Spiculae 1 aut rare 2-flos-
culatae, 9-10 mm longae, fusiformes, lateraliter com-
pressae; flosculus inferior bisexualis, 9-10 mm longus vel
rudimentalis. Gluma inferior 2 mm longa; gluma superior
7 mm longa, 9-11 -nervia; lemma 7 mm longum, 9-
nervium; palea 7 mm longa, late sulcata. Caryopsis ellip-
tica, 6 mm longa, pallide brunnea.

Habit unknown. Fragments of the culm inter-
nodes from the tip of the culm delicate, with thin
pendent or clambering tips. Intemodes fistulose, ca.
1 cm diam., glabrous, smooth, but not lustrous,
opaque, finely mottled with very small, dark green,
rounded spots (not stripes) and with a glaucous-
whitish touch toward the upper part. Upper culms
sometimes rugose with small, sharp, retrorse prickle
hairs. Culm walls 1 mm thick. Nodes salient with
white retrorse hairs below. Culm leaves not seen.
Branch complement (upper portion of plant) with
40—50 branches, these 10—15 cm long, 1 mm thick,
glabrous, 1-2-noded before the leaves. Branch leaves
3-4 per complement: sheaths tight, glabrous, adax-
ially dark-spotted, the overlapping margin finely cil-
iate or glabrous, the intercostal zone finely trans-
versally rugose; oral setae 13-15 cm long, dimin¬
ishing in size downward, along the margins of the
sheath; pseudopetiole dark, rugose, with a few fine
prickle hairs at the base adaxially; inner ligule 0.5
mm long; outer ligule a salient rim; blade 2-7 cm
long, 5-12 mm wide, L:W = 4-5.8, lanceolate,
with an acuminate apex and slightly asymmetrical
at the base, glabrous on both surfaces except for
the base abaxially, the hairs whitish, hyaline, ap-
pressed, on one side of the midnerve; adaxial surface
with 2-3 marginal nerves on one side with rows of
fine prickle hairs; margins finely scabrous. Inflores¬
cences spikelike, pectinate, 2-3 cm long, half in¬
serted in ultimate sheath. Rachis and pedicels pilose.
Spikelets 1- or rarely 2-flowered. Glumes 2, unequal.
Lower glume 2 mm long, 1 mm wide, acuminate,
carinate, 1-nerved, puberulent, ciliate at the mar¬
gins. Upper glume 7 mm long, 3 mm wide, broadly
lanceolate, 9-11 -nerved, finely puberulent, ciliate
at the margins, the nerves anastomosing. Lower
floret bisexual, 9-10 mm long, 2-3 mm wide, fu¬
siform, laterally compressed. Lemma 7 mm long, 4

mm wide, broadly lanceolate, finely puberulent, 9-
nerved, the nerves anastomosing, the margins cili¬
ate. Palea 7 mm long, glabrous, 2-keeled, broadly
sulcate between the keels, the sulcus finely pilose,
the keels ciliate. Rachilla extension flat, ca. 0.5 mm
wide, puberulent and ciliate laterally, bearing one
of several different stages of development of the
upper floret, these varying in size and shape, from
a small rudiment to a fully formed fertile floret,
identical to the lower one. Rachilla extension of the
upper floret bristle-like, with a small rudiment at the
tip. Lodicules 3, 2 mm long, finely ciliate apically.
Ovary not seen. Caryopsis 6 mm long, 1.5 mm wide,
ellipsoid, rostrate, light brown, shiny, the embryo
and the hilum scarcely manifest externally.

Phenology. The only available flowering speci¬
men of this species is the holotype.

Distribution. Known only from the type locality,
in the State of Sao Paulo, Brazil.

Merostachys caucaiana was found growing in a
secondary forest in the Municipality of Cotia, ca.
40 km from Sao Paulo.

Merostachys filgueirasii Sendulsky, sp. nov.
TYPE: Brazil. Distrito Federal: Catetinho,
15°57'S, 47 <> 59 , W, 7 Jan. 1982 (fl), Filgueiras
& Pereira 953 (holotype, IBGE; isotypes, CCN,
MG, SP, UEC). Figure 3.

Culmi 2-10 m longi, 1-1.5 cm diametro; internodia
fistulosa, scabra. Nodi glabri; crista supranodali tubiformi.
Foliorum culmorum laminae angustae, 4-5 cm longae et
4 mm latae. Rami in complemento 20-50. Foliorum
ramorum laminae lanceolatae, 4-9 cm longa, 8-12 mm
latae. Inflorescentia spiciformis, pectinata, 3.5-4.5 cm
longa, ramorum foliarum terminans. Pedicelli pilosi, base
dilata in projecturam nudam, globosam. Spiculae 13.5-
14 mm longae, fusiformes, 1-flosculatae. Gluma infera 2
mm longa; gluma supera 8-10 mm longa, 7 - 9 -nervia,
aristulata; lemma 8-9 mm longum, 21-23-nervium, sub-
tiliter puberulum, ad marginem strigosum, pilis fuscis;
palea 11 mm longa, 12-nervia, anguste sulcata. Caryopsis
elliptica, 7.5 mm longa, cinereo-brunnea.

Caespitose, woody bamboo, forming dense clumps.
Rhizomes pachymorph, sympodial, densely covered
by yellow scales. Culms 2-10 m long, 1-1.5 cm
diam., flexuous at the tips, according to Filgueiras
(1988). Intemodes fistulose, scabrous, green when
young, yellowish when mature, finely mottled with
small, dark green stripes. Culm walls of the middle
intemodes 1-1.5 mm thick. Nodes glabrous, salient,
supranodal ridge tubiform, 1-2 mm long, especially
evident in young culms. Culm leaves deciduous:
sheaths 12-15 cm long, 5-6 cm wide, rounded at
the summit, fringed at the apex with dense tufts of

Volume 5, Number 1
1995

Sendulsky

Merostachys from Brazil

81

Figure 3. Merostachys filgueirasii Sendulsky. —a. Flowering branches. —b. Base of the branch complement. —
c. Lower glume. —d. Upper glume. —e. Lemma. —f. Palea. —g. Spikelet. —h. Apical part of the inflorescence.
—i. Caryopsis, embryo view. —j. Caryopsis, hilum view. —k. Rhizome. —1. Upper portion of the branch leaf sheath
and base of the blade (abaxial surface). —m. Culm leaf. Based on Filgueiras <£ Pereira 953 except for k and m,
which are based on Filgueiras & Brochado 2009.

hairs; hairs 12 mm long; abaxial surface glabrous,
densely nerved, dark; adaxial surface somewhat
shiny, dark, striate, densely nerved; margins gla¬
brous or finely ciliate; inner ligule thick, finely ciliate;
outer ligule a short, inconspicuous rim; blades re¬
flexed, deciduous, narrow, 4-5 cm long, 4 mm wide.

Branch complement with 20-50 branches, these
30-45 cm long, 1 mm thick, glabrous, smooth,
yellowish, 5-6-noded before the leaves, the nodes
with fine, whitish pubescence below. Branch leaves
4-7 per complement: sheaths tight, glabrous, the
overlapping margin finely ciliate toward base; oral

82

Novon

setae fine, 5-7 mm long, minutely scabrous, crisped
distally; inner ligule solid, finely ciliate, ca. 0.5 mm
long; outer ligule a short inconspicuous rim; pseu¬
dopetiole flat, grayish yellow, slightly puberulent, 3
mm long; blades 4-9 cm long, 8-12 mm wide, L:
W = 5-7.5, lanceolate, light green, finely pilose on
abaxial surface, more densely pilose at the base, the
hairs whitish; adaxial surface glabrous, 2-3 marginal
nerves on one side with rows of fine prickle hairs;
margins and the apex finely scabrous. Inflorescence
terminal on leafy branches, spikelike, pectinate, se-
cund, 3.5-4.5 cm long, with 16-20 spikelets. Ra-
chis densely pilose. Pedicel pilose, at the base swollen
into a lateral, globose, tuberous, naked projection.
Spikelets 13.5-14 mm long, 3 mm wide, solitary,
1-flowered, bisexual, attenuate, fusiform. Glumes 2,
unequal. Lower glume 1-nerved, carinate, mucro-
nate, finely puberulent, 2 mm long, ca. 1 mm wide
at the base. Upper glume 8-10 mm long, 3 mm
wide, 7-9-nerved, broadly lanceolate, finely puber¬
ulent, the apex scabrous, aristulate, the awn ca. 1
mm long; adaxial surface shiny, dark-spotted. Lem¬
ma 8-9 mm long, 6 mm wide, broadly lanceolate,
21-23-nerved, the nerves anastomosing in the up¬
per part; abaxial surface puberulent, apically densely
strigose near the margin, the hairs thick, short, dark;
adaxial surface dark-spotted. Palea 11 mm long,
ca. 5 mm wide, glabrous, 2-keeled, narrowly sulcate
abaxially, 12-nerved, the nerves obliquely anasto¬
mosing; keels finely ciliate; adaxial surface shiny.
Rachilla extension bristle-like, 7 mm long, with a
minute rudiment. Lodicules not seen. Ovary elon¬
gate, style single, the stigmas 2, plumose, the sta¬
mens 3, the anthers 6 mm long, dark. Caryopsis
7.5 mm long, ca. 3 mm wide, ellipsoid, rostrate,
grayish brown, finely striate, the embryo and the
hilum scarcely manifest externally.

Phenology. According to Filgueiras (1988 and
pers. comm.) two populations of Merostachys fil-
gueirasii bloomed and died gregariously in 1980-
1982, at the Reserva Ecologica do IBGE (Brazilian
Institute of Geography and Statistics), and no re-
growth was observed. A few months later, hundreds
of seedlings could be seen in the area; these were
collected by Filgueiras (Filgueiras 1021, IBGE).
This collection contains seedlings in different stages
of growth. The smallest specimen is 10 cm long,
with 5 leaves and the caryopsis still attached. The
seedlings of M. filgueirasii grew fast and occupied
the bare ground left by the parent clump very quick¬
ly. By 1992, the two populations had completely
re-established themselves at approximately the same
site as the previous ones. No previous flowering is
recorded.

Distribution. Brazil. Distrito Federal: grounds of
the Reserva Ecologica do IBGE, gallery forest in
cerrado vegetation, 850-1340 m, and grounds of
Catetinho. However, the population of the type spec¬
imen at Catetinho was completely wiped out because
the site was cleared for tourist interests.

The swollen lateral projections at the base of the
pedicels are very peculiar; these are also found in
M. plurijlora Munro ex Camus and in M. skvort-
zovii (described later in this paper). In the former
they are rather prominent, whereas in the latter
they are small and are located on the inner side of
the pedicel.

This species was cited by Tarciso S. Filgueiras
(1988) as Merostachys multiramea and is named
for him, a friend and colleague, for his contributions
to the knowledge of the Brazilian grasses.

Paratypes. BRAZIL. Distrito Federal: Reserva
Ecologica do IBGE, 15°57'S, 47“52'W, proximo a cerca
W da Recor, 26 Sep. 1980 (fl), Heringer et al. 5812
(IBGE, K, US), 18 Jan. 1982 (fl), Filgueiras 955 (IBGE),
8 Jun. 1982 (seedlings), Filgueiras 1021 (IBGE); Reserva
Ecologica do IBGE, mata do ribeirao Taquara, 20 Nov.
1990 (st), Filgueiras & Brochado 2009 (IBGE, SP).

Merostachys glauca McClure & L. B. Smith.
TYPE: Brazil. Estado de Santa Catarina: Pal-
hoga, Piloes, mata, 200 m, 24 feb. 1956 (st),
Reitz <6 Klein 2737 (holotype, US). Figure 4.

Habit unknown. According to McClure & L. B.
Smith (1967) only fragments of the midculm are
known. Internodes ca. 30 cm long, 1.2-2.5 diam.,
glabrous and smooth, greenish yellow, whitish
glaucous upward, the thinner internodes greenish,
smooth, somewhat shiny, finely mottled with very
small, dark green rounded spots. Culm walls 1-3
mm thick. Nodes salient, with retrorse white hairs
below; supranodal ridge tubiform. Culm leaves de¬
ciduous: sheaths oblong, 29 cm long, 11 cm wide,
striate, rather rough, with a trace of wax and a few
scattered white hairs, the blades not seen. Branch
complement with 30-60 branches, these 1 - 2 -noded
before the leaves, glabrous. Branch leaves 5—7 per
complement: sheaths tight, striate; inner ligule dark,
0.5 mm long, ciliate at the margin; outer ligule a
salient rim; pseudopetiole 3-4 mm long, dark, gla¬
brous; oral setae 5 mm long, deciduous; blades nar¬
rowly linear-lanceolate, attenuate at the apex, 5-
13 cm long, 6-12(-17) mm wide, L:W = 8.3-
10.8; adaxial surface glabrous, with 1-2 marginal
nerves on one side with rows of fine, sharp, sparse
prickle hairs; abaxial surface glabrous except for
densely pubescent area at the base beside the mid-

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Merostachys from Brazil

83

Figure 4. Merostachys glauca McClure & L. B. Smith. —a. Flowering branch complement. —b. Part of the
inflorescence. —c. Spikelet, showing lower glume and lemma. —d. Spikelet, showing upper glume and lemma. —e.
Lower glume. —f. Upper glume. —g. Rachilla segment. —h. Lemma, adaxial surface. —i. Lemma, abaxial surface.
—j. Palea with rachilla and rudiment. —k. Upper part of the branch leaf sheath and base of the blade (abaxial
surface). Based on Klein & Sousa Sob. 10480.

nerve; margins sparsely scabrous. The epidermis of
the sheath and the blade presents extremely small,
rounded, flat papillae and some small scattered,
pointed or unpointed prickle hairs. Inflorescences
terminal on leafy branches, spikelike, pectinate, se-
cund, 1.5-3 cm long, with 8-10 spikelets. Rachis

and pedicels pilose. Spikelets 1-flowered, solitary,
fusiform, 10 mm long, 2.5 mm wide. Glumes 2,
unequal. Lower glume 2.5 mm long, 1 mm wide,
1 -nerved, finely puberulent, ciliate at the upper mar¬
gin. Upper glume lanceolate, apiculate, finely pu¬
berulent, 6.5-7 mm long, 3 mm wide, 10-nerved,

84

Novon

some nerves anastomosing; adaxial surface shiny,
brownish, with a few dark spots; margins ciliate.
Lemma broadly lanceolate, 7.5-8 mm long, 4 mm
wide, 9-nerved, adaxial surface shiny; abaxial sur¬
face densely pilose, the hairs ca. 0.5 mm long, silver-
grayish. Palea 8 mm long, ca. 2 mm wide (folded),
finely puberulent, 8-nerved, 2-keeled, broadly sili¬
cate abaxially, 8-nerved, the keels ciliate, the sulcus
puberulent; adaxial surface shiny, light brown, in¬
conspicuously anastomosing. Rachilla extension about
half the length of the palea, finely ciliate. Caryopsis
not seen.

Phenology. According to Smith et al. (1981)
flowering collections are known from 1971-1973.

Distribution. Brazil, State of Santa Catarina.

Merostachys glauca was originally described from
vegetative material only (McClure & L. B. Smith,
1967). Later, the same species, but in flowering
stage, was described again (Smith et al., 1981),
although without a Latin diagnosis for the floral part.
Merostachys glauca is distinguished by an unusu¬
ally densely hirsute lemma and by the long, narrow
branched leaf blades. The smooth intemodes, mot¬
tled with dark green rounded spots, the glaucous-
whitish touch upward, the dark-spotted adaxial sur¬
face of the branch leaf sheath, and the size of the
1-flowered spikelet suggest that this species resem¬
bles M. caucaiana.

Additional specimens examined. BRAZIL. Santa
Catarina: Paulo Lopes, Bom Retiro, mata, 400 m, 13
Dec. 1972 (fl), Klein & Sousa Sob. 104HO f\Y); Joinville,
Estrada Dona Francisca, mata, 15 Jan. 1974 (st), Klein
& Klein 10996 (NY).

Merostachys kleinii Sendulsky, sp. nov. TYPE:
Brazil. Santa Catarina: Papanduva, E.R.F., 1
km ao sul da entrada para Papanduva, 800 m,
16 Jan. 1974 (fr), Klein & Klein 11008 (ho-
lotype, RB; isotype, US). Figure 5.

Habitus ignotus. Fragmenta culmorum internodiorum
2-3.5 cm diametro, fistulosa, scabra, maculis et striis
atro-viridibus. Nodi pilis densis appressis, retrorsis, albis
ad basin. Folia culmorum ignota. Rami in complemento
50-120. Foliorum ramorum laminae 3-9 cm longae, 7-
15 mm latae, ianceolatae. Inflorescentia spiciformis, se-
cunda, 3-6 cm longa. Spiculae 2-flosculata, usque 16
mm longa, congestae, dense aggregatae, 2-seriatae; gluma
inferior 4 mm longa; gluma superior 12-14 mm longa,
15-nervia, apiculata; flosculi bisexuales, late fusiformes,
10-12 mm longi; lemma 10 mm longum, 16-17-nervium;
palea fusiformis, 11-12 mm longa, 12-14-nervia, late
sulcata. Rachillae projectura plana, ca. 1 mm lata; pro-
jectura flosculi superioris setiformis, rudimento ornata.
Caryopsis late ovoidea, 6-7 mm longa, atro-lutea aut
brunnea.

Habit unknown. Fragments of the culm inter¬
nodes 2-3.5 cm diam., fistulose, scabrous, gray-
greenish, with dark green spots and stripes. Culm
walls of the middle intemodes 2-3.5 mm diam.
Nodes salient, prenodal zone with dense, appressed,
retrorse, whitish silky hairs below. Culm leaves not
seen. Branch complement dense, with 50-120
branches, these 15-40 cm long, 1-1.5 mm thick,
glabrous, smooth, grayish green, 2-3-noded before
the leaves, nodes dark. Branch leaves 4-5 per com¬
plement: sheaths tight, glabrous, the margins gla¬
brous; oral setae fine, finely scabrous, deciduous;
inner ligule short, dark, finely ciliate; outer ligule a
salient rim; pseudopetiole dark, 2.5 mm long, 1 mm
wide; blades 3-9 cm long, 7-15 mm wide, L: W =
4.2-6, lanceolate, apically attenuate, adaxially
somewhat rough, 2-3 marginal nerves on one side
with rows of fine prickle hairs, abaxially glabrous
except for the densely pubescent base, the hairs
long, villous, whitish, beside the midnerve. Inflores¬
cences terminal on leafy branches, spikelike, secund,
congested, 3-6 cm long, 2 cm wide, with 28-32
densely agglomerated spikelets in double rows. Ra-
chis pilose. Pedicels short, pilose. Spikelet always 2-
flowered, 16 mm long, florets bisexual, broadly fu¬
siform, 10-12 mm long, 4 mm wide (with mature
fruit), glabrous. Glumes 2, very unequal. Lower
glume ca. 4 mm long, 1.5 mm wide, 1-nerved,
puberulent, ciliate at the margins. Upper glume 12-
14 mm long, 5 mm wide, broadly lanceolate, apic-
ulate, finely puberulent, 15-nerved, some nerves
anastomosing; adaxial surface shiny, brownish,
densely dark-spotted. Lemma 10 mm long, 6 mm
wide, broadly lanceolate, puberulent, 16-17-nerved,
some nerves anastomosing; adaxial surface shiny,
brownish, densely dark-spotted. Palea fusiform, 11"
12 mm long, ca. 6 mm wide, glabrous, 12-14-
nerved, broadly sulcate abaxially, the keels ciliate,
some nerves anastomosing; adaxial surface shiny,
golden. Rachilla extension between the florets flat,
ca. 1 mm wide, finely puberulent, hairs whitish,
bearing a fertile upper floret identical to the lower
one. Rachilla extension of the upper floret bristle¬
like, with a small rudiment at the tip. Caryopsis
broadly ovoid, basally and apically rounded, rostrate,
dark yellow, 6-7 mm long, 4-4.5 mm wide, the
embryo and the hilum scarcely manifest externally-

Phenology. Flowering collections are known only
from 1974.

Distribution. Southern Brazil.

Merostachys kleinii seems to be a robust plant,
with thick culms. This species can be found in several
herbaria, but complete vegetative collections are still

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1995

Sendulsky

Merostachys from Brazil

85

Figure 5. Merostachys kleinii Sendulsky. —a. Flowering branch complement. —b. Caryopsis, hilum view. —c.
Caryopsis, embryo view. —d. Spikelet with two fertile florets and a rudiment. —e. Palea. —f. Lemma. —g. Upper
glume. —h. Lower glume. —i. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based
on Klein & Klein 11008.

not available. The species is distinct in its 2-flowered
spikelet, the stout appearance of its branch com¬
plements, inflorescences and spikelets, and in having
the largest caryopsis in the multiramea group. For
many years this species was conceived as being a

2-flowered form of “multiramea. ” The true M. mul¬
tiramea Hackel already is a 2-flowered species. Mer¬
ostachys kleinii is named in honor of the late Rob¬
erto M. Klein, a distinguished botanist from Itajai,
Santa Catarina, Brazil.

86

Novon

Paratype. BRAZIL. Santa Catarina: 2 km a leste
de Bocaina do Sid, 900 m, 22 Jan. 1974 (fr), Klein &
Klein 11062 (SP).

Merostachys magellanica Sendulsky, sp. nov.
TYPE: Brazil. State of Sao Paulo: Municipio
Anhembi, Fazenda Barreiro Rico, 22°40'S,
48°09'W, 510 m, 11 Jan. 1988 (fr), Reis de
Magalhaes s.n. (holotype, SP 248339; iso¬
types, BLA, BR, IBGE, ICN, K, MO, NY, P,
R, RB, UB, UEC, US, W, WIS). Figure 6.

Culmi ca. 6-8 m alti, 12-14 mm diametro, fistulosi;
internodia 60-80 cm longa; infera laevia, luteo-cinerea,
striis maculisque viridi-fuscis minutis picta; supra cinereo-
viridia, aculeis retrorsis munita; nodi prominentes, supra
subterque nodum pilis lanatis densis brevibus vestita. Folia
culmorum laminae reflexae, ca. 5-12 cm longae, 12-15
mm latae. Rami verticillati, in complemento 12-130.
Folia ramorum laminae 8-17 cm longae, 1.5-2 cm latae,
lanceolatae. Inflorescentia spiciformis, pectinata, imbri-
cata, 6-8 cm longa, ramum foliorum terminans. Spiculae
11-14 mm longae, fusiformes ubi juvenes, ad maturita-
tem ovoidae, apicibus attenuatis, duabus lineis albis par-
allelis, longitudinalibus, carenis palearum formatis. Car-
yopsis oblonga, ovoidea, asymmetries, lageniformis, ros-
trata, 6-7 mm longa, castanea.

Rhizomes sympodial, pachymorph. Culms woody,
ca. 6-8 m tall, 12-14 mm diam., fistulose, erect;
intemodes 60-80 cm long; lower intemodes grayish
yellow, with minute dark green stripes and spots,
smooth and shiny above the nodes; upper intemodes
greenish gray, with small, retrorse prickle hairs;
culm walls 2 mm thick; nodes salient, with a band
of short, dense, woolly hairs above and below. Culm
leaves deciduous: sheath 25-30 cm long, 5-7 cm
wide, rounded and asymmetrical at the summit,
fringed with fine, wavy hairs, these reddish brown,
but white at the base, 5—6 mm long, diminishing in
size downward along the margins of the sheath;
abaxially olive-green, many-nerved, the nerves fine,
light-colored, smooth, shiny; blade reflexed, 5-12
cm long, 12-15 mm wide, attenuating gradually to
a fine point. Branch complement with 12-130 sub¬
equal branches, these 15-65 cm long, 1-2 mm
thick, 2-3-noded before the leaves, yellowish, nearly
glabrous and shiny, with green stripes, sometimes
with minute retrorse prickle hairs, the base with a
few densely strigose, deciduous scales, these 3-20
mm long; first intemode above the scales densely
and retrorsely strigose. Foliage leaves 4-7 per com¬
plement. Sheath tight, glabrous, striate, the lower
sheath aphyllous or with a rudimentary leaf and a
few oral setae, the uppermost a little loose at the
summit. Oral setae erect or crisped distally, 3-10
mm long, dark yellow or red-brown, slightly dimin¬
ishing in size downward along the upper margin of

the sheath, then giving way to a fine row of short,
silver cilia. Inner ligule a thick, dark yellow struc¬
ture, ca. 0.5 mm long, finely ciliate on the margin;
outer ligule a salient, fight-colored rim. Pseudope¬
tiole dark, dorsiventrally compressed, adaxially sca-
berulous and waxy, glabrous on abaxial surface.
Blades linear-lanceolate, 8-17 cm long, 1.5-2 cm
wide, L:W = 5.3-8.5 (the lowermost and the up¬
permost smaller, 2.5-6 cm long, 3-4 mm wide),
the base slightly asymmetrical, rounded to cuneate,
gradually attenuate toward a scabrous apex, sub-
glabrous on both surfaces, the adaxial surface some¬
times finely hirsute near the base and also bearing
2-8 rows of hyaline, antrorse-strigose prickle hairs
along the nerves near one margin; the nerves in¬
conspicuous on the adaxial surface, weakly salient
on the abaxial surface. Inflorescences terminal on
leafy branches, spikelike, falcate, congested, second,
6-8 cm long, with distinctive row of upper glumes
when young or in early flowering stage, when mature
and fruiting, 12-18 cm long, with loose, open,
broadly spreading spikelets; rachis finely strigose,
hairs white, sometimes with a rudimentary leaf
emerging at the top of the oblique scar at the base
of the inflorescence and subtending the lowermost
spikelet. Spikelets 11-14 mm long, 1.8-2(-4) mm
wide, 1 -flowered, slightly curved, attenuate, narrow¬
ly fusiform when immature, ovoid when mature,
swollen at the base, attenuate at the apex, grayish
olive-green, except for two parallel, longitudinal,
white, evident lines formed by the keels of the palea.
Pedicels short, densely hispid. Glumes 2, unequal.
Lower glume 3-3.5 mm long, 1 mm wide, trian¬
gular, 1-nerved, slightly curved, shortly strigose at
the apex, ca. '/ t the spikelet length, the mid-nerve
spotted adaxially, the spots dark, purplish. Upper
glume 5.5-7.5 mm long, 3 mm wide, lanceolate,
apiculate, 9-nerved, shortly strigose, % the spikelet
length, the adaxial surface shiny, with large, dark
spots; when young, glumes forming a clear fine along
the row of densely imbricated spikelets. Rachilla
segment between the upper glume and the lemma
2.5 mm long, cup-shaped, 1.2 mm diam., finely
strigose, the hairs white. Lemma 10-11.5 mm long,
5-5.5 mm wide, oblong, lanceolate, 14 - 15 -nerved,
finely strigose abaxially, shiny, dark-spotted adaxi¬
ally, ciliate at the margins, embracing the palea (at
maturity only at the base). Palea a little longer than
the lemma, 10.5-12.5 mm long, 4 mm wide, curved,
8-12-nerved, with oblique transverse veinlets, the
dorsal keels white, minutely ciliate, the adaxial sur¬
face shiny, dark-spotted. Prolongation of the rachilla
bristle-like, nearly as long as the palea, pubescent
near the minute rudiment. Lodicules 3, 3.3 mm
long, finely ciliate at the apex, faintly 2 - 3 -nerved.

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Merostachys from Brazil

87

Figure 6. Merostachys magellanica Sendulsky. —a. Lodicules. —b. Lower glume. —c. Upper glume. —d. Detail
of the abaxial surface of the upper glume. —e. Lemma. —f. Palea. —g. Spikelet, lemma side view. —h. Spikelet,
palea side view. —i. Spikelet, lateral view. —j. Bisexual flower. —k. Palea. —1. Mature inflorescence, fruiting stage.
—m. Base of the spikelet. —n. Oral setae. —o. Caryopsis hilum view. —p. Caryopsis, embryo view. —q. Caryopsis,
lateral view. —r. Apex of a blade. —s. Base of a blade, adaxial surface. —t. Branch complement of the upper and
slender part of a culm, with young inflorescences. —u. Mature spikelet, with parallel, white keels on the palea. All
illustrations based on J. G. Kuhlmann s.n. (US 1255439) except I, o, p, q, and u, which are based on Reis de
Magalhaes s.n. (SP 248339).

Stamens 3, the anthers 9 mm long. Ovary oblong,
subcylindrical; style single; stigmas 2, long-fringed
when mature. Caryopsis oblong, 6-7 mm long, (la-
genoid) curved or asymmetrical in the dorsiventral

plane, sulcate, apically long rostrate; pericarp gla¬
brous, of uniform thickness, light brown; embryo
small, basal; hilum linear, but the embryotegium
weakly manifested externally.

88

Novon

Phenology. Merostachys magellanica has been
collected in its fertile state in 1891, 1921-1922,
1956, and 1988-1990. These collections suggest
a flowering cycle of 30-34 years.

Distribution. Brazil, in the States of Rio de Ja¬
neiro and Sao Paulo, from sea level to ca. 700 m.

The first known collection of Merostachys ma¬
gellanica was made in Peruibe, Sao Paulo, in 1891,
followed by the collections in 1921 and 1922 in
Rio de Janeiro. Mature caryopses were collected in
1956 by Jose Carlos Reis de Magalhaes on his ranch,
“Barreiro Rico,” in the State of Sao Paulo (this is,
in fact, a well-kept private forest reserve). Werner
Bockermann, a zoologist at the Sao Paulo Zoological
Garden, has studied these caryopses as a food supply
for birds in the reserve.

When Merostachys magellanica flowered again
in 1988 and set seed in quantity, a large collection
was made by Reis de Magalhaes and sent to the
Institute of Botany in Sao Paulo, together with ac¬
curate data and observations concerning the habitat:
soil analysis, rainfall, climate, behavior of the bam¬
boos, etc. According to Reis de Magalhaes, the poor¬
est soil was exactly in the area where the species
was collected. The adjacent forest area, 500 m
away, with much better soil, has few or no bamboos.
Thus, this new species shows a distinct preference
for poor soil.

Merostachys magellanica is characterized by the
spikelets, which have two white, longitudinal lines
formed by the keels of the palea, and by the general
aspect of the congested inflorescences when young,
which present upper glumes in a distinctive row.
When mature, the inflorescences are twice as long
as wide, with loose, open, broadly spreading spikelets
and have light brown, dorsiventrally asymmetrical
caryopses.

Merostachys magellanica is named for Jose Car¬
los Reis de Magalhaes, collector of the type collec¬
tion, who in his respect for and love of nature knows
few equals.

Paratypes. BRAZIL. Rio de Janeiro: Botafogo,
Mundo Novo, 11 Nov. 1921 (early fl), J. G. Kuhlmann
s.n. (RB 79198, US 1447816k Corcovado, Morro do
Mundo Novo, Oct. 1922 (fl), J. G. Kuhlmann s.n. (US
1255439); Reserva Florestal da Fabrica Alian^a, Sep.
1922 (fl-abnormal), J. G. Kuhlmann s.n. (RB 3852, US
1447817). Sao Paulo: Peruibe, 1891 (fl), Loefgren &
Edwall s.n. (US 1021511); Municipio Itatins, Iguape,
Esta^ao Ecologica Jureia, 19 Nov. 1987 (fl), Catharino
1334 (SP); Municipio Anhembi, Fazenda Barreiro Rico,
21 fev. 1956 (fr), Reis de Magalhaes s.n. (SP 248383);
Cananeia, Out. 1956 (fr), Pereira s.n. (SP 169270);
Parque Estadual da Ilha do Cardoso, 8 Dec. 1987 (fl),
Kirizauia 1984 (SP); Ilha do Cardoso, estrada para tur-
bina, 3 Dec. 1990 (fl), Barros 2024 (SP); Ilha do Cardoso,
Cambriu, 23 Aug. 1988 (late fr), Rossi et al. 486 (SP).

Merostachys multiramea Hackel. In Feddes Re-
pert. Nov. Spec. Regni Veg. 7: 326. 1909.
TYPE: Brazil. Rio Grande do Sul: Municipio
Rio Pardo, Fazenda Itacolomi, elev. 70 m, Apr.
1906 (fl), Jurgens s.n. (label G-308) (holotype,
W). Figure 7.

Merostachys anomala Dutra, Revista Sudamer. Bot. 5:
151. 1938. Fig. 2, p. 149, under M. burchellii
Munro emend. Dutra (figures misplaced). Syn. nov.
TYPE: Brazil. Rio Grande do Sul: Sao Leopoldo,
Fazenda da Pedreira, 6 Oct. 1906 (fl), Dutra 518
(holotype, BLA; isotype, US).

Culms woody, fistulose, terete, yellowish gray,
glabrous, smooth, but densely pilose near the nodes,
12 m long, according to C. Jurgens s.n. (label G-
308); pendulous above, 6-9 m high, 3-4 cm diam.
below, according to Dutra (1938); nodes prominent,
glabrous, densely pilose above, pilose and also with
a row of white, dense, brushlike, retrorse hairs below.
Culm leaves of an incompletely developed shoot (see
discussion): sheaths 20-25 cm long, 5-6 cm wide,
symmetrical, papery, opaque, rugose adaxially,
fringed with dense, wavy, reddish hairs at the sum¬
mit, diminishing in size downward along the upper
margin; inner ligule a thick membrane, 1-2 mm
long, opaque; outer ligule a salient rim; blades lan¬
ceolate, asymmetrical, ca. 7 cm long, 1 cm wide,
gradually attenuate to a very fine point. Branch
complement with 30-70 unequal leafy branches,
these 5-50 cm long, 1-1.2 mm thick, glabrous,
smooth, grayish green, 2-3-noded before the leaves;
nodes prominent, dark, the base densely covered
with small, deciduous, apiculate, glabrous scales.
Foliage leaves 3-8 per complement, the leaves of
the flowering branches smaller than those of the
sterile ones. Sheaths tight, glabrous, striate, the mar¬
gins finely ciliate or glabrous. Oral setae 3-4 mm
long, straight, white, extremely deciduous. Inner
ligule papery, finely ciliate on the margin, 0.5 mm
long. Outer ligule a salient rim. Pseudopetiole gla¬
brous, dark, flat, 2.5-4 mm long. Blades 3-20 cm
long, 5-20 mm wide, lanceolate or narrowly lan¬
ceolate, attenuate to a very narrow, filiform point,
the base rounded to cuneate, slightly asymmetrical,
subglabrous on both surfaces except for a small area
on the abaxial surface near the base, bearing short,
white, dense hairs mixed with wax and for 2-3
lateral nerves on the adaxial surface near one mar¬
gin, bearing parallel rows of antrorse, strigose, hy¬
aline prickle hairs. Inflorescence 5—7(—9) cm long,
straight or subfalcate, secund, terminal on a leafy
branch, usually fully exserted from the uppermost
sheath when mature. Peduncle of the inflorescence
and the rachis densely strigose, the hairs white.

Volume 5, Number 1
1995

Sendulsky

Merostachys from Brazil

89

Figure 7. Merostachys multiramea Hackel. —a. Part of a branch complement with some young inflorescences,
—a,. Mature inflorescence. —b. Part of the spike. —c. Lower glume. —d. Upper glume. —e. Lemma. —f. Palea,
abaxial surface. —g. Palea, adaxial surface. —h. Spikelet, glumes removed. —i. Bisexual flower. — j. Lodicules. —
k. Base of a leaf, adaxial surface. —1. Caryopsis, hilum view. —m. Caryopsis, lateral view. —n. Caryopsis, embryo
view. —o. Base of a leaf, adaxial surface, showing ligule and a few oral setae. —p. Culm leaf. All illustrations based
on Dutra 518 , except a,, which is based on Dutra 517.

Spikelets 2-flowered, 11-12 mm long, grayish green,
with wrinkled surface, both florets bisexual, 7-9 mm
long, 3 mm wide, laterally strongly compressed when
immature, prominently keeled, imbricate at an angle

of 45°. Pedicel ca. 1 mm long, densely hispid. Glumes
2, unequal. Lower glume 2.5-3 mm long, 1 mm
wide, lanceolate, 1-nerved, the abaxial surface pu-
berulent, the margins densely ciliate. Upper glume

90

Novon

7- 9 mm long, ca. 5 mm wide, half as long as upper
floret, carinate, apiculate, asymmetrical, 9-nerved,
the nerves anastomosing; abaxial surface finely hir-
sute-appressed, the margins finely ciliate. Rachilla
segment between the lower and the upper florets 2-
3.5 mm long, flattened, narrow below, dilated at the
apex, finely appressed-pubescent. Lemma 8 mm long,
5 mm wide, carinate, adaxially slightly dark-spotted,
9-11-nerved. Palea 8 mm long, curved, bicarinate,
broadly canaliculate between the finely ciliate keels,

8- nerved, the veinlets anastomosing in the upper
part. Prolongation of the rachilla (on the upper floret
only) bristle-like, flattened, half as long as the palea
or a little over, bearing a minute rudiment. Lodicules
3, membranous, ca. 0.7 mm long, ciliate at the
apex. Stamens 3, the anthers 4-5.5 mm long. Ovary
oblong, subcylindrical; style single; stigmas 2, plu¬
mose. Caryopsis 6-7 mm long, 3 mm wide, conical,
attenuate at the apex, rostrate, dark brown, nearly
black, shiny; embryo small, basal; hilum linear, but
the sulcus and the embryo weakly manifested ex¬
ternally.

Phenology. Flowering collections are known from
1876 (Dutra, 1938), 1906-1907, 1914-1915,
1937-1939, and 1974-1978.

Distribution. Southern Brazil.

The description of the culm leaves of Merosta-
chys multiramea is based on Prance et al. 6887,
the only collection in which these leaves are present.
This material is not very typical of the species;
together with Gibbs et al. 6672 and Ihering s.n.
(NY, SP 10315) it makes up a group of three similar
specimens, all collected in the State of Sao Paulo.
If these specimens can be admitted as good M.
multiramea, the description of the culm leaves is
valid. The only difference between these collections
and the others is that the spikelets are not wrinkled
and the short hairs that cover the spikelets are black
and not so worn out and colorless as on the spikelets
of the specimens from the State of Rio Grande do
Sul.

Additional specimens examined. BRAZIL. Parana:
Ipiranga, Dusen 14367, 5 Jan. 1914 (fl) in US sheet,
and 15 Jan. 1915 (fl) in WIS sheet. Rio Grande do
Sul: Valle do Rio Taquari, Sao Leopoldo, 1907 (fr), Dutra
s.n. (NY, SP 10313, US 1503502); Suburbios, Oct. 1906
(fl), Dutra 151 (RB); Bom Jesus, Fazenda Carauna, 1907
(fl and fr), Dutra 517 (BLA); Sao Leopoldo, Quilombo,
Nov. 1937 (fl), Dutra 1634, (ICN); Panambi, 4 Oct.
1970 (fl), Pott s.n. (BLA 8363). Sao Paulo: Reserva
Carlos Boteiho, entre S. Miguel Arcanjo e Sete Barras,
elev. ca. 800 m, 31 Jan. 1978 (fl), Prance et al. 6887
(RB, US); Rodovia Jaquia a Piedade, 29 Oct. 1977 (fl),
Gibbs et al. 6672 (SP); Sao Bernardo, Oct. 1914 (fl),
Ihering s.n. (NY, SP 10315).

Merostachys pilifera Sendulsky, sp. nov. TYPE:
Brazil. Rio Grande do Sul: Mun. Sao Francisco
de Paula, Primeiro Distrito, Sao Francisco de
Paula, Morro da Pera (uppermost part of Serra
Geral), 29°27'S 50°35'W, alt. 970 m, 11 May
1972 (fl), Soderstrom & Sucre 1987 (holotype,
RB). Figure 8.

Culmi usque 20 m longi, fistulosi ad extremitates solidi.
Culmi internodia dense pilosa supra et infra nodos, pilis
translucidis, appressis, ca. 1 nun longis. Folia culmorum
ignotae. Crista supranodalis tubiformis. Rami in comple-
mento 50 ad extremitates culmorum. Foliorum ramorum
laminae lanceolatae, 2-8 cm longae, 4-12 mm latae.
Inflorescentia spiciformis, pectinata. Spiculae 1-flosculata,
bisexualis, late elliptica, 13-14 nun longa; gluma infera

2.5 mm longa; gluma supera 9 mm longa; 11-nervia;
lemma 10 mm longum, 15-nervium; palea fusiformis, 11
mm longa, 10-12-nervia, anguste sulcata. Caryopsis non
visa.

Woody, according to the collectors’ note, culms
erect, up to 20 m tall, 5 cm diam., closely clumped
and forming a thicket; internodes fistulose, 60-65
cm long at base, with thin walls, becoming shorter,
successively thinner and solid above; the tips whip¬
like, arching. New culm intemodes densely pilose
above and below the nodes, the hairs sericeous,
translucent, appressed, thick, ca. 1 mm long, these
substituted in some occasionally hairless parts of the
intemode areas by dense, sharp, retrorse prickle
hairs with globose, yellowish bases, together with
some scattered sericeous hairs. The old thick culms
still display prickles and areas with worn and dam¬
aged hairs. Nodes salient, supranodal ridge tubiform,
especially evident in new culms, prenodal zone with
dense, appressed, retrorse hyaline hairs. Culm leaves
not seen. Branch complement from tip of the culm
(intemode 5 mm diam.) with 50 branches, these
15-20 cm long, ca. 0.5 mm thick, glabrous, smooth,

1- 2-noded before the leaves. Branch leaves 2-3
per complement: sheaths tight, finely puberulent;
the overlapping margin glabrous; adaxial surface
shiny, golden, dark-spotted; oral setae fine, ca. 5-
6 mm long, straight; inner ligule short, finely ciliate;
outer ligule an inconspicuous rim; pseudopetiole dark,
finely puberulent, 2-2.5 mm long; blades lanceolate,

2- 8 cm long, 4-12 mm wide, L:W = 6.2-6.6,
glabrous, the abaxial surface with a very few fine
hairs. Inflorescences terminal on leafy branches, 2-

3.5 cm long, spikelike, pectinate. Rachis and ped¬
icels pubescent. Spikelets solitary, 1-flowered, bi¬
sexual, broadly ellipsoid, somewhat inflated, 13-14
mm long, 4 mm wide. Glumes 2, unequal. Lower
glume 2.5 mm long, ca. 1 mm wide, shortly awned,
1-nerved. Upper glume broadly lanceolate, 9 mm
long, 4 mm wide, shortly awned, 11 -nerved, some
nerves anastomosing, the adaxial surface dark-spot-

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Sendulsky

Merostachys from Brazil

91

Figure 8. Merostachys pilifera Sendulsky. —a. Flowering branch complement. —b. Lodicules. —c. Hermaphroditic
flower. —d. Spikelet. —e. Apical portion of the inflorescence. —f. Culm hairs. —h. Lower glume. —i. Upper glume.
—j. Lemma. —k. Palea. —1. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based
on Soderstrom & Sucre 1987.

ted. Lemma very broadly lanceolate, deeply con¬
cave, enclosing a palea, 10 mm long, 7 mm wide,
15-nerved, finely puberulent abaxially, the adaxial
surface dark-spotted, the margins densely ciliate
upward. Palea fusiform, 11 mm long, 2.5 mm wide

(folded), 10-12-nerved, 2-keeled, narrowly sulcate,
the keels finely ciliate, abaxially puberulent at the
apex and along the sulcus; some nerves anastomos¬
ing; adaxial surface shiny. Lodicules 3, subequal,
ca. 2.5 mm long, finely ciliate. Stamens 3, the

92

Novon

anthers dark, ca. 10 nun long. Ovary elongate, style
single, stigmas 2. Caryopsis not seen.

Phenology. The only known specimen of this
species is the holotype, collected in 1972.

Distribution. Known only from the type locality.

Merostachys pilifera is known only from the type
locality in the State of Rio Grande do Sul. The
description presented above was based solely on
Soderstrom & Sucre 1897, 1972. The label states
that the entire colony of this bamboo flowered after
this date and is represented by Soderstrom 2040,
2 Feb. 1973 (US). Merostachys pilifera is distin¬
guished by its pilose culm internodes, the character
from which the name is derived. It is somewhat
similar to M. abadiana in the details of the spikelet
structure.

Merostachys scandens Sendulsky, sp. nov. TYPE:
Brazil. State of Sao Paulo: Municipio de Sao
Paulo, 10 km S of center of city of Sao Paulo,
in grounds of the Instituto de Botanica, Parque
do Estado, 6 Dec. 1973 (fl), Sendulsky 1319
(holotype, SP; isotypes, BLA, BR, IBGE, ICN,
K, MO, NY, P, R, RB, UB, UEC, US, W,
WIS). Figure 9.

Culmi ca. 6 m longi, 1-3 cm diametro; internodia
fistulosa, subtiliter rugulosa; nodi pilis densis, appressis,
retrorsis. Foliorum culmorum laminae angustae, 8-12 cm
longae, 4-5 mm latae. Rami in complemento 40-60(-
100). Foliorum ramorum laminae lanceolatae, glauco-
virides, 3.5-10 cm longae, 8-14 mm latae. Inflorescen-
tiae spiciformes, 5-6 cm longae, ramorum foliorum ter-
minantes. Spiculae ellipticae, 1-flosculatae, 13-14 mm
longae, distichae, lateraliter dispositae; gluma infera 4.5-
5 mm longa; gluma supera 9-10 mm longa, 15-nervia;
lemma 11 mm longum, 19—23-nervium, ad marginem
superiorem strigosum; palea fusiformis, 11 mm longa, 12-
nervia, anguste sulcata. Caryopsis ovoidea ad apicem sub¬
tiliter attenuata, 5.5-6.5 mm longa, cinereo-lutea.

Caespitose, woody bamboo. Culms ca. 6 m long,
1-3 cm diam., flexuous, with long, attenuate,
threadlike, pendent or clambering tips. Internode
fistulose, finely rugose, dark gray, with dull green
stripes. Culm walls ca. 2 mm thick. Nodes salient,
prenodal zone with dense appressed whitish, retrorse
hairs. Culm leaves deciduous: sheaths 15-25 cm
long, 4-8 cm wide, narrowing toward the summit,
fringed at the apex with 1-cm-long, golden reddish,
distally crisped hairs; abaxial surface glabrous,
densely nerved; adaxial surface somewhat shiny or
dark, apically tessellate; margins glabrous; inner lig-
ule 1 mm long, finely ciliolate; outer ligule a salient
rim; blades reflexed, deciduous, narrow, 8-12 cm
long, 4—5 mm wide, attenuate, forming a fine point.
Branch complement with 40-60(-100) branches.

these 30-60 cm long, 1-1.5 mm thick, grayish,
glabrous or puberulent, 2-3-noded before the leaves,
nodes glabrous. Branch leaves 5-6 per complement:
sheaths tight, glabrous, the intercostal zone finely
transversely rugose, the overlapping margin gla¬
brous; oral setae fine, deciduous, ca. 3 mm long;
inner ligule ca. 0.8 mm long, finely ciliolate; outer
ligule a salient rim; pseudopetiole flat, slightly pu¬
berulent, 2.5 mm long; blades 3.5-10 cm long, 8-
14 mm wide, L: W = 4.4-7.1, lanceolate, glaucous-
green, adaxially glabrous, 2-3 marginal nerves on
one side with rows of very fine prickle hairs; abaxial
surface glabrous except for finely strigose base, the
margins and the apex very finely scabrous. Inflo¬
rescences terminal on leafy branches, spikelike, 5-
6 cm long, with 17-18 spikelets, these distichously
arranged, distinctly separate and laterally outspread
along the narrow, obscurely triquetrous, slightly sin¬
uous, densely pilose rachis or rarely pectinate. Spike-
lets solitary, 1-flowered, bisexual, ellipsoid, 13-14
mm long, 3 mm wide. Glumes 2, unequal. Lower
glume 1 -nerved, carinate, attenuate toward the apex,
finely puberulent, 4.5-5 mm long, ca. 1 mm wide
at the base. Upper glume 9-10 mm long, 4 mm
wide, broadly lanceolate, carinate, finely puberulent,
15-nerved, some nerves anastomosing, the apex sca¬
brous, aristulate, ca. 1 mm long; adaxial surface
shiny, dark-spotted; margins finely ciliate. Lemma
11 mm long, 5-5.5 mm wide, lanceolate, 19-23-
nerved, the nerves anastomosing; abaxial surface
somewhat strigose, the hairs appressed, longer and
denser toward the upper margins, these densely
ciliate; adaxial surface shiny, dark-spotted. Palea
fusiform (folded), 11 mm long, ca. 4 mm wide,
glabrous, opaque, 2-keeled, narrowly sulcate abax-
ially, 12-nerved, the nerves obliquely anastomosing
in the upper part; keels finely ciliate, the hairs dark
red; adaxial surface shiny. Rachilla extension bristle¬
like, 9 mm long, with a minute rudiment. Lodicules
3, subequal, 2.5 mm long, finely ciliate at the apex.
Stamens 3, the anthers 12 mm long, dark. Ovary
elongate, the style single, the stigmas 2, plumose;
stigmatic papillae sharply pointed. Stamens 3, the
anthers 12 mm long, dark. Caryopsis 5.5-6.5 mm
long, ca. 3 mm wide, ovoid, apically slightly atten¬
uate, rostrate, grayish yellow, smooth, apically slightly
attenuate, the embryo and the hilum scarcely man¬
ifested externally.

Phenology. The flowering collections are known
from 1941 and 1972-1974, thus suggesting a flow¬
ering cycle of 31-33 years.

Distribution. Southern Brazil.

Merostachys scandens was in bloom at the same
time as M. skvortzovii (described later in this paper)

Volume 5, Number 1
1995

Sendulsky

Merostachys from Brazil

93

Figure 9. Merostachys scandens Sendulsky. —a. Flowering branch complement. —b. Gynoecium with two lodicules.
—c. Stigmatic papillae. —d. Spikelet in bloom (flowering cycle 1973). —e. Lodicules. —f. Apical part of the
inflorescence. —g. Culm leaf. —h. Caryopsis, hilum view. —i. Caryopsis, embryo view. —j. Lower glume. —k.
Upper glume. —1. Lemma. —m. Palea. —n. Upper portion of the branch leaf sheath and base of the blade (abaxial
surface). Based on Sendulsky 1319.

and in the same forest of the Biological Reserve, on
the grounds of the Institute de Botanica, Sao Paulo.
The plants did not form dense clumps, but were
slender and leaning on vegetation. The glaucous-
green leaves are distinct from the green leaves of

M. skvortzovii. Because the plant was deep in the
shaded forest, the visiting insects were not observed
(see the discussion in M. skvortzovii). The habit of
the inflorescence, its widely separated distichous
spreading spikelets, is of some interest. This orga-

94

Novon

nization of the inflorescence is quite rare in the genus
and contradicts the significance of the name Mer-
ostachys, formed from the Greek meros, part (par¬
tial or incomplete), and stachys, spike, alluding to
the secund (one-sided) profile of the inflorescence
(McClure, 1973). In M. scandens the inflorescence
may sometimes also be pectinate, when for some
unknown reason the very narrow rachis bends lon¬
gitudinally over the ventral surface, bringing to¬
gether its margins and the spikelets of two opposite
rows.

Paratypes. BRAZIL. Sao Paulo: Municipio de Sao
Paulo, 10 km S of center of city of Sao Paulo, in grounds
of the Instituto de Botanica, Parque do Estado, 6 Dec.
1973 (fl), Sendulsky 1317 (SP), 6 Dec. 1973 (fl), Sen-
dulsky 1320 (SP, US), 17 Dec. 1973 (fl), Sendulsky
1323 (SP, US); Reserva Biologica, Parque Estadual das
Fontes do Ipiranga, 8 Mar. 1974 (fr), Sendulsky 1365
(SP); Sao Paulo, nativa no Jardim Botanico, 1 Dec. 1941
(fl), O. Handro s.n. (SP sheet 79756).

Merostachys skvortzovii Sendulsky, sp. nov.

TYPE: Brazil. Estado de Sao Paulo: Sao Paulo,

Parque do Estado e Jardim Botanico, 20 Oct.

1973, Skvortzov s.n. (holotype, SP 120954).

Figure 10.

Culmi 4-6 m longi, 2-3 cm diametro; internodia fis-
tulosa, scabra; nodi subpilosi, pilis albis. Foliorum cul-
morum laminae lanceolatae, 6-10 cm longae, 10 mm
latae. Rami in complemento 150-200. Foliorum ramo-
rum laminae lanceolatae, 3-11 cm longae, 6-16 mm
latae. Inflorescentia spiciformis, pectinata, 1.5-3 cm lon-
ga, ramorum foliorum terminans. Spiculae 10-12 mm
longae, fusiformes, 1-flosculatae; gluma infera 1.5 mm
longa; gluma supera 8-9 mm longa, 9-11-nervia, aris-
tulata; lemma ca. 9 mm longum, 16-18-nervium, pub-
erulum, ad apicem strigosum; palea ca. 10 mm longa,
13-14-nervia, anguste sulcata. Caryopsis ovoidea, ad ap¬
icem globosa, 5 mm longa et ca. 3 mm lata, cinereo-latea.

Caespitose, woody bamboo, forming dense clumps.
Culms 4-6 m long, 2-3 cm diam., erect except for
the flexuose tips. Internodes fistulose, scabrous, with
dark green stripes. Culm walls of the middle inter-
nodes 1-1.5 mm thick. Nodes salient with a touch
of whitish pubescence above and below, younger
nodes with short, tubiform supranodal ridge. Culm
leaves deciduous: sheath 15-20 cm long, 5-6 cm
wide, auriculate, fringed at the apex with a few hairs,
abaxial surface glabrous, densely nerved, dark; ad-
axial surface opaque, dark, striate, tessellate above;
inner ligule 1 mm long, finely ciliate at the margin,
the cilia white; outer ligule a short inconspicuous
rim; margins glabrous; blades lanceolate, reflexed,
deciduous, 6-10 cm long, 10 mm wide. Branch
complement with 150-200 slender branches, these
20-50 cm long, 1 mm thick, glabrous or finely
puberulent, the intemodes ca. 4-6 cm long, 2-3-

noded before the leaves. Branch leaves 5-7 per
complement: sheaths tight, glabrous, striate; oral
setae fine, 5-6 mm long; inner ligule a finely ciliolate
membrane, ca. 1 mm long; outer ligule a short rim;
pseudopetiole flat, 3 mm long, finely puberulent;
blades lanceolate, 3-11 cm long, 6-16 mm wide,
L:W = 5-6.9, adaxially glabrous, slightly pilose
abaxially, densely pilose at the base; when old, gla¬
brous on both surfaces except at the base abaxially;
adaxial surface with 3-4 marginal nerves on one
side with rows of fine prickle hairs; margins finely
scabrous. Inflorescences terminal on leafy branches,
spikelike, pectinate, 1.5-3 cm long, with 16-18
spikelets. Rachis and pedicels pilose, some of the
pedicels present at base on inner side small, swollen,
whitish, rugose projections. Spikelets 10-12 mm
long, 3 mm wide, solitary, 1-flowered, acuminate,
fusiform. Glumes 2, unequal. Lower glume 1-nerved,
carinate, acuminate, finely puberulent, 1.5 mm long,
0.8 mm wide at the base. Upper glume 8-9 mm
long, 3.5 mm wide, broadly lanceolate, finely pu¬
berulent, 9-11 -nerved, the nerves anastomosing in
the upper part, the apex scabrous, aristulate, ca. 1
mm long; adaxial surface shiny, dark-spotted. Upper
glume sometimes subtending a rudimentary bud
(vestige of a depauperate floret attached to rachilla
segment). Lemma ca. 9 mm long, 6 mm wide, broad¬
ly lanceolate, 16-18-nerved, the nerves obliquely
anastomosing in the upper part; abaxial surface pu¬
berulent, somewhat strigose in the upper part; ad¬
axial surface dark-spotted; margins ciliate. Palea ca.
10 mm long, 5 mm wide, glabrous, 13-14-nerved,
2-keeled, narrowly sulcate abaxially, the keels finely
ciliate; adaxial surface shiny. Rachilla extension bris-
tle-like, 8 mm long, with a minute rudiment. Lod-
icules 3, subequal, 3.2 mm long. Ovary elongate;
stigmas 2, plumose, the stigmatic papillae elongate,
rounded at the tips; stamens 3, the anthers 6 mm
long, dark. Caryopsis 5 mm long, ca. 3 mm wide,
ovoid, apically rounded, rostrate, grayish yellow,
smooth; hilum linear, the embryo hardly visible.

Phenology. Flowering collections are known from
1941 and 1972-1974, suggesting a flowering cycle
of 31-33 years.

Distribution. Southern Brazil.

Most collections of Merostachys consist of frag¬
mentary flowering branches. The vegetative parts
of the plants are nearly always omitted. Conse¬
quently, the descriptions and measurements of the
leaves, branch complements, culm and culm leaves,
when available, are based on flowering material;
usually they are gathered from the easily reachable
upper branches of the plant, which are always small¬
er in size and therefore less representative of the

Volume 5, Number 1
1995

Sendulsky

Merostachys from Brazil

95

Figure 10. Merostachys skvortzovii Sendulsky. —a. Flowering branch complement. —b. Gynoecium with two
lodicules. —c. Stigmatic papillae with pollen grains. —d. Bisexual flower with one lodicule. —e. Lodicules. —f.
Spikelet in bloom (flowering cycle 1973). —g. Apical portion of the inflorescence. —h. Lower glume. —i. Palea.
—j. Lemma. — k. Upper glume. —1. Rudimentary bud attached to rachilla segment. —m. Upper portion of the
branch leaf sheath and base of the blade (abaxial surface). —n. Caryopsis, embryo view. —o. Caryopsis, hilum view.
— p. Culm leaf. Based on the type specimen Skvortzov s.n. (SP 120954).

plant as a whole. Deciduous and decaying culm
leaves from basal or midculm nodes, originated dur¬
ing the initial growing period, are lost by the time
of flowering. The plant stops producing any new
vegetative organs and turns to flowering. When it

is possible to find the vegetative, but complete col¬
lection, made from the same plant, the branch leaves
are twice as large and the culm leaves are remark¬
ably larger. Unfortunately, it is very seldom that a
correctly identified collection displays both vegeta-

96

Novon

tive and flowering material. The vegetative mate¬
rials, Soderstrom 2033 (US 2909096) and Sod-
erstrom 2035 (US 2909100), show large culm
leaves, probably of M. skvortzovii, under the name
M. multiramea. The culm leaf sheaths are 25-30
cm long, 18-19 cm wide, culm leaf blades 8.5 cm
long, 25 mm wide.

During the 1972-1974 flowering cycle, the flow¬
ers showed signs of being of interest to a variety of
insects.

This species is named in honor of the late Boris
V. Skvortzov, Professor of Botany, Chinese Forest
Academy, Harbin, Manchuria, who dedicated his
later years to the study of the Brazilian flora, col¬
lector of the type material, and the beloved father
of the author.

Paratypes. BRAZIL. Parana: Municipio Palmas, Es¬
trada Palmas-Ponte Serrado, 5 Dec. 1971 (fl), Hatsch-
bach et al. 28267 (NY, SP). Sao Paulo: Sao Paulo,
Reserva Biologica, Parque Estadual das Fontes do Ipi-
ranga, 7 fev. 1974 (fl), Sendulsky 1341 (SP, WIS); 10
km S of center of city of Sao Paulo, in grounds of the
Instituto de Botanica, Parque do Estado, 10 Oct. 1973
(fl), Sendulsky 1314 (SP), 20 Oct. 1973 (fl), Sendulsky
1315 (SP, WIS), 6 Dec. 1973 (fl), Sendulsky 1318 { SP,
WIS), 17 Dec. 1973 (fl), Sendulsky 1321 (SP, WIS),
17 Dec. 1973 (fl), Sendulsky 1322 (SP, WIS), 23 Dec.
1973, Sendulsky 1324 (SP, WIS); nativa no Jardim
Botanico, 21 Oct. 1941 (fl), Gehrl. s.n. (SP 46002). Rio
Grande do Sul: Municipio Colorado, km 251-252 on
road between Carazinho and Bonambi, near Carazinho,
555 m, 30 Jan. 1973 (fl), Soderstrom 2029 (US); Depois
de Caracol, ca. 10 km N de Canela, 17 dez. 1972, Porto,
Girardi & Lindeman s.n. (ICN); Municipio tenente Por-
tela, 3 km S of Derrobadas, 31 Jan. 1973, Soderstrom
2033 (US); Reserva Florestal do Turvo, 31 Jan. 1973,
Soderstrom 2035 (US).

Acknowledgments. This study has been sup¬
ported by the Conselho Nacional e Desenvolvimento
Cientifico e Tecnologico (CNPq). I thank the staff

of the U.S. National Herbarium, Smithsonian In¬
stitution, for assistance and for making their collec¬
tions available for study, Tarciso S. Filgueiras (IBGE)
for the Latin diagnoses, A. G. Burman for interest
and collaboration in the initial phase of this study,
Fabio de Barros for helpful comments on the manu¬
script, and the anonymous reviewers for their critical
readings of the manuscript. Special gratitude is ex¬
pressed to Marshall R. Crosby for helpful sugges¬
tions, and to Amy Scheuler McPherson (MO) and
Lynn G. Clark, Department of Botany, Iowa State
University, for kind assistance and revision of the
original manuscript. The plates were drawn by the
author.

Literature Cited

Dutra, J. 1938. Les Bambusees de Rio Grande du Sud.

Revista Sudamer. Bot. 5(5/6): 145-152.
Filgueiras, T. S. 1988. Bambus nativos do Distrito
Federal, Brasil (Gramineae: Bambusoideae). Revista
Brasil. Bot. 11: 47-66.

Hackel, E. 1909. Gramineae Novae. VI. Repertorium
Specierum Novarum Regni Vegetabilis 7: 311-327.
Martius, C. F. P. von. 1840-1906. Flora Bras-
iliensis. . .14 volumes (in 40), published in 130 fas¬
cicles. Munich, Vienna. [Bambuseae by J. C. Doell,
1880: volume 2(3): 161-219 + plates 44-56.]
McClure, F. A. 1973. Genera of bamboos native to the
New World (Gramineae: Bambusoideae). [Edited by
T. R. Soderstrom.] Smithsonian Contr. Bot. 9: 1-
148.

-& L. B. Smith. 1967. Bambuseae. Pp. 1-78

in P. R. Reitz (editor). Flora Ilustrada Catarinense,
Pt. 1 Gramineas—Suplemento. Herbario “Barbosa
Rodrigues,” Itajai, Santa Catarina, Brasil.

Munro, W. 1868. A monograph of the Bambusaceae,
including descriptions of all the species. Trans. Linn.
Soc. London 26: 1-157, 6 plates.

Smith, L. B., D. C. Wasshausen & R. M. Klein. 1981.
Gramineas. In: P. R. Reitz (editor), Flora Ilustrada
Catarinense, Vol. 1. Herbario “Barbosa Rodrigues,’
Itajai, Santa Catarina, Brasil.

New Combinations in North American Schoenoplectus,
Bolboschoenus, Isolepis, and Trichophorum (Cyperaceae)

S. Galen Smith

Department of Biology, University of Wisconsin-Whitewater, Whitewater,
Wisconsin 53190, U.S.A.

Abstract. Correct names are provided for Vol¬
ume 11 of the Flora of North America for taxa
segregated from Scirpus sensu lato. Schoenoplectus
validus is placed in synonymy under Sch. taber-
naemontani. New combinations made herein are:
Sch. acutus var. occidentalism Sch. x steinmetzii
(pro sp.; = Sch. heterochaetus x tabernaemon-
tani); Sch. pungens var. longispicatus; Sch. pun-
gens var. badius (Scirpus americanus subsp. mon-
ophyllus var. monophyllus, misapplied); Sch.
x contortus (= Sch. americanus x pungens); Sch.
hallii; Bolboschoenus glaucus (Lamarck) (Scirpus
tuberosus, misapplied); Isolepis molesta; Tricho¬
phorum dementis; and T. clintonii.

The following new combinations, synonymies, and
typifications provide correct names in Schoeno¬
plectus, Bolboschoenus, Isolepis, and Trichopho¬
rum, all segregated from Scirpus sensu lato (Cy¬
peraceae) for the upcoming Volume 11 of the Flora
of North America (Morin et al. [in prep.]). Descrip¬
tions and keys for identifying the taxa included here
will be provided in the upcoming Flora. Types ex¬
amined for this study by A. E. Schuyler, who pro¬
vided me with information on them, are indicated
by !Schuyler. Types that I have seen only as pho¬
tocopies or photographs provided by herbarium cu¬
rators or microfiches in libraries are indicated by
Iphoto.

1. Schoenoplectus acutus (G. H. E. Muhlenberg
ex J. Bigelow) A. Love & D. Love, Bull. Torrey
Bot. Club 81: 33. 1954. Scirpus acutus G. H.
E. Muhlenberg ex J. Bigelow, FI. Boston.: 15.
1814. Schoenoplectus lacustris L. subsp. acu¬
tus A. Love & D. Love, Taxon 30: 849. 1981.
TYPE: U.S.A. Massachusetts: Fresh Pond [near
Cambridge], July 1853, Herb. William Boott,
s.n. (neotype, GH, selected here).

I have selected as neotype the only known spec¬
imen of Scirpus lacustris as traditionally defined
that was collected at Fresh Pond. No specimens

from the Boston area that could have been seen by
Bigelow have been located (Pamela White, GH, pers.
comm.). Also, there is no direct evidence that Big¬
elow saw the Muhlenberg herbarium specimens; these
add further confusion because each of the two sheets
(labeled maculatus) in Folder 63 bears a mixed
collection of seven or eight culms of Schoenoplectus
acutus, Sch. tabernaemontani, and putative hy¬
brids as described by me (Smith, 1969).

Schoenoplectus acutus is part of a difficult world¬
wide group that includes Sch. lacustris (L.) Palla,
Sch. tabernaemontani (C. C. Gmelin) Palla, and
Sch. heterochaetus (Chase) Sojak. Koyama (1962)
treated the entire complex as the single species
Scirpus lacustris and placed Scirpus acutus in syn¬
onymy under Scirpus lacustris subsp. glaucus (J.
E. Smith) C. J. Hartman (see Sch. tabernaemontani
below).

Two varieties are here recognized for North
America:

a. Schoenoplectus acutus var. acutus.

Scirpus lacustris L. var. condensatus C. H. Peck, Rep.
(Annual) Regents, Univ. State New York, New York
State Mus. 53: 853. 1900. Syn. nov. Scirpus val¬
idus Vahl var. condensatus (C. H. Peck) Beetle,
Univ. Wyoming Pubis. 13: 6. 1948. TYPE: U.S.A.
New York: [Cattaraugus Co.], Lime Lake, 5 and 20
Aug. 1899, F. E. Fenno 17 (holotype, NYS, ’Schuy¬
ler).

Distribution. Eastern North America.

b. Schoenoplectus acutus var. occidentalis (S.

Watson) S. G. Smith, comb. nov. Basionym:
Scirpus lacustris var. occidentalis S. Watson,
Bot. California 2: 218. 1880. Scirpus occi¬
dentalis (S. Watson) Chase, Rhodora 6: 68.
1904. Scirpus acutus var. occidentalis (S.
Watson) A. A. Beetle, Univ. Wyoming Pubis.
13: 5. 1948. TYPE: U.S.A. New Mexico:
1851-1852, C. Wright 1940 (mature achenes)
(lectotype, NY, selected here; isolectotype, UC).

Novon 5: 97-102. 1995.

98

Novon

Scirpus malheurensis L. F. Henderson, Rhodora 32: 20.

1930. Scirpus acutus var. malheurensis (L. F. Hen¬
derson) A. A. Beetle, Univ. Wyoming Pubis. 13: 5.
1948. TYPE: U.S.A. Oregon: Harney Co., in water
or on wet shores of Malheur Lake, 15 July 1927,
L. F. Henderson 8655 (holotype, ORE, ISchuyler;
isotype, CAS).

Scirpus rubiginosus A. A. Beetle, Amer. J. Bot. 28: 697.
1941. Syn. nov. Schoenoplectus rubiginosus (A.
A. Beetle) Sojak, Cas. Nar. Mus., Odd. Prir. 140:
127. 1972. TYPE: U.S.A. California: Marin Co.,
along tide flat N of Stinson Beach, 9 May 1941,
Beetle <£ Constance 2686 (lectotype [imm.], UC,
selected here, with specimen of Sch. acutus x cal-
ifornicus on sheet designated by Beetle as holotype;
isolectotypes [imm.], CAS, GH, NY—on sheets with
specimen of Sch. acutus x californicus). [Other
sheets from the type collection at CAS, F, KSC,
RSA, and US are Sch. acutus x californicus .]

Chase (1904: 68)included all North American Scir¬
pus acutus in Scirpus occidentalis (S. Watson) Chase.

I have seen nine specimens that Watson annotated
as Scirpus lacustris var. occidentalis. The two not
cited by Chase (1904: 69) are: (1) the lectotype of
Schoenoplectus acutus var. occidentalis, and (2)
U.S.A. California: Sacramento Valley, Wilkes Ex¬
pedition [1842], (no collector given) #1788 (NY).
Of all, only the Wright and Wilkes Expedition spec¬
imens bear achenes. I selected the Wright 1940
specimen as the lectotype because it is typical of Sch.
acutus var. occidentalis throughout its range in its
lenticular achenes and mostly bifid styles, whereas
the mostly trigonous achenes and mostly trifid styles
of the Wilkes Expedition specimen are known only
from low elevations in California.

The following collections from the type locality
closely match the type collection of Scirpus rubi¬
ginosus: U.S.A. California: Marin Co., S end of
Bolinas Lagoon ca. V 2 mi. N of village of Stinson
Beach, fresh water seepages at edge of tidal salt
marsh. Schoenoplectus acutus var. occidentalis
(achenes present): 20 Aug. 1969, S. G. Smith 4713
(MO, PH, UC, US), 24 July 1973, S. G. Smith
5557 (GH, NY). Schoenoplectus acutus x cali¬
fornicus (achenes lacking): 20 Aug. 1969, S. G.
Smith 4714 (PH, UC, US, UW), 24 July 1973, S.
G. Smith 5556 (GH, MO, NY).

For the lectotype of Scirpus rubiginosus I se¬
lected the specimen of Schoenoplectus acutus rath¬
er than the hybrid because: (1) Beetle’s protologue
clearly indicates that his name applies to plants
correctly placed in Sch. acutus rather than in the
hybrid; and (2) Beetle described the achenes, which
are lacking in the type specimens collected in May
but are present in my collections of Sch. acutus ,
cited above, collected in July and August.

2. Schoenoplectus tabernaemontani (C. C.

Gmelin) Palla, Verh. K. K. Zool.-Bot. Ges. Wien
38, Sitzb.: 49. 1888. Basionym: Scirpus ta¬
bernaemontani C. C. Gmelin, FI. Bad. 1: 101.
April or May 1805. Scirpus lacustris L. subsp.
tabernaemontani (C. C. Gmelin) Syme, Engl.
Bot. ed. 10, 3: 63. 1870. Scirpus lacustris
var. tabernaemontani (C. C. Gmelin) Doell,
Rhein. FI. 165. 1943. Schoenoplectus lacus¬
tris subsp. tabernaemontani (C. C. Gmelin)
Love & Love, Folia Geobot. Phytotax. 10: 275.
1975. TYPE: Europe: in Rheni vivinia (holo¬
type, KR, Iphoto, det. on sheet by Schuyler in
1974).

Scirpus validus Vahl, Enum. PI. 2: 268. Oct. 1805 or
later. Scirpus lacustris var. validus (Vahl) Kuken-
thal, Feddes Repert. 23: 200. 1926 or 1927.
Schoenoplectus validus (Vahl) A. Love & D. Love,
Bull. Torrey Bot. Club 81: 33. 1954. Scirpus la¬
custris subsp. validus (Vahl) T. Koyama, Canad. J.
Bot. 40: 927. 1962. Schoenoplectus lacustris (L.)
Palla subsp. validus (Vahl) T. Koyama, Occas. Pap.
Pauahi Bishop Mus. 29: 128. 1989. TYPE: e Car-
ibaeis ded. Dr. Banks, Herb. Vahl s.n. (holotype,
C, Iphoto, ISchuyler).

Scirpus glaucus J. E. Smith, Engl. Bot. 33, pi. 2321.
1811. nom. illeg., not Scirpus glaucus Lamarck,
Tabl. Encycl. 1: 142. 1791. Scirpus lacustris subsp.
glaucus (J. E. Smith) C. J. Hartman, Sv. Norsk
Exc.-Fl.: 10. 1846. Schoenoplectus lacustris (L.)
Palla subsp. glaucus (J. E. Smith) Becherer, Feddes
Repert. 25: 11. 1928. TYPE: Scotland. Galloway,
on the sides of a saltmarsh on the W of Ardbigland,
1800, J. Mackay 6 (lectotype, LINN—Herb. Smith
105.25, selected here, Iphoto, det. by Clive Jermy
(BM)).

Scirpus tabernaemontani forma luxurious Miquel, Ann.
Mus. Bot. Lugd.-Batav. 2: 143. 1865. Scirpus la¬
custris subsp. validus var. luxurious (Miquel) T.
Koyama, Canad. J. Bot. 40: 927. 1962. Scirpus
lacustris subsp. creber var. luxurians (Miquel) T.
Koyamya, Phytologia 8: 458. 1963. Schoenoplec¬
tus validus subsp. luxurians (Miquel) Sojak, Cas.
Nar. Mus., Odd. Pfir. 141: 62. 1972. TYPE: Japan.
In ripa fluminis Jahaki, Gawa ins Nippon, #633
(lectotype, L, selected here—not seen, ISchuyler).
Scirpus validus var. creber Fernald, Rhodora 45: 283.
1943. Schoenoplectus validus (M. Vahl) A. Love
& D. Love var. creber (Fernald) A. Love & D. Love,
Bull. Torrey Bot. Club 81: 33. 1954. Scirpus val¬
idus subsp. creber (Fernald) T. Koyama, Phytologia
8: 458. 1963. Schoenoplectus lacustris subsp. cre¬
ber (Fernald) A. Love & D. Love, Taxon 30: 849.
1981. TYPE: U.S.A. New York: Fisher’s Island,
10-15 Aug., H. St. John 2581 (holotype, GH).

The typification of Scirpus glaucus J. E. Smith
is important because it is the basionym for the above
widely used names Scirpus lacustris subsp. glaucus
and Schoenoplectus lacustris subsp. glaucus. Nich-

Volume 5, Number 1
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Smith

North American Cyperaceae

99

olas Turland (BM, pers. comm. 1994) provided the
following information: Neither the collection by Bor-
rer from Sussex, England, used to produce the il¬
lustration of Scirpus glaucus in English Botany,
nor any material from Norfolk, England, as men¬
tioned in the protologue to plate 2321, appears to
exist at BM. The only other specimen mentioned in
the protologue is Mackay 6, which was examined
for me by Clive Jermy (BM); it has bifid styles and
dark brown papillae on the floral scales and is a
typical example of northwestern European S. ta-
bernaemontani.

My studies and those of A. E. Schuyler (pers.
comm.) indicate that Schoenoplectus tabernaemon-
tani and Sch. validus are definitely conspecific.
They differ by very few characteristics and inter¬
grade completely in both North America and Eur¬
asia. When Raymond (1957: 140) united these two
species he expressed doubts: “1 have tentatively
referred [a single specimen from Indo-China] to S.
tabernaemontani. . . . S. tabernaemontani and S.
validus are very closely related if not conspecific.”
Koyama (1962) combined European S. tabernae¬
montani with the North American S. acutus Muhl¬
enberg ex Bigelow in S. lacustris L. subsp. glaucus
(J. E. Smith) C. J. Hartman, placed Western Hemi¬
sphere and Hawaiian plants in S. lacustris L. subsp.
validus (M. Vahl) T. Koyama var. validus, and
placed Asian S. tabernaemontani in S. lacustris
subsp. validus var. luxurious (Miquel) T. Koyama.
Later, however, Koyama (1978: 209) treated S.
tabernaemontani and S. validus as separate but
doubtfully distinct species each occurring in both
North America and Eurasia. The known distribution
of Sch. tabernaemontani as here treated is the
boreal to tropical zones of most of Eurasia and North
America, the Caribbean, parts of Africa and tem¬
perate South America, the Hawaiian and other Pa¬
cific Islands, Australia, and New Zealand. Variation
in Sch. tabernaemontani sensu lato is complex, and
my attempts to clearly delimit infraspecific taxa have
led me to the opinion that delimitation of infraspecific
taxa is premature. Northwestern European plants
have densely to sparsely reddish scabrous floral scales,
especially near the bases of the spikelets; I have
seen very similar plants from scattered localities in
northern Canada, along both Atlantic and Pacific
coasts of North America, and southern Africa. Scir¬
pus validus var. creber is distinguished only by its
short floral scales, which partially expose the achenes,
and intergrades with forms with longer scales over
most of North America, and I have seen similar
forms from northwestern Europe.

3. Schoenoplectus x steinmetzii (Fernald) S. G.
Smith (pro sp.), comb. nov. Basionym: Scirpus
steinmetzii Fernald, Rhodora 45: 286 and plate
766. 1943. TYPE: U.S.A. Maine: Penobscott
County, bank of Passadumkeag Stream [ca. 2
mi. E of the village of], Passadumkeag, 12 Aug.
1937, F. H. Steinmetz 355 (holotype, GH;
isotypes, MAINE, NY, US).

I consider Schoenoplectus x steinmetzii to be a
hybrid between Schoenoplectus heterochaetus
(Chase) Sojak and Schoenoplectus tabernaemon¬
tani (C. C. Gmelin) Palla.

Koyama (1962: 927) placed Scirpus steinmetzii
in synonymy under S. lacustris subsp. validus (Vahl)
T. Koyama var. validus and treated S. hetero¬
chaetus Chase as S. lacustris subsp. lacustris var.
tenuiculmis (Sheldon) T. Koyama.

Scirpus steinmetzii has previously been reported
only from Passadumkeag Stream in the immediate
vicinity of the type locality, where the plants in the
several known colonies are very similar to each other
and are intermediate between Schoenoplectus het¬
erochaetus and Sch. tabernaemontani in leaf, culm
air cavity, floral scale, style, achene, and perigonial
bristle characters. The protologue of S. steinmetzii
erroneously describes the styles as bifid, the peri-
gonial bristles as 0-2, and the achenes as plano¬
convex. The plants at the type locality produce very
few achenes, in which the seeds are often aborted,
and most of the pollen grains are abnormal when
stained by cotton blue in lacto-phenol (confirmed by
Arthur Haines, MAINE, pers. comm. 1994). Al¬
though Sch. tabernaemontani is common in the
immediate vicinity of the type locality, the closest
known stations of Sch. heterochaetus are each ca.
360 km distant (Lake Champlain and vicinity, where
it is common and hybridizes with Sch. acutus, and
eastern Massachusetts, where it may be extirpated).
The region of the type locality, however, includes
vast roadless areas with numerous wetlands that
have been little explored botanically.

Other specimens from Ontario, Quebec, Iowa,
Massachusetts, New York, Minnesota, and Wiscon¬
sin are similar to specimens from the type popula¬
tion, but identification of most is uncertain because
they lack achenes. The following collection from a
population I have studied is nearly indistinguishable
from specimens from the type population except
that it produces many achenes containing fully
formed seeds: U.S.A. Wisconsin: Rock Co., ca. 1.5
mi. SW of Lima Center along Wis. Hwy. 59, T4N,
R14E, section 29, many clones in ca. 1 m of water.

100

Novon

in undrained depression with Schoenoplectus het-
erochaetus and Sch. tabernaemontani, 26 Aug.
1993, S. G. Smith 5208 ( GH, MAINE, NU, WIS).

4. Schoenoplectus pungens (M. Vahl) Palla, Bot.
Jahrb. Verh. K. K. Zool.-Bot. Ges. Wien 38,
Sitzb.: 49. 1888. Basionym: Scirpus pungens
M. Vahl, Enum. PI. 2: 255. 1805. TYPE:
Europe?: Erhart misit, herb. Vahl s.n. (holo-
type, C, Iphoto, ISchuyler).

Schoenoplectus pungens was long known by the
name Scirpus americanus Persoon, misapplied
(Schoenoplectus americanus (Persoon) Volkart ex
Schinz & R. Keller). The type of Scirpus ameri¬
canus Persoon is conspecific with plants long known
by the later synonym Scirpus olneyi A. Gray but
correctly known as Sch. americanus (Schuyler,
1974).

I agree with Koyama (1963) that North American
and European Schoenoplectus pungens can be di¬
vided into three varieties, although the character¬
istics of these are somewhat different from those he
gave, and in my opinion the differences are too small
for the recognition of subspecies.

4a. Schoenoplectus pungens var. pungens.

Synonymy and distribution as given by Koyama
(1963: 1117) for Scirpus americanus Persoon subsp.
americanus var. americanus.

4b. Schoenoplectus pungens var. longispi-
catus (N. L. Britton) S. G. Smith, comb. nov.
Basionym: Scirpus americanus Persoon var.
longispicatus N. L. Britton, Trans. New York
Acad. Sci. 11: 78. 1892. Scirpus longispi¬
catus (N. L. Britton) Smyth, Trans. Kansas
Acad. Sci. 16: 163. 1899. Schoenoplectus
americanus subsp. longispicatus (N. L. Brit¬
ton) Sojak, Cas. Nar. Mus., Odd. Prir. 141:
62. 1972. Scirpus pungens var. longispicatus
(N. L. Britton) Cronquist, Intermountain Flora
6: 74. 1977. Schoenoplectus pungens subsp.
longispicatus (N. L. Britton) A. Love & D.
Love, Taxon 30: 849. 1981. TYPE: U.S.A.
New Mexico: 1851 or 1852, C. Wright 1963
(lectotype, NY, selected here, ISchuyler; iso-
lectotypes, PH, US, GH 2 sheets).

Scirpus solispicatus Lunell, Amer. Midi. Naturalist 4:
230. 1915. TYPE: U.S.A. North Dakota: Benson
Co., Butte, 18 Aug. 1907, J. Lunell s.n. (lectotype,
MIN, selected here, ISchuyler).

The lectotype sheet of Schoenoplectus pungens
var. longispicatus , and the isolectotype sheets at
US and GH, all bear mixed collections of Schoen¬
oplectus pungens and putative Sch. americanus x
pungens hybrids; and sheet #28000 at GH also
bears a specimen of Sch. americanus (which a note
handwritten on the sheet suggests belongs to a col¬
lection from Nevada distributed by Wheeler in 1871).
I selected the specimens of Sch. pungens as the
lectotypes because they are typical of most western
North American Sch. pungens and bear mature
achenes, which are described in the original de¬
scription, whereas the putative hybrid specimens are
intermediate between Sch. americanus and Sch.
pungens and lack achenes. The other collection
(Colorado: Platte R., Dr. Geo. Smith) that Britton
cited in the protologue has not been located.

Three other specimens of Scirpus solispicatus
collected by Lunell at Butte in 1906 (GH, MIN,
NY) are very similar to the lectotype of Scirpus
solispicatus , which I chose because of its mature
achenes with intact bristles.

Schoenoplectus pungens var. longispicatus is
composed of at least the North American plants
included by Koyama (1963: 1118) in Scirpus amer¬
icanus Persoon subsp. americanus var. longispi¬
catus (N. L. Britton) T. Koyama.

4c. Schoenoplectus pungens (M. Vahl) Palla
var. badius (K. Presl) S. G. Smith, comb, et
stat. nov. Basionym: Scirpus badius K. Presl,
Reliq. Haenk. 1: 193. 1828. TYPE: U.S.A.
California: hab. ad Monte-Rey, Haenke s.n.
(holotype, PR, ISchuyler).

Schoenoplectus pungens var. badius is com¬
posed of at least the North American plants included
in Scirpus americanus subsp. monophyllus (K.
Presl) T. Koyama var. monophyllus (Koyama, 1963:
1118).

My examination of the types and original de¬
scriptions indicate that Scirpus badius K. Presl
rather than Scirpus monophyllus K. Presl is the
correct basionym for a taxon including the North
American Pacific Coastal plants of Schoenoplectus
pungens. Koyama (1963) placed Scirpus badius
in synonymy under Sch. americanus subsp. mon¬
ophyllus var. monophyllus , based on Scirpus mon¬
ophyllus K. Presl, Reliq. Haenk. 1: 193. 1828,
TYPE: Peru: Montana Peruvia, Haenke s.n. (ho¬
lotype, PR, ISchuyler). All three specimens on the
type sheet of S. monophyllus (photo in Koyama,
1963) are typical of Sch. americanus rather than

Volume 5, Number 1
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Smith

North American Cyperaceae

101

the Pacific Coastal North American Sch. pungens
in their leaves, floral scales, achenes, and perigonial
bristles. The styles are a mixture of bifid and trifid,
which can occur in either species. The culms, how¬
ever, have more-or-less flat sides and are very firm,
retaining their trigonous shape on pressing, which
is fairly common in Sch. pungens but rare in Sch.
americanus, in which the culm sides are typically
deeply concave and readily flattened on pressing.
Except for the style number, the pertinent parts of
the original description of Scirpus monophyllus,
“Colmo triquetro inferne monophyllo, . . . stylo bi-
fido, . . . caryopsidem obovatum plano-convex-
am . . .” agree with the type. Thus, the name Scir¬
pus monophyllus K. Presl applies to plants belong¬
ing to Sckoenoplectus americanus rather than to
Sch. pungens.

In the case of Scirpus badius, the type is a fine
specimen with a rhizome and four complete shoots
bearing inflorescences with some intact styles and
mature achenes. It is an excellent example of the
North American Pacific Coastal plants of Schoen-
oplectus pungens. The original description of S.
badius, which reads in part: “Colmo triquetro fol-
iato, . . . stylo trifido, . . . Setae sex retrorsum sca-
brae, caryopsidae una quarta parte breviores. Car-
yopsis obovata trigona. . . ,” agrees well with the
type.

5. Schoenoplectus x contortus (E. H. Eames)

S. G. Smith, comb. nov. Basionym: Scirpus
olneyi A. Gray var. contortus E. H. Eames,
Rhodora 9: 220. 1907. Scirpus x contortus
(E. H. Eames) T. Koyama, Canad. J. Bot. 41:
1123. 1963. TYPE: U.S.A. Connecticut: Mil¬
ford, plentiful in a brackish marsh at Pond
Point, 28 Sep. 1907, E. H. Eames 5847 (ho-
lotype, GH, ex herb. Eames; isotype, MO).

6. Schoenoplectus hallii (A. Gray) S. G. Smith,

comb. nov. Basionym: Scirpus hallii A. Gray,
Man. ed. 3, addend.: xcvii. 1862. Scirpus su-
pinus L. var. hallii (A. Gray) A. Gray, Man.
ed. 5: 563. 1867. Scirpus uninodis (Defile)
Boissier var. hallii A. A. Beetle, Amer. J. Bot.
29: 656. 1942. TYPE: U.S.A. Illinois: Mason
Co. or Menard Co., wet banks of pond, [Aug.
1860 or 1861], E. Hall [& D. Vasey & M.
S. Bebb?] s.n. (holotype, GH).

Raynal (1976: 141) placed Schoenoplectus hal¬
lii in synonymy under Sch. erectus (Poiret) Palla
ex J. Raynal.

The type locality, date, names of collectors, and
number of collections are all in doubt, as indicated
by differences among the protologue, Gray’s an¬

notation on the holotype, labels on the holotype and
several possible isotypes (F, NY, US), and a letter
from Gray [to Torrey?] at NY.

7. Bolboschoenus glaucus (Lamarck) S. G. Smith,

comb. nov. Basionym: Scirpus glaucus La¬
marck, Tabl. Encycl. 1: 142. 1791. Scirpus
maritimus L. [var.] glaucus (Lamarck) Nees
in Wight, Contr. Bot. India 111. 1834. TYPE:
Senegal, herb. Lamarck sheet 673/14, Rous¬
sillon s.n. (holotype, P, Iphoto, '.Schuyler; det.
as Scirpus maritimus L. by J. Raynal in 1966).

Examination of types and protologues indicates
that Scirpus glaucus Lamarck is the correct name
for the species to which Beetle (1942: 85) and
Koyama (1962: 936) misapplied the later name
Scirpus tuberosus Desfontaines, FI. Atlant. 1: 50.
1798; synonym: Scirpus maritimus L. [var.] tuber¬
osus (Desfontaines) Roemer & Schultes, Syst. Veg.
2: 139. 1817; Reigera maritimus [var.] tuberosus
(Desfontaines) Opiz, Seznam: 83. 1852; Bolbo¬
schoenus tuberosus (Desfontaines) Hadac, Bull. Coll.
Sci. Baghdad 6: 13. 1961. The type of S. tuberosus,
[Algeria]: habitat in lacubus prope La Calle, Herbier
de la Flore Atlantique donne au Museum, par M.
Desfontaines, s.n. (holotype, P, Iphoto; det. as Scir¬
pus maritimus L. by A. E. Schuyler in 1966), is
probably S. maritimus or a close relative in having
most of the spikelets in a sessile mass, bifid and trifid
styles, and lenticular achenes. The protologue, which
reads in part: “spicufis congestis. . .sessifibus aut
brevio pedunculatis,” agrees with the type. How¬
ever, the type of 5. glaucus is typical of the entity
sometimes known as S. tuberosus in its achenes and
trifid styles (Schuyler, pers. comm.) and its panic¬
ulate inflorescence with most of the spikelets on rays.
The protologue of S. glaucus, which reads in part
“umbella composita subpaniculata, spicufis pedicel-
latis,” agrees with the type.

Bolboschoenus glaucus is widely confused with
Scirpus maritimus L. (Schoenoplectus maritimus
(L.) Lye, Bolboschoenus maritimus (L.) Palla). Its
achenes were described by Browning & Gordon-
Gray (1993) under “Bolboschoenus maritimus en¬
tity 2.” It is widespread in the warmer parts of the
Old World and sparingly introduced into North
America.

8. Isolepis molesta (M. C. Johnston) S. G. Smith,

comb. nov. Basionym: Scirpus molestus M. C.
Johnston, Southw. Naturalist 9: 310. 1964.
TYPE: U.S.A. Texas: Walker Co., Huntsville,
27 Apr. 1944,5. R. Warner 20,000 (holotype,
TEX not seen, ISchuyler).

102

Novon

9. Trichophorum dementis (M. E. Jones) S. G.

Smith, comb. nov. Basionym: Scirpus demen¬
tis M. E. Jones, Contr. W. Bot. 14: 21. 1912.
TYPE: U.S.A. California: Rae Lake, King’s
River, alpine, 20 July 1910, Mrs. Clements
s.n. (lectotype, selected here, POM 111694,
ISchuyler; isolectotypes, DS, POM 111695).

Scirpus yosemitanus F. J. Smiley, Univ. Calif. Publ. Bot.
9: 108. 1921. TYPE: U.S.A. California: Yosemite
National Park, Tuolumne Meadows, soda springs, 8
Aug. 1898, /. 11. Congdon s.n. (holotype, GH).

10. Trichophorum clintonii (A. Gray) S. G.
Smith, comb. nov. Basionym: Scirpus clintonii
A. Gray, Amer. J. Sci., ser. 2, 38: 290. 1864.
TYPE: U.S.A. New York: between Buffalo and
Williamsville, June 1864, G. W. Clinton s.n.
(holotype, GH, ISchuyler; isotypes, NY,
ISchuyler; US, not seen; K not seen, ISchuyler).
Two possible isotypes at F bear insufficient data.

Acknowledgments. Alfred E. Schuyler (PH) pro¬
vided much essential information and counsel over
many years. Fred R. Barrie (MO), Jane Browning
(NU), K. D. Gordon-Gray (NU), Hugh H. litis (WIS),
Dan H. Nicolson (US), Anton A. Reznicek (MICH),
Mark T. Strong (US), and many other colleagues
generously gave help and advice. Nicholas Turland
and Clive Jermy (BM), Alicia Lourteig (P), G. Thijsse
(L), and Pamela White (GH) kindly provided infor¬
mation on types. Curators of the following herbaria
made specimens available for study, and many of
them were very helpful during my visits to their
herbaria: ALA, ALU, ARIZ, ASC, BHO, CAN, CAS,
CLEMS, DAV, F, FLAS, FTU, GA, GH, ISC, JEPS,
K, KANU, KSC, LAF, LL, LSU, MAINE, MICH,
MIN, MO, NEB, NFLD, NLU, NMW, NO, NY,
OS, OSC, PH, PR, RM, RSA, SD, TAES, TEX,
TRT, UC, UCSB, UNA, UNM, USCH, US, USF,

UTC, WIS, WS, WTU. My work was greatly aided
by a Smithsonian Institution research fellowship, by
research grants from the Flora of North America
Project, the Jepson Herbarium, and the University
of Wisconsin, and by the facilities generously pro¬
vided since my retirement by the Department of
Biology, University of Wisconsin-Whitewater.

Literature Cited

Beetle, A. A. 1942. Studies in the genus Scirpus L.
IV. The section Bolboschoenus Palla. Amer. J. Bot.
42: 82-88.

Browning, J. & K. D. Gordon-Gray. 1993. Studies in
the Cyperaceae in southern Africa. 21: The taxo¬
nomic significance of the achene and its embryo in
Bolboschoenus. S. African J. Bot. 59: 311-318.
Chase, A. 1904. The North American allies of Scirpus
lacustris. Rhodora 6: 65-71.

Koyama, T. 1962. The genus Scirpus Linn. Some
North American aphylloid species. Canad. J. Bot. 40:
913-937.

-. 1963. The genus Scirpus Linn. Critical spe¬
cies in the section Pterolepis. Canad. J. Bot. 41:
1107-1131.

-. 1978. Cyperaceae. In: Hui-Lin Liu et al.

(editors), Flora of Taiwan. Vol. 5: 191-372. Epoch
Publishing, Taipei, Taiwan.

Morin, N. R., Convening Editor, and Flora of North
America Editorial Committee. [In prep.] Flora of
North America. Vol. 11. Oxford Univ. Press, New
York.

Raymond, M. 1957. Some new or critical Scirpus from
Indo-China. Le Naturaliste Canadien 84: 111 149.
Raynal, J. 1976. Notes Cyperologiques: 26. Le Genre
Schoenoplectus II. L’amphicarpie et la sect. Supini.
Adansonia, ser. 2, 16: 119-155.

Schuyler, A. E. 1974. Typification and application of
the names Scirpus americanus Pers., S. olneyi Gray,
and S. pungens Vahl. Rhodora 76 (805): 51-52.
Smith, S. G. 1969. Natural hybridization in the Scirpus
lacustris complex in North Central United States.
Pp. 175-199 in J. E. Gunckel (editor), Current
Topics in Plant Science. Academic Press, New York
and London.

A New Nerine Species (Amaryllidaceae tribe Amaryllideae) from the
Koup Karoo, South Africa

D. A. Snijman

Compton Herbarium, National Botanical Institute, Private Bag X7,

Claremont 7735, South Africa

Abstract. Nerine marincowitzii is a rare new
species from the Koup Karoo, Cape Province, South
Africa. Most closely related to the eastern Cape N.
huttoniae Schonland, N. marincowitzii is distin¬
guished by hysteranthous leaves, a sigmoid scape
that detaches at ground level when dry, flowers that
turn brown at senescence, and a wind-dispersed in-
fructescence. It grows on exposed seasonal washes
and flowers at the end of the autumn rainfall season
in May.

Nerine Herbert (1820), comprising approxi¬
mately 22 species, is a horticulturally important
southern African genus of Amaryllideae (Dahlgren
et al., 1985). Of the ten genera in this mainly sub-
Saharan African tribe, Nerine is most closely allied
to Brunsvigia Heister (1755), sharing with it apo-
morphic fasciculate stamens. Traub (1967) has sug¬
gested that the two genera should be united. Until
strong phylogenetic evidence for this becomes avail¬
able, however, Nerine is retained as a distinct genus.
Nerine is distinguished from Brunsvigia by some¬
what smaller flowers, mostly with undulate tepal
margins, and distintegrating, globose, membranous
capsules with bulging, exposed seeds (Dyer, 1976).

Traub (1967) recognized four sections in Nerine.
Section Laticomae Traub, the most widespread in
southern Africa, comprises species with short, stout
scapes whose length more or less approximates the
width of the flower cluster. According to Obermeyer
(1993) the section includes two species, N. laticoma
(Ker-Gawler) Durand & Schinz and N. huttoniae
Schonland. The newly described N. marincowitzii
(Fig. 1) adds a further species to section Laticomae.

Nerine marincowitzii was collected southeast of
Leeu-Gamka in the Koup Karoo, Cape Province,
widely disjunct from N. laticoma and N. huttoniae
(Fig. 2). Unlike the highly seasonal summer rainfall
region occupied by these two species, the semiarid
area occupied by N. marincowitzii has an extended
rainfall season, and although annual precipitation in
the area is low and highly variable it benefits from
summer and autumn rainfall (Zucchini & Adamson,
1984).

Nerine marincowitzii is distinguished from N.
laticoma and N. huttoniae by hysteranthous, nar¬
row leaves (3-4 mm wide), a scape that is elliptical
rather than compressed in transverse section, and
by plane-edged tepals. The new species is most sim¬
ilar to N. huttoniae and shares with it a many-
flowered inflorescence (up to 40 flowers) and rigid,
widely radiating pedicels, which bestow a hemi¬
spherical shape on the flower head. Nerine marin¬
cowitzii is unique in the genus in having an apo-
morphic curved scape and tepals that turn brown
and remain outspread with age. These specialized
floral changes also occur in the closely allied Hessea
Herbert and differ from the usual pattern of senes¬
cence in Amaryllideae, where the floral pigmentation
becomes accentuated and the tepals ultimately col¬
lapse (Snijman, 1994).

The wind-dispersed infructescence of Nerine
marincowitzii is a further specialization not before
known in Nerine. When dry, the scape detaches at
ground level, the curvature of the scape confers a
somewhat spherical configuration on the infructesc¬
ence, and the entire structure tumbles away in the
wind. This derived mechanism, known as anemo-
geochory and reported to occur in arid areas world¬
wide (Van der Pijl, 1982), is also well developed in
Brunsvigia, Boophone Herbert, and Cybistetes
Milne-Redhead & Schweickerdt in Amaryllideae
(Snijman & Williamson, 1994).

Nerine marincowitzii Snijman, sp. nov. TYPE:
South Africa. Cape Province: 3222 (Beaufort
West) Koup Karoo, farm Vredendal, SE of
Leeu-Gamka, in washes along Kleinwaterval-
rivier (CD), 19 May 1990, Snijman 1245 (ho-
lotype, NBG; isotypes, MO, PRE). Figure 1.

Inter species sectionis Laticomarum Traub inflores-
centia multiflora (floribus usque 40) et pedicillis rigidis
radiatim ad Nerine huttoniam Schonland accedens sed
foliis hysteranthis, scapo sigmoideo, floribus brunneis ae-
tate provecta et infructescentia anemodispersa ab ea re-
cedens.

Plants small, to 20 cm high. Bulb solitary, de¬
pressed-globose, 45-55 mm across, covered with

Novon 5: 103-105. 1995.

104

Novon

Figure 1. Nerine marincowitzii Snijman. —1. Inflorescence. —2. Mature leaves. —3. Mature bulb. —4. Flower
at antbesis. 5. Anther after dehiscence. —6, 7. Dorsifixed anthers before dehiscence. —8. Mature seeds in
disintegrating capsule. Scale bars: 1-3, 8 = 2 cm; 4 = 7 mm; 5-7 = 3 mm. Original painting by E. Ward-Hilhorst
from Vlok s.n.

Volume 5, Number 1
1995

Snijman

Nerine marincowitzii

105

Figure 2. Known distribution of Nerine sect. Laticom-
ae: N. laticoma (A), N. huttoniae (★), N. marincowitzii
(•).

thinly fibrous tunics, cream-colored and fleshy with¬
in, extended into a slender neck to 60 mm long.
Foliage leaves 5-6, absent at anthesis, distichous,
suberect to recurved, narrowly ligulate, tapering
gradually distally, canaliculate, somewhat succulent,
to 30 cm x 3-4 mm, bright green, glabrous. In¬
florescence 20-40-flowered, hemispherical, com¬
pact, 10-16 cm across; scape sigmoid, rigid, ellip¬
tical in cross section, 11-14 cm x 2.5-3.5 mm,
reddish brown, breaking at ground level when dry;
spathe valves 2, narrowly lanceolate, 35-55 x 3-
5 mm, papery, wine-red; bracteoles filiform, to 10
mm long; pedicels radiating widely, straight, terete,
40-75 mm long, shiny green, turning straw-colored
and rigid in fruit. Perigone strongly zygomorphic,
pink, initially with whitish throat, aging to brown,
scentless; tepals shortly connate proximally, oblong-
lanceolate, falcate and upturned at anthesis, 17-22
x 4-6.5 mm, plane; filaments slightly declinate,
filiform, 13-17 mm long, expanded and connate
proximally into a short tube to 0.25 mm long; an¬
thers dorsifixed, ca. 5 mm long and wine-red before
opening; pollen whitish. Ovary dark wine-red; ovules
1 or rarely 2 per locule, absent from innermost
flowers; style slender, declinate, ca. 20 mm long;
nectar present around style base; stigma shortly
trilobed. Capsule small, papery, loculicidally dehis¬
cent, rapidly outgrown by seeds. Seeds ellipsoidal,
succulent, 6-7 mm across, wine-red; testa with sto¬
mata.

Phenology. Flowering is limited to approximately
three weeks in May depending on the advent of rain
in autumn (probably until the end of April). The

leaves appear in spring when growth is rapid and
die back during a dry spell in summer.

Distribution and habitat. Nerine marincowitzii
is known from a single population about 40 kilo¬
meters southeast of Leeu-Gamka, near the Klein-
watervalrivier, a tributary of the Gamka River. The
population, comprising approximately 400 plants, is
confined to sparsely vegetated, seasonal washes with
a north aspect, where the bulbs are lodged in fight
clay soil, covered with sharp blue-gray slate chips.

Etymology. The specific epithet honors C. P.
Marincowitz of the farm Kleinsleutelfontein, Koup
Karoo, whose original knowledge and appreciation
of this remarkable species led to its identification as
new.

Paratype. SOUTH AFRICA. Cape: 3222 (Beaufort
West) Koup Karoo, farm Vredendal, SE of Leeu-Gamka
(CD), 10 May 1992, Vlok s.n. (NBG).

Acknowledgments. I thank Jan Vlok of Cape
Nature Conservation and Museums and Colin Pat-
erson-Jones for their assistance in the field, and
Adela Romanowski of PRE and artist Ellaphie Ward-
Hilhorst for the botanical illustration.

Literature Cited

Dahlgren, R. H. T., H. T. Clifford & P. F. Yeo. 1985.
The Families of the Monocotyledons. Springer-Ver-
lag, Berlin.

Dyer, R. A. 1976. Amaryllidaceae. Pp. 947-954 in

R. A. Dyer (editor). The Genera of Southern African
Flowering Plants 2. Department of Agricultural Tech¬
nical Services, Pretoria.

Heister, L. 1755. Beschreibung eines neuen Ges-
chlechts. Waysenhause, Braunschweig.

Herbert, W. 1820. Nerine rosea. Rose-coloured Nerine
in J. Sims (editor), Bot. Mag. 47: 2124. Sherwood,
Neely & Jones, London.

Obermeyer, A. A. 1993. Nerine Herb. P. 162 in T.
H. Arnold & B. C. De Wet (editors), Plants of Southern
Africa: Names and Distribution. Mem. Bot. Survey

S. Africa 62.

Snijman, D. A. 1994. Systematics of Hessea, Stru-
maria and Carpolyza (Amaryllideae: Amaryllida¬
ceae). Contrib. Bolus Herb. 16: 1-162.

- & G. Williamson. 1994. A taxonomic re-as-

sessment of Ammocharis herrei and Cybistetes lon-
gifolia (Amaryllideae: Amaryllidaceae). Bothalia 24:
127-132.

Traub, H. P. 1967. Review of the genus Nerine Herb.
PL Life 23: 3-32.

Van der Pijl, L. 1982. Principles of Dispersal in Higher
Plants. Springer-Verlag, Berlin.

Zucchini, W. & P. J. Adamson. 1984. The Occurrence
and Severity of Droughts in South Africa. Water
Research Commission Report No. 91/1/1984. De¬
partment of Civil Engineering, University of Stellen¬
bosch.

Two New Species of Combretaceae from Puntarenas, Costa Rica

Clive A. Stace

Department of Botany, University of Leicester, Leicester LEI 7RH, England

ABSTRACT. Two new species of Combretaceae are
described from Costa Rica. Combretum graciliflo-
rum is related to C. cacoucia Exell and is only the
second species of section Cacoucia (Aublet) Engler

6 Diels known in America. Buchenavia costari-
censis is the second species of the genus from Cen¬
tral America and is evidently not closely related to
the other (B. tetraphylla (Aublet) R. A. Howard).
Combretum assimile Eichler is also reported as new
to Costa Rica, from the same area as the above two
new species (Osa Peninsula, Puntarenas).

Among specimens collected in Costa Rica in 1988-
1991 and recently sent to me for naming by the
Missouri Botanical Garden are three taxa new to
Costa Rica, two of them new to science. The two
latter are the first new Central American Combre¬
taceae to be discovered for many years, indicating
the need for further detailed studies of the area in
question (Osa Peninsula, Puntarenas).

Combretum graciliflorum Stace, sp. nov. TYPE:
Costa Rica. Puntarenas: Canton de Osa/Bue-
nos Aires, W part of main ridge of Fila Retinto,
along and near trail from Palmar Norte to Jal¬
isco, 8°59.5'N, 83°28’W, 780-960 m, 9 Dec.
1988, Grayum & Herrera 9131 (holotype,
MO).

A speciebus consectionibus floribus extra dense pubes-
centibus minoribus angustioribusque differt. Pedicelli 3-

7 mm; hypanthium inferius (cum ovario) 3-4 mm; hy-
panthium superius anguste curvato-infundibuliforme 10-
17 mm longum (sepalis 2.5-3.5 mm longis inclusis), 3.5-
6 mm latum; petala usque ad 3 mm exserta, ca. 7x5
mm, late elliptica.

Liana. Minute stalked glands frequent on leaves
and inflorescences. Leaves opposite, 3.5-11.5 x
2.5-6 cm, ovate-oblong, abruptly and shortly acu¬
minate at apex, cordate to rounded-cordate at base,
with 5-6 pairs of strongly curved lateral veins,
densely reddish brown appressed-pubescent on veins,
otherwise sparsely pubescent. Petiole 3-6 mm,
densely reddish brown appressed-pubescent. Inflo¬
rescence a simple, terminal raceme 6-14.5 cm (not
fully elongated). Young stems, rachis, bracts, and
outside of flowers densely reddish brown appressed-
pubescent. Lowest bracts ca. 14 x 5 mm, narrowly

Novon 5: 106-108. 1995.

elliptic, decreasing in size acropetally. Flowers (from
base of pedicel to tip of petals, excluding stamens
and style) 18-28 mm. Pedicel 3-7 mm. Lower
hypanthium (including ovary) 3-4 mm. Upper hy¬
panthium (including sepals) 10-17 mm, narrowly
curved-infundibuliform, gradually widening from
base, 3.5-6 mm across at base of sepals. Sepals
2.5-3.5 x 2-3 mm, triangular, acute at apex.
Petals ca. 7 x 5 mm, broadly elliptic, pubescent,
subacute at apex, exceeding sepals by up to 3 mm.
Stamens exserted (but broken off). Style exserted at
least 10 mm when mature.

The label describes the plant as a coarse liana
swooping low in forest, with yellowish green petals.
However, the petals have dried very dark, and they
and the stamens may have been red, as in most
members of the same section; the collectors might
have been referring to the sepals.

I have seen only one sheet of this species, but it
is clearly a member of section Cacoucia (Aublet)
Engler & Diels (Exell, 1953). This section contains
one American species (C. cacoucia Exell) and prob¬
ably four species in West Africa (Fig. 1). Combretum
graciliflorum is readily separable from C. cacoucia
because of its much narrower flowers, and comes
closer to some of the West African species in flower
shape, notably C. mooreanum Exell, C. stenopterum
Exell, and some plants of C. bracteatum (M. A.
Lawson) Engler & Diels (especially plants formerly
segregated as C. gentilii De Wildeman). However,
it is distinct from all of these in details of flower
structure. The flowers of section Cacoucia are the
largest and most highly developed in the genus, with
a long curved upper hypanthium, attaining 4.5 cm
long in some plants of C. bracteatum (e.g., the type
specimen of C. bucciniflorum Exell). They are most¬
ly red or orange in color, have long-exserted stamens
and style, and are probably bird-pollinated. The flow¬
ers of the new C. graciliflorum are the smallest in
the section.

The fruits of the West African species of the
genus are 5-winged, the wings being wide and thin
in C. bracteatum (including C. gentilii and C. buc¬
ciniflorum) and C. platypterum (Welwitsch) Hutch¬
inson & Dalziel, and very narrow and corky in C.
stenopterum and C. mooreanum. In contrast, the

Volume 5, Number 1

1995

Stace

Combretaceae from Costa Rica

107

Figure 1. Distribution of Combretum sect. Cacoucia. African range covers four species; it coincides closely with
the original area of tropical rainforest. American range covers C. cacoucia and (•) C. graciliflorum sp. nov. Continents
are drawn to the same scale and correct relative latitudes.

fruits of C. cacoucia are larger (5-8 x 1.7-3 cm)
and unwinged, with five strong longitudinal ridges.
It is thus very desirable that fruiting material of C.
graciliflorum be discovered. If it is a riverine spe¬
cies, like the three last-mentioned, then it too prob¬
ably has ridged or very narrowly winged fruits.

Buchenavia costaricensis Stace, sp. nov. TYPE:
Costa Rica. Puntarenas; Reserva Forestal Golfo
Dulce Aguabuena, Sector Cuenca Oeste,
8°42'2'N, 83°31'30'W, 50-150 m, 28 Sep.
1991, Aguilar 502 (holotype, MO; isotypes,
CR, LTR).

Ab aliis speciebus petiolis longis (1.3-3.5 cm longi)
gracilibus (laminae saepe solum duplo longae), inflores-
centiae elongato-spicatae, fructibus glabris (apice suba-
cuto ad rotundato), hypanthiis superis extus glabris, foliis
maturis glabris, hypanthiis inferis rhachidibusque sparse
pubescentibus differt.

Tree to 35 m, with long, thin, straight buttresses.
Leaves spirally arranged, crowded on branchlet tips;
lamina 2.5-13 x 1.5-5 cm, obovate or narrowly
so, with obtuse or rounded apex, with acute decur¬
rent base, with pubescent prominent midrib and
lateral veins on lower surface at flowering, otherwise
glabrous; petiole 1.3-3.5 cm, slender, eglandular.
Inflorescence an elongated spike; peduncle and ra-
chis 6-12 cm, slender, sparsely pubescent at flow¬

ering. Flowers 4-5 mm from base of ovary to tip
of calyx, yellowish green; lower hypanthium 2.5-3
mm, slender, puberulous in lower ovary-bearing part,
glabrous in upper part; upper hypanthium 1.5-2 x
4-5 mm, glabrous on outside, pubescent on inside,
± entire at margin; stamens 10, exserted up to 1.5
mm; style exserted up to 1.5(-2) mm. Fruit 2-3
x 1.2-1.9 cm, ellipsoid or obovoid, subacute to
rounded at apex, yellowish green when ripe, gla¬
brous; endocarp irregularly but strongly longitudi¬
nally furrowed, slightly compressed.

The holotype is a tree 20 m x 40 cm DBH in
fruit.

Buchenavia costaricensis is recognizable on the
basis of its leaves with long thin petioles (many about
half as long as the lamina); glabrous fruits with
obtuse to rounded apex; glabrous upper hypanthia
and mature leaves; and leaves at flowering time,
inflorescence axis, and lower hypanthium only
sparsely pubescent. The leaf shape, texture, and
venation is also quite characteristic.

This is only the second species of Buchenavia to
be found outside mainland South America (Exell &
Stace, 1963). Buchenavia tetraphylla (Aublet) R.
A. Howard (B. capitata (Vahl) Eichler) occurs in
the West Indies (Cuba to Trinidad) and in Costa
Rica and Panama, as well as widely throughout
South America, where a further 20 species occur

108

Novon

(Fig. 2). Buchenavia tetraphylla is distinct from
B. costaricensis in its thicker shorter petiole (0.4-
1.5(-2) cm) and capitate inflorescence.

Paratypes. COSTA RICA. Puntarenas: Canton de
Osa, Playa Campanario o San Josesito, Sierpe, primary
forest, 8°39'39"N, 83°42'50‘ , W, 1-10 m, 5 Jan. 1991
(fl), Harmon 192 (CR, LTR, MO); Canton de Golfito
Jimenez, Alto de la Carbonera, camino a Cerro de Osa,
8°25'30"N, 83°19'00"W, 200-350 m, 18 Sep. 1990
(fr), Chacon 1046 (CR, LTR); primary forest, Parque
Nac. Manuel Antonio, 9°23'N, 84°09'W, 1-10 m, 11
Apr. 1990 (fr), Harmon 95 (CR, LTR); Canton de Golfito
La Palma, en pequenas areas de bosque alrededor del
pueblo, 8°38'30"N, 83°23'00"W, 50 m, 14 Sep. 1992
(fr), Aguilar 1302 (CR, LTR).

The following two specimens have been brought
to my attention by Barry Hammel. COSTA RICA.
Puntarenas: Parque Nac. Manuel Antonio, woods
near ANEP road, Punta Catedral, Puerto Escondido,
and Lotz Road, 9°04'N, 84°10'W, 1-100 m, 3 Jan.
1990 (in bud), Harmon 40 (CR not seen); Canton
de Osa, Rincon de Osa, entrada a Chocuaco,
8°43'52"N, 83°27'15"W, 100-200 m, 27 Aug.
1992 (fr), Zamora, Aguilar, Ramirez & Martin
1864 (CR not seen).

COMBRETUM ASSIM1LE ElCHLER

Combretum assimile is a member of section Com-
bretum, represented mainly by C. frulicosum (Loe-
fling) Stuntz and C. farinosum Kunth in Central
America (including Costa Rica). Combretum assi¬
mile differs from these two species in its larger
flowers and fruits and in its dark red scales on the
flowers, fruits, inflorescence axis, leaves, and young
stems (golden in C. fruticosum, gray in C. farinos¬
um). It is primarily a plant of the Amazon basin,
especially on the Peru-Brazil border. Combretum
sambuense Pittier was described from the old Pan¬
ama Canal Zone, and there are also records from
Colombia and Venezuela (Exell, 1953), but in my
opinion that species should be included in C. assi¬
mile. The following collection therefore extends the
known distribution of C. assimile westward into Cos¬
ta Rica: COSTA RICA. Puntarenas: Reserva For-
estal Golfo Dulce Aguabuena, Sector Sur, 8°42'20"N,

Figure 2. Distribution of Buchenavia. Continuous line:
B. tetraphylla. •: B. costaricensis. Broken line: 20 other
species.

83°31'30"W, 50-150 m, 26 Dec. 1991, Aguilar
& Herrera 793 (LTR, MO).

Combretum fruticosum occurs in the same area,
but the southern limit of C. farinosum is in northern
Costa Rica.

Acknowledgments. I am grateful to Michael Gra-
yum (MO) for information concerning the holotype
of Combretum graciliflorum, to Barry Hammel (MO)
for data on and specimens of Buchenavia costari¬
censis, and to Carel Jongkind (WAG) for valuable
data on the African species of section Cacoucia.

Literature Cited

Exell, A. W. 1953. The Combretum species of the New
World. J. Linn. Soc. (Bot.) 55: 103-141.

- & C. A. Stace. 1963. Revision of the genera

Buchenavia and Ramatuella. Bull. Brit. Mus. (Nat.
Hist.), Bot. 3: 3-46.

A New Combination in Merremia (Convolvulaceae) for the
Flora of China

George W. Staples

Department of Natural Sciences, B. P. Bishop Museum, Honolulu, Hawaii
96817-0916, U.S.A.

Fang Rhui-cheng

Kunming Institute of Botany, Academia Sinica, Heilongtan, Kunming, Yunnan,
People’s Republic of China

Abstract. On the basis of pollen morphology and
floral characters, Ipomoea caloxantha is trans¬
ferred to Merremia.

While revising the Convolvulaceae for the Flora
of China, the name Ipomoea caloxantha Diels came
to our attention. Fang & Huang (1979) listed /.
caloxantha at the end of their treatment for Ipom¬
oea with a Chinese translation of Diels’s original
description, but did not include it in their key to
species of the genus nor give it any further discus¬
sion.

Based on Diels’s description of the corolla as
“intense lutea” and “flowers bright yellow,” one of
us (GWS) suspected this to be a Merremia, in which
genus yellow corollas are fairly common, whereas
very few Ipomoea species have them. The holotype
of I. caloxantha was examined, and the suspicion
proved correct. A quick check of the pollen grains
under the dissecting microscope showed them to be
nonspinulose, more or less spherical, and apparently
12-rugate. This conforms to Type 3 pollen described
for Merremia in the terminology of Ferguson et al.
(1977). A new combination is therefore necessary.

Merremia caloxantha (Diels) Staples & R. C.
Fang, comb. nov. Basionym: Ipomoea calox¬
antha Diels, Notes Roy. Bot. Gard. Edinburgh
5: 203. 1912. TYPE: China. Yunnan (west¬
ern): valley of the Mekong, at the crossing of
the Teng-Yueh-Talifu road, lat. 25°18'N, 4200
ft., Sep. 1903, G. Forrest 1111 (holotype, E).

An amplified description of Merremia caloxantha
will be published in the Flora of China. We maintain
this species until further study can be devoted to
Merremia in Asia, where the species are inade¬
quately known, at best, and it is quite possible that
other species remain to be described, or may have
already been described under Ipomoea.

Acknowledgments. We thank the Keeper of the
Herbarium at the Royal Botanic Garden, Edinburgh,
for the loan of the holotype and Ihsan Al-Shehbaz
for expediting the publication of this brief note.

Literature Cited

Fang Rhui-cheng & Huang Shu-hua. 1979. Convol¬
vulaceae. In: Wu Cheng-yi (editor), FI. Reipubl. Po-
pularis Sin. 64(1): 1-153.

Ferguson, I. K., B. Verdcourt & M. M. Poole. 1977.
Pollen morphology in the genera Merremia and Op-
erculina (Convolvulaceae) and its taxonomic signif¬
icance. Kew Bull. 31: 763-773.

Novon 5: 109. 1995.

Miconia amplinodis (Melastomataceae: Miconieae), una Nueva
Especie para Costa Rica

Gina Umaha Dodero

Herbario Nacional de Costa Rica, Museo Nacional de Costa Rica, Apartado 749-1000,
San Jose, Costa Rica

Frank Almeda

Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco,
California 94118-4599, U.S.A.

RESUMEN. Se describe y se ilustra Miconia am¬
plinodis, una nueva especie de Costa Rica. Su ca-
racteristica mas sobresaliente es el poseer unas pro-
yecciones nodales tanto en tallos como en inflores-
cencia. Se comenta su relacion con otras especies
del section Cremanium, al cual se adscribe.

ABSTRACT. Miconia amplinodis, a new species
from Costa Rica, is described and illustrated. The
most outstanding feature of this species is its suc¬
culent auriculate projections at branch and inflo¬
rescence nodes. The relationships of M. amplinodis
to other species in section Cremanium are also dis¬
cussed.

Miconia, el genero mas grande de la familia Me¬
lastomataceae, se encuentra distribuido en la region
neotropical con aproximadamente 1000 especies.
Estas han sido agrupadas en 11 diferentes secciones
en base a la gran variedad presentada en la mor-
fologia del androceo.

En Costa Rica, este genero se halla representado
por unas 100 especies aproximadamente; numero
que se ha incrementado durante los ultimos aiios
debido a una mayor actividad de colecta en bosques
de mediana elevation, de donde se han descrito
varias especies nuevas (Almeda & Umana, 1993).

La especie que aqui se describe es conocida de
la Cordillera de Cuanacaste, Cordillera de Tilaran
y la Cordillera Volcanica Central en elevaciones de
1100 a 2000 m.s.n.m.

Miconia amplinodis Umana & Almeda, sp. nov.
TIPO: Costa Rica. Alajuela: Bajos del Toro
Amarillo, 10°12'10'N, 84°18'43"W, 1800 m,
15 jun. 1992, Umana et al. 495 (holotipo,
CR; isotipos, CAS, COL, F, MEXU, MO, US,
USJ). Figura 1.

Novon 5: 110-113. 1995.

Frutex 0.5-2 m altus. Rami ramulique sicut petioli
inflorescentiaque glabri, nodis tumidis et auriculis 2.5-

3.5 mm longis infra petiolorum insertiones armatis. Petioli
1.3-4.8 cm longi; lamina 8-11.7 x 3.3-5.7 cm elliptica,
apice breviter hebeti-acuminato basi acuta vel obtusa, 3-
nervata. Panicula 7-17 cm longa multiflora; bracteolis
1-2 x 0.75-1 mm deciduis. Hypanthium (ad torum) 2-

2.5 mm longum; calycis tubus 0.20-0.25 mm longus,
lobis usque ad 0.5 mm longis et 0.75 mm latis. Petala
1.5-2 x 1-1.5 mm. Stamina 10, isomorphica glabra;
filamenta 2 mm longa; antherarum thecae 1-1.4 x 0.25-
0.50 mm obovato-oblongae 4-porosae, connectivo paulo
(0.4 mm) prolongato. Stigma truncatum; stylus 3-3.5 x
0.2 mm glaber; ovarium % inferum, 3(?)-loculare, apice
truncato-conico pilis paucis glanduliferis.

Arbusto de 0.5-2 m de altura; entrenudos, ramas,
peciolos e inflorescencia glabros; nudos del tallo e
inflorescencia con unas proyecciones persistentes,
glabras, redondeadas y carnosas de 2.5-3.5 mm de
largo; hojas de un par iguales en tamano, elipticas,
algo coriaceas, margen aserrado, 8-11.7 cm de
largo y 3.3-5.7 cm de ancho, apice corto acumi-
nado, base anchamente aguda a obtusa; 5 nervios
(incluyendo los dos nervios laterales tenues); peciolos
1.3-4.8 cm de largo. Inflorescencia una panicula
terminal de muchas flores, 7-17 cm de largo; brac-
teas pares, ovadas, 2-6 mm de largo y 2.5 mm de
ancho, horde irregular, glabras, sesiles, caducas;
bracteolas pares, ovadas, 1-2 mm de largo y 0.75-
1 mm de ancho, borde irregular, glabras, sesiles,
caducas; pedicelos glabros, 0.25-0.5 mm de largo;
hipantio (en la floracion), campanulado, glabro, con
10 costillas, 2-2.5 mm de largo hasta el toro; tubo
del caliz 0.20-0.25 mm de largo; lobulos del caliz
glabros, anchamente redondeados, 0.5 mm de largo
y 0.75 mm de ancho; dientes externos persistentes,
sobrepasados en tamano por los lobulos del caliz;
petalos 5, glabros, concavos, suborbiculares, blanco-
verdosos, isomorficos, 1.5-2 mm de largo y 1-1-5
mm de ancho; filamentos glabros, distalmente ge-
niculados, 2 mm de largo; anteras 1-1.4 mm de

Volume 5, Number 1
1995

Umana & Almeda
Miconia amplinodis

111

Figura 1. Miconia amplinodis Umana & Almeda. —A. Habito. B. Superficie abaxial de la hoja. C. Detalle
de las proyecciones nodales. —D. Flor. —E. Petalo. —F. Estambres, vista ventral (izquierda), vista dorsolateral
(centro) y vista lateral (derecha). —G. Bractea. —H. Fruto. I. Semillas. (A - F de Umana et al. 495; H, I de
Umana et al. 465.)

112

Novon

largo, cuneadas, distalmente truncadas con 4 poros
terminales ventralmente inclinados; conectivo en-
grosado dorsalmente y prolongado bajo las tecas 0.4
mm. Ovario 3(?)-loculare, infero, estilo glabro, recto,
3-3.5 mm de largo; estigma truncado. Fruto una
baya globosa, 6 mm de diametro, morado cuando
inmaduro, bianco en la madurez. Semillas nume-
rosas, cafe-rojizas, ovoides, 1.3 mm de largo, la testa
papilosa.

Distribution y fenologia. Se conoce de la Cor¬
dillera de Guanacaste, Cordillera Volcanica Central
(Alajuela: Bajos del Toro Amarillo, Fila Volcan Viejo
cercanias del Volcan Poas; Heredia: Parque Nacio-
nal Braulio Carrillo), y de la Cordillera de Tilaran,
Reserva de Bosque Nuboso Monteverde, en la parte
noroeste de Costa Rica en elevaciones de 1100-
2000 m. Esta especie ha sido colectada en sitios
abiertos y soleados, en orillas de caminos y en po-
treros (Fig. 2). Los especimenes con flor han sido
colectados en los meses de abril, mayo, junio, julio
y septiembre; muestras con fruto en febrero, abril,
septiembre, noviembre y diciembre.

Miconia amplinodis fue colectada por primera
vez en Monteverde en 1979; esta muestra presen-

taba solamente frutos lo mismo que posteriores co-
lectas hechas en la misma localidad y en Fila Volcan
Viejo y Bajos del Toro. No fue hasta 1992 cuando
se logro realizar una colecta de material plenamente
florecido.

Miconia amplinodis se caracteriza por ser un
arbusto glabro, con hojas glabras de 5 nervios con
borde ligeramente aserrado de color morado, y con
flores de 5 partes y anteras cuneiformes de 4 loculos.
Su caracteristica mas sobresaliente, y unica dentro
de las especies de Miconia conocidas para el pais,
es el poseer unas proyecciones persistentes, glabras,
redondeadas y carnosas a nivel de nudos tanto en
tallos como en inflorescencias. Estas proyecciones
nodales engrosadas son raras en Miconia y en la
familia Melastomataceae en general, y hasta el mo¬
menta no se conoce su significancia funcional. Cu-
riosamente estas estructuras presentan un color di-
ferente dependiendo de si estan expuestas o no a la
luz solar; cuando es asi, el color varia entre rojo y
morado, cuando se hallan en sitios sombreados estas
presentan un color verde. Generalmente se presen¬
tan 2 por nudo con excepcion de una muestra que
presentaba 3 de estas estructuras por nudo.

Esta especie, asignable a la seccion Cremanium

Volume 5, Number 1
1995

Umana & Almeda
Miconia amplinodis

113

en virtud de sus anteras cuneiformes de 4 loculos,
se halla relacionada cercanamente a Miconia re-
sima Naudin, especie conocida solamente para Co¬
lombia. Las caracteristicas que ambas comparten
son el poseer flores de 5-partes y androceos muy
similares. A diferencia de Miconia amplinodis, M.
resima carece de las proyecciones nodales de la que
parten los peciolos y las ramificaciones de las inflo-
rescencias; posee hojas de mayor tamano (15-20
x 6-7 cm), y presenta un estigma subpeltado.

Entre las especies de Miconia, dentro de la sec-
cion Cremanium, que poseen proyecciones anula-
riformes conspicuas en cada nudo, M. amplinodis
es tambien similar a M. manicata Cogniaux & Glea¬
son de Colombia, en el tamano de las hojas, mor-
fologia del hipantio, y forma de los petalos. Miconia
manicata difiere de nuestra nueva especie en poseer
lamina foliar ciliada (los cilios de approx. 1 mm de
largo) y base largo cuneada, tecas de las anteras de
4 poros incompletos con el conectivo basalmente
trilobulado, ovario medio infero, y estigma capitado.

El nombre dado a esta especie hace alusion a las
proyecciones conspicuas que presente a nivel de
nudos en tallos e inflorescencias, caracteristicas que
la hace facil de reconocer.

Paratipos. COSTA RICA. Alajuela: Volcan Poas,
faldas del Cerro innominado ubicado al N del crater, 5
mayo 1992, Ballestero 148 (CR); San Carlos, Fila Volcan
Viejo, 1800-2000 m, 11-14 feb. 1986, Gomez-Laurito
11021 (CR); San Carlos, Fila Volcan Viejo, 1800-2000
m, 13 nov. 1986, Gomez-Laurito 11251 (CR); Bajos del
Toro Amarillo, 10°12'10'N, 84°18'43"W, 1800 m, 10
abr. 1989, Umana etal. 276 (CR), 28 abr. 1992, Umana
et al. 465 (CR), 14 mayo 1992, Umana et al. 484 y

488 (CR). Alajuela/Puntarenas/Guanacaste: Cor¬
dillera de Tilaran, Reserva Biologica Monteverde, 1540-
1600 m, 21 feb. 1986, Almeda et al. 4999 (CAS, CR);
Reserva Biologica Monteverde, 10°18'N, 84°46'W, 1100
m, 1 jul. 1989, Bello 996 (CAS, CR); Reserva Biologica
Monteverde, 1500-1600 m, Haber ex Bello 5190 (CAS,
CR, MO); Reserva Biologica Monteverde, 10°18'N,
84°47'W, 1500-1600 m, 16 jul. 1990, Haber & Zu
chowski 10014 (CAS, CR, MO); Reserva Biologica Mon¬
teverde, 10°18'N, 84°47' r W, 1500-1600 m, 31 die.
1990, Haber & Zuchowski 10378 (CAS, CR, MO); Re¬
serva Biologica Monteverde, 10°18'N, 84°47'W, 1520-
1600 m, 14 sep. 1991, Haber & Zuchowski 10872
(CAS, CR, MO); Reserva Biologica Monteverde, 10°18'N,
84°48'W, 1600 m, 17 jun. 1992, Haber et al. 11206
(CR); Reserva Biologica Monteverde, 3 jul. 1979, Lumer
1086 (CAS). Heredia: Parque Nacional Braulio Carrillo,
San Isidro, Cerro Zurqui, 10°03'30*’N, 84°01T5*’W, 1800
m, 16 jul. 1992, Ballestero et al. 177 (CR). Guana-
caste: Parque Nacional Rincon de la Vieja, Sector Las
Pailas en la cima del Volcan, 10°49'30*’N, 85°19 / 30"W,
1700 m, 9 enero 1994, Morales 2243 (CR).

Agradecimientos. Queremos hacer publico nues-
tro agradecimiento a Pablo Sanchez V. y a Jorge
Gomez-L. por su colaboracion en la descripcion la-
tina de la especie que aqui se describe; a Jorge
Gomez-L., Miguel Ballestero y William Haber por
el comentario sobre las colectas hechas de la especie
en los diferentes lugares; y a Rafael Chacon por su
colaboracion en las giras realizadas para la colecta
del material.

Literatura Citada

Almeda, F. & G. Umana D. 1993. New species of
Miconia (Melastomataceae) from Costa Rica. Novon
3: 5-10.

Justicia nevlingii (Acanthaceae), a New Species from Mexico

Dieter C. VTasshausen

Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Thomas F. Daniel

Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco,
California 94118, U.S.A.

ABSTRACT. A new species of Justicia with heter-
omorphic bracts, J. nevlingii , is described from
southern Mexico. The species is compared to two
other heteromorphically bracted species from Mex¬
ico and Central America, J. chol and J. costaricana.
It differs from these in several features, including
its 4-aperturate pollen.

In 1974, Larry Nevling of the Field Museum of
Natural History called to the attention of the senior
author an unusual collection of Acanthaceae from
Veracruz representing an unknown species. Other
collections resembling this species from Veracruz
and Chiapas have since been assembled and studied.
They are described below as a new species of Jus¬
ticia with heteromorphic bracts.

Justicia nevlingii Wasshausen & T. F. Daniel,
sp. nov. TYPE: Mexico. Veracruz: Hidalgoti-
tlan, 0-2 km del Plan de Arroyo-Alvaro Obre-
gon, 17°01'N, 94°40'W, 14 Apr. 1974, J.
Dorantes et al. D-2799 (holotype, US; isotype,
F). Figures 1, 2.

Herba perennis usque ad 1.5 m alta. Folia petiolata,
laminae ovatae, 6.5-18.5 cm longae, 3-10.4 cm latae.
Spicae terminales et axillares, densae, pedunculis et rhach-
idibus dense pilosis. Bracteae heteromorphae; bracteae
fertiles petiolatae, obovatae vel ellipticae, 5.5-12 mm
longae, 2.7-6.5 mm latae; bracteae steriles lineares-lan-
ceolatae vel lineares vel subulatae, 2-5 mm longae, 0.3-
0.5 mm latae. Calyx 5-lobus. Corolla alba vel flavida vel
cremea, 9-10 mm longa, extus pubescens trichomatibus
eglandulosis. Thecae antherarum 2, superpositae, glabrae,
theca inferiore basi calcarata; pollen 4-apertum. Capsula
7-8.5 mm longa, pilosa. Semina 2.

Erect, perennial herbs to 1.5 m tall. Stems
subquadrangular, quadrifariously pilose with tri-
chomes eglandular, appressed and spreading, 0.7-
2 mm long, multi-septate with maroon septae. Leaves
petiolate, petioles 1.5-7.2 cm long, densely pilose,
blades ovate to elliptic, 6.5-18.5 cm long, 3-10.4

Novon 5: 114-117. 1995.

cm wide, short-acuminate at apex, obtuse or rounded
to subattenuate at base, entire to sinuate, upper
surface glossy, sparingly pilose with trichomes ap¬
pressed, lower surface more densely pilose (especial¬
ly the central and lateral veins) with trichomes ap¬
pressed and spreading, to 1 mm long. Inflorescence
of (axillary and) terminal dense panicles of spikes
to 15 cm long and 1.5-4.4 cm diam., erect or
ascending, rachises densely pilose with trichomes
sordid, erect, to 1 mm long. Floral bracts hetero¬
morphic; fertile bracts petiolate, obovate to elliptic,
5.5-12 mm long, 2.7-6.5 mm wide, acute to trun-
cate-apiculate at apex, abaxial surface pubescent
with antrorse-appressed eglandular trichomes 0.1-
0.5 mm long and glandular trichomes to 0.2 mm
long, margin ciliate with cauline type trichomes;
sterile bracts linear-lanceolate to linear to subulate,
2-5 mm long, 0.3-0.5 mm wide. Bracteoles oblan-
ceolate, 4-9 mm long, 1.2-2.2 mm wide. Calyx
deeply 5-lobed, 3-5 mm long, lobes lanceolate, 2.5-
4 mm long, 0.5-0.6 mm wide, puberulous. Corolla
white to yellowish to cream, 9-10 mm long, exter¬
nally pubescent with eglandular trichomes 0.2-0.5
mm long; tube 5-5.5 mm long, 0.8 mm diam. at
base, 3 mm diam. at apex; limb 2-lipped, upper lip
erect, rugulate, ovate-lanceolate, 3-4.5 mm long,
2 mm wide, 2-lobed at apex, lobes ca. 0.25 mm
long and wide, rounded, lower lip ± spreading, 3-
lobed, 3.5-6 mm long, 3 mm wide at base of lobes,
lobes obovate, 1.2-1.5 mm long, 1.2-1.5 mm wide,
rounded at apex. Stamens inserted near apex of
corolla tube, 4 mm long, filaments glabrous, thecae
superposed, glabrous, upper theca ca. 1 mm long,
lower thecae ca. 0.8 mm long, calcarate with blunt
appendage ca. 0.3 mm long; pollen prolate 4-aper¬
turate, apertures flanked on each side by a ± con¬
tinuous band of exine and a pseudocolpus (Fig- 2);
2 staminode-like protrusions of corolla present near
base of filaments, 0.4-1.2 mm long, densely villous.
Style 6.5 -7 mm long, glabrous, stigma 0.2-0.3 mm
long. Capsules 7-8.5 mm long, pilose. Seeds 2,

Volume 5, Number 1
1995

Wasshausen & Daniel
Justicia nevlingii

115

ED C

Figure 1. Justicia ne vlingii Wasshausen & T. F. Daniel (Dorantes et al. D-2799). A. Habit. B. Bract.

C. Bracteoles. — D. Calyx lobes. — E. Corolla and stamens. —F. Expanded corolla, stamens, and staminode-like
protrusions.

116

Novon

Figure 2. Scanning electron (SEM) photomicrographs of pollen of Justicia nevlingii (Dorantes et al. D-2799). —
A. Equatorial view, x2800. —B. Portion of aperture, x7000.

spherical to discoid, flattened, 2.1-2.8 mm long, 2-
2.5 mm wide, smooth.

Distribution. Southern Mexico (Chiapas and Ve¬
racruz); plants occur in lowland and lower montane
rainforests at elevations from 130 to 470 m.

Paratypes. MEXICO. Veracruz: Mpio. Hidalgotitlan,
0-2 km road Plan de Arroyo-Arroyo Alegre, 17°15'N,
94°35'W, 23 Apr. 1974, Jesus Dorantes et al. D-2964
(F, US); Mpio. Hidalgotitlan, between Hnos. Cedillo-Au¬
gustin Melgar, 17°13'30' r N, 94°35'W, 26 Apr. 1974,
Mario Vazquez et al. 411 (F, US); Las Choapas, Las
Cruces, 14 July 1970, A. Gomez-Pompa & L. Nevling
1522 (F, MEXU); Mpio. Hidalgotitlan, La Laguna,
Uxpanapa, 29 Mar. 1974, A. Gomez-Pompa & L. Nevl¬
ing 5183 (F). Chiapas: Mpio. Ixtacomitan, above Ixta-
comitan, 10 May 1973, D. Breedlove 35058 (DS,
MEXU); St. Bartolo, Apr. 1840, J. Linden s.n. (G, P);
along Rio Pichucalco, 3 km NE of Pichucalco toward
Villahermosa, 17°32'N, 93°04'W, 2 July 1969, B. Marchs
& C. Marcks 913 (LL, WIS); 37 km NW of Bonampak,
6 km NNW of Nuevo Guerrero, G. Davidse et al. 20534
(BM, LL, MEXU); between Pichucalco and Rayon near
Solosuchiapa, cuadrante 57-A, J. Chavelas P. et al 442
(MEXU).

Justicia nevlingii resembles both J. chol T. F.
Daniel (Chiapas) and J. costaricana Leonard (Costa
Rica) by its heteromorphic bracts and staminode-

like protrusions of the corolla. It may be distin¬
guished from these species by the following key:

la. Bracts eciliate or inconspicuously ciliate with

trichomes to 0.2 mm long; calyx 5-7 mm long;
corolla externally glandular; cauline trichomes
lacking maroon septae; cloud forests at 900-
1550 m in Costa Rica. J- costaricana

lb. Bracts conspicuously ciliate with trichomes to
2 mm long; calyx 2.5-5 mm long; corolla ex¬
ternally eglandular; cauline trichomes with con¬
spicuous maroon septae; lowland to lower mon¬
tane rainforests at 50-470 m in Mexico.

2a. Young stems with trichomes concentrated
in 2 lines; inflorescence of axillary spikes;
bracteoles 0.4-1 mm wide; style pubes¬
cent; seeds 4, 1.3-1.6 mm long, covered
with low rounded papillae or ridges; pollen

3-aperturate. J- c h°l

2b. Young stems with trichomes restricted to
4 lines; inflorescence of (axillary to) ter¬
minal panicles of spikes; bracteoles 1.2-
2.2 mm wide; style glabrous; seeds 2, 2.1 -
2.8 mm long, smooth; pollen 4-
aperturate. J. nevlingii

Graham (1988) stated that the pollen of Justicia
is either 2- or 3-aperturate but that 4-aperturate
pollen (not confirmed by her study) had been re¬
ported in a few New World species. Daniel (1990,

Volume 5, Number 1
1995

Wasshausen & Daniel
Justicia nevlingii

117

1993) documented 4-aperturate pollen in several
species of Justicia (i.e., J. angustijiora D. N. Gib¬
son, J. tabascina T. F. Daniel, and J. valvata T.
F. Daniel) and has since observed it in several others
(e.g., J multicaulis Donnell Smith, J. silvicola D.
N. Gibson, and /. warnockii B. L. Turner).

Graham (1988) also noted that all the species of
Justicia that she had observed have an androecium
of two stamens and no staminodes. Justicia nevlin¬
gii and the two other heteromorphically bracted
species of Justicia noted above exhibit two stami-
node-like protrusions near the base of the filaments.
It is not known whether these protrusions technically
are staminodes or whether they are merely thick¬
enings of the corolla near the point of insertion of
the filaments. Whatever their origin, they appear
to be present in species of Justicia with hetero-
morphic bracts. Heteromorphically bracted species
of Justicia are now known from Mexico and Central

America, and they undoubtedly occur in South
America as well. Determining whether they consti¬
tute a monophyletic infrageneric taxon within Jus¬
ticia deserves additional study.

Acknowledgments. We thank Alice Tangerini for
skillfully preparing the line drawings, and Stephen
F. Smith and the staff of the National Museum of
Natural History SEM Laboratory for providing the
high-quality pollen photomicrographs.

Literature Cited

Daniel, T. F. 1990. New and reconsidered Mexican
Acanthaceae. IV. Proc. Calif. Acad. Sci. 46: 279-
287.

-. 1993. New and reconsidered Mexican Acan¬
thaceae. V. Contr. Univ. Michigan Herb. 19: 271-
291.

Graham, V. A. 1988. Delimitation and infra-generic
classification of Justicia (Acanthaceae). Kew Bull.
43: 551-624.

Errata

Croat, T. B. & M. H. Grayum. 1994. Philodendron
Subgenus Pteromischum (Araceae) from Saul, French
Guiana. Novon 4: 211-219.

On page 216, second column, third paragraph,
in the discussion of Pteromischum placidum, the
following should be changed: “It [ Pteromischum
placidum ] is very easily confused with P. rud-
geanum Schott, which differs in climbing directly
up the side of the tree, and in having blades that
are matte rather than glossy on the lower blade
surfaces, a slender spadix, and in generally flowering
ahead of P. placidum. It was rarely at anthesis
during mid-February, whereas many plants of P.
placidum were in flower.” It should read: “It [ Pter¬

omischum placidum ] is very easily confused with
P. rudgeanum Schott. . . and in generally flowering
later than P. placidum. Philodendron placidum
was rarely in flower in mid-February, whereas many
plants of P. rudgeanum were in flower.”

On page 219, the first paragraph of the second
column, the following should be changed: “In ad¬
dition, Philodendron placidum flowers later than
P. rudgeanum , which was only rarely in flower in
mid-February when P. placidum was in full flower."
It should read: “In addition, P. rudgeanum flowers
later than P. placidum, which was only rarely in
flower in mid-February when P. rudgeanum was in
full flower.”

Volume 5, Number 1, pp. 1-118 of NOVON was published on 27 March 1995.

Novon 5: 118. 1995.

Volume 5
Number 2
1995

NOVON

MISSOURI BOTANICAL

SEP 1 2 1995

GARDEN LIBRARY

A Note on the Kazakh Endemic Lap pula zaissanica (Boraginaceae)

N. K. Aralbaev

Main Botanical Garden, Timiryazev St., 480070 Almaty, Republic of Kazakhstan

Abstract. Studies of the Kazakh endemic Lap-
pula coronifera (M. Popov) subsp. zaissanica Ar¬
albaev show that it is a distinct species. The new
combination Lappula zaissanica (Aralbaev) Aral¬
baev is proposed.

Recent collections of an interesting Lappula from
the Zaissan depression of eastern Kazakhstan were
referred to Lappula coronifera by Kudabaeva &
Aralbaev (1989). Subsequent collections showed that
the plants represent a distinct subspecies, L. coron¬
ifera subsp. zaissanica (Aralbaev, 1990). Further
studies and critical comparison with authentic plants
of L. coronifera, as well as additional fieldwork,
revealed that the plants in question represent a
distinct species quite common in the Zaissan de¬
pression. A new combination is herein proposed.

Lappula zaissanica (Aralbaev) Aralbaev, comb,
nov. Basionym: Lappula coronifera M. Popov
subsp. zaissanica Aralbaev, Bot. Zhurn. 75:
707. 1990. TYPE: E Kazakhstan. Northern
Zaissan: above Schubarschilik, sandy-pebbled
desert, 8 June 1984, Aralbaev s.n. (holotype,
Altay Botanical Garden, Leninogorsk (AIC); iso¬
type, LE).

Herbs annual, 7-15(-20) cm tall. Stems 1 or
several, branching at base or apex. Basal leaves
rosulate, dried at fruiting time; cauline leaves sessile,
lanceolate, 9-12 mm long, 1-2.5 mm wide, some¬
what longitudinally folded, sparsely tomentose and
greenish adaxially, moderately tomentose abaxially.
Bracts shorter than fruits, lowermost oblanceolate,
upper ones narrowly lanceolate. Pedicels tomentose,

as long as or slightly longer than calyx, slightly
reflexed in fruit. Calyx tomentose, with bristly tu¬
bercles; lobes in fruit narrowly lanceolate, 3.5-4
mm long, slightly reflexed. Corolla blue, 3 cm long,
with broadly campanulate limb. Fruit broadly ovoid;
nutlets similar, with a smooth crownlike cup turning
from yellow to brown as nutlets mature, each side
with 6 long, glochidiate spines shorter than or nearly
as long as the cup; disc not keeled, smooth, bright;
gynophore 2-2.5 mm long; style exserted from nut¬
lets by 0.2-0.3 mm.

Lappula zaissanica, which is a Kazakh endemic
known thus far only from the Zaissan depression,
is most closely related to L. coronifera, from which
it differs by having nutlets with smooth, not keeled,
bright-colored discs and broader wings with shorter
teeth.

Additional specimens examined. KAZAKHSTAN. N
Zaissan Region: in valleys above Kiin-Kerish, 18 May
1983, Aralbaev s.n. (paratype; AIC); Kara-Biryuk, peb¬
bled desert, 12 June 1993, Aralbaev s.n. (Main Botanical
Garden, Almaty).

Acknowledgments. I am grateful to Ihsan Al-
Shehbaz and Larissa A. Karavaeva for their help
with the manuscript.

Literature Cited

Aralbaev, N. K. 1990. A note on Lappula coronifera
(Boraginaceae). Bot. Zhurn. (Moscow & Leningrad)
75: 707-708.

Kudabaeva, G. M. & N. K. Aralbaev. 1989. Genus
Lappula Moench. in flora of north part of Zaissan’s
depression. Mat. Kazakh. Acad. Bot. Inst. Herb.
Alma-Ata 1989: 34-42,

Novon 5: 119. 1995.

New Species of Gibsoniothamnus (Bignoniaceae: Schlegelieae)

Kerry Barringer

Brooklyn Botanic Garden, 1000 Washington Avenue,

Brooklyn, New York 11225-1099, U.S.A.

Alwyn Gentryf

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT. Three new species of Gibsoniotham¬
nus L. O. Williams, G. grandiflorus, G. stellatus,
and G. versicolor, are described as new.

Gibsoniothamnus L. 0. Williams is a genus of
about 14 species of epiphytic shrubs, variously clas¬
sified in the Bignoniaceae or Scrophulariaceae. This
unusual group of Central American epiphytes is most
closely related to the neotropical genus Schlegelia
Miquel, currently classified in the Bignoniaceae tribe
Schlegelieae. Undescribed species have been filed
with both Scrophulariaceae and Bignoniaceae, so a
collaborative project was begun to identify and de¬
scribe the new species. Collaborative work was cut
short by the untimely death of A1 Gentry. The fol¬
lowing species were discovered during the study.

Gibsoniothamnus grandiflorus A. Gentry &
Barringer, sp. nov. TYPE: Panama. Veraguas:
ridge of Cordillera de Tute along trail to Cerro
Tute, ca. 3-4 km past Escuela Agricola Alto
de Piedra, just W of Santa Fe, 8°32'N,
81°07'W, 20 Mar. 1982, Knapp & Kress
4389 (holotype, MO; isotype, PMA). Figure 1.

Species fioribus grandis, a Gibsoniothamno truncato
lobis calyce triangulatis longe acuminatis, pedicellis lon-
gioribus differt.

Epiphytic shrub to 3 m tall; bark gray-brown;
stems angled, glabrous, to 5 mm diam., with raised
leaf bases at the nodes. Leaves opposite, subequal
to equal across the node; petioles 5-14 mm long,
glabrous or sparsely pubescent; blade ovate to ob-
ovate or lanceolate, 6.5-14.5 cm long, 3.0-6.2 cm
wide, coriaceous, glabrous, shiny above, the base
cuneate, the apex acuminate, secondary veins 3 or
4 per side, impressed above, prominent below, glan¬
dular pits in fields between the veins below, pocket
domatia occasionally present in the axils of the mid¬
vein and secondary veins toward the base of the
blade. Flowers in congested, axillary panicles; pri¬
mary peduncle to 1.5 cm long; bracts narrowly

Novon 5: 120-124. 1995.

triangulate, 3-5 mm long, pubescent; the secondary
peduncles to 7 mm long; bracteoles to 3 mm long,
ovate, acuminate; pedicels 2.5-3.5 cm long, thicker
above; calyx funnelform, the tube 5-9 mm long,
red to red-purple, the lobes narrowly triangulate,
unequal, 7-13 mm long, long-acuminate, glabrous
to minutely ciliate; corolla cylindric, slightly ampliate
above, 3.5-4.0 cm long, 6-7 mm diam., reddish
pink to red, fleshy, the upper lobes rounded, 3-4
mm long and wide, the lateral lobes rounded to
ligulate, 4-5 mm long, 4 mm wide, the median lobe
ligulate, 4-6 mm long, 3-4 mm wide; stamens at¬
tached 6-9 mm from the base of the corolla tube,
filaments 20-26 mm long, glabrous; staminode 10-
20 mm long, filiform, the apex clavate; pistil gla¬
brous, the ovary globose to oblate, 2-3 mm diam.,
the style 30 mm long, the stigma slightly bilobed.
Berry globose, 8-10 mm diam., green.

This species is most easily distinguished by its
large, slightly ampliate flowers. It appears to be most
closely related to Gibsoniothamnus truncatus and
G. latidentatus . It differs from G. truncatus by its
larger, ampliate corollas, pedicels more than 2 cm
long, and narrowly triangulate calyx lobes. It differs
from G. latidentatus by its corollas more than 3
cm long, reddish calyx, short staminode, and calyx
lobes 7-13 mm long. This species has an interesting
disjunct distribution: the plants from Panama, col¬
lected along the El Llano-Carti road, have narrower,
more acuminate leaves than those from Code, most¬
ly collected on Cerro Tute.

Paratypes. PANAMA. Code: vicinity of Escuela
Agricola Alto de Piedra, 3200-3400 ft., 3 Apr. 1980,
Antonio 3961 (MO); about 16 km NW of Santa Fe on
descent to Rio Caloveborita, 650 m, 4 Sep. 1975, Dress¬
ier 5142 (MO); near third branch of Rio Santa Maria,
10-14 km NW of Santa Fe, 650-750 m, 10-11 Oct.
1975, Dressier 5171 (MO); shoulder of Cerro Tute, 25
May 1977, Folsom & Edwards 3345 (MO); ridge of
Cerro Tute, along trail to Cerro Tute, 3-4 km past Escuela
Agricola Alto de Piedra, 8°32'N, 81°07'W, 800-1400
m, 20 Mar. 1982, Knapp & Kress 4363 (MO); Cerro
Tute, E slopes, 1 km beyond Escuela Agricola Alto Piedra

Volume 5, Number 2
1995

Barringer & Gentry

New Species of Gibsoniothamnus

121

above Santa Fe, 900-1200 m, 14 May 1981, Sytsma
& Andersson 4655 (MO, PMA). Panama: El Llano-
Carti road, 10 km N of Pan Am Highway, 340 m, 15
Feb. 1975, Gentry & Mori 14216 (F, MO); 6-7 mi.
from PanAmerican highway on El Llano-Carti road,
9°12'N, 79°00'W, 400 m, 26 Feb. 1982, Knapp &
Mallet 3829 (MO, PMA); 8.2 mi. from Panamerican
highway on the El Llano-Carti rd„ 9°14’N, 79°00'W,
450 m, 24 Mar. 1982, Knapp & Huft 4415 (MO, PMA);
toad between El Llano and Carti-Tupile road, 12 mi.
above Pan American highway, 200-500 m, 30 Mar.
!973, Liesner 1321 (F, MO, NY, PMA); El Llano-Carti
td-, 10.8 km from interamerican highway, 1100-1200
ft-, 27 Dec. 1974, Mori, Kallunki & Hansen 4121 (MO);
El Llano-Carti road, 21.6 mi. from Inter-American high¬

way, 350 m, 20 Mar. 1975, Mori & Kallunki 5104
(MO, PMA); El Llano-Carti road, 9 mi. from Pan Amer¬
ican highway, 350—400 m, 28 Apr. 1981, Sytsma 4146
(NY).

Gibsoniothamnus stellatus A. Gentry & Barrin¬
ger, sp. nov. TYPE: Panama. Chiriqui; Cerro
Colorado, along road to copper mine 33.1 km
beyond the bridge over the Rio San Felix, 13.1
km beyond turnoff to Escopeta, 1400 m, 15
July 1976, Croat 37200 (holotype, MO; iso¬
type, MO). Figure 2.

122

Novon

A Gibsoniothamno alato foliis minoribus, petiolis 3-
5 ram longis, tubis calyce 6-8 mm longis, staminibus 17—
18 mm longis differt.

Epiphytic shrub to 1.5 m tall; bark pale gray-
brown; stems angled, glabrous, the leafy portions to
2 mm diam., the older portions with raised leaf bases.
Leaves opposite, unequal to subequal; petioles 3-5
mm long, glabrous, unwinged; lamina ovate-lanceo¬
late, coriaceous, glabrous, gland-dotted, the base
cuneate, the apex acuminate to acute, the midvein

impressed above, prominent below, the secondary
veins not impressed above, prominent below, 2-3
per side, domatia occasionally found on the under¬
side of the lamina near the base, the larger leaves
15-25 mm long, 9-16 mm wide, the smaller leaves
5-12 mm long, 4-8 mm wide. Flowers borne near
the tips of the branches, solitary, axillary; pedicels
10-12 mm long, thick above, bracteolate at base,
glabrous; calyx campanulate, purple, the tube 6-8
mm long, the lobes 8-9 mm long, triangulate, the
wings broad, 3-4 mm wide; corolla tubular, white.

Volume 5, Number 2
1995

Barringer & Gentry

New Species of Gibsoniothamnus

123

2.0-2.5 cm long, 4-5 mm diam., the upper and
lateral lobes rounded to ligulate, 3-4 mm long and
wide, the margin ciliate, the median lobe ligulate, 3
mm long, 1.5-2.0 mm wide; stamens attached 5
mm above the base of the corolla tube, with a ring
of trichomes at the point of insertion, the filaments
17-18 mm long; staminode 5-6 mm long; pistil

glabrous, the ovary globose, 1.5 mm diam., the style
14 mm long, the stigma capitate. Immature berries
globose, green.

This species is distinguished by its small leaves
and large, winged calyx lobes. It appears to be most
closely related to Gibsoniothamnus alatus, native

124

Novon

to southern Darien and adjacent Choco, because
both species have strongly alate calyces and white
corollas. It can be distinguished from that species
by its smaller leaves, calyx tube more than 5 mm
long, and stamens less than 18 mm long.

Paratypes. PANAMA. Bocas del Toro: region of
Cerro Colorado, 7.5 mi. from Chami camp, 8°35'N,
81°45'W, 1220-1250 m, 13 Apr. 1986, McPherson
8882 (MO, NY, PMA).

Gibsoniothamnus versicolor A. Gentry & Bar¬
ringer, sp. nov. TYPE: Panama. San Bias:
Playon Chico, Rio Ukupseni, Campamento Neba
Dummat hasta las cascada, 50-100 m, 9°15'N,
78°15'W, 30 Oct. 1991, H. Herrera, J. Mor¬
ris & J. Mojica 1022 (holotype, MO; isotypes,
BKL, PMA). Figure 3.

A Gihsoniothamno allenii foliis majoribus anguste el-
lipticis, petiolis ciliatis, floribus paniculatis, lobis calyce
majoribus differt.

Epiphytic shrub or climber; bark gray-brown;
stems angled, glabrous to sparsely pilose, with raised
leaf bases at the nodes. Leaves opposite, equal to
subequal; petioles 5—12 mm long, pilose, unwinged;
blade narrowly elliptic, 4-11 cm long, 1.7-4.0 cm
wide, subcoriaceous, dull olive gray above, tannish
below, sparsely pilose above, with scattered tri-
chomes along the midvein below, strongly lepidote-
glandular below, domatia absent. Flowers in terminal
panicles; pedicel 2-3 cm long, sparsely pilose, slight¬
ly thickened above; calyx campanulate, pink to
mauve, the tube 3—5 mm long, the lobes nearly
linear, 7-12 (-15 in fruit) mm long, porrect, then
reflexed after anthesis; corolla tubular, white or red¬

dish violet, 2 cm long, 4 mm wide at the base, 3
mm wide near the apex, the lobes ovate to round,
1.5-3.0 mm long; stamens inserted 4-5 mm from
the base of the corolla tube, with a basal ring of
trichomes, the filaments 1 cm long; staminode 5
mm long, filiform; ovary depressed-globose, 2 mm
long, glabrous, the style 15 mm long, the stigma
subcapitate. Young fruit green, subglobose, 6 mm
long.

This species differs from Gibsoniothamnus alatus
A. Gentry, the only other species in this genus known
from eastern Panama, by its linear, unwinged calyx
lobes. It appears to be most closely related to G.
allenii A. Gentry, which grows in the hills around
El Valle. This species differs from G. allenii by its
longer calyx lobes, 7-15 mm long, and its larger,
subcoriaceous leaves. D’Arcy (1979) listed Croat
27281 under G. cornutus (Donnell Smith) A. Gen¬
try, but that species differs by its short, triangulate
calyx lobes and coriaceous leaves.

Paratype. PANAMA. Darien: headwaters of the Rio
Tuquesa, gold mining camp of Tyler Kitteridge, ca. 2 air
km from the continental divide, 26 Aug. 1974, Croat
27281 (MO).

Acknowledgments. I thank the curators of F,
MO, NY, and US for allowing us to study their
collections.

Literature Cited

D’Arcy, W. G. 1979. Scrophulariaceae. In: R. E.
Woodson & Shery, Flora of Panama. Ann. Missouri
Bot.Gard. 77:173-274. [Gibsoniothamnus pp. 220-
227.]

Two New Species of Weinmannia (Cunoniaceae) from the
Venezuelan Guayana

Paul E. Berry and Jason Bradford

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Abstract. Two new species of Weinmannia (Cu¬
noniaceae) from the highlands of southern Venezuela
are described and illustrated. Weinmannia ilute-
puiensis P. E. Berry & J. Bradford is a simple¬
leaved, serrate-margined species known only from
Ilu-tepui in Estado Bolivar, and Weinmannia coro-
coroensis J. Bradford & P. E. Berry is a pinnate¬
leaved species with tiny, revolute-margined leaflets
currently known only from Cerro Coro Coro in Es¬
tado Amazonas.

While preparing the floristic treatment of Cu¬
noniaceae for the Flora of the Venezuelan Guayana
and as part of a broader study of neotropical Wein¬
mannia by the second author, two distinctive new
species of Weinmannia were found among the col¬
lections examined from the tepuis, or tabletop moun¬
tains, of southern Venezuela. Both species were col¬
lected from relatively poorly explored mountains,
one from Ilu-tepui in Estado Bolivar, and the other
from Cerro Coro Coro in Estado Amazonas. These
species are described and illustrated below.

Weinmannia ilutepuiensis P. E. Berry & J.
Bradford, sp. nov. TYPE: Venezuela. Estado
Bolivar: Ilu-tepui, lower plateau, with varied
habitats, rocky, boggy, and short forest to 5 m
tall, 05°25'03"N, 60°29'W, 2500 m, 16 Apr.
1988, Liesner 23413 (holotype, MO; isotypes,
NY, US, VEN). Figure 1A-D.

Frutex vel arbor l-5(-10)-metralis, foliis simplicibus,
laminis foliorum infra dense ochraceo-lanatis, ellipticis,
coriaceis, marginibus serratis, 1. 5-6.0 x 0.8-3. 5 cm;
pseudoracemis 4-6 cm longis, lanatis, pedicellis sub an-
thesi 1-2.5 mm longis post anthesin 2.5-4 mm longis;
capsulis villoso-lanatis, 3-3.5 mm longis.

Shrub or small tree 1 -5(-10) m tall; young stems
lanate, nodes hirsute. Leaves opposite, simple, ellip¬
tical, subcoriaceous, 1.5-6.0 cm long, 0.8-3.5 cm
wide, broadly acute to rounded at the apex, broadly
acute at the base, sparsely pilose to glabrescent and
dark green on the upper surface, light brown and
densely lanate-villous on the lower surface; second¬
ary veins 10-15 per side, impressed on the upper
surface and prominent on lower surface, subparallel

from midvein until divaricating within ca. 4 mm of
the margin with each branch joining an adjacent
branch at a marginal sinus; margins serrate, planar
to revolute, with 10-15 glandular teeth per side
between successive sinuses or adjacent to a sinus;
petioles 3-4 mm long, lanate; stipules broadly ellip¬
tical to suborbicular, 3-6 mm long, 3-5 mm wide,
hirsute on the outside, dark brown and glabrous
adaxially. Pseudoracemes in pairs at the branch
apex, exceeding the leaves, 4-6 cm long, axis lanate.
Flowers arranged in fascicles along the inflorescence
axis with 1-several flowers per fascicle; bracteoles
subtending fascicles, linear to obovate, 1.5-2 mm
long, with long hairs on the dorsal surface; pedicels
1.0-2.5 mm long in flower, 2.5-4.0 mm long in
fruit, lanate; sepals four, elliptic to narrowly deltoid,
acute, 1-1.5 mm long, lanate abaxially, glabrous
adaxially, persistent; petals four, narrowly obovate,
1.5-2.5 mm long, ca. 1 mm wide, glabrous, off-
white; filaments eight, 2.5-4.0 mm long, anthers
oblong, ca. 0.4 mm wide, glabrous; ovary lanate,
ca. 1.5 mm long, the 2 styles divergent and each
1.5 mm long, basally pubescent but glabrous toward
the tips. Capsules 3.0-3.5 mm long (not including
style), lanate-villous; seeds oblong, ca. 1.0 mm long,
sparsely covered with simple hairs 0.5 mm long.

Ecology and distribution. Occurring in patches
of short forest on lower summit plateau overlying
sandstone, known only from Ilu-tepui in the eastern
tepui chain of Estado Bolivar, Venezuela, between
2500 and 2630 m elevation.

In the size and shape of leaves, inflorescences,
flowers, and fruits, Weinmannia ilutepuiensis most
closely resembles the Andean W. rollottii Killip,
especially variety testudineata (Cuatrecasas) Ber¬
nards Both species belong to series Dryadifoliae
Bernardi (Bernardi, 1963). However, W. ilute¬
puiensis differs in its dense, lanate-villous indumen¬
tum, including the calyx and the capsule (which are
glabrous in W. rollottii).

On the same helicopter trip to Ilu-tepui during
which Ronald Liesner collected the type specimen,
he also made three other collections of Weinmannia
(numbers 23422, 23417, and 23366, all at MO).
None of these additional collections fits easily into

Novon 5; 125-127. 1995.

126

Novon

D 8 Detail of ^r‘»7T a i au ‘:f ul(,n r P , E - Berr y & J - Bradf °rd- -A. Habit. -B. Flower. -C. Fruit. -

LctLL'f Sft.SE? E BSrirSStf ^ 2 f 38 ^nmannia

fruit. Drawn from Huber 12296. Ha b.t. F. Deta.1 of leaf pa.r and stipule. -G. Flower. -H. Young

Volume 5, Number 2
1995

Berry & Bradford
Weinmannia from Venezuela

127

any known species, and they appear to be inter¬
mediates between W. ilutepuiensis and W. bra-
chystachya Willdenow ex Engler. Since these two
species are so different phenotypically ( W. bra-
chystachya is characterized by small, compound,
glabrous leaves with pseudo-umbellate inflores¬
cences), the apparent hybrids were only recognized
when all Liesner collections from the area were seen
together. Weinmannia brachystachya is not yet
known from Ilu-tepui, but it does occur on the near¬
by Roraima-tepui.

Paratypes. VENEZUELA. Bolivar: Cumbre del Ilu-
(Uru-)tepui, sector centro-meridional, vegetacion herba-
ceo-fruticosa sobre superficies poco disectadas, 5°25'N,
60°59'W, 2630 m, 29 Apr. 1984, Huber 9520 (MYF,
US); Ilu-tepui, lower plateau, with varied habitats, rocky,
boggy, and short forest to 5 m tall, 05°25'03"N, 60°29'W,
2500 m, 15 Apr. 1988, Liesner 23338 (MO, US, VEN);
Ilu-tepui, cumbre slopes, saddle between N peak and
central plateau, 7000-7800 ft., 15 Mar. 1952, Maguire
33426 (NY, US); Ilu-tepui, slopes below uppermost
W-facing escarpment, 7500-8000 ft., Maguire 33508
(NY, US).

Weinmannia corocoroensis J. Bradford & P.
E. Berry, sp. nov. TYPE; Venezuela. Estado
Amazonas: cumbre del Cerro Coro-Coro, en las
cabeceras nor-occidentales del Rio Manapiare
(sector NW de la Serranxa Yutaje), 05°46'N,
66°12'W, 2200 m, 12 Nov. 1987, Otto Huber
12296 (holotype, MO; isotypes, MYF, VEN).
Figure 1E-H.

A Weinmannia guyanensi Klotzscb foliolis subintegris
valde revolutis, foliolo terminali lateralibus simili, inffo-
rescentiis congestioribus, pubescentia densiore longio-
reque differt.

Shrub 0.5-1.5 m tall; young stems villous, the
hairs 1-1.5 mm long. Leaves opposite, imparipin-
nate, rachis winged, rigid-coriaceous, leaflets 1- or
2-paired (occasionally unifoliate), the terminal and
lateral leaflets elliptical to obovate, similar in size
and shape, 5—12 mm long, ca. 3—5 mm wide when
flattened but lateral margins strongly inrolled, lateral
margins entire, apex generally with three small teeth,
upper surface glabrous with hairs occasional along
midvein, lower surface glabrous except for strigose
hairs 1-1.5 mm long along midvein especially to¬
ward the base; secondary veins 6—9 per side, barely
visible below, subparallel from midvein until divar¬
icating within ca. 1 mm of the margin with each
branch joining an adjacent branch and together
forming a vein which extends to the margin; petioles
1-2 mm long, densely villous; stipules obovate, cil-
iate, hairs sparse on outer surface, glabrous on inner
surface, 2-4 mm long, 1.5-3.5 mm wide. Pseu¬

doracemes in axillary pairs near branch tips, ex¬
ceeding the leaves, 20-30 mm long in flower, axis
villous, the basal 8—12 mm sterile, the apical portion
densely covered with flowers. Flowers arranged in
fascicles along the inflorescence axis with 1-several
flowers per fascicle; bracteoles subtending fascicles,
rectangular, apex blunt, ciliate, 0.7-1 mm long,
0.5-0.8 mm wide; pedicels 1.5-3.0 mm long in
flower, with sparse, shaggy hairs; sepals four, nar¬
rowly deltoid, acute, 1.0-1.5 mm long, hairs on
midvein and apex; petals four, obovate, 1.5-2.0
mm long, ca. 1 mm wide, glabrous, white, with a
prominent midvein divaricating near the apex; fil¬
aments eight, 2.0-2.8 mm long, anthers oblong, ca.
0.4 mm long, glabrous; gynoecium 1.5-1.8 mm
long, glabrous, the two styles divergent, glabrous,
ca. 1 mm long. Mature fruits not seen, recorded on
labels as being red.

Ecology and distribution. Locally frequent in
tepui scrub communities overlying sandstone and in
high-tepui meadows along streams; known only from
the summit of Cerro Coro Coro in Estado Amazonas,
Venezuela, at 2200 m elevation.

Weinmannia corocoroensis resembles W. guy-
anensis Klotzsch, especially variety quinata Cua-
trecasas, in its habit and leaf venation pattern, but
differs in having denser, longer pubescence along
the stems, nodes, and inflorescence structures;
smaller, more revolute and nearly entire leaflets;
terminal and lateral leaflets of the same size; and
shorter, more congested inflorescences. This species
was noted in the field by Otto Huber as being com¬
mon, while another specimen ( Huber 12332, MO)
from the same location and apparently belonging to
W. guyanensis was rare. Weinmannia corocoroen¬
sis is also similar to W. brachystachya Willdenow
ex Engler in the few, apical teeth of the leaflets and
in its shrubby habit, but differs in its more elongate
inflorescence, denser pubescence (on stems, mid¬
veins of leaves, inflorescence axis, and petioles),
more regularly elliptical leaflets, and more congested
nodes on the stem.

Acknowledgments. This work was supported in
part by National Science Foundation grant DEB-
9201044. We thank Jose Cuatrecasas for sharing
his knowledge of the genus Weinmannia with us
and first suggesting that W. ilutepuiensis was a new
species. Bruno Manara made the line drawings of
the two species, and Roy Gereau kindly reviewed
the Latin diagnoses.

Literature Cited

Bernardi, L. 1963. Revisio generis Weinmannia. Pars
II: Sectio Simplicifoliae. Candollea 18: 285-334.

New Combinations in Eastern North American Elymus (Poaceae)

Julian J. N. Campbell

The Nature Conservancy, 642 W. Main St., Lexington, Kentucky 40508, U.S.A.

ABSTRACT. Two new varietal combinations are made
in the grass genus Elymus: E. glabriflorus (Vasey)
Scribner & Ball var. australis (Scribner & Ball) J.
J. N. Campbell, and E. villosus Muhlenberg var.
arkansanas (Scribner & Ball) J. J. N. Campbell.

The following new combinations have been found
necessary in preparing the treatment of eastern Ely¬
mus species for the forthcoming Manual of North
American Grasses (M. E. Barkworth & K. Capels,
editors, in prep.).

Elymus glabriflorus (Vasey) Scribner & Ball var.
australis (Scribner & Ball) J. J. N. Campbell,
comb. nov. Basionym: Elymus australis Scrib¬
ner & Ball, U.S.D.A. Div. Agrostol. Bull. 24:
46. 1901. TYPE: U.S.A. North Carolina: Bilt-
more Herbarium 411b (holotype, US).

Elymus villosus Muhlenberg var. arkansanas
(Scribner & Ball) J. J. N. Campbell, comb. nov.
Basionym: Elymus arkansanas Scribner & Ball,
U.S.D.A. Div. Agrostol. Bull. 24: 45. 1901.
TYPE: U.S.A. Arkansas: Harvey 7 in part
(holotype, US).

In most recent treatments, these taxa have gen¬
erally been recognized at the forma level or not at
all. The morphological differences are largely a mat¬
ter of presence or absence of pubescence in the
spike. However, each taxon is locally more frequent
than typical plants of the species, and there appear
to be some associations with different habitats and
some geographic trends.

Novon 5: 128. 1995.

Festuca diclina (Poaceae), a New Species from Northwestern Mexico

Stephen J. Darbyshire

Centre for Land and Biological Resources Research, Agriculture and Argi-Food Canada,
Wm. Saunders Bldg., Central Experimental Farm, Ottawa, Ontario, K1A 0C6, Canada

ABSTRACT. A series of collections from Guadalupe
y Calvo in southern Chihuahua State, Mexico, is
described as a new species, Festuca diclina (Po¬
aceae), and compared with similar species from Cen¬
tral America.

Festuca diclina S. J. Darbyshire, sp. nov. TYPE:
Mexico. Chihuahua: Mpio. Guadalupe y Calvo,
Cerro de Mohinora, S de Guadalupe y Calvo,
3300 m, alpina-subalpina, en ladera rocosa, 27
Aug. 1987, McDonald & Martinez 2392 (ho-
lotype, DAO 661232; isotypes, TEX (ex DAO
591464), MEXU (ex DAO 592610)). Fig¬
ure 1.

Plantae perennes caespitosae, innovationibus intrava-
ginalibus praeditae. Vaginae foliorum ima basi tantum
integrae, vertice scabrae; ligulae 0.2-0.5 mm longae;
laminae foliorum plicatae vel convolutae, 1.2-1.5 mm
latae, abaxialiter ima basi scabrae, adaxialiter sparsim
scabrae, 5-7-costatae, fasciculis sclerenchymaticis cum
nervis infra et saepe supra conjunctis, non confluentibus.
Culmi 30-75 cm longi, glabri. Paniculae 8-15 cm longae,
plus minusve laxae, pyramidales, axe et ramulis glabris
vel trichomatibus sparsis, ramulis interdum tortuosis. Spic-
ulae 8-12 mm longae, (2-)3-4(-5) flosculis; glumae in-
feriores 2.4-4 mm longae, 1-nerviae; glumae superiores
3.5-5 mm longae, (l-)3-nerviae; rachillae trichomatibus
0.1-0.3 mm longae praeditae; lemmata lanceolata, glabra
vel apice scabra, l-3(-5)-nervia, 6-8 mm longa, inter¬
dum arista ad 1.0 mm longa; paleae 5.5-7 mm longae,
bidentatae, apice piloso-hirsutae inter nervos, nervis cum
trichomatibus 0.1-0.2 mm longis prope apicem; antherae
interdum abortivae praesertim flosculis inferioribus, 3-
4.2 mm longae; ovariorum apex dense pubescens.

Caespitose, loosely tufted perennial (Fig. 1A);
shoots intravaginal. Vegetative shoots to 25 cm tall,
light green (not glaucous); sheaths rapidly decaying
to fibers, the margins closed only at the base, the
upper parts of the sheaths, collars, and lower abaxial
blade surface retrorsely scabrous; auricles absent;
ligules short, 0.2-0.5 mm long, acute to truncate,
sometimes lacerate, puberulent apically and abaxi-
ally, membranous; blades plicate to convolute, adax-
ially sparsely scabrous, 1.2-1.5 mm wide on flat¬
tened blades (about 0.4-0.8 mm in the longest di¬
mension of a cross section), with 7(-9) veins and
5-7 adaxial ribs; sclerenchyma in discrete fascicles
opposite veins abaxially and usually adaxially (Fig.

2A-C), the fascicles usually joining veins abaxially
and sometimes adaxially thus forming a girder be¬
tween the epidermises through the veins (Fig. 2A).
Culms 30-75 cm tall, glabrous; nodes black or brown,
glabrous. Flag leaf sheath sometimes loose. Panicle
open, pyramidal, 8-15 cm long; rachis glabrous
below and sparsely scabrous on the angles above;
branches ± flexuous and sometimes sinuously un¬
dulating, 1 or 2 per node, glabrous or sparsely
scabrous, the lowest 5-9 cm long. Spikelets 8-12
mm long, distal on the branches, with (2-)3-4(-5)
florets; glumes chartaceous, glabrous or scabrous on
the keel (especially apically), ovate-lanceolate to lan¬
ceolate with wide hyaline margins; lower glume 2.4-
4 mm long, 1-nerved; upper glume 3.5-5 mm long,
(l-)3-nerved; rachilla with hairs 0.1-0.3 mm long
(Fig. 1C); lemmas 6-8 mm long, narrowly lanceolate
(Fig. IB), glabrous or scabrous-hirsute apically, 1-
3(-5)-nerved (the lateral ones faint), the apex at¬
tenuate, sometimes with a terminal awn or mucro
to 1.0 mm long; paleas subequal to the lemmas,
5.5-7 mm long, bidentate, apically pilose-hirsute
between the nerves and with hairs 0.1-0.2 mm long
on the nerves (Fig. ID, E); anthers sometimes abor¬
tive especially in lower florets, 3-4.2 mm long when
developed; ovary apex densely pubescent (Fig. IF);
styles 2. Caryopses not seen.

Specimen label data from the type collection de¬
scribe the culms as up to 2 m high, although all
specimens examined were much less.

Distribution. Collections of Festuca diclina are
known only from the vicinity of Guadalupe y Calvo,
Sierra Madre Occidental, Mexico, in montane pine
forests at altitudes of 2950-3300 m.

Festuca diclina belongs to Festuca subg. Festuca
and is similar to F. hintoniana E. B. Alexeev (Figs.
1G-K, 2D) but differs primarily in the loose tufts
with non-persistent sheaths decaying to separate fi¬
bers (densely tufted with persistent chartaceous
sheaths in F. hintoniana), scabrous upper sheaths,
collars, and lower leaf blades (not glabrous), nar¬
rower leaf blades (1.6-1.9 mm on flattened blades
and 0.7-1.0 mm in longest dimension of transverse
section in F. hintoniana ), blade sclerenchyma in
discrete fascicles (more or less continuous abaxial

Novon 5: 129-132.1995.

130

Novon

Figures A, G
Figures B, D, H, J
Figures C, E, F, I, K

1 cm
1 mm
1 mm

Volume 5, Number 2
1995

Derbyshire
Festuca diclina

131

C D

Figure 2. Camera lucida drawings of cross sections of mature leaves from vegetative shoots. A C. Festuca diclina
S. J. Darbyshire. —D. F. hintoniana E. B. Alexeev. (A based on McDonald & Martinez 2392 (TEX), B based on
McDonald & Nesom 2475 (ARIZ 285274), C based on Nesom & McDonald 6473 (ARIZ 281336), D based on
Hinton 17243 (K).)

Figure 1. Festuca species. A-F. F. diclina S. j. Darbyshire. —A. Habit. —B. Lower floret. —C. Rachilla. —D.
Palea. — E. Palea apex. —F. Ovary. G K. F. hintoniana E. B. Alexeev. —G. Habit. —H. Lower floret. —I.
Rachilla. —J. Palea. —K. Palea apex. (A composite, B-F based on McDonald & Nesom 2489 (DAO 592613), G-
K based on Hinton 17243 (K).)

132

Novon

band in F. hintoniana; Fig. 2D), which extend in¬
ward to meet vascular bundles, sclerenchyma fas¬
cicles of the blades visible from the abaxial surface
(invisible or very faint in F. hintoniana ), longer
rachilla hairs (0.05-0.07 mm in F. hintoniana; Fig.
11), longer hairs on the palea veins (scabrous prickles
0.02-0.05 mm in F. hintoniana; Fig. IK) and
pilose-hirsute apically between the palea veins (gla¬
brous in F. hintoniana). The Mexican species F.
coahuilana Gonzalez-Ledesma & S. D. Koch is dis¬
tinguished from F. diclina by the longer ligules (to
1 mm), which form erect auricular projections, wider
leaf blades (1.5-2.2 mm) with 8-11 nerves, erect
inflorescence branches, shorter lemmas (5.0-6.2
mm), and shorter anthers (2.5-2.7 mm). Two taxa
recently described from Costa Rica, F. herrarae
Davidse and F. talamancensis Davidse, are both
readily distinguished from F. diclina by their gla¬
brous ovaries and shorter anthers (less than 2.1 mm
long). Festuca diclina is known from the Sierra
Madre Occidental of northwestern Mexico, while
both F. hintoniana and F. coahuilana are known
only from the Sierra Madre Oriental in northeastern
Mexico.

This species is so named because the lower florets
within a spikelet frequently contain abortive anthers,

while the upper florets frequently contain apparently
abortive ovaries.

Paratypes. MEXICO. Chihuahua: Mpio. Guadalupe
y Calvo, NW side of Cerro Mohinora, ca. 13 mi. SW of
Guadalupe y Calvo, 25°57'N, 107°03'W, open pine-fir
woods with scattered spruce, top and near top of summit,
open rocky slopes at lip of precipice with scattered krum-
holz [sic] pines, 3250 m, 21 Aug. 1988, McDonald &
Nesom 2489 (ARIZ 285277, DAO 592612, 592613);
NW side of Cerro Mohinora, ca. 13 mi. SW of Guadalupe
y Calvo, 25°57’N, 107°03’W, open pine-fir woods with
scattered spruce, top and near top of summit, open rocky
slopes at lip of precipice with scattered krumholz pines,
3250 m, 21 Aug. 1988, McDonald & Nesom 2475
(ARIZ 285274, DAO 592611); N side of Cerro Mohi¬
nora, ca. 13 mi. SW of Guadalupe y Calvo, 25°57'N,
107°03'W, open pine-fir woods with scattered spruce,
nearly vertical N-facing rock wall, very moist with many
bryophytes and rich herbaceous flora, 2950 m, 20 Aug.
1988, Nesom & McDonald 6473 (ARIZ 281336, TEX).

Acknowledgments. I am grateful to Andy
McDonald for sending material from his collections,

J. Cayouette and John and Charlotte Reeder for
useful comments, James E. Darbyshire for assistance
with the Latin description, Marcel Jomphe for the
illustrations, and herbarium curators at ARIZ, DAO,

K, and TEX for making specimens available for
study.

Three New Species of Rustia (Rubiaceae, Condamineeae) from
Panama and Ecuador

Piero G. Delprete

Botany Department, The University of Texas at Austin, Austin, Texas 78713, U.S.A.

Abstract. Three new species of Rustia are de¬
scribed, two from Ecuador ( R. alba and R. viri-
diflora) and one from Panama (R. dressleri ). Illus¬
trations of the three species are provided.

RESUMEN. Se describen tres especies nuevas de
Rustia, dos de Ecuador (R. alba y R. viridiflora)
y una de Panama (R. dressleri). Se incluyen ilus-
traciones de las tres especies.

Rustia (Cinchonoideae, Condamineeae; Hooker,
1873; Robbrecht, 1988; Delprete, in prep.) is a
genus of trees and shrubs that was founded by Jo¬
hann Friedrich Klotzsch, curator of the Berlin her¬
barium in the mid-1800s. Klotzsch (1846) separated
Exostema formosum Chamisso & Schlechtendal to
establish Rustia, in honor of Dr. Rust (“Dem An-
denken des verstorbenen Prasidenten Dr. Rust gew-
idmet”). Rustia, along with Tresanthera H. Kar-
sten, is unique in the woody Rubiaceae in having
poricidal anthers and pellucid-punctate leaves. Rus¬
tia comprises approximately 15 species endemic to
tropical America, from Nicaragua to southern Brazil.
The only previous work on Rustia was published
by Donald R. Simpson (1976), but this was restricted
to only a few species of western South America.

During preparation of a monographic treatment
of the tribe Condamineeae (Delprete, in prep.), I
examined several herbarium specimens from poorly
collected areas of Central and South America, many
of them unidentified or incorrectly identified. Among
these collections, I encountered three species new
to science, two from Ecuador and one from Panama.

In the following descriptions I use the abbrevia¬
tions L/W to represent the length to width ratio of
the leaf blades, and BA to represent the basal angle
from the leaf margin to the midrib.

Rustia alba Delprete, sp. nov. TYPE: Ecuador.
Carchi: environs of Maldonado, wet montane
forest, 1450-1650 m, 31 June 1978, M. T.
Madison 4808 with T. C. Plowmann, H. A.
Kennedy & L. Besse (holotype, AAU; isotypes,
F, QCA, SEL not seen, US). Figure 1.

Arbores ve! frutices glabri; stipulae interpetiolares cad-
ucae. Foliorum laminae 23-28 cm longae, 13-16 cm
latae, glanduli-punctatae; domatia desunt. Inflorescentia
40-46 cm longa; alabastra floralia cylindrica aestivatione
valvati-reduplicata; calyx cupulatus ac membranaceus 1.5-
3.0 mm longus; corolla alba 1.5-1.9 cm longa ad basim
constricta lobis % longitudinis corollae; antherae exsertae
dorsifixae concavae poricidales, thecae ad basim subae-
qualis; filamentis basiliter dilatatis ascendenti-strigosisque.
Capsulas non visi.

Shrub or tree, with several straight trunks 6 m
tall (probably stump sprouts). Young branches
smooth, glabrous, terete; older branches rugose, te¬
rete, grayish to pale brown and with sparse lenticels.
Stipules interpetiolar, free at base, present only in
bud and overlapping, narrowly triangular, glabrous
outside, glabrous and with few basal colleters inside,
2.8-3.7 mm long, 7-9 mm wide at base, reddish
green, leaving a linear scar ca. 0.5 mm wide of the
same color as the stem. Leaves petiolate, petioles
2-3 cm long, 3.0-4.5 mm thick, adaxially narrowly
concave to flattened, glabrous; basally thickened,
not pulvinate; blade 23-28 cm long, 13-16 cm
wide, L/W = 1.6:1 to 1.7:1; widely elliptic to
widely obovate, acute to rounded at base (BA =
30-45°), obtuse and short-acuminate at apex; the
acumen up to 1 cm long; blade when fresh pale
green above and yellowish green below, subcoria-
ceous, drying pale green, glabrous above and below,
pellucid punctate (see Fig. 1). Primary and second¬
ary veins glabrous and prominent below, secondary
veins 16-21 each side; tertiary veins starting sub¬
parallel and openly reticulate in the center, faintly
evident above and below; domatia absent. Inflores¬
cences terminal, laxly paniculate and pyramidal,
40-46 cm long, basal branches 22-30 cm long,
lateral branches 6-8 pairs, decussate; basal portion
of axis not branched, 7-11 cm long. Rachis terete,
rachis and branches glabrous to minutely hirtellous;
flowers on distal branches in cymules (rarely alter¬
nate). Distal bracts 4-5 mm long, 4-5 mm wide,
deltoid; bracteoles subtending the flowers 2-3 mm
long, 2-3 mm wide, deltoid, glabrous. Flowers pro-
tandrous, pedicellate, pedicel 4-6 mm long, glabrous
to minutely puberulent; hypanthium narrowly ob-
conical, 4-6 mm long, 2-3 mm wide, glabrous.
Calyx cupular with small lobes, membranous and

Novon 5: 133-139. 1995.

/ cm

134

Novon

Figure 1. Rustia alba Delprete. —A. Habit of inflorescence with mature leaf. —B. Abaxial view of blade showing
peUueid punctation. —C. Flower bud showing cupular calyx. —D. Anther. —E. Detail of anther external surface.
— F. Style. (Drawn from type: Madison et al. 4808.)

2 mm

Volume 5, Number 2
1995

Delprete

Rustia from Panama and Ecuador

135

glabrous, 1.5-3.0 mm long, calyx lobes 5, 1.0-2.5
mm long and 1.5-2.0 mm wide at base, glabrous
and ciliolate; calyx persistent. Corolla tubular, 1.5-
1.9 cm long, with a basal constriction and reflexed
lobes, flower bud cylindrical and pointed at apex,
aestivation valvate with contact zone, white, and
semifleshy when fresh; tube subcylindrical with basal
constriction, 5-6 mm long, ca. 2 mm wide at base
and 3.5-4.0 mm wide at the orifice; glabrous and
minutely verrucose outside; microscopically (40 x)
granulose inside, without a ring of hairs; corolla lobes
5, 9-11 mm long, ca. 3 mm wide, narrowly tri¬
angular and reflexed, glabrous and minutely ver¬
rucose outside; densely ascending short-strigose at
base, minutely papillose throughout and microscop¬
ically (40 X) papillose at margins inside. Lobes % of
corolla length. Stamens 5, fully exserted, subequal,
filaments attached 4-5 mm from the base of the
tube, anthers cylindrical and subsagittate at base
(see Fig. 2), rectilinear to slightly curved, 6.5-7.0
mm long, ca. 1.5 mm wide, dorsifixed near the base,
the thecae subequal at base, papillose throughout,
dehiscing by two pores at apex; filaments ca. 3 mm
long, flattened and widened at base, with a basal
tuft of ascending strigose hairs 0.3-0.5 mm long.
Style exserted, 16-20 mm long, glabrous, style
branches ovate, 3.0-3.5 mm long and 1.0-1.3 mm
wide at base, stigmatic surface papillose. Ovary two-
celled, placentation axile, turbinate to narrowly ob¬
long, glabrous, with many ovules in each locule
horizontally inserted. Capsule not seen, but probably
obconical to obovoid, like the hypanthium.

Rustia alba is similar to R. formosa (Chamisso
& Schlechtendal) Klotzsch but differs mainly in hav¬
ing a bigger inflorescence (exceptionally up to 36
cm long in R. formosa), smaller flowers; calyx cu¬
pular, more expanded and membranous; shorter
styles; and longer membranous style lobes (ca. 3
mm long in R. alba vs. ca. 1 mm long in R.
formosa ).

Rustia alba is known only from the type, which
was collected in the cloud forest of northern Ec¬
uador, around Maldonado; in contrast, R. formosa
is endemic to central and southern Brazil. Rustia
alba is easily distinguishable from all other species
of Rustia in western South America by its white
erect flowers (hence the epithet) and by its widely
obovate leaves.

Rustia viridiflora Delprete, sp. nov. TYPE: Ec¬
uador. Morona-Santiago: Achutza, Jibaro set¬
tlement in the vicinity of Macuma, ca. 50 km
N of Macas, 23 Mar. 1973, H. Lugo 3674
(holotype, GB sheets A and B). Figure 2.

Arbores usque ad 16 m altae; stipulae interpetiolares
caducae. Foliorum laminae 34-62 cm longae, 9.5-20.5
cm latae, glandulis punctatae; domatia desunt. Inflores-
centia 22-54 cm longa, paniculata et secundiflora; ala-
bastra floralia cylindrica acuminata, aestivatione valvati-
reduplicata; calyx valde redactus 0.3-0.5 mm longus
truncatus; corolla viridis 1.9-2.2 cm longa, lobis
longitudinis corollae ad basim bulbosa; antherae rectae
pilosae inclusae poricidales basaliter subcaudatae dorsifix-
ae prope basim; filamenta tereta ad basim aurei-pilosa.
Capsulae angustissime obconicae 18-23 mm longae disco
conico.

Tree 6-16 m tall. Young branches glabrous to
minutely puberulent, terete; older branches becom¬
ing glabrous, rugulose, terete, grayish to pale brown
and with sparse punctiform to linear lenticels, up to
1.0 cm long. Stipules not seen, probably interpe-
tiolar and free at base (observed from stem scars),
leaving a linear scar 1.0-1.5 mm wide, of the same
color as stem. Leaves petiolate, petiole 4.5-7.0 cm
long, 3-5 mm thick, subterete, glabrous; basally
thickened, not pulvinated; blade 34-62 cm long,
9.5-20.5 cm wide, L/W 3:1 to 3.5:1; lanceolate
to elliptic, acute to obtuse (BA = 35-65°) at base,
acute to attenuate at apex, sometimes tapering to
an acumen 2-3 cm long; blade when fresh dark
green above and dull green below; drying pale brown,
stilf-chartaceous; glabrous above and below, pellucid
punctate (see Fig. 2). Primary and secondary veins
glabrous and prominent below, secondary veins 19-
25 each side; tertiary veins starting subparallel and
openly reticulate in the center, faintly evident above
and fairly evident below; domatia absent. Inflores¬
cences terminal, 22-54 cm long, 9-22 cm wide at
base, laxly paniculate and pyramidal, with decussate
racemoid lateral branches; lateral branches 3-7 pairs,
basal portion of axis not branched, 10-20 cm long.
Rachis terete to decussately obtuse-compressed, ra-
chis and branches glabrous to minutely puberulent;
flowers on distal racemoid branches or on racemoid
cymules. Bracts subtending lateral branches 1-3
mm long, 2-3 mm wide, deltoid; bracteoles sub¬
tending the flowers 0.5-1.0 mm long, 0.7-1.0 mm
wide, deltoid, glabrous. Flowers protandrous, pedi¬
cellate, pedicel 6-9 mm long, glabrous to minutely
puberulent; hypanthium narrowly obconical, 4-6
mm long, 2-4 mm wide, glabrous to minutely golden
hirtellous. Calyx reduced to a wavy margin with
barely distinguishable lobes, 0.3-0.5 mm long and
persistent. Corolla tubular with bulbous base and
small constriction just above, 1.9-2.2 cm long, flow¬
er bud cylindrical, aestivation valvate with contact
zone, color green throughout (rarely reddish with
green lobes), semifleshy when fresh; tube subcylindr¬
ical and striate, 1.4-1.8 cm long, 4-5 mm wide at
base and 2.0-2.5 mm wide at the orifice, glabrous

136

Novon

show n. f v " ld jfi° ra Delprete. -A Habit of inflorescence with mature leaves. -B. Abaxial view of blade

showing 7 C ['?' er bu d showing bulbous base. -D. Anther. -E. Style. -F. Mature capsule

OTmT " ° type: 80 3674 ' SheetS A and B; capsule drawnfr ° m paratype: F. Prieto

Volume 5, Number 2
1995

Delprete

Rustia from Panama and Ecuador

137

or golden hirtellous outside; ascending strigose at
base inside, without a ring of hairs, the remaining
medial and superior zone glabrous or with sparse
hairs; corolla lobes 5, 3-5 mm long, ca. 2.5 mm
wide at base, triangular to ovate, glabrous outside,
glabrous and with minute glandular hairs (hairs up
to 0.5 mm long) at margins inside. Lobes V t to % of
corolla length. Stamens 5, included (only the very
tip exserted), subequal, filaments attached 3.5-5.5
mm from the base of the tube, anthers elongated
and rectilinear, 11-12 mm long, 1.2—1.3 mm wide,
dorsifixed near the base, base subcaudate, with sharp
ridges, papillose-echinate to ascending-strigulose and
descending glandular-pilose at base; dehiscing by two
pores at apex; filaments 6-7 mm long, sparse as¬
cending-pilose and with a tuft of golden pilose hairs
at base. Style exserted, 22-28 mm long, glabrous
and minutely verrucose throughout, style branches
ovate and acute at apex, 1.0-1.3 mm long, stigmatic
surface smooth to minutely papillose. Ovary two-
celled, placentation axile, turbinate to narrowly ob-
conical, glabrous, with many ovules in each locule
horizontally inserted. Immature fruits green to red¬
dish. Capsules very narrowly obconical (see Fig. 2),
apex shortly conical, (14—)18—23 mm long, 4-6
mm wide, black and without lenticels, capsule and
disk black and glabrous, disk obviously exceeding
the calyx; dehiscing loculicidally, disk-septicidal de¬
hiscence absent even in old capsules.

Rustia viridiflora is unique in the genus in having
basally bulbous green flowers and capsules very nar¬
rowly obconical with a conical disk. It is known from
the eastern slopes of the southern Ecuadorian Andes
(Prov. Morona-Santiago), at the western margin of
the Amazon forest.

Paratype. ECUADOR. Morona-Santiago: low hills
W of Rio Chupiangas, 20-22 Nov. 1944, F. Prieto
CHUP-20 (NY [2], US).

Rustia dressleri Delprete, sp. nov. TYPE: Pan¬
ama. Panama: El Llano-Carti Highway, 8-10
km N of El Llano, 31 Aug. 1974, R. Dressier
4703 (holotype, MO). Figure 3.

Arbores vel frutices; stipulae interpetiolares caducae.
Laminae 30-45 cm longae, 9.5-13.0 cm latae, glanduli-
punctatae; domatia desunt vel tricomata pauca brevia
praesentia. Inflorescentia 12-21 cm longa paniculata;
alabastra floralia clavata, aestivatione valvati-reduplicata;
calyx valde redactus ca. 0.5 mm longus truncatus; corolla
alba 1.2-1.4 cm longa lobis l / 2 longitudinis corollae; an-
therae exsertae poricidales concavae basaliter rotundatae
dorsifixae prope basim; filamenta basaliter dilatata ad bas-
im descendens-pilosa. Capsulas non visi.

Shrub or tree (specimen collected from fallen
branches). Young branches grayish, glabrous, te¬

rete; older branches glabrous, rugose, terete, pale
brown-grayish, lenticels absent; leafy branchlets
semisucculent, 7-9 mm wide. Stipules not seen,
interpetiolar, free at base (observed from stem scars),
leaving a linear scar ca. 1 mm wide, of the same
color as stem. Leaves petiolate, petioles 28-32 mm
long, ca. 3.0 mm thick, adaxially concave to flat¬
tened, glabrous; thickened and becoming corky at
base; blade 30-45 cm long, 9.5-13.0 cm wide, L/
W 3:1 to 4:1; narrowly elliptic to oblanceolate,
cuneate (BA = 20-25°) at base, acute and acu¬
minate at apex, the acumen ca. 1 cm long; blade
when fresh shiny grass-green above and pale grayish
green below, subcoriaceous; drying olive-green, semi-
coriaceous; glabrous above and below; evidently pel¬
lucid punctate (see Fig. 3). Primary and second¬
ary veins glabrous and prominent below, secondary
veins 24-30 each side, subparallel; tertiary veins
starting subparallel and openly reticulate in the cen¬
ter, very evident above and faintly evident below;
domatia absent or a tuft of few hairs. Inflorescences
terminal, 12-21 cm long, reduced panicles with
opposite decussate branches; basal branches 6.5-
11.0 cm long, lateral branches 4-5 pairs, basal
portion of axis not branched, 0.9-2.0 cm long.
Rachis decussately compressed and terete at base,
rachis and branches semisucculent, glabrous
throughout and moderately puberulent at nodes;
flowers in opposite cymules on rachis and lateral
branches. Distal bracts subtending lateral branches
2.5-3.0 mm long, 3-4 mm wide, deltoid, glabrous;
bracteoles subtending the flowers 1.0-1.5 mm long,
1-2 mm wide, deltoid, glabrous. Flowers protan-
drous, subsessile or pedicellate, pedicel 0.5-4 mm
long, glabrous; hypanthium obconical, 2.5-3.5 mm
long, 2.0-2.5 mm wide, glabrous. Calyx extremely
reduced, with a wavy margin or with barely distin¬
guishable lobes, ca. 0.5 mm long, sometimes cilio-
late, persistent. Corolla tubular with spreading lobes,
1.2-1.4 cm long; flower bud clavate, ca. 2.5 mm
at base and 3.5-4.0 mm at bulge; aestivation valvate
with contact zone, color white, semifleshy when fresh;
tube 7-8 mm long, 2.5 mm wide at base and 3.5-
4 mm wide at the orifice, glabrous outside, glabrous
inside and with a ring of white-pilose retrorse to
erect hairs, 5-6 mm from the base, at the point of
attachment of the filaments, which is also the base
of the corolla lobes; corolla lobes 5, 5-6 mm long,
ca. 3 mm wide, triangular, semifleshy, glabrous out¬
side, white-strigose at base and margins microscop¬
ically (40 x) papillose inside. Lobes ca. V 2 of corolla
length. Stamens 5, partially exserted because of
narrowly spreading lobes, subequal, filaments at¬
tached 5-6 mm from the base of the tube, anthers
concave, yellow and banana-shaped, 4-5 mm long.

138

Novon

Figure 3. Rustia dressleri Delprete. —A. Habit of inflorescence with mature leaf. —B. Abaxial view of blade
showing pellucid punctation. C. Detail of open flower showing callous base and descending pilose tufts. —D. Flower
bud. (Drawn from holotype: Dressier 4703.)

1.8-2.0 mm wide, dorsifixed near the base, bases
of thecae rounded and subequal, microscopically
(40 x) papillose-echinate throughout, dehiscing by
two pores at apex; filaments ca. 2 mm long, distally
terete and basally flattened, adnate to tube, glabrous
throughout and barbate at base, with a tuft of de¬

scending white-pilose hairs 0.1-0.2 mm long (see
Fig. 3). Style exserted, 13-16 mm long, glabrous,
style branches ovate to narrowly ovate, ca. 1.5 mm
long, stigmatic surface smooth. Ovary two-celled,
placentation axile, obconical, with many ovules in
each locule horizontally inserted. Capsule not seen.

Volume 5, Number 2
1995

Delprete

Rustia from Panama and Ecuador

139

Rustia dressleri is very closely related to R.
occidentalis (Bentham) Hemsley in having fleshy
clavate buds, but differs in having white flowers
(flesh-red to purple in R. occidentalis), filaments
with basal tuft of descending white-pilose hairs (pu-
berulent in R. occidentalis), larger inflorescences
(up to 11 cm long in R. occidentalis), young bran-
chlets thick and semisucculent, the whole plant gla¬
brous, leaf blades 30-45 cm long (exceptionally up
to 30 cm in R. occidentalis), and with 24-30
secondary veins each side (11-16 in R. occiden¬
talis).

In his treatment of Rubiaceae for the Flora of
Panama, Dwyer (1980) erroneously cited the para-
type of Rustia dressleri (Dressier 4749) as R.
panamensis Dwyer. The latter was subsequently
reduced to synonymy under the new combination
R. costaricensis (Standley) Lorence (Burger & Tay¬
lor, 1993).

Rustia dressleri differs from R. costaricensis in
its inflorescence 12-21 cm long (vs. 20-33 cm long
in R. costaricensis), twice compound (vs. thrice
compound in R. costaricensis), with 4-5 pairs of
lateral branches (vs. 8-11 pairs in R. costaricensis),
flower bud clavate (vs. pointed in R. costaricensis),
corolla tubular (vs. cupular in R. costaricensis) and
12-14 mm long (vs. 3-4 mm long in R. costari¬
censis).

Known only from two collections, this species was
first encountered by Robert L. Dressier as fallen
branches along a stream. Dressier is a well-known
orchidologist, and I am pleased to honor him with
this eponym.

Paratype. PANAMA. Panama: El Llano-Carti High¬
way, 10-12 km N of El Llano, 12 Sep. 1974, R. Dressier
4749 (MO).

Acknowledgments. I am grateful to the directors
and staff of the following institutions for the loan of
material and for help in fieldwork: AAU, B, BR,
CAS, CHOCO, COL, CR, DAV, DS, F, G, GB, GH,
JBGP, JEPS, K, LL, MO, NY, P, QCA, QCNE, S,
TEX, UC, US. I thank Lennart Andersson, John
Dwyer, and Charlotte Taylor for the stimuli toward
my research. I especially thank Billie Turner and
Guy Nesom for reviewing the manuscript, and David
Lorence for helpful comments. Guy Nesom also
helped me with the Latin descriptions. The illustra¬
tions are provided by the author.

Literature Cited

Burger, W. & C. M. Taylor. 1993. Rubiaceae. In:

Flora Costaricensis. Fieldiana, Bot. 33: 306-307.
Dwyer, J. D. 1980. Rubiaceae. In: Flora of Panama.

Ann. Missouri Bot. Gard. 67: 483-485.

Hooker, J. D. 1873. Rubiaceae. In: G. Bentham & J.

D. Hooker, Genera Plantarum 2: 7-151.

Klotzsch, J. F. 1846. In: F. G. Hayne, Getreue Dar-
stellung und Beschreibung der in der Arzneykunde
Gebraiichlichen Gewachse. 14: 14-15. Nicolai, Ber¬
lin.

Robbrecht, E. 1988. Tropical Woody Rubiaceae. Opera
Bot. Belgica 1: 1-271.

Simpson, R. D. 1976. Studies in neotropical Rubiaceae.
Rustia. Phytologia 33: 4-8.

New Orchid Species from Costa Rica

Robert L. Dressier

Missouri Botanical Garden. Mailing address: Rt. 2, Box 565C,
Micanopy, Florida 32667, U.S.A.

Abstract. Five new orchid species from Costa
Rica are described and illustrated from living or
liquid-preserved material: Encyclia ortizii, Malaxis
monsviridis, M. talamancana, Sobralia dissimilis,
and S. doremiliae. The orthography of Sobralia
“corazoi ” is discussed, and S. carazoi is thought
to be correct.

Costa Rica has a long tradition of resident nat¬
uralists and orchidophiles collecting and pressing
plants for study. The orchid flora of Costa Rica is
thus much better sampled than those of neighboring
countries. Even so, the preparation of the orchid
treatment for Manual de la Flora de Costa Rica
continues to reveal unnamed species, especially
among terrestrials and smaller-flowered epiphytes.
Five of these are here named and described.

Encyclia ortizii Dressier, sp. nov. TYPE: Costa
Rica. Prov. Alajuela: Reserva San Ramon,
aprox. 30 km NNO de San Ramon, 8, 9 die.
1984, R. L. Dressier y Biologta 350 no. 288
(holotype, USJ). Figure 1.

Pseudobulbi pyriforme-ovoidei unifoliati; folia oblan-
ceolato-oblonga; sepala oblongo-oblanceolata acuta; petala
anguste oblanceolata; labello unguiculato trilobato lobi
laterales parvi oblongi lobo intermedio subquadrato-ovato
callo alto trigono carinato ornato.

Epiphytic, caespitose; pseudobulbs unifoliate,
narrowly pyriform-ovate, somewhat flattened, 3.7
x 0.8-1.2 cm; leaves oblanceolate-oblong, nar¬
rowed basally, obtuse to subacute, 8.2-10 x 2.3-
2.6 cm; inflorescence racemose, 12-24 cm, bracts
deltoid, acute, 3-4 x 1.8-2 mm; pedicel with ovary
10-12 mm; sepals oblong-oblanceolate, acute, 9 x
2.8-3 mm; petals narrowly oblanceolate, 8x2
mm; lip unguiculate, claw ca. 4 mm, basally adnate
to column, blade 3-lobed, lateral lobes oblong, por-
rect, clasping column apex, 2 x 1.3 mm; midlobe
at right angle to column, ovate, 4 x 3.8 mm, callus
prominent, trigonous, 1.5 x 1.5 mm, with laminar
keel ca. 1.3 mm high; column ca. 7 mm, dorsal
profile concave, midtooth ovate-elliptic, 1.7 x 0.8
mm, surpassing anther and lateral teeth, lateral teeth
deltoid, 0.8 x 0.6 mm.

Encyclia ortizii is a member of subgenus Os-
mophyta and is closely allied to E. campylostalix
(Reichenbach f.) Schlechter, E. fortunae Dressier,
and E. glauca (Knowles & Westcott) Dressier &
Pollard. The relatively slender column resembles
that of E. campylostalix, while the sharply defined,
rounded lateral lobes of the lip resemble those of E.
glauca and E. fortunae. It is quite distinctive in the
high, keeled callus and the orientation of the lip.
The epithet honors Rodolfo Ortiz, director of the
San Ramon nature reserve, in whose company the
type material was collected.

Malaxis monsviridis Dressier, sp. nov. TYPE:
Costa Rica. Puntarenas: Monteverde Reserve,
near Alajuela boundary, Cerros Centinelas, elev.
1550-1600 m, 10°18'N, 84°47'W, 19 Aug.
1984, M. H. Grayum & P. Sleeper 3842
(holotype, MO). Figure 2A, B.

Herba perennis erecta; pseudobulbis epigaeis ovato-
conicis; folia duo late ovata breviter acuminata; inflores-
centia subumbellata; bracteae deltoideo-lanceolatae; se¬
pala oblongo-ovata obtusa; petala filiformia; labello
subquadrato profunde hastato apice subquadrato trilob-
ulato profunde excavato.

Pseudobulbs above ground, 1.5-3 x 0.6-1.2
cm, approximate or up to 3.5 cm apart on rhizome.
Leaves 2, petioles 5-6 cm, blades broadly ovate,
short-acuminate, 5-7.5 x 3.2-5 cm. Inflorescence
subumbellate, scape 10-15 cm, floral bracts deltoid-
lanceolate, to 1.5 mm; pedicel with ovary 6.5-10
mm; sepals oblong-ovate, obtuse, dorsal sepal 3-3.5
x 1.3-1.6 mm, lateral sepals 2.7-3 x 2mm; petals
filiform, 2-3 mm; lip hastate, subquadrate, 2.5-3
x 2-2.6 mm, basal auricles subdeltoid, retrorse,
prominent; cavities 2, about half length of blade,
cavity apically projecting under blade of lip; apex
of lip 3-lobulate, lobules subequal, lateral lobes
rounded, midlobule rounded or deltoid; column about
1 mm long; color: pale green or lip brownish green.
Flowering July through September.

Malaxis monsviridis may be recognized by the
prominently hastate lip with short, deep cavities and
wide, trilobulate apex. Both M. carpinterae
(Schlechter) Ames and M. hastilabia (Reichenbach

Novon 5: 140-145. 1995.

Volume 5, Number 2
1995

Dressier

Orchids from Costa Rica

141

view.

B

no. 288). —A. Flower. —B. Column and lip, lateral

lateral view.

142

Novon

f.) Kuntze have lips with retrorse lobules and tri-
lobulate apices, but the apices are much narrower
and the cavity either narrower or longer.

Paratypes. COSTA RICA. All from the area of
Monteverde: J. T. Atwood 89-319 (SEL), K. Barrin¬
ger,, E. Christenson & B. Perez 4168 (F), V. J. Dryer
699 (F), W. A. Haber & E. Bello C. 2423 (MO), W.
Haber & W. Zuchowski 9315 (MO), ft. 0. Lawton 1329
(F).

Malaxis talamancana Dressier, sp. nov. TYPE:
Costa Rica. San Jose: Villa Mills, Cordillera de
Talamanca, ca. 3000 m, 28 mar. 1993, R. L.
Dressier & D. E. Mora s.n. (holotype, USJ).
Figure 2C, D.

Herba perennis, erecta; pseudobulbis epigaeis, conicis;
folia duo deltoideo-ovata cuneata vel subcordata breviter
acuminata; inilorescentia subumbellata; bracteae delto-
ideo-lanceolateae; sepala ovata vel elliptica obtusa; petala
sublinearia; labello deltoideo-ovato hastato ad apicem min¬
ute tridentato auricula retrorsa deltoidea vel denticulata.

Terrestrial, pseudobulbs above ground, conic, 2-
4.5 x 0.8-2 cm, to 2.5 cm apart on rhizome.
Leaves 2, petioles 5-7 cm; blades deltoid-ovate,
cuneate to subcordate, apices short-acuminate, 5.5-
8 x 3-4.2 cm. Inflorescence subumbellate, scape
15-20 cm, bracts deltoid-lanceolate, ca. 2 x 1 mm,
ovary with pedicel 17-21 mm; sepals ovate to el¬
liptic, narrowly obtuse, 3-6 X 1.5-2.3 mm; petals
sublinear, 3-4 x 0.6 mm; lip 3-5 x 2-2.5 mm
(in natural position), hastate, deltoid-ovate, apex mi¬
nutely 3-dentate, lateral lobules folded upward, au¬
ricles retrorse, rounded, deltoid or denticulate, 0.8-
1 x 0.8 mm; basal cavity deep, 1.5-2 mm, with
rounded median keel, distal margin emergent.

Malaxis talamancana may be recognized by the
deltoid-ovate, hastate lip minutely 3-dentate api-
cally. Superficially, the flowers are similar to those
of M. brachyrrhynchos (Reichenbach f.) Ames, but
that species has subterranean corms and the details
of the lip are quite distinct.

Paratypes. COSTA RICA. San Jose: trail from Ca¬
naan to Cerro Chirripo, between 2000 and 3000 m elev.,
20 & 25 Aug. 1971, W. C. Burger & L. Gomez P.
8358 (F). Limon: Cordillera de Talamanca, Atlantic
slope, between Rio Terbi and the Rio Sini, elev. 2400-
2750 m, 13 Sep. 1984, G. Davidse , G. Herrera Ch. &
M. H. Grayum 29048 (MO).

Sobralia dissimilis Dressier, sp. nov. TYPE: Cos¬
ta Rica. Puntarenas: Monteverde Reserve, TV
towers at 1750 m elev., 26 June 1989, J. T.
Atwood 89-243 (holotype, MO; isotypes, CR,
FLAS, SEL). Figure 3.

Herba epiphytica; folia elliptica; flores e bracteis erectis
imbricantibus nonullis erumpentis; sepala ovato-oblonga
obtusa mucronata; petala ovato-oblonga; labellum ovatum
margine fortiter undulatum; discus carinis biformis un-
decim ornatus.

Epiphytic, caespitose, stems stout, 35-75 cm;
sheaths glabrous, verruculose. Leaves several, ellip¬
tic, acuminate, 13-15 x 4.5-5.2 cm, thin, not
markedly corrugated. Inflorescence bract cluster el¬
lipsoid, 5-6 cm, loose, glabrous, with 2 flowers at
once; sepals ovate-oblong, obtuse, mucronate, 3.1-
3.3 X 1.4 cm, lateral sepals basally connate for 6-
7 mm; petals oblong-ovate, 3.1 x 1.3 cm; lip ovate,
about 3 x 2.3 cm, tube about 1 cm, then flaring,
exposing column, with 2 fleshy calli about 8 mm
from base, with 11 keels from near center, outer 6
keels high, curved, not undulate, outermost short,
next intermediate, then reaching to within 5 mm of
apex, inner 5 keels strongly undulate, reaching 2-
5 mm from apex; column 2.5 cm, basally slender,
column wings narrowly deltoid, 4.5 x 3 mm, an-
trorse.

The epithet dissimilis , or different, refers both
to the fact that the species is strikingly distinct from
all others and to the two different types of keel on
the Up. Sobralia dissimilis is clearly a close ally of
S. undatocarinata C. Schweinfurth, of which there
is an excellent photograph in Orquidea 5: 209,
1975 (as S. mucronata Ames & C. Schweinfurth;
Horich, 1975). Sobralia dissimilis is quite distinct
in the shorter, rounded limb of the lip (not pandurate
or deeply bifid), in the wider, antrorse column wings,
and in the two sharply different types of keel on the
lip.

Sobralia doremiliae Dressier, sp. nov. TYPE:
Costa Rica. Cartago: slopes and steep roadside
banks along the Rio Grande de Orosi about 11
km SSE of bridge at Tapanti, elev. about 1500
m, 10 July 1977, R. L. Wilbur, F. Almeda
<fe 7. Daniel 22470 (holotype, DUKE). Figure
4.

Herba epiphytica; folia elliptica; flores e bracteis brev-
issimis nonnullis erumpente; sepala oblanceolata mucron¬
ata; petala oblongo-elliptica obtusa; labellum subquadrato-
obovatum retusum; discus basi carinis humilis quindecim
ornatus.

Epiphytic or on steep slopes, caespitose, stems
10-25 cm; sheaths glabrous or slightly verruculose
and scurfy. Leaves eUiptic or ovate-eUiptic, short-
acuminate, 4.5-9.5 x 1.7-3.4 cm. Inflorescence
subtended by short, wide, divergent bracts, the floral
bracts wide, retuse, scarcely emergent from the
inflorescence bracts; flowers subcampanulate, white

Volume 5, Number 2
1995

Dressier

Orchids from Costa Rica

143

Figure 3. Sobralia dissimilis Dressier (Atwood 89-243). —A. Habit. —B. Flower. —C. Lip, lateral view. —D.
Lip, natural position. —E. Perianth, flattened. —F. Column, ventral view. G. Column, lateral view.

144

Novon

Figure 4. Sobralia doremiliae Dressier (Dressier & Mora s.n.). —A. Inflorescence, showing short bracts. B.
Perianth, flattened. —C. Column.

or pale green, base of lip orange or orange-brown;
sepals united for 3 mm, oblanceolate or elliptic-
oblanceolate, mucronate, 17-20 x 5-6 mm; petals
oblong-elliptic, obtuse, 16-18 x 4-5 mm; lip ba-
sally cuneate, subquadrate-obovate, refuse, 17-18
x 11-12 mm, margins lacerate-dentate above mid¬
dle, base with about 15 keels, most shorter than
column, central 4 subequal to column; column 8-
9 mm, wings falcate, surpassing anther; capsule
4.5-5 x 0.8-1.1 mm.

The epithet doremiliae honors Dora Emilia Mora
de Retana, director of the Lankester Garden, with
whom we collected and photographed this species
near the type locality. The specimen chosen as type
includes much more flowering material than our own
scanty material (at USJ). I have purposely omitted
the “a” of Dora to avoid a seeming diphthong that
could lead to mispronunciation (the alternative being
hyphenation).

Sobralia doremiliae ranges at least from the
San Ramon area of Costa Rica to Cerro Colorado
in Panama. It may be recognized by the small,
subcampanulate flower and the form of the inflo¬
rescence. It somewhat resembles S. carazoi Lankes¬
ter & Ames (see note below); Smith 1109 is a

mixture of these species. Sobralia doremiliae, how¬
ever, is smaller in all parts, and the ornamentation
of the lip is quite different, as are the bracts of the
inflorescence. The floral bracts of S. carazoi are
elliptic and densely scurfy.

The inflorescence of S. doremiliae resembles that
of S. lindleyana Reichenbach f. and S. macro-
phylla Reichenbach f. in that the sheaths of the
inflorescence bracts and the uppermost leaves are
markedly infundibuliform, the bracts, themselves,
are short and wide, and the floral bracts are quite
short, remain green, and do not project as an ellip¬
soid bract cluster (Fig. 4A).

Paratypes. COSTA RICA. Alajuela: Los Angeles de
San Ramon, 21 VII 1932, A. M. Brenes 16124 (F), 2
VIII 1932, A. M. Brenes 16246 (F); La Pena de Zarcero,
alt. 1450 m, 22 Aug. 1938, A. Smith 1109 (AMES,
SEL). Cartago: weedy roadsides and steep heavily veg¬
etated banks along the ICE road 9 km NW of the Tapanti
Dam or 9 km SE of the Tapanti Bridge across the Rio
Grande de Orosi, alt. ca. 1300 m, H. L. Wilbur 30765
(DUKE). San Jose: La Hondura de San Jose, alt. 1300-
1700 m, 16, 19 Mar. 1924, P. C. Standley 377IS
(AMES), alt. 1300 m, 15 agosto 1933, M. Valerio 798
(F). PANAMA. Chiriqui: Cerro Colorado, ca. 50 km N
of San Felix, elev. ca. 1400 m, 17 Aug. 1975, R- L.
Dressier 5092 (FLAS), elev. 1400-1500 m, 20 Sep.
1977, R. L. Dressier 5708 (FLAS, PMA).

Volume 5, Number 2
1995

Dressier

Orchids from Costa Rica

145

Orthography of Sobralia carazoi

Sobralia corazoi was published by Lankester and
Ames (Ames, 1924). The surname Corazo, if such
exists, is not common in Costa Rica, but Carazo is
well known. Rafael Lucas Rodriguez investigated this
in some detail (letter to L. A. Garay, 22 Dec. 1978,
Microfiche of Ames Herbarium, Inter Documenta¬
tion Co.) and found that the person honored by the
epithet was Enrique Carazo Quezada, concluding
that the spelling “ corazoi ” was an error. That the
error had been recognized at the Ames Orchid Her¬
barium is suggested by specimens clearly labeled as
“ carazoi ” by Schweinfurth in 1938.

Acknowledgments. I am indebted to Wendy
Zomlefer and Willow Zuchowski for the preparation
of the drawings used here. Recent fieldwork has
been supported by the National Science Foundation
and the Office of Forestry, Environment and Natural
Resources, Bureau of Science and Technology, of
the U.S. Agency for International Development un¬
der NSF Grant DEB-9200812 (to John Atwood).

Literature Cited

Ames, 0. 1924. Additions to the orchid flora of tropical
America. Sched. Orch. 7: 1-36.

Horich, C. K. 1975. Las sobralias, encanto lindo pero
fugaz. Orquidea 5: 204-212.

Two New Species of Paspalum (Poaceae: Paniceae) from Brazil

Tarciso S. Filgueiras

Reserva Ecologica do IBGE, Caixa Postal 08770, Brasilia, DF 70314-970, Brazil

Gerrit Davidse

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. Two narrow endemics, Paspalum cam-
pinarum (group Corcovadensia) from Amazonas,
Brazil, and P. zuloagae (group Quadrifaria) from
Minas Gerais, Brazil, are described, illustrated, and
compared with putatively related species. Paspalum
zuloagae is easily distinguished from all members
of the Quadrifaria group by its prominent sheath
auricles, and P. campinarum is distinguished from
South American species of the Corcovadensia group
by its upper glume %-% as long as the upper floret
and the possession of axillary inflorescences.

While studying Poaceae collections from Brazil,
some specimens of Paspalum L. were encountered
that could not be assigned to any species in that
genus and are described here as new. Paspalum is
a largely American genus of approximately 330
species. As the largest grass genus in Brazil, many
of its species are ecological dominants of savannas
and grasslands. The monograph by Chase (1929)
of North American species, and her unpublished
work of the South American species [accessible from
the Hitchcock and Chase Library in the Botany
Department of the Smithsonian Institution], are es¬
sential references for studies in the genus. Chase
proposed a largely informal infrageneric classifica¬
tion in these two works, and it has been extensively
used by many contemporary agrostologists.

Paspalum campinarum Filgueiras & Davidse,
sp. nov. TYPE: Brazil. Amazonas: Transama¬
zon Highway, 53 km W of Aripuana River,
“campina” region, common but not abundant
in open campina of white sandy soil, 29 June
1979, C. E. Calderon, 0. P. Monteiro & J.
Guedes 2715 (holotype, INPA; isotypes, MO,
US). Figure 1.

Ab omnibus speciebus austroamericanis Corcovaden-
siae inflorescentia terminali ac axillari, gluma superiore
Vs-'A longitudinis spiculae differt.

Caespitose perennials; rhizomes short, knotty.
Culms 40-75 cm long, erect to semi-decumbent at
the base, branched just above the lowest nodes but

occasionally also with flowering branches from the
middle or upper nodes, with 6-8 elongate inter-
nodes; basal internodes pilose, the upper ones gla¬
brous; basal nodes pilose to glabrescent, the upper
ones glabrous. Leaves cauline only; basal bladeless
sheaths densely pilose, the upper ones glabrescent
to glabrous, striate, one margin long ciliate, the other
glabrous; auricles usually 2, 0.5-1.5 mm long, de¬
veloped primarily on the upper sheaths; ligule 0.2-
0.3 mm long, membranous, glabrous; blades 4-9.9
cm long, 9-19 mm wide, lanceolate, slightly cordate
at the base, glabrous on both surfaces, strongly
striate, especially on the adaxial surface, the margins
scabrous, usually ciliate at the base and pubescent
near the margins of the very short pseudopetiole
with tuberculate, white hairs 2-5 mm long. Inflo¬
rescences 10.5-13 cm long when fully exserted,
terminal and axillary, with 5-25 ascending racemes
in terminal inflorescences, fewer in axillary inflo¬
rescences; common rachis 9-12 cm long, glabrous;
racemes 1.5-6.5 cm long, terminating in a spikelet;
raceme rachis 0.3-0.6 mm wide, triquetrous, with
a few scattered, tuberculate, white hairs 4-8 mm
long. Pedicels 0.5-1 mm long, of unequal length,
paired, the apex discoidal. Spikelets 2.3-2.9 mm
long, 1-1.2 mm wide, elliptical in outline, paired,
pubescent, tan-brown; lower glume absent; upper
glume %-% as long as the spikelet, 3-nerved, the
surface appressed pilose, the margins densely ciliate;
lower floret sterile, consisting only of a lower lemma;
lower lemma as long as the spikelet, 3-nerved, the
surface glabrous, the margins ciliate; lower palea
absent; upper floret bisexual, minutely longitudinally
striate, pale, glabrous; upper lemma 5-nerved, gla¬
brous, minutely longitudinally striate, the germina¬
tion flap evident at the base; upper palea similar to
the upper lemma in size and texture, 2-nerved,
minutely longitudinally striate; lodicules 2, condu-
plicate; stamens 3, the anthers 2-2.2 mm long, tan
to brownish; ovary 0.3-0.4 mm long; styles 2, sep¬
arate; stigmas 2, plumose, tan. Caryopsis not found.

In the following differentiating characters—pe¬
rennial fife form, relatively broad leaf blades, pan-

Novon 5: 146-151. 1995.

Volume 5, Number 2
1995

Filgueiras & Davidse
Paspalum from Brazil

147

Figure 1. Paspalum campinarum Filgueiras & Davidse, illustrated from the type collection (Calderon et al 2715).
-A. Habit. -B. Terminal portion of a raceme. -C. Dorsal (left) and ventral (nght) views of a secondary sp,kelet.
D. Dorsal (left) and ventral (left) views of a primary spikelet.

148

Novon

icles with several to many racemes, a narrow rachis,
paired, undifferentiated plano-convex spikelets, pale
upper florets, and minutely longitudinally striate tex¬
ture of the upper floret—P. campinarum agrees
with those of group Corcovadensia , of which three
other species occur in Brazil: P. corcovadense Rad-
di, P. loefgrenii Ekman, and P. mandiocanum Tri-
nius. It is also worth noting that in both P. corco¬
vadense and P. mandiocanum the upper glume is
distinctly shorter than the spikelet, but not nearly
as short as in P. campinarum. We, therefore, be¬
lieve that its true affinity is with this species group.

We also compared Paspalum campinarum to P.
inaequivalve Raddi because the latter is a perennial
species with relatively short, broad leaf blades, ter¬
minal and axillary panicles with several to many
racemes, a narrow rachis, paired, undifferentiated
spikelets, pale upper florets, and, especially, a short¬
ened upper glume that exposes the upper floret.
Chase (unpublished) included two species in the group
Inaequivalvia: Paspalum inaequivalve Raddi and
P. microstachyum J. S. Presl. The latter is an annual
with long-pedicellate, irregularly arranged spikelets
with the upper glume as long as the lower lemma,
and is clearly not closely related to P. campinarum
or P. inaequivalve. Paspalum inaequivalve differs
from P. campinarum in its weak, trailing to de¬
cumbent culms, glabrous, noncordate basal leaves,
3 to 15 racemes with glabrous rachis and, especially,
a smooth, shining upper floret that is not minutely
longitudinally striate.

It is perhaps unwise to differentiate species groups
in Paspalum primarily on the basis of relative length
of the upper glume, since reduction of the upper
glume seems to have evolved repeatedly in obviously
different groups of Paspalum , for example: P. in¬
aequivalve (group Inaequivalvia), P. usteri Hackel
[sect. Anachrysis (Nees) Pilger], P. marmoratum
Kuhlmann (group Plicatula ), P. delicatum Swallen
(group Parviflora), and P. decumbens Swartz (group
Decumbentes). Reduction has been carried to the
extreme in species without any upper glume, as in:
P. gardnerianum Nees (sect. Eremachyrion Doell,
group Gardneriana), P. nudatum Luces (group
Parviflora ), and P. pulchellum Kunth (group Rei-
maria).

Paspalum campinarum is known only from a
single collection from the white sandy “campina”
in the Brazilian Amazonia, where it was found to be
“common but not abundant.” The specific epithet
alludes to this campinas habitat. It is notable that
this is the same locality in which a highly distinctive
grass genus, Arundoclaytonia Davidse & Ellis, was
discovered at the same time (Davidse & Ellis, 1987).
We assume that like A. dissimilis Davidse & Ellis,

Paspalum campinarum is probably restricted to the
white-sand soils of Amazonian campinas.

The four Brazilian species of the Corcovadensia
group may be distinguished as follows:

la. Panicle rachis 15-25 cm long; spikelets 2.5-
2.9 mm long; upper glume as long as the spike-

let; lower glume frequently present .. P. loefgrenii

lb. Panicle rachis usually less than 15 cm long;
spikelets 1.8-2.5 mm long; upper glume short¬
er than the upper floret; lower glume always
absent.

2a. Upper glume ‘/m-flo as long as upper

floret .P. campinarum

2b. Upper glume as long as the upper floret.

3a. Spikelets 1.4-1.5 mm wide, elliptic-
oblanceolate; racemes relatively thick;
culms frequently decumbent at the
base and rooting at the lower nodes

.P. mandiocanum

3b. Spikelets 0.9-1.0 mm wide, broadly
elliptical; racemes relatively slender;
culms generally erect and not rooting
at the lower nodes.P. corcovadense

Paspalum zuloagae Davidse & Filgueiras, sp.
nov. TYPE: Brazil. Minas Gerais: Serra da
Gramma, summit, mossy rock, 19-25 Apr.
1925, A. Chase 9578 (holotype, US; isotype,
MO). Figure 2.

Ab omnibus speciebus Quadrifariae auriculis conspi-
cuis, 5-13 cm longis (auriculae ligulae saepe connatae)
differt. Paspalo coryphaeo Trinius simile sed rhizoma-
tibus nullis, laminis 5-17 cm longis, basin versus atten-
uatis, inflorescentia 8-17 cm longa, 12-27 racemis dif¬
fert.

Perennial, caespitose; rhizomes weakly developed
or absent. Culms 65-90 cm long, erect, frequently
branched at the base, sometimes with aggregate
branches from the middle nodes; nodes glabrous;
internodes glabrous. Leaves mostly basal, but old
culms without basal leaves; basal leaves with im¬
bricate sheaths; sheaths strongly striate, glabrous
on the back, often purplish, the margins long-ciliate;
auricles 2, 5-13 mm long, conspicuous, glabrous
to sparsely long-pilose; ligule 4-11 cm long, mem¬
branous, mostly adnate with the auricles, but dis¬
tinguishable by its membranous texture; ligular area
glabrous, rarely with a few yellowish hairs 1-2 mm
long; blades 5-17 cm long, 6-12 mm wide, linear-
lanceolate, attenuate at the base, acuminate at the
apex, glaucous, glabrous on both surfaces or with a
few scattered, whitish hairs on the adaxial surface,
the margins prominently ciliate with tuberculate-
based, whitish or slightly yellowish hairs 3-5 mm

Volume 5, Number 2
1995

Filgueiras & Davidse
Paspalum from Brazil

149

Figure 2. Paspalum zuloagae Davidse & Filgueiras, illustrated from the type coUection (Chase 9578). —A. Habit.
—B. Ligular area of a leaf. —C. Portion of a raceme with paired spikelets. D. Dorsal (left) and ventral (right)
views of secondary spikelet. —E. Dorsal (left) and ventral (right) views of a primary spikelet.

long, the cilia less frequent in the uppermost leaves;
epidermal cells visible on the abaxial surface of
blades, especially on the older leaves. Inflorescence
8-17 cm long, terminal, with 12-27 ascending ra¬

cemes; common rachis 6.5-14 cm long, glabrous;
racemes 2-12 cm long, with a tuft of whitish or
slightly yellowish hairs at the base, decreasing in
size toward the apex, terminating in a spikelet; ra-

150

Novon

ceme rachis ca. 0.2 mm wide, glabrous. Pedicels
0.3-1 mm long, of unequal length, paired, glabrous,
waxy, the apex discoidal. Spikelets 2-2.5 mm long,
0.8-1 mm wide, narrowly elliptic in outline, paired,
glabrescent to puberulent, tan to yellowish; lower
glume absent; upper glume %-% as long as the
spikelet, 3-nerved, glabrescent to puberulent; lower
floret sterile, consisting only of a lower lemma; lower
lemma as long as the spikelet, 3-nerved, glabrous
to puberulent; lower palea absent; upper floret bi¬
sexual, smooth, pale, glabrous; upper lemma
5-nerved, glabrous, longitudinally striate; upper pa¬
lea similar to the upper lemma in texture and color,
2-nerved, longitudinally striate; lodicules 2, con-
duplicate; stamens 3, the anthers 1-1.5 mm long,
deep purple; ovary 0.4-0.5 mm long; styles 2, sep¬
arate; stigmas 2, plumose, purplish. Caryopsis not
found.

This new species belongs to the informal Quad-
rifaria group of Chase by virtue of its perennial life
form, terminal panicles with numerous racemes, nar¬
row raceme rachis, small, paired, undifferentiated,
tan to green, elliptical spikelets, and pale upper
floret. Paspalum zuloagae is distinguishable from
all the other species of group Quadrifaria by the
presence of two conspicuous auricles on either side
of the sheath apex. The auricles are mostly her¬
baceous in texture with well-developed veins that
are continuous with those of the sheath. This grades
into a membranous apex near the tip of the auricle.
In fully developed leaves from the middle nodes of
the culms, the auricles are almost completely adnate
with the equally long membranous ligule. In basal
leaves and leaves from the upper nodes, the auricle
may be considerably larger than the ligule and the
upper half of the auricle is therefore free. Both
auricles and ligule are typically greatly reduced in
the flag leaf. The delicate connection between the
ligule and auricle is readily torn so that the ligule
and auricles often seem superficially separate in
older leaves.

The spikelets are slightly dimorphic in that the
upper glume is shorter in the secondary or short-
pedicellate spikelets compared to those of the pri¬
mary or long-pedicellate spikelets.

The specific epithet honors Fernando O. Zuloaga,
prominent Argentinean agrostologist, long-time
friend, and valued collaborator of both authors. The
collections Chase 9562 and 9578 were cited in the
index to Chase’s unpublished South American manu¬
script as the basis for the establishment of a new
species, “ Paspalum auritum C., ined.” However,
no description was provided and no annotation on
the labels was made.

Paratypes. BRAZIL. Minas Gerais: Serra da Gram-
ma, open summit, 19-25 Apr. 1925, A. Chase 9562
(MO, US 2 sheets); Serra do Espinhajo, ca. 20 km E of
Diamantina, elev. 900 m, cerrado, culms to ca. 1.5 m
tall, 22 Mar. 1970, H. S. Irwin, S. F. da Fonseca, R.
Souza, R. Reis dos Santos & J. Ramos 28072 (UB).

The following key is provided to distinguish all
the species that we include in group Quadrifaria :

la. Leaf sheaths with 2 conspicuous auricles 5-13

mm long . P. zuloagae

lb. Leaf sheaths without auricles or, if present,
inconspicuous and 2 mm long or less.

2a. Raceme rachis with ciliate hairs 1-3 mm
long; cauline leaf blades with the basal 14—

14 strongly narrowed, folded, and pseudo-
petiolate.

3a. Leaf blades 4-8 mm wide .

. P. quadrifarium

3b. Leaf blades 12-15 mm wide .

. P. brunneum

2b. Raceme rachis glabrous or scaberulous, long
hairs restricted to the base and axils of the
racemes, rarely the rachis shortly pubes¬
cent; cauline leaf blades rounded to nar¬
rowed but not pseudopetiolate.

4a. Basal nodes, internodes, and leaf
sheaths densely villous; sheaths
strongly keeled; spikelets glabrous ..

. P. dasytrichium

4b. Basal nodes, internodes, and leaf
sheaths glabrous to pilose, but never
densely villous; sheaths slightly keeled
to weakly rounded; spikelets pubes¬
cent . P. coryphaeum

Paspalum brunneum Mez, Repert. Spec. Nov. Reg-
ni Veg. 15: 74. 1917. TYPE: Brazil. Guana-
bara: Glaziou 9050 (lectotype, designated by
Smith et al. (1982), US).

Distribution: Argentina, Brazil, Paraguay, Uru¬
guay.

Paspalum coryphaeum Trinius, Gram. Panic. 114.
1826. TYPE: Brazil. In humidis prope Ytu,
Langsdorff s.n. (holotype, LE not seen).

Paspalum chapadense Swallen, Phytologia 14: 363.
1967. TYPE: Brazil. Maranhao: collected in cha-
pada between Caxias and Barro do Corda, 18-26
Feb. 1934, Swallen 3508 (holotype, US; isotype,
K not seen).

Paspalum familiare Steudel, Syn. PI. Glumac. 1: 24.
1853. TYPE: Venezuela. Sucre: Funck & Schlim
228 (holotype, P?, fragments, US).

Paspalum indutum Luces, J. Wash. Acad. Sci. 32: 162,
f. 6. 1942. Syn. nov. TYPE: Venezuela. Merida:
Pozo Hondo, near Egido, 960 m, Apr. 1940, Ser-
gent 37 (holotype, VEN, fragments, MO, US).
Paspalum pruinosum Trinius, Sp. Gram. 3: t. 272. 1836.
TYPE: Brazil. Pr. Cachoeira, Anon. s.n. (LE not
seen).

Volume 5, Number 2
1995

Filgueiras & Davidse
Paspalum from Brazil

151

Paspalum violascens Mez, Repert. Spec. Nov. Regni
Veg. 15: 73. 1917. TYPE: Trinidad: in loco ac-
curatione haud indicata, Broadway 2175 (holotype,
B?, fragment, US; isotype, US).

Distribution: Argentina, Belize, Brazil, Colombia,
Costa Rica, Guianas, Honduras, Mexico, Panama,
Trinidad, Venezuela.

Paspalum dasytrichium Dusen ex Swallen, Phy-
tologia 14: 363. 1967. TYPE: Brazil. Parana:
Villa Velha, collected among rocks, 21 Oct.
1914, Dusen 15700 (holotype, US).

Paspalum dasytrichium var. dasytrichium.

Paspalum dasytrichium var. glabrum Lyman B. Smith
& D. C. Wasshausen, Bradea 2: 247. 1978. TYPE:
Brazil. Santa Catarina: Mun. Lajes, moist ground in
shade, woods borders, Coxilha Rica, 6 Jan. 1946,
Swallen 8173 (holotype, US).

Distribution: Brazil (Parana, Santa Catarina, Sao
Paulo).

Paspalum quadrifarium Lamarck, Tabl. Encycl. 1:
176. 1791. TYPE: Argentina. Buenos Aires,
Commerson s.n. (lectotype, designated by Par-
odi (1937), P not seen).

Paspalum ferrugineum Trinius, Sp. Gram. 2: 136. 1829.
TYPE: Chile, Lindley s.n. (syntype, LE not seen).
Uruguay, Otto s.n. (syntype, fragment, US).
Paspalum quadrifarium var. ferrugineum (Trinius) Her-
ter, Anales Mus. Hist. Nat. Montevideo, ser. 2, 3:
55. 1929.

Distribution: Argentina, Brazil, Paraguay, Uru¬
guay.

Paspalum zuloagae Davidse & Filgueiras.

Distribution: Brazil (Minas Gerais).

Acknowledgments. We thank Paul M. Peterson
of the Smithsonian Institution for providing us with
photocopies of Agnes Chase’s unpublished manu¬
script of the South American species of Paspalum.
We thank Linda S. Ellis for skillfully rendering the
illustrations. Filgueiras thanks CAPES (Brazilian Post-
Graduation Agency) and IBGE for a post-doctoral
fellowship that enabled him to work at the Missouri
Botanical Garden. He also thanks the Missouri Bo¬
tanical Garden for making their facilities available
to him for his research. He especially thanks his
friend Clarence Lovejoy and family for their friend¬
ship and support during his stay in St. Louis.

Literature Cited

Chase, A. 1929. The North American species of Pas¬
palum. Contr. U.S. Natl. Herb. 28(1): i-viii, 1-310,
ix-xvii.

Davidse, G. & R. P. Ellis. 1987. Arundoclaytonia, a
new genus of the Steyermarkochloeae: Arundino-
ideae from Brazil. Ann. Missouri Bot. Gard. 74: 479-
490.

Parodi, L. 1937. Contribucion al estudio de las gra-
mineas del genero “Paspalum.” Revista Mus. La
Plate, n.s., Bot. 1: 211-251.

Smith, L. B., D. C. Wasshausen & R. M. Klein. 1982.
Gramineas. In R. Reitz (editor). Flora Ilustrada Ca-
tarinense. Herbario “Barbosa Rodrigues,” Itajai.

The Genus Polycarpon (Caryophyllaceae) in South America,
with One New Species from the Norte Chico of Chile

Roy E. Gereau

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Clodomiro Marticorena

Departamento de Botanica, Universidad de Concepcion, Casilla 2407, Concepcion, Chile

Abstract. Polycarpon coquimbense Gereau &
Marticorena (Caryophyllaceae) is described from
central Chile, a key to and a brief conspectus of the
South American species of Polycarpon are provid¬
ed, and P. australis Britton is reduced to synonymy
under P. suffruticosum Grisebach.

RESUMEN. Se presentan la descripcion de Poly¬
carpon coquimbense Gereau & Marticorena (Car¬
yophyllaceae) proveniente de Chile central, una cla¬
ve y una sinopsis de las especies sudamericanas de
Polycarpon ; y P. australis Britton esta ubicado en
la sinonimia de P. suffruticosum Grisebach.

Polycarpon Loefling ex L. (Caryophyllaceae, Pa-
ronychioideae, Polycarpeae; Pax & Hoffmann, 1934)
is a genus of some 16 species of warm and warm-
temperate regions of the world (Pedersen, 1987).
The following combination of characters distinguish¬
es Polycarpon from the other genera of Caryophyl¬
laceae in South America (Reiche, 1896; Macbride,
1937; Anon, 1967; Pedersen, 1984, 1987): stipules
present, interpetiolar, connate (though sometimes
deciduous and detectable only as an interfoliar rim);
flowers in cymes; sepals free or connate only at
extreme base, usually keeled, rarely cucullate to
subcucullate; petals present, entire to emarginate;
style undivided at least in lower half; stigma 3-lobed
to 3-parted; fruit a several-seeded capsule; and seeds
not winged.

Macbride (1937) mistakenly treated Drymaria
engleriana (Muschler) Baehni & J. F. Macbride as
Polycarpon englerianum Muschler (Duke, 1961);
P. urbanianum Muschler is here considered a prob¬
able synonym of Drymaria engleriana. Examina¬
tion of an isotype of Polycarpon australis Britton
(Morong 933, MO) has shown it to be synonymous
with P. suffruticosum Grisebach. With these exclu¬
sions, four species of Polycarpon are known from
South America. Three of these species are native,
including the one newly described here. The four
species may be distinguished as follows:

Key to the Species of Polycarpon in South America

la. Root slender, delicate; base of stem herbaceous.

2a. Stems with a line of pubescence on one

side of apical internodes; leaves opposite
or in verticils of 5-10, the blades linear to
oblanceolate; sepals not winged on keel ..

.1. P. apurense

2b. Stems glabrous; leaves opposite or in ver¬
ticils of 3 or 4, the blades elliptic to obo-

vate; sepals winged on keel .

. 3. P. tetraphyllum

lb. Root thick, fleshy or woody; base of stem a
woody caudex.

3a. Leaves strictly opposite, the blades broadly
obovate to orbicular, 5-9 mm long, obtuse
to rounded at base; cymes dense; sepals
1-nerved, keeled; stamens 5; capsule ca.

1.6 mm long; seeds 3-7 .. 4. P. coquimbense
3b. Leaves opposite or verticillate, the blades
spatulate to oblong- or linear-lanceolate,
10-40 mm long, attenuate at base; cymes
lax; sepals 3-nerved, subcucullate; stamens
3; capsule ca. 5 mm long; seeds 25-35
. 2. P. suffruticosum

Enumeration of the South American Species

1. Polycarpon apurense Kunth in Humb., Bonpl.

& Kunth, Nov. Gen. Sp. 6: 40. 1823.

Synonymy in Pedersen (1987).

ARGENTINA (Corrientes) and URUGUAY (Peder¬
sen, 1987). BRAZIL. Goias: Irwin et al. 17682 (MO,

NY). Mato Grosso: Harley & Souza 11101 (K, MO).

Para: Berg & Henderson 649 (MO, NY). Sao Paulo:

Eiten & Campos 3473 (MO). VENEZUELA. Amazo¬
nas: Carnevali et al. 1503 (MO, VEN). Apure: Davidse

& Gonzalez 12877 (MO).

2. Polycarpon suffruticosum Grisebach, Abh. Kon-

igl. Ges. Wiss. Gottingen 19: 29. 1874.

Polycarpon australis Britton, Ann. New York Acad. Sci.

7: 53. 1892 [sub “ Polycarpa ”]. Syn. nov. TYPE.

Paraguay. Pilcomayo River, 1888-1890, Morong

933 (holotype, NY not seen; isotype, MO).

Further synonymy in Pedersen (1987).

Novon 5: 152-155. 1995.

Volume 5, Number 2 Gereau & Marticorena 153

1995 Polycarpon in South America

Figure 1. Polycarpon coquimbense Gereau & Marticorena. — A. Flowering plant. —B. Open flowrer. —C. Dehiscing
capsule. —D. Seed, dorsal view. —E. Seed, lateral view. —F. Sepal, dorsal view. —G. Petal, dorsal view. — H.
Stamens, lateral and ventral views. Illustrations based on: A-G, Ricardi et al. 1135; H, Ricardi et al. 1802.

154

Novon

ARGENTINA. Buenos Aires: Anon (1967). Cor¬
doba, Jujuy, Tucuman, Santa Fe: Martlnez-Crovetto
(1967). Entre Rios: Pedersen (1987). Chaco: Jorgen¬
sen 2366 (MO). Corrientes: Krapovickas et al. 27409
(CTES, MO). Formosa: Morel 4593 (LIL, MO). Salta:
Rosa 264 (MO). PARAGUAY. Gran Chaco: Hassler
2481 (MO). Presidente Hayes: Schinini & Palacios
25831 (CTES, MO).

3. Polycarpon tetraphyllum (L.) L., Syst. Nat. ed.

10. 881. 1759. Mollugo tetraphylla L., Sp.
PI. 1: 89. 1753.

Native to the Mediterranean region of Eurasia
and North Africa; introduced in southern Brazil,
Uruguay, coastal Argentina (Pedersen, 1984), and
central provinces of Chile (Reiche, 1896). Also col¬
lected in:

CHILE. Juan Fernandez Islands: Meyer 9373 (MO,
NA). PERU. Lima: Valencia & Cano 1718 (MO). VEN¬
EZUELA. Distrito Federal: Croat 21820 ( MO).

4. Polycarpon coquimbense Gereau & Marti-

corena, sp. nov. TYPE: Chile. Prov. Coquimbo:
Dpto. Elqui, Camino Intemacional a San Juan
[Argentina], Puente Las Terneras, 22 km al
interior de Guanta, 1650 m, 29°59'S, 70°14'W,
7.1.1967, Ricardi, Marticorena & Matthei
1802 (holotype, CONC; isotype, OS). Figure 1.

Fruticulus humilis, ramis basalibus lignosis tortis, ra-
mulis floriferis sublignosis, usque ad 15 cm longis. Folia
opposita; stipulis caducis (nobis non observatis), sed fo-
liorum basibus crista stipulari manifeste unitis; petiolis
0.4-1.0 mm longis, crassiusculis; laminis 5-9 mm longis
x 4-9 mm latis, late obovatis ad orbiculatis, apice ro-
tundatis, basim versus obtusis ad rotundatis, coriaceis,
nervo unico longitudinali vix manifesto. Cymae densae,
terminales, dichotomae; bracteis scariosis ovatis 0.6-0.9
mm longis. Flores subsessiles; sepalis 5, 1.9-2.4 mm
longis, ovato-orbiculatis, obtusis, uninervibus, nervo viri-
dibus carinatisque; petalis perigynis, 1.3-1.5 mm longis,
oblongis, obtusis, integris; staminibus 5, perigynis, fila-
mentis 1.0-1.1 mm longis, antheris ca. 0.5 mm longis;
ovario trigono-ovoideo ca. 0.8 mm longo x 1.0 mm lato,
stylo ca. 0.7 mm longo, indiviso, stigmatibus 3 liberis.
Capsula trigono-ovoidea ca. 1.6 mm longa x 1.1 mm
lata; seminibus 3-7, 0.90-1.05 mm longis x 0.35-0.40
mm latis, erectis, lunato-fusiformibus, superficie lucente,
fere laevi (minute celluloso-papillosa).

Shrublet, very low, the basal branches woody,
twisted, the flowering branchlets subligneous, to 15
cm long, yellowish to olive-green, longitudinally sul-
cate, glabrous to minutely pulverulent with very
short fasciculate trichomes. Leaves opposite, some¬
times with an axillary pair of smaller leaves; stipules
caducous (none observed), but leaf bases conspic¬
uously united by a stipular ridge; petioles 0.4-1.0
mm long, rather stout, adaxially canaliculate; blades
5-9 x 4-9 mm, broadly obovate to orbicular.

rounded at apex, obtuse to rounded at base, with a
single, inconspicuous longitudinal nerve, tomentu-
lose above and beneath when young with branched
fasciculate trichomes. Cymes dense, terminal, di¬
chotomous, 4-6-branched, the basal internodes 4-
10 mm long, the upper very short; bracts scarious,
ovate, 0.6-0.9 mm long; branches tomentulose as
in young leaves. Flowers subsessile; sepals 5, 1.9-
2.4 mm long, ovate-orbicular, obtuse, 1-nerved,
green and keeled on nerve, broadly hyaline on mar¬
gins, united for ca. 0.5 mm at base and forming an
hypogynous disk, tomentulose externally as in
branches of cyme; petals perigynous, 1.3-1.5 mm
long, oblong, obtuse, entire; stamens 5, perigynous,
the filaments 1.0-1.1 mm long, the anthers ca. 0.5
mm long; ovary trigonous-ovoid, ca. 0.8 mm x 1.0
mm, unilocular, with 9 ovules inserted at base of
central column; style ca. 0.7 mm long, contiguous
with column of ovary, undivided, with 3 free stigmas.
Capsule trigonous-ovoid, ca. 1.6 x 1.1 mm; seeds
3-7, 0.90-1.05 x 0.35-0.40 mm, erect, lunate-
fusiform; embryo dorsal, curved, brown; perisperm
ventral, yellowish brown; seed surface shining, near¬
ly smooth (minutely cellular-papillose).

Polycarpon coquimbense resembles P. suffruti-
cosum in its habit; the differences are noted in the
key above. In this respect it also resembles P. po-
lycarpoides (Bivona-Bernardi) Zodda ex Jahandiez
& Maire subsp. herniarioides (Ball) Maire & Weill-
er of dry montane habitats in northwestern Africa
and the southeastern Iberian Peninsula (Amich &
Pedrol, 1990), and P. sauvagei Mathez of the Cen¬
tral Plateau of Morocco (Mathez, 1983). Polycar¬
pon sauvagei is readily distinguished by its sessile,
narrowly elliptic leaves measuring only 2.2-5.0 x
0.5-1.5 mm and its large flowers (sepals 3.5-4.0
mm long, petals ca. 3.5 mm long). Polycarpon
polycarpoides subsp. herniarioides is superficially
quite similar to P. coquimbense, but the former
taxon may be distinguished by the following char¬
acters: leaf blades 3-7 x 1-2.5 mm, oval to ob¬
ovate, acute at apex; sepals 1.5-2.0 mm long, only
slightly cucullate (not keeled); seeds 0.6-0.8 mm
long, nearly white. [Note: Amich & Pedrol (1990)
mistakenly ascribed the combination Polycarpon
polycarpoides to Fiori (Nuovo Giorn. Bot. Ital., nov.
ser. 15: 347. 1908), who specifically rejected the
name as “una logomachia senza senso comune.
Maire (1963) cited its authorship as Zodda ex Fiori
(Nuov. FI. Italia 1: 451. 1923), but Fiori there used
polycarpoides only as an unranked infraspecific
epithet under P. peploides DC. Jahandiez & Maire
(Cat. PI. Maroc 2: 212. 1932) validly transferred
the specific epithet from Hagaea polycarpoides

Volume 5, Number 2
1995

Gereau & Marticorena
Polycarpon in South America

155

Bivona-Bemardi to Polycarpon , crediting Zodda with
the combination. The correct author citation is
therefore P. polycarpoides (Bivona-Bernardi) Zod¬
da ex Jahandiez & Maire.]

Paratypes. CHILE. Prov. Coquimbo: Quanta, entre
Rivadavia y Guanta, 1215 m, 29°50'S, 70°25'W,
4.II.1963, Ricardi, Marticorena & Matthei 677 (CONC,
MO); camino de La Serena a Banos del Toro, 23 km al
interior de Guanta, 1650 m, 29°59'S, 70°15'W,
27.X. 1964, Ricardi , Marticorena & Matthei 1135
(CONC, MO).

Acknowledgments. We thank R. K. Rabeler and
C. M. Taylor for critical reading of the manuscript,
T. F. Stuessy for loan of specimens from OS, and
N. Moya for the fine illustration.

Literature Cited

Amich, F. & J. Pedrol. 1990. Polycarpon Loefl. ex L.
In: S. Castroviejo, M. Lainz, G. Lopez Gonzalez, P.
Montserrat, F. Munoz Garmendia, J. Paiva & L.
Villar (editors), Flora Iberica 2: 161-167. Real Jar-
din Botanico, Madrid.

A non Suarez de Cullen, D. 1967. Caryophyllaceae. In:
A. L. Cabrera (editor). Flora de la Provincia de Bue¬

nos Aires 3: 192-237. Instituto Nacional de Tec-
nologia Agropecuaria, Buenos Aires.

Duke, J. A. 1961. Preliminary revision of the genus
Drymaria. Ann. Missouri Bot. Gard. 48: 173-268.

Macbride, J. F. 1937. Caryophyllaceae. In: J. F. Mac-
bride, Flora of Peru, Field Mus. Nat. Hist., Bot. Ser.
13(2): 578-638.

Maire, R. 1963. Caryophyllaceae. In: R. Maire, Flore
de PAfrique du Nord 9: 5-300, Editions Paul Lech-
evalier, Paris.

Martinez-Crovetto, R. 1967. Catalogo preliminar de las
Caryophyllaceae de la Argentina y del Uruguay. Bon-
plandia (Corrientes) 2: 187-264.

Mathez, J. 1983. Une espece nouvelle du genre Po¬
lycarpon: (Caryophyllaceae): P. sauvagei. Bull. Soc.
Bot. France 130, Lett. Bot.: 357-362.

Pax, F. & K. Hoffmann. 1934. Caryophyllaceae. Nat.
Pflanzenfam. ed. 2, 16c: 275-364.

Pedersen, T. M. 1984. Caryophyllaceae. In: M. N.
Correa (editor), Flora Patagonica 4a: 196-276. In¬
stituto Nacional de Tecnologia Agropecuaria, Buenos
Aires.

-. 1987. Caryophyllaceae, Cariofilaceas. In: N.

Troncoso de Burkart & N. M. Bacigalupo (editors).
Flora Ilustrada de Entre Rios (Argentina) 3: 251-
291. Instituto Nacional de Tecnologia Agropecuaria,
Buenos Aires.

Reiche, C. 1896. Cariofilaceas. In: C. Reiche, Flora de
Chile 1: 173-212. Imprenta Cervantes, Santiago.

New Species of the Southern African Genus
Geissorhiza (Iridaceae: Ixioideae)

Peter Goldblatt

B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299,
St. Louis, Missouri 63166, U.S.A.

John C. Manning

National Botanical Institute, P. Bag X7, Claremont 7735, South Africa

ABSTRACT. Three new species are recognized in
the southern African Geissorhiza, a genus now in¬
cluding 84 species. Geissorhiza silenoides and G.
rupicola are members of subgenus Geissorhiza and
occur in the Ceres District of the southwestern Cape,
South Africa. Both have pale pink flowers with un¬
usually long perianth tubes for the genus. Geissor¬
hiza silenoides is additionally distinguished by hav¬
ing unequal stamens, each filament being of a dif¬
ferent length, scabrid-puberulent stems, and leaves
with heavily thickened margins and midribs. Geis¬
sorhiza rupicola has unusual soft-textured corm
tunics and, like G. silenoides, has leaves with strong¬
ly thickened margins and midribs. The third new
species, G. uliginosa, is restricted to the Swartberg
Mountains of the southern Cape and grows on wet
cliffs and under waterfalls. It stands out in the genus
in having the flowers solitary on the stems, and very
soft-textured corm tunics that rapidly decay, some¬
times leaving the corms apparently without tunics.
All three new species occur in the southern and
southwestern part of southern Africa, a region known
for the richness and diversity of its flora, particularly
of geophytes.

More than half of the estimated 1750-1800 spe¬
cies of the petaloid monocot family, Iridaceae, occur
in southern Africa (Goldblatt, 1994). The region is
well known botanically, yet despite over 250 years
of botanical exploration, novelties continue to be
discovered there. Although the family is a numeri¬
cally significant element of the flora of a large part
of southern Africa, Iridaceae are most diverse in
the southern and western coastal belt and adjacent
mountains of the subcontinent, a region of mediter¬
ranean-type climate with winter rainfall and summer
drought. Three new species of Geissorhiza, a genus
virtually restricted to the winter rainfall zone of
southern Africa, are described here. Some 81 spe¬
cies were admitted to the genus when it was last
revised (Goldblatt, 1985), and the new species de-

Novon 5: 156-161. 1995.

scribed here thus bring the total to 84. Geissorhiza
is a member of Iridaceae subfamily Ixioideae, the
largest of the four subfamilies currently recognized
in Iridaceae (Goldblatt, 1990, 1991). It is one of
the largest genera of the subfamily and one of the
larger genera in the southern African flora. One of
the new species, G. uliginosa, belongs in subgenus
Weihea, which is characterized by concentric corm
tunics, and G. silenoides and G. rupicola are mem¬
bers of subgenus Geissorhiza, species of which have
the characteristic imbricate corm tunics that in¬
spired the generic name, a Greek word meaning
tiled root.

1. Geissorhiza silenoides Goldblatt & J. C. Man¬
ning, sp. nov. TYPE: South Africa. Western
Cape: Ceres District, lower slopes of Gydo Pass,
20 Sep. 1993, Goldblatt & Manning 9739
(holotype, NBG; isotypes, K, MO, PRE, WAG).
Figure 1A.

Plantae 20-30 cm altae, cormo 5-7 mm diametro
tunicis duris imbricatis, foliis linearibus 1.2-1.5 mm latis
marginibus costisque incrassatis, spica 4-7 florum, flori-
bus pallide roseis, tubo perianthii cylindrico 15-17 mm
longis, tepalis subaequalibus 12.5-14 x 6-7 mm, fila-
mentis inaequalibus 4, 7 et 9 mm longis, antheris ca. 4.5
mm longis, stylo eccentrico.

Plants 20-30 cm high. Corm ± ovoid, 5-7 mm
diam., flattened slightly below on one side, the tunics
woody, blackish, imbricate, the layers breaking be¬
low into triangular points. Cataphylls membranous,
the upper light purple above the ground. Leaves 4,
the lower 2 basal and longest, reaching to at least
the middle of the stem, sometimes to the base of
the spike, straight and erect or trailing, the blades
linear, 1.2-1.5 mm wide, oval in section with 2
narrow grooves on each surface, the midrib much
thickened, the margins raised and arched over the
surface, the upper leaves cauline, sheathing for at
least half their length, the blades like the basal, the
uppermost sometimes only 9-12 mm long and en-

Volume 5, Number 2
1995

Goldblatt & Manning
Geissorhiza from Southern Africa

157

158

Novon

tirely sheathing. Stem erect below, flexed above the
sheaths of the 2 upper leaves and inclined above,
unbranched, minutely scabrid-puberulent. Spike
flexuose, drooping in bud, 4-7-flowered; bracts un¬
equal, the outer pale green, the upper margins pink¬
ish purple, 7-10 mm long, the inner about two-
thirds as long, transparent with 2 green keels. Flow¬
ers hypocrateriform, actinomorphic, pale pink,
darker toward the tepal margins, the center becom¬
ing whitish, unscented; perianth tube cylindric, 15-
17 mm long; tepals subequal, obovate, 12.5-14 x
6-7 mm, when fully open spreading at right angles
to the tube. Filaments unequal, inserted ca. 2 mm
below the apex of the tube, 4, 7, and 9 mm long,
straight and erect; anthers ca. 4.5 mm long, cream,
the pollen pale yellow. Ovary ovoid, ca. 2 mm long;
style exceeding the anthers by 2-4 mm, the branch¬
es unusually long, 5-6 mm long, recurving. Cap¬
sules and seeds unknown.

Flowering September to mid October.

Etymology. The species is named for the resem¬
blance of the flowers to those of some southern
African species of Silene (Caryophyllaceae), a spe¬
cies of which grows near Geissorhiza silenoides and
has flowers of nearly the same pink color.

Distribution and habitat. Geissorhiza sile¬
noides is evidently restricted to the Ceres District
of Western Cape Province, South Africa, an area
that is relatively well explored botanically. Only two
collections of the species are known, suggesting that
it is rare and extremely local in distribution. The
type collection was made along a roadside cutting
on the south-facing lower mountain slopes of Gydo
Pass, and the other collection, made in 1924, is
without precise location. The slopes of Gydo Pass
have a relatively high rainfall and soils of heavy
clay derived from shales of the Bokkeveld Shale
Series. These slopes are normally densely clothed
with fynbos vegetation dominated by the arborescent
Protea laurifolia Thunberg, and a dense ground
cover and little open ground where geophytes can
grow. Geophytes are, however, fairly common and
diverse there and appear in profusion after fire or
clearing of the bush, such as along the road verges,
where there were extensive populations of G. sile¬
noides growing together with Hesperantha bach-
mannii Baker, also Iridaceae. We suspect that G.
silenoides may occur elsewhere in similar habitats
along the lower mountain slopes of the ranges north
and east of Ceres, where the same vegetation and
soils occur. After fire, the slopes adjacent to Gydo
Pass should be explored for additional populations
of G. silenoides.

The blackish imbricate corm tunics regularly

notched below into blunt triangular segments place
Geissorhiza silenoides in subgenus Geissorhiza , one
of two subgenera recognized by Goldblatt (1985).
Within the subgenus, this species stands out in the
scabrid-puberulent stems, short leaves with heavily
thickened margins and midribs, thus with two narrow
grooves on each surface, and in the flowers. The
pale pink perianth color is unusual in subgenus Geis¬
sorhiza , although it is known in a few species of
sections Geissorhiza and Monticola. The most dis¬
tinctive floral features of G. silenoides are the long
perianth tube combined with the unequal stamens,
each filament being of a different length. Unequal
filaments occur sporadically in subgenus Geissor¬
hiza , although typically only one filament is short
and the remaining two are equal. The combination
of long tube and unequal stamens is known only in
section Monticola , but most members of this section
have soft-textured corm tunics and only one, G.
scopulosa Goldblatt, has puberulous stems (Gold¬
blatt, 1985). In section Geissorhiza unequal fila¬
ments also occur in some species, and although hard-
textured corm tunics and leaves with thickened mar¬
gins and midribs are typical of this section, a long
perianth tube is not. Sectional placement of G. sil¬
enoides is somewhat uncertain, but we tentatively
suggest that G. silenoides has its closest relatives
among the species of section Monticola. Because
of its unusual combination of derived features, its
immediate relatives are uncertain.

Geissorhiza silenoides is undoubtedly Geissor¬
hiza sp. 1 (Goldblatt, 1985), listed as such because
of the inadequate material known at the time. The
single sheet, collected by F. A. Rogers in 1924,
comprises two specimens that are rather poorly
pressed. Nevertheless, the scabrid-puberulent stems,
leaves with thickened margins and midribs, and flow¬
ers with long perianth tubes are clearly evident.

Paratype. SOUTH AFRICA. Western Cape: 3319
(Worcester) “Ceres,” (possibly the town itself or the dis¬
trict) (?AD), Sep. 1924, Rogers 28746 (Z).

2. Geissorhiza rupicola Goldblatt & J. C. Man¬
ning, sp. nov. TYPE: South Africa. Western
Cape: Mosterthoek Twins, Green Gullies Kloof,
ca. 1250 m, 9 Dec. 1984, Esterhuysen 36197
(holotype, BOL; isotypes, MO, NBG, PRE,
WAG). Figure IB.

Plantae (10-)15-20 cm altae, cormo 9-12 mm dia-
metro tunicis mollibus, foliis (2—)3 linearibus 1.5-3 mm
latis marginibus costisque forte incrassatis, spica (1 —)2—
6 florum, floribus pallide roseis, tubo perianthii cylindricis
(16-)19-23 mm longis, tepalis subaequalibus 12-15 x
4.5-5.5 mm, filamentis ca. 8 mm longis aequalibus, an-
theris ca. 3.5 mm longis, stylo eccentrico.

Volume 5, Number 2
1995

Goldblatt & Manning
Geissorhiza from Southern Africa

159

Plants (10-)15-20 cm high. Corm 9-12 mm
diam., globose, the tunics initially unbroken, soft
and papery, becoming tom into narrow vertical strips,
ultimately almost fibrous. Cataphylls usually dry
and brown at flowering time, usually persisting and
accumulating around the stem base as a neck of
dry brittle fibers. Leaves (2-)3, the lower one or
two basal, the lowermost longest, reaching to be¬
tween the upper third of the stem and the base of
the spike, the second of the basal leaves usually
much shorter, and sheathing in the lower half, the
blades linear, 1.5-3 mm wide, the margins and
midribs heavily thickened and raised into wings over¬
arching the laminar surfaces, uppermost leaf in¬
serted on the mid to upper part of the stem, smaller
than the basal, sheathing for at least half its length,
sometimes entirely. Stem ascending or drooping from
cracks in steep rock faces, smooth, unbranched.
Spike (l-)2-6-flowered, flexuose; bracts subequal,
the outer pale green entirely or purple toward the
apices, 8-11 mm long, the inner membranous nar¬
rower than the outer, two keeled, forked in the upper
1-2 mm. Flowers pale pink, actinomorphic, un¬
scented; perianth tube cylindric, (16-)19-23 mm
long; tepals subequal, ovoid-oblong, spreading at
right angles to the tube, 12-15 x 4.5-5.5 mm.
Filaments ca. 8 mm long, exserted ca. 6 mm from
the tube; anthers ca. 3.5 mm long, cream, the pollen
yellow. Ovary oblong, 2-3 mm long; style eccentric,
dividing at or beyond the anther apices, ultimately
exceeding the anthers by ca. 4 mm, the branches
ca. 2 mm long. Capsules globose, 4-6 mm long;
seeds prismatic, the angles ridged, brown, ca. 1.5
mm at the longest axis.

Flowering late November to early December.

Etymology. From the Latin “rupicola,” favoring
rocky habitats.

Distribution and habitat. Geissorhiza rupicola
is known only from moist shady sites on Mosterthoek
Twins, a mountain at the western end of the Lan-
geberg Mountains, in the Ceres District of W estern
Cape Province, South Africa. Plants are confined
to moist sites, where they grow in thin soil on rocks,
sometimes in cracks in steep cliffs. Such habitats
are uncommon as late in the year as November and
December in the summer dry Western Cape. Geis¬
sorhiza rupicola joins a small number of species of
this genus, mostly of section Monticola, that favor
montane habitats and flower late in the season in
sheltered sites. The species was discovered by the
Cape botanist Elsie Esterhuysen, who has made an
extraordinary contribution to the knowledge of the
mountain flora of the southern and western Cape.

Geissorhiza rupicola stands out in the genus in

its unusually long perianth tube, 2-2.5 times as long
as the tepals and 16-22 mm long. A tube of this
length is also known in G. grandiflora Goldblatt,
but this species has longer tepals and unequal sta¬
mens, one filament of which is shorter than the other
two. Geissorhiza silenoides also has an unusually
long perianth tube but can immediately be distin¬
guished from G. rupicola by its puberulous stem,
hard imbricate corm tunics, and unequal stamens.

Paratype. SOUTH AFRICA. Western Cape: 3319
(Worcester) Mosterthoek Twins, Green Gullies Kloof, ca.
1250 m (4000 ft.) (AD), 18 Dec. 1988 (fr), Goldblatt
7483 (BOL, MO, NBG).

3. Geissorhiza uliginosa Goldblatt & J. C. Man¬
ning, sp. nov. TYPE: South Africa. Western
Cape: Oudtshoorn District, Swartberg Moun¬
tains, stream E of Gouekrans Hiking Trail hut
(AC), 15 Jan. 1989, Vlok 2075 (holotype, NBG;
isotype, MO). Figure 2.

Plantae 15-30 cm altae, cormo reducto 3-5 mm dia-
metro, foliis 5, laminibus planis marginibus costisque lev-
iter incrassatis, floris solitariis, floribus roseis vel purpureis,
tubo perianthii infundibuliformis 9-13 mm longis, tepalis
subaequalibus 20-22 x 9-11 mm, filamentis 12-14 mm
longis aequalibus, antheris 7-9 mm longis, stylo declinato.

Plants 15-30 cm high. Corm very reduced, 3-
5 mm diam., the tunics light brown, soft-textured,
not accumulating with age. Cataphylls rapidly de¬
caying, membranous. Leaves usually at least 5, the
lower 3 basal and longest, usually shorter than the
stems but trailing, the blades plane with lightly thick¬
ened margins and midribs, the cauline leaves shorter
than the basal, largely or entirely sheathing. Stem
drooping, usually with at least one, sometimes up
to four branches. Spike 1-flowered; bracts green,
the margins often red, the outer 16-20 mm long,
the inner about as long as or slightly shorter than
the outer. Flowers funnel-shaped, actinomorphic,
pale pink to purplish, cream in the throat, the base
of the tepals dark pink, unscented; perianth tube
narrowly funnel-shaped, 9—13 mm long; tepals
ovate-oblong, subequal, 20—22 x 9-11 mm, sub¬
patent when fully open. Filaments 12-14 mm long,
equal in length, unilateral; anthers 7-9 mm long,
dark brown, the pollen brownish. Ovary ovoid, 3-
4 mm long; style declinate, extended below the
filaments, the branches ca. 4 mm long. Capsules
and seeds unknown.

Flowering December and January.

Etymology. Named for the moisture-loving con¬
ditions the species requires, “ uliginosa ,” the Latin
adjective describing such habitats.

Distribution and habitat. Geissorhiza uliginosa

160

Novon

Volume 5, Number 2
1995

Goldblatt & Manning
Geissorhiza from Southern Africa

161

is restricted to waterfalls and wet cliffs in the Swart-
berg Mountains of the southern part of Western
Cape Province, South Africa. Plants are known from
only a few localities in this semiarid area of low and
predominantly winter rainfall. The species joins a
handful of others in the genus that favor this spe¬
cialized habitat, e.g., G. cataractarum Goldblatt, G.
elsiae Goldblatt, and G. outeniquensis Goldblatt.
The species was discovered by the South African
botanist J. H. J. Vlok, who made the first two col¬
lections in the summer of 1984-1985.

Despite the similar habitats favored by Geissor¬
hiza uliginosa and the species mentioned above,
the relationships of G. uliginosa appear to lie not
with any of these other moisture-loving, cliff-dwelling
species, but with another species of the southern
Western Cape, G. fourcadei (L. Bolus) G. Lewis.
That species grows in thin soils on rock outcrops in
the mountains of the southern Cape, and it flowers
mainly in the autumn, February to May. Like G.
fourcadei , G. uliginosa has pale pink flowers and
branched stems, each of which bears just a single
flower. Geissorhiza uliginosa differs in having flow¬
ers with a shorter perianth tube, 9-13 mm long,
plane leaves, and soft-textured short-lived corm tu¬
nics. This contrasts with the longer perianth tube,
(8-) 17-20 mm long, xeromorphic terete, four-
grooved leaves, persistent fibrous cataphylls, and

larger corms with persistent firm-papery corm tunics
of G. fourcadei.

Paratypes. SOUTH AFRICA. Western Cape: 3321
(Ladismith) Swartberg Mountains, along road to Gamkak-
loof at Waterfall Kloof (DC), 5 Dec. 1985, Goldblatt
7455 (MO). 3322 (Oudtshoorn) upper N-facing slopes of
Waboomsberg in perennial waterfall, ca. 4800 ft. (AC),
26 Dec. 1984, Vlok 881 (MO, NBG); lower southern
slopes of the Swartberg at Rust-en-Vrede, cliffs next to
waterfall (AD), 20 Jan. 1985, Vlok 894 (MO).

Acknowledgments . We thank Elsie Esterhuysen
and Jan Vlok for assistance with fieldwork and for
drawing our attention to G. rupicola and G. uli¬
ginosa, respectively.

Literature Cited

Goldblatt, P. 1985. Systematics of the southern African
genus Geissorhiza (Iridaceae-Ixioideae). Ann. Mis-
souri Bot. Gard. 72: 277-447.

-. 1990. Phytogeny and classification of Irida-

ceae. Ann. Missouri Bot. Gard. 77: 607-627.

-. 1991. An overview of the systematics, phy-

logeny and biology of the southern African Iridaceae.
Contr. Bolus Herb. 13: 1-74.

-. 1994. Geography of African Iridaceae. Pp.

931-940 in J. H. Seyani & A. C. Chikuni (editors),
Proceedings of the XIHth plenary Meeting AETFAT,
Malawi, Volume 2. National Herbarium and Botanic
Gardens of Malawi, Zomba.

Nolina cismontana (Nolinaceae), a New Species
Name for an Old Taxon

William J. Hess

The Morton Arboretum, Lisle, Illinois 60532, U.S.A.
James C. Dice

California Department of Fish and Game, P.O. Box 2537,
Borrego Springs, California 92004, U.S.A.

ABSTRACT. Nolina parryi subsp. woljii is shown
to be a synonym of N. parryi subsp. parryi. Con¬
sequently, N. parryi subsp. parryi sensu Munz has
been misinterpreted and is without a name. Nolina
cismontana is proposed as the name for this un¬
described taxon.

It is necessary to publish a new name for Nolina
parryi S. Watson sensu Munz before volume 11 of
the Flora of North America is published. The junior
author studied the California Nolina species complex
for his M.S. thesis (Dice, 1988). He determined that
the name N. parryi was assigned by western bot¬
anists to the wrong plants, and consequently these
plants are unnamed.

Munz (in Munz & Roos, 1950) described Nolina
parryi subsp. wolfii based on plants from the Kings¬
ton Range in the eastern Mojave Desert and the
San Jacinto/Santa Rosa Mountains along the north¬
western edge of the Colorado Desert. Plants fitting
this description are now known to extend from the
Kern Plateau in southern Tulare County south to
the Laguna and Pinyon Mountains of San Diego
County, California. The taxon is found in the drier
parts of the Peninsular Ranges, along the western
edge of the Colorado Desert and in the desert ranges
of the Mojave Desert. This means that by this in¬
terpretation subspecies parryi occurs in the coastal
drainages below 3000 feet from Ventura to San
Diego counties and west of subspecies wolfii. There
is no evidence that Munz ever examined the type
of N. parryi. Dice examined the holotypes and
isotypes of subspecies parryi and subspecies wolfii.
The holotype of subspecies parryi and Watson’s
(1879) publication of its description indicate that N.
parryi came from the desert east of San Bernardino
and not from the coastal side of the mountains. The
leaves on this holotype and isotype are wider than
what Munz allows for his subspecies parryi and
would key out in his keys to subspecies wolfii. Sub¬
species wolfii is a synonym for Watson’s typical N.

Novon 5: 162-164. 1995.

parryi, and the plants from the coastal drainages
that Munz was calling subspecies parryi are without
a name.

Dice (1988) showed that there are quantitative
morphological traits such as leaf number per rosette,
stem length, leaf width, panicle length, panicle stalk
diameter, and bract size that help to differentiate
Nolina parryi from N. cismontana. Besides the
geographical difference stated above, N. parryi is
found on granite and granodiorite-derived substrates
in the xeric pinyon-juniper woodlands, while N. cis¬
montana occurs principally on sandstone and shale
substrates of chaparral vegetation. The morpholog¬
ical, geographical, and ecological differences to us
warrant the recognition at the species level of the
coastal foothills and valleys taxon.

Nolina cismontana Dice, sp. nov. TYPE: U.S.A.
California: Orange Co., Santa Ana Mts., Tra-
buco Canyon, 350 m, 1986, J. Dice & T.
Oberbauer 650 [staminate] (holotype, SD
121705; isotypes, ARIZ, NY, RSA, UC).

A specie Nolina parryi caulibus brevibus, generaliter
< 0.4 m longis; rosulis amplitudine mediocribus, gener¬
aliter ex 30-90 foliis constantibus; foliis latitudine me¬
diocribus, 12-30 mm latis accurate super basem expan¬
sum; pedunculo inflorescentiae 14-35 mm diametro basi
differt.

Shrubs to subshrubs, 0.5—1.5 m tall, few- to
many-branched from a woody caudex, branching
both above and below ground, with l-30(-75) leaf
rosettes; mature rosettes 0.7-1.6 m diam. with 30-
90 leaves per rosette. Leaves green (occasionally
glaucous) lance-linear, 0.5-1.4 m long at maturity
and 1.2—3 cm wide just above the expanded leaf
base; leaf bases white, deltoid-rhomboidal, spoon¬
shaped, 4-11.5 cm long and 3-8.5 cm wide at
base; leaf margins serrulate. Inflorescences large,
1.3-3.1 m tall, 10-40 cm wide, with 25-75 sec¬
ondary panicle branches, longest secondary branch
13-35 cm long, longest tertiary branch 1-10 cm

Volume 5, Number 2
1995

Hess & Dice
Nolina cismontana

163

long; panicle stalks 1.4-3.5 cm diam. at base, bar¬
ren the lowermost 0.4-1.5 m; bracts and bractlets
large, conspicuous, persistent, lance-linear to del¬
toid-lanceolate, 10-40 cm long, 1.5-5 cm wide at
base (bract subtending lowermost secondary panicle
branch); pedicels 2-4 mm long, jointed near middle.
Tepals cream to white, ovate, apiculate-mucronate
to rounded and introrsely papillate at tip. Staminate
flowers broadly campanulate to recurved at anthesis,
with outer tepals 3-5 mm long, 1.5-2.5 mm wide;
stamens 2-4 mm long; ovaries on short stalks above
perianth, 0.5-1.5 mm tall and 1-1.5 mm diam.;
styles and stigmatic surfaces greatly reduced or
wanting. Carpellate flowers with outer tepals 3-4
mm long, 2 mm wide, and erect to campanulate at
anthesis; staminodes 1-2 mm long; ovary on short
stalk above perianth, 3 mm tall and 2-3 mm diam.;
styles 3, closely appressed but distinct, < 0.5 mm
long; stigmatic surface well developed. Fruits pa¬
pery, 3-lobed capsules, 8-12 mm tall and slightly
broader, emarginate at base and apex, bursting ir¬
regularly. Seeds ovoid to oblong, 4-5 mm long, 3-
4 mm diam.; seed coats reddish brown. N = 19
(Lenz, 1950).

Distribution. Cismontane southern California, on
dry chaparral-covered slopes extending from the
foothills of the Santa Ynez Mountains in western
Ventura County, south through the Simi Hills and
Santa Ana Mountains to the foothills west of the
Palomar and Cuyamaca Mountains in San Diego
County. Elevation 230-1275 m.

Etymology. The specific epithet “cismontana”
means this side of the mountain and refers to the
hillside slopes of the foothills of the southern coastal
range, where it commonly occurs.

Common name. Chaparral nolina, chaparral
beargrass.

Paratypes. U.S.A. California: Orange Co., Santa
Ana Mountains, elev. 365 m, 1986, J. Dice & T. Ob-
erbauer 651 [pistillate] (ARIZ, NY, RSA, SD, UC).

With the recognition of Nolina cismontana, the
synonymy for Nolina parryi has to be updated.

Nolina parryi S. Watson, Proc. Amer. Acad. Arts
14: 247. 1879. Nolina bigelovii (Torrey) S.
Watson var. parryi (S. Watson) L. D. Benson,
Bull. Univ. Arizona 15: 384. 1945. Nolina
bigelovii subsp. parryi (S. Watson) E. Murray,
Kalmia 13: 10. 1983. TYPE: U.S.A. Califor¬
nia: [probably Riverside Co.] desert E of San
Bernardino, Parry s.n. (holotype, GH; isotype,
MO).

Nolina parryi S. Watson subsp. wolfii Munz in Munz &
Roos, Aliso 2(3): 221. 1950. Nolina bigelovii var.
wolfii (Munz) L. D. Benson, in Benson & Darrow,
Trees Shrubs Southwest. Deserts 72. 1954. Nolina
wolfii (Munz) Munz, FI. S. Calif. 865. 1974. Nolina
bigelovii (Torrey) S. Watson subsp. wolfii (Munz)
E. Murray, Kalmia 13: 10. 1983. TYPE: U.S.A.
California: eastern San Bernardino Co., Mojave Des¬
ert, Kingston Mountains, 0.5 mi. W of Beck Spring
(Crystal Spring on herbarium labels), 1 Volf 7655
(holotype, RSA; isotypes, A, DS, F, GH, JEPS, LA,
MO, NY, POM, SD, UC, US).

The key to Nolina in The Jepson Manual (Dice,
1993) does not reflect the presence of N. cismon¬
tana, although there is mention of an undescribed
species under the description of N. parryi. The key
below is adapted from Dice (1988).

Key to the Species of Nolina in California

la. Old leaf margins fibrous-shredding; inflores¬
cence bracts deciduous; seeds gray.

. Nolina bigelovii

lb. Leaf margins minutely and persistently serrate;
inflorescence bracts persistent; seeds reddish
brown.

2a. Aboveground stems not obvious; mature
rosettes with < 45 leaves; leaf bases barely
expanded; inflorescences < 1.6 m, base of
stalk 0.5-1.8 cm diam.; bracts small, in¬
conspicuous . Nolina interrata

2b. Aboveground stems obvious; mature ro¬
settes with 45-200 leaves; leaf bases
strongly expanded; inflorescences 1.6-4
m, base of stalk 1.4-9 cm diam.; bracts
large, papery.

3a. Stems 0.3-2.1 m long; rosettes with
65-200 leaves; leaves 20-40 mm
wide just above expanded leaf base;
inflorescence stalks 2.6-9 cm diam.

at base . Nolina parryi

3b. Stems up to 0.4 m long; rosettes with
30-90 leaves; leaves 12-30 mm wide
just above expanded base; inflores¬
cence stalks 1.4-3.5 cm diam. at base
. Nolina cismontana

Conservation status. Nolina cismontana was re¬
cently added to the U.S. Fish and Wildlife Service’s
(USFWS) list of plant taxa native to the United
States that are being reviewed for possible addition
to the List of Endangered and Threatened Plants
under the Endangered Species Act of 1973, as
amended (Department of the Interior, Fish and
Wildlife Service, 1993). In USFWS’s most recent
review, N. cismontana (as Nolina sp. nov. ined.,
chaparral bear-grass from California) is treated as
a Category 2 candidate species for listing under the
Federal Endangered Species Act. Category 2 can¬
didate species are “taxa for which information now
in the possession of the Service indicates that pro-

164

Novon

posing to list as endangered or threatened is possibly
appropriate, but for which sufficient data on biolog¬
ical vulnerability and threat are not currently avail¬
able to support proposed rules.” Nolina cismontana
is also deserving of consideration from the California
Department of Fish and Game for possible inclusion
in their list of plant species of special concern and
by the California Native Plant Society for inclusion
in their Inventory of the Rare and Endangered Vas¬
cular Plants of California.

Throughout most of its range, from Ventura
County through northern San Diego County, Nolina
cismontana is threatened by residential and com¬
mercial land development. At present the only pro¬
tected populations of significant sizes are the Viejas
Mountain population in southern San Diego County,
which lies within the U.S. Forest Service’s Cleveland
National Forest; the California Department of Fish
and Game’s Coal Canyon Ecological Reserve in the
Santa Ana Mountains of eastern Orange County;
and scattered holdings within the Cleveland National
Forest in the Santa Ana Mountains.

Acknowledgments. The authors thank Ian
MacPhail for providing the Latin descripton of No¬
lina cismontana. The junior author thanks R.
Mitchel Beauchamp, Ann Croft, Michael Donoghue,
Steve Junak, Deborah Kinsinger, Karlin Marsh, Tom
Oberbauer, Fred Roberts, John Taft, Tim Thomas,
Harold Wier, and Kirsten Winter for information
on populations of N. cismontana and their conser¬
vation status; Michael J. Donoghue, Geoffrey A.

Levin, and Michael G. Simpson for their inspiration,
thoughtful suggestions, encouragement, and insight
throughout the course of his thesis project; the mem¬
bers of the Biology Department at San Diego State
University and their Systematics and Evolutionary
Biology Program; and the San Diego Natural History
Museum for providing the necessary facilities and
equipment to complete this study. The senior author
is most appreciative of Floyd Swink, who reviewed
the manuscript for grammar and other errors. We
thank the unknown reviewers of this manuscript.
We gratefully acknowledge the curators of the her¬
baria for the loans of Nolina specimens.

Literature Cited

Department of the Interior, Fish and Wildlife Service.
1993. 50 CFR Part 17: Endangered and Threat¬
ened wildlife and plants; Review of plant taxa for
listing as Endangered or Threatened species. Federal
Register 58(188): 51144-51190. Thursday, Sep¬
tember 30, 1993.

Dice, J. C. 1988. Systematic Studies in the Nolina
bigelovii-N. parryi (Nolinaceae) Complex. M.S. the¬
sis, San Diego State University, San Diego, Califor¬
nia.

-. 1993. Nolina. P. 1203 in J. C. Hickman

(editor), The Jepson Manual: Higher Plants of Cali¬
fornia. Univ. California Press, Berkeley, California.
Lenz, L. W. 1950. Chromosome numbers of some west¬
ern American plants, I. Aliso 2(3): 317-318.
Munz, P. A. & J. C. Roos. 1950. California miscellany
II. Aliso 2(3): 217-238.

Watson, S. 1879. XV. Contributions to American bot¬
any. I. Revision of the North American Liliaceae.
Proc. Amer. Acad. Arts 14: 213-303.

Nomenclatural Changes in Polygonum, Persicaria, and
Aconogonon (Polygonaceae)

Harold Hinds

Biology Department, University of New Brunswick, Fredericton,

New Brunswick, Canada, E3B 5A3

Abstract. Polygonum caurianum subsp. hud-
sonianum is raised to species rank. Polygonum
bistortoides is transferred to Persicaria, and Po¬
lygonum alaskanum var. glabrescens is transferred
to Aconogonon.

For the forthcoming treatment of Polygonum s.l.
for the Flora of North America , several new no¬
menclatural combinations are required to reflect new
knowledge in the taxonomy of this group.

Polygonum caurianum B. Robinson subsp. hud-
sonianum S. J. Wolf & McNeill is raised to species
rank as Polygonum hudsonianum (S. J. Wolf &
McNeill) Hinds, and the new combinations Persi¬
caria bistortoides (Pursh) Hinds and Aconogonon
alaskanum (W. Wight ex Hulten) var. glabrescens
(Hulten) Hinds are made.

Polygonum hudsonianum (S. J. Wolf & McNeill)
Hinds, stat. et comb. nov. Basionym: Polyg¬
onum caurianum B. L. Robinson subsp. hud¬
sonianum S. J. Wolf & McNeill, Rhodora 88:
856. 1986. TYPE: Canada. Quebec, Fort
George, Baie James, 30 Aug. 1950, Lepage
12863 (holotype, DAO).

Polygonum hudsonianum forms part of a com¬
plex of species within section Polygonum that in¬
cludes P. fowled B. L. Robinson and P. caurianum
B. L. Robinson. Examination of relevant collections
has led me to the conclusion that P. hudsonianum
is more closely related to P. fowled than to P.
caurianum. There appears to be a clear distinction
between P. caurianum and P. hudsonianum based
partly on size but also on features of the perianth.

Table 1. Morphological comparison between Polygonum fowled , P. hudsonianum, and P. caurianum.

P. fowleri

P. hudsonianum

P. caurianum

Fruiting perianth

Size (mm)

3.0-4.5, mean 3.75

(2.3-)2.5-3.2(-3.5), mean
2.9

± pouched at base

1.6-2.3(-3), mean 2.3

Shape

broadly pouched

tapering to base of lobes

Color

light green

greenish to roseate

roseate

Lobes

cucullate; overlapping ex¬
cept late season ones

cucullate; ± overlapping

not cucullate; ± flat, flared
and curved outward

Achenes

Size (mm)

(2.5-)3.0-3.7(-4.0), mean
3.3

(1.8-)2.0-2.5(-3.1), mean
2.2

1.4-1.6(-2.2), mean 1.7

Shape, surface tex-

broadly ovate, beaked;

broadly ovate-lanceolate;

ovate-lanceolate; not or

ture, and cross

granular roughened; 2

± beaked; roughened to

only obscurely beaked;

section

equal and 1 broader
side, many biconvex

definitely striate-papil¬
late; variable in cross
section, many biconvex

rough to faintly striate-
papillate; mostly 1 nar¬
row and 2 broader sides

Included or
excluded

included except late season
ones all exserted

included or slightly exsert¬
ed; late season ones all
exserted

all exserted

Leaves

light to dark green or ±
reddish

reddish tinged

reddish to purplish tinged

Distribution

NE & NW coastal US &
Canada; S coastal Alas¬
ka; James Bay, W Hud¬
son Bay

centered E of Hudson Bay
& N of St. Lawrence
River, Quebec

centered in INW Alaska,
Yukon & NW North¬
west Territories

Novon 5: 165-166. 1995.

166

Novon

achene, and distribution (Table 1). Separation be¬
tween P.fowleri and P. hudsonianum, however, is
more difficult. Many characteristics are shared by
these two taxa, including the expanded base of the
perianth (pouched), cucullate three outer sepals, and
beaked achenes. As Table 1 indicates, however,
there are enough quantitative and qualitative dif¬
ferences between the two taxa upon which to rec¬
ognize P. hudsonianum as a species.

The description in Wolf & McNeill (1986) of
Polygonum caurianum subsp. hudsonianum should
be amended to include characteristics as indicated
in Table 1.

Persicaria bistortoides (Pursh) Hinds, comb. nov.
Basionym: Polygonum bistortoides Pursh, FI.
Amer. Sept., 1: 271. 1814. Bistorta bistor¬
toides (Pursh) Small, Bull. Torrey Bot. Club
33: 57. 1906. TYPE: [U.S.A. Idaho:] “Quam-
ash Flats, banks of the Missouri,” now called
Weippe Prairie, Clearwater Co., 20 June, M.
Lewis, 1806 (holotype, PH).

Persicaria (sect. Bistorta) bistortoides is closely
related to Persicaria bistorta (L.) G. Sampaio. It is
recognized based on the research of Ronse Decraene
& Akeroyd (1988), whose studies of floral mor¬

phology in the Polygonaceae support including the
Bistorta group in the genus Persicaria.

Aconogonon alaskanum (W. Wight ex Hulten)
Sojak var. glabrescens (Hulten) Hinds, comb,
nov. Basionym: Polygonum alaskanum W.
Wight ex Hulten var. glabrescens Hulten, Acta.
Univ. Lund. N.F. Avd. 2, 40(1) 4: 612. 1944.
TYPE: U.S.A. Alaska: Bonanaza Creek, 19
June 1924, Eastwood 298 (holotype, GH).

Chambers (1992) clarified the correct name in
Aconogonon for this species, but because he pre¬
ferred to include these taxa in Polygonum, left the
publication of this new combination to someone else.

Acknowledgment. I wish to acknowledge the con¬
siderable assistance of George H. Flanders, Jr.

Literature Cited

Chambers, K. L. 1992. Choosing the correct name for
Aconogonon (Polygonum sect. Aconogonon) in
Alaska. Rhodora 94: 319-322.

Ronse Decraene, L.-P. & J. R. Akeroyd. 1988. Generic
limits in Polygonum and related genera (Polygona¬
ceae) on the basis of floral characters. Bot. J. Linn.
Soc. 98: 321-371.

Wolf, S. J. & J. McNeill. 1986. Synopsis and achene
morphology of Polygonum section Polygonum (Po¬
lygonaceae) in Canada. Rhodora 88: 457-479.

Una Especie Nueva de Rhamnus (Rhamnaceae) del Cerro Salomon,
Sierra Atravesada, Oaxaca, Mexico

Mario Ishiki Ishihara

Department of Biological Science, Faculty of Science, Hiroshima University,
Kagami-yama 1-3-1, Higashi-hiroshima-shi, Hiroshima-ken, 724, Japan

RESUMEN. Se describe Rhamnus wendtii del Cer¬
ro Salomon, Sierra Atravesada, Oaxaca, Mexico,
como especie nueva que parece estar emparentada
a R. mcvaughii. Se conoce solamente de la localidad
tipo, de selvas bajas perennifolias y bosques cer-
canos.

Abstract. Rhamnus wendtii is described as a
new species from Cerro Salomon, Sierra Atravesada,
Oaxaca, Mexico, with apparent affinities to R.
mcvaughii. It is known only from the type locality,
in elfin forest and surrounding vegetation.

El Cerro Salomon se encuentra ubicado en el
limite entre los municipios de Santa Maria Chima-
lapa y San Miguel Chimalapa en el distrito de Ju-
chitan, Oaxaca, Mexico (Fig. 1), al noreste de la
Congregacion de Benito Juarez y aproximadamente
a 44 km en linea recta al norte de San Pedro
Tapanatepec, a 16°46'15" latitud norte y 94°11 '45"
longitud oeste. La altitud de la cresta es entre 1700
a 1950 m. Fisiograficamente el Cerro Salomon per-
tenece a la Sierra Madre de Chiapas, que se extiende
en forma paralela a la Costa del Pacifico y corre
desde el Istmo de Tehuantepec a traves de Chiapas
hasta Guatemala. Las rocas de la Sierra Madre de
Chiapas son muy antiguas, siendo plutones del pa-
leozoico, compuesto de rocas graniticas, granodio-
nticas y dioriticas, con rocas metamorficas aun mas
viejas de edades precambricas y paleozoicas; la si¬
erra ha estado elevada sobre el nivel del mar en
forma continua por lo menos desde principios del
Terciario (Ferrusquia-Villafranca, 1993). El Cerro
Salomon se encuentra en el extremo noroccidental
dentro de este macizo montanoso, ya cerca del Istmo
de Tehuantepec, formando parte de la Sierra Atra¬
vesada, en un area de suelos arenosos provenientes
de viejas rocas metamorficas (esquistos y gneis) (Fe¬
rrusquia-Villafranca, 1993; Lopez Ramos, 1974).

La cubierta vegetal del lugar esta constituido por
varios tipos de bosque mesofilo de montana altos y
bajos, con bosque de pino en los sitios mas expuestos;
en la parte superior del cerro se establecen dos
manchones de vegetacion de baja estatura (selva

baja perennifolia) rodeados por comunidades mas
altas. Ishiki (1988) publico una lista floristica de las
selvas bajas perennifolias del Cerro Salomon, pero
el listado de los bosques mesofilos que la rodean no
es completo. Dentro de las novedades encontradas
durante el trabajo de campo tenemos el siguiente.

Rhamnus (subgen. Frangula) wendtii Ishiki, sp.
nov. TIPO: Mexico. Oaxaca: Mpio. San Miguel
Chimalapa, Cima del Cerro Salomon, al NO de
la Congregacion Benito Juarez, ca. 44 km en
linea recta al N de San Pedro Tapanatepec,
filo del Cerro con selva baja perennifolia,
16°46'15"N, 94°11'45’'0, 1830 m, 16 abr.
1986 (ft, fr), Ishiki 1575 (holotipo, MEXU;
isotipo, CHAPA). Figura 2.

Ex affinitate R. mcvaughii L. A. & M. C. Johnston.
Cataphylla nula; stipulae persistentes 2-3 mm Iongae;
petioli 6.0-12.5 mm longi; laminae membranaceae, co-
riaceae, ellipticae, oblongae, (3.6-)5.8-7.0(-8) cm lon-
gae, (1.4-)1.9-2.3(-2.5) cm latae, (2.6-)3(-3.2)-plo lon-
giores quam latiores, basi cuneatae, apice acuminatae,
margine crenatae; nervis lateralibus utroque latera (4-)5-
6(-7) sub angulo circa (35-)40-50° abeuntibus, supra
ad costam laete pubescentes, subtus ad costam et nervos
laterales pubescentes pilis 0.4 mm longis; flores 5-meri;
pedicelli fructiferi demum 12-15(-16) mm longi; cupula
floralis sub fructa demum integra; drupa 6-7 mm dia-
metro; putamina 3.

Arbusto o arbol pequeno de 1 -2 m de alto, corteza
obscura un poco rojiza en las partes viejas; ramitas
con abundantes pelos adpresos de 0.4 mm de largo
en las partes jovenes y con pelos erectos de
(0.1-)0.15-0.2 mm de largo a glabras enlas partes
viejas, en la parte intermedia con ambos tipos de
tricomas; escamas de la yema ausente. Hojas alter-
nas, lamina membranosa a coriacea, eliptica a oblon-
ga, (3.6-)5.8-7.0(-8) cm de largo, (1.4-)1.9-
2.3(-2.5) cm de ancho, (2.6—)3(—3.2) veces mas
largo que ancho, base cuneada, apice acuminado,
margen crenado con (4-)5-6(-8) dientes dispuestos
en los dos tercios distales de la lamina, a intervalos
de (2.5-)4.0-8.5(-14.5) mm y 1 —3(—4) dientes por
cm, haz verde obscuro, glabro, ligeramente pubes-
cente en el nervio central, enves verde amarillento-

Novon 5: 167-170. 1995.

168

Novon

cafesoso, glabro, ligeramente pubescente con pelos
adpresos y aislados de 0.4 ram en el nervio central
y lateral, y sericeo con pelos de 0.4 mm de largo
en las axilas de los nervios, venacion eucamptodro-
ma con (4-)5-6(-7) pares de nervios laterales con
angulo de (35-)40-50° al nervio central; peciolo de
6-12.5 mm de largo, con abundantes pelos adpresos
de 0.4 mm de largo; estipulas de 2-3 mm de largo,
pubescentes, obscuras, persistentes. Flores 1 o 2 por
axila, pedunculo ausente o muy pequeno; pedicelo
de 11-12 mm de largo, pubescente, con pelos erec-
tos de 0.1 mm de largo y en ocasiones algunos pelos
adpresos de 0.5 mm de largo; copa floral de 1.3-
1.5 mm de largo, 2-2.5 mm de diametro, poco
pubescente; sepalos 5, deltoideos, triangulares, 1.5—
1.8 mm de largo, 1.4-1.5 mm de ancho en la base,
interior glabro, exterior con pelos aislados de 0.1-
0.2 mm de largo; petalos 5, cuculados y hendidos,
1-1.4 mm de largo, 1-1.5 mm de ancho, altemos
a los sepalos; estambres 5, filamentos de 0.5-0.85
mm de largo, triangulares con la base ancha (0.3—
0.4 mm), opuestos a los sepalos; anteras 0.8 mm
de largo, introrsas, dorsiiijas, cada una abrazada por
un petalo, con dehiscencia longitudinal; polen tri-
colpado, heteropolar con simetrfa bilateral, angu-
laperturado de 9.5 /zm de diametro ecuatorial, su-
perficie ondulada y perforada (foveolada) (Fig. 3);
ovario glabro, 1-1.25 mm de diametro, 0.7 mm de

alto, estilo 0.8-0.95 mm de largo, estigma trilobado.
Frutos 1 a 2 por axila; pedicelo en la fructificacion
de 12— 15(— 16) mm de largo; copa floral circular,
borde entero, sin restos del perianto, de 1.5-3 mm
de diametro, 1 mm de profundidad; fruto drupaceo,
globoso, glabro, de 6-7 mm de diametro en fresco,
cafe obscuro, casi negro al madurar (pasando por
un color rojo-vino), con 3 pirenos; mericarpios ehp-
tico-ovados, de 4-5 mm de largo, 4 mm de ancho,
endocarpio de 0.15-0.2 mm de grosor; semillas una
por mericarpio, testa negra, ligeramente fusionada
con el endocarpio, muy delgada; endosperma una
capa amarilla hasta de 0.3 mm de grosor; cotile-
dones suborbiculares, engordados, ca. 4 mm de lar¬
go, ca. 4 mm de ancho cada uno, ca. 0.6 mm de
grosor; germinacion criptocotilar y epigea.

Habitat. En selvas bajas perennifolias con Wein-
mannia glabra L.f., Gaultheria odorata Kunth,
Rapanea juergensenii Mez, R. myricoides (Sch-
lechtendal) Lundell, Cavendishia bracteata( Ruiz &
Pavon ex J. St.-Hilaire) Hoerold, Satyria warsze-
wiczii Klotzsch, Disterigma, Clethra, Liquidambar
y abundantes lauraceas y melastomataceas; con es-
trato superior abierto de Pinus oocarpa Schiede ex
Y. Schechter var. trifoliata Martinez. Suelo limo-
arenoso, con abundante hojarasca y musgo (Sphag¬
num), y altitudes de 1800 a 1950 m.

Volume 5, Number 2
1995

Ishiki

Rhamnus wendtii

169

Figure 2. Rhamnus wendtii Ishiki. —A. Rama con inflorescencia. —B. Detalle de la rama y de la estipula. — C.
Flor con los sepalos y petalos anteriores parcialmente eliminados. —D. Rama con fructificacion. E. Fruto maduro
con la copa floral circular. —F. Corte transversal del fruto que muestra las tres semillas y semilla (A, Ishiki 1527 ;
B-C, Ishiki 1338 ; D-F, Ishiki 1575).

170

Novon

Figura 3. Microfotografia electronica de barrido (SEM)
del grano de polen de Rhamnus wendtii Ishiki (barra =
3.2 /un).

Distribution. Unicamente se ha localizado en la
localidad tipo, en comunidades de bosque de pino-
encino y selva baja perennifolia. Por ser una region
muy poco conocida botanicamente, es posible que
existen otras poblaciones en otros cerros cercanos
de Oaxaca y Chiapas.

Paratipos. MEXICO. Oaxaca: Mpio. San Miguel Chi-
malapa, Localidad tipo, 11 feb. 1986 (fl), Ishiki 1263 ;
(fr), Ishiki 1338; 16 abr. 1986 (fr), Ishiki 1466; (fl, fr),
Ishiki 1506; (fl, fr), Ishiki 1527; (fr), Ishiki 1545; (fl),
Ishiki 1603; (fr), Ishiki 1644; 20 ago. 1986 (fr), Ishiki
1710; 15 oct. 1986 (fl, fr), Ishiki 1791 (MEXU, CHAPA).

Siguiendo la clasificacion de Wolf (1938), Grubov
(1949), Suessenguth (1953) y Johnston & Johnston
(1978), R. wendtii pertenece al subgenero Fran-
gula por la ausencia de escamas en la yema, inflo-
rescencia una umbela con pedunculo o sesil, flores
perfectas, la presencia de sepalos, frutos de color
negro o purpura al madurar, fruto indehicente, y
endosperma escaso o ausente. Johnston & Johnston
(1978) no reconocen grupos de especies o series
dentro de las especies Neotropicales de Rhamnus
subgenero Frangula y consideran que forman una
unidad filetica compacta y la necesidad de realizar
mas estudios para dilucidar el problema. La especie
mas emparentada probablemente es R. mcvaughii,
especie distribuida desde Hidalgo hasta el centro de
Oaxaca. Rhamnus mcvaughii difiere de la nueva
especie en las hojas mas pequenas (1.6-5.5(-7.5)
x 0.8-1.9(-2.6) cm) con apice frecuentemente ob-
tuso o agudo, las venas secundarias mas numerosas
(6-12 pares) formando un angulo mas abierto (may¬
or de 50°), y con mayor numero de dientes mar-

ginales por centimetre ((4-)5-8(-9)). Habita alti¬
tudes superiores a los 2000 msnm.

En la clave de Johnston y Johnston (1978), la
especie nueva frecuentemente se determinaria como
R. mucronata Schlechtendal, especie distribuida
desde Michoacan hasta El Salvador, en matorrales,
bosque de pino-encino, bosque de oyamel, a altitudes
de (1500-)1900-2800(-3000) m. Rhamnus mu¬
cronata difiere de la especie nueva en los pedicelos
mas cortos (5-7(-13) mm) y las hojas ligeramente
mas grandes (1.9—8.5(— 13) x (0.7-)0.9-3.2(-4)
cm) con venas laterales mas numerosas (6— 10(—13))
en un angulo mas abierto y con mayor numero de
dientes marginales por centimetro (3-6(-8)), y la
inflorescencia con mayor numero de flores (1-
5(-10)).

El epiteto de la especie nueva honra a Thomas
L. Wendt, estudioso de la Flora Tropical de Mexico,
quien fungio en una parte importante de mi for-
macion botanica.

Agradecimientos. Agradezco a Thomas L. Wendt
y a Jerzy Rzedowski R., sus valiosos comentarios;
a Jorge Valdez Carrasco por la toma de fotografia
de los granos de polen en el microscopio electronico
de barrido del Centro de Entomologia y Acarologia
del Colegio de Postgraduados; a Eduardo Merino
del Centro de Estudios del Desarrollo Rural del Co¬
legio de Postgraduados y a Erika Ishiki, la ilustracion
de la nueva especie; y a CONACyT por el apoyo
financiero para el trabajo de campo, a traves del
proyecto Flora Mesoamericana (Colegio de Postgra¬
duados).

Literatura Citada

Ferrusquia-Villafranca, I. 1993. Geology of Mexico: A
Synopsis. Pp. 3-107 in T. P. Ramamoorthy, R. Bye,
A. Lot & J. Fa (editores), Biological Diversity of
Mexico: Origins and Distributions. Oxford Univ. Press,
New York.

Grubov, V. I. 1949. Rhamnales. Pp. 485-516 in B.
K. Shishkin (editor), Flora of the U.S.S.R. Volume
14. Moskva-Leningrad.

Ishiki I., M. 1988. Las Selvas Bajas Perennifolias del
Cerro Salomon, Region de Chimalapa, Oaxaca: Flora,
Comunidades y Relaciones Fitogeograficas. Tesis de
Maestria. Centro de Botanica, Colegio de Postgrad¬
uados, Chapingo, Mexico.

Johnston, M. C. & L. A. Johnston. 1978. Rhamnus.
Flora Neotropica 20: 1-96.

Lopez Ramos, E. 1974. Carta Geologica del Estado de
Oaxaca, 2° Ed. [Provisional]. Instituto de Geologia,
U.N.A.M., Mexico, D.F.

Suessenguth, K. 1953. Rhamnaceae. Pp. 7-173 in A.
Engler & K. Prantl (editores). Die Natiirlichen Pflan-
zenfamilien. Band 20D. Wilhelm Engelmann, Leip¬
zig- . r

Wolf, C. B. 1938. The North American species ol
Rhamnus. Rancho Santa Ana Bot. Gard. Monogr.,
Bot. Ser. 1: 1-136.

New Species and Combinations in Millettia Wight & Arnott and Pon-
gamiopsis R. Viguier (Leguminosae-Papilionoideae-Millettieae) from

Madagascar

Jean-Noel Labat

Museum national d’Histoire naturelle, Laboratoire de Phanerogamic, UMS 826 CNRS-MNHN,
16 rue Buffon 75005 Paris, France

David J. Du Puy

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, England

ABSTRACT. Morphological characters support the
description of five new species of Millettia, M. ca-
puronii Du Puy & Labat, M. hitsika Du Puy &
Labat, M. nathaliae Du Puy & Labat, M. orientalis
Du Puy & Labat, and M. taolanaroensis Du Puy
& Labat, and one new species of Pongamiopsis, P.
viguieri Du Puy & Labat, from Madagascar. The
genus Neodunnia R. Viguier is considered to be a
synonym of Millettia, and two new combinations,
M. aurea (R. Viguier) Du Puy & Labat and M.
richardiana (Baillon) Du Puy & Labat, are made.

R£sum£. L’etude des caracteres morphologiques
permet la description de cinq nouvelles especes de
Millettia: M. capuronii Du Puy & Labat, M. hit¬
sika Du Puy & Labat, M. nathaliae Du Puy &
Labat, M. orientalis Du Puy & Labat, et M. tao¬
lanaroensis Du Puy & Labat, et une nouvelle espece
de Pongamiopsis, P. viguieri Du Puy & Labat, de
Madagascar. Le genre Neodunnia R. Viguier est
considere comme etant un synonyme de Millettia
et deux nouvelles combinaisons, M. aurea (R. Vi¬
guier) Du Puy & Labat and M. richardiana (Baillon)
Du Puy & Labat, sont faites.

Prior to completion of an account of the subfamily
Papilionoideae in Madagascar, six new species are
described and illustrated and two new combinations
are made in the closely related genera Millettia and
Pongamiopsis.

Millettia Wight & Arnott, with about 100 spe¬
cies, is one of the largest genera of the tribe Mil-
lettieae. The genus is mainly paleotropical, occurring
in Asia and Africa, with only about five species in
California and Mexico (formerly regarded as a sep¬
arate genus Hesperothamnus T. S. Brandegee, see
Geesink, 1984: 102-106). Prior to the present pub¬
lication, only one variable species, M. lenneoides
Vatke, a small tree, shrub, or liane was recognized

in Madagascar: a total of eight species, all endemic,
are now known to occur on the island. They are
widely distributed throughout lowland Madagascar,
but do not occur on the Central Plateaux.

Hutchinson (1964) considered Neodunnia R. Vi¬
guier to be a synonym of Millettia, but it was
regarded as a separate genus by Geesink (1981,
1984) on the basis of its solitary flowers produced
from the bud scales or lowest nodes of new, emerging
shoots. In fact, in M. (Neodunnia) richardiana,
the flowers are produced in a cluster of 2-4 on a
very short peduncle, the inflorescence being a re¬
duced pseudoraceme. The second species, M. (Neo¬
dunnia) aurea, has a similar inflorescence but with
the axis somewhat elongated and the flowers in two
or three distant clusters: this species and the newly
described M. nathaliae are intermediate with the
main group of species with multi-flowered pseudor¬
acemes. These species are otherwise very similar to
other, more typical, Millettia species in Madagas¬
car, and Neodunnia is not considered here to be
distinct.

Pongamiopsis R. Viguier is a genus of three
species endemic to Madagascar (including the new
species P. viguieri, described here). It is closely
allied to Millettia, differing in its very swollen, el¬
lipsoid pods, which do not taper toward the base,
and its seeds, which are spherical or reniform, not
flat and discoid as in Millettia species. The seeds
also lack the rim aril extended into a short flap on
one side, which is characteristic of Millettia in Mad¬
agascar.

Millettia capuronii Du Puy & Labat, sp. nov.
TYPE: NE Madagascar. [Near Sambava], S
de la basse Bemarivo, versant oriental de l’Am-
batobiribiry, 100-330 m, 9 Avr. 1967 (fl).
Service Forestier de Madagascar 27670-SF,
R. Capuron (holotype, P; isotypes, K, L, MO,
P, TEF). Figure 1.

Novon 5: 171-182. 1995.

172

No von

Figure 1. Millettia capuronii Du Puy & Labat. —A. Habit. —P, C. Leaflet undersurface. — D. Flower. —E.
Calyx. F. Young pod. G. Pod. —H. Seed. (A F drawn lV.>ni Service Forestier de Madagascar SF 27670, and
G and H from Service Forestier de Madagascar SF 10027. )

Species apice foliolorum contracto, foliis et leguminibus
glabris, et inflorescentiis cum axibus secundariis brevibus
et gracilibus pares apicales florum gerentibus distincta.

A deciduous tree 4-8 m tall, flowering on young,
developing shoots; [DBH not known]; twigs glabrous
wih raised lenticels and small, spherical dormant
buds. Leaves with 5-11 opposite leaflets; rachis
glabrous; stipels absent. Leaflets ovate, 25-55 x
15-40 mm, obtuse to rounded basally, the apex
acuminate and tapering into a short to long drip tip,

finally obtuse and mucronate, entirely glabrous, pal¬
er beneath. Pseudoracemes delicate, ca. 4-6 cm
king, from the leaf axils near the base of new growths,
the flowers toward the base of the inflorescence
paired on short, slender secondary axes up to 6 mm
lung; pedicels with a pair of minute, filiform brac-
teol B s pear the middle which are caducous before
the flower opens. Flowers ca. 12-15 mm long, violet.
Calyx 3~4 mm long, subglabrous with a minutely
c iliate margin, with 5 short, broadly triangular teeth,

Volume 5, Number 2
1995

Labat & Du Puy
Millettieae from Madagascar

173

the upper pair smallest and close together. Standard
limb ca. 10 x 10 mm, cuneate basally, finely and
sparsely appressed pubescent behind. Wings as long
as the keel. Staminal sheath ca. 9 mm long. Pods
obovate to oblong-obovate, flat, 65-80 x 15-20
mm, tapering toward the base, the apex rounded
and beaked, glabrous, splitting into 2 twisted, woody
valves, with 1-4 seeds. Seeds oblong-discoid, ca. 10
x 10 mm, chestnut brown, the aril extended into
a short flap on one side.

Millettia capuronii is recognizable by its tapering
leaflet apices, its glabrous leaves and pods, and its
distinctive inflorescences with short, slender sec¬
ondary axes bearing apical pairs of flowers.

Distribution. Eastern Madagascar, collected in¬
frequently but in widespread localities along the east¬
ern coastal region including Fort Dauphin (Taolan-
aro), Farafangana, and Sambava.

Habitat. Lowland evergreen forest near the coast,
at low altitudes.

Flowering time. Only recorded in April.

Vernacular name. Sikidihazo.

This species is dedicated to Rene Capuron, col¬
lector of the type specimen, in recognition of his
outstanding contributions to the knowledge of the
forest flora of Madagascar.

Paratypes. MADAGASCAR. Prov. de Toliara, Pre¬
fecture Taolaiiaro (Fort Dauphin), foret a 5 km de Ste.
Luce, au N de Maliaforaky, 24°46'S, 47°09'E, 0-10 m,
28 Avr. 1989(st), Rabevohitra, Dumetz & Gereau 1946
(MO, P, TAN); Prov. de Toliara, Prefecture de Fort
Dauphin, foret ombrophile de Manatantely, 50-250 m,
9 Nov. 1990 (jfr), Rabevohitra 2426 (MO, P, TAN);
Dct. de Farafangana, Andafia, 20 Avr. 1953 (fr), Service
Forestier de Madagascar 10027-SF (P, TEF).

Millettia hitsika Du Puy & Labat, sp. nov. TYPE:
E Madagascar. Foret d’Analalava, Fenoarivo,
4 km W de Foulpointe, 20—50 m, 3 Dec. 1958
(fl). Service Forestier de Madagascar 20154-
SF, R. Capuron (holotype, P; isotypes, K, L,
MO, P, TEF). Figure 2.

Species habitu tenui et delicato et foliolis cum apice
acuto, supra glabris et subtus appresso-pubescentibus dis-
tincta; a M. orientalis differt foliis, foliolis, inflorescentibus
at floribus magnopere parvioribus.

A deciduous tree or shrub to 10 m tall, flowering
on new, developing shoots soon after they emerge
from the buds; DBH 18 cm (one record only); twigs
brownish pubescent when young, glabrescent with
rounded dormant buds when mature. Leaves with
9-13 opposite leaflets; rachis pubescent when young;
stipels absent. Immature leaflets narrowly elliptic,
[mature leaflets not known], acute and mucronate

apically, glabrous except for the midvein above,
densely covered in short, appressed hairs beneath.
Flowers in delicate pseudoracemes 4-9.5 cm long
from the lowest nodes and leaf axils on very young
shoots with developing leaves, the flowers solitary
or paired on very short side spurs up to 1.5 mm
long; pedicels with a pair of very small, filiform
bracteoles up to 3 mm long which are caducous
before the flower opens. Flowers 10-12 mm long,
[pink or violet]. Calyx 2.5-4 mm long, brownish
pubescent, with 5 short, broadly triangular teeth,
the upper pair smallest and close together. Standard
limb ca. 10 x 10 mm, thinly pubescent behind.
Wings as long as the keel. Staminal sheath 8-9 mm
long. [Pods and seeds not known.]

Millettia hitsika is characterized by its fine, del¬
icate habit and by its narrow (immature) leaflets with
acute apices, which are glabrous above and ap-
pressed-pubescent beneath. It is only known in flow¬
er with immature foliage, in which it somewhat re¬
sembles Pongamiopsis pervilleana (Baillon) R. Vi-
guier. It is easily distinguished from M. orientalis,
the only other Millettia species that occurs in the
humid, eastern forests, a much more robust species
with much larger leaves, leaflets, inflorescences, and
flowers.

Distribution. Eastern Madagascar, on the coastal
plain, from Ambila-Lemaitso to Foulpointe.

Habitat. In evergreen forest on sand, at up to
ca. 50 m altitude.

Flowering time. October-March.

Local uses. The wood is used in construction.

Vernacular name. Hitsika.

The species name is derived from the vernacular
name for this species.

Paratypes. MADAGASCAR. Ambila Lemaitso, 13 Mar.
1951 (fl), Service Forestier de Madagascar 3260-SF
(K, P, TEF); Ambila Lemaitso, 15 Jan. 1953 (fl). Service
Forestier de Madagascar 7257-SF (P, TEF); Dct. de
Brickaville, Ambila Lemaitso, sous parcelle F2K, 20 Fev.
1958 (fl), Service Forestier de Madagascar 19002-SF
(P, TEF).

Millettia nathaliae Du Puy & Labat, sp. nov.
TYPE: N Madagascar. Track to Irodo along
the northern ridge above the Irodo River, ca.
5 km E of Route Nationale 6 to Antsiranana
(Diego-Suarez), the nearest village Andrano-
mena, Sarainana River, 12°37'S, 49°26'E, ca.
235 m, 26 Nov. 1992 (fl, fr), D. J. Du Puy,
G. Lewis & B. Schrire M564 (holotype, K;
isotypes, P, MO, TAN). Figure 3.

174

Novon

Figure 2. Millettia hitsika Du Puy & Labat. —A. Habit. —B. Leaflet upper surface. —C. Leaflet lower surface.
—D. Flower. —E. Calyx. (A-E drawn from Service Forestier de Madagascar SF 20154.)

Species M. richardiana affinis, inflorescentia elongata
cum fasciculis pluribus florum, floribus minoribus, pedi-
celbs multo brevioribus, et leguminibus glabris.

A shrub or small tree ca. 3-15 m tall, flowering
on new shoots as they emerge from the buds and
whilst the leaves are still very immature, DBH up
to 40 cm; bark smooth with shallow vertical fissures,
dark gray; young shoots densely golden pubescent;
mature twigs glabrous, gray, with pale lenticels and
large spherical dormant buds. Leaves with (13-) 15-
25 opposite leaflets; rachis slightly pubescent; stipels
absent. Leaflets narrowly oblong-elliptic, 13-37 x
6-11 mm, with a long apical mucro (up to ca. 2
mm long), subcoriaceous and with decurved mar¬
gins, densely golden pubescent when young, becom¬
ing glabrous when mature except for the midvein
beneath. Pseudoracemes l-3.5(-5.5) cm long, from
the axils of very immature leaves, golden-velvety
pubescent when still in flower, the axis often angular.

the flowers in 1-3 clusters, often paired, but the
bottom group may have 2-4 flowers; pedicels short,
2-4 mm long, with a pair of linear bracteoles ca.
3 mm long, which are caducous as the flowers open.
Flowers 8-10 mm long, bright fuchsia pink fading
to pale mauve, the standard pale green at the base.
Calyx 4.5-5 mm long, pale pubescent with occa¬
sional brown hairs, purple, with 3 narrowly trian¬
gular lower teeth with slender, tapering tips, as long
as or shorter than the calyx cup, and with 2 small,
very closely spaced upper teeth. Standard limb ca.
6-8 x 8-10 mm, shallowly notched at the apex,
white pubescent behind. Wings longer than the keel,
both slightly pubescent on the outer surfaces. Sta-
minal sheath ca. 7 mm long. Pod obovate to oblong,
flat, 40-70 x 1 6- 1 8 mm, tapering toward the base,
rounded and with a decurved beak apically, gla¬
brous, splitting into two woody valves with l - 3
seeds. Seeds disc-shaped, 8-9 x 8—9 mm, finely

Volume 5, Number 2
1995

Labat & Du Puy
Millettieae from Madagascar

175

D M-leUa a*rtdM. v A

Figure 3. Millettia nathaliae Du Puy & Labat. —A. Flowering stem. —B. Leaf. —C. Flower. —D. Calyx.
E. Pod. —F. Seed. (A-F drawn from Du Puy , Lewis & Schrire M564.)

mottled dark brown, the aril extended into a short
flap on one side.

Millettia nathaliae is closely related to M. ri-
chardiana but is distinguished by its elongated in¬

florescence with several clusters of flowers, its small¬
er flowers, its much shorter pedicels, and its glabrous
pods. It also closely resembles Pongamiopsis amyg-
dalina (Baillon) R. Viguier when in flower and before

176

Novon

the leaflets develop, but its much smaller flowers are
characteristic.

Distribution. Northern Madagascar, Ankarana
Massif, around Diego-Suarez and Sahafary forest
north of the Irodo River.

Habitat. In deciduous woodland and shrubland,
especially on limestone, but also on sand, at ca.
200-250 m altitude.

Flowering time. Only recorded in November.

This species is dedicated to Nathalie Labat.

Paratypes. MADAGASCAR. Ankarana Massif, SE
margin, about 25 km NE of Ambilobe, near the village
of Ambilomagodro, 13 o 01'S, 49°07'E, 200 m, 17 Nov.
1992 (fl), Du Puy, Lewis & Schrire M559 (K, MO, P,
TAN); Foret d’Andranomavo, Canton d’Anamakia, Dis¬
trict de Diego-Suarez, 24 Juin 1951 (fr), Service Forestier
de Madagascar 3596-SF (K, P, TEF); Bassin de la
Saharaina, foret de Sahafary, 27 Nov. 1958 (fl), Service
Forestier de Madagascar 20112-SF, Capuron (K, MO,
P, TEF).

Millettia orientalis Du Puy & Labat, sp. nov.
TYPE: E Madagascar. Baie d’Antongil, Nosy
Mangabe, 5 km from Maroansetra, 15°33'S
49°46'E, 0-330 m, 9 Jan. 1989 (fl), G. Schatz
& J. Miller 2495 (holotype, MO; isotypes, K,
P, TAN). Figure 4.

Species habitu robusto ramulis crassis et gemmis mag-
nis, foliis longis (ca. 25-30 cm), foliolis numerosis (11-
21) grandibus et inflorescentiis longis (9-20 cm) densis
floribus numerosis grandibus (15-20 mm longis) distincta.

A large, deciduous tree ca. 15-30 m tall, flow¬
ering on new, developing shoots; [DBH not known];
young shoots densely velvety-pubescent, becoming
glabrous and gray, the older twigs thick and blunt-
ended with large, subspherical dormant buds. Stip¬
ules very large and conspicuous on emerging shoots,
obovate, ca. 20-25 x 8-10 mm, soon caducous.
Leaves large, ca. 25-30 cm long, with (11—)15—
21 opposite or subopposite, subequal leaflets; rachis
velvety-pubescent when young; stipels absent. Leaf¬
lets very large, oblong-elliptic to obovate, usually
narrow, 60-110 x 20—35 mm, acuminate apically,
obtuse to rounded basally, glabrous above, densely
velvety beneath especially while young. Flowers in
robust pseudoracemes 9-20 cm long from the axils
of developing leaves near the base of new shoots,
the flowers often paired and the lower ones on short
secondary axes up to 5 mm long; pedicels with 2
slender, soon caducous bracteoles ca. 4 mm long.
Flowers 15-20 mm long, pink fading to pale lav¬
ender or mauve, the standard green at the base,
sweetly fragrant. Calyx 4-7 mm long, densely pu¬
bescent, purplish, with 3 broadly triangular lower

teeth, the upper pair short and much reduced. Stan¬
dard limb 11-15 X 12-14 mm, pubescent behind.
Wings as long as the keel, finely pubescent outside.
Staminal sheath 14-17 mm long, the free portion
of the filaments rather long. Pods narrowly oblong
to obovate, flat, ca. 70-150 x 20-30 mm, slightly
thickened along both margins, velvety pubescent,
splitting into 2 twisted, woody valves with 2-6 seeds.
Seeds oblong to discoid, flat, 13-16 x 8-11 mm,
chestnut brown, the aril extended into a short flap
on one side.

Millettia orientalis is the only Millettia that
forms a canopy tree in the humid eastern forests.
It is characterized by its robust habit with thick twigs
and large buds, its long leaves with many large
leaflets, and its long, dense inflorescences with nu¬
merous large flowers.

Distribution. Known from around the Bay of
Antongil and in the region of Taolaiiaro (Fort Dau¬
phin).

Habitat. In lowland humid evergreen forest, up
to ca. 300 m altitude.

Flowering time. October-January.

Vernacular name. Fanamoakondro.

The specific epithet indicates the provenance of
this species, in the eastern forests of Madagascar.

Paratypes. MADAGASCAR. Prov. of Toliara, Fort
Dauphin [Taolaiiaro] Region, NW of town near road to
Andohahela Reserve, along trail W of road, 24°46'S,
46°53'E, 100-200 m, 11 Nov. 1989 (fl), McPherson
14456 (K, MO, P, TAN); Emafotra, Mahatalaky, Fort
Dauphin, 27 Mai 1954 (fr). Service Forestier de Mad¬
agascar 10084-SF (K, MO, P, TEF); Beandri, Maha¬
talaky, Fort Dauphin, 23 Oct. 1954 (st), Service Forestier
de Madagascar 11139-SF (P, TEF); Amboangy, Ma¬
hatalaky, Fort Dauphin, 20 Oct. 1955 (fl), Service Fo¬
restier de Madagascar 14861-SF (K, MO, P, TEF); E
Madagascar, environs de la Baie d’Antongil, massif de
Farankaraina, entre Navana et Andranofotsy, 0-150 m,
16 Sep. 1957 (fr). Service Forestier de Madagascar
18307-SF, Capuron (K, P, TEF).

Millettia taolanaroensis Du Puy & Labat, sp.
nov. TYPE: SE Madagascar. SW de Fort Dau¬
phin [Taolaiiaro], Baie de Ranofotsy, colline
pres d’ltaly, 10 Jan. 1963 (fl). Service Fores¬
tier de Madagascar 22367-SF, R. Capuron
(holotype, P; isotypes, K, MO, P, TEF, WAG).
Figure 5.

Species foliolis subglabris nitidis obtusis ad rotundatis
subtus cinerascentibus, pubescentia aperta pilis nitidis cu-
preis brevibus praesertim in rachidi, petiolulis, axe inflo-
rescentiae, pedicellis et calycibus, et leguminibus glabris
secus marginem superiorem incrassatis distincta.

Volume 5, Number 2
1995

Labat & Du Puy
Millettieae from Madagascar

177

Figure 4. MiUettia orientals Du Puy & Labat. -A. Habit. -B. Leaflet. -C. Leaflet undersurface. -R Young
stem with stipules, leaf and inflorescence. —E. Flower. —F. Calyx. —G, H. Pod. I. Seed. (A, E, and F drawn
from Schatz & Miller 2495; B and C from Service Forestier de Madagascar SF 18307; D from Service horestier
de Madagascar SF 11139; G-I from Service Forestier de Madagascar SF 10084.)

178

Novon

Figure S. Millettia taolanaroensis Du Puy & Labat. —A. Habit. —B. Petiole and first leaflet pair. —C. Leaflet
apex. —D. Leaflet undersurface. —E. Flower. —F. Calyx. —G, H. Pod. —I. Seed. (A, C-F drawn from Service
Forestier de Madagascar SF 22367 ; B and G from Phillipson & Milijaona 3526 ; H and I from Service Forestier
de Madagascar SF 8503.)

A deciduous shrub or small tree 4-6 m tall,
flowering on young shoots with fully expanded leaves;
twigs sparsely pubescent when young, glabrescent,
with small dormant buds. Leaves with 7-11 opposite
leaflets; rachis thinly coppery-pubescent; stipels ab¬
sent. Leaflets elliptic, 12-30(-36) x 9-19 mm,
rounded basally, obtuse or rounded to notched and
mucronulate apically, dark green, glossy and gla¬
brous above, gray-green and almost glabrous with
very scattered, short brown hairs beneath. Flowers
in delicate pseudoracemes ca. 4-6 cm long from

the axils of leaves near the base of new shoots, the
flowers solitary or paired on small knobs; pedicels
with 2 slender, soon caducous bracteoles near the
middle. Flowers 10-12 mm long, mauve or violet.
Calyx 3-4 mm long, densely coppery-pubescent,
with 5 short, broadly triangular teeth, the upper
pair smallest and close together. Standard limb ca.
9x9 mm, cuneate basally, finely pubescent behind.
Wings as long as the keel. Staminal sheath ca. 8
mm long. Pods obovate to oblong, flat, (40-)50-90
x (12—)14— 19 mm, tapering basally, the apex

Volume 5, Number 2
1995

Labat & Du Puy
Millettieae from Madagascar

179

rounded and beaked, slightly thickened along the
upper margin, glabrous or with very scattered brown
hairs, splitting into 2 twisted, woody valves, with 1-
5 (usually 1-3) seeds. Seeds oblong-discoid, 9-10
x 7-8 mm, chestnut brown, the aril extended into
a small flap on one side.

Millettia taolanaroensis is recognizable by its
subglabrous, glossy, obtuse to rounded leaflets, which
are grayish beneath, by its thin pubescence of lus¬
trous, copper-colored short hairs most noticeable on
the leaf rachis and petiolules, the inflorescence axis
and pedicels and the calyces, and by its glabrous
pods thickened along the upper margin. It appears
to flower when the leaves are almost full-sized rather
than as new growth emerges from the buds.

Distribution. Southeastern Madagascar, only
known from around Taolafiaro (Fort Dauphin), on
the coastal plains to the north and on the lower
slopes of the mountains to the west (including An-
dohahela, parcelle 3).

Habitat. In coastal, evergreen forest on sand,
and in transitional, deciduous woodland often near
the coast.

Flowering time. Only recorded in January.

Local uses. Used as a fish poison.

Vernacular name. Anakaraka.

The specific epithet indicates the provenance of
this species, in the southern region of Madagascar,
near the town of Taolafiaro.

Paratypes. MADAGASCAR. Maravato pres de Fort
Dauphin, Avr. 1972 (fr), Boiteau s.n. (P); Prov. de Fort
Dauphin, Andrahomana, 24 Juin 1926 (fr), Decary 3993
(P); Baie des Galions (Ranofotsy) au SW de Fort Dauphin,
1-100 m, 18-21 Fev. 1955 (fr), Humbert & Capuron
29019 (L, MO, P); Falaise cohere NW du Cap Sainte-
Marie, vers Lavanono, 1-150 m, 8 Mar. 1955 (fr),
Humbert & Capuron 29323 (K, P); Fort Dauphin region,
W of town along highway on southern boundary of Re¬
serve Integrale no. 11, Andohahela, parcel 3, dry forest
W of Col des Ranopiso, 120-140 m, 27 Jan. 1990 (fr),
McPherson & Pigeon 14911 (K, MO, P, TAN); 5 km
S of Manambaro, 23 km W of Fort Dauphin, 25°05'S,
46°49'E, 150 m, 30 Mar. 1991 (fr), Miller & Randri-
anasolo 6 224 (K, MO, P, TAN); Reserve d'Andohahela,
parcelle 3, 25°02'S, 46°40'E, 100 m, 17 Feb. 1990 (fr),
Phillipson & Milijaona 3526 (K, MO, P, TAN); Pre¬
fecture de Fort Dauphin, Petriky, 25°05'S, 46°51'E, 0-
10 m, 16 Oct. 1989 (fr), Rabevohitra 2018 (K, MO,
P, TAN); Mandena, Fort Dauphin, 8 Avr. 1953 (fr).
Service Forestier de Madagascar 8172-SF (P, TEF);
Bords d'un cours d'eau. aux environs de Ranopiso, route
Ambovombe-Fort Dauphin, 23 Sep. 1953 (fr). Service
Forestier de Madagascar 8503-SF, Capuron (K, MO,
P> TEF); entre km 454 et 455 de la route de Fort
Dauphin, 19 Sep. 1954 (fr), Service Forestier de Mad¬
agascar 10911-SF (P, TEF).

Millettia aurea (R. Viguier) Du Puy & Labat,
comb. nov. Basionym: Neodunnia aurea R.
Viguier, Notul. Syst. (Paris) 14: 73, 1950.
TYPE; (based on three collections) W Mada¬
gascar. Ambongo, Manongarivo [S of Soalala],
Oct. 1903 (fl, jfr), H. Perrier de la Bathie
1617 (lectotype, chosen here, P; isolectotypes,
K, L, MO, P, TAN).

R. Viguier cited three syntypes in the original
description, including H. Perrier de la Bathie 4831
and 4879, in addition to the lectotype selected here.

Millettia aurea may be confused with M. ri-
chardiana, which differs in its much shorter inflo¬
rescences with a single cluster of (larger) flowers,
its smaller and less pubescent calyx with shorter
teeth, especially the upper pair, which are very short
and connate to near the tip, its much less pubescent
petals, and its usually smaller leaflets.

Millettia aurea is an uncommon species of west¬
ern Madagascar, with a restricted distribution in the
Boina and the Ambongo, south to Maintirano, most
frequently recorded from the region of Ankara-
fantsika, and from Soalala to Mitsinjo. It occurs in
deciduous woodland, on sand, often near the coast
but also inland at up to ca. 300 m altitude.

Millettia richardiana (Baillon) Du Puy & Labat,
comb. nov. Basionym: Mundulea richardiana
Baillon, Bull. Soc. Linn. Paris 1: 389-390,
1883, pro parte. TYPE: (based on three col¬
lections) N Madagascar. [Diego-Suarez (Antsi-
ranana)], Baie des Amis (11), C. Richard 158
(lectotype, chosen here, P).

Mundulea hysterantha Baker, J. Linn. Soc., Bot. 25:
309, 1890. Syn. nov. TYPE: W Madagascar. An-
drona (fl, jfr), R. Baron 5444 (holotype, K; isotype,

P).

Millettia baroni Drake in Grandidier, Hist. Phys. Nat.
Pol. Madagascar, Plantes, Texte 1: 141, 1903. Syn.
nov. TYPE: Madagascar (fr), R. Baron 4952 (ho¬
lotype, P; isotypes, K, P).

Neodunnia atrocyanea R. Viguier, Notul. Syst. (Paris)
14: 72, 1950. Syn. nov. TYPE: NW Madagascar.
Riviere de Mananjeba, Oct. 1909 (fl), H. Perrier
de la Bathie 4845 (holotype, P; isotype, P).
Neodunnia edentata R. Viguier, Notul. Syst. (Paris) 14:
73, 1950, pro parte. Syn. nov. TYPE: (based on
four collections) NW Madagascar. Majunga, Dec.
1924 (fl), H. Perrier de la Bathie 16800 (lectotype,
chosen here, P; isolectotypes, K, P, MO).

H. Baillon cited three syntypes in the original
description of Mundulea richardiana, including
Boivin 2712 and 2713, in addition to the lectotype
selected here. Of the two Boivin specimens (both
mounted on the same herbarium sheet) only 2712

180

Novon

corresponds with M. richardiana; Boivin 2713 is
Pongamiopsis amygdalina (Baillon) R. Viguier.

In the original description of Neodunnia eden-
tata, R. Viguier cited four syntypes: H. Perrier de
la Bathie 1601 is a specimen of Pongamiopsis
pervilleana (Baillon) R. Viguier; H. Perrier de la
Bathie 4076, 16800, and Louvel 53 are all spec¬
imens of Millettia richardiana. H. Perrier de la
Bathie 16800 is composed of two elements: the first
was collected in December 1924, in flower, and is
selected here as the lectotype; the second was col¬
lected in January 1926, in fruit, and is considered
here to be a separate specimen.

Millettia richardiana is especially variable in the
number of leaflets, the form and pubescence of the
leaflets, and the pubescence of the pods. Variants
can be constant within certain localities. However,
none of these variants seem distinct enough to be
recognized as separate taxa.

Millettia richardiana is very widely distributed
throughout western Madagascar from Antsiranana
(Diego-Suarez) to Toliara (Tulear) and southwestern
Madagascar, including the forest of Zombitsy, the
Bemaraha Massif, the Ambongo, the Boina and the
Ankarana Massif. It occurs in deciduous woodland,
mainly on calcareous or sandy soils, on limestone,
sand over limestone or on coastal dunes, at up to
ca. 300 m altitude.

Pongamiopsis viguieri Du Puy & Labat, sp. nov.
TYPE: W Madagascar. Tsingy de Bemaraha
(Reserve Naturelle no. 9), environs d’Antsa-
lova, 5 km a l’E de Bevitika, 18°38'S, 44°43'E,
150-250 m, 30 Nov. 1992 (fr), J.-N. Labat,
R. Edmond & O. Laivao 2243 (holotype, P;
isotypes, K, MO, P, TAN). Figure 6.

A P. amygdalina differt inflorescentiis brevibus con-
densatis, dentibus calycis latioribus, apicibus tenuibus des-
titutis, vexillo, alis et carina dense pubescentibus, vagina
staminea breviori, et leguminibus plerumque elongatiori-
bus seminibus numerosioribus,

A deciduous tree (2-4-)10 m tall, flowering on
new shoots as they emerge from the buds and whilst
the leaves are still very immature; young shoots
densely pubescent, glabrescent; twigs with large leaf
scars and large, spherical dormant buds. Leaves
becoming very large, with 7-11 opposite leaflets;
stipels absent. Leaflets oblong, oblong-elliptic or ob-
long-obovate, 50-200 x 20-80 mm, with a broad,
V-shaped apical notch with a mucro at the base of
the notch, silky pubescent at first but becoming
glabrous above and pubescent mainly on the veins
beneath, the veins strongly raised beneath, the
straight, parallel secondary veins extending to the

slightly thickened margin. Racemes produced in the
axils of the lowest leaves of developing shoots, short
and condensed, the axis ca. 1-3 cm long, with
numerous flowers in closely spaced clusters of 2 or
3 on short spurs; pedicels with a pair of small,
caducous bracts toward the apex. Flowers 13-15
mm long, white washed with pale lilac, the standard
with a pale green basal eye. Calyx 6-8 mm long,
densely appressed-hairy, with 3 triangular, acute
lower teeth, the upper pair partially connate but
with free, widely separated, recurved tips. Standard
limb 12-14 x 12-13 mm, densely silky pubescent
behind. Wings about as long as the keel, narrow,
both appressed-pubescent outside and upcurved api-
cally, the keel shortly beaked. Staminal sheath 12-
15 mm long. Style short-hairy, hooked apically. Pod
strongly inflated, ellipsoidal to oblong-ellipsoidal, 45-
70 x 25-35 x 23-35 mm, obliquely beaked api¬
cally, the upper margin broad and thickened, vel¬
vety-pubescent, wrinkled, thick and woody, inde-
hiscent or eventually splitting into 2 valves, usually
with 3-5 seeds (occasionally some pods with fewer
seeds). Seeds subspherical, compressed, ca. 13 mm
diam., becoming dark gray, the hilum raised and
surrounded by a narrow aril.

Pongamiopsis viguieri is closely related to P.
amygdalina, with similar leaves, pod type, and
growth habit, but differs in its short, condensed
inflorescences, its broader calyx teeth lacking the
very slender tips, its densely pubescent standard,
wings, and keel, its shorter staminal sheath, and its
usually more elongated pods with more numerous
seeds. It occurs in western rather than northern
Madagascar, which is the range of P. amygdalina.

Distribution. Western Madagascar, uncommon,
mainly from the Bemaraha Massif, but also collected
at Amboanio south of Mahajanga (Majunga) and at
Andranomavo near Namoroka.

Habitat. In deciduous woodland, always over
limestone outcrops or “tsingy,” from near sea level
to ca. 500 m altitude.

Flowering time. October.

This species is dedicated to Rene Viguier, in rec¬
ognition of his extensive research on the Legumi-
nosae of Madagascar.

Paratypes. MADAGASCAR. Vallee de la Tsiribihina
dans le Bemaraha, 20-350 m, Oct. 1933 (fl, jfr), Hum¬
bert 11480 (K, MO, P); Antsingy de Bemaraha, vers
Anbodiriana (E d’Antsalova), NE du pont, 100-150 m,
10 Dec. 1952 (fr), Leandri, Capuron & Razafindrakoto
2102 (BR, K, L, MO, NY, P, PRE, WAG); Antsingy de
Bemaraha, vers Anbodiriana (E d’Antsalova), 100-150
m, 21-27 Jan. & 8-11 F6v. 1960 (fr), Leandri &
Saboureau 3071 (K, L, MO, NY, P, PRE); Ambongo,
environs d’Andranomavo, Jan. 1905 (jfr), Perrier de la

Volume 5, Number 2
1995

Labat & Du Puy
Millettieae from Madagascar

181

Figure 6. Pongamiopsis viguieri Du Puy & Labat. A. Habit with mature leaf and pod- j j Habit with

leaf and flowers. —C. Leaflet undersurface. —D. Leaflet apex. —E. Flower. —F. Calyx. — G. Pod. — H. seed (A,
C, and D drawn from Labat, Edmond & Laivao 2243; B, E, and F drawn from Humbert 11480; G and H from
Leandri & Saboureau 3071.)

182

Novon

Bathie 1801 (P); Rochers calcaires du bord de mer a
Amboanio pres de Majunga, Fev. 1901 (fr), Perrier de
la Bathie 1828 (P); Antsingy de Bemaraha, route An-
tsalova-Tsiandro, aux environs de la clairiere d’Ambo-
diriana, 10 Dec. 1952 (fr), Service Forestier de Mad¬
agascar 6808-SF, Capuron (K, L, MO, P, TEF).

Acknowledgments. We thank Frangoise Theu-
reau and Margaret Tebbs for the illustrations and
R. Hoogland for assistance with the Latin descrip¬
tions. D. Du Puy thanks the Royal Society for the
opportunity to undertake collaborative research in
the Laboratoire de Phanerogamie, Paris, and the
Weston Foundation for the support of his continued
research in Madagascar and Kew. We thank the
Directors and staff of the Laboratoire de Phanero¬
gamic, Paris, the Herbarium, Royal Botanic Gar¬

dens, Kew, the Parc de Tzimbazaza, Antananarivo,
and the Centre National de la Recherche Ecologique,
Antananarivo. We are also grateful to the National
Geographic Society for supporting our recent field¬
work in Madagascar.

Literature Cited

Geesink, R. 1981. Tephrosieae. Pp. 245-260 in R.
M. Polhill & P. H. Raven (editors), Advances in
Legume Systematics, Part 1. Royal Botanic Gardens,
Kew.

-. 1984. Scala Millettiearum. A survey of the

genera of the tribe Millettieae (Legum.-Pap.) with
methodological considerations. E. J. Brill/Leiden Univ.
Press, Leiden Botanical Series 8.

Hutchinson, J. 1964. The Genera of Flowering Plants.
Vol. 1. Oxford Univ. Press, Oxford.

Another New, Nearly Extinct Species of Hibiscadelphus (Malvaceae)
from the Hawaiian Islands

David H. Lorence

National Tropical Botanical Garden, P.0. Box 340, Lawai, Hawaii 96765, U.S.A.
Warren L. Wagner

Department of Botany, National Museum of Natural History, Smithsonian Institution, MRC-
166, Washington, D.C. 20560, U.S.A.

ABSTRACT. Four individuals of a new species of
Hibiscadelphus (Malvaceae: Hibisceae) have been
discovered on Kaua'i, the oldest of the major Ha¬
waiian Islands. Hibiscadelphus woodii is described
and illustrated, and its status and basal relationship
within the genus are discussed. The imperiled status
of the other six species is reviewed, and a key to
the genus is given.

Hibiscadelphus Rock consists of seven species
endemic to the Hawaiian Islands (Bates, 1990), in¬
cluding the one described herein. The genus was
originally established by Joseph F. Rock (Radlkoffer
& Rock, 1911) to accommodate three species of
shrubs or small trees that he had collected from
dryland forests: two species on the island of Hawai‘i,
H- giffardianus Rock (Mauna Loa) and H. huala-
laiensis Rock (Hualalai), and H. wilderianus Rock
from Auwahi on East Maui. Four additional species
were subsequently described: H. bombycinus C.
Forbes from the Kohala Mountains on the island of
Hawai‘i (Forbes, 1920), H. distans L. Bishop &
Herbst from the Koai‘e Canyon of Kaua'i (Bishop
& Herbst, 1973), H. crucibracteatus Hobdy from
Puhielelu Ridge on Lana'i (Hobdy, 1984), and the
new species, H. woodii, described herein from Ka-
lalau Valley, Kaua'i.

Hibiscadelphus is a member of the tribe Hibis¬
ceae, represented in the Hawaiian Islands by the
indigenous genus Hibiscus L., following Fryxell’s
(1968) classification. Although closely related to Hi¬
biscus (the generic name means “brother of Hibis-
cus ), Hibiscadelphus is clearly a distinct mono-
phyletic group based on its curved and narrowly
convolute zygomorphic corollas with the lower two
lobes shorter than the other three, calyces that are
circumscissily deciduous in fruit, and relatively thick,
woody capsules (Hobdy, 1984; Bates, 1990). The
green, yellow, or magenta flowers with tubular,
curved corollas are presumably adapted to pollina¬
tion by native Meliphagid and Drepanidine birds.

known as the honeyeaters and honeycreepers (Hob¬
dy, 1984). The Hawaiian name for members of the
genus is hau kuahiwi, meaning upland or mountain
hau, because of their resemblance to the common
lowland hau, Hibiscus tiliaceus L. (Rock, 1913;
Bates, 1990).

Due to the destruction and severe alteration of
the diverse dry and mesic forest habitats and loss
of all of the Meliphagid birds in the Hawaiian Islands
(Hobdy, 1984), five of the seven Hibiscadelphus
species have become extinct in the wild, although
at least two of these are maintained through culti¬
vated plants: H. giffardianus and H. hualalaiensis.
Only H. distans, a U.S. federally fisted endangered
species, and the new species described in this paper
have naturally occurring wild populations (Table 1;
Bates, 1990). The status of the genus is extremely
precarious at present.

In March 1991 an unusual species of Hibisca¬
delphus not corresponding with any of the described
taxa was discovered on Kaua'i by Kenneth R. Wood,
staff member of the National Tropical Botanical
Garden, along with M. Query and S. Montgomery.
Four plants were found growing on a steep slope
below the rim of Kalalau Valley in northwestern
Kaua'i (K. R. Wood, 1992, unpublished). Study of
flowering specimens has revealed that they represent
a new species.

Hibiscadelphus woodii Lorence & W. L. Wag¬
ner, sp. nov. TYPE: Hawaiian Islands. Kaua'i:
Hanalei District, Kalalau Rim, N of Kahuama'a
Flat, steep diverse lowland mesic forest, 990-
1020 m, 3 Mar. 1991, K. R. Wood, M. Query
& S. Montgomery 629 (holotype, PTBG, a
flower also in spirit collection; isotypes, BISH,
K, MO, NY, US). Figure 1.

Species foliis glabratis vel venis sparsim stellato-pub-
erulis, pagina abaxiali laminae axillis venarum trichom-
atibus stellatis instruetis, calyce trichomatibus stellatis cum
radiis 5-8 ornatis, corolla cerina cupreo-suffusa atropur-
purescentia distinguibilis.

Novon 5: 183-187. 1995.

184

Novon

Table 1. Status of species of Hibiscadelphus (D. Ragone, unpublished observations, 1993). DOFAW — Division
of Forestry and Wildlife, Department of Land and Natural Resources, State of Hawaii (plantings at Pu‘uwa‘awa a,
Kona, Hawaii Island); HVNP = Hawaii Volcanoes National Park (planting at Kipuka Puaulu, Hawaii Island); NTBG

= National Tropical Botanical Garden (plantings at
(plantings at Haleiwa, 0‘ahu).

Lawai and Limahuli Gardens, Kaua‘i); WA = Waimea Arboretum

Species

Distribution

Wild

populations

Wild

Known individuals

Cultivated

H. bombycinus

Hawaii

Extinct

—

—

(Kohala Mts.)

H. crucibracteatus

Lana‘i

Extinct

—

—

(Puhielelu Ridge)

H. distans

Kaua‘i

2

137

39 NTBG

(Koai‘e Canyon)

14 WA

H. giffardianus

Hawai‘i

Extinct

—

7 HVNP

(Mauna Loa)

1 NTBG

8WA

H. hualalaiensis

Hawai‘i

Extinct

—

3 NTBG

(Hualalai)

20-30 DOFAW

H. wilderianus

East Maui

Extinct

—

—

(Auwahi)

H. woodii

Kaua‘i

i

4

—

(Kalalau Valley)

Small branched trees 2.5-5 m tall with dense
rounded crown, trunk with DBH 10-20 cm, the
twigs glabrate. Leaves with petioles 2.8-5.8 cm long
bearing scattered stellate trichomes when young,
glabrate; stipules subulate, 3-5 mm long, sparsely
stellate-pubescent; blade broadly ovate to very
broadly ovate, 7-9 cm long, 6.5-8.4 cm wide, thin-
coriaceous, drying chartaceous; adaxially glabrate
or with scattered small, translucent stellate tri¬
chomes on principal veins, abaxially with few scat¬
tered stellate trichomes on principal veins or gla¬
brate, with sparse tufts of stellate trichomes in major
vein axils, when young with scattered stellate tri¬
chomes along margins and at junction with petiole,
the base deeply cordate, occasionally shallowly so,
the apex acute to weakly acuminate or broadly ob¬
tuse, the principal veins 5 (7), the margins irregu¬
larly and coarsely crenate-dentate. Flowers solitary,
the peduncles 1.4-2.1 cm long, sparsely stellate
pubescent; involucral bracts 4-6, free essentially to
their base, 11-15 mm long, 1.8-4 mm wide, linear-
oblong; calyx tubular, 1.3-1.5 cm long (up to 1.7
cm when fresh), green, shallowly 5-lobed, the lobes
2-3 mm long, sometimes in age splitting further,
moderately densely pubescent with stellate tri¬
chomes composed of 5-9 horizontal radiating arms
free to their base; corolla 45-47 mm long, stellate
pubescent, yellow with coppery tinge when fresh,
rapidly becoming purplish maroon, the lobes 35-
40 mm long, 10-12 mm wide, with conspicuous
veins, moderately stellate pubescent with trichomes

becoming restricted to veins in inner % of lobes
abaxially; staminal column exserted ca. 7 mm be¬
yond corolla lobes, stigmas ca. 1 mm long, subglo-
bose, reddish brown, villosulous, exserted 14 mm
beyond corolla lobes; style villous; ovary 7 mm long,
4-5 mm wide, 5-angled, stellate pubescent, endo-
carp segments apparently 10 (in cross section). Fruit
unknown.

Distribution and habitat. The new species is
known only from a population of four trees at the
type locality—accessible only with climbing ropes.
It occurs on a steep slope of predominantly basalt
talus with patches of brown soil and leaf litter in
remnant diverse lowland mesic forest. Associated
native taxa include Melicope pallida (Hillebrand)
T. Hartley & B. Stone, Nototrichium sp. nov., Hed-
yotis sp. nov., Chamaesyce sp. nov., Lysimachia
glutinosa Rock, L. kalalauensis Skottsberg, Poa
mannii Munro ex Hillebrand, Stenogyne campan-
ulata Weller & Sakai, and Lobelia niihauensis St.
John. The alien daisy fleabane, Erigeron karvin-
skianus DC., is an extremely invasive perennial herb
that threatens to overrun and smother the native
cliff habitat in this region. Accessible areas above
these cliffs have been severely degraded by feral
goats and pigs. The Hibiscadelphus woodii plants
are also susceptible to damage by falling rocks dis¬
lodged by these animals, which aggravate erosion.

Reproduction and conservation. Flowering ma¬
terial has been collected in March, April, and Sep-

Volume 5, Number 2
1995

Lorence & Wagner
Hibiscadelphus

185

Figure 1. Hibiscadelphus woodii Lorence & W. L. Wagner. Habit. Scale 1 cm.

tember, but no fruit set has been observed in spite
of efforts to manually outcross and bag the flowers.
Introduced birds, i.e., the Japanese white eye (Zos-
terops japonicus), were seen piercing the corollas
above the calyx, presumably robbing nectar (note
on Wood et al. 640). A liquid-preserved flower of
Wood 726 contained three adult Nitidulidae beetles
probably representing an endemic species (D. Ja¬

mieson, pers. comm.). The flower also contained two
half-grown larvae of the endemic moth Crocidosema
marcidella (Walsingham) (Lepidoptera: Tortrici-
dae), which had completely eaten the style down to
the ovary. Larvae of Hawaiian Crocidosema species
are known to feed on indigenous Malvaceae fruits
and foliage (Zimmerman, 1978), and damage by
these larvae may be responsible for the observed

186

Novon

lack of fruit set in H. woodii. Attempts to propagate
the new species by air layering, cuttings, grafts, and
tissue culture thus far have failed.

We take pleasure in naming this new species for
Kenneth R. Wood, in recognition of his continued
efforts to understand and protect the rare and en¬
dangered plants of the Hawaiian Islands, especially
those of Kalalau Valley.

Paratypes. HAWAIIAN ISLANDS. Kaua'i: Hanalei
District, Kalalau Rim, N of Kahuama'a Flat, steep diverse
lowland mesic forest, 990-1000 m, 6 Mar. 1991, K. R.
Wood et al. 640 (BH), 15 Apr. 1991, K. R. Wood et
al. 726 (PTBG), 23 Sep. 1991, K. R. Wood & S. Perl¬
man 1600 (PTBG).

Affinities. Hibiscadelphus woodii differs from its
congeners in having leaves that are glabrate except
for rare stellate trichomes adaxially and sparse stel¬
late trichomes on the veins and in the principal vein
axils abaxially, stellate trichomes on the calyx with
rays free to the base, and yellow corollas with a
coppery tinge, rapidly becoming purplish maroon
with age. Hibiscadelphus distans differs in its in-
volucral bracts connate for about one-third of their
length, multi-rayed calyx trichomes of two sizes, a
smaller yellowish green corolla rapidly becoming dull
reddish, and densely stellate pubescence on the co¬
rolla with the trichomes becoming restricted to the
veins on the inner one-third of the lobes abaxially.
Hibiscadelphus giffardianus differs from If. woodii
in having filiform involucral bracts up to 1.1 mm
wide, whereas those of H. hualalaiensis are short,

triangular teeth only 0.5-3 mm long. Hibiscadel¬
phus crucibracteatus differs in having longer in¬
volucral bracts (20-30 mm), whereas H. wilderi-
anus has a longer calyx (2.3-2.5 cm). Hibisca¬
delphus bombycinus differs from H. woodii in hav¬
ing leaves with the lamina densely pubescent beneath
and a shorter corolla (3.3 cm). Fruiting material,
unknown for H. woodii, is required to further assess
its affinities. Examination of transverse hand-cut
sections of a flowering ovary of H. woodii suggests
the endocarp will separate into ten segments in fruit
as in H. bombycinus, H. crucibracteatus, H. gif¬
fardianus, H. hualalaiensis, and H. wilderianus,
rather than into five boat-shaped segments as in H.
distans.

The phylogenetic relationships of this unique Ha¬
waiian endemic genus were recently analyzed (Funk
& Wagner, 1995). In the single cladogram resulting
from their morphological study the position occupied
by Hibiscadelphus woodii is basal. Because fruit
characters are among the most important ones for
determining the tree topology, the placement of H.
woodii is in doubt. Their analysis estimated that the
fruit character of H. woodii is plesiomorphic (i.e.,
chartaceous, not woody) as in H. distans. In order
to investigate the stability of the basal position of
H. woodii in the phylogeny they recoded the missing
character as apomorphic. Even with this change,
however, H. woodii remained in the basal position.
Species of Hibiscadelphus may be separated by
characters given in the following key (adapted from
Bates, 1990).

Key to the Species of Hibiscadelphus

la. Involucral bracts connate ca. V, of their length; mesocarp weakly developed and usually adnate to exocarp;

endocarp segments 5, boatlike; Koai‘e Canyon, Kaua'i. II. distans

lb. Involucral bracts free or connate slightly at base; reticulate mesocarp strongly developed; endocarp segments

2a. Involucral bracts filiform or obsolete, up to 1.1 mm wide.

3a. Involucral bracts 0.5-2(-3) mm long; corolla yellowish green, fading to purplish within, 2-5(-5.5)

cm long; Hualalai, Hawai'i. H. hualalaiensis

3b. Involucral bracts 18-35 mm long (hybrids between H. giffardianus and other species tend to
have bracts 5-15 mm long); corolla grayish green externally, dark magenta within, (5-)6-7 cm

long; Mauna Loa, Hawai'i . H. giffardianus

2b. Involucral bracts linear to spatulate, 1-7 mm wide.

4a. Involucral bracts 20-30 mm long, 3-7 mm wide; calyx 4.2-4.9 cm long; corolla 5-6.5 cm long;

Puhielelu Ridge, Lana'i. H. crucibracteatus

4b. Involucral bracts 9-18 mm long, 1.5-4 mm wide; calyx 1.3-2.5 cm long; corolla 3.3-4.7 cm
long.

5a. Leaf lamina adaxially glabrate or with minute, scattered, stellate trichomes only on principal
veins, abaxially glabrate or with scattered stellate trichomes only on principal veins, the

trichomes sparsely tufted in principal vein axils; Kalalau Valley, Kaua'i . H. woodii

5b. Leaf lamina adaxially sparsely stellate pubescent, abaxially sparsely to densely stellate pu¬
bescent, the trichomes densely tufted in principal vein axils; Mauai, Hawai'i.

6a. Involucral bracts 9-10 mm long; calyx ca. 1.2 cm long; Kohala Mts., Hawai'i .

. H. bombycinus

6b. Involucral bracts 11-18 mm long; calyx 2.3-2.5 cm long; Auwahi, East Maui ......

. H. wilderianus

Volume 5, Number 2
1995

Lorence & Wagner
Hibiscadelphus

187

Acknowledgments. We thank Anna Asquith for
preparing the skillfully rendered illustration. Dean
Jamieson and Laura Ishi kindly identified the insect
specimens. We are also grateful to Kenneth R.
Wood and Diane Ragone for providing data and
field observations, Robynn Shannon for a careful
review of a draft of the manuscript, and to Loyal
Mehrhoff and an anonymous reviewer for valuable
comments on the manuscript.

Literature Cited

Bates, D. M. 1990. Malvaceae. Pp. 868-903 in W.
L. Wagner, D. R. Herbst & S. H. Sohmer, Manual
of the Flowering Plants of Hawaii. Univ. Hawaii
Press & Bishop Museum Press, Honolulu.

Bishop, L. E. & D. R. Herbst. 1973. A new Hibis¬
cadelphus (Malvaceae) from Kauai. Brittonia 25:
290-293.

Forbes, C. N. 1920. New Hawaiian plants. Occas. Pap.
Bernice Pauahi Bishop Mus. 7(3): 33-39.

Fryxell, P. A. 1968. A redefinition of the tribe Gos-
sypieae. Bot. Gaz. 129: 296-308.

Funk, V. A. & W. L. Wagner. 1995. Biogeography
of seven ancient Hawaiian plant lineages. In: W. L.
Wagner & V. A. Funk (editors), Hawaiian Bioge¬
ography: Evolution on a Hot Spot Archipelago.
Smithsonian Institution Press, Washington, D.C.

Hobdy, R. W. 1984. A re-evaluation of the genus
Hibiscadelphus (Malvaceae) and the description of
a new species. Occas. Pap. Bernice P. Bishop Mus.
25(11): 1-7.

Radlkoffer, L. & J. F. Rock. 1911. New and noteworthy
Hawaiian plants. Hawaiian Board of Agriculture and
Forestry Botanical Bull. 1: 1-15.

Rock, J. F. 1913. The Indigenous Trees of the Hawaiian
Islands. Charles E. Tuttle, Rutland, Vermont.

Wood, K. R. 1992. New Hibiscadelphus found on
Kauai. Hawaii’s Forests and Wildlife 7(1): 15-117.

Zimmerman, E. C. 1978. Insects of Hawaii. Vol. 9,
Microlepidoptera, Part 1. The Univ. Press of Hawaii,
Honolulu, Hawaii.

A New Species of Tournefortia (Boraginaceae) from
La Planada, Colombia

James S. Miller

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT. Tournefortia restrepoae J. S. Miller,
sp. nov., is described from the Reserva Natural La
Planada in Narino, Colombia. The species is distinct
in its evidently and evenly pustulate leaf surface
with depressed tertiary venation and elevated are-
oles, and in its widely separated petiolate flowers. It
is currently known only from the type locality.

The genus Tournefortia comprises about 150
species distributed throughout tropical and subtrop¬
ical parts of the world (Miller, 1988). The vast
majority of the species occur in the Neotropics, and
the genus is only sparsely represented in the Old
World. South America, particularly the northern
Andes, is the region with the greatest number of
species and morphological diversity (Killip, “The
Andean species of Tournefortia ,” Smithsonian In¬
stitution Archives, unpublished; Miller, 1988). The
species described below is one of many narrowly
distributed endemics in mid-elevation Andean for¬
ests.

Tournefortia restrepoae J. S. Miller, sp. nov.
TYPE: Colombia. Narino: Mpio. de Ricaurte,
Reserva Natural La Planada, elev. 1800 m,
18 Nov. 1993 (young fruits), Carla Restrepo
778 (holotype, MO 4619857; isotype, PSO not
seen). Figure 1.

Frutex sciophilus. Folia alterna; lamina elliptica ad el-
liptico-oblonga vel leviter obovata, 12-24 cm longa, 6-
11 cm lata, apice acuminata, base acuta, supra hirsuta,
infra strigosa, adaxialiter manifeste aequaliter pustulata,
nervis tertiariis depressis, areolis elevatis; petiolo 1.7-5
cm longo, supra canaliculate, dense brunneo-pubescenti.
Inflorescentia intemodalis, cymosa; pedicellis 1.5-2(-4)
mm longis inter se 1-7 mm distantibus. Flos sepalis lan-
ceolatis, 4-6 mm longis, pubescentibus; corolla tubiformi,
ca. 6 mm longa, lobulis anguste ovatis, ca. 2 mm longis;
staminibus ignotis; stylo 4 mm longo. Fructus late ovoi-
deus, albus, glaber.

Understory shrub. Leaves alternate, blades ellip¬
tic to elliptic-oblong or slightly obovate, 12-24 cm
long, 6-11 cm wide, the apex acuminate, the base
acute, sometimes asymmetrically so, the margin en¬
tire, somewhat undulate, the adaxial surface prom¬

inently and evenly pustulate with the tertiary ve¬
nation depressed and the areoles elevated, hirsute
with several erect, siliceous hairs arising from each
areole, the abaxial surface strigose with the slightly
appressed hairs mostly restricted to the veins, each
areole with one to several sunken pits, the venation
brochidodromous with 9-16 secondary veins; peti¬
oles 1.7-5 cm long, canaliculate on the adaxial
surface, densely brown pubescent. Inflorescence in-
temodal, a dichotomously branched cyme, the
branches 6-10 cm long, the peduncle 7.5 cm long,
both densely brown pubescent. Flowers on 1.5-
2(-4)-mm-long pedicels, 1-7 mm apart, bisexual;
sepals green, lanceolate, ± equal, 4-6 mm long,
0.4-1.3 mm wide, sparsely but obviously pubescent;
corolla tubular, ca. 6 mm long, the lobes narrowly
ovate, ca. 2 mm long; stamens unknown; ovary
ovoid, ca. 1 mm long, ca. 0.5 mm broad, borne on
an annular nectariferous disk, the style ca. 4 mm
long, the stigma conical. Fruits white, drupaceous,
the exocarp glabrous, the mesocarp thin, the en-
docarp distinctly 4-parted, broadly ovoid, ca. 5 mm
long, ca. 4 mm wide.

Distribution. Tournefortia restrepoae is known
only from the type collection from cloud forest at
1800 m at La Planada in Narino, Colombia.

Tournefortia restrepoae is a distinctive member
of Tournefortia sect. Tournefortia, which is by far
the largest section of the genus, containing the ma¬
jority of the neotropical species, and is characterized
by having free anthers and fruits that are unlobed
and lack a corky mesocarp. E. P. Killip’s unpub¬
lished treatment of the Andean species of the genus
recognized 38 species in the section, none of which
appear very close to T. restrepoae. The species is
distinct in its alternate, distinctly pustulate leaves
and widely spaced, pedicellate flowers. The leaf sur¬
face is unique in the genus. The type collection is
mostly in young fruit with only the old remains of
a few floral parts, thus better knowledge of rela¬
tionships will require the collection of better flow¬
ering material. The species is named in honor of
Carla Restrepo, who has done much to improve our
knowledge of the plants of the Andean Cordillera in
Colombia.

Novon 5: 188-189. 1995.

Volume 5, Number 2
1995

Miller

Tournefortia restrepoae

189

Figure 1. Tournefortia restrepoae J. S. Miller (Restrepo 778). —A. Fruiting branch. B. Section of infructescence
showing fruiting calyx. —C. Lateral and apical view of fruit. —D. Upper leaf surface. E. Lower leaf surface.

Paratypes. COLOMBIA. IMarino: Mpio. de Ricaurte,
Reserva Natural La Planada, elev. 1800 m, Restrepo
730 (PSO not seen).

Acknowledgments. I thank John Myers for the
illustration, Joan W. Nowicke and the Smithsonian
Institution for making E. P. Killip's unpublished

manuscript available, Roy E. Gereau for help with
the Latin description, and Gordon McPherson for
review comments.

Literature Cited

Miller, J. S. 1988. A revised treatment of Boraginaceae
for Panama. Ann. Misssouri Bot. Gard. 75: 456-
521.

Aristida tuitensis (Poaceae: Aristideae), a New Species from
El Tuito, Jalisco, Mexico

Jorge G. Sanchez-Ken and Patricia Davila A.

Instituto de Biologla, U.N.A.M., Depto. de Botanica, Herbario Nacional, Apartado Postal 70-
233, 04510 Mexico, D.F., Mexico

Abstract. A new species, Aristida tuitensis from
El Tuito, Jalisco, Mexico, is described and illustrat¬
ed, and its distribution and ecology are discussed.
It belongs to Aristida sect. Streptachne and is dis¬
tinguished from A. schiedeana by its spreading in¬
florescence, shorter spikelets, and the lack of dentate
glumes, and from A. hintonii by its inflorescence
shape, the internode, and sheath length. A table
summarizing the differences and similarities among
the three species is presented.

As part of research to revise the genus Aristida
L. section Streptachne (R. Brown) Domin, some
unusual specimens of Aristida were collected in the
area of El Tuito, Jalisco, Mexico, in early 1993.
The material was initially identified as A. schie¬
deana, a common grass in the area. However, with
additional studies of these specimens, as well as other
collections from IBUG and COCA, it was realized
that these specimens represent a new species.

Aristida tuitensis Sanchez-Ken & P. Davila, sp.
nov. TYPE: Mexico. Jalisco: Municipio El Tui¬
to, El Tuito, 9 a 10 km sobre la brecha a las
Minas del Cuale (E de El Tuito), elev. 1900
m, 13 ene. 1991, J. Sanchez-Ken, M. May-
field & B. Westlund 500 (holotype, MEXU;
isotypes, ENCB, IBUG, MICH, MO). Figure 1.

Gramen perenne caespitosum; internodia ramosa, 9-
15 infra inflorescentiam; vaginae coriaceae internodiis
longiores; ligulae externae presentes; spiculae effusae bin-
atae; glumae infernae glumis supernis longiores; glumae
infernae no dentatae, 3-nervatae, nervis centrales con-
spicui, nervi laterales inconspicui; calli 0.7-1.0 mm longi;
lemmata glumis breviores vel eas aequantes raro; col-
umnae lemmatum 6-10 mm longae; aristae laterales ab-
sentes; antherae 3-4 mm longae.

Perennial tufted grass, erect; pseudorhizome
present, 80-120 cm long. Internodes 2.0-
9.0(-12.0) cm long, 1-2 mm diam., glabrous,
straight, terete, bamboo-like, branched, with 9-15
intemodes below the inflorescence; nodes glabrous,
inconspicuous. Leaf sheaths 3-9 cm long, 4-6 mm
wide, longer than the internodes, strongly overlap¬
ping, glabrous, not striate, strongly coriaceous, in-

Novon 5: 190-192. 1995.

eluding the margins; inner ligule ciliate, 0.2-0.3
mm long; auricles lacking, sometimes a tuft of lateral
hairs 2 mm long instead of the auricles in juvenile
stages; outer ligule or contra ligule present; blade
10-50 cm long, 2-4 mm wide, linear, flat, straight
to slightly flexuous when dry, adaxially scaberulous
with scattered hairs toward the base, abaxially smooth
or rarely scaberulous, with 7 main conspicuous
nerves, but with 21, 23, or 25 nerves in total.
Inflorescence an open panicle, 14-35 cm long, 12-
20 cm wide; basal internode 18-70 cm long; pe-
duncule 11-30 cm long, glabrous or scaberulous;
branches 9— 13(— 15) cm long, branching until 5th
order, ascendant, strongly spreading, with the spike-
lets located at the apex; pulvinus present in the axils
of all branches and pedicels, glabrous; pedicels 5-
18 mm long. Spikelets usually paired, 20-28 mm
long from the base of glumes to the tip of the awn,
spreading; glumes varying in size, usually the first
larger than the second one, sometimes the second
glume larger, persistent; first glume 4-8 mm long,
to 1 mm wide, 3-nerved, the two lateral nerves
inconspicuous, glabrous, keeled and scabrous in the
keel, apically acute and awned, not dentate, awn
0.5-2.0 mm long; second glume 6.0-7.5 mm long,
to 1.0 mm wide, 1-nerved, keeled and glabrous in
the keel, apically obtuse and awned, not dentate,
the awn 0.5-1.0 mm long; calluses 0.7-1.0 mm
long; lemma shorter than or sometimes as long as
the glumes, 4.5-6.0 mm long without the column,
1.0-1.3 mm wide, convolute, 3-nerved, terete, sca¬
berulous in the upper third, not articulate with the
column, not articulated; column 6-10 mm long,
twisted, abaxially scaberulous, adaxially muricate,
basally geniculate; awn 1, 8-10 mm long, straight,
basally geniculate, thin; lateral awns lacking; palea
0.5-0.7 mm long, not keeled, smooth; lodicules 2,
1.0-1.2 mm long, hyaline; stamens 3; anthers 3-
4 mm long; stigma 1-2 mm long. Caryopsis not
seen.

Distribution. Aristida tuitensis is endemic to the
area of El Tuito in the state of Jalisco, northeast of
Chamela. It occurs in a pine and oak forest that
also contains some elements that are characteristic

Volume 5, Number 2
1995

Sanchez-Ken & Davila
Aristida tuitensis

191

Figure 1. Aristida tuitensis Sanchez-Ken & P. Davila. -A. Habit. -B. Stem. -C. Adaxial and abaxial view of
the leaf sheath. —D. Inflorescence. —E. Spikelet (all from Sanchez-Ken et al. 500).

of a tropical rainforest. Altitudinally, it ranges from
1300 to 1900 m. The soils are reddish and usually
sandy to clayish. Flowering occurs between Septem¬
ber and January.

According to Henrard’s (1929) criteria, this spe¬
cies belongs to Aristida sect. Streptachne. As shown
by the herbarium annotations, this species was con¬
fused with A. schiedeana in Flora Novo-Galiciana

192

Novon

Table 1. Morphologic

comparison among Aristida tuitensis, A. hintonii, and A. schiedeana.

Character

A. tuitensis

A. hintonii

A. schiedeana

Branching

basal, middle

basal

basal

Number of internodes

9-15

2-10

2-7

Stem

shorter

longer

longer

Sheath length compared
with internode length

longer

shorter

shorter

Leaf shape

straight to flexuous

straight to flexuous

flexuous to coiled

Inflorescence

few-flowered

flowered

few-to many-flowered

Inflorescence branches

strongly spreading

spreading

appressed

Pulvinus

all branches

usually all branches

usually only first-order
branches

Spikelet

usually paired

paired and triads

paired and triads

Larger glume

usually the first

usually the second

second or first

Glumes

not dentate

dentate

dentate

Lateral awns

lacking

0.1-1.0 mm

0.1-2.0 mm

Anthers

3-4 mm

1.5-2.0 mm

1.5-2.0 mm

(McVaugh, 1983). However, A. tuitensis has reli¬
able diagnostic characters by which it can be dis¬
tinguished from A. schiedeana and A. hintonii,
probably the two most closely related species (Table
1). A notable character of A. tuitensis is the rela¬
tively long anthers compared to the lemma body.
To our knowledge, such long anthers have not been
reported in any other species of the genus.

Artificial Key to Three Related Species of Aristida
Sect. Streptachne

la. Anthers 3-4 mm long; sheaths not striate; in¬
florescence branches strongly spreading.

. A. tuitensis

lb. Anthers 1.5-2 mm long; sheaths striate; inflo¬
rescence branches slightly spreading or ap-
pressed.

2a. Spikelets usually spreading; pulvinus usu¬
ally in ail branches; leaf blades straight to
flexuous; lateral awns up to 1.0 mm long

. A. hintonii

2b. Spikelet appressed to branches; pulvinus
in the branches of 1st order; leaf blades
flexuous to coiled; lateral awns up to 2.0
111111 l° n 8 .. A. schiedeana

The specific epithet refers to the area in which
the species was discovered, El Tuito. Many other
taxa have recently been discovered in this region.

Paratypes. MEXICO. Jalisco: Mpio. Cuautitlan, Cer-
ca de 10 km al sur-suroeste de las Joyas, Sierra de
Manantlan, R. Guzman 6120 (COCA); Mpio. El Cuale,

Lado este de las Minas de Zimapan, brecha a Cuale, R.
Guzman M. 6101a (IBUG); Mpio. El Tuito, 30 mi. S of
Puerto Vallarta, A. A. Beetle et al. M-3610 (COCA); Al
sur-sureste de Puerto Vallarta, 14 km por la brecha a
Minas de Zimapan, 1 km al oeste de Providencia por la
brecha El Tuito-El Cuale, R. Guzman M. 6094 (COCA);
Km 2 por la brecha a Minas de Zimapan, F. J. Santana
M. 1196 (IBUG); Km 14 de la brecha El Tuito-Minas
Zimapan, 1 km al oeste del Aserradero Provincia, F. J.
Santana M. 1204 (IBUG); Mpio Talpa, Entre Cuale y
Talpa, J. de la Torre V. s.n. (IBUG).

Acknowledgments. We thank Jose Luis Villa¬
senor and Oswaldo Tellez for reviewing the manu¬
script. We thank the McArthur Foundation, the
Consejo Nacional de Ciencia y Tecnologia (CONACyT
CPECBN-021759), and the Direccion General de
Asuntos del Personal Academico (D.G.A.P.A.-
U.N.A.M.) for financial support for fieldwork and
botanical research in different parts of Mexico. A
special thanks to Eloisa Duarte for the elaboration
of the drawing. Finally, we thank the Instituto de
Biologia, U.N.A.M., where this research was carried
out.

Literature Cited

Henrard, T. 1929. A monograph of the genus Aristida.

Meded. Rijks-Herb. Leiden no. 58: 1-156.
McVaugh, R. 1983. Gramineae. In: Flora Novo-Gali-
ciana: A Descriptive Account of the Vascular Plants
of Western Mexico. Vol. 14. The Univ. Michigan
Press, Ann Arbor.

Novelties in the Neotropical Genus Ouratea Aublet (Ochnaceae)

Claude Sastre

Laboratoire de Phanerogamic, Museum National d’Histoire Naturelle, 16 rue Buffon, F-75005,

Paris, France

ABSTRACT. A new section of Ouratea, section Cad-
vcae Sastre, and eight new species of Ouratea from
Venezuela and adjacent countries are described: O.
pendulosepala Sastre (sect. Cardiocarpae), O. oli-
gantha Steyermark ex Sastre (sect. Cardiocarpae),
0. apurensis Sastre (sect. Ouratea), 0. paratatei
Sastre (sect. Ouratea), O. cidiana Sastre (sect. Our-
atella), O. guriensis Sastre (sect. Ouratella), O.
pseudoguildingii Sastre (sect. Ouratella), and O.
squamata Sastre (sect. Ouratella).

During preparation of the Ochnaceae treatment
for the Flora of the Venezuelan Guayana, approx¬
imately 20 undescribed species of Ouratea Aublet
from southern Venezuela and adjacent areas have
been found. In order to properly place these new
species within the subgeneric classification, a brief
review and update of the previous system (Sastre,
1988) is needed. In the previous classification, I
recognized six sections: Kaieteurea (Dwyer) Sastre,
Cardiocarpae (Engler) Sastre, Polyouratea (Tiegh-
em) Sastre, Persistens Sastre, Ouratella (Tieghem)
Sastre, and Ouratea, which is the type section of
the genus (with O. guianensis Aublet as the type).
The new system also has six sections, but section
Persistens is now placed in synonymy of section
Ouratea, and section Caducae is newly described
below.

Aublet’s (1775) description of Ouratea guianen¬
sis was based on a flowering sheet (in BM), illustrated
by plate no. 152, and all additional material of this
species studied for my 1988 classification was either
in flower or old fruit, making it impossible to assess
the structure of the carpids (fruiting carpels) with
precision. Recent study of herbarium specimens of
O. guianensis from CAY, MO, NY, and VEN clearly
shows, however, that its carpids are similar to those
of the species previously included in section Per¬
sistens. This necessitates the transfer of section
Persistens into the type section, and the recognition
of a new section to include species previously (and
erroneously) thought to be closely related to 0.
guianensis, but whose fruit structure is now seen
to be different.

Ouratea sect. Caducae Sastre, sect. nov. TYPE:
Ouratea longifolia (Lamarck) Engler in Mart.,
FI. Bras. 12(2): 316. 1876. Ochna longifolia
Lamarck, Encycl. Meth. 4: 511. 1798.

Inflorescentia terminalis, flores 5 sepalis, 5 carpellis,
fructus 5 sepalis caducis, 1-3 carpellis verticalibus.

Inflorescence terminal, flowers with 5 sepals and
5 carpels, sepals caducous in fruit, fruiting carpels
1-3, vertical.

Key to the Sections of Ouratea

1. Flowers with 2-4 sepals united in 2-3 parts ..

.1. sect. Kaieteurea

1. Flowers with 5 distinct sepals . 2

2(1). Fruits with the carpels borne horizontally ..

.2. sect. Cardiocarpae

2. Fruits with the carpels borne vertically . 3

3(2). Flowers with (5)6-10 carpels.

.3. sect Polyouratea

3. Flowers always with 5 carpels . 4

4(3). Fruits with persistent sepals ... 4. sect. Ouratea

4. Fruits without sepals . 5

5(4). Inflorescences axillary, rarely terminal ....

.5. sect. Ouratella

5. Inflorescences always terminal 6. sect. Caducae

Description and Nomenclature of the Sections

1. Ouratea Aublet sect. Kaieteurea (Dwyer) Sastre,

Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B
Adansonia 1: 50. 1988. Kaieteurea Dwyer,
Bull. Torrey Bot. Club 70: 50. 1943. TYPE:
Ouratea gillyana (Dwyer) Sandwith & Ma¬
guire.

Inflorescence terminal or axillary; flowers with
2-4 sepals united in 2-3 parts, and 5 (-7 in O.
articulata Sastre) carpels; fruits with sepals cadu¬
cous or persistent and the carpels vertical.

2. Ouratea Aublet sect. Cardiocarpae (Engler)

Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10,
sect. B Adansonia 1: 55. 1988. Ouratea sect.
Eugomphia (Planchon) Engler, ser. Cardi¬
ocarpae Engler in Mart., FI. Bras. 12(2): 307.
1876. TYPE: Ouratea cardiosperma (DC.)
Engler.

Novon 5; 193-200. 1995.

194

Novon

Diouratea Tieghem, Ann. Sci. Nat. Bot. ser. 8, 16: 227.
1902.

Inflorescence terminal; flowers with 5 sepals and
5 carpels; fruit with the carpels horizontal.

3. Ouratea Aublet sect. Polyouratea (Tieghem)

Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10,
sect. B Adansonia 1: 55. 1988. Polyouratea
Tieghem, Ann. Sci. Nat. Bot. ser. 8, 16: 268.
1902. TYPE: Ouratea hexasperma (A. St.
Hilaire) Baillon.

Inflorescence terminal; flowers with 5 sepals and
6-10 carpels; fruit with the carpels vertical.

4. Ouratea Aublet sect. Ouratea. TYPE: Ouratea

guianensis Aublet.

Ouratea sect. Persistens Sastre, Bull. Mus. Natl. Hist.
Nat. 4°. ser. 10, sect. B Adansonia 1: 59. 1988.

Inflorescence terminal; flowers with 5 sepals and
5 carpels; fruits with sepals persistent and the carpels
vertical.

5. Ouratea Aublet sect. Ouratella (Tieghem) Sastre,

Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B
Adansonia 1: 59. 1988. Ouratella Tieghem,
Ann. Sci. Nat. Bot. ser. 8, 16: 289. 1902.
TYPE: Ouratea mexicana (Humboldt & Bonp-
land) Engler.

Inflorescence axillary, rarely terminal, spicate or
paniculate; flowers with 5 sepals and 5 carpels, fruit
with carpels vertical.

6. Ouratea Aublet sect. Caducae Sastre. TYPE:

Ouratea longifolia (Lamarck) Engler.

Ouratea Aublet sect. Ouratea, pro parte fide Sastre (1988)
excluding O. guianensis.

See description above.

Inflorescence terminal; flowers with 5 sepals and
5 carpels; fruit with carpels vertical.

Following are descriptions of eight new species in
sections Cardiocarpae (two species), Ouratea (two
species), and Ouratella (four species). The descrip¬
tion of new species in section Caducae will be pre¬
sented in a future paper.

Ouratea Section Cardiocarpae

Ouratea pendulosepala Sastre, sp. nov. TYPE:
Brazil. Amazonas: Mun. Pres. Figueredo, 8 km
da Vila Residencial Atrari, Canteiro de obras
de Balbina. 1°30'-2°S, 59°30'-60°W, mata de
terra firme, solo argiloso, 19 sep. 1986 (fr), C.
A. Cid Ferreira et al. 8219 (holotype, P; is¬
otypes, INPA not seen, NY not seen). Figure
1 .

Ab Ouratea cerebroidea Sastre, foliis nervis non im-
pressis, paucis visibilibus, margine sub-integra, sepalis 12
x 2 mm, persistentibus pendusque in fructu, staminibus
papillatis, stylo 10 mm longo, differt.

Tree 12 m tall, branches glabrous. Leaves cori¬
aceous, 12-16 cm long, 5-6 cm wide, obovate,
apex acuminate, base attenuate, margin undulate,
subentire, principal vein salient on lower surface,
lateral veins faintly evident. Inflorescence panicu¬
late, terminal, 14 cm long. Peduncle 13-15 mm
long, weakly papillate; sepals 5, 10-11 x 2 mm,
ovate; petals 5, obovate, 15-16 x 6-7 mm; sta¬
mens 10, papillate, 10 mm long. Young fruits with
peduncle 15 mm long, sepals 5, pendent and per¬
sistent in older fruits, 12 x 2 mm, ovate. Fruits
with 1-4 horizontal carpels, the carpophore sub-
cylindric.

Until now, section Cardiocarpae was restricted
to the Guianas and had only two species: Ouratea
cardiosperma (DC.) Engler and O. cerebroidea
Sastre. The pendent, persistent sepals are charac¬
teristic of the new species and are unknown in the
other species of Ouratea.

Paratypes. BRAZIL. Amapa: Camipi, EMBRAPA
reserve and vicinity, 00°10'N, 51°37'W, 6 Sep. 1983,
S. Mori et al. 15864, 15865, 15868, 15869, 15874
(MG not seen, NY not seen, P). Amazonas: Mun. Bar-
celos, 00°30'N, 63°30'W, rio Araca, 3 jul. 1985, E.
Sette Silva et al. 214 (INPA not seen, NY not seen, P).
Para: Mun. de Almeirin, Mte. Dourado, estrada MTD-
oeste em dire$ao a serra da mina de bauxita, 17 sep.
1985 (fl), M. J. P. Pires et al. 627 (K not seen, P).

Ouratea oligantha Steyermark ex Sastre, sp. nov.
TYPE: Venezuela. Bolivar: Alto Rio Cuyuni,
SE escarpment of Cerro Pitin, Cordillera Epi-
cara, 400 m, B. Maguire, J. A. Steyermark
& C. Maguire 53644 (holotype, NY). Figure
2 .

Ab Ouratea pendulosepala Sastre, inflorescentiis par¬
vis (4-10 cm longis), laminis parvis (7-9 cm longis) et
sepalis caducis in fructus, differt.

Shrub 1.5—4 m tall, branches glabrous. Petiole
1 cm long, leaves coriaceous, 7-9 x 3-3.5 cm.

TYPlj

inntnit mil rwsimn

CU-tiM-h^- f+-d~Ln[A. £»£>«.

Figures 14. — 1 . Ouratea pendulosepala Sastre; Cid Ferreira et al. 821 9 (holotype, P). —2. Ouratea oligantha

'^mark ex Sastre; Maguire et al. 53644 (holotype, NY). —3. Ouratea apurensis Sastre; Steyermark et al.
Oh 62 (holotype, P). —4. Ouratea paratatei Sastre; Blanco 525 (holotype, P).

196

Novon

elliptic, apex acuminate, base obtuse, margins
toothed, undulate, principal vein salient on lower
surface, with 8-12 unpaired lateral veins on each
side of the principal vein, ascending, salient on lower
surface between the small, subparallel veins, a few
of which are also salient. Inflorescences usually ax¬
illary, sometimes terminal, spicate or sometimes pa¬
niculate, 4-10 cm long. Pedicel 12 mm long, sepals
5, 7 x 2 mm, oblong; petals 5, 15 x 6 mm, ovate;
stamens 10, 1 cm long, carpels 5, horizontal; style
single, 5.5 mm long. Young fruit with peduncle 15
mm long, sepals late-caducous, carpels 1-3, hemi¬
spheric, carpophore conical.

Distribution. Known only from the type locality.

The type and paratypes cited here were annotated
by Steyermark at different times as “O. oligantha
sp. nov.” and “O. soderstromii Sastre.” I have
chosen to use Steyermark’s name for the new spe¬
cies. The material cited clearly does not belong to
Ouratea soderstromii; the new species belongs to
section Cardiocarpae, while O. soderstromii is a
member of section Ouratella. The name used by
Steyermark is validated here. Ouratea oligantha is
most closely related to O. pendulosepala, described
above, but differs by the short inflorescences (4-10
cm), that are usually spicate (vs. largely paniculate),
short leaves (7—9 cm long), and the sepals caducous
in fruit. Ouratea pendulosepala has longer inflo¬
rescences (14 cm long), longer leaves (12-16 cm
long), and persistent sepals. Ouratea oligantha is
the first species of section Cardiocarpae known in
Venezuela.

Paratypes. VENEZUELA. Bolivar: Alto Rio Cuyuni,
Rio Chicanan, Cerro Pitin, Cordillera Epicara, 3 sep.
1962, Maguire et al. 53568 (NY), 9-11 sep. 1962,
Maguire et al. 53679 (NY).

Ouratea Section Ouratea

Ouratea apurensis Sastre, sp. nov. TYPE: Ven¬
ezuela. Apure: Reserva Forestal San Camilo,
selva siempreverde a lo largo de la Quebrada
de la Cristalina hasta la vega del Cerro la Nulita,
250-280 m, 2 apr. 1968, Steyermark et al.
101762 (holotype, P; isotypes, NY, VEN). Fig¬
ure 3.

Ab Ouratea marahuacensi Maguire et Steyermark,
folia petiolo 2-3 mm longo (non 5-9 mm) nervis later-
alibus prominentibus, sepalis 8 x 2-3 mm, petalis 10 x
5 mm, staminibus 7.5 mm longis, stylo 7.5 mm longo,
differt.

Tree 7 m tall, branches glabrous. Petioles 2-3
mm long, leaf blades elliptic, undulate, 6.5-9 x 2-
2.5 cm, apex acuminate, base attenuate, margin

crenulate, principal vein salient on the lower surface,
lateral vein prominent, transversal tertiary veins
parallel. Inflorescence paniculate, terminal, 8-11
cm long. Peduncle 5-6 mm long, with short hairs
0.2-0.3 mm long; sepals 5, oblong-lanceolate, 8 x
2-3 mm; petals 5, obovate, 10 x 5 mm; stamens
10, 7.5 mm long; style 7.5 mm long. Young fruits
with sepals persistent.

Both Ouratea marahuacensis Maguire & Stey¬
ermark and O. apurensis occur in Venezuela, the
former on Cerro Marahuaca, Estado Amazonas, at
1150 m elevation, the latter in the Reserva Forestal
San Camilo, Estado Apure. Ouratea apurensis dif¬
fers from 0 . marahuacensis by the shorter petiole
(2-3 mm long vs. 5-9 mm long), salient lateral
veins, longer sepals (8 mm long vs. 6-7.5 mm long),
longer petals (10 mm long vs. 6-7.5 mm long), and
longer stamens (7.5 mm long vs. 5-6 mm long).

Ouratea apurensis is known only from the type
collection.

Ouratea paratatei Sastre, sp. nov. TYPE: Ven¬
ezuela. Bolivar: Campamentos “El Paraiso” y
“La Yagua” a 48 km NE del caserio Las Rosas,
este ultimo a 17 km de Upata (carretera nueva
Upata-San Felix), jun. 1966, Blanco 525 (ho¬
lotype, P; isotypes, NY, VEN). Figure 4.

Ab Ouratea tatei Gleason, lamina elliptica viridi, nervo
principali prominenti pagina superiore, nervis lateralibus
non impressis, differt. Ab O. pseudotatei Maguire et Stey¬
ermark, lamina nervo principali prominenti pagina su¬
periore, nervis tertiaris paucis visibilibus (non impressis),
fructus sepalis persistentibus, differt.

Tree 7 m tall, branches glabrous. Petiole 1 cm
long, leaf blade coriaceous, elliptic-ovate, 14-19 x
4.5-7 cm, apex acuminate, base obtuse to round,
margin undulate-denticulate, principal vein raised
on upper surface, lateral veins subequal, tertiary
veins anastomosing, faintly evident. Inflorescence
paniculate, terminal, 12 cm long. Flowers not known.
Fruits with 5 persistent sepals, these 7 x 2.5 mm,
lanceolate, acuminate, the carpels 1-3, obovate, 8-
9 x 5.6-6 mm, carpophore conical, 5-6 mm long,
peduncle 1 cm long.

Ouratea tatei Gleason, O. pseudotatei Maguire
& Steyermark, and O. paratatei are similar, but
the fruits of O. pseudotatei do not have persistent
sepals and so belong to section Caducae; O. pseu¬
dotatei is endemic to Guyana. Ouratea tatei and
O. paratatei are Venezuelan species, the former
occurring in the highlands of Estado Bolivar (Ro-
raima-tepui, Chimanta-tepui), the latter from the

Volume 5, Number 2
1995

Sastre

Ouratea

197

lowlands of Estado Bolivar in the Sierra de Imataca.
The differences of the Venezuelan species in section
Ouratea are outlined in the key below.

Key to the Venezuelan Species of Ouratea Sect.
Ouratea

1. Leaf blades with 2 types of secondary veins:
large ascending ones terminating along the blade
margin, and between these and the midvein,
some smaller, horizontal, ± parallel veins . 2

1. Leaf blades with subequal secondary veins .... 4
2(1). Leaves 18-22 cm long, sepals 9-10 mm long

. 0. guianensis Aublet

2. Leaves 5-9 cm long, sepals 5-8 mm long ... 3
3(2). Petiole 2-3 mm long; sepals 8 mm long;

petals 10 mm long. O. apurensis Sastre

3. Petiole 5-9 mm long; sepals 6-7.5 mm long;

petals 5-6 mm long .

.... 0. marahuacensis Maguire & Steyermark

4(1). Leaves 12-19 cm long . 5

4. Leaves 3-12 cm long . 6

5(4). Leaf blades oblong, veins on the lower surface

of the blade impressed . 0. tatei Gleason

5. Leaf blades elliptic-ovate; veins on the lower

surface of the blade not impressed .

. O. paratatei Sastre

6(4). Leaf margin clearly undulate and toothed ..

. 0. superba Engler

6. Leaf margin few-toothed . 7

7(6). Leaf blades 3-6 x 1. 7-2.7 cm .

. O. longistyla Maguire & Steyermark

7. Leaf blades 6-12 x 3-5 cm.

. 0. spruceana Engler

Ouratea Sect. Ouratella

Ouratea cidiana Sastre, sp. nov. TYPE: Brazil.
Para: Mun. Oriximina, Rio Trombetas a mon-
tante do Cachoeira Portura prox. as ilhas Res-
posta, 00°52'S, 57°03'W, 22 aug. 1986, Cid
Ferreira et al. 7971 (holotype, P; isotypes,
INPA not seen, NY not seen). Figure 5.

Ab 0. soderstromii Sastre basi foliae obtusa vel ro-
tundata, margine serrata, inflorescentia 2-8 cm longa
differt.

Shrub 3 m tall, stems glabrous. Leaves coria¬
ceous, elliptic, 6-9.5 x 1.5-4 cm; base rounded;
margin undulate and denticulate; apex acuminate;
principal vein slightly prominent above, with 9-11
unpaired, arcuate lateral veins on each side of prin¬
cipal vein, with fine and parallel intersecondaries;
petiole 5-6 mm long. Inflorescences axillary, spi-
cate, 2-8 cm long, the peduncle 4 mm long. Sepals
5, obovate, 6x3 mm; petals 5, obovate, 7x4
mm; stamens 10, anthers sessile, 5 mm long; style
6 mm long. Fruit not seen.

Distribution. Known only from the type locality,
in Rio Trombetas, Para, Brazil.

Ouratea cidiana differs from O. soderstromii by
its rounded leaf bases (vs. obtuse); serrate margins
(vs. denticulate), and inflorescences 2-8 cm long
(vs. 8-10 cm).

Paratype. BRAZIL. Para: Rio Trombetas, near Cach¬
oeira Porteira, 28 May 1974, Campbell et al. P.22383
(COL, INPA not seen, NY).

Ouratea guriensis Sastre, sp. nov. TYPE: Ven¬
ezuela. Bolivar: Represa Guri, islands and W
side of river, 1 Apr. 1981, Liesner & Gonzalez
11081 (holotype, VEN; isotypes, MO, NY).
Figure 6.

Ab 0. soderstromii Sastre nervis II et III anastomo-
santibus, inflorescentiis 2-8 cm longis, carpophoro fructi
discoideo 3 mm alto (non conico 10-13 mm alto) differt.

Shrub 1.5 m tall, stems glabrous. Leaves coria¬
ceous, unequal, elliptic, 6-9.5 x 1.5-4 cm; base
obtuse or rounded, margin undulate and serrate,
apex acuminate; principal vein slightly prominent
on lower surface, the secondary veins subequal and
anastomosing with the tertiary; petiole 5-7 mm long.
Inflorescences axillary, spicate, 2-8 cm long with
1-3 leaves in the basal part, the peduncle 6 mm
long. Sepals 5, coriaceous, triangular, 7x3 mm;
petals 5, obovate, 9-10 x 1.5-2 mm; stamens 10,
6 mm long. Fruits with carpophore discoid, 3x8
mm, carpels 7-9 x 4-5 mm.

Distribution. Known only from the type locality
in Estado Bolivar near the Guri Dam.

Ouratea guriensis differs from 0. soderstromii
by its anastomosed secondary and tertiary veins,
inflorescence 2-8 cm long (vs. 8-10 cm), and car¬
pophore 3 mm long and discoid (vs. 10-13 mm
long and conic). It differs from O. cidiana by its
anastomosed secondary and tertiary veins.

Paratype. VENEZUELA. Bolivar: Represa Guri, ca.
0.5 km SSW of dam, on a steep forested slope with open
rocks at top, ca. 250-350 m, 7°46'N, 63°00'W, 31 Mar.
1981, Liesner & Gonzalez 11034 (MO, NY, VEN).

Ouratea pseudoguildingii Sastre, sp. nov. TYPE:
Venezuela. Bolivar: Rio Horeda, 1-4 km above
mouth, 90 m, 15 Dec. 1955, Wurdach &
Monachino 39901 (holotype, P; isotype, NY).
Figure 7.

Ab O. guildingii (Planchon) van Tieghem, foliis apice
obtusa (non acuta), nervis II subparallelis (et non inae-
qualibus cum 8-12 magnis inter nervis subparallelis ten¬
ues transversales), differt. Ab O. grosourdyi Van Tiegh¬
em, foliis apice obtusa (non acuta et mucronata), nervis
II et III anastomosantibus, inflorescentiis paniculatis (et
non spiciformibus), differt.

Figures 5-8. —5. Ouratea cidiana Sastre; Cid Ferreira et al. 7971 (holotype, P). —6. Ouralea guriensis Sastre;

Liesner & Gonzalez 11081 (holotype, VEN). —7. Ouratea pseudoguildingii Sastre; Wurdack & Monachino 39901
(holotype, P). —8. Ouratea squamata Sastre; Steyermark et al. 123963 (holotype, VEN).

Volume 5, Number 2
1995

Sastre

Ouratea

199

Tree 3-5 m tall, stems glabrous. Leaves coria¬
ceous, elliptic, 3-5.5 x 1.5-2.5 cm; base rounded;
margin denticulate; apex obtuse; principal vein
prominent below, the secondary veins subparallel,
the secondary and tertiary veins anastomosed; pet¬
iole 2 mm long. Inflorescence axillary, paniculate,
5-10 cm long, the peduncle 7-8 mm long. Sepals
5, coriaceous, obovate, 8x2 mm; petals 5, spa-
thulate, 9x6 mm; stamens 10, a few ornate,
anthers subsessile, 5 mm long; carpels 5, style 5
mm long. Fruit not seen.

Ouratea pseudoguildingii differs from O. guil-
dingii by its obtuse leaf apices (vs. acute) and sub¬
parallel venation. It differs from O. grosourdyi by
its obtuse leaf apices (vs. acute and mucronate),
secondary and tertiary venation anastomosed (vs.
not anastomosed), and paniculate inflorescence (vs.
spicate).

Distribution. Middle Rio Orinoco, Venezuela.

Paratypes. VENEZUELA. Bolivar: Puerto Ordaz,
San Felix, Apr. 1964, Aristiguieta 5280 (NY, VEN).
Guarico: Dto. Infante, Municipio Cabrita, laguna del
Caiman, Finca los Becerros, 21 mar. 1990, S. Tillelt el
al. SPB 279 (P, UCU).

Ouratea squamata Sastre, sp. nov. TYPE: Ven¬
ezuela. Bolivar: Distr. Heres, cerro Marutani,
cumbre, afloramiento de piedra arenisca en la
altiplanicie a lo largo del Rio Carla, afluente de
las cabeceras del Rio Paragua, 3°50'N,
62°15'W, 1200 m, 11-14 jan. 1981, Stey-
ermark et al. 123963 (holotype, VEN; isotype,
NY). Figure 8.

Ab O. poeppigii van Tieghem foliis inaequalibus, nervis
lateralibus pagina superiore impressis, pagina inferiore
non visibilibus, nervis II et III anastomosantibus, differt.

Tree, 10 m tall, young stems with scales. Leaves
coriaceous, unequal, elliptic, sometimes folded, 10-
14 x 4-6 cm; base rounded; margin denticulate;
apex acuminate; principal vein prominent below,
secondary veins subparallel, impressed above, not
apparent below, secondary and tertiary veins anas¬
tomosed, petiole 3-4 mm long. Inflorescence axil¬
lary, paniculate, 16-26 cm long with 1-3 leaves at
the basal part, axes with scales. Flowers not seen.
Fruit with conical carpophore, 5-6 x 5 mm, carpels
5-6 x 4 mm.

Distribution. Known only from the type locality
on Cerro Marutani, Bolivar, Venezuela, near the
headwaters of the Rio Paragua.

Known only from the type specimen; Ouratea
squamata differs from O. poeppigii, an Amazonian
Brazil species, by its unequal leaves, lateral veins

impressed above and not apparent below, and the
secondary and tertiary venation anastomosed.

Key to the Species of Ouratea Sect. Ouratella from
Venezuela and Adjacent Areas

1. Inflorescence spicate, not or few-branched .... 2

1. Inflorescence paniculate .. 10

2(1). Inflorescence 11-30 cm long . 3

2. Inflorescence 2-10 cm long. 4

3(2). Leaves 10-15 x 4-6.5 cm.

. 0. croizatii Steyermark & Maguire

3. Leaves 6-10 x 3-6 cm .

. O. grosourdyi (Tieghem) Steyermark

4(2). Leaf blade with the margin rolled and few¬
toothed ... O. yapacana Sastre

4. Leaf blade flat with the margin toothed . 5

5(4). Leaves 20-60 cm long . 6

5. Leaves less than 15 cm long . 7

6(5). Leaf blade oblong, 30-60 cm long .

. O. ramiflora Sastre

6. Leaf blade ovate, 20-35 cm long .

. O. ornata Maguire & Steyermark

7(5). Leaf blade 3-5 x 2-5 cm.

.O. guaiquinimensis Sastre

7. Leaf blade 6-15 x 1.5-7 cm. 8

8(7). Leaf blade membranous, inflorescences 8-10

cm long . O. soderstromii Sastre

8. Leaf blade coriaceous, inflorescences 2-8 cm

long . 9

9(8). Secondary and tertiary leaf venation reticu¬
late . O. guriensis Sastre

9. Secondary and tertiary leaf venation not re¬
ticulate . O. cidiana Sastre

10(1). Leaf blades with length/width ratio 1-2.5 11

10. Leaf blades with length/width ratio > 2.5 14

11(10). Leaf blades 3-5.5 x 1.5-2.5 cm .

. O. pseudoguildingii Sastre

11. Leaf blades 6-15 x 3.5-6 cm. 12

12(11). Leaf blade with 2 types of lateral veins;

inflorescence 4-7 cm long.

. O. caracasana (Planchon) Engler

12. Leaf blade with lateral veins subequal, in¬
florescence 7-17 cm long . 13

13(12). Venation of the leaf blade not impressed,

inflorescence 7-10 cm long .

. 0. ramossisima Maguire & Steyermark

13. Venation of the leaf blade impressed, in¬
florescence 16-17 cm long .

.O. timehriensis Sastre

14(10). Inflorescences 3-5 cm long.

. O. guildingii (Planchon) Urban

14. Inflorescences > 5 cm long . 15

15(14). Leaf blade with lateral veins subequal, sec¬
ondary and tertiary veins anastomosed .. 16

15. Leaf blade with 2 types of lateral veins,

secondary and tertiary veins not anasto¬
mosed . 1 ‘

16(15). Leaf blades 5.5-8 x 2-3 cm; young stems
without scales; inflorescence axis papillate
. O. papillata Maguire & Steyermark

16. Leaf blades 10-14 x 4-6 cm; young stems

with scales; inflorescence axis not papillate
.... O. squamata Sastre

200

Novon

17(15). Leaf blades coriaceous .

. O. duidae Steyermark

17. Leaf blades membranaceous .

. O. poeppigii Van Tieghem

Ouratea cidiana, 0. ramiflora , and O. poep¬
pigii occur in Amazonian Brazil, and 0. timehrien-
sis occurs in Guyana.

Acknowledgments. I thank Bruce K. Holst for
assistance with the manuscript.

Literature Cited

Aublet, F. 1775. Histoire des plantes de la Guyane
fran£aise 1: 1-261. Paris.

Engler, A. 1876. Ochnaceae. In: Martius, Flora Bras-
iliensis 12(2): 301-332, pi. 62-77. Monachii.

Kanis, A. 1968. A revision of the Ochnaceae of the
Indo-Pacific area. Blumea 16: 1-82.

Sastre, C. 1988. Synopsis generis Ouratea Aublet (Och¬
naceae). Bull. Mus. Natl. Hist. Nat. Paris 4°. ser.,
sect. B Adansonia 1: 47-67.

Tieghem, P. van. 1902. Sur les Ochnacees. Ann. Sci.
Nat. Bot., ser 8, 16: 161-416.

New Species and Combinations in Rubiaceae from
Costa Rica and Panama

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT. The new combinations Psychotria ne-
bulosa (Dwyer) C. M. Taylor and P. roseocrema
(Dwyer) C. M. Taylor are made, and the following
new species are described: Chiococca caputensis
D. H. Lorence & C. M. Taylor, Manettia longi-
pedicellata C. M. Taylor, Pentagonia lobata C.
M. Taylor, Psychotria burgeri C. M. Taylor, Psy¬
chotria saltatrix C. M. Taylor, and Rudgea laevis
C. M. Taylor.

During review of Rubiaceae materials for prep¬
aration of the Manual to the Plants of Costa Rica,
the following new species and taxonomic problems
were discovered. These comments supplement the
treatment by Burger & Taylor (1993).

Chiococca caputensis D. H. Lorence & C. M.
Taylor, sp. nov. TYPE: Panama. Panama: Cer-
ro Jefe, NE of Panama City, 9°15'N, 79°30'W,
12 July 1986, G. McPherson 9744 (holotype,
MO 3624428; isotype, PMA not seen). Figure
IB, C.

Chiococcae belizensi Lundell similis, sed ab ea corollis
albis etiam nervis secundariis brochidodromis semel ar-
cuatis differt.

Glabrous low shrubs or usually woody vines.
Leaves 2-8 cm long, 1.2-4 cm wide, elliptic, at
apex acute to slightly acuminate or sometimes ob¬
tuse, at base cuneate, coriaceous; secondary veins
3-5 pairs, brochidodromous, looping once to inter¬
connect, above and below plane, with the costa plane
or prominulous on both surfaces; petioles 4— 10 mm
long; stipules interpetiolar, triangular, 2-3 mm long,
acuminate to usually apiculate. Inflorescences ax¬
illary, 1 per axil, racemose to sometimes 1-2 times
branched, 3-6 cm long; peduncles 5-15 mm long;
bracts subtending pedicels deltoid to narrowly tri¬
angular, 0.5-1.5 mm long; pedicels 1-4 mm long;
calyx glabrous, green, with hypanthium ellipsoid,
somewhat flattened laterally, 2-2.5 mm long, limb
1-1.3 mm long, divided for ca. 14 its length, lobes
deltoid, entire to ciliolate; corolla glabrous through¬
out, white becoming cream to pale yellow, funnel-
form, tube 6.5-7 mm long, ca. 1 mm diam. at base
and 4-5 mm diam. at apex, lobes narrowly trian¬

gular, 3-4 mm long, acute; stamens included, fil¬
aments ca. 1 mm long, pilose, anthers very narrowly
oblong, 3.5-4 mm long; style 11-11.5 mm long,
twisted, white, papillose throughout, at apex clavate.
Fruit ellipsoid, strongly laterally compressed, 8-12
mm diam., white, fleshy.

Habitat and phenology. Known only from wet
forest on the upper slopes and summit of Cerro Jefe
in central Panama, at 550-1000 m. Collected in
flower in July, in mature fruit September through
February.

Chiococca caputensis is similar in its included
stamens and climbing habit to C. alba (L.) A. S.
Hitchcock, which can be distinguished by its pa¬
pyraceous leaf blades and usually smaller mature
fruits, 4-8 mm in diameter. This new species has
also been confused with C. belizensis Lundell, which
has coriaceous leaves, included stamens, and fruit
size similar to that of C. caputensis, but which can
be distinguished from C. caputensis by its lemon
yellow corollas and secondary leaf veins connected
by one to three additional smaller loops outside the
principal loops. Additionally, C. belizensis is found
from sea level to 1500 m but has been collected
predominantly below 400 m elevation.

Chiococca caputensis has been confused with C.
durifolia Dwyer, a synonym of C. belizensis, and
also with C. jefensis Dwyer, which is a synonym of
Elaeagia nitidifolia Dwyer. The specific epithet
refers to the type locality, Cerro Jefe, which trans¬
lated from the Spanish means “chief’ mountain.
Several other species of Rubiaceae, for example,
Palicourea tubuliflora Dwyer and Psychotria ol-
gae Dwyer & M. V. Hayden, appear to be endemic
to this mountain, and C. caputensis may be a similar
local endemic.

Paratypes. PANAMA. Panama: beyond Goofy Lake
on road to Cerro Jefe, Correa & Dressier 467 (MO),
Croat 15235 (MO); Cerro Jefe, NE of Panama City,
Correa et al. 4576 (MO, PMA), Croat 35865, D’Arcy
12192 (MO), Elias & Hayden 1799 (MO), Folsom &
Harp 1373 (MO), Folsom 3854 (MO), Gentry 2878
(MO), Gentry & Dwyer 5540 (MO), Gentry 6133 (MO),
Kirkbride & Crobbs 9 (MO), McDonagh et al. 47 (BM,
MO), McPherson 9733 (MO, PMA), 10012 ( MO, PMA),
10032 (MO, PMA, PTBG), 15855 (MO, PMA), Mori

Novon 5: 201-207. 1995.

202

Novon

6527 (MO), 7994 (MO), Sytsma 1490 (MO), Tyson et
al. 3207 (MO), Tyson 3374 (MO), Witherspoon 8554
(MO).

Manettia longipedicellata C. M. Taylor, sp. nov.
TYPE: Costa Rica. Limon: Canton de Matina,
Cordillera Talamanca, 200 m aguas abajo de
la confluencia de la quebrada Canabral con el
rio Barbilla, 10°00'10''N, 83°25'30"W, 5 Nov.
1988, G. Herrera 2290 (holotype, CR; isotype,
MO 3649361). Figure 1A.

Flores in cymas paucifloras pedunculis 1-2.5 cm longis
insidentes aggregati, pedicellis 1-2.5 cm longis; lobis ca-
lycinis 4, 2 mm longis; corolla viridula ad rosea, tubo 6-
10 mm longo, lobis 4, 3-4 mm longis.

Glabrescent to retrorsely hispidulous, extensively
twining, herbaceous to suffrutescent vines. Leaves
elliptic to ovate or broadly lanceolate, 3-12 cm long,
1.5-5 cm wide, acuminate at apex, cuneate to trun¬
cate at base, membranaceous; secondary veins 4-
6 pairs; petioles 5-15 mm long; stipules interpetio-
lar, triangular, 1 mm long, with 4-8 cartilaginous
teeth or triangular appendages, these often caducous
and the stipules appearing truncate. Inflorescences
axillary and terminal, cymose, with 1-4 flowers;
peduncles 1-2.5 cm long; bracts triangular, to 2.5
mm long, scarious; pedicels 1-2.5 cm long, often
thickened near apex and flowers and fruits appearing
stipitate; calyx glabrescent, limb 2-4 mm long, di¬
vided to base, lobes 4, narrowly triangular, acute,
reflexed; corolla tubular, externally glabrous, inter¬
nally pilose, green to white or usually pink with
exserted yellow trichomes in throat, tube 6-10 mm
long, lobes ovate, 3-4 mm long, acute. Capsules
obconic, 5-9 mm long, frequently somewhat dicoc-
cous, compressed laterally; seeds orbicular to some¬
what irregular, ca. 2 mm diam.

Habitat and phenology. Wet Caribbean slopes
of Nicaragua and Costa Rica at 100-1100 m, most
frequently collected as a vine climbing on thickets
along streams below 300 m elevation. Collected in
flower and fruit usually concurrently, February to
May and in December.

This new species is similar to Manettia reclinata
L., which can be distinguished in southern Central
America by its six to eight calyx lobes 4-7 mm
long, dark red corollas, and pedicels usually 1 cm
long or shorter, in contrast to four calyx lobes 2-
4 mm long, green to white or pink corollas, and
pedicels 1-2.5 cm long in M. longipedicellata.
Plants of M. reclinata from Mexico and northern
Central America may have peduncles and pedicels
to 4 cm long, but differ in the other characters.

Paratypes. COSTA RICA. Alajuela: Piedades de San
Ramon, Brenes 4555 (CR). Cartago: Florencia Norte,
CATIE, Turrialba, Poveda 3109 (CR). Heredia: Finca
La Selva, the OTS field station near Puerto Viejo de
Sarapiqui, near the junction of the Rios Puerto Viejo and
Sarapiqui, Almeda & Anderson 5130 (F), Folsom 8851
(DUKE), 10032 (DUKE), Grayum 1144 (DUKE), 2711
(DUKE), Hammel 8099 (DUKE), 11752 (DUKE), Opler
1645 (CR), Wilbur 28181 (DUKE), 28308 (DUKE),
38626 (DUKE), 39485 (DUKE). Limon: Canton de
Pococi, along road between Mata de Limon (on quadrangle
map as “Milloncito”) and Millon, 10°24'30"N, 83°37'W,
Grayum & Hammel 8407 (CR, MO). NICARAGUA.
Zelaya: Cano Monte Cristo, “Las Faldas,” 11°36'N,
83°51'W, Moreno 14776 (MO); Montecristo, al N de
Barra Punta Gorda, 11°34'N, 83°48'W, Sandino 2232
(MO); Cano Costa Riquita, ca. 1.8 km SW of Colonia
Naciones Unidas, above road between there and Colonia
Nueva Leon, 11°43'N, 83°18'W, Stevens 4942 (MO).

Pentagonia lobata C. M. Taylor, sp. nov. TYPE:
Costa Rica. Puntarenas: Canton de Golfito, Re-
serva Natural de Vida Silvestre Golfito, final
del camino hacia las torres, 8°39'52"N,
83°10'00"W, 7 Dec. 1992, B. Hammel 18619
(holotype, MO 4355639; isotypes, CR, F, MO
4355638). Figure ID.

A speciebus ceteris Pentagoniae Bentham foliis pin-
natifidis etiam calyce spathaceo differt.

Monocaulous shrub or small tree to 7 m tall;
stems lenticellate. Leaves 62-99 cm long, 60-90
cm wide, deeply pinnatifid, with sinuses extending
to within 3-5 cm of costa, with lobes 4-6 on each
side, these acute to somewhat acuminate, 7-14 cm
wide, the basalmost lobes usually ca. % as long and
wide, the entire blade chartaceous to subcoriaceous,
glabrous and smooth adaxially, abaxially glabrescent
to usually spreading-puberulous or shortly pilosulous
on principal veins and sometimes also on lamina,
abaxially lamina also finely striate, costa and longer
secondary veins prominent above and below, shorter
secondary veins slightly thickened, minor venation
not visible; petioles 12-22 cm long; stipules ligu-
late, 6-8 cm long, 2.5-3.5 cm wide, coriaceous,
acute, with costa thickened near base, finely ap-
pressed-puberulous to sericeous. Flowers hermaph¬
roditic, glomerulate in groups of ca. 12-20 in leaf
axils, each inflorescence enclosed by 1 bract resem¬
bling a stipule, these bracts 8-9 cm long, 3-7 cm
wide, each flower with a stipe 1-3 mm long and
subtended by 1 floral bract, these floral bracts 15-
25 mm long, 6-12 mm wide, ovate, acute; hypan-
thium glabrous, cylindrical to ellipsoid, 3—5 mm
long, 2-3 mm diam.; calyx limb red, coriaceous,
glabrous, 2.5-4 cm long, spathaceous, splitting for
ca. % of its length, acute at apex; corolla green to
cream, funnelform, carnose, fibrous, glabrous

Volume 5, Number 2
1995

Taylor

Rubiaceae from Costa Rica and Panama

203

Figure 1. —A. Manettia longipedicellata C. M. Taylor (from Grayum 1144, DUKE), habit. B, C. Chiococca
caputensis C. M. Taylor & D. H. Lorence. —B. Habit (from McPherson 10012, MO). —C. Flower (from McPherson
9744, MO). —D. Pentagonia lobata C. M. Taylor, flower (from Hammel 18619, MO). —E. Psychotria burgeri
c. M. Taylor, habit (from Haber 9778, F). F, G. Rudgea laevis C. M. Taylor. —F. Flower (from Haber 452, MO).
—G. Habit (from Haber 1070, MO). A, B, D, E, G to 5-cm scale; C, F to 5-mm scale.

204

Novon

throughout, tubes 38-46 mm long, 3-5 mm diam.
at base, 10-15 mm diam. at apex, curved, lobes 5,
deltoid, 4-8 mm long; stamens 5, included, fila¬
ments attached ca. V 3 of length of tube above base,
ca. 15 mm long, swollen and densely pilose to villous
at attachment point, glabrous otherwise, anthers
narrowly oblong, ca. 3.5 mm long; style included,
ca. 2 cm long, stigmas 1, lanceolate, ca. 2 mm long;
disk bilobed, ca. 1.5 mm high. Fruit not seen.

Habitat and phenology. Moist forest in the vi¬
cinity of Golfito and on the Osa Peninsula in the
Golfo Dulce region of Costa Rica at 300-500 m.
Collected in flower in July and December.

This species is included in Pentagonia based on
its fleshy monocaulous habit (characteristic though
not exclusive in this genus), relatively large, firm-
textured leaves with the abaxial surface finely stri¬
ate, and five-merous, relatively large, fleshy flowers
borne in contracted axillary inflorescences. The fine¬
ly striate texture of the abaxial leaf surfaces with
the minor venation not at all visible seems to be
unique to this genus.

Pentagonia lobata is distinguished within Pen¬
tagonia by its combination of pinnatifid leaves and
a spathaceous calyx. Pinnatifid leaves also charac¬
terize P. tinajita Seemann of Costa Rica and Pan¬
ama (here circumscribed to include P. alfaroana
Standley and P. gymnopoda Standley), which can
be separated by its regularly lobed calyx limb. A
spathaceous calyx also characterizes P. spathicalyx
K. Schumann of Amazonian Brazil, Ecuador, and
Peru, which has elliptic entire leaves.

Pentagonia is poorly known taxonomically and
ecologically. Individuals of most species are appar¬
ently infrequent, or at least infrequently collected,
and good specimens are difficult to make due to the
large fleshy leaves and flowers. Some species cannot
be separated vegetatively. Flowering characteristics
appear to be reliable for separating species, but
flowering specimens are sometimes difficult to relate
to fruiting specimens (the majority of collections).
The range of size and shape variation seen in her¬
barium material is notable in some species, to 100%
in some features (see descriptions in Burger & Tay¬
lor, 1993): this may be real morphological variation,
or it may reflect variable shrinkage or other artifacts
of the preparation of dried specimens. Some species
of Pentagonia were originally separated based on
details of leaf shape such as acute versus rounded
leaf bases, but with more material available these
conditions often show continuous variation and are
not adequate to distinguish species. The deeply pin¬
natifid leaves found in some Pentagonia species are
here regarded as a consistent character at the spe¬

cies level, although this has not been verified with
field observations. Good taxonomic knowledge of
Pentagonia will require extensive field-based study.

Paratypes. COSTA RICA. Puntarenas: Canton de
Osa, Reserva Forestal Golfo Dulce, Rancho Quemado,
8°42'00"N, 83°35'40”W, Hammel et al. 18309 (CR, F,
MO).

Psychotria burgeri C. M. Taylor, sp. nov. TYPE:
Costa Rica. Puntarenas: Canton de Puntarenas,
Monteverde, along Quebrada Maquina,
10°18’N, 84°48’W, 12 Mar. 1990, W. Haber
9778 (holotype, CR; isotypes, F, MO 4619858,
US). Figure IE.

A Psychotria pleuropoda J. Donnell Smith foliis sub-
sessilibus basim rotundatis ad truncatis differt.

Small trees to 3 m tall, glabrous. Leaves narrowly
elliptic to oblanceolate, 7-13 cm long, 1.4-3.6 cm
wide, at apex acute to slightly acuminate, at base
truncate to rounded, chartaceous, glabrous above
and below; secondary veins 10-12 pairs, looping to
interconnect in a rather straight submarginal vein,
plane above, prominulous below, without domatia;
petioles stout, 2-3 mm long, glabrous; stipules unit¬
ed around the stem, deciduous below distalmost 1-
3 nodes, ovate, split along one side, 6-15 mm long,
acute or frequently bilobed with the lobes acuminate,
sometimes strongly costate, glabrous, ciliate. Inflo¬
rescences terminal, pyramidal; peduncles 2-4.5 cm
long; panicles 2-3 cm long and wide, branched once,
with bracts 3-4 mm long, triangular to ovate, ciliate,
acute; flowers sessile in 3 glomerules of 3-5; hy-
panthium not seen; calyx limb green, glabrous, 1-
1.5 mm long, lobed for ca. 14 °f its length, lobes
deltoid, acute, ciliolate; corollas not seen. Infruc-
tescences similar to inflorescences; fruit ellipsoid,
8-11 mm long, 6-12 mm diam., becoming red;
pyrenes with 4-5 low rounded longitudinal ridges.

Habitat and phenology. Wet premontane forest
at ca. 1400 m in the Cordillera de Tilaran, Costa
Rica. Collected in flower and fruit in March.

Psychotria burgeri is placed in subgenus Psy¬
chotria based on its deciduous stipules, red fruits,
and distinctive gray drying color. Subgenus Psy¬
chotria is well known in Central America (Hamilton,
1989), and even though its flowers have not been
seen, this new species is clearly undescribed. It is
similar to P. pleuropoda J. Donnell Smith of north¬
ern Central America, which can be separated by its
leaves with acute to tapered bases and secondary
veins 15-18 pairs, petioles 2-10 mm long, and
elevational range of 50-500 m; P. pleuropoda is
so far not known from Costa Rica. Range disjunc-

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Taylor

Rubiaceae from Costa Rica and Panama

205

tions and disjunct species pairs between northern
Central America and the Cordillera de Tilaran in
Costa Rica are frequent in Costa Rican Rubiaceae
(Burger & Taylor, 1993), although few are accom¬
panied by habitat shifts such as this. The specific
epithet honors William C. Burger, a long-time stu¬
dent of the flora and phytogeography of Costa Rica.

Psychotria nebulosa (Dwyer) C. M. Taylor, comb,
nov. Basionym: Coussarea nebulosa Dwyer,
Ann. Missouri Bot. Gard. 67: 131. 1980.
TYPE: Panama. Chiriqui: Monte Rey near Bo-
quete, 20 July 1971, T. B. Croat 15868 (ho-
lotype, MO 2162999; isotype, MO 4043108).

Rudgea chiriquiensis Dwyer, Ann. Missouri Bot. Gard.
67: 476. 1980. Syn. nov. Not Psychotria chiri¬
quiensis (Standley) C. M. Taylor & W. C. Burger,
1993. TYPE: Panama. Chiriqui: Cerro Colorado,
along road above San Felix 29 km above bridge over
Rio San Felix (7.9 km above turnoff to Escopeta),
14 July 1976, T. B. Croat 37071 (holotype, MO
2798805; isotype, MO 2389189).

Study of additional material available since the
publication of Coussarea nebulosa and Rudgea chi¬
riquiensis shows that they refer to the same species,
and that this species is best placed in Psychotria.
The inflorescences on the type specimens of R.
chiriquiensis are older and consequently larger and
relatively more open than those on the type speci¬
mens of C. nebulosa, but additional material shows
this to be a developmental series, and no other
features separate these specimens. The mature fruits
are fleshy drupes 12-14 mm long with two hard,
longitudinally ridged pyrenes. This fruit structure is
characteristic of both Psychotria and Rudgea but
excludes this species from Coussarea, which has
smooth, thin-walled, usually solitary seeds in fleshy
to leathery fruits. Rudgea is separated from Psy¬
chotria by its glandular-fimbriate stipules. The stip¬
ules of this species are shortly bidentate and not at
all glandular, and it is here transferred to Psycho¬
tria.

Psychotria nebulosa belongs to subgenus Het-
eropsychotria Steyermark, which is characterized
by green or gray-green drying color, persistent usu¬
ally bidentate stipules, and blue or black mature
fruits. The mature fruit color of P. nebulosa is not
known. Based on inflorescence, stipule, and fruit
morphology it seems closely related to P. eurycarpa
Standley, which can be separated by its smaller
leaves that are usually shiny on the adaxial surfaces
and pyrenes that are longitudinally angled rather
than ridged, and to P. umbelliformis Dwyer & M.
V. Hayden, which is densely pilosulous throughout.
Psychotria nebulosa is also similar in general aspect

to Coussarea austin-smithii Standley, which can be
distinguished by its subglobose to ellipsoid fruits with
smooth to slightly angled, usually solitary seeds.
Psychotria nebulosa appears to be restricted to
Chiriqui and Veraguas provinces in western Panama.

Psychotria roseocrema (Dwyer) C. M. Taylor,
comb. nov. Basionym: Coussarea roseocrema
Dwyer, Ann. Missouri Bot. Gard. 67: 133.
1980. TYPE: Panama. Panama: El Llano-
Carti highway, 6-10 km N of El Llano, 13
Apr. 1973, R. L. Dressier 4334 (holotype,
MO 2126258).

Additional material collected since the publication
of this name shows that the fruits of this species are
fleshy and blue-black at maturity with two hard,
longitudinally ridged pyrenes. As discussed above,
this fruit structure is characteristic of Psychotria
but is not found in Coussarea, and accordingly this
species is here transferred to Psychotria. Psycho¬
tria roseocrema belongs to subgenus Heteropsy-
chotria.

Psychotria roseocrema is similar in general as¬
pect to P. umbelliformis, which can be distinguished
by its pilosulous pubescence throughout, and to P.
eurycarpa and Coussarea austin-smithii, both of
which lack the strong looping submarginal leaf veins
found in P. roseocrema.

Psychotria saltatrix C. M. Taylor, sp. nov. TYPE:
Costa Rica. Cartago: Refugio Nacional Tapanti,
por la orilla del Rio Grande de Orosi, 9°45'N,
83°47'W, 14 Aug. 1991, B. Hammel et al.
18328 (holotype, CR; isotypes, F, MO
4619859). Figure 2.

A speciebus ceteris Psychotriae L. subg. Psychotriae
foliis stipulis bracteis calycibusque majoribus etiam in-
fructescentiis ut videtur axillaris differt.

Soft shrubs to 2 m tall; stems puberulous to hir-
tellous becoming glabrescent with age. Leaves ellip¬
tic, 18-32 cm long, 6.5-13.5 cm wide, at apex
acute to acuminate with the tip ca. 1 cm long, at
base acute to usually tapered, drying membrana¬
ceous, above glabrous, below glabrous on lamina but
moderately to usually densely puberulous to pilo¬
sulous on veins; secondary veins 18-24 pairs, di¬
verging from costa at 80-90° or more, usually loop¬
ing to interconnect in an undulating to rather straight
submarginal vein, with 1-2 secondary loops as well,
plane above, prominulous below, without domatia;
petioles 1.5-8 cm long; stipules interpetiolar, de¬
ciduous after 1-2 distal nodes, obovate, 25-35 mm
long, 12-16 mm wide, acute, basal and central

206

Novon

Figure 2. Psychotria saltatrix C. M. Taylor (from Hammel 18030 , MO).

portion chartaceous to subcoriaceous and smooth,
marginal portions membranaceous and with fine par¬
allel veining, both portions puberulous or pilosulous
becoming glabrescent. Inflorescences terminal, py¬
ramidal; peduncles 3-60 mm; panicles 2-10 cm
long, 4-12 cm wide, with branches spreading at ca.
80-90° or occasionally more, with 2 branches at
the basalmost nodes or on larger inflorescences fre¬
quently 4 branches and then these generally similar
in size but 2 spreading and 2 reflexed, puberulous
to hirtellous, with bracts lanceolate, membrana¬
ceous, puberulous to glabrescent, ciliolate, acute, 2-
10 mm long, those subtending primary branches 8-
10 mm long and those subtending flowers 2-4 mm
long; flowers sessile or very shortly stipitate in glom-
erules of 3-5, each subtended by 1-3 bracts; hy-
panthium turbinate, 1 — 1.2 mm long, glabrescent to
puberulous; calyx limb green, glabrous to puberu¬
lous, 1.5-2 mm long, divided for ca. %-% its length,
lobes 5, deltoid to ovate, ciliolate, acute; corolla
salverform, white, externally glabrous or usually
densely puberulous at top of tube, internally glabrous
except densely pilose in 10 tufts at stamen attach¬
ment, tubes 2-3 mm long, 1-1.2 mm diam., lobes
5, ligulate, 2-3 mm long, somewhat thickened at
apex in bud, becoming acute at anthesis; stamens
5, exserted, attached just below top of corolla tube,
filaments 0.5-0.8 mm long, anthers oblong, ca. 1
mm long; style exserted, 3.5-4.5 mm long, stigma
lobes 2, linear, ca. 0.5 mm long; disk annular,
smooth, ca. 0.3-0.5 mm long. Infructescences over¬
topped by sympodial stem growth and appearing

pseudoaxillary, as fruit matures usually displaced to
3rd or 4th node below apex, more expanded than
inflorescences, to 12.5 cm long and 16 cm wide,
bracts often deciduous ; fruit with stipe ca. 0.5 mm
long, obovoid, 6-6.5 mm long, 4-4.5 mm wide,
puberulous to glabrescent, becoming orange to red;
pyrenes with 4—5 rather sharp longitudinal ridges.

Habitat and phenology. Costa Rica to Colombia,
in wet premontane and montane forest at 1250-
2100 m. Collected in flower in May and August, in
fruit January, July, September, and October.

Psychotria saltatrix is placed in subgenus Psy¬
chotria (Hamilton, 1989) based on its deciduous
interpetiolar stipules, orange to red mature fruits,
and distinctive gray-green to brown drying color.
Within this subgenus it is distinguished by its ap¬
parently pseudoaxillary inflorescences, relatively
large leaves and stipules, secondary veins that often
diverge from the costa at more than a 90° angle
and usually form a distinct submarginal vein, well-
developed inflorescence bracts, relatively long calyx
limbs, obovoid fruits, and very wet premontane or
montane habitats. The inflorescences are terminal
in position but the infructescences are typically over¬
topped by sympodial growth from one axil. This
growth pattern is seen in other species of subgenus
Psychotria, but is rarely as strongly or consistently
developed as in this new species. Relatively few other
species of subgenus Psychotria inhabit wet pre¬
montane or montane forest. In the two flowering
collections seen, both the stamens and the style are

Volume 5, Number 2
1995

Taylor

Rubiaceae from Costa Rica and Panama

207

exserted in all of the flowers, and P. saltatrix may
be homostylous. This condition (if actually present)
would be highly unusual in the subgenus. The spe¬
cific epithet refers to the occurrence of this species
in widely separated wet cool sites in the Andean
Cordillera, western Panama, and Costa Rica.

Three fruiting specimens from similar habitats in
southern Colombia and Ecuador, de Benavides 9292
(MO), Sparre 17686 (S), and Hoover 1993 (MO),
are similar to P. saltatrix but can be separated by
their shorter calyx limbs and bracts subtending the
flowers, both ca. 1 mm long, secondary leaf veins
14-15 pairs, and stipules 5.5-6 cm long. These
specimens may represent a separate, apparently un¬
named taxon, or they may reflect clinal variation in
P. saltatrix', no conclusion can be made without
flowering material.

Paratypes. COLOMBIA. Antioquia: municipio de
Valdivia, vereda San Feraiin, sitio “Alto de Ventanas,”
2 km al NE de la via Medellin-Valdivia en la via que
conduce al municipio de Briceno, 7°50'N, 75 C 32 W, Cal-
lejas et al. 10735 (F, HUA, MO); municipio de Valdivia,
vereda San Fermin, cerca 120 km NE de Medellin, en
la via a Briceno, finca de Los Restrepo, 07°06'N, 07°32'W,
Callejas & Gomez 10913 (HUA, MO); municipio de
Urrao, Parque Nacional Natural “Las Orquideas,” cam-
ino de Calles hacia Venados arriba. Sector Bellavista,
Cogollo et al. 5028 (JAUM, MO). Narino: municipio
Ricaurte, Reserva Natural La Planada, Restrepo 637
(MO). COSTA RICA. Cartago: Tapanti, /. Chacon et
al. 1432 (CR, MO). San Jose: Parque Nacional Braulio
Carrillo, Estacion Zurqui, 10°03'30"N, 84°01'W, Ham-
mel et al. 18030 (CR, F, MO). PANAMA. Chiriqui:
Cerro Colorado Mine, Antonio 4878 (MO). Darien: Cer-
ro Pirre, valley between Pirre and next most southerly
peak, Folsom 4441 (MO); Cerro Pirre, top of mountain
and ridge just S of Pirre, Folsom 4521 (MO).

Rudgea laevis C. M. Taylor, sp. nov. TYPE: Costa
Rica. Puntarenas: Monteverde, Bajos de La
Lindora, rio Guacimal, 10°18'N, 84°48’W, 19
Feb. 1988, W. Haber & E. Bello 8271 (ho-
lotype, CR; isotype, MO 3646480). Figure IF,
G.

A Rudgea laurifolia (Kunth) Steyermark venulis pau-
cioribus calycibus corollisque minoribus differt.

Shrubs or small trees to 6 m tall; stems to 15
cm DBH, glabrous. Leaves elliptic, 5.5—12 cm long,
1 -8 5.5 cm wide, at apex acute, at base cuneate
to obtuse, drying chartaceous, glabrous above and
below; secondary veins 4-7 pairs, not looping to
interconnect, above plane to somewhat sulcate, be¬
low prominulous, usually with foveolae in axils; pet¬
ioles glabrous, 4-12 mm long; stipules glabrous,
united around stem into a persistent continuous trun¬
cate sheath 1.5—2 mm long bearing on each side 1
caducous deltoid gland ca. 1 mm long and wide.
Inflorescences terminal, pyramidal to somewhat

rounded, green, pilosulous to glabrescent; peduncles
1.5-3 cm long; panicles 2.5-3 cm long, 4-8 cm
wide, with bracts 0.5-1 mm long, deltoid, obtuse;
flowers sessile in dichasial cymes of 5-9, with hy-
panthium turbinate, ca. 1 mm long, glabrous; calyx
limb green, glabrous, 0.3-0.5 mm long, subtruncate
to 5-denticulate; corolla salverform, white, glabrous
externally and internally except barbate in throat;
tube 3-4 mm long, 1-2 mm diam.; lobes 5, ligulate,
3-4 mm long, acute, unappendaged; stamens 5,
exserted, attached just below top of corolla tube,
filaments 0.8-1 mm long, anthers ca. 2 mm long,
narrowly oblong; style exserted, 6-6.5 mm long,
puberulous in distal portion, stigma lobes 2, linear,
ca. 1 mm long; disk annular, smooth, ca. 0.5 mm
high. Infructescences similar in size, proportion, and
color to inflorescences; fruit globose, 6-10 mm diam.,
glabrous, white; pyrenes smooth.

Habitat and phenology. Costa Rica, in wet pre-
montane and lower montane forest of Cordillera de
Tilaran at 1000-1400 m. Collected in flower in
February and April, in fruit June, September, and
November.

This new species is similar in aspect to Rudgea
laurifolia (Kunth) Steyermark, which can be dis¬
tinguished from R. laevis by its several to numerous
acicular stipular glands, more numerous secondary
leaf veins (9-11 pairs), longer calyx limb (2.5-2.7
mm long), longer corolla tube (ca. 7.5 mm long),
shortly appendaged corolla lobes, and lowland wet
forest habitat. Rudgea laurifolia is known only from
the basins of the Rios Negro and Orinoco in South
America.

Paratypes. COSTA RICA. Alajuela: Reserva Biolo-
gica Monteverde, Finca Ston’s, 10°19'N, 84°43'W, Ob-
ando 140 (CR, MO). Puntarenas: Monteverde comu-
nidad, 10°18'N, 84°48'W, Bello 4 (CR), Haber 452
(MO), 1070 (MO).

Acknowledgments. I thank the curators of CR,
DUKE, F, and HUA for making specimens available
for study; R. Callejas for facilitating fieldwork in
Antioquia, Colombia; R. E. Gereau for revision of
the Latin diagnoses; and W. C. Burger, M. Grayum,
B. Hammel, D. Lorence, and W. D. Stevens for
helpful comments. Travel related to this work was
supported in part by the National Science Foun¬
dation (BSR 87-00068).

Literature Cited

Burger, W. B. & C. M. Taylor. 1993. Flora Costari-
censis: Rubiaceae. Fieldiana, Bot. n.s. 33: 1-333.
Hamilton, C. 1989. A revision of Mesoamerican Psy-
chotria subg. Psychotria (Rubiaceae). Ann. Missouri
Bot. Card. 76: 67-111, 386-429, 886-916.

Erratum

Ten, Chi & Jun-liang Yang. 1994. Roegneria triden-
tata, a new species of Triticeae (Poaceae) from Qing-
hai, China. Novon 4: 310-313.

On page 313, first column, four lines from bottom

should read “Chromosome 19 has a large satellite,
and chromosome 21 has a very short arm connected
with a small satellite.”

Volume 5, Number 2,

pp. 119-208 of NOVON was published on 21 June 1995.

Volume 5
Number 3
1995

_-JS0VON

$£p 121995

pMtDEN UBRARt,

Novelties and Notes in North American Aristida (Gramineae)

Kelly W. Allred

Range Science Herbarium, Department of Animal & Range Sciences,

New Mexico State University, Las Cruces, New Mexico 88003, U.S.A.

Jesus Valdes-Reyrm

Departamento de Botanica, Universidad Autonoina Agraria “Antonio Narro,”
Saltillo, Coahuila 25315, Mexico

Abstract. New taxa of Aristida include three re¬
lated to A. schiedeana Trinius & Ruprecht (A. pe-
tersonii sp. nov., A. eludens sp. nov., and A. spa-
nospicula sp. nov.) and one variety of A. purpurea
Nuttall (var. perplexa var. nov.). Two species are re¬
duced in rank (A. schiedeana var. orcuttiana [Vasey]
comb. nov. and A. purpurea forma brownii [War-
nock] comb, nov.), and one species is reinstated (A.
curvifolia Fournier). A key is given for the A. schie¬
deana complex.

Resumen. Nuevos taxa de Aristida que incluyen
tres especies con las aristas laterales reducidas, re-
lacionadas con A. schiedeana Trinius & Ruprecht
(A. petersonii sp. nov., A. eludens sp. nov., y A. spa-
nospicula sp. nov.) y una variedad con las aristas
laterales desarrolladas de A. purpurea Nuttall (var.
perplexa var. nov.) son descritos. Dos especies son
reducidas en rango (A. schiedeana var. orcuttiana
[Vasey] comb. nov. y A. purpurea forma brownii
[Wamock] comb, nov.) y una especie es reinstalada
(A. curvifolia Fournier). Una clave para el complejo
A. schiedeana es presentada.

The genus Aristida (Arundinoideae: Aristideae)
comprises approximately 300 species found
throughout the world, but its members are partic¬
ularly frequent in warm, semiarid environments.
Many species have three awns terminating the sin¬
gle floret, but it is not uncommon to find single-
awned relatives of three-awned taxa. Species are

notorious for their taxonomic difficulty, due partly,
we believe, to undescribed variation that obscures
the taxonomic boundaries. Major revisionary efforts
include those of Henrard (1927, 1928, 1929) and
Hitchcock (1924, 1935).

Aristida petersonii Allred & Valdes-Reyna, sp.
nov. TYPE: Mexico. Oaxaca: 7.7 km NW of
Tlaxiaco on road to San Juan Mixtepec, steep
slopes with Pinus, Quercus, and Arbutus, elev.
2020 m, 2 Sep. 1990, P. M. Peterson & A.
Campos-Villanueva 9731 (holotype, NMCR; is¬
otypes, ANSM, US). Figure 1.

Species nova propria vaginis lanosis, glumis glabris et
valde inversis (primis secundis longioribus), paniculis an-
gustis, rostris tortis lemmatum, et aristis singularibus a
congeneribus Americanae borealis diversa.

Plants perennial, tufted; culms 50-100 cm tall,
erect, unbranched except for basal tillering; inter¬
nodes glabrous, striate, terete. Sheaths striate,
shorter than the intemodes, rounded on the back,
loosely lanose, the hairs curling and tangled, the
upper sheaths sparsely lanose to glabrous; collars
often with a tuft of longer cobwebby hairs at the
comers; throat glabrous except for the ligule. Lig-
ules a minute fringe of hairs less than 0.5 mm long.
Blades glabrous, flat when fresh and mature (new
leaves convolute), folded upon drying, 15-30 cm
long, 2-2.5 mm wide when flat, the lateral nerves
generally coalescent and the margin thickened, the

Nnvniv 5- 900-999. 1QQ5

210

Novon

F**"® 1 - Aristida petersonii Allred & Valdfe-Reyna (Peterson & Campos-Villanueva 9731). —A. Vegetative habit.
—B. Sheath pubescence. —C. Inflorescence. —D. Spikelet: glumes left, floret right.

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

211

Figure 2. Geographic distributions of Aristida petersonii Allred & Valdes-Reyna (stars), A. eludens Allred & Valdes-
Reyna (closed circles), and A. spanospicula Allred & Valdes-Reyna (squares).

marginal veins ca. 0.5 mm wide. Panicles 20—30
cm long, narrow, contracted, erect, the nodes gla¬
brous; primary branches few, single or paired at the
nodes, erect to appressed, 4- to 8-flowered, the low¬
er ones 6—12 cm long; secondary branches weakly
developed, 2- to 3-flowered, appressed; pulvini ab¬
sent in the axils of the primary and secondary
branches and pedicels, the spikelets thus ap¬
pressed along the branches and main axis. Glumes
glabrous, inverse, membranous and sub-hyaline, 1-
nerved, light tan to purplish, the apices acute but
not awned, the second glume Vi to % the length of
the first; first glume (9—)10—13 mm long; second
glume 6—7 mm long. Lemmas glabrous, mottled, 7—
8 mm long from the base of the callus to the twisted
portion of the beak; beak prominently twisted, (4—)
5—6 mm long; central awn not twisted, geniculate-
bent, 7—9 mm long; lateral awns completely re¬
duced, represented by tiny points ca. 0.1 mm long
at the end of the beak; callus ca. 0.5 mm long, with
straight hairs to 1 mm long. Palea completely en¬
closed by the lemma, 2-nerved, hyaline, obtuse to
rounded, 1—1.5 mm long. Lodicules 2, flabellate, 1—

1.5 mm long. Caryopsis fusiform, chestnut-brown,
4.5-5.5 mm long.

Flowering known only from September, but pre¬
sumed August-October. Distribution. Pine/oak/ju¬
niper woodlands; Mexico, known only from south-
central Oaxaca (Fig. 2).

The specific epithet, petersonii, honors the col¬
lector of this species, Paul M. Peterson of the
Smithsonian Institution.

Aristida petersonii is one of three perennial spe¬
cies in North America with lanate or floccose
sheaths, the other two being A. lanosa Muhlenberg
ex Elliott and A. scribneriana A. S. Hitchcock. Ar¬
istida lanosa, of the southeastern United States, dif¬
fers in having sheaths mostly longer than the in-
temodes, blades 2-6 mm wide, panicles mostly 30-
70 cm long with lanose tufts in the branch axils,
longer glumes (though they share the inverse po¬
sition), and three well-developed awns. Aristida
scribneriana, of the central highlands of Mexico,
differs in having lanose intemodes (including the
peduncles) and blades (both surfaces), sparsely vil-

212

Novon

lous glumes that are nearly equal in length, and
normally three well-developed awns. Other single-
awned Aristida with narrow panicles that have gla¬
brous (or very sparsely pilose) sheaths and equal
glumes are A. eludens, A. spanospicula, and epul-
vinate forms of A. pansa. Aristida petersonii is the
only North American Aristida with lanose sheaths,
narrow panicles, glabrous glumes in the inverse po¬
sition, and single-awned spikelets.

Paratypes. MEXICO. Oaxaca: 24.5 km W of Tlax-
iaco and 7.4 km NE of San Juan Mixtepec, gravelly slopes
with Pinus, Quercus, Arbutus and scattered Juniperus, elev.
2220 m, 3 Sep. 1990, P. M. Peterson & A. Campos-Vil-
lanueva 9746 (ANSM, NMCR, US).

Aristida schiedeana Trinius & Ruprecht
Complex

The Aristida schiedeana complex includes A.
schiedeana, A. orcuttiana Vasey, A. laxa Cavanilles,
and the two new species A. eludens and A. spano¬
spicula. This complex is characterized by having
flat blades often curling like wood shavings, gen¬
erally equal glumes, a twisted lemma beak, and
single awns (except A. laxa). Its members are cus¬
tomarily found as sparse understory in pine/oak for¬
ests of the Mexican and Central American central
cordilleran mountain ranges.

Aristida eludens Allred & Valdes-Reyna, sp. nov.
TYPE: Mexico. Coahuila: San Lorenzo Can¬
yon, ca. 8 km S of Saltillo, mountain scrub
grassland vegetation with Berberis, Bouteloua,
Hilaria, Dasylirion, very rocky sandy loam, flat
bajada on N-facing side of canyon, 6000 ft.
(1830 m), 3 Sep. 1991, J. Valdes-Reyna 2254
(holotype, ANSM; isotype, NMSU). Figure 3.

Species nova laminis ubi vivis planis vel plicatis laxe,
paniculis angustis absque pulvinis axillaribus praeter in-
terdum ramos infimos, glumis subaequalibus, et aristis
singularibus a congeneribus Americanae borealis diversa.

Plants perennial, tufted; culms (35-)40-65(-80)
cm tall, erect, branching only at the base; inter-
nodes terete, glabrous, leaves basal and cauline,
light green. Sheaths longer than the intemodes, gla¬
brous or scaberulous except for the summit, round¬
ed on the back; collars with a tuft of cobwebby hairs
at the comers (becoming glabrous upwards), some¬
times with a line of minute hairs across the back;
throat glabrous or puberulent. Ligules a fringe of
hairs 0.2-0.3 mm long. Blades flat or loosely folded
when fresh, rolled upon drying, glabrous or sca¬
berulous abaxially, puberulent or glabrate adaxially
especially toward the base, 12-35 cm long, 1-1.5
mm wide when flat. Panicles 10-22 cm long and

1— 2 cm wide, with 16 or more spikelets, narrow,
contracted, the nodes glabrous; primary branches
erect, appressed to the main axis except occasion¬
ally the lowermost branch divaricate, naked at the
base, the lower ones 5—9 cm long; pulvini absent,
the spikelets appressed, only occasionally present
in the axil of the lowermost branch and causing it
to spread from the axis. Glumes glabrous, brownish,
subequal or the first slightly shorter or longer, 1-
nerved, 8-13 mm long, the apices acute and often
with a mucro. Lemmas glabrous, mottled, 10-13
mm long from the base of the callus to the diver¬
gence of the awns; beak twisted, (3-)4-5 mm long;
central awn not twisted, geniculate-bent, 5-10 mm
long; lateral awns highly reduced, erect, 0.1—3 mm
long; callus 0.5—1 mm long with stiff hairs ca. 1
mm long. Paleas completely enclosed by the lem¬
ma, 2-nerved, hyaline, rounded, ca. 1 mm long,
shorter than the lodicules. Stamens 3, the anthers

2- 2.5 mm long. Lodicules 2, flabellate, 1—1.5 mm
long. Caryopses fusiform, chestnut-brown, ca. 6 mm
long.

Flowering April—May, August—November. Distri¬
bution. Desert scrub/grassland habitats in calcare¬
ous soil, but extending up into the pine/oak zones;
Mexico, in the states of Chihuahua, Coahuila,
Durango, Guanajuato, Nuevo Leon, Oaxaca, Que-
retaro, and San Luis Potosf (Fig. 2).

The epithet eludens refers to the tendency of
plants of this species to grow only within the
branches and foliage of protective shrubs, evading,
as it were, grazing animals.

Aristida eludens is the desert scrub/grassland
representative of the A. schiedeana complex. Aris¬
tida eludens is distinguished most easily from the
other species of the complex by possessing con¬
tracted panicles where the primary branches lack
pulvini; it also differs by having cauline leaves with
non-coiling blades and by the habitat. Aristida elu¬
dens occurs mostly in the desert scrub and semi-
desert grasslands of northern Mexico, and only in¬
frequently extends into the lower reaches of
adjacent pine/oak communities. It has not been
found to grow sympatrically with any of the other
species of the complex. Consistent with the other
members of the complex, its blades are nearly al¬
ways flat or only loosely folded in the natural state
(this inay be obscure in dried specimens), though
they do not coil like wood shavings. The flat blades
contrast with the tightly involute blades of sympat-
ric A. pansa Wooton & Standley and A. purpurea
Nuttall s.l. Although the panicles of A. eludens usu¬
ally lack pulvini and are thus narrow and contract¬
ed, it is not uncommon to find an isolated plant or

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

213

Figure 3. Aristida eludens Allred & Valdfe-Reyna (Valdes-Reyna 2254). -A. Habit. -B. Spikelet: glumes left, floret
right.

214

Novon

two in each population with only the lowermost
branch spreading from a single axillary pulvinus.
Only one individual from northern Coahuila (Ibarra
s.n.) has been found to be completely pulvinate
(pulvini in all axils).

Paratypes. MEXICO. Chihuahua: Sierra Carrasco,
ca. 31 mi. NW of Julimes, 6000 ft., 15 Sep, 1973, J.
Henrickson 12965.5 (TEX/LL); Santa Eulalia Mts., 10 Sep.
1885, C. G. Pringle 386 (TEX/LL); La Campana, 1650 m,
7 Sep. 1973, J. ValcUs-Reyna 148 (TAES). Coahuila: Si¬
erra de la Paila (lado norte), Canada becerros, 1700 m, 13
Oct. 1989, J. A. Villarreal 5469 (ANSM); 12 km al W de
Saltillo, carr. 40, 2000 m, 9 July 1983, ,/. Valdds-Reyna
& /.. Rodriguez 1522 (ANSM); San Lorenzo Canyon, ca. 8
km S of Saltillo, mountain scmb grassland vegetation with
Berber! s, Bouteloua , Hilaria, Dasylirion, very rocky sandy
loam, flat bajada on N-facing side of canyon, anthers 1.5
mm, 6000 ft. (1829 m), 3 Sep. 1991, K. W. Allred & J.
Valdds-Reyna 5488, 5491, 5492, 5495, 5497 (ANSM,
NMCR); Est. Cameros, Cameros Pass, steep hills W of
town, very rocky calcareous soil along cobble road to mi¬
crowave tower, desert grassland/scrub vegetation with Yuc¬
ca carnerosana. Agave, Nassella, Aristida curvifolia, 6750
ft. (2057 m), 5 Sep. 1991, K. W. Allred & J. Valdds-Reyna
5527, 5528 (ANSM, NMCR); Sierra Garcia, San Lazaro
Pass, desert mountain scrub vegetation with ocotillo, le-
chuguilla. Acacia, and Opuntia, limestone N-facing steep
slopes, 1200 ft. (366 m), 6 Sep. 1991, K. W. Allred & J.
Valdds-Reyna 5540 (ANSM, NMCR); San Lorenzo Canyon,
25 Aug. 1981, S. L. Hatch 4501 (ANSM, TAES); Cameros,
camino a torre de microondas, 3 km al poniente de la
estacidn, 30 km al S de Saltillo, matorral de Yucca car¬
nerosana, Pinus cembroides, Dasylirion cedrosanum, Cea-
nothus greggii, Nassella tenuissima, 17 Oct. 1986, J. Es-
pinosa-A. 97, 186 (ANSM, NMCR); San Lorenzo Canyon,
22 Aug. 1980, M. Madrigal-A. s.n. (TAES); Sierra de Par¬
ras en Los Chupaderos, 14 Apr. 1981, A. Rodriguez s.n.
(CIID1R-DURANGO); Estacidn Cameros, camino torre de
microondas, 2100 m, 24 May 1982, J. Valdds-Reyna 1456
(TAES, US); 18 mi. S of Saltillo on hwy. 54, just W of Est.
Cameros, 2250 m, 18 Oct. 1989, P. Peterson 8416
(ANSM, US); Canon San Lorenzo, 1900 m, 22 Aug. 1980,
[no collector] (TAES); Mpio. Arteaga, 10 mi. NE of hwy!
57 at Los Alpes, with pinyon. Yucca, Agave, 24 Aug. 1983
S. L Hatch 5022 (ANSM, NMCR, TAES); Mpio. Saltillo’,
Cerro del Pueblo, al W de la ciudad de Saltillo, 11 Sep.
1990, J. Valdds-Reyna 2055 (ANSM, NMCR); Mpio.
Ocampo, Rancho La Rueda, D. Ibarra s.n. (ANSM). Du¬
rango: 45 km WNW of Huejuquilla El Alto, 1830-2150
m, 23 Oct. 1983, D. E. Breedlove 59212 (CAS, MO). Gua¬
najuato: 30 km SE of San Felipe on hwy. 37 to Ledn,
rocky slopes in open grassland, 2290 m, 30 Aug. 1990
P. M. Peterson 9693 (ANSM, NMCR, US); oak slopes ca!
12 mi. on rd. between Guanajuato and Santa Rosa 30
Sep. 1946, H. E. Moore, Jr. 1351 (US). Oaxaca: 19.5 km
NW of Tlaxiaco and 12.3 km NE of San Juan Mixtepec,
near jet. of road to Yosonama, sandy slopes with pine, oak
and manzanita, 2410 m, 3 Sep. 1990, P. M. Peterson 9736
(ANSM, NMCR, US). Nuevo Leon: near Puerto Mexico
along hwy. 57, 22 Aug. 1983, S. L. Hatch 5011 (ARIZ,
TAES). Queretaro: 5 km al NE de Bernal, sobre la car-
retera a Tolim4n, ladera caliza con vegetacidn de matorral
xerftfilo, 1950 m, 10 Nov. 1988, Rzedouiski s.n. (ANSM,
NMCR). San Luis Potosi: Mpio. Villa de Arriaga, Cerro
del Gallo, 2500 m, 3 Oct. 1962, E. Hemdandez-X. 1413

(ANSM); mountain slopes, 13 mi. S of San Luis Potost,
7100 ft., 2 Sep. 1939, F. Shreve 9307 (US); near village
of San Francisco in Sierra de San Miguelito, ca. 25 km
SW of San Luis Potosf, pine/oak, 22—2400 m, 5 Sep.
1954, E. R. Sohrts 1062 (US); Cnyn. del Muerto, ca. 3 km
W of Rodrigo, in Sierra de San Miguelito, 1800-2200 m,
18 Sep. 1954, E. R. Sohns 1317 (US); ca. 5 mi. SW of
city in hills, 18 July 1950,,/. R. Reeder 1372 (ARIZ).

Several specimens from central Mexico approach
Aristida eludens in having single awns and narrow
panicles, but differ in being very tall (to 1.5 m)
robust plants with knotty bases and curling blades,
and in occurring in oak and pine/oak forests. They
are here excluded from A. eludens and are tenta¬
tively considered aberrant forms of plants normally
with spreading panicle branches. Duplicates of
some of these were referred by McVaugh (1983) to
A. schiedeana or A. laxa.

Excluded from A. eludens: MEXICO. Jalisco: Autldn,
1290 m, 5/XI/1975, C. Castro 84 (MEXU); near Villa
Guerrero, Cerro del Aquila, 25 Sep. 1980, A. Beetle 5811
(MEXU); Paso de la Troje, near km 36, SW of Ojuelos on
rd. to Aguascalientes, 2100—2300 m, 9-12 Aug. 1958, R.
McVaugh 16818 (US); Rio Blanco, 6 Oct. 1886, E. Palmer
476 (MO); Rio Blanco, June—Oct. 1886, /'.. Palmer 517.5
(US); Rio Blanco, June-Oct. 1886, E. Palmer 769 (US);
Mpio. Mezquitic, 5 km E of Rancho El Mortero, pine/oak,
2200 m, 5 Nov. 1963, Rzedouiski 17687 (US). Morelos:
Alarc an. 31 Aug. 1910, C. Orcutt 3863 (MO). Puebla:
35 km de Tetela, 1550 m, 1/VI11/1981, S. Contreras 534
(MEXU).

Aristida spanospicula Allred, Valdes-Reyna &
Sdnchez-Ken, sp. nov. TYPE: Mexico. Chihua¬
hua: Sierra Madre Occidental, at Cuesta Prie-
ta, along road from San Juanito to Creel, 3.1
mi. S of San Juanito, pine forest with scrub
oak understory of Lupinus, Muhlenbergia, and
Trisetum, thin forest loam calcareous soil, nu¬
merous clumps among the scrub oak, 7500 ft.
(2280 m), 22 Sep. 1992, K. W. Allred & J-
Valdds-Reyna 5787 (holotype, NMCR; iso¬
types, ANSM, NMC, TAES). Figure 4.

Species nova Aristidae schiedeanae affinis laminis plan¬
ts saepe lorsivis scobiformibus, paniculis angustis absque
pulvinis axillaribus, spiculis paucis dispersis late, glumis
magnis spadiceis, rostris exsertis tortis lemmatum, et ar-
istis singularibus a congeneribus Americanae borealis div-
ersa.

Plants perennial, tufted; culms (20-)25-55(-60)
cm tall, ascending, slender, not branching above
the base; internodes terete, glabrous to minutely
scaberulous. Leaves basal, light green to yellowish
green. Sheaths longer than the intemodes, glabrous
to sparsely pilose, rounded on the back; collars gla-
brous to sparsely pilose at the comers, lacking a
line of hairs across the back; throats glabrous ex-

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

215

Figure 4. Aristida spanospicula Allred & Valdfc-Reyna (AUred 5787). -A. Habit. -B. Spikelet: glumes left, floret
right.

216

Novon

cept for the ligule. Ligules a fringe of hairs less than
0.5 mm long. Blades flat to loosely folded when
fresh, rolled upon drying, usually coiling like wood
shavings in age, glabrous abaxially, glabrous to pu-
berulent adaxially, 14-25 cm long, 1.5-2 mm wide
when flat, the margins thickened by the coales¬
cence of the outer veins. Panicles (8-)12-22 cm
long, 1—2 cm wide, few flowered with 3—8(—12)
spikelets, often drooping, narrow but loose, the
nodes glabrous; primary branches weakly devel¬
oped, appressed-erect, bearing 1—2(-4) spikelets;
pulvini absent. Spikelets few and widely spaced,
scarcely overlapping, the pedicels often drooping
and somewhat capillary. Glumes glabrous to sca-
berulous, mostly 1-nerved, occasionally a short lat¬
eral nerve developed, brownish, subequal or the
first 1-2 mm shorter or longer, 9—15 mm long. Lem¬
mas glabrous, (16—)18—22(—25) mm long from the
base of the callus to the divergence of the awns;
beak twisted, exserted beyond the glumes, (6—)8—
10(—15) mm long; central awn not twisted, genic¬
ulate-bent at ca. 90°, 10-16(—20) mm long; lateral
awns highly reduced, erect, less than 0.5 mm long;
callus 0.5-1 mm long with stiff hairs less than 1
mm long. Paleas completely enclosed by the lem¬
mas, 2-nerved, membranous, ca. 1 mm long, shorter
than the lodicules. Stamens 3, the anthers light
brown, ca. 2 mm long. Lodicules 2, flabellate, 1-
1.5 mm long. Caryopses fusiform, brown, 7—9 mm
long.

Flowering August-October. Distribution. As yet,
found only in pine/oak forests of the Sierra Madre
Occidentale, often on nearly bare ground, usually
rocky or crumbly calcareous soil; Mexico, in the
states of Chihuahua, Durango, and Sonora (Fig. 2).

We are pleased to cooperate with Jorge SAnchez-
Ken, of the Herbario Nacional de Mexico (MEXU),
in describing this new species. The epithet spano-
spicula alludes to the panicles with few spikelets.

Aristida spanospicula differs from others in the
complex by the epulvinate panicle, fewer spikelets,
and longer lemmas. It is apparently restricted to
the pine/oak forests of the Sierra Madre Occidental
in northern Mexico. Aristida spanospicula occasion¬
ally grows intermingled with A. schiedeana, and
some of the sympatric populations contain inter¬
mediate plants suggesting hybridization or incom¬
plete morphological delineation of the species. The
latter would not be uncommon in Aristida.

Paratypes. MEXICO. Chihuahua: ca. 6 mi. S of
Creel, rocky outcrops near small stream, in ponderosa
pine community with 5-needle pine and juniper, sandy
loam soil, 22 Aug. 1985, K. W. Allred 3074 (NMCR); Cas-
cada Basaseachic Overlook on S side of canyon, very steep
N-facing slopes in oak/pine forest, with Quercus crassifol-

ia, Arbutus, 6600 ft. (2012 m), 11 Sep. 1987, K. W. Allred
4607 (NMCR); 4 mi. W of Cuesta Blanca in the Sierra
Brena, approximately 32 road mi. SW of Col. Juarez, pine-
oak woodland with a very sparse understory, rocky gravelly
soil, 6500 ft. (1981 m), 21 Sep. 1992, K. W. Allred & J.
Valdts-Reyna 5745, 5746 (NMCR); Sierra Madre Occi¬
dental, 11 mi. W of Babicora on hwy. 180 to Madera, pine-
oak forest with sparse grass understory of Bouteloua, Muh-
lenbergia, Piptochaetium, rocky loam soil, 6800 ft. (2072
m), 22 Sep. 1992, K. W. Allred & J. Valdes-Reyna 5776
(ANSM, NMCR); Sierra Madre Occidental, at Cuesta Prie-
ta, along road from San Juanito to Creel, 3.1 mi. S of San
Juanito, pine forest with scrub oak understory of Lupinus,
Muhlenbergia, and Trisetum, thin forest loam calcareous
soil, approaching A. schiedeana , 7500 ft. (2280 m), 22
Sep. 1992, K. W. Allred & J. Valdes-Reyna 5792 (ANSM,
NMCR); Cusarare, S of Creel, 14 Sep. 1973, Bye, R. 5047
(TEX/LL); S&nchez, rocky pine woods, 8000 ft., 12 Oct.
1910, A. S. Hitchcock 7671 (US); Chuhuichupa, Aug.-Sep.
1936, Lesueur 6112 (CAS, MO, TEX/LL); Colonia Cum-
bres de Majalca, approx. 32 km W of hwy. 45 N of Chi¬
huahua, table rock and rocky slopes with oak, pine, ju¬
niper, and Arctostaphylos, 23 Sep. 1988, P. M. Peterson
5810 (ANSM, NMCR, US); 43.5 km W of Balleza and
51.6 km E of Guachochi, sandy, clay flats with pine, oak,
and Arctostaphylos pungens, 2320 m, 18 Sep. 1991, P. M.
Peterson et al. 10756 (ANSM, NMCR, US); Parque Na¬
cional Barranca del Cobre, 24.8 km NE of La Bufa on
road to Samachic, gentle slopes in forest of pine, oak, and
Arbutus, 2440 m, 20 Sep. 1991, P. M. Peterson et al.
10799 (ANSM, NMCR, US); pine-oak region, Sierra Ma¬
dre occidental, W of Casas Grandes, 3 mi. W of Cuesta
Blanca, 2073 m, 4 Sep. 1958, J. R. & C. G. Reeder 3214
(ARIZ, US). Durango: Sierra Madre Occidental, moun¬
tains E of Durango City, approximately 30 mi. E of the
city at the movie set for “Fat Man and Little Boy,” deep
barrancas in pine/oak/manzanita communities, soil chalky,
calcareous, very crumbly, 7500 ft. (2286 m), 24 Sep.
1992, K. W. Allred & J. Valdes-Reyna 5831, 5837 (ANSM,
NMCR); Sierra Madre Occidental, along hwy. 40 from Du¬
rango City to El Salto, 30 mi. W of Durango, pine/oak/
arctostaphylos woodland with thick grass understory of
Muhlenbergia, Trachypogon, slight slope, loam soil, 8400
ft. (2560 m), 24 Sep. 1992, K. W. Allred & J. Valdes- Reyna
5845 (ANSM, NMCR); Sierra Madre along hwy. 40 from
Durango City to El Salto, 32 mi. W of Durango, adjacent
to Parque El Tecuan, pine savannah parkland with 5-nee¬
dle pine, gently W-facing slope, grass understory of Aris¬
tida, Panicum, forest loam soil, 8000 ft. (2438 m), 24 Sep.
1992, K. W. Allred & J. Valdts-Reyna 5851 (ANSM,
NMCR). Sonora: E of Cananea, pine woods. Sierra de los
Ajos, 26 Sep. 1981, A. A. Beetle 7859 (MO).

The relationship of A. schiedeana and A. orcut-
tiana has long been problematical. D4vila & S£n-
chez-Ken (1994), Gould & Moran (1981), and Pohl
& Davidse (1994) treated the two as conspecific
without distinction; Henrard (1927, 1928, 1929)
treated the two as distinct species; Hitchcock treat¬
ed them both ways (conspecific in 1924, distinct in
1935); and Beetle (1983) presumably treated the
two as conspecific, though he does not mention A.
orcuttiana in the text at all, even though the type
is from Baja California Norte. Table 1 presents the
salient differences between the two taxa. Although

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

217

the morphological distinctions are incomplete and
difficult to interpret, they are separated geograph¬
ically (Fig. 5). Aristida orcuttiana is here relegated
to varietal status under A. schiedeana:

Aristida schiedeana Trinius & Ruprecht var. or¬
cuttiana (Vasey) Allred & Valdes-Reyna,
comb. nov. Basionym: Aristida orcuttiana Va¬
sey, Bull. Torrey Bot. Club 13: 27. 1886. SYN-
TYPES: Mexico. Baja California Norte: Han¬
sen’s Ranch, 6000 ft., 30 July 1883, C. R.
Orcutt 507 (lectotype, selected here, US).
U.S.A. Arizona: M. E. Jones (not seen).

Key to the Members of the Aristida schiedeana Com¬
plex:

la. Panicles open, all the primary branches with ax¬
illary pulvini and spreading sharply outward
from the main axis upon exsertion of the panicle
from the sheath and throughout maturity.

2a. Lemmas with three well-developed awns, the
lateral awns at least V 4 the length of the cen¬
tral awn. A. laxa Cavanilles

(We use the name A. laxa Cavanilles in its
traditional sense, although S4nchez-Ken
(pers. comm.) informs us that in the type of
A. laxa , the lateral awns are nearly absent;
there may be difficulties with the application
of this name.)

2b. Lemmas with a single well-developed awn,
the lateral awns minute or at most V 4 the

length of the central awn.

. A. schiedeana Trinius & Ruprecht

3a. First glume usually equal to or shorter
than the second; glumes usually sca¬
brous; blade above ligule usually with
scattered pilose hairs; collar usually
with a line of short hairs . .var. schiedeana
3b. First glume usually longer than the sec¬
ond; glumes usually glabrous; blade
above ligule usually glabrous; collar
usually lacking a line of hairs ....

.var. orcuttiana (Vasey)

Allred & Vald6s-Reyna

lb. Panicles closed, contracted, none of the primary
branches (or occasionally only the lowermost)
with axillary pulvini and thus not spreading from
the axis, though the branch tips or spikelets may
droop outward.

4a. Lemmas (16-)18-22(-25) mm long (from the
base of the callus to the divergence of the
awn); panicles few flowered, with 3—12

loosely disposed spikelets.

. A. spanospicula Allred & Valdes-Reyna

4b. Lemmas less than 10-13 mm long (from the
base of the callus to the divergence of the
awn); panicles densely flowered, with 16 or

more congested spikelets.

. A. eludens Allred & Valdes-Reyna

Aristida purpurea Nuttall Complex

The Aristida purpurea complex has tightly invo¬
lute to loosely folded blades, panicles without ax-

Table 1. Comparison of Aristida schiedeana Trinius &
Ruprecht and A. orcuttiana Vasey.

A. schiedeana

A. orcuttiana

Pilose hairs above

Usually present

Usually absent

ligule

Collar hairs

Usually present
in a line

Usually absent

Glume pubescence

Usually scabrous

Usually glabrous

Glume 1 length

6—15 usually

6-14 usually

(mm)

equal to

longer than

glume 2

glume 2

Glume 2 length

8-14

5—10(—13)

(mm)

Central awn length

(5—)7—15

5-14

(mm)

Lateral awn length

0.3-1.5(-4)

0-3(-7)

(mm)

Distribution (Fig.

North-central

Southwestern

5)

Mexico to

United States

Nicaragua

to northern
Mexico

illary pulvini (except in two varieties), and strongly
unequal glumes, the first much shorter than the
second. Its members are common in arid and semi-
arid plains and deserts, and include A. purpurea
s.l. (Allred, 1984), A. brownii Wamock, and A. cur-
vifolia Fournier.

Aristida purpurea Nuttall var. perplexa Allred
& Valdes-Reyna, var. nov. TYPE: U.S.A. New
Mexico: Dona Ana County, USDA Jornada Ex¬
perimental Range, ca. 16 mi. NE of Las Cru¬
ces, pasture 12, ca. 4800 ft., sandy soil, very
common, 15 July 1986, K. W. Allred 4035 (ho-
lotype, NMCR). Figure 6.

Varietas nova Aristidae pansae et A. purpureae similis;
ab A. pansa arachnoideis destitutis angulis vaginarum,
glumis inaequalibus (primis secundis brevioribus), et ar-
istis longis ascendentibus differt; ab A. purpurea paniculis
apertis habentibus pulvinos axillares differt.

Plants perennial, tufted; culms (30-)40—65 cm
tall, erect, simple or sparsely branched at the base;
intemodes terete, glabrous to puberulent. Leaves
basal and cauline, yellowish to pale green. Sheaths
rounded on the back, glabrous to puberulent, with
longer hairs at the summit; collars usually with a
line of short stiff hairs across the back, the comers
glabrous or with a tuft of erect, straight to somewhat
crinkly hairs (but not cobwebby) ca. 1 mm long;
throats pubemlent in addition to the ligule. Ligules
a fringe of hairs ca. 0.5 mm long. Blades rolled
when fresh, sometimes only loosely folded, usually
glabrous abaxially and pubemlent adaxially, mostly

218

Novon

8-20 cm long and less than 1 mm wide, arcuate to
curling upon drying. Panicles (8-)10-26 cm long
and (4-)6-12 cm wide, open, pyramidal; primary
branches stiffly ascending to widely spreading from
axillary pulvini, sometimes somewhat flexuous or
capillary, naked at the base, the lower ones 3—6
(-8) cm long; pulvini present in the axils of the
primary branches but usually absent from the ped¬
icels, the spikelets thus appressed along the
branches. Glumes glabrous, membranous to scari-
ous, light tan, strongly unequal and the first Vi to
% the length of the second, 1-nerved, the first (4.5-)
5-7(-7.5) mm long, the second 8—11(—12) mm long.
Lemmas glabrous to scabrous, often mottled, (8-)
10-12(-13) mm long from the base of the callus to
the divergence of the awns; beak usually not twist¬
ed, reaching about to the tip of the second glume
or slightly beyond, 2.5-3.5(-4) mm long and 0.1-
0.2 mm wide; awns subequal, mostly spreading 40-
50°, rarely approaching 90° from the vertical, the
central awn (1.5-)2-3 cm long, the lateral awns 2-
4 mm shorter; callus 0.5-1 mm long with short stiff
hairs ca. 0.5 mm long. Paleas completely enclosed

by the lemmas, membranous, acute, 2-nerved, 1—
1.3 mm long. Stamens 3, the anthers 0.7—1.5 mm
long. Lodicules 2, flabellate, 1-1.4 mm long. Car-
yopses fusiform, light brownish.

Flowering July-October. Distribution. Sandy to
rocky plains and mesas in desert grassland/scrub
communities, often calcareous soil; Mexico, in the
states of Chihuahua, Coahuila, and San Luis Potosf;
United States, in the states of Arizona, New Mexico,
and Texas (Fig. 7).

The epithet perplexa refers to the prior confusion
of this variety with Aristida pansa.

Because of its spreading panicle branches, spec¬
imens of variety perplexa have long been confused
with Aristida pansa Wooton & Standley. The new
variety differs from this latter species by its lack of
cobwebby hairs at the collar (a feature consistent
throughout the A. pansa complex), by the acuminate
unequal glumes (the first %-% the length of the
second), and by the longer awns (mostly 2—3 cm
vs. 1.5 cm or less), which generally ascend at an
angle of 40-50° rather than being nearly horizontal.

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

219

Figure 6. Aristida purpurea Nut,all var. perplexa Allred & Valdfe-Reyna (Peterson 10632). -A. Habit. -B. Spikele,
glumes left, floret right.

220

Novon

The spikelets of variety perplexa are essentially
identical to spikelets of A. purpurea var. purpurea.
Variety perplexa may grow in homogeneous isolated
populations (the type locality) or be sympatric with
A. pansa and other varieties of A. purpurea, as is
the case at the type locality of A. pansa in Dona
Ana County, New Mexico. Variety perplexa is the
second variety in the A. purpurea complex with
spreading, pulvinate panicle branches. The other,
variety parishii (A. S. Hitchcock) Allred, which is
restricted to Sonoran Desert habitats, is similar to
variety wrightii (Nash) Allred and often lacks the
axillary pulvini so that the panicle is narrow or spi-
cate. The panicles of variety perplexa, however, are
consistently pulvinate.

Paratypes. MEXICO. Chihuahua: just S of New Mex¬
ico border near Columbus, rocky hills, 15 Oct. 1983, K.
W. Allred 2629 (NMCR); Sierra Las Pampas, N end of the
sierra along dirt road from Las Pampas to jet. with road
to Camargo, 6.2 mi. SE of jet., Chihuahuan Desert Scrub,
rocky calcareous slope, 4500 ft. (1372 m), 23 Sep. 1992,
K. W. Allred & J. Valdes-Reyna 5806 , 5807 (NMCR); near
Chihuahua, 13 Oct. 1885, C. G. Pringle s.n. (POM/RSA).
Coahuila: 16.9 km NE of San Miguel on road towards

Boquillas, 1180 m, 13 Sep. 1991, P. M. Peterson 10608
(ANSM, NMCR, US); Sierra El Pino, 9.2 km SW of Ran¬
cho El Cimarron along the eastern slope, calcareous rocky
slopes, 1500 m, 14 Sep. 1991, P. M. Peterson 10632
(ANSM, NMCR, US); near La Rosa, 5000 ft., 18 Aug.
1940, F. Shreve & E. R. Tinhham 4593 (ARIZ). San Luis
Potosi: along hwy. 49 from Zacatecas to San Luis Potost,
32.4 mi. E of Salina, 21 mi. W of San Luis Potosi, desert
thorn scrub community, rocky soil, 6200 ft. (1890 m), 26
Sep. 1992, K. W. Allred & J. Valclts-Reyna 5873 , 5874,
5875 (NMCR); along hwy. 57 at jet. to San Jose del Re¬
fugio, gypsum, 1340 m, 20 Oct. 1978, J. R. Reeder 7052
(ARIZ). U.S.A. Arizona: Cochise Co., along Cascabel to
Willcox road, near 8 mi. marker, rocky limestone slope
with ocotillo, Agave and grasses, 1450 m, 25 Aug. 1992,
J. R. & C. G. Reeder 8878 (ARIZ, NMCR); 1 mi. NW of
Naco (Waco?), 6 Oct. 1936, L A. Goodding 389 (ARIZ);
Apache Pass between Dos Cabezas and Chiricahua Mts.,
Siphon Cnyn., 1450 m, 23 Aug. 1993, J. R. Reeder 9020
(ARIZ). Pima Co., Santa Rita Mts., hwy. jet. E of Vail,
3000 ft., 7 Sep. 1938, L. Benson 9121 (ARIZ). New Mex¬
ico: Bernalillo Co., 3 mi. W of Albuquerque, 9 Oct. 1945,
R. A. Darrow 3367 (ARIZ). Chaves Co., Hwy. 380 ca. 1^
mi. W of Caprock, 8 July 1982, K. W. Allred 231
(NMCR); 3.5 mi. E of Elkins on hwy. 70, sandy, 03 "
Artemisia-grassland, 26 Aug. 1974, L C. Higgins 917
(NMC). Dona Ana Co., Ropes Spring, San Andres Mts^,
17 Aug. 1933, [collector unknown] 89 (NMCR); 7 mi-

Volume 5, Number 3
1995

Allred & Valdes-Reyna
North American Aristida

221

of Hatch at jet. of hwy. 26 and 85, 2 Oct. 1982, T. Potter
17 (NMCR); Las Cruces, 6 Oct. 1904, D. Griffiths 7417
(NMCR); near Tortugas Mt., 24 Oct. 1904, E. 0. Wooton
s.n. (NMC); White Sands Missile Range, 5 km E of head¬
quarters, 200 m S of Nike Ave., granitic sandy bajada, 23
Sep. 1990, R. Spellenberg 10678 (NMC). Eddy Co., abt.
40 mi. E of Roswell, 7 mi. N of hwy. 380, Mather Natural
Area, sandy, shin oak, 3 Aug. 1979, R. Spellenberg 5249
(NMC). Guadalupe Co., 2 mi. S of Vaughn, 15 July 1979,
L Rockhill 20 (NMCR). Luna Co., Cooke’s Range, Prov-
inger Draw, 12 Aug. 1986, ]. T. Columbus 419 (NMCR);
5 mi. W of Columbus, 9 July 1984, J. S. Trent 107
(NMCR). Socorro Co., ca. 12 mi. E of San Antonio along
hwy. 380, 5 July 1984, K. W. Allred 2700 (NMCR); road¬
side E of Belen on hwy. 60, 7 Apr. 1952, E. F. Costetter
7962 (NMC). Valencia Co., 20 mi. NE of Suwanee, 1 Sep.
1935, K. W. Parker 633 (NMCR). Texas: Culberson Co.,
deep sand 21 mi. W of Kent, 4000 ft., 17 Aug. 1950, B.
H. Wamock 9317 (SR); dunes near Salt Flats, 4000 ft., 29
Aug. 1965, B. H. Warnock 20678 (SR); at Salt Flat Station,
E of Salt Flat, 4000 ft., 29 Aug. 1965, B. H. Warnock
20674 (SR). Pecos Co., limestone hills 20—35 mi. S of Ft.
Stockton, along Sanderson hwy., 3100 ft., 1 July 1955, B.
H. Warnock 13266 (SR).

Aristida brownii Wamock (Wamock, 1982) was
described to accommodate single-awned Aristida
(the lateral awns highly reduced) otherwise identi¬
cal to A. purpurea var. wrightii. Since its descrip¬
tion from Brewster County, Texas, individuals of
this entity have been found from other Texas lo¬
calities, several locations in southern New Mexico,
and one locality in Coahuila, Mexico. Plants of A.
brownii are always found intermingled with variety
wrightii, usually in sparse numbers, and never as
isolated homogeneous populations. In addition, it is
not uncommon to find in the same population with
typical A. brownii plants with lateral awns 14—% the
length of the central awn, an attribute intermediate
between A. brownii and the sympatric variety
wrightii. We find the recognition of this taxon un¬
tenable at the species level. Because the single-
awned feature is unique in A. purpurea s.l. and eas¬
ily confused with other taxa, we propose its
recognition at the level of forma. We have placed
the name in a 3-level hierarchy to indicate its af¬
finities within A. purpurea, i.e., A. purpurea var.
wrightii forma brownii.

Aristida purpurea Nuttall forma brownii (War¬
nock) Allred & Valdes-Reyna, comb. nov. Bas-
ionym; Aristida brownii Wamock, Sida 9: 358.
1982. TYPE: U.S.A. Texas: Brewster County,
widespread perennial on limestone hills of Del
Norte Mountains, elevation 4600 ft. or more,
27 June 1981, Warnock 141 (holotype, Lajitas
Museum, not seen; isotypes, SMU, TEX).

Allred (1984) reduced Aristida curvifolia Four¬
nier to a variety of A. purpurea because its strongly

unequal glumes are also characteristic of the other
varieties of A. purpurea. Further field studies during
the subsequent 10 years have shown that disposi¬
tion to be unwarranted, and we propose the rein¬
statement of this taxon at the species rank, as did
Divila & Sanchez-Ken (1994). Aristida curvifolia is
consistently distinguished from all other desert Ar¬
istida by the following features: blades stiff, tightly
involute, yellow-green, the epidermis nearly smooth
and the veins hardly noticeable; glumes broad,
blunt, unequal; lemma beak (apex) 1-2 mm long,
0.2-0.5 mm wide, straight to only slightly twisted
even when mature; panicles spicate, always epul-
vinate. Aristida curvifolia often grows with both A.
purpurea and A. partsa, and no evidence of hybrid¬
ization or intermediacy has been noted. Indeed, this
taxon turns out to be one of the more uniform spe¬
cies of Aristida in North America. It is presently
known from 12 states in Mexico: Aguascalientes,
Baja California Norte, Chihuahua, Coahuila, Gua¬
najuato, Nuevo Le6n, Oaxaca, Puebla, San Luis Po-
tosf, Tamaulipas, Veracruz, and Zacatecas.

Acknowledgments. Portions of this research were
supported by the New Mexico State University Ag¬
ricultural Experiment Station. We thank Paul Pe¬
terson for sharing his collections, Jorge Sdnchez-
Ken for his cooperation and hospitality, Richard
Spellenberg for allowing us use of his maps, Miguel
A. Carranza-P. for help in fieldwork, Cathy Montes
of the Department of Agricultural Communications
(New Mexico State University) for digitizing and
printing the maps, anonymous reviewers for their
helpful comments, and the staffs of the following
institutions for the generous use of their specimens
and facilities: AR1Z, ASU, CHAPA, COCA, ENCB,
JEPS, MEXU, MO, POM/RSA, TAES, TEX/LL,
UC, US. The illustrations were rendered by Cuauht¬
emoc Gonzfilez de Le6n.

Literature Cited

Allred, K. W. 1984. Morphologic variation and classifi¬
cation of the North American Aristida purpurea complex
(Gramineae). Brittonia 36: 382—395.

Beetle, A. A. 1983. Las Gramfneas de Mexico. Secretaria
de Agricultura y Recursos Hidriulicos, Mexico City.
D4vila, P. & J. SSnchez-Ken. 1994. Flora del Valle de
Tehuacta-Cuicatlan. Fasc. 3: 7-22. Inst. Biol. Univ.
Nac. Aut. Mexico, Mexico D.F.

Gould, F. W. & R. Moran. 1981. The Grasses of Baja
California, Mexico. San Diego Society of Natural His¬
tory, Memoir 12.

Henrard, J. T. 1927. A critical revision of the genus Ar¬
istida. Meded. Rijks-Herb. 54: 221-464.

-. 1928. A critical revision of the genus Aristida.

Meded. Rijks-Herb. 54: 465-701.

-. 1929. A monograph of the genus Aristida. Med¬
ed. Rijks-Herb. 58: 1-153.

222

Novon

Hitchcock, A. S. 1924. The North American species of
Aristida. Contr. U.S. Natl. Herb. 22: 517-586.

-. 1935. Aristida. North American Flora 17: 376—

406.

McVaugh, R. 1983. Flora Novo-Galiciana, Vol. 14, Gra-
mineae. Univ. Michigan Press, Ann Arbor.

Pohl, R. W. & G. Davidse. 1994. Aristida. Pp. 253-257

in: G. Davidse, M. Sousa S. & A. O. Chater (editors),
Flora Mesoamericana, Vol. 6. Alismataceae a Cypera-
ceae. Universidad Nacional Aut6noma de Mexico, Mex¬
ico D.F.; Missouri Botanical Garden, St. Louis; The Nat¬
ural History Museum, London.

Wamock, B. H. 1982. A new three-awn grass from trans-
Pecos Texas. Sida 9: 358-359.

Two New Species of the Genus Sterigmapetalum (Rhizophoraceae)
from the Venezuelan and Brazilian Amazonian Region

Gerardo A. Aymard C. and Nidia Cuello A.

UNELLEZ-Guanare, Programa de R.N.R. Herbario Universitario (PORT),

Mesa de Cavacas, Edo. Portuguesa, Venezuela 3323

Abstract. Two new species of Sterigmapetalum
subg. Sterigmapetalum are described from Vene¬
zuela and Brazil: 5. chrysophyllum and S. plum-
beum. Relationships with related species are dis¬
cussed, and an illustration for each species is
provided.

Resumen. Se describen dos nuevas especies de
Sterigmapetalum subgenero Sterigmapetalum de
Venezuela y Brasil: S. chrysophyllum y .S', plum-
beum. Se discuten sus relaciones con las especies
afines y se presentan ilustraciones de cada una de
ellas.

Sterigmapetalum Kuhlmann is a small neotropi¬
cal genus with nine species in the Rhizophoraceae.
Sterigmapetalum and Cassipourea Aublet were seg¬
regated by Steyermark & Liesner (1983) into a new
tribe, the Hypogyneae. Subsequently, Tobe & Ra¬
ven (1987), based on a detailed study of the mor¬
phology of the seed, flower, and fruit, placed these
two genera in the Macarisieae, a treatment origi¬
nally employed by Schimper (1893) a century ear¬
lier.

Eight of the nine species of Sterigmapetalum oc¬
cur in northern South America (Colombia, Vene¬
zuela, Guyana, and Surinam) and one (5. obovatum
Kuhlmann) is restricted to the western and central
Amazonian region of Peru and Brazil (Prance et al.,
1975).

Sterigmapetalum chrysophyllum Aymard &
Cuello, sp. nov. TYPE: Venezuela. Amazonas:
Dpto. Atabapo, Rio Ventuari, bosques en pla¬
teau, 04°14'N, 65°S8'W, 1250 m, Feb. 1992
(fl), Proyecto Inventario de los Recursos Na-
turales de la Guayana Venezolana-CVG-TEC-
MIN, A. Chaviel 411 (holotype, PORT; iso¬
types, MO, VEN). Figure 1.

Species nova quae a S. exappendiculato Steyermark &
Liesner stipulis triangularibus, acutis, 3—5 mm longis; fo-
liis coriaceis, 3—8 cm longis, supra glabris, pagina infema
tomento flavo omnino occulta; petiolo 2-4 mm longo, ju-
ventute dense flavo-tomentoso; petalis extus epapillatis,
fimbriis lateralibus munitis differt.

Trees 15—20 m high; branches 3-verticillate, te¬
rete, covered completely by a yellow-ocher pubes¬
cence when young, glabrous or sparsely adpressed-
pubescent when mature. Stipules triangular,
acuminate, 3—5 mm long, with yellow pubescent
trichomes on both surfaces, internally glandular at
the base. Leaves 3-whorled, the blades coriaceous,
elliptical or oblong-elliptical, 3-8 X 1.5-5.5 cm,
obtuse to truncated at the base, retuse at apex, mar¬
gin subrevolute, crenate, blades glabrous on upper
surface, covered completely by yellow tomentum on
under surface, lateral nerves 8-14 pairs, tertiary
venation more evident on upper surface than on
under surface; petiole 2-A X 1-2 mm, densely yel¬
low-ocher pubescent when young, glabrous or
sparsely adpressed-pubescent when mature. Inflo¬
rescences solitary in the leaf axils, 1-2 cm long; 1
to many flowers, peduncle striate, adpressed-pu¬
bescent with yellow trichomes; pedicels 1 mm long,
adpressed-pubescent with yellow trichomes on both
surfaces; flowers subtended by bractlets, lanceolate,
1-2 mm long, adpressed-pubescent with yellow tri¬
chomes on both sides; staminate flowers with 5-
lobulate calyx, obovate, 3-5 X 2-3 mm (calyx and
lobes), adpressed-pubescent with yellow trichomes
externally, adpressed-pubescent with sericeous tri¬
chomes internally; petals 5, 2-3 mm long, unguic-
ulate, externally adpressed-pubescent with yellow
trichomes from the middle to the apex, glabrous
internally, with three apical appendices and two lat¬
eral ligules, the ligules 1-1.5 mm long, glabrous,
4-5-laciniate; stamens 8-10, united at the base,
filaments glabrous, 1-2 mm long, anthers 1 mm
long, oblong, glabrous; pistilodium rudimentary,
conic, hispid with yellow trichomes; pistillate flow¬
ers not seen. Fruit obovate, 8—12 X 4—7 mm, com¬
pletely covered by an adpressed yellow-ocher in-
dument, seeds 4-5 mm long, oblong, glabrous and
reticulate.

Distribution and common names. Sterigmape¬
talum chrysophyllum is only known from the middle
and upper Ventuari River basin, Atabapo Depart¬
ment, Amazonas State, Venezuela, at 1250 m. It is

Novon 5: 223-226. 1995.

224

Novon

Figure 1. Sterigmapetalum chrysophyllum Aymard & Cuello. —A. Inflorescence with open flowers. —B. Fruiting
branch. —C. Flowering branch. —D. Flower with 1 sepal removed leaving a horizontal scar. —E. Young fruit. Scale:
A, D = 0.5 cm; B, C = 5 cm; E = 1 cm. A, C, and D drawn from holotype (Chaviel 411); B and E drawn from
paratype (Chaviel 406).

Volume 5, Number 3
1995

Aymard C. & Cuello A.
Amazonian Sterigmapetalum

225

Figure 2. Sterigmapetalum plumbeum Aymard & Cuello. —A. Pistillate flower showing the ligules. -B. Flowering
branch. —C. Base of the under surface of the leaf lamina. —D. Pistillate inflorescence. Scale: A, C, and D - 1 cm;
B — 5 cm. All from Daly et al. 5608.

known by the common names of “kasemani” and
“tadiji” (Y’ekuana).

Ecology. Mesothermic humid forests of medium
height (18-25 m), on sandstone surface outcrop¬
pings.

The absence of resinous exudate and the pres¬
ence of 3-whorled leaves and laciniate petal ligules

indicate that Sterigmapetalum chrysophyllum be¬
longs to subgenus Sterigmapetalum (Steyermark &
Liesner, 1983). This new species is related to 5.
exappendiculatum Steyermark & Liesner, but it dif¬
fers in its triangular-acuminate stipules that are 3-
5 mm long (vs. stipules ovate-lanceolate, subob-
tuse, and 7-10 mm long); coriaceous leaves that

226

Novon

are 3-8 cm long, glabrous or sparsely adpressed-
pubescent above and completely covered by yellow
indumentum below (vs. leaves subcoriaceous, 7—14
cm long, sparsely pilose on the midrib above, mod¬
erately to densely adpressed-sericeous below); pet¬
ioles 2-4 mm long, completely covered by a yellow-
ocher pubescence when young, and glabrous or
sparsely adpressed-pubescent when mature; and
petals that are not papillate externally, with two lat¬
eral ligules 4—5-laciniate (vs. petiole 5-11 mm
long, sericeous-tomentose, and petals that are pa¬
pillate externally and without lateral ligules).

Paralype. VENEZUELA. Amazonas: Dpto. Atabapo,
Rto Ventuari, bosques en plateau, 04°14'N, 65°58'W,
1250 m, Feb. 1992 (fl), Proyecto Inventario de los Re-
cursos Naturales de la Guayana Venezolana-CVG-TEC-
MIN, A. Chaviel 406 (MO, PORT, VEN).

Sterigmapetalum plumbeum Aymard & Cuello,
sp. nov. TYPE: Brazil. Amazonas: Mun. Sao
Gabriel da Cachoeira, Upper Rio Negro, near
Comunidade Aparecida, opposite mouth of
Igarape Tuari, 0°20'N, 67°19'W, 6 Nov. 1987
(fl), D. C. Daly, P. J. Maas & J. Ramos 5608
(holotype, MO; isotypes, INPA, NY). Figure 2.

A S. colombiano Monachino stipulis triangularibus, 5-
6 mm longis; foliis coriaceis, oblongis, 8-11 cm longis,
5—7.5 cm latis, utrinque glabris, nervis lateralibus 17-24-
jugatis; inflorescentiis 1.4-2.2 cm longis; floribus 4-5 mm
longis, sepalis intus indumento grisaceo nitido obtectis,
petalis oblanceolatis differt.

Trees 8 m high; branches 3-verticillate, lenticel-
late, sparsely adpressed-pubescent when young,
glabrous when mature. Stipules triangular, 5-6 mm
long, densely adpressed-pubescent on both sur¬
faces, barbate and glandular internally at the base.
Leaves 3-whorled, the blades coriaceous, oblong,
8-11 X 5-7.5 cm, base rounded, apex emarginate,
margins subrevolute, glabrous on both surfaces, lat¬
eral nerves 17—24 on both sides, tertiary venation
very evident on both surfaces, petioles 5-7 X 2-3
mm, sparsely adpressed-pubescent on young
leaves, glabrous on mature leaves. Inflorescences
solitary in the axils, 1.4—2.2 cm long; peduncles
adpressed-pubescent, especially at the base; pedi¬
cels 1-2 mm long, adpressed-pubescent, subtended
by 3 bracts, lanceolate-triangular, the bracts 3 mm
long, sparsely adpressed-pubescent externally,
densely adpressed-pubescent internally; pistillate
flowers with a 5-lobulate calyx, 4—5 mm long, 3-4
mm wide, glabrous to sparsely pilose externally,
densely adpressed-pubescent internally with a sil¬
very gray indumentum; petals 5, white, 5-5.5 mm
long, unguiculate, oblanceolate, externally adpres¬

sed-pubescent from the middle to the apex, inter¬
nally glabrous, with five lateral ligules, the ligules
3 mm long, glabrous, 5-laciniate; staminodes 10,
united at the base, filaments glabrous, 1 mm long;
ovary densely pubescent, style glabrous, 3 mm
long, stigma capitate, staminate flowers and fruits
not seen.

Distribution. Only known in the upper Rio Ne¬
gro region, Municipio San Gabriel de Cachoeira,
Amazonas, Brazil.

Ecology. Upper Amazonian caatinga, on sandy
soil.

Sterigmapetalum plumbeum is related to S. col-
ombianum Monachino (restricted to mesothermic
humid forests in the department of Magdalena,
northwestern Colombia; Barbosa, 1988), but it can
be distinguished from this species by the triangular
stipules, 5-6 mm long, coriaceous leaves, oblong,
8-11 cm long, 5-7.5 cm wide, glabrous on both
sides, lateral nerves 17—24 (vs. ovate stipules, 1.5-
2 mm long, subcoriaceous, obovate leaves, 4.5-6.7
cm long, 2-4 cm wide, glabrous to sparsely pilose
on the main vein and lateral ones below, lateral
nerves 8-12); inflorescences 1.4—2.2 cm long, pis¬
tillate flowers 4—5 mm long, with sepals internally
covered by a silvery gray indumentum and petals
oblanceolate (vs. inflorescences 8-10 mm long, pis¬
tillate flowers 2-3 mm long with sepals internally
sericeous and the petals subobovate).

Acknowledgments. We thank Paul Berry and
Bruce Holst for their collaboration during our stay
at the Missouri Botanical Garden, Roy Gereau (MO)
for the verification of the Latin descriptions, Laur¬
ence Dorr (US) for valuable comments on the
manuscript, and John Myers for the excellent illus¬
trations.

Literature Cited

Barbosa, C. 1988. Rhizophoraceae en el Herbario Na¬
tional Colombiano. Trianea 2: 251—263.

Prance, G. T., M. Freitas da Silva, B. W. Alburquerque,
I. de J. da Silva Araujo, L. M. Medeiros Carreira, M.
M. Nogueira Braga, M. Macedo, P. N. Da ConceicSo, P.
L. Braga Lisboa, P. I. Braga, R. C. Lobato Lisboa & R.
C. Queiroz Vilhena. 1975. RevisSo Taxon6mica das es-
pecies amazfinicas de Rhizophoraceae. Acta Amazonica

5: 5-22.

Schimper, A. F. W. 1893. Rhizophoraceae. In: A. Engler
& K. Prantl (editors). Die Natlirlichen Pflanzenfamilien.
Wilhelm Engelmann, Leipzig. 3(7): 42—56.

Steyermark, J. A. & R. Liesner. 1983. Revision of the
genus Sterigmapetalum (Rhizophoraceae). Ann. Missou¬
ri Bot. Gard. 70: 179-193.

Tobe, H. & P. H. Raven. 1987. The embryology and re¬
lationships of Cassipourea and Sterigmapetalum (Rhi¬
zophoraceae—Macarisieae). Opera Bot. 92: 253-264.

Taxonomic Changes in Ixonanthaceae from the
Venezuelan Guayana

Paul E. Berry

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Nelson Ramirez

Centro de Biologfa Tropical, Universidad Central tie Venezuela, Apartado 20513,
San Martin, Caracas, Venezuela

ABSTRACT. Cyrillopsis micrantha (Steyermark) P.
E. Berry & N. Ramirez is a new combination based
on Ochthocosmus micranthm Steyermark and con¬
stitutes only the second known species of Cyrillop¬
sis (Ixonanthaceae). The number of varieties in
Ochthocosmus roraimae Bentham is reduced from
three to two, and Ochthocosmus parvifolius Hallier
f. is treated as a species of uncertain status, with
no known extant type material.

The family Ixonanthaceae, which is sometimes
treated as a subfamily of the Linaceae, includes two
genera in the Neotropics, Cyrillopsis and Ochtho¬
cosmus. These two genera are distinguished mainly
by fruit and pedicel characters. Ochthocosmus has
a 5-locular ovary, the fruit is a prismatically cylin-
dric, elongate, 5-valved capsule, and the seeds
have a conspicuous, thin, lateral wing. Further¬
more, the pedicels are non-articulated, that is, they
abscise flush with the rachis. In contrast, Cyrillopsis
has a 2-locular ovary, its fruit is an elongate, slight¬
ly asymmetrical, 2-valved capsule, and the seeds
lack wings but are partly covered at the upper end
by a 2-lobed membranaceous aril; its pedicels are
articulated, with an abscission zone and bracteoles
located roughly one-third of the way up the pedicel
from the rachis.

Until now, Cyrillopsis was considered to have a
single species, C. paraensis Kuhlman, and Ochtho¬
cosmus to have eight recognized neotropical species
(Steyermark & Luteyn, 1980; Steyermark, 1984,
1988). Although some botanists, such as Mabberley
(1990), consider Ochthocosmus to include species
both in South America and in Africa, Forman
(1965) presented convincing arguments to place the
African species in the genus Phyllocosmus
Klotzsch, thus leaving Ochthocosmus as a strictly
neotropical group.

While preparing the treatment of the Ixonantha¬
ceae for the Flora of the Venezuelan Guayana, we
examined recently collected fruiting material of

Ochthocosmus micranthus Steyermark, a species
that was described from flowering specimens only
(Steyermark, 1988). The articulated pedicels, aril-
late seeds, and 2-locular fruits of these new collec¬
tions place the species clearly in Cyrillopsis rather
than in Ochthocosmus. The transfer of O. micran¬
thus to Cyrillopsis brings the number of currently
recognized species in Cyrillopsis to two and in
Ochthocosmus to seven. We also reduce the number
of varieties in O. roraimae from three to two by
removing 0. roraimae Bentham var. parvifolius
(Hallier f.) Steyermark & Luteyn as a variety of this
species.

Cyrillopsis micrantha (Steyermark) P. E. Berry &
N. Ramirez, comb. nov. Basionym: Ochthocos¬
mus micranthus Steyermark. Ann. Missouri
Bot. Gard. 75: 318. 1988. TYPE: Venezuela.
Bolivar: Ayavaparu, 10 km SW of Wadacapia-
pue, 5°18'N, 60°58'W, 1100 m, 13 Nov. 1986,
Herndndez 348 (holotype, VEN; isotype, NY).
[Steyermark (1988) was incorrect in citing the
holotype at MO; rather, the holotype is at VEN
and is so labeled, and there is no type material
at MO.] Figure 1.

Steyermark described this species based on
two flowering specimens that strongly resemble
other species of Ochthocosmus from the Guayana
region. Three more recent collections from the
type locality, however, Huber 11958, Huber &
Herndndez 11948, and Herndndez 395, all have
nearly mature fruits, and they conform to the
characteristics of Cyrillopsis described above
(see Fig. 1). Also, the type collection and the
other specimens cited below all have the char¬
acteristic articulated pedicels of Cyrillopsis that
do not occur in Ochthocosmus.

This second species of Cyrillopsis has much
smaller leaves than the previously known species,
C. paraensis Kuhlman. Whereas C. paraensis is pri-

Novon 5: 227-229. 1995.

228

No von

Figure 1. Details of an inflorescence and fruit of Cyrillopsis micrantha (Steyermark) P. E. Berry & N. Ramirez. —A.
Section of inflorescence after most of the flowers have dehisced. Note the articulated pedicels and the bracteole at the
point of articulation. Drawn from Herndndez 348 (VEN). —B. Mature fruit, a somewhat fleshy capsule splitting into
two halves. —C. Seed with arillate structure at the apex. B and C drawn from Huber & Herndndez 11948 (MO).

marily a lowland species distributed from south¬
eastern Venezuela to French Guiana and northern
Brazil, C. micrantha is an upland species endemic
to the southeastern part of the Gran Sabana at
around 1100 m elevation. It occurs in an unusual
shrub savanna (described in Folster, 1992: 37) that
is dominated by a sympatric population of Ochtho-
cosmus roraimae.

Additional specimens examined. VENEZUELA. Boli¬
var: Ayavaparu, 10 km SW of Wadakapiapue, 5°18'N,
60°58'W, 1100 m, 13 Nov. 1986, Herndndez 359 (US,
VEN); Ayavapard, 15 km al W de Wadakapiapud-tepui,
5 16 30'N, 61°00'W, 1110 m, 6 Mar. 1987, Herndndez
395 (herbarium at San Ignacio de Yuruant, Venezuela);
ca. 10 km SW of Wadakapiapuc-tepui. N of the junction
of the Rfo Caraurfn and the Rfo Yuruant, ca. 1100 m,
5 15 N, 60 58 W, 18 Feb. 1987, Huber & Herndndez
11948 (MO, US, VEN); ca. 10 km SW of Wadakapiapud-
tepui, N of the junction of the Rto Caraurfn and the Rfo
Yuruanf, ca. 1030 m, 5°14'N, 60°58'W, 19 Feb 1987
Huber 11958 (MO, MYF, US—2 sheets, VEN).

The label of Herndndez 395 cites “Wontai” as a
common name for this species (in Taurepan, a di¬
alect of the Pemon Amerindian group).

Ochthocosmus roraimae Bentham in Hooker, Lon¬
don J. Bot. 2: 366. 1843. TYPE: Guyana or
Venezuela (“British Guiana” on label), large
shrub 12—16 ft. high, banks of rivers, near Mt.
Roraima, Oct.-Nov. 1842, Richard Schom-
burgk 1037 (holotype, K, photo, MO).

This species is widespread over much of the Ven¬
ezuelan Gran Sabana and extends into western
Guyana and adjacent Roraima state in Brazil. It
occurs between 800 and 1300 m elevation; field
observations by Gabriel Pic6n and Nelson Ramfrez
(pers. comm.) have shown 0. roraimae to be a vari¬
able taxon occurring in a variety of local habitats,
from fairly open shrublands to low or moderately
tall and dense forest.

Steyermark & Luteyn (1980) recognized three va¬
rieties under 0. roraimae: var. roraimae, var. parvi-
folius (Hallier f.) Steyermark & Luteyn, and var.
grandifolius (Steyermark) Steyermark & Luteyn.
Ochthocosmus roraimae var. grandifolius, which is
known only from the type collection, is unique in its
large, stiff, and virtually entire-margined leaves, so
it is maintained here as a distinct variety. However,
Steyermark and Luteyn employed an extremely nar¬
row concept for the typical variety roraimae, char¬
acterizing it by a shiny and reticulately raised-
veined upper surface and assigning to it only two
additional specimens besides the type (of which they
examined a reduced black-and-white photograph).
They then relegated to variety parvifolius all remain¬
ing specimens of the species, encompassing a wide
range of variation in leaf size, crenulation of the mar¬
gins, and length of the inflorescence. After examin¬
ing a full-sized color Cibachrome print of the type
of 0. roraimae, kindly sent by the Kew herbarium.

Volume 5, Number 3
1995

Berry & Ramirez 229

Ixonanthaceae from Venezuelan Guayana

and studying the full range of variation of the spe¬
cies, there is no consistent way to distinguish the
specimens of variety roraimae from variety parvifol¬
ius, and all specimens cited by Steyermark & Luteyn
(1980) under variety parvifolius are now treated as
belonging to variety roraimae. The status of the bas-
ionym of variety parvifolius is discussed following the
key to the varieties of O. roraimae.

Key to the Varieties of Ochthocosmus roraimae

1. Leaf blades of fertile branches oval, 7-13 X 4—8
cm, margins entire or nearly so; panicles 11—14
cm long, 10-12 cm wide; known only from the

upper slopes of Cerro Venamo.

. 0. roraimae var. grandifolius

1. Leaf blades of fertile branches mostly obovate, 3—

10 X 2—6 cm, margins usually noticeably crenate
or toothed; panicles 3—8 cm long, 2—5 cm wide;
widespread throughout much of the Gran Sabana,
also in western Guyana and adjacent Brazil . . .
. 0. roraimae var. roraimae

Ochthocosmus parvifolius Hallier f., Beih. Bot. Cen-
tralbl. 39 (2). 15. 1921. Ochthocosmus rorai¬
mae var. parvifolius (Hallier f.) Steyermark &
Luteyn, Brittonia 32: 135. 1980. TYPE: Guy¬
ana or Venezuela (“British Guiana” on label),
undershrub 6—10 ft. high, sandstone region,
Mt. Roraima, Oct.—Nov. 1842, Richard Schom-
burgk 1046 (holotype, B destroyed, photo, US,
VEN).

The holotype of Ochthocosmus parvifolius Hallier
f. at B was destroyed in 1943, and there is appar¬
ently no extant duplicate material (inquiries were
made at B, BM, G, K, and W). The type photo¬
graphs are dark and much reduced in size from the
original specimen, with no evidence of flowers or
inflorescences. Steyermark & Luteyn (1980) treated
this as a variety of O. roraimae, but the inflores¬
cence, described as just 2.5 cm long, is very short
for that species. According to the protologue of 0.
parvifolius and Schomburgk’s collection notes in the
New York Botanical Garden archives, the type
specimen came from a low shrub smaller than the
more robust and often tree-like O. roraimae.
Ochthocosmus parvifolius was also described as
having unequal sepals, which are characteristic of

species such as 0. floribundus Gleason, but not of
O. roraimae, which has equal-sized, glandular-mar-
gined sepals. Ochthocosmus floribundus is not
known to occur around Mt. Roraima and is known
primarily from areas farther north and west in Es-
tado Bolivar such as AuyAn-tepui and Cerro Guia-
quinima.

The possibility also exists that Ochthocosmus
parvifolius is conspecific with a later-described spe¬
cies such as 0. attenuatus Steyermark & Luteyn,
which is known so far only from the type locality
near San Rafael de Camoirdn in the northern part
of the Gran Sabana. Given the current uncertainties
concerning the identity of O. parvifolius, and until
the issue can be better resolved, we recommend
that this species be treated as one of uncertain po¬
sition (incertae sedis).

Acknowledgments. We thank Bruno Manara for
making the illustrations, the Royal Botanic Gardens
at Kew for the Cibachrome type photograph of
Ochthocosmus roraimae, The New York Botanical
Garden for providing a loan of their specimens and
copies from the Schomburgk collection notes, and
H. Sipman (B), S. Knapp (BM). F. Jacquemoud (G),
G. Lucas (K), J. Pruski (US), and H. Riedl (W) for
checking for possible type material of 0. parvifol¬
ius. This work was supported in part by National
Science Foundation grant BSR-9201044 to the
Missouri Botanical Garden. CVG-EDELCA (Elec-
trificacidn del Caronf, C.A.) kindly provided logis¬
tical support for field studies related to this paper.

Literature Cited

Folster, H. 1992. Holocene autochthonous forest degra¬
dation in southeast Venezuela. Pp. 25-A4 in J. G. Gol-
dammer (editor). Tropical Forests in Transition. Birk-
hauser Verlag, Basel.

Forman, L. 1965. A new genus of Ixonanthaceae, with
notes on the family. Kew Bull. 19: 517-526.

Mabberley, D. J. 1990. The Plant-Book. A Portable Dic¬
tionary of the Higher Plants. Cambridge Univ. Press,
Cambridge.

Steyermark, J. A. 1984. Flora of the Venezuelan Guaya¬
na. I. Ann. Missouri Bot. Gard. 71: 297-340.

-. 1988. Flora of the Venezuelan Guayana. IV.

Ann. Missouri Bot. Gard. 75: 311—351.

-& J. L. Luteyn. 1980. Revision of the genus

Ochthocosmus. Brittonia 32: 128—143.

Three New Combinations in Pfaffia (Amaranthaceae) from the
New World Tropics

Thomas Borsch

Forschungsinstitut Senckenberg, Senckenberganlage 25,

60325 Frankfurt am Main, Germany

Abstract. The new combinations Pfaffia aurata
(Martius) Borsch, Pfaffia completa (Uline & W. L.
Bray) Borsch, and Pfaffia costaricensis (Standley)
Borsch are made. In analyzing the pollen morphol¬
ogy and floral structures of the three species, it is
shown that they belong to Pfaffia Martius and not
to Iresine P. Browne, nom. cons.

Pfaffia belongs to Amaranthaceae subfamily
Gomphrenoideae (Schinz, 1934; Townsend, 1993).
It comprises about 35 species, all characterized by
long trichomes arising in tufts from the pedicel or
the base of the perianth, a bilobate stigma, and the
flowers all being hermaphrodite. The genus is dis¬
tributed in the New World, according to Stiitzer’s
(1935) monograph, ranging from Mexico south¬
wards throughout the tropics, including the Amazon
basin, to the region of Bahia Blanca, Argentina.

During preparation of the Amaranthaceae treat¬
ment for Flora de Nicaragua and subsequent stud¬
ies in the genera Pfaffia and Iresine, three new
combinations turned out to be necessary.

Pfaffia aurata (Martius) Borsch, comb. nov. Bas-
ionym: Trommsdorffia aurata Martius, Nova
Genera et Species Plantarum, Vol. 2; 41. 1826.
TYPE: Brazil. Provinciae Rio Negro: habitat
in argillosis praeruptis oparatis flu[. . .] Japura
ripis ad Mar[ibis], Dec.-Jan., C. F. P. v. Mar¬
tius s.n. (holotype, M).

Pfaffia completa (Uline & W. L. Bray) Borsch,
comb. nov. Basionym: Iresine completa Uline
& W 1,. Bray, Bot. Gaz. (Crawfordsville) 21:
349. 1896. TYPE: Honduras. Santa Barbara:
San Pedro Sula, 200 m, C. Thieme 338 (ho¬
lotype, US; isotypes, BM, K). Figure 1.

Iresine wrightii Standley, Contr. U.S. Natl. Herb. 18: 97.
1916. Syn. nov. TYPE: Nicaragua [Herbarium of the
U.S. North Pac. Expl. Exped. under Commanders
Ringgold and Rodgers 1853-1856], C. Wright s.n.
(holotype, US).

Iresine ealea (Ibanez) Standley, Contrib. U.S.
Natl. Herb. 18: 94. 1916.

Iresine completa Uline & W. L. Bray f. glabriuscula Sues-
senguth, Repert. Spec. Nov. Regni Veg. 39: 11.
1935. Syn. nov. TYPE: Mexico. Mazatl&n: Felsige
Abhange, Jan.-Mar. 1902, C. A. Purpus 362 (holo¬
type, WU).

There has been some confusion about the iden¬
tity of this species because John Donnell Smith re¬
numbered Thieme’s specimen ( Thieme 338) with
his own number, 5443. There is a second collection
by Thieme from the same locality in Honduras, re¬
numbered 5447. Suessenguth (1934) erroneously
cited “Donnell-Smith n. 5447" as the type. This
specimen (BM, US) is a functionally male plant of
Iresine calea (Ibanez) Standley. It has small pistil-
lodes and might have been interpreted as being
hermaphroditic for that reason, but the plant lacks
the long basal trichomes that in hermaphroditic or
functionally female flowers of all species of Iresine
serve for wind dispersal. Obviously, this specimen
cannot be the type mentioned by Uline & Bray
(1896). In line with his misinterpretation, Suessen¬
guth (1935) described Iresine completa Uline & W.
L. Bray f. glabriuscula Suessenguth from a speci¬
men that is a functionally male plant of Iresine ca¬
lea.

When publishing Iresine completa, Uline & Bray
(1896) stressed the complete hermaphroditism of
their plant, a remarkable character in contrast to
the many species belonging to Iresine which are
dioecious or polygamous. It is obvious that they
called it “ completa ” for that reason.

Pfaffia costaricensis (Standley) Borsch, comb,
nov. Basionym: Iresine costaricensis Standley,
Contrib. U.S. Natl. Herb. 18: 94. 1916. TYPE:
Costa Rica. Tucurrique, Buissons a Las Vuel-
tas, 635 m. Mar. 1899, A. Tonduz 13183 (ho¬
lotype, US; isotypes, K, US). Figure 2.

Pfaffia aurata, P. completa, and P. costaricensis
show the bilobate stigma, which is a distinguishing
character of Pfaffia. In P. aurata the lobes have
almost the outline of an equilateral triangle. In P-
completa the lobes are narrowly triangular, and in
P. costaricensis the stigma is shaped more or less

Novon 5; 230-233. 1995.

Volume 5, Number 3
1995

Borsch

New Combinations in Pfaffia

231

Figures 1, 2. Scanning electron (SEM) photomicrograph of air-dried pollen. —1. Pfaffia completa (Thieme 338, BM).
—2. P. costaricensis (Tonduz 13183, K).

like a mushroom cap and the lobes are very short
and broad. All three species have subequal, 3-
veined tepals with a tuft of long, straight and stiff
simple trichomes arising abaxially at the base of
the tepal and from the pedicel. The flowers are al¬
ways hermaphroditic. Duke’s (1961) suspicion that
P. completa could be a polygamous species with the
pistillate flowers perfect and the staminate flowers
bearing pistillodes is a result of the above-men¬
tioned confusion about the species’ identity. In all
three species the filaments are united at the base
into a shallow cup. Broadly rounded pseudostami-
nodia are present, which are distinctly shorter than
the free part of the filament. Sometimes they are
notched at the apex or they are indistinct and the
margin of the cup is only slightly raised.

The pollen grains of Pfaffia aurata, P. completa
(Fig. 1), and P. costaricensis (Fig. 2) are pantopor-
ate, having 12—14, 5-angular lumina and sunken
pores. They represent the Gomphrena- type, one of
two major pollen types in the Amaranthaceae (Erdt-
man, 1952). The Gomphrena-type is placed in a
specialized category in Nowicke’s (1975) study of
centrospermous pollen. Pfaffia has pollen of the
Gomphrena-type, whereas pollen of lresine is sim¬
ilar to the Amaranthus- type of Erdtman, which cor¬

responds to Pollen-Type II of Nowicke (1975). In
Pfaffia aurata, P. completa, and P. costaricensis, the
tectum appears to cover the columellae completely
and the muri have more or less vertical sides. The
extraporal sexine bears distinct elongate spinules
in all three species. This is exactly the same struc¬
ture as in species of Pfaffia (see illustrations of
Pfaffia pollen by Nowicke & Skvarla (1979), Cuad-
rado (1988), Eliasson (1988), and Townsend
(1993)). There is only variation in aperture number,
which is known to occur between species within
many genera of Amaranthaceae. Eliasson (1988)
showed Pfaffia- pollen to have a row of warts or spi¬
nules on the distal part of their tectum. In P. au¬
rata, a species illustrated by Eliasson as figure 34
under the misapplied name Pfaffia paniculala
(Martius) Kuntze, these rows are not very straight.
In investigating more specimens of P. aurata from
different localities the same observation was made.
Sometimes the spinules were in an undulate row,
some of them even side by side. The pollen of P.
costaricensis (Fig. 2) also has some of the spinules
side by side, but there is a certain amount of vari¬
ation, from a side by side arrangement of the spi¬
nules to straight rows, even within a number of
grains from a single anther. In contrast to lresine.

232

Novon

the spinules are never evenly spread over a more
or less convex distal part of the tectum. In Iresine
the pores are small and apertures cover a minor
part of the surface of the grain. Iresine angustifolia
Euphrasen has more specialized pollen with wider
pores, compared to the other species of the genus,
but the distal parts of the tectum are still convex
and the spinules are evenly spread.

Pfaffia aurata, P. completa, and P. costaricensis
are closely related and belong to a group that was
described as Trommsdorffia by Martius (1826). In
1839 Dietrich transferred Trommsdorffia aurata
Martius (the type species of Trommsdorffia ), T. ar-
gentata Martius, and T. pulverulenta Martius to Ire¬
sine P. Browne. Similarities in habit (all species be¬
ing scandent shrubs or lianas), several times
branched racemose inflorescence structures, and
the common occurrence of long trichomes serving
for dispersal were the decisive factors for merging
the two genera. My own studies in connection with
a revision of Iresine revealed that Trommsdorffia is
different from Iresine not only in pollen morphology
but also in floral structure, most strikingly in the
morphology of the stigma (two more or less elon¬
gated, filiform branches). Iresine and Trommsdorffia
belong to completely different evolutionary lines.
This is also supported by the strong trend to the
evolution of polygamous or dioecious species in Ire-
sine, whereas the relatives of Pfaffia aurata and all
other species of Pfaffia are generally hermaphro¬
ditic. Consequently, Trommsdorffia has to be ex¬
cluded from Iresine. Trommsdorffia pulverulenta
Martius (= Pfaffia densipellita Borsch), a morpho¬
logically well defined species also allied to P. au¬
rata from Peru, was excluded from Iresine earlier
(Borsch, 1993).

Pfaffia aurata and its allies form a small group
of closely related species within the genus Pfaffia.
Differences from the other species of Pfaffia are
merely a matter of degree. The occurrence of more
or less broadly rounded pseudostaminodia in P. au¬
rata and its relatives is not sufficient to support
generic status. Their position within the genus Pfaf¬
fia and the generic concept of Pfaffia will be dis¬
cussed in detail in a separate paper. It is question¬
able whether Trommsdorffia argentata Martius is
specifically distinct from Pfaffia aurata, and sev¬
eral other morphologically very similar taxa (e.g.,
Iresine argentata (Martius) D. Dietrich var. ama-
zonica Seubert = Iresine aurata (Martius) D. Die¬
trich var. amazonica (Seubert) Suessenguth) are be¬
ing studied more thoroughly before a taxonomic
decision is made as to their status among the rel¬
atives of P. aurata.

Pfaffia aurata has tepals that are ovate-oblong.

3-veined, 1.3-1.8 mm long and stigma lobes that
have more or less the outline of an equilateral tri¬
angle. Pfaffia completa is characterized by its te¬
pals, which are narrowly triangular, prominently 3-
veined, about 2.5 mm long, narrowly triangular
stigma lobes becoming almost cylindrical at top,
and broadly lanceolate blades, which are glabrous
on the upper and sparsely pubescent on the lower
surface. Pfaffia costaricensis has extremely small
floral parts and conspicuously finely branched in¬
florescences, representing 3—4 times branched rac¬
emose structures with strictly opposite branches.
The tepals are ovate, 1.1—1.3 mm long, and indis¬
tinctly 3-veined. The stigma lobes are very broadly
rounded, even shorter and broader than in P. au¬
rata. The anthers have a length of 0.2 mm, in con¬
trast to the other species, which have anthers that
are about 0.4 mm long. Burger (1983) described
the characteristic elliptic or broadly elliptic leaf
blades of P. costaricensis, with up to 12 pairs of
major secondary veins.

Pfaffia aurata probably ranges throughout the
Neotropics. In Ecuador the species is well estab¬
lished in virgin forests as well as rastrojos, accord¬
ing to the Flora of Ecuador (Eliasson, 1987). There
the species is erroneously called Pfaffia paniculata
(Martius) Kuntze, a name of a different taxon be¬
longing to a group of closely related species that
were described as Hebanthe by Martius (1826).
Suessenguth (1934) described Pfaffia completa as
occurring from Guatemala to Costa Rica. This range
is partly a consequence of the misinterpretation of
the species’s identity. In herbaria many older spec¬
imens identified as Iresine completa Uline & W. L.
Bray belong in fact to Iresine calea (Ibdnez) Stan-
dley. Apart from the type locality in Honduras,
Pfaffia completa is at the moment proven to occur
in Nicaragua and Panama. Two specimens exist
from Nicaragua. One has been collected recently
on Isla de Ometepe in Lago de Nicaragua, and the
other is the type of Iresine wrightii. The single spec¬
imen from Panama has been collected in the Darien
and is also cited by Duke (1961). Pfaffia costari¬
censis is endemic to Costa Rica and is known only
from the central part of the country (Burger, 1983).
A recent collection has been made in a remote area
in Puntarenas, where the plant was climbing on
secondary woody vegetation (Hammel, pers.
comm.).

Additional specimens examined. Iresine calea (Iba¬
nez) Standley (specimens incorrectly identified as Iresine
completa Uline & W. L. Bray): COSTA RICA. Puntar¬
enas: Santo Domingo de Golfo Dulce, Tonduz 9861 (BM).
HONDURAS. Santa Barbara: San Pedro Sula, Thieme
s.n., renumbered by Donnell Smith 5447 (BM,US). NIC-

Volume 5, Number 3
1995

Borsch

New Combinations in Pfaffia

233

ARAGUA. Managua: S of Managua, J. M. & M.T. Green-
man 5725 (MO). Pfaffia aurata (Martius) Borsch: NIC¬
ARAGUA. Boaco: San Jos6 de los Remates, Moreno
20337 (MO). Rivas: Isla de Ometepe, al NE del Volc&n
Concepcidn, Sandino 4233 (FR, HNMN not seen). Ze-
laya: La Pimienta, Pipoly 6291 (FR); along new road from
Siuna to Matagalpa, ca. 31.4 km beyond Rfo Uli, near
Wani, Stevens 7493 (FR). Pfaffia completa (Uline & W.
L. Bray) Borsch: NICARAGUA. Rivas: Isla de Ometepe,
Volc£n Madera, Robleto 291 (FR, MO). PANAMA. Dari¬
en: Paca, Williams 706 (US). Pfaffia costaricensis
(Standley) Borsch: COSTA RICA. Puntarenas: Canton de
Osa, Fila Costena, Fila Cruces, cabezeras del Rfo Piedras
Blancas, Hammel 19289 (MO).

Acknowledgments. I thank the curators and di¬
rectors of the herbaria MO, US, and WU for loaning
specimens, of BM and K for loaning specimens and
permitting the removal of pollen for SEM studies,
and of M for allowing me to study the herbarium
of C. F. P. v. Martius. I am grateful to Alan J. Paton
for his kind help during my work on the collections
at Kew, to Troels M. Pedersen, Mburucuya, Argen¬
tina, for hinting to me the true identity of Hebanthe
paniculata Martius, and to Barry Hammel, Missouri
Botanical Garden, for information on Pfaffia cos¬
taricensis. The SEM photographs were taken at the
Forschungsinstitut Senckenberg using a CamScan
CS 24 Compact scanning electron microscope. I am
especially grateful to the research staff of the Mis¬
souri Botanical Garden, who first drew my attention
to the Amaranthaceae.

Literature Cited

Borsch, Th. 1993. Amaranthaceae. Pp. 18-26 + Appen¬
dix I in L. Brako & J. L. Zarucchi, Catalogue of the
Flowering Plants and Gymnosperms of Peru/Cat£logo de
las Angiospermas y Gimnospermas del Perti. Monogr.
Syst. Bot. Missouri Bot. Gard. 45.

Burger, W. C. 1983. Amaranthaceae. In: Flora Costari¬
censis. Fieldiana, Bot. 13: 142-180.

Cuadrado, G. A. 1988. Granos de polen de Amarantha¬
ceae del nordeste Argentino: 2. G£nero Pfaffia. Bol.
Soc. Argent. Bot. 25(3—4): 385-394.

Dietrich, D. 1839. Synopsis Plantarum. Vol. 1. Bernhard
Friedrich Voigt, Weimar.

Duke, J. A. 1961. Amaranthaceae. In: Flora of Panama.
Ann. Missouri Bot. Gard. 48: 6-50.

Eliasson, U. H. 1987. Amaranthaceae. Pp. 1-138 in G.
Harling & L. Andersson, Flora of Ecuador, Vol. 28. Ber-
lings, Alov.

-. 1988. Floral morphology and taxonomic rela¬
tions among the genera of Amaranthaceae in the New
World and the Hawaiian Islands. J. Linn. Soc., Bot. 96:
235-283.

Erdtman, G. 1952. Pollen Morphology and Plant Taxon¬
omy. Angiosperms. Almqvist & Wiksell, Stockholm.

Martius, C. F. P. von. 1823—1832. Nova Genera et Spe¬
cies Plantarum. 3 Vols. Lindauer et Wolf, Monachii.

Nowicke, J. W. 1975. Pollen morphology in the order
Centrospermae. Grana 15: 51-77.

-& J. J. Skvarla. 1979. Pollen morphology: The

potential influence in higher order systematics. Ann
Missouri Bot. Gard. 66: 633-700.

Schinz, H. 1934. Amaranthaceae. Pp. 7-85 in A. Engler
& K. PrantI, Die natiirlichen Pflanzenfamilien, ed. 2,
16c. Leipzig.

Stutzer, 0. 1935. Die Gattung Pfaffia. Repert. Spec. Nov.
Regni Veg. Beih. LXXXVIII, 1-46. Berlin.

Suessenguth, K. 1934. Neue und kritische Amarantaceen
aus Siid-und Mittelamerika. Repert. Spec. Nov. Regni
Veg. 35: 298-337.

-. 1935. Neue und kritische Pflanzen aus Suda-

merika, insbesondere Amarantaceen, sowie eine neue
Gattung der Podostemonaceae. Repert. Spec. Nov. Reg¬
ni Veg. 39: 1—20.

Townsend, C. C. 1993. Amaranthaceae. Pp. 70-91 in K.
Kubitzki et al., Families and Genera of Flowering
Plants, Vol. 2. Springer, Berlin.

Uline, E. B. & W. L. Bray. 1896. Synopsis of North Amer¬
ican Amaranthaceae V. Bot. Gaz. (Crawfordsville) 21:
348-356.

Paspalum morichalense (Poaceae: Paniceae), a New Aquatic
Species from South America

Gerrit Davidse

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Fernando 0. Zuloaga

Instituto de Botanico Darwinion, Casilla de Correo 22, San Isidro, 1642, Argentina
Tarciso S. Filgueiras

Reserva Ecologica do IBGE, Caixa Postal 08770, 312-970 Brasilia, DF, Brazil

Abstract. Paspalum morichalense is described,
illustrated, and compared to related species of the
group Dissecta. It occurs in Venezuela, Surinam,
Brazil, and Bolivia. Its confusion with Paspalum
commutatum Nees and Paspalum lacustre Chase ex
Swallen is discussed.

We have been aware for some time of the exis¬
tence of a possibly undescribed, aquatic species of
Paspalum L. However, because of its confusion
with two other species, P. commutatum and P. la¬
custre, it was not possible to describe it until the
types of those two names had been studied. Pas¬
palum is a largely American genus with approxi¬
mately 330 species. The group names used in this
paper are the ones used by Chase (1929, ined.) in
her infrageneric classification of the genus.

Paspalum morichalense Davidse, Zuloaga & Fil¬
gueiras, sp. nov. TYPE: Venezuela. Guarico:
ca. 32 km SSE of Calabozo on Finca Becerra,
in and around morichal with standing water,
morichal dominated by Mauritia flexuosa, 65
m, 6 Nov. 1973, Davidse 3770 (holotype, MO;
isotypes, AAU, K, LE, MEXU, MG, PRE, SI,
US, VEN). Figure 1.

Gramen perenne, repens vel fluitans, aquaticum; culmi
13-25 cm alti, ramosi. Folia caulina; ligulae 0.6-2.0 mm
longae, glabrae. Racemi 1-2, 3-6 cm longi; rhachis 1.1-
3.5 mm lata, alata, herbacea, scaberula. Spiculae 2.1—2.9
X 0.9-1.1 mm, ellipticae, singulares, non imbricatae, gla¬
brae, acutae. Antherae 3, 0.O—0.8 mm longae, purpureae.

Creeping or floating, aquatic perennial. Culms
many-noded, decumbent and rooting at the lower
nodes, freely branching, the erect portion of the
culms 13—25 cm tall; intemodes hollow, glabrous;
nodes dark, glabrous or sparsely pilose. Leaves
cauline. Sheaths 2-8 cm long, glabrous, slightly

Novon 5; 234-237. 1995.

keeled; collar dark glabrous. Ligules 0.6—2.0 mm
long, membranous, truncate, glabrous. Blades 3.5-
11 cm long, 1.5-3 mm wide, linear to linear-lan¬
ceolate, flat, narrowed at the base, widely spreading
to reflexed, glabrous to scabrous on the adaxial sur¬
face and glabrous on the abaxial surface, the mar¬
gins scaberulous. Inflorescence terminal, partially
included in the upper leaves; peduncle 2 cm long,
cylindric; racemes 1-2, 3-6 cm long; rachis 1.1—
3.5 mm wide, winged, herbaceous, green, incon¬
spicuously veined, glabrous, terminating in a spike-
let, the margins minutely scaberulous, usually fold¬
ed and partially embracing the spikelets, the
spikelets not crowded, not overlapping on the same
side of the rachis; pedicels solitary, 0.3-0.5 mm
long, scaberulous. Spikelets 2.1-2.9 mm long, 0.9—
1.1 mm wide, elliptic, concave-convex, broadly
acute, whitish green, glabrous. Lower glume absent.
Upper glume and lower lemma subequal or the up¬
per glume 0.1-0.2 mm shorter than the upper flo¬
ret, hyaline, 3—5-nerved, with one central nerve
and with the submarginal and marginal nerves
close together. Lower floret sterile. Lower lemma 5-
nerved. Lower palea absent. Upper floret 2.1—2.8
mm long, 0.9—1.1 mm wide, as long as the spikelet,
bisexual, elliptic, acute, whitish, indurate, smooth
and shining, papillose and with prickle hairs and
bicellular microhairs at the apex; lodicules 2, 0.3
mm long, truncate. Stamens 3; anthers 0.5-0.8 mm
long, purple. Styles 2, separate; stigmas ca. V4 as
long as the styles, plumose, purple. Caryopsis 1.6—
1.8 mm long, 0.9 mm wide, elliptic, brownish; hi-
lum ca. % as long as the caryopsis, linear; embryo
ca. % as long as the caryopsis.

This species is known from Venezuela to Brazil
and Bolivia. We have studied it in the field in GuA-
rico, Venezuela, and Bahia, Brazil. In GuArico it
grows on mud and in shallow water of swamps and

Volume 5, Number 3
1995

Davidse et al.

Paspalum morichalense

235

Figure I. Pnnicum morichalense Davidse, Zuloaga & Filgueiras (Davidse 3770 MO). —A. Habit. B. Inflorescence.
-C. Spikelet, upper glume view. -D. Spikelet. lateral view. E. Spikelet, lower lemma vtew. F. Uppeir floret,
lemma view. -G Upper floret, palea view. -H. Detail of apex of the upper lemma. -I. Caryops.s, embryo s.de. -
J. Caryopsis, hilum side.

ponds at an elevation between 65 and 125 m, es¬
pecially in morichales, palm swamps dominated by
Mauritia flexuosa L. f. (Arecaceae). The Bahian
gathering was collected at an elevation of approx¬
imately 450 m in a pond ca. 20 m across and ca.
1-2 m deep. Here it grew at the edges and at the
center of the pond and was much sought after by

cattle and horses. The latter entered into the deeper
waters of the pond to reach the soft, floating culms.

The specific epithet is derived from the word
morichal, which is the Venezuelan designation for
a Mauritia swamp. The equivalent Brazilian vege¬
tation type is commonly known as vereda.

Because of its few racemes, foliaceous, winged

236

Novon

rachis, aquatic habit, creeping or floating culms
with many elongated nodes, and glabrous spikelets,
Paspalum morichalense belongs to the informal
group Dissecta of Chase (1929). Within this group
it is most closely related to P. lacustre and P. dis-
sectum (L.) L.

Paspalum lacustre differs from P. morichalense
by its more numerous racemes ('1—6 vs. 1—2), spike-
lets sharply acute (vs. broadly acute), the upper
floret shorter than the upper glume and lower lem¬
ma (0.4—0.9 mm shorter vs. as long as or only 0.2
mm shorter), longer pedicels (ca. 1.0 mm vs. 0.3-
0.5 mm), and rachis scabrous versus minutely sca-
berulous. Paspalum lacustre is known only from the
type collection [BRAZIL. Amapa: Mun. Macapa,
Ambe, localidade no afluente do Rio Pedreira, em
lago fundo. 8 jul. 1951, R. L. Frdes & G. A. Black
27312 (holotype, INPA not seen, fragment US; iso¬
type, US)].

Paspalum dissectum differs from P. morichalense
in its more numerous racemes (2-A vs. 1-2), spike-
lets generally smaller (1.8-2.2 X 1.2-1.5 vs. 2.1-
2.9 X 0.9-1.1 mm), obovate (vs. elliptic), and more
densely arranged (i.e., overlapping in the same row
on one side of the rachis vs. not overlapping in one
row), veins more strongly developed in the rachis
wings, and the rachis wing margins scabrous (vs.
minutely scaberulous).

Although specimens of Paspalum morichalense
have been known at least since the 1864 collection
of Warming in Brazil, the species has remained for¬
mally unrecognized until now and has been con¬
fused with two other species. As noted by Chase
(ined.), Doell (1877) in Flora Brasiliensis partially
misapplied the name Paspalum commutatum. Un¬
der this species he correctly cited a Riedel collec¬
tion, but also included the Warming collection.
Paspalum commutatum, as shown by an examina¬
tion of a fragment of the holotype [BRAZIL: pr.
Porto Est. locis tempore pluviore inundatis . . .
Langsdorff s.n. (holotype, LE not seen, fragment
US)], belongs to the group Plicatula of Chase (1929)
because it has compressed sheaths, 4-5 racemes,
broadly obovate spikelets, and a dark brown, shin¬
ing upper floret. There is no close relationship be¬
tween groups Dissecta and Plicatula.

As shown by her annotations and her unpub¬
lished manuscript. Chase (ined.) recognized that P.
morichalense was an undescribed species, but she
did not differentiate it from P. lacustre, a concept
also followed by Judziewicz (1990). On the other
hand, Swallen (1967), who effectively published P.
lacustre, apparently did distinguish between P. mor¬
ichalense and P. lacustre, since he cited only the
type collection, Frdes & Black 27312, under his

new species. Unfortunately, he mistakenly classi¬
fied this species as a member of the Plicatula
group. In annotations of the US specimens Chase
had equated the two collections of Frdes and Black
( 27195 and 27312 ) under the same, unpublished
name. We, on the other hand, include Frdes &
Black 27195 under P. morichalense, and Frdes &
Black 27312 under P. lacustre.

As noted earlier, the description of P. lacustre in
Judziewicz (1990: 484) is based on a mixture of P.
morichalense and P. lacustre, but the illustration
(fig. 84A—C) represents only P. morichalense.

These species of group Dissecta may be distin¬
guished through the following key:

la. Rachis of the raceme extending beyond the
spikelets as a naked point.

2a. Spikelets 0.6-0.8 mm wide, in 2 rows; Unit¬
ed States to Argentina. P. repens Bergius

2b. Spikelets 0.8-1.2 mm wide, in 1 row; Mex¬
ico (Nayarit to Chiapas) . . . P. longicuspe Nash

lb. Rachis ending in a spikelet.

3a. Upper floret as long as the lower lemma and
upper glume or at most 0.1-0.2 mm shorter;
Venezuela, Surinam, Brazil, Bolivia . . .

P. morichalense Davidse, Zuloaga & Filgueiras
3b. Upper floret at least 0.2 mm shorter than the
lower lemma and upper glume.

4a. Spikelets on the same side of the rachis
not overlapping; Brazil (Amapa) . .

. P. lacustre Chase ex Swallen

4b. Spikelets on the same side of the rachis
overlapping, especially toward the base
of the rachis.

5a. Spikelets 1.8-2.2 mm long, obtuse;
eastern United States, Cuba . .

. P. dissectum (L.) L.

5b. Spikelets 3—3.9 mm long, acumi¬
nate.

6a. Racemes 2, rarely 1; blades
mostly 3-6 mm wide; Cuba,

Jamaica .

. . P. serratum Hitchcock & Chase
6b. Racemes 3-5, rarely 2; blades
mostly more than 8 mm wide;
southern United States to Ar¬
gentina . . . P. acuminatum Raddi

Paratypes. VENEZUELA. Bolivar: small swamp in
open savanna at base of Serranta de Pijiguao, 160 km SW
of Caicara del Orinoco, 6°35'N, 66°45'W, 100—125 m, 12
Sep. 1985, Steyermark, Holst & Manara 131802 (MO,
VEN). Guariro: 48 km N of Santa Rita along Hwy. 12
between Cabruta and Chagauramas, shallow pond with
standing water in Trachypogon savanna with Curatella and
Bowdichia, 110 m, 21 Nov. 1973, Davidse 4287 (COL, K,
MO, P, RB, US, VEN). SURINAM. Sipaliwini, savannah,
30 Jan. 1965, Fraser 3 (US). BRAZIL. Amapa: Mun. de
Macapa. Rio Macaeoari, Sao Benedito do Bra?o, 4 Jan.
1951, Frdes & Black 27195 (US). Bahia: natante na lagoa
Pipiri, Rio Santa Clara, 17 Dec. 1987, Filgueiras 1295
(CEN, IBGE, K, SP). Minas Gerais: Lagoa Santa, in
aquis stagnantibus, fluitans, 5 Apr. 1864, Warming s.n.
(US). Mato Grosso: just S of the Rio Suia Missu Ferry,

Volume 5, Number 3
1995

Davidse et al.

Paspalum morichalense

237

ca. 40 km NW of base camp at 12°49'S, 51°46'W, edge
of small lake in shallow water, grass with weak procum¬
bent rooting stems with ascending culms, 24 Nov. 1968,
Harley 11213 (MO, UB, US). Mato Grosso do Sul: Mun.
de Corumbti, Fazenda Santa Delfina, erva prostrada, pouco
arraigada ao solo, frequente em va^ante seca, solo aren-
oso, 3 Dec. 1978, Allem, Vails, Vieira & Comastri 2557
(CEN, MO); Mun. de Corumb£, Fazenda Sao Bento, erva
prostrada, pouco arraigada ao solo, frequente em vazcante
seca, 2 Nov. 1978, Allem, Vails, Vieira & Comastri 2529
(CEN, MO). Para: Gavinho, Maraj6 Island, Mar. 1920,
Goeldi 284 (US). BOLIVIA. Rfo Guarap6, Santo Antonio
de Manoelito, 25 Jun. 1952, Black & Cordeiro 52-15320
(US).

Acknowledgments. We thank Vladimiro T. Du-
das for the excellent illustration and Robert J. So-
reng for checking the identity of the types at LE.

Fieldwork by Davidse was made possible through
the support of NSF grant GB40630. Filgueiras
thanks CNPq (Brazilian Research Council), Brasi¬
lia, for a fellowship.

Literature Cited

Chase, A. 1929. The North American species of Paspa¬
lum. Contr. U.S. Natl. Herb. 28: 1-310, i-xvii.

-. Ined. Paspalum of South America. Unpublished

manuscript. Hitchcock and Chase Library, Botany De¬
partment, Smithsonian Institution, Washington, D.C.
Doell, J. C. 1877. Paniceae. In: C. F. P. Martius, FI. Bras.
2(2): 33-358.

Judziewicz, E. J. and collaborators. 1990. Poaceae (Gra-
mineae). FI. Guiananas, Ser. A: Phanerogams 8: 1-727.
Swallen, J. R. 1967. New species of Paspalum. Phyto-
logia 14: 35B-389.

A New Species of Brachynema (Olacaceae) from South America

Rodrigo Duno de Stefano

Fundacion Institute) Botanico de Venezuela, Apartado 2156, Caracas 1010-A, Venezuela

Paul E. Berry

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Giovannina Orsini Veldsquez

Universidad Central de Venezuela, Facultad de Ciencias, Escuela de Biologfa,
Apartado 20513, Caracas 1040, Venezuela

Abstract. A new species is described and illus¬
trated in the previously monotypic genus Brachy¬
nema (Olacaceae). Brachynema axillare is known
from the base of Sierra de la Neblina in southern
Venezuela and northern Brazil, as well as in Am¬
azonian Peru; it differs from B. ramiflorum Ben-
tham in its axillary inflorescences, non-striped co¬
rolla tube, glabrous corolla lobes, and mostly
4-merous flowers.

Resumen. Se describe una nueva especie en el
genero Brachynema (Olacaceae), previamente con-
siderado monotfpico. Brachynema axillare se cono-
ce de la base de la Sierra de la Neblina en el sur
de Venezuela y en el norte de Brasil, as! como en
la Amazonia peruana. Difiere de B. ramiflorum en
sus inflorescencias axilares, tubo de la corola no
rayado, ldbulos de la corola glabros, y las flores
mayormente 4-meras.

In the most recent revision of neotropical Ola¬
caceae (Sleumer, 1984), Brachynema was consid¬
ered monotypic and endemic to central Amazonian
Brazil. In a series of expeditions to the Sierra de
la Neblina in southern Venezuela between 1984
and 1987 (Brewer-Carfas, 1988), numerous addi¬
tional collections of the genus were made and sub¬
sequently determined as Brachynema ramiflorum
Bentham. However, more detailed study revealed
that the Venezuelan specimens are significantly dif¬
ferent from this species and merit recognition as a
distinct species, due to their axillary (vs. ramiflo-
rous), often tetramerous (vs. strictly pentamerous)
flowers, tricolpate instead of triporate pollen (Lob-
reau-Callen, 1980), larger fruits, smaller calyx at
anthesis, glabrous corolla lobes (vs. internally
hairy), and the base of the corolla tube purple and
lacking the characteristic “zebra” stripes of B. ram¬
iflorum.

Brachynema axillare R. Duno & P. E. Berry, sp.
nov. TYPE: Venezuela. Estado Amazonas: Dpto.
Rio Negro, 0 to 2 km W of Cerro La Neblina
Base Camp, along Rio Mawarinuma, 0°50'N,
66°10'W, 140 m, 7 Feb. 1984, Liesner 15707
(holotype, VEN; isotype, MO). Figure 1.

Haec species Brachynemati ramifloro Bentham affinis,
sed ab eo infloreseentia axillari non ramiflora, floribus
plerumque tetrameris, corollae lobulis glabris etiam tubo
basaliter purpureo, polline tricolpato differt.

Sparsely branched shrub or small tree 3-8 m tall.
Leaves chartaceous, simple, alternate, glabrous, es-
tipulate; blade narrowly to broadly elliptic or ovate,
20-40 cm long, 5-20 cm wide, base rounded to cu-
neate, apex long-acuminate to caudate, the tip often
somewhat curved to one side, penninerved, with 9-
13 subopposite to alternate pairs of parallel second¬
ary veins looping near the margin to form a sub¬
marginal vein, prominent on the lower surface, sunk¬
en on the upper surface, tertiary veins conspicuous
and running perpendicular to the secondary veins;
margin shallowly to prominently sinuate-dentate,
each tooth ending in a gland; petiole (1—)3—25 cm
long, terete, noticeably thickened at both the base
and the apex. Inflorescence an axillary fascicle or
dense corymb of 10-20 flowers on a thickened, gla¬
brous to puberulous axis 5-10 mm long. Flowers
actinomorphic, bisexual, 4- or 5-merous; pedicels 1—
2.5 mm long. Calyx cupular, greenish white when
fresh, shallowly lobed, ca. 0.5 mm tall and 2 mm
wide at anthesis. Corolla tube 16-25 mm long at
anthesis, firm-fleshy when fresh, 2-4 mm wide at the
base, glabrous to puberulous, uniformly violet-purple
at the base, turning white toward the rim, the lobes
white, glabrous, 4-6 mm long. Stamens 4 or 5, epi-
petalous, included at the base of the tube, filaments
ca. 1 mm long; anthers basifixed, ca. 3 mm long,
narrowly pointed at the apex, longitudinally dehis-

Novon 5: 238-240 1995

Volume 5, Number 3
1995

Duno de Stefano et al.
Brachynema axillare

239

cent; pollen suboblate, polar axis 22.5 /am long,
equatorial axis 28.7 /am long, circular or subtrian-
gular in polar view, tricolpate, the colpi broadened,
14 /am long by 5 /am wide, exine very finely retic¬
ulate or psilate, 3 /am thick, without apertural thick¬
enings. Ovary subglobose, 3—5-locular; ovules 1 per
locule, pendulous; stigma sessile, capitate. Drupe
globose or depressedly globose with a pointed apical
projection, 2—2.5 cm diam., green to purple-black

when fresh, the calyx enlarged and persistent, sur¬
rounding the lower part of the fruit. Seed 1. subglob-
ular, longitudinally striate, 1.5-2 cm diam.

Ecology and distribution. Evergreen lowland
forest on slopes and along gravelly and silty river-
banks, 100-200 m elevation, known mainly from
the base of Sierra de la Neblina in adjacent parts
of Estado Amazonas, Brazil, and Estado Amazonas,

240

Novon

Venezuela; also known in Amazonian Peru from
Departamento Loreto near the confluence of the
Rios Maranon and Ucayali.

Brachynema is still of uncertain placement in the
Olacaceae and is unique in the family in its glan¬
dular-dentate leaves and pulvinar thickenings at
both the base and apex of the petiole (Sleumer,
1984). The leaves of B. axillare are very similar to
those of B. ramiflorum, but the flowers and fruits
are always borne near the ends of young branches
in the leaf axils, rather than lower down on the
trunk or on branches where the leaves have already
dehisced. Neither the distinctive “zebra” striping
of the lower part of the corolla of of B. ramiflorum
nor the lines of hairs present on the inside of the
corolla lobes have been observed in any specimen
of B. axillare.

Paratypes. VENEZUELA. Amazonas: 0-2 km E of
Cerro La Neblina Base Camp, on Rfo Mawarinuma,
0°50'N, 66°10'W, 140 m, 7 Feb. 1984, Liesner 15707
(MO), 20 Feb. 1984, Liesner 16136 (MO, VEN); along Rfo
Baria (= Rfo Mawarinuma) just upstream from Base
Camp, SW side of Cerro de La Neblina, 0°49'50"N,
66°09'40"W, 140 m, 10 Apr. 1984, Gentry & Stein 46512

(MO), 11 Apr. 1984, Thomas & Plowman 3002 (MO, NY),
16 Apr. 1984, Gentry & Stein 46668 (MO), 4-5 July 1984,
Davidse & Miller 27009 (MO), 27 Nov. 1984, Croat 59373
(MO), 4 Dec. 1984, Anderson 13480 (MICH, MO), 17 Feb.
1985, Nee 30915 (MO, VEN), 18 Feb. 1985, Boom &
Weitzman 5894 (MO, NY), Stannard 459 (VEN), Stannard
460 (VEN); upper Rfo Barfa, ca. 0°55'N, 66°15'W, Dav¬
idse & Miller 26908 (MO); Cerro de la Neblina, lowland
and slope forests, 170 m, 29 Jan. 1954, Maguire , Wurdack
& Bunting 37401A (MO, NY). BRAZIL. Amazonas: Ser-
ra da Neblina Base Camp on Cano Tucano, Rio Cauaburf,
125 m, 12 Nov. 1965, Maguire, Steyermark & Maguire
60160 (MO, NY). PERU. Loreto: Dtto. Nauta, Rfo Mar¬
anon, Quebrada de Sapira del caserfo Florida, ca. 8 km
above Nauta, 13 June 1979, Rimachi 4465 (MO).

Acknowledgments. We thank Roy Gereau for
revising the Latin diagnosis.

Literature Cited

Brewer-Carfas, C. (Editor). 1988. Cerro de la Neblina.
Resultados de la expedici6n 1983-1987. Fundacidn
para el Desarrollo de las Ciencias Ffsicas, Matem4ticas
y Naturales, Caracas, Venezuela.

Lobreau-Callen, D. 1980. Caractferes compares du pollen
des Icacinaceae et des Olacaceae. Adansonia II, 20(1):
29-80.

Sleumer, H. O. 1984. Olacaceae. Flora Neotropica 38.
New York Botanical Garden, New York.

Dos Especies Nuevas del Genero Drosera (Droseraceae) en
Venezuela y Otros Comentarios Taxonomicos

Rodrigo Duno de Stefano

Fundacion Instituto Botanico de Venezuela, Apartado 2156, Caracas 1010-A, Venezuela

Alastair Culham

Department of Botany, Reading University, Reading, Whiteknights, RO. Box 221,
Reading RGG 2AS, United Kingdom

Resumen. Se describen dos especies nuevas del
genero Drosera provenientes de la Guayana Vene-
zolana: Drosera hirticalyx y D. yutajensis. Las es¬
pecies D. colombiana A. Fernandez y D. panamen-
sis M. D. Correa & A. S. Taylor son consideradas
sin6nimos de D. cayennensis Sagot ex Diels, mien-
tras que D. esmeraldae (Steyermark) Maguire &
Wurdack es citada de Colombia y D. communis A.
Saint-Hilaire es excluida de Venezuela.

Abstract. Two new species of Drosera are de¬
scribed from the Venezuelan Guayana: D. hirticalyx
and D. yutajensis. Drosera colombiana A. Fernan¬
dez and D. panamensis M. D. Correa & A. S. Taylor
are considered nonspecific with D. cayennensis Sa¬
got ex Diels. Drosera esmeraldae (Steyermark) Ma¬
guire & Wurdack is reported for Colombia, and the
occurrence of D. communis A. Saint-Hilaire in Ven¬
ezuela is disputed.

Se describen dos especies nuevas del genero
Drosera (Droseraceae), las cuales presentan hojas
simples estipuladas, y gineceo con 3 estilos bipar-
tidos desde la base, caracterfsticas dstas tfpicas del
subg6nero Drosera secci6n Drosera.

Drosera hirticalyx R. Duno & Culham sp. nov.
TIPO: Venezuela. Amazonas: Dpto. Atabapo,
Cerro Marahuaca-Huha, cumbre altiplanicie
no arbolada, m&s o menos plana, con piedras
expuestas, rfo abajo, 2580 m, 3°40'N,
65°30'0, 31 ene. 1982, Steyermark, Guariglia,
Holmgren, Luteyn & Mori 125982 (holotipo,
NY; isdtipo, VEN). Figura 1A-E.

Speciei Drosera roraimae affinis, sed foliis brevioribus.
Lamina basi gradatim attenuata; petiolus glaber vel subtus
pilis raris, unicellularibus, praeditus. Inflorescentia 4-15
cm, (1—)2—6-flora; pedunculus glandulis sessilibus. Sem¬
ina elliptica, faveolata, 100—150 X 50-100 /cm.

Planta herbdcea arrosetada. Rafces fibrosas. Tallo
corto. Hojas espatuladas, altemas, simples, 5—12

mm de largo; estfpulas intrapeciolares, 2-4 mm de
largo, 4-6 partidas, soldadas hasta 1.0-1.5 mm de
su base; el peci'olo ensanchdndose hacia la ldmina
y poco diferenciado, 4—6 mm de largo, glabro o con
escasos pelos simples en el envds; lamina obovada
o muy ampliamente obovada, 2.5-5.0 mm de largo
y 2.0-4.0 mm de ancho, glabra o con pocos pelos
simples en el enves; pelos glandulares (tentdculos)
en el centra y margen de la Mmina, hasta 2 mm de
largo. Escapo 16 2 por planta, 4—15 cm de largo.
Inflorescencia monocdsica, terminal. Flores (1—)2—6;
pedunculos 4—13 cm de largo, con pubescencia muy
variable a lo largo del mismo: en el extremo proximal
con glandulas sdsiles diminutas, pelos glandulares y
a veces glabros o pelos simples, mientras que en el
extremo terminal con gldndulas sesiles y pelos glan¬
dulares; brdcteas lanceoladas 2 mm de largo y 1 mm
de ancho, con pelos glandulares; pedicelos 0.5-1.0
mm de largo, con pelos glandulares. S6palos 5, lan-
ceolados, soldados hasta 1.5 mm de su base, 5.0 mm
de largo y 2.0-2.2 mm de ancho, acuminados, con
pelos glandulares, margen entero. P6talos 5, espa-
tulados, 6 mm de largo y 2 mm de ancho. Ovario
tricarpelar, 1-locular, rudimentos seminales nume-
rosos; estilo 3, bipartido, 2 mm de largo; estigmas
como un abanico. Fruto tipo capsula que se abre por
3 valvas. Semillas faveoladas, elfpticas, 100-150
gm de largo y 50-100 ptm de ancho, con abundan-
tes granos de cera en la superficie.

Drosera hirticalyx estd presente en la Guayana
Venezolana (Cerro Marahuaca) y crece en aflora-
mientos de arenisca entre 1200 y 2700 m sobre el
nivel del mar en lugares humedos.

Drosera hirticalyx es afin a D. roraimae (Klotzsch ex
Diels) Maguire & J. R. Laundon; esta ultima especie
dene las hojas nris laigas (10-20 mm), el pectolo muy
pubescente y claramente diferenciado de la lamina, el
numero de floras es mayor (4-20), los pelos glandu-
losos en el caliz son mds pequeiios (comparar Fig. ID
y F). Por uldmo la semilla es mas larga (400-600 pm)

Novon 5: 241-245. 1995.

242

Novon

Figura 1. A-E. Drosera hirticalyx R. Duno & Culham Steyermark & Holst 130701 (VEN). —A. Habito. —B. Hoja
cara superior. —C. Hoja cara inferior. —D. CSliz. —E. Semilla. F, G. Drosera roraimae (Klotzsch ex Diels) Maguire
& J. R. Laundon (Liesner 23990, VEN). —F. CAliz. —G. Semilla.

y tiene una forma angosto-oblonga con los extremos de
la testa inflados (comparar Fig. IE y G).

Pardtipos. VENEZUELA. Amazonas: Dpto. Atabapo,
open rocky plateau of Cerro Marahuaca above Salto los
Monos on tributary of headwaters of rfo Iguapo, 3°37'N,
65°23’W, 2555 m, near stream, flowers white, 26 Feb. 1985,
Liesner 17998 (MO); Cerro Marahuaca, summit on SE cor¬
ner, open rocky plateau and ravines, in small pool of water,
3°37'N, 65°21'W, 2700 m, 12 Oct. 1988, Liesner 24685
(MO); Cerro Marahuaca, summit on SE comer, open rocky
plateau and ravines, forming cushions on edge of pond
3°37'N, 65°21'W, 2700 m, 12 Oct. 1988, Liesner 24689
(MO); Cerro Marahuaca, summit, SW side of center, open
rocky plateau and ravines, on slope, buds pink, 3°39'N,
65°26’W, 2660 m, 24 Oct. 1988, Liesner 25286 (MO); Cerro
Marahuaca, summit on undulating plateau with island of
shrubs and Marahuacaeae with stream branch leading to
Salto de Monos and southeastern edge to Cano Iguapo, S

and SE of summit camp, on moist ground, growing with
130700, leaves dense, short, spreading, 337 3. 65°23 W,
2520-2650 m, 26-27 Feb. 1985, Steyermark & Holst
130701 (MO, VEN); Cerro Marahuaca, summit on undu¬
lating plateau with island of shrubs and Marahuacaeae with
stream branch leading to Salto de Monos and southeastern
edge to Cano Iguapo, S and SE of summit camp, petals
white, 3°37'N, 65°23'W, 2520-2650 m, 26-27 Feb. 1985,
Steyermark & Holst 130753 (MO, VEN).

Drosera yutajensis R. Duno & Culham, sp. nov.
TIPO: Venezuela. Amazonas; Dpto. Atures, val¬
ley of rfo Coro-Coro, W of Serrania de \utaje,
sandstone plateau W of river valley, along trick¬
ling stream in slightly depressed area, flowing
across exposed sandstone, 1100 m, 5 41 N,
66°09’W, Feb. 1987, Holst & Liesner 3277 (holo-
tipo, MO; isotipos, RNG, VEN). Figura 2.

Volume 5, Number 3
1995

Duno de Stefano & Culham
Drosera de Venezuela

243

Figura 2. Drosera yutajensis R. Duno & Culham (Huber 12636, VEN). -A. Hibito. -B. Hoja cara superior. -C.
Hoja cara inferior. —D. C4liz. —E. Semilla.

Speciebus Drosera villosa atque D. arenicola affinis. fo¬
lia lineari-spathulata 10—20 mm, distincte petiolata, pe-
tiolis dense pubescentibus. Inflorescentia 20—25 cm, 16-
30-flora; pedunculus glandulis raris praeditus. Petala 5,
spathulata, 4.5-5 X 2.5 mm; semina faveolata, ovoidea,
400-430 X 230-250 pm.

Planta herb&cea arrosetada. Rafces fibrosas. Ta-

Ho corto. Hojas linear-espatuladas, 10-20 mm de
largo; estipulas intrapeciolares, 5-6 mm de largo,
5-6 partidas, soldadas hasta 1 mm de su base;
pecfolo ensanch&ndose hacia la lamina pero bien
diferenciado, 5-7 mm de largo y 1-2 mm de ancho,
pelos simples en el enves; laminas oblanceoladas

244

Novon

u obovadas, 6-12 mm de largo y 2-4 mm de ancho,
pelos simples en el env6s, pelos glandulares (ten-
taculos) en centra y margen de la lamina, hasta 4.0
mm de largo. Escapos 1-3 por planta, 20-25 cm
de largo. Inflorescencia monoeasica, terminal. Flo¬
res 16-30; pedunculo 10—12 cm de largo, erecto
con pelos glandulares; braeteas lineares, 2 mm de
largo y 0.1 mm de ancho, con pelos glandulares
escasos; pedicelos 2-3 mm de largo, con pelos
glandulares. Sepalos 5, ovados, 3—5 mm de largo y
1 mm de ancho, con pelos glandulares en el env6s,
margen entero. Petalos 5, espatulados, 4.5—5.0 mm
de largo y 2.5 mm de ancho. Ovario tricarpelar, 1-
locular, rudimentos seminales numerosos; estilos 3,
bipartidos, 2 mm de largo; estigmas como un aban-
ico. Fruto tipo cApsula que se abre por 3 valvas.
Semillas faveoladas, ovadas, 400-430 pm de largo
y 230-250 pm de ancho.

Drosera yutajensis es solo conocida de tres co-
lecciones provenientes del cerro Yutaje, en la cuen-
ca del rio Manapiare. Crece en afloramientos de
arenisca entre 600 y 1800 m sobre el nivel del mar,
en lugares humedos.

Drosera yutajensis es afm a D. villosa A. Saint-
Hilaire (presente solo en Brasil), ambas con hojas
linear-espatuladas, pero mAs largas en la especie
brasilera (2-5 cm), donde adernas el pecfolo apenas
se diferencia de la ldmina; la semilla en este ultimo
caso es tambiAn faveolada pero angostamente ob-
longa y mucho rods larga (600-700 pm). Tambien
es afln a D. arenicola Steyermark, las hojas con la
misma forma y pubescencia pero mds pequefias en
D. arenicola (8—12 mm); en esta ultima el pedun¬
culo es mAs corto (1-8 cm), el numero de floras
menor (1-10) y la semilla es casi elfptica.

Pardtipos. VENEZUELA. Amazonas: Dpto. Atures,
Rto Coro-Coro, W of Serrania de Yutajg, 8.5 km N of
settlement of Yutaj6, small tributary, E of river, semi-ex¬
posed creek running over igneous fault horded by sand¬
stone, vegetation low, in sandy areas; 650 m, 5°42'N,
66°08’W, 25 Feb. 1987, Holst & Liesner 3179 (MO); Cum-
bre del cerro Yutaj6, sector oriental de la Serrania Yutaje,
herbazal inundable en planicie aluvial, bosquetes de pal-
mas al pie de las vertientes, arbustal y herbazal denso
sobre las vertientes rocosas, ocasionalmente en sitios
abiertos del arbustal y herbazal, 5°45'N, 66°03'0, 1800
m, 21 mar. 1989, Huber 12636 (MYF, PMA, VEN).

TambiAn fueron estudiadas las especies D. ca¬
yennensis Sagot ex Diels (1906), D. colombiana A.
FemAndez (1965) y D. panamensis M. D. Correa &
A. S. Taylor (1976). Estas especies comparten im-
portantes atributos como son: forma, tamano y pu¬
bescencia de las hojas; tipo (pelos simples) y den-
sidad de la pubescencia en el pedunculo y sepalos;
la condici6n de sepalos libras desde la base y el

tipo de dehiscencia de la capsula. A pesar de que
las semillas encontradas en todas las muestras eran
inmaduras, presentaban apariencias muy similares.
Todas estas similitudes sugieren que nos encontra-
mos en presencia de una sola especie. Previamente
Brummer-Dinger (1955) consider6 D. sanariapoana
Steyermark coespeclfica a D. cayennensis.

Drosera cayennensis Sagot ex Diels, Pflanzenfam.
4(112): 86. 1906. TIPO: French Guiana, Sagot
s.n. (P).

Drosera sanariapoana Steyermark, Fieldiana Bot. 28(2):
243. 1952. TIPO: Venezuela. Amazonas: collected
on wet savanna, vicinity of Sanariapo, near Rfo Sana-
riapo, tributary of Rfo Orinoco, 100 m, “petal rose,”
8 Sep. 1944, Steyermark 58472 (hol6tipo, NY).
Drosera colombiana A. Fernandez. Caldasia 9(43): 226,
fig. 1 1965. Sin. nov. TIPO: Colombia. Meta: Llanos
de San Martin, en sabana, “pelos finos rojos sobre
fonde verde, floras blancas,” jul. 1948, Dryander
3019 (holntipo. COL; isotipo. US).

Drosera panamensis M. D. Correa & A. S. Taylor, Ann.
Missouri Bot. Gard. 63: 390. 1976. Sin. nov. TIPO:
Panam4. La Yeguada, altos de Baltazar y el Veladero,
650 m, 4 ago. 1976, Correa et al. 2215 (holdtipo,
PMA; isotipo-. CHR no visto, COL, DUKE no visto,
F no visto, K, MEXU no visto, MO no visto, NSW
no visto, U no visto, US no visto).

Material adicional estudiado. GUAYANA FRANCESA:
Leprieuer 145 (parAtipo de D. cayennensis, M), Spruce
1068 (parAtipo de D. cayennensis, M). VENEZUELA.
Amazonas: Guanchez 1083 (MO, VEN).

Drosera esmeraldae (Steyermark) Maguire &
Wurdack fue conocida anteriormente solo de las
tierras bajas de Guayana (Sabana de Esmeralda,
Estado Amazonas). Material coleccionado reciente-
mente en Colombia (Vaupes y Amazonas) pertenece
a este tax6n.

Drosera esmeraldae (Steyermark) Maguire & Wur¬
dack, Mem. New York Bot. Gard. 9(3): 335.
1957. Drosera tenella var. esmeraldae Steyer¬
mark, Fieldiana Bot., 28(2): 244. 1952. TIPO:
Venezuela. Amazonas: wet places in savanna,
between Esmeralda savanna and southeastern
base of Cerro Duida, 200 m, 22 Aug. 1944,
Steyermark 60920, (holdtipo, F; isdtipo, NY).

Material adicional estudiado. COLOMBIA. Vaupes:
Schultes 22610 (COL). Amazonas: Idrobo 8970 (COL),
Sastre & Reichel 5209 (COL). VENEZUELA. Amazonas:
Clark 6475 (NY), Holst & Liesner 3275 (MO), Liesner &
Holst 21800 (MO), Huber 2697 (MO), Maguire et al.
29616 (BM, FM, NY), Stergios & Aymard 4277 (PORT).

En relacion con Drosera communis A. Saint-Hi-
laire, Maguire & Wurdack (1957) indican la pre¬
sencia de esta especie para Venezuela; sin embargo
no incluyen lista de exsiccatas. Los pocos espect-

Volume 5, Number 3
1995

Duno de Stefano & Culham
Drosera de Venezuela

245

raenes recolectados posteriormente en Venezuela e
identificados por J. A. Steyermark: (Krai 72009
(VEN), Steyermark 128551 (MO, VEN) y Steyer¬
mark 128978-A (MO, VEN)) difieren claramente
del material tipo de esta especie: Saint-Hilaire 695
(holotipo, P); estas muestras fueron identificadas
como D. roraimae (Klotzsch ex Diels) Maguire & J.
R. Laundon. Por lo tanto consideramos que esta
espeeie no esta presente en Venezuela ni Colombia
y su Area de distribucion queda limitada al hemis-
ferio Sur (Brasil meridional y Uruguay).

Drosera communis A. Saint-Hilaire, PI. Usual. Bras.
XV plate 15. 1824. TIPO: Brasil. Minas Ger¬
ais: Saint Hilaire 695 (holotipo. P).

Drosera parvifolia A. Saint-Hilaire, Hist. PL Remarq. Bre-
sil, p. 263, t. 25. 1824. TIPO: Brasil. Minas Gerais:
Saint Hilaire D377 (hol6tipo, P).

Drosera communis var. pauciflora Eichler en Martius, FI.
Bras., 14: 394. 1872. TIPO: B.

Material adicional estudiado. BRASIL. Minas Ge¬

rais: Glaziou 10258 (P). Rio de Janeiro: Glaziou 14483
(P). Sao Paulo: Saint Hilaire D 722 (P), Gardner 3573
(K), Luetzelburg 21485 (M), Luetzelburg 21509 (M). PA¬
RAGUAY: Hassler 5271 (K, P), Hassler 5634 (K, P).

Agradecimientos. Expresamos nuestro mds sin¬
cere agradecimiento a los curadores de los siguien-
tes herbarios: B, BM. COL. K, F. M. MA. MO. MYF,
P. PORT. US. Asf como a J. FemAndez-Casas (MA)
por sus comentarios y descripciones en latin. tam-
bi£n a Otto Huber, Paul Berry y Bruce Holst por
la correcion del manuscrito.

Literatura Citada

Brummer-Dinger, C. H. 1955. Notes on Guiana Drosera-
ceae. Acta Bot. Neerl. 4(1): 136-138.

Correa, M. D. & A. S. Taylor B. 1976. Droseraceae. Ann.

Missouri Bot. Gard. 63: 389—392.

Diels, L. 1906. Droseraceae. En: Engler, Das Pflanzen-
reich 26 (4): 1-136.

Femfindez-PArez, A. 1965. Plantas Insectfvoras, II Dro¬
seraceae de Colombia. Caldasia 11(43): 219-232.
Maguire, B. & J. J. Wurdack. 1957. The Botany of the
Guayana Highland—Part II. Mem. New York Bot. Gard.
9(3): 331-336.

Palisota flagelliflora (Commelinaceae), a New Species from
Cameroon with a Unique Habit

Robert B. Faden

Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Abstract. Palisota flagelliflora Faden (Comme¬
linaceae), a rosette species, is unique in Palisota
in bearing its inflorescences on axillary, flagelliform
shoots that lie on the forest floor. These shoots con¬
tinue to grow sympodially and give rise to addi¬
tional inflorescences or viviparously produce a new
rosette. Other rare or unique features include: in¬
florescences composed of single cincinni, long ped¬
icels, often vertical flowers, yellow staminode fila¬
ment hairs, bearded filament and connective in the
unpaired stamen, and bearded style.

The genus Palisota Reichenbach f. ex Endlicher,
with ca. 18 species (Faden & Hunt, 1991), is the
largest African genus of Commelinaceae occurring
predominantly in forests (Faden, 1983). Its center
of diversity is Central Africa, especially Cameroon
and Gabon. Until recently, all described species
were either acaulescent rosette plants or herba¬
ceous shrubs, and all had terminal or terminal and
axillary, thyrsiform inflorescences. Recently de¬
scribed species (Brenan, 1984) have habits and in¬
florescences that depart from these types, including
a decumbent habit and strictly axillary inflores¬
cences. A unique habit, rosettes producing flagel¬
liform flowering shoots, is described below in the
new species P. flagelliflora.

Palisota flagelliflora Faden, sp. nov. TYPE: Cam¬
eroon. South Prov. [South-Central Prov.]: Kri-
bi-Lolodorf road 3.5 km along the road to Edca
that branches off to the N at km 18 (just past
Bissiang), 200 m on a track that branches off
the Bissiang-Edea road, 3°02'N, 10°02'E, ex¬
ploited forest with much of the structure and
understory vegetation remaining, nearby pure
stand of Cecropia ( Musanga ) cecropioides along
the Bissiang-Edea road, 29 Jan. 1986, Faden
& Mbamba 86/50 (holotype, US sheet no.
3264040; isotypes, B, BR, C, COI, G, GC,
HBG, IEC, K(3), LISC, MA, MO, P, S, UPS,
US(4), WAG, YA(2)). Field-collected inflores¬
cences and fruits, preserved in FAA (US vial
B470), are part of the type collection. Figures
1 , 2 .

Herbae perennes foliis rosulatis, 14-50 cm longis, 3.5—
10(—11) cm latis, lamina elliptica vel oblongo-elliptica ad
oblanceolatam vel obovatam, 10-40 cm longa; inflores-
centiis terminalibus et bracteas oppositis in surculis ax-
illaribus, flagelliformibus (1.5—)2S—100(—175) cm longis,
cincinnis solitariis et contractis, bracteis binatis subtentis
compositis; floribus bisexualibus et masculis, stamine im-
pari apice filamenti et in connective barbato, sacculo an-
therae in flore bisexuali indehiscenti, in flore masculo de-
hiscenti, ovulis biseriatis; fructibus triangulari-ellipsoideis
ad triangulari-ovoideos, apice acuto ad rostratum, 1.4—2.3
cm longis, 0.9—1.2 cm latis, coccineis vel rubiginosis,
dense pilosis; seminibus uni- vel biseriatis.

Perennial with a rhizome to 15 cm long, fre¬
quently producing offsets, leaves all or mainly in
rosettes, petiolate, 14—50 cm long, 3.5-10(-ll) cm
wide, petiole 4—19 cm long, appressed pubescent,
lamina elliptic or oblong-elliptic to oblanceolate or
obovate, 10-40 cm long, apex abruptly acute or
acuminate, base cuneate to attenuate, surfaces dis-
colorous, adaxial green, glabrous or appressed
whitish sericeous and glabrescent (this pubescence
± persistent in cultivation), abaxial persistently ap¬
pressed whitish sericeous. Inflorescences terminal,
usually becoming bract-opposed, pedunculate,
borne on axillary, leafless, bracteate, flagelliform
shoots (1.5-)25-100(-175) cm long, at first ascend¬
ing, soon becoming prostrate and creeping through
the leaf litter, rarely obviously branched, rarely
rooting, many-noded, extending sympodially from
the axil of the bract subtending the peduncle, the
flagelliform shoot in this manner often producing a
succession of terminal/bract-opposed inflores¬
cences, often viviparously producing a new rosette
distally; intemodes appressed sericeous, nodes cov¬
ered by persistent maroon bracts 1.5-4 cm long,
appressed whitish sericeous; peduncle 1-8.5 cm
long, usually appressed sericeous, sometimes
densely lanate; each inflorescence consisting ol a
single contracted, several-flowered cincinnus sub¬
tended by a pair of maroon bracts 1.5-2.5 cm long,
similar in shape, color, and pubescence to those of
the flagelliform shoot. Flowers bisexual and male,
predominantly vertical or with the unpaired (outer)
sepal uppermost, occasionally horizontal, ca. lb-
18 mm wide, odorless; pedicels erect in flower, be-

Novon 5: 246-251. 1995.

Volume 5, Number 3
1995

Faden

Palisota flagelliflora

247

Figure 1. Palisota flagelliflora Faden. —A. Habit, showing flagelliform flowering shoot. —B. Inflorescence terminal
on flagelliform flowering shoot, the shoot growth continuing sympodially from the axil of the bract at the base of the
peduncle. —C. Unpaired stamen, bisexual flower, ventral view. —D. Unpaired stamen, bisexual flower, dorsal view. -
E. Paired stamen, dorsal view. —F. Paired staminode. —G. Staminode filament hairs. —H. Gynoecium, lateral view.
—I. Fruit, showing contorted pedicel. —j. Seeds, dorsal and ventral views. All from Faden & Mbamba 86/50.

248

Novon

f r, re 2 ‘ Pa l“ 0 ^ l fl a Z dl f ora Fade »- —A. Male flower, front view. —B. Male flower, side view. —C. Bisexual flower,
front View. —D. Bisexual flower, side view. All from Faden & Mamba 86/50.

coming spirally contorted in fruit, 1.5-3.5 cm long,
white, sometimes tinged with reddish purple, pi¬
lose; sepals subequal, boat-shaped, lanceolate-ob¬
long to oblong-elliptic, (3-)6-7.5 mm long, 2-3 mm
wide, reddish purple outside, the color shining

through, densely hirsute with reddish hairs forming
an apical tuft, usually intermixed with finer, shorter,
white hairs; petals subequal, outer slightly broader
than the others, elliptic, (4—)6 mm long, (2—)3—3.5
mm wide, white, inner petals oblong to spathulate-

Volume 5, Number 3
1995

Faden

Palisota flagelliflora

249

oblong, (4—)6.5—7 mm long, (1.5-)2-2.5 mm wide,
usually white, sometimes maroon with white mar¬
gins; unpaired staminode lacking or with a filament
1.5—2 mm long and bearded subapically with ap¬
parently moniliform, white-based yellow hairs;
paired staminodes with filaments 2—3 mm long,
white, bearded in the distal half with hairs like
those of the outer staminode; paired stamens with
filaments (2-)3—3.5 mm long, glabrous, white, an¬
thers broadly elliptic to oblong-elliptic or ovate-el¬
liptic, (1—)1.5—1.75 mm long, 0.7—1 mm wide, de¬
hiscence extrorse, connective yellow, glabrous,
pollen yellow; unpaired stamen dimorphic, in bi¬
sexual flower slightly shorter to slightly longer than
the paired stamens, filament (1.5—)2.5-3.5 mm
long, strongly declinate, white, usually densely
bearded with minute yellow hairs just below the
anther, anther broadly ovoid, ca. (1—)1.8 mm long
and wide, yellow, pollen sacs C-shaped, indehis-
cent, connective usually densely bearded with min¬
ute yellow hairs; in male flower unpaired stamen
distinctly longer than the paired, more central in
the flower, usually not declinate, filament ca. 5 mm
long, white, bearded at apex, as in bisexual flower,
anther similar to that of bisexual flower, including
densely bearded connective, but smaller (ca. 1.4
mm long and wide) and pollen sacs dehiscent, in-
trorse, with pollen concolorous with that of the
paired stamens; gynoecium reduced and apparently
abortive in the male flower, in the bisexual: ovary
ovoid, deeply 3-lobed, ca. 2-2.5 mm long, densely
pubescent with forward-directed, white or reddish
hairs, locules 5-7-ovulate, ovules biseriate, style
exceeding the stamens, 3.2—4 mm long, straight or
declinate, yellow, sparsely bearded basally with ap-
pressed hairs like those of the ovary, stigma deeply
3-lobed, yellow with a central brownish area. Fruits
triangular-ellipsoid to triangular-ovoid berries,
acute to beaked at the apex, 1.4—2.3 cm long, 0.9—
1.2 cm wide, scarlet or reddish brown, densely red¬
dish pilose. Seeds uni- or biseriate, 1—4 per locule,
ovate-triangular in outline, ca. 4—5 mm diam.

Habitat. Secondary and exploited forest, 10—
520 m.

Flowering. January, February, July, September.
In cultivation the flowers open at about 0940 and
begin to fade at 1230-1330.

Distribution. Endemic to Cameroon.

Palisota flagelliflora is unique in Palisota be¬
cause of its flagelliform shoots, bract-opposed inflo¬
rescences, and vivipary. It may further be distin¬
guished by its inflorescences composed of a single
cincinnus, long pedicels, large, often vertical flow¬
ers, yellow staminode filament hairs, bearded sta¬

men filament and connective in the unpaired sta¬
men, and bearded style. It also has the earliest
flower opening time for any Palisota species thus
far recorded or observed.

In the above description the dimensions of the
floral parts were measured from preserved flowers
of a cultivated plant of Faden & Mbamba 86/50.
The much smaller dimensions for some parts, listed
parenthetically, came from a pressed flower on a
small plant of Leeuivenberg 8296 (WAG). No field-
preserved flowers were available. The larger flowers
were the largest observed in Palisota.

The staminode filament hairs were described
above as “apparently moniliform” because of the
beadlike appearance of the cells at low magnifica¬
tion. However, higher magnification (Fig. 1G) re¬
vealed that the cells of the hairs were individually
dumbbell-shaped, not spherical or ellipsoid, as in
typical moniliform hairs.

The nature of the flagelliform shoots requires
some comment. Morphologically they could be con¬
sidered one of the following: long, axillary inflores¬
cences; axillary shoots bearing terminal inflores¬
cences; inflorescence-shoots; or stolons bearing
inflorescences. Their actual structure is clear, so
the problem is more semantic than morphological.
Forman (1962) tried to distinguish between an ax¬
illary inflorescence and a reduced axillary shoot
bearing a terminal inflorescence, which he termed
an inflorescence-shoot. Faden (1991: 11) found the
difference between the two, which was based on the
absence (inflorescence) or presence (inflorescence-
shoot) of a basal prophyll, not very useful because
the whole range from leafy lateral shoots with a
terminal inflorescence to reduced lateral shoots
with only bracts (inflorescence-shoot) could be
found within a single plant in some Aneilema spe¬
cies. The axillary inflorescence might have evolved
by further reduction, namely loss of the prophyll.

Many forest Commelinaceae produce stolons: ax¬
illary, elongate, horizontal shoots with only leaf
sheaths basally (Faden, 1988). Distally stolons pro¬
duce new vegetative shoots, not inflorescences. The
inflorescence-terminated shoots of Palisota flagel¬
liflora are dissimilar because they bear only bracts
and they are primarily for sexual, not vegetative,
reproduction. When they viviparously produce a
new plant, they are more like the reproductive
shoots of the epiphytic Belosynapsis vivipara (Dal-
zell) C. E. C. Fischer than like stolons of other
terrestrial species. Because they are unique in Pal¬
isota, and perhaps within the family, I have used
the term “flagelliform shoots” to describe them.

The regularly biseriate ovules were a surprising
find in view of the fact that the seeds may be uni-

250

Novon

seriate. The alignment of the seeds in the fruit
seems to be a function, at least partly, of seed num¬
ber per locule. When three or fewer the seeds ap¬
pear uniseriate, when four or more, biseriate. Only
two fruits were available for dissection, so which of
these arrangements predominates and whether
more seeds are ever present is unknown.

One of the two sheets of Leeuwenberg 8296
(WAG) bears two small plants each with a short
inflorescence (only 1.5 cm long in one case). Had
these been the sole plants collected, treating them
as Palisota flagelliflora would have had to have
been based largely on the very long pedicels and
bearded connective of the unpaired stamen. The
second sheet, as well as specimens of this collec¬
tion seen elsewhere, clearly demonstrate that the
small plants are merely young P. flagelliflora with
short inflorescences.

Faden & Mbamba 86/47 was noted in the field
as differing from Faden & Mbamba 86/50 in not
forming dense stands, in being sterile (it had pro¬
duced new rosettes at the ends of axillary, flagel-
liform shoots), less robust, with a reddish petiole
and smaller, less discolorous lamina more abruptly
narrowed into the petiole, and with smaller bracts
on the flagelliform shoots. Although exact chromo¬
some counts were not obtained from either popu¬
lation, 2 n = ca. 40 (approximately diploid) was
counted for Faden & Mbamba 86/47 and 2n = ca.
60 for Faden & Mbamba 86/50 (approximately trip-
loid). Faden & Mbamba 86/50 was selected as the
type, despite its being possibly triploid, because
abundant reproductive material was present in the
field, including ripe fruits and buds, and that pop¬
ulation also flowered in cultivation, whereas Faden
& Mbamba 86/47 did not flower before it died.

The inclusion of Bos 3877 in Palisota flagelliflo¬
ra was questionable. Although it came from very
close to the type locality, it differed most obviously
from the other collections by its broader leaves,
more robust flowering shoots that perhaps were not
lying in the leaf litter, peduncles completely cov¬
ered by lanate, brownish hairs, and by lacking ster¬
ile bracts between the bracts at the bases of the
peduncles. Dissection of flower buds, nearly all
partly eaten or bored, revealed that all three stamen
filaments and connectives were glabrous, differing
from all other collections of P. flagelliflora. In the
absence of a larger number of collections, Bos 3877
is provisionally included in P. flagelliflora.

Bos 3877 approached Palisota lagopus Mild-
braed in robustness and peduncle pubescence, dif¬
fering most obviously by its longer, axillary, sym-
podial flowering shoot with several inflorescences
and longer, spirally contorted fruiting pedicels. Its

somewhat intermediacy between this species and P.
flagelliflora suggested a possible hybrid origin, but
it could just as readily represent an undescribed
species or a form of P. flagelliflora.

Palisota flagelliflora is an understory species
that has been collected in secondary and exploited
primary forest. It should also occur in undisturbed
primary forest. In Littoral Province the primary veg¬
etation at the locality for this species would be “At¬
lantic forest of the northwestern type with Caesal-
piniaceae rare” (Letouzey, 1985). At the localities
east of Kribi the primary forest type would be “At¬
lantic littoral forest with Caesalpiniaceae relatively
rare, with Saccoglottis gabonensis ” (Letouzey,
1985).

In the field Palisota flagelliflora may form dense
stands because of its ability to reproduce vegeta-
tively. That tendency, along with the small size of
the rosettes, compared to other species, allows it to
be spotted from a distance. Closer up, the flagelli¬
form shoots and vivipary are very distinctive, as is
the presence of hairs on the connective and fila¬
ment apex of the unpaired stamen.

Palisota flagelliflora appears related to the red-
fruited species of Palisota because of its fruit color,
hairy fruits (glabrous in some species), persistent
faded flowers, and long, spirally contorted fruiting
pedicels. Most of those species, such as P. barteri
Hooker and P. mannii C. B. Clarke, have uniseriate
seeds and would thereby belong to section Monos-
tichos C. B. Clarke. Palisota flagelliflora, however,
with its biseriate ovules and sometimes biseriate
seeds, would have to be placed in section Palisota
(synonym: section Distichos C. B. Clarke, 1881:
133) with the blue- and black-fruited species such
as P. ambigua (P. Beauvois) C. B. Clarke and P.
hirsuta (Thunberg) K. Schumann. This apparent
anomaly may be explained either by Clarke’s sec¬
tions being unnatural, which well may be the case,
or by seed arrangement not always reflecting ovule
arrangement, about which we have insufficient
data.

Palisota flagelliflora is probably most closely re¬
lated to P. satabiei Brenan and P. bogneri Brenan,
small, red-fruited species with axillary inflores¬
cences, biseriate ovules, and, like P. flagelliflora, a
restricted distribution in the species-rich coastal
forests of Cameroon and Gabon. Palisota flagelli-
fora uniquely shares with P. satabiei the features of
the inflorescence consisting of a single cincinnus,
yellow staminode filament hairs, bearded anther
connective, and bearded style. Preserved fruits of
P. satabiei (Satabid & Letouzey 384, YA) showed
partly biseriate seeds in one locule, as in P. fla¬
gelliflora.

Volume 5, Number 3
1995

Faden

Patisota flagelliflora

251

Palisota flagelliflora, P. satabiei, and P. bogneri
extend the range of habit and inflorescence types
previously known in Palisota : decumbent habit (P.
satabiei), vivipary, strictly axillary inflorescences,
inflorescences composed of single cincinni, and fla-
gelliform flowering shoots. My research on the
Commelinaceae of Cameroon has revealed addi¬
tional habit types in the genus: scandent (to 15 m
in length) in P. thollonii Hua, and decumbent with
a terminal thyrse in an undescribed species from
Cameroon and Gabon. The diversity of habit types
now known within Palisota rivals that in the much
larger neotropical genus Dichorisandra, which is
the ecological counterpart of Palisota in South
America.

It should be noted that the bisexual flower de¬
picted in Figure 2 has the lower (unpaired) stamen
longer and less declinate than in the most extreme
cases. Insufficient flowers have been examined to
generalize about the “typical” curvature and length
of the filament.

Paratypes. CAMEROON. Littoral Prov.: Foret du
Bukaka, 3 km E of kbcnr. a village on km 11 Nkongsam-
ba-Loum road, 4°50’N, 9°54'E, alt. 520 m, 8 Sep. 1971,
Leeuwenberg 8296 (US, WAG—2 sheets, YA). South
Prov.: ca. 18 km from Kribi, Lolodorf road, 3°00'N,
10°02'E, 5 Feb. 1969, Bos 3877 (WAG, YA); 4 km N of
km 20 Kribi-Lolodorf, 3°02'N, 10°03'E, 4 July 1970, Bos
7014 (US, WAG—3 sheets, YA); Kribi-Lolodorf road, km
20, end of track ca. 2 km N of the road, ca. 3°01'N,

10°03'E, alt. 10 m, 29 Jan. 1986, Faden & Mbamba 86/
47 (P, US, YA).

Acknowledgments. I thank B. Satabid for facil¬
itating my fieldwork in Cameroon, Alice Tangerini
for the line drawings, and my wife, Audrey J. Fa¬
den, and other volunteers for taking care of my re¬
search plants for many years.

Literature Cited

Brenan, J. P. M. 1984. Two new species of Palisota (Com¬
melinaceae) from West Africa. Kew Bull. 39: 829-832.
Clarke, C. B. 1881. Commelinaceae. Pp. 113—324 in A.
& C. de Candolle, Monographiae Phanerogamarum 3.
Sumptibus G. Masson, Paris.

Faden, R. B. 1983. Phytogeography of African Comme¬
linaceae. Bothalia 14: 553—557.

-. 1988. Vegetative and reproductive features of

forest and nonforest genera of African Commelinaceae.
Pp. 521—526 in: P. Goldblatt & P. P. Lowry II (editors).
Modem Systematic Studies in African Botany. Monogr.
Syst. Bot. Missouri Bot. Gard. 25.

-. 1991. The morphology and taxonomy of Aneile-

ma R. Brown (Commelinaceae). Smithsonian Contr. Bot.
76: 1-166.

-& D. R. Hunt. 1991. The classification of the

Commelinaceae. Taxon 40: 19—31.

Forman, L. L. 1962. Aetheolirion, a new genus of Com¬
melinaceae from Thailand, with notes on allied genera.
Kew Bull. 16: 209-221.

Letouzey, R. 1985. Notice de la carte phytog£ographique
du Cameroun: 1:500,000. Institut de la Recherche
Agronomique (Herbier National), Yaounde, and Institut
de la Carte Internationale de la Vigdtation, Toulouse.

Typification of Pinus apulcensis Lindley (Pinaceae), a
Misinterpreted Name for a Latin American Pine

Alps Farjon

Oxford Forestry Institute, Department of Plant Sciences, University of Oxford,
South Parks Road, Oxford 0X1 3RB. England, U.K.

Abstract. The Mexican taxon Pinus apulcensis
Lindley is lectotypified with an ovuliferous cone
found among carpological material in the herbar¬
ium of the Natural History Museum in Vienna (W),
which was sent to Endlicher by Lindley. This cone
confirms treatments of that name in publications
on Mexican pines up to 1945—1948, when Marti¬
nez interpreted it in a different sense, using a
specimen he had received from the putative type
locality. The resulting confusion has been cor¬
rected, and a revised synonymy of Lindley’s taxon
is presented.

During work being carried out to complete the
monographic study of pines for Flora Neotropica
started by Brian T. Styles, a problem was encoun¬
tered regarding the current interpretation of one of
John Lindley’s species from Mexico, Pinus apulcen¬
sis. This interpretation, commonly accepted since
Martinez (1945, 1948), and followed by Stead &
Styles (1984) in their revision of the “ pseudostro-
bus" group, was due to the lack of a type specimen,
and was based on what may be described as the
topotype method: the assumption that whatever
pine grows today at the type locality must be Lin¬
dley’s taxon.

Lindley (1839) published two new species of
pine based on collections made by C. T. Hartweg
in Mexico in 1838: Pinus pseudostrobus from An-
gangueo (“Anganguco,” Michoacdn) and P. apul¬
censis from “ravines near Apulco” (Hidalgo). (The
confusion of Apulco with Acapulco seems to have
originated with Endlicher (1847: 154).) The origi¬
nal descriptions of both are concise. Apart from
short leaves and glaucous shoots, P. apulcensis is
said to differ from P. pseudostrobus mainly in its
cones, “covered closely with pyramidal elevations,
which are sometimes prolonged and contracted in
the middle, especially those near the points of the
cones.”

Lindley’s herbarium is largely at CGE; a few
types are also at BM and K (Stafleu & Cowan,
1981). A catalog of conifers (Diimmer, 1913) lists
what was in the Lindley herbarium (CGE) early in

Novon 5: 9r>9-95f> 1QQ5

the twentieth century. Neither of the two taxa with
which we are concerned here is mentioned in that
catalog. Searches for original material of Pinus
apulcensis in the above-mentioned herbaria were
unsuccessful (Stead & Styles, 1984).

Loudon (1842: 1014—1015) translated Lindley’s
diagnosis and repeated his English description of
Pinus apulcensis but stressed even more the “backs
of the scales. . .sometimes prolonged into a hook,
particularly those nearest the base and the point.”
He stated that the species was introduced in 1839,
“by cones sent home by Hartweg, from which many
plants have been raised.” And finally, he presented
figures 1899 and 1900, “from specimens sent home
by Hartweg.” Figure 1899 is a natural-size (12 cm
long) line drawing of a closed (green?) cone clearly
showing what is meant by “prolonged and contract¬
ed pyramidal elevations” (Lindley, 1839) and
“scales prolonged into a hook” (Loudon, 1842).
There can be little doubt that this drawing repre¬
sents both Hartweg’s original collection and Lin¬
dley’s P. apulcensis (Fig. 1).

Shaw (1909) reduced Pinus apulcensis Lindley to
P. pseudostrobus var. apulcensis, citing the basio-
nym, Loudon’s (1842) figure 1899, Endlicher
(1847: 153), and three Mexican collections: Nelson
985 (US) and Nelson 2539 (US), both from Oaxaca,
and Pringle 8788 (MEXU, US) from Mexico, as his
references. In his plate 12, figures 8, 7, and 6 rep¬
resent cones of these collections respectively. Like
Lindley and Loudon, Shaw emphasized the “pe¬
culiar development of the apophyses of the cone,
which may attain remarkable prominence.” These
three collections present considerable variation in
development of the apophysis, with Nelson 2539
closely resembling figure 1899 in Loudon (1842),
and Nelson 985 attaining much longer apophysal
elongations and very strongly developed umbos.
The present study, based on many more collections,
demonstrates that Nelson 2539 and Nelson 985 rep¬
resent the two extremes of a cline in these char¬
acters. Pringle 8788 has strongly developed umbos,
but the apophyses of the scales are flat and the

Volume 5, Number 3
1995

Farjon

Pinus apulcensis

253

1914 ARBORETUM ET FRUTICETUM BRITANNICUM.

t 48. P. apulce'nsis Lindl. The Apulco Pine.

Identification. Lindl. in Bot. Reg. M. Cbron., 1039, No. 100.

Synonyme. P. acapulcensis G. Don in Sweet's Hort. Brit. ed. 3. p. 769.
Engravings. Our figs. 1899,1900. from specimens sent home by Hartweg-

5 cm
(cone)

Figure 1. Illustration of Pinus apulcensis Lindley (Fig. 1899) in Loudon’s An Encyclopaedia of Trees and Shrubs; Being
the Arboretum et Fruticetum Abridged (1842); drawn “from specimens sent home by Hartweg.”

scales rather thin, which places it outside this
cline.

Martinez (1945: 192; see also 1948: 199) refuted
Shaw’s inclusion of the two Nelson cones, in the
belief that the specimens from Oaxaca were not

conspecific with those from Apulco: “The speci¬
mens from Apulco and surrounding areas lack the
characteristic elongations seen in specimens found
further south, that is, in Veracruz and Oaxaca”
[translated from Spanish]. He observed that Lindley

254

Novon

did not mention these elongations (“prolonga-
ciones,” meaning the elongated apophyses) but only
“a recurved tip,” which sounds like an attempt to
bend Lindley’s description into shape with what
Martinez saw from Apulco. His subsequent “new”
combination Pinas pseudostrobus Lindley var. apul-
censis Martinez is superfluous because it repeats
Shaw (1909) by including Lindley’s type, the iden¬
tity of which was totally unknown to Martinez. The
specimens he referred to (e.g., Martinez 3434 from
Apulco, Hidalgo “var. nov.” MEXU; Martinez,
1948: 201, f. 162) indeed lack any pronounced
elongation of apophyses and/or umbos. However,
they do not represent Lindley’s species, but Mar¬
tinez’s interpretation of it, based on the cited lo¬
cality. Collected around 1939, almost exactly a cen¬
tury after Hartweg made his collections, they are
not necessarily from the same location. North of the
city of Tulancingo there are two localities with the
name Apulco, one at the railwayhead and another
ca. 4 km further down a road. To the east of these
are mountains (Cerro Viego) and streams (canyons)
with pine forests. This constitutes a large area, and
it is impossible to know exactly where Hartweg
made his collection.

Given the depletion of pine forests in many parts
of Mexico in the last 100 years, it may well be that
the trees Hartweg saw and collected are no longer
there.

For the collections from Oaxaca that Shaw had
cited under Pinus pseudostrobus var. apulcensis
(Nelson 985 and Nelson 2359), Martinez (1945:
195; see also 1948: 202) proposed a new combi¬
nation, P. pseudostrobus var. oaxacana, citing in
synonymy P pseudostrobus var. apulcensis (Lindley)
Shaw. This was an error: he clearly meant to ex¬
clude Lindley s type (Mirov, 1958) and to describe
a new variety based on the two Nelson collections.
Further, the name was not validly published be¬
cause he failed to provide a Latin diagnosis. Mirov
(1958), referring to the specimens cited by Shaw
(1909) and the description given by Martinez
(1945), and excluding P. apulcensis Lindley, de¬
scribed it as a new species: P. oaxacana Mirov.
Harrison (1965) then validated Martinez’s earlier
combination under P. pseudostrobus at the rank of
variety.

Martinez, in the absence of Lindley’s type and
with no easy access to European herbaria or nine¬
teenth-century botanical literature (he studied
these pines during the Second World War), relied
on the topotype method to establish the identity
of Pinus apulcensis Lindley. Understandable as
this may be in the case of Martinez, given the
circumstances under which he worked, it is re¬

markable that later authors (Loock, 1950; Mirov,
1958; Harrison, 1965; Stead, 1983; Stead &
Styles, 1984) simply accepted his judgment and
never pursued the identity of Lindley’s name by
typification.

Stead (1983) undertook a numerical study of
variation in which one of his collecting sites (No.
14) was the putative type locality of Pinus apul¬
censis Lindley at Apulco, Hidalgo. In his Principal
Component Analysis (PCA), the Apulco material
(from 25 trees) groups with sites 13 (P. pseudo¬
strobus var. estevezii Martinez) and 15 (P. pseudo¬
strobus var. coatepecensis Martinez) on character
states such as “increasing cone size, needle width
and stomatal lines,” but without any clear discon¬
tinuity. Cone size also increased for site 8 (rep¬
resenting P. pseudostrobus var. oaxacana (Mirov)
Harrison), but no other characters, including its
“very distinctive cones” would separate out this
taxon.

The conclusions outlined in Stead (1983) were
adopted by Stead & Styles (1984), who identified
the material from sites 13-15 as Pinus pseudostro¬
bus subsp. apulcensis (Lindley) Stead. The two va¬
rieties of Martinez (from sites 13 and 15) were cited
as synonyms, as well as P. pseudostrobus var. apul¬
censis “sensu Shaw pro parte” (which, as can be
understood from the synonymy under the previous
taxon treated in their revision, means as to type,
not to specimens cited). The type, however, was
cited as “Apulco, Hartweg s.n. (not seen).” (Lindley
(1839) did not indicate any lack of a collection
number and only mentioned that it was “found by
Mr. Hartweg”).

It is, of course, the identity of Pinus apulcensis
Lindley that needs to be established before adopt¬
ing this epithet. From the above it is obvious that
an ovuliferous cone is needed; there is general
agreement that foliage characters within Pinus
pseudostrobus s.l. show considerable overlap and
that leaves alone are inadequate for certain deter¬
mination (Stead & Styles, 1984; Perry. 1991; Car-
vajal & McVaugh, 1992). Lindley, when he was
Secretary of the Horticultural Society of London,
distributed material collected by Hartweg to various
botanists in Europe, among whom were Parlatore in
Florence and Endlicher in Vienna. On a recent visit
to Vienna (W), a number of conifer cones of Hart-
weg’s gatherings were found. Among them are two
cones of a Mexican pine with Lindley’s labels: P-
apulcensis Lindley” and a vial with seeds and
scales. The cones are 9.5 X 6 cm (closed) and 10
X 8 cm (open) and strongly resemble figure 1899
in Loudon (1842). They are original material and

Volume 5, Number 3
1995

Farjon

Pinus apulcensis

255

5 cm

Figure 2. Closed ovuliferous cone of Pinus apulcensis
Lindley, with a label written by John Lindley; it is part of
the lectotype in the carpological collection of the Natural
History Museum in Vienna (W).

are here designated as the lectotype of Pinus apul¬
censis Lindley (Fig. 2).

The synonymy given here pertains only to Pinus
apulcensis Lindley; no full synonymy is given for P.
pseudostrobus Lindley.

Pinus pseudostrobus Lindley var. apulcensis
(Lindley) Shaw, Publ. Arnold Arbor. 1: 19, t.
12, f. 6—8. 1909. Pinus apulcensis Lindley, Ed¬
wards’s Bot. Reg. 25: 63. Aug. 1839 [& Allg.
Gartenzeitung 7: 325. 1839]. Pinus pseudostro¬
bus subsp. apulcensis (Lindley) Stead, Bot. J.
Linn. Soc. 89; 269. 1984. TYPE: Mexico. Hi¬
dalgo: ravines near Apulco, C. T. Hartweg s.n.,
ex Hort. Soc. London, 2 ovuliferous cones with
labels: P. apulcensis Lindley in Lindley’s hand¬
writing, vial with seeds and scales (lectotype,
here designated, W).

Pinus pseudostrobus var. oaxacana Martfnez, Anales Inst.
Biol. Univ. Nac. Mexico 16: 1945, f. 164—168. 1945,
nom. inval.

Pinus oaxacana Mirov, Madrono 14: 145. 1958. Pinus
pseudostrobus var. oaxacana (Mirov) S. G. Harrison,

Taxon 14: 247. 1965. TYPE: Mexico. Oaxaca: La
Parada, E. W. Nelson 985 (holotype, US).

Pinus pseudostrobus Lindley is a widely distrib¬
uted species with polymorphic ovuliferous cones,
occurring from Sinaloa and Nuevo Le6n in Mexico
to Honduras (Stead & Styles, 1984). Pinus pseu¬
dostrobus var. apulcensis (Lindley) Shaw is a cone
variant distinguished by more or less clinal char¬
acters becoming more abundant in southern Mex¬
ico. It is usually sympatric with P. pseudostrobus
Lindley var. pseudostrobus but apparently absent
from large parts of the range of the species (Car-
vajal & McVaugh, 1992). Acceptance at any other
rank than variety therefore seems unwarranted for
this taxon. Current use of the epithet apulcensis in
the sense of Article 57 (formerly 69.4) of the Code
(Greuter et al., 1994) is only at infraspecific rank.
The name P. oaxacana is occasionally used, but
this taxon, too, is treated in most of the literature,
especially forestry literature, at infraspecific rank.
There is therefore only a weak case for a proposal
either to reject P apulcensis Lindley or to conserve
it with a conserved type, and I strongly prefer to
correct the errors made before I take up the name
in the sense of its type in Flora Neotropica.

Acknowledgments. Thanks are due to staff at
the Herbarium of the Museum of Natural History
in Vienna, in particular B. Wallnofer, who alerted
me to the existence of old carpological material in
W despite its presumed destruction during World
War II, and who granted a loan to Oxford (FHO) of
the relevant material.

Literature Cited

Carvajal, S. & R. McVaugh. 1992. Pinus L. Pp. 32-100
in: R. McVaugh, Flora Novo-Galiciana 17. Gymno-
sperms and Pteridophytes. Univ. Michigan Herbarium.
Ann Arbor.

Dfimmer, R. A. 1913. The conifers of the Lindley her¬
barium; Botany School, Cambridge. J. Roy. Hort. Soc.
39 (1): 63-91.

Endlicher, S. L. 1847. Synopsis Coniferarum. Scheitlin
& Zollikofer, Sangalli (Sankt Gallen).

Greuter. W„ F. R. Barrie. H. M. Burdet, W. G. Chaloner,
V. Demoulin, D. L. Hawksworth, P. M. Jorgensen. D. H.
Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors.
1994. International Code of Botanical Nomenclature
(The Tokyo Code), Adopted by the XVth International
Botanical Congress, Yokohama, August-September,
1993. Regnum Veg. 131.

Harrison, S. G. 1965. Note on gymnosperm nomencla¬
ture. Taxon 14: 247.

Lindley, J. 1839. Miscellaneous notices: Mexican pines.
Edwards’s Bot. Reg. 25: 63.

Loock, E. E. M. 1950. The pines of Mexico and British
Honduras. S. Africa Dept. Agric. and Forest. Bull. 35.
Loudon, J. C. 1842. An Encyclopaedia of Trees and
Shrubs; Being the Arboretum et Fruticetum abridged.
Published by the author, London.

256

Novon

Martinez, M. 1945. Las pinaceas mexicanas. Anales Inst.
Biol. Univ. Nac. Mexico 16: 1-345.

-. 1948. Los Pinos Mexicanos, ed. 2. Edicionas

Botas, Mexico City.

McVaugh, R. 1972. Botanical exploration in Nueva Ga¬
licia, Mexico, from 1790 to the present time. Contr.
Univ. Michigan Herb. 9: 205—357.

Mirov, N. T. 1958. Pinus oaxacana, a new species from
Mexico. Madrono 14: 145-150.

Perry, J. P. 1991. The Pines of Mexico and Central Amer¬
ica. Timber Press, Portland, Oregon.

Shaw, G. R. 1909. The pines of Mexico. Publ. Arnold
Arbor. 1.

Stafleu, F. A. & R. S. Cowan. 1981. Taxonomic Literature
3 [TL-2]. Regnum Veg. 105, Bohn, Scheltema & Hol-
kema, Utrecht.

Stead, J. W. 1983. Studies of variation in Central Amer¬
ican pines V: A numerical study of variation in the
Pseudostrobus group. Silvae Genet. 32: 101-114
[115].

-& B. T. Styles. 1984. Studies of Central Amer¬
ican pines: A revision of the “pseudostrobus” group
(Pinaceae). Bot. J. Linn. Soc. 89: 249—275.

Aureliana darcyi, a New Species of Solanaceae from Brazil

L. d’A. Freire de Carvalho and Massimo G. Bovini
Jardim Botanico do Rio de Janeiro, Segao de Botanica Sistematica, Rua Pacheco Leao 915,
CEP 22460-030, Rio de Janeiro, RJ, Brazil

Abstract. A new species of the genus Aureliana
from the coastal vegetation and secondary forest of
Paraty Municipality is described. It is distinguished
from other species in this genus by the long-acu¬
minate calyx lobes and sparse pubescence.

The genus Aureliana Sendtner was recently re¬
vised by Hunziker & Barboza (1991), who recog¬
nized eight taxa, all found in Brazil and concen¬
trated in the central-western, southeastern, and
southern regions.

During a floristic survey carried out by the Rio
de Janeiro Botanical Garden in the Cairugu Envi¬
ronmental Protection Area, Praty Municipality, Rio
de Janeiro state, a morphologically distinct species
of Aureliana was discovered.

Aureliana darcyi Carvalho & Bovini, sp. nov.
TYPE: Brazil. Rio de Janeiro: Paraty, Apa-
Cairugu, ilha das Almas, na orla da ilha, 15
Mar. 1989, A. L Almeida, B. Kurtz et al. 7
(holotype, RB; isotype, MO). Figures 1—7.

Arbor ca. 2—3 m alta; ramis dichotomis puberulis. Folia
solitaria, membranacea, glabrescentia, trichomatibus sim-
plicibus vestita. Inflorescentiae ex fasciculis 4—12-floris
constantes, receptaculo recto; pedicellis filiformibus apice
vix dilatato. Flos in alabastra globosus; calyce campanu-
lato, lobulis longe acuminatis aequalibus aestivatione lib-
eris; corolla viridi; staminum filamentis ca. 2.5 mm longis,
antheris ca. 1.5 mm longis. Bacca globosa; placenta ex-
pansa protuberationes formante; seminis testa reticulata.

Shrub or small tree ca. 2—3 m, branches dichot¬
omous with distinct longitudinal ribs, puberulent,
with simple hairs and lenticels. Leaves solitary,
membranaceous, dark green above and light green
underneath, 3.0-12.0 X 0.8-4.5 cm, lanceolate,
acuminate, base slightly decurrent, venation camp-
todromous, glabrate to pubescent, hairs simple; pet¬
iole flattened, 1.0-1.5 cm, puberulent. Inflores¬
cence 4—12-fasciculate, receptacles straight,

peduncle 0.3—1.2 cm. Flowers green, axillary. Ca¬
lyx whitish green, campanulate, ca. 3 mm, lobes
1.0-2.0 mm, regularly long-acuminate, glabrate ex¬
ternally. Corolla green, when mature white with
green macula, 1.7-1.9 cm, rotate-stellate; lobes
5.0-6.0 mm, lanceolate, reticulate-veiny, margins
ciliate. Pedicel 0.9—1.0 cm, filiform, slightly en¬
larged at the apex. Anthers light brown, ca. 1.5
mm, basifixed, filaments white, ca. 2.5 mm, filiform.
Ovary less than 1.0 mm, globose to ovoid; style ca.
1.0 mm, stigmatic region apical. Fruit green, glo¬
bose, ca. 1.0 cm diam., pericarp thick, placenta
with protuberances. Calyx not persistent in fruit, ca.
70 seeds, 0.2-0.3 mm long, oblong, testa reticulate.

This plant was growing in the sun near sea level
on an island (ilha das Almas) in the Atlantic Ocean.
At the time of collection, it appeared to be in full
flower with different stages of flowering present.
The type specimen was collected in secondary for¬
est, in full sunlight. It is hairier and the leaves are
wide-lanceolate.

The characters that distinguish A. darcyi from
other species of the genus are the long-acuminate
calyx lobes and the absence of indumentum on the
exterior of the calyx.

Aureliana darcyi is closely related to A. wettstein-
iana (Witasek) A. T. Hunziker & Barboza, as shown
by characteristics of the calyx lobes and flower bud.
Overall appearance plus the sparse pubescence
place it near A. fasciculata (Vellozo) Sendtner var.
fasciculata.

The specific epithet honors William G. D’Arcy,
eminent botanist from the Missouri Botanical Gar¬
den, for his important work on Solanaceae.

Paratypes. BRAZIL. Rio de Janeiro: caminho para
a Toca do Ouro, pr6x. 4 estrada Parati-Cunha, 03 Jul.
1992, L. C. Giordano, E. F. Guimaraes et al. 1312 (RB);
passando a 18 ponte entrar & direita (sentido RJ/SP) i ca.
de 13 km do trevo de Praty, subindo o Rio Corisco, 29
Jun. 1993, R. C. C. Reis, R. Marquete et al. 32 (RB).

Novon 5: 257-258. 1995.

258

Novon

F'guresl 7 AureUana darcyi Carvalho & Bovini (holotype). —1. Habit. —2. Receptacle of the inflorescence. —3.
R U . ' - <- , V '^' s owm 8 e ong-acuminate lobes. —5. Relation of the calyx to the corolla, external face. —6.

Relation ol the filaments to the anthers, internal face. —7. Globose fruit.

Acknowledgments. The authors thank Angela
M. Sturdart Vax of the Brazilian Institute of Geog¬
raphy and Statistics (IBGE) for the Latin descrip¬
tion, and Dorothy Sue Dunn de Araujo of the State
Environmental Protection Foundation (FEEMA) for
translating the text into English.

Literature Cited

Hunziker, A. T. & G. E. Barboza. 1991. Estudios sobre
Solanaceae XXX. Revisibn de Aureliana. Darwiniana
30(1-4): 95-113.

Carex radfordii (Section Laxiflorae: Cyperaceae), a New
Species from the Southern Appalachians

L. L. Gaddy

The Wild Wood Associates, Inc., The Wild Wood Way, Walhalla,

South Carolina 29691, U.S.A.

Abstract. Carex radfordii Gaddy is a new spe¬
cies of Carex sect. Laxiflorae from the southern Ap¬
palachian Mountains of North Carolina, South Car¬
olina, and Georgia. It is a robust species most
closely allied to Carex purpurifera Mackenzie and
Carex laxflora Lamarck. Its long awns, glaucous
blue-green blades, and brownish white basal
sheaths distinguish it from other species of Carex.

Carex sect. Laxiflorae has been studied by vari¬
ous investigators in this century. Recently, Bryson
(1980) carried out a morphological analysis, and
Manhart (1984, 1985, 1986) conducted studies on
the morphology, chemotaxonomy, and cytology of
the section.

Carex laxflora Lamarck, the namesake for sec¬
tion Laxiflorae, ranges across eastern North Amer¬
ica from Maine and southern Quebec south to North
Carolina west to Wisconsin and Indiana (Gleason
& Cronquist, 1991). It is probably the most variable
and complex species in section Laxiflorae and is
easily confused with other species in this section.
Cariciologists have often wondered whether Carex
laxflora was one species or a complex of species.
Mackenzie (1935) described C. purpurifera Mac¬
kenzie from specimens that were previously thought
to be C. laxflora, and Brysons (1985) description
of C. manhartii Bryson indicates that C. purpurifera
is, in fact, made up of at least two distinct elements.

The following species was first seen in the spring
of 1979. Since then, I have observed the plant at
numerous sites and mapped its distribution. I now
conclude that it is a new taxon.

Carex radfordii Gaddy, sp. nov. TYPE: U.S.A.
South Carolina: Oconee County, in shallow
soil, over calcareous rocks. Station Cove, 25
Apr. 1986, L. L Gaddy s.n. (holotype, CLEMS;
isotypes, GA, MO. NCU, NY. US, USCH,
WCUH, herbarium of C. T. Bryson). Figure 1.

A Carex laxflora aristis squamis pistillatis longis et
laminis venetis glaucis differt; a Carex purpurifera vaginis
basalis albidis brunneis differt.

Plant caespitose perennial, culms 4—6 dm tall.

strongly aphyllopodic, triangular, usually 1.5 X
longer than blades. Basal sheaths brownish white,
3—9 cm long, cataphylls well developed, 1-7 cm
long. Blades blue-green, glaucous on undersides,
2—5.5 cm long, 7-15 mm wide, midrib well devel¬
oped, with small teeth dorsally. Staminate spikes
2-A cm long, 3-6 mm wide, usually solitary,
peduncles 4—12 cm long. Pistillate spikes 3-5, pe¬
duncles 4—12 cm long, bracts 4—10 cm long, 1-2
mm wide, apices acuminate, sheaths 1-3 cm long,
ligules scarious, 4—6 mm long. Perigynia 3—10 per
spike. 3-A.5 mm long, obovoid, light green, beak
slightly recurved, to 1.5 mm long; scales acute to
aristate, often asymmetrical, rarely retuse, serrulate
awns 1—5 mm long, usually overtopping the beak
of the perigynia, total scale and awn length 3—9
mm, midrib green, margins scarious and hyaline.
Achenes obovoid, filling body of the perigynia, 1—
2.5 mm long, 0.8-1.2 mm wide, styles exserted,
stigmas 3.

I take pleasure in naming this species after Al¬
bert E. Radford. Professor Emeritus, University of
North Carolina (Chapel Hill), and a lifelong student
of Carex. In the mid 1970s, Dr. Radford told me
that the Brevard Belt of the Carolinas and Georgia
was a rich floristic zone that offered many potential
discoveries. I honor him with this discovery.

Some of the essential characters of C. radfordii,
C. purpurifera, C. laxflora, C. striatula Michaux.
and C. manhartii are compared in Table 1. As may
been seen in the table, C. radfordii shares many
characters with C. purpurifera and C. laxflora, and
fewer characters with C. striatula and C. manhartii.
Furthermore, measurements of the height of the
flowering culms, pistillate scale body length, and
pistillate scale awn length overlap somewhat in
these five species. In height of flowering culm, C.
radfordii is a robust species that can only be com¬
pared to C. purpurifera. Its blue-green blades (not
obvious in herbarium specimens), which are glau¬
cous on the undersides, are similar to those of C.
purpurifera, except for the lack of purplish pigment
in the basal sheaths. Its pistillate scale awns are
usually much longer than those of C. laxflora, C.

Novon 5: 2.59—261. 1995.

260

No von

Figure 1. Variation in pistillate scale shape and awn length in Carex radfordii Gaddy: from lower and upper pistillate
spikes of the type specimen. 9X. Illustration by Hu Ye.

striatula, and C. manhartii. Occasionally, the awns
of C. purpurifera' s pistillate scales approach those
of C. radfordii in length; however, on a given plant,
the mean pistillate scale awn length is always great¬
er in C. radfordii. The shape of the pistillate scale
awn in C. radfordii is also distinct from that of other
related species. In C. laxiflora, C. striatula, and C.
manhartii, the pistillate scales are acute to awned;
in C. purpurifera, the scales are more variable, of¬
ten being retuse; in C. radfordii, the scales are
long-awned to rarely retuse, with a majority usually
asymmetrical. Chromosome number in C. radfordii
has been determined to be n = 23, which separates
it from all related species (Robert Naczi, unpubli¬
shed data; Manhart, 1986).

Carex radfordii can be delimited from related
species by several characters, as may be seen in

the key below. From Carex manhartii and C. pur¬
purifera, to which it appears to be most closely re¬
lated, it can be separated by its brownish white
culm bases. From C. laxiflora and C. striatula, it
can be distinguished by its blue-green blades with
glaucous undersides and its asymmetrical pistillate
scale bodies with extremely long awns.

Upon first examination, C. radfordii appears to
be an “albino” C. purpurifera. Closer inspection,
however, reveals that the staminate spikes of C.
radfordii are significantly longer than those of C.
purpurifera and that the pistillate scales are more
variable in shape in C. radfordii. Other good field
characters for C. radfordii are its erect flowering
culms (those of C. laxiflora and C. striatula usually
lean over) and its strongly aphyllopodic culm bases.

Carex radfordii is found in rich, often rocky, cal-

Table 1. Carex radfordii and its allies.

Character

C. radfordii

C. purpurifera

C. laxiflora

C. striatula

C. manhartii

Height of flow¬
ering culms

4-6.5 dm

3.7-6.7 dm

3.1—5 dm

3-5.5 dm

2.1-3.9 dm

Blade color

Blue-green,

glaucous

underside

Blue-green,

glaucous

underside

Dark green

Whitish blue-
green

Dark green

Basal sheath
color

Brownish

white

Purple

Brownish white

Brownish white

Reddish purple

Shape of pis¬
tillate scale

Long-awned to
asymmetrical
to retuse

Long-awned to
retuse

Acute to awned

Acute to awned

Acute to awned

Scale length

2-4 mm

1.7—5 mm

2-4.5 mm

1.5-5.7 mm

3.5-4 mm

Awn length

1-5 mm

1—5 mm

0.5-2.3 mm

0.8-3.7 mm

0.1-3.5 mm

Chromosome

number

n = 23

n = 17, 18,

19

n = 20

n = 18, 20

n = 14

Volume 5, Number 3
1995

Gaddy

Carex radfordii

261

careous woodlands on well-drained soils. The plant
is found from about 300 m to just under 650 m in
elevation. Of the closely related species discussed
in Table 1, Carex radfordii is sympatric only with
C. manhartii (see below). Most known sites are lo¬
cated in the Brevard or Chauga Belt, a belt of meta¬
sedimentary rock along a transition zone between
the Piedmont and the Blue Ridge of the Carolinas
and Georgia. The rocks of this belt have weathered
to produce soils with high concentrations of calci¬
um and magnesium (Hatcher, 1969). The soil pH
at C. radfordii sites is fairly high for the generally
acidic southern Blue Ridge region; at Station Cove,
the type locality, the pH is 6.0 (Gaddy, 1990). Al¬
though it occurs in rich coves with C. austro-caro-
liniana Bailey, Carex radfordii is usually found at
rockier, better-drained microsites than those occu¬
pied by C. austro-caroliniana. Below Whitewater
Falls, North Carolina, C. radfordii grows alongside
of C. manhartii. When a plant of C. radfordii from
the type locality was experimentally grown in low
calcium, low pH soils with a plant of C. laxiflora
from the Blue Ridge of North Carolina and a plant
of C. striatula from the Piedmont of South Carolina,
all three species retained their distinguishing mor¬
phological characteristics.

Key to C. radfordii and Related Species

la. Base of culms brown, brownish white, or white.

2a. Blades whitish green to blue-green, under¬
sides white striolate; pistillate scales sym¬
metrical, awns 0.5-3.7 mm long.

3a. Blades whitish green to blue-green . .

. C. striatula

3b. Blades dark green. C. laxiflora

2b. Blades blue-green, undersides glaucous;
pistillate scale bodies often asymmetrical,
awns 1—5 mm long. C. radfordii

lb. Base of culms reddish, reddish purple, or pur¬

plish.

3a. Blades green, undersides not glaucous . . .

. C. manhartii

3b. Blades blue-green, undersides glaucous . .

.C. purpurifera

Paratypes. U.S.A. Georgia: Stephens County, Cedar
Creek, Camp Mikell Rd. off Ga. Rt. 184, just W of Camp
Mikell, N of Toccoa, elev. ca. 300 m, deep gorge, 19 June
1975, D. E. Boufford 16552 & E. Wood (GA). North Car¬
olina: Macon County, Whitewater Falls, just below falls
on lower portion of rich slopes, 15 May 1994, L L Gaddy

s.n. (CLEMS). South Carolina: Greenville County, with
Carex austro-caroliniana and Cladrastis kentuckea off
Bentfield Rd. N side of Saluda River, 17 May 1993, S. R.
Hill 25034 (CLEMS); Oconee County, in rich, calcareous
woods along Ramsey Creek below Rich Mountain, 20 Apr.
1992, L L Gaddy s.n. (CLEMS); Rich woods, Station
Cove, 22 Apr. 1986, L. L Gaddy s.n. (CLEMS); NE side
of ridge at Tamassee Creek, Rich cove woodland, Comp.
17, Stnd. 16, 8 May 1993, S. R. Hill 24994 (CLEMS).
Pickens County, Rich woods, below “the Narrows” in the
Eastatoe Gorge, 23 Apr. 1987, L L Gaddy s.n. with D.
A. Rayner (CLEMS); 1.3 mi. S of NC line, N of Rocky
Bottom, elev. ca. 2100 ft., diverse mesic forest slope near
creek crossing, soil a rich alkaline or neutral loam, oc¬
casional to frequent, conspicuously glaucous perennial, 10
Apr. 1988, S. R. Hill 19102 (CLEMS); shaded rich mesic
woodland, 1.3 mi. S of NC line, N of Rocky Bottom, elev.
ca. 2100 ft., occasional glaucous herb on steep slope, 9
May 1988, S. R. Hill 19271 with C. Horn (CLEMS).

Acknowledgments. I thank individuals who
commented on drafts of this paper over the last two
years. Special thanks go to A. A. Reznicek, Charles
T. Bryson, and Robert Naczi. I also acknowledge
the curators of the following herbaria for loans and
assistance: CLEMS, GA, MICH, NCU, OKL,
UARK.

Literature Cited

Bryson, C. T. 1980. A Revision of the North American
Carex Section Laxiflorae (Cyperaceae). Ph. D. Disser¬
tation, Mississippi State University, Mississippi State.
-. 1985. A new species of Carex (section Laxiflo¬
rae) from the southern Appalachians. Castanea 50: 15—
18.

Gaddy, L. L. 1990. Glade Fern Ravine, a rich fern site
in the Blue Ridge Province of South Carolina. Castanea
55: 282-285.

Gleason, H. A. & A. Cronquist. 1991. Manual of the
Vascular Plants of Northeastern United States and Ad¬
jacent Canada. Van Nostrand, 2nd ed.. The New York
Botanical Garden, Bronx, New York.

Hatcher, R. D. 1969. Stratigraphy, petrology, and struc¬
ture of the low rank belt of the Blue Ridge of the north-
westernmost South Carolina. Geologic Notes 13: 105-
141.

Mackenzie, K. K. 1935. Carex purpurifera and Carex lax¬
iflora. In: North American Flora 18: 253—255.

Manhart, J. R. 1984. A biosystematic study of Carex (sec¬
tion Laxiflorae). Ph.D. Dissertation, University of Geor¬
gia, Athens.

-. 1985. Foliar flavonoids of the North American

members of Carex section Laxiflorae (Cyperaceae). Bio-
chem. Syst. and Ecol. 14: 85-90.

-. 1986. Cytology of Carex purpurifera Mack. (Cy¬
peraceae). Rhodora 88: 141—147.

New Species of Southern African Moraea (Iridaceae—Iridioideae),
and the Reduction of Rheome

Peter Goldblatt

B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299,

St. Louis, Missouri 63166, U.S.A.

John C. Manning

National Botanical Institute, P. Bag X7, Claremont 7735, South Africa

Abstract. Two new species of the large African
genus Moraea (125 species), M. rivulicola and M.
regalis, both members of subgenus Viemseuxia, are
described here. Moraea rivulicola has cream to
greenish flowers, unusually long capsules, and is a
robust plant restricted to the banks and beds of
seasonal streams in Namaqualand, in the north¬
western part of South Africa. Moraea regalis is
known from one locality, near De Rust in the Little
Karoo, South Africa. Both species occur in the
semiarid winter rainfall part of southern Africa, a
region known for the richness and diversity of its
flora. The genus Rheome, now believed to be most
closely related to species of Moraea subg. Moraea,
is reduced to synonymy, and the three species of
Rheome are transferred to Moraea as M. maximi-
liani, M. nana, and M. umbellata.

Over half of the estimated 1750-1800 species of
the petaloid monocot family, Iridaceae, occur in
southern Africa (Goldblatt, 1991, 1994). The area
is well known botanically, yet despite more than
250 years of botanical exploration, novelties con¬
tinue to be discovered there. Although the family
is a significant element of the flora throughout the
subcontinent, Iridaceae are most diverse in the
southern and western parts of South Africa, a region
of winter rainfall and summer drought. Two new
species of Moraea are described here, both from
this portion of southern Africa, and the genus
Rheome is reduced to synonymy in Moraea. Moraea
is the largest African genus of subfamily Iridoideae,
and the central genus of Irideae-Homeriinae (Gold¬
blatt, 1976, 1986a). At the last revising, Goldblatt
(1986a) admitted 119 species to the genus. Since

then one species, M. vlokii (Goldblatt, 1992), has
been described. With the addition of the two new
species described here and three transferred from
Rheome, there are now 125 species in the genus.
According to the revised taxonomy, some 110 spe¬
cies of Moraea occur in southern Africa, that is,
south of the Zambezi—Cunene axis, most of them
endemic, and by far the majority restricted to the
semiarid and arid southwestern portion of the re¬
gion.

New Species

1. Moraea rivulicola Goldblatt & J. C. Manning,
sp. nov. TYPE: South Africa. Northern Cape
Province: Namaqualand, Springbok District,
streambed at the eastern end of Spektakel
Pass, 11 Sep. 1993, Goldblatt & Manning
9710 (holotype, NBG; isotypes, K, MO, PRE,
WAG). Figure 1A-D.

Plantae 50-80 cm altae, cormo globoso ca. 14—18 mm
in diametro tunicis reticulatis, folio solitario lineare can-
aliculato ca. 10 mm lato, caule ramoso, floribus cremeis
vel virescentibus vel pallide brunneis, tepalis exterioribus
unguiculatis atropunctatis ad bases limborum 20—24 mm
longis, limbis lanceolatis ca. 14 mm longis, unguibus can-
aliculatis 7—10 mm longis, tepalis interioribus 13—15 mm
longis tricuspidatis, filamentis ca. 8 mm longis in columno
ca. 5.5 mm longo connatis, antheris 4—5 mm longis, ramis
styli ca. 6 mm longis, cristis 3—5 mm longis erectis, cap-
sulis cylindricis 25-32 mm longis.

Plants 50-80 cm high. Corm globose, 14—18 mm
diam., the tunics of pale, coarse vertical libers con¬
nected by fine cross strands in herringbone pattern.
Cataphylls usually dry and brown at flowering time,
extending 5-10 mm above the ground. Foliage leaf

Figure 1. Morphology of Moraea rivulicola Goldblatt & J. C. Manning ( Goldblatt & Manning 9710 ) and M. unguiculata
Kew Gawler ( Goldblatt & Manning 9711). M. rivulicola: —A. Corm and basal part of stem. —B. Upper part of stem.

C. Outer and inner tepal. D. Capsule. M. unguiculata: —E. Leaf and upper part of stem. —F. Outer and inner
tepal. —G. Capsule.

Novon 5: 262-269. 1995.

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1995

Goldblatt & Manning
Southern African Moraea

263

264

Novon

Figure 2. Distribution ranges of Moraea rivulicola (closed circles), M. regalis (open circle), and M. unguiculata
(delimited by heavy lines).

solitary, basal, 2-2.5 times as long as the stem,
channeled, 6-10 mm wide (when opened flat). Stem
erect, consisting of 4(-5) intemodes, straight and
erect, the intemodes 10-15 cm long, simple or with
1-3 branches, the branches held close to the axes,
bearing an entirely sheathing bractlike leaf at each
of the Sf-A) nodes, sheathing leaves 50-65 mm
long, the apices attenuate-filiform. Inflorescences
rhipidia, these single and terminal on each branch,
several-flowered; spathes enclosing the rhipidia un¬
equal, green, firm-textured, with dry brown atten¬
uate apices, the outer ca. 45 mm long, the inner
55-67 mm long. Flowers cream, greenish or beige,
with large nectar guides at the bases of the outer
tepal limbs, these consisting of dark spots on a dull
yellow field, the claws similarly dotted, both guide
and outer tepal claws glabrous, inner tepals light
to reddish brown, also densely spotted; outer tepals
20—24 X 10—13 mm, the claws ascending, chan¬
neled, 7-10 mm long, each with a large basal nec¬
tary, the nectary opposed by a lobe ca. 2 mm long,
the limbs reflexed 45°-60° below horizontal, 14 X
10 mm; inner tepals 13-15 X 4—5 mm, the claws
ascending, ca. 8 mm long, channeled, the margins
usually partly conduplicate, the limbs trilobed, the
median lobe linear, curving inward laxly and asym¬

metrically coiled, lateral lobes broad and weakly
defined, obtuse, 1—2 mm long. Filaments ca. 8 mm
long, united in a cylindric column ca. 5.5 mm long,
free and diverging in the upper 2.5 mm; anthers 4—
5 mm long, each appressed to the opposed style
branch, shortly exceeding the stigma lobes, orange.
Ovary oblong-trigonous, 11—14 mm long, exserted
from the spathes; style dividing at the apex of the
filament column, the branches ca. 6 X ca. 2 mm,
diverging, each with paired terminal lanceolate
crests 3—5 mm long. Capsules cylindric, 25—32 mm
long; seeds angular, only known in the immature
state.

Flowering mid August to late September.

Etymology. From the Latin rivulieola, “living in
streams,” referring to the specialized habitat of the
species.

Distribution and habitat. Moraea rivulicola is a
fairly rare endemic of northern Namaqualand,
Northern Cape Province, South Africa. It extends
from the Spektakel Mountains west of Springbok
southward to the Kamiesberg (Fig. 2). Plants appear
to be restricted to the beds of seasonal streams and
along drainage lines, usually with their corms deep¬
ly wedged in crevices in the granite bedrock. Re-

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Goldblatt & Manning
Southern African Moraea

265

cords are all from areas of fairly high elevation, and
for Namaqualand, relatively high rainfall.

Although Moraea rivulicola was first collected in
1883 by Harry Bolus and in the past was thought
to be no more than a large-flowered variant of the
widespread and fairly common M. unguiculata Ker
Gawler (Goldblatt, 1976, 1986a), field observations
in Namaqualand in 1993 and 1994 show that this
treatment is incorrect. At two sites some 100 km
apart, M. rivulicola was found growing along and in
small seasonal streams, while fairly typical M. un¬
guiculata grew on adjacent well-drained slopes.
Seen at the same time and in quite different habi¬
tats, it became evident that these were different
species, not merely ecotypes or local variants of a
single polymorphic one. Critical comparison of liv¬
ing plants showed that not only is M. rivulicola
more robust vegetatively, and has larger flowers, but
that there are several critical qualitative differences
between it and M. unguiculata.

The flowers of Moraea rivulicola (Fig. 1A—C) are
cream with a pale greenish or beige tinge, and the
outer tepals are ca. 24 mm long with the limbs
typically held at 60° below the horizontal. The nec¬
tar guides, located at the base of the tepal limb,
are whitish covered with small dark brown to black¬
ish spots, and both the nectar guide area and the
tepal claws are smooth. The inner tepals are also
distinctive: the central lobe of the tripartite limb is
drawn into a slender obliquely coiled cusp, while
the lateral lobes are obtuse and very short. The
claws of the inner tepals are channeled and the
margins are partly conduplicate, thus forming a
nearly closed tube. The ovary is oblong-cylindric
and 11—14 mm long, and gives rise to cylindrical
capsules 25-32 mm long. Ripe seeds are unknown,
but are probably larger than those of M. unguicu¬
lata. Moraea unguiculata, both in Namaqualand
and elsewhere across its range, has smaller flowers
with the outer tepals seldom exceeding 16 mm (Fig.
1E-G). Larger dimensions for the species given by
Goldblatt (1976, 1986a) included plants now re-
ferred to M. rivulicola. The tepal limbs in Nama¬
qualand populations of M. unguiculata are 8-10
mm long, and are held slightly below the horizontal,
or flexed up to 45°, but in populations in the south¬
ern Cape, the limbs may be fully reflexed. Both the
small nectar guide area and the outer tepal claws
are densely pubescent. The inner tepals are 7—10
mm long, and also tripartite. The central cusp is
long and symmetrically coiled inward, but the outer
lobes are relatively longer and more distinctly
formed than in M. rivulicola, and may be subacute.
The inner tepal claws are more or less terete or
slightly flattened on the adaxial surface. Flower col¬

or in M. unguiculata varies, ranging from pure
white to cream, brownish, and even blue, and floral
form (but not size) is unusually plastic (Goldblatt,
1986a). The outer tepal limbs may extend outward
or, in some populations outside Namaqualand, may
be completely reflexed.

The distinctive pale green leaves of Moraea ri¬
vulicola are relatively long, sometimes up to 2.5
times as long as the stem, thus over 1 m in length,
broadly channeled, and 6-10 mm wide. The stems
are 4—5 intemodes long and bear 3-4 sheathing
leaves 50-65 mm long. Leaves of M. unguiculata
are darker gray-green, shorter, seldom more than a
third again as long as the stem, and they are nar¬
rowly channeled, 2—3(-4) mm wide. The stems are
3 intemodes long and bear 2 sheathing leaves TO¬
SS mm long. Thus the two species differ both in leaf
and stem morphology as well as in floral structure.

Differences in the leaf texture and venation of
the two species reflect notable anatomical differ¬
ences (Fig. 3). In transverse section the leaf of Mo¬
raea unguiculata (Fig. 3B) is relatively thin. The
heavily sclerified sheaths of the major bundles
touch those of the opposing minor bundles and
there is a large strand of subepidermal marginal
sclerenchyma, not apparently associated with a vas¬
cular bundle (although a small central cavity sug¬
gests the presence of minor veins in early ontoge¬
ny). In comparison, the leaf of M. rivulicola is much
thicker in transverse section (Fig. 3A), the sheaths
of the major bundles are less heavily sclerified and
the opposing minor bundles are well separated from
them, and most significantly, there is a small mar¬
ginal vein with a small sclerified phloem cap. These
differences in leaf anatomy between the two species
are of the kind most commonly associated with xer-
ic versus more mesic habitats.

Habitat differences between Moraea rivulicola
and M. unguiculata appear to be absolute. The lat¬
ter grows on open exposed rocky sites, or on stony
slopes, in both cases in well-drained and often arid
situations. Moraea rivulicola has, however, so far
only been found in seasonally wet habitats, water¬
logged or even inundated for part of the growing
season. In Namaqualand M. unguiculata may be
found either on soils derived from shale, as in the
Spektakel Mountains or near Steinkopf (Goldblatt,
1986a), or on granite-derived soils. Moraea rivuli¬
cola has only been found on pockets of granite-
derived sand in rocky granite on the banks or in
the beds of streams.

Both Moraea rivulicola and M. unguiculata be¬
long to subgenus Vieusseuxia, members of which
share the following synapomorphies: long-lasting
flowers, a solitary foliage leaf, distinctive reticulate

266

Novon

corm tunics composed of pale, soft-textured fibers,
and an asymmetric karyotype with x = 6. Within
the subgenus, M. rivulicola and M. unguiculala ap¬
pear to be allied to those species that have similar
trilobed inner tepals, inconspicuous nectar guides,
and a long filament column. Other species in the
alliance include the southwestern Cape species M.
tricuspidata (L. fil.) G. Lewis and M. bellendenii
(Sweet) N. E. Brown, and the eastern southern Af¬
rican M. trifida R. Foster, M. pubiflora N. E. Brown,
and M. brevistyla (Goldblatt) Goldblatt.

Paratypes. SOUTH AFRICA. Northern Cape: 2917
(Springbok) 22 km W of Springbok, eastern slopes of
Spektakel Mountains, in streambed (DA), 26 Sep. 1974,
Goldblatt 2791 (MO, PRE); streambed at the eastern end
of Spektakel Pass, 4 Sep. 1994, Goldblatt & Manning
9971 (MO, NBG); rocky places near Modderfontein (DB),

Sep. 1883, H. Bolus 6618 (BOL). 3018 (Kamiesberg) east¬
ern slopes of Rooiberg, ca. 1050 m (AC), 14 Oct. 1976,
Goldblatt 4313 (MO, PRE); Kamiesberg Mountains, farm
Welkom, streambed near the top of Langkloof, 14 Sep.
1993, Goldblatt & Manning 9770 (MO).

2. Moraea regalis Goldblatt & J. C. Manning, sp.
nov. TYPE; South Africa. Western Cape: Little
Karoo, De Rust, rocky hillside near old cem¬
etery, 19 Sep. 1994, /. A. Vlok & A. L Schutte
345 (holotype, NBG; isotypes, K, MO, PRE,
WAG). Figure 4.

Plantae 18-25(—35) cm altae, cormo globoso 12—14 mm
in diametro tunicis reticulatis, folio solitario lineare can-
aliculato, caule eramoso, floribus violaceis, tepalis exter-
ioribus albis notatis unguiculatis (26—)30-A0 mm tongis,
limbis lanceolatis 13—20 mm longis, tepalis interioribus
20—24 mm longis erectis saepe unguiculatis et limbis tn-

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Goldblatt & Manning
Southern African Moraea

267

cuspidatis, filamentis 12—15 mm longis connatis in col-
umno 9—10 mm longo, antheris 6-9 mm longis, ramis styli
10—12 X 2 mm divergentibus, cristis anguste lanceolatis
vel linearibus 5-8 mm longis.

Plants 18-25(-35) cm high. Corms globose, 12-
14 mm diam., the tunics of medium to coarsely
textured reticulate fibers, light brown. Cataphylls
dry and brown at flowering time. Foliage leaf sol¬
itary, basal, half again to twice as long as the stem,
linear and canaliculate, ca. 3 mm wide (when
opened flat), upright below, trailing above. Stem
erect, 3 intemodes long, straight and erect, un¬
branched, the lowermost internode shortest and the
uppermost longest, bearing an entirely sheathing
bractlike leaf at each node, these 25-40 mm long,
the apices attenuate. Inflorescence a single, usually
2-flowered terminal rhipidium; enclosing spathes
unequal, green, firm-textured, with dry brown at¬
tenuate apices, the outer 25-40 mm long, the inner
45-65 mm long. Flowers violet, the limbs of the
outer tepals each marked at the base with a small
white triangular nectar guide outlined with a dark
purple zone, the base of the outer tepal limb mi¬
nutely pubescent, the claw heavily pubescent, the
hairs dark violet, evidently without nectar, scent¬
less; outer tepals (26—)30-40 mm long, the claws
ascending, 12-16 mm long, with a smooth deltoid
zone at the base, evidently a vestigial perigonal
nectary, the limbs 13—20 mm long, directed down¬
ward at right angles to the ground; inner tepals erect
and ± linear to slightly wider just below the mid¬
line, thus obscurely clawed, 20-24 mm long, the
claw ca. 12 mm long, the limb aristate or occasion¬
ally tricuspidate, then the lateral lobe 1-2 mm long,
and central cusp ca. 10 mm long. Filaments 12-15
mm long, united below in a cylindric column 9-10
mm long, free and diverging in the upper 3-5 mm;
anthers 6-9 mm long, each appressed to the op¬
posed style branch, just shorter than or just ex¬
ceeding the stigma lobes, the pollen orange. Ovary
oblong-trigonous, 8-10 mm long, usually fully ex-
serted from the spathes, style dividing just above
the apex of the filament column, the branches 10-
12 X 2 mm, diverging, each with paired terminal,
narrowly lanceolate to linear crests 5-8 mm long.
Capsules ovoid-oblong, 12-16 mm long; seeds an¬
gular and prismatic, ca. 1.5 mm long.

Flowering mid August to mid September.

Etymology. From the Latin regalis, “royal,” re¬
ferring to the dark blue-purple or violet color of the
flowers.

Distribution and habitat. Moraea regalis ap¬
pears to have been collected for the first time in
1993, when it was found near the Little Karoo town

268

Novon

of De Rust (Fig. 2). So far it is known only from a
single locality, south-trending slopes of a rocky hill
south of De Rust. The substrate is sandy soil de¬
rived from Enon Conglomerate.

Like Moraea rivulicola, described above, M. re-
galis is a typical member of subgenus Vieusseuxia.
It is distinguished in the genus and subgenus by
the striking dark violet flower color, the relatively
large flower size with the outer tepals 26-40 mm
long, the long tepal claws, and the narrow, either
aristate and more or less linear or tricuspidate inner
tepals (Fig. 4). Its affinities lie with M. unguiculata
and M. algoemis Goldblatt, both of which have
smaller flowers and trilobed inner tepals with a
large incurved central cusp and small but broadly
obtuse lateral lobes. The slender filament column
and narrow style branches are also typical of this
group of species.

Paratype. SOUTH AFRICA. Western Cape: 3322
(Oudtshoom) Little Karoo, De Rust, hill S of the town,
stony S-facing slopes (BC), Sep. 1993, J. A. Vlok & A. L
Schutte s.n. (MO).

Reduction of Rheome Goldblatt

The genus Rheome was erected by Goldblatt
(1980) to accommodate two species of Homeria, H.
umbellata (Thunberg) G. Lewis and H. maximiliani
Schlechter. These differed from other species then
included in Homeria in their brown, nonreticulate
corm tunics, apically diverging outer inflorescence
spathes, and apically crowded rhipidia (inflores¬
cence units) on the stem. The remaining species of
Homeria have reticulate corm tunics composed of
coarse black fibers, sheathing outer rhipidial
spathes, and rhipidia that are not crowded apically.
The basic chromosome number also differs in the
two groups. Species of Homeria have x = 6, and
secondarily 2 n = 10, 9, or 8, whereas H. umbellata
and H. maximiliani have x = 10, considered to be
the plesiomorphic chromosome number for Hom-
eriinae (Goldblatt, 1976, 1980; Rudall & Goldblatt,
1993). Moreover, a series of crossing experiments
showed that although most of the species of Hom¬
eria could be crossed with one another, none could
be crossed with H. maximiliani. The latter, how¬
ever, could be crossed with one species of a second
genus Hexaglottis, H. nana L. Bolus, which also
has the corm tunics, rhipidial spathes, and arrange¬
ment of the rhipidia characteristic of Rheome. Nei¬
ther Homeria maximiliani nor Hexaglottis nana
could be crossed with a selection of species of Hex¬
aglottis. It was evident on the basis of morpholog¬
ical, cytological, and crossing evidence that Hom¬
eria umbellata, H. maximiliani, and Hexaglottis

nana were a monophyletic group parallel to Hex¬
aglottis and Homeria and that the close floral sim¬
ilarity to the latter two genera was due to conver¬
gence, most likely through the parallel reduction of
the more complex Moraea- type flower, plesio¬
morphic in subtribe Homeriinae in its dimorphic
inner and outer tepals and narrow style branches
either bearing a reduced crest or lacking them en¬
tirely.

What was not evident when Rheome was de¬
scribed was that the distinctive corm tunics, inflo¬
rescence spathes, and arrangement of rhipidia
characteristic of that genus have a parallel in Mo¬
raea longiflora Ker Gawler (subg. Moraea sect.
Tubiflora). The second species of section Tubiflora,
M. coopen Baker, also has brown, nonreticulate
corm tunics, but it has entirely sheathing rhipidial
spathes and laxly disposed rhipidia. Basic chro¬
mosome number in subgenus Moraea is x — 10,
which also accords with the number in Rheome.
The discovery of M. linderi Goldblatt (Goldblatt,
1982) provided the missing link between Rheome
and Moraea. This species has a flower typical of
Moraea, having large outer tepals with conspicuous
nectar guides, narrow inner tepals, broad style
branches with paired petaloid crests, and a trans¬
verse stigma, but corms, corm tunics, rhipidial
spathes and rhipidial arrangement of the type char¬
acteristic of Rheome, and a basic chromosome num¬
ber of x = 10 (Goldblatt, 1986b).

Finally, a phylogenetic study of subtribe Hom¬
eriinae (Rudall & Goldblatt, 1993), using several
leaf anatomical characters as well as morphology
and chromosome cytology, provided evidence that
Rheome, Moraea sect. Tubiflora, and two other spe¬
cies of section Moraea, M. linderi and M. margar-
etae Goldblatt, comprise a clade. The study did not
include enough species to show, with any degree of
confidence, the species or section most closely al¬
lied. Nevertheless, the evidence now available
makes it clear that the species of Rheome and Hex¬
aglottis nana are nested within a small clade of
species of Moraea subg. Moraea. Under these cir¬
cumstances, continued recognition of Rheome, a
small genus nested within a section of the large and
diverse Moraea, is clearly inconsistent with modem
phylogenetic classification (Hennig, 1966; Dono-
ghue & Cantino, 1988). We therefore propose to
synonymize Rheome in Moraea. As noted above,
Rheome, H. nana, M. linderi, and M. margaretae
comprise a clade with the following apomorphic
characteristics: nonreticulate, brown corm tunics;
apically diverging outer rhipidial spathes, and a
tendency for the crowding of rhipidia and foliage
leaves near the stem apex. Some of these characters

Volume 5, Number 3
1995

Goldblatt & Manning
Southern African Moraea

269

are also shared by the members of section Tubiflora
which, in addition, have the tepals united in a tube,
apomorphic in Moraea. The absence of some of the
above synapomorphies in M. cooperi is puzzling,
and we suspect may be due to character reversal.
The taxonomy of species transferred to Moraea is
outlined below.

1. Moraea maximiliani (Schlechter) Goldblatt &

J. C. Manning, comb. nov. Basionym: Homeria
maximiliani Schlechter, Bot. Jahrb. Syst. 27:
94. 1900. Rheome maximiliani (Schlechter)
Goldblatt, Bot. Not. 133: 92. 1980. TYPE:
South Africa. Western Cape: Hoek, Clanwil-
liam District, Schlechter 8700 (lectotype, des¬
ignated by Goldblatt (1980), B; isotypes, BOL,

K. MO, S).

2. Moraea nana (L. Bolus) Goldblatt & J. C. Man¬

ning, comb. nov. Basionym: Hexaglottis nana

L. Bolus, Kew Bull. Misc. Inf. 1932: 326.
1932 [1933]. TYPE: South Africa. Western
Cape: Clanwilliam District, without precise lo¬
cality, Weintraub s.n. (holotype, BOL 19972).

3. Moraea umbellata Thunberg, Dissertatio de

Moraea no. 16. 1787. Homeria umbellata
(Thunberg) G. Lewis, J. S. African Bot. 14: 86.
1948. Rheome umbellata (Thunberg) Gold¬
blatt, Bot. Not. 133: 92. 1980. TYPE: South
Africa. Western Cape: near Piketberg, Thun¬
berg s.n. (holotype. Herb. Thunberg 1230 at
UPS).

Acknowledgments. We thank Jan Vlok, Western

Cape Department of Nature Conservation, for his

assistance with fieldwork and for bringing M. re¬
gain to our attention.

Literature Cited

Donoghue, M. J. & P. D. Cantino. 1988. Paraphyly, an¬
cestors and the goals of taxonomy: A botanical defense
of cladism. Bot. Rev. 54: 107—127.

Goldblatt, P. 1976. The genus Moraea in the winter rain¬
fall region of southern Africa. Ann. Missouri Bot. Card.
63: 657-786.

-. 1980. Redefinition of Homeria and Moraea (Ir-

idaceae) in the light of biosystematic data, with Rheome
gen. nov. Bot. Not. 133: 85-95.

-. 1982. A synopsis of Moraea (Iridaceae) with

new taxa, transfers and notes. Ann. Missouri Bot. Gard.
69: 351-369.

-. 1986a. Convergent evolution of the Homeria

flower type in six new species of Moraea (Iridaceae) in
southern Africa. Ann. Missouri Bot. Gard. 73: 102—116.

-. 1986b. The Moraeas of Southern Africa. Ann.

Kirstenbosch Bot. Gard. 14: 1—224.

-. 1991. An overview of the systematics, phylogeny

and biology of the southern African Iridaceae. Contr.
Bolus Herb. 13: 1—74.

-. 1992. New species, chromosome cytology and

notes on the southern African Iridaceae—Irideae: Mo¬
raea, Roggeveldia and Homeria. S. African J. Bot. 58:
209-214.

-. 1994. Geography of African Iridaceae. Pp. 931—

940 in J. H. Seyani & A. C. Chikuni (editors). Pro¬
ceedings of the XHIth plenary Meeting AETFAT, Ma¬
lawi, Volume 2. National Herbarium and Botanic
Gardens of Malawi, Zomba.

Hennig, W. 1966. Phylogenetic Systematics. Translated
by D. D. Davis & R. Zangerl. Univ. Illinois Press, Ur-
bana.

Rudall, P. & P. Goldblatt. 1993. Leaf anatomy and sys¬
tematics of Homeriinae (Iridaceae). Bot. J. Linn. Soc.
Ill: 379-397.

A New Rhynchospora (Cyperaceae) from the Cordillera de
Talamanca, Costa Rica

Jorge Gdmez-Laurito

Escuela de Biologfa, Universidad de Costa Rica, Ciudad Universitaria 2060,
San Jose, Costa Rica

Abstract. Rhynchospora cabecarae G<5mez-Lauri-
to (Cyperaceae), a new species belonging to section
Racemosae C. B. Clarke, is described and illustrated.

A new species of Rhynchospora Vahl (Cypera¬
ceae) is described as a result of revision of material
recently collected during a field trip to the Carib¬
bean slope of the Cordillera de Talamanca in Costa
Rica.

Rhynchospora cabecarae Gbmez-Laurito, sp.
nov. TYPE: Costa Rica. Li rnon: Talamanca, N
flank of Cerro Casma, along Ujarras-San Jose
Cab^car trail. Cordillera de Talamanca,
9°20'30"N, 83°13'30"W, 2250-2270 m, 17
Mar. 1993, M. Grayum 10354 (holotype, USJ).
Figure 1.

Ex affinitate R. boliviensis C. B. Clarke atque R. dissi -
tiflorae Steudel ex Bockeler, ab utroque characteribus in-
florescentiarum et achaeniarum praecipue differt.

Perennial, loosely caespitose with short rhizomes
and coarse roots. Culms covered by old sheaths,
erect, stiff, ca. 60 cm tall, 2.5 mm thick, nodose,
leafy with many cauline leaves. Leaves 8-20 cm
long, erect, somewhat wiry; blades linear, 2-2.8
mm wide, gradually narrowed apically, the margins
ciliate to scabrous, channeled, the midrib adaxially
pilose, raised and ciliate abaxially. Sheaths striate,
reddish brown, villous with soft, curled hairs. Inflo¬
rescence paniculate, of ca. 5 partial panicles, the
panicles not exceeding the leaves, 1.5—2.5 cm long,
1.3—2 cm wide, of 11-23 ascending spikelets; the
peduncles 1.3-2 cm long, terete to complanate, hir¬

sute, the axis sparsely hirsute. Bracts leaflike.
Bractlets subulate, 4—6 mm long, hirsute. Spikelets
6-7 mm long, 1—1.3 mm wide, acute, reddish
brown, 2-fruited. Lower glumes empty, ovate, 2.5-
3 mm long, mucronate, other flower-bearing glumes
lanceolate, 4 mm long, 1.5 mm wide, membranous,
mucronate at the tip. Achenes ovate-elliptic to el¬
liptic, tumid, 1.2 mm long, 0.7 mm wide, with a
conspicuous basal stipe 0.5 mm long, shiny-casta-
neous, cancellate with minute isodiametrical cells.
Style-base conical, 1 mm high, 0.8 mm broad at
base, dorsiventrally compressed, brown. Hypogy-
nous bristles 5, upwardly scabrous, rather slender,
shorter than the achene body, or often only 2 very
short rudimentary ones. Stamens 3.

Etymology. This species is named after the Ca-
becar people, who inhabit the Talamanca region.

Rhynchospora cabecarae is a member of section
Racemosae C. B. Clarke because of its short rhi¬
zome, culms covered by old sheaths, many cauline
leaves, inflorescence paniculate, number of hypog-
ynous bristles shorter than the achene body, often
only 2 vestigials, and cancellate achenes. It is ap¬
parently related to R. boliviensis C. B. Clarke and
R. dissitiflora Steudel ex Bockeler. As has recently
been stated by Thomas (1993), this group is quite
variable and rare. Rhynchospora cabecarae is
known only from the type collection. It is distin¬
guished from the above-mentioned species by the
combination of characters in Table 1.

Rhynchospora cabecarae was found in an open
scrubland with Sphagnum and shrubs and treelets
of several families, including Clusia (Clusiaceae),

Table 1. Comparison of three related species of Rhynchospora.

R. boliviensis

R. dissitiflora

R. cabecarae

Inflorescences A single open panicle with

slender peduncles 1—5 cm
long with solitary spikelets
Achenes Widely obovate, lenticular to

plano-convex; brownish gray
to straw color. Stipe none

3-5 racemes, each with 1-4
spikelets with slender and
pendant peduncles

Widely obovate, lenticular,
pale brownish gray. Stipe
none

5 partial small panicles, each
with 11-23 spikelets; pe¬
duncles short, 1—3 cm long
Ovate to ovate-elliptic, tumid,
shiny-castaneous, with a
distinct stipe 0.5 mm long

Novon 5: 270-271. 1995.

Volume 5, Number 3
1995

Gomez-Laurito 271

Rhynchospora cabecarae from Costa Rica

figure 1. Rhynchospora cabecarae Gdmez-Laurito (Grayum 10354, USJ). a. Habit. b. Partial panicle.
Achene.

Prunus (Rosaceae), Vaccinium (Ericaceae), Ilex
(Aquifoliaceae), Miconia (Melastomataceae), Pali-
courea (Rubiaceae), Gaiadendron (Loranthaceae),
Monnina (Polygalaceae), Neomirandea (Astera-
ceae), and Phyllonoma (Grossulariaceae) (M. Gra¬
yum, in litt. 1994).

Acknowledgments. I am grateful to Michael H.
Grayum for reviewing the manuscript and to Judy
Hoomans for the excellent illustration.

Literature Cited

Thomas, W. W. 1993. A synopsis of Rhynchospora (Cy-
peraceae) in Mesoamerica. Brittonia 44: 1 4-44 .

Two New Species in Cuphea (Lythraceae), and a
Note on Alzateaceae

Shirley A. Graham

Department of Biological Sciences, Kent State University, Kent, Ohio 44242 U.S.A.

Abstract. Cuphea (sect. Melvilla) alaniana from
central Mexico and C. (sect. Diploptychia) armata
from Central America are described and illustrated.
The monotypic family Alzateaceae, long known
from Peru and Bolivia, and more recently discov¬
ered in Costa Rica, Panama, and Colombia, is here
reported from Ecuador for the first time. Of the two
Ecuadorian collections, one belongs to the north¬
ern, broad-leaved Alzatea verticillata subsp. ampli-
folia; the other is most similar to the southern sub¬
species verticillata. Apical leaf glands are a newly
recognized feature of the genus. They are inter¬
preted as extra-floral nectaries.

In revisionary studies of Cuphea, two new North
American species were discovered among collections
on loan from a number of North American herbaria.
Cuphea alaniana is a large-flowered perennial of
section Melvilla, subsection Polyspermum (Koehne,
1903). It is one of four species of the subsection,
which is endemic to the western and southern moun¬
tains of Mexico. Cuphea armata, discovered in
neighboring areas of Guatemala, El Salvador, and
Honduras is a new species of section Diploptychia.

Cuphea alaniana S. A. Graham, sp. nov. TYPE:
Mexico. Guerrero: Distrito Adama, Temisco,
Barranca del Consuelo, 500 m, 15 Nov. 1937,
Mexia 8810a (holotype, UC; isotype, GH). Fig¬
ure la-d.

Suffmtices vel frutices, probabiliter ad 1.5 m, caulis
simplex, erectus, viscosissimus. Petioli 5-10 mm longi, ad
summum caulis sessiles; folia 25-100 mm longa, 5-35
mm lata, anguste lanceolata vel linearia. Flores 25-32 mm
longi. rubro-purpurei, glanduloso-hispidi; lobus dorsalis
calycis ceteris paullo major; appendices prominentes, se-
tosae. Petala 6, roseo-purpurea, inaequalia, 2 dorsalia 7-
9 mm longa, manifeste unguiculata, 4 ventralia 1.0-2.5
mm longa. Ovula 40-50. Discus 2-3 mm longus, 2.5 mm
latus, 2 mm crassus, basi ovarii late affixus.

Suffrutescent or shrubby perennials, probably
0.5-1.5 m tall, stems simple, erect, from a fibrous-
rooted base, branched distally in the inflorescence;
stems and pedicels highly viscous, the vestiture
densely, minutely glandular-puberulous and glan¬
dular-hispid, the hairs thin, erect, colorless or red-

Novon 5: 272-277. 1995.

dish purple, bulbous-based, minute (< 0.25 mm)
or 0.5-1 mm long; intemodes up to half as long as
the subtending leaves. Leaves with petioles 5—10
mm long, becoming subsessile or sessile distally in
the inflorescence; blades 25-100 mm long, 5-35
mm wide, membranous, crowded on the upper
stem, narrowly lanceolate-linear, base acute, apex
acute; blade surface minutely scabrous, or scabrous
with sparse to abundant white, weak, slightly ap-
pressed hairs to 1 mm long, the margins of young
leaves densely ciliate, the hairs erect, glandular-
viscous, to 1 mm long; upper leaves very gradually
diminishing in size to become the lanceolate-linear
bracts of the inflorescence. Inflorescences densely
flowered, terminal, leafy racemes; flowers solitary
at the nodes, interaxillary; pedicels 4—6 mm long;
bracteoles 1.5 mm long, lanceolate-linear or re¬
duced to a few fleshy hairs, positioned 1.0-1.5 mm
below the apex of the pedicel. Floral tubes 25-32
mm long, including a rounded, horizontal base ex¬
tending 1 mm beyond the pedicel, 4—5 mm wide in
lateral view at anthesis, dorsally slightly convex,
the neck slightly or not at all narrowed, the mouth
flaring, oblique in lateral view, the ventral side ex¬
tended 1 inin or less beyond the dorsal side, the
floral tube more distinctly convex and distended in
fruit, the neck contracted; outer surface reddish
purple, straw-colored ventrally at the mouth, glan¬
dular-puberulous and glandular-hispid with abun¬
dant, erect, thin, colorless or reddish purple hairs
to 1 mm long; inner surface pale, neither bialate
nor vesiculate, glabrous or lightly villous above the
stamens, glabrous below; calyx lobes subequal, the
dorsal lobe deltoid, 2X3 mm, the others 1X2
mm, margins not ciliated; appendages at the sinus¬
es of the lobes prominent, 1 mm long, equaling the
calyx lobes, thick, green or reddish purple, the
apex bulbous, crowned by 5-8 setae 1—2 mm long.
Petals 6, rose-purple, the 2 dorsal 7—9 mm long
including a slender claw 1-2 mm long, narrowly
obovate, the 4 ventral 1.0-2.5 mm long, linear-spa-
thulate, caducous. Stamens 11, all inserted at % the
length of the floral tube, the 2 dorsalmost shortest,
included, the 5 antesepalous exserted, the 4 ante-
petalous included to subexserted; filaments gla-

Volume 5, Number 3
1995

Graham

Cuphea

273

Figure 1. a-d, Cuphea alaniana S. A. Graham, —a. Habit. —b. Lateral view of flower. —c. Nectariferous disc at
base of the ovary within the floral tube. —d. Floral tube opened dorsally, ovary removed to show position and number
of stamens. —e. Cuphea armata S. A. Graham, lateral view of flower. Drawn from the holotypes.

brous to lightly villous, reddish; anthers purple.
Pollen triangular in polar view; tricolporate, syn-
colpate, pores not protruding; exine uniformly fine¬
ly striate, the striae extending ca. Vi the distance
to the poles, partially anastomosing; diameter 24-
28 gm in lactic acid. Style exserted at maturity,
glabrous; stigma punctiform; ovary gibbous dorsal¬

ly, the gibbosity pronounced in flower; ovules 40-
50. Seeds numerous, 2 mm long, 1.5 mm wide. Disc
massive, 2-3 mm long, 2.5 mm wide, 2 mm thick,
horizontal, oblong-rectangular in dorsal view,
broadly attached at the base of the ovary.

Phenology. Collected in flower and fruit in No¬
vember.

274

Novon

Figure 2. Distribution of Cuphea alaniana and C. armata in Mexico and Central America.

Distribution and habitat (Fig. 2). In southern
Mexico state and adjacent Guerrero; low spiny, de¬
ciduous woods, dry slopes, and sandy streamsides;
500-700 m.

Etymology. The species is named after Alan
Graham, who has generously and ably assisted on
field excursions many times since our first trip to
Mexico for Cuphea in 1961.

The recognition of Cuphea alaniana adds a
fourth species to subsection Polyspermum of section
Melvilla. Species of the subsection are endemic to
the western and southern mountains of Mexico. The
subsection is characterized by spurless, deeply col¬
ored, thick-bodied floral tubes 20-32 mm long and
ovules numbering 40-120. In C. alaniana, the red¬
dish purple floral tube with two large erect, clawed
dorsal petals, four small ventral petals, and a mas¬
sive horizontal nectary termed the “disc” serve best
to distinguish it from C. micropetala var. hirtella
Koehne, to which it is surely most closely related.
The latter differs in having six minute petals of
equal size and a small disc. Both species share an
unusual, dorsally gibbous ovary in addition to sim¬
ilar vegetative, trichome, and pollen morphology.
Within the subsection, C. alaniana also resembles
the poorly known C. heteropetala Koehne from Oa¬

xaca by its slightly enlarged dorsal lobe and large
petals, but C. heteropetala lacks a gibbous ovary
and has a small, erect disc. Attempts to relocate C.
alaniana in the states of Mexico and Guerrero were
unsuccessful. The type area, along the river near
Temisco, Guerrero, is now occupied by dwellings
and cultivated fields. Exploration further upstream
in less disturbed sites also failed to locate the spe¬
cies. The paratype locality, directly north of Tem¬
isco in Mexico, was not accessible by road in 1989,
and was not visited by us.

Paratypes. MEXICO. Mexico: Cerro de San Antonio,
Tlatlaya, Matuda 32110 (GH, MEXU).

Cuphea armata S. A. Graham, sp. nov. TYPE: El
Salvador. Chalatenango: 4 km SSE of La Pal¬
ma, pine forest, 950 m, 11 June 1970, G. Dav-
idse & R. Pohl 2073 (holotype, F; isotype,
MO). Figure le.

Suffrutices, 0.5-1 m, caules erecti vel suberecti, pilis
minutis albidis antrorsis adpressisque. Folia sessilia vel
subsessilia, 20-50 mm longa, 5-12 mm lata ad medium
caul is, ovata, oblonga, vel sublinearia, basi obtusa vel cor-
data, utrinque scabrella. Inflorescentia sat distincta, ra-
cemosa, interdum subcomposita; pedicelli gracili, 2-5 mm
longi, persistentes; bracteolae 0.5 mm longae, lineari-lan-

Volume 5, Number 3
1995

Graham

Cuphea

275

ceolatae. Flores 12—15 mm longi, pallidi purpureo-rosei,
basi puberuli saepe prominente bulboso-setosi, calcari 4
mm longo, distincte ascendente etiam conico; intus infra
stamina bialatus, alae glabrae. Petala 6, 2 dorsalia intense
violacea, majora, 7—9 mm longa; 4 ventralia minora 3-5
mm longa, pallide violacea. Stamina 11, 9 longiora exser-
ta. Stylus demum exsertus; ovula 8—10. Semina 2 X 1.5
mm, oblonga, margine obtusa. Discus crassus, deflexus.

Semi-shrubby perennials, 0.5—1 m, erect or
weakly erect and sprawling, irregularly branched
from above the base, branches shorter than to lon¬
ger than the main stem, bearing minute white, op¬
pressed, antrorse, sharp-pointed hairs and a lon¬
gitudinal row of minute, falcate, antrorse hairs;
intemodes mostly equal to or shorter than the sub¬
tending leaves. Leaves sessile or subsessile, petioles
0-1 mm; blades at midstem 20-50 mm long, 5—12
mm wide, decreasing in size above and below the
midstem, membranous, ovate, oblong, or sublinear,
base rounded to cordate, apex acute to acuminate;
surfaces finely scabrous, appearing glabrous, the
hairs minute, appressed, the margins of the upper
leaves and bracts ciliolate, bearing sparse bulbous-
based setae, upper surface darker green than the
lower; upper leaves gradually diminishing in size
to become the bracts of the inflorescence. Inflores¬
cences terminal bracteolate racemes or loose, short-
branched thyrses; one flower interaxillary at a node,
1—3 additional flowers sometimes present on slen¬
der, alternate, axillary branchlets ca. 1 cm long;
pedicels 2—5 mm, slender, persistent; bracteoles
0.5 mm, linear-lanceolate, persistent at the apex of
the pedicel. Floral tubes 12—15 mm long, including
an ascending, conical spur 4 mm long, the apex
sometimes slightly descending, floral tubes dorsally
concave, 1-2 mm wide at anthesis, in lateral view
the base wider than the midportion, the neck not
strongly contracted at anthesis, the mouth oblique,
the ventral side extended 1 mm or less beyond the
dorsal, the dorsal and adjacent lateral ribs diverg¬
ing approximately 1 mm; outer surface pale rose-
purple, darkest dorsally, puberulent with sparse but
prominent red-purple, bulbous-based setae present
on the dorsal and lateral ribs, especially toward the
base of the tube, rarely individual flowers lacking
setae, setae 0.5 mm long; inner surface lightly vil¬
lous to glabrous above the stamens in a shallow
ventral pouch, glabrous below, the alae glabrous;
calyx lobes subequal, broadly triangular, 1.0 X 1.5
mm, margins of the dorsal lobe setulose-ciliolate,
rarely glabrous, the others glabrous; appendages
0.5 mm long, terminating in a short bristle. Petals
6, unequal, the 2 dorsal largest, 7—9 mm long, ob¬
long to nearly orbicular, not clawed, deep purple,
held erect or reflexed slightly back over the dorsal
side of the tube, subtended at the base by a tri¬

angular to comiculate, thickened scale free at the
tip for 0.2 mm, the 4 ventral petals 3-5 mm, oblong
to obovate, pale purple, held horizontally. Stamens
11, the 2 dorsalmost stamens shortest, included, 9
inserted at ca. 2-3 mm below the rim of the floral
tube, exserted; filaments probably pale purple, gla¬
brous; anthers purple. Pollen oval-triangular to
nearly spherical, pores very slightly protruding; tri-
colporate, syncolpate; exine distinctly and uniform¬
ly striate, striae covering the surface of the grain,
and possibly extending to the poles without disag¬
gregation; diameter 18-20 gm in lactic acid. Style
ultimately well-exserted; stigma punctiform; ovules
8-10. Seeds 2 X 1.5 mm, oblong, brown, margin
thick, rounded. Disc 1 mm long, 0.5 mm wide at
base of ovary, thick, triangular in outline, deflexed
into the spur.

Phenology. Collected in flower and fruit in
June, September, and December.

Distribution and habitat (Fig. 2). Southeastern
Guatemala, Honduras, and El Salvador; pine for¬
ests; 900—1640 m.

Etymology. The specific epithet refers to the
red-purple, bulbous setae that arm the floral tube
and upper bracts of the inflorescence.

The species is placed in section Diploptychia by
the presence of the key character of the section,
the alae, two wings of tissue that extend from the
base of the dorsalmost stamens in the floral tube to
the base of the tube (Koehne, 1903). Cuphea ar-
mata bears close resemblance to Cuphea pinetorum
Bentham and C. avigera Robinson & Seaton but
differs from both by its unequal, bicolored petals.
The prominent reddish, bulbous-based setae and
pale purple floral tube further distinguish it from
C. pinetorum, which has a white to rose, hirsute and
pubescent floral tube. Cuphea armata resembles C.
avigera especially by its conical, ascending spur,
but lacks the glandular-viscous patches on the
stems that distinguish C. avigera. Neither C. pine¬
torum nor C. avigera has been collected within the
range of C. armata. Because agamospermy with ac¬
companying infertile pollen is present in several
species of the section, pollen of C. armata was test¬
ed for pollen viability. In cotton blue-lactic acid
after 24 hours, 99% of all pollen stained deeply,
indicating viable protoplasm. The pollen exine
seems to lack the meshwork of striae at the poles
that is typical of the pollen group to which C. avi¬
gera and C. pinetorum pollen types belong (Gra¬
ham, in prep.), but this needs to be confirmed by
SEM observations when more abundant material
becomes available.

Paratypes. EL SALVADOR. Chalatenango: carretera

276

Novon

Figure 3. Distribution and variation in leaf shape, size, and petiole length in Alzatea verticillata. Examples 1-5
represent subspecies amplifolia, characterized by large, broad, oblong, sessile or subsessile leaves. Examples 6-8
represent subspecies verticillata, characterized by small, elliptical, oblong or obovate, distinctly petiolate leaves. Petioles
on plants represented by examples 5 and 6 are approximately intermediate between the two subspecies. The major
distinction remaining between the subspecies is leaf size, especially leaf width.

a La Palma, Montalvo 3682 (MO); Rfo Nunuapa near La
Palma, R. Seiler 574 (F). GUATEMALA. Chiquimula:
pine forest between Guatemala-Honduras border and Atu-
lapa, Molina R. * Molina 25260 (F, MO). HONDURAS.
Intibuca: 12.7 mi. N of Marcala along the road from Mar-
eala to La Esperanza, Davidse et al. 35191 (MO).

Note on Alzateaceae

The New World tropical family Alzateaceae is
represented by the single species Alzatea verticil¬
lata Ruiz & PavtSn. The species had been placed
in eight families in five orders, including the Lyth-
raceae, before its segregation to the monotypic fam¬
ily Alzateaceae (Graham, 1984). Alzatea verticillata
was described first from the eastern side of the Pe¬
ruvian Andes in 1798 (Ruiz & Pavdn, Syst. Veg.
FI. Peruv. Chil. 1: 72). It was considered restricted
to Bolivia and Peru until 1936, when it was col¬
lected in the low montane forests of Costa Rica. In
1978, it was discovered in the elfin cloud forest of
Veraguas Province, Panama, and subsequently, in

1986, was found in Colombia on the Cerro del Tor-
rd, Departamento del Choco (Silverstone-Sopkin &
Graham, 1986). The distribution of the species is
now further enlarged by collections made by the
late Alwyn Gentry on his last field expedition to
Ecuador. Two collections. Gentry 80008 and 80334
(MO), from the Cordillera del Condor, near the dis¬
puted Ecuador-Peru border in Morona-Santiago
Province, represent the first record of the family in
Ecuador, and tenuously establish its continuous
distribution in mid- to low-montane forests along
the eastern slopes of the mountains from Bolivia to
Costa Rica (Fig. 3). The species is one of an esti¬
mated 26% of Ecuadorian species restricted to
mid-elevations (900-3000 m) with widespread dis¬
tribution in the Neotropics (Balslev, 1988).

The two Ecuadorian collections differ in leaf
shape and size, and represent two subspecies. Gen¬
try 80334, collected from an 18-m-tall tree at 2100
m, is typical of the elliptical to oblong or obovate,

Volume 5, Number 3
1995

Graham

Cuphea

277

petiolate, small-leaved subspecies verticillata from
Peru and Bolivia (Fig. 3). Gentry 80008, collected
from a 5-m-tall tree at 1350 m, has the broad, ob¬
long, large leaves of subspecies amplifolia, until
now known from Colombia, Panama, and Costa
Rica. Both collections have short-petiolate leaves
and are intermediate for this character between the
distinctly petiolate-leaved subspecies verticillata
and the sessile-leaved subspecies amplifolia. The
distinctions between the subspecies, which were
described before the species was known from Co¬
lombia or Ecuador (Graham, 1984), are diminished
by the partial intermediacy of the Gentry collec¬
tions. If future collections of Alzatea from northern
Peru and Ecuador sample a fuller range of leaf
shape and size, it should be easy to decide whether
or not to continue recognition of these intraspecific
taxa.

Leaves of Gentry 80334, among all collections of
Alzatea, are the first to display apical leaf glands.
The glands are similar to the hydathodes and extra¬
floral nectaries described for several other genera
of the Myrtales (Ross & Suessenguth, 1926; Rao &
Chakraborti, 1982; Turner & Lersten, 1983; Gra¬
ham, 1984). The shiny, hardened secretion at the
Alzatea leaf apex suggests the secretory product is
a sugar solution and the gland is probably an extra¬
floral nectary.

Specimens examined. ECUADOR. Morona-Santia-
go: Gualaquiza Cantdn, Cordillera del Cdndor, ridge top
above Banderas, primary forest, 03°28'S, 78°22'W, pri¬
mary forest, 17 July 1993, A. Gentry 80008 (MO); Cam-

pamento Achupalla, 15 km E of Gualaquiza, 03 ; 27'S.
78°22'W, dense tangled scrubby forest, 22 July 1993, A.
Gentry 80334 (MO).

Acknowledgments. I thank Ishmael Calzada for
his energetic assistance in the field during our un¬
successful attempts to relocate C. alaniana. I am
also grateful to Ronald Liesner, who first deter¬
mined the Alzatea collections, and whose intimate
knowledge of plant families has resulted in spe¬
cialists everywhere receiving many interesting and
important specimens that otherwise might have
gone undetermined or unnoticed among exsiccatae.
Wendy Mahon-Hils prepared the illustrations in
Figure 1. This research was supported by NSF
grant DEB 8806523.

Literature Cited

Balslev, H. 1988. Distribution patterns of Ecuadorean
plant species. Taxon 37: 567—577.

Graham, S. A. 1984. Alzateaceae, a new family of Myr¬
tales in the American tropics. Ann. Missouri Bot. Gard.
71: 757-779.

Koehne, E. 1903. Lythraceae. Pp. 1-326 in A. Engler
(editor). Das Pflanzenreich, IV. 216. Heft 17. H. R. En-
gelmann, Leipzig.

Rao, T. A. & S. Chakraborti. 1982. A little looked at
attribute of the leaves of Sonneratia caseolaris (L.). Curr.
Sci. 51: 303-305.

Ross, H. & K. Suessenguth. 1926. Das Apikalorgan der
Blatter von Lafoensia. Flora 120: 1—18.
Silverstone-Sopkin, P. A. & S. A. Graham. 1986. Alza¬
teaceae, a plant family new to Colombia. Brittonia 38:
340-343.

Turner, G. W. & N. R. Lersten. 1983. Apical foliar nec¬
tary of pomegranate (Punica granatum: Punicaceae).
Amer. J. Bot. 70: 475-480.

Muhlenbergia aguascalientensis (Poaceae: Chloridoideae:
Eragrostideae), a New Species from Mexico

Yolanda Herrera-Arrieta

CIIDIR Unidad Durango, COFAA-Instituto Politecnico Nacional, Apartado Postal 738,
Durango, Dgo., 34000 Mexico

Margarita De la Cerda-Lemus

Centro Basico, Universidad Autonoma de Aguascalientes Avenida Universidad s/n,
Aguascalientes, Ags., 20100, Mexico

Abstract. Muhlenbergia aguascalientensis, from
Sierra Frfa, Aguascalientes, Mexico, is described
and illustrated. A tabular comparison with its re¬
lated species, M. durangensis Herrera and M. mich-
isensis Herrera & Peterson, is given.

The genus Muhlenbergia Schreber (Poaceae,
Chloridoideae) is comprised of over 160 species
(Clayton & Renvoize, 1986), most of which are dis¬
tributed in North and South America. In the south¬
western United States and northern Mexico, species
of Muhlenbergia make up about 25% of the grass¬
land flora and provide a considerable amount of
forage for livestock (Cronquist et al., 1977). The
majority of species are xeromorphic, and a large
percentage of the Mexican species are endemic
(44% of the M. montana (Nuttall) Hitchcock com¬
plex; Herrera-Arrieta & Grant, 1993). A distinct
taxon belonging to this complex was found during
an examination of herbarium specimens at HUAA
to complete the monographic descriptions of the
Muhlenbergia montana complex, and to make a Ho¬
listic study of the grasses from Aguascalientes. Ad¬
ditional field collections were made in the Sierra
Fria, Aguascalientes, Mexico, where the species
was first collected. The new species is named after
this region.

Muhlenbergia aguascalientensis Herrera & De
la Cerda, sp. nov. TYPE: Mexico. Aguasca¬
lientes: Mpio. de San Jos6 de Gracia, NW of
Aguascalientes, 12 km NW from La Congoja,
Playa Mariquitas, Monte Grande de la Sierra
Frfa, 2850 m, 22°15'6.7"N, 102°37'24.9"W, 8
Nov. 1986, G. Garcia 2801 (holotype, HUAA;
isotypes, CIIDIR, ENCB. IEB). Figure 1.

A Muhlenbergia michisensis culmis 30-58 cm altis, lig-
ulis 0.5-0.7(-l) mm longis, auriculis 1—2(—6) mm longis;
laminis basalis 5-10(-15) cm longis; paniculis 5-8(-15)

cm longis; spiculis 5-6.5 mm longis; aristis 2—3.5 cm lon¬
gis; antheris 2.4—3 mm longis differt.

Caespitose perennial, rhizomatous, 30-58 cm
tall. Sheaths up to 1.5 cm long, glabrous to sca-
berulous, striate, shorter than the intemodes, with
hyaline margins, elongated toward the blade form¬
ing auricles l-2(-6) mm long. Ligules 0.5-0.7(-l)
mm long, membranous, hyaline, fimbriate, rounded.
Basal blades 5—10(—15) cm long, 1-2.5 mm wide,
scaberulous, flat to folded, the margins and central
rib sclerosed, scabrous. Panicles 5—8(—15) cm long,
2-4 cm wide, open, loosely flowered, pyramidal,
largely exserted, olive, the ascending branches ap-
pressed to spreading at maturity, 20—40° from the
culm axis, 2—5 cm long; pedicels 1-2.5 mm long,
scabrous; inflorescence branches 2—5 cm long; cen¬
tral axis 8-ribbed at the base of the panicle, sca¬
brous. Spikelets 5—6.5 mm long, erect, 1-flowered.
Glumes 5-6.5 mm long, oblanceolate, entire, longer
than the lemma, usually equal in length, the apex
sharply apiculate to mucronate, the mucron 0.5-1
mm long, 1-nerved, olivaceous; second glume with
the 2 lateral nerves inconspicuous, scabrous along
the entire length. Lemma 4.5-5 mm long, oblong-
lanceolate, hyaline to yellowish, awned, glabrous to
shortly pilose on the lateral nerves at base; apex
acute, minutely bifid, the lobes to 0.2 mm long; awn
2-3.5 cm long, flexuous, yellowish. Palea 4.2-4.8
mm long, oblanceolate, entire, glabrous to pilose on
the nerves at the base. Anthers 2.4-3 mm long,
purplish. Caryopsis not seen. Chromosome number
2n = 20.

Distribution and habitat. Muhlenbergia aguas¬
calientensis is known from northwestern Aguasca¬
lientes, northwest of La Congoja, San Jose de Gra¬
cia, at 2850 m. It occurs in grassland in open forest
of Juniperus deppeana Steudel and Quercus side-
roxyla Humboldt & Bonpland.

Muhlenbergia aguascalientensis resembles M.

Volume 5, Number 3
1995

Herrera-Arrieta & De la Cerda-Lemus
Muhlenbergia aguascalientensis

279

Figure 1. Muhlenbergia aguascalientensis Herrera & De la Cerda ( G. Garcia 2801). A. Habit. B. Ligule. C.
Inflorescence branch. —D. Spikelet. —E. Glumes. —F. Lower glume, dorsal view. —G. Floret. —H. Lemma, lateral
view. —I. Palea, dorsal view.

280

Novon

Table 1. Characters distinguishing Muhlenbergia aguascalientensis, M. durangensis,

and M. michisensis.

Characters

M. durangensis

M. aguascalientensis

M. michisensis

Rhizomes

Present

Present

Absent

Ligule apex

Truncate-fimbriate

Truncate-fimbriate

Acuminate

Ligule length

0.5—0.7(-l) mm

0.5—0.7(-l) mm

4—7 mm

Leaves

Caulinar

Basal

Caulinar

Panicle length

10—25 cm

5—8(—15) cm

12-17 cm

Spikelet length

6-7(-7.5) mm

5-6.5 mm

4.5-5 mm

Glume length

(5—)6—7(—7.5) mm

5-6.5 mm

4.5—5 mm

Lemma length

5-6.7(-7) mm

4.5-5 mm

3.5—4 mm

Lemma apex

Entire

Minutely bifid

Minutely bifid

Lemma lobes length

0.0 mm

0.2 mm

0.2 mm

Lemma awn length

15—20 mm

20-35 mm

12-13 mm

Palea length

5-6 mm

4.2-4.8 mm

2.8-3.5 mm

Palea apex

Entire-apiculate

Entire

Bilobed

Anther length

(2.5—)3—3.5 mm

2.4—3 mm

2—2.4 mm

michisensis Herrera & Peterson and M. durangensis
Herrera; however, these two species differ in many
morphological characteristics (Table 1). Muhlenber-
gia aguascalientensis is similar to M. flaviseta
Scribner, a species of restricted distribution in the
Sierra Madre Occidental, sharing with it macro-
morphological features such as basal leaves and a
long-exserted panicle, but differing in most of the
spikelet features.

The flavonoid chemistry of M. aguascalientensis
is very similar to M. eriophylla Swallen and M.
michisensis Herrera & Peterson. They share: quer¬
cetin 3-0 rhamnosilxyloside, luteolin 6 galactoside,
luteolin 6 glucoside, luteolin 8 glucoside, apigenin
7-0 arabinoside, apigenin 7-0 diarabinoside, api¬
genin 7-0 glucoside, tricin 5-0 glucoside, tricin 7-
0 glucuronide, and 4'hydroxyflavone 7-0 glucoside
(Herrera & Bain, 1991). Muhlenbergia aguascalien¬
tensis also contains luteolin 6 xylosilglucoside,
which was previously thought to be a unique com¬
pound of M. durangensis.

Paratypes. MEXICO. Aguascalientes: Playa Mari-
quitas. Sierra Frfa, 12 km NW from La Congoja, San Jos6
de Gracia, 22°15'6.7"N, 102°37'24.9"W, 2850 m, 25 Nov.
1993, Herrera & De la Cerda 1185 (CIIDIR, HUAA, IEB,
ENCB, MEXU, US).

Acknowledgments. This study was partially
supported by the Banco de Mexico, COFAA-IPN
and UAA.

Literature Cited

Clayton, W. D. & S. A. Renvoize. 1986. Genera Gramin-
um. Grasses of the World. Kew Bull., Addit. Ser. XIII:
1—389. Her Majesty’s Stationery Office, London.
Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L.
Reveal & P. K. Holmgren. 1977. Intermountain Flora.
Vol. 6, The Monocotyledons. Columbia Univ. Press,
New York.

Herrera A., Y. & J. F. Bain. 1991. Flavonoid profiles in
the Muhlenbergia montana complex (Poaceae). Bio-
chem. Syst. Ecol. 19: 665-672.

Herrera-Arrieta, Y. & W. F. Grant. 1993. Correlation be¬
tween generated morphological character data and fla¬
vonoid content of species in the Muhlenbergia montana
complex. Canad. J. Bot. 71: 816-826.

New Taxa and Combinations in the Tribe Juanulloeae (Solanaceae)

Sandra Knapp

Department of Botany, The Natural History Museum, Cromwell Road,
London SW75BD, United Kingdom

ABSTRACT. A new species of Trianaea from east¬
ern Ecuador, T. naeka, is described and illustrated.
Several new combinations in the genera Trianaea,
Schultesianthus, and Juanulloa are made in prep¬
aration of a larger work on the phylogeny of the
tribe Juanulloeae.

A recent phylogenetic study of the tribe Juan¬
ulloeae (Persson et al., 1994) has necessitated sev¬
eral new combinations in the genera Trianaea,
Schultesianthus, and Juanulloa. These nomencla-
tural changes are made here in preparation for a
conspectus of the tribe and a more ample discus¬
sion of relationships within the groups. In addition
to the nomenclatural changes identified, several
new taxa were discovered, one of which is de¬
scribed here. Other new taxa will be described as
more material becomes available.

Trianaea naeka S. Knapp, sp. nov. TYPE: Ecuador.
Morona Santiago: Colinas, above the Mision Sa-
lesiana-Shuar, 800-1000 m, 3°25'S, 78°35'W,
6 Nov. 1986, Neill & Cerdn 7449 (holotype,
MO; isotypes, NY, QCNE). Figure 1.

Species Trianaeae nobili Planchon & Linden similis,
sed foliis linearis pubescentibus, inflorescentiis uniflori-
bus differt.

Epiphytic shrub or liana at forest edge; stems
densely hirsute, the trichomes 4—5 mm long, con¬
sisting of a simple, 2-4-celled, uniseriate tip aris¬
ing from a multicellular, multiseriate, pustular
base, the apex minutely glandular, golden brown,
the uniseriate tips of the trichomes often deciduous,
leaving the pustulate base; bark of older stems pale
tan, loose and somewhat exfoliating. Leaves 0.4—1
X 5-10 cm, linear, coriaceous, drying dark olive
green, the venation obscure, ca. 8—10 primary
veins, densely hirsute with simple, uniseriate, mi¬
nutely glandular trichomes like those of the stems,
but lacking the pustular bases, arising directly from
the epidermis, the base attenuate, the margins rev¬
olute, the apex acuminate; petiole 5—8 mm, some¬
what winged from the attenuate leaf base, drying
dark brown. Inflorescence terminal, consisting of a
single flower. Buds not known. Pedicel at anthesis

18-20 cm long and flexuous, 2 mm diam. at the
base, ca. 10 mm diam. at the apex, densely pubes¬
cent with simple uniseriate 2—4-ee]|ed trichomes,
the trichomes 4—5 mm, less rigid than those of the
stems, lacking pustulate bases. Flowers with the
calyx tube 1.5-2 cm, conical, the lobes 4—5 cm,
triangular, pubescent with simple uniseriate tri¬
chomes like those of the pedicel, sparsely pubes¬
cent within; corolla campanulate, greenish brown
or purplish (“clear lila” fide van der Wetff & Gudifio
11246), the tube flaring, ca. 5 cm long, the lobes
rounded, 1.5—2 cm long, reflexed at anthesis,
sparsely pubescent with trichomes like the calyx,
the trichomes weak and crumpling; stamens with
the filaments ca. 3.5 cm, inserted in the basal Vs of
the corolla tube, the anthers ca. 7X3 mm, ven-
trifixed; ovary glabrous, the style 5.5-6 cm long,
glabrous, the stigma globose, ca. 5 mm diam., pa¬
pillose. Fruit and seeds not known.

Distribution. Known only from wet premontane
rainforest in eastern Ecuador, from 800 to 1000 m
elevation.

Trianaea naeka is a striking plant and cannot be
confused with any other species of Trianaea, or with
any other species of epiphytic Solanaceae. The nar¬
rowly linear, densely pubescent leaves are found
nowhere else in the tribe. Pollen morphology is of
the Trianaea nobilis type (Persson et al., 1994),
where the exine ornamentation is reticulate at the
equator and at the poles but the muri are not joined
in the mesocolpium to form a tectum.

The specific epithet, naeka, is the Shuar name
(given by Antonio Wampash, Shuar informant) for
this plant.

Paratype. ECUADOR. Morona Santiago: along new
road Mendez-Morona, km 30—35, 800 m, 19 Aug. 1989,
van der Werjf A' Gudifio 11246 (QCNE).

Trianaea brevipes (Cuatrecasas) S. Knapp, comb,
et stat. nov. Basionym: Trianaea spectabilis
Cuatrecasas var. brevipes, J. Wash. Acad. Sci.
49: 271. 1959. TYPE: Colombia. Cundina-
marca: Monterredondo between Guaybetal and
Limoncito, Cordillera Oriental, forests, 1800—
2100 m, 6 July 1955, Vogel 6 (holotype, US;
isotype, M).

282

Novon

MISSOURI

BOTANICAL GARDEN
HERBARIUM

N? 4355395

Figure 1. Holotype of Trianaea naeka S. Knapp (Neill & Cerdn 7449, MO).

Volume 5, Number 3
1995

Knapp

Tribe Juanulloeae (Solanaceae)

283

Schultesianthus venosus (Standley & C. V. Mor¬
ton) S. Knapp, comb. nov. Basionym: Markea
verwsa Standley & C. V. Morton, Publ. Field
Mus. Nat. Hist., Bot. 18: 1065. 1938. TYPE:
Costa Rica. Alajuela: Vara Blanca de Sarapi-
quf, 1770 m, Oct. 1937, Skutch 3355 (holo-
type, US 1643419).

Schultesianthus crosbianus (D’Arcy) S. Knapp,
comb. nov. Basionym: Markea crosbiana
D’Arcy, Ann. Missouri Bot. Gard. 60: 654.
1973 [1974], TYPE: Panama. Chiriquf: Cerro
Horqueta NW of Boquete, 5000-5800 ft.,
Dwyer et al. 430 (holotype, MO).

Schultesianthus uniflorus (Lundell) S. Knapp,
comb. nov. Basionym: Markea uniflora Lun¬
dell, Phytologia 1: 340. 1939. TYPE: Mexico.
Chiapas: Volcan Tacana, 2800 m, 27 Mar.
1939, Matuda 2816 (holotype, MICH; iso¬
types, MEXU, MO).

Juanulloa wardiana (D’Arcy) S. Knapp, comb,
nov. Basionym: Rahowardiana wardiana
D’Arcy, Ann. Missouri Bot. Gard. 60: 671.
1973 [1974]. TYPE: Panama. Cocld beyond
La Mesa towards El Llano and the Panama/
Code border N of El Valle de Anton, 800-860
m, Luteyn 3150 (holotype, DUKE, isotype,
MO).

Juanulloa globifera (S. Knapp & D’Arcy) S.
Knapp, comb. nov. Basionym: Rahowardiana
globifera S. Knapp & D’Arcy, Novon 3: 429.
1993. TYPE: Colombia. Antioquia: Parque
Nacional Natural “Las Orquideas,” Sector
Venados, 890 m. 6°33'N, 76°19'W, 1 Apr.
1988, Cogollo et al. 2895 (holotype, JAUM-
018167; isotypes, COL, FMB, JAUM, MO).

Acknowledgments. I thank Viveca Persson and
Steve Blackmore, my coworkers on pollen mor¬
phology of Solanaceae; Roy Vickery and the cura¬
tors of the herbaria mentioned in the text for help
with the loan of specimens; Tamara Nunez and Da¬
vid Neill for asistance at QCNE; and the Photo¬
graphic Unit of the Natural History Museum for the
photographic plate.

Literature Cited

Persson, V., S. Knapp & S. Blackmore. 1994. Pollen mor¬
phology and systematics of tribe Juanulloeae A. T. Hun-
ziker (Solanaceae). Rev. Palaeobot. Palynol. 83: 1-30.

A New Species of Sicydium Schlechtendal
(Cucurbitaceae: Zanonioideae, Zanonieae, Sicydiinae) for the
Flora Mesoamericana

Rafael Lira

Herbario Nacional de Mexico, Departamento de Botanica, Instituto de Biologfa, UNAM,
Apdo. Postal 70-367, Delegacion Coyoacdn, 04510, Mexico, D.F.

Abstract. Sicydium daviliae, from the region of
Ocozocuautla, Chiapas, Mexico, is described as
new. It can be distinguished from the other species
in the genus by the presence of a soft and dense
pubescence composed of long, multicellular, non-
glandular hairs covering stems, blade surfaces, pet¬
ioles and tendrils, by filaments 2—3 times longer
than anthers, by petal margins entire, and by sta-
minate pedicels articulate at the middle or further
up.

Resumen. Se describe como nueva especie a Si¬
cydium daviliae, de la region de Ocozocuautla,
Chiapas, Mexico. Esta nueva especie se distingue
de las otras especies del genero por la pubescencia
densa y suave, compuesta de tricomas largos, mul-
ticelulares y no glandulares, la cual cubre los tal-
los, ambas superficies de la lamina, los peciolos y
los zarcillos, ast como tambi£n por presentar fila-
mentos 2-3 veces mas largos que las anteras, pe¬
tal os con los mdrgenes enteros y pedicelos estam-
inados articulados en la mitad o poco mas arriba.

As a result of examination of herbarium speci¬
mens for the treatment of Sicydium for Flora Me¬
soamericana, one specimen was found whose sta¬
men and vestiture features set it apart as a new
species, described below.

Sicydium daviliae Lira, sp. nov. TYPE: Mexico.
Chiapas: Mpio. Oeozocuauda, Camino Horizon¬
te a Santa Laura, 12 June 1983, F. Vdzquez B.
1024 (male plant) (holotype, XAL). Figure 1.

Sicydio tamnifolio (HBK) Cogniaux similis, sed differt
staminum filamentis quam antheris 2—3 plo longioribus,
foliorum, caulium, petiolorumque indumento densiore, tri-
chomatibus longis apice non glandularibus constato, pe-
talorum marginibus integris, florum masculorum pedicellis
in parte media vel altiore articulatis.

Herbaceous vine, perennial, dioecious, drying
yellowish green; stems, petioles, upper and lower
surfaces, and tendrils densely pubescent to tomen-
tose, with long (1—2 mm), soft multicellular, non-

glandular hairs. Stems angulate-sulcate. Petioles
angulate-sulcate, l-1.5(-2) cm long; blades
subcoriaceous, ovate-cordate, 3.5—5.5 cm long, 3—
5 cm broad, entire, apex acute to shortly acuminate,
margins entire to slightly undulate-dentate, basal
sinus cordate. Tendrils simple. Flowers small, ar¬
ranged in loose panicles. Staminate panicles 4-6
cm long, the branches commonly with a reduced
leaf in the axils; pedicels filiform, 1—1.3 mm long,
articulate at the middle or a little above the middle,
subtended by a lanceolate reduced leaf; receptacle
disk-shaped, less than 0.2 mm long, 1-1.2 mm
wide, puberulent; sepals 5, triangular, ca. 0.6 mm
long, 0.3 mm wide, puberulent; petals 5, ovate-lan¬
ceolate, spreading, greenish cream or greenish
white, 1—1.3 mm long, 0.5-0.6 mm wide, puberu¬
lent, the margins entire; stamens 3, totally free, two
with bithecial anthers, one with a monothecial an¬
ther; filaments inserted in the base of the recepta¬
cle, 0.4-0.6 mm long, slightly bent inward, forked
toward the apex in the stamens with bithecial an¬
thers; anthers oblong-elliptic, 0.2—0.3 mm long,
opening by a longitudinal suture. Plants with pis¬
tillate flowers and fruits unknown.

Sicydium daviliae is known only from the type,
collected in the Municipio Ocozocuautla, Chiapas,
Mexico, growing in disturbed tropical rainforest at
900 m, on sandy, clayish, and rocky soils.

The approximately eight species of Sicydium are
distributed in the Neotropics. They have been tra¬
ditionally distinguished mainly on the basis of sta¬
men structure, and presence/absence and distri¬
bution of vestiture (Cogniaux, 1881, 1916; Dieterle,
1976; Jeffrey & Trujillo, 1992; Nee, 1993; Schle¬
chtendal, 1832; Wunderlin, 1978). So far, three
species were found in Mexico—Mesoamerica: S.
tamnifolium (HBK) Cogniaux, a species ranging
from central Mexico to South America, and S.
tuerckheimii J. Donell-Smith and S. schiedeanum
Schlechtendal (= S. glabrum Standley & Steyer-
mark), both ranging at least from southern Mexico
to Central America.

Volume 5, Number 3
1995

Lira

Sicydium daviliae

285

0.5 mm

Figure 1. Sicydium daviliae Lira. —A. Habit. —B. Staminate flower. —C. Comparison among the tnchomes present
on the leaves of S. daviliae (1), S. tuerckheimii (2), and S. tamnifolium (3). Based on type specimen F. Vdzquez B. 1024
(XAL).

286

Novon

Sicydium daviliae can readily be distinguished
from the above-mentioned species by its unique
combination of stamens with filaments 2-3 times
longer than the anthers; petals with entire margins;
a dense, soft indument made up of long, multicel¬
lular, nonglandular hairs; and staminate pedicels
that are articulate at or above the middle. In con¬
trast, Sicydium tamnifolium includes populations
with leaf blades, stems, and petioles densely and
softly pubescent, but the pubescence is less dense
and the hairs are much shorter and gland-tipped.
In addition, the stamens of S. tamnifolium have
very short or obsolete filaments, the petal margins
are somewhat fimbriate or laciniate, and the pedi¬
cels of the staminate flowers are articulate near the
base.

A species with a similar vestiture to that of Si¬
cydium daviliae is S. araguense Steyermark & Tru¬
jillo from Venezuela (Jeffrey & Trujillo, 1992).
However, in S. araguense the stems are only pu-
berulent with age, in adult leaves the pubescence
is not as dense or soft, and the leaf blades are nar¬
rowly lanceolate and longer. The new species is
named to honor Patricia Davila, Mexican agrostol-
ogist.

A Key to the Mexican/Mesoamerican Species of
Sicydium

la. Plants essentially glabrous or only slightly pu-

berulent; leaves always unlobed, usually lustrous
in both surfaces; filaments 2-2.5 times longer
than anthers.5. schiedeanum Schlechtendal

lb. Plants conspicuously soft-puberulent, pubescent,
or tomentose at least on the lower surface of the
leaves; leaves unlobed to angulate or slightly lo-
bate, not lustrous; filaments obsolete to 3 times
longer than anthers.

2a. Leaves pubemlent and rough-scabrous on
the upper surface, densely and softly pubes¬
cent on the lower surface, the pubescence
commonly dark reddish brown when dry; fil¬
aments 2-3 times longer than anthers . . .

.S. tuerckheimii J. Donell-Smith

2b. Leaves pubescent, puberulent to tomentose
on both surfaces, the pubescence yellowish

or whitish in dry specimens; filaments ob¬
solete to 2—3 times longer than anthers.

3a. Plants puberulent to villous, the vesti¬
ture mainly of glandular hairs; filaments
obsolete; margins of the petals slightly
fimbriate or laciniate; pedicels of the
staminate flowers articulate near the
base . ... S. tamnifolium (HBK) Cogniaux
3b. Plants densely and softly pubescent to
tomentose, the vestiture mainly of non¬
glandular hairs; filaments 2—3 times
longer than anthers; margins of the pet¬
als entire; pedicels of the staminate
flowers articulate in the middle or
slightly above. S. daviliae Lira

Acknowledgments. I thank the curators of BM,
BR, CHAPA, ENCB, F, HUH, K, LL, MICH, MO,
NY, P. TEX, US, and XAL for allowing me to review
collections of Sicydium, especially Sergio Avendano
(XAL) for making available the specimen on which
the description of the new species is based. Eloisa
Duarte prepared the excellent drawings, and David
Sutton (BM), Fernando Chiang, and two anonymous
reviewers commented critically on the manuscript;
Fernando Chiang also prepared the English version
of the paper, as well as the Latin diagnosis.

Literature Cited

Cogniaux, A. 1881. Cucurbitacfes. Pp. 325-951 in A.
de Candolle & C. de Candolle (editors), Monographiae
Phanerogamarum. G. Masson, Paris.

-. 1916. Cucurbitaceae—Fevilleae et Melothrieae.

Pp. 1—277 in A. Engler (editor). Das Pflanzenreich.
Regni Vegetabilis Conspectus. Verlag von Wilhelm En-
gelmann, Leipzig.

Dieterle, J. V. A. 1976. Cucurbitaceae. In: Flora of Gua¬
temala. Part XL Fieldiana Bot. 24: 306-395.

Jeffrey, C. & B. Trujillo. 1992. Cucurbitaceae. Pp. 11-
201 in G. Morillo (editor). Flora de Venezuela. Fondo
Editorial Acta Cientffica Venezolana, Caracas, Venezue¬
la.

Nee, M. 1993. Cucurbitaceae. In: V. Sosa (editor). Flora
de Veracruz. Fasctculo 74. Instituto de Ecologfa A.C./
University of Riverside, Xalapa, Veracruz.
Schlechtendal. D. F. L. 1832. De Plantis Mexicanis. Lin-
naea 7: 380-400.

Wunderlin, R. P. 1978. Cucurbitaceae. In: R. E. Wood-
son, Jr. & R. W. Schery (editors). Flora of Panama. Part
IX. Ann. Missouri Bot. Card. 65: 285—366.

Conceveiba parvifolia (Euphorbiaceae), a New Species from
Panama and Colombia

Gordon D. McPherson

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ABSTRACT. Conceveiba parvifolia from Darien,
Panama, and Antioquia, Colombia, is described
and illustrated.

Two wind-thrown trees encountered on the for¬
ested slopes above the abandoned Cana gold mine
in Darien are apparently the first known represen¬
tatives of the genus Conceveiba Aublet from Pana¬
ma, although the genus was previously known from
both neighboring Costa Rica and Colombia. Spec¬
imens from these two trees key readily to the sub¬
tribe Conceveibinae of the Alchomeae (Acalyphoi-
deae) in Webster’s synopsis (1994). Within that
subtribe, however, the Panamanian collections
combine characters currently used to distinguish
Conceveiba and Gavarretia Baillon, i.e., on the one
hand the sepals of the pistillate flower are separate
(usually), the style branches blunt, and the anthers
muticous (as in Conceveiba), while on the other
hand the sepals of the pistillate flower are eglan-
dular, and the ovary is 2-locular (as in Gavarretia,
although some species of Conceveiba are similarly
eglandular). The generic boundaries were not clear
in this subtribe even before this intermediate taxon
further blurred them, and I have chosen to name
the new species in Conceveiba since I agree with
Webster (1994) that Mueller’s inclusion (1874) of
Gavarretia as a section of Conceveiba may be jus¬
tified.

Conceveiba parvifolia McPherson, sp. nov.
TYPE: Panama. Darien: near Cana mine S of
El Real (Cerro Pirre massif), 7°45'N, 77°40'W,
ca. 900 m, 26 Aug. 1987. McPherson 11607
(holotype, PMA; isotypes, BM, COL. CR, DAV.
EAP, F, G, GH. HUA. K, MEXU, MICH, MO,
NY, P, US). Figure 1.

Species foliis minoribus (2.7—16.5 cm X 0.8-5.5 cm),
infiorescentiis masculis brevibus (10 cm) fere eglandula-
tis, floribus masculis noil fasciculatis, sepalis floris femi-
nei 4(—5) eglandulatis, ovario biloculari, stylis 2 semicon-
natis a congeneribus diversa.

Tree (6-) 15-20 m; colored exudate absent; youn¬
gest branches stellate-puberulent, without lenticels,
terete or somewhat ridged, the older ones glabrate

and bearing elongated lenticels. Leaf blade (2.7—)

3.5- 8.0(-16.5) cm long, (0.8-)1.8-4.0(-5.5) cm
wide, elliptical-oblong, obtuse at the base, acumi¬
nate at the apex, the margin crenate (sometimes
obscurely so) with 5—8 mucronate teeth, the upper
surface sparsely stellate-puberulent before maturi¬
ty, rapidly glabrate, the lower surface densely stel¬
late-puberulent before maturity, eventually gla¬
brate, often with 1—several embedded glands,
secondary veins 6—10 on each side of the midrib,
the reticulum slightly raised on the upper surface
and more strongly raised on the lower; petiole 1.3—
4.0(—4.8) cm long, 0.5-1.0 mm diam., semicircular
in cross section, sparsely stellate-puberulent, a dis¬
tal pulvinus usually discolored and thereby distin¬
guishable, 2-3 mm long, sometimes slightly swol¬
len, a smaller proximal pulvinus often present;
stipules subulate, 2—4 mm long, up to 0.5 mm wide,
stellate-puberulent, quickly caducous. Plants di¬
oecious; inflorescences terminal or from the axils
of the most distal leaves. Staminate inflorescences
paniculate or less often racemose, up to 10 cm long,
the axes stellate-puberulent, bracts triangular, up
to 2 mm long, stellate-puberulent, each bract often
accompanied by two smaller bracteoles, both bracts
and bracteoles sometimes bearing weakly differ¬
entiated glabrous glandular patches; pedicels 1—1.5
mm, not fasciculate, glabrous; buds ca. 2 mm
diam., apiculate; sepals valvate, 2 or 3 in number,

2.5- 3.5 mm long, up to 3 mm wide, concave, eglan¬
dular, green, glabrous without, somewhat granular-
puberulent within at base of filaments; petals and
disk absent; stamens 35-42, filaments up to 3 mm
long, free or the central ones connate basally, gla¬
brous, anthers ca. 0.8 mm long, rounded, somewhat
puberulent; pistillode absent. Pistillate inflores¬
cences racemose, up to 5 cm long (occasionally the
most proximal flowers subtended by only slightly
modified leaves), the axes stellate-puberulent,
eglandular; bracts not seen, rapidly caducous; ped¬
icels 4—13(—18) mm long, somewhat inflated dis-
tally, stellate-puberulent, usually bearing two
bracteoles ca. 2 mm long, these caducous, usually
attached near the pedicel base but sometimes at¬
tached subapically and then mistakable for sepals;

Nnvniv 5- 9R7-98Q 1QQ5

288

Novon

Figure 1. Conceveiba parvifolia McPherson. —A. Twig bearing staminate inflorescences. —B. Portion of stammate
inflorescence. —C. Staminate flower. —D. Pistillate inflorescence. —E. Pistillate flower with 4 sepals and a bracteole.
(A—C based on McPherson 11621; D, E based on McPherson 11607.)

Volume 5, Number 3
1995

McPherson
Conceveiba parvifolia

289

sepals 4(-5) in number, 2-5 mm long, subulate,
free or occasionally two of them partially connate,
eglandular, pale green, stellate-puberulent; petals,
disk, and staminodes absent; pistil 6-9 mm long,
stellate-puberulent, ovary ca. 3 mm long, 2-locular;
styles 2 in number, fused basally into a column
1.5—3 mm long, free and bifid distally, these free
portions stigmatic adaxially and ca. 3 mm long in
total, the ultimate branches ca. 2 mm long, blunt-
tipped; ovules 1 per locule. Fruit unknown.

From the seven or eight other members of its
genus and from the monotypic Gavarretia, Concev¬
eiba parvifolia can be separated by its smaller
leaves, shorter and only obscurely glandular sta-
minate inflorescences, non-fasciculate staminate
flowers, and pistillate flower with 4(—5) eglandular
sepals, a bilocular ovary, and 2 styles connate half
their length.

At the suggestion of Grady L. Webster, I exam¬
ined two additional specimens that proved to be¬
long to the new species. These collections, made

by R. Callejas, F. J. Roldan, and J. Betancur of
HUA, extend the distribution of Conceveiba parvi¬
folia from easternmost Central America well into
the northwestern corner of South America.

Paratypes. PANAMA. Darien: near Cana mine S of
El Real (Cerro Pine massif), 7°45'N, 77°40'W, ca. 900 m,
26 Aug. 1987, McPherson 11621 (BM, COL, CR, DAV, F,
HUA, K, MEXU, MICH, MO, NY, PMA, US). COLOM¬
BIA. Antioquia: Municipio Anon, via Dos Bocas-Provi-
dencia, 7°26'N, 74°56'W, 100-120 m, 9 jul. 1987, Cal¬
lejas et al. 4477 (MO); Municipio Remedies, 10-12 km
NW de Remedios en la via a Zaragoza, 7°05'N, 74°15’W,
300 m, 16 sep. 1987, Callejas et al. 5261 (MO).

Acknowledgments. I thank David Olson, who
accompanied me to Cana to collect ants, and John
Myers, who drew the illustration.

Literature Cited

Mueller, J. 1874. Euphorbiaceae [part 2], Pp. 293-750
in Martius, Flora Brasiliensis, Vol. 11. Cramer, Wein-
heim.

Webster, G. L. 1994. Synopsis of the genera and supra-
generic taxa of Euphorbiaceae. Ann. Missouri Bot.
Card. 81: 33-144.

A New Votomita (Melastomataceae) from Panama

Thomas Morley

Department of Plant Biology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A.

Frank Almeda

Department of Botany, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118-4599, U.S.A.

Abstract. A new species of Votomita, V. cupuli¬
formis, has been found on the north side of western
Panama, the first member of the genus known to
occur in Central America. The new species is most
similar to V. roraimensis Morley of central Roraima,
Brazil. Distinguishing features are discussed, and
a key to all nine species of the genus is provided.

Votomita cupuliformis Morley & Almeda, sp.
nov. TYPE: Panama. Bocas del Toro: Laguna
de Chiriqui, rincon Sur-Este Nuri, campamen-
to cerca pequena poblacion Guaymi, 15 km
Oeste de Punta Cricamola, entrando Ensenada
de Catavela, y subiendo Quebrada Nuri,
8°55'N, 81°49'W, unos 5 km al Sur del cam-
pamento, 10-25 m, 18 mar. 1993, Robin Fos¬
ter, A. Herre, E. Kalko <£ C. Handley 14563
(holotype, CAS; isotypes, MO, PMA, SCZ, US).
Figures 1, 2.

Arbor glabra usque 10 m alta: cryptae stomatophorae
Typi 11. parvae, simplices; epidermis supera folii stratiis
1 cellularum; limbus calycis cupuliformis lobis apiculatis;
stamina monadelpha; ovarium 2-loculare ovuliis 16 axil-
laribus.

Glabrous tree 10 m high; young twigs rounded.
Petioles 4—6 mm long; blades 6.5-11.9 cm long,

2- 4-4.5 cm wide, elliptic to elliptic-ovate or seldom
slightly obovate, abruptly acuminate at apex, acute
to abruptly short-attenuate on the petiole at base;
midrib grooved above, narrowly 2-winged below;
lateral nerves obscurely visible to invisible above
and below when dry. Midrib xylem tubular; sto-
matal crypts Type II or borderline I—II (see Morley,
1976); upper epidermis one cell thick, lacking mu¬
cilaginous walls; hypodermis none; foliar sclereids
all terminal, variable in shape, mostly columnar
with large branches at each end, sometimes slant¬
ing, sometimes with a horizontal central body at
veinlet level bearing branches upward and down¬
ward. Peduncles terminal, axillary, and at the up¬
permost leafless nodes, 1 per side, of 1 intemode

3- 7 mm long, 1-flowered; bracts narrowly

Novon 5; 290-293. 1995.

triangular, acute, entire, 1.2—1.5 mm long, decid¬
uous at anthesis. True pedicels 8-11 mm long;
flowers known only from buds, the buds yellow;
flowers 4-merous; calyx including inferior ovary
4.5—5 mm long, obconic to slightly campanulate,
1.8-2.2 mm long, the surface flecked with scurfy
patches at least when moist; calyx limb abruptly
expanded and shallowly cup-shaped, 2.5 mm long,
ca. 3.8 mm diam.; calyx lobes 0.7 mm long, 2.5
mm wide, broadly rounded-triangular and promi¬
nently apiculate. Immature petals ovate, acute, ses¬
sile, adherent to each other in bud, 8 mm long, ca.
3.5 mm wide. Stamens monadelphous, 4.24.5 mm
long, 0.9-1.0 mm wide, adherent on the back to
the petals in bud, this condition at anthesis un¬
known; filaments ca. 0.4 mm long; anthers 3.74.2
mm long; thecae 2.4—2.7 mm long; gland adaxial
at the apex, shallowly concave, 0.7 mm long, 0.4—
0.5 mm wide. Ovary 2-locular with axile placen-
tation, each locule with ca. 8 ovules crowded at
different levels on the placenta, the ovules 16 in
all. Calyx prominent on the immature fruit; ripe
fruit and seed unknown.

Distribution. Known only from the type locality
near the southeast comer of the Laguna de Chiriqui
on the north side of western Panama. “Terra firme,
bosque alto, humedo sobre filos y pendientes.”

The new species is the first member of Votomita
known to occur in Central America. Its leaves are
overall the simplest in the genus in their average
size and shape, lack of hairs, undivided upper epi¬
dermis, strictly terminal foliar stone cells with
only a moderate columnar tendency, and small
simple stomatal crypts. The flowers, however, are
of an intermediate level of reduction from the pre¬
sumed ancestral type: in the nine species of Vo¬
tomita there is a general trend from large-flowered
plants with 4-locular ovaries and numerous ovules
to small-flowered ones with 1 locule and 5—10
ovules. Votomita cupuliformis has a 2-locular ova¬
ry with 16 ovules; in this feature the plant is most
similar to V. roraimensis of central Roraima, Bra-

Volume 5, Number 3
1995

Morley & Almeda

Votomita cupuliformis from Panama

291

Figure 1. Votomita cupuliformis Morley & Almeda, habit drawing (by Ellen del Valle).

zil, which has 4 locules and 15-16 ovules. The
new species is easily distinguished from V. rorai-
mensis, however, by the former’s longer petioles,
simpler leaf anatomy, larger petals and stamens,
and 2-locular ovary.

A key is provided to all nine species to illus¬
trate how the three named since the latest mono¬
graph (V. cupuliformis, V. pubescens, V. roraimensis)
compare to the six included in the monograph
(Morley, 1976):

la. Calyx including inferior ovary 7.5-10.5 mm long;
ovules 20-48.

2a. Leaf blades pubescent beneath, 10.5—15.5

cm long. V- pubescens Morley

2b. Leaf blades (excluding the midrib) glabrous
beneath, 7—12 cm long.

за. Petioles 3—5 mm long, placentation

axile. V. plerocarpa (Morley) Morley

3b. Petioles 7-12 mm long; placentation

basal in each locule .

. V. monadelpha (Ducke) Morley

lb. Calyx including inferior ovary 2.2—5.6 mm long;
ovules 5—16 and 36.

4a. Calyx including inferior ovary 5-5.6 mm
long; stamens 5.“T-6.1 mm long; ovules ca.

зб, in 4 whorls. V. orbinaxia Morley

292

Novon

F igure 2. Votomita cupuliformis. —A. Leaf shapes (traced). B—I (camera lucida): —B. Cleared portion of leaf blade
showing veins and terminal sclereids. —C. Cross section of leaf blade showing sclereids, stomatal crypts, and upper
epidermis. D. Cross section of leaf midrib. —E. Large flower bud. —F. Longisection of flower before anthesis. G.
Petal. —H. Anthers, face and side views. —1. Immature fruit. (All by TM.)

Volume 5, Number 3
1995

Morley & Almeda

Votomita cupuliformis from Panama

293

4b. Calyx 2.2-5 mm long; stamens 2.9—5.1 mm
long; ovules 5-16.

5a. Calyx including inferior ovary 4.5—5
mm long; ovules 15-16.

6a. Petioles 1.5-2.5 mm long; petals
ca. 4.7-5 mm long; anthers 2.6-2.8
mm long; thecae 1.2-1.3 mm long;
ovary 4-locular. . . V. roraimensis Morley
6b. Petioles 4—6 mm long; petals ca. 8
mm long; anthers 3.7—4.2 mm long;
thecae 2.4-2.7 mm long; ovary 2-

locular .

. . . V. cupuliformis Morley & Almeda
5b. Calyx 2.2^1 mm long; ovules 5-10.

7a. Apex of blade abruptly acuminate;
petioles 4—9 mm long; blades (7—)
9—12 cm long . . . V guianensis Aublet

7b. Apex of blade rounded to acute and
often mucronulate; petioles 1.5-4
mm long; blades 2.5-7.5 cm long.

8a. Blades 22—42 times as long as
the petiole; pedicels 1—1.3 mm
diam. 2 mm below the calyx

teeth when dry .

. V. orinocensis Morley

8b. Blades 10-20 times as long as
the petiole; pedicels 1.5—2.3
mm diam. 2 mm below the ca¬
lyx teeth when dry .

. . . .V. monantha (Urban) Morley

Literature Cited

Morley, T. 1976. Memecyleae (Melastomataceae). FI.

Neotrop. 15: 1—295.

New Names in Papaver Section Meconella (Papaveraceae)

David F. Murray

Herbarium (ALA), University of Alaska Museum, Fairbanks, Alaska 99775-6960, U.S.A.

Abstract. Two combinations ( Papaver macounii
subsp. discolor and P. radicatum subsp. kluanensis)
and a subspecies ( P. nudicaule subsp. americanum)
are published.

To complete a treatment of the scapose poppies
(section Meconella of Papaver) for Flora of North
America, it is necessary to publish one new com¬
bination and to validate one combination and a
subspecies that have been previously proposed, but
not in compliance with the International Code of
Botanical Nomenclature (Greuter et al., 1994).

Papaver macounii subsp. discolor (Hulten) Ran-
del ex D. F. Murray, stat. nov. Basionym: Pa¬
paver macounii var. discolor Hulten in Hulten,
FI. Alaska Yukon 5: 803. 1945. TYPE: U.S.A.
Alaska: Nome, Anderson 3250 (holotype, S).

This combination was made by Randel (1977:
425) without a full and direct reference to the bas-
ionym’s author and place of valid publication as
required by Article 32.2 (Greuter et al., 1994),
which I have now supplied.

Papaver nudicaule subsp. americanum Randel
ex D. F. Murray, subsp. nov. TYPE: [Canada.]
Klotassin area Yukon T[erritory], SW of Yukon
River, between Selkirk and White River, Aug.
1916, D. D. Cairnes 91890 (holotype, CAN).

Randel (1977: 437) determined that the speci¬
mens from Alaska and Yukon represented a taxon
distinct from the Asiatic material she had studied.
She proposed Papaver nudicaule subsp. american¬
um, for which she supplied a diagnosis but failed
to select a type as required by Article 37.1 (Greuter
et al., 1994). As holotype I have chosen a specimen
complete with leaves, flowers, and fruits, which
Randel had seen and annotated as subspecies
americanum.

Papaver radicatum subsp. kluanensis (D. Love)
D. F. Murray, comb, et stat. nov. Basionym:
Papaver kluanensis D. Love in Love & Freed¬
man, Bot. Not. 109: 178-180. 1956. TYPE:
[Canada], Yukon [Territory]: N of Quill Creek
camp, 20 mi. W of Burwash landing, alt. ca.
5000 ft., 19 June 1953, Freedman s.n. (holo¬
type, MAN).

Doris Love described Papaver kluanensis based
on specimens from southwestern Yukon, which she
later (1969) showed to be more widely distributed
southward along the Rocky Mountains. Whereas
she maintained this taxon as distinct from the Utah
populations in this complex, I am, at least for the
time being, including them within P. radicatum as
subspecies kluanensis. Specimens of subspecies
kluanensis from the type locality and elsewhere in
the very northern part of its range are taller, with
larger flowers and ellipsoid-subglobose capsules.
Plants from the southernmost part of the range (Col¬
orado, Utah, and New Mexico) are consistently
shorter, with smaller flowers and oblong-obconic
capsules. Taken alone, the Utah specimens are uni¬
form and distinct; nevertheless, I agree with Randel
(1975) that the northern and southern populations
are linked in Colorado and Wyoming by interme¬
diate forms.

Welsh (1986) has applied the name Papaver rad¬
icatum var. pygmaeum (Rydberg) Welsh to the Utah
specimens that I have determined as P. radicatum
subsp. kluanensis. I view P. pygmaeum as morpho¬
logically distinct from the P. radicatum complex,
being more closely related to the European P. al-
pinum group and different from the Utah plants in
several respects. Papaver pygmaeum has leaves
sparsely strigose to commonly glabrous, leaf lobes
short and orbicular, and capsules with whitish (ivo¬
ry) trichomes, whereas P. radicatum subsp. klu¬
anensis has leaves coarsely strigose, lobes obovate,
and capsules with light (straw colored) to dark
brown trichomes.

Papaver pygmaeum is a diploid (2 n = 14; Pack¬
er, 1968). Faberge (1942, 1944) applied the name
P. pygmaeum to a specimen from which he obtained
a chromosome count of n = 21. The origin for this
material was given by Faberge as “Pike’s Peak
Utah.” Clearly, this count was actually derived from
a specimen of P. radicatum subsp. kluanensis from
Pike’s Peak, Colorado, sent to Faberge by [W. P]
Cottam from his home base in Utah. Papaver rad¬
icatum subsp. kluanensis has a chromosome num¬
ber of 2 n = 42 (Love, 1969, based on Colorado
material) or 2 n = 56 (fide Moss & Packer, 1983,
based on material from Alberta).

Volume 5, Number 3
1995

Murray

Papaver Section Meconella

295

Acknowledgments. My thanks to Robert Kiger
and John McNeill for reviewing the manuscript and
for suggesting improvements; and to the curators of
CAN, MAN, NY, and S for loans or their many
courtesies during visits to view types and other
specimens.

Literature Cited

Faberge, A. C. 1942. Genetics of the Scapiflora section
of Papaver 1. The garden Iceland poppy. J. Genetics
44: 169-193.

-. 1944. Genetics of the Scapiflora section of Pa¬
paver 3. Interspecific hybrids and genetic homology. J.
Genetics 46: 125-149.

Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner,
V. Demoulin, D. L. Hawksworth, P. M. J0rgensen, D. H.
Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors.
1994. International Code of Botanical Nomenclature
(The Tokyo Code), Adopted by the XVth International

Botanical Congress, Yokohama, August-September,
1993. Regnum Veg. 131.

Love, D. 1969. Papaver at high altitudes in the Rocky
Mountains. Brittonia 21: 1-10.

Moss, E. H. & J. G. Packer. 1983. Flora of Alberta, ed.
2. Univ. Toronto Press, Toronto.

Packer, J. G. 1968. IOPB chromosome number reports
XVII. Taxon 17: 285-288.

Randel, U. 1975. Die Beziehungen von Papaver pyg -
maeum Rydb. aus den Rocky Mountains zum norda-
merikanischen P. kluanense D. Love sowie zu einigen
nordost-asiatischen Vertretern der Sektion Scapiflora
Reichenb. im Vergleich mit P. alpinum L. (Papavera-
ceae). Feddes Repert. 86(1-2): 19—37.

Randel, U. 1977. Ober Sippen des subarktisch-arktisch-
en Nordamerikas, des Beringia-Gebietes und Nordost-
Asiens der Sektion Lasiolrachyphylla Bernh. (Papaver-
aceae) und deren Beziehungen zueinander und zu
Sippen anderer Arealteile der Sektion. Feddes Repert.
88(7-8): 421^50.

Welsh, S. L. 1986. New taxa and combinations in the
Utah Flora. Great Basin Naturalist 46: 254-260.

Validation of a New Species of Diospyros (Ebenaceae) from
Yunnan, China

Wu Zhengyi (C. Y. Wu) and Zhu Hua (H. Chu)

Kunming Institute of Botany, Academia Sinica, Kunming 650204, Yunnan, China

ABSTRACT. Diospyros xishuangbannaensis, previ¬
ously invalidly described, is validated.

Diospyros xishuangbannaensis was described in
1990 (Zhu & Wu, 1990), but two types, represent¬
ing flowering and fruiting states, were designated,
thus contravening Article 37 of the International
Code of Botanical Nomenclature (Greuter et al.,
1994) and making the name invalid. The species
is herein validated by designating the fruiting ma¬
terial as the holotype.

Diospyros xishuangbannaensis C. Y. Wu & H.

Chu, sp. nov. TYPE: China. Yunnan: Xish-
uangbanna, Mengla, Bubeng, 20 July 1988
(fr), Chu Hua & Wang Hong 2451 (holotype,
KUN).

Proxima Diospyroti martabanicae Clarke, sed ab ea flore
masculo solitario, corolla breviore, staminibus 8, endos-
permio laevigato differt; a D. pilosula (A. de Candolle)
Hiem foliis subtus persaepe dense pubescentibus, nervis
lateralibus utrinsecus 10-15, tertiariis utrinque conspi-
cuis, flore femineo 5-mero, flore masculo solitario, antheris
attenuato-ovatis apice acutis differt.

Trees ca. 20 m tall; branchlets densely brown
pubescent. Leaves oblong or oblong-lanceolate, 8-
18 cm long, 3-4.5 cm wide, stiffly chartaceous to
subcoriaceous, somewhat pubescent adaxially only
on midvein when young, soon completely glabrous,
abaxially with pubescence along midvein like that
of young branchlets, base rounded or subcordate,
apex acuminate; midvein sunken adaxially, elevat¬
ed abaxially; lateral veins 10-15 on each side, flat

adaxially, elevated abaxially, tertiary veins conspic¬
uous on both surfaces; petiole 2—3 mm long, pu¬
bescent as in branchlets. Flowers solitary; stami-
nate flowers 4-merous, on pubescent pedicels 8—10
mm long; calyx 6-7 mm long, divided nearly to
base into linear-lanceolate lobes, appressed seri¬
ceous outside, glabrous inside; corolla salverform,
sericeous outside as in calyx, glabrous inside, tube
ca. 7 mm long, lobes ca. 9 mm long; stamens 8,
unequal, glabrous; anthers attenuate-ovate, acute at
apex; rudimentary ovary small villous; pistillate
flowers 5-merous, on pubescent pedicels 10-16
mm long; calyx and corolla as in staminate flowers;
ovary ovoid, pubescent, 4-loculed. Fruit globose,
2—2.5 cm diam., densely sericeous; fruiting pedicel
1—1.6 cm; fruiting calyx reflexed outward; seeds 4;
endosperm smooth.

Paratype. CHINA. Yunnan: Xishuangbanna, Mengla,
Manzhuan, 7 Mar. 1983, Zhou Shochin & Wang Hong
2452 (KUN).

Acknowledgments. Michael G. Gilbert is
thanked for pointing out the problem and Ihsan A.
Al-Shehbaz for helping with the manuscript.

Literature Cited

Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner,
V. Demoulin, D. L. Hawksworth, P. M. Jorgensen, D. H.
Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors.
1994. International Code of Botanical Nomenclature
(The Tokyo Code), Adopted by the XVth International
Botanical Congress, Yokohama, August—September,
1993. Regnum Veg. 131.

Zhu Hua & Wu Zhengyi. 1990. Contributions to the Flora
of Xishuangbanna (1). Acta Bot. Yunnan. 12: 375—380.

Lectotypification of Kengyilia mutica
(Keng ex Keng & S. L. Chen)

J. L. Yang, Yen & Baum (Poaceae), and Change of Circumscription

Chi Yen and Jun-Liang Yang

Triticeae Research Institute, Sichuan Agricultural University, Dujiangyan City 611830,

Sichuan, China

Bernard R. Baum

Centre for Land and Biological Resources Research, Agriculture and Agri-Food Canada,
Central Experimental Farm, K.W. Neatby Building, Ottawa, Ontario, K1A 0C6, Canada

Abstract. A lectotype of Kengyilia mutica (Keng
ex Keng & S. L. Chen) J. L. Yang, Yen & Baum is
here designated based on an illustration of the ho-
lotype; the latter has been lost. A more complete
description than the original, which was based on
a single and incomplete specimen, is provided.

Professor Y. L. Keng identified a single specimen
obtained from the Province of Qinghai, China, as
Roegneria mutica Keng, nomen nudum. During
1944, Y. L. Keng collected grasses in Qinghai with
his son, P. C. Keng. At that time Guide County was
occupied by a military lord, Ma Bu-Fong and his
herds of horses. Keng did not obtain permission to
collect in this region. A friendly person who worked
on the horse farm brought Keng a specimen. Keng
examined the plant, decided that it differed from
related species, and called it Roegneria mutica (P.
C. Keng, pers. comm. 1992). He subsequently pub¬
lished Roegneria mutica Keng as a nomen nudum
(Keng et al., 1957) and later provided a description
(Keng et al., 1959), but in Chinese only, which re¬
sulted therefore in an illegitimate name. The Latin
description was finally published in 1963 (Keng &
Chen, 1963), giving it invalid status.

The single specimen on which this species was
based may have been destroyed during the “Cul¬
tural Revolution” (Song Gui-Qing, pers. comm.
1992), or may not have been returned to the her¬
barium when it was lent for drawing the figure in
Keng et al. (1959, and P. C. Keng, pers. comm.).
This specimen is “No. 4, Military Horse Farm,
Guide, Qinghai”—the precise locality is unknown;
however, it was collected in an area near the two
subsequent known locations (Fig. 1) and was con¬
served in the herbarium of Nanjing University (N).
The only remains of the type collection are thus the
illustration, figure 337 in Keng et al. (1959), be¬

cause the protologue (Keng & Chen, 1963) referred
to this figure. This interpretation is in accord with
the definition of what constitutes type material in
the Code (Greuter et al., 1994; Article 9.7 Foot¬
notes). Thus, figure 337 in Keng et al. (1959) is
here designated as the lectotype (Fig. 2).

Kengyilia mutica (Keng ex Keng & S. L. Chen)
J. L. Yang, Yen & Baum, Hereditas 116: 27.
1992. Basionym: Roegneria mutica Keng ex
Keng & S. L. Chen, Acta Nanjing Univ. 1: 87.
1963. TYPE: China. Qinghai Province: Guide
County, Military horse farm, 4 [“No. 4, Military
Horse Farm, Guide, Qinghai”] (holotype, N
now lost). Lectotype, here designated: figure
337 in Keng et al. (1959).

Roegneria mutica Keng in Keng et al., Clav. Gen. & Sp.
Gram. Sin. 75, 188. 1957. Nomen nudum. Roegneria
mutica Keng in Keng et al., FI. lllustr. Plant. Prim.
Sin. Gram. 408, figure 337. 1959. Nomen inval.,
based only on Chinese description.

An expedition to Guide County, Qinghai Prov¬
ince, was conducted in 1993 in order to study the
morphological variation at the type locality. There,
we realized that what was described as an awnless
species was in fact an unusual type and that most
of the individuals of Kengyilia mutica have short
lemma awns. Although we collected awnless spec¬
imens in 1986 on the Lajishan mountains, at 3200
m, we found in the 1993 collections only a few
specimens with awnless florets intermixed with
short-awned florets on the same spike. Most indi¬
viduals of this species, based on study of extensive
collections, are short-awned. Totally awnless plants
have yet to be found. We provide here a revised
description of Kengyilia mutica and a drawing (Fig.
3).

Perennial with fibrous roots. Culms 40-70 cm
Novon 5: 297-300. 1995.

298

Novon

Distribution of KengyiNa mutica

Figure 1. Distribution of Kengyilia mutica (Keng ex
Keng & S. L. Chen) J. L. Yang, Yen & Baum.

tall, puberulent below the spikes; 3-noded, genic¬
ulate at the lowest node (Fig. 3A). Leaf sheath
smooth, glabrous or puberulent; ligules scabrous,
ca. 1 mm long; leaf blades flat or involute, glabrous
or puberulent, 4—6 mm wide; tillers with narrow
leaf blades, usually up to 19 cm long. Spikes erect
or slightly curved, 5-9 cm long, 8 mm wide (Fig.
3A); rachis intemodes puberulent, the upper ones
3-4 mm long, the lower ones 8-9 mm long. Spike-
lets yellow-green, 12 mm long, with 4—6 florets
(Fig. 31), the uppermost not well developed; rachilla
intemodes puberulent, ca. 1.5 mm long; glumes ob¬
long ovate, 5-6 mm long, smoothly glabrous, acute,
usually 3-nerved with the midnerve prominent (Fig.
3F, G); lemmas 5-nerved, densely hirsute (Fig. 3H),
with rounded back, tips acute, mucronate (Fig. 3B)
or ending in an awn 1.5-5 mm long (Fig. 3H); 1st
lemma ca. 9.5 mm long; palea equal to or shorter
than lemma, with an obtuse to slightly emarginate
apex, ciliate toward tip on the two keels (Fig. 3C);
anthers yellow or black, ca. 3 mm long. Caryopsis
tip hirsute (Fig. 3E).

Distribution in China. Qinghai Province, Guide
County (Fig. 1), endemic.

Specimens examined. All collected in 1993: CHINA.
Qinghai Province: Guide County, I^ajishan Mountain,
alt. 3200 m, in Stipa steppe, J. L Yang & R. von Bothmer
860020, 860021, 930002, 930007 (all SAUTI, DAO).

Figure 337 Roegneria mutica Keng sp. nov.

1. Adult plant; 2. Spikelet;

3. Florelet; 4. Dorsal view of palea;

5. Caryopsis; 6. Lodicules, anthers with filament

ovary with stigmata.

(No. 4, Guide Military Ranch, Qinghai, Holotype)

Figure 2. Illustration of the lectotype of Kengyilia mu¬
tica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen &
Baum.

1

Volume 5, Number 3
1995

Yen et al.
Kengyilia mutica

299

H

Figure 3. Detailed drawing of Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Aang. Aen & Baum. A. Mature
plant. —B. Dorsal view of floret, showing an awnless lemma. —C. Ventral view of floret. —D. Ventral view of palea.
—E. Caryopsis. —F. Lower glume. —G. Upper glume. —H. Dorsal view of an awned lemma. —I. Spikelet.

300

Novon

Literature Cited

Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner,
V. Demoulin, D. L. Hawksworth, P. M. j0rgensen, D. H.
Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors.
1994. International Code of Botanical Nomenclature
(The Tokyo Code), Adopted by the XVth International
Botanical Congress, Yokohama, August-September,
1993. Regnum Veg. 131.

Keng, Y. L. & S. L. Chen. 1963. A revision of the genus
Roegneria C. Koch of China. Acta Nanjing University

(Biology) 1: 1-92. [In Chinese, with Latin descriptions
of new species, and with an English summary.]

-(editor), P. C. Keng, Y. C. Tang, B. Z. Gao, S. L.

Chen & L. Liu. 1957. Claves Generum et Specierum
Graminearum Primarum Sinicarum, Appendice Nomen-
clatione Systematica. Science Press, Beijing [in Chi¬
nese]. *

-& S. L. Chen, J. R. Feng, Z. Y. Feng, B. Z. Gao,

L. Z. Jia, X. Q. Jiang, P. C. Keng, L. Liu, W. Q. Shi, Y.
C. Tang & S. Q. Zhong. 1959. Flora Illustrata Plantar-
urn Primarum Sinicarum, Gramineae. 403, f. 337. Sci¬
ence Press, Beijing [in Chinese].

Dracontium croatii (Araceae), a New Species from the Western
Slopes of the Andes in Ecuador

Guanghua Zhu

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. and
Department of Biology, University of Missouri-St. Louis, 8001 Natural Bridge Road,
St. Louis, Missouri 63121, U.S.A.

Abstract. Dracontium croatii G. Zhu, a new spe¬
cies from the western slopes of the Andes in Ec¬
uador, is described and illustrated. Morphological¬
ly, this species does not appear to be particularly
close to any member of the genus and is readily
distinguished by its showy greenish spathe, large
and globose berries, rounded and strongly inter¬
rupted irregular warty seeds, and matte upper blade
surface.

Resumen. Dracontium croatii G. Zhu, una espe-
cie nueva de las laderas occidentales de los Andes
de Ecuador se describe e ilustra. A1 parecer, la
nueva especie no esta cercanamente relacionada
con otros taxa en el g^nero. Es fdcilmente recono-
cible por la siguiente combinacion de caracteres:
la espata es vistosa y verdosa, la superficie adaxial
de las hojas es opaca y las semillas y los frutos son
redondeados.

Dracontium L. (Araceae), with 22 species and 2
varieties, is known exclusively from the New World
tropics. One of the most poorly known genera of the
Araceae, it is represented in Ecuador by at least 3
species, 1 of which is new to science. This new
species was detected in the course of a taxonomic
revision of the genus.

Dracontium croatii G. Zhu, sp. nov. TYPE:
U.S.A. Missouri: St. Louis, cultivated plant in
the Missouri Botanical Garden, Zhu 1493 (ho-
lotype, MO); originally collected along unfin¬
ished road between Lita and San Lorenzo, 15.5
km W of Lita, 705 m, 0°55'N, 78°28'W, San
Lorenzo Canton, Esmeraldas, Ecuador, Croat
72368. Figures 1, 2.

Herba terrestris, magna, tuberosa, unifoliata, usque
plus quam 3 m alia; tuber hemisphaericum, usque 15 cm
diam., 7 cm crassum, apice planum interdum tuberculis;
folium bene ante vel post anthesin prodiens; petiolus 1 1
3 m longus, cylindricus, camosus, atro aut purpureo-vir-
idibus maculis, guttatus pallide viridis aut sordide albus;
lamina tripartita usque 20 cm longis, 9 cm latis, apicibus
acuminatis, interdum bifidis; pedunculus 1-1.7 m longus,
1.3—2 cm diam., cylindricus; spatha 23—50 cm longa, 8—

20 cm lata, cucullata basi, obovata aspectu frontali, extus
impolita, atroflava-viridis suffusa purpurea, intus velutina,
flava-viridis apice et secus margines exteriores, cum ner-
vis atropurpureis, viridi-alba basi usque 3 cm prope mar¬
gines et usque 7 cm in medio; spadix 3—5 cm longus, 1.3
cm diam.; tepala 7-8, subcucullata apice; baccae globo-
sae, usque 1—2 cm diam.; semina rubra, suffusa aurantia-
ca, plus minusve rotunda, usque 0.8-1 cm diam., 0.5 cm
crassa.

Terrestrial herb, perennial, to more than 3 m tall.
Tuber hemispherical, to 15 cm diam., 7 cm thick,
20-30 cm underneath the ground, flat at apex, with
or without a few tubercles among numerous roots,
underneath smooth, convex, without tubercles and
roots. Leaf 1, arising from tuber 4—30 cm below the
ground level; petiole 1.4—3 m long, 3.5-6 cm diam.
at base, 1.5-2.5 cm diam. at apex, cylindric, fleshy,
dark or purplish green, mottled pale green or dirty
white forming a reptilian pattern, distal half
smooth, proximal half with many protuberances;
major divisions of blade 3, each to 1 m long, matte,
sometimes fenestrate, upper surface green, lower
surface slightly paler; ultimate segments to 20 cm
long, 9 cm wide, oblanceolate, apex gradually acu¬
minate or caudate, or bilobed, lobes gradually acu¬
minate; smaller leaflets triangular, or ± oblanceo¬
late, 3.5-15 cm long, 2.5-6 cm wide; midrib and
major veins convex, sunken adaxially, round-raised
abaxially, secondary veins impressed on upper sur¬
face and raised on lower surface; rachises pale
green, smooth, faintly mottled as petiole, rachis
(from petiole apex to the first major subdivision) of
the middle division up to twice as long as that of
lateral divisions. Inflorescence arising from apex of
tuber before or after the leaf development; cata-
phylls to 20 cm long, unribbed, drying light brown
toward base, darker brown toward apex; peduncle
1-1.7 m long, 1.3-2 cm diam. at mid point, cylin¬
dric, fleshy, dark blackish purple, sparsely and
weakly streaked, violet or dark yellow-green, be¬
coming somewhat more brownish with larger
swatches of yellow-green above middle, these be¬
coming obliquely arranged, larger and paler toward
apex, beset with irregular protuberances through-

302

Novon

Figure 1. Dracontium croatii G. Zhu (Croat 72368). —A. Leaf. —B. Peduncle (left) and petiole (right).

out; spathe 23-50 cm long, 8—20 cm wide when
open, conspicuously hooded, convolute at base, ob-
ovate in front face view, outside matte, green tinged
purple, apex and the lower Ps of spathe densely
purplish speckled, midrib broadly convex, pale yel¬
lowish green in lower half of spathe, purplish on
upper half of spathe, with 11-12 dark purplish ma¬
jor veins on either side, these veins interconnected
by oblique cross veins, inner surface velvety, tri¬
color: apex and along outer margins yellow-green,
with dark purple veins, area 4-6 cm from the outer
margins dark purple and becoming more velvety,
area 3 cm high near the margins and 7 cm high
medially from the base solid greenish white, the
border between the white and purple portions
abrupt but with smoothly feathered margin, apex
probably rounded or weakly cuspidate (apparently
somewhat damaged and split upon opening), outer
margins thin and weakly curled back, moderately
undulate toward base; spadix 3-5 cm long, 1.3 cm
diam. excluding protruding styles, purplish, axis of
spadix pinkish within, aerenchymatous. Flowers ir¬
regularly rounded, 2.5-3 mm diam.; tepals 7-8 per
flower, 1.8—2.2 mm long, 1-1.5 mm wide, somewhat
cucullate at apex, inner margin broadly rounded,
purplish with paler, weakly raised projections, pal¬
er and somewhat scalloped along the edges; styles
1-1.2 mm long, 0.8-1 mm diam., spindle-shaped,
purplish, covered with weakly raised projections as
tepals; stigmas somewhat cushion-shaped, 2- or
rarely 3-lobed, 0.4 mm high, 0.8 mm diam., cov¬
ered with brushlike translucent bristles, becoming
purplish after anthesis; stamens not emerging above
tepals; filaments ca. 1.5 mm long; anthers brown¬
ish, clearly visible between tepals and style prior

to anthesis. Berries globose, 1—2 cm diam., olive-
green with purplish spots, becoming deep red;
seeds 2 per berry, red tinged orange, with thick,
strongly interrupted, irregular warts, round, 0.8-1
cm diam., 0.5 cm thick.

Dracontium croatii is apparently restricted to the
western slopes of the Ecuadorian Andes between
450 and 1330 m. In addition to the type locality,
collections have also been made from Carchf Prov¬
ince. The species may also be expected in Los Rios
Province in Ecuador, as well as adjacent areas of
Colombia.

Morphologically, Dracontium croatii does not ap¬
pear to be particularly close to any member of the
genus. It is readily distinguished by its showy
spathe with a unique coloration: outer surface
greenish with raised, dark purple veins, inner sur¬
face velvety, dark purple with 4—6 cm wide, yellow-
green outer margins and a solid greenish white area
3-7 cm high at the base. The globose berries of the
species, at 1-2 cm diam., are the largest in Dra¬
contium. Other species, such as D. spruceanum,
may also have globose berries, but they never ex¬
ceed 1 cm diam. Seeds of D. croatii are definitely
the largest known in the genus; the distinctive seed
morphology is illustrated on one of the cover pages
of Aroideana (Madison, 1980). Dracontium croatii
is also often marked by a matte upper blade sur¬
face, which is glossy or semiglossy in other species.

Inflorescences of Dracontium croatii have been
collected in November, fruits in July. Under culti¬
vation, D. croatii flowers in September and from
November to January. It produces inflorescences
before or immediately after new leaf development.

Volume 5, Number 3
1995

Zhu

Dracontium croatii

303

Figure 2. Dracontium croatii G. Zhu. —A. Infructescence (Croat 72368). —B, C. Inflorescence (Zhu 1493, type). —
D. Inflorescences opened to show the spadix and the creamy-white area at the base of the inner spathe surface (Zhu
1493, type).

304

No von

Dracontium croatii is named in honor of Thomas
B. Croat, from whose collection of living Araceae
in the Missouri Botanical Garden I recognized this
new species and prepared the holotype. Two other
live collections are known: one from the Marie Sel¬
by Botanical Gardens (75-491), the other from the
Missouri Botanical Garden (received from the Selby
Gardens, 78-2173); both were collected by Calaway
Dodson in Pichincha Province, Ecuador, in 1975
and 1978. Other plants of this species have been
distributed by the Selby Gardens to various other
places under the name D. ornatum. One of these
plants is cultivated by Dewey Fisk in Florida.

Paratypes. ECUADOR. Carchf: Forestry Reserve, vi¬
cinity of encampment in Aw4 Ethnic, 0°58'N, 78°16’W,
1330 m. Hoover 3279 (QCA); trail from San Marcos to
Tobar Donoso, 660 m, 1°07'N, 78°20'W, Barfod 41597
(AAU, MO, QCA). Esmeraldas: along unfinished road
between Lita and San Lorenzo, 15.5 km W of Lita, 705
m, 0°55'N, 78°28'W, Croat 72368 (MO). Pichincha:
bridge over Rfo Chiguilpe near junction with Rfo Baba, 7
km from junction of entrance road at 7 km from Santo
Domingo on highway to Quevedo, 450 m, Dodson 5951
(SEL); La Centinela, at crest of Montanas de Ila on road
12 km from Patricia Pilar to 24 de Mayo, 600 m, Dodson
7288 (MO, SEL), Dodson & Fallen 7762 (MO, QCNE,
SEL); La Centinela, 0.2 km past Escuela Mixta La Cen¬
tinela, along trail to left of road, exactly 13 km E from
main Santo Domingo-Quevedo Highway in Patricia Pilar,
1000 m, 0°32'S, 79°11'W, Croat 73040 (MO). Cultivated
plants: Marie Selby Botanical Gardens (75-461, vouch-

ered by Dodson 5951), Christenson 1576 (SEL), Croat
71798 (MO), Madison 4140 (SEL); Marie Selby Botanical
Gardens (78-2173, vouchered by Dodson 7288), Christen¬
son 1154 (SEL), Plowman 10926 (F); Missouri Botanical
Garden (vouchered by Croat 72368), Zhu 1518 (MO);
Missouri Botanical Garden (received from the Marie Selby
Botanical Gardens, 78-2173, vouchered by Dodson 7288),
Zhu 1453 (MO).

Acknowledgments. This paper is part of my
doctoral dissertation to be submitted to the Uni¬
versity of Missouri-St. Louis Department of Biology.
I thank my major advisor, Thomas B. Croat, for
discussions and critical suggestions on this subject.
I also thank Mike Grayum, Sue Thompson, German
Camevali, Eleanor Sauer, P. Mick Richardson, Da¬
vid Boufford, Bruce Bartholomew, Petra Malesev-
ich, Donna Atwood, Kathy Upton, and Hong Song
for comments on the manuscript and other assis¬
tance. I extend my appreciation to Dewey Fisk, Ju¬
lius Boos, and curators of the Marie Selby Botanical
Gardens and Herbarium for providing specimens
and photos. The research was conducted at the Mis¬
souri Botanical Garden and was partially supported
by NSF doctoral dissertation research grant DEB-
9310171 to T. B. Croat and G. Zhu, and by a grant
from the Andrew W. Mellon Foundation to G. Zhu.

Literature Cited

Madison, M. 1980. Cover photo. Aroideana 3(1): 1-2.

Volume 5, Number 3, pp. 209-304 of NOVON was published on 05 September 1995.

Volume 5
Number 4
1995

NOVON

MISSOURI BOTANIC"!

DEC 2 7 1995

GARDEN LlbRrtRt'

A New Blakea (Melastomataceae: Blakeeae) from
Tapantf National Park, Costa Rica

Frank Almeda

Department of Botany, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118-4599, U.S.A.

Gina Umaha Dodero

Herbario Nacional de Costa Rica, Museo Nacional, Apartado 749-1000,
San Jos6, Costa Rica

Abstract. Blakea tapantiana, a new Costa Rican
endemic, is described, illustrated, and compared
with B. grandiflora, an allopatric Costa Rican en¬
demic that occurs at higher elevations and flowers
sporadically throughout the year. The new species
is distinguished by its short-pedicellate flowers,
caudate-acuminate outer floral bracts, white petals,
and unappendaged anther thecae.

Blakea, a neotropical genus with over 100 de¬
scribed species, continues to yield new Mesoamer-
ican taxa of local distribution (Umana & Almeda,
1991). Unlike many species of Blakea, which tend
to flower sporadically throughout the year, the spe¬
cies proposed here appears to flower for a short
period during the rainy season followed by a rather
lengthy period of fruit maturation. This probably
accounts, in part, for the paucity of flowering and
mature fruiting specimens in herbaria and empha¬
sizes the need for repeated visits to mid-elevation
montane forests in Costa Rica and elsewhere in
tropical America.

Blakea tapantiana Umana & Almeda, sp. nov.
TYPE: Costa Rica. Cartago: Parque Nacional
Tapantf, Sendero Oropdndola, 9°48'18"N,
83°57'12"W, 1200 m, 3 Sep. 1992 (fl), Umana
et al. 497 (holotype, CR; isotypes, CAS, COL,
F, K, MEXU, MO, US, USJ, WIS). Figure 1.

Frutex epiphyticus vel terrestris. Ramuli primum sub-
quadrangulati demum teretes sicut foliorum subtus venae
primariae et secundariae pedicelli bracteaeque primum
densiuseule furfuracei, demum glabrescentes. Lamina el-
liptiea vel oblongo-elliptiea, apice caudato-acuminato.
Flores 6-meri in quoque nodo superiore 1—3, pedicellis
0.6-1 cm longis: bracteae exteriores ca. 3-6 mm coalitae
apice hebeti-acuminato. Antherae 4.5-5 X 1—1.5 mm in¬
ter se non cohaerentes; connectivum dorsaliter paulo ele-
vatum ecalcaratum.

Terrestrial or epiphytic shrubs 2-6 m tall. Upper
branchlets rounded-quadrate to somewhat sulcate,
becoming rounded and glabrous with age. Vegeta¬
tive buds, young branchlets, young petioles, pri¬
mary and secondary abaxial leaf veins, and imma¬
ture inner and outer floral bracts densely to
moderately covered with dendritic and scurfy hairs.
Mature leaves of a pair equal to somewhat unequal
in size, essentially glabrous adaxially or sparingly
beset with a mixture of paleaceous, dendritic, and/
or conic hairs along the elevated veins; petioles
1.5-3.5 cm long; blades coriaceous, 8.3-17 cm
long and 4.1-7.6 cm wide, elliptic to oblong-ellip¬
tic, rarely elliptic-obovate, apex acuminate to cau¬
date-acuminate, base acute varying to obtuse, mar¬
gin denticulate to subentire, 5-nerved or
5-plinerved with the innermost pair of primary
veins diverging from the median vein 3—6 mm
above the blade base, the striolate transverse sec¬
ondary veins spaced 1.5—3 mm apart at the widest

Novon 5: 305-308. 1995.

306

Novon

igure 1. Blakea tapantiana Umana & Almeda. —A. Habit. —B. Representative leaf (abaxial surface). —C. Enlarge¬
ment of a primary leaf vein (abaxial surface). —D. Immature fruit enveloped by outer and inner floral bracts. —E.
\oung fruit (top view) showing ovary summit, torus, and calyx lobes. —E Fused inner floral bracts. —G. Petal. —H.
Stamens, lateral view (left) and ventral view (right). (A—C from Almeda et al. 6860 ; D—H from Umana et al. 497.)

Volume 5, Number 4
1995

Almeda & Umana
Blakea from Costa Rica

307

portion of the blade. Flowers erect, 1-3 in each leaf
axil of uppermost branches; pedicels 0.6-1 cin
long, thick but somewhat compressed and 2-edged,
glabrous and lenticellate. Floral bracts sessile, en¬
tire; outer bracts 1.8-2.5 X 0.8-1.1 cm, fused ba-
sally for 3-6 mm, ovate, apex acuminate to
caudate-acuminate, margin entire, 3-veined with
only the median vein usually prominulous, glabrous
throughout; inner bracts 1—1.2 cm long, connate
basally for nearly half their length to form a 2-lobed
cuplike collar that envelops the hypanthium but is
somewhat concealed by the outer bracts, the free
portions of the inner bracts rounded-triangular and
glabrous, 0.4-0.7 X 0.7-0.8 cm. Hypanthium (in
young fruit) campanulate, 0.8-1 cm long to the to¬
rus and 0.6—0.8 cm diam., glabrous. Calyx tube
2.5-3 mm long; calyx lobes 4.5-6 mm long and 5-
6 mm wide basally, rounded-triangular, margin ±
entire and conspicuously callose-thickened, gla¬
brous on both surfaces. Petals 6, glabrous, 17—21
X 7-10 mm, white, narrowly obovate, apically
rounded (sometimes obliquely so) and often abrupt¬
ly and bluntly acute, essentially entire and cilio-
late. Stamens 12, isomorphic; filaments 5-7 mm
long, complanate, glabrous and declined to one side
of the flower opposing the style; anthers 4.5-5 mm
long, 1—1.5 mm wide, yellow, oblong, laterally com¬
pressed with two well-separated pores at the trun¬
cate apex; connective slightly dilated dorso-basally
ca. 0.5 mm above the filament insertion into a low
blunt thickening. Ovary wholly inferior, 6-celled,
glabrous at the slightly elevated bluntly conic apex
but lacking a collar. Style somewhat declinate, gla¬
brous, 10-11 mm long; the expanded stigma sub¬
truncate to narrowly conic. Mature berry and seeds
not seen.

Distribution. This little-collected cloud forest
species is known only from Cartago Province, Costa
Rica, in a region extending from the recently des¬
ignated (1993) Tapantf National Park (formerly Re¬
fugio Nacional de Vida Silvestre) to Valle Escon¬
dido on the Atlantic slope of the Cordillera de
Talamanea at 1150-1450 m.

Blakea tapantiana is notable for its caudate-acu¬
minate leaf apices, short-pedicellate flowers, acu¬
minate to caudate-acuminate outer floral bracts,
white petals, and unappendaged anther thecae.

Among the few white-flowered Costa Rican spe¬
cies of Blakea that exhibit both an epiphytic and
free-living shrubby habit, B. tapantiana is likely to
be confused only with B. grandiflora Hemsley. The
latter species is also endemic to Costa Rica, but it
is most commonly encountered in the wet wooded

slopes and valleys of the Cordillera Central and
Cordillera TilarAn.

Many of the diagnostic features that can be used
to separate Blakea tapantiana from B. grandiflora
are quantitative. The leaf blades of B. grandiflora
are consistently larger (13-25.5 X 7.7-14.7 cm),
as are the pedicels (2.5-4.5 cm), outer floral bracts
(1.3-2 cm wide), and fully expanded petals (25-35
X 25-32 cm). In B. grandiflora the outer floral
bracts are fused basally for 8-15 mm, but they are
fused for only 3-6 mm in B. tapantiana. Flower
color also appears to provide a diagnostic field
character for distinguishing the two species. The
apex of floral buds in B. tapantiana is pale pink,
but the petals are completely white when fully ex¬
panded. In B. grandiflora, however, the buds are
deep pink, and the expanded petals are flushed
pink apically on one or both surfaces.

Other more subtle but consistent differences can
be used to distinguish these two species. In overall
aspect the vegetative and floral organs of Blakea
grandiflora are coarser and thicker, but these are
difficult to quantify. The anthers of B. grandiflora
have a distinctive blunt appendage borne dorsally
on the connective about 1 mm above the base of
the thecae. The anther connective in B. tapantiana
is merely thickened and not elaborated into a knob¬
by appendage. Blakea grandiflora, like II. tapan¬
tiana, has a sparse to dense indument on upper¬
most branchlets, vegetative and young floral buds,
and petioles that consists of coarse dendritic hairs
that vary toward or are intermixed with paleaceous
hairs that have somewhat lacerate margins. In B.
tapantiana the dendritic hairs are not as coarse or
as well developed and vary toward a furfuraceous
indument on abaxial leaf surfaces.

Taken together these consistent differences, cou¬
pled with the elevational and geographical sepa¬
ration between B. tapantiana and B. grandiflora,
provide a compelling rationale for recognizing two
species. This taxonomic interpretation is consistent
with a pattern of sharply discontinuous quantitative
differences seen in other species pairs such as B.
austin-smithii Standley and B. chlorantha Almeda
and in B. calycosa Gleason and B. tuberculata Don¬
nell Smith.

Paratypes. COSTA RICA. Cartago: Refugio Nacional
de Fauna Silvestre Tapantf, Sendero de la Orop^ndola bor¬
dering Rfo Macho, 22 Mar. 1986 (fr), Almeda et al. 5699
(CAS, CR, MO, NY); Refugio Nacional de Vida Silvestre,
Sendero de la (iropcndola about 1.2—2.7 km from the re¬
serve entrance, 1 Mar. 1991 (fr), Almeda et al. 6860 (CAS,
CR, F, US); Refugio Nacional de Vida Silvestre, Sendero
de la I iropcndola. 13 Feb. 1992 (fr), Almeda et al. 6971
(CAS, CR, MO, US); bank of Rfo Grande de Orosf, 4 km
S of Tapantf, 15 Sep. 1967 (fl). Lent 1280 (F); Valle Es-

308

Novon

condido, 15 Apr. 1966 (fr), Schnell 689 (F); Parque Na-
cional Tapantf. Sendero (Iropdndola, 9°48'18''N,
83°57'12"W, 1200 m, 20 Aug. 1992 (fl), Umana et al. 496
(CR), 29 Sep. 1992 (fl), Umana et al. 498a (CR), 24 Sep.
1993 (fl), Umana et al. 498b (CR).

Acknowledgments. We thank Colleen Sudekum
for preparing the line drawings, Barrett Anderson,
Bruce Bartholomew, Thomas F. Daniel, Leticia Pa¬
checo, and Nelson Zamora for assistance in the

field, the staff of the Herbario Nacional de Costa
Rica for logistical support, and the Servicios de
Parques Nacionales de Costa Rica for permission
to collect in protected areas of Costa Rica.

Literature Cited

Umana D., G. & F. Almeda. 1991. Blakea costaricensis
(Melastomataceae): A remarkable new epiphyte from
southeastern Costa Rica. Selbyana 12: 1-4.

Placement of Arabidopsis parvula in Thellungiella (Brassicaceae)

Ihsan A. Al-Shehbaz and Steve L. O’Kane, Jr.

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. The genus Thellungiella is recog¬
nized, and the characters separating its two species
from Arabidopsis are given. Arabidopsis parvula
(Schrenk) O. E. Schulz is recognized in Thellun¬
giella.

The limits of Arabidopsis have been the subject
of considerable controversy, and as many as 50 spe¬
cies were previously placed in the genus (Al-Sheh-
baz, 1988; Price et al., 1994). Detailed morpholog¬
ical and molecular studies by the authors reveal
that Arabidopsis includes less than ten species,
most of which were previously assigned to other
genera, and that the remaining species are more
appropriately placed in five or six genera. The re¬
sults of these on-going studies will be published
elsewhere.

The present paper deals with two species here
recognized as members of Thellungiella O. E.
Schulz. Thellungiella parvula has generally been
accepted in Arabidopsis in all recent floras (e.g..
Ball, 1964; Busch, 1939; Hedge, 1965, 1968; Pav¬
lov, 1961) that cover the areas of its range. The
generic placement of T. salsuginea (Pallas) 0. E.
Schulz has been more controversial; although sev¬
eral authors followed Schulz (1924) in treating it in
Thellungiella, Al-Shehbaz (1988) and Rollins
(1993) retained the species in Arabidopsis. Schulz
(1924, 1936) placed Arabidops is and Thellungiella
in different subtribes separated solely on the basis
of presence versus absence of seed-coat mucilage.
As shown by Vaughan and Whitehouse (1971), the
presence or absence of mucilage in wetted seeds
can be observed in various species of the same ge¬
nus and, therefore, is not a reliable taxonomic char¬
acter. Although the seeds of T. parvula produce mu¬
cilage upon wetting and those of T. salsuginea do
not, it is abundantly clear that this and other dif¬
ferences given in the key below do not justify the
placement of the two species in different genera.

In addition to Thellungiella salsuginea, Schulz
(1924), Busch (1939), and Pavlov (1961) recog¬
nized another species, T. halophila (C. A. Meyer)
D. E. Schulz, which they separated from T. salsug¬
inea primarily by having coarsely dentate to divid¬
ed instead of entire to repand basal leaves. It is
highly unlikely that these differences alone justify

the recognition of T. halophila as distinct from T.
salsuginea. Because the types of these two species
were not available for this study, we refrain from
formally reducing T. halophila to the synonymy of
T. salsuginea.

As here delimited, Thellungiella consists of two
(or perhaps three) species that are centered pri¬
marily in Kazakhstan and neighboring Russia.
Thellungiella parvula is distributed in Kazakhstan,
southern Russia, Turkmenistan, and Turkey, where¬
as T. salsuginea is widespread in Kazakhstan,
southern Russia, Mongolia, northern China, Cana¬
da, and the United States. Both species are restrict¬
ed to strongly saline soils, and such edaphic ad¬
aptation to salinity, which is rare elsewhere in the
Brassicaceae, is not exhibited by any species of
Arabidopsis. In addition, both species are glabrous
throughout, are glaucous on stems and leaves, and
do not produce well-defined basal rosettes. All spe¬
cies of Arabidopsis produce basal rosettes that often
persist well after fruit maturity, and none has glau¬
cous stems and leaves. The characteristic indu¬
mentum of mixed simple and stalked furcate tri-
chomes should readily distinguish species of
Arabidopsis from members of Thellungiella. The two
species of Thellungiella are easily separated as fol¬
lows:

la. Cauline leaves oblong to ovate or cordate, not
fleshy, auriculate to rarely amplexicaul at base;
infructescence axis straight; petals much longer
than sepals; seeds not mucilaginous when wetted

. T. salsuginea

lb. Cauline leaves linear to linear-oblong, fleshy, not

auriculate at base; infructescence axis flexuous;
petals subequaling sepals or absent; seeds mu¬
cilaginous when wetted . T. parvula

Thellungiella parvula (Schrenk) Al-Shehbaz &
O’Kane, comb. nov. Basionym: Diplotaxis?
parvula Schrenk in Fischer & C. A. Meyer,
Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint
lY'tcrsbourg ser. 2, 2: 199. 1844. TYPE: [Ka¬
zakhstan]. “In desertis salsuginosus ad Ters-
akkan,” A. Schrenk s.n. (holotype, LE; iso¬
type?, G).

Acknowledgments. This research was supported
by the National Science Foundation grant DEB-

Novon 5: 309-310. 1995.

310

Novon

9208433 and the National Geographic Society
grant 5068-93, both of which were awarded to Al-
Shehbaz. We thank Barbara Schaal (Washington
University) for her support in conducting the mo¬
lecular studies leading to these results.

Literature Cited

Al-Shehbaz, I. A. 1988. The genera of Sisymbrieae (Cru-
ciferae; Brassicaceae) in the southeastern United States.
J. Arnold Arbor. 69: 213-237.

Ball, P. W. 1964. Arabidopsis. In: T. G. Tutin et al. (edi¬
tors), FI. Europaea 1: 267-268.

Busch, N. A. 1939. Arabidopsis. In: V. L. Komarov (edi¬
tor), FI. URSS 8: 76-80.

Hedge, 1. C. 1965. Arabidopsis. In: P. H. Davis (editor),
FI. Turkey 1: 489-490.

-. 1968. Sisymbrieae. In: K. H. Rechinger (editor),

FI. Iranica 57: 309-342.

Pavlov, N. V. 1961. Flora Kazakhstana 4: 1—545.

Price, R. A., J. D. Palmer & I. A. Al-Shehbaz. 1994.
Systematic relationships of Arabidopsis: A molecular
and morphological perspective. In: E. M. Meyerowitz &
C. R. Sommerville (editors), Arabidopsis. Cold Springs
Harbor Laboratory Press, New York.

Rollins, R. C. 1993. The Cruciferae of Continental North
America. Stanford Univ. Press, California.

Schulz, O. E. 1924. Cruciferae-Sisymbrieae. In: A. En-
gler (editor), Pflanzenreich IV. 105(Heft 86): 1-388.

-. 1936. Cruciferae. In: A. Engler & K. Prantl (ed¬
itors), Natiirlichen Pflanzenfamilien ed. 2. 17B: 227—
658.

Vaughan, J. G. & J. M. Whitehouse. 1971. Seed structure
and the taxonomy of the Cruciferae. Bot. J. Linn. Soc.
64: 383^09.

New Species of Cestrum (Solanaceae) and Synonymy under
Two Widespread Species

Carmen Benitez de Rojas

Facultad Agronomfa, Universidad Central de Venezuela, Maracay,
Apartado Correos 4597, Aragua, Venezuela

William G. D’Arcy

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Abstract. Three new species of Cestrum, C. til-
lettii, C. jaramillanum, and C. ruizteranianum, are
newly described from Venezuela. Cestrum tomen-
tosum L. f. and C. strigilatum Ruiz & Pav6n have
been identified as the correct names for two spe¬
cies for which many names have been proposed,
and synonymies for these two species are pre¬
sented.

Three new species are described from different
parts of Venezuela: Sierra de Perija in northeastern
Zulia State, the western, Andean region of Merida
State, and the more or less central part of the coast¬
al range in the Federal District. The last species is
also found in Ecuador.

Two other widespread species have been known
under many different names, and because con¬
cepts of their ranges have been fragmented with
so many names in use, we present the synonymy
here, rather than waiting until a revision of the
genus can be published for some major region.
One of these, C. tomentosum L. f., ranges from
Mexico to Peru and Venezuela; the other, C. stri¬
gilatum Ruiz & Pavdn, ranges from Costa Rica to
Venezuela, Peru, Brazil, and Argentina. Cestrum
embraces many species with local or regional dis¬
tributions, but these two add to the short list of
species that are widespread in both Central and
South America: C. alternifolium (Jacquin) O. E.
Schulz, C. latifolium Lamarck, C. megalophyllum
Dunal, C. microcalyx Francey, C. rejlexum Sendt-
ner, C. racemosum Ruiz & Pav6n, and C. scandens
Vahl.

Cestrum jaramillanum Benttez & D’Arcy, sp.
nov. TYPE: Ecuador. Pichincha: Centenela,
Montafias de Ila, 12 km E of Patricia Pilar,
virgin rain forest, 550-650 m, 79°19'W,
0°34'S, 10 July 1979 (fl). Lojtnant & Molau
15835 (holotype, AAU; isotype, CB). Figure 1.

Arbor frutexve 2—3 m, foliis ovatis, basi rotundatis,
truncatis, subcordatisve, membranaceis, glabrescentis, fo¬
liis minoris ovatis. Inflorescentia axillaria, subumbellata,
axibus 6-7 mm longis, tomentulosis glabratisve. Bracteae
sessiles foliaceae, lanceolatae, ad maturitatam fructus per¬
sisted, lateraliter calycem involvens.

Shrub or small tree 2-3 m, branched, the
branches slender, at first compressed, sometimes
drying grooved, later terete, tomentulose with
mostly reduced, crinkled ascending hairs, the in-
temodes 3-4.5 cm long. Leaves ovate, 10—14 X
4-7.5 cm, basally rounded, truncate, or subcor-
date, apically short-acuminate, membranous or
chartaceous, glabrous above on emerging, the bas¬
al half beneath tomentulose on emerging, glabres-
cent, the veins inconspicuous above, the costa and
major veins slightly reddish, somewhat elevated
and finely puberulent beneath, the lateral veins 7-
8 on each side, ascending, distally arcuate or
looping and forming a submarginal vein 0.5—1 cm
from the margin, the margin plane; petiole 5—10
mm long, finely pubescent; minor leaves ovate, 7
X 1.5 mm, sessile, glabrate. Inflorescences axil¬
lary, the peduncle ca. 1 mm long, longer in fruit,
subumbellate with 3 axes 6-7 mm long, tomen¬
tulose to glabrate, in fruit 15 mm long. Flowers
sessile; bracts foliaceous, narrowly lanceolate, 10
X 3.5 mm, persistent in fruit and partially envel¬
oping the calyx laterally, sessile, acuminate, gla¬
brate or pubescent on both sides; calyx tubular, 4
X 2.5 mm, drying stramineous, basally rugulose,
glabrate, 5-lobed, the lobes deltoid to broadly acu¬
minate, faintly 5-costate; corolla pale green, 29-
30 mm, the tube 26 mm long, gradually expanded
upward, the mouth not contracted, 2.5—3 mm
wide, 5-lobed, the lobes 6-6.5 X 1 mm, oblong,
the apex acute, the pleated margin short-pilose;
stamens 22-23 mm long, the filaments straight,
smooth, glabrous, basally adnate for 21 mm, free
1 mm; anthers orbicular, 1 mm across; ovary 0.7
mm diam., slightly rugose, glabrous; ovules 5-6;

Novon 5: 311-317. 1995.

312

Novon

Figure 1. Oestrum jaramillanum Benitez & I)'A re v. -—A. Flowering branch. —B. Flowers. —C. Flower opened lon¬
gitudinally showing corolla, anthers, and pistil. —D. Fruit. (A, B, C after iMtnant & Molau 15835; D after Steyermark
92147.)

style 23 mm long; stigma slightly bilobed, includ¬
ed. Fruit ovoid, 8-10 X 6-7 mm, the pericarp
thin, about equaling and loosely enclosed in the
bract; seeds 3—4, dark brown, 5-6.5 mm long.

Cestrum jaramillanum is found in moist forests
on the western slopes of the Ecuadorean Andes and
on slopes of the Venezuelan coastal cordillera at
550-1300 m elevation.

Volume 5, Number 4
1995

Benitez & D’Arcy

New Species and Synonymy in Oestrum

313

This species is named in honor of Jaime Lucio
Jaramillo Asanza, professor and active plant col¬
lector at the Pontificia Universidad Catolica del Ec¬
uador in Quito, who has greatly assisted many bot¬
anists visiting his country.

This new species is suggestive of Oestrum reflex-
um Sendtner and C. amictum Schlechtendal, both
described from Brazil and having conspicuous
bracts, but in C. reflexum the corolla tube is abrupt¬
ly expanded at the throat, and the plant is scandent;
in C. amictum the leaves are basally acute or acu¬
minate and decurrent on the petiole, and the minor
leaves are reniform. This new species is apparently
a plant of lower or middle elevations, the two col¬
lections being made at 550-1300 m.

Paratype. VENEZUELA. Distrito Federal: Cerro
Naiguata, laderas pendientes del Norte, arriba del pueblo
de Naiguatd, bosque humedo denso, Lomas de Las Deli-
cias entre Quebrada de Basenilla y Quebrada Guayoyo,
9—12 km suroeste de Hacienda Cocuizal, 1000—1300 m,
15-19 Nov. (fl, fr), Steyermarh 92147 (NY, US, VEN).

Cestrum ruizteranianum Benitez & D’Arcy, sp.
nov. TYPE: Venezuela. Merida: Distrito Ran¬
gel, trail from La Negrita downstream towards
Puente de La Escalera, montane cloud forest,
2550-2950 m, 1-2 Nov. 1978 (fl), Luteyn et
al. 6171 (holotype, NY). Figure 2.

Flores copiosi, calyce atrobrunneo, extus glanduloso,
pilos sparsos ferens, corolla 20-26 mm longa, dilute viridi
ex purpurea. Filamenta 7—9 mm ad corollam coalita, ad
basim parte adnata pubescenti, in loco conjunetionis tum-
idis, geniculatis, parte libera 7.5—9.5 mm. Fructus haud
suppetunt.

Shrub 1-4 m tall, erect, branched, young stems
angular, terete when mature, scurfy pubescent with
dark, crinkled, perhaps branched hairs. Leaves
narrowly elliptical to elliptical, 4—6 X 1.5-3 cm,
basally obtuse, apically acute or obtuse, firmly
membranous, glabrous on both sides except for
some scurfy hairs on minor veins, the major veins
impressed above, elevated beneath, 8-10 on each
side, ascending, looping and uniting near the mar¬
gins, the margins slightly revolute; petioles 6-8 mm
long, slender, inrolled, tomentulose above; minor
leaves sometimes present, ovate, 10—13 X 5—7 mm,
with petioles 0.5-1 mm long. Inflorescences axil¬
lary racemes, sometimes appearing as terminal
panicles, 2.5—10 cm long; peduncles 0.7—5 cm
long; bracts 1.5-2.5 mm, linear. Flowers with ped¬
icels obsolete; calyx drying dark brown, tubular, 6—
8 X 3—3.5 mm, faintly striate, thick, glandular and
with sparse hairs outside, pubescent within and
with glandular hairs halfway down, 5-lobed, the
lobes 2X2 mm; corolla pale green, purplish out¬

side, pale yellowish green inside, 20—26 mm long,
exserted ca. 19 mm from the calyx, the tube 17—
20 mm long, 3-3.5 mm at its widest, contracted
around the ovary and then gradually expanded up¬
ward, the throat not constricted, 5 mm wide at the
mouth, 5-lobed, the lobes 3-5 X 1.5-2 mm, nar¬
rowly triangular-acuminate, sometimes sparingly
pubescent, the pleated margin indexed and tomen-
tose; stamens 15.5-19 mm long, adnate 7-9 mm,
the adnate portion pubescent for the basal 3—1 mm,
free part 7.5—9.5 mm, the insertion geniculate-tu¬
mid, 1.5-2.5 mm long; anthers orbicular, 1 mm
across; ovary lobed, 0.7 diam., glabrous, papillose
4-5 mm below the stigma, exceeding the stamens
by 1.5 mm; style 15-19 mm, stigma subcapitate,
slightly bilobed, included; ovules 16-18. Fruit not
seen.

Cestrum ruizteranianum is found in cloud forest
ravines in the Venezuelan Andes at elevations of
2500-2900 m.

This species is named in honor of Luis Rufz-
Teran, a botanist who for many years collected in¬
teresting plants in Andean northwestern Venezuela.

As a herbarium specimen, this species resembles
showy yellow-flowered species from upland Central
America, e.g., C. laxum Bentham, but the flowers
are reported as pale green and purple. It differs also
in its glandular calyx.

Although three collections are cited, they were
made the same day by the same collecting party,
perhaps at the same place.

Paratypes. VENEZUELA. Merida: Distrito Rangel,
Gavidia, orillas del camino entre el Hato de La Escalera
y el puente sobre la quebrada La Escazes, 2550—2950 m,
1 Nov. 1978 (fl), RuCz-Terdn et al. 16171 (MERF, MY),
16154 (MY).

Cestrum strigilatum is one of the most wide¬
spread species of Cestrum. It ranges from coastal
Brazil and northern Argentina to Costa Rica, oc¬
curring from sea level to about 1600 m. In his
monograph of the genus, Francey (1935) used the
later name C. calycinum for this species. His ra¬
tionale for use of names of Willdenow in Roemer
and Schultes (1819) instead of those of Kunth
(1818) was stated on p. 142 of the monograph,
which cites Kuntze’s Revisio Generum and Sendtner
in Flora Brasiliensis. Some of the following synon¬
ymy was expressed by D’Arcy (1972) and Cabrera
(1983), but not by Smith and Downs (1966). We
have chosen not to lectotypify the species at this
time.

Francey included Cestrum lundianum Dunal (in
DC., Prodr. 13(1): 658) in synonymy under this spe¬
cies. Photographs of the type (Brazil, Sao Paulo,

314

Novon

figure 2. Cestrum ruizteranianum Benftez & D’Arcy. —A. Flowering branch. —B. Flower. —C. Flower opened
longitudinally showing corolla, anthers, and pistil. (After Luteyn 6191.)

2cm

Volume 5, Number 4
1995

Benitez & D’Arcy

New Species and Synonymy in Oestrum

315

Lund 134 (G-DC, -IDC microfiche, =F photo

006898 (MO))) did not permit verification, but its

identity with C. strigilatum seems unlikely.

Cestrum strigilatum Ruiz & Pav6n, FI. Peruv. 2:
29, t. 156. 1799. TYPE: Peru. Nemoribus ver¬
sus Pozuzo, Chinchao et Cuchero vicos, Ruiz
& Pavdn s.n. (holotype, M; isotypes, B, HAL,
fide Francey, neither seen, F photo of element
once at B, 018394).

Cestrum calycinum Kunth, Nov. Gen. Sp. 3: 58. 1818.
TYPE: Peru. Prope Gonzanama, 1180 hex, Bonpland
(B-W 4461 not seen, microfiche). Synonymized by
D’Arcy (1974); Cabrera (1983: 471).

Cestrum viridiflorum Hooker, Bot. Mag. t. 4022. 1843.
TYPE: Brazil. Porto Alegre, Tweedie (K). Synony¬
mized by Francey (1935: 137); Cabrera (1983: 471).

Cestrum unibracteatum Dunal in DC., Prodr. 13(1): 656.
1852. SYNTYPES: Peru. Sylvis Cochero, Dombey
s.n. (G-DC, MPU, neither seen), Poeppig s.n. (sub
“96 C. longium, Ruiz & Pav6n 1246”) (G-DC not
seen, =IDC microfiche, F photo 006899 MO). Syn¬
onymized by Francey (1935: 142).

Cestrum unibracteatum var. [p] brachystachys Dunal in
DC., Prodr. 13(1): 657. 1852. TYPE: Brazil. Circa
Cujaba uhique, da Silva Manso 34 (G-DC not seen,
= IDC microfiche, =F photo 023178 MO). Synony¬
mized by Francey (1935: 142).

Cestrum cancellatum Dunal in DC., Prodr. 13(1): 657.
1852. TYPE: Peru. Poeppig 3080 (G-DC not seen,
= IDC microfiche, =F photo of B, not extant 002969
MO).

Cestrum strigilatum (strigillatum) var. [a] laxiflorum Kun-
tze. Rev. Gen. PI. 3: 220. 1893. SYNTYPES: Argen¬
tina. Salta: Or£n, Lorenz & Hieronymus s.n. (NY);
Bolivia. Cochabamba: Yungas “Juntas,” Kuntze s.n.
(NY). Synonymized by Francey (1935: 142); Cabrera
(1983: 471).

Cestrum calycinum Kunth, Nov. Gen. Sp. 3: 58. 1818.
Cestrum strigilatum (strigillatum ) var. [/3] calycinum
(Kunth) Kuntze, Rev. Gen. PI. 3: 220. 1893. SYN¬
TYPES: Bolivia. Matto Grosso: West-Velasco bis
Tunarigebirge, 200-2400 m, collector not named.

Cestrum impressum Rusby, Bull. New York Bot. Gard. 4:
425. 1905. TYPE: Bolivia. La Paz: Bang 2516 (NY
not seen). Synonymized by Francey (1935: 137);
Smith & Downs (1966: 230).

Sessea rugosa Rusby, Bull. New York Bot. Gard. 8: 119.
1912. TYPE: Bolivia. Apolo, 4800 ft., R. S. Williams
2449 (NY not seen). Synonymized by Francey (1935:
142).

Cestrum calycinum var. tenuijlorum F rancey, Candollea 6:
142. 1935. Syn. nov. SYNTYPES: Paraguay. Villa
Fmcamaci6n, Bettfreund 131; Argentina. Yaguara-
Zapa, Niederlein 268b (G not seen, F photo 002998
MO).

Cestrum strigilatum var. tenuijlorum Francey, Candollea 6:
144. 1935. Syn. nov. SYNTYPES: Ecuador. Balao,
Eggers 14274 (lectotype, designated here, W; isolec-
totype, US fragment). Guayaquil, Jameson 57 (B, M,
W, none seen).

Cestrum aristeguietae Steyermark, Acta Bot. Venez. 6(1-
4): 86. 1971. TYPE: Venezuela. Carabobo: wooded
slopes along Rfo San Gian, 2 km below Planta Elec¬

trics, S of Borburata, alt. 350 m, J. & C. Steyermark
95463 (holotype, VEN; isotypes, NY, P, US).
Cestrum longifolium Ruiz & Pavdn ex Dunal in DC.,
Prodr. 13(1): 657. 1852. Invalid name, cited in syn¬
onymy of C. unibracteatum.

Cestrum tiilettii Benftez & D’Arcy, sp. nov. TYPE:
Venezuela. Zulia: Distrito Perijfi, headwaters of
Rfo Guasare, Sierra de PerijA, Serranfa de Va-
lledupar, environs of Campamento Frontera V,
along international boundary, 2700-3300 m,
10°23'07.8"N, 72°52'42.5"W, 10-19 July
1974 (fl), lillett 747-1021 (holotype, MY; is¬
otypes, AAU, MO, MYF, VEN). Figure 3.

Arbor 2—3 m alta, foliis glabris coriaceis, marginibus
revolutis, foliis minoribus carentibus. Inflorescentia axil¬
laris, congesta, tomentosa. Flores fragrantes diurni casta-
nei, sessiles, calyce viride apice purpureo tubuloso, co¬
rolla 15—18.5 mm longa, staminibus 10-13 mm longis,
geniculatis tumidisque ad insertionem; ovula 5. Fructus
ovoideus 9X6 mm.

Tree 2-3 m tall; stems brown, glabrous, striate
and scarred. Leaves firm, subcoriaceous, narrowly
elliptical, basally narrowly cuneate, slightly acute
near the top, the tip obtuse, 4-6 X 1.3-1.5 cm,
dark green, shiny above, dull green beneath, the
major veins sunken above, salient beneath, 6—8
on each side, the minor veins impressed beneath,
the margin revolute, glabrous on both sides; pet¬
iole canaliculate, 3—7 mm long, glabrous. Minor
leaves wanting. Inflorescences axillary, congested,
the main axes tomentose, bright matte green, 1.5—
3 cm long, the peduncles 5—7 mm long, pubes¬
cent, thickened, with circular scars from fallen
flowers, flowers few per node. Flowers sessile
(pedicels obsolete) with faint, sweet fragrance dur¬
ing the day; calyx yellow-green, flushed distally
with dark purple, tubular, thick, costate, the cos¬
tas especially conspicuous distally, slightly pu¬
bescent outside with hairs slightly thickened near
the base, 4.5-5.5 mm long, 2-2.5 mm thick, 5-
lobate, the lobes 1.5-2.5 mm long, narrowly tri¬
angular, the apex pubescent; corolla 15-18.5 mm
long, the tube 13—16 mm long, slightly expanded
toward the apex, the throat not noticeably con¬
tracted, the mouth ca. 2 mm wide, 5-lobate, the
lobes 2-2.5 mm long, the folded margins puber-
ulent; stamens 10-13 mm long; filaments white,
adnate 6-9 mm, free 4.5 mm, the point of inser¬
tion geniculate-tumid, slightly pilose, 1 mm long;
anthers brown, spherical, 0.5 mm diam.; ovary
ovoid, glabrous, smooth, 1 mm diam., 5-ovulate;
style 12-13.5 mm long, papillose 1-2 mm below
the stigma, exceeding the stamens by 1.5 mm;
stigma green. Fruit ovoid, 9X6 mm, pericarp
thick, opaque; seeds 6—7, 3—3.5 mm long.

316

Novon

Figure 3. Cestrum tillettii Benftez & D’Arcy. —A. Flowering branch. —B. Flower. —C. Flower opened longitudinally.
—D. Fruits. —E. Leaf, abaxial side. (A, B, C after Tillet 747-1021; E after Tillet 747-929.)

Cestrum tillettii occurs in cloud forests at the
headwaters of rivers at elevations of 2700-3650 m.
It has been found on the eastern side of the Sierra
de PerijA, a mountain range shared by Venezuela
and Colombia.

This species is named in honor of Stephen S.
Tillett, professor in the Faculty of Pharmacy at the
Universidad Central de Venezuela, Caracas. He was
the leader of the expedition that collected the only
known specimens of this species.

1

1cm

Volume 5, Number 4
1995

Benitez & D’Arcy

New Species and Synonymy in Oestrum

317

Paratypes. VENEZUELA. Zulia: Distrito Perij4,
headwaters of Rfo Ap6n, Sierra de Perij£, Serranfa Valle-
dupar. Hacienda Buena Vista to N side of Cerro Lami-
nado, ca. 5 km N of Buena Vista, along international
boundary, 3300-3650 m, 10°20'23"N, 72°54'14"W, 9-10
July (fr), Tillett & Honig 747-929 (MY); Dist. Perijd,
headwaters of Rfo Guasare along international boundary,
“Campamento Frontera V,” 2700-3300 m, 10°23'7.8"N,
72°52'45.5"W, 20-23 July (fl, fr). Wood & Berry 88 (MO,
VEN).

Oestrum tomentosum is one of the most wide¬
spread species of Oestrum , and in many places it
is common, especially as a successional element,
occurring from about 1000 to 2600 m elevation.
The species has been collected from as far north
as Sonora, Mexico, and as far south as extreme
southern Peru. It is also found in northwestern Ven¬
ezuela but is generally absent from the Amazon re¬
gion of South America. It is a shrub or small tree,
usually erect, with pubescent leaves, calyces, and
corolla tubes, and it has a shiny, purple-black,
fleshy fruit.

The identity of many of the names cited here was
suggested by Nee (1986: 49), who also provided a
good description (as C. lanatum).

Cestrum tomentosum L. f., Suppl. PI. 150. 1781.
TYPE: Colombia. Mutis 94 (holotype, LINN
258.6 not seen, =IDC microfiche).

Oestrum hirsutum Jacquin, PL Hort. Schoenbr. 3: 41, t.
324. 1798. Syn. nov. TYPE: Indias Occidentales:
cult. hort. Schoenbrunn (W, B-W 4449, not seen,
=IDC microfiche, =F photo 033032 MO).

Cestrum lanuginosum Ruiz & Pav6n, Fl. Peruv. 2: 30, tab.
157. 1799. Syn. nov. TYPE: Peru. Prov. Arequipa:
Camana, collibus arenosis, Ruiz s.n. (lectotype, des¬
ignated here, G not seen, =F photo 002984; isolec-
totype, F).

Cestrum lanatum Martens & Galeotti, Bull. Acad. Roy.
Sci. Bruxelles 12(2): 146. 1845. TYPE: Mexico. Ve¬
racruz: dans les bois et haies de la colonie de Mir-
ador, 3000 pieds, Galeotti 1208 (holotype, BR not
seen; isotypes, G, K, NY, US, W).

Cestrum moritzii Dunal in DC., Prodr. 13 (1): 619. 1852.
Syn. nov. TYPE: Venezuela. Caracas, Moritz 309 (ho¬
lotype, G-DC not seen, =IDC microfiche, =F photo
002988; isotypes, BM, MO).

Cestrum miersianum Weddell, Chlor. And. 2: 97. 1857.
TYPE: Colombia. Sierra Nevada de Santa Marta,
3,300 m, Linden 1615 (holotype, G-DC not seen,
= IDC microfiche).

Cestrum miersianum Pittier, J. Wash. Acad. Sci. 22(2): 37.
1932, not Weddell (1857). Syn. nov. TYPE: Vene¬

zuela. Merida: San Rafael de Mucuchfes, 3150 m,
A. Jahn 767 (holotype, VEN; isotype, MO). Cestrum
neomiersianum Benftez, Rev. Fac. Agron. (Maracay)
7: 90. 1974. New name.

Cestrum diasae Pittier, J. Wash. Acad. Sci. 22 (2): 29.
1932. Syn. nov. TYPE: Venezuela. M6rida: Misintd,
arriba de Mucuchfes, 3500 m, Pittier 12919 (holo¬
type, VEN; isotype, US).

Cestrum meridanum Pittier, J. Wash. Acad. Sci. 22(2): 36.
1932. Syn. nov. TYPE: Venezuela. Vecinidades de
Merida, 1700 m, Pittier 12858 (holotype, VEN; iso¬
type, US).

Cestrum densiflorum Francey, Candollea 6: 195. 1935.
Syn. nov. SYNTYPES: Venezuela. Merida: entre
Chachopo y Timotes, Pittier 13294 (lectotype, des¬
ignated here, NY; isolectotypes, F, MO, US, VEN);
Pittier 13323 (NY, VEN).

Cestrum densiflorum Francey var. puberulum Francey,
Candollea 6: 196. 1935. Syn. nov. TYPE: Venezuela.
Trujillo. San Pablo de Mendoza, Pittier 13323 (ho¬
lotype, VEN; isotype, US).

Cestrum verbascifolium Zuccarini ex Francey, Candollea
6: 191. 1935. Syn. nov. TYPE: cultivated in Berlin
and Munich (B-W 4449 not seen, =IDC microfiche).
Cestrum ambatense Francey, Candollea 6: 169. 1935. Syn.
nov. TYPE: Ecuador. Prov. Tungurahua: vicinity of
Ambato, Pachano 75 (holotype, NY; isotype, US
fragment).

Acknowledgments. The authors acknowledge
grants from the Consejo Nacional de Investiga-
ciones Cientfficas y Tecnoldgicos, CONICIT (PI-
056) and the National Science Foundation, NSF
(INT-9116039). Michael Nee, The New York
Botanical Garden, reviewed the manuscript and
augmented parts of it.

Literature Cited

Cabrera, A. L. 1983. Flora de la Provincia de Jujuy, Re-
publica Argentina. Colecci6n Cientffica del INTA, Bue¬
nos Aires.

D’Arcy, W. G. 1972 [1973]. Solanaceae. In: R. E. Wood-
son et al., Flora of Panama. Ann. Missouri Bot. Gard.
60: 573-870.

Francey, P. 1935-1936. Monographic du genre Cestrum
L. Candollea 6: 46-398; 7: 1-132.

Kunth, C. S. 1818. Solaneae. In: Nova Genera et Species
Plantarum, Vol. 3. Paris.

Nee, M. 1986. Solanaceae. In: Flora de Veracruz. 49: 1-
191. Inst. Nac. Invest. Recursos Bi6t., Xalapa.

Roemer, J. J. & J. A. Schultes. 1819. Caroli a Linn6
equitis Systema vegetabilium secundum classes ordines
genera species, Vol. 4. Stuttgart.

Smith, L. B. & R. J. Downs. 1966. Solan£ceas. Parte I.
In: P. R. Reitz (editor). Flora Illustrada Catarinense.
Herb£rio “Barbosa Rodrigues,” Itajai, Santa Catarina,
Brazil.

Two New Species of Fuchsia Section Fuchsia (Onagraceae) from
Southern Ecuador

Paul E. Berry

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.

Abstract. Two new species of Fuchsia sect.
Fuchsia (Onagraceae) from the Andes of southern
Ecuador are described and illustrated. Fuchsia
campii P. E. Berry is known from mid-elevation
cloud forest of Azuay and Loja Provinces, and
Fuchsia summa P. E. Berry is a high-elevation spe¬
cies currently known only from Loja Province. Both
occur in Podocarpus National Park, together with
several other rare species of Fuchsia.

Studies and collections of plants from southern
Ecuador have increased dramatically in the past de¬
cade, especially in areas such as Podocarpus Na¬
tional Park in Loja Province. Many novel plants have
been located, and this paper describes two new spe¬
cies of Fuchsia that occur in cloud forest areas on
the slopes of Cerro Toledo, one of them also known
further north in Azuay Province. Three other species
previously thought to be extremely rare in southern
Ecuador are also known from Cerro Toledo: F. andrei
I. M. Johnston, F. scherffiana Andre, and F. steyer-
markii P. E. Berry. All these taxa belong to the larg¬
est section of the genus, section Fuchsia, which now
approaches 65 species. They are also examples of
the high degree of local endemism in cloud forests
of southern Ecuador, where moist montane habitats
are often separated by dry valleys and show a wide
range of precipitation regimes.

Fuchsia campii P. E. Berry, sp. nov. TYPE: Ec¬
uador. Azuay: Quebradas leading into the Rfo
Collay, 3—8 km N of Sevilla de Oro, 7000—
8300 ft., 27 Aug. 1945, W. H. Camp E-4994
(holotype, RSA herbarium # 71952; isotypes,
G-DEL, MO, NY, UC, US). Figure 1.

Frutex 1.5-4 m altus, foliis 3-vel 4-verticillatis, inter-
dum oppositis, 3—9 X 1—3 cm, margine subintegro vel 7—
10-dentato, floribus axillaribus apicem versus, pedicellis
18-30 mm longis, tubo florali roseo 29-45 mm longo, se-
palis lanceolatis 12-15 mm longis, petalis auriantiacis 8-
11 mm longis, 4—6 mm latis.

Erect to recumbent shrub 1.5-4 m tall, well
branched with mosdy ascending branches; young
growth pubescent, the trichomes 0.1-0.5 mm long,
appressed to erect; branchlets terete to slighdy an-

Novon 5: 318-322. 1995.

gled, older branches with purplish, exfoliating bark.
Leaves mostly in whorls of 3 or 4, sometimes opposite
or subopposite; blade membranous, narrowly elliptic,
3—9 cm long, 1-3 cm wide, acute to acuminate at the
apex, acute at the base, subglabrous to sparsely stri-
gose when young on both surfaces and margin, tri¬
chomes persisting mainly along the midvein and sec¬
ondary nerves on lower surface when fully expanded,
upper surface smooth, shiny, and light to dark green,
lower surface paler green; margin nearly entire, usu¬
ally with 7-10 glandular teeth projecting slightly from
the edge especially in the upper Vi to % of the blade;
secondary nerves (5-)8-10 on either side of the mid¬
vein; petiole strigose, (4—)6-25 mm long; stipules nar¬
rowly lanceolate when young, 1—1.5 mm long, ca. 0.2
mm wide, the tip quickly abscising and the thicker
lower third persisting or abscising later. Flowers ax¬
illary in the upper leaf axils, 2-several per node, ped¬
icel 18-30 mm long; floral tube narrowly funnelform,
(29-)32-45 mm long, 2.5-3.5 mm wide at the base,
narrowed to 2—2.5 mm wide in the lower V4, then
gradually widened until 5-8 mm wide at the rim,
sparsely strigose outside, densely pubescent inside in
the lower narrowed portion; nectary a doughnut¬
shaped ring 1—2 mm high and 2—3 mm wide at the
base of the tube surrounding the style; ovary 4-angled,
5-6 mm long, 2-3 mm thick; sepals narrowly trian¬
gular or lanceolate, 12-15 mm long, 4—5 mm wide at
the base; petals elliptic to obovate, 8-11 mm long, 4—
6 mm wide, broadly acute at the apex, sometimes with
a small mucro; tube and sepals bright pink except for
the green sepal tips, petals orange and darker than
the tube; style pink, 40-64 mm long, densely pubes¬
cent in lower third, stigma pink, capitate, ca. 2 mm
high, ca. 2.5 mm wide; filaments 10-12 mm and 7—
9 mm long, anthers oblong, 2-3 mm long, 1-1.5 mm
wide. Fruit oblong-ellipsoid before maturity, subro¬
tund when ripe, 9-15 mm long, 5-9 mm thick; seeds
flattened, obtriangular in outline, 1.8-2.2 mm long,
1.2—1.6 mm wide, ±200 per fruit. Gametic chromo¬
some number n = 11.

Ecology and distribution. Occurring in thick¬
ets, along streams, and along roadsides in pasture
zones on moist mountain slopes, 2300—2700
(—3350) m, in Azuay and Loja Provinces, Ecuador.

Volume 5, Number 4
1995

Berry

Fuchsia from Southern Ecuador

319

Figure 1. Fuchsia campii P. E. Berry. —A. Habit. -B. Detail of stipules. -C. Flower at anthesis. —D. Flower in
longitudinal cross section. —E. Petal. —F. Anther. Drawn from Berry & Brako 4646.

Etymology. Named for Wendell H. Camp
(1904—1963), a botanist who made extensive plant
collections in southern Ecuador for the New York
Botanical Garden in the mid 1940s.

The type specimen of Fuchsia campii was treated
by Munz (1974) under Fuchsia ayavacensis HBK,

but that species has wider and shorter petals, larger
and more pubescent leaves, and occurs in drier
habitats in southern Ecuador and in northern Peru.
One of the paratypes cited here (Camp E-4257) was
listed by Munz under F. canescens Bentham, but
that species is confined to southern Colombia and

320

Novon

has thick, firm flowers, short pedicels, trowel¬
shaped petals, tetragonous sepals in bud, and most¬
ly 4-whorled leaves. Fuchsia campii might be con¬
fused with F. vulcanica Andre (sensu lato) but
differs in its narrower, orange petals contrasting
with the pink floral tube and sepals (the sepals also
turn green toward the apex) and in the narrower
leaves with margins entire or else with small teeth.
It is apparently diploid, as chromosome counts of
n = 11 and 2n = 22 were obtained from Escobar
1543, and three collections identified as F. vulcan¬
ica from Azuay Province were tetraploid (Berry,
1982).

Paratypes. ECUADOR. Azuay: E Cordillera, 1—8 km
N of the village of Sevilla de Oro, 8000-9000 ft., 27 July
1945, Camp E-4257 (NY, RSA); quebradas leading into
the Rfo Collay, 3—8 km N of Sevilla de Oro, 7000-8000
ft., 27 Aug. 1945, Camp E-4994 (MO); Partidero Llantera-
Chiquintad-Saucay-Guandum, 10,000 ft., 27 May 1979,
Jaramillo 1068 (AAU, QCA). Loja: road from Yangana to
Cerro Toledo, 2360 m, 13 Nov. 1988, Berry & Brako 4646
(MO, QCA); between Yangana and Toledo, 7700-8000 ft.,
26 July 1979, Escobar 1543 (MO, TEX), 1544 (MO, TEX),
1545 (MO, TEX).

Fuchsia summa P. E. Berry, sp. nov. TYPE: Ec¬
uador. Loja: shrub patches in open pArarno
above tree line, road from Yangana to Cerro
Toledo, 3150 m, 13 Nov. 1988. P. E. Berry &
L Brako 4644 (holotype, MO; isotypes, QCA,
US). Figure 2.

Frutex 30—100 cm altus, ramulis erectis foliis apicem
versus congestis 3-vel 4-verticillatis, 15-35 mm longis,
petiolis 5-8 mm longis. Flores axillares, 4-angulati, ped-
icellis 4—7(—10) mm longis, tubo florali 18-26(-32) mm
longo, sepalis 10—14 mm longis, petalis suborbicularibus
8-12 X 7-9 mm.

Erect shrub 30—100 cm tall; branches erect,
mostly bare except for the leafy tips, with short
intemodes mostly 6-12(-25) mm long and pro¬
truding leaf base scars, branchlets appressed-pu-
bescent with whitish trichomes. Leaves mostly in
whorls of 3 or less often 4, densely clustered at
the stem apices; blade subcoriaceous, broadly el¬
liptic, 15—35 mm long, 12—18 mm wide, broadly
to narrowly acute or rounded at the apex and base,
usually with scattered appressed trichomes along
the midvein on both surfaces, upper surface dark
green, lower surface paler green; margin gland-
denticulate with trichomes between the teeth; sec¬
ondary nerves 4—6 on either side of the midvein;
petiole 5-8 mm long; stipules narrowly lanceolate,
1—2 mm long, 0.3—0.5 mm wide. Flowers axillary
and pendent in the upper leaf axils, 2-several per
node, pedicel 4—7 mm long in flower, to 10 mm

long in fruit; floral tube narrowly funnelform, 18—
26(-32) mm long, 2-2.5 mm wide at the base, 4—
6 mm wide at the rim, 4-angled in cross section,
appressed-pubescent to nearly glabrous in the
lower half outside, densely villous in lower half
inside; nectary a doughnut-shaped ring ca. 1.5
mm high and 2 mm wide at the base of the tube
surrounding the style; ovary 4-angled, 5—6 mm
long, 2-3 mm thick; sepals broadly elliptic, acute
to subacuminate at the apex, 10—14 mm long, 5—
6 mm wide at the base, quadrangular in cross sec¬
tion when joined before opening; petals suborbi-
cular, slightly spreading, 8-12 mm long, 7—9 mm
wide, broadly acute to mucronate at the apex,
broadly acute to spathulate at the base; tube pale
red, sepals pale green in bud but becoming
flushed with red by anthesis, petals slightly darker
red than the tube; style red, 28—35 mm long, vil¬
lous in lower half, stigma pink, capitate, ca. 2 mm
high by 2 mm wide; filaments 5—7 mm and 4-5
mm long, anthers broadly oblong, ca. 2 mm long
and 1.5 mm wide. Fruit 4-angled before fully ma¬
ture, 9-12 mm long, ca. 5 mm thick; seeds flat¬
tened, obtriangular in outline, 1.9—2.1 mm long,
ca. 1.5 mm wide.

Ecology and distribution. Occurring in low
shrub patches among open grassy areas at and
above tree line, on very humid upper slopes of Cer¬
ro Toledo east of Yangana in Loja Province, Ecua¬
dor, 3100-3450 m elevation.

Etymology. From the Latin summus, meaning
highest or uppermost, in reference to this species
occurring at tree line and at higher elevations than
any other species of Fuchsia on the same mountain.

This species forms small, dense thickets in ex¬
posed areas close to tree line. Its leaves are small,
with few secondary veins, and are tightly congest¬
ed toward the stem apices. The flowers are dis¬
tinctive in their short pedicels that angle sharply
downward alongside the stems and in the strongly
quadrangular ovary, floral tube, and sepals in bud.
Fuchsia summa appears to be morphologically
most similar to F. loxensis HBK, a broadly circum¬
scribed species in Berry (1982). Besides the char¬
acters listed above, F. summa differs in the green¬
ish coloration of the sepals and generally more
pubescent flowers.

Paratypes. ECUADOR. Loja: road from Yangana to
Cerro Toledo, 3140 m, 12 Nov. 1988, Berry & Brako 4643
(MO, QCA); Cerro Toledo, Parque Nacional Podocarpus,
3000-3400 m, 27 Nov. 1988, Madsen 75617 (AAU, MO),
1 Dec. 1988, Madsen et al. 75640 (AAU), 21 July 1989,
Madsen et al. 86119 (AAU, MO), 30 Oct. 1989, Madsen
86320 (AAU), 30 Oct. 1989, Madsen 86343 (AAU); Cerro

Volume 5, Number 4 Berry

1995 Fuchsia from Southern Ecuador

e

E

io

I-1

321

Figure 2. Fuchsia summa P. E. Berry. A. Habit. B. Detail of stipules. —C. Leaf and detail of margin. —D. Flower at anthesis. —E. Flower in longitudinal cross section with
details of the stigma and nectary; at lower left a cross section of the ovary. —F. Detail of a sepal, petal, and two stamens. Drawn from Berry & Brako 4644.

322

Novon

Toledo E of Yangana, Parque Nacional Podocarpus, 3400-
3450 m, 26 Feb. 1985, 0llgaard et al. 58186 (AAU).

Acknowledgments. This work was supported by
grant 3827-88 of the National Geographic Society.
The illustrations were prepared by Bobbi Angell,
and Roy Gereau kindly reviewed the Latin diag-

Literature Cited

Berry, P. E. 1982. The systematics and evolution of Fuch¬
sia sect. Fuchsia (Onagraceae). Ann. Missouri Bot.
Gard. 69: 1—198.

Munz, P. A. 1974. Flora of Ecuador. 141. Onagraceae.
Opera Bot. Ser. B no. 3, 3—46.

New Combinations in Chamaesyce (Euphorbiaceae) from
Queensland, Australia

Paul 1. Forster and Rodney J. F. Henderson
Queensland Herbarium, Queensland Department of Environment & Heritage,
Meiers Road, Indooroopilly, Queensland 4068, Australia

Abstract. The new combinations Chamaesyce
carissoides (F. M. Bailey) D. C. Hassall ex P. I. For¬
ster & R. J. F. Henderson, C. centralis (B. G.
Thompson) P. I. Forster & R. J. F. Henderson, C.
coghlanii (F. M. Bailey) D. C. Hassall ex P. I. For¬
ster & R. J. F. Henderson, C. petala (Ewart & L.
R. Kerr) P. I. Forster & R. J. F. Henderson, and C.
psammogeton (P. S. Green) P. I. Forster & R. J. F.
Henderson are made.

Recognition of Chamaesyce S. F. Gray as a genus
distinct from Euphorbia L. has received consider¬
able support (e.g., Hassall, 1976; Koutnik, 1984,
1987; Smith, 1981; Webster, 1994) or rejection
(e.g., Radcliffe-Smith, 1980; McPherson & Tirel,
1987; Oudejans, 1990) among taxonomists. Revi¬
sions of Chamaesyce and Euphorbia s. str. in Aus¬
tralia were undertaken by Hassall (1977a). These
were unfortunately never published in full; only a
revision of the latter genus (Hassall, 1977b), valid
transfers of some names into Chamaesyce (Hassall,
1976), and discussion of arid zone speciation pat¬
terns (Hassall, 1982) appeared in print.

Recognition of Chamaesyce in Australia subse¬
quent to Hassall’s work has not been widespread,
with most authors opting for a broadly circum¬
scribed Euphorbia (e.g., Forster, 1993, 1994;
OGreen, 1993; Forster & Henderson, 1994; Thomp¬
son, 1992; Weber, 1986; Wheeler, 1987) as op¬
posed to the narrower view of James and Harden
(1990). Despite this lack of local support, we are
now of the opinion that the arguments proposed by
Hassall (1976), Koutnik (1984, 1987), and Webster
(1994) for recognizing Chamaesyce as a genus in
its own right are convincing and that their views
should be supported. As a new edition of “Queens¬
land Vascular Plants Names and Distribution” the
Queensland Herbarium (BR1) census of Queens¬
land Plants in which the authors intend to recog¬
nize Chamaesyce as a distinct genus is soon to be
prepared, new names are required for certain taxa.
While valid combinations exist in Chamaesyce for
the majority of taxa that occur in Queensland, no
combinations exist for five species. Therefore, these

combinations are made here. Two of these combi¬
nations were listed in the unpublished thesis of
Hassall (1977a) but never published.

1. Chamaesyce carissoides (F. M. Bailey) D. C.

Hassall ex P. I. Forster & R. J. F. Henderson,
comb. nov. Basionym: Euphorbia carissoides F.
M. Bailey, Queensland Agric. J. 16: 449.
1906. TYPE: Australia. Queensland: Cook
District, Herberton, Ringrose [AQ342536] (ho-
lotype, BRI).

2. Chamaesyce centralis (B. G. Thompson) P. I.

Forster & R. J. F. Henderson, comb. nov. Bas¬
ionym: Euphorbia centralis B. G. Thompson,
Nuytsia 8: 353. 1992. TYPE: Australia. North¬
ern Territory: 3 km SW of Alice Springs, 18
Jan. 1990, Thompson 3408 (holotype, DNA not
seen; isotypes, BRI; AD & CANB not seen).

3. Chamaesyce coghlanii (F. M. Bailey) D. C.

Hassall ex P. I. Forster & R. J. F. Henderson,
comb. nov. Basionym: Euphorbia coghlanii F.
M. Bailey, Bot. Bull. 13: 12. 1896. TYPE:
Australia. Queensland: Gregory North District,
Roxborough, Georgina River, Dec. 1895, Bai¬
ley [AQ342538] (holotype, BRI).

4. Chamaesyce petala (Ewart & L. R. Kerr) P. I.

Forster & R. J. F. Henderson, comb. nov. Bas¬
ionym: Euphorbia petala Ewart & L. R. Kerr,
Proc. Roy. Soc. Victoria, New Series 39(1): 1.
1926. TYPE: Australia. Northern Territory:
Wycliffe Well, June 1924, Ewart (holotype,
MEL not seen).

5. Chamaesyce psammogeton (P. S. Green) P.

I. Forster & R. J. F. Henderson, comb. nov.
Basionym: Euphorbia psammogeton P. S.
Green, Kew Bull. 48: 314. 1993. TYPE: Aus¬
tralia. Lord Howe Island: Blinky Beach, 13
Nov. 1963, Green 1625 (holotype, K not seen;
isotype, A not seen).

Literature Cited

Forster, P. I. 1993. Rediscovery of Euphorbia carissoides
(Euphorbiaceae). Austral. Syst. Bot. 6: 269—272.

Novon 5; 323-324. 1995.

324

Novon

-. 1994. Tropical beaches and Euphorbia atoto in

Australia and Papuasia. Cact. Succ. J. (U.S.) 66: 7-10.

-& R. J. F. Henderson. 1994. Euphorbiaceae. Pp.

107—117 in: R. J. F. Henderson (editor), Queensland
Vascular Plants: Names and Distribution. Queensland
Department of Environment & Heritage, Brisbane.

Green, P. S. 1993. Notes relating to the floras of Norfolk
& Lord Howe Island, IV. Kew Bull. 48: 307-325.

Hassall, D. C. 1976. Numerical and cytotaxonomic evi¬
dence for generic delimitation in Australian Euphor-
bieae. Austral. J. Bot. 24: 633-640.

-. 1977a. Systematic studies in Australian Eu-

phorbieae (Euphorbiaceae). Doctor of Philosophy The¬
sis, University of Queensland. St Lucia, Brisbane.

-. 1977b. The genus Euphorbia in Australia. Aus¬
tral. J. Bot. 25: 429-453.

-. 1982. Distribution and evolution of Euphorbia

and Chamaesyce (Euphorbiaceae) in the arid zone of
Australia. Pp. 323-328 in W. R. Barker & P. J. M.
Greenslade (editors), Evolution of the Flora and Fauna
of Arid Australia. Peacock Publications, Frewville.

James, T. A. & G. J. Harden. 1990. Euphorbiaceae. In:
G. J. Harden (editor), FI. New South Wales 1: 389—430.
Univ. New South Wales Press, Sydney.

Koutnik, D. L. 1984. Chamaesyce (Euphorbiaceae) — A
newly recognised genus in southern Africa. S. African
J. Bot. 3: 262-264.

-. 1987. A taxonomic revision of the Hawaiian

species of the genus Chamaesyce (Euphorbiaceae). Al-
lertonia 4: 331-388.

McPherson, G. & C. Tirel. 1987. Flore de la Nouvelle-
Cal6donie et Dependences. 14. Euphorbiaceae I. Mu¬
seum National D’Histoire Naturelle, Paris.

Oudejans, R. C. H. M. 1990. World Catalogue of Species
Names Published in the Tribe Euphorbieae (Euphorbi¬
aceae), With Their Geographical Distribution. R. C. H.
M. Oudejans, Utrecht.

Radcliffe-Smith, A. 1980. Euphorbia. In: A. K. Airy
Shaw, The Euphorbiaceae of New Guinea. Her Majesty’s
Stationery Office, London.

Smith, A. C. 1981. Euphorbiaceae. In: FI. Vit. Nova 2:
439-575. Pacific Tropical Botanical Gardens, Lawai.

Thompson, B. G. 1992. Three new species of Euphorbia
L. subgenus Chamaesyce Rafinesque (Euphorbiaceae)
from central and northern Australia. Nuytsia 8: 351—
360.

Weber, J. Z. 1986. Euphorbiaceae. In: J. P. Jessop & H.
R. Toelken (editors), FI. South Australia 2: 735-768.
Government Printer, Adelaide.

Webster, G. L. 1994. Synopsis of the genera and supra-
generic taxa of Euphorbiaceae. Ann. Missouri Bot.
Gard. 81: 33-144.

Wheeler, J. R. 1987. Euphorbiaceae. In: N.G. Marchant
et al., FI. Perth Region, 444-455. Western Australian
Herbarium, South Perth.

Gladiolus somalensis (Iridaceae), a New Species from
Northeastern Somalia

Peter Goldblatt

B. A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Mats Thulin

Department of Systematic Botany, University of Uppsala, Villavagen 6,
S-752 36 Uppsala, Sweden

Abstract. Gladiolus somalensis is a new species
restricted to the limestone escarpments of the Sa-
naag and Bari Regions of northeastern Somalia. Al¬
though Gladiolus is the largest genus of Iridaceae
subfamily Ixioideae and has some 83 species in
tropical Africa, only 4 species occur in Somalia and
only G. somalensis is endemic there. The affinities
of this new species are uncertain, but the relatively
small flowers with a short perianth tube appear to
place the species in subgenus Gladiolus, a taxon
that is most diverse and speciose in southern Af¬
rica.

The Old World genus Gladiolus, a member of
Iridaceae subfamily Ixioideae and one of the larger,
if not the largest genus of the family, consists of
some 250 species (Goldblatt, in press; Goldblatt &
Manning, ms.). The center of Gladiolus in terms of
species numbers and taxonomic diversity is tem¬
perate southern Africa, more particularly the winter
rainfall region of southern Africa (Lewis et al.,
1972; Goldblatt, 1991). The genus is, however, well
represented in tropical Africa, where some 83 spe¬
cies occur south of the Sahara and north of the
borders of Namibia, Botswana, and South Africa
(Goldblatt, in press), 75 of them endemic. At least
150 species occur in southern Africa (Goldblatt &
Manning, in prep.), another 8 in Madagascar (Gold¬
blatt, 1989), and perhaps 10 more in Europe, North
Africa, and the Middle East. Here we describe a
new species of Gladiolus restricted to northeastern
Somalia, one of only four species of the genus re¬
corded from that country, and the only one endemic
there (Goldblatt, 1995).

Although first collected in 1956, Gladiolus so¬
malensis remained until now too poorly known to
be described or even assigned with confidence to
genus. The type collection made in January 1995,
however, is well preserved, and is accompanied by
photographs and spirit material. This has made it

possible to draw up a description, formally name
the species, and have an illustration made.

Gladiolus somalensis Goldblatt & Thulin, sp. nov.
TYPE: Somalia. Sanaag Region, escarpment S
of Laasqoray, near Ragad, 11°00'N, 48°29'E,
evergreen bush on limestone, 16 Jan. 1995,
Thulin, Dahir & Hassan 9079 (holotype, UPS).
Figure 1.

Plantae (7—)12—30 cm altae, eormo ca. 12 mm in dia-
metro, foliis 4—5 linearibus (1-12—1 mm latis, spica 2—10
florum, bracteis extemis viridibus (7—)12—17(-23) mm
longis internis minoribus, flos aurantiacus, tepalis later-
alibus infemis infra flavis (raro tepalo inferiore infra flavo),
tubo perianthii infundibuliformi 6-8 mm longo, tepalis
lanceolatis tribus supemis majoribus 16-18 X ca. 8 mm,
infemis 12—15 X 5—5.5 mm, filamentis 8-10 mm longis,
antheris 3—5 mm longis, ramis styli filiformibus ca. 2.5
mm longis.

Plants (7-)12-30 cm high. Corm obconic, ca. 12
mm diam., the tunics of softly textured layers, these
decaying with age into fine netted fibers. Leaves
four or five, the lower three basal and longest,
reaching at least to the base of the spike and one
or more often slightly exceeding the spike, the
blades ± linear, (l-)2-4 mm wide, the upper one
or two leaves inserted on the lower half of the stem,
smaller than the basal leaves. Stem erect, simple
or with one or two branches, ca. 1.2 mm diam. be¬
low the base of the spike. Spike lightly flexuose, 2-
10-flowered; bracts green and soft-textured, the out¬
er (7-)12-17(-23) mm long, the inner about
two-thirds as long as the outer. Flowers zygomor-
phic, orange, the lower lateral tepals (rarely the
lower median tepal) bright yellow in the lower half;
perianth tube funnel-shaped, 6-8 mm long; tepals
unequal, lanceolate, the upper three larger than the
lower, the dorsal inclined over the stamens, 16-18
X 8 mm, the upper laterals about as long, the lower
tepals ± parallel to the ground, the lower lateral

Novon 5: 325-328. 1995.

326

Novon

Figure 1. Gladiolus somalensis Goldblatt & THulin
size. (Drawn by John C. Manning.)

(from Thulin, Dahir & Hassan 9079). Scale: approximately life

Volume 5, Number 4
1995

Goldblatt & Thulin
Gladiolus somalensls

327

tepals ca. 15 X 5.5 mm, the margins raised below
and the surface channeled in the lower half, the
lower median ca. 12 X 5 mm, not normally chan¬
neled. Filaments 8—10 mm long, exserted 1—6 mm
from the tube; anthers 3—5 mm long, yellow. Ovary
ovoid, 2—3 mm long; style arching over the stamens,
dividing ca. 1.5 mm beyond the anther apices, the
branches ca. 2.5 mm long, filiform, evidently not
expanded apically. Capsules and seeds unknown.

Flowering January and February.

Distribution and Habitat

Still poorly known, Gladiolus somalensis appears
to be restricted to the limestone escarpments of the
Sanaag and Bari Regions facing the Gulf of Aden
in northeastern Somalia. In the type locality, in the
eastern part of the Cal Madow Range in Sanaag,
only a single plant was seen despite extensive
searching. This was growing in evergreen bushland
dominated by Buxus hildebrandtii Baillon (Buxa-
ceae), a native boxwood, and Cadia purpurea Forss-
kal (Fabaceae) at 1350 m elevation. Gladiolus
schweinfurthii Baker, a species of moderate eleva¬
tions in Eritrea, Ethiopia, and Kenya, is a fairly
common and conspicuous plant in the area. Cal
Madow is a major center of endemism in Somalia,
described in detail by Thulin (1994a).

Two early collections are referred to Gladiolus
somalensis, both from the Cal Miskat Range in Bari
Region, some 150 km to the east of the type local¬
ity. Nothing is known about the habitat of the spe¬
cies in this area, one that is generally drier than
Cal Madow, and from where neither Buxus or Cadia
is known.

Diagnosis, Variation, and Relationships

Gladiolus somalensis is readily distinguished
from all other tropical African species of the genus
by its small, bright orange flowers with a conspic¬
uous yellow nectar guide located on the lower lat¬
eral tepals (or sometimes, and presumably abnor¬
mally, only on the lower median tepal) and fairly
short perianth tube, 6—8 mm long (Fig. 1). The flow¬
ers have a superficial resemblance to species of
Tritonia, a genus in which most species have bright
orange flowers with yellow markings on the lower
tepals. A characteristic feature of most orange-flow¬
ered species of Tritonia is the presence of a large,
tooth-like callus in the lower center of each of the
lower tepals, and this feature is lacking in G. so¬
malensis. Gladiolus and Tritonia also differ funda¬
mentally in their seeds and in the nature of the
floral bracts. The seeds of Tritonia are prismatic to

globose, smooth, and have the vascular trace ex¬
cluded (Goldblatt & Manning, 1995), and the
bracts of the genus are typically fairly short, more
or less membranous to scarious, and have bifurcate
or trifurcate apices. The capsules and seeds of G.
somalensis are unknown and cannot be used to as¬
sist in generic placement. The bracts, however, are
relatively large, soft-textured, and green, hence
quite typical of Gladiolus. The style branches are
linear and do not appear to be apically expanded
as is the case with most species of Gladiolus, but
this minor difference in style branch structure does
not seem particularly significant in relation to ge¬
neric placement.

The two collections from Cal Miskat both consist
of small plants up to 12 cm high with leaves 1-2
mm wide, whereas the type specimen is 30 cm high
and has leaves up to 4 mm wide. Sacco s.n. is ster¬
ile, but the inflorescence of Azzaroli 6 is unbran¬
ched and has only two flowers. The type, from Cal
Madow, has a branched inflorescence with 10 flow¬
ers on the main axis. Also, the flowers are some¬
what smaller in Azzaroli 6 than in the type. To prop¬
erly evaluate these differences further field studies
are needed. For the present we think these three
collections are best regarded as representing a sin¬
gle variable species.

The relationships of Gladiolus somalensis within
the genus are uncertain. It bears a fair resemblance
to the Ethiopian G. calcicola Goldblatt and to the
Eritrean G. mensensis Baker (both subgenus Ophio-
lyza), largely because both these species have rel¬
atively small flowers and are fairly small plants with
several well-developed foliage leaves (Goldblatt, in
press). It is more likely that the affinities of G. so¬
malensis lie with southern African species of sub¬
genus Gladiolus. Members of that subgenus usually
have short-tubed flowers of a size comparable to
those of G. somalensis. Most southern African spe¬
cies of subgenus Gladiolus have ellipsoid, apically
acute capsules, unlike the ovoid to oblong, apically
three-lobed capsules of G. calcicola and G. men¬
sensis, more characteristic of subgenus Ophiolyza.
Until more is known about G. somalensis, especially
the nature of its capsules and seeds, its affinities
remain speculative at best.

Although a relationship with southern African
species seems less likely because of the distance
involved, there are phytogeographical connections
between the Somalian region and arid parts of
southern Africa (de Winter, 1974; Thulin, 1994b).
In Iridaceae the distribution of Babiana is a strik¬
ing example. One species of this largely South Af¬
rican genus occurs on Socotra, an island off the
coast of Somalia (Lewis, 1959). Another example

328

Novon

involving a petaloid monocot concerns a new spe¬
cies of Trachyandra (Asphodelaceae) that grows in
the same area as Gladiolus somalensis, but in a
different habitat, crevices in shady limestone rocks
(Thulin, 1995). Trachyandra is otherwise a mainly
southern African genus with a few species extend¬
ing into tropical Africa.

Paratypes. SOMALIA. Bari Region: Wadi Hantara,
Candala. 6 Feb. 1956, Azzaroli 6 (FT); Azienda Uar Ma¬
han, Jan.-Feb. 1959, Sacco s.n. (FT).

Acknowledgments. Mats Thulin’s fieldwork in
Somalia was supported by SAREC (Swedish Agen¬
cy for Research Cooperation with Developing
Countries). We thank John Manning for the illus¬
tration reproduced here.

Literature Cited

Goldblatt, P. 1989. Systematics of Gladiolus (Iridaceae)
in Madagascar. Bull. Mus. Natl. Hist. Nat., S6r. 4, Sect.
B, Adansonia 11: 235-255.

-. 1991. An overview of the systematics, phylogeny

and biology of the southern African Iridaceae. Contrib.
Bolus Herb. 13: 1-74.

-. 1995. Iridaceae. Pp. 62-67 in M. Thulin (edi¬

tor), Flora of Somalia, volume 4. Royal Botanic Gar¬
dens, Kew. In press.

-. In press (1996). Gladiolus in Tropical Africa.

Timber Press, Portland, Oregon.

-& J. C. Manning. 1995. Phylogeny of the African

genera Anomatheca and Freesia (Iridaceae—Ixioideae),
and a new genus Xenoscapa. Syst. Bot. 20: 161-178.

Lewis, G. J. 1959. The genus Babiana. J. S. African Bot.,
Suppl. 3.

-. A. A. Obermeyer & T. T. Barnard. 1972. A re¬
vision of the South African species of Gladiolus. J. S.
African Bot., Suppl. 10: 1-316.

Thulin, M. 1994a. Cal Madow, Somalia. Pp. 194—195 in
S. D. Davis, V. H. Heywood & A. C. Hamilton (editors).
Centre of Plant Diversity, a Guide and Strategy for their
Conservation, Volume 1. Information Press, Oxford,
UK.

-. 1994b. Aspects of disjunct distributions and en¬
demism in the arid parts of the Horn of Africa, partic¬
ularly Somalia. Pp. 1105-1119 in J. H. Seyani & A. C.
Chikuni (editors). Proceedings of the Xlllth plenary
Meeting AETFAT, Malawi, Volume 2. National Herbar¬
ium and Botanic Gardens of Malawi, Zomba.

-. 1995. Trachyandra triquetra (Asphodelaceae), a

new species from the Horn of Africa. Nordic J. Bot. 16.
In press.

Winter, B. de. 1974. Floristic relationships between the
northern and southern arid areas of Africa. Mitt. Bot.
Staatssamml. Miinchen 10: 424-^137.

Some Taxonomic Changes in Syringa L. (Oleaceae), Including a
Revision of Series Pubescentes

P. S. Green

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom

M. C. Chang

Shanghai Museum of Natural History, 260 Yanan Road E., Shanghai,
People’s Republic of China

Abstract. A number of species of Syringa are
reviewed, and series Pubescentes is revised. At in¬
fraspecific rank, the following new combinations in
Syringa from Asia are proposed: S. komarowii
subsp. reflexa, S. oblata subsp. dilatata, S. reticu¬
lata subsp. amurensis, S. reticulata subsp. pekinen-
sis, and S. pubescens subsp. microphylla var. potan-
inii. Identification keys are presented.

The preparation of an account of Syringa for the
forthcoming English version of the Flora of China ,
combined, for the senior author, with the prepara¬
tion of an account for the European Garden Flora,
has led to a review of the genus and a reassessment
of the appropriate rank for some of the previously
recognized species or varieties. The following are
the taxonomic and nomenclatural consequences.

A. Syringa komarowii

Syringa komarowii Schneider subsp. reflexa
(Schneider) P. S. Green & M. C. Chang, stat.
nov. Basionym: Syringa reflexa Schneider, Re-
pert. Spec. Nov. Regni Veg. 9: 80. 1910. SYN-
TYPES: “C.-China. Hupeh: 8-9000'”, Henry
6819 (K); “W. Hupeh,” Wilson 2078 (K).

Although McKelvey (1928: 77) stated, “The re¬
lationship of Syringa komarowi to S. reflexa Schnei¬
der is exceedingly close, and it is possible that at
some future time S. komarowi may be classified as
an extreme form of the Hupeh [Hubei] plant,” it
was not until 1990 that the two were formally unit¬
ed at the varietal rank (Chang & Chen, 1990: 35).
However, while bearing the morphological differ¬
ences in mind, because S. komarowii and S. reflexa
occupy distinct geographical areas, the former in
southern Gansu, southern Shaanxi, Sichuan, and
northern Yunnan, the latter in western Hubei and
northeastern Sichuan, we believe that subspecies is

the appropriate infraspecific catagory. They may be
distinguished as follows:

la. Corolla lobes somewhat erect; inflorescence usu¬
ally ± compact .subsp. komarowii

lb. Corolla lobes spreading; inflorescence somewhat

pyramidal, often interrupted.subsp. reflexa

B. Syringa oblata

Syringa oblata Lindley subsp. dilatata (Nakai) P.
S. Green & M. C. Chang, stat. nov. Basionym:
Syringa dilatata Nakai, Bot. Mag. (Tokyo) 32:
128. 1918. SYNTYPES: Korea. “Pon-san,”
Kamibayashi s.n., “Pan-syu-non,” Uchiyama
s.n., “in summo montis Nonensan,” Nakai
2209, “Kosui,” Nakai 2602, “Su-hun,” Nakai
2610, and “Chokoku,” Kin-o-sho 268 (synty-
pes, TI not seen).

As well as exhibiting morphological distinctness,
Syringa dilatata and S. oblata have different dis¬
tributions, the former in Korea and northeastern
China, and the latter in northern China. The rank
of subspecies seems therefore to be the more ap¬
propriate one. They may be distinguished as fol¬
lows:

la. Leaves usually slightly broader than long, 2.5-7

X 3—8 cm, base truncate to usually slightly cor¬
date; corolla tube 6-11 mm long, corolla lobes
4—6 mm long.subsp. oblata

lb. Leaves usually slightly narrower than long, 3—7

X 2.5-6 cm, base truncate; corolla tube 11—14
mm long, corolla lobes 5—8 mm long.

.subsp. dilatata

C. Syringa reticulata

Syringa reticulata (Blume) Hara subsp. amuren¬
sis (Ruprecht) P. S. Green & M. C. Chang, stat.
nov. Basionym: Syringa amurensis Ruprecht,
Bull. Cl. Phys.-Math. Imp. Acad. Sci. St. P6-
tersb. 15: 371. 1857. TYPE: China/Russia.
Ussuri River area, Maack s.n. (syntypes, ?LE
not seen).

Novon 5: 329-333. 1995.

330

Novon

Syringa reticulata (Blume) Hara subsp. pekinen-
sis (Ruprecht) P. S. Green & M. C. Chang, stat.
nov. Basionym: Syringa pekinensis Ruprecht,
Bull. Cl. Phys.-Math. Imp. Acad. Sci. St. Pe-
tersb. 15: 371. 1857. TYPE: China. Hebei:
?Kirilov s.n. (holotype, ?LE not seen).

Once again, and for similar reasons, it is be¬
lieved that subspecific rank is the appropriate one
for these taxa. Syringa reticulata subsp. reticulata
is native to Japan, subspecies amurensis to north-
easternmost China, adjacent Russia, and Korea,
and subspecies pekinensis to northern China and
Mongolia. Although in most of the relevant litera¬
ture, the last has been treated at specific rank, it
is noteworthy that Maximowicz (1859: 194), only
two years after the plant was first described as a
species, reduced it to varietal rank. Furthermore,
Hemsley (in Forbes & Hemsley, 1889: 82) sank S.
pekinensis under S. amurensis and commented,
“The differences between the Mandshurian, Japa¬
nese, and Chinese specimens are slight.” They may
be keyed out as follows:

la. Leaves mostly longer than 7 cm, hairy, especially
on the midrib and main veins on the under sur¬
face; capsules blunt.subsp. reticulata

lb. Leaves mostly less than 7 cm long, glabrous
above.

2a. Petioles stoutish, 1—2 cm long; veinlets

slightly sunk; capsule apex blunt.

.subsp. amurensis

2b. Petioles slender, 1.5-3 cm long; veinlets not
sunk on the upper surface of the leaf; cap¬
sule apex acute .subsp. pekinensis

D. A Revision of Series Pubescentes

Chang and Chen (1990; see also Chang & Qui,
1992: 63-71) have proposed changes in rank for
some of the taxa in series Pubescentes (C. K.
Schneider) Lingelsheim, which, following the clas¬
sic monograph of McKelvey (1928), have previously
been treated as species by botanists and gardeners
alike. The taxa in this section are closely related,
and, for some, classification at the rank of species
does not seem justified. However, examination of
numerous collections of these plants has led to cer¬
tain modifications to the classification proposed by
Chang and Chen. These are set out below, following
an identification key to the revised series. The nu¬
merous synonyms cited by McKelvey (1928) are not
repeated here.

Key to Syringa Series Pubescentes

la. Leaves with lateral veins ± pinnate, the 2 lowest pairs of veins not closely adjacent.

2a. Leaves usually manifestly longer than broad, sometimes almost as broad as long; corolla lobes 2—4 mm
long.

3a. Upper (adaxial) surface of leaves glabrous or glabrescent, except sometimes for a slight pubescence
on midrib and main veins on the under surface, rarely pilose.

4a. Leaves usually 1.5-3.5(—7) cm long, I 2.51—1) cm broad.1. S. pubescens

5a. Young stems and inflorescence axes ± 4-angled; corolla tube 10-15 mm long; north-central

China .la, subsp. pubescens

5b. Young stems and inflorescence axes — terete; corolla tube 8—10 mm long; northwestern and

west-central China.Id. subsp. microphylla var. microphylla

4b. Leaves 5—11 (rarely less) cm long, 2.5-6 (rarely less) cm broad .lb. subsp. patula

3b. Upper surface of leaves finely, often scattered, pilose.

6a. Leaves (2.5—)3—5(—7) cm long, (1.5—)2—2.5(—3.5) cm broad, densely pilose to villous below.

7a. Young stems and inflorescence axes puberulent to densely pilose.

8a. Corolla tube 7—8 mm long; petioles 2—12 mm long .lc. subsp. julianae

8b. Corolla tube 8—10 mm long; petioles 2—5 mm long . . Id. subsp. microphylla var. polaninii

7b. Young stems and inflorescence axes ± densely villous.2. S. mairei

6b. Leaves 1.5-3 cm long, 0.6—1.8 cm broad, glabrous except for a pilose midrib and primary veins

below.3. 5. pinetorum

2b. Leaves as broad as long, 1.2—2.2 cm broad, or slightly broader than long, almost circular; corolla lobes

4-5 mm long.4 5. wa rdii

lb. Leaves with 2 pairs of veins ± palmately arranged at the lamina base, blade usually 1.5—3(—4) cm long, 1—

2.7(—3) cm broad.5 5. meyeri

1. Syringa pubescens Turczaninow, Bull. Soc.
Nat. Moscou 13: 73. 1840. TYPE: China. He¬
bei: 1831, Kirilov s.n.? (holotype, LE, photo¬
graph, K).

Erect or spreading shrubs to 5 m tall, young

stems glabrous or pubescent. Leaves narrowly to
broadly ovate, elliptic or broadly elliptic, 1.5-9(—
11) cm long, l-5(-6) cm broad, glabrous to densely
villous, 3 primary veins not closely adjacent at the
lamina base. Inflorescence dense or open, glabrous
to pilose, 3—15 cm long. Corolla tube 4—15 mm

Volume 5, Number 4
1995

Green & Chang
Syringa

331

long, lobes 2—4 mm long. Capsule slender or stou-
tish, 0.7—1.7 cm long, smooth or with a few lenti-
cels.

la. Syringa pubescens subsp. pubescens

Erect shrubs to 5 m tall, young stems glabrous.
Leaves ovate to usually broadly ovate, sometimes
elliptic, (1.5-)2.5-3.5(-7) cm long, (l-)1.7-2.5(-4)
cm broad, apex acute to obtuse, very slightly acu¬
minate, glabrous above, glabrous to pilose below,
especially on the midrib and primary veins toward
their bases. Inflorescence ± dense, 4—8(-10) cm
long, axes glabrous to pilose. Corolla tube 10-15
mm long, lobes 2-4 mm long, purplish lilac, paler
within. Capsule stoutish, 9-10 mm long, smooth
with a few lenticels.

This subspecies is recorded (Chang in Chang &
Qui, 1992) from the following provinces of China,
all in the northern part of the country: Hebei, Hen¬
an, eastern Shaanxi, western Shandong, and
Shanxi.

lb. Syringa pubescens subsp. patula (Palibin)
M. C. Chang & X. L. Chen, Invest. Stud. Nat.
10: 34. 1990. Ligustrum patulum Palibin, Acta
Hurt. Petrop. 18: 146 (Conspec. FI. Korea 2:
10), 1900. Syringa patula (Palibin) Nakai, Bot.
Mag. (Tokyo) 40: 148. 1926. TYPE: Korea,
Sontag s.n. (holotype, ?LE not seen).

Syringa velutina Komarov, Acta Hort. Petrop. 18: 428.
1901. TYPE: Korea, Komarov s.n. (syntype, ?LE not
seen).

Syringa palibiniana Nakai, Bot. Mag. (Tokyo) 27: 32.
1913. TYPE: Korea, Faurie s.n. (holotype, TI not
seen).

Erect shrubs to 3 m tall, young stems slightly
pubescent to glabrous. Leaves ovate or broadly
ovate to usually elliptic or broadly elliptic, (3—)5—
9(—11) cm long, (2-)2.5—5(-6) cm broad, apex
slightly acuminate, glabrous above, rarely scat¬
tered short pilose, pilose to usually glabrous below
except for the midrib and primary veins toward
their bases. Inflorescence ± dense, 5-9(—15) cm
long, pilose. Corolla tube 7-8(-10) mm long, lobes
2(—3) mm long, vinaceous lilac, white within. Cap¬
sules slender, curved, 1—1.5 cm long, with a few
lenticels.

Recorded from northeastemmost China (Jilin
and Liaoning) and Korea. This is a hardy plant
that has proved its worth as a garden plant under
rigorous climatic conditions. In the West it is per¬
haps best known in cultivation in the form of cv.
‘Miss Kim.’

lc. Syringa pubescens subsp. julianae (C. K.
Schneider) M. C. Chang & X. L. Chen, Invest.
Stud. Nat. 10: 34. 1990. Syringa julianae C.
K. Schneider, Ill. Handb. Laubholzk. 2: 777.
1912 & Bull. Misc. Inform. Kew 1912: 37.
1912. TYPE: cultivated, seed from China,
Hubei, Wilson 1220A (holotype, ?W not seen;
isotype, K).

Spreading shrubs to 2 m tall, young stems pu-
berulent. Leaves narrowly ovate to elliptic, (2.5—)4—
5(-7) cm long, (1.5—)2.3—2.5(-3) cm broad, apex
acute, slightly long-acuminate, scattered pilose
above, pilose below, densely so on midrib and main
veins; petioles 2-12 mm long. Inflorescence ±
open, (3-)4—6(—10) cm long, densely pilose. Corolla
tube 7—8 mm long, lobes 2-3 mm long, violet-pur¬
ple, paler within. Capsules slender, 0.7—1 cm long,
smooth.

This subspecies has been recorded only from the
province of Hubei.

l d. Syringa pubescens subsp. niicrophylla

(Diels) M. C. Chang & X. L. Chen, Invest.
Stud. Nat. 10: 34. 1990. Syringa microphylla
Diels, Bot. Jahrb. Syst. 29: 531. 1900. SYN-
TYPES: China. Shaanxi: Giraldi 1644 & 1645
(B destroyed).

i. Syringa pubescens [subsp. niicrophylla] var.
microphylla

Spreading shrubs to 2 m tall, young stems finely
puberulent to rarely glabrous. Leaves ovate to el¬
liptic. narrowly to broadly so, apex obtuse to acute,
very slightly acuminate, glabrous above, rarely
scattered pilose, glabrous below, except the midrib
and primary veins toward their bases, rarely pilose.
Inflorescence ± dense, 4—10f—12) cm long, pilose.
Corolla tube 8-10 mm long, lobes 2-3 mm long,
pinkish lilac, slightly paler within. Capsules ±
slender, 1.2-1.5 cm long, smooth with a few lenti¬
cels.

Recorded (see Chang in Chang & Qui, 1992: 67)
from the Chinese provinces of Gansu, Hebei, Hen¬
an, Hubei, Ningxia, Qinghai, Shaanxi, Shanxi, and
Sichuan. Variety flavoanthera (X. L. Chen) M. C.
Chang has been recognized, but the taxonomic val¬
ue of anther color is uncertain, and the recording
of it can be suspect, seeing that the pollen of pur¬
plish anthers can make them appear yellow after
dehiscence.

332

Novon

ii. Syringa pubescens [subsp. microphylla] var.
potaninii (C. K. Schneider) P. S. Green & M.
C. Chang, comb, et stat. nov. Basionym: Syrin¬
ga potaninii C. K. Schneider, Repert. Spec.
Nov. Regni Veg. 9: 80. 1910. TYPE: “Nord-
China, O-kansu, am Flusse Tschi lo ku im Ge-
birge,” 1885, Potanin s.n. (holotype, ?LE not
seen; isotype, K).

More or less erect shrubs to 4 m tall, young
stems finely puberulent. Leaves ovate to elliptic,
(2.5—)3—5(—6) cm long, (1.5—)2—2.5(—3.5) cm
broad, apex acute, somewhat acuminate, puberu¬
lent to glabrous above, rarely scattered pilose,
scattered to densely pilose below, especially on
the midrib and main veins toward the base; peti¬
oles 2-5 mm long. Inflorescence ± dense, 6-10
cm long, finely puberulent. Corolla tube 8-10 mm
long, lobes 2—3.5 mm long, pinkish lilac, paler
within. Capsules slender, 1.5—1.7 cm long, smooth
with a few lenticels.

Known only from the Chinese province of Gansu.

2. Syringa mairei (H. Lcveille) Rehder, J. Arnold

Arbor. 15: 302. 1934. Ligustrum mairei H.
L6veill6, Cat. PI. Yun-Nan 181. 1916. TYPE:
China. Yunnan: Maire s.n. (hclotype, E, pho¬
tograph, K).

Syringa rugulosa McKelvey, J. Arnold Arbor. 6: 152.
1925. TYPE: China. Yunnan: Maire “169/1914” (ho¬
lotype, E, photograph, K).

Shrubs to 4 m tall, young stems densely villous.
Leaves ovate to elliptic, somewhat broadly so, (2—)
4-^L5(-6) cm long, (1.5—)2—2.5(—3) cm broad, apex
acuminate, pilose above, densely pilose below. In¬
florescence dense, 7—12 cm long, villous. Corolla
tube 7—9 mm long, lobes 2—3 mm long, “rose-vio¬
let.” Capsule stoutish, 1 cm long, smooth with a
few lenticels.

This is a little known plant from Yunnan. The
two names, Syringa mairei and S. rugulosa, were
almost certainly based on duplicates of the same
collection made by E. E. Maire.

3. Syringa pinetorum W. W. Smith, Notes Roy.

Bot. Card. Edinburgh 9: 132. 1916. TYPE:
China. Yunnan: Forrest 12472 (holotype, E,
photograph, K).

Slender shrubs to 2.5 m tall, young stems pi¬
lose. Leaves narrowly ovate to elliptic, 1.5-3 cm
long, 0.6-1.8 cm broad, apex acute, acuminate,
scattered pilose above, glabrous below except for
the midrib and primary veins toward their bases.

Inflorescence ± dense, 8-12 cm long, pilose. Co¬
rolla tube 9—10 mm long, lobes 3 mm long, “pale
lavender-rose.” Capsule with inconspicuous len¬
ticels.

A little known plant from Tibet and the province
of Yunnan. Plants that have been cultivated under
this name have been misidentified. They have fre¬
quently been found to be S. yunnanensis Franchet.

4. Syringa wardii W. W. Smith, Notes Roy. Bot.

Card. Edinburgh 9: 132. 1916. TYPE: China.
Yunnan: Kingdon Ward 312 (holotype, E, pho¬
tograph, K).

Shrubs to 5 m tall, young stems pilose. Leaves
± circular, about 1.2-2.2 cm long and broad, apex
rounded, extremely shortly acuminate, glabrous
above and below, 3 primary veins on each side, not
adjacent at the base of the lamina. Inflorescence
somewhat dense, 6-9 cm long, pubescent. Corolla
tube 13—14 mm long, lobes 4—5 mm long, pale
pinkish lilac? Capsule with inconspicuous lenti¬
cels.

This species is only known from the type collec¬
tion and was previously treated as a synonym of
Syringa pinetorum by Chang (in Chang & Qui,
1992: 71). However, the small, more or less circular
leaves and long corolla tube mark it out as distinct.

5. Syringa meyeri C. K. Schneider in Sargent,

PI. Wilson. 1: 301. 1912. TYPE: cultivated at
the Arnold Arboretum, origin from cultivation
in China, Hebei, Meyer, under Dept, of Agri¬
culture number 23032 (holotype. A).

Compact shrub to 1.5 m tall, young stems finely
puberulent. Leaves ovate to very broadly ovate-el¬
liptic, (1.5-)2-3(-4) cm long, (l-)1.8-2.7(-3) cm
broad, apex obtuse, rarely somewhat acute, very
slightly and shortly acuminate, glabrous above, gla¬
brous below except the midrib and the primary
veins toward their bases, 2 (or 3) primary veins on
each side, subpalmate, arising within 2 mm of the
base of the lamina. Inflorescence dense, 5—11 cm
long, puberulent. Corolla tube 9-12 mm long, lobes
2-3 mm long, purplish lilac, paler within. Capsules
slender, 1-1.2 cm long, lenticular.

This compact, relatively dwarf species was first
described from cultivated material. It is widely
grown in northern China, and in the West under
the name ‘Palibin’ (see Green, 1979). It was
known only as a cultivated plant until discovered
in the wild by Chang as recently as 1989 in the
province of Liaoning. This plant has been de¬
scribed as:

Volume 5, Number 4
1995

Green & Chang
Syringa

333

Syringa meyeri var. spontanea M. C. Chang, Invest.
Stud. Nat. 10: 33. 1990. TYPE: China. Liaon¬
ing: M. C. Chang & X. K. Qin 2872 (holotype,
SHM; isotype, K).

The variety is distinguished by its looser inflo¬
rescence, somewhat shorter corolla tube, and small¬
er leaf.

Literature Cited

Chang, M. C. & X. L. Chen. 1990. Studies on Chinese
Syringa (1). Invest. Stud. Nat. 10: 32—U). [Chinese with
English summary.]

-& L. Q. Qui (Editors). 1992. Oleaceae. FI. Rei-

publ. Popularis Sin. 61: 2—222.

Forbes, F. B. & W. B. Hemsley. 1889. Index florae si¬
nensis 2. J. Linn. Soc., Bot. 26: 1—561.

Green, P. S. 1979. Syringa meyeri Palibin. Bot. Mag. 182:
t. NS. 778.

McKelvey, S. D. 1928. The Lilac, a Monograph. London.

Maximowicz, C. J. 1859. Primitiae florae amurensis.
Mdm. Acad. Imp. Sci. St. Petersburg Divers Savants 9.

Homalia pennatula (Musci: Neckeraceae), a New Combination
from Southeast Asia, with a Key to the Species of Homalia

Si He

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Johannes Enroth

Botanical Museum, P.0. Box 47, FIN-00014 University of Helsinki, Finland

Abstract. Symphyodon pennatulus (Dixon) Dix¬
on from Southeast Asia is transferred to the genus
Homalia of the Neckeraceae (Musci). A key to the
five species and one variety of Homalia is provided.

Homalia (Bridel) Schimper is primarily distrib¬
uted in the northern temperate regions and nearby
subtropical borders, where it grows on boulders,
rock outcrops, or bases of trees. It is characterized
by strongly complanate plants with leaves arranged
in distinct ranks; by oblong-ovate to spatulate
leaves with broad apices and smooth leaf cells; and
by smooth capsules with a “hypnoid” peristome.
Symphyodon Montagne has 14 species worldwide
and is primarily distributed in the tropical and sub¬
tropical regions of Southeast Asia, with one species
in Hawaii, South Africa, and Central and South
America, respectively. In Southeast Asia, it grows
exclusively on tree trunks and tree branches. Be¬
sides its habitat distinction, taxonomic characters
of generic importance for Symphyodon are found in
the capsule ornamentation and peristomes. It has
echinate capsules and an “isobryoid” peristome.
Other secondary gametophyte features include:
leaves often appressed and imbricate or spreading;
leaf apices narrowed; and leaf cells prorate.

Symphyodon pennatulus (Dixon) Dixon was ex¬
cluded from the genus Symphyodon, because it is
epilithic and has strongly complanate to sub-disti¬
chous leaves (He & Snider, 1992). The type spec¬
imen of Symphyodon pennatulus is meager and con¬
tains no sporophytes. Since no other specimens of
this species were available. He & Snider (1992)
were uncertain of its generic placement. Recently,
numerous collections of this moss have come from
Thailand, and though they consistently lack spo¬
rophytes, we now have enough material to make
detailed observations on the gametophytes. Sym¬
phyodon pennatulus exhibits many aspects of Hom¬
alia, such as a strongly complanate habit, a hom-
alioid-type leaf insertion showing an eight-ranked
phyllotaxy (see He, 1992), oblong-ovate to spatulate

Novon 5: 334-335. 1995.

leaves with broad and rounded apices, and a vari¬
able costal morphology. It is similar to the widely
distributed Homalia trichomanoides (Bridel)
Schimper in overall similarities of leaf shape and
the absence of pseudoparaphyllia, but it differs in
having distinct double costae and linear, projecting
median leaf cells. On the basis of the double leaf
costae and elongate median leaf cells, Symphyodon
pennatulus appears most closely related to Homalia
glabella (Hedwig) Schimper from Central and South
America, except that it lacks pseudoparaphyllia.
Although the narrowly linear and projecting median
leaf cells of Symphyodon pennatulus and its pres¬
ence in southern Asia differ from those of previ¬
ously recognized Homalia species, the overall ga¬
metophyte features of this moss support its
placement in Homalia. Hence, we are making the
following new combination. A key to the five spe¬
cies and one variety of Homalia now recognized is
provided. A description and an illustration of the
species treated here will be published in the forth¬
coming revision of Homalia by the first author.

Homalia pennatula (Dixon) He & Enroth, comb,
nov. Basionym: Stereodon pennatulus Mitten ex
Dixon, Rec. Bot. Surv. India 6: 66. 1914. Sym¬
phyodon pennatulus (Dixon) Dixon in Broth-
erus, Nat. Pflanzenfam., ed. 2, 11: 267. 1925.
TYPE: Myanmar (Burma). Moulmein, Parish
s.n. (holotype, NY; isotype, BM).

In his book entitled Burma, Its People and Pro¬
ductions, Mason (1883) published a list of plants,
including mosses. Homalia erosa Hampe and Hom¬
alia pennatula Mitten were listed in this publica¬
tion. Both names are apparently nomina nuda,
since no descriptions or basionyms were given. The
former is clearly a herbarium name, because we
have seen several herbarium specimens annotated
with the name Homalia erosa Hampe. However, we
have seen no specimens bearing the name Homalia
pennatula Mitten, nor has it ever been published

Volume 5, Number 4
1995

He & Enroth

Homalia pennatula from Southeast Asia

335

validly. Although it could be based on Mitten’s her¬
barium name Stereodon pennatulus, Homalia pen¬
natula Mitten has no nomenclatural bearing in our
paper.

In recognizing that Homalia trichomanoides var.
japonica (Besch.) He, stat. nov., cited in He (1992)
has not been validly published, we make the fol¬
lowing new combination:

Homalia trichomanoides var. japonica (Bes-
cherelle) He, comb. nov. Basionym: Homalia
japonica Bescherelle, J. Bot. (Morot) 13: 39.
1899. TYPE: Japan. Sengantoge, Faurie
14984 (holotype, PC; isotype, KYO).

Additional selected specimens examined. MYANMAR.
Moulmein, Kurz 2930 (BM). INDIA. Kerala, Norhett
13377 (BM). THAILAND. Changburi: Larsen et al. 1857
(AAU, MO). Chiangmai: Mt. Suthep, 98°55'E, 18°50'N,
Touw 8395 (BM, L, MO, NY). Krabi: Khao Phanom Ben-
cha Park, Schafer-Verwimp & Verwimp 16230 (H, MO).
Nakhon Nayok: Larsen et al. 3380 (AAU, MO). Nakhon
Sawan: 98°55'E, 16°40'N, Touw 8319 (BM, L, MO, NY).
Prachinburi: 101°30'E, 14°30'N, Touw 12215 (BM, L,
MO). Trat: 102°49'E, 11°59'N, Charoenphol et al. 4982
(AAU, BM, H, MO, NY); Ko Kut, 101°31'E, 12°35'N,
Charoenphol et al. 5129 (AAU, BM, H, MO, NY).

Key to the Species and Varieties of Homalia

1. Costa single, extending Vi—Y 5 the leaf length ... 2

1. Costa double or forked, less than Vs the leaf

length or indistinct. 4

2(1). Pseudoparaphyllia present; costa stout, extend¬
ing 3 A~y s the leaf length; leaf apices dentate,
with large teeth; western Europe, northern Af¬
rica, Canary Islands ... 1. H. lusitanica Schimper

2. Pseudoparaphyllia absent; costa slender, ex¬

tending Vi-V a the leaf length; leaf apices mod¬
erately serrate, with small teeth. 3

3(2). Autoicous; secondary stems often blunt near
the apex; flagelliform branches uncommon;
eastern Asia, Russia, Europe, North America,
Mexico.

.2a. H. trichomanoides var. trichomanoides

(Hedwig) Schimper

3. Dioicous; secondary stems often attenuate near

the apex; flagelliform branches commonly pres¬
ent; Japan, Korea, Taiwan .

.2b. H. trichomanoides var. japonica

(Bescherelle) He

4(1). Median leaf cells rhombic, 8.5-17.5 pm long;
Azores, Maderia, Canary Islands, Caucasus . .

.3. H. webbiana (Montagne) Schimper

4. Median leaf cells oblong-rhomboidal to narrow¬
ly linear, 35-100 pm long. 5

5(4). Leaves 1.25-3.25 mm X 0.75-1.25 mm; pseu¬
doparaphyllia present; Mexico, West Indies,
Central America, Brazil, Venezuela .

.4. H. glabella (Hedwig) Schimper

5. Leaves 0.8-1.2 mm X 0.5-0.65 mm; pseudo¬
paraphyllia absent; India, Myanmar, Thailand

.5. H. pennatula (Dixon) He & Enroth

Acknowledgments. We thank Bruce Allen (MO)

and Marshall R. Crosby (MO) for constructive com¬
ments on the manuscript.

Literature Cited

He, Si. 1992. A worldwide taxonomic revision of the gen¬
era Homalia, Pendulothecium (Musci: Neckeraceae)
and Symphyodon (Musci: Symphyodontaceae). Doctoral
Dissertation, University Microfilms, Ann Arbor.

-& J. A. Snider. 1992. A preliminary survey of

the moss genus Symphyodon (Symphyodontaceae, Mus¬
ci). Bryobrothera 1: 283—287.

Mason, F. 1883. Burma, Its People and Productions. Vol.
II (Botany). Stephen Austin & Sons, Hertford.

Polygonum hickmanii (Polygonaceae), a New Species from
California

Harold Hinds

Biology Department, University of New Brunswick, Fredericton,

New Brunswick, E3B 6E1 Canada

Randall Morgan

3500 N. Main Street, Soquel, California 95073, U.S.A.

Abstract. A new endemic species of Polygonum
is described from Scotts Valley, California. Differ¬
entiation from closely related taxa P. parryi and P.
heterosepnlum is presented.

Polygonum (sect. Duravia) hickmanii H. R.
Hinds & R. Morgan, sp. nov. TYPE: U.S.A.
California: Santa Cruz County, Scotts Valley,
grassland N of Navarra Drive, W of Carbonero
Creek, 1 July 1993, R. Morgan 2119 (holo-
type, CAS; isotypes, MO, RSA, UC, UNB).
Hickman’s knotweed.

Herba annua. Folia sessilia, 0.5-3.5 cm X 1—1.5 mm,
revoluta. Ocrea divisa ad basem in longas rectas libras.
Flores singuli in axillis omnium paene bracteorum folior-
um. Achenia laevia, (2—)2.3 X 1.6 mm, duobus lateribus
concavis et uno latere latiore planioreque.

Erect taprooted annual, 2-5 cm, simple in de¬
pauperate plants to profusely branched from near
the base. Leaves sessile, linear, acuminate, mucro-
nate, 0.5—3.5 cm X 1—1.5 mm, revolute and wing-
nerved below, closely investing and with the ocreae
± concealing the stems. Ocrea 6 mm long, silvery,
divided nearly to base into long straight filaments,
adnate to the base of the leaves and appearing al¬
most as long as the apical leaves. Flowers single in
axils of almost all bracteal leaves, tepals white, with
two outer tepals cucullate and mucronate, 3 mm
long, three inner tepals rounded to obtuse 2 mm
long, anthers conspicuous, orange-pink 0.4 mm.
Achenes smooth, (2-)2.3 X 1.6 mm, light to dark
olive brown, lighter on the angles, with two concave
sides and one broader ± flat side; not exserted.
Styles 0.3 mm, persistent. Flowering from late May
to August (October).

Polygonum hickmanii is known from two sites
about one mile apart, at the northern end of Scotts
Valley, Santa Cruz County, California, at about
700—800 feet elevation and occurs only in very re¬
stricted microhabitats within an isolated relictual
grassland. The sites are gently sloping to nearly
level, with fine-textured shallow soil over outcrops
of Santa Cruz mudstone and Purisima sandstone,
sparsely vegetated with small annual herbs. This
habitat is shared with a more abundant but equally
narrow endemic, Chorizanthe robusta C. Parry var.
hartwegii (Bentham) J. Reveal & R. Morgan. The
population of Polygonum hickmanii fluctuated from
about 200 to 400 plants in 1990-1992 to 1000 or
more in 1993 following a relatively wet winter.

Polygonum hickmanii is sympatric with no other
members of the section Duravia. The nearest lo¬
cation of a closely related species, P. parryi Greene
(itself disjunct), is at Mt. Hamilton about 50 miles
inland.

Polygonum hickmanii differs from P. parryi in its
larger white flowers, longer leaves, larger anthers
and achenes, and longer, straight ocreae. It differs
from P. heterosepalum M. E. Peck & Ownbey in
achene shape (more rounded at base as opposed to
more tapering), size (1.5-2 mm), and flower struc¬
ture (perianth segments very unequal).

Polygonum hickmanii is named for James C.
Hickman (1941-1993), distinguished botanist, who
confirmed the uniqueness of this taxon and whose
advice and inspiration is here acknowledged.

Acknowledgments. The authors thank George
Flanders, Irene Johnston, and John Geyssen for
their considerable assistance with manuscript prep¬
aration.

Novon 5: 336. 1995.

Una Nueva Especie de Marsdenia (Asclepiadaceae) de Mexico

Veronica Judrez-Jaimes

Herbario National, Instituto de Biologfa, U.N.A.M., Apartado Postal 70-233,
Delegation Coyoacan 04510, Mexico D.F., Mexico

W. Douglas Stevens

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Resumen. Marsdenia rzedouiskiana es descrita
como nueva. Esta especie ha sido confundida con M.
macrophylla (Humboldt & Bonpland ex Schultes)
E. Fournier pero esta mds relacionada con M. prin-
glei S. Watson. La distribution de M. rzedowskiana
esta restringida al noreste de Puebla y partes ad-
yacentes de Veracruz, hacia el sur de la distribu-
ci6n conocida de M. pringlei y en altitudes super-
iores a las de M. macrophylla.

Abstract. Marsdenia rzedowskiana is described.
This new species has been confused with M. ma¬
crophylla, but is most closely related to M. pringlei.
Marsdenia rzedowskiana seems to be restricted to
northeastern Puebla and adjacent Veracruz, to the
south of the known distribution of M. pringlei and
at higher elevations than M. macrophylla.

El genero Marsdenia R. Brown se encuentra bien
representado en Mexico por 24—26 especies que se
distribuyen en todo el pais. Durante la revision
para el tratamiento del genero para Mexico y Cen-
troamerica. se encontro material que inicialmente
se incluyo en la section Ruehssia, subsection Ma-
crophyllae (Rothe, 1915); sin embargo, pertenecia
mas bien a la section Rostrata, subsection Edules,
y adernas correspondfa a una especie no descrita.

Marsdenia rzedowskiana Judrez-Jaimes & W. D.
Stevens, sp. nov. TIPO: Mexico. Puebla: Mun-
icipio Teziutlan, Agua de Obispo, 30 mar.
1973 (fl), F. Ventura 8102 (holdtipo, MEXU;
isotipos, ENCB, MO). Figura 1.

Marsdeniae pringlei S. Watson similis sed floribus mi-
noribus, apice styli perbreviore et integro recedit.

Plantas trepadoras, con ldtex bianco. Tallos gla-
bros, gruesos, 5—7 mm de ancho. Pecfolos 1—3 cm,
glabrescentes; ldminas 5.0-15.5 X 2.6-10.5 cm,
elfpticas, con 7—8 nervios secundarios, base redon-
deada a obtusa, apice acuminado a cortamente ac-
uminado, coridceas, glabras. Inflorescencias 3—6 X
2-4 cm, con 10-22 flores, extra-axilares, laxamente

ramificadas; brdcteas elfpticas y acuminadas, 2 mm
de largo, pubescentes; pedunculos 1.0-2.5 cm de
largo, ligeramente pubescentes; pedicelos 2—4 mm
de largo, pubescentes. Ldbulos del cdliz 2.6-3.0 X
2.2-2.8 mm, orbiculares, dpice algo agudo, ligera¬
mente pubescentes por fuera, glabros por dentro,
mdrgenes ciliados, los senos con 1—3 gldndulas por
dentro; corola campanulada, 5.0-6.5 X 4.8-6.6
mm, glabra por fuera, ldbulos 3.0-3.2 X 1.8-2.3
mm, oblongos, erectos en la antesis, la superficie
interna pilosa en el centra, apices ligeramente
emarginados, tubo 2.5-3.0 mm de largo, piloso en
la garganta, los pelos largos y blancos; ldbulos de
la corona 1.5—1.7 X 1.0-1.5 mm, camosos, redon-
damente deltoides, con los mdrgenes laterales sal-
ientes, la punta libre y terminando al mismo nivel
de la parte apical de las anteras, ginostegio 2—3
mm de largo, membranas de las anteras 0.9-1.2
mm, ampliamente ovadas, anteras 0.4—0.5 mm de
largo, elipsoides; corptisculos 0.3-0.4 mm de largo,
elfpticos, polinios 0.3-0.4 mm de largo, casi glo-
bosos, apice del estilo 2.0-2.5 mm de largo, cdni-
co-alargado, con una pequena Ifnea de bifurcacion
en la parte apical; carpelos glabros. Folfculos des-
conocidos.

Marsdenia rzedowskiana estd muy relacionada
con M. pringlei S. Watson, puesto que ambas tienen
hojas ampliamente elfpticas, cimas ramificadas,
flores amarillas, garganta de la corola con pelos
blancos, ldbulos de la corona deltoides, dpiee del
estilo alargado y polinios casi globosos. Sin embar¬
go, M. rzedowskiana se diferencia por poseer flores
mds pequenas, ldbulos de la corola mds o menos
del mismo tamano que el tubo, tubo de la corola
no urceolado, apice del estilo 3 veces mds corto y
presentando apenas una Ifnea de bifurcacidn en la
parte apical. Por otra parte, M. rzedowskiana solo
es conocida de la Sierra Norte de Puebla y la regidn
adyacente a Jalapa, Veracruz, en lo que Rzedowski
(1978) considera bosques mesofilos de montana,
mientras que M. pringlei se distribuye en un drea
mds extensa de la Sierra Madre Oriental.

Novon 5: 337-339. 1995.

338

Novon

El material de Marsdenia rzedmvskiana tradi-
cionalmente ha sido determinado como M. macro-
phylla principalmente por su aspecto general, pero
la presencia en M. rzedowskiana de una inAores-

cencia muy ramificada de Acres amarillas, lobules
de la corona deltoides y dpice del estilo cdnico-
alargado la separa definitivamente de esta; ademas,
M. rzedowskiana crece en bosques superiores a los

Volume 5, Number 4
1995

Juarez-Jaimes & Stevens
Marsdenia rzedowskiana

339

1300 m, mientras que M. macrophylla es de tierras
bajas en Mexico.

Marsdenia rzedowskiana Judrez-Jaimes & W. D.
Stevens se dedica a Jerzy Rzedowski por sus im-
portantes aportes al estudio de la vegetacion y la
botariica de Mexico, trabajos por los cuales los au-
tores tienen una gran admiracion.

Pardtipos. MEXICO. Veracruz: Mpio. Aratlan. El Ce-
rro, 3 jun. 1977 (fl), F. Ventura 14030 (XAL); Mpio. Ye-
cuatla, along winding road from Naolinco to Misantla, just
above Santa Rita and 3 km by road from Paz de Enriquez,
1550 m (fr), M. Nee et al. 26425 (MO). Puebla: Mpio.
Hueytamalco, El Popual, 1500 m, F. Ventura 1038 (US).

Agradecimientos. Agradecemos a Hector Her¬
nandez, Francisco Lorea, Fernando Chiang y Agus-
tln Rudas la revision crftica del manuscrito. Henk
van der Werff escrihio la diagnosis en latln; John
Myers realizo la ilustracion de la planta.

Literatura Citada

Rothe, W. 1915. Uber die Gattung Marsdenia R. Br. und
die Stammpflanze der Condurangorinde. Bot. Jahrb.
Syst. 52: 354-434.

Rzedowski, 1. 1978. Vegetaci6n de Mexico. Limusa, Max-

New Taxa of Corispermum L. (Chenopodiaceae), with Preliminary
Comments on the Taxonomy of the Genus in North America

Sergei L. Mosyakin

N. G. Kholodny Institute of Botany, 2 Tereshchenkivska Str., Kiev, 252601 Ukraine

ABSTRACT. Five new species and three varieties
of Corispermum L. (Chenopodiaceae) are described
from North America. Corispermum pacificum Mo¬
syakin and C. pallidum Mosyakin are restricted to
the Pacific Northwest; C. welshii Mosyakin is
known from Utah and adjacent parts of Wyoming;
C. navicula Mosyakin occurs in Colorado and prob¬
ably adjacent states; C. hookeri Mosyakin is known
from Canada. Corispermum pacificum is closely re¬
lated to the Asian taxa C. crassifolium Turczaninov
and C. maynense Ignatov; C. pallidum is possibly
related to C. macrocarpum Bunge ex Maximowicz;
the three remaining new taxa and all other native
North American taxa are allied to the C. pallasii
Steven aggregate. The nomenclature of C. pallasii
[— C. leptopterum (Ascherson) Iljin = C. sibiricum
Iljin subsp. baicalense Iljin = C. bjelorussicum Klo¬
kov & Krasnova], C. villosum Rydberg, and C.
americanum (Nuttall) Nuttall is discussed. Three
new varieties are described: C. americanum var.
rydbergii Mosyakin, C. hookeri var. pseudodeclina-
tum Mosyakin, and C. ochotense var. alaskanum
Mosyakin. Corispermum ochotense Ignatov, a spe¬
cies recently described from the northeastern part
of the Russian Far East, is reported from Alaska
and northwestern Canada, where it is represented
by two varieties. Corispermum ochotense var. alas¬
kanum differs from the typical variety in having
larger fruits. Preliminary results of a study of the
genus in North America are discussed.

Corispermum has the reputation of being a prob¬
lem for taxonomists and may be regarded as one of
the most taxonomically complicated groups within
Chenopodiaceae. As an example, the name Coris¬
permum hyssopifolium L., which refers to a species
restricted mostly to a small area in southeastern
Europe and adjacent parts of western Asia, has
been misapplied to almost all other species of the
genus. For a long time the representatives of this
genus were believed to have been introduced to
North America from Eurasia. In the treatment by
Maihle and Blackwell (1978) only three species
were reported for North America: C. hyssopifolium
L., C. nitidum Kitaibel ex Schultes, and C. orientale

Lamarck (all Eurasian). However, it is now clear
from the fossil evidence that Corispermum was
present in North America at least 38,000 years be¬
fore the present (for details see Betancourt et ah,
1984). Native American species of Corispermum
are evidently closely related to eastern Asian (es¬
pecially Siberian) ones (see discussion below).

I strongly believe that even less than 10,000
years was sufficient for Asian ancestors of North
American taxa to evolve into several distinct races
(species or subspecies) because the high rate of
microevolutionary processes is crucial for a suc¬
cessful life strategy of such annual psammophytes
of pioneer and marginal habitats. The important mi¬
croevolutionary patterns within the genus include
both allogamy (performed by wind pollination) and
autogamy (especially under unfavorable environ¬
mental conditions), high degree of morphological
variability (including so-called “cryptic variabil¬
ity”), and high possibility of hybridization (es¬
pecially between closely related species), which is
typical for many other chenopods, including Che-
nopodium L. However, polyploids, which are fairly
common in Chenopodium and many other genera of
the family, seem to be absent, or at least extremely
rare, in Corispermum: all taxa studied until present
were diploids with 2 n = 18 (for additional discus¬
sion, see Adamkiewicz, 1970; Strazdin’sh, 1985;
Mosyakin, 1988a).

In the course of preparation of a taxonomic treat¬
ment of Corispermum for the Flora of North Amer¬
ica, it has become evident that several new species
and varieties should be described. Before describ¬
ing these taxa I will briefly discuss the taxonomy
of other Corispermum species from North America.
However, I should mention that the taxonomic
treatment of the genus in North America is not yet
complete, and more efforts, especially study of vari¬
ability in the natural habitats, are needed. At pres¬
ent my experience in North American Corispermum
is based exclusively on herbarium studies, plus
some experience with Eurasian species, which was
especially crucial for this work, and four weeks of
fieldwork in the southwestern United States (Utah
and Nevada). Because of this, some of the following

Novon 5; 340-353. 1995.

Volume 5, Number 4
1995

Mosyakin

Corispermum from North America

341

considerations (especially concerning the C. pal-
lasii Steven— C. americaniirn (Nuttall) Nuttall aggre¬
gate) should be regarded as preliminary, a kind of
brief introduction to the taxonomic problems in
North American Corispermum. A history of taxo¬
nomic study of the genus in North America, as well
as literature citations, may be found in Maihle and
Blackwell (1978); thus, some of the dates and ref¬
erences are not mentioned below in order to avoid
repetition.

Discussion

The first taxon described from North America
was C. hyssopifolium L. var. americanum Nuttall [=
C. americanum (Nuttall) Nuttall], Unfortunately, its
authentic description is insufficient, and the lec-
totype specimen (Nuttall s.n., PH; labeled “C. hys¬
sopifolium var. pubescens ”) is immature. In my opin¬
ion, this specimen cannot be assigned with
certainty to any native or introduced species. Be¬
cause of this, in the early stage of my work on North
American Corispermum I regarded the name C.
americanum as a nomen dubium. Fortunately, there
is another authentic specimen, which evidently was
annotated by Nuttall. This specimen is deposited
in the Besser memorial collection at KW. It is la¬
beled “Corispermum hyssopifolium * americanum
m. Nuttall”; as in other similar cases, an asterisk
indicates the taxon described by Nuttall himself.
The herbarium sheet contains an immature plant,
which is almost identical to the PH specimen, and
a branch from another plant, which is more mature.
Corispermum americanum was cultivated by Nuttall
(see Nuttall, 1818), and probably the more mature
specimen represents a fragment collected from a
cultivated plant. Because of this find the long-ne¬
glected name C. americanum can be restored.

Judging from the description of Corispermum pi-
losum Rafinesque (1836) (since no authentic her¬
barium material has yet been traced), this taxon
does not belong to Corispermum at all. It was de¬
scribed from Florida, where no species of Corisper-
mum are (or have been) known. The name C. pi-
losum probably refers to the hispid form of Salsola
kali L. aggregate (immature specimens of Corisper¬
mum and Salsola L. are often confused in herbaria).
Moreover, the name is a later homonym of C. pi-
losum Pallas.

In my opinion, Corispermum villosum Rydberg
(1897) should be regarded as a distinct native spe¬
cies, which is closely related to the North American
c. americanum and Asian-North American C. pal-
lasii Steven (see below), and possibly also to C.
chinganieum Iljin (more distantly). Intermediate

forms between C. americanum and C. villosum oc¬
casionally occur in regions where these species are
sympatric. The characteristic features of C. villosum
s. str. are small (usually 1.8-3.0 X 1.5-2.0 mm)
fruits, almost wingless, or with a narrow wing up to
0.1(-0.15) mm broad; both wingless and narrow¬
winged forms are sympatric, but wingless plants
seem to be much more common in the Rockies. In
the taxonomy of Corispermum, as well as of many
other Chenopodiaceae, the most reliable characters
are those of the fruits. The degree of pubescence
is an unstable character in most representatives of
the genus: it largely depends on the age of the plant
(plants often become glabrous at maturity). Because
of this I can find no sound justification for main¬
taining C. villosum and C. emarginatum Rydberg
(1904) as separate species. The lectotype of the first
species (Montana: Manhattan, 1895, Rydberg 2623,
US; isolectotype at NY) is younger and, conse¬
quently, more pubescent; the type of C. emargina¬
tum (Wyoming: Laramie, 15 Sep. 1897, Aven Nelson
4282, RM; isotypes at GH, MO, NY) is completely
mature and almost glabrous. However, the fruit
characters are essentially the same in both species,
and, in my opinion, these taxa should be united
under the name C. villosum, which has priority.

Corispermum imbricatum A. Nelson (1909) and
C. simplicissimum Lunell (1910), despite their dif¬
ferences in habit, both have small narrow-winged
fruits of similar shape and dimensions. The type of
C. imbricatum (Wyoming: Laramie Hills, 29 Aug.
1902, Elias Nelson 733, RM) represents a morpho-
type, with stems branched from the base, and in¬
florescences distinctly clavate and dense at the
apex. This morphotype is somewhat transitional to¬
ward C. villosum. The authentic specimens of C.
simplicissimum (type: North Dakota: Pierce Co.,
Barton, on a lake shore, 26 Aug. 1890, Lunell 395,
MIN) are branched above the base, and their inflo¬
rescences are more lax and narrow: judging from
available herbarium specimens, this morphotype
seems to be common in the Great Plains region. It
differs from typical C. americanum only in branch¬
ing habit. In both C. imbricatum and C. simplicis¬
simum, branches of inflorescences are simple or
nearly so. These taxa probably represent extremes
of the variability of C. americanum s.l.

Corispermum americanum has the largest area of
distribution among the native species of Corisper¬
mum, centered mostly in the Rocky Mountains and
adjacent portions of the Great Plains, and it is very
variable, especially in its vegetative characters.
Plants occurring in the Great Plains region usually
have winged fruits and often show some degree of

342

Novon

morphological transition toward C. pallasii (see be¬
low).

Specimens related to the Corispermum pallasii-
C. americanum aggregate with unusually large (up
to 4.5-5.0 mm long) wingless or narrow-winged
fruits are known from the southwestern United
States (mostly Texas. Oklahoma, Colorado, New
Mexico), and from western central Canada (mostly
Alberta and Saskatchewan). The Canadian plants
are evidently more closely related to C. pallasii s.
str. and C. ochotense Ignatov var. alaskanum Mo-
syakin described below, but they are very different
in habit: inflorescences are usually short-clavate,
dense and thick, with strongly imbricate broad
bracts. The large-fruited Canadian plants are prob¬
ably identical to “C. hyssopifolium ” var. robustius
Hooker (see Hooker, 1840). However, I have not
seen the type of Hooker’s variety. Because of this I
am not convinced that it is the same as the large-
fruited Canadian plants, and prefer to describe the
latter as C. hookeri Mosyakin (see below).

The large-fruited plants from the southwestern
United States are more closely related to Corisper¬
mum americanum s. str. They differ from northern
large-fruited specimens in having fruits more elon¬
gate and obovate (more distinctly broadest above
the middle than in the Canadian specimens), often
with white warts (formed by loose portions of peri¬
carp) and reddish brown spots (a character that oc¬
curs in several species, including C. pallasii), usu¬
ally with a more prominent triangular “beak”
(formed by the style bases in mature fruits) at the
apex. Some of these forms remain enigmatic, and
more information is needed to determine their tax¬
onomic status. The most striking and recognizable
morphotype is described below as C. navicula Mo¬
syakin.

Another complicated case is Corispermum mar¬
ginal Rydberg. In the protologue Rydberg (1903)
cited one collection from New Mexico and three
localities from Colorado. His description matches
the characters of some plants occurring in the
southwestern United States and northern Mexico.
These plants differ from C. americanum s. str. and
C. villosum s. str. in having fruits larger, up to 4.0(-
4.5) mm long, and more prominently winged (wing
usually ca. 0.2-0.4 mm broad), and usually a more
interrupted inflorescence. Thus, Rydberg’s concept
of C. marginale as a species distinct from C. vil¬
losum was correct, and it is unfortunate that he
cited as the holotype the New Mexican specimen
(Albuquerque, 1884, C. L. Herrick s.n., NY) with
small narrow-winged fruits. This specimen differs
from C. americanum s. str. only in having slightly
more lax spikes, which is not sufficient for regard¬

ing it as specifically distinct. As is evident from his
comments, Rydberg (1897) believed that C. hysso¬
pifolium s. str. was a species with large fruits (up
to 5 mm long); this erroneous concept probably af¬
fected his decision, since he apparently tried to
stress the difference between his C. marginale and
C. hyssopifolium (sensu Rydberg). The name must
stand or fall together with the type, and because of
this the epithet “marginale” should be regarded as
a synonym of C. americanum s. str. However, I be¬
lieve that other specimens cited by Rydberg (para-
types) represent a distinctive variety (or possibly
subspecies), and it is validated below as C. amer¬
icanum var. rydbergii Mosyakin. This variety, as
well as some other forms of C. americanum, usually
has lax spikes, and due to this was constantly mis-
identified by North American authors as C. nitid-
um.

For a better understanding of these taxa, it is
necessary to comment briefly on Corispermum ni-
tidum Kitaibel ex Schultes (or “so-called C. nitid-
um”). I have seen several herbarium specimens of
the true C. nitidum collected in North America re¬
cently. Judging from the fruit morphology, almost
all American representatives of the “C. nitidum”
morphotype belong to the C. pallasii—C. american¬
um aggregate. Because of this I strongly believe
that most of the records of this species from North
America are referable to C. americanum. The first
collections of the “C. nitidum ” morphotype were
made as early as 1847 in New Mexico (A. Fendler
702, GH, MO); 1852 in Chihuahua, Mexico (G.
Thurber 811, GH); 1873 (71 S. Brandegee 841, MO);
1878 (Marcus A. Jones 655, MO); 1881-1882 in
Colorado (“Common along Pueblo,” R. W. Wood¬
ward s.n., GH); 1879 in Texas (E. Palmer 1165,
GH); these are only examples based on GH and
MO collections. This list may be greatly extended
by inclusion of references to specimens from other
herbaria. In the earliest collections most of the
mentioned localities were on the edge of civiliza¬
tion, and I cannot believe that European C. nitidum
had already been widespread in the relative “wil¬
derness” of the North American West. In the mid¬
dle of the nineteenth century this species was poor¬
ly known even in Europe. Its natural area of
distribution originally covered only the Pannonian
(southeastern Central Europe, mostly along the
Danube River and some of its tributaries in Hun¬
gary, Austria, Romania, and Bulgaria) and Pontic
(southern Ukraine and southwestern Russia) flo-
ristic regions, with a small extension into the Tu¬
ranian area (northwestern Kazakhstan). Besides, C.
nitidum does not seem to be a markedly invasive
species. It is definitely introduced in some locali-

Volume 5, Number 4
1995

Mosyakin

Corispermum from North America

343

ties in Europe (e.g., in Poland, Byelorussia, central
Russia, etc.), but it cannot be regarded as wide¬
spread in the Old World. At present I know only
two cases of successful and comparatively wide ter¬
ritorial expansion of Corispermum species: C. pal-
lasii (= C. leptopterum) in central and western Eu¬
rope, and C. declinatum Stephan ex Iljin in eastern
Europe and southwestern Siberia.

Amazing morphologic parallelism is common in
Corispermum. For example, typical specimens of C.
heptapotamicum Iljin (central Asia) may be distin¬
guished from C. nitidum by their broader, usually
linear-oblanceolate lower leaves, and fruits some¬
what similar to those of C. lehmannianum Bunge;
but in many cases herbarium specimens are almost
indistinguishable. Such species as C. filifolium C.
A. Meyer ex A. Becker, C. aralocaspicum Iljin, and
C. declinatum also can be confused with C. nitid¬
um. Fortunately, in all these cases the fruit char¬
acters help to distinguish the taxa. Another exam¬
ple of this striking parallelism between some New
World and Old World taxa of Chenopodiaceae is
Chenopodium novopokrovskyanum (Aellen) Uotila
(see Uotila, 1993), a species of the C. strictum Roth
aggregate restricted to arid regions of southeastern
Europe and the Irano-Turanian portion of Asia,
which is similar in habit (but not closely related)
to native North American taxa of the C. leptophyl-
lum Nuttall ex S. Watson aggregate.

The species that seems to be common in the
Great Lakes region is conspecific with Corispermum
leptopterum (Ascherson) Iljin, a taxon described
from central Europe. However, in Europe this spe¬
cies is evidently introduced; the possible means of
its introduction will be discussed in the forthcom¬
ing review of the European species of the genus.
Voss (1985) was the first to suggest that the Great
Lakes plants may be conspecific with C. leptopter¬
um.

Forms identical with Corispermum leptopterum
are known from Siberia (especially southern and
eastern parts, plus adjacent portions of Mongolia
and China), where they are commonly referred to
as C. sibiricum subsp. baicalense Iljin (see Straz-
din’sh, 1985). At present I cannot find evident dif¬
ferences between most of the Siberian, European,
and North American plants of this aggregate, except
for usually narrower leaves, more lax and narrow
spikes, and slightly smaller fruits in average North
American specimens. I believe that this species
may also be native in North America, at least in
part.

Deciding which name has priority for Siberian
“Corispermum leptopterum ,” one must consider at
least two older names: C. patens Fisher ex Roemer

& Schultes and C. pallasii Steven (both described
from Siberia; see Steven, 1817). I have not yet
traced any mature and unquestionably authentic
specimens of the first species, and it still remains
a nomen dubium. The type of C. pallasii (Dahuria,
Helm s.n., H) belongs to the group that includes
plants previously referred to mostly as C. sibiricum
s.l. The holotype specimen is not completely ma¬
ture, and because of this I previously hesitated to
assign it to a particular race of this group. More¬
over, some presumably authentic specimens of C.
pallasii in Besser and Turczaninov memorial col¬
lections at KW belong to C. redowskii Fischer ex
Steven, a distinctive Siberian species with pubes¬
cent undulate-winged fruits. However, Steven knew
C. redowskii well (see Steven, 1817), and it is im¬
probable that he could confuse it with another spe¬
cies. He mentioned only one collection of his C.
pallasii (“Semina in specimine juniore a D. Helm
mihi dato, videntur emarginata”), which should be
consequently regarded as the holotype. Study of
several young fruits of the type of C. pallasii, and
their comparison with immature fruits of European
C. leptopterum and Siberian C. sibiricum subsp.
baicalense, revealed no evident difference between
them. Thus, the earliest available name for this spe¬
cies is C. pallasii.

Corispermum pallasii may be distinguished from
other species occurring in the United States and
Canada by its comparatively large (usually 3.0-4.5
X 2.0-3.5 mm), obovate or obovate-elliptic fruits
(normally distinctly broadest above the middle);
thick wing (translucent only along margins) (0.2-)
0.3-0.4 mm broad, which is usually long-adnate to
the style bases, forming a broadly triangular or
rounded-triangular apex; style bases often separat¬
ed to below the edge of the wing; and inflorescences
usually clavate and rather dense, or occasionally
clavate-linear (the last character is very variable).
Judging from the morphology of the fruits, this spe¬
cies is closely related to C. americanum.

Some European forms of the Corispermum pal¬
lasii aggregate, known mostly as C. membranaceum
Iljin or C. leptopterum var. membranaceum (Iljin)
Aellen, have smaller and less winged fruits, mor¬
phologically approaching C. americanum. The
Asian relatives of C. pallasii are: C. stauntonii Mo-
quin-Tandon, C. elongatum Bunge ex Maximowicz
s.l. (including C. confertum Bunge ex Maximowicz),
C. candelabrum Iljin, C. bardunovii M. Popov ex
M. Lomonosova, and some other species. Corisper¬
mum sibiricum Iljin s. str. (subsp. sibiricum =
subsp. jenissejense Iljin) is also related to this ag¬
gregate, but more distantly. Corispermum sibiricum
subsp. amurense Iljin is conspecific with C. elon-

344

Novon

gatum. For additional information about C. pallasii
(= C. leptopterum) see Iljin (1929), Aellen (1961,
1964), Strazdin'sh (1985), Kock (1986), and Mo-
syakin (1988b).

In the Atlas Florae Europaeae (Jalas & Suomi-
nen, 1980) and the second edition of Flora Euro-
paea (Aellen & Akeroyd, 1993), Corispermum pal¬
lasii (= C. leptopterum) and another distinct
species, C. gallicum Iljin, were erroneously sub¬
merged in C. intermedium Schweigger. The latter is
a distinctive endemic of the southeastern coast of
the Baltic Sea, and its differences from C. pallasii
are numerous and evident: constant absence of
perianth segments, broadly ovate or almost orbic-
ulate fruits with broader wing rounded at the apex,
etc. In spite of some similarity between the habits
of C. intermedium and the C. canescens Kitaibel ex
Schultes-C. marschallii Steven aggregate, their his¬
tory and development were independent, repre¬
senting two hypothetical migration routes in the
westward European expansion of the genus: north¬
ern route, affected mostly by glacial factors of the
Pleistocene, especially Riss-Wiirm/Mikulino inter¬
glacial; and southern route, affected mostly by the
pre-Pleistocene and Pleistocene transgressions and
regressions of inland seas. The first migration path
may be traced from east to west by the present
distribution of C. algidum Iljin (northeastemmost
European Russia), C. intermedium (Baltic Sea
coasts), and C. gallicum (southern France). Addi¬
tional evidence supporting this geohistorical recon¬
struction may be found in the discovery of fossil
fruits of Corispermum in the Late Weichselian de¬
posits in England (ca. 42,000-45,000 years BP)
and Belgium (see Godwin, 1975; Bell, 1969). The
genus was not known from the British Isles in his¬
torical times until the very recent discovery of in¬
troduced C. pallasii reported by Gibson (1992) as
C. leptopterum. The end of the second (southern)
migration path of Corispermum in Europe is marked
by the westernmost area of natural distribution of
C. canescens s. str. and C. nitidum in the Pannonian
floristic region.

Because of some parallels between arid and nu¬
tritionally deficient habitats (see Small, 1973), the
xeromorphic species of Corispermum are well
adapted to both hot deserts of central Asia and
southwestern North America, and cool river shores
of subarctic Siberia and Alaska. Usually they are
most abundant in pioneer, marginal, and naturally
and/or artificially disturbed habitats. In the Qua¬
ternary the most widescale pre-anthropogeic dis¬
turbance was caused by Pleistocene glaciations
(see, e.g., Pielou, 1991); the glacial events affected
both distribution and evolution of many species

with “weedy” life strategies (see Harlan & de Wet,
1965). An excellent example of a response of
“weedy” species to the glacial and post-glacial
events of the late Pleistocene and Holocene is Am¬
brosia L. (see Bassett & Terasmae, 1962).

I believe that the Pleistocene glaciations and
early Holocene post-glacial events played an im¬
portant role in the history of Corispermum. Expan¬
sions and retreats of glaciers, as well as eustatic
and isostatic sea level changes, created many pio¬
neer, marginal habitats, especially fluvioglacial, al¬
luvial, littoral deposits in coastal and limnetic ar¬
eas, river valleys, etc. Abundance of such habitats
provided excellent migration routes for ancestors of
modem species of Corispermum. Since some evi¬
dence exists that paleo-Americans collected and
consumed Corispermum fruits (see Betancourt et
al., 1984), the possible role of prehistoric people in
the local dispersal of seeds should also be taken
into consideration.

More detailed reconstruction of the history of
distribution and migration of Corispermum during
the Pleistocene and Holocene will be published lat¬
er. However, it seems to be useful to mention some
basic considerations concerning the history of the
genus in North America.

Corispermum was present in North America at
least 38,000 years BP (before present) in Alaska
and the Yukon, 11,000-14,000 years BP in Arizona
and Utah, ca. 4,000 years BP in New Mexico (for
details see Betancourt et al., 1984; Matthews,
1982; Rosendahl, 1948; Young, 1982).

In my opinion, ancestors of the modem native
North American taxa of this genus migrated to non-
glaciated parts of Alaska and the Yukon through
the Bering Land Bridge after the Pelukian (Kazan-
tsevo/Sangamon) sea transgression, most possibly
after Zyryanka/Early Wisconsin glaciation, during
the Kargin (Zyryanka-Sartan) interstadial. Their
further southward expansion became possible after
the retreat of Late Wisconsin glaciers and separa¬
tion of Laurentide and Cordilleran ice sheets. There
is also the possibility of their migration to more
southern parts of North America during the Middle
Wisconsin interstadial; this helps to explain the
most ancient records of fossil Corispermum fruits in
the southwestern United States.

Judging from the present distribution of Coris¬
permum species and available fossil records of the
genus, there were at least three main paleo-migra-
tion routes of ancestors of modem taxa in North
America. All these routes started in non-glaeiated
parts of Beringia. The Pacific route followed the
coastal line of the Pacific Ocean and main river
valleys of northwestern North America. This route

Volume 5, Number 4
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Mosyakin

Corispermum from North America

345

seems to be the main migration path for ancestors
of C. pacificum and C. pallidum. The two other mi¬
gration corridors are marked by the present distri¬
bution of species of the C. pallasii—C. americanum
aggregate. One of these reconstructed routes went
southward and partly southwestward along the
Rocky Mountains and adjacent plains, and another
crossed Canada in a southeastern direction toward
the Great Lakes. During their expansion, species of
Corispermum probably followed shorelines of Pleis¬
tocene glacial and post-glacial paleo-lakes (Agas¬
siz, McConnell, precursors of the Great Lakes, etc.)
and ancient river valleys, as well as fluvioglacial
deposits exposed after the retreat of glaciers.

Validation of New Taxa

As stated above, Corispermum is represented in
North America by several native species belonging
to at least three species aggregates (species groups).
Five of these species are described below.

Corispermum pacificum Mosyakin, sp. nov.
TYPE: U.S.A. Washington: Wawawai, appar¬
ently introduced, Oct. 1893, C. V. Piper 1770
(Native ! 1900 C. V. P.) (holotype, GH; iso¬
types, F, GH, MIN, MO, NY, UC). Figure 1A.

Planta annua, ca. 15-40 cm alta, a basi ramosa, glabra
vel sparsim stellato-pilosa (saepissime in partibus juven-
alibus). Folia 2—5(—7) cm longa, 0.2-0.6 mm lata, anguste
lanceolata, lineari-lanceolata vel linearia, plana, apice
brevissime mucronulata. Inflorescentia plerumque com-
pacta, densa vel rarius plus minusve remotiflora, apice
solum congesta: ovoidea, elongato-ovoidea, elongato-cla-
vata, rarius late linearia. Bracteae saepissime (1.0—)1.5—
2.5 cm longae, 0.3-0.7(-0.9) cm latae; ovato-lanceolatae,
anguste ovato-lanceolatae, ovatae, lanceolatae, apice ac-
uminatae; saepissime late membranaceo-marginatae, im-
bricatim dispositae. Perianthium monophyllum, apice
denticulatum. Fructus maturi 3.0-4.0 X 2.7—3.8 mm, sub-
rotundati vel rotundato-obovati, per lineam mediam latis-
simi vel rarius leviter supra medium, in fronte paulo con-
vexi vel rarius subplani, a tergo plerumque plani vel
leviter concavi, nitiduli, alati; ala plerumque (0.2—)0.3—
0.6 mm lata, membranacea, subpellucida, margine undu-
lala vel irregulariter minutissime denticulata (rarius inte-
gerrima), apice rotundata vel rarius subemarginata; nucle¬
us maturus saepissime niger, rarius atro-olivaceus.

Annual plant ca. 15-40 cm tall (or occasionally
more?), usually branched from the base; glabrous,
or sparsely covered with branched and almost stel¬
late hairs (especially when young). Leaves usually
2—5(—7) cm long, 0.2-0.6 mm broad, narrowly lan¬
ceolate, linear-lanceolate, or linear; flat or nearly
so; short-mucronulate at apex. Inflorescences usu¬
ally compact, rather dense, or sometimes ± lax,
condensed only near the apex; ovate, ovate-elon¬
gate, obovate, elongate-clavate, or occasionally

broadly linear. Bracts (1.0-)1.5-2.5 cm long, 0.3—
0.7(—0.9) cm broad; ovate-lanceolate, narrowly
ovate-lanceolate, almost ovate, or lanceolate, acu¬
minate at apex; usually with broad membranous
margins; overlapping. Perianth segment 1, with
erose or denticulate apex. Mature fruits usually
3.0-4.0 X 2.7-3.8 mm, almost orbicular or orbic-
ular-obovate in outline, broadest near the middle
(or rarely slightly above the middle); slightly convex
(rarely almost flat) at the abaxial side, flat or slight¬
ly concave at the adaxial side; shiny; winged. Wing
usually (0.2—)0.3-0.6 mm broad, thin, translucent;
slightly undulate or indistinctly erose-denticulate at
margins; rounded or rarely indistinctly emarginate
(notched) at apex. Mature fruit body (nucula, “ker¬
nel”) in most cases black (rarely deep olive green),
sharply contrasting with greenish semi-transparent
wings.

Corispermum pacificum is not closely related to
other native or introduced North American taxa. Its
closest allies are Siberian C. crassifolium Turczan-
inov and C. maynense Ignatov. The latter species
occurs in the northeastern Russian Far East and
may be expected to occur in Alaska and/or north¬
western Canada. Corispermum pacificum differs
from C. maynense by its usually more robust habit,
and its wing rounded (rarely rounded-truncate or
indistinctly emarginate, but not triangular) at apex.
From C. crassifolium it may be distinguished by the
constant presence of perianth segments, and more
flattened black mature fruits. At present C. pacifi¬
cum is known to me only from cited localities in
Washington, Oregon, and northwestern Idaho, but
it probably also occurs in adjacent regions of Brit¬
ish Columbia.

The specimens of this species were misidentilied
by Maihle and Blackwell (1978) partly as Corisper¬
mum hyssopifolium, and partly as C. nitidum. I do
not agree with these determinations, nor with rea¬
sons for splitting C. pacificum into two species.

Paratypes. U.S.A. Idaho: Nez Perce Co., sandy island
in Clearwater River, opposite Lewiston, 12 Oct. 1924, H.
St. John 6792 (UC). Oregon: Howell’s Pacific Coast
Plants, Josephine Co., Grant’s Pass, Sep. 1887, Thomas
Houiell s.n. (OSC); Multnomah Co., sandy bar on the Co¬
lumbia River just E of the “Interstate Bridge.” 23 Sep.
1915, M. A. Flinn s.n. (OSC); Plants of Oregon. Sand bar
on Hayden Island opposite Vancouver, Washington, 11
Oct. 1919, J. C. Nelson 2955 (GH); Whitman Co., Wa-
wawaii, Snake River, dry river sands, 13 Oct. 1923, F. A.
Warren 168 (NY); Flora of Northwest America. Sandy
ground, Hayden Island, Oregon, 8 Oct. 1927, J W.
Thompson 3915 (MO, PH); Hood River Co., sandy muddy
bank of the Columbia River, with Riccia and Cyperus.
about 15-20 mi. W of the Dalles, Oregon. 22 Oct. 1961,
Georgia Mason 4201 (OSC). Washington: Flora of Wash¬
ington Territory. Northern Transcontinental Survey, Yaki-

346

Novon

E

E

Figure 1. Fruits of Corispermum : comparison between North American and Asian species. —A. Corispermumpacificum (North America). —B. Corispermumpallidum (North America).
—C. Corispermum platypterum (Asia). —D. Corispermum maerocarpum (Asia). —E. Corispermum welshii (North America).

Volume 5, Number 4
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Mosyakin

Corispermum from North America

347

ma Region, sands of the Columbia, Oct. 1882, Frank
Tweedy s.n. (NY); W. Klickitat Co., sandy banks of the
Columbia River, 11 Sep., 29 Oct. 1893, W. N. Suksdorf
1385 (F, GH, MIN, MO, NY, UC) [A mixed collection; see
note below—S.M.]; Klickitat Co., Spedio, sandy shores
and sandy slopes, very common, 19 Sep. 1925, M. W.
Gorman s.n. (OSC); Whitman Co., sandy bank of the
Snake River at Wawawai, 15 Oct. 1939, Marion Ownbey
& Ruth P. Ownbey s.n. (CAN, GH, OSC, PH, RM, RSA,
TEX, UC); Whitman Co., Wawawai—Lewiston road along
Snake River, on sandy beach at edge of river, 15 Sep.
1956, R. T. Ogilvie s.n. (DAO, UBC).

Some degree of reproductive isolation between
the sympatric species of Corispermum exists be¬
cause of different phenology, especially different
flowering period. However, occasional hybrids
probably occur between partly sympatric C. paci-
ficum and C. villosum. These hybrids are similar in
habit to C. pacificum: they have usually broad
leaves and dense inflorescences. However, many of
their fruits (up to 50—70%) are abortive, almost
lacking a perisperm. Occasional well-developed
fruits are polymorphic and show transitional char¬
acters: they are usually narrow-winged or almost
wingless, strongly convex at the abaxial side (char¬
acters of C. villosum), black or nearly so, broadly
ovate to orbiculate in outline (characters of C. pa¬
cificum). The high percentage of abortive fruits in¬
dicates that C. pacificum and C. villosum are tax-
onomically rather distant species, because hybrids
between closely related taxa of Corispermum (i.e.,
belonging to the same species aggregate) usually
show no evident loss of fertility. The studied spec¬
imens of C. pacificum X C. villosum are cited be¬
low:

U.S.A. Oregon: Umatilla Co., mouth of Umatilla River,
Umatilla, 11 July 1915, M. E. Peck 6667 (OSC); along
Columbia, opposite Dalles, Oregon, 1915, J. Evans s.n.
(OSC); Baker Co., 28 Nov. 1952, LeRoy Wright s.n. (OSC);
Hood River Co., drifting sand of the Columbia River, 10
Sep. (fl), 1 Oct. 1923 (fr), L F. Henderson 344 (of 1924)
(MO). Washington: Wallula, sandy soil, alt. 220 m, 17
Sep. 1894, J. B. Leiberg 912 (OSC).

Corispermum pallidum Mosyakin, sp. nov.
TYPE: U.S.A. Washington: Douglas Co., in
drifting sand, Wilson Creeks, 25 July 1893. J.
II. Sandberg & J. B. Leiberg 309 (holotype,
MO; isotypes, CAN, F, OSC, PH. UC, US). Fig¬
ure IB.

Planta annua, ca. 5—25 cm alta, a basi ramosa, sub¬
glabra vel in partibus juvenalibus stellato-pilosa et min-
utissime papillosa. Folia 2—5 cm longa, 0.1—0.2(-0.3) cm
lata, linearia, anguste linearia vel filiformia, plana vel par-
tim convoluta, apice brevissime mucronulata. Inflorescen-
tia plerumque laxiuscula, rarius densiuscula; linearia vel
elongato-linearia. Bracteae (0.5—)1.0—1.5(—2.0) cm longae,
0.2—0.5 cm latae; plerumque anguste ovato-lanceolatae

vel lanceolatae, apice acuminatae; saepissime membran-
aceo-marginatae, imbricatim dispositae vel plus minusve
remotae. Perianthium monophyllum, apice denticulatum.
Fructus maturi saepissime 2.8-3.5(-3.8) X 2.4—2.8(-3.3)
mm, rotundato-obovati vel obovati, plerumque supra me¬
dium vel rarius per lineam mediam latissimi, in fronte
subplani vel subplano-convexi, a tergo plerumque plani
vel leviter concavi, subnitidi vel opaci, alati; ala pler¬
umque 0.7—1.0 mm lata, membranacea, subpellucida,
margine erosa vel irregulariter eroso-denticulata (rarius
subintegra), apice stylorum basi longe adnata, emarginata
vel rotundata; incisura inter basin stylorum margine ala¬
rum profundiore; nucleus maturus pallidus, stramineus vel
brunneus, saepe maculatus.

Annual plant ca. 5-25 cm tall (or more?),
branched from the base, almost glabrous, or cov¬
ered with branched and almost stellate hairs and
minute papillae (especially on veins of young bracts
and upper leaves). Leaves 2-5 cm long, 0.1-0.2(-
0.3) cm broad, linear, narrowly linear, or filiform;
flat or occasionally convolute (especially at matu¬
rity); acuminate or short-mucronulate at apex. In¬
florescences usually lax, rarely rather dense; linear
or elongate-linear. Bracts usually (0.5—)1.0-1.5(-
2.0) cm long, 0.2-0.5 cm broad, usually narrowly
ovate-lanceolate or lanceolate, acuminate at apex;
in most cases with distinctly membranous margins;
overlapping or slightly remote. Mature fruits 2.8-
3.5(—3.8) X 2.4—2.8(—3.3) mm, rounded-obovate or
obovate in outline, usually broadest above the mid¬
dle or occasionally near the middle; almost flat or
slightly convex at abaxial side, plane or slightly
concave at adaxial side; slightly shiny or dull;
winged. Wing normally 0.7—1.0 mm broad, thin,
translucent, with erose or irregularly erose-dentic-
ulate (rarely almost entire) margins; long-adnate to
the style bases, emarginate (notched) or rounded at
apex. Style bases unusually long (ca. 0.7-1.0 mm,
including their parts adnate to wing), in most cases
divided to below the edge of the wing. Mature fruit
body (“kernel,” nucula) flattened, pale, usually
straw-colored or yellowish brown, occasionally with
reddish brown spots.

Corispermum pallidum seems to be a well-de¬
fined species. The combination of its characters is
very distinctive: pale, flattened, and small fruit
body; very broad (especially as compared to the
fruit dimensions), thin translucent wing with erose
margins; and long style bases distinctly divided in
their upper parts. Young bracts and upper leaves
of C. pallidum are often papillose on margins and
veins, in combination with typical branched tri-
chomes. This character (papillosity) is quite rare in
Corispermum, since most of its species are normally
covered only with branched or almost stellate hairs.
The same combination of branched hairs and min-

348

Novon

ute papillae occasionally occurs in some Asian spe¬
cies. It is hard to believe, but C. pallidum seems
to be most closely related to the eastern Asian spe¬
cies C. macrocarpum Bunge ex Maximowicz, which
occurs at present in the southern Russian Far East
and northeastern China, mostly along the Amur
River and some of its tributaries. Two Manchurian
species, C. sterwlepis Kitagawa and C. platypterum
Kitagawa, also belong to the same aggregate (for
additional information about these Asian taxa, see
Kitagawa, 1935; Baranov, 1969; Tsien Cho-po &
Ma Cheng-gung, 1979). Fruits of C. pallidum are
very similar to those of C. macrocarpum in their
morphological features, but not in their size; the
latter species has much larger fruits 5.0-6.0(-6.5)
X 3.8-5.5 mm (Fig. 1B-D).

Maihle and Blackwell (1978) placed rather uni¬
form specimens of Corispermum pallidum into two
taxa, C. hyssopifolium sensu Maihle & Blackwell
(Sandberg & Leiberg 309, Thompson 6764) and C.
nitidum sensu Maihle & Blackwell (St. John, Court¬
ney & Parker 4948).

Paratypes. U.S.A. Washington: Flora of Eastern Wash¬
ington. Wilson Creek, July 1893, J. H. Sandberg & J. B.
Leiberg s.n. (MIN); Grant Co., dry sand, SW of Moses
Lake, alt. 1000-1150 ft., 25 June 1921, Harold St. John,
W. D. Courtney & Charles S. Parker 4948 (GH, UC); Grant
Co., sandy sagebrush plains near Quincy, 15 June 1931,
J. William Thompson 6764 (GH, MO).

Corispermum welshii Mosyakin, sp. nov. TYPE:
U.S.A. Utah: Kane Co., sand dunes, with pon-
derosa pine, willow, and gambel oak, ca. 1900
m. Coral Pink Dunes, T43S, R8W, S8, ca. 8
mi. due W of Kanab, 25 Sep. 1992, S. L. Welsh
& K. H. Thorne 25170 (holotype, MO; iso¬
types, BRY, NY). Figure IE.

Planta annua, ca. 10-35 cm alta, a basi ramosa, dense
vel sparsim stellato-pilosa. Folia 2-6 cm longa, 0.2-0.4(-
0.5) cm lata, lineari-lanceolata vel linearia, fere plana vel
rarissime partim subconvoluta, apice brevissime mucron-
ulata. Inflorescentia plerumque compacta, densa vel rarius
plus minusve laxiuscula, apice congesta; ovoidea, elon-
gato-obovata vel elongato-clavata. Bracteae 1.0-3.0 cm
longae, 0.3-0.8 cm latae; ovatae vel ovato-lanceolatae,
rarius anguste ovato-lanceolatae, apice acuminatae; late
membranaceo-marginatae, imbricatim dispositae. Perian-
thium monophyllum, apice denticulatum. Fructus maturi
saepissime (3.3—)3.7-4.6 X (2.7—)3.0—3.6 mm, obovati vel
rotundato-obovati, plerumque supra medium vel rarius per
lineam mediam latissimi, in fronte convexi, a tergo pler¬
umque leviter concavi, subnitidi vel opaci, alati; ala pler¬
umque (0.3—)0.4—0.6 mm lata, membranacea, subpellu-
cida (rarius valdiuscula, margine solum subpellucida),
margine integra vel irregulariter eroso-denticulata, apice
rotundata, truncata vel subemarginata; nucleus maturus
stramineus, fulvus vel brunneus, saepissime maculatus et
albido-verrucosus.

Annual plant ca. 10—35 cm tall (or more?),
branched from the base or nearly so, densely or
sparsely covered with branched or stellate hairs.
Leaves 2-6 cm long, 0.2-0.4(-0.5) cm broad, lin¬
ear-lanceolate or linear, usually plane, or rarely
slightly convolute or folded (especially in dried
plants), short-mucronulate at apex. Inflorescences
usually rather compact and dense, rarely ± lax,
condensed only at apex; ovate, oblong-obovate, or
oblong-clavate in outline. Bracts 1.0—3.0 cm long,
0.3-0.8 cm broad, ovate or ovate-lanceolate (rarely
narrowly ovate-lanceolate), acuminate at apex, with
broad membranous margins. Perianth segment 1,
minutely dentate at apex. Mature fruits normally
(3.3-)3.7—4.6 X (2.7-)3.0-3.6 mm, broadly obovate
or obovate-orbicular in outline, in most cases
broadest above the middle (rarely near the middle);
convex at abaxial side, usually concave at adaxial
side; slightly shiny or dull; winged. Wing usually
(0.3—)0.4-0.6 mm broad, thin, translucent (or oc¬
casionally more thick, translucent only at margin),
with entire or irregularly erose-denticulate margins;
rounded, truncate, or indistinctly emarginate at
apex. Mature fruit body (nucula, “kernel”) straw-
colored (yellowish brown), light brown or brown,
usually with reddish brown spots and whitish warts.

This species is probably closely related to the
broad-winged and large-fruited plants of the south¬
western U.S.A. ( Corispermum americanum var. ryd-
bergii Mosyakin), which Rydberg included in his C.
marginale (excluding the type of C. marginale).
However, C. welshii differs from the mentioned va¬
riety, as well as from C. americanum sensu proprio,
in having shorter, broader, and denser spikes, dis¬
tinctly broader leaves and bracts, more rounded
(occasionally almost orbicular), and often more
prominently winged fruits. Its difference from C.
villosum is also evident, since it has larger fruits
approximately twice as long and broad as the latter.
From C. pallasii it may be distinguished by its more
orbicular fruits having membranaceous wings,
which are rounded, rounded-truncate, or indistinct¬
ly emarginate at apex. In general appearance C.
welshii resembles some eastern Asian taxa, for ex¬
ample, C. elongatum s.l. It is also somewhat similar
in habit to European C. marschallii but is not close¬
ly related to it. At present C. welshii is known to
me only from Utah and Wyoming, but similar forms
(which are in need of additional study) occur also
in other southwestern states, particularly in Colo¬
rado and western Texas. For example, the following
collection from western Texas probably also belongs
to C. welshii (it differs from typical plants in having
more lax spikes and prostrate habit, but the fruits

Volume 5, Number 4
1995

Mosyakin

Corispermum from North America

349

are almost identical to those of the type): Flora of
Texas. Gypsum, Hudspeth Co., from 13 mi. E of
Dell City on nearly barren gypsum dunes, 14 Oct.
1984, R. D. Worthington 12752 (TEX).

Paratypes. U.S.A. Utah: Millard Co., sand dunes, alt.
4500 ft., 18 Sep. 1926, collected by Cottam, determined
by Garrett 1010 (BRY); Millard Co., juniper association,
sand dunes, 18 Sep. 1926, coll. A. 0. Garrett 1010 (RM);
Salt Lake Co., South Salt Lake City, just S of intersection
of 7400 South and 20th East in a sand dune area, alt.
4650 ft., T. 25, R. 1 E„ SE ii of Sec. 28, 29 Oct. 1967,
L. Arnou 805 (GH), 805a (COLO); Kane Co., Halls Creek
Bay, Lake Powell, sandy benchland, 4 June 1972, S. L
Welsh & G. Moore 11797 (BRY) [this plant evidently was
buried by sand, and because of this it has developed atyp-
ically long branches—S.M.]; Kane Co., Coral Pink Sand-
dunes State Park S of Mt. Carmel Junction, elev. 5500 ft.,
11 Oct. 1972, B. Albee 1215 (UT); Uintah Co., T3S, R22E,
SW Sec. 4, Red Fleet Reservoir, ca. 6000 ft. elev., opens
in juniper community, sandy soil, 3 Oct. 1980, E. Neese
& M. Chatterley 9894 (NY, RM) [these specimens have
fruits slightly smaller than in the type—S.M.]; Kane Co.,
Navajo Sandstone, Ponderosa, Oakbrush community, T43S
R7W S30, Moquith Mountain, ca. 7 mi. W of Kanab, 2142
m elev., 25 Sep. 1992, K. Thorne & S. L. Welsh 15192
(BRY) [also a specimen with slightly smaller fruits—
S.M.]. Wyoming: Sweetwater Co., sand dunes, near
Steamboat Mt., 1 Sep. 1936, Marion Ownbey & Robert L.
Lang 1122 (COLO); Niobrara Co., T34N R61W Sec 36
SE V *, sandy roadside, associates: Cenehrus , Salsola ; elev.
4700 ft., 26 Sep. 1986, R. Dorn 4549 (NY) [inflorescences
of this specimen are more lax than in the type, showing a
transition toward C. americanum —S.M.].

Corispermum navicula Mosyakin, sp. nov.
TYPE: U.S.A. Colorado: Jackson Co., North
Park, North sand dunes due E of Dowell
Ranch, T10N, R78W, Sec. 6, between Govern¬
ment Creek and North Sand Creek, Eagle Hill
Quadr., 29 Sep. 1976, F. Martin Brown s.n.
(holotype, COLO).

Planta annua, ca. 5—15(—25?) cm alta, a basi ramosa,
sparsim stellato-pilosa vel glabriuscula. Folia (1.5—)2—4
cm longa, 0.1-0.5 cm lata, anguste lanceolata vel iineari-
lanceolata, plana, rarissime partim convoluta. Inflorescen-
tia plerumque compacta, densa, ovoidea vel elongato-ovo-
idea. Bracteae 0.5-2.0 cm longae, 0.2-0.6 cm latae;
imbricatim dispositae. Perianthium monophyllum. Fructus
maturi (4.2-)4.5-5.0(-5.2) X 2.5-3.0 mm, elongato-obov-
ati, supra medium latissimi; in fronte valde convexi. a
tergo plerumque distinctissime concavi; apteri vel anguste
alati. Ala plerumque 0.1—0.2(—0.3) mm lata, valdiuscula.
non pellucida vel semipellucida; margine integerrima vel
minutissime eroso-denticulata, saepe involuta; apice ros-
trata, triangularia, stylorum basi longe adnata. Nucleus
maturus atro-brunneus vel atro-viridis, plerumque macu-
latus et/vel albido-verrucosus.

Annual plant ca. 5—15(—25?) cm tall, branched
from the base, sparsely covered with branched and
almost stellate hairs, or almost glabrous. Leaves
(1.5-)2-4 cm long, 0.1-0.5 cm broad, narrowly lan¬

ceolate or linear-lanceolate, flat or rarely convolute.
Inflorescences usually compact and dense, ovate or
oblong-ovate in outline. Bracts 0.5-2.0 cm long,
0.2-0.6 cm broad, strongly imbricate. Perianth seg¬
ment 1. Mature fruits (4.2-)4.5-5.0(-5.2) X 2.5-
3.0 mm, elongate-obovate or narrowly obovate-el-
liptic, broadest above the middle; strongly convex
at abaxial side, usually prominently concave at ad-
axial side; narrowly winged or almost wingless.
Wing usually 0.1-0.2(—0.3) mm broad, rather thick,
not translucent, or semi-translucent; with entire or
indistinctly erose margins, often involute toward the
adaxial side; rostrate-triangular and long-adnate to
style bases at apex. Mature fruit body (nucula,
“kernel”) usually dark brown or dark green, usually
with dark spots and whitish warts.

This species is amazingly similar in its fruit mor¬
phology to the Siberian species Corispermum bar-
durwvii M. Popov ex M. Lomonosova (see M. N.
Lomonosova, 1992). Corispermum navicula and C.
bardunovii probably represent results of parallel
evolution (or parallel variability?) within North
American and Asian representatives of the C. pal-
lasii aggregate, respectively. It is also worth men¬
tioning that the most distinctive characters of C.
navicula and C. bardunovii, such as elongated fruit
body with almost parallel margins in the middle
portion and distinctly triangular apex, show a tran¬
sition (or a parallel evolutionary trend?) toward the
representatives of Corispermum sect. Declinata Mo¬
syakin (1994). Additional study of C. navicula is
desirable for clarification of its relationships with
other species. Some specimens from Oklahoma
(e.g., Stillwater, Oklahoma, in sand on bank of Ci¬
marron, 6 Nov. 1899, E. E. Bogue s.n., MIN) prob¬
ably also belong to C. navicula, or at least approach
this species most closely.

Paratype. U.S.A. Colorado: Jackson Co., North Sand
Hills, E of Cowdrey, T10N, R78W, S6 NE W, 2575 m, 1
Oct. 1982, D. H. Wilken 13924 (NY, RM).

Corispermum hookeri Mosyakin, sp. nov. TYPE:
Canada. Saskatchewan: District de Moose Jaw,
palier sablonneux au pied du Coteau du Mis¬
souri, 5-6 mi. 4 Test de Mortlach, dune ev-
entrce, 11 Sep. 1960, B. Boivin & G. F. Led-
ingham 14079 (holotype, DAO; isotypes, NY,
TRT, UBC).

Planta annua, ca. 10-40 cm alta, a basi ramosa, spar¬
sim stellato-pilosa vel glabriuscula. Folia 2—5 cm longa,
(0.1-)0.2-0.5(-0.6) cm lata, anguste lanceolata vel li-
neari-lanceolata, plana. Inflorescentia plerumque densa
ovoidea, ovato-clavata vel ovato-cylindrica, rarissime in-
terrupta. Bracteae 0.5-1.5(-2.0) cm longae, 0.3-1.0 cm
latae, ovatae vel ovato-lanceolatae, imbricatim dispositae.

350

Novon

Perianthium monophyllum. Fructus maturi (3.2—)3.5-
4.5(—5.0) X 2.2—3.3(-3.5) mm, elongato-obovati vel obov-
ati, per lineam mediam latissimi vel leviter supra mediam
latiores; in fronte valde convexi, a tergo concavi vel sub-
plani; apteri vel angustissime alati. Ala ad 0.2 mm lata,
semipellucida; margine integerrima, apice rotundata vel
leviter triangularia. Nucleus maturus atro-olivaceus, brun-
neus vel rarissime niger, plerumque non maculatus et non
verrucosus, rarius maculatus.

Annual plant ca. 10-40 cm tall (or occasionally
more?), branched from the base, sparsely covered
with branched and almost stellate hairs. Leaves
usually 2-5 cm long, (0.1-)0.2-0.5(—0.6) cm broad,
narrowly lanceolate or linear-lanceolate, plane. In¬
florescences usually dense, ovate, ovate-clavate, or
ovate-cylindric, rarely interrupted (especially near
the base). Bracts 0.5-1.5(—2.0) cm long, 0.3-1.0
cm broad, ovate or ovate-lanceolate, imbricate
(usually strongly overlapping). Perianth segment 1.
Mature fruits (3.2-)3.5-4.5(—5.0) X 2.2-3.3(-3.5)
mm, oblong-obovate or obovate, broadest near the
middle or slightly above the middle; strongly con¬
vex at abaxial side, prominently concave to sub¬
plane at adaxial side; wingless or narrowly winged.
Wing (when present) up to 0.2 mm broad, semi¬
transparent; margins entire, apex rounded or indis¬
tinctly triangular. Mature fruit body (nucula, “ker¬
nel”) usually deep olive green, brown, or rarely
almost black, normally without spots and/or warts,
or occasionally spotted.

This species seems to be most closely related to
Corispermum pallasii s. str. and C. ochotense var.
alaskanum (see discussion above and comments on
C. ochotense below). It is also similar to the small-
fruited species, C. villosum, and to the large-fruited
C. navicula described above. Some specimens of C.
hookeri from British Columbia have unusually elon¬
gated fruits somewhat similar to those of Asian spe¬
cies of Corispermum sect. Declinata. These plants
are described below as a new variety.

Paratypes. CANADA. Alberta: Lethbridge, 27 Aug.
1911, M. 0 . Malt s.n. (CAN); Battle River, S of Wain-
wright, sandy soil, near river bank, 8 Sep. 1951, E. H.
Moss 9725 (DAO) [the voucher specimen: 2n = 18, de¬
termined by G. A. Mulligan]; SW of Orion, sand dunes,
grazed by livestock in the area, 28 Aug. 1952, E. H. Moss
10110 (CAN). British Columbia: very common on Dryas
Island in Fraser River at Hope, 19 July 1914, T. L Thack¬
er s.n. (UBC); Lytton, 26 Sep. 1922, W. B. Anderson 6678
(V); in sand landfill near Alex Fraser Bridge, Annacis
Island, many plants in sand, 7 Sep. 1988, Frank Lomer
s.n. (UBC); sandy ground under the Pattullo Bridge, Sur¬
rey, a single plant, 7 Sep. 1991, Frank Lomer s.n. (UBC);
Steveston, Richmond, Lulu Island, 49°07'N, 123°10'W, 7
Sep. 1992, Huber Moore & Terry Taylor s.n. (UBC); sand
landfill dredged from Fraser River, near Fraser Surrey
Docks, Surrey, many plants in sand, 27 Aug. 1993, Frank
Lomer s.n. (UBC) [this plant is transitional toward C. hook¬

eri var. pseudodeclinatum Mosyakin described below—
S.M.]. Manitoba: sand hill 35 mi. SE of Brandon, 23 Sep.
1960, G. A. Stevenson 2255 (CAN, DAO). Northwest
Territories: Mackenzie District, in dry sand near shore¬
line, sand hills on S shore of Mackenzie River about 16
mi. downstream from Fort Simpson, 61°58'N, 121°45'W,
25 July 1955, W. J. Cody & J. M. Matte 8944 (DAO) [the
specimen is immature; fortunately, authors also collected
some fragments of dry plants remaining from the previous
year—S.M.]; Mackenzie District, occasional in disturbed
sand along roadside on mainland about 1 mi. S of Fort
Simpson island, 61°52'N, 121°22'W, 3 Aug. 1955, W. J.
Cody & J. M. Matte 9130 (DAO) [the specimen is im¬
mature, but very similar to the previous one—S.M.]. On¬
tario: Sarnia, Oct. 1936, W. A. Dent s.n. (DAO); Norfolk
Co., Long Point, 22 Sep. 1946, F. A. Clarkson 6684 (DAO)
[a specimen with the same collection number at TRT ap¬
proaches C. pallasii —S.M.]; Long Point Village, sand on
open shore, 24 Sep. 1966, R. Beschel 15722 (CAN); Nor¬
folk Co., Long Point, South Walsingham Tp., 12 Oct.
1969, P. M. Catling & J. E. Cruise s.n. (TRT); Norfolk
Co., Long Point, along S shore E of Courtright Ridge, Lake
Erie, blow-out on sand dune, 23 Sep. 1976, K. M. Lindsay
& 1. D. MacDonald 1092 (CAN); Bruce Co., boundary
between Chiefs’ pt. I.R. and Amabel Twp., on sand, beside
road at edge of pine plantation, 29 Sep. 1977, Joseph W.
Johnson (TRT); Norfolk Co., Long Point, Squires Ridge to
Little Creek Ridges, 12 x /i to 14 Vi mi. from base of Point,
on S beach, open, active sand dunes along beach, nu¬
merous, with Ammophila breviligulata and Andropogon
scoparius, 20 Sep. 1979, A. A. Reznicek & P. M. Catling
5255 (DAO). Saskatchewan: Beaver Creek, sandy hill¬
side, 9 Sep. 1937, W. J. White s.n. (TRT); Saskatoon, on
sands by river, Sep. 1931, A. W. Anderson s.n. (CAN);
Saskatoon, 20 mi. SW at Pike Lake, partially covered sand
dunes, 10 Sep. 1951, G. F. Ledingham 1219 (DAO);
Moose Jaw Region, Caron, SW Va 16-17-29 W. 2nd, shift¬
ing sand in and around blow-out spots, long-abandoned
land, 24 Sep. 1955, T. H. Hudson 1768 (DAO); District
de Rosetown-Biggar, coulee Agassiz, Canton Bernard, 12-
15 mi. au sud-est de Demaine, Platiferes de la Saskatch-
ewan-Sud, dune en mouvement, 18 Sep. 1960, B. Boivin
14156 (DAO); Saskatoon District, Saskatoon, sandflat be¬
tween natural levee and cutbank of S Sask. River, local,
with Elymus canadensis, Psoralea lanceolata, 13 Sep.
1964, J. H. Hudson 2214 (DAO); Lake Atabasca, E of
William Point, Salix, Myrica on dry, eroding sand beach
on shore, 6 Aug. 1975, G. W. Argus & D. J. White 9750
(CAN) [inflorescence interrupted—S.M.].

Corispermum hookeri Mosyakin var. pseudo¬
declinatum Mosyakin, var. nov. TYPE: Can¬
ada. British Columbia: beside Burnaby Lake,
sand dune, 3 Oct. 1965, /. Bayly 83 (holotype,
UBC).

Fructus elongati vel oblongo-obovati, longitudine lati-
tudinem duplo vel subduplo superanti; in fronte valde
convexi, a tergo concavi. Inflorescentia elongata, interrup-
ta, apice solum congesta.

Fruits elongate, oblong-obovate, ca. 2 times as
long as broad; prominently convex at abaxial side,
concave at adaxial side. Inflorescences elongate,
interrupted, condensed only at apex.

Volume 5, Number 4
1995

Mosyakin

Corispermum from North America

351

Paratype. CANADA. British Columbia: Okanagan,
Quilchena, 30 July 1906, E. Wilson 776 (UBC).

Corispermum americanum (Nuttall) Nuttall var.
rydbergii Mosyakin, var. nov. TYPE: Mexico.
Chihuahua: sand dunes, 10-19 Oct. 1935,
Harde LeSueur 285 (MO).

Fructus maturi (3.2 -)3.3 4.0(- 4.3) X 2.3-3.2 mm, sae-
pissime late alati, supra medium latissimi. Ala 0.3-0.4(—
0.5) mm lata. Inflorescentia plerumque angusta, laxius-
eula, non denst, plus minusve remotiflora. Ceterum typo
respondet.

Mature fruits (3.2-)3.5-4.0(-4.3) X 2.3-3.2 mm,
usually broadly winged, broadest above the middle.
Wing 0.3-0.4(—0.5) mm broad. Inflorescences nor¬
mally narrow, lax; flowers/fruits ± remote. Other
characters as in C. americanum var. americanum.

Paratype. U.S.A. Colorado: Huerfano Valley, near
Gardner, sandy creek bottom, alt. 7000 ft., 10 Sep. 1900,
F. K. Vreeland 657 (NY, CAN) [this specimen is also a
paratype of C. marginale Rydberg; see discussion above—
S.M.],

Because the Corispermum americanum group is
currently under revision, I refrain now from citing
additional paratypes of this variety. However, it
seems to occur in several localities in Arizona, Col¬
orado, New Mexico, Texas, and Utah. New collec¬
tions of mature specimens from this region would
be extremely helpful for detailing its distribution.
Due to its characteristic narrow and interrupted in¬
florescence, this variety, as well as many specimens
of C. americanum s. str., was constantly misiden-
tified as C. nitidum.

Corispermum ochotense Ignatov var. alaskan-
um Mosyakin, var. nov. TYPE: U.S.A. Alaska:
Onion Portage, Upper Kobuk River, 67°06'N,
158°15'W, in sand dunes, open site, 10 Aug.
1967, Charles Schuieger 196 (holotype, ALA).

Fructus maturus 3.2 -1.0 mm longus, 2.2—2.7 mm latus.
Ceterum ut in var. ochotense.

Mature fruit 3.2-4.0 mm long, 2.2-2.7 mm wide.
Other characters as in variety ochotense.

Paratypes. U.S.A. Alaska: Yukon River between
Rampart and Tanana, 9 Aug. 1932, L J. Palmer 13
(CAN); Ester Siding Field, July 1936, G. IF Gasser s.n.
(ALA); Kobuk River valley, forming dense “stands’ on
sandy river bar at Point Goldie on Kobuk River, 2 Aug.
1974, C. H. Racine 732 (ALA); Kobuk River valley, low,
abundant plant covering sand surface of dunes on S side
of Kobuk River opposite mouth of Hunt River, 12 Aug.
1974, C. H. Racine 731 (ALA).

Corispermum ochotense, a comparatively small
(often dwarf) arctic and subarctic plant, has been

described recently from the northeastern Russian
Far East (Ignatov, 1986). It is normally floriferous
almost to the base, often developing fruits even in
the axils of lower leaves and branches. The lower
and middle bracts in the inflorescence are usually
linear or linear-oblanceolate (often broadest toward
the apex, as well as some leaves), rather long, leaf¬
like, almost not different from cauline leaves in
their shape and size. In addition, plants often turn
deep beet red at maturity. The illustration of “C.
hyssopifolium” in the Flora of Alaska (Hulten,
1968) gives an accurate impression of the habit of
C. ochotense. The type of C. ochotense (“regio Ma¬
gadan, distr. Ola, pagus Talon, syrtis arenosa ripar-
ia, 28—30 Aug. 1971, A. P. Khokhrjakov”), which
I have studied at MHA, is eonspecific with the
Alaskan and northwestern Canadian plants. How¬
ever, some of the Alaskan specimens differ from
their trans-Beringian relatives in having larger
fruits. The typical variety of C. ochotense also oc¬
curs in North America. The species is known to me
from the following additional localities in Canada
(as many specimens were not completely mature,
the varieties are not indicated; however, most of the
specimens seem to belong to var. ochotense):

CANADA. Northwest Territories: District of Mac¬
kenzie, Aklavik River near Aklavik, on the section of slit
beach, 6 Sep. 1948, Margaret E. Oldenburg 48—1241
(GH); Mackenzie District, Norman Wells, forming thick
mats on eroding heavy soil on slope by airstrip, 22 July
1953, W. J. Cody & R. L Gutteridge 7466 (COLO, DAO,
GH, MICH, MO, OSC, TRT, UBC); Mackenzie District,
rare on gravel beach of Mackenzie River, Royal Canadian
Corps Signals Transmitters, 5 mi. upstream from Norman
Wells, 27 July 1953, W. ]. Cody & R. L Gutteridge 7603
(DAO, F); Mackenzie District, 13.0 km WSW of Norman
Wells, N.W.T. Site No. 8-10-13, beach ridge, sandy, with
Populus tremuloides , 10 Aug. 1974, N. J. Walker 2303
(DAO); Mackenzie District, E bank of MacKenzie River,
Norman Wells, N.W.T. Site No. 8-18-1, riverbank, many
disturbance species, rare, 18 Aug. 1974, N. J. Walker
2386 (DAO).

The following collection from northern Saskatch¬
ewan probably also belongs here (although the
plants were not completely mature):

CANADA. Saskatchewan: S shore of Lake Atabasca
at Beaver Point, lower sandy beach line at, and just above,
shoreline, scattered, but frequent, 8 Aug. 1979, V. L
Harms & R. A. Wright 27215 (DAO, GH, RM).

As outlined above, the native North American
species of Corispermum are comparable to most of
the widely recognized Eurasian taxa of the genus,
regarding the species concept applied to them and
morphological differences between species. They
are also comparable to some other readily recog¬
nized taxa of Chenopodiaceae, e.g., numerous seg-

352

Novon

regate species of such North American aggregates
as Chenopodium leptophyllum s.l., or C. fremontii
S. Watson s.l. However, especially for followers of
a broader species concept, I would like to outline
the species aggregates (groups) of closely related
taxa of North American Corispermum. Because
North American representatives of this genus were
commonly misidentified and misunderstood, this
explanation would be helpful in preventing possible
further mistakes.

The closest Asian allies of American Corisper¬
mum are C. pallasii (for almost all native taxa), C.
macrocarpum (for C. pallidum ), and C. crassifolium
(for C. pacificum). In my opinion, in the future C.
pacificum and C. pallidum could not be lumped
together with other North American taxa. However,
using a very broad species concept, ore may regard
C. pacificum as a subspecies of C. crassifolium.
Corispermum pallasii is most closely related to C.
americanum, C. navicula, and C. welshii; under a
broad species concept the three latter species could
fit at the subspecies level as well. Corispermum
ochotense, C. hookeri, and C. villosum form another
unit, which is, however, connected to the previous
one by occasional forms transitional between C.
americanum and C. villosum, as well as by transi¬
tions between C. pallasii and C. hookeri. Using the
ultra-broad species concept, one could unite the
mentioned taxa under the priority name C. pallasii.
However, I believe that at the present stage of our
knowledge, such an approach could only obscure
the true relationships within this group.

At present I prefer to place the native North
American representatives of Corispermum into
three species aggregates (groups):

the Corispermum pallasii aggregate: C. pallasii s.
str., C. americanum, C. navicula, C. welshii, C.
hookeri, C. villosum, C. ochotense; also the Asian
C. sibiricum, C. stauntonii, C. elongatum, C. bar-
dunovii, and probably some others.

the Corispermum crassifolium aggregate: C. pa¬
cificum plus the Asian C. crassifolium s. str. and C.
maynense.

the Corispermum macrocarpum aggregate: C. pal¬
lidum and the Asian C. macrocarpum s. str., C.
platypterum, and C. stenolepis.

A key to the species and species aggregates of
Corispermum occurring in North America will be
published later, in a treatment of the genus for the
fourth volume of the Flora of North America. Nat¬
ural arrangement of some Eurasian species and
species aggregates will be discussed in an article
submitted to the Ukranian Botanical Journal
(Ukrajins’kyj botanicnyj zurnal).

New, more complete (and preferably serial) her¬

barium material of North American Corispermum,
as well as field and experimental studies, would be
helpful for further progress in understanding the
complicated patterns of taxonomy and distribution
of the genus in North America.

Acknowledgments. The present study of North
American Corispermum was possible due to the
support provided by the Missouri Botanical Garden
(MO). I thank all members of the staff of the Flora
of North America Project for their extremely kind
help and advice. The kind help of the staff of many
American, Canadian, Russian, Ukrainian, Finnish,
Austrian, Bulgarian, Chinese, and Japanese her¬
baria (listed alphabetically: ALA, BKL, BRY, CAN,
COLO, DAO, F, GH, H, KW, KWHA, LE, MHA,
MICH, MIN, MO, MU, MW, NY, OSC, PE, PH, RM,
RSA, SOM, TEX, TI, TRT, UBC, UC, US, UT, UTC,
V, W, WU) is especially greatly appreciated. I thank
Charlie Jarvis (BM) and Konstantin Dobolyi (BP)
for sending me excellent photographs of type spec¬
imens and fruits of Corispermum; thanks are also
due to Amy Scheuler McPherson, Diana Gunter,
and Helen Jeude (MO) for their skillful comments
on the style of the manuscript. The fieldwork in the
southwestern United States (Utah and Nevada) dur¬
ing August and September 1994 was made possible
by support from the National Science Foundation
and kind help from Noel Holmgren (NY), Stanley
L. Welsh (BRY), and Steven E. Clemants (BKL).
Special thanks are due to Nancy R. Morin (MO),
Leila M. Shultz (GH), Clifford W. Crompton (DAO),
and Ihsan Al-Shehbaz (MO) for productive discus¬
sion, and to Yevonn Wilson-Ramsey (MO) and
Anne Keats Smith (MO) for their kind assistance
in producing illustrations and obtaining necessary
literature.

Literature Cited

Adamkiewicz, E. 1970. Studia cyto-taksonomiczne i em-
briologiczne nad czterema gatunkami rodzaju Corisper¬
mum L. (Cytotaxonomic and embryological studies on
four species of the genus Corispermum L.). Acta Soc.
Bot. Poloniae 39(4): 626-652.

Aellen, P. 1961. Corispermum L. Pp. 716-723 in: G.
Hegi, Illustrierte Flora von Mitteleuropa. Aufl. 2. Bd.3.
T.2. Munchen.

-. 1964. Corispermum L. Pp. 99—100 in: T. G. Tu-

tin et al.. Flora Europaea. Vol. 1. Cambridge Univ.
Press, Cambridge.

-(revised by J. R. Akeroyd). 1993. Corispermum

L. Pp. 119-120 in: T. G. Tutin et al.. Flora Europaea.
Ed. 2. Vol. 1. Cambridge Univ. Press, Cambridge.
Baranov, A. I. 1969. The species of Corispermum (Che-
nopodiaceae) in Northeastern China. J. Jap. Bot. 44:
161-169; 195-206.

Bassett, I. J. & J. Terasmae. 1962. Ragweeds, Ambrosia
species, in Canada and their history in postglacial time.
Canad. J. Bot. 40: 141-150.

Volume 5, Number 4
1995

Mosyakin

Corispermum from North America

353

Bell, F. G. 1969. The occurrence of southern, steppe and
halophyte elements in Weichselian (last-glacial) floras
from southern Britain. New Phytol. 68: 913-922.

Betancourt, J. L., A. Long, D. J. Donahue, A. J. T. Jull &
T. H. Zabel. 1984. Pre-Columbian age for North Amer¬
ican Corispermum L. (Chenopodiaceae) confirmed by
accelerator radiocarbon dating. Nature 311: 653-655.

Gibson, Ch. 1992. Corispermum leptopterum (Aschers.)
Iljin in North Essex. B.S.B.I. News, No. 62: 41—42.

Godwin, H. 1975. The History of the British Flora: A
Factual Basis for Phytogeography. Ed. 2. Cambridge
Univ. Press, Cambridge.

Harlan, J. R. & J. M. J. de Wet. 1965. Some thoughts
about weeds. Econ. Bot. 19: 16-24.

Hooker, W. J. 1840. Flora Boreali-Americana, or the Bot¬
any of the Northern Parts of British America. 2 vols.
London.

Hult6n, E. 1968. Flora of Alaska and Neighboring Ter¬
ritories. A Manual of the Vascular Plants. Stanford Univ.
Press, Stanford, California.

Ignatov, M. S. 1986. Chetyre novykh vida s Dal’nego Vos-
toka (Four new species from the Soviet Far East). Byull.
Moskovsk. Obshch. Isp. Prir., Otd. Biol. 91 (3): 111-
114.

Iljin, M. M. 1929. Novye vidy roda Corispermum L. (Cor-
ispermi generis species novae). Izv. Glavn. Bot. Sada
SSSR 28: 638-654.

Jalas, J. & J. Suominen. Editors. 1980. Atlas Florae Eu-
ropaeae. Distribution of vascular plants in Europe. Vol.
5. Helsinki.

Kitagawa, M. 1935. Corispermum in Manshuria & Korea.
Pp. 99—105 in: Report of the First Scientific Expedition
to Manchoukuo. Sect. 4, part 2. Tokyo.

Kock, U.-V. 1986. Verbreitung, Ausbreitungsgeschichte,
Soziologie und Okologie von Corispermum leptopterum
(Aschers.) Iljin in der DDR. I. Verbreitung und Aus¬
breitungsgeschichte. Gleditschia 14: 305-325.

Lomonosova, M. N. 1992. Corispermum L. Pp. 166-171
in: I. M. Krasnoborov & L. I. Malyschev (editors). Flora
Sibiri (Flora Sibiriae). Vol. 5 (Salicaceae-Amarantha-
ceae). Novosibirsk.

Lunell, J. 1910. New plants from North Dakota. Amer.
Midi. Naturalist 1(8): 204-208.

Maihle, N. J. & W. H. Blackwell. 1978. A synopsis of
North American Corispermum (Chenopodiaceae). Sida
7(4): 382-391.

Matthews, J. V. 1982. East Beringia during Late Wiscon¬
sin time: A review of the biotic evidence. Pp. 127—150
in D. M. Hopkins et al., Paleoecology of Beringia. Ac¬
ademic Press, London.

Mosyakin, S. L. 1988a. Pryrodno-vydova dyfferentsiatsiya
rodu Corispermum L. na terytoriyi Ukrayiny. (The nat¬
ural-specific differentiation of the genus Corispermum
L. in Ukraine.) Ukrayins’k. Bot. Zhum. 45(3): 37-40.

-. 1988b. Krytychnyy pereglyad vydiv rodu Cor¬
ispermum L. flory Ukrayiny. (A critical revision of Cor¬
ispermum species in Ukraine.) Ukrayins’k. Bot. Zhurn.
45(5): 19-23.

--. 1994. New infrageneric taxa of Corispermum L.

(Chenopodiaceae). Novon 4: 153-154.

Nelson, A. 1909. Coulter’s New Manual of Botany of the
Central Rocky Mountains. Revised ed. American Book,
Chicago.

Nuttall, T. 1818. The Genera of North American Plants,
and a Catalogue of the Species to the Year 1817. Vol.
2. Philadelphia.

-. 1834. Collections toward a flora of Arkansas.

Amer. Philos. Soc. Trans. 5: 139—203.

Pielou, E. C. 1991. After the Ice Age: The Return of Life
to Glaciated North America. Univ. Chicago Press, Chi¬
cago & London.

Rafinesque, C. S. 1836. New Flora and Botany of North
America. Part 4. Philadelphia.

Rosendahl, C. 0. 1948. A contribution to the knowledge
of the Pleistocene flora of Minnesota. Ecology 29: 284—
315.

Rydberg, P. A. 1897. Rarities from Montana. I. Bull. Tor-
rey Bot. Club. 24(4): 188-192.

-. 1903. Studies on the Rocky Mountain flora. X.

Bull. Torrey Bot. Club. 30(4): 247-262.

-. 1904. Studies of the Rocky Mountain flora—XI.

Bull. Torrey Bot. Club 31: 399-410.

Small, E. 1973. Xeromorphy in plants as a possible basis
for migration between arid and nutritionally-deficient
environments. Bot. Not. (Lund) 126: 534—539.

Steven, C. 1817. Observationes in Plantas Rossicas et
descriptiones specie mm novarum. M£m. Soc. Imp. Na-
turalistes Moscou 5: 334—338.

Strazdin’sh, Yu. G. 1985. Rod Corispermum L. Pp. 154—
166 in: L. V. Tabaka & al. Flora i rastitel’nost’ Latviys-
koy SSR. Vostochno-Latviyskiy geobotanocheskiy ray¬
on. (Flora and vegetation of the Latvian SSR. The
Eastern Latvian geobotanic region). Zinatne, Riga.

Tsien Cho-po & Ma Cheng-gung. 1979. Corispermum L.
Pp. 50-75 in: Flora Reipublicae Popularis Sinicae. To-
mus 25(2). Beijing.

Uotila, P. 1993. Taxonomic and nomenclatural notes on
Ckenopodium in the Flora Iranica area. Ann. Bot. Fenn.
30(3): 189-194.

Voss, E. 1985. Michigan Flora. Part 2. Dicots (Saurura-
ceae-Comaceae). Ann Arbor, Michigan.

Young, S. B. 1982. The vegetation of Land-Bridge Ber¬
ingia. Pp. 179-191 in: D. M. Hopkins et al., Paleo¬
ecology of Beringia. Academic Press, London.

Note added in proof.

Since this article was submitted for publication, some
additional isotypes and paratypes have been found in the
herbaria at US and LE: Corispermum pacificum: C. V. Piper
1770 (isotype, US); W. N. Suksdorf 1385 (paratype, US);
C. pallidum: J. H. Sandberg & J. B. Leiberg 309 (isotype,
LE); C. hookeri: B. Boivin & G. F. Ledingham 14079 (iso¬
type, LE); C. ochotense var. alaskanum: L J. Palmer 13
(paratype, US).

Corispermum welshii is now known from two additional
states of the U.S.A.: Arizona: Coconino Co., Teetso
Spring, sandy soil, 13 Nov. 1971, N. D. Atwood & D.
Kaneko 3420 (US); New Mexico: Thornton, 19 Oct.
1903, D. Griffiths 6209 (US).

A New Species of Geonoma (Arecaceae) from Panama

Gregory C. de Nevers

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A.

Michael H. Grayum

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. Geonoma mooreana is described from
wet forests on the Atlantic slope of western Pana¬
ma.

Review of herbarium material for floristic and
monographic projects involving Central American
palms (Arecaceae) has revealed the following new
Panamanian species of Geonoma (Arecoideae:
Geonomeae).

Geonoma mooreana de Nevers & Grayum, sp.
nov. TYPE: Panama. Veraguas: vicinity Santa
Fe, along road between Santa Fe and Calove-
bora, 1.7 mi. past Alto Piedra School, 1.5 mi.
beyond Quebrada Cosilla (previously referred
to as Rio Primero Braso), 8°33'N, 81°08'W.
570 m, 13 July 1994 (fruit), Croat & Zhu
76826 (holotype, MO; isotypes, CAS, PMA).

Species cum G. tenuissima H. E. Moore optime con-
gruens, sed differt foliis regulariter pinnatis pinnis angus-
tis, inflorescentiis majoribus, labio infero fovearum flor-
alium bifidoque.

Stems caespitose, with to at least 4 stems per
plant, 2-3.5 m high, slender, to at least 1.7 cm
diam. at nodes; leaves regularly pinnate; sheath 9—
18+ cm; petiole 41—52 cm, broadly channeled
adaxially, rounded abaxially; rachis 62-81 cm, oc¬
casionally terminating in a slender filament to at
least 1.7 cm; pinnae 25—35(—51) per side, narrowly
linear with one prominent raised vein adaxially, one
prominent and two less prominent raised veins
abaxially, 9—42 X 0.4—1.5(—3.1) cm, glabrous or
essentially so; inflorescence infrafoliar, 2—4 nodes
below current leaves, paniculate, branched to three
orders, 24—32 X 18-38 cm; peduncle 6-17 cm;
prophyll and peduncular bract inserted close to¬
gether at base of peduncle; prophyll 11—15.5(-21)
X 2.8—5.0 cm, slightly swollen in bud, obscurely
striate; peduncular bract slightly smaller than pro¬
phyll, included in it; inflorescence rachis 6-20.5
cm, with 4—7 primary branches; rachillae ca. 110-
165, reddish, 13—19 cm X 0.5—1 mm, sparsely to
moderately setulose at anthesis (later glabrescent).

drying rugose; floral pits spirally arranged, ca. 1—2
mm apart, the lips prominently exserted from the
rachillae, the lower lip bifid, the orifice ca. 1 X 1
mm; staminate flowers ca. 3 mm, sepals and petals
subequal, free, erect at anthesis, slightly imbricate
below; stamens 6, filaments spreading, anthers
sharply reflexed from the filaments; pistillate flow¬
ers ca. 2 mm, sepals and petals subequal, barely
exserted, styles 3, reflexed, staminodial tube trun¬
cate; fruit oblong to globose, green, drying brown,
5-6 X 4—5 mm, surface minutely verrucose (peb¬
bled); seed black, oblong, 4—5 mm; germination un¬
known. Figure 1.

Geonoma mooreana occurs in wet forests on the
Atlantic slope of western Panama, at elevations of
ca. 100-1000 m. It has been collected about 30
km from the Costa Rican border in Bocas del Toro
Province. The apparent geographic discontinuity
between the Bocas del Toro and Veraguas stations
is probably spurious, as the area in question is one
of the most poorly collected in Panama.

Geonoma mooreana is most similar phenetically
to G. scoparia Grayum & de Nevers, G. concinna
Burret, and G. tenuissima H. E. Moore. Geonoma
scoparia, with which specimens representing G.
mooreana were compared by Grayum and de Ne¬
vers (1988: 113), shares the highly branched,
broomlike inflorescence and pebbled fruit, but dif¬
fers in its consistently solitary habit, generally
smaller dimension?, trijugate leaves, and crenate
(vs. truncate) staminodial tube. Geonoma concinna
has the truncate staminodial tube and pebbled
fruits of G. mooreana, but differs in its consistently
trijugate leaves, much smaller inflorescence bracts,
and fewer, thicker rachillae. The inflorescence of
G. mooreana also resembles that of G. tenuissima
of Ecuador, which differs in its simple leaf blades,
shorter, more delicate inflorescence rachillae, and
floral pits with the lower lip entire rather than bifid.

Several collections of Geonoma mooreana bear
incomplete or partial inflorescences. Bulky palm
inflorescences usually must be pruned for pressing,
and the severed branches are often used to make

Novon 5: 354-356. 1995.

Volume 5, Number 4
1995

de Nevers & Grayum
Geonoma mooreana

355

Figure 1. Geonoma mooreana de Nevers & Grayum (A [except fruits]. Nee 11252; B, Hammel 3459; A [fruits], C,
Kirkbride & Duke 649). —A. Section of stem with leaf base and inflorescence (many rachillae removed for clarity)
(X0.50). —B. Medial portion of leaf (X0.75). —C. Section of infructescence rachilla with fruits (X1.75).

356

Novon

duplicate specimens. Thus, the PMA and CAS is¬
otypes of G. mooreana include the main basal and
suprabasal branches (respectively) that are missing
from the holotype infructescence. Isolated branches
may easily be mistaken for entire inflorescences or
infructescences in the herbarium, yielding substan¬
tially false impressions of such attributes as pedun¬
cle length, overall inflorescence size, and the num¬
ber of rachillae.

The specific epithet commemorates the late Har¬
old E. Moore, Jr. (1917—1980), leading palm au¬
thority of his generation, who realized this was a
new species and annotated two of the paratypes
(Hammel 3459 and Nee 11252) as “Geonoma sp.
nov.”

Paratypes. PANAMA. Bocas del Toro: between Q.
Lugron and Cerro Bonyic, nr. Rfo Teribe, 90-270 m, 13
Apr. 1968 (fmit), Kirkbride & Duke 649 (MO, NY). Code:
Atlantic slope along Rfo San Juan above its fork with Rfo
Tife, ca. 1200 ft., 12 June 1978 (fruit), Hammel 3459

(MO). Veraguas: Caribbean slope above Rfo Primero Bra-
zo 5 mi. NW of Santa Fe, 700-1200 m, 18-19 Mar. 1973
(flower), Croat 23120 (MO 2 sheets); along road between
Escuela Agrfcola and Alto Piedra (above Santa Fe) and
Rfo Dos Bocas ca. 5-8 km from Escuela, 730—770 m, 26
July 1974 (bud), Croat 25931 (MO); rd. from Santa Fe to
(ialovehora. creek about 5 km beyond Escuela Agrfcola,
ca. 500 m, 8°40'N, 81°5'W, 1 Mar. 1995 (past fruiting),
de Nevers et al. 10601 (CAS); lower montane wet forest 7
km W of Santa Fe on new road past agricultural school,
ca. 2900 ft., 12 Apr. 1974 (flower/young fruit). Nee 11252
(MO).

Acknowledgments. We are indebted to Kathy
Bell for the excellent line drawings of Geonoma
mooreana, and to John Dransfield and an anony¬
mous reviewer for valuable comments on the manu¬
script.

Literature Cited

Grayum, M. H. & G. C. de Nevers. 1988. New and rare

understory palms from the Peninsula de Osa, Costa

Rica, and adjacent regions. Principes 32: 101—114.

Nomenclatural Notes on the Myrsinaceae of China

John J. Pipoly III

Botanical Research Institute of Texas, 509 Pecan Street,

Fort Worth, Texas 76102-4060, U.S.A.

Chen Chieh

Kunming Institute of Botany, Academia Sinica, Kunming, People’s Republic of China

Abstract. The following nomenclatural changes
are proposed: Maesa brevipaniculata (C. Y. Wu &
C. Chen) Pipoly & C. Chen, M. confusa (C. M. Hu)
Pipoly & C. Chen, Embelia laeta (L.) Mez subsp.
papilligera (Nakai) Pipoly & C. Chen, E. ribes Bur-
man f. subsp. pachyphylla (C. Y. Wu & C. Chen)
Pipoly & C. Chen, Ardisia obtusa Mez subsp. pa¬
chyphylla (Dunn) Pipoly & C. Chen, Myrsine cica-
tricosa (C. Chen) Pipoly & C. Chen, M. verruculosa
(C. Chen) Pipoly & C. Chen, M.faberi (Mez) Pipoly
& C. Chen, and M. kwangsiensis (E. H. Walker)
Pipoly & C. Chen. Twenty-three taxa in Embelia
and Ardisia are reduced to synonymy for the first
time. Twelve names in Embelia, Ardisia, and Myr¬
sine are lectotypified.

In order to make the names available for the
forthcoming treatment of Myrsinaceae for the Flora
of China, the following nomenclatural changes in
the genera Maesa, Embelia, Ardisia, and Myrsine
(including Rapanea) are herein proposed. The au¬
thors have studied type or authentic material from
China, the Pacific, and eastern Malesia and are
broadening species concepts based on those stud¬
ies. Type specimens seen from KUN have the ac¬
ronym HY marked on them.

Maesa ForsskAl

Maesa is the only genus in the Myrsinaceae with
a semi-inferior to inferior ovary and a berrylike
fruit. This paleotropical genus contains nearly 200
species, of which 28 are known from China.

Maesa brevipaniculata (C. Y. Wu & C. Chen) Pi¬
poly & C. Chen, comb. nov. Basionym: Maesa
parvifolia A. DC. var. brevipaniculata C. Y. Wu
& C. Chen, FI. Yunnan. 1: 324. 1977. TYPE:
China. Yunnan: Malipo, 26 Aug. 1962, K. M.
Feng 22805 (holotype, KUN).

Maesa brevipaniculata is distinguished from M.
parvifolia by having a velutinous brown tomentum.

instead of a brown hirtellous-tomentose and rufous
glandular-granulose tomentum on the branchlets,
petioles, and inflorescence rachises. Therefore, it is
more consistent to recognize M. brevipaniculata as
a distinct species.

Maesa confusa (C. M. Hu) Pipoly & C. Chen,
comb. nov. Basionym: Maesa consanguinea
Merrill var. confusa C. M. Hu, Bot. J. South
China 1: 12. 1992. TYPE: China. Hainan:
Bao-ting Xian, Tong-za Mao-an, 23 Feb. 1957,
L. Tang 3789 (holotype, IBSC).

Maesa confusa is a poorly known taxon, but its
densely and minutely glandular-granulose inflores¬
cence rachises, long pedicels, highly angulate
branchlets, and densely black punctate-lineate
fruits readily separate it from M. consanguinea.

Embeua Burman f.

Embelia contains approximately 150 species of
lianas (at maturity) that are distributed in the Pa-
leotropics from southern China to Malesia, west
through Indochina to India, and on to Madagascar
and continental tropical Africa. Embelia is a mem¬
ber of the tribe Myrsineae and is the sister genus
of Cybianthus Martius (Pipoly, 1987). It is the only
lianous member of the family and is distinguished
by its plagiotropic branches and free petals and
stamens. Walker (1940) recognized 18 species in
China, of which 12, as well as 2 additional species
hitherto unknown in China, are recognized by the
authors.

Embelia pauciflora Diels, Bot. Jahrb. 29: 517.
1900. TYPE: China. Sichuan: Nanchuan, Bock
& von Rosthom 25 (holotype, B presumed de¬
stroyed; fragment, US).

Embelia blinii H. L£veill6, Repert. Spec. Nov. Regni Veg.
10: 375. 1912. Syn. nov. TYPE: China. Guizhou:
near Mt. College, to Kematong, Feb. 1898 (pistillate
fl), J. Chattanjon & E. Bodinier 2081 (holotype, E;
isotypes. A, P).

Novon 5: 357-361. 1995.

358

Novon

Embelia dielsii H. L6veill6, Repert. Spec. Nov. Regni Veg.
10: 374. 1912. Syn. nov. TYPE: China. Guizhou:
Pin-Fa, 17 Feb. 1903 (staminate fl),./. Cavalerie (ho-
lotype, E; isotypes. A, P).

Embelia kaopoensis H. I/veiHc, Repert. Spec. Nov. Regni
Veg. 12: 186. 1913. Syn. nov. TYPE: China. Guizh¬
ou: Kao-Po, 15 Dec. 1911 (pistillate fl, fr), E. Bod-
inier s.n. (holotype, E).

Embelia pauciflora apparently exhibits extreme
local variation in its precocious staminate flower¬
ing, which causes abrupt quantitative variation for
those plants bearing essentially “juvenile” leaves.
Differences among the populations represented by
the type specimens of Embelia blinii, E. dielsii, E.
kaopoensis, and E. schlechteri are entirely quanti¬
tative and represent at best localized variation.

Embelia laeta (L.) Mez in Engler, Pflanzenr. 9(IV.
236): 326. 1902.

Embelia laeta is readily recognized from the oth¬
er members of Embelia by the minute emargination
at the leaf apex and gray to reddish bark. It consists
of two subspecies, of which subspecies papilligera
is distributed in China (Guangdong, Jiangxi, Tai¬
wan) and Japan, whereas subspecies laeta is dis¬
tributed in China (Fujian, Guangdong, Guangxi,
Hainan, Jiangxi, Taiwan, Yunnan), Cambodia, Laos,
Thailand, and Vietnam. It appears that subspecies
papilligera grows in somewhat more humid habitats
than subspecies laeta. The two subspecies may be
easily distinguished by the following key.

la. Branchlets straight to somewhat flexuous, gla¬
brous; leaves chartaceous, margins flat; pedicels
and calyx glabrous; petals glandular-ciliate along

the margin.subsp. laeta

lb. Branchlets geniculate, minutely and densely ru¬

fous glandular papillose-tomentose; leaves cori¬
aceous, margins revolute; pedicels and calyx
glandular-puberulent; petals glabrous along the
margin.subsp. papilligera

Embelia laeta (L.) Mez subsp. papilligera (Nakai)
Pipoly & C. Chen, stat. nov. Basionym: Sam¬
ara laeta var. papilligera Nakai in Nakai &
Honda, Nov. Fl. Japan 9: 24. 1943. TYPE: Ja¬
pan. Mt. Arisan, between Heishana & Niman-
daira, to 7000 ft. alt., Apr. 1914 (fl, fr), Hayata
s.n. (holotype, TI).

Embelia parviflora Wallich ex A. de Candolle,
Trans. Linn. Soc. 17: 130. 1834. TYPE: India.
Assam: Wallich 2307 (holotype, K-W).

Embelia pulchella Mez in Engler, Pflanzenr. 9(IV. 236):
324. 1902. Syn. nov. SYNTYPES: India. Assam:
Griffith s.n. (B destroyed, K); Manipur: Watt 7277
(lectotype, here designated, K).

Embelia myrtifolia Hemsley & Mez, Notizbl. Bot. Gart.
Berlin 3: 107. 1901. SYNTYPES: China. Yunnan:
Mentze, S mountains, (pist. fl), A. Henry 12826; Chi¬
na. Yunnan: Mentze, S mountains, (stam. and pist.
fls), A. Henry 9384 (lectotype, here designated. A;
isolectotypes, MO, US (pist. fl)).

Embelia ribes Burman f., Fl. Ind. 62. pi. 23. 1768.

As a result of a detailed study of Embelia ribes
throughout its geographic range, it became appar¬
ent that two distinct entities exist. One grows in
forests on well-drained soils, while the other is re¬
stricted to water-logged habitats. These are recog¬
nized as subspecies that can be distinguished as
follows:

la. Branchlets lenticellate; leaf blade chartaceous,
sordid adaxially; floral bracts subulate; petals
erect in anthesis; stamens and staminodes equal
in size, subequaling petals; anthers longer than
wide; fruit 3—4(—5) mm diam. when dried; plants
of forests, on well-drained sands or sandy

loams.subsp. ribes

lb. Branchlets without lenticels; leaf blade coria¬

ceous to cartilaginous, nitid adaxially; floral
bracts lanceolate; petals highly reflexed in an¬
thesis; stamens slightly exserted ca. 1 mm in an¬
thesis, staminodes ca. Vs petal length; anthers
wider than long; fruit 2—3 mm diam. when dried;
plants of poorly drained clay areas, swamps, and
forests.subsp. pachyphylla

Embelia ribes subsp. ribes is distributed in China
(Fujian, Guangdong, Guangxi, Guizhou, Hainan,
Xizang, and Yunnan), Sri Lanka, India, Myanmar,
Laos, Thailand, Cambodia, Vietnam, Malaysia, Su¬
matra, Java, and Kalimantan.

Embelia ribes Burman f. subsp. pachyphylla (C.
Y. Wu & C. Chen) Pipoly & C. Chen, stat. nov.
Basionym: Embelia ribes var. pachyphylla
Chun ex C. Y. Wu & C. Chen, Fl. Yunnan. 1:
364. 1977. TYPE: China. Guangdong: Le-
chang Xian, Zuo Jing-lie 20451 (holotype,
KUN).

Embelia tenuis Mez in Engler, Pflanzenr. 9(IV. 236): 314.
1902. Syn. nov. TYPE: Malaysia. Sarawak: Hullet
271 (holotype, K).

Subspecies pachyphylla is distributed in China
(Guangdong, Guangxi, Hainan, Yunnan), Sarawak,
Sumatra, and Vietnam.

Embelia vestita Roxburgh, FL Ind. ed. Carey 2:
288. 1824.

Volume 5, Number 4
1995

Pipoly & Chen
Myrsinaceae of China

359

Embelia prunifolia Mez in Engler, Pflanzenr. IV (236):
316. 1902. SYNTYPES: China. Yunnan: Mentze,
1700-2000 m, (stam. fl), A. Henry 9380C (B de¬
stroyed, A); 10260 (B destroyed, A), 11327A,
11327B (B destroyed. A); China. Yunnan: 5500', A.
Henry 9380 (lectotype, here designated. A; isolec-
totype, B destroyed).

Embelia oblongifolia Hemsley, J. Linn. Soc., Bot. 26: 62.
1882. Syn. nov. TYPE: China. Guangdong: Lofau-
shan. Ford 90 (holotype, K).

Embelia bodinieri H. Levoillf'. Repert. Spec. Nov. Regni
Veg. 9: 327. 1911. Syn. nov. TYPE: China. Guizhou:
./. Esquirol 743 (holotype, E; isotype, P).

Embelia rudis Handel-Mazzetti, Anz. Akad. Wiss. Wien.
Math.-Nat. 59: 108. 1922. Syn. nov. TYPE: China.
Jiangxi: Handel-Mazzetti 157 (holotype, W).

Embelia nigroviridis C. Chen, Acta Phytotax. Sin. 16(3):
81. 1978. Syn. nov. TYPE: China. Xizang: Cha-yu,
Tsinghai-Tibet Expedition 73-678 (holotype, HP).

Embelia vestita is here treated as a polymorphic,
broad-ranging species with numerous locally vari¬
able populations. It grows in mixed forests, ever¬
green, broad-leaved forests, shrubby areas, and
limestone hillsides in China (Fujian, Guangdong,
Guangxi, Guizhou, Hainan, Hunan, Sichuan, Tai¬
wan, Xizang, Yunnan, Zhejiang), India, Myanmar,
Nepal, and Vietnam.

The type of Embelia rudis represents populations
whose tertiary veins are extremely thick and whose
adaxial leaf surface is more nitid than the type of
E. vestita. The type of Embelia oblongifolia repre¬
sents populations with remotely serrate leaves and
fewer, coarse secondary veins than those of the type
of E. rudis. The variation noted above precluded
segregation of even infraspecific groupings. There¬
fore, the species is treated as a polymorphic och-
lospecies (sensu White, 1962; Prance, 1972, 1982;
Pipoly, 1983). The highly nitid leaf blades and
inconspicuously pellucid or brown punctate sepals
easily separate E. vestita from E. undulata and E.
scandens, the most closely related congeners.

A finisi a Swartz

Ardisia is a pantropical genus of about 500 spe¬
cies, including many as yet undescribed from Viet¬
nam, Sabah, Sarawak, Sulawesi, New Caledonia,
Madagascar, Colombia, Ecuador, and Peru. It is
represented in China by 64 species, most of which
are members of subgenus Crispardisia.

Ardisia obtusa Mez in Engler, Pflanzenr. 9(IV.
236): 104. 1907. TYPE; China. Hainan: Henry
7990 (holotype, B destroyed; photo and frag¬
ment, US).

Ardisia obtusa is a member of subgenus Akosmos
Mez, which includes about 50 species and has its
centers of diversity in Borneo, Celebes, and Mo¬

luccas. It is represented in China by two subspe¬
cies: subspecies obtusa is endemic to China
(Guangdong and Hainan) and grows in thickets,
broad-leaf forests, hilly areas, and along creek mar¬
gins; subspecies pachyphylla grows in dense ever¬
green broad-leaf forests, steep valley slopes, and in
places with poor drainage in China (Guangxi, Hai¬
nan) and Vietnam. The two subspecies can be sep¬
arated as follows:

la. Leaves oblanceolate to obovate; petioles broadly
marginate, 1.2—1.5 cm long; sepals 1.2-1.5 mm
long; petals 3-6 mm long, inconspicuously pellucid
punctate; 0-100 m, lowland forests . . . subsp. obtusa

lb. Leaves narrowly oblanceolate or linear-lanceolate;

petioles maiginate and canaliculate, 0.6-1 cm long;
sepals 0.8-1 mm long; petals 2-2.5 mm long,
densely and minutely orange punctate, 400-700 m,
montane forests.subsp. pachyphylla

Ardisia obtusa Mez subsp. pachyphylla (Dunn) Pi¬
poly & C. Chen, comb, et stat. nov. Basionym:
Ardisia pachyphylla Dunn, Bull. Misc. Inform.
1912: 368. 1912. TYPE: China. Hainan: Five
Finger Mountains, Katsumata s.n, (HK No. 6680)
(holotype, HK; photo and fragment. A, K).

Ardisia gracUiflora Pitard in Lecomte, FL Gen. Indo-Chine
3: 832.1930. Syn. nov. TYPE: [Vietnam], Annam: Phu-
Hu, prov. Nhatiang, 19 Jan. 1923 (fl), E. Poilane 1362
(holotype, P; isotypes, A 2 sheets).

Ardisia graciliflora differs from subspecies pachy¬
phylla by a combination of largely quantitative char¬
acters, and it is considered to be a diminutive form
of subspecies pachyphylla, perhaps a juvenile flow¬
ering precociously.

Our study of specimens most commonly referred to
Ardisia yurmanensis Mez has revealed that the earliest
name for the taxon is Ardisia thyrsiflora D. Don.
Quantitative differences in leaf length and number of
flowers per corymb has caused taxonomic confusion.
This species, as now interpreted, is most closely re¬
lated to A. amherstiana A. DC., a taxon most fre¬
quently encountered in Vietnam. The revised synon¬
ymy follows.

Ardisia thyrsiflora D. Don, Prodr. Fl. Nep. 148.
1825. TYPE: Nepal, Wallich s.n. (BM).

Ardisia tenera Mez in Engler, Pflanzenr. 9(TV. 236): 104.
1902. Syn. nov. SYNTYPES: China. Yunnan: Szemao,
1500 m, A. Henry 12123A (B destroyed, A); China.
Yunnan: Szemao, 1500 m, A. Heruy 12094 (holotype,
B destroyed; lectotype, here designated. A; isolecto-
types, MO, US).

Ardisia yurmanensis Mez in Engler, Pflanzenr. 9(TV. 236):
107. 1902. Syn. nov. TYPE: China. Yunnan: Szemao,
5000', tree 30', fls. white, A. Henry 13095 (holotype, B
destroyed; lectotype, here designated. A; isolectotypes,

MO, NY, US).

360

Novon

Ardisia penduliflora Mez in Engler, Pflanzenr. IV(9). 236:
150.1902. Syn. nov. SYNTYPES: China. Hubei: 1885—
1888 (11), A. Henry 4314 (B destroyed); China. Hubei:
1885-1888 (fl), A. Henry 6365 (lectotype, here desig¬
nated, US).

Ardisia undulata Mez in Engler, Pflanzenr. 9(IV. 236): 146.
1902. Syn. nov. TYPE: China. Yunnan: Mephra, 1700
m, A Henry 10779 (holotype, B destroyed; lectotype,
here designated. A; isolectotypes, MO, US). Later hom¬
onym, not C. B. Clarke (1882).

Ardisia thyrsiflora grows on woody hills and in
broad-leaf forests, valleys, and damp places in China
(Guangxi, Hainan, Xizang, Yunnan), India, Myanmar.
Nepal, and Vietnam. It is a highly variable species
closely related to A. amherstaiana A. DC., A. waitakii
C. M. Hu, A. baotingensis C. M. Hu, and A. quinque-
gona Blume. These four taxa, along with Ardisia fordii
Hemsley, a diminutive plant, form a natural group
within subgenus Akosmos. The separation of these
taxa in the flora is at best artificial, and further studies
are needed to resolve the limits in this complex.

Ardisia corymbifera Mez in Engler, Pflanzenr. 9(IV.
236): 149. 1902. SYNTYPES: China. Yunnan:
Simao, 1250 m, (fl), A. Henry 11724 (B de¬
stroyed); China. Yunnan: Simao, (fl), A. Henry
12,000A (holotype, B destroyed; lectotype, here
designated. A; isolectotype, MO).

Ardisia corymbifera Mez var. tuberifera C. Chen, Acta Phy¬
totax. Sin. 16(3): 80. 1978. Syn. nov. TYPE: China.
Guangxi: Lung Chow, S. Chun 13096 (holotype, HK).
Ardisia maculosa Mez in Engler, Pflanzenr. 9(IV. 236): 146.
1902. Syn. nov. SYNTYPES: China. Yunnan: Szemao,
6000 ft. elev. (fl, fr), A. Henry 12088D (B destroyed);
China. Yunnan: Szemao, shrub 8', fls. pink, fr. red. (fl,
fr), A. Henry 12088E (holotype, B destroyed; lectotype,
here designated, US 458840; isolectotypes, A
00025251, 00025252; MO 126457, 126458). NOTE:
Herbarium accession numbers are included here be¬
cause many sheets contain two accession numbers with
two specimens, one of which may not have been cited
in the protologue.

Ardisia radians Hemsley & Mez in Engler. Pflanzenr. 9(IV.
236): 146. 1902. Syn. nov. Blahfya radians (Hemsley
& Mez) Masamune, Short PL Form. 162. 1936. SYN¬
TYPES: China Yunnan: Szemao, 4500 ft. elev., A. Hen¬
ry 535, 9226A (B destroyed, K); China. Yunnan: Sze¬
mao. 4500 ft. elev., shrub 8', fls white (fl), A. Henry
11624C (lectotype, here designated, K; isolectotypes. A,
B destroyed, MO).

Ardisia patens Mez in Engler, Pflanzenr. 9(IV. 236): 149.
1902. Syn. nov. TYPE: China. Yunnan: Szemao, 5000
ft. elev., shrub 3', fr. red (fr), A. Henry 12088B (holo¬
type, B destroyed; lectotype, here designated, MO; iso¬
lectotype, A).

Ardisia tonkinensis A. DC., Repert. Spec. Nov. Regni Veg. 8:
354. 1910. Syn. nov. SYNTYPES: Vietnam. Tonkin:
Kirn kibe. in fields of Dong Ham & Dong Bau, H. Bon
2142. 2541. 2912 (G, P); Vo-Xa on Mt. Chuir-Hac, H.
Bon 2902 (G, P). Lectotypification of this binomial will
be made after further study of all duplicates.

Ardisia rectangularis Hyata, J. Coll. Sci. Univ. Tokyo 30: 182.
1911. Syn. nov. SYNTYPES: Japan. Shintiku: Gozhi-
zann, June 1906, T. Kawakuimi & U. Mori 1435 (TI);
Kusshaku, 1905, S. Nagasawa 358 (TT); Taito: Muri-
murisha, Dec. 1899, K. Miyake s.n. (TI). This binomial
will be lectotypified when all duplicates of the collec¬
tions cited in the protologue are examined.

Ardisia stellifera Pitard, Fl. Indo-Chine 3: 863. 1930. Syn.
nov. TYPE: [Vietnam]. Annam: Thua-tien, Thuy-cam,
Eberhardt s.n. (holotype, P; fragment. A).

Ardisia virens Kurz var. annamensis Pitard, Fl. Indo-Chine 3:
868. 1930. Syn. nov. TYPE: (Vietnam], Annam: Quang-
tri: Dent du Tigre, 800 m, Poilane s.n. (holotype, P;
fragment, A).

Ardisia maculosa Mez var. symplocifolia C. Chen, Act. Phy¬
totax. Sin. 16(3): 79. 1978. Syn. nov. TYPE: China.
Yunnan: Hokao, S. Hsuan 610078 (holotype, KUN).

Ardisia virens is closely related to A. elegantissima
H. Leveille. The separation of the two is based on the
presence or absence of papillae on the inflorescence
rachises and branchlets. Ardisia virens grows in dense
evergreen broad-leaf forests, hillsides, dark damp
places, humus-rich soils in China (Guangxi, Guizhou,
Hainan, Taiwan, Yunnan), India, Myanmar, Thailand,
Vietnam, Kalimantan, Brunei, and Sabah.

Myrsine L.

As here delimited, Myrsine L. (including Rapanea
Aublet) is a pantropical genus of about 300 species,
of which 11 grow in China. Recent studies of tropical
American taxa (Pipoly, 1991, 1992a, b, c) have shown
that many taxa have staminate flowers with anthers
on filaments, while the pistillate flowers have the fil¬
aments reduced or barely detectable. Pipoly (1992a)
provided a complete rationale for including Rapanea
as a synonym of Myrsine. Our treatment of Myrsine to
include species formerly placed in Rapanea requires
several new combinations.

Myrsine cicatricosa (C. Chen) Pipoly & C. Chen,
comb. nov. Basionym: Rapanea cicatricosa C. Y.
Wu & C. Chen ex C. Chen, Fl. Yunnan. 1: 381.
1977. TYPE: China. Vietnam, Shaba, Huan-
glianshan, Sino-Vietnamese Expedition 55 (holo¬
type, KUN).

Myrsine verruculosa (C. Chen) Pipoly & C. Chen,
comb. nov. Basionym: Rapanea verruculosa C. Y.
Wu ex C. Chen, Fl. Yunnan. 1: 380. 1977.
TYPE: China. Yunnan: Xichou Xian, 26 Sep.
1947, K. M. Feng 12062 (holotype, KUN).

Myrsine faberi (Mez) Pipoly & C. Chen, comb. nov.
Basionym: Rapanea faberi Mez in Engler, Pflan¬
zenr. 9(IV. 236): 358. 1902. SYNTYPES: China.
Yunnan: Mentze, 2000 m, A. Henry 9173A (B
destroyed, W); China. Guangdong: Faber 132 (B
destroyed; lectotype, here designated, US; iso¬
lectotype, W).

Volume 5, Number 4
1995

Pipoly & Chen
Myrsinaceae of China

361

Myrsine kwangsiensis (E. H. Walker) Pipoly & C.
Chen, comb. nov. Basionym: Rapanea kwang-
siensis E. H. Walker, J. Wash. Acad. Sci. 21: 479.
1931. TYPE: China. Guangxi: Lanlow, E Linyen,
Ching 6657 (holotype, US; isotype, A).

Rapanea kwangsiensis E. H. Walker var. lanceolata C. Y. Wu
& C. Chen, FI. Yunnan. 1: 383. 1977. Syn. nov. TYPE:
China. Yunnan: Jinping Xian, 8 Jan. 1933, H. T. Tsai
52550 (holotype, KUN).

Acknowledgments. Our studies for the Flora of
China have been greatly facilitated by the Missouri
Botanical Garden. We thank Ihsan Al-Shehbaz, An¬
thony Brach, and Michael Gilbert for assistance. We
also thank the curators and staff of the Harvard Uni¬
versity Herbaria, particularly Emily Wood and David
Boufford, and those of the Smithsonian Institution,
particularly George Russell, Vicki Funk, and Larry
Skog, for hospitality and help during our visits to
those institutions.

Literature Cited

Pipoly, J. 1983. Contributions toward a monograph of Cy-
bianthus (Myrsinaceae): III. A revision of subgenus Laxi-
ftorus. Brittonia 35: 61-80.

-. 1987. A systematic revision of the genus Cybian-

thus subgenus Grammadenia (Myrsinaceae). Mem. New
York Bot. Gard. 43: 1—76.

-. 1991. Systematic studies in the genus Myrsine L.

(Myrsinaceae) in Guayana. Novon 1: 204—210.

-. 1992a. A further note on Myrsine perpauciflora Pi¬
poly (Myrsinaceae). Novon 2: 176.

-. 1992b. Estudios en el g&iero Myrsine (Myrsina¬
ceae) de Colombia. Caldasia 17(1): 3—9.

-. 1992c. Notes on the genus Myrsine (Myrsinaceae)

in Peru. Novon 2: 392-^107.

Prance, G. 1972. Chrysobalanaceae. Flora Neotropica 9:
1-410.

-. 1982. Forest refuges: Evidence from woody an-

giosperms. Pp. 137-156 in G. Prance (editor), Biolog¬
ical Diversification in the Tropics. Columbia Univ.
Press, New York.

Walker, E. H. 1940. A revision of the Eastern Asiatic
Myrsinaceae. Philipp. J. Sci. 73: 1-258.

White, F. 1962. Geographic variation and speciation in
Africa with particular reference to Diospyros. Pp. 71-
103 in D. Nichols (editor). Taxonomy and Geography.
The Systematics Association Publ. No. 4, London.

New Andean Zanthoxylum (Rutaceae) with Distinctive
Vegetative Characters

Carlos Reynel

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ABSTRACT. Three new Andean species of Zan¬
thoxylum (= Fagara), Z. gentryi Reynel, Z. lenti-
culare Reynel, and Z. lepidopteriphilum Reynel, are
described and illustrated. Vegetative characters of
taxonomic and practical identification value are
briefly discussed. Some of these features relate to
insect—plant interactions.

RESUMEN. Tres nuevas especies de Zanthoxylum (=
Fagara ), Z. gentryi Reynel, Z. lenticulare Reynel y
Z. lepidopteriphilum Reynel, son descritas e ilustra-
das. Se discuten brevemente caracteres vegetativos de
utilidad taxonomica y prdctica en la ideritificaeion de
especies en el genera. Algunos de estos caracteres
tienen relacidn con interacciones insecto-planta.

The mainly pantropical and subtropical genus
Zanthoxylum L. s.l. comprises about 200 woody
species, mostly trees. Engler (1874, 1896, 1931)
regarded Zanthoxylum s. str. as separate from Fa¬
gara L. and proposed a sectional and subsectional
scheme for the latter. Waterman (1975a, 1976)
combined the two genera and transferred the spe¬
cies from Fagara to Zanthoxylum s.l.; evidence has
accumulated in recent years supporting that unifi¬
cation, on the grounds of floral morphology (Bri-
zicky, 1962; Hartley, 1966; Beurton, 1994), floral
anatomy (Yamazaki, 1988), wood anatomy (Reynel
& Miller, in prep.), and phytochemistry (summa¬
rized in Waterman, 1975b, 1977, 1983).

In this paper I describe three new Andean species
of Zanthoxylum that have distinctive vegetative fea¬
tures: interspaced, short zones of clustered leaf and
bud-scale scars (Z. gentryi. Fig. 1A, 2B); hollow
myrmecophilous apical branchlets (7. lenticulare,
Fig. IB); and twice-revolute basal leaflet margins (Z.
lepidopteriphilum, Fig. 2A, 2C). These new species
belong to the group of New World species with 5-
merous, differentiated perianths placed in Fagara
subsection Paniculatae-Neogaeae by Engler (1896).

Zanthoxylum gentryi Reynel, sp. nov. TYPE: Co¬
lombia. Valle: Municipio Restrepo, Vereda de
Riogrande, abajo del puente, en predios de la
familia Reyna, en la via Pavas-Restrepo, 1200
m, 10 Feb. 1990 (fr), J. Ramos & C. Reyna 2551
(holotype, MO; isotype, CUVC). Figure 1A, 2B.

Novon 5: 362-367. 1995.

Haec species Z. petiolari A. St.-Hilaire & Tulasne af-
finis, sed ab eo foliis 15—45 cm longis, interjugis 3.5—9
cm longis, foliolis lateralibus ovatis vel oblongis, sessili-
bus, (5-)7—17 cm longis, marginibus integris, panicula
ampla, 7-20 X 6—16 cm (infructescentia usque ad 25 X
35 cm); petalis 2.6-3.5 mm longis, coccis 4—7 mm longis
et seminibus 4—6 mm longis differt.

Dioecious treelets or trees to 10 m tall; apical
branchlets 5-10 mm thick, with interspaced, short
zones of clustered leaf and bud-scale scars, also
with sparse, straight to incurved, compressed-con¬
ical spines to 6 mm long and sparse lenticels to 1
mm long, the branchlets covered with occasional
white simple hairs to 0.1 mm long, the periderm
usually peeling when dry. Leaves paripinnate or im-
paripinnate, 4—7(-9)-foliolate, 15-45 cm long, the
terminal leaflet in imparipinnate leaves promptly
deciduous; petiole 4—13 cm long; rachis terete, 2-
3 mm thick, glabrescent, the interjuga 3.5-9 cm
long, sometimes with an abaxial spine to 5 mm
long; leaflets chartaceous, rigid, usually drying ol¬
ive green and shiny on both sides, laterals opposite,
sessile, terminal leaflet with petiolule 0—3 cm long,
ultimate pair usually larger than rest, all leaflets
ovate to oblong, (5—)7—17 X 3.5—9.5 cm, the apex
acuminate, with acumen to 10 X 6 mm, base acute
to obtuse or truncate; margin entire or subentire;
leaflets glabrescent; midvein adaxially prominu-
lous, secondary veins 10—16 pairs, prominulous on
both sides, tertiary veins subreticulate. Inflores¬
cences terminal or subterminal, multiflorous pani¬
cles 7-20 X 6-16 cm (-25 X 35 cm in fruit),
branched 3-4 times; main axis 2—3 mm thick (-4
mm in fruit), secondary axes to 3—5 cm long (—20
cm in fruit), mostly forming a 60°-90° angle with
main axis, the inflorescence axes covered with
white, sparse simple hairs to 0.1 mm long; inflo¬
rescence bracts ovate to deltoid, 0.6-1 (-3) X 0.7—
1.2(—2) mm, glabrescent. Flowers 5-merous; floral
bracts 1-3 at base of pedicel, deltoid to ovate, 0.6-
1 X 0.5-1 mm, covered with indument similar to
inflorescence axes or glabrescent; pedicel 0-1 X
1.2—2 mm (-4 mm long in fruit); calyx 0.5—1.4 X
1—3 mm, sepals free or connate to Vi their total
length, ovate to deltoid, 0.8-1.2 mm broad, acute,
glabrescent; petals saccate, ovate, 2.6-3.5 X 1.2—

Volume 5, Number 4
1995

Reynel

New Andean Zanthoxylum

363

Figure 1. -A. Zanthoxylum gentryi Reynel, apical branchlet with leaf and inflorescence (Ramos 2334). -B. Zan¬
thoxylum lenticulare Reynel, apical branchlet with leaf and infructescence (Soeyarto et al. 4058).

1 .4 mm, the apex rounded to acute, strongly in-
flexed, usually with an ellipsoid resin gland to 0.2
mm diam., the petals glabrous. Staminate flowers
with 5 stamens, exserted Vy-Vi their total length;
filaments slender, 2.8-4 mm long, anthers 1.2—1.8

mm long, thecae oblong, the connective with an
apical ellipsoid gland 0.2-0.3 mm long; nectary
discoid to conical-truncate, 5-lobate, 0.4—0.5 X 1-
1.3 mm; pistillodes 1-3, distinct to totally connate,
elongate-pyriform, 0.7—1.5 X 0.2—0.5 mm. Pistil-

364

Novon

late flowers with 0—5 staminodes, membranous,
ovate to oblong; gynophore conical, 1.2—1.8 X 0.7—
1.6 mm; carpels 3—5, free except in the common
stigma, ovaries obovoid, 0.5—0.8 X 1—1.5 mm,
styles attached at adaxial upper half of ovaries, to
0.2 X 0.2 mm, stigma discoid, 3—5-lobed, 1.8-2.1
mm diam., 0.2-0.4 mm thick. Fruit apocarpous,
(2—)3—4(—5)-follicular, frequently 1—3 follicles
aborting; follicles subovoid, slightly laterally com¬
pressed, 4—7 mm long, glabrescent; seed ellipsoid
to subreniform, 4-6 X 2.5—5 mm, the funicular
scar rounded, 1.5-2 X 0.8-1.2 mm.

Distribution. Endemic to the Department of
Valle in Colombia, at 600-1100 m, in wet primary
or secondary forests.

Within neotropical species of Zanthoxylum with
differentiated perianths, the presence of inter¬
spaced, short zones of clustered leaf and bud-scale
scars on apical branchlets (Fig. 2B) is character¬
istic of a small group: Z. petiolare A. St. Hilaire &
Tulasne (syn. Z. naranjillo Grisebach), a common
species from southern Brazil, Bolivia, Argentina,
and Paraguay; Z. pentandrum (Aublet) R. A. How¬
ard (syn. Z. hermaphroditum Willdenow) from
northern Brazil, Venezuela, and the Guianas; Z.
verrucosum (Cuatrecasas) P. G. Waterman from Co¬
lombia; and Z. pucro from Panama. Zanthoxylum,
gentryi can be distinguished from the species of the
Zanthoxylum petiolare group by the following key:

Key to Species in the Zanthoxylum petiolare Group

la. Leaves with lateral leaflets sessile. Z. gentryi

lb. Leaves with lateral leaflets petiolulate, petiolules
2—10 mm long.

2a. Interjuga 5-12 cm long; leaflets 12-22 cm
long.

3a. Leaves imparipinnate, with ovate leaf¬
lets; inflorescence axes glabrous; pistil¬
late flowers with petals 3-3.5 mm long;
follicles 5-6 mm long .... Z. verrucosum
3b. Leaves paripinnate, with oblong leaflets;
inflorescence axes densely and finely
pubescent; pistillate flowers with petals
1.7-2.4 mm long; follicles 6-9 mm

long. Z. pucro

2b. Interjuga 1.5-5 cm long; leaflets 3—12(—16)
cm long.

4a. Petals 1.3—1.7 mm long; anthers 0.4—

0.6 mm long. Z. pentandrum

4b. Petals 1.9—2.2 mm long; anthers 0.9-

1.1 mm long.Z. petiolare

The new species is named after Alwyn H. Gentry
(1947—1993), botanist and ecologist, whose work and
knowledge of the Andean-Amazonian flora and its
diversity inspired a generation of tropical biologists.

Paratypes. COLOMBIA. Valle: Municipio Toro, que-
brada San Lazaro (fr), Devia 328 (MO, T1JLV); near Yo-

toco, N of Cali, 3°45'N, 76°30'W, Gentry et al. 54073
(COL, MO); El Silencio, Hacienda Himalaya, W of Yum-
bo, 3°50'N, 76°40'W (fr). Gentry et al. 65516 (COL, MO);
Finca El Espinal, 25 km N of Cali (fr), Hughes 129 (FHO,
MO); Dagua Valley, 15.5 km below Dagua and 1.5 km
below Lobo Guerrero, quebrada de la Chapa, upper end
El Boquerfin of rfo Dagua (fl), Hutchinson et al. 3102
(COL, F, G, K, MO, NY, UC, US); Municipio Restrepo,
Ramos 2334 ( 5), 2281 (d) (MO, CUVC); Municipio El
Cerrito, pueblo de Guabas, cerca de la frontera con Mun¬
icipio Guacari, Silverstone-Sopkin et al. 2294, 3662 (fr)
(CUVC, MO).

Zanthoxylum lenticulare Reynel, sp. nov. TYPE:
Colombia. Antioquia: Municipio de Anorf,
Corregimiento de Providencia, entre Dos Bo-
cas y Anorf, Buenos Aires, 4 km from Provi¬
dencia, 400-900 m, aprox. 7°N, 75°W, May
1973 (fr), Soejarto et al. 4058 (holotype, COL;
isotypes, F, GH, MO). Figure IB.

Haec species Z. compacto (Huber ex Albuquerque) P.
G. Waterman affinis, sed ab eo foliis 40-70 cm longis 13-
21-foliolatis, foliolis 8-12 cm longis, subcaudatis, margi-
nibus minute crenulatis; paniculis amplissimis congestis
15-35 cm longis, 3( -4)-ramificatis, axibus primariis non
fistulosis, axibus secundariis 4—13 cm longis; petalis 1.5-
1.9 mm longis; coccis discoideis laevibus 3-4 mm longis,
seminibus 2.5—3 mm longis differt.

Dioecious, myrmecophilous treelet or tree to 12 m
tall; apical branchlets hollowed, 4—8 mm thick, with
sparse, straight, compressed-conical spines to 6 mm
long and lenticels to 0.5 mm long, the branchlets cov¬
ered with sparse to dense, white, simple hairs to 0.1
mm long or glabrescent. Leaves evenly distributed on
branchlets, imparipinnate, (11—)13—21(—25)-foliolate,
40-70 cm long; petiole (5.5—)7-10(—15) cm long; ra-
chis terete, narrowly and deeply canaliculate, 2-4 mm
thick, interjuga (2.5-)3-4.5 cm long, sometimes with
1(2) abaxial spines to 5 mm long, the rachis covered
with indument similar to the branchlets; leaflets char-
taceous to submembranaceous, drying pale green to
reddish brown above, pale green beneath, laterals op¬
posite to alternate, sessile, terminal leaflet with peti-
olule 2—10 mm long, ultimate pair or terminal leaflet
usually larger than rest, all leaflets oblong to subovate,
(2.5—)8—12(—15) X 2-4.5 cm, the apex acuminate to
caudate, the acumen to 15 X 5 mm, base acute; mar¬
gin crenate, the teeth separated 1—1.5 mm at the mid¬
dle of leaflets; leaflets without hairs; midvein adaxially
prominulous; few rounded glands 0.2-0.3 mm diam.
usually present on abaxial blade near midvein; sec¬
ondary veins 16-22 pairs, prominulous on both sides,
tertiary veins usually prominulous and subreticulate
on both sides. Inflorescences terminal or subterminal,
multiflorous, congested panicles (8-)15-35 X (8-)15-
22 cm, branched 3(-4) times; main axis 2-4 mm
thick, secondary axes 4—13 cm long, mostly forming

Volume 5, Number 4
1995

Reynel

New Andean Zanthoxylum

365

a 60°-90° angle with main axis, the inflorescence axes
covered with sparse to dense, white, simple hairs to
0.05(-0.1) mm long or glabrescent; inflorescence
bracts deltoid to ovate, 0.2-0.5(-l) X 0.3-0.5 mm,
covered with indument similar to inflorescence axes,
usually ciliate. Flowers 5-merous; floral bracts (1)2—5
at base of pedicel, ovate to lanceolate, 2-3 X 1 mm,
covered with indument similar to inflorescence axes
or glabrescent; pedicel terete, 0.2-0.7 X 0.1-0.2 mm
(—2.5 X 0.5 mm in fruit), glabrescent; calyx 0.5-0.7
X 0.&-1 mm, sepals connate Vi their total length,
ovate, 0.3-0.4 mm broad, rounded to acute, glabres¬
cent; petals saccate, ovate, 1.5—1.9 X 0.8-1.1 mm,
the apex rounded, usually apiculate with apiculum to
0.2 mm long, also with an ellipsoid gland to 0.2 mm
diam., the petals glabrous. Slaminate flowers with sta¬
mens 5, exserted Vs to Vi their length, the filaments
slender, 2—3 mm long, anthers 0.8-1 mm long, thecae
oblong; nectary conical-truncate, 5-lobate, 0.3-0.4 X
0.3-0.6 mm; pistillodes 1-3, elongate-pyriform, dis¬
tinct to totally connate, 0.2-0.5 X 0.2 mm. Pistillate
flowers with no staminodes; gynophore conical, 0.2-
0.3 X 0.4—0.6 mm; carpels (2)3, distinct except in
the common stigma, ovaries oblongoid, 0.4—0.6 X
0.4—0.6 mm; styles terete, attached at adaxial upper
half of ovaries, 0.1-0.2 X 0.1-0.2 mm; stigma dis¬
coid. Fruit apocarpous, (2-)3-follicular, frequently 1-
2 follicles aborting; follicles discoid, 3-4 X 3 mm,
glabrous; seed subdiscoid, 2.5-3 X 0.6-0.9 mm, the
funicular scar linear, 1—2 X 0.3-0.4 mm. Common
name: “Sando.”

Distribution. Colombia (Antioquia) and Vene¬
zuela (TAchira and Zulia), between 90 and 900 m,
in wet primary forests.

Three other New World species of Zanthoxylum
also have hollow, myrmecophilous apical branch-
lets, which are very consistent and of practical val¬
ue for identification. The following key allows the
comparison of related species.

Key to Myrmecophilous Neotropical Species of Zan-
thoxylum

la. Apical branchlets with enlarged, interspaced
zones of clustered leaf scars; panicles clustered

at the apex of branchlets . . Zanthoxylum setulosum

lb. Apical branchlets uniformly terete; panicles sol¬
itary, terminal.

2a. leaves velutinous-pubescent; leaflets 5-12

cm broad. Zanthoxylum formiciferum

2b. leaves glabrous or glabrescent; leaflets 1—5
cm broad.

3a. Panicles compact, 3—10 cm broad; fruit
with follicles ovoid to obovoid, 6-9 mm

long. Zanthoxylum compactum

3a. Panicles expanded, 15-22 cm broad;
fruit with follicles discoid, 4—4.5 mm
long. Zanthoxylum lenticulare

Paratypes. COLOMBIA. Antioquia: Municipio San
Luis, carton del rfo Claro (3), Cogollo 831 (COL, MO);
Municipio lani/a. Rfo Cauca, right margin on road from El
Doce to Barro Blanco, 8°35'N, 75°25'W (fr), Daly et al.
5253 (COL, F, NY); Rio Claro, camino al Refugio (9, fr),
Renteria et al. 2855 (COL, MO); Municipio Anori, Rio An-
ori valley, near Planta Providencia, 7°30'N, 74°50'W, Shep¬
herd 374 (COL, MO, WIS), 6 Aug. 1977, 913 (COL, MO,
WIS); Corregimiento de providencia, entre Dos Bocas y An¬
ori, Buenos Aires, ca. 7°N, 75°W (3), Soejarto 3922 (COL,
F(2), MO). VENEZUELA. Carabobo: vicinity of Las
Trincheras, near Valencia, H. Pittier 7655 (GH). Tachira:
ca. 10.5 km NE of La Fria, 90 m, 8°16'N, 72‘TO'W, 16
Nov. 1979 (3), J. Steyermark et al. 120511 (MO, NY, VEN).
Zulia: Distrito of Coldn, aldea Querrequerre, 12—15 km N
of Casigua (fr). Bunting et al. 7427 (NY, VEN).

Zanthoxylum lepidopteriphilum Reynel, sp. nov.
TYPE: Peru. Piura: Prov. Huancabamba, Canc-
haque, between “Chorro Blanco” and “War-
war,” 2000-2500 m, 18 Jan. 1989 (fr), C. Diaz,
T. D. Pennington & C. Reynel 3192 (holotype,
MO; isotypes, K, MOL, USM). Figure 2A, C.

Haec species Z. mantaro (J. F. Macbride) J. F. Mac-
bride affinis, sed ab eo foliis 32 45 cm longis, foliolis 9—
17(—25), 7— 10(—13) cm longis, basibus abaxialibus bis re-
volutis cylindrum marginalem formantibus 8-10(—14) mm
longis; panicula ampla 9—14 cm longa (infructescentia
usque ad 20 cm longa); petalis 2—3 mm longis; coccis 5-
8(-10) mm longis, seminibus 5-6 mm longis differt.

Dioecious trees to 20 m tall; apical branchlets
terete, 7-10 mm thick, unarmed, covered with
sparse to dense, white or yellow, stellate, bifid and
simple hairs 0.2-0.6 mm long. Leaves evenly dis¬
tributed on branchlets, imparipinnate, 32—45 cm
long, 9-17(-25)-foliolate; petiole (3.5-)4—10 cm
long; rachis terete, obscurely canaliculate, 2-4 mm
thick, interjuga 3-5 cm long, the rachis covered with
indument similar to branchlets or glabrescent; peti-
olules 3-5 mm long; leaflets chartaceous, drying ol¬
ive green to reddish brown on both sides, laterals
opposite, ultimate or medial pair larger than rest, all
leaflets oblong, 7—10(—13) X 2.5—3(—3.5) cm, the
apex emarginate, rounded or acuminate, with acu¬
men to 6 X 4 mm, base acute, oblique in laterals,
tubular-twice-revolute 8-10(-14) mm or more along
margin, each side of the midvein; margin entire to
slightly crenate, with teeth separated 2-3 mm at the
middle of leaflets; leaflets covered with sparse to
dense, white to yellow, stellate, bifid and simple
hairs to 0.2-0.5 mm especially along midvein and
abaxial base, or glabrescent; midvein adaxially nar¬
rowly and deeply impressed; secondary veins 12-16
pairs, prominulous on both sides, tertiary veins usu¬
ally prominulous and subreticulate on both sides.
Inflorescences terminal or subterminal panicles,
branched 1-2 times, pauciflorous to submultiflorous,
9-14 X 2.S-5.5 cm (-20 X 20 cm in fruit); main

366

Novon

3 cm

-B r zLwfrtt7 /U R R ^ el - a P' ca ^ branchlet with leaf and infructescence (Diaz et al. 3192).

(sLfs“^SoaZ!tai^2^i n ^r aP 7 ! )ra,, ; et , Sh r ng short zones of clus '™ d «"<! bud-scale sears (arrows)

low™. f“£r5tS7"“" “»• h ™"' *■ b -' ™*i"

Volume 5, Number 4
1995

Reynel

New Andean Zanthoxytum

367

axis 2-3 mm thick (—5 mm in fruit), secondary axes
1—2 cm long (-6 cm in fruit), the inflorescence axes
covered with sparse to dense, stellate, bifid and sim¬
ple hairs 0.2-0.8 mm long; inflorescence bracts del¬
toid to lanceolate or ovate, 0.5-1.3 X 0.5-0.7 mm,
covered with indument similar to inflorescence axes.
Flowers 5-merous; floral bracts l(-3) at base of ped¬
icel, ovate to deltoid, 0.4-0.6 X 0.3-0.6 mm, with
indument similar to inflorescence axes; pedicel sub-
terete, 0.6—1.2 X 0.6-0.8 mm; calyx with sepals ba-
sally connate to Vs their total length, ovate to deltoid,
0.8-1.2 X 0.5-0.8 mm, covered with sparse, stellate,
bifid and simple hairs 0.3-0.5 mm long, usually cil-
iate; petals oblong to ovate, 2-3 X 1.2-1.6 mm, sac¬
cate, the apex inflexed, rounded, usually apiculate,
with apiculum 0.2—0.4 mm long, petals covered with
scattered, stellate, bifid or simple hairs 0.2-0.4 mm
long, subciliate. Staminate flowers with 5 stamens,
exserted to Vs their total length, filaments slender,
2.5—3.8 mm long, anthers oblongoid, 0.8-1 mm long,
the connective with an obscure apical gland 0.1-0.2
mm diam.; nectary conical-truncate to discoid, ba-
sally 5-lobed, 0.4-0.6 X 0.9-1.2 mm; pistillodes 2-
3, distinct to connate, pyriform-elongate, 0.6-0.9 X
0.3-0.4 mm. Pistillate flowers with staminodes 0-5,
membranous, ovate to oblong, 0.4-0.6 X 0.2-0.3
mm; gynophore conical, 0.6-1.2 X 0.7—1.4 mm; car¬
pels 3—5, free except in the common stigma, ovaries
obovoid, 0.7—1.3 X 0.8-1.3 mm; styles attached at
adaxial upper half of ovaries, 0.3-0.5 X 0.2-0.3
mm, stigma discoid, 3—5-lobed, 1.5-2.1 mm diam.,
0.2-0.4 mm thick. Fruit apocarpous, 2-3(-5)-foli-
cular, frequently 1-3 follicles aborting; pedicel 3-5
X 1.7—2.3 mm, with indument similar to inflores¬
cence axes; follicles globose to obovate-compressed,
5-8(-10) mm long, glabrescent; seed globose to
ovoid-compressed, 5-6 X 4—5 mm, the funicular
scar linear, 4—5 X 0.4—0.7 mm.

Distribution. Ecuador (Imbabura, Loja, Pichin-
cha) and Peru (Piura, Cajamarca), 1000-2500 m,
in montane cloud forests.

The New World species of Zanthoxylum with dif¬
ferentiated and 5-merous perianths, placed in Fa-
gara subsection Paniculatae-Neogaeae by Engler
(1896), frequently have lateral leaflets with basally
slightly to strongly revolute margins. Zanthoxylum
lepidopteriphilum is unique in having the basal
margin twice-revolute, forming a conspicuous cyl¬
inder 8-10(-14) mm long along the margin on each
side of the midvein (Fig. 2C). The larvae of an un¬
identified lepidopteran have been found to inhabit
the interior of this marginally formed cylinder. Zan¬
thoxylum lepidopteriphilum is similar to Z. mantaro
(J. F. Macbride) J. F. Macbride in its apical bran-

chlets and leaves with indument of simple, bifid
and stellate hairs and 5-merous, 3-5-carpellate
flowers, but differs in its markedly larger terminal
buds, leaves, leaflets, and inflorescences; sizes of
these organs for both species compare as follows
(first measurement for Z. lepidopteriphilum, second
for Z. mantaro): leaves 32-45 cm long vs. 9-15 cm
long; leaves 9-17(-25)-foliolate vs. 5-7(-9)-folio-
late; petioles 4—10 cm long vs. 1.5-3.5 cm long;
leaflets 7-10(-13) cm long vs. (2-)3-6(-9) cm long;
and inflorescences 9-14 cm long (-20 cm in fruit)
vs. (2.5-)4—6 cm long (-9 cm in fruit).

Paratypes. PERU. Cajamarca: San Miguel, Distrito La
Florida, Agua Azul (fr), Diaz 2095 (MO, MOL, TEX,
USM). Piura: Cerro Aypate, 49 k E of Ayabaca, in dis¬
turbed dry cloud forest, 4°35'S, 79°32'W, Gentry et al.
75045 (MO, USM). ECUADOR. Imbabura: Cantdn Ur-
cuquf, Balneario Cachimbio, 0°27’N, 78°14W (d), W. Pa¬
lacios 5241 (MO, QCA). Loja: Cant6n Sozoranga, slope
of San Fernando, exiting Las Cochas, 4°18'S, 79°57’W
(fr), Palacios 3321 (MO, QCA). Pichincha: Reserva Geo-
botdnica Pululahua, between La Reventazdn and Pulula-
hua, 0°05'N, 78°30'W, (9, fr) C. Ceron et al. 5673 (MO,
QCA).

Acknowledgments. I thank R. Gereau, J. Hunt,
P. M. Richardson, and C. M. Taylor for their critical
reading of the manuscript. Funds from the Andrew
W. Mellon Foundation made this work possible.

Literature Cited

Beurton, C. 1994. Gynoecium and perianth in Zanthox¬
ylum s. 1. (Rutaceae). PI. Syst. Evol. 189: 165-191.
Brizicky, G. 1962. Taxonomical and nomenclatural notes
on Zanthoxylum and Glycosmis (Rutaceae). J. Arnold
Arbor. 43: 80-93.

Engler, A. 1874. Rutaceae. In: C. Martius, Flora Bras-
iliensis 12(2): 7-196.

-. 1896. Rutaceae. In: A. Engler & K. Prantl, Die

Natiirlichen Pflanzenfamilien 3(4): 114—119.

-. 1931. Rutaceae. In: A. Engler & K. Prantl, Die

NatUrlichen Pflanzenfamilien, Ed. 2. 19a: 214—224.
Hartley, T. 1966. A revision of the Malesian species of
Zanthoxylum. J. Arnold Arbor. 47: 171—221.

Waterman, P. G. 1975a. New combinations in Zanthox¬
ylum L. Taxon 24: 361—366.

-. 1975b. Alkaloids of the Rutaceae: Their distri¬
bution and systematic significance. Biochem. Syst.
Ecol. 3: 149-180.

-. 1976. Further new combinations in Zanthoxy¬
lum L. Taxon 25: 594.

-. 1977. A review of the chemosystematics of the

genus Zanthoxylum L. (Rutaceae). Pp. 109-129 in B.
Stone (editor), The Role and Goals of Tropical Botanic
Gardens. Univ. of Malaya, Kuala Lumpur.

-. 1983. Phylogenetic implications of the distri¬
bution of secondary metabolites within the Rutales. Pp.
377-400 in P. G. Waterman & M. Grundon (editors).
Chemistry and Chemical Taxonomy of the Rutales. Ac¬
ademic Press, London.

Yamazaki, T. 1988. Floral anatomy of the genus Zan¬
thoxylum L. J. Jap. Bot. 63: 7—16.

New Combinations and a New Variety in Mesoamerican Dalbergia
(Fabaceae: Papilionoideae)

Velva E. Rudd

California State University, Northridge, California 91330, U.S.A.

Abstract. Six new combinations with change of
status are proposed to reflect the interrelationship
as well as the singularity of the taxa: Dalbergia
glabra var. chontalensis, D. glabra var. paucifolio-
lata, D. retusa var. cuscatlanica, D. retusa var. hy¬
poleuca, D. retusa var. lineata, and D. retusa var.
pacifica. One variety, D. calderonii var. molinae, is
described as new.

Most species of Mesoamerican Dalbergia were
originally described from one or a few specimens.
As more material has become available gradation
of characters has become evident. Many of the dif¬
ferences between taxa are subtle, and there is con¬
siderable intergradation. In some cases reduction
in rank appears to be warranted; in one, differences
are deemed worthy of recognition as a new variety.

Dalbergia glabra (Miller) Standley var. chontal¬
ensis (Standley & Williams) Rudd, comb, et
stat. nov. Basionym: Dalbergia chontalensis
Standley & Williams, Ceiba 1: 81. 1950.
TYPE: Nicaragua. Chontales: vecindad de Jui-
galpa, en matorrales de una cuesta rocosa, alt.
160 m, 4—13 June 1947, Standley 9296 (ho-
lotype, F).

Dalbergia glabra (Miller) Standley var. paucifo-
Uolata (Lundell) Rudd, comb, et stat. nov.
Basionym: Dalbergia paucifoliolata Lundell,
Contr. Univ. Michigan Herb. 4: 8. 1940. TYPE:
Mexico. Tabasco: Reforma on the Rio San
Marti r above Balancan, on river bank, 22—26
May 1939, Matuda 3167 (holotype, MICH).

Dalbergia retusa Hemsley var. cuscatlanica

(Standley) Rudd, comb, et stat. nov. Basionym:
Amerimnon cuscatlanicum Standley, J. Wash.
Acad. Sci. 13: 442. 1923. TYPE: El Salvador.
San Salvador: 1923, Calderdn 1557 (holotype,
US).

Dalbergia retusa Hemsley var. hypoleuca (Pittier)
Rudd, comb, et stat. nov. Basionym: Dalbergia
hypoleuca Pittier, J. Wash. Acad. Sci. 12: 62.
1922. TYPE: Costa Rica. Alajuela: El Escobal,
near Atenas, Golcher 1747 (holotype, US, orig¬
inally in John Donnell Smith herbarium).

Dalbergia retusa Hemsley var. lineata (Pittier)
Rudd, comb, et stat. nov. Basionym: Dalbergia
lineata Pittier, J. Wash. Acad. Sci. 12: 63.
1922. Amerimnon lineatum (Pittier) Standley,
J. Wash. Acad. Sci. 13: 442. 1923. TYPE:
Costa Rica. Guanacaste: Nicoya, Apr. 1900,
Tonduz 13969 (holotype, US).

Dalbergia retusa Hemsley var. pacifica (Standley
& Steyermark) Rudd, comb, et stat. nov. Bas¬
ionym: Dalbergia pacifica Standley & Steyer¬
mark, Field Mus. Nat. Hist., Bot. Ser. 22: 236.
1940. TYPE: Guatemala. Suchitepdquez: Co-
cales, roadside, 215 m, 5 Jan. 1939, Standley
62109 (holotype, F).

Dalbergia calderonii Standley var. molinae
Rudd, var. nov. TYPE: Honduras. Morazan:
floresta de nubes area Rancho Quemado, sur-
oeste de San Juancito, 2000 m, 19 June 1948,
Molina 1040 (holotype, F).

Arbores nubiculae a varietate typica foliis plerumque
longis, foliolis parviis numerosisque differunt.

Trees 4—8 m high. Leaves (11—)13—15-foliolate,
the axis about 15-20 cm long. Leaflets 4—6 X 1.5-
2.5 cm, elliptic, obtuse to acute. Flowers cream to
orange-yellow, 4.5—5 mm long. Fruit brown-veluti-
nous, 7—8 X 1.5 cm, the stipe about 8-10 mm long.

Dalbergia calderonii var. calderonii occurs at
lower elevations and different habitat than variety
molinae (350-850 m in low to medium deciduous
forest vs. ca. 1300-2100 m at the edge of cloud
forest); it bears leaves with fewer leaflets ((3—)5-
9(— 1 l)-foliolate, with an axis 6-10 cm long vs. (11—)
13—15-foliolate, with an axis ca. 15—20 cm long);
and its fruits have a shorter stipe (ca. 5 mm long
vs. ca. 8-10 mm long).

Paratypes. HONDURAS. Morazan: bosque de nu¬
bes, entre Santa Lucfa y Valle de Angeles, 1800 m, Mo¬
lina 3301 (F); Montana La Tigre, suroeste de San Juancito,
2100 m, Molina 10661 (F); edge of cloud forest, moun¬
tains above San Juancito, 2000 m, Williams & Molina
13328 (F). NICARAGUA. Jinotega: bosque de la entrada
a Aranguez, 1300-1400 m, Araquistain 3640 (MO). Ma-
tagalpa: road to Aranguez, pasture and patches of cloud
forest, 1360—1380 m, Stevens et al. 17067 (MO); cloud

Novon 5: 368-369. 1995.

Volume 5, Number 4
1995

Rudd

Mesoamerican Datbergia

369

forest area, Sta. Maria de Ostuma, between Matagalpa and
Jinotega, 1300—1500 m, Williams el al. 23442 (NY).

This variety is named in honor of Antonio Molina
R., Honduran botanist, following a suggestion by P.
H. Standley and L. O. Williams in an unpublished
herbarium name.

Acknowledgment. I am indebted to Joy Nishida
for her help in preparing this manuscript for pub¬
lication.

Three New Species of Clavija (Theophrastaceae)

Bertil Stahl

Department of Botany, Stockholm University, S-106 91 Stockholm, Sweden
Present address: Herbario QCA, Departamento de Ciencias Biologicas,
Pontificia Universidad Catolica del Ecuador,

Apartado 17-01-2184, Quito, Ecuador

Abstract. Three new species of Clavija, C. la-
planadae (Narino, Colombia), C. subandina (Napo,
Ecuador), and C. obtusifolia (Huanuco—San Martin,
Peru), are described and illustrated, and their af¬
finities are briefly discussed.

Since the completion of my revision of Clavija
(St&hl, 1991) many additional collections of the ge¬
nus have been made or brought to my attention. In
this material several undescribed species have
been encountered. The three species described in
this paper were all gathered in premontane or lower
montane Andean forest, the type of habitat richest
in species of Clavija.

Clavija laplanadae StShl, sp. nov. TYPE: Colom¬
bia. Narino: La Planada Nature Reserve, S of
Ricaurte, 1800 m, 78°02'W, 01°10'N, 28 Feb.
1992 (fl), Stahl & Eriksen 652 (holotype, COL;
isotypes, GB, PSO, herbarium of the La Plan¬
ada Nature Reserve). Figure 1.

Arbor parva ad 4 m alta, surculis glabris. Folia pe-
tiolis 7—11 cm longis; lamina oblanceolata vel anguste
oblanceolata, 60—120 X 12—28 cm, absque sclerenchy-
mate subepidermali, basi angustata, apice obtuso vel ro-
tundato, margine serrulato vel integro, venis lateralibus
conspicuis. Racemi ad 45 cm longi; (lores 15-40, 5-
meri. Calyx lobis 2.S-2.7 X 2.5-3 mm. Corolla tubo 4
mm longo, lobis 7.0-8.0 X 7.5-8.5 mm. Ovula 3-10.
Fructus ignotus.

Treelet to 4 m high, unbranched or rarely sparse¬
ly branched, stem to ca. 8 cm diam., young shoots
glabrous, brown or light brown. Leaves arranged in
1, 2, or rarely 3 pseudowhorls toward the summit
of main stem or branches; blade oblanceolate to
narrowly oblanceolate, 60-120 X 12—28 cm, pa¬
pyraceous when dry, 0.25-0.30 mm thick, some¬
what bullate, glabrous, dark green, without subepi-
dermal, extraxyllary sclerenchyma, base narrowly
attenuate, apex rounded or obtuse, often somewhat
retuse, margins entire or sparsely and minutely ser¬
rulate with teeth usually most numerous toward
base, lateral veins conspicuous, particularly be¬
neath, surfaces between veinlets smooth; petiole 7—

11 cm long, 5-8 mm thick, glabrous. Racemes pen¬
dent, to 45 cm long with 15-40 flowers, rachis 2—
3 mm thick at base, glabrous, pedicels 4—5 mm
long, bracts 0.6-0.8 mm long, inserted at the nodes.
Flowers 5-merous, morphologically bisexual (uni¬
sexual flowers not seen); calyx seemingly glabrous,
lobes broadly ovate, 2.5—2.7 X 2.5—3.0 mm, mar¬
gins erose; corolla pale orange, tube 4 mm long,
lobes very broadly ovate or subrotund, 7.0-8.0 X
7.5—8.5 mm. Staminodes ovoid-gibbous, completely
fused with corolla, alternating with the lobes. Sta¬
men filaments glabrous, fused into a tube 1.5 mm
long. Pistil narrowly ovoid, 3.0 X 1.0 mm, stigma

2- lobed; ovules 3-10 in 1 or 2 rows. Fruits not
seen.

This new species, distinguished by its huge
leaves without extraxyllary subepidermal scleren¬
chyma, was collected during fieldwork in southern
Colombia a few years ago. However, at the La Plan¬
ada Field Station it had been known for a long time
as one of the plants exhibited along their nature
trail. It shows clear affinities with Clavija eggersi-
ana Mez, a species of the moist, semideciduous
forests of coastal Ecuador, differing primarily in its
more rigid leaves with some subepidermal scleren¬
chyma and its much smaller flowers.

Clavija laplanadae is so far known solely from
the type locality, but it is quite possible that a pop¬
ulation at the Rio Guajalito Reserve in Prov. Pi-
chincha, Ecuador, should be referred to this spe¬
cies. The extant material from the Rio Guajalito
population is lacking flowers and was tentatively
included in C. eggersiana in my previous treat¬
ments (St4hl, 1990, 1991).

Clavija subandina St&hl, sp. nov. TYPE: Ecuador.
Napo: Canton Archidona, S slopes of Vole An
Sumaco, on trail between El Pacto and Hu-
ahua Sumaco, 1600 m, 00°37'S, 77°35'W, 24
Oct. 1989 (fl), Palacios & Neill 4725 (holo¬
type, MO; isotype, GB). Figure 2.

Arbor parva ad 10 m alta, surculis glabris. Folia petiolis

3- 7 cm longis; lamina oblanceolata, 35-60 X 7—12 cm.

Novon 5: 370-374. 1995.

Volume 5, Number 4
1995

SMhl

Clavija from South America

371

Figure 1. Clavija laplanadae Stahl. —A. Habit. —B. Leaf. —C. Inflorescence. —D. Flower in apical view showing
corolla lobes, staminodes, and anthers. —E. Pedicel and calyx. A, unvouchered, drawn from photographic slides; B-
E, based on Stdhl & Eriksen 652 (GB).

basi angustata, apice acuto vel breviter acuminato, mar-
gine integro alboque, sclerenchymate in fasciculis subep-
idermalibus disposito, fasciculis adaxialibus immersis.
Racemi ad 35 cm longi; flores 30—55, 4-meri. Calyx lobis
3.0-3.5 X 4.0-4.2 mm. Corolla tubo circiter 3 mm longo,
lobis 6-7 X 6-7 mm. Fructus ignotus.

Tree to 10 m high, sparsely branched or un¬
branched, trunk ca. 10 cm diam., young shoots gla¬
brous, brown. Leaf blade oblanceolate, 35—60 X 7—
12 cm, coriaceous, 0.30-0.40 mm thick, glabrous,
light green beneath, dark green above, with sub-

372

Novon

Figure 2. Clavija subandina Sthhl. —A. Part of stem with Leaves. —B. TS of leaf blade, showing immersed adaxial
bundles of sclerenchyma. —C. Inflorescence. —D. Flowers, in bud and at the beginning of anthesis. —E. Staminate
flower with one corolla lobe removed showing stamen tube and staminodes. Based on Palacios & Neill 4725 (MO).

epidermal bundles of extraxyllary sclerenchyma,
the adaxial bundles somewhat immersed into the
mesophyll, base narrowly attenuate, apex acute or
shortly acuminate, margins entire with a rather con¬
spicuous pallid border, lateral veins rather incon¬
spicuous, surfaces between veins sparsely striate;
petiole 3—7 cm long, 3-6 mm thick, glabrous. Ra¬
cemes pendent, to 35 cm long with 30-55 flowers,
rachis ca. 1.5 mm thick, glabrous, pedicels 2-3 mm
long, bracts ca. 1 mm long, inserted at the nodes.
Flowers 4-merous (only functionally staminate flow¬
ers seen); calyx lobes very broadly ovate, 3.0-3.5
X 4.0-4.2 mm, margins erase; corolla orange-red,
tube ca. 3 mm long, lobes suborbicular, 6.0-7.0 X
6.0-7.0 mm. Stamen filaments glabrous, fused into
a tube 2 mm long. Pistil very narrowly ovoid, ca. 3
X 0.5 mm, stigma entire. Fruits not seen.

Clavija subandina seems to be most closely re¬

lated to C. repanda St hill, a species restricted to a
small area of semi-deciduous, lower montane forest
in southwestern Ecuador (Sthhl, 1986). That spe¬
cies differs from C. subandina in its repand leaf
margins and by lacking or having very little ex¬
traxyllary foliar sclerenchyma; C. repanda seems
also to be a smaller and more slender plant than
C. subandina. An interesting leaf anatomical fea¬
ture of C. subandina is the disposition of the ad¬
axial sclerenchyma, which is not located adjacent
to the epidermis as in most other species of the
genus but slightly immersed into the subjacent me¬
sophyll (Fig. 2B). This arrangement of the foliar
sclerenchyma was previously known only in C. ven-
osa Sthhl and C. clavata Decaisne (Sthhl, 1991),
neither of which can be considered closely related
to C. subandina.

Clavija subandina has been collected at about

Volume 5, Number 4
1995

StShl

Clavija from South America

373

1600-1800 m on the southern slopes of Volcan Su-
maco. The discovery of a new species of Clavija at
this elevation in the Ecuadorian Oriente is rather
unexpected. On the eastern side of the Ecuadorian
Andes the genus has previously been recorded only
once from above 1000 m altitude ( C. weberbaueri
Mez, up to 1300 m). However, at least two species
of Clavija are known from localities at or slightly
above 2000 m on the Pacific slopes.

Despite the relatively isolated location of Volcan
Sumaco, its paramo vegetation has been found to
be floristically very similar to other pAramo areas
in Ecuador, including very few endemics, if any
(Lpjtnant & Molau, 1982). The discovery of Clavija
subandina, as well as a recently described species
of Menispermaceae from the same general area
(Bameby, 1990), indicates that additional narrow
endemics can be expected from the premontane
and lower montane forests surrounding Volcan Su¬
maco.

Paratypes. ECUADOR. Napo: the Pacto Area, 40 km
on road from 1 lolh'n to Loreto, 1800 m, 24—25 Aug. 1988
(ster.), Jaramillo 023, 055 (both MO).

Clavija obtusifulia StAhl, sp. nov. TYPE: Peru.
HuAnuco: Prov. Leoncio Prado, Dtto. Hermilio
Valdizan, near La Divisoria, 1500-1600 m, 27
June 1976 (fl & young fr), Schunke 9424 (ho-
lotype, MO). Figure 3.

Frutex ad 1 m altus, surculis puberulis. Folia petiolis
4—5 cm longis; lamina oblanceolata vel elliptica, 30—33
X 10-13 cm, basi angustata, apice obtuso, margine in-
tegro, paginis subtiliter striatis. Racemi 4—6 cm longi, as-
cendentes; flores 6—12, 5-meri. Calyx lobis 3.5-4.0 X
4.0-4.5 mm. Corolla tubo 2.5 mm longo, lobis 5.0 X 5.0—
5.5 mm. Ovula circiter 15. Fructus 2.5—3.5 cm diam.,
seminibus 7-12.

Shrub to 1 m high, young shoots puberulous.
Leaf blade oblaneeolate to elliptic, 30-33 X 10-
13 cm, subcoriaceous, 0.30-0.35 mm thick, gla¬
brous, green above, pale green beneath, with sub-
epidermal bundles of extraxyllary sclerenchyma ad¬
jacent to the epidermis on both sides, base
attenuate, apex obtuse, margins entire, lateral veins
rather inconspicuous, surfaces between veins stri¬
ate; petiole 4—5 cm long, ca. 2 mm thick, glabrous.
Racemes ascending, 4—6 cm long with 6—12 flow¬
ers, rachis 0.5 mm thick, sparsely and minutely
puberulous, pedicels 4—5 mm long, distally thick¬
ened, bracts ca. 0.5 mm long, inserted at the nodes.
Flowers 5-merous, morphologically bisexual (uni¬
sexual flowers not seen); calyx lobes very broadly
ovate, 3.5-4.0 X 4.0-4.5 mm, margins vaguely
erose; corolla orange-red, tube 2.5 mm long, lobes
5.0 X 5.0-5.5 mm. Stamen filaments glabrous.

Figure 3. Clavija obtusifolia Stahl. —A. Branchlet with
leaves and inflorescence. —B. Inflorescence, flowers in
late bud and (third flower from base) without corolla and
stamens. —C. Corolla cut open to show staminal tube. —
D. Placenta with ovules. Based on Schunke 9424 (MO).

fused into a tube 1.5-1.7 mm long. Pistil ovoid, 3.0
X 1.5 mm, stigma indistinctly 2-lobed; ovules ca.
15 in two rows. Fruits 2.5—3.5 cm diam., pericarp
0.3 mm thick, smooth; persisting calyx lobes re¬
flexed. Seeds 7-12, irregularly depressed-oblong,
8-11 X 7-8 mm.

In shape and size of the leaves Clavija obtusifolia
is similar to C. weberbaueri Mez, a species of wide
distribution in the western Amazon. From C. we¬
berbaueri, C. obtusifolia differs primarily in its
much larger flowers. Other distinguishing features
are the obtuse leaf apex, pale green lower leaf sur¬
face, and thick pedicels. In addition, morphologi¬
cally bisexual flowers of C. weberbaueri have 1—3
ovules, whereas those of C. obtusifolia have ca. 15.

374

Novon

Paratypes. PERU. San Martin: Prov. Mariscal C4-
ceres, Dtto. Uchiza, Cachiyacu de Lopuna, 450-500 m,
11 July 1974 (fr), Schunke 7323 (MO); Dtto. Tocache Nue¬
vo, Quebrada de Challua-Gacu, ca. 500 m, 1 Feb. 1979
(fr), Schunke 10757 (MO).

Literature Cited

Bameby, R. C. 1990. Two new species of Disciphania
(Menispermaceae: Tinosporeae) from subandean Ecua¬
dor and Peru. Brittonia 42: 178-181.

L0jtnant, B. & U. Molau. 1982. Analysis of a virgin par¬
amo plant community on Volc6n Sumaco, Ecuador. Nor¬
dic J. Bot. 2: 567-574.

St&hl, B. 1986. Two new species of Clavija (Theophras-
taceae) from NW South America. Nordic J. Bot. 6: 769-
772.

-. 1990. Theophrastaceae. Pp. 1—21 in G. Harling

& L. Andersson (editors), Flora of Ecuador 39. Dept, of
Systematic Botany, University of Goteborg, Goteborg.

-. 1991. A revision of Clavija (Theophrastaceae).

Opera Bot. 107: 1-77.

Eltroplectris brachycentron Szlachetko (Orchidaceae), a
New Orchid Species from Bolivia

Dariusz L. Szlachetko

Gdansk University, Department of Plant Ecology and Nature Protection,
Al. Legionow 9, 80-441 Gdansk, Poland

Abstract. Eltroplectris brachycentron Szlachetko
from Bolivia is described and illustrated, and its
taxonomic position is briefly discussed. An illus¬
trated key to the Bolivian species of Eltroplectris is
provided.

The genus Eltroplectris Rafinesque, with 12 spe¬
cies, is comprised of terrestrial orchids from Cen¬
tral and South America. The characteristic feature
of this genus is the presence of a threadlike spur,
formed by lateral sepals fused to each other in their
lower parts and the basal part of the lip. At first
glance, Eltroplectris appears similar to Pelexia Po-
iteau ex Lindley. The two genera differ, however, in
the structure of the rostellum and viscidium (Fig.
1). In Eltroplectris, the viscidium is semi-sheathlike
and slips from the rostellum, leaving the subulate,
sharp core of the rostellum. In Pelexia, the visci¬
dium originates on the external surface of the ros¬
tellum, is massive and nodular; the rostellum rem¬
nant is ribbonlike and obtuse at the apex.
Eltroplectris also appears to be related to Ptero-
glossa Schlechter. In both genera, the rostellum and
viscidium are similar in structure; the differences
primarily concern the structure of the spur. In El¬
troplectris the lateral sepals are fused into a tube
along the entire length of the free section of the
spur, whereas in Pteroglossa the lateral sepals re¬
main free to the apex of the spur. In Eltroplectris,
the free section of the spur is more or less equal to
the length of the part adhering to the ovary; in Pte¬
roglossa the spur is only free at the apex and fre¬
quently forms a chin at the base of the ovary. In
view of such clear criteria differentiating the two
genera, the fact that Garay (1982) transferred El¬
troplectris lurida (Correa) Pabst, a species closely
related to E. musera (Kraenzlin) Szlachetko, to Pte¬
roglossa is striking.

While studying herbarium material of Eltroplec-
tris at BM, I found a new species from Bolivia.

Eltroplectris brachycentron Szlachetko, sp. nov.
TYPE: Bolivia. Choreti, 3 mi. from Comiri, a
pumping station on the pipeline above Rio
Perapeti, 20°00'S, 63°30'W, 13 Sep. 1949,
Brooke 5634 (holotype, BM). Figure 2.

Species haec E. miserae propinqua, sed pede columnae
et calcare ovario manifeste breviori et labello ad clinan-
drium adnato sub apicem constricto jam diagnoscenda.

Stem 28 cm long, 2 mm diam. at base, 1 mm
diam. below inflorescence, erect, delicate, glabrous.
No leaves at flowering time. Cauline bracts 8, her¬
baceous, transparent, membranous, acute, adnate to
the stem, lower bracts longer, upper bracts shorter
than intemodes. Inflorescence 12.5 cm long, 25- to
30-flowered, multilateral, lax. Flowers small, arcu¬
ate, glabrous. Floral bracts 9 mm long, ovate-lan¬
ceolate, acuminate, 3-nerved, herbaceous, trans¬
parent, membranous, in lower flowers longer than
ovaries, in upper ones equal in length to ovaries.
Ovary 6 mm long, twisted at base, ovary neck about
1 mm long. Dorsal sepal 5 mm long, 1.8 mm wide,
ovate-lanceolate, acuminate, 3-nerved, slightly con¬
cave in the center, fleshy, thickened at the apex.
Free part of the lateral sepals 4 mm long, 1.3 mm
wide, lanceolate, acuminate, 3-nerved, delicate,
rather fleshy, slightly thickened at apices. Petals 4
mm long, 1 mm wide, lanceolate-falcate, subobtuse,
3-nerved, thin, slightly adnate to dorsal sepal. Lip
subsessile, constricted at the apex, thin, delicate,
papillate, distinctly adnate to the clinandrium; bas¬
al part of hypochile 3.2 mm long, 0.8 mm wide,
linear, apical part of hypochile 3.2 mm long, 2.5
mm wide, oval; epichile 1 mm long, 0.9 mm wide,
ovate, obtuse. Spur partially free, formed from the
column foot, lip and basal parts of lateral sepals,
as long as one-half of the ovary. Free part of the
spur 1 mm long, subulate. Gynostemium 3 mm
long, erect, short, relatively massive. Column foot
3 mm long, with the apical 1 mm free. Anther 1.8
mm long. Rostellum remnant 0.4 mm long, subu¬
late, acute.

Etymology: brachys (Greek), short; cenlron
(Greek), spur; an allusion to the short and sharply
pointed spur.

Eltroplectris brachycentron Szlachetko is closely
related to E. misera (Kraenzlin) Szlachetko. Both
of the species share a similar habit and gynostem¬
ium structure, but differ in the length of the spur,
lip form, and adnation between lip and clinandri-

Novon 5: 375-378. 1995.

376

Novon

Figure 1. Gynostemium structure and spur of Pelexia (a-d), Eltroplectris (e—g, h) and Pteroglossa (e-g, i). —a, e.
Rostellum with viscidium, bottom view. —b, f. Rostellum remnant. —c, g. Longitudinal section of rostellum and
viscidium. —d, h, i. Spur, pedicel, and ovary.

um. In E. brachycentron the spur reaches to the
mid point of the ovary, the lip is distinctly con¬
stricted at the apex, and the side lobes of the lip
are agglutinate with the clinandrium. In E. misera
the spur is as long as the ovary, the lip is not
divided into a hypochile and epichile, and it is
completely free from the clinandrium. Eltroplectris
misera (Kraenzlin) Szlachetko is well illustrated
by Dodson & Vasquez (1989a: 214) (as E. travas-
sosii (Rolfe) Garay).

It is worth noting that apart from E. brachycen¬
tron and E. misera, three other species of the ge¬
nus are known so far from Bolivia: E. longicornu
(Cogniaux) Pabst, described and known hitherto
only from southern Brazil; E. triloba (Lindley)
Pabst, illustrated by Dodson & Vasquez (1989b:
321) as E. calcarata (Swartz) Garay & Sweet; and
E. roseo-alba (Reichenbach f.) Hamer & Garay.

The Bolivian species of Eltroplectris can be sep¬
arated as follows (Fig. 3):

la. Lip elliptic in general outline, usually constricted
near the middle, flowers large, leaves gathered in
the basal rosette, obovate, subsessile . . E. roseo-alba

lb. Lip of different shape, constricted near the apex
if at all, flowers small, leaves petiolate if present
at flowering time.

2a. Hypochile linear in the basal part, ± dilated
in the apical part, side lobes rounded.

3a. Lip constricted near the apex, apical half
of hypochile elliptic, spur half as long as

the ovary. E. brachycentron

3b. Lip not constricted, the apical half nar¬
rowly elliptic, spur nearly as long as the

ovary. E. misera

2b. Hypochile triangular, side lobes triangular,
epichile ligulate to lanceolate.

4a. No leaves at flowering time, spur longer
than ovary and pedicel, lateral sepals

lanceolate. E. longicornu

4b. Leaves long petiolate, forming the basal
rosette at flowering, spur as long as the
ovary and pedicel or shorter, lateral se¬
pals strongly falcate. E. triloba

Acknowledgments. I am grateful to the curator
of BM for the hospitality during my visit, and Ry-
szard Ochyra for the latinization of the diagnosis.

Literature Cited

Dodson, C. H. & R. Vasquez Ch. 1989a. Orchids of Bo¬
livia. leones Plantarum Tropicarum. Ser. 2, fasc. 3:
201—300. Missouri Botanical Garden, St. Louis.

Volume 5, Number 4
1995

Szlachetko

New Orchid from Bolivia

377

Figure 2. Eltroplectris brachycentron Szlachetko. —a. Flower and floral bract. —b. Dorsal sepal.
Lateral sepal. —e. Lip. Drawn from the holotype.

c. Petal. —d.

378

Novon

Figure 3. An illustrated key to the Bolivian species of Eltroplectris. —a. E. roseo-alba (Reichenbach f.) Hamer &
Garay. —b . E brachycentron Szlachetko. —c. E. misera (Kraenzlin) Szlachetko. —d. Lip shape of E. longicornu
(Cogmaux) Pabst (e) and E. triloba (Lindley) Pabst (f).

ol ^ 1989b. Orchids of Bolivia. leones Garay, L. A. 1982. A generic revision of the Spiranthinae.

Plantarum Tropicarum. Ser. 2, fasc. 4: 301-400. Mis- Bot. Mus. Leafl. Harvard Univ. 28(4): 277-425.
souri Botanical Garden, St. Louis.

New Taxa of Rubiaceae from Amazonian Colombia,
Ecuador, and Peru

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

Abstract. The new taxa Coussarea linearis C. M.
Taylor, Palicourea macarthurorum C. M. Taylor,
and P. quadrifolia subsp. leticiana C. M. Taylor are
described based largely on material collected dur¬
ing floristic inventories of several biological re¬
serves in northwestern Amazonia.

During preparation of floras of the Parque Na-
cional Natural Amacayacu in southern Amazonas
Department, Colombia, and several biological re¬
serves near Iquitos, Peru, the following unde¬
scribed taxa were discovered.

Coussarea linearis C. M. Taylor, sp. nov. TYPE:
Peru. Loreto: Maynas, Iquitos, Allpahuayo, Es-
tacion Experimental del Instituto de Investi-
gaciones de la Amazonia Peruana (IIAP),
04°10'S, 73°30'W, 150-180 m, 18 Jan. 1991
(fl), R. Vdzquez 15853 (holotype, MO 4069962;
isotype, AMAZ). Figure 1.

Haec species a congeneribus pubescentia adpresso-
puberula, inflorescentia fasciculata reducta bracteis li-
nearibus 4—8 mm longis munita ac lobulis calycinis li-
nearibus 4—8 mm longis distinguitur.

Shrubs or subshrubs flowering at 0.5 m tall, to 4
m tall; stems terete, densely appressed-pilosulous
sometimes becoming glabrescent with age. Leaf
blades elliptic, 7.5—15 cm long, 1.5-8 cm wide, at
apex acute to acuminate with slender tips 8—15 mm
long, at base acute, papyraceous, sparsely to mod¬
erately appressed-pilosulous on lamina above and
below and similarly but more densely so on costa
and secondary veins on both sides; secondary veins
8-12 pairs, usually looping to interconnect, without
or with 1(—2) intersecondary veins present between
pairs of secondary veins, without domatia, the costa
and secondary veins plane to thickened above and
prominulous below; petioles densely appressed-pi¬
losulous, 3-23 mm long; stipules in bud ovoid, del¬
toid to rounded, persistent or falling off before the
leaves, leaving a truncate remnant, moderately to
densely appressed-pilosulous, becoming glabres¬
cent and indurate with age, 1.5-3.5 mm long, entire
to somewhat erose, enclosing colleters 0.5-2 mm
long. Inflorescences terminal, congested to some¬

what open, densely appressed-pilosulous, pedun¬
cles 1—3, fasciculate, 2—8 mm long, the flowers 1—
3 on each peduncle, sessile, bracts 2-4 per flower,
narrowly triangular to linear, appressed-pilosulous,
acute, 4—8 mm long; flowers with hypanthium cy¬
lindrical to ellipsoid, densely appressed-pilosulous
to -pilose, ca. 1.3 mm long; calyx limb densely
appressed-pilosulous, with tube 1.8-2 mm long,
lobes 4, linear, 4—8 mm long, acute; corollas sal-
verform, in bud yellow-green to whitish, externally
densely appressed-pilosulous, tubes ca. 12 mm
long, lobes 4, narrowly triangular to lanceolate, ca.
13 mm long, acute; style ca. 9 mm long, stigmas 2,
linear, ca. 3 mm long. Fruits ellipsoid to obovoid,
11-13 mm long, 7-9 mm diam., appressed-pilo¬
sulous, white, probably spongy; seed solitary, ellip¬
soid.

Distribution, habitat, and phenology. Amazo¬
nian Colombia and Peru, in terra firme forest at
100-180 m. Collected in flower in January and Oc¬
tober, in fruit January through September.

This species is distinguished by its densely ap¬
pressed-pilosulous pubescence on the stems,
leaves, stipules, and inflorescences; fasciculate and
rather reduced inflorescences; externally ap¬
pressed-pilosulous to -pilose corollas; numerous
linear inflorescence bracts; and linear relatively
long stipule lobes. The corollas are known only in
bud; because in many species of Coussarea the
flowers are nocturnal and the corolla tubes and
sometimes also the lobes elongate markedly just
before anthesis, the sizes given above are mini-
mums, and no reliable estimate of the mature flower
size can be made at present.

Although the stipules of this species may appear
glandular-fimbriate, the condition that distinguishes
Rudgea, the glands in Coussarea linearis are col¬
leters borne internally to the stipules and exposed
when the stipules fall, rather than glands attached
to the stipule apices as in Rudgea. The placement
of this species in Coussarea is indicated by the
four-parted calyx limb and corolla, stipules that en¬
close the buds in an imbricated apically rounded
conformation, and spongy white fruits containing
one ellipsoid seed with a papyraceous testa, in con-

Novon 5: 379-383. 1995.

380

No von

Figure 1. Coussarea linearis C. M. Taylor. —A. Habit. —B. Flower bud. A to 5-cm scale; B to 5-mm scale. A from
Vdsquez & Jaramillo 16520 (MO); B from Vdsquez 15853 (MO).

trast to the usually five-parted calyx and corolla and
fruits with two hard planoconvex pyrenes in Rudgea
and Psychotria, and aristate or mucronate stipules
with their apical projections usually crossed in Far-
amea.

This species can be confused with several other
pubescent species of Coussarea with well-devel¬
oped calyx lobes, in particular C. flava Poeppig &
Endlicher, which also has subsessile and congested
to subcapitate inflorescences, but which can be
separated by its oblanceolate to narrowly elliptic
leaves 8-28 cm long and 4—6.5 cm wide, and its
stems and flowers with spreading pilosulous tri-
chomes to 3 mm long; and C. enneantha Standley,
with the calyx limb composed of a tube 4—12 mm
long and narrowly triangular lobes 2-6 mm long,
the stem and inflorescence pubescence spreading,
the inflorescences branched at least once, and the
bracts 3 mm long or shorter or sometimes lacking.

Paratypes. COLOMBIA. Amazonas: Parque Nacion-
al Natural Amacayacu, near Tikuna village of Mocagua,
03°48'S, 70°18'W, Gentry & Villa-Lopera 60830 (COL,
MO); Parque Amacayacu, Centro Administrative Amacay¬
acu, Pipoly et al. 15080 (COL, MO), 15117 (COL, MO);
Parque Amacayacu, Centro Administrativo Mata-matA Pi¬
poly et al. 15681 (COL, MO), Rudas et al. 1297 (COL,
MO), 1579 (COL, MO); Parque Amacayacu, Vdsquez et al.
12598 (AMAZ, MO). PERU. Loreto: Maynas, Iquitos,
Allpahuayo, Estacion Experimental del Instituto de Inves-
tigaciones de la Amazonia Peruana (IIAP), Rueda & RuCz
648 (AMAZ. MO), Vdsquez & Jaramillo 16520 (AMAZ,
MO), Vdsquez 16798 (AMAZ, MO).

Palicourea inacarthuroruin C. M. Taylor, sp.
nov. TYPE: Colombia. Amazonas: Municipio
de Leticia, Corregimiento de TarapacA, Parque
Nacional Natural Amacayacu, Cabana Pamatd
(extremo nor-occidental del Parque), trocha
que sale de la cabana, paralela al Cano Pa-
mat£, 03°11'S, 70°20'W, 100 m, 1 July 1991
(fl), A. Rudas et al. 2725 (holotype, COL; iso¬
type, MO). Figure 2.

Haec species a Palicourea longiflora (Aublet) L. C.
Richard foliis utrinque glabris, pedunculo 0.5-3 cm lon-
go. inflorescentiae bracteis reductis, corolla extus glabra
ac fructu 10-11 mm longo distinguitur.

Shrubs to 3 m tall; stems glabrous, becoming te¬
rete. Leaves paired, blades elliptic to elliptic-oblong
or somewhat ovate, 10-16 cm long, 2.8-6 cm wide,
at apex acute to often acuminate with slender tips
1-2 cm long, at base acute to usually cuneate, pa¬
pyraceous, glabrous throughout; secondary veins 7—
12 pairs, looping broadly to interconnect, with 1-2
weak intersecondary veins usually present between
pairs of secondary veins, costa and secondary veins
thickened to prominulous above and below, the
lesser venation sometimes slightly thickened; peti¬
oles glabrous, 8—12 mm long; stipules glabrous,
united around the stem into a low continuous
sheath 0.5—1 mm long, entire, broadly concave, in-
terpetiolarly bilobed, lobes deltoid to ligulate, ob¬
tuse to rounded, 0.8-1.5 mm long, entire. Inflores¬
cences corymbiform, erect; peduncles 0.5—3 cm
long; panicles 2-4 cm long, 3-A.5 cm wide, with 1

Volume 5, Number 4
1995

Taylor

New Rubiaceae from South America

381

Figure 2. Palicourea macarthurorum C. M. Taylor. —A. Habit. —B. Flower bud. A to 5-cm scale; B to .V mm scale.
A from Rubio 94 (MO); B from Rudas et al. 2725 (MO).

pair of weakly developed lateral branches, these
and pedicels ascending, with flowers pedicellate in
open cymules of 2-5 or occasionally one flower per
cymule subsessile; bracts rudimentary or lacking,
those subtending larger branches usually displaced
from the branching point, 0.2—0.5 mm long, round¬
ed; pedicels 0.5—11 mm long; peduncle, axis,
branches, bracts, and pedicels glabrous to minutely
puberulous, yellow; flowers with hypanthium ca. 1
mm long, cylindrical to turbinate, minutely puber¬
ulous; calyx limb ca. 0.3 mm long, puberulous,
broadly shallowly lobed, ciliolate; corollas funnel-
form, yellow, swollen and gibbous at base, strongly
bent here but generally straight in the tube, gla¬
brous externally, internally glabrous except for a
densely pilose ring 1—1.5 mm wide just above the
basal swelling, tube ca. 17 mm long, ca. 1.5 mm
diam. just above the pilose ring, ca. 3 mm diam. at
mouth, lobes narrowly triangular, ca. 3 mm long,
acute and slightly thickened at apex; disk ca. 0.8
mm high. Infructescences similar in size to inflores¬
cences, the branches and pedicels often more
spreading, becoming red; fruits ellipsoid, 11-13
mm long, 10-12 mm wide, laterally somewhat flat¬
tened, glabrous; pyrenes with 3—5 broadly angled
low ridges.

Distribution, habitat, and phenology. Amazo¬
nian Colombia and Ecuador, in wet forest at 100-
1200 m. Collected in flower March, April, and July,
in fruit April and November.

This species is distinguished within Palicourea
by its relatively small stipules, short corymbiform
inflorescences with well-developed pedicels and ru¬
dimentary bracts, yellow and externally glabrous
corollas, and distinctive, large fruits with the py¬

renes nearly smooth. The fruit color is unknown but
those of most Palicourea species are blue-black.
This species is similar to P. longiflora (Aublet) L.
C. Richard, which is common further east in Ama¬
zonia and can be distinguished by its leaves pu¬
berulous to pilosulous abaxially, inflorescences with
2-3 pairs of lateral branches and bracts 0.5-7 mm
long borne on peduncles 2.5-10 cm long, corollas
puberulous externally, and fruits 4-5 mm long and
5-6 mm wide with the pyrenes distinctly and some¬
times rather sharply longitudinally ridged.

The specific epithet commemorates John D. and
Catherine T. MacArthur, whose foundation has
sponsored floristic inventories of several important
sites in Amazonian Peru and Colombia, including
the Parque Amacayacu.

Paratypes. COLOMBIA. Amazonas: Municipio de
Leticia, Parque Nacional Natural Amacayacu, Pipoly et al.
15319 (COL, MO), 15299 (COL, MO), 15353 (COL, MO),
15902 (COL, MO), Rudas et al. 3442 (COL, MO). Putu¬
mayo: Vereda Medio Alan, camino Medio Af4n—Churutr.
belo, ca. 2.5 horas de Mocoa hacfa el oriente, Garcia et
al. 64 (CUVC, MO). ECUADOR. Morona-Santiago:
Cordillera Cutucii, toward the Itzintza, ca. 02°40'S, 78°W,
Camp E-1352 (S). Napo: carretera Holltn-Loreto, Km 40,
00°43'S, 77°36'W, Hurtado et al. 208 (MO, QCNE), car¬
retera Hollfn—Loreto, Km 40-50, Hurtado 550 (MO,
QCNE). Pastaza: vfa Auca, 115 km al sur de Coca, cerca
del rfo Tigtiino, carretera de PETRO-CANADA en con¬
struct: i6n. 01°15'S, 76°55'W, Rubio 94 (MO, QCNE).

Palicourea quadrifolia (Rudge) de Candolle,
Prodr. 4: 529. 1830.

Steyermark (1972: 760) presented synonymy and
a discussion of this species.

382

Novon

Key to the Subspecies of

PAUCOUREA QUADR1FOUA

1. Leaves quadrate; floral bracts narrowly triangular
to linear, 0.2—0.5 mm wide; Guyana, French Gui¬
ana, Venezuela, Brazil, southern Peru (Madre de

Dios).subsp. quadrifolia

1. Leaves paired; floral bracts narrowly triangular to
elliptic, 0.3—1.2 mm wide; Colombia, northern
Peru (Loreto).subsp. leticiana

Palicourea quadrifolia subsp. leticiana C. M.

Taylor, subsp. nov. TYPE: Colombia. Amazo¬
nas: Municipio de Leticia, Corregimiento de
Tarapaca, Parque Nacional Natural Amacaya-
cu. Cabana Lorena (Rfo Cothu<5), 03°02'S,
70°00'W, 100 m, 24 June 1991, A. Rudas et
al. 2437 (holotype, COL; isotypes, MO
3933506, MO 4258973).

Palicourea lucentifolia Standley, Publ. Field Columbian
Mus., Bot. Ser. 8: 223. 1930. Syn. nov. TYPE: Peru.
Loreto: La Victoria on the Amazon River, 29 Aug.
1929, L. Williams 2906 (holotype, F 604457).

Haec varietas a varietate typica foliis binatis ac bracteis
floralibus ex linearibus anguste ellipticis 0.3—1.2 mm latis
differt.

Flowering at 1.5 m tall, to 5 m tall; stems gla¬
brous to sparsely puberulous. Leaves paired, with
blades narrowly elliptic to elliptic, 10.5—35 cm
long, 3-11.5 cm wide, acute to usually acuminate
at apex with slender tips 5-28 mm long, cuneate
to usually acute and sometimes tapering at base,
papyraceous, glabrous and usually shiny above,
glabrous to minutely puberulous below; secondary
veins 11-28 pairs, sometimes looping to intercon¬
nect distally or usually uniting with the margin,
usually with 1—3 weak intersecondary veins be¬
tween each pair of secondary veins; petioles gla¬
brous to sparsely puberulous, 3-15 mm long; stip¬
ules glabrous to sparsely puberulous, united around
the stem into a triangular to truncate continuous
sheath 2-6 mm long, enclosing and often exceeded
by colleters and appearing ciliate, sometimes split¬
ting to base, interpetiolarly bilobed, the lobes tri¬
angular to usually subulate, 0.5-3 mm long. Inflo¬
rescences corymbiform, flat-topped to broadly
rounded, erect; peduncles 4—11.5 cm long, sub¬
tended by a truncate stipule sheath that lacks
leaves or bears reduced leaves and terminates a
reduced or unexpanded intemode; panicles 4.5-9
cm long, 6-12 cm wide, lowermost branches
paired, ascending; flowers sessile in congested to
open and dichasial cymules of 3-7, each subtended
by l-2(-3) bracts, these free or slightly fused, lin¬
ear to narrowly elliptic, acute, 1.2—3 mm long, 0.3-
1.2 mm wide, ciliolate, on principal branches the

bracts absent or displaced markedly from the
branching nodes; peduncle, axis, branches, and
bracts yellow, minutely puberulous to pilosulous
with trichomes ca. 0.1-0.3 mm long; flowers disty-
lous, with hypanthium moderately to densely pu¬
berulous, turbinate to hemispherical, 0.3—0.5 mm
long, calyx limb densely puberulous, subtruncate
to dentate, ca. 0.2-0.3 mm long; corollas tubular,
yellow, externally glabrescent to densely puberu¬
lous and usually sparsely hirtellous on lobes, short¬
ly swollen at base, generally straight there and in
tube, internally glabrous except for a densely vil¬
lous ring ca. 1.5 mm wide just above the basal
swelling, tubes 8-12 mm long, 2.5—3 mm diam.,
lobes triangular to deltoid, 1—2 mm long; disk ca.
0.5 mm high. Infructescences similar in size and
proportion to inflorescences or sometimes more ex¬
panded with the branches spreading more widely,
becoming orange to red; fruits ellipsoid, ca. 3 mm
long, ca. 5 mm wide, somewhat didymous, glabres¬
cent; pyrenes 2, hemispherical to subglobose,
smooth.

Distribution, habitat, and phenology. Amazo¬
nian Colombia and Peru, in wet forest at ca. 100
m. Collected in flower in June and August to Sep¬
tember.

Palicourea quadrifolia subsp. leticiana differs
from subspecies quadrifolia only its paired rather
than quadrate leaves and wider floral bracts; oth¬
erwise it falls within the range of variation seen in
the latter subspecies. Paired leaves are occasion¬
ally found in some verticillate-leaved species (see
descriptions in Steyermark, 1974), and may be¬
come fixed in local populations, as in P. triphylla
de Candolle in Cuba (Taylor, 1993). This feature,
together with the consistently relatively broad floral
bracts, suggests that the plants segregated here as
subspecies leticiana are isolated from more eastern
and southern populations of P. quadrifolia. The
lack of any observable difference in reproductive
characteristics between these two morphological
forms and their complementary geographic ranges
suggest that subspecific status is more appropriate
than varietal or specific recognition. The epithet re¬
fers to the geographic area from which these plants
were first known.

Standley described Palicourea lucentifolia based
on a single specimen that was collected in the stage
in which the corollas have all fallen and no fruit
development is evident. Because conclusive iden¬
tification of Palicourea species is based on corolla
characteristics, it is preferable to have a type spec¬
imen that bears mature flowers, as does Rudas et
al. 2437, which also has several duplicates.

Volume 5, Number 4
1995

Taylor

New Rubiaceae from South America

383

Paratypes. COLOMBIA. Amazonas: municipio de
Leticia, Parque Nacional Natural Amacayacu, Rudas el al.
2352 (COL, MO); Rto Amazonas, vicinity of Leticia,
Schultes el al. 24023 (ECON, MO), 24032 (ECON). Ca-
queta: Florencia, cerro de La Sardina, Cuatrecasas 8891
(COL).

Acknowledgments. I thank A. Rudas and J. Pi-
poly for facilitating access to specimens from
Parque Amacayacu and the Iquitos Reserves, the
curators of F and US for the loan of specimens, and
R. E. Gereau 'or preparation of the Latin diagnoses.
This work was supported in part by a grant from
the John D. and Catherine T. MacArthur Founda¬

tion’s World Environment and Resources Program,
“Documentation of Neotropical Diversity and Bio¬
geography.”

Literature Cited

Steyermark, J. A. 1972. Palicourea. In: B. Maguire &
Collaborators, Flora of the Guayana Highlands. Mem.
New York Bot. Card. 23: 717-777.

-. 1974. Palicourea. In: T. Lasser (editor). Flora de

Venezuela 9: 1683—1830. Instituto Bot^nico, Direcci6n
de Recursos Naturales Renovables, Ministerio de Agri¬
culture y Crfa. Caracas, Venezuela.

Taylor, C. M. 1993. Revision of Palicourea (Rubiaceae:
Psychotrieae) in the West Indies. Moscosoa 7: 201-241.

Inga megaphylla (Leguminosae, Mimosoideae), a New Species
from Western Amazonia (Colombia, Peru), with Comments on
Architectural Features Unusual in the Genus Inga

Hans Vester

Hugo de Vries Laboratorium Universiteit van Amsterdam (The Netherlands Centre for
Geo-ecological Research. ICG), Kruislaan 318, 1098 SM Amsterdam,

The Netherlands

Odile Fancy

Museum national d’Histoire naturelle, Laboratoire de Phanerogamie, 16, rue Buffon,
F-75005 Paris, France

Abstract. A new species of Inga, discovered by
one of the authors (HV), is described and named
as Inga megaphylla Poncy & Vester. The architec¬
ture of this small understory cauliflorous tree is de¬
scribed and compared with that of another under¬
story species recently described from French
Guiana, Inga retinocarpa Poncy.

Inga (sect. Inga ser. Pilosulae) megaphylla Poncy
& Vester, sp. nov. TYPE: Colombia. Amazonas:
Rio Caquetd, Araracuara, 16 Oct. 1990 (fl),
Vester, Castro & Cleef 126 (holotype, P; iso¬
type, COAH). Figures 1-8.

Arbor parva, foliis amplis, foliolis (2—)3—4(—5)-jugis,
rachide alata. Inflorescentiae spicatae, cauliflorae, floribus
pubescentibus. Calyx irregulariter fissus; corolla ca. 14
mm longa; tubus staminalis exsertus. Legumen rectum,
glabrum, seminibus prominentibus.

Treelet, the fertile plants collected 6-10 m high,
with a trunk 3—9 cm diam.; bark light gray, scarcely
lenticelled. Young twigs rusty pubescent. Leaves
very large, pinnate with (2—)3—4(—5) pairs of leaf¬
lets. Stipules early caducous, seen on the seedling
only. Petiole 7—10(—15) cm long, thick (up to 8 mm
diam.) terete at length or distally narrowly winged,
glabrous or slightly tomentose, lignified on old
leaves. Segments of the rachis 6-10 cm each,
winged, oblong, up to 2 cm wide, the main nerve
tomentose on both faces. Nectaries orbicular, 12
mm diam. Leaflets elliptic and almost symmetrical,
those of the proximal pair ca. 16 X 6 cm, of the
distal one up to 40 X 15 cm; base acute to cuneate,
apex acute to obtuse, acuminate. Inflorescence a
cauliflorous spike, arising from the trunk or older
lateral branches up to 6-10 m from the ground,
solitary or fascicled in the axil of prominent and
large (ca. 1.5 cm) triangular leaf scars; peduncle

slender, 1.5—5(-7)cm long; rachis 1.5-2 cm long;
bracts (very few seen) caducous, or if persistent in¬
conspicuous (1 mm long), rigid, scaly; flowers ped¬
icellate, pedicel ca. 1.5 mm long; calyx funnelform,
yellowish, very light-rusty, 6-7 mm long, irregularly
cleft, glabrous to lightly pubescent, and more so on
the margins of the teeth, these acute, ca. 2 mm
long; corolla pale yellowish green, 13—14 mm long,
covered with dense appressed shiny hairs, the lobes
acute ca. 2.5 mm long; androecium white, the sta-
minal tube exserted, up to 17 mm long, the free
part of the filaments ca. 30 mm; bases of the sta-
minal tube and corolla fused in a 2-mm-long ste-
monozone; nectariferous dots are visible inside the
staminal tube at its base, surrounding the base of
the ovary. Ovary glabrous, ovules 20—24 (4 flowers
dissected). Pod straight, hanging, 15—20(—22) cm
X ca. 1.7 cm, glabrous; the sutures slightly prom¬
inent, 2—3 mm wide, the valves transversally rip¬
pled, dark green at maturity, the seeds not contig¬
uous, prominent. Seeds 14—16 (26 for the specimen
from Peru), 1.5 X 7.8 mm, the tegument a white
edible pulp, the cotyledons bright black. Seedling:
epicotyl 4 cm; first leaves opposite, bifoliolate (i.e.,
with one pair of leaflets), the petiole ca. 0.5 cm;
next leaves alternate with a longer and winged pet¬
iole, the first 2—5 next ones also bifoliolate, then
with two, then three pairs of leaflets; axis, terminal
bud, and veins green tomentose; stipules ca. 0.5
cm long, linear.

The specific epithet refers to the very large
leaves.

Distribution. Lowland rainforest of western Ama¬
zonia (southern Colombia and northern Peru).

Local name. Jiteinicoai (huitoto).

Inga megaphylla is referable to series Pilosulae

Novon 5: 384-387. 1995.

Volume 5, Number 4
1995

Vester & Poncy

Inga megaphylla from Western Amazonia

385

Figures 1-8. Inga megaphylla Poncy & Vester. —1. Leaf. —2. Inflorescence. —3. Fruit. —4. Flower. —5. Longi¬
tudinal section of the base of the flower, the ovary removed, the calyx dotted, the corolla, staminal tube, and stemonozone
hatched. —6. The dorsifixed anther, lateral and vertical views. —7. Stigma. —8. Seedling (only one leaf of the first
opposite pair of leaves is drawn).

386

Novon

Figures 9-12. Architecture of Inga megaphylla (9, 10) in Colombia and /. relinocarpa (11, 12) in French Guiana. —
9. Inga megaphylla, branching pattern of a ca. 10-m-high tree. —10. Schematic architectural pattern drawn from
observation of several individuals. —11. Inga retinocarpa, branching patterns of an individual 5.50 m high. —12.
Habit of an individual 3.50 m high: all leaves are represented; branching was sylleptic although the primary axis
continues living without developing further.

Volume 5, Number 4
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Vester & Poncy

Inga megaphylla from Western Amazonia

387

of section Inga. Its floral characters suggest affini¬
ties with I. splendens Willdenow, which, however,
has sessile flowers and a larger calyx.

Inga megaphylla is an understory, cauliflorous
treelet with very large leaves that simulate lateral
branches. Its architecture can be described as fol¬
lows (Figs. 9, 10): main axis unbranched or sparse¬
ly branched often up to 4 m high; branches ortho¬
tropic like the main axis; branching not rhythmic
or not clearly so; sylleptic branching very scarce
and only one branch at a time; most branching pro-
leptic. The individual illustrated in Figure 9 exhib¬
its these features, while Figure 10 integrates all the
observations made. The architectural model must
be related to Champagnat’s model (Halle & Olde-
man, 1970; Halle et ah, 1978), although the axes
do not bend in the way suggested in the definition
of that model. In a somewhat open undergrowth
with more light available, the trunk and branches
can be slender and bending.

The habit of our new species can be compared
with that of another understory, cauliflorous species
described from French Guiana, Inga retinocarpa
Poncy (Poncy, 1991). This species appears to be
quite common around the Field Research Station
“Les Nouragues” (French Guiana), near the Ara-
taye River, where the type specimen was collected.
This is also a small tree, although it seems to reach
a greater size than I. megaphylla. However, small
individuals only 4 m high consisting of one ortho¬
tropic, very scarcely branched primary axis can be
fertile. All samples observed have orthotropic lat¬
eral branches with spirally arranged leaves (Figure
12). Branching is mainly sylleptic, and as in I. me¬
gaphylla not clearly rhythmic, but certainly not
continuous. The result of this sylleptic branching
is that I. retinocarpa forms a broader crown. The
basic architectural model is nevertheless difficult

to determine for such plants in very constraining
living conditions. The species is now grown in the
nursery of the camp, but only the early stages have
been observed; indeed, I. retinocarpa seems to grow
extremely slowly, so that trees of 5 cm diameter
could be several decades old.

Such architectural features are surprising in a
genus where so many species exhibit the same
Troll’s model (Poncy, 1985). Inga megaphylla and
I. retinocarpa, which are remotely allopatric and not
closely related taxonomically, seem, however, to
have adapted in very similar ways to understory
conditions.

Paratypes. COLOMBIA. Caqueta: Rfo Caqueta. Ar-
aracuara, 2 Feb. 1991 (fr), Vester & Castro 315 (COAH,
P), 28 Sep. 1993 (fl), Vester & Roman 805 (COL, COAH,
P, AMD). PERU. Loreto: rio Yubineto, 15 Feb. 1978 (fr).
Barrier 668 (P).

Acknowledgments. The first author thanks Tro-
penbos, ICETEX, and the Dutch Canopy Founda¬
tion for the financial support of fieldwork carried
out in Araracuara. He also thanks his field guides:
Bartolome Castro, Oscar Roman, Anfbal Matapi,
Jesus Manaidego, Simon Roman, and Miguel Guer¬
rero.

Literature Cited

Halle F. & R. A. A. Oldeman. 1970. Essai sur (’archi¬
tecture et la dynamique de croissance des arbres tro-
pieaux. Masson, Paris.

-,-& P. B. Tomlinson. 1978. Tropical Trees

and Forests: An Architectural Analysis. Springer-Ver-
lag, Berlin.

Poncy, O. 1985. Le genre Inga (L£gumineuses, Mimo-
soideae) en Guyane frangaise: Svstcnial lquc. morphol-
ogie des formes juveniles, ccologie. Mem. Mus. Natl.
Hist. Nat. (Paris), s6r. B, Bot. 31: 1—124.

Poncy, O. 1991. Deux nouvelles esptVcs de Inga (Mi-
mosaceae), et notes nomenclaturales et taxonomiques
sur trois autres espoces Bull. Mus. Hist. Nat. (Paris),
4e srr. 13, sect. B, Adansonia, 3 4: 147—154.

Encephalartos imbricans (Zamiaceae): A New Species from Uganda

P. Vorster and P. Heibloem

Department of Botany, University of Stellenbosch, Private Bag XI, 7602 Matieland,
South Africa. Current address of P. Heibloem: P.0. Box 700, Nambour,
Queensland 4560, Australia

ABSTRACT. Encephalartos imbricans, a new spe¬
cies from Uganda, is described, illustrated, and cir¬
cumscribed in terms of other tropical African ar¬
borescent species. It is a large arborescent plant,
with leaflets mostly strongly incubously overlap¬
ping. The strobili are green and glabrous; the mega-
strobilus is ovoid, shortly pedunculate, with the
surfaces of the bullae wrinkled; the microstrobili
emerge in succession, being narrowly ovoid, long-
pedunculate, with the surfaces of the bullae
smooth.

Encephalartos is widespread in the eastern parts
of Africa, from the Eastern Cape Province of South
Africa to southern Sudan, extending westward
through equatorial Africa to Ghana and Nigeria
(Goode, 1989: 244; Fig. 1). Nowhere is it common;
it occurs mostly as relic populations, of which a
considerable percentage must be considered to be
in danger of extinction. The description of E. im¬
bricans brings the number of described and rec¬
ognized taxa to 57.

Encephalartos imbricans Vorster, sp. nov. TYPE:
Uganda. Near northern shore of Lake Victoria,
Oct. 1994, P. Heibloem sub P. Vorster 3034 (ho-
lotype, K). Figures 2-5.

Planta magna arborescens. Foliola plerumque valde in-
cuba, minime leviter falcata, dentibus in marginibus am-
bobus ad spationibus 25-50 mm et dentibus 3-4 confertis
concentratis ad basin marginis superioris. Strobili virides,
glabri, bullarum porcis mediolateralibus integris; mega-
strobili ovoidei, breve pedunculati, bullarum faciebus ra-
dialiter rugosis; microstrobili in successione emergentes,
anguste ovoidei, longe pedunculati, bullarum faciebus lae-
vigatis.

Large arborescent, unbranched plant; suckering
profusely from base. Stem up to 6 m long, pros¬
trate after 1.5—2.0 m from base, 400-650 mm
diam., covered with old leaf bases. Fronds nu¬
merous, spreading, straight, emerging fresh me¬
dium green and maturing to dark glossy green,
about 2(-3.5) m long, leaflets gradually reduced
to prickles toward proximal part of petiole to leave
no prickle-free petiole; median leaflets orientated

at angle of about 45° with rachis toward apex of
frond, opposing leaflets are in the same plane as
seen in cross section of leaf, very narrowly ovate
and very slightly falcate, with teeth 25-50 mm
apart on both margins and often 3-4 closely
spaced teeth near base of upper margin, mostly
strongly incubously overlapping, 200-250 mm
long and up to 22 mm wide. Megastrobili 2-3,
ovoid with apical sporophylls sterile, shortly pe¬
dunculate with exposed peduncle 30-40 mm long,
glaucous green, glabrous, about 300 mm long and
180 mm diam.; bullae of sporophylls moderately
projecting, terminal facet centrally situated and
about % the horizontal diameter of bulla, median
facet differentiated in most sporophylls, ridges
separating facets clearly differentiated and simple,
terminal facet smooth but other facets radially ru¬
gose. Microstrobili 5—8, emerging and maturing in
succession, each carried on a stout peduncle up
to 300 mm long; strobili glaucous green, glabrous,
about 300 mm long and 90 mm diam. at maturity;
bullae of sporophylls moderately projected, slight¬
ly drooping, only terminal facet differentiated and
about V 3 the horizontal diameter of bulla, smooth.
Seed covered with orange (not red) sarcotesta,
about 45—50 mm long and 25-30 mm wide; with
sarcotesta removed about 40 mm long and 18-24
mm wide.

Variation. A concentration of 3 to 4 closely
spaced teeth near the base of the upper margin is
present in most (Fig. 2B), but not all (Fig. 3), leaf¬
lets. Likewise, the majority of plants have strongly
overlapping leaflets; but through a quirk of fate the
frond collected for the type specimen has well¬
spaced leaflets—it probably developed in a situa¬
tion where it was shaded by other plants and thus
became etiolated.

Phenology. During October the plants had
fresh male and female cones, and seeds from the
previous season were scattered about. A mature
megastrobilus collected at this time disintegrated
in mid-November, suggesting that the megastrobili
take slightly more than a year to mature.

Geographical distribution and habitat. Known

Novon 5: 38B-394. 1995.

Volume 5, Number 4
1995

Vorster & Heibloem
Encephalartos imbricans

389

Figure 1. Encephalartos imbricans Vorster. Known geographical distribution in relation to distributions of other tropical
African arborescent species. Based on Stevenson et al. (1990b) and Goode (1989). Scale bar — 1000 km.

Total distribution of
genus Encephalartos

Encephalartos bubalinus
E. gratus
E. hildebrandtii
E. imbricans
E. ituriensis
E. kisambo
E. laurentianus
a E. sclavoi
▼ E. septentrionals

K1 J7

only from two populations on the northern shore of
Lake Victoria, where it occurs on two adjacent low
hills in otherwise fairly flat country (Fig. 1). It
grows on bare granite outcrops, wedged in crevices
from which the plants cannot escape, in minimal
soil, and under intense insolation. These outcrops
(Fig. 6) have minimal vegetation, yet the charred
trunks of Encephalartos imbricans testify that the
plants are subject to periodic burning, and suitable
niches for seedling establishment must be very
scarce.

Conservation status. The population visited
consisted of about 300 individuals. It appeared to
be vigorous and was coning, yet no young plants or

evidence of seedling regeneration were seen. It
seems likely that the number of plants is in equi¬
librium with the limited extent of the specialized
habitat. Seed dissected contained no embryos, and
no insects that could be suspected of being pollen
vectors were found in either male or female strobili.
We therefore conclude that these populations are
vulnerable to overcollecting, and, accordingly, the
precise locality is not made public.

Affinities. Within Encephalartos, interspecific
relationships are still poorly understood. For many
years species were separated solely on vegetative
characteristics (Vorster, 1993). There does indeed
seem to be a considerable amount of convergence

390

Novon

Figure 2. Encephalartos imbricans Vorster. —A. Proximal part of petiole. —B. Median leaflet. —C. Megasporophyll
in ventral view. —D. Seed with sarcotesta. —E. Seed with sarcotesta removed. —F. Microsporophyll in ventral view.
Scale bar = 50 mm. (Heibloem sub Vorster 3034.)

Volume 5, Number 4
1995

Vorster & Heibloem
Encephalartos imbricans

391

Figure 3. Encephalartos imbricans Vorster. Part of me¬
dian zone of frond, showing incubously overlapping leaf¬
lets. Scale bar = 100 mm. (Plant in habitat.)

in cone characteristics, yet these are often useful
to circumscribe species, if not always reflecting
phylogenetic relationships. It would appear that re¬
curring vegetative characteristics may well be cryp¬
tic signposts of evolutionary kinship. The following
is therefore intended to identify E. imbricans, but
is certainly not intended to reflect phylogenetic re¬
lationships.

Encephalartos imbricans resembles E. bubalinus
Melville (Melville, 1957) in its green, ovoid, shortly
pedunculate megastrobili (Fig. 4E), but differs by
the somewhat wrinkled instead of smooth facets of
the megastrobilate bullae (Fig. 4F), and by the
long-peduncled instead of almost sessile microstro-
bili. In both species the leaflets are overlapping,
but in E. imbricans the leaflets taper gradually to
their apices (Fig. 2B), whereas in E. bubalinus they
are uncinate.

Encephalartos imbricans resembles an unde¬
scribed species from western Uganda in its green,
ovoid megastrobili, but is readily distinguished by
the somewhat wrinkled instead of smooth facets of
the megastrobilate bullae, and the mostly strongly

overlapping (Fig. 3) instead of well-spaced leaflets.
The concentration of 3—4 teeth near the base of the
upper leaflet margin was not seen in the western
Ugandan species.

Encephalartos imbricans resembles E. laurenti-
anus De Wildeman (De Wildeman, 1903a, b, c,
1907; Bois, 1907; Gentil, 1904a, b; Lebrun, 1930;
Robyns, 1948) by its shortly pedunculate, ovoid
megastrobili, of which the facets of the bullae are
somewhat wrinkled, as well as by the large number
of long-pedunculate microstrobili; it is distin¬
guished by the absence of a characteristic thin in¬
dumentum of reddish brown trichomes, and green
rather than greenish yellow pigmentation of the
strobili. Vegetatively, E. laurentianus is a very dif¬
ferent species, with much larger (up to 500 mm
long and 50 mm wide) and softer-textured leaflets.

The new species resembles Encephalartos septen-
trionalis Schweinfurth (Goode, 1989: 236) by its
pedunculate, ovoid megastrobili, and long-pedun¬
culate microstrobili, but differs in the very short
instead of very long (up to 300 mm) peduncled me¬
gastrobili (Goode, 1989: 239) and the glaucous
green instead of greenish yellow pigmentation of
the strobili. Both species tend to have a concentra¬
tion of up to 4 teeth near the base of the upper
margin of the leaflet, but in E. septentrionalis the
leaflets are not overlapping but well spaced. De¬
spite being an old species, E. septentrionalis is not
well known, and in view of its unusually wide geo¬
graphical distribution (according to the map in Ste¬
venson et al., 1990b: 157) more than one species
may be involved. In this respect it is worth noting
that the very long megastrobilate peduncle depicted
by Goode (1989: 239) is not mentioned elsewhere
in the literature.

Encephalartos imbricans resembles E. sclavoi
De Luca, Stevenson & Moretti (Stevenson et al.,
1990a, b) by the ovoid, shortly pedunculate mega¬
strobili as well as by the overlapping leaflets; it
differs in that both mega- and microstrobili are
green with centrally placed terminal facets on ru¬
gose megastrobilate bullae instead of orange-yel¬
low with terminal facets of the smooth megastro¬
bilate bullae situated toward the proximal margins
of the bullae, the long (up to 300 mm) instead of
very short-peduncled (20-40 mm) microstrobili,
and the narrower (up to 22 mm vs. up to 40 mm)
leaflets. Encephalartos sclavoi lacks the concen¬
tration of 3 to 4 teeth near the base of the upper
leaflet margin.

Both Encephalartos imbricans and E. turneri La-
vranos & Goode (Lavranos & Goode, 1985) have
imbricate leaflets, but in E. turneri they are suc-
cubous instead of incubous (Fig. 3), they are wider

392

Novon

E

Figure 4. Encephalartos imbricans Vorster. A. Microstrobilus in immature stage. —B. Portion of immature micro¬
strobilus, showing centrally situated terminal facet. —C. Microstrobilus at anthesis. —D. Portion of microstrobilus at
anthesis, showing drooping terminal facet. —E. Megastrobilus. —F. Portion of megastrobilus showing rugose facets of
bullae. Scale bars (at the right in each photograph): A, C, E = 100 mm; B, D, F = 10 mm. ( Heibloem sub Vorster
3034.)

Volume 5, Number 4
1995

Vorster & Heibloem
Encephalartos imbricans

393

Figure 5. Encephalartos imbricans Vorster. Plant in hab¬
itat, showing habit. Scale bar = 1 m.

(up to 30 mm vs. up to 22 mm), and characteris¬
tically the apices are uncinate instead of tapering
to a fine point. In E. turneri the megastrobilus is
narrowly ovoid instead of ovoid, completely sessile
instead of shortly peduncled, and the bullae project
more. It bears fewer microstrobili (up to 3 vs. 5-8)
on shorter peduncles (up to 120 vs. to 300 mm). In
both sexes the cones are not green but yellowish
with a conspicuous pink bloom (Lavranos & Goode,
1985).

Encephalartos imbricans resembles E. ituriensis
Bamps & Lisowski (Bamps & Lisowski, 1990) by
its long-peduncled microstrobili. However, in E.
ituriensis the megastrobilus is shortly cylindrical
and the strobili of both sexes are yellow instead of
green (Goode, 1989). Moreover, the type collection
of E. ituriensis (Lisowski 41057, BR) leaves the im¬
pression of relatively soft textured and flaccid leaf¬
lets as in E. laurentianus, in marked contrast to the
hard and rigid leaflets of E. imbricans.

Other east African arborescent species with
which we compared Encephalartos imbricans are
E. hildebrandtii A. Braun & Bouche (Praia, 1915;

Stapf, 1914), E. kisambo Faden & Beentje (Faden
& Beentje, 1989; Moretti et al., 1989), and E. te-
gulaneus Melville (Melville, 1957). From all these
it differs by its glaucous green instead of yellow
cones, and the ovoid instead of cylindrical mega-
strobili. Yet in various respects it resembles these
species, which probably indicates a near relation¬
ship. In all four species the microstrobili are borne
on very long peduncles, which can be up to 300
mm long. In both E. imbricans and E. hildebrand¬
tii the microstrobili emerge and mature in succes¬
sion rather than simultaneously. This trait is
shared by the undescribed species from western
Uganda, by E. gratus Prain (Prain, 1916) from Ma¬
lawi, and E. ferox G. Bertoloni (pers. obs.) from
southern Mozambique and South Africa, and is
suspected to be present in at least some other
tropical African species including E. septentrion-
alis and E. laurentianus. In the South African spe¬
cies, in the E. manikensis (Gilliland) Gilliland
complex, the E. poggei Ascherson & Graebner
group, and in E. barteri Carruthers, the cones
emerge simultaneously (teste Goode, 1989; pers.
obs.). The conspicuous grouping of 3 to 4 teeth
near the base of the upper leaflet margin is shared
by E. hildebrandtii (some specimens, e.g., Faden,
Faden & Rathbun 72 136, K), E. kisambo, E. te-
gulaneus, E. bubalinus (where it is poorly ex¬
pressed), the Malawian E. gratus, and the South
African E. woodii Sander, and it is tempting to
read phylogenetic significance in it.

Melville (1957: 248) briefly described a variety
dentatus of E. hildebrandtii, based on cultivated
specimens. Heenan (1977: 281—282) reported that
he had rediscovered it near Jinja in Uganda. This
is not the same as E. imbricans, because E. hilde¬
brandtii var. dentatus is distinguished by divari¬
cate-dentate medio-lateral ridges on both the mi¬
cro- and megasporophyllate bullae.

Acknowledgments. The research of which this pa¬
per is a result, was financed by the University of
Stellenbosch and the South African Foundation for
Research Development. We thank the Keeper of
Botany at K for granting to P. V. the opportunity to
examine material of Encephalartos in K, and for
providing us with photocopies of rare literature. For
assistance with the latter we also wish to record our
appreciation to the South African liaison officer at
Kew, Miss M. Koekemoer. P. H. wishes to express
his gratitude to Paul C. Mackenzie of Nairobi, Ke¬
nya, for logistic and other support during the course
of his fieldwork.

394

Novon

Literature Cited

Bamps, P. & S. Lisowski. 1990. A new species of Ence-
phalartos [E. ituriensis ] (Zamiaceae) from northeastern
Zaire. Mem. New York Bot. Gard. 57: 152—155.

Bois, D. 1907. Les Encephalartos lemarinelianus et lau-
rentianus. Rev. Hort. 79: 177-179.

Faden, R. B. & H. J. Beentje. 1989. Encephalartos kis-
ambo, a new cycad from Kenya, with a note on E. te-
gulaneus. Utafiti 2: 7-10.

Gentil, L. 1904a. Encephalartos from the Congo. Gard.
Chron., June 1904: 370-371.

-. 1904b. Les cycad6es au Congo. Rev. Hort. Beige

£trang£re 30: 7—9.

Goode, D. 1989. Cycads of Africa. Struik Winchester,
Cape Town.

Heenan, D. 1977. Some observations on the cycads of
central Africa. J. Linn. Soc., Bot. 74: 277-288.

Lavranos, J. & D. Goode. 1985. Encephalartos turned
(Cycadaceae), a new species from Mozambique. Garcia
de Orta 7: 11—14.

Lebrun, J. 1930. Note sur VEncephalartos laurentianus
De Wild., de la valine du Kwango. Rev. Zool. Bot. Af-
ricaines 19: 384—390.

Melville, R. 1957. Encephalartos in central Africa. Kew
Bull. 2: 237-257.

Moretti, A., P. de Luca, J. P. Sclavo & D. W. Stevenson.
1989. Encephalartos voiensis (Zamiaceae), a new east
central African species in the E. hildebrandtii complex.
Ann. Missouri Bot. Gard. 76: 934r-938.

Prain, D. 1915. Encephalartos hildebrandtii. Bot. Mag.
141: tt. 8592 & 8593.

-. 1916. Diagnoses Africanae LXVII. Kew Bull. 7:

180-182.

Robyns, W. 1948. Cycadaceae. Pp. 1^1 in Flore du Con¬
go beige et du Ruanda-Urundi, Spermatophytes, vol. 1.
I.N.E.A.C., Bruxelles.

Stapf, O. 1914. Encephalartos hildebrandtii. Kew Bull.
10: 386-392.

Stevenson, D. W., A. Moretti & P. de Luca. 1990a. En¬
cephalartos sclavoi De Luca, D. Stevenson & Moretti
(Zamiaceae), a new species from Tanzania. Delpinoa,
n.s. 29-30: 3-5.

-, -& -. 1990b. A new species of

Encephalartos [E. sclavoi ] (Zamiaceae) from Tanzania.
Mem. New York Bot. Gard. 57: 156—161.

Vorster, P. 1993. Taxonomy of Encephalartos (Zamiaceae):
Taxonomically useful external characteristics. Pp. 294—
299 in D. W. Stevenson & K. J. Norstog (editors). Pro¬
ceedings of CYCAD 90, the Second International
Conference on Cycad Biology, Palm & Cycad Societies
of Australia, Milton, Queensland.

De Wildeman, E. 1903a. Etudes de syst^matique et de
g6ographie botaniques sur la Flore du Bas- et du Moy-
en-Congo. Ann. Mus. Congo, S6r. 5, Bot. 1: 9-10, tt.
23-25.

-. 1903b. Leones Selectae Horti Thenensis 4: 183.

-. 1903c. Notices sur des plantes utiles ou int6r-

essantes de la Flore du Congo [1 J: 386-394.

-. 1907. Pp. 364—370 in Mission Emile Laurent.

Vanbuggenhout, Bruxelles.

Kengyilia guidenensis (Poaceae: Triticeae), a
New Species from Western China

Chi Yen and Jun-Liang Yang

Triticeae Research Institute, Sichuan Agricultural University, Dujiangyan City 611830,
Sichuan, People’s Republic of China

Bernard R. Baum

Centre for Land and Biological Resources Research, Agriculture and Agri-Food Canada,
Central Experimental Farm, Ottawa, Ontario, Canada, K1A 0C6

Abstract. Kengyilia guidenensis, a new species
from western China, is described and illustrated.
Cytological observations are provided, and compar¬
isons with similar species are made.

During an expedition to western China, a new
species of Kengyilia was found. It was growing on
a mountain slope close to the highway from Guide
to Guomaying, between the 150 and 151 Km road
poles. When collected it was identified as K. nana
J. L. Yang, Yen & Baum; however, the habitat of K.
nana is alpine steppes above 4000 m, whereas the
Guide population was found just below 3100 m al¬
titude and in stony slopes. The habitat of the Guide
population is similar to that of K. kokonorica (Keng)
J. L. Yang, Yen & Baum, but differs by being dwarf
in appearance and by having ovate, dense, small
spikes. In the same site another species of Kengyi¬
lia was found. This was A. mutica (Keng ex Keng
& S. L. Chen) J. L. Yang, Yen & Baum, from which
it readily differs in gross morphology by its dwarf
habit and by its short ovate spike with long lemma
awn. Kengyilia mutica does not have awns, and its
spike is long and cylindrical.

These preliminary observations led to the sus¬
picion that a new species was on hand. The purpose
of the present paper is to describe this species and
to detail differences from the species most similar
to it.

Materials and Methods

Herbarium specimens and seeds were collected
in 1993 in Guide County, Qinghai Province, China,
and compared with specimens of Kengyilia nana
and A. kokonorica (including their types) in N and
SAUTI. Cytological observations were made using
the methods detailed in Yen and Yang (1990).

Description

Kengyilia guidenensis Yen, J. L. Yang & Baum,
sp. nov. TYPE: China. Qinghai Province:
Guide County, Guide to Guomaying highway
between markers 150 and 151 Km, alt. 3100
m, 35°89'N, 100°26'E, 11 Sep. 1993, C. Yen,
J. L. Yang & G. L Sun 930001 (holotype,
SAUTI; isotype, DAO). Figure 1.

Kengyiliae kokonoricae (Keng) J. L. Yang, Yen & Baum
affinis, sed laminis superioribus albo-villosis; spicis bre-
vioribus, 3—3.5 cm, et angustioribus, 5 mm; spiculis bre-
vioribus, 7 mm, et plurifloribus, 5—7; glumis longioribus,
4-7 mm, lanceolatis; aristis lemmatum brevioribus, 1.5—
4 mm; carinis palearum ominino ciliatis; antheris longior¬
ibus, 3 mm, differt. Kengyiliae nanae J. L. Yang, Yen &
Baum affinis, sed culmis longioribus, 35-50 cm longis;
foliorum vaginis glabris, laminis longioribus, 8.5-9 cm
longis; spicis angustioribus, 5 mm latis; glumis lanceola¬
tis; aristis lemmatum brevioribus, 1.5-4 mm longis; paleis
ominino ciliatis differt.

Caespitose perennial, with fibrous roots (Fig.
1A). Culms caespitose, erect, 35—50 cm tall, 1.5
mm diam., 2—3-noded, pubescent just below the
spike for 1—3 cm, geniculate at the base. Leaf
sheaths glabrous, rarely pubescent in the lowermost
sheaths; ligules scarious, membranous, truncate,
premorse, ca. 0.5 mm long; leaf blades flat or
slightly involute, narrowly lanceolate, 8.5-9 cm
long, 3 mm wide, their adaxial surface with dense
1-mm-long white hairs emerging from the lateral
nerves, their abaxial surface glabrous; leaf blades
of the tillers densely pubescent, up to 12 cm long.
Spikes erect, short and dense, ovate, 3-3.5 cm
long, 5 mm wide; rachis intemodes 1.5-2 mm long,
densely white-pilose; spikelets oblong-ovate (Fig.
1J), greenish purple, with 5-7 florets 7 mm long
(excluding awns); rachilla intemodes 1—1.2 mm
long, densely hirtulous; glumes unequal, wide lan¬
ceolate, with herbaceous membranous margins (Fig.
IB, C), 1(—3)-nerved, with midrib pronounced and

Novon 5: 395-397. 1995.

396

Novon

Dorsad L K ?f ' ll T ^(T m “, Yen ’ l h Yang & Baum ' - A - Plant - - B ' « lume - —C. Upper glume. -D.

J Tpfkelet " V ' eW ° rel - ~ F ' Pa ' ea - Anther - ~ H L^icu^s. -I. Young gynoecium. -

10 cm

Volume 5, Number 4
1995

Yen et al. 397

Kengyilia guidenensis from Western China

thus appearing as keels, densely white-hirsute,
acuminate or mucronate, the 1st glume 3—5 mm
long, the 2nd glume 4—7 mm long; glumes of the
uppermost spikelets membranous and 2-lobed, very
unequal; lemmas ovate, 7—9 mm long (excluding
awns), densely beset with 1-mm-long white hairs,
ending with a 1.5-4-mm-long scabrous awn (Fig.
ID, E); the lowermost lemma 7 mm long; callus
with 0.8-mm-long hairs on both sides; palea equal
to lemma, with emarginate apex, keels ciliolate in
their entire length, densely hirsute between the two
keels (Fig. IF); anthers yellow, 3 mm long (Fig.
1G). Lodicules triangular (Fig. 1H) with a distinct
side lobe in their lower half. Caryopsis brown, 5
mm long.

Kengyilia guidenensis grows on stony slopes, be¬
tween broken granite rocks, around 3100 m.

Kengyilia guidenensis has 2 n — 42. It carries the
SPY combination of haplomes, typical of Kengyilia
by their relative chromosome length and arm ratio
(Fig. 2), similar to the karyotypes of K. gobicola
Yen & J. L Yang (Yen & Yang, 1990), K. habahe-
nensis Baum, Yen & J. L. Yang (Baum et al., 1990),
and K. nana J. L. Yang, Yen & Baum, K. tahela-
cana J. L. Yang, Yen & Baum, and K. zhaosuensis
J. L. Yang, Yen & Baum (Yang et al., 1992).

Acknowledgments. We are very grateful to J.
Cayouette, Centre for Land and Biological Re¬
sources Research, Ottawa, for comments on the
manuscript, especially for improving the Latin. We
are indebted to G. Q. Song, Dept. Biology, Nanjing
University (N) herbarium, for her facilitating our
examination of the holotype of K. kokonorica.

7<Y

o/»

'V* s

V v r - n

/V

/

Mu >

.<0
4 L?

}q |( |i IV CO 7Cfl

?? ij 7 { >( 3! H ?c

? I If f f •» < )\ « *'»

Figure 2. Karyotype of Kengyilia guidenensis Yen, J. L.
Yang & Baum. First row illustrates the P haplome; second
and third rows illustrate the S and Y haplomes mixed.

Literature Cited

Baum, B. R., C. Yen & J. L. Yang. 1990. Kengyilia ha-
bahenensis (Poaceae: Triticeae)—A new species from
the Altai mountains, China. PI. Syst. Evol. 174: 103-
108.

Yang, J. L., C. Yen & B. R. Baum. 1992. Three new
species of the genus Kengyilia (Poaceae: Triticeae) from
West China and new combinations of related species.
Canad. J. Bot. 71: 339-345.

Yen, C. & J. L. Yang. 1990. Kengyilia gobicola, a new
taxon from West China. Canad. J. Bot. 68: 1894—1897.

Volume 5, Number 1, pp. 1-118 of NOVON was published on 27 March 1995.
Volume 5, Number 2, pp. 119-208 of NOVON was published on 21 June 1995.
Volume 5, Number 3, pp. 209-304 of NOVON was published on 05 September 1995.
Volume 5, Number 4, pp. 305-398 of NOVON was published on 15 December 1995.