Volume 15
Number 1

2005

NOVON

Desideria mieheorum (Brassicaceae), a New Species from Tibet

Ihsan A. Al-Shehbaz

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

ihsan.al-shehhaz@mobot.org

Abstract. I lie new Tibetan Desideria mieheorum
(Brassicaceae) is described and illustrated. It dif¬
fers from the closely related D. Jlabellata by having
slender styles 5-7 mm long, entire stigmas, blue
petals, and 10 to 16 ovules per ovary. Desideria
Jlabellata has obsolete styles, prominently 2-lobed
stigmas, purple petals, and 14 to 24 ovules per loc-
ule.

Key words: Brassicaceae, China, Desideria, Tibet.

Desideria Pampanini is a genus of I 1 species
distributed in the Himalayas and adjacent central
Asia (Al-Shehbaz, 2001). 1 rom its nearest relatives,
Eurycarpus Botschantzev, Christolea Cambessedes,
and Ijeiopora (C. A. Meyer) Dvorak, Desideria is
easily distinguished by having its fruits rectangular
in cross section, dentate leaves, valves with prom¬
inent marginal veins, and valve apices persistently
united with the replum. A detailed account of these
genera is presented in Al-Shehbaz (2001) and is
not repeated here. China has eight species (two en¬
demic) in Desideria distributed in Qinghai, Tibet
(Xizang), and Xinjiang autonomous regions (Zhou
et al„ 2001).

I he new species was discovered among a collec¬
tion of 52 mustards kindly sent by Georg and Sa¬
bine Miehe (University of Marburg, Germany) for
determination. It is named in their honor and in
recognition of their extensive collections of alpine
Tibetan plants.

Desideria mieheorum Al-Shehbaz, sp. nov.
TYPE: China. Tibet (Xizang): Chang La, IN of
Sangsang, 29°41'N, 86°43'E, pioneer on frost
debris, 5480-5600 m, 15 Sep. 2003, Georg &
Sabine Miehe 03-112-21 (holotype, MO). Fig¬
ure 1.

Herba perennis dense pilosa. Folia late obovata vel spa-
t h n lata, 1.5—3 X 0.7— 1.5 cm, dentata. Racemi 5— 9-flori;
pedicel I i fructiferi tenues, 0.8-2.5 cm longi. Sepala ob-
longa, dense pilosa, 5-6 X 1.5-2 mm; pelala caerulea,
late spathulata, 10-13 X 3-1 mm; ovula 10-16. Fructus
lanceolati vel laneolato-lineari, 1.5—4 cm X 5—8 mm,
eompressi, dense pilosi; septum completum; stylus 5—7
mm longus, glaber; stigma integrum; semina ovata, 2.5-3
X 1.5-2 mm, biseriata; cotyledones accumbentes.

Perennial herbs 5-15 cm tall, densely pilose
throughout; trichomes simple, straight, 1—2 mm
long; caudex thick, covered with leaf remains of
previous years; stems simple. Leaves basal and
eauline, not fleshy; petiole 2—4.5 cm long, pilose;
leaf blade broadly obovate to spatulate, 1.5—3 X
0.7— 1.5 cm, pilose, base cuneate to attenuate, mar¬
gins dentate, apex acute. Kacemes 5- to 9-flowered,
ebracteate; fruiting pedicels slender, ascending,
straight to curved, 0.8— 2.5 cm long, spreading pi¬
lose. Sepals free, oblong, 5-6 X 1.5-2 mm, often
persistent to fruit maturity, densely pilose, base not
saccate, margins membranous; petals blue, broadly
spatulate, 10—13 X 3—4 mm, apex obtuse; claw 6—
7 mm long; filaments blue, slightly dilated at base,
median pair 5—6 mm long, lateral pair 3—4 mm
long; anthers oblong, 1—1.3 mm long; ovules 10 to
16 per ovary. Fruit lanceolate to lanceolate-linear,
1-5—4 cm X 5—8 mm, strongly flattened parallel to
septum; valves densely pilose, distinctly veined;
septum complete, membranous; style slender, 5-7
mm long, glabrous; stigma entire; seeds dark
brown, ovate, 2.5-3 X 1.5-2 mm, biseriate, mi¬
nutely reticulate; cotyledon accumbent.

Desideria mieheorum is readily distinguished
from all species of Desideria by its slender styles
5—7 mm long. In the remaining species, the style

Novon 15: 1-3. 2005.

2

Novon

Figure I. Desideria mieheorum Al-Shehbaz. — A. Plant. — H. Sepal. — C. Petal. — D. Dehisced fruit. Seale. A. D
1 cm; M, C = I mm. Drawn by Al-Shehbaz from the holotype (G. & .S’. Miehe O.'i-1 12-21, MO).

is obsolete or rarely (as in I). haranensis Al-Sheh¬
baz and / ). baiogoinenis (K. C. Kuan & Z. X. An)
Al-Shehbaz) reaches I to 1.5 mm in length. The
new species is most closely related to D. jlubelluta.

(Hegel) Al-Shehbaz, which it resembles in having
similar flower and fruit size, pilose indumentum
consisting of long simple trichomes, ebracteate ra¬
cemes. and leaves obovate to broadly spatulate. It

Volume 15, Number 1
2005

Al-Shehbaz

New Species from Tibet

3

differs by having slender styles 5—7 mm long, entire
stigmas, blue petals, and 10 to 16 ovules per ovary.
By contrast, D. flabellata has obsolete styles, prom¬
inently 2-lobed stigmas, purple petals, and 14 to
24 ovules per locule.

As in all other species of Desideria, /). mieheo-
rum has fruits that readily detach at maturity from
the fruiting pedicels. T he fruits have papery valves
that readily dehisce at base but remain united with
the replum at apex. Therefore, the detached fruit
acts as a dispersal unit, and it releases the seeds
as it tumbles on the ground.

Acknowledgments. I am most grateful to Georg

and Sabine Miehe for sending their Tibetan collec¬
tions of Brassicaceae for my determinations. I thank
Victoria Hollowed and an anonymous reviewer for
their critical comments on the manuscript.

Literature Cited

Al-Shehbaz, I. A. 2001. A review of gamosepaly in ihe
Brassicaceae and a revision of Desideria, with a critical
evaluation of related genera. Ann. Missouri Bot Card
87: 549-563.

Zhou, T. Y., L. L. Lu, G. Yang & 1. A. Al-Shehbaz. 2001.
Brassicaceae. In: Z. Y. Wu & P. H. Raven (editors),
flora of China 8: 1-193. Science Press, Beijing, and
Missouri Botanical Garden, St. Louis.

Thirty-Five New Species of Eugenia (Myrtaceae)
from Mesoamerica

Fred R. Barrie

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Address for correspondence: Botany Department, llie Held Museum,

1400 S. Lake Shore Drive, Chicago, Illinois 60605, U.S.A. fred.barrie@mobot.org

Abstract. Preparation of a treatment of Eugenia
(Myrtaceae) for the Flora Mesoamericarui resulted
in the discovery of many taxa heretofore unde¬
scribed. Mere, 35 new species from all parts of Me-
soamerica arc described and illustrated: Eugenia
belloi, E. breedlovei, E. cararaensis, E. cerroca-
caoensis, E. chavarriae, E. cintalaparui, E. cocosen-
sis, E. coibensis, E. corusca, E. esteliensis, E. fari-
nacea, E. gomezii, E. grayumii , E. grijalvae, E.
hammellii , E. hartshornii , E. herrarae, E. intibu-
cana, E. lempana, E. liesneri, E. lithosperma, E.
locuples, E. magnijlora, E. mcphersonii, E. molinae,
E. monleverdensis, E. paloverdensis, E. quercetorum,
E. riosae, E. sarwarlosensis, E. selvana, E. shinnsb-
ito, E. tilarana , E. verruculata, and E. zuchowskiae.

Key wards: Eugenia, Mesoamerica, Myrtaceae.

The Flora Mesoamericarui Project has stimulated
an increased interest in Central American Myrta¬
ceae. Recent research has resulted in the discovery
of one new genus (Chamguava, Landrum, 1991)
and numerous now species in established genera
such as Mosiera (Landrum, 1992), Calyptranthes
and Myrcia (Moist, 1999; Moist & Kawasaki, 2004),
and Blinia (Barrie, 2004).

This paper presents descriptions and illustra¬
tions for 35 new species of Eugenia. The type col¬
lections of the majority, 21, were made in Costa
Rica. An additional 6 come from Honduras, 3 from
Nicaragua, 2 each from Panama and Chiapas, Mex¬
ico, and I from Kl Salvador.

This publication increases to 140 the number ol
Eugenia species known from Mesoamerica. A com¬
prehensive identification key will be published in
volume four of Flora Mesoamericana.

Kugcnia belloi Barrie, sp. nov. TYPh: Costa Bica.
Alajuela: Reserva Forestal de Arenal, Quebra-
da San Gerardo, Rfo Cano Negro, finea de
Lnrique Ouesada y Ceferino Gonzales, 19 tel).
RXX) (hr), E. Bello I92() (holotype, 1NB; iso-
types, I', MO). Figure 1.

Eugeniae skulchii aemulans, sc<l foliis coriaceis, nervis

Novon 15: 4-49. 2005.

paucioribus et basibus late cuneatis vel rotundatis, flori-
bus majoribus differt.

Trees to 30 m X 20 cm; branchlets, emergent
leaves, buds, inflorescences, and young fruits ap-
pressed -pubescent with rufous, dibrachiate, and in¬
fundibular hairs; branchlets compressed; bark red¬
dish. Leaves e lliptic, the blades 8—17 X 3.5— 6.5
cm, 1.5— 2.5 times as long as wide, coriaceous, dry¬
ing dark, dull green above, dull, yellowish brown
below; midvein impressed above, raised below, ta¬
pering distally; lateral veins 12 to Ml per side, plus
intermediates, raised on both surfaces; marginal
veins similar to the laterals and weakly arched be¬
tween them, 2—3 mm from the margins; upper sur¬
face ultimately glabrate, the midvein persistently
and densely pubescent with sordid or cinereous
hairs, the lower surface and veins persistently and
densely covered with yellow-brown, infundibular
hairs; glands on the upper surface numerous and
minute or these obscure, glands on the lower sur¬
face present but obscured by the vestiture; base
broadly cuneate or rounded; margins recurved, oi-
ten revolute near the base, merging into the inner
edge of the petiole; apex acute or acuminate; pet¬
ioles 10—20 mm long, channeled dorsally, ap-
pressed-pubescent. Inflorescences axillary, race¬
mose, solitary; axis 5—10 X 2—3 nun, terete; bracts
1-1.5 X 1 nun, persistent, cymbiform, pubescent
on both surfaces, the margins searious, the apex
acute; buds not seen. Flowers pedicellate, the ped¬
icels, bracteoles, hypanthium and calyx uniformly
appressed-pubescent; pedicels stout, 7—10 X 2
mm; bracteoles fused into a cymbiform involucre,

3— 5 mm diain., pubescent on both surfaces, the
margins searious, the apex rounded; hypanthium
globose, 4-5 mm diam., glandular; calyx lobes in
subequal pairs, depressed ovate, 4—6 X 4—7 mm,
pubescent on both surfaces; petals not seen; disk

4— 5 mm diam., pubescent with erect, rufous hairs;
stamens ea. 2(H), 5-6 mm long; style 6 mm long,
glabrous. Fruits 30-40 X 30-40 mm, globose; peri¬
carp fleshy, ca. 10 mm thick, the surface verrucose
but not costate, uniformly and minutely pubescent.

Mouy

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

5

Figures 1, 2. —1 (left). Eugenia belloi Barrie (. E . Bello 1926 ; holotype, 1NB). —2 (right). Eugenia breedlovei Barrie ( Breedlove & R. Thorne 30778: holotype.

6

Novon

the braeteoles persistent, the involucre separating
along the marginal sutures, widely ovate, 2.5 X 5
mm; calyx erect in developing fruit but probably
absent at maturity; mature fruit tan.

Habitat, distribution, and phenology. Known
from the wet montane forests along the Atlantic
slope, Costa Rica, at elevations of 700— 1300 m.
Collected in flower in December; collected in Iruil
February-April.

Eugenia belloi is similar to E. skutchii C. V. Mor¬
ton & Standley. Rolli are large trees with leaves of
similar size and overall shape and the underside
uniformly covered with persistent, infundibular
hairs. However, the leaves of E. belloi are coria¬
ceous, with 12 to 18 pairs of lateral veins, plus an
indeterminate number of less well-defined inter¬
mediate veins and with the base broadly cuneate
to rounded. In A. skutchii, the leaves are charta-
ceous, with 16 to 20 well-defined lateral veins and
cuneate leaf bases. Moreover, the flowers of E. skut¬
chii are smaller; the hypanthium is 2-4 mm long
and campanulate rather than globose, and costate
rather than unribbed; the opposing pairs of calyx
lobes are markedly unequal in size, with the lobes
of the smaller pair 2-8 X 2-5 mm and those of the
larger pair 3-4 X 4—6 mm; the disk is 3-4 mm in
diameter (vs. 4—5 mm in E. belloi)-, arid the style,
which is glabrous in E. belloi, bears a few contorted
hairs.

Etymology. Named in honor of Eric Hello, not¬
ed authority on the vegetation of the Monteverde
region, coauthor of An Introduction to the Cloud
Eorest Trees of Monteverde, and the collector of the
type.

Paralypes. COSTA RICA. Guanaeaste: San Gerardo
Abajo, Rfo Cano Negro, l i liras Quesada v Arce, E. Hello
& E. Cruz 4277 (I IN It); Tilaran, Cordillera de Tilaran. I'al-
mital, Quebrada Malanga, Rfo Cano Negro, E. Hello <£■ E.
Cruz 4890 (E, INR. MO); Upper Rfo Chiquito valley, 5 km
N of Santa Elena, Atlantic slope rain forest, W. 4. Haber
et al. 4877 (CR, K).

Eugenia breedlovei Barrie, sp. nov. 1 't PE; Mex¬
ico. Chiapas: Barriozabal, 13 km N of Ber-
riozabal near Pozo Turipache & finca El Sus¬
pire, 25 Dec. 1972 (fr). I). E. Breedlove tA R.
Thorne 30778 (holotype, DS; isotypes, E.
MEXU, MICH, MO. NY, TEX). Figure 2.

Ah Eugenia uliginosa inflorescentiis axillaribus, flori-
bus et fructibus non stipitatis, pedicellis brevioribus dif-
fert.

Trees to ca. 10 m tall; branchlets, buds, emergent
leaves, and inflorescences minutely puberulent, the
hairs ca. 0.1 mm, erect, cinereous; branchlets com¬

pressed at nodes; bark gray. Leaves ellipt ic, the
blades 13-23 X 5.5-11 cm, 2-2.8 times as long
as wide, coriaceous, drying olive above, lighter
green below; midvein Hat or weakly convex on the
upper surface, elevated on the lower surface; lateral
veins 14 to 20, plus intermediates, raised on the
upper and lower surfaces, downwardly curved at
the midvein; marginal veins straight, 1-2 mm from
tht‘ margins; upper and lower surfaces glabrate or
with a few hairs near the base, the midvein glabrate
above, minutely puberulent below; glands obscure
on the upper surface, scattered to numerous dark
or pellucid glands below; base cuneate; margins
revolute; apex bluntly acute; petioles 10-17 mm
long, corky, scaly, or fissured transversely, flattened
or depressed dorsally, minutely puberulent. Inflo¬
rescences cauliflorous, solitary, racemose; axis 1— 6
X 2-4 mm. stout, about as wide as long; flowers 6
to 10; bracts 0.5 mm long, persistent, triangular,
glabrate in fruit, the margins ciliolate, the apex
acute; buds 7-10 mm long, obovoid. Flowers ped¬
icellate but sometimes appearing sessile, the ped¬
icels 1 X 2—3 mm; bracteoles 0.5—1 X 0.5—1 mm,
persistent, similar to the bracts, the base free, tlx*
margins ciliolate, the apex acute or rounded; hy¬
panthium 3—5 mm, campanulate or ureeolate, mi¬
nutely puberulent; calyx lobes in unequal pairs, tlx*
inner aixl outer surfaces puberulent, the margin cil¬
iolate, the apex rounded, the smaller pair 3—4 X
4—5 mm, the larger pair 4—5 X 5—6 mm; petals not
seen; disk 4-5 mm diam., minutely puberulent; sta¬
mens 180 to 200, 7—10 mm long; style El — 16 mm
long, puberulent. Fruits 18—30 X 18—30 mm, glo¬
bose or subglobose. constricted apically beneath
the persistent calyx; pericarp thick, fleshy, glan¬
dular, glabrate or puberulent al the base and apex;
immature color yellow-green.

Habitat, distribution, and phenology. Lower
montane rain forests, Chiapas, Mexico, at eleva¬
tions of 500-1000 m. Collected in flower March-
May; collected in fruit September-December.

Specimens of Eugenia breedlovei have been
identified as E. uliginosa Lundell, which has leaves
half the size of this species, cauliflorous inflores¬
cences, and flowers and fruits with longer pedicels
(3-5 mm). In E. uliginosa. the base of the hypan¬
thium and of the fruit is elongated to form a stipe
3—4 mm long.

Etymology. The epithet honors Dennis Breed¬
love. authority on the Chiapan flora and tlx* found¬
ing editor of the Elora de Cluapas. I he type col¬
lection is but one of his tens ol thousands from the
slate.

Paralypes. MEXICO. Chiapas: M| )io. Orozocoautla

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

7

de Espinosa, just above the community & lake of Malpaso,
45 km N of Ocozocoautla, I). E. Breedlove 20734 (MICH.
MO, NY); 13 km N of Berriozabal. I). E. Breedlove 2 / 053
(MEXU, MO); mpio. Berriozabal, 13 km N of Berriozabal.
near Pozo Turipaehe & Finca El Suspiro, D. E. Breedlove
35317 (DS); mpio. Ocozocoautla, 46 km N of Ocozocoau¬
tla on road to Mai Paso, I). E. Breedlove 38285 (MEXU,
MO, TEX); mpio. San Fernando, steep slopes with sea¬
sonal evergreen forest at the tunnel on the road from Tux-
tla Gutierrez to the Chicoasen Dam, I). E. Breedlove
40029 (DS). I). E. Breedlove 41549 (DS); mpio. Ocozo¬
coautla de Espinosa, Canada La Palma, SE del Cerro el
perico 1.5 km, A. M. A. 9541 (CAS); mpio. Ocozocoautla
de Espinosa, El Ocote, Canada el Palmar, M. Gonzdlez-
Espinosa et al. 1 181 (MEXU); Noroeste de San Fernando,
E Miranda 5244 (MEXU): Encanada de La Chacona a El
Aguacate, al Noroeste de Tuxtla Gutierrez, F. Miranda
0128 (MEXU, MICH); Finca La Chacona (Noroeste Tuxtla
Gutierrez), E Miranda 6869 (MEXU); mpio. San Fernan¬
do, Canada Muniz, G. Rodriguez- ( hiilldn 107 (CAS); mpio.
Ocozocoautla de Espinosa, SW side of the Presa de Mal¬
paso, A. S. Ton 3320 (DS. F, LL. MICH, NY).

Eugenia oararaensis Barrie & Q. Jimenez, sp.
nov. TYPE: Costa Rica. Puntarenas: Reserva
Bioldgica Carara. Montanas Jamaica, Sendero
a hijagual, 27 Sep. 1990 (ft), (). Jimenez 909
(holotype, INB; isotype, F). Figure 3.

Ab Eugenia truncata ramulis el foliis glabris, floribus
et fructibus minoribus, ealycibus pallidis differt.

Irees to 15 m X 20 cm; young shoots apparently
minutely puberulent. the hairs ca. 0.2 mm, erect,
glabrous otherwise except for the ciliate margins of
the bracts and bracteoles and the densely pubes¬
cent staminal ring; branchlets terete or compressed;
bark tan or brown soon glabrate or with patches of
persistent, erect hairs. Leaves elliptic to obovate,
the blades 8—20 X 3—8 cm, 2—2.8 times as long as
wide, ehartaeeous, drying a eoncolorous dull, lighl
green; midvein convex, typically between parallel
grooves and ridged above, convex below; lateral
veins 10 to 18 per side, plus intermediates; mar¬
ginal veins similar to the lateral veins, weakly
arched, 1—3 mm from the margins; glands numer¬
ous and raised above, scattered and dark below;
base cuneate; margins somewhat revolute, decur¬
rent into the raised inner edge of the petiole; apex
abruptly acuminate or acuminate-apiculate; peti¬
oles 4—5 mm long, impressed dorsallv. Inflores¬
cences axillary or terminal, solitary, fasciculate or
short-racemose, the axis less than 2 mm long; flow¬
ers I to 10; bracts 0.5 mm, persistent, cymbiform,
the margins scarious or scarious-ciliate, otherwise
glabrous, the apex acute; buds not seen. Flowers
pedicellate, the pedicels 10-40 mm long, graeile,
conspicuously glandular; bracteoles 0.5—1 X 0.5—

I mm, persistent, ovate, the base free, the margins
scarious or scarious-ciliate, the apex rounded to

acute; hypanthium 1.5 mm long, globose; calyx
lobes in equal pairs, 2.5-3 X 2.5-3 mm, pale in
color with conspicuous dark glands, the apex
rounded, the margins entire; petals not seen; disk
2.5-3 mm diam., the staminal ring densely pubes¬
cent. the hairs 0.3 mm; stamens ca. 80 to 100, 5-
6 mm long; style 5 mm long, glabrous. Fruits 10-
20 X 10—20 mm, globose; pericarp thin-walled,
conspicuously glandular, glabrous; calyx persistent,
erect, pale; mature fruits dark red.

Habitat, distribution, and phenology. Primary
forests of the Pacific slope in Costa Rica at eleva¬
tions ol 0—300 m. Collected in flower December—
June; collected in fruit August-Oetober.

Eugenia cararaensis is distinguished by the ax¬
illary, fasciculate or short-racemose inflorescences
with flowers and fruits borne on graeile pedicels
10—10 mm long. The persistent, pale calyx lobes
offer a color contrast to the dark red body of the
fruit. I his species is similar to E. truncata 0. Berg,
a species with hirsute young growth and flowers
typically with one or both bracteoles attached 1—5
mm beneath the hypanthium.

Paratypes. COSTA RICA. Puntarenas: Palmar Nor¬
te, el Bosque lluvioso de las montanas bajas cerca de la
quebrada Benjamin, al norte del pueblo, M. Bohlke 595
(CR); PN Corcovado, Carate, Est. La Leona. Bosque ale-
daiio. ,4. Cliacdn 990 (INB); Golfito. Jimenez. Alto de Car-
bonera, camino a Cerro Osa, G. Herrera 4311 (F, INB);
Golfito PN Corcovado, Peninsula de Osa, Sirena, Bosque
primario, 6. Maass 41 (1. INB); R. K Golfo Dulce, pen¬
insula de Osa. Rancho Quemado, Finca de Juan Marin,
Cerca a Guerra, J. Marin 83 (F, INB); Osa. Este “ro Guerra,
Sierpe, f ineas que se encuentran al E del Estero, ./. Mann
173 (F. INB); Reserva Biologica Carara, Sector Quebrada
Bonita, Sitio Quebrada Bonita, R. Zuniga 311 (F. INB).
San Juan: Reserva Bioldgica Carara, Sitio Sendero Lalo
Barboza, It. Zuniga &: G. Varela 288 (F, INB).

Eugenia cerrocacaoensis Barrie, sp. nov. TYPE:
Costa Rica. Guanacaste: Liberia. Parque Na¬
tional Guanacaste, Cordillera de Guanacaste.
Estacion de Cacao, ID Feb. 1995 (II. fr), E.
Alfaro 92 (holotype, INB; isotypes, CR. F. MO.
SEE). Figure 4.

Ab aliis Eugeniae costaricensibus foliis brevioribus, el-
lipticis vel obovatis, coriaceis, floribus pusillis differt.

Small trees or shrubs, to 5 m; young growth gla¬
brous except for the ciliate margins of the bracts,
bracteoles, and calyx; branchlets stout, terete; bark
gray, fissured horizontally. Leaves elliptic or ob¬
ovate. the blades 2-5 X 1.5—3 cm. 1.5—2 times as
long as wide, coriaceous, drying brown above, tan
or olive below; midvein sulcate above, convex be¬
low; lateral veins 6 to 8, plus a few intermediates,
obscure on both surfaces or prominent below; mar-

8

Novon

gures 3, 4. — 3 (left). Eugenia cararaensis Barrie & Q. Jimenez (, Bohlke 595; paratype, CH). — f (right). Eugenia cerrocacaoensis Barrie (Alfaro 92; holotype. INB).

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

9

ginal veins similar to the laterals and arched be¬
tween them, 1-2 mm from the margins; glands ob¬
scure above, numerous and dark below; base
cuneate; margins revolute, if only on the lower half
of the blade, decurrent along the petiole; apex
bluntly acute or rounded, the tip ultimately round¬
ed or retuse; petioles 3-4 mm long, flattened dor-
sally. Inflorescences axillary or cauliflorous, race¬
mose, solitary; axis 1—5(8) mm long; flowers 2 to 8;
bracts 0.5-1 mm long, persistent through anthesis,
ovate, the margins eiliate, the apex acute; buds 2
mm long, globose. Flowers pedicellate, the pedicels
1-2 mm long, slender in flower, stout in fruit; brae-
teoles persistent, ovate, fused and forming an in¬
volucre I mm diam. in bud and flower, reduced and
separate in fruit, the margins scarious-ciliolate, the
apex rounded; hypanthium 1 mm diam., globose;
calyx lobes depressed ovate, in subequal pairs, 0.5
X I mm, the margins scarious-ciliolate, the apex
rounded; petals not seen; disk 1 — 1.5 mm diam.;
stamens 25 to 40, 3—4 mm long; style 4 mm long,
glabrous. Fruits 10-15 X 8-10 mm, ellipsoid to
obovoid; pericarp thin-walled, glandular; calyx ab¬
sent; mature color red.

Habitat, distribution, and phenology. All known
collections are from secondary forests on Cerro Ca¬
cao. Guanacaste Province, Costa Rica, at 1100 m
elevation. Collected in flower in February; collected
in fruit February-November.

Eugenia cerrocacaoensis is a distinctive small
tree or shrub with stout branchlets, elliptic to ob-
ovate, coriaceous leaves with bluntly acute or
rounded apices and small flowers, with a globose
hypanthium ea. I mm in diameter, small, depressed
ovate calyx lobes and a disk 1—1.5 mm in diameter,
on short pedicels, 1-2 mm long. Vegetatively, E.
cerrocacaoensis resembles the coriaceous-leaved
form of Siphoneugena dussii (Krug & Urban) Proen-
ya. However, in that species the flowers are borne
in fascicles or on a raehis less than 1 mm long, the
pedicels are variable in length, up to 6 mm long,
and the calyx is fused and poricidal in bud.

Paratypes. COSTA RICA. Guanacaste: Liberia, Ks-
tacion Cacao, PN Guanacaste, R. Espinoza 55 (CR, INR);
I’N Guanacaste. Kstaeion Cacao Cordillera de Guanacaste.
E Quesada 3 (INB).

Eugenia chavarriae Barrie, sp. nov. TYPE: Costa
Rica. Guanacaste: Reserva Bioldgica, Lomas
de Barbudal, Valle del Tempisque; Lomas Bar-
budal, horde de camino al Ref. Rafael Lucas
y orilla del R. Gabuyo. Sendero Bagaces, 25
Mar. 1993 (fr), U. Chavarria 760 (holotype,
INB; isotypes, CR, F, MO). Figure 5.

Eugeniae principium aemulans, sed foliis nervis later-
alibus obscuris superficialibus, inflorescentiis subsessili-
bus, fruetibus 4—6 mm diametro differt.

Trees to 6 m or more; branchlets terete, puber-
ulent. the hairs 0.2 mm, simple, cinereous, erect or
recurved. Leaves elliptic and tapering symmetri¬
cally to base and apex or slightly obovate, the
blades 2.5— 6.5 X 1—3 cm, 2.3—3 times as long as
wide, coriaceous, drying dark green above, paler
below; midvein concave or flat above, convex be¬
low; lateral veins 4 to 6 per side, straight and as¬
cending, obscure above; marginal veins similar to
the laterals, straight or weakly arched, 0.5—1 mm
from the margins; both surfaces with scattered,
erect or recurved hairs, ultimately glabrate; glands
small, numerous on both surfaces or obscure above;
base cuneate; margins decurrent into the petiole;
apex bluntly acute; petioles 2-4 mm long, flattened
or concave dorsally, glabrous or pubescent. Inflo¬
rescences axillary or cauliflorous, short -racemose,
solitary; axis ca. 1 mm long, flowers I to 4; bracts
0.5 mm, persistent, elliptic, puberulent, the mar¬
gins eiliate, the apex acute; buds 3 mm long, ob¬
ovoid. puberulent. Flowers pedicellate, the pedicels
6-10 mm long, gracile, the pedicels, braeteoles, hy¬
panthium, and calyx puberulent, the hairs 0.2 mm,
cinereous, erect or recurved; braeteoles persistent,
0.5—1 mm long, ovate, the base free, the margins
eiliate, the apex acuminate; hypanthium 1-1.5 mm
diam., globose; calyx lobes in subequal pairs, 0.5—

1 X 1 mm, the margins eiliate, the apex acute or
acuminate; petals 3-3.5 X 2-2.5 mm. elliptic or
ovate, the margins eiliate, the apex rounded; disk
1.5-2 mm diam., puberulent; stamens 30 to 40, 3-
4.5 mm long; style 4—5 mm long, glabrous. Fruits
4-6 mm diam., globose; pericarp thin-walled, con¬
spicuously glandular, glabrate or with a few scat¬
tered hairs; calyx persistent, inconspicuous; mature
color reddish purple.

Habitat, distribution, and phenology. Pacific
slope dry forests in northwestern Costa Rica at el¬
evations of 0—800 m. Collected in flower Decem-
ber-January; collected in fruit January-March.

Eugenia chavarriae is distinguished by the
leaves with short petioles and elliptic blades taper¬
ing symmetrically to the base and apex. The habit
and vestiture are similar to E. principium McVaugh,
a species scattered along the Pacific coast from Co¬
lima, Mexico, to Panama. In that species, however,
the blades of the leaves, while similar in size, have
a cuneate base, an acute or acuminate apex, and
persistent, recurved hairs along the margins on the
upper surface, 'flic flowers and fruits of both spe¬
cies are similar in size, but in E. principium the

4261401

10

Novon

Figures 5. 6. — 5 (left). Eugenia chavarriae Barrie (Chavarria 760: holotype, INB). — 6 (right). Eugenia cintalapana Barrie ( Breedlove 49604: holotype. MO).

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

11

calyx lobes are pink at anthesis and persistent and
erect on mature fruits.

Etymology. The epithet honors the collector of
the type. Ulises Chavarria, a parataxonomist with
the Institute Nacional de Biodiversidad, Costa
Rica, a park official at the Parque Nacional Palo
Verde, and the coauthor of Arboles Comunes del
Parque Nacional Palo Verde.

Paratypes. COSTA RICA. ( >uanacaste: Comelco
property, near Bagaces, K. S. Bawa 2054 (MO); OTS re¬
search area. Stewart property (28 km N of Gafias), G. If
Frankie 236 (F, MO); Sta. Rosa NR. 30 km NW of Liberia,
D. H. Janzen 10994 (MO), l). H. Janzen 11715 (MO), D.
//. Janzen 11759 (MO), D. //. Janzen 12169 (MO), D. 11.
Janzen 12423 (MO); PN Sta. Rosa, 2 km E of park head¬
quarters, A'. Liesner 4233 (CR, MO); Comelco property,
near Bagaces, Area E, Transect R2, P. A. Opler 502 (CR,
F); PN Sta. Rosa, L J. Poveda 332 (F); Santa Rosa Na¬
tional Park, Siempre Verde, G. Schofield 76-104 (CR);
lower slopes of Volcan Rincdn de la Vieja, along R. Col¬
orado, A. O. Williams et al. 26646 (CAS, CR, K|2|).

Eugenia cintalapana Barrie, sp. nov. TYPE: Mex¬
ico. Chiapas: Cintalapa de Figueroa, 5 km W
of Rizo de Oro along Mex. Hwy. 190, 20 Jan.
1981 (fr), D. E. Breedlove 49604 (holotype,
MO; isotype, CAS). Figure 6.

Eugeniae avicenniae aemulans, sed indumento tantum
simplici, foliis, floribus et fructibus parvioribus differt.

Shrubs to 7 m or more; branchlets and emerging
leaves and inflorescences minutely puberulent, the
hairs ca. 0.2 mm, erect, or plants glabrous; branch-
lets terete or flattened at the terminal nodes; bark
silvery-gray. Leaves narrowly rhombic or narrowly
elliptic, the blades 2.5-4 X 0.5-1 cm, 4-5 times
as long as wide, chartaeeous, drying a concolorous
green; midvein flat or slightly convex on both sur¬
faces. tan in color; lateral veins 5 to 8, barely dis-
cernable above, straight and ascending; marginal
veins straight, ca. 1 mm from the margins, obscure
above; both surfaces glabrate; glands numerous and
dark on the upper and lower surfaces; base narrow¬
ly cuneate; margins cartilaginous, decurrent into
the inner edge of the petiole; apex bluntly acute or
rounded; petioles 1—2 mm long, channeled dorsally,
glabrate. Inflorescences axillary or eauliflorous,
glomerate or short-racemose, solitary or 2 super¬
posed; axis to I mm, glabrous or minutely puber¬
ulent; flowers I to 6; bracts 0.5 mm long, persistent,
triangular, glabrate or witb a few persistent hairs,
the margins entire or ciliate, the apex acute; buds
1.5— 2.5 mm, pyriform. Flowers short-pedicellate,
the pedicels 1-3 mm long, glabrate or minutely pu¬
berulent; bracteoles 0. 2-0.5 X 0.2 mm. persistent,
triangular, glabrate or with a few' persistent hairs,
the base free, the margins scarious, the apex acu¬

minate; hypanthium 0.7-1 mm long, eampanulate;
calyx lobes in subequal pairs, 0.2— 0.5 X 0.6—1
mm. depressed ovate, conspicuously white at an¬
thesis, glabrate, the margins entire, the apex round¬
ed; petals 2—2.5 X 1 .5—2 mm, elliptic, the margins
entire, the apex rounded; disk ca. 1 mm diam., gla¬
brous; stamens 20 to 25, 2 mm long; style 4 mm
long, glabrous. Fruits 5-7 X 4-6 mm. globose or
subglobose; pericarp thin-walled, rarely obscurely
ribbed, prominently glandular, glabrous or with
scattered, appressed hairs; calyx persistent, erect;
mature color unknown.

Habitat , distribution, and phenology. Known
from tropical deciduous forests in eastern Oaxaca
and western Chiapas at elevations of 800-1000 m.
Collected in flower in October; collected in fruit in
January.

Collections of Eugenia cintalapana have been
identified as E. avicenniae Standley, a species of
western Oaxaca and Guerrero. That species, how¬
ever, is much more coarsely pubescent, w ith flowers
and fruits that are nearly twice as large as are found
in this species. The flowers of E. cintalapana are
similar in size to those of E. capuli (Schlechtendal
& Chamisso) Hooker & Arnott, a species of wetter
montane habitats ranging from eastern Mexico to
Guatemala, but E. cintalapana differs in having
bracteoles that are free at the base, rather than con¬
nate, and conspicuous, white calyx lobes that con¬
trast with the green hypanthium. In E. capuli , as in
most Eugenia, the calyx and the hypanthium are
concolorous.

Paratypes. MEXICO. Chiapas: mpio. Cintalapa de
Figueroa, 3 km W of Rizo de Oro, I). E. Breedlove 36751
(OS, MICH) , 1). E. Breedlove & F. Almeda 64899 (CAS,
CR, MICH). Oaxaca: slopes of Guiengola, E. J. Alexan¬
der 346 (MICH).

Eugenia cocosensis Barrie, sp. nov. TYPE: Costa
Rica. Puntarenas: Isla de Cocos, high plateau
SE of Chatham Bay, 9-1 1 Apr. 1979 (fr), R.
Foster 4129 (holotype, CR: isotypes, CAS, F,
MO). Figure 7.

Ab Eugenia pacifica foliis et floribus paucioribus, fruc--
tibus late ellipsoideis vel obovoideis differt.

Small trees or shrubs, 4-5 m; buds, inflores¬
cences, and emergent leaves thinly appressed-pu-
bescent with cinereous or pale coppery, dibrachiate
hairs, 0.2 mm long; branchlets compressed; bark
reddish, glabrate. Leaves lanceolate or ovate, the
blades 3.5—5 X 1—2 cm, 2.5—4 times as long as
wide, coriaceous, drying a concolorous olive, the
petioles and midveins reddish; midvein impressed
above, raised below; lateral veins 4 to 6, straight.

12

Novon

Figures 7. 8. — 7 (left). Eugenia cocosensis Barrie R. (Foster 4129: holotype, CR). — 8 (right). Eugenia roibensis Barrie (Buyer 1639: holotype, F).

Volume 15, Number 1

2005

Barrie

Eugenia from Mesoamerica

13

obscure on the upper surface; marginal veins sim¬
ilar to the laterals and arched between them, 1—2
mm from the margins; both surfaces glabrate;
glands numerous, conspicuous, usually strongly
punctate above, dark below; base cuneate or slight¬
ly rounded; margins revolute near the base, decur¬
rent into the inner edge of the petiole; apex acu¬
minate; petioles 2-3 mm long, channeled dorsally,
glabrate. Inflorescences axillary, racemose, solitary,
appressed-pubescent with pale coppery, dibrachia-
te hairs; axis 1—3 mm long; (lowers 2 to 6; bracts
0.5 mm long, persistent, ovate, the margins scari-
ous, the apex acute; buds 2 mm long, turbinate.
Flowers pedicellate, the pedicels 2 mm long, ap-
pressed-pubescent; bracteoles fused, forming an in¬
volucre 1 mm diam., dilated at anthesis, appressed
pubescent on the outer surface only, the margins
scarious, the apex rounded; hypanthium 1 mm long,
obconic, appressed pubescent; calyx lobes in un¬
equal pairs, 0.5—1 X 1 mm, elliptic, appressed pu¬
bescent on the outer surface, the margins scarious-
eiliate, the apex rounded; petals 1 X I mm, ovate,
tin* margins entire, the apex bluntly acute; disk ea.

1 mm diam., glabrous; stamens 25 to 30, 1 mm
long; style 1.5 mm long, glabrous. Fruits 9—10 X
7—8 mm, widely ellipsoid or obovoid; pericarp thin-
walled. glabrous, glandular; calyx persistent, erect
but inconspicuous; mature color black.

Habitat, distribution , and phenology. Known
only from Isla de Cocos, Costa Ivica, in wet forests
at elevations of 100—500 m. Collected in fruit in
April.

Apparently endemic to the Isla de Cocos, Eu¬
genia cocosensis is similar to E. pacifica Bentham,
which is also endemic to the island. Phis species
has leaves with blades hall the si/e of E. pacifica,
with petioles about one-third as long, 2—3 mm ver¬
sus 6—10 mm. The flowers of E. pacifica are twice
the size of those of E. cocosensis, and the ellipsoidal
fruits are longer, 10—13 mm, and narrower, 4—6
mm.

Paratype. COSTA RICA. Puntarenas: Cocos Island,
high plateau SK of Chatham bay. It. lb Foster 41 73 (K,
OH, MO).

Eugenia eoibensis Barrie, sp. nov. TYPE: Pana¬
ma. Veraguas: Isla de Coiba, 19 Aug. 1961 (fl).
./. D. Dwyer 1639 (holotype, F; istoype, (ill).
Figure 8.

Habitu cum Eugenia rhombea similis, sed indumento
persistenti, pedicellis brevioribus et fructibus grandiori-
bus differt.

Shrubs to 5 m; young growth tomentose with

coarsely appressed, simple and dibraehiate hairs,
coppery or rufous, becoming olive or gray with age;
branchlets terete; bark reddish, with persistent ap¬
pressed and erect hairs. Leaves ovate, elliptic or
obovate, the blades 4-7 X 2—3.5 cm, 2—3 times as
long as wide, chartaeeous, drying dark green or
brown above, gray or olive from the persistent hairs
below; midvein sulcate to impressed above, elevat¬
ed below; lateral veins 5 to 7 per side, often ob¬
scure above, slightly raised below; marginal veins
similar to the laterals and weakly arched between
them, 1—2 mm from the margins; upper surface with
the hairs appressed, cinereous, persistent but thin¬
ning with age, the lower surface persistently vested
with a dense covering of gray or olive dibraehiate
and infundibular hairs; glands numerous and mi¬
nute above, somewhat larger below, visible through
the vestiture; base cuneate; margins revolute near
the base, decurrent along the inner edge of the pet¬
iole; apex acuminate or caudate-acuminate; peti¬
oles 5—7 mm long, channeled dorsally, appressed-
pubescent. Inflorescences axillary, racemose;
solitary or 2 superposed, axis l^f mm long, cop¬
pery appressed-pubescent; flowers 2 to 6; bracts ea.
0.5 mm long, persistent but reduced in fruit, ovate,
vestiture similar to the axis and pedicels in early
bud, thinning by anthesis, prominently glandular;
buds 2 mm long, obovoid. Flowers pedicellate, the
pedicels 3—6 mm long, coppery appressed-pubes¬
cent; bracteoles similar to the bracts, persistent, the
pair usually fused and forming a well-defined in¬
volucre ea. 1 mm diam., dilated at anthesis, ap-
pressed-pubescent on the outer surface, the hairs
thinning by anthesis, coarsely glandular, the mar¬
gins eiliolate, the apex acute; hypanthium 1.5 mm
long, campanulate, coarsely coppery appressed-pu¬
bescent; calyx lobes in markedly unequal pairs, el¬
liptic, appressed-pubescent on both surfaces, the
margins eiliate, the apex rounded, the smaller pair
1 — 1.5 X 1 — 1.5 mm, the larger 2— 2.5 X 2—3 mm;
petals 1—4.5 X 3—4 mm, elliptic, the margins cil-
iate, the apex rounded; disk 2—2.5 mm diam.,
sparsely pubescent near the style base; stamens ca.
50, 5—6 mm long; style 5—6 mm long, typically with
a lew scattered hairs. Fruits 6—9 X 6—9 mm, glo¬
bose or ellipsoid; pericarp thin-walled, glandular,
persistently pubescent; calyx persistent, the lobes
curled; mature color purple-black.

Habitat, distribution , and phenology. Known
only from coastal scrub forest on the Isla de Coiba,
off the Pacific coast of Panama, at elevations below
100 m. Collected in flower August— January; col¬
lected in fruit October— March.

Eugenia eoibensis is distinguished by the persis-

14

Novon

tent pubescence, coppery on yownp; growth but
graying with age. The underside of the leaves have
a dense covering of dibrachiate and infundibular
hairs, much like that of the Costa Rican species E.
lepidota (). Berg and E. belloi. The habit and leaf
shape are similar to those of E. rhombea (0. Berg)
Krug & Urban, a species widely distributed in
coastal areas around the Caribbean basin. Unlike
E. coibensis, E. rhombea is glabrous except for scat¬
tered, simple hairs on tin* flower parts and its leaves
are rigidly coriaceous.

Paratypes. PANAMA. Veraguas: Isla Coiba, desde el
Campamento de Las Salinas, 2 km al NO y posteriormente
al SSO hacia la cima de Cerro San Juan, ./. Aranda et al.
2196 (F); Montijo, Knsenada Santa Cruz, a lo largo de la
costa (derecha a izquierda), ./. Aranda et al. 2469 (US),
A. Espinosa et al. 7831MV (F); Rio Escondido, cerca de
la desenibocadura, junto al rfo, J. Aranda et al. MBI290
(F), A. Espinosa et at. 527MB (F); Santa Cruz, eamino de
el Catival, .S'. Castroviejo et id. 71 1 1 MV ( F); Punta Cirilo,
a 3 km de la desenibocadura del rfo, .N. Castroviejo et id.
13543 (F); playa del rfo de la Boa, ./. Cuadras et al.
79B9MV (F); borde del rfo Playa Hermosa, ./. Cuadras et
al. 803 1 MV (F); ensenada de Santa Cruz, A. Norte de la
isla, Orilla de playa en Ensenada Tigrito y Tigmn, C. Gal-
i lames et al. 2156 (F. MO); rfo Escondido, entre la primera
y la segunda cascada, E M. Garmandia A' M. Velayos
8973MV (F); Sendero de Sla. Cruz, ca. 500 m de la F.s-
taeidn Bioldgica, .4. Ibanez et al. 960AI (F); Isla de Coiba,
Rfo Negro, cauce principal, A. Ibanez et al. 8915MV (F).

Kiigcniu corusca Barrie, sp. nov. TYRE: Costa
Rica. Alajuela: San Ramon, Bosque Eterno ile
los Ninos, Cordillera de Tilaran, 5 km W of
Chachagua, Quebrada Chachaguita drainage,
17 July 1993 (fr), W. Haber et al. 1 1542
(holotype, INB: isotypes, CR, F, MO. SEL).
Figure 9.

Ab Eugenia sarapiquensi foliis subficialibus coruseis.
floribus el fructibus pedicellis differt.

Trees to 10 m; young growth densely appressed-
pubeseent with coppery, simple hairs, the stems
and upper surfaces of the leaves soon glabrate;
branchlets compressed; bark gray, smooth. Leaves
elliptic or obovate, the blades 7—15 X 2.5—5 cm,
2—3 times as long as wide, chartaceous, drying
green above and a glittery, silver-green below; mid¬
vein impressed or suleate above, raised below; lat¬
eral veins 5 to 9 per side, curving-ascending, raised
on the lower surface, interconnected by numerous,
netted veins; marginal veins often poorly defined or
obscure above, raised and similar to the laterals
below and strongly arched between them, 2-5 mm
from the margins; upper surface glabrate, the lower
surface persistently covered with a dense mat of
minute, simple, colorless or pale coppery hairs, the
individual hairs barely distinguishable, producing

a reflective, glittery surface; glands on the upper
surface numerous and small, glands below ob¬
scured by the vestiture; base cuneate or narrowly
cuneate; margins decurrent along the petiole; apex
acuminate lo caudate-acuminate; petioles 5—8 mm
long, depressed or suleate dorsally, glabrate or with
a few hairs. Inflorescences axillary or possibly cau-
liflorous, fasciculate; bracts not seen; buds not
seen. Flowers not seen. Fruits 20—25 X 25—40 mm,
irregularly obovoid or transversely ellipsoid; peri¬
carp relatively fleshy, glandular-rugose, glabrous;
pedicels 4—9 X 2 mm long, sometimes swollen be¬
neath the fruit, the bractcoles less than 0.5 mm.
persistent but reduced, triangular, the base free, the
apex acute; calyx apparently absent; mature color
orange-yellow.

Habitat, distribution, and phenology. Known
from wet forests along the Atlantic slope in Costa
Rica and Nicaragua at elevations of 100-1000 m.
Collected in fruit February-August.

Eugenia corusca is one of a group of Costa Rican
species, including E. sarapiquensis R. E. Sanchez,
E. grayumii, and E. gomezii, characterized by char¬
taceous, elliptic or ovate leaves with blades 10—25
cm long and curving ascending lateral veins. In this
species, the underside of the leaves is persistently
and densely covered with tightly oppressed hairs,
silvery or grayish green in color, producing a glit¬
tery, or coruscate, metallic sheen. In the other Costa
Rican species ol Eugenia in which the undersides
of tin1 leaves are persistently pubescent, the vesti¬
ture is dull and nonreflective.

Etymology. The epithet refers lo tin* coruscate
leaf undersides.

Paratypes. COSTA RICA. Alajuela: San Ramon, I .os
Angeles, Reserva Biologica de San Ramon, Las Rocas, 2
km al Oeste de Finca de Carlos Gonzdlez. C. Ilerrera &
V. Mora 6876 (CR); Cuatuso, Cordillera de Tilardn, 5 km
N del Lago Arenal, Cercanfas del Lago Coter, Sendero
Montana, Finca Coter-Lago Coter, ,/. Sanchez & A. Cas-
cante-M. 39 6 (CR, F). Giianueaste: Tilardn, Zona Pro-
teetora Tenorio, Tierras Morenas, Rfo San Lorenzo, C. Al¬
varado 131 (CR[2], INB); Tilardn, Corillera de Tilardn, I-
2 km Vi of Lago de Cote, 13 km N of Tilardn, Continental
Divide on SE slope of Volcdn Tenorio, Eeo-lodge, IT Haber
& W. /in lion ski 11634 (CR, F. INR); Tilaran ZP Tenorio,
Cordillera de Guanacaste, Tierras Morenas. alrededores
Rfo San Lorenzo, C. Rodriguez 148 (CR, F, INR. MO).
Heredia: PN Rraulio Carillo, Puesto El Ceibo, gently
sloping area just above the steep rim of the Rfo Peje gorge
on the W side, along side path that goes from guard house
to water supply intake, about 15 minutes uphill, B. Boyle
1395 (F. INR); Sarapiquf, l.a Selva Riological Station, lie-
side CC trail at 1200 m, tropical wet forest, on well-
drained upland soils, tenth hectare transect LS-2. B. Boyle
5280 (INR); Sarapiquf, Finca La Selva, Puerto Viejo, lean¬
ing over Quebrada Esquina. ea. 2 km upstream from S
boundary of La Selva, G. .S’. Hartshorn 1315 (CR, F, MO).

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

15

Figures 9, 10. —9 (left). Eugenia corusca Barrie ( Haber et al. 11542 ; holotype, 1I\B). —10 (right). Eugenia esteliensis Barrie ( Neill I22H ; holotype, MO).

16

Novon

1 .ilium: hills .'i.S kin (air) S of Islas Buena Vista on the
Kfo Colorado, 16 km (air) S of Barra del Colorado, G.
Davids,- & G. Herrera 31176 (MO). NICARAGUA. Rio
San Juan: Castillo, Keserva Indio-Mafz, a 8 km de la
eaheeera del R fo Bartola, en direecion haeia el Cerro el
Diablo, R. Rueda el al. 5288 (F, MO); Castillo, Keserva
Indio-Maiz, Cerro el Diablo, R. Rueda el al. 5441 (MO);
Castillo, Keserva Indio-Mafz, a lo largo del Cano Chon-
talefto, R. Rueda el al. 6163 (F, MO).

Eugenia esteliensis Barrie, sp. nov. TYPE; Nica¬
ragua. Estelf: summit of Cerro Quialm. 7 km
W of Kstelf, 2 Nov. 1976 (IV), 11 Neill 1228
(holotype, MO; isotypes, MICH, NY). Figure
10.

Ab Eugenia guatemalensi loliis august ioribus, indu¬
menta eupreo differ!.

Trees to 20 m; the young branches, leaves, buds,
and inflorescences appressed-pubescent with cop¬
pery, dibraehiate hairs; branchlets weakly com¬
pressed; bark reddish, glabrate or with some per¬
sistent hairs. Leaves elliptic or obovate, the blades
4-7 X 1-2 mm, 8—1' limes as long as wide, coria¬
ceous, drying green above, coppery from the ap-
pressed hairs below, eventually glabrate and gray;
midvein silicate above, elevated below and tapering
distally; lateral veins in 6 to 12 pairs, straight and
ascending; marginal veins similar to the lateral
veins, straight or weakly arched, ea. I mm from the
margins; upper surface of mature leaves glabrate or
with a lew, scattered coppery hairs, the lower sur¬
face initially densely coppery, appressed-pubes¬
cent, glabrate with age, the hairs commonly patch¬
ily deciduous; glands numerous above and below;
base narrowly cuneate; margins revolute, somewhat
thickened, deeurrent into the splayed inner edges
of the petiole; apex caudate-acuminate, the acumen
10—15 mm long; petioles V) — 1 2 mm long, chan¬
neled. Inflorescences axillary, racemose, solitary or

2 or 3 superposed; axis 8-10 mm long, coppery
appressed-pubescent in flower, glabrate in fruit;
flowers 5 to 9, a terminal flower usually present;
bracts 0.5 mm long, persistent, triangular, coppery
appressed-pubescent, the margins eiliate, the apex
sharply acute; buds 3—3.5 mm long, obconic, cop¬
pery appressed-pubescent. Flowers pedicellate, the
pedicels 2—6 mm long; braeteoles 0.3—1 mm long,
persistent, triangular, coppery appressed-pubes¬
cent, the base free, the margins eiliate, the apex
sharply acute; hypanthium ea. 2 mm long, obconic
or campanulate, sometimes weakly costate, coppery
appressed-pubescent; calyx lobes in markedly un¬
equal [tails, widely ovate, coppery appressed-pu¬
bescent on both surfaces, the margins eiliate. the
apex rounded, the outer pair 0.7— 1.5 X 2 mm, the
inner pair 2—2.5 X 2—3 mm; petals 3—3.5 X 2.5—

3 mm, widely ovate or elliptic, the margins eiliate.
the apex rounded; disk 2.2— 2.5 mm diam., with a
few erect, coppery hairs al the base of the style;
stamens 80 to 100, ca. 5 mm long; style ea. 5 mm
long, coppery-pubescent along the lower half.
Fruits 0— 8 X 6— 8 mm, oblate or globose, substip-
itale; pericarp slightly fleshy, entire or weakly cos¬
late; appressed pale, dibraehiate hairs scattered
over the surface, concentrated at the base and apex;
calyx persistent, erect or curled; mature fruit black.

Habitat , distribution , and phenology. Fine-oak
forests and cloud forests, western Nicaragua at el¬
evations of I 1(H)— 1600 m. Collected in (lower.
May— September; collected in fruit October— March.

Eugenia esteliensis is similar to E. guaternalensis
Donnell Smith, but differs in vesliture color, cop¬
pery versus sordid or cinereous, and branchlet col¬
or, reddish versus gray. The leaves, though similar
in size, are narrower, 3—4 versus 2—3 times as long
as wide, while the undersides, ultimately glabrate
in this species, retain persistent, scattered hairs in
E. guaternalensis.

Paratypes. NICARAGUA. Estelf: Mesas Moropoten-
le, G. Davidse el al. 30623 (F, MO); Cerro Omaha, situado
a nnos 8 km al NK de la eiudad de Kstelf, A. Grijalva N
M. Araquistain 666 (MO); Cerro Quiabu, al NO de Kstelf,
P. P. Moreno 1316 (MO); San Jos6 de la Laguna, P. P.
Moreno 8005 (MO); al pie del Cerro Omaha. P. P. Moreno
8101 (MO); a 8 km de la Carretera Panameriea, camino
a San Nicolas, P. Moreno 8730 (MO); 1.5 km al N del
valle San Jos6 de la laguna, camino a San Nicolas, l‘.
Moreno 11331 (MO); ft km sobre la carretera a San Ni¬
colas, San Jose de la laguna, P. Moreno 12677 (MO); fal-
das del cerro Tisey, P. P. Moreno I2726l> (MO); La La¬
guna, 500 m al N del valle, P. P. Moreno 15755 (MO); I
km al N de San Jose de la Laguna, P. Moreno 16738 (MO);
Sail Jose de la Laguna, P. Moreno 17374 (MO); Llano el
I’ozo, Iff km de Kstelf, camino a la Laguna de Miraflores,
por Paso de Lertn, P. P. Moreno 17456 (MO); Cerro Quia-
I u1. P. P. Moreno 10262 (MO); LI Chayote P. P. Moreno
21767 (MO); Mesas Plan Helado, 25 km al NL de Lslelf.
P. P. Moreno 24156 (MO); near Cerro Sta. Rosa de Apa-
guaji; 7 km NW San Nicolas, Dr. Davila Rolano's farm.
I). Neill 714 (MO|2|); Cerro Quiabu (= Cerro Las Brisas),
W. I). Stevens 16013 (MICH, MO); I 1.7-13.7 km NL of
Hwy. I at Lslelf on road to Yah', Mesas Meroponte, IF. D.
Stevens 15603 (MICH. MO); en San Jose, a 10 km al SL
de Kstelf, 0. Tellez & P. P. Moreno 4800 (ML, XU. MO).
Jiuotegu: region of La Montanita & Las Mesitas, in sierra
W of Jiuotegu, P. G. Standley 10332 (L). Leon: near Ls-
telf border at Finea Quetzal Cayan. ea. 15 km SW of Ks¬
telf, J. T. Atwood & I ). A. Neill ANI54 (MICH|2], MO).
IVIudriz: Carretera a Cusmapa, a 17 km de Somoto, LI
Sileneio, P. P. Moreno 17627 (MO). Mulugalpa: Lsqui-
pulas, Cerro Santa Maria, 3 km al SL de Ksquipulas, P.
P. Moreno 25375 (MO).

Eugenia farinacea Barrie, sp. nov. TYPE: Nica¬
ragua. Matagalpa: El Picacho, E of Sta. Marfa
dc Ostuma, Cordillera Daricuse, 26 May 1974
(fr), A. Molina R. & A. R. Molina 30513 (ho¬
lotype, MO; isotype, F). Figure I I.

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2005

Barrie

Eugenia from Mesoamerica

17

11 (left). Eugenia farinacea Barrie (Molina A; Molina 30513', holotype, MO). — 12 (right). Eugenia gomezii Barrie (Gdmez et al. 22695; holotype, CR).

18

Novon

Alt Eugenia yunckeri indumento non tanlum simplici,
foliis ritTvis lateral i l>ns 10—14, petiolis longioribus, flori-
l)ns majoribus diflert.

Small trees, to 10 m; young stems, petioles, in-
Horescenees densely pubescent wit!) mixture of mi¬
nute sordid or cinereous, dibrachiate, infundibular
and simple hairs; branehlets compressed, ridged;
bark tan. Leaves elliptic to obovate, the blades 9.5—
16 X 3.5— 5.5 cm, 2.5— 3.5 times as long as wide,
coriaceous, drying dull green above, sordid or ci¬
nereous below; midvein depressed between 2 par¬
allel ridges, lateral veins 10 to 14 per side, straight
and ascending, marginal veins similar to the lat¬
erals and weakly arched between them. 1—2 mm
from the margins; upper surface and midvein gla-
brate or with scattered, dibrachiate hairs, the lower
surface densely covered with minute, cinereous or
sordid, infundibular and appressed, dibrachiate
hairs, ea. 0.1 mm long; glands obscure above,
glands dark and scattered below, typically obscured
by the vestiture; base cuneate; margin somewhat
thickened and revolute; apex acute to acuminate;
petioles 10-20 mm long, sulcate dorsally, pubes¬
cent with a mixture of appressed, dibrachiate hairs
and erect, simple hairs. Inflorescences stout axil¬
lary racemes, the axis, bracts, bracteoles, hypan-
thium, and calyx uniformly coppery-pubescent with
a dense covering of appressed. dibrachiate hairs in
bud; axis 1-6 X 2-2.5 mm; flowers 4 to 8; bracts
1 X 1 mm, persistent at least through flowering,
the margins entire, the apex rounded; buds 5 mm,
obovoid. Flowers pedicellate, the pedicels 2-5 X
2-3 mm long, stout, sometimes wider proximally,
appressed coppery-pubescent; bracteoles 1—2 X
1.5—2 mm, persistent, sometimes wider than long,
appressed-pubescent in bud with coppery or sordid,
dibrachiate hairs on the inner and oilier surfaces,
at anthesis the outer surface glabrate and dark,
contrasting with the coppery pedicel and hypanthi-
um, the base free, the margin filiate, the apex
rounded; hypanthium 3—5 mm long, campanulate,
densely coppery-pubescent; calyx lobes in sube¬
qual pairs, 6-7 X 5—6 mm, elliptic, densely cop¬
pery-pubescent on the inner and outer surfaces, at
anthesis the outer surface glabrate and dark, the
margins ciliolate, the apex rounded; petals 6—8 X
4—6 mm, oblong, the margins ciliate, the apex
rounded; disk ea. 5 mm diam., densely coppery-
tomentose; stamens ca. 200, 6—7 mm long; style 7—
10 mm long, glabrous or w ith scattered simple hairs
on lower half. Fruits 20-40 X 18—35 mm, globose
or obovoid, the base slightly attenuate; pericarp 2—
3 mm thick on nearly mature fruit, fleshy; the sur¬
face verruculate and farinaceous, uniformly cov¬
ered with minute, sordid or cinereous dibrachiate

and simple hairs. 0. 1 mm long: calyx absent; ma¬
ture color unknown.

Habitat , distribution , and phenology. Wet pre-
montane forests, montane forests, cloud forests in
Honduras and Nicaragua at elevations of 1000-
1600 m. Collected in flower August-September;
collected in fruit September— May.

Along with Eugenia belloi , E. lepulota, and E.
hartsbornii , E. farinacea is among a group of spe¬
cies in which the undersides of the leaves are uni¬
formly covered with a persistent, dense mixture of
dibrachiate and infundibular hairs. In this species,
the fruits are 2-4 cm in diameter, with a fleshy,
rugose pericarp. When immature, they are uniform¬
ly covered with minute, sordid or cinereous hairs,
giving them a farinaceous texture. Whether the ves¬
titure persists in fully mature fruit is unknown. Col¬
lections of this species have been identified as Eu¬
genia yunckeri Standley, which differs in having
generally smaller leaves with half as many lateral
veins and shorter petioles (5—9 mm), pedicels less
than 2 mm long that are often as wide or wider than
long, and globose fruits approximately 2 cm in di¬
ameter. In E. yunckeri , the vestiture is composed of
simple, erect hairs only.

Etymology. The specific epithet refers to the lar-
inaceous fruits.

Paratypes. HONDURAS. CumuYugiiu: between San
Antoina Siguatepeque & F,1 Cedral, A. Molina R. 225ft 6
(F, M()[2], NY); FI Cedral on Cordillera Monteeilos. 16
km SW of Siguatepeque, A. Molina II. 25519 (h NY);
Catnino al Cerro La Carlota, a 2 km de la carretera al
Tuma, P. P. Moreno 15660 (MO). NICARAGUA. Mata-
galpa: Kl Porvenir, Cordillera Central. A. Molina R.
20511 (F. NY, US); Finea Sta. Marfa de Ostuma, Cordil¬
lera Dariense, I,. <). Williams A- A. Molina R. 12646 (f);
Fuente Pura, Km 142, carretera Matagalpa-Jinotega, P. P.
Moreno 17057 (MO); Hacienda Sta. Marfa de Ostuma,
Cordillera Dariense, .S'. Tomlin 84 (MO); Sta. Marfa de
Ostuma, Cordillera Central, /, (). Williams et al. 25065
(K|2), LI, NY, US).

Eugenia gomezii Barrie, sp. nov. T\ PE: Costa
Rica. Puntarenas: path from Bellavista to Co-
toncito, I I Mar. 1984 (fl), /, I). Gdmez et id.
22695 (holotvpe, CR; isotypes, CR, MO, TEX).
Figure 1 2.

Eugeniae glandulosopuiwtatae aemulans, sed foliis pe¬
tiolis longioribus et apieibus brevioribus, floribus oauli-
floris et pedicellatis differt.

Trees to 15 in or more; young growth glabrous or
with a few simple hairs on buds and the margins
of the bracteoles; branehlets compressed; bark gray,
smooth and glandular. Leaves elliptic, the blades
4—8 X 2-4 cm. 1.5— 2.5 times as long as w ide, ehar-
taceous, drying a concolorous, bright green; mid-

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Eugenia from Mesoamerica

19

vein flat or concave above, convex below; lateral
veins 8 to 10 per side, pins intermediates, promi¬
nent on both surfaces; marginal veins similar to the
lateral veins and weakly arched between them, 1-
3 mm from the margins; glands large, pellucid,
prominent on both surfaces and along the midvein;
base cuneate; margins entire; apex acuminate, the
tip ultimately rounded; petioles 7—10 mm long, sul-
cate dorsally, prominently glandular. Inflorescences
cauliflorous, fasciculate; llowers 2 to 10; bracts
0.3— 0.5 mm long, deciduous past anthesis, cymbi-
form, the margins entire, the apex rounded; buds 2
mm long, obovoid. Flowers pedicellate, the pedicels
3-5 mm long, prominently glandular; bracteoles I
X 0.5 mm, deciduous after anthesis, ovate, prom¬
inently glandular, the base free, the margins sear-
ious with or without a few simple hairs, the apex
acuminate; hypanthium 1.5—2 mm long, campanu-
late, prominently glandular; calyx lobes widely
ovate or elliptic, petaloid, white, in unequal pairs,
the margins entire, the apex rounded, the smaller
pair 1—2 X 1—1.5 mm, the larger 2—3 X 1.5— 2.5
mm; petals 2.5—3 X 2—2.5 mm, elliptic, the mar¬
gins entire, the apex rounded; disk 2-3 mm diam.,
glabrous; stamens 30 to 50, 3—4 mm long; style 4
mm long, glabrous. Fruits 20-25 X 18-22 mm,
broadly ellipsoid or obovoid; pericarp fleshy. 1—2
mm thick, glandular, glabrous; calyx absent; mature
color red-purple.

Habitat , distribution, and phenology. Known
from the montane forests of western Panama and
eastern Costa Rica at elevations of 1000-1800 m.
Collected in flower January-April; collected in fruit
August-January.

A nearly glabrous, medium-sized tree, Eugenia
gomezii is similar to E. glandulosopunctata P. E.
Sanchez & Poveda in having prominent glands on
the branchlets, leaves, anil inflorescences. The lat¬
ter species, however, has more prominent leaf api¬
ces that are 10—20 mm long and much shorter pet¬
ioles, 2-3 mm long. They also differ in the position
of the inflorescence, axillary in E. glandulosopunc¬
tata, cauliflorous in E. gomezi, the hypanthium in
flower, clavate in E. glandolosopunctata, campan-
ulate here and in the shape of the fruit, which are
longer (20—35 mm) and narrower (10—20 mm) in E.
glandulosopunctata. Eugenia siggersii Standley is
another species with similarly shaped, prominently
glandular leaves and cauliflorous inflorescences.
That species, however, has massive, globose fruits,
3-6 cm in diameter, with a thick, fleshy pericarp.

Etymology. Named for the co-collector of the
type, l.uis Diego Gomez, noted expert on the flora

of Costa Rica and director of the Fas Cruces and
Finca La Selva biological stations.

Paratypes. COSTA RICA. C. Laskowski 1442 (CR. F).
Puntarenas: Osa Aguabuena, oeste, Rincon, It. Aguilar
1359 (K 1NR[2], MO); Coto Brus, Cuenca Terraba-Sierpe,
Coma Indio, E. Alfaro 1881 (F, INR); Coto B rus, Cuenca
Terraba-Sierpe, Las Mellizas. Cafrosa. E. Alfaro 1931 (F.
INB); Coto Brus, Cuenca Terraba-Sierpe, Z.P. Las Tablas,
Est. Biol. Las Alturas, collectando sobre el carril, E. Al¬
faro 2361 (F, INB); Cerro Pando, on road through ridge
forest above the Rio Coton & Rio Negro, K. Barringer &
L. I). Gdmez 1634 (CR, F. MO); vicinity of first large con¬
crete culvert before Finca Las Alturas at Coton, R. Coton,
/. B. (.root 44369 (MO); Lila LI I igre, SE of Las Alturas,
G. Davidse 24198 (BM. MO); forest along trail betw. Las
Alturas & Cotonsito, along R. Coton, G. Davidse 24370
(CAS. MO); foothills of the Cordillera de Talamanca, for¬
ested valley of the R. Coton between Sitio Cotdn (Colon-
sito) & Sitio Tablas, G. Davidse 24422 (MO); Talamanca
along Quebrada Kuisa (tributary of Rfo Lori) near crossing
of UjarnSs-San Jos£ Cabecar trail, Cordillera de Talaman-
ea, M. Grayum 10286 (F); above La Tigra. near Panama
border, D. Hazlett 5048 (F); Osa Palmar Norte, bosque
aledano a la toma de agua potable, faldas de fila Retinto,
Q. Jimenez 1078 (F, INB, MO); Est. Boscosa, Rincbn de
Osa, 28 Sep. 1991, L. Martinez s.n. (INB); Zona Protectora
Las Tablas, San Vito Coto Brus-Sabalito Rfo Negro, G.
Mora 21 (CR, MO); Coto Brus ZP Las Tablas, Cuenca
Terraba-Sierpe, sendero Huacas Cotoncito a 2 km E del
Cerro Chiva, E. Navarro V. 716 (F, INB, MO). San Jose:
Zona Protectora Las Tablas, R. Cotoncito, /. A. Chacdn et
al. 1791 (MO, TEX, WIS). PANAMA. Chiriqih: upper
small house of Finca Ojo de Agua, E of R. Chevo, .S'.
Knapp 1542 (MO).

Eugenia grayumii Barrie, sp. nov. TYPE: Costa
Rica. Cartago: Coto Brus, along R. Jaba, 1 km
SW of Las Cruces Botanical Garden, 10 Mar.
1987 (fr), 17. Grayum & J. Ajfolter 8129 (ho-
lotype, INB; isotype. MO). Figure 13.

Eugeniae truncatae aemulans, sed foliis basibus cunea-
tis, petiolatis 6-8 mm, fructibus majoribus, ellipsoidalibus
vel obovoideis differt.

Small trees to 15 m; young stems, petioles, and
pedicels moderately to densely puberulent, the
hairs simple, erect or recurved, 0.1— 0.2 mm;
branchlets compressed; bark tan or gray. Leaves
elliptic, the blades 6-13 X 2-5 cm, 2-2.6 times
as long as wide, chartaceous, drying a concolorous
olive, the veins a contrasting tan; midvein indented
or depressed above, weakly convex below; lateral
veins 5 to 6 per side, impressed above or prominent
on both surfaces, curving-ascending, with numer¬
ous, reticulate intermediate veins; marginal veins
well defined, similar to the laterals and arched be¬
tween them, 2-4 mm from the margins; upper and
lower surfaces glabrate or with a few scattered
hairs, the margins ciliate, the hairs recurved, 0.1—
0.2 mm long, the midvein puberulent or glabrate

20

Novon

Figures 13, 14. — 13 (left). Eugenia grayumii Barrie (Grayum & Affolter 8120: holotype, INB). — 1 1 (right). Eugenia grijalvae Barrie {Grijalva 324: holotype, MO).

Volume 15, Number 1
2005

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Eugenia from Mesoamerica

21

above and below; glands convex, distributed uni¬
formly over both surfaces; base cuneate; margins
entire; apex acuminate; petioles 6—8 mm long, flat¬
tened dorsally, puberulent. Inflorescences axillarv
or canliflorous, sessile, solitary or 2 superposed;
flowers 4 to 10; bracts 0.5 mm long, persistent, tri¬
angular. puberulent, the apex acute. Flowers not
seen. Fruits 12—15 X 10—12 mm, ovoid; pericarp
somewhat fleshy, glandular-verrucose, glabrous or
with scattered, minute hairs; pedicels 5—20 mm
long, puberulent; bracteoles 0.7—1 X 0.5 mm, el¬
liptic, persistent, opposite or subopposite, one or
both adnate or up to 1 mm below the hypanthium.
puberulent. the base free, the margins ciliate. the
apex acute; calyx persistent, erect, the lobes in sub¬
equal pairs, 2—8 X 2—3 mm, triangular, glabrous,
the margins ciliate, the apex rounded or bluntly
acute; disk 3—4 nun diam., glabrous; mature color
red.

Habitat, distribution, and phenology. Known
from primary and secondary forests, eastern and
central Costa Rica, at elevations of 500—1300 m.
Collected in fruit January, March, September, De¬
cember.

Eugenia grayumii is one of three species (the
other two are E. truncata 0. Berg of Costa Rica and
E. jutiapensis Standley & Steyermark of eastern
Guatemala and FI Salvador) in which the bracteoles
of many of the flowers are not paired and opposite
each other immediately beneath the hypanthium as
is typical in Eugenia, but which are instead offset,
attached singularly and remotely at different points
along the pedicel. This species differs from E. trun-
cata in having leaves with cuneate, rather than
truncate or abruptly cordate, bases and longer pet¬
ioles, 6—8 mm versus 2—3 mm. The pedicels are
typically longer, 15—22 mm. and more gracile in E.
truncata than in E. grayumii, where they are usu¬
ally between 5 and 15 mm long, but in the latter
species individual flowers or fruits may have ped¬
icels up to 20 mm long.

Etymology. The epithet honors the collector of
the type, Michael Grayum. noted authority on Ar-
aceae and co-editor of the Manual de Plantas de
Costa Rica.

Paratypes. COSTA RICA. Cartago: Coto Brus,
Slopes of Cerro Paraguas, P. Aldrich 02011 (CR); ca. 17
km N of San Vito on dirt road to Potrero Grande, C. Da¬
vidson 7161 (F. MICH. MO); Cana Gordas Fas Cruces
Botanical Garden, 6’. S. Hartshorn 2165 (CR[2], F. MO.
NY); Turrialba, Monumento Nacional Guayabo, camino ar-
riba de la cada de la Administracion, G. Rivera et al. 1008
(CR, F, K, MO[2]). San Jose: Reserva Biologiea Carara
Rijagual. R. Hammel et id. 17708 (F. k|2|); Turrubares,
RB Carara, Valle del Taracoles. Sector Bijugaf, hordes del

bosque 2° en el lfniite de la Reserva, (A Jimenez & V. //.
Ramirez 1722 (F. INB. MO).

Eugenia grijalvae Barrie, sp. nov. TV PE: Nica¬
ragua. Atlantico Norte: Cerro FI Hormiguero,
14 Apr. 1979 (fr), A. Grijalva 324 (holotype,
MO; isotype, MICH). Figure 14.

Ab aliis Eugeniae nicaraguensibus indumento coffeato
et foliis laneeolatis vel anguste ellipticis differt.

Trees to 10 m or more; young stems, leaves, and
inflorescences densely appressed-pubescent, the
hairs dibrachiate, coffee-brown; branchlets terete;
bark brown, persistently pubescent. Leaves narrow¬
ly elliptic or lanceolate, the blades 7-12 X 1.5—
2.5 cm, 4—7 times as long as wide, coriaceous, dry¬
ing dark green above, coffee-brown from the
persistent hairs below; midvein flat or concave be¬
tween 2 parallel ridges above, convex below; lateral
veins 12 to 18 per side, plus intermediates, prom¬
inent on both surfaces, though somewhat obscured
by the vestiture below; marginal veins similar to the
laterals and weakly arched between them, ca. I mm
from the margins; upper surface glabrate or with a
fewr, scattered hairs, the lower surface persistently
covered w ith a dense mat of appressed, coffee-col¬
ored hairs; glands on the upper surface a mixture
of large, raised glands and, on mature leaves, nu¬
merous minute, punctate glands on the lower sur¬
face typically obscured by the vestiture; base nar¬
rowly cuneate; margins revolute, abruptly merging
into the inner edge of the petiole; apex acuminate
to caudate-acuminate, the acumen straight to fal¬
cate; petioles 9—10 mm long, channeled dorsally,
thinly appressed-pubescent. Inflorescences axillary,
racemose, solitary, all parts appressed pubescent;
axis 3—5 X 1—2 mm; flowers 4 or more; bracts 1
mm long, persistent or deciduous, cymbiform, pu¬
bescent on both surfaces, the margin entire, the
apex acute; buds not seen. Flowers not seen, ap¬
parently large, pedicellate, the pedicels 4—5 mm
long, the bracteoles 1—2 X 2—3 nun, persistent,
widely ovate, pubescent on the inner and outer sur¬
faces, the base free, the margins entire, the apex
rounded, the hypanthium and calyx densely ap-
pressed-pubescent, the disk ca. 5 mm diam., sta¬
mens ca. 130 to 150. Fruits 20—30 X 20—30 mm,
globose; pericarp fleshy, at least 2—4 mm thick,
glandular and at times weakly costate, persistently
appressed-pubescent; calyx persistent, erect, the
lobes 2—3 X 3—4 mm, convex, widely ovate, ap¬
pressed-pubescent on the inner and outer surfaces;
immature color coffee-brown from the vestiture,
mature color unknown.

Habitat, distribution, and phenology. Known

22

Novon

only from cloud forests in central Nicaragua at el¬
evations of 800—1200 m. Collected in fruit in April.

I’lie distinctive, coffee-brown vestiture and nar¬
rowly elliptic or lanceolate leaves distinguish t li is
species from all others in Nicaragua. The persis¬
tently pubescent undersides ol the leaves lighten
with age. as though a bit of cream had been added.

Etymology. The epithet honors the collector ol
the type, Alfredo Grijalva, curator of the Herbario
Nacional de Nicaragua.

Paratypes. NICARAGUA. Malagulpa: Cerro El Hor-
miguero, western range, ./. Pipoly 5175 (MICH, MO). Ze-
laya: Cerro La Pimienta numbers 1 & 2, eastern range,
summit ol 2 peaks, northernmost and central, J. Pipoly
5255 (MICH. MO).

Eugenia linmmclii Barrie, sp. nov. T\ PE: Costa
Rica. Puntarenas: Golfito, R. F. Golfo Dulce,
Osa Peninsula, Rancho Quemado, ca. 15 km
Oeste de Rincon, en horde de bosque en fila
antes de llegar a Rancho, 6 May 1992 (b, (I.
Ir), B. Hummel, G. Galeano & A. Henderson
18505 (holotype, I NR; isotypes, E. MO). Fig¬
ure 15.

Eugeniae glandulosopunctatae aemulans, sed indumen-
lo sericeo, infloreseentiis raeemosis. hypanthiis urceolatis
et calycibus lobis anguste ellipticis differ!.

Trees to 25 m; young stems sericeous, the hairs
0.5 mm, simple, white; branchlets terete or com¬
pressed at the nodes; bark gray, flaking. Leaves el¬
liptic, the blades 4—9 X 2-3 cm, 2—3 times as long
as wide, chartaceous, drying concolorous or brown-
olive above, pale green or tan below; midvein sul-
cate above, convex below; lateral veins 7 to 10 per
side, plus intermediates; marginal veins similar to
the laterals and arched between them, 1—2 mm
from the margins; both surfaces of young leaves
thinly pubescent with pale, appressed, silky hairs,
glahrate at maturity; glands on the upper surface
obscure, the lower surface with a few, scattered pel¬
lucid glands; base cuneate; margins revolute, de¬
current along the inner edge of the petiole; apex
acuminate, the ultimate tip blunt or rounded; pet¬
ioles 5—7 mm long, suleate dorsallv, with appressed
hairs, similar to those of the stems. Inflorescences
axillary, of single flowers or more commonly race¬
mose, solitary, the pedicels, bracts, and buds thinly
pubescent, the hairs 0.5 mm long, white, simple,
appressed; axis 10-30 X 1 mm, terete; flowers I
to 6; bracts 1-1.5 X 1 mm. present at anthesis,
cymbiform, the margins ciliate, the apex acute;
buds 5 mm long, obovoid. Flowers pedicellate, the
pedicels 4—12 mm long, thinly pubescent, the hairs
white, appressed; bracteoles ca. 2 X 0.5 mm, de¬

ciduous prior to anthesis, narrowly elliptic, thinly
pubescent on inner and outer surfaces, the base
free, the margins ciliate, the apex acute; hypanthi-
nm 2-2.5 mm long, narrowly urceolate, sericeous,
the base attenuate; calyx lobes in unequal pairs,
narrowly elliptic*, connate near the base, thinly pu¬
bescent on inner and outer surfaces, the margins
entire, the apex rounded, the smaller pair ca. 2.5
X 1 mm, the larger pair ca. 5 X 1 mm; disk 2—2.5
mm diam., glabrous or with a few, scattered hairs;
stamens 50 to 60, 6—6.5 mm long; style 6-7 mm
long, glabrous or with a few, scattered hairs. Fruits
20—25 X 10—14 mm, ellipsoid or ovoid; pericarp
moderately fleshy, glandular, glabrous; calyx per¬
sistent, the lobes spreading; mature color yellow-
orange'.

Habitat, distribution, and phenology. Primary
forests, central Costa Rica to western Panama at
elevations of 200-500 m. Collected in flower
March— December; collected in fruit September—
June.

Eugenia hammelii is distinguished most readily
from other Costa Rican Eugenia by flower charac¬
ters: the narrowly urceolate hypanthium and the
narrowly elliptic calyx lobes, in unequal pairs,
which persist on the maturing fruit. In bud, the
larger lobes are straight and extend well beyond the
unopened corolla. Herbarium collections might be
confused with E. glandulosopunctata, a small tree
of the Atlantic slope, which has branchlets and
leaves of similar shape and texture. The two differ,
however in vestiture; E. glandulosopunctata is gla¬
brous except for the margins of the bracts and the
flower parts, while the young growth of E. hammelii
is thinly sericeous. The two species differ in several
floral characters, also. In E. glandulosopuntacta the
bracteoles are triangular, about 0.5 mm long and
persistent; the hypanthium is clavate and covered
with numerous, prominent glands; the calyx lobes
are 1 .2-2 mm long, ovate, with a rounded apex.
I'he flowers of E. hammelii have elliptic bracteoles
ca. 2 mm long that are deciduous prior to anthesis,
a narrowly urceolate hypanthium that is sericeous
with white hairs and the distinctive, narrowly ellip¬
tic- calyx lobes.

Etymology. The epithet honors Barry Hammel,
noted expert on the Costa Rican flora and co-editor
of the Manual de Plantas de Costa Rica.

Paratypes. COSTA RICA. Heredia: Chilamate de
Sarapiquf, S end of Cerros Sardinal, on bills S side* ol R.
Sarapiquf, li. Hummel et id. 14186 (MO, TEX); Sarapiquf,
Finca La Selva, Puerto Viejo, squatter clearing near SW
corner, G. S. Hartshorn 1853 (CR, MO); Sarapiquf, Finca
Fa Selva, Puerto Viejo, Vargas clearing near W boundary,
G. S. Hartshorn 1568 (CR, MO). Puntarenas: Osa, IFF.

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Eugenia from Mesoamerica

23

Figures 15, 16. — 15 (left). Eugenia hammelii Barrie ( Hammel 18505: holotype, INB). — 16 (right). Eugenia hartshornii Barrie ( Hartshorn 1249 ; holotvpe. MO).

24

Novon

Golfo Dulce, Penfnsula de Osa, Agaubuena, sector oeste,
A’. Aguilar 694 ( 1 1\ li ): Golfito, R.K Golfo Dulce, Serranias
de Golfito, Sendero San Joseeito, R. Aguilar 5121 (INB,
MO); Golfito, R.K Golfo Dulce, Penfnsula <le Osa, Puerto
Jimenez, R. Aguilar 5156 (INB. MO); Golfito, R.K Golfo
Dulce, Penfnsula de Osa, Estacidn Agujas, Rio Agujas,
Sendero Bonanza, I. Azofeifa -16ft (INB); Puerto Cortez.
Palmar Norte, Quebrada Benjamin, Subiendo l ila Retinto,

Herrera 6775 (F); Osa, R.F. Golfo Dulce, Peninsula de
Osa, Rancho Quemado, Fila Guerra, Rinc6n, M. Segura
17 (CR. I', INR. MO). PANAMA. Boras del Toro: sig-
uiendo el camino que va a Quebrada Bonyic, I). Santa-
mar(a & II. Ixira 1 103 (MO).

Eugenia liarlslioriiii Barrie, sp. nov. I ) PE: Costa
Rica. Heredia; Sarapiquf, Finca La Selva,
Puerto Viejo. 29 June 1973 (11), G. I lari shorn
1249 (holotvpe. MO; isotypes. CR. F, N't ). fig¬
ure 16.

Eugeniae skutchii aemulans, sed floribus paucioribus,
fructibus ellipsoiilalibus dilfert.

Small trees, to 5 m or more; branchlets, buds,
and inflorescences densely appressed-pubescent
with ochraceous, dibrachiate hairs; brancldets te¬
rete or weakly 4-angled; bark fissured, persistently
pubescent. Leaves elliptic or obovate, the blades
13-20 X 6-8 cm. 2. 3-2.8 limes as long as wide,
chartaeeous, drying a concolorous light to dark ol¬
ive, or paler below; midvein impressed above, el¬
evated below; lateral veins 16 to 18 per side, plus
intermediates, prominent on the lower surface; mar¬
ginal veins similar to the laterals and weakly
arched between them, 2—3 mm from the margins;
upper surface ultimately glabrate or with a few scat¬
tered hairs, the midvein persistently pubescent, the
lower surface densely and uniformly covered with
ochraceous infundibular hairs; glands on the upper
surface numerous, minute, raised, the glands on the
lower surface larger, typically obscured by the ves-
titure; base euneate; margins merging into the inner
edge of the petiole; apex acuminate; petioles 5—6
mm long, sulcate dorsally. persistently appressed-
pubescent. Inflorescences axillary, racemose, soli¬
tary, the axis, bracts, and buds persistently ap¬
pressed-pubescent with coppery, dibrachiate hairs;
axis 5—10 X 1—1.5 mm, 4-angled; flowers 4 to 8;
bracts 1 .5 mm long, persistent, cymbiform, pubes¬
cent on both surfaces, the margins scarious, the
apex rounded; buds 4 mm long, obovoid. Flowers
pedicellate, the pedicels 5—10 mm long, the vesti-
ture of the pedicels, involucre, hypanthium, and
calyx similar to that of the axis; bracteoles fused,
forming an involucre 2 X I mm diam., dilated in
bud and flower, pubescent on both surfaces, the
margins scarious, the apex acute; hypanthium 2
mm long, campanulate, costate; calyx lobes round¬

ed or ovate, in unequal pairs, densely pubescent
on both surfaces, the margins ciliatc, the apex
rounded, the outer pair 2 X 2-3 mm, the inner pair
3—4 X 3 mm; petals 5 X 3 mm, obovate, the mar¬
gins ciliate, the apex rounded or bluntly acute; disk
3-3.5 mm diam., bearing erect, minute, coppery
hairs; stamens 80 to 100, 4-5 mm long; style 7 mm
long, with a few. contorted hairs. Mature fruits not
seen; immature fruits ca. 15 X 10 mm, ellipsoid;
pericarp fleshy, rugose, uniformly pubescent.

Habitat, distribution, and phenology. Lowland
forests and premonlane wet forests along the Atlan¬
tic slope, Costa Rica, at elevations of 0—300 m.
Collected in flower in June; collected in fruit in
January and March.

Collections of Eugenia hartshornii are very sim¬
ilar to those of E. skutchii, a species of tin1 montane
forests at elevations of 1000—1800 m. While E.
hartshornii has much shorter petioles, 5—6 mm ver¬
sus 10-20 mm, the leaf blades of the two species
are of similar size and shape, with the underside
densely and persistently covered with infundibular
hairs. The inflorescences are similar, although in K.
skutchii they are typically longer, with an axis up
to 25 mm long. The flowers in E. hartshornii are
rather smaller; the hypanthium is 2 mm long, ver¬
sus 2—4 mm as in E. skutchii, and the larger set of
calyx lobes in the latter species are often much
broader than long, 3-4 mm long by 3-6 mm wide.
In both species, flowers and fruits are subtended
by a prominent, dilated involucre formed by the
fused bracteoles.

Etymology. The epithet honors the collector of
the type. Gary Hartshorn, the CEO of the Organi¬
zation for Tropical Studies and a noted authority on
the ecology of Costa Rica.

Puratypes. COSTA RICA. Heredia: Orilla del R.
Guacimito, Magsaysay, /. A. Chacdn 152 (CR, MO); Center
Arboretum, Finca La Selva, Puerto Viejo, G. W. Frankie
185 (F, MO); Sarapiquf, Llanura de San Carlos, La Virgen,
N. Zamora el al. 2527 (INB). Union: Hacienda Tapezeo-
Hda. La Suerte. 29 air km \\ of Tortuguero, C. Davidson
el al. 6905 (MO); Pueblo Nuevo, 17 km NE of Gudeimo,
on the farm of Bernardo Herrera M., A. Tlwmsen 1017
(CR).

Eugenia herrerae Barrie, sp. nov. TYPE: Costa
Rica. Union: Cuapiles Los Angeles. 24 Feb.
1990 (b. fl). G. Herrera & .S. Schik 3790 (ho¬
lotvpe, CR: isotypes, F, K. MO). Figure 17.

Al> Eugenia siggersii loliis majoribus. petiolis rubris el
floribus majoribus differt.

frees to 10 m or more; young growth reddish,
glabrous or with a few scattered simple hairs, all

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2005

Barrie

Eugenia from Mesoamerica

25

figures 17. 18. —17 (left). Eugenia herrerae Barrie (Herrera & Schik 3790: holotvpe, CR). —18 (right). Eugenia intibucana Barrie (. Molina R. 22596: holotype, MO).

26

Novon

parts ultimately glabrate except for the ciliate mar¬
gins of the bracts, bracteoles, calyx lobes, and pet-
als; branchlets terete; bark reddish. Leaves narrow¬
ly elliptic or narrowly oblanceolate to el I i | >t ic or
obovate, the blades 12—16 X 3—7 cm, 2— 4 times
as long as wide, coriaceous, drying a eoncolorous
olive; midvein incised above, elevated below; lat¬
eral veins It) to 12, eurving-ascending, raised on
the lower surface; marginal veins similar to the lat¬
eral veins, straight, ea. I mm from the margins;
glands on both surfaces pellucid, prominent and
raised; base narrowly cuneate; margins revolute
near the base, decurrent into the inner edge of the
petiole; apex abruptly acuminate; petioles 6—9 mm
long, reddish, sulcate dorsally. Inflorescences cau-
liflorous, fasciculate; flowers 4 to 10; bracts 0.5
mm. scaly, the margins scarious, the apex acute or
rounded; burls 5—7 mm, globose. Flowers pedicel¬
late, the pedicels 8-12 mm, prominently glandular;
bracteoles fused, forming an involucre ea. I mm
across, the margins scarious, with a few, minute
hairs, the apex acute; hypanthium 2—3 mm long,
obconic; calyx lobes in unequal pairs, depressed
ovate, the margins ciliolate, the apex rounded, the
outer pair 2—3.5 X 3.5—4 mm, the inner pair 4—5.5
X 4—5 mm; petals 5-6 X 4-5 mm, elliptic or ovate,
the margins ciliate, the apex rounded; disk 4-4.5
mm diam.. the staminal ring eiliate; stamens 90 to
100, 5—7.5 mm long; style 8—9 mm long, glabrous.
Fruits 35 X 35 mm, globose; pericarp fleshy, ea. 5
mm thick, prominently glandular; calyx persistent,
erect; immature color yellow, mature color un¬
known.

Habitat , distribution , and phenology. Montane
forests along the Atlantic slope ol western Panama
and eastern Costa Rica at elevations of 700—1200
m. Collected in flower in February— March; collect¬
ed in fruit in September— October.

Eugenia herrerae is distinguished by its red new
growth, the large, elliptic leaves with reddish pet¬
ioles anil abruptly acuminate apices, the cauliflo-
rous flowers with involucrate bracteoles, and the
globose fruits with a thick anti fleshy pericarp. Eu¬
genia siggersii has similar, though often larger,
fruits but differs in having leaves with shorter
blades (5.5-12 cm) that dry a bright green, with
green petioles and more prominent veins and
glands, and flowers with the bracteoles free at the
base.

Etymology. The epithet honors the collector ol
the type, Gerardo Herrera, who has contributed
many valued specimens of Costa Rican plants,
among them numerous previously undescribed spe¬
cies.

Paratype. PANAMA. Veraguas: N ol Sta. Fe, ea. 2
km N ol Escuela Agneola Alto <le Piedra, .S'. Mori & ,/.
Kallunki 2617 (F. MO).

Eugenia iiitibiicana Rarrie, sp. nov. TYPE: Hon¬
duras. Intibuca: vicinity of Mazaguara, Valley
de Jesus de Otoro, 3 Sep. 1968 (b, fl), A. Mo¬
lina R. 22596 (holotype, MO; isotypes, RM,
I)S, F, NY). Figure 18.

Ah Eugenia hypargyreu foliis plus uervis lateralibus et
petiolis duplo longioribus, infloresoentiis longioribus dif¬
fer!.

Shrubs or small trees to 8 m; stems, young
leaves, inflorescences densely covered with op¬
pressed. reddish, dibrachiate hairs, the hairs paling
with age; branchlets compressed, weakly chan¬
neled: bark gray, pubescent with appressed, red¬
dish hairs. Leaves obovate to elliptic, the blades 7-
12 X 4—6 cm, 1.5—3 times as long as wide,
coriaceous, drying a bright green above, yellow or
white below from the dense mat ol appressed hairs;
midvein concave above, convex below; lateral veins
6 to 10 per side, straight and ascending; marginal
veins strongly arched, 3—4 mm from the margins;
upper surface pubescent with a mixture ol simple
and dibrachiate hairs, the lower surface covered
with a dense mat of tightly appressed, white or sor¬
did hairs; glands obscure on the upper surface, typ¬
ically obscured by vestiture on the lower surface;
base acute or cuneate; margins revolute; apex
bluntly acute or rounded; petioles 10-17 mm long,
terete, densely appressed-pubescent, the hairs sor¬
did. Inflorescence axillary, racemose, solitary or 2
or 3 superposed, the axis, bracts, anil buds uni¬
formly appressed-pubescent with reddish, dibra¬
chiate hairs; axis 10—25 mm long; flowers 4 to 10;
bracts 1-2 X I mm. persistent at least through
flowering, triangular, the apex acute; buds 4—6 mm
long, obovoid. Flowers pedicellate, the pedicels 4—
8 mm. reddish appressed-pubescent; bracteoles
0.7-1. 5 X 0. 5-0.7 mm, persistent, triangular, ini¬
tially appressed-pubescent, tin1 hairs thinning, ex¬
posing the prominent black glands, the bracteoles
then contrasting with the pedicel and hypanthium,
the base Iree, the apex acuminate; hypanthium 1.5—
2.5 mm long, obconic, weakly costate, calyx lobes
in markedly unequal pairs, reddish appressed-pu¬
bescent on tin- inner and outer surfaces, the margin
ciliate, the apex rounded, the smaller pair 1.5—2 X
2.5 mm, the larger pair 3—4 X 3.5-4 mm; disk 3—
4 mm, the staminal ring tufted-eiliate; stamens 130
to 160. 5—6 mm long; style 7-9 mm long, scattered,
simple hairs along the lower third, f ruits not seen.

Habitat, distribution, and phenology. Endemic

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2005

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Eugenia from Mesoamerica

27

to the pine-oak forests of central Honduras, 600-
1500 m. Collected in flower August— February.

Most of the specimens cited under Eugenia in-
tibucana were originally identified as E. hypargyrea
Standley, apparently on the basis of general leaf
shape and similar vestiture. However, a suite of
characters distinguish the two. The leaves of E. in-
tibucana have more lateral veins (6 to 10 per side
vs. 5 to 7) and longer petioles (10—17 vs. 5—10
mm). While in both species the undersides of tin;
leaves are covered with a dense mat of appressed,
dibrachiate hairs, in E. hypargyrea the upper side
is glabrate except lor a few hairs near the base of
the blade, while in E. intibucana it is persistently
pubescent. The inflorescence axis is longer in this
species, 10—25 mm versus 5-10 mm. The hypan-
thium, calyx, and staminal disk are similar in size
in both species, but E. intibucana has longer (5-6
vs. 4-4.5 mm), more numerous stamens (130 to 160
vs. 85 to 100) and a longer style (7-9 vs. 6-7 mm)
that is pubescent along its lower third, not wholly
glabrous.

Paratypes. HONDURAS. Choluteca: vicinty of San
Marcos de Colon. P. C. Standley 15926 (F). Comayagua:
near Kl Negrito, Comayagua Valley, I). Hazlett s.n. (MO);
La Piramide, betw. Comayagua & F. Morazon border, A.
Molina R. & A. R. Molina 227/6 (RM. DS, F, NY). Co¬
pan: 2 km N of Sta. Rosa de Copan. I). Hazlett 762 (MO);
along El Higuerito R., 6 km from San Juan Opoa, A. Mo¬
lina R. A’ A. R. Molina 31 122 (F, MO). Lempira:
Matorrales de Ouebrada Banaderos cerca de Lepaera, A.
Molina R. 12995 (F[2], NY). Francisco Morazan: Oril-
las del R. Jalan, a 12 km al O de Guaimaca, A. Molina
R. 2930 (F, Gil); edge of R. Guaeerique betw. Los Lau-
reles & Las Tapias, NW of Tegucigalpa, A. Molina R.
18596 (F, GH, NY); along Amarateea creek, Amarateca
valley between Zambrano & Amarateca, A. Molina R.
22648 (F, MO, NY); rocky roadside lulls of Tamara Valley,
vicinity of R. de FI Hombre, A. Molina R. Ad. R. Molina
24542 (F, NY); along road betw. Tegucigalpa & Puente
Colorado, near Choluteca R. on way to Talanga, A. Molina
R. A A. R. Molina 25829 (RM. F, MO, NY). Santa Bar¬
bara: 4 km NF of La Arada, 1L E. Harmon A ./. I). Dwyer
3819 (MO).

Eugenia lempana Barrie, sp. nov. TYPE: Hon¬
duras. Ocotepeque: thickets of the Lempa Riv¬
er, road to Esquipulas, 29 Aug. 1968 (fr), A.
Molina R. 22427 (holotype, F; isotypes, BM.
GH, NY[2|). Figure 19.

Eugeniae guatemalemi aemulans, sed ramulis grosse
tomentosis, mox glabratis, foliis glabratis, floribus brac-
teolis caducis differt.

Trees to 5 m; young growth coarsely tomentose
with coppery, contorted, dibrachiate hairs; branch-
lets terete or somewhat compressed, soon glabrate
or nearly so; bark initially reddish, then white or

gray. Leaves narrowly elliptic to narrowly obovate,
the blades 3—7 X 1—3 cm, 2—4 times as long as
wide, coriaceous, drying concolorous or slightly
paler green below; midvein sulcate above, convex
below; lateral veins 10 to 14 per side, straight and
ascending, prominent below; marginal veins similar
to the laterals and arched between them, 1-2 mm
from the margins; upper surface soon glabrate or
with a few, scattered hairs near the base, the lower
surface more persistently pubescent with coppery,
contorted hairs, ultimately glabrate; glands numer¬
ous, small and convex on both surfaces; base cu-
neate; margins cartilaginous, weakly revolute; apex
bluntly acute to acuminate, the tip rounded; peti¬
oles 5-7 mm long, sulcate dorsally, persistently ap-
pressed-pubescent, glandular. Inflorescences axil¬
lary, racemose, solitary or 2 superposed; axis 5—15
mm long, coarsely coppery tomentose; flowers to
II. a terminal flower often present; bracts 1 — 1.5
nun long, deciduous soon after anthesis, ovate, gla¬
brate and prominently glandular, the base clasping,
the margins scarious, the apex acute; buds 3 mm
long, obovoid. Flowers pedicellate or the terminal
flower sessile, the pedicels 2-5 mm long, terete or
weakly 4-angled, coarsely coppery tomentose; brac-
teoles 2—4 X 0.5 mm, deciduous at or soon after
anthesis, narrowly elliptic to lanceolate, glabrate
and prominently glandular in bud, the base free,
the margins scarious, the apex acute; hypanthium
1-2 mm long, campanulate, coarsely coppery to¬
mentose; calyx lobes in unequal pairs, appressed-
pubescent on the inner and outer surfaces, the mar¬
gins ciliate, the apex rounded, the smaller pair
triangular, 1.5-2 X 2 mm, the larger pair ovate,
2.5—3 X 3 mm; petals not seen; disk 3 mm diam.,
pubescent near style base; stamens and style not
seen. Fruits 7-15 X 5-9 mm, ellipsoid; pericarp
thin-walled, glandular, persistently appressed-pu-
bescent; calyx persistent, erect; mature color un¬
known.

Habitat, distribution, and phenology. Dry thick¬
ets in the Rfo Lempa drainage, Honduras, 600— 900
m elevation. Collected in flower in June; collected
in fruit in August and September.

Eugenia lempana is a small tree with leaves, in¬
florescences, and flowers similar in size and shape
to E. guatemalensis, which differs in having closely
appressed vestiture that is thinning but persistent
on the undersides of the leaves, branehlets, and
inflorescences, persistent bracteoles that are 1 mm
long or less, and smaller, globose or ovoid fruits.
The coarse, coppery pubescence on the emergent
growth is similar to that of E. salamensis Donnell
Smith, as are the large, narrowly elliptic bracteoles

28

Novon

Figures 19. 20. — 19 (left). Eugenia lempana Barrie ( Molina R. 22427; holotype, F). — 20 (right). Eugenia liesneri Barrie {Liesner 26428; holotype, F).

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

29

that are deciduous at or soon after anthesis. In that
species, however, the leaves, flowers, and fruits are
twice the size of E. lempnna s, and the inflores¬
cences axes are shorter, 3—7 cm long.

Paratypes. HONDURAS. Comayagua: (Juebrada
Los Mercedes, Valle Comayagua, A. Molina l\. 14379 (F).
Ocotepeque: Llano del Ojo de Agua, 2 km W above
Antigua Ocotepeque, drainage Lempa R., A. Molina R.
24168 (F, MO); thickets ol Lempa R. between Sta. Anita
& Sta. Fe, .4. Molina R. & A. R. Molina 31061 (F, MO).

Eugenia liesneri Barrie, sp. nov. TYPE: Hondu¬
ras. La Paz: Las Trancas, 5 km NW of Guaji-
quiro, 22 May 1993 (fl). R. Liesner 26428 (ho-
lotype, F; isotype, MO). Figure 20.

Ab Eugenia pittieri fforibus pedicellatis, bracteolis cad-
ucis et fruetibus grandioribus differt.

Shrubs or small trees, to 6 m; young stems,
leaves, and inflorescences minutely puberulent
with simple, erect or recurved hairs, 0.1 mm; veg¬
etative buds sericeous, the hairs silvery; branchlets
compressed; bark gray, smooth. Leaves elliptic or
ovate, 2.5— 4.2 X 1-1.5 cm, 2.5—4 times as long as
wide, coriaceous, drying nearly concolorous or a
slightly paler green below; midvein flat or weakly
convex above, weakly convex below; lateral veins
5 to 8 per side, straight and ascending, marginal
veins weakly arched, ca. I mm from the margins;
glabrate or with scattered, recurved hairs along the
lower third of the margin above, the midvein gla¬
brate or puberulent above and below, the hairs
sometimes restricted to the lower third; glands ap¬
parent on both surfaces; base euneate; margin rev¬
olute, decurrent along the petiole; apex caudate-
acuminate, the tip ultimately rounded; petioles 2—
3 mm long, flattened or concave dorsally, minutely
puberulent. Inflorescences axillary, racemose or
flowers solitary, the axis, bracts, and buds uniformly
puberulent; axis 3—8 mm long; flowers I to 6; bracts
0.5 mm long, deciduous near anthesis, triangular,
apex acute; buds 3—3.5 mm long, obovoid. Flowers
pedicellate, the pedicels, braeteoles, hypanthium,
and calyx uniformly puberulent; pedicels 4—8 mm
long, noticeably longer than the internodes; brac-
teoles 0.5—1 mm long, spreading or divaricate in
bud, deciduous in flower, the base free, the margins
ciliolate, the apex acute; hypanthium 1 — 1.5 mm
long, obconic, sub-stipitate; calyx lobes in mark¬
edly unequal pairs, puberulent on outer surface, the
margins ciliate, the apex rounded, the smaller pair
0.8— 1.3 X 0.8— 1.5 mm, the larger pair 1.5— 1.8 X
1.5— 1.8 mm; petals 3-4.5 X 3-3.5 mm, widely el¬
liptic or ovate, the margin ciliate at the apex, oth¬
erwise entire, the apex rounded; disk 1 .8-2 mm

diam., glabrous; stamens 50 to 65, 4—5 mm long;
style 5—6 mm long, glabrous. Fruits 8—10 mm, glo¬
bose; pericarp thin-walled, glandular, glabrous; ca¬
lyx persistent, erect; mature color dark red to black.

Habitat, distribution, and phenology. Known
from the cloud forest regions of Honduras, El Sal¬
vador, and eastern Guatemala, at elevations of
1500—2300 m. Collected in flower March— May; col¬
lected in fruit July— March.

In habit and its short-petiolate, elliptic or ovate
leaves, Eugenia liesneri resembles Eugenia pittieri
Standley, a species of wet forests at elevations be¬
low' 1 200 m that ranges from Guatemala to Panama.
In the latter species, however, the flowers are ses¬
sile or with pedicels no more than 1 mm long, the
braeteoles are persistent and sometimes connate
near the base, and the floral disk is 0.5-1 mm in
diameter. In E. liesneri, the flowers are borne on
pedicels 4—8 mm long, the braeteoles are decidu¬
ous around the time the flowers open, and the floral
disk is 1.8—2 mm in diameter. Many collections of
E. liesneri have been identified as E. tikalana Lun-
dell, a lowland species of Belize, the Peten. and
adjacent areas of Mexico. Eugenia tikalana differs
in having somewhat longer, to 7 cm, and broader,
2-2.5 times as long as wide, and smaller flowers
with the disk 1 — 1.5 mm in diameter and the sta¬
mens less than 4 mm long.

Etymology. The epithet honors the collector of
the type, Ronald Liesner of the Missouri Botanical
Garden.

Paratypes. EL SALVADOR. Santa Ana: Bosque de
Montecristo, A. Molina R. & A. R. Molina 12626 (F, LL,
NY, US); Bosque Nebuloso de Monte Cristo, M. L. Reyna
944 (MICH); near Montecristo, I). Witsberger 839 (MICH).
GUATEMALA. Cliiquimulu: middle slopes of Motana
Norte to FI Jutal on Cerro Brujo, SF of Concepcion de las
Minas, ./. 4. Steyermark 31067 (F, NY). HONDURAS. In-
tihuca: Quebrada de Pelon de Guise, A. Molina R. 6411
(F, GH); bosque de nubes de Calaveras, Cord, de Opaluca,
4. Molina R. 6416 (F); bosque nebuloso de Calaveras,
cordillera Opalaca, 6 km NO de La Esperanza, A. Molina
R. 7977 (BM. F, WIS); bosque de montana Calaveras,
Cord. Opalaca, 8 km de La Esperanza, A. Molina R. 8717
(F, MO, NY); bosque abierto, 25 km de La Esperanza,
Montana Opalaca, 4. Molina R. 10912 (F); cloud forest of
Cordillera Opalaca betw. Calaveras & Pela Narfz, road to
La Esperanza, 4. Molina R. 22608 (BM. I)S, F. NY 1 2 ]);
El Duraznillo on Cordillera Opalaca, 4. Molina R. & A.
R. Molina 24491 (F[2]); open forest of Ouebrada Lejarsia
between km 9 and 1 1 on La Esperanza— Marcala rd., 4.
Molina R. & A. R. Molina 24302 (F); Pela Narfz on Cor¬
dillera Opalaca, 4. Molina R. A 4. R. Molina 25581 (F.
MO, NY); Sierra Opalaca above FI Duraznillo, A. Molina
R. & A. R. Molina 26080 (BM. F. NY). La Paz: 20 km
SO Opatorao, /. Colindres 68 (CAS, MO); Las Trancas,
near Guajiquiro— San Isidro Road, ca. 1 1 km by road (ca.
5 km air) NW of Guajiquiro, Reserva Biologica Guajiqui¬
ro, R. Evans 1713 (F); 20 km SO de Opatoro, M. Martinez

30

Novon

M. 59 (MO); Cordillera Guajiquiro, 5 km a Sabanetas ,4.
Molina R. & A. R. Molina 13876 (F, LL, NY); cut over
mixed forest between l,as Marias & El Cerron on Cordil¬
lera Guajiquiro, A. Molina R. <£• A. R. Molina 24227 (BM.
I)S|2|, F, (41); open forest of Montana Verde on Cordillera
Guajiquiro, A. Molina R. <V A. R. Molina 24388 (MM, DS,
F, (ill).

Eugenia lithospcrma Barrie, sp. nov. TV PE: Cos-
la Rica. Alajuela: San Ramon, RB Montever¬
de, Cortlillera de Tilaran, Rfo Penas Blancas,
Refugio Aleman’s Quebrada Rojas, 29 Feb.
1992 (fr), E. Hello 4446 (holotype, INB; iso¬
types, F, MO). Figure 21.

Eugeniae hiraeifoliae aemulans, sed foliis paueioribus,
ovatis vel elliptieis, fructibus globosis, testis induratis dif-
fert.

Trees to 15 m X 30 em; young buds, braneblets,
leaves, and inflorescences coarsely tomentose with
yellow-brown, contorted, dibrachiate hairs, to 0.5
mm; braneblets compressed at the terminal node;
bark tan, pubescent, flaking. Leaves ovate to ellip¬
tic. tin* blades 7—14 X 3—6 cm, 2—3 times as long
as wide, chartaceous, drying a concolorous, dull
green; midvein impressed above, elevated below;
lateral veins 8 to 14 per side, slightly raised to
weakly inscribed in the upper surface, prominent
below and often tan in color; marginal veins similar
to the laterals and weakly arched between them, 3—
5 nun from the margins; upper surface soon gla-
brate or more commonly with a few, scattered, ap-
pressed hairs, the midvein glabrate or persistently
pubescent, the lower surface and midvein persis¬
tently covered with appressed hairs, these thinning
with age; glands on both surfaces numerous and
small, usually dark below; base cuneate; margins
decurrent into the inner edge of the petiole; apex
acuminate to caudate-acuminate; petioles 8-12 mm
long, suleate dorsally, with scattered, appressed
hairs. Inflorescences axillary, solitary, racemose,
coarsely tomentose; axis 15—40 mm long; flowers 7
to 13, a terminal flower present; bracts 1—3 mm,
deciduous by anthesis, cymbiform, ovate or lance¬
olate, tomentose on outer surface only, the base
clasping, the margins ciliate, the apex acute; buds
2 mm long, obovoid. Flowers pedicellate, the ped¬
icels 2—5 mm long, coarsely tomentose; bracteoles
1.5—2 X 1 mm, deciduous at or before anthesis,
ovate to elliptic, tomentose on the outer surface
only, the base free, the margins ciliate, the apex
acute; hypanthium campanulate, 2—3 mm long,
coarsely tomentose; calyx lobes widely ovate, in
subequal pairs, 2.5—3 X 3—3.5 mm, coarsely to¬
mentose' on the outer surface and on the upper V3
to Va of the inner surface, the apex rounded to

bluntly acute; petals 4.5— 5.5 X 4—5 mm, widely
elliptic, the margins ciliate; disk 3-4.5 mm diam.,
coarsely pubescent; stamens 100 to 130, 4—8 mm
long; style 6—7 mm long, with scattered, contorted
hairs. Fruits 20—35 X 20-35 mm, globose; pericarp
fleshy, 1—2 mm thick, tesla indurate, 1—2 mm thick;
calyx present or absent, the lobes curled, the outer
surface glandular, with a few, relietual appressed
hairs; mature color unknown.

Habitat , distribution, and phenology. Wet mon¬
tane forests along the Atlantic slope in Costa Rica
at elevations of 100— 12(H) m. Collected in flower
April— June; collected in fruit September— February.

Ibis species is distinguished by the thick, in¬
durate seed coat, a character otherwise unknown in
Mesoamerican Eugenia. The coarse indumentum
composed of simple and dibrachiate hairs, the elon¬
gate inflorescence, and the prominent, decidous
bracteoles are similar to E. hiraeifolia Standley,
which differs in having larger, obovate leaves, el¬
lipsoidal fruits, and seeds with a thin, membranous
testa. The leaves are similar those of E. grayumii,
which bears only simple hairs and subsessile inflo¬
rescences.

Paratypes. COSTA RICA. Alajuela: Reserva Biolo-
gica Monteverde, Rfo Renas Blancas, Eladio’s y alrede-
dores, E. Bello 1175 (INB); Reserva Bioldgica Montever¬
de, Rfo Renas Blancas, E. Bello 531 (F, INB. MO);
Reserva Bioldgica Monteverde, Rfo Pefias Blancas, Finca
Tofio Gonzalez. E. Bello 1624 (INB, MO); San Ramdn,
Reserva Forestal de San Ramon, I. Wallenberg & H.
Schafer 59 (CR). Cartago: Chitara, Turrialba, I,. ./. Po-
veda 186 (CR, MO). Guanaeasle: Cordillera de Tilaran,
Hotel Ecolodge, sendero de las cabinas, 6'. Rivera 3054
(CR, F). Heredia: Finca La Selva, Sarapiquf, K. S. Bawa
594 (MO); Finca La Selva, Puerto Viejo, G. W. Erankie 78
(F, MO). Puntarenas: Runtarenas Cordillera de Tilaran,
Monteverde San Gerardo Biological Station. Sendero Ta-
bac6n, I). Pennys 448 (F, INB).

Eugenia locuples Standley & L. (). Williams ex
Barrie, sp. nov. TYPE: Honduras. FI Parafso:
orilla de Quebrada Siete Pasos y vecindades
NE de Dan If, 14 Mar. 1963 (ft), A. Molina R.
I I3B7 (holotype, F; isotype, NY). Figure 22.

Eugeniae quercelorum aemulans, sed rhachidibus 1-2
mm longis, pedicellis 2—4 mm longis et fructibus globosis
vel oblatis, 7—9 mm diametro differt.

Shrubs or small trees, to 5 m; young stems,
leaves, and inflorescences coarsely rufous-tomen-
tose, the hairs 0.2-0. 5 mm, dibrachiate and con¬
torted; branchlets terete or weakly compressed;
bark flaky, tan or gray appressed-pubescent, the
hairs cinereous. Leaves elliptic, the blades 4—7.5
X 1—3 cm, 2.2—3 times as long as w ide, coriaceous,
drying a dark green or olive above, lighter below;

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

31

Figures 21. 22. — 21 (left). Eugenia luhosperma Barrie ( Bello 4-146: holotype. INB). — 22 (right). Eugenia locuples Standley & L. 0. Williams ex Barrie ( Molina It 11387 ; holotvpe. F).

32

No von

midvein impressed above, elevated below; lateral
veins 8 to 12 per side, plus intermediates; marginal
veins similar to the lateral veins and weakly arched
between them, 1-2 mm from the margins; upper
surface of mature leaves with a few scattered hairs,
eventually glabrate and nitid. the lower surface re¬
maining villosulous much longer, the hairs cinere¬
ous, hut ultimately glabrate; glands obscure on both
surfaces, or witli scattered, dark glands on the lower
surface only; base cuneate; margins revolute; apex
acute to acuminate; petioles 4—5 mm long, im¬
pressed dorsally, ultimately glabrate. Inflorescences
axillary, short-racemose, solitary, the axis, bracts,
and buds rufous-tomentose; axis 1-2 mm long, of¬
ten obscured by t lit* buds and flowers; flowers 5 to
7, a terminal flower sometimes present; bracts 1—2
mm long, persistent through anthesis, absent in
fruit, eymhiform, t lie apex acute; buds 4—5 mm
long, globose or obovoid. Flowers pedicellate, the
pedicels, bracteoles, hypanthium, and calyx rufous-
tomentose; pedicels 2—4 mm long; bracteoles 1.5—

2 X 0.5 mm, deciduous at anthesis or soon after,
elliptic, the inner surface glabrous, the base free,
the margins obscured by the vestiture, the apex
acute; hypanthium 1.5—2 mm long; obeonic or cam-
panulate, calyx lobes in subequal pairs. 2—3 X 2—

3 nun. the outer surface densely rufous-tomentose,
the inner surface glabrous, the apex bluntly acute
or rounded; petals 4—5 X 3-4 mm, elliptic or ob-
ovate, the margins ciliate, the apex rounded; disk
2.5—3 mm diam., with scattered, erect hairs; sta¬
mens 70 to 100, 5—7 mm long; style 6—8 mm long,
glabrous. Fruits 7—9 X 7-9 nun. globose or oblate,
tin* base entire or substipitate; pericarp thin-walled,
glandular, weakly ribbed, thinly vested with pale,
dibrachiate hairs, more densely so at the base and
apex; calvx persistent, erect, tomentose without,
glabrous within; mature color red.

Habitat, distribution, and phenology. Open,
mixed forests in Honduras and Nicaragua at ele¬
vations of 500— 900 m. Collected in flower January—
April; collected in fruit April— July.

Between 1950 and 1970. several sets of speci¬
mens were widely distributed under the names
“Eugenia locuples Standley & Williams and “Eu¬
genia quercetorum Standley & Williams.” Never
validly published, the names did appear in print at
least once as nornina nuda (Molina, 197 ’5: 82). Al¬
though two species are clearly involved, the names
were' not applied consistently and specimens of ei¬
ther species may be labeled with either name. Both
species are shrubs or small trees of 5 m or less,
with the emergent growth coarsely tomentose with
dibrachiate, coppery-golden hairs, which are ulti¬

mately shed. Eugenia locuples differs from E. quer¬
cetorum Barrie in having narrower leaves (2.2-3 vs.
1.3— 2.3 times as long as wide), much shorter inflo¬
rescence axes (1—2 mm vs. 5-25 mm), shorter ped¬
icels (2-4 mm vs. 3—10 mm), and smaller fruits,
fhe juvenile vestiture tends to persist on the un¬
derside of the leaves in E. locuples, though it is
ultimately lost.

Etymology. The epithet translates as “rich” or
“opulent” and is probably a reference to the luxu¬
riant, reddish gold tomentum that covers the emer¬
gent growth.

Paratypes. HONDURAS. El Paraiso: en matorrales
de If. Vallecillo, NE de Danlf, A. Molina R. 7591 (E[2j);
orillas del R. Teupasenti ca. del pueblo de Teupasenti, A.
Molina R. 11900 (F, (41. NY); Sierra El Chile entre El
Robledal y El Junquillo, A. Molina R. 14190 (E. I.I.. NY);
km 73 entre El Ojo de Agua y Rfo California, A. Molina
R. 14485 (E, NY); along or near R. California, betw. If.
Choluteca & Jacaleapa, P. C. Standley 17182 (F); It. Cal¬
ifornia, base of Sierra de la Villa Santa, P. C. Standley
28112 (F); canyon ea. 3 km W of Danlf, /,. O. Wdliams
18170 (F, GH). Francisco Morazon: a orillas de la Que-
brada Cantarranas NE de Cantarranas, A. Molina R. 7804
(E, (41). NICARAGUA. Esteli: along Rfo Estelf. ca. 8 km
SW of Estelf, A. Molina R. & /.. 0. Williams 20180 (E.
NY. US, WIS[2|). Jinotega: along trail betw. Jinotega &
Las Mesitas. W of Jinotega. P. C. Standle > 9722 (E); along
trail betw. Jinotega & Las Mesitas, W of Jinotega. P. C.
Standley 9748 (F); region ol La Montanita & Las Mesitas.
in sierra \X ol Jinotega, I1. C. Standley 10898 (E. MICH).

Eugenia mugniflora Barrie, sp. nov. TY l’E: Costa
Rica. Limon: Pococi, R.N.F.S. Barra del Col¬
orado, Elanura del Tortuguero, Sardinas. 1 Jan.
1995 (b, fl), F. Araya 705 (holotype, INB: iso¬
type, F). Figure 23.

Ab aliis Eugeniae mesoamericanae floribus maximis et
antheris linearis differt.

Shrubs or trees to 7 in or more; branehlets. buds,
emerging leaves, and inflorescences tomentose, the
hairs simple and dibrachiate, contorted, ochra-
ceous; branehlets compressed, glabrate; bark white,
flaky. Leaves obovate to elliptic, the blades 20—35
X 9—16 cm, 2—2.3 times as long as wide, charta-
ceous. drying dark green above, tan below; midvein
flat above, with one or several parallel, irregular
ridges, elevated below; lateral veins 13 to 15, flat
or inscribed above, raised below; marginal veins
similar to lateral veins, weakly arched, ca. 5 mm
from the margins; upper and lower surfaces sparse¬
ly pubescent or glabrate, the midvein persistently
pubescent above and below; glands obscure or nu¬
merous. minute and pitted above, scattered and
dark below; base cuneate; margins entire; apex
abruptly acuminate; petioles 10—15 mm long, flat¬
tened or weakly indented dorsally, pubescent. In-

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Barrie

Eugenia from Mesoamerica

33

23, 24. -23 (left). Eugenia magniflora Barrie (Araya 705 ; isotype. F). -24 (right). Eugenia mcphersonii Barrie [McPherson 13579 : holotype, MO).

34

Novon

florescences cauliHnmus, fasciculate; bracts 4-3 X
1 nun. deciduous around anthesis, linear, tomentose
on the inner and outer surfaces, the margins ciliate.
the apex acute; huds 20 X 20 mm, globose, to¬
mentose. Flowers extremely large, pedicellate, the
pedicels, bracteoles, hypanthium, and calyx dense¬
ly tomentose with ochraeeous, contorted, simple
and dibrachiate hairs: pedicels 6— 10 X 2—2..') mm;
bracteoles fused, forming a 4-lobed involucre, ea.
10 mm diam., tlx- inner and outer surfaces tomen¬
tose, the margins ciliate. the apices of the lobes
acute; hypanthium 5-6 mm long, obconic, promi¬
nently tube inflate, the base attenuate; calyx lobes
in unequal pairs, the larger pair 20—25 X 20—25
mm, hooded, enveloping the expanding bud, the
inner and outer surfaces tomentose, the margins
fused approximately half their length, splitting at
anthesis, the apex rounded; petals not seen; disk
12—15 mm diam., pubescent at the base of the
style; stamens ea. 600, 10 mm long, the anthers
2.5-3 mm long, orange; style 15 mm long, glabrous.
Fruits 35 X 35 mm, subglobose; pericarp fleshy,
luberculate but not noticeably costate, uniformly
covered with ochraeeous hairs when immature; ma¬
ture fruits not seen.

Habitat, distribution, and phenology. An un¬
derstory tree or shub of the wet lowland forests
along the Atlantic slope in Costa Rica at elevations
of 0-100 m. Collected in flower in January and
February.

Eugenia rnagniflora is readily distinguished from
other Mesoamerican Eugenia by the large eauliflo-
rous flowers with the bracteoles fused, forming a 4-
lobed involucre, the tuberculate hypanthium, the
partially fused, hooded calyx lobes, the linear an¬
thers, and tuberculate fruits. Though in all ways
larger, the flowers are somewhat similar to those of
E. feijoi (). Berg, a predominately South American
species that has been collected in Panama, notably
in partially fused calyx lobes and the linear an¬
thers, here 2.5—3 mm long, with a terminal ap¬
pendage. In most species of Eugenia the anthers
are oblong, less than I mm long, and terminated
by a gland. According to the label on the type of
E. rnagniflora. the anthers are orange, an unusual
color in Eugenia, where the anthers are typically
white or pale yellow.

Paratype. COSTA KICA. Heredia: along trail
through forest between finca Bella Vista (a.k.a. Plastieo)
on the Quebrafla l igre, and Las Horquetas, ea. 6 km SVi
of Las Horquetas, M. Grayum et id. 5068 (CR, MO. f).

Eugenia mcpliersonii Barrie, sp. nov. I 'i PL:
Panama. Chiriquf: Fortuna Dam region, along
trail to Cerro Hornito (Pate de Macho) on S
ridge of watershed, I < Jan. 1689 (b, fl), G.
McPherson 13579 (holotype, MO; isotype, CR).
Figure 24.

Al> Eugenia costaricensi florilms majoribus, braeteolis
eaducis et fructibus majoribus, ellipsoidalibus vel ovo-
ideis differt.

frees to 10 in; young growth apparently glabrous
except for a few simple hairs, ea. 0.1 mm, on the
buds, emergent leaves, and the margins of the
bracts, bracteoles, and calyx lobes; branchlets com¬
pressed; bark chestnut brown to gray. Leaves ellip¬
tic to obovate, the blades 3—6 X 1—2.5 cm, 2—3
times as long as wide, coriaceous, drying dull dark
brown above, reddish brown below; midvein flat or
convex above and below; lateral veins 10 to 14 per
side, straight and ascending; marginal veins ea. I
mm from the margins; upper and lower surfaces
glabrate; glands obscure above, obscure or scat¬
tered below; base cuneate; margins decurrent into
the splayed inner edge of the petiole; apex abruptly
acuminate to caudate-acuminate; petioles 2-5 mm
long, silicate, glabrate. Inflorescences axillary or at
leafless nodes, sessile, solitary; flowers 1 to 4;
bracts deciduous; buds 3-4.5 mm long, pyriform.
Flowers pedicellate, the pedicels 5-9 mm long,
weakly compressed, glabrous; bracteoles 0.3— 0.5 X
0.7— 0.7 mm, deciduous in flower, the base free, the
margins scarious and occasionally ciliate, the apex
rounded, glabrous or with tufted hairs; hypanthium
1.5— 2.5 mm long, campanulate, glabrous, calyx
lobes in unequal pairs, the margins scarious, gla¬
brate or with a few scattered hairs, the apex round¬
ed. the smaller pair 1.5-3 X 2-2.5 mm, the larger
pair 3-4.5 X 2-3 mm; petals 4-5 X 3.5-4 mm,
elliptic, the margins entire, the apex rounded; disk
2-3 mm diam., with scattered, erect, coppery hairs;
stamens 70 to 90, 4-4.5 mm long; style 5 mm long,
glabrous. Fruits 20-25 X 10-15 mm, ellipsoid or
ovoid; pericarp thin-walled, glandular, glabrous; ca¬
lyx persistent, erect; mature color red to black.

Habitat, distribution, and phenology. Known
only from the montane rain forests and cloud forests
in western Panama at elevations of 1 100— 2200 m.
Collected in flower March— May; collected in fruit
November-February.

The leaves and inflorescences of Eugenia
mcpliersonii are similar to tlx* more widespread E.
costaricensis 0. Berg, differing in the slightly larger
flowers with deciduous bracteoles and ellipsoid
rather than globose fruits. The flowers and fruits of

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

35

h. costaricensis are commonly stipitate or substip-
itale; in this species, they are sessile.

Etymology. Named for the collector of the type,
Gordon McPherson, noted authority on the Pana¬
manian flora.

Paratypes. PANAMA. Boras del Toro: Germ Colo¬
rado mine area, divide road; along trail into Bocas «X in
woods on Pacific slope, from Chami Sta. to ca. 9 mi. along
rd., B. Hammel & J. Trainer 14920 (F). Chiriqui: 3.5 mi.
NK of Boquete, end of road along B. Palo Alto. B. A.
Hammel 5699 (MO); end of rd. past Palo Alto E of Bo-
quete. B. A. Hammel 6068 (MO); Fortuna Dam region,
along trail to Gerro Hornito (Pate de Macho) on S ridge of
watershed, G. McPherson 13570 (CR, F); N of San Felix
at Chiriqui— Bocas del Toro border, on Gerro Colorado cop¬
per mine rd. along Continental Divide, S. Mori & J. Kal-
Innki 5805 (MO), S. Mori & J. Kallunki 5936 (MO); near
Gerro Colorado, ca. 4.2 mi. from Chami Camp, G. Mc¬
Pherson 8961 (CR, F, MO); Boquete, Cerro Horqueta,
along path, C. von Hagen & IT. von Hagen 2030 (MO
NY).

Eugenia molinae Barrie, sp. nov. TYPE: Hondu¬
ras. El Parafso: Matorrales de Quebrada El
Coyol entre El Junquillo y El Robledal, 12
June 1964 (fl), A. Molina R. 14138 (holotype,
E; isotypes, E, GH, NY). Figure 25.

Eugeniae rhombeae aemulans, sed ramulis, foliis et in-
florescentiis pilosulis, foliis ellipticis et fructihus ellipso-
idalibus differt.

Shrubs or small trees, to 5 m; stems and leaves
pilosulous, tlu1 hairs simple, erect or recurved, 0.3—
0.8 mm; branchlets compressed, hirsute; bark
smooth, tan. Leaves elliptic, the blades 3-5 X 1.8-
3 cm, 1.6— 2.2 times as long as wide, coriaceous,
concolorous; midvein flat above, convex below; lat¬
eral veins 6 to 8 pairs, obscure above, slightly
raised on the lower surface; marginal veins arched,
1~2 mm from the margins; upper and lower surfac¬
es uniformly pilosulous, the hairs persistent, widely
spaced; glands few, pellucid, scattered on the lower
surface only; base cuneate to rounded; margins car¬
tilaginous, decurrent along the petiole; apex bluntly
acute or rounded; petioles 2—3 mm long, flattened
or depressed dorsally, hirsute. Inflorescences axil¬
lary or eauhflorous, sessde or short-racemose, or
flowers solitary; axis 0-1.5 mm long, pilosulous;
flowers 1 to 6; bracts 1.5— 2.5 mm, deciduous, cym-
biform. hirsute, the margins ciliate, the apex acute;
buds 3—3.5 mm long, obconic, hirsute. Flowers
pedicellate, the pedicels 5—9 mm long; bracteoles
0.5-0. 9 X 0.3 mm, persistent, elliptic, hirsute, the
base free, the margins ciliate, the apex acuminate;
hypanthium 1.5 mm long, campanulate, glabrous or
with scattered, erect hairs, contrasting with the* hir¬
sute pedicel and bracteoles, the base attenuate or

substipitate; calyx lobes ovate, concave, in unequal
pairs, vestiture similar to the hypanthium, the mar¬
gins scarious-ciliate, the apex rounded, the smaller
pair 1.5 X 2 mm the larger pair 2.5-3 X 2 mm;
petals 3.5—6 X 2-2.5 mm, the margins ciliate, the
apex rounded; disk 2—2.5 mm diam., glabrous or
the staminal ring hirsute; stamens 60 to 70. 4.5-
5.5 mm long; style ca. 6 mm long, glabrous. Fruits
8-10 X 7-8 mm. broadly ellipsoid; pericarp thin-
walled, glandular, glabrous; calyx persistent, erect;
mature color black.

Habitat, distribution, and phenology. Xerophytic
forests, matorrales in Honduras at elevations of
600—1300 m. Collected in flower in June; collected
in fruit in September.

Eugenia molinae may be distinguished from oth¬
er Honduran Eugenia by the elliptic, short-peti-
oled. coriaceous leaves having both surfaces per¬
sistently and uniformly pilosulous with a mixture of
widely spaced, straight and recurved hairs.

Etymology. The epithet honors Antonio Molina
R. of the Escuela Agrfcola Panamericana, the noted
authority on the flora of Honduras and adjacent re¬
gions, the author of numerous new taxa. and a pro¬
lific collector. Among his collections are the types
for six of the species described in this publication.

Paratypes. HONDURAS. Intibuca: jet. of Jicarito &
Gallo Creeks, drainage of (lie R. Yeguare, A. Molina R.
152 (BM, F, GH, MO, US, WIS); vicinity of Mazaguara,
valley of Jesus de Otoro, A. Molina R. 22598 (F, MO, NY).

Eugenia monteverdensis Barrie, sp. nov. TYPE:
Costa Rica. Puntarenas: in Mata’s pasture.
Monteverde, 24 Aug. 1975 (fr), G. Hartshorn
1778 (holotype, CR; isotypes, CR, F, MO). Fig¬
ure 26.

Ah Eugenia tilarana foliis cum basibus revolutis, flo-
ribus majoribus, fructihus majoribus, ochraceis differt.

Trees to 20 m or more; young growth coppery
appressed-pubescent, the hairs dibrachiate;
branchlets compressed, weakly 4-angled; bark gray,
glabrate. Leaves elliptic or obovate, the blades 8-
12 X 2.5—6 cm, 2—2.7 times as long as wide, cliar-
taceous, drying dark green above, pale green or
brown below; midvein sulcate or impressed above,
raised below; lateral veins 8 to 10 per side, mod¬
erately curving-ascending, raised on the lower sur¬
face; marginal veins similar to the laterals and
arched between them, 1-2 mm from the margins;
upper and lower surfaces uniformly pubescent with
appressed coppery or cinereous hairs, the upper
surface soon glabrate, tlu* vestiture more persistent
on the lower surface, but it, too, ultimately glabrate
or with a few, scattered hairs; glands obscure above,

36

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Volume 15, Number 1
2005

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Eugenia from Mesoamerica

37

dark and numerous below: base cuneate, typically
strongly revolute near the petiole, often appearing
"pinched"; margins attenuate along tbe petiole;
apex acuminate; petioles 8-15 mm long, sulcate
dorsally, appressed-pubescent. Inflorescences axil¬
lary, racemose, solitary or more commonly 2 or 3
superposed, uniformly coppery appressed-pubes¬
cent; axis 10—20 X 1—2 nun, 4-angled; flowers 10
to 16; bracts persistent, 0.5-1. 5 mm long, eymbi-
form, glabrate in bud. dark and contrasting with the
coppery axis and pedicel, prominently glandular-
buds 3.5-5 mm long, obovoid. Flowers pedicellate,
tbe pedicels 4—10 mm long, 4-angled; braeteoles
persistent, separate or fused and forming an invo¬
lucre 2 mm across, dilated at anthesis, if separate
then 1—1.5 X I mm, ovate, glabrate and promi¬
nently glandular, dark and contrasting with the cop¬
pery pedicel and hypanthium, the base free, the
margins st ations, the apex acute; hypanthium 2—4
mm long, campanulate, entire or weakly ribbed; ca¬
lyx lobes widely ovate, in markedly unequal pairs,
coppery appressed-pubescent on both surfaces, tilt'
margins ciliolate, the apex rounded, the smaller
pair I X 2 mm, the larger pair 3—3.5 X 4 mm;
petals 4—4.5 X 3.5—4 mm, ovate, the margins en¬
tire, the apex rounded; disk 3—3.5 mm diarn., gla¬
brous or with a few, scattered, erect hairs; stamens
ca. 100. 5—7 mm long; style 7—9 mm long, glabrous,
fruits 25—35 X 20—22 mm. irregularly obovoid or
oblong, the base elongate; pericarp coriaceous, ru¬
gose, persistently dull brown-tomentose but ulti¬
mately glabrate; calyx absent; mature color dull
yellow.

Habitat, distribution, and phenology. Montane
forests of central Costa Rica at elevations ol 1000—
2000 m. Collected in flower January— November;
collected in fruit January— August.

Eugenia monteverdensis is one of three newly de¬
scribed species from tbe Monte verde region, the
others being E. tilarana and E. zuchowskiae, which
have been persistently identified as either E. octo-
pleura Krug & Urban or E. guatemalensis. All have
axillary, racemose inflorescences, solitary or several
superposed, with 4-angled axes, densely vested
with appressed, coppery, dibrachiate hairs. Eugenia
monteverdensis is distinguished by the elliptic or
obovate leaves with the margins strongly revolute
near the petiole and the yellow, irregularly obovoid
or ellipsoid fruits that lack ribbing. The fruits of E.
octopleura, E. tilarana, and E. zuchowskiae are
ribbed; the fruits of E. guatemalensis are unribbed,
but much smaller (5-12 X 5—9 mm).

Earatypes. COS1A RICA. Reserva Monteverde, Cor¬
dillera de Tilaran [covers parts of Alajuela, Puntarenas,

and Guanacaste], V. J. Dryer 1358 (CR, F), V J. Dryer
1626 (CR. I. M0). Alajuela: Reserva Forestal San Ra¬
mon. trail S of the station, ./. Bittner A G. Herrera 2028
(CR); I. a Palma de San Ramon. A. Brenes 4625 (F); Mon¬
teverde Cloud Forest Reserve, Penas Blancas R. valley on
Atlantic slope, IF A. Haber & E. Bello 5816 (MO); San
Ramon, Cordillera de Tilaran, El Castillo, primary forest
patch near Arenal Vista Podge, lado SF del lago, B. Har¬
mon 349 (IMF MO); Zarcero, A. Smith A239 (A, F, MO.
US); arbre au bord du R. Segundo, A. Tonduz 1712 (HR).
Cartago: entre Turrialba y Florencia, A. Jimenez M. 3318
(CR, F, MO); vicinity of La Congreja, ca. 10 km S of El
Tejar. Cord, de Talamanca, L 0. Williams et al. 24111
(F). Guanacaste: Monteverde, Transect 6, ,4. Gentry et al.
71589 (F. MO). Heredia: Zurquf. B. C. Standley & M.
Valerio 48254 (F). Puntarenas: ca. 2 km SW of Montev¬
erde, on the Pacific watershed, lb C. Burger & J. I,. Gentry
Jr. 8801 (F); Cordillera de Tilaran, camino que va de la
Resi jrva Ahsoluta de Cabo Blanco subiendo por alto San
Luis, A. Eerndndez 1386 (CR. INB); Monteverde, A. Gen¬
try A; lb A. Haber 48764 (CR, MO); Monteverde, com¬
munity. lb A. Haber 206 (CR, MO); Monteverde, near lab,
lb 4. Haber 224 (CR, MO); Monteverde. upper commu¬
nity. W. A. Haber 758 (CAS. MO), W. 4. Haber 764 (BM,
MO), lb A. Haber 765 (MO), lb A. Haber 766 (MO); Re¬
serva Bioldgica Monteverde, Pacific slope. Research Trail,
lb Haber et al. 10810 (CR, F, MO): Reserva Bioldgica
Monteverde, Pacific slope, road from field station to di¬
vide, lb Haber A1 lb Zuchowski 10852 (CR[2|, k). San
Jose: La Palma, A. Tonduz 12558 (A. F. OH. NY); San
Pedro de Coronado, M. Valerio 1643 (F).

Eugenia paloverdensis Barrie, sp. nov. TYPE:
Costa Rica. Guanacaste: Bagaces, P.N. Palo
Verde, Valle de Tempisque, Est. Palo Verde,
Cerro Guayacan, 23 June 1993 (fl). U. Cha¬
varria 826 (holotype, INB; isotypes, CR, F,
MO). Figure 27.

Eugeniae principium aemulans, sed indumentis ex parte
dibrachiatis, floribus et fructibus majoribus, subsessilibus
differt.

Shrubs or small trees; young growth coarsely ap¬
pressed-pubescent, the hairs simple and dibrachi¬
ate, coppery; branchlets and leaves soon glabrate;
branchlets somewhat flattened at the nodes; bark
brown to gray. Leaves elliptic to ovate or lanceolate,
the blades 3-6.5 X 1 .5-2.2 cm, 1.6-3 times as
long as wide, coriaceous, drying green to olive, con-
colorous or somewhat paler below, the midvein a
contrasting white or brown; midvein flat or concave
anil between 2 parallel ridges above, convex below;
lateral veins 6 to 8, straight; marginal veins similar
to the lateral veins, straight, ca. I mm from the
margins; both surfaces with appressed, cinereous
dibrachiate hairs somewhat persistent, but eventu¬
ally glabrate; glands pellucid, numerous and prom¬
inent on both surfaces; base rounded; margins
merging abruptly into the inner edges of the petiole;
apex acute or acuminate; petioles 1.5— 2.5 mm long,
pubescent with appressed, cinereous hairs or gla-

38

Novon

!7 (left). Eugenia pa l oven lens is Barrie (U. Chavarria 8'2f)\ holotype. INB). — 28 (right). Eugenia quercetorum Standley & L. 0. Williams ex Barrie (Molina

Volume 15, Number 1
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Eugenia from Mesoamerica

39

brate, channeled on die upper surface. Inflores¬
cences axillary, racemose, solitary, uniformly ap-
pressed-pubescent, coppery but soon cinereous;
axis L-3 mm long; flowers 2 to 6; bracts 1 X 0.5
mm, persistent, cymbiform, glabrate before the axis
and then darker in color and contrasting with it,
the margins entire, the apex acute; buds 2 mm,
obovoid. Flowers pedicellate, the pedicels 1-2 mm
long, approximately equal to the axis, bracteoles I
X 0.3 mm, persistent, lanceolate, conspicuously
glandular, glabrate before the hypanthium and then
darker in color and contrasting with it, the base
free, the apex acute; hypanthium 1 mm long, cam-
panulate, appressed pubescent; calvx lobes in un¬
equal pairs, the inner and outer surfaces ap-
pressed-pubescent. the margin entire, the apex
rounded to bluntly acute, the smaller pair 1-1.5 X
15—2 mm, the larger pair 1.5—3 X 1.5—2 mm; pet¬
als ovate, 4-4.5 X 4 mm. the margin ciliate near
the apex, the apex rounded; disk 2.5—3 mm diam.,
glabrous or with a few hairs at the base of the style;
stamens ea. 80, 5—8 mm long; style 6—7 mm long,
with a few basal hairs. Fruits 10-14 X 10-12 mm,
broadly ellipsoid to globose; pericarp thin-walled,
glandular, glabrous; calyx persistent, erect; mature
color purple-black.

Habitat, distribution, and phenology. Known
only from the lowland dry forest on the Pacific coast
in Guanaeaste Province, Costa Rica, at elevations
of 0—100 m. Collected in flower in June; collected
in fruit in October.

A small shrub or tree with leaves similar in size
and shape to Eugenia principium, E. paloverdensis
may be distinguished by its vestiture, a mixture of
simple and dibrachiate hairs, the flowers with ped¬
icels 1—2 mm long, and calyx lobes sinular in color
to tbe hypanthium. In Eugenia principium, the hairs
are simple, and the flowers are borne on pedicels
6-14 mm long and have pink, petaloid calyx lobes.

Paratypes. COSTA MICA. Guanaeaste: Bagaees.
PN. Palo Verde, Valle de Tempisque, Est. Palo Verde,
creciendo entre F i la Guayacan y Fila Cueva del Tigre, IJ.
Chavarria 850 (CM. K I [Nil, MO); Est. Mioldgiea Palo
Verde, M. E Quigley 937 (MO).

Eugenia quercetorum Standley & L. O. W il barns
ex Barrie, sp. nov. TY PE: Honduras. El Par-
aiso: drainage of the Rio Yeguare, en el Rib
Eizapa, Llano Lizapa, 17 June 1951 (b), A.
Molina R. 3946 (holotype, F; isotypes, RM,
GH, MO). Figure 28.

Eugeniae locuplete aemulans, sed infloreseentiis rhaeh-
idibus 5-25 mm longis, pedicellis 3-10 mm longis, frue-
libus ellipsoidalibus vel subglobosis, 13-17 mm longis
differt.

Shrubs or small trees 2—5 in; young stems, buds,
leaves, and inflorescences densely tomentose, the
hairs 0.3—1 mm, dibrachiate, contorted, rufous or
cinereous; branchlets weakly compressed; bark tan
or gray, flaky, ultimately glabrate. Leaves elliptic* or
widely elliptic to obovate, the blades 3.5-8 X 2-5
cm, 1.3-2. 3 times as long as wide, chartaceous,
drying dark green and nitid above, olive below;
midvein depressed or suleate above, convex below;
lateral veins 5 to 8 per side, obscure above on ma¬
ture leaves, raised below; marginal veins similar to
the laterals and arched between them. 1-2 mm
from the margin, obscure above; surfaces of young
leaves villosulous, the hairs rufous, dibrachiate,
surfaces of mature leaves glabrous or nearly so;
glands convex on the upper surface of young leaves,
obscure or rarely punctate on mature leaves, dark
and densely to thinly distributed over the lower sur¬
face; base cuneate to broadly cuneate or rounded;
margins revolute, decurrent into the inner edge of
the petiole; apex bluntly acute; petioles 4-10 mm
long, suleate dorsally, glabrate or with cinereous or
rufous, dibrachiate hairs. Inflorescences axillary,
racemose, solitary, the axis, bracts, and buds uni¬
formly rufous-tomentose, the hairs dibrachiate; axis
5—25 X 1 mm; flowers 4 to 11, a terminal flower
often present; bracts 1—2 mm long, deciduous, cym¬
biform, the margin scarious, the apex rounded.
Flowers pedicellate, the pedicels, hypanthium. and
calyx uniformly rufous-tomentose, the hairs dibra¬
chiate: pedicels 3—10 mm long; bracteoles 1.5— 2.5
X 1 mm, deciduous at or prior to anthesis, elliptic
or ovate, initially rufous-tomentose, becoming gla¬
brate and contrasting with the hypanthium before
falling, the base free, the margins ciliate, the apex
acute; hypanthium 1 .5—2 mm long, eampanulate;
calyx lobes in unequal pairs, the inner and outer
surfaces densely vested with contorted, rufous or
cinereous, dibrachiate hairs, the apex rounded, the
smaller pair 2-2.5 X 2-3 mm, the larger pair 3-
3.5 X 3—4 mm; petals 3.5— 4.5 X 4—4.5 mm, de¬
pressed ovate, glabrous or with a few scattered
hairs, the apex bluntly acute, the margins ciliate;
disk 3—4 mm diam., rufous-tomentose at base of
style; stamens ca. 80, 4—6 mm long; style 6—7 mm
long, with hairs scattered along entire length. Fruits
13-17 X 10 — 1 1 mm, sul)glol)osc or ellipsoid; pon-
carp thin-walled, glandular, weakly and irregularly
costate, with scattered, short hairs or a few dibra¬
chiate hairs at base and apex; calyx persistent, pu¬
bescent within and without; mature color deep pur¬
ple: or black.

Habitat, distribution, and phenology. Known
from oak forests and cloud forests at elevations of

40

Novon

400-1500 m. Collected in flower in July; collected
in fruit September— January.

The name Eugenia quercetorum, like E. locuples,
appeared on several sets of specimens widely dis¬
tributed 30 to 50 years ago. Though never validly
published, the name appeared in print at least once
as a nomen nudum (Molina, 1975). As noted in the
discussion of E. locuples, E. quercetorum differs
most notably in having broader leaves, shorter in¬
florescence axes, shorter pedicels, and larger fruits.

Etymology. The epithet refers to the species
oak forest habitat.

Earn types. HONDURAS. El Paraiso: drainage of the
K. Yeguare, Quebrada Cfiupadero, N de Yusear&n, A. Mo¬
lina R. 1684 (F, GH), A. Molina R. 1687 (F, GH. MO), A.
Molina R. 1665 (F, GH); drainage of die R. Yeguare, en
la Quebrada F.l Mum, entre F.l Empalme & El Mum. 20
km al N de Yusearan. A. Molina R. 1018 (F, Gil): orillas
de la quebrada, Valle Jainastran. A. Molina l\. 7488 (f );
Valle JamastrAn, A. Molina R. 7468 (F); a orillas del R.
Oropolt eerca de Oropolf, A. Molina R. 10078 (f[2|. NY );
entre San Lucas y Manzaragua, A. Molina R. 10750 (I5M.
F, GH, MO. NY. WIS); I’inares y colinas roeosas de Chu-
padero, earn a Yuscaran. A. Molina R. 14426 (F. IT. NY);
Quebrada do Dantas. on rd. between Las Mesas & Yus-
cantn. R. C. Standley 14971a (F). R. (■■ Standley 14973a
(F. MICH), R. C. Standley 14685 (F); Germs el /apotillo,
road betw. Zarnorano & Gufnope, (•. E. Webster el al.
11678 (F, GH, MICH|2|, MO); valley of R. Dantas. bar-
ranco El Mum. G. E. Webster et al. 12042 (MICH|2|); Que¬
brada El Mum botw. Las Mesas & Yusearan. /,. 0. Wil¬
liams 42806 (F). Francisco Mora/.an: along paved b\\\.
between Tegucigalpa N lalanga. 35 km N of Iegucigalpa,
T. II. Groat & I). R. Hannon 03608 (MO); entre Cuesta de
Los Mucrtos y Monte Obseuro, Cordillera A/.acualpa, A.
Molina R. 14516 (F. NY); between Tegucigalpa & Puente
Colorado, near Choluteea R. on way to lalanga, A. Molina
R. A- A. R. Molina 25827 (F).

Eugenia riosae Barrie, sp. nov. I Y PE: Costa Hica.

Guanacaste: PN Guanacaste, Estacidn Piti 1 la.

Sendero El Mismo, Finca La Pasompa, 9 Dec.

1990 (II). Rios 257 (holotvpe, INB: isotypes.

CB. F, MO. SKI.). Figure 29.

Habitu Eugenia resca similis. sed bracteolis parviorilnis
cl cadueis, hypanthio conspicue glanduloso, fructibus ir-
regulariter ellipsoidalibus vcl globosis, 10— 2. > nun longis
differt.

Trees 5—10 m; young growth apparently coarsely
tomentose, the hairs ca. 0.5 mm, simple or dibra-
cliiate, soon reduced to rclictual hairs in the axis
of the leaves and hranchlets only; hranchlcts flat¬
tened at nodes; hark gray, puberulent. the hairs
0.1— 0.2 mm, simple, erect, or glabrous. Leaves el¬
liptic to obovate, the blades 7-12 X 3—6 cm, 1.6—
2.3 limes as long as wide, ehartaeeous, drying olive
above, yellow-green below; midvein flat to silicate
above, convex below; lateral veins 12 to 14 per

side, plus intermediates; marginal veins similar to
the laterals and weakly arched between them, 1-2
mm from the margins; both surfaces of mature
leaves glabrous; glands pellucid, prominent, large
and small on both surfaces; base euneate; margins
decurrent into the inner edge of the petiole; apex
abruptly acuminate; petioles 5—7 mm long, concave
dorsally, minutely puberulent. Inflorescences axil¬
lary and cauliflorous, solitary, fasciculate; flowers 8
to 10; bracts 0. 2-0.3 mm long, deciduous prior to
anthesis, minute, triangular, minutely puberulent;
buds 2-3 mm long, turbinate. Flowers sessile or
nearly so. the pedicels less than 0.5 mm long, con¬
spicuously glandular; bracteoles 0.5— 0.7 X 0.5
mm, deciduous soon after anthesis, ovate, conspic¬
uously glandular, the margins scarious-ciliate, the
apex acute or rounded; hypanthium 1.5—2 mm long,
campanulate, with or without an extremely short
stipe, glabrous or with a few' hairs, conspicuously
glandular; calyx lobes in markedly unequal pairs,
white or cream, the color contrasting with the dark¬
er hypanthium. glabrous, conspicuously glandular,
the margins entire, the apex rounded, the outer pair
1 — 1.5 X 1.5 mm, the inner pair 2—3.5 X 2.5 mm;
petals 5 X 2—3 mm. oblong, the margins entire or
with a few hairs, the apex rounded; disk 2 mm. with
a few, tufted hairs; stamens 40 to 50, 4.5-7 mm
long; stvle 7-8 mm long, glabrous. Fruits 10—25 X
10-14 mm. irregularly ellipsoid or globose, the
base often substipitate; pericarp thick, fleshy,
prominently glandular, glabrous or with a few hairs,
the calyx persistent; mature color red.

Habitat, distribution, and phenology. Dry for¬
ests along the Pacific slope in Costa Hica at ele¬
vations of 500-1000 m. Collected in flower in De¬
cember— January; collected in fruit March— April.

The elliptic to obovate leaves and fasciculate in¬
florescences of Eugenia riosae are similar to those
of tin1 Guatemalan species E. vesca Lundell. How¬
ever. in E. vesca the vestiture is composed solely
of simple hairs, the flowers and fruits are subtended
by broadly ovate bracteoles that are connate at the
margins, and the fruits are globose. 5-8 mm in di¬
ameter.

Etymology. The epithet honors Petrona Bios,
the collector of the type, a parataxonomist for the
Instituto Nat ional de Biodiversidad. Costa Rica.

Raratypes. COSTA Bit. A. Alajuela: PN Kmcon tie
La Vieja, Camino a Colonia Blanca, entre Quebrada Ran¬
cho Grande y finca Los Mora, G. Ruera 1 184 (CR, I'. INB.
MO); Upala, PN Guanacaste, Cordillera de Guanacaste,
Estacidn San Ramon, Sendero el Nispero, l\ Quesada 288
(CR. E. INB. MO). Guanacaste: La Cruz, PN Cuanacasle,
Est. Piti I la. Sendero los Memos, E. Alfaro 182 (CR, 1.
INB. MO); Tilaran ZP Tenorio, Berras Morenas, Rio San

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

41

Figures 29. 30. — 29 (left). Eugenia riosae Barrie (Rios 257: holotype, I\B|. — 30 (right). Eugenia sancarlosensis Barrie ( Jimenez et al. 1023: holotvpe. CR).

42

Novon

I .orenzo, C. Alvarado 115 (CR, INB. K); Tilanln, Cordil¬
lera dc Tilaran. Palmital, Quebrada Malanga, Rfo Cano
Negro, E. Hello A hi. Cruz 4884 (INB), hi. Hello & E. Cruz
4898 (CM, INB): Liberia, BN Guanaeaste, Nueva Xelan-
dia, Est . San Ramon, Sendero al Nfspero, I). Garcia 153
(INB, MO); BN Guanarasle. Estaeidn Bitilla, Eila Orosililo
y Sendero El Misnio. I\ Rios 322 (CB, INB. MO): lilaran
ZB Tenorio TilartSn. Cord, de Guanaeaste, lierras Moren-
as, alrededores Rfo San Lorenzo, G. Rodriguez 141 (CB.
E, INB, MO); Tilardn ZB I’enorio lilaran. Cord. V. lilanin,
Tierras Morenas, Rfo San Lorenzo, G. Rodriguez A Q.
Jimdnez 82 (E INB. MO). Heredia: Sarapiquf llorquetas,
Est. El Plastieo, 15 km de Horquetas, Sendero El Tigre,
J. Sanchez et al. 31 1 (CB, E).

Eugenia sancarlosensis Barrie, sp. nov. TYPE:
Costa Rica. Alajuela: San Carlos, 7 km al N
de Boca Tapada, Finca Hogar de Ancianos, 24
Sep. 1991 (fl). (J. Jimenez el al. 1028 (holo-
tyj>e, CR; isotypes, CR, F, INB. MO). Figure
30.

Ab Eugenia herrerae foliis cum costa planiusenla, flo-
ribus cum hypantio globoso, braeteolis apicibus eaducis,
basibus eonnatis, persistentibus, inerassatis differt.

Large trees, to 35 m or more; young growth not
seen; branchlets terete, the nodes swollen; bark
drying dark reddish purple, thinly pubescent with
erect, simple hairs, ea. 0.5 mm. Leaves narrowly
elliptic, the blades 13—20 X 3—6 cm, 3 to 4 times
as long as wide, chartaceous or subcoriaceous, dry¬
ing a dull olive above, yellow-green below, the mid¬
vein tan. Hat or with I or 2 thin, parallel ridges
above, convex below; lateral veins 10 to 18 per
side, raised below; marginal veins similar to the
laterals and arched between them, 3-4 mm from
the margins; both surfaces of mature blades gla¬
brous; glands obscure above, small anti scattered
below; base cuneate; margins entire; apex acumi¬
nate; petioles 5—8 mm long, reddish purple, similar
to the branchlets, flattened or concave dorsally. In¬
florescences axillary or possibly cauliflorous, fas¬
ciculate; (lowers 8 to 12; bracts reduced or absent;
buds not seen. Flowers pedicellate, the pedicels
10—15 mm long, glabrous; braeteoles connate and
thickened at the base, the basal portion ultimately
forming a ring beneath the hypanthium and devel¬
oping fruit, the apex 0.5 X 0.5 mm. triangular, de¬
ciduous at anthesis, the margins ciliate, the apex
acute; hypanthium ea. 4 mm long, globose, proba¬
bly glabrous; calyx lobes in subequal pairs. 3—3.5
X 2 mm, triangular, fused basally, forming a short
tube ca. I mm long, the outer surface glabrous, the
inner surface densely pubescent, the hairs 0.1— 0.2
mm, appressed, the margins entire, the apex acute;
corolla not seen; disk 3—3.5 mm long, glabrous; sta¬
mens ea. 150; style ca. 5 mm long, glabrous. Im¬
mature fruits 10 X 10 mm, globose, constricted

beneath the calyx; pericarp thick and fleshy, gla¬
brous; calyx persistent, spreading, mature color un¬
known.

Habitat, distribution, and phenology. Known
from the wet lowland forests on the Atlantic slope
in northeastern Costa Rica and adjacent Nicaragua
at elevations of 0—300 m. Collected in flower in
September.

Eugenia sancarlosensis is similar to E. herrerae
in having red young growth and petioles, this is a
much larger tree, however, with branchlets that are
often swollen at the nodes and leaves with a flat,
rather than incised, midvein. The globose hypan¬
thium is subtended by connate braeteoles with de¬
ciduous tips. After the tips have fallen, the persis¬
tent bases form a distinctive ring beneath the
flower.

Paratypes. COSTA RICA. Alajuela: San Carlos,
Llanura de Sail Carlos, Bilal. Boca Tapada, Finca Hogar
de Ancianos, A. Zamora A- A. Zeleddn 2352 (INB); San
Carlos Llanura de San Carlos, Boca Tapada, Finca Aser-
radero San Jorge, A. Zamora 2357 (INB). NICARAGUA.
Atluntico del Sur: Bluefields. Sitio Fonseca, aprox. ()2
km al NE a lo largo del Rfo Kama, A. Zamora A M.
Castrillo 2049 (INB).

Eugenia selvana Barrie, sp. nov. TV PE: Costa
Rica. Heredia: Finca La Selva, the OTS field
station on the R. Puerto Viejo just E of con¬
fluence with R. Sarapiquf, betw. Arboretum &
Near Loop Trail, 14 May 1980 (fl). II. Hummel
8662 (holotype, MO: isotype, F). Figure 31.

Eugeniae venezuelensi aemulans, sed indumento pub-
erulo, lobis ealyeis anguste triangulis, pallidis. fructibus
12-15 mm diametro differt.

Trees to 10 m or more; branchlets and inflores¬
cences minutely puberulent, the hairs 0.1— 0.2 mm.
simple, erect; emergent leaves puberulent along the
midvein and margins; branchlets compressed, slen¬
der; bark brown, smooth, puberulent. Leaves ellip¬
tic to ovate, the blades 4—9 X 1.5—3 cm. 2.5—3
times as long as wide, chartaceous, drying dark
green above, pale green below; midvein depressed
above, convex below; lateral veins 6 to t per side,
plus a lew intermediates, strongly curving-aseend-
ing, prominent on the lower surface; marginal veins
similar to the lateral veins and strongly arched be¬
tween them, 2-3 mm from the margins; upper sur¬
face persistently puberulent along the midvein, the
lower surface with scattered hairs along the mid¬
vein and the lateral veins, the margins ciliate,
sometimes along the lower half only; glands obscure
above, numerous, small and dark below; base
cuneate; margins entire; apex acuminate to cau-

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

43

31 (left). Eugenia selvana Barrie ( Hammel 8662: holotype, MO). — 32 (right). Eugenia shimishito Barrie ( Reyna 733: holotype, MICH).

44

Novon

date-acuminate; petioles 1—3 mm long, flattened or
concave dorsally, persistently puberulent. Inflores¬
cences axillary, solitary, short-racemose, persistent¬
ly puberulent; axis 1-4 mm long; flowers 2 to ( >;
bracts 0.5 mm long, persistent at least through an-
tliesis. elliptic, puberulent on the outer surface, the
margins ciliate, the apex acute; buds 1.5—2 mm
long, globose. Flowers short-pedicellate, the pedi¬
cels I mm long, persistently puberulent; bracteoles
0.3— 0.7 X 0.3 mm, persistent, ovate or ellipsoid,
puberulent, the base free, the margins ciliate, the
apex acute; hypanthium 1 mm long, cainpanulate;
calyx lobes in subequal pairs, 1.5— 2.5 X 0.5 mm,
narrowly triangular, white and somewhat petaloid,
glabrate, the margins ciliate, the apex acuminate,
the tip sharply pointed; petals 2—3 X 1—2 mm, el¬
liptic, concave, the margins ciliate, the apex round¬
ed; disk 1—1.5 mm diam., with scattered, minute
hairs; stamens 20 to 25, 3—5 mm long; style 5-7
mm long, glabrous. Fruits 12—15 X 12—15 mm. glo¬
bose or subglobose; pericarp thin-walled, glandular,
glabrous; calyx persistent but inconspicuous; ma¬
ture color unknown.

Habitat, distribution, and phenology . Lowland
wet forests on the Atlantic slope of Costa Rica at
elevations below 100 m. Collected in flower May-
June; collected in fruit July-February.

The leaves and short-racemose inflorescence
with nearly sessile flowers are similar to those of
Eugenia venezuelensis (). Berg, a species found in
Panama and from Nicaragua to Chiapas, Mexico,
but which is not known from Costa Rica. Eugenia
selvana differs in the minutely puberulent vestiture
of hairs 0. 1-0.2 mm long and the narrowly trian¬
gular, sharply pointed calyx lobes. I he vestiture of
E. venezuelensis is a denser covering of longer, 0.5—
I mm, straight and recurved hairs, and the calyx
lobes are rounded.

Haratypes. COSTA RICA. Heredia: Sendero entre el
campamento Canta Rana y R. I’eje, Magsasay, I. A. Cha-
c6n 79 (CR, MO); S of Puerto Viejo. 2 km S of Magsasay
Penal Colony, W of the road, N. Garwood et id. 1 155 (BM.
CR, K MFXU. MO); Finca I a Selva, Puerto Viejo de Sar-
apiquf, at edge of forest along W boundary, W of Research
Trail, M. II. Gray am 1990 (F. MO); La Selva, Rfo Sara-
piquf near Puerto Viejo, jet. SSO & LOG trails, A. Gentry
tV R. Ortiz 78554 (F); Magsasay, near La Selva, Transect
10, A. //. Gentry et al. 7I700A (MO); Magsasay, near La
Selva, Transect 10. A. //. Gentry 7I778A (MO); Finca l.a
Selva, the OTS field station on the R. Puerto Viejo just F,
of confluence with R. Sarapiquf, F River Rd., 620 S X
1220 K m grid, R. Ilammel 9181 (F, MO); Sarapiquf. Fin¬
ca La Selva, Puerto Viejo, W boundary, along Hunters
pejibayes, G. S. Hartshorn 1241 (F, MO. NY); Sarapiquf,
Finca La Selva, Puerto Viejo, W boundary, along Hunter's
pejibayes, G. 8. Hartshorn 1800 (CR, F, MO. NV ); along
Starkey Road. 4.5 km SW of bridge at Puerto Viejo, 14.

I). Stevens 18488 (CR. MO). Liiuon: Bri-Bri. /.. ./. Hove da
1158 (F. MO).

Kiigeniu sliiniisliilo Barrie, sp. nov. 1 4 PL: 1.1 Sal¬
vador. Santa Anti: cloud forest of Monteeristo,
5 Jan. 1979 (fl. fr). M. E. Reyna 788 (holotvpe.
MICH; isotype. MICH). Figure 32.

Ab Eugenia glandulosopunctata indumentis sericeis,
fruet i bus dupio longioribus differt.

Shrubs or small trees; new growth sparsely se¬
riceous. with simple, silky hairs; branchlets terete
or weakly compressed; bark tan or gray, glabrate.
Leaves ovate, the blades 4—7 X 2—3 cm, 2.2— 2.5
times as long as wide, ehartaceous, drying eonco-
lorous green, the midvein tan below; midvein sili¬
cate above, convex below; lateral veins 12 to 14
per side, plus intermediates, prominent on both
surfaces; marginal veins similar to the laterals and
arched between them, 1-2 mm from the margins;
glabrate; glands obscure on both surfaces; base rn-
neate; margins decurrent into the inner edge of the
petiole; apex acuminate or caudate-acuminate; pet¬
ioles 5—7 mm long, channeled or sulcate dorsally,
glabrate or with a few, scattered hairs. Inflores¬
cences axillary, fasciculate or very short-racemose,
solitary, glabrate or with a few, persistent hairs, axis
0 I mm long; flowers 2 to 4; bracts to 0.5 nun long,
probably persistent but minute, glabrate, the mar¬
gins entire, the apex acute; buds 4—5 mm long,
obovoid. Flowers pedicellate, the pedicels 10—20
mm long, gracile, with a few appressed hairs; brac¬
teoles 1-1.5 X 0.3 mm. persistent through anthesis
but absent in fruit, lanceolate, the back keeled, the
base free, the margins ciliate, the apex acute; hy¬
panthium 1—2 mm long, cainpanulate. sparsely se¬
riceous; calyx lobes in subequal pairs, 2-2.5 X 2
mm, triangular, glabrate, the margins ciliate, the
apex bluntly acute; petals 3-5 X 2.5-3.5 mm.
ovate, the margins entire, the apex rounded; disk
2—2.5 mm diam., coarsely pubescent; stamens ca.
80, 4—5 mm long; style 5 mm long, glabrous. Fruits
35—45 X 15—20 mm, ellipsoid, the base and apex
abruptly attenuate; pericarp thin-walled, glandular,
glabrous; calyx lobes usually persistent, reflexed;
mature color pale yellow.

Habitat, distribution, and phenology. Known
only from the cloud forest atop Cerro Monteeristo,
Santa Ana province, Fl Salvador, at elevations of
1900-2300 m. Collected in flower in January; col¬
lected in fruit in September.

Eugenia shimishito is distinguished by the seri¬
ceous emergent growth, the few-flowered inflores¬
cences, and the flowers and fruits on pedicels 10-
20 mm long. The fruits are similar in shape to those

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

45

of the Costa Rican species E. glandulosopunctata ,
Rut twice as large. That species also has prominent,
pellucid glands on the leaves, flowers, and fruits.

Etymology. The epithet “shimishito” is, ac¬
cording to the label on the type, the local vernac¬
ular name for this species. As a noun in apposition,
it retains its original gender.

Paratypes. EL SALVADOR. Santa Ana: San Jose In-
genio, I’.N. Montecristo, la quebrada del pico gordo, V. M.
Martinez 599 (LAGU, MO); San .lost'- Ingenio, P.N. Mon¬
tecristo, parqueo, V. M. Martinez 754 (LAGIJ, MO);
bosque nebuloso Montecristo, sendero a Gerro Brujo, M.
L. Reyna 1408 (MO).

Eugenia tilaraiia Barrie, sp. nov. TYPE: Costa
Rica. Puntarenas: Monteverde, Reserva Biol-
dgica, 8 Aug. 1971 (b, fl), E. ./. Poveda II II
(holotype, CR; isotypes, F, MO). Figure 33.

Ab Eugenia guatemalensi inflorescentiis majoribus,
fructibus ellipsoidalibus vel obovoideis, costatis differ!.

Frees to 20 m or more; young growth coppery
appressed-pubescent, the hairs dibraehiate;
branchlets compressed; bark tan or brown, smooth,
glabrate. Leaves elliptic or obovate, the blades 5—
9 X 2.5— 3.5 cm, 2.0— 2.6 times as long as wide,
chartaceous, drying a nearly concolorous, dark
green or paler below; midvein sulcate above, con¬
vex or raised below; lateral veins 8 to 10 per side,
prominent on both surfaces, ascending and weakly
if at all curved; marginal veins similar to the lat¬
erals and arched between them, 1—2 mm from the
margins; both surfaces soon glabrate; glands on
both surfaces raised, numerous and small, or ob¬
scure above; base eiineate; margins entire, flat or
weakly down-turned near the base, decurrent along
the petiole; apex acuminate, the acumen 5—15 mm
long; petioles 5—10 mm long, sulcate dorsally, gla¬
brate or with a few persistent hairs. Inflorescences
axillary, racemose, solitary or 2 superposed, the
axis, bracts, and buds uniformly coppery ap-
pressed-pubescent; axis 5—15 X 1—2 mm long, 4-
angled in (lower, terete in fruit; (lowers 6 to 10;
bracts 0.5—1 mm long, persistent through anthesis,
cymbiform, spreading, the apex acute; buds 3—4
mm long, obovoid, costate. Flowers pedicellate, the
pedicels 4—8 mm long, 4-angled, the pedicels,
bracteoles, hypanthium, and calyx uniformly ap¬
pressed-pubescent; bracteoles persistent, free or
more commonly partially or wholly fused, forming
an involucre 1 — 1.5 mm diam., dilated at anthesis,
the inner surface glabrate, the margins scarious, the
apex acute; hypanthium 2.5—3 mm long, campan-
ulate, costate; calyx lobes in strongly unequal pairs,
widely ovate, appressed-pubescent on both surfac¬

es, the margins eiliate, the apex rounded, the small¬
er pair 0.7—1 X 2—3 mm, the larger pair 2—3 X
2.5—3 mm; petals 2.5— 3.5 X 1.5— 2.5 mm, elliptic
or ovate, convex, the margins eiliate, the apex
rounded; disk 2.5—3 mm diam., glabrous or with
scattered, erect hairs; stamens 70 to 100, 3—6 mm
long; style 4—6 mm long, glabrous. Fruits 25—35 X
15—20 mm, ellipsoid or obovoid, the base common¬
ly attenuate, the apex obtuse; pericarp thin-walled,
ribbed, glandular, uniformly but thinly pubescent;
disk expanded, 5—6 mm diam.; calyx typically ab¬
sent; mature color black.

Habitat , distribution, and phenology. Montane
forests along the Continental Divide in central and
northern Costa Rica at elevations of 900— 1600 m.
Collected in flower June— November; collected in
fruit July— December.

A large* tree of montane forests. Eugenia tilarana
is the second of the three species newly described
here that have been commonly identified as either
E. octopleura or E. guatemalensis. It differs from
the latter in having larger (25—35 vs. 5—12 mm
long), ribbed, ellipsoid or obovoid fruits. The fruits
also distinguish it from E. monteverdensis, which
has unribbed, tan fruits.

Paratypes. COSTA RICA. Vertiente Pactfico, Reserva
Monteverde, Cordillera de Tilardn [covers parts of Alajue-
la, Puntarenas, and Guanacaste], V. J. Dryer 1057 (CR,
F). Alajuela: San Ramon, RB Monteverde, Cordillera de
Tilaran, Valle del Rio Penas Blancas, Quebrada Fl Valle,
E. Hello A: E. Cruz 5204 (CR, F, INR, MO); Monteverde
Cloud Forest Reserve, Penas Blancas R. valley on Atlantic
slope, W. A. Haber & E. Hello 5817 (MO). Guanacaste:
R. Chiquito de Arenal, Tilaran, Altos de San Bosco, It)
km NW Monteverde, Atlantic slope. W. A. Haber et al.
5563 (MO). Puntarenas: San Luis, Monteverde, Fincade
Chepe Rojas, al Oeste del pueblo, E. Hello et al. 61 (CR,
MO), E. Hello et al. I!7 (CR, F, MO); iri & around the
Monteverde Nature reserve, mostly on the Pacific water¬
shed, IF C. Hurger A" R. Haber 9653 (CAS, CR, F); Mon¬
teverde, upper community nr. Casona, Tract X, Pacific
slope, W. Haber A W. Zuchowski 10814 (CR, F, MO);
Mata’s Pasture, Monteverde, G. S. Hartshorn 1781 (F. MO,
NY).

Eugenia verruculata Barrie, sp. nov. TYPE: Cos¬
ta Rica. Puntarenas: Monteverde, Reserva
Biologica, 4 Apr. 1990 (fl), IF Haber 9833 (ho-
lotype, INB; isotypes, F. MO). Figure 34.

Eugeniae eostaricensi acmulans, sed floribus majoribus,
fructibus verruculatis, carnosioribus differt.

Trees to 7 m or more; young branches, emergent
leaves, and developing inflorescences coarsely pu¬
bescent with simple, straight or contorted, coppery
hairs, 0.2— 0.5 mm; branchlets terete; bark Ian or
brown, quickly graying, glabrate, slightly flaky.

46

Novon

Figures 33. 34. — 33 (left). Eugenia tilarana Barrie ( Poveda 7/7/; holotvpe. CR). — 34 (right). Eugenia verruculata Barrie (Haber 9833 ; holotvpe. INB).

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

47

Leaves elliptic, the blades 2—5 X 1-2.5 cm, twice
as long as wide, coriaceous, drying dark green
above, paler below; midvein weakly impressed or
sulcate above, convex below; lateral veins 7 to 9
per side, often partial ly obscure above; marginal
veins similar to the laterals and arched between
them, ca. 1 mm from the margins; both surfaces
glabrate; glands typically obscure on the upper sur¬
face, small, pellucid on the lower surface; base cu-
neate; margins somewhat cartilaginous, decurrent
into the inner edge of the petiole; apex acute to
acuminate; petioles 4 mm long, channeled dorsal ly,
becoming corky ami fissured with age, glabrate. In¬
florescences axillary or at leafless nodes, racemose,
solitary; axis 1—2 mm, glabrate or with a few, scat¬
tered hairs; flowers (1)2 to 8; bracts 0.5 mm long,
persistent, cymbiform, glabrate, the margins eiliate.
the apex acute; buds 2.5—3 mm long, obconic.
Flowers pedicellate, the pedicels 1—4 mm long, gla-
brale, glandular, in bud and flower; bracteoles ca.
1 X 1 mm, persistent, ovate, glabrate, the base free,
commonly clasping the stipe, the margins eiliate,
the apex rounded or obtuse; hypanthium 1.5 mm
long, campanulate, the base commonly elongated
into a stipe, ca. 1 mm, glabrate; calyx lobes in sub¬
equal pairs, 1-1.5 X 1.5-2 mm, widely elliptic, the
margins eiliate, the apex rounded; petals 3.5 X 3
mm, elliptic, the margins eiliate, the apex rounded;
disk 2-2.5 mm diam., glabrous; stamens 70 to 80,
3-5 mm long; style 5—6 mm long, glabrous. Fruits
10—15 X 10—15 mm, globose, with or without a
short stipe; pericarp somewhat fleshy, prominently
glandular-verruculate; calyx persistent but incon¬
spicuous; mature color purple-black.

Habitat, distribution, and phenology . Known
only from the montane wet forests of central Costa
Rica at elevations of 1500 m. Collected in flower
April-May; collected in fruit May-January.

Eugenia verruculata is similar to E. costaricensis,
hut may he distinguished by several floral and fruit
characters. Although both species have sessile or
short-racemose inflorescences, the pedicels of E.
verruculata are generally shorter, 1—4 mm versus
2—8 nun. The bracteoles are ovate with rounded or
obtuse apices and the hypanthium is campanulate.
In E. costaricensis the bracteoles are commonly
keeled, with acuminate apices, and the hypanthium
is urceolate. The flints of E. verruculata are % the
size of those of E. costaricensis, and have a thicker,
fleshier pericarp with a warty texture.

Etymology. The epithet refers to the glandular
verruculate fruits of this species.

Paratypes. COSTA RICA. Reserva Monteverde, Cor¬
dillera de Tilaran [covers parts of Alajuela, Puntarenas,

and Guanacaste], V. ./. Dryer 1353 (K, MO). Puntarenas:
Guacirnal, Altos de Rio Veracruz, Finca Brenes, E. Hello
1798 (CR, F, MO); San Luis, Monteverde, Caniino a Ve¬
racruz, A. Ferndndez 430 (F INB. MO); Monteverde, hull-
pen on John Campbell farm, W. Haber <£■ ,/. Atwood 96 13
(CR, K); Monteverde, upper community W. Haber 292
(MO); Monteverde, upper community near reserve. Pacific
slope wet forest, bullpen W. Haber 10610 (CR, F, MO,
SEL); Monteverde, upper community, on farms of Wolf
Guindon & John Campbell, Pacific slope, W. Haber el al.
10988 (INB).

Eugenia zuchowskiae Barrie, sp. nov. TYPE:
Costa Rica. Puntarenas: Monteverde, Pacific
slope in community, 24 July 1990 (fr), 1L Ha¬
ber & W. Zuchowski 10036 (holotype, CR; iso¬
types, CR, F, MO). Figure 35.

Eugeniae tilaranae aemulans, sed foliis cum petiolis
longioribus, apicibus caudalo-acuminatis vel falcatis,
fructibus minoribus differt.

Trees to 20 m or more; young growth coppery
appressed-pubescent, the hairs dibrachiate;
branchlets terete; bark dark reddish brown, smooth,
glabrate. Leaves elliptic to obovate, the blades 4.5—
9 X 1 .5-5 cm, 2—3 times as long as wide, subco-
riaceous, drying dark, glossy green above, pale
green or brown below; midvein sulcate above,
raised below; lateral veins 5 to 7 per side, straight
and ascending, typically obscure above on fresh
material but apparent on dried leaves; marginal
veins similar to the laterals and arched between
them, 1-2 mm from the margins; upper surface gla¬
brate, coppery hairs persistent on the lower surface,
more densely near the base; glands typically ob¬
scure above, minute and numerous below; base eu-
neate; margins cartilaginous; apex caudate-acumi¬
nate, up to Vs of the total blade length, twisted or
falcate; petioles 10—20 mm long, sulcate dorsally,
glabrate or with a few, persistent hairs. Inflores¬
cences axillary, racemose, solitary or rarely 2 su¬
perposed, the axis, bracts, and buds uniformly op¬
pressed coppery-pubescent; axis 4—20 mm long,
4-angled in flower, terete in fruit; flowers 4 to 10;
bracts 0.5—1 mm, persistent through anthesis, cym¬
biform. spreading, the apex acute; buds obovoid,
3—4 nun long, costate. Flowers pedicellate, the ped¬
icels 3—5 mm long, 4-angled; bracteoles 0.5—1 X
0.5—1 mm, persistent, dilated at anthesis, triangu¬
lar, coppery appressed-pubescent, the base free or
weakly connate, the margins scarious, the apex
acute; hypanthium 1.5—2 mm long, obconic, cos¬
tate; calyx lobes in unequal pairs, widely ovate,
convex, appressed-pubescent on both surfaces, the
margin entire, the apex rounded, the smaller pair
1-1.5 X 1-2 mm, the larger pair 2—3.5 X 2.5— 3.5

48

Novon

NTARBNAS: Cant6n de Puntarenas
nteverde, Pacific *lop« >n noaaunity.
10* 18 ' N 84’48’W 1400-1500 ■

ee, 18 ■, forest edge. Fruit orange tc
d to purple black aatura , to 1.5 x

HOLOTY PI", of

Eugenia zuchowskiae Barne

IN8TITVT0 NACIONAL DH BIODI VKRS1DAD
MYRTACBAB lilliaa la Ur IMH

Eugenia octopleura Krug k Urb. ex Urb.

Mtilifioa: P. I. Uickti-Yndii. If/llJJ

Hlliu Haber 10036 Jl Jilf M*

Willow Zuchowski _

ItKMHARIO NACIONAL DR COSTA H1CA (CR)

nun mo ucioui di iionra»i»i»//iiiwii wratcu cum l»)

figure .'55. Eugenia zuchowskiae Barrie (llaber & Zuchowski 10036 ; holotype, CK).

mm; petals 4 X 3 mm long, elliptic, the margins
ciliate, the apex rounded; disk 2.5—3 mm diam.,
glabrous; stamens 75 to 100, 5-5.5 mm long; style
4—5.5 mm long, glabrous. Fruits 25—30 X 10—15
mm, ellipsoid; pericarp thin-walled, partially or
wholly 8-ribbed, glandular, glabrate or with a lew
hairs; calyx persistent but reduced; mature color
purple-black.

Habitat , distribution , and phenology. Montane
forests along the Continental Divide in central Cos-
la Kica at elevations of 1400-1600 m. Collected in
flower May— October; collected in Iruit March— Au¬
gust.

The third of the Monterverdean species often
identified as cither Eugenia octopleura or E. gua-
temalensis, Eugenia zuchowskiae is most easily dis¬
tinguished from these two species, as well as from
E. monteverdensis and E. tilarana , by the glossy
leaves with caudate or falcate, somewhat twisted,
apices. The lateral veins of fresh leaves are typi¬
cally obscure on the upper surface, though they
may be visible in dried specimens. The fruits are
ellipsoidal and usually 8-ribbed.

Etymology. The epithet honors Willow Zu¬
chowski, the co-collector of the type and a noted
specialist on the flora of the Monteverde region.

Volume 15, Number 1
2005

Barrie

Eugenia from Mesoamerica

49

Paratypes. COSTA RICA. Puntarenas: Reserva
Biologica Monteverde, Rio Guacimal, Comunidad-Reser-
va. Vertiente Pacifica, E. Hello 2086 (CR. F, MO); Reserva
Monteverde, Cordillera de Tilaran, en portrero ca. de Re-
serva, V. J. Dryer 90S (CR, F, MO); Reserva Monteverde,
Cordillera de Tilaran, on faja de bosque perturbado ca. dc
Reserva, V. J. Dryer 1443 (CR); Reserva Monteverde, Cor¬
dillera de Tilaran. en portrero ca. de Reserva, V. J. Dryer
1580 (CR. F, MO); Monteverde, upper community, IT A.
Haber 554 (MO); Monteverde, upper community. Pacific
slope, T. Guindon farm and bullpen, IT Haber et al. 996 0
(1NR); Monteverde, upper community nr. Casona, Tract X.
IT Haber & IT Zuchowski 10802 (CR. F, MO); in Mata’s
pasture, Monteverde, S. Hartshorn 1776 (CR[3|, F, MO.
NY), G. S. Hartshorn 1781 (CR); Tos Ayotes, near Tilaran,
P.C. Standley & ./. Valeria 45461 (F).

Acknowledgments. The research underlying
this paper was conducted while in residence in the
Botany Department of The Field Museum, Chicago,
Illinois. 1 thank the department for its generous,
continuing support. The Institute Nacional de Bio-
diversidad supplied financial and logistical support
for research in Costa Rica. Thanks to all the INBio
staff, especially Nelson Zamora, Francisco Morales,
and Barry Haminel. Bill Haber generously shared

his knowledge of the Myrtaceae of the Monteverde
region. Thanks to Lucia Kawasaki, Bruce Holst,
and Les Landrum for many enlightening and con-
structive discussions of Neotropical Myrtaceae.
And many thanks to the curators of the herbaria
who loaned specimens for this study, including A,
BM, BR. CAS, CR, F, GH. HAL, INB. K. MEXU,
MICH. MO, NY, P. SEL. TEX-I.L. UC, and US.

Literature Cited

Barrie, F. R. 2004. Synopsis of Plinia (Myrtaceae) in Me¬
soamerica. Novon 14: 380—400.

Holst, B. K. 1999. A new species of Calyptranthes (Myr¬
taceae) from Nicaragua. Novon 9: 517—519.

- & M. L. Kawasaki. 2004. New species of Calyp¬
tranthes and Myrcia (Myrtaceae) from Central America.
Selbyana 25: 87—100.

Landrum, L. R. 1991. Chamguava: A new genus of Myr¬
taceae (Mvrtinae) from Mesoamerica. Svst. Bot. 16: 21 —
29.

- . 1992. Mosiera (Myrtaceae) in Mexico and Me¬
soamerica. Novon 2: 26—29.

Molina R., A. 1975. Fnumeracion de las plantas de Hon¬
duras. Ceiba 19: 1 — 118.

Two New Cauliflorous Species of Daphnopsis (Thymelaeaceae) from

French Guiana and Surinam

Kerry Barringer

Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 11225-1099,

U.S.A. kerryharringer@bbg.org

John F. Pruski

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

john.pruski@mobot.org

Abstract. Two new cauliflorous species of
Daphnopsis (Thymelaeaceae) from French Guiana
and Surinam are described, illustrated, and their
relationships discussed. A key to the species of
Thymelaeaceae in French Guiana and Surinam is
provided. Daphnopsis granitica Pruski & Barringer
and D. granvillei Barringer grow on the granitic
inselbergs of northern to southwestern French (ini-
ana and adjacent Surinam. They are each charac¬
terized by brachyblasts from older stems, the ab¬
sence of petaloid appendages, and the presence of
orange fruits. Daphnopsis granitica is distinctive
because of its narrowly oblanceolate leaves, while
D. granvillei has broader, oblanceolate to obovate
leaves.

Resumkn. Se describe y ilnstra dos nuevas es-
peeies eaulifloras de Daphnopsis (Thymelaeaceae)
de la Guyana Francesa y Surinam y se discute sus
relaciones. Se presenta una clave para las especies
de Thymelaeaceae de la Guyana Francesa y Suri¬
nam. Daphnopsis granitica Pruski & Barringer y I).
granvillei Barringer erecen sobre lajas gramticas
del norte al suroeste de la Guyana Francesa y la
region adyacente de Surinam. Ambas especies se
caracterizan por tener braquiblastos de los tallos
viejos, por la auseneia de apendices petaloides y
por la presencia de frutos anaranjados. Daphnopsis
granitica se diferencia por tener las hojas angos-
tamente oblanceoladas, mientras tpie D. granvillei
tiene las hojas mas anchas y oblanceoladas a obova-
das.

Key words: brachyblast. cauliflory, Daphnopsis ,
French Guiana, Schoenohiblns, Surinam. Thymelae¬
aceae. Thymelaeoideae.

Daphnopsis Martins (Thymelaeaceae: Thyme¬
laeoideae) is a genus of 50 to 65 species native to
the Neotropics ( Barringer & Nevling, 2004; Herber,
2003 [2002]). Species are shrubs or small trees

with small, tubular or campanulate, essentially
apetalous, unisexual flowers with eight stamens,
short calyx lobes, and a pseudomonomerous ovary.
The genus was most recently revised by Nevling
(1959, 1963).

The following species were discovered during ex¬
ploration of isolated, granitic mountains of French
Guiana by Jean-Jacques de Granville and his as¬
sociates. In February 2003, a team from Utrecht
University collected Daphnopsis granvillei in the
Nassau Mountains of Surinam, 150 km west of
Montagues de la Trinite, French Guiana. These two
new1 species are the first species of Daphnopsis de¬
scribed from French Guiana and Surinam. They art1
also the sole species of Daphnopsis with orange
fruits, a feature previously considered diagnostic
for the genus Schoenobiblus Martins.

Most specimens of each species described herein
have been distributed to specialists using the fam¬
ily identifications of either Myrsinaceae or Ery-
throxylaceae. Daphnopsis granitica was known
solely as a herbarium name of the second author,
then at US, where it was revived by the first author.
Daphnopsis granvillei Barringer was treated as
Daphnopsis sp. 1 by Kawasaki and Mori (2002).
I'he only other Thymelaeaceae reported from
French Guiana or Surinam, Schoenobiblus daph-
noides Martins, is distinguished by its rotate, deep¬
ly lobed calyces, and its four stamens. The key in
Kawasaki and Mori (2002) is modified below.

Key to the Species ok Thymelaeaceae in French
Guiana and Surinam

la. Flowers anil fruits in terminal cymes; calyx tube

shorter than the lobes; stamens 4 .

. Schoenobiblus daphnoides Martius

1 b. Flowers and fruits in brachyblasts on older stems;
calyx tube longer than the lobes: stamens 8.

2a. Leaves narrowly lanceolate, 1.7—3 cm wide;

petioles to 2 mm long .

. Daphnopsis granitica Pruski & Barringer

Novon 15; 50-54. 2005.

Volume 15, Number 1
2005

Barringer & Pruski

New Species of Daphnopsis

51

2b. Leaves oblaneeolate to obovate, 5—9 cm

wide; petioles 4—7 mm long .

. Daphnopsis granvillei Barringer

We place Daphnopsis granitica and D. granvillei
in subgenus Neivira (Grisebach) Nevling (Nevling,
1959), where they resemble the cauliflorous, Ve¬
nezuelan I). steyermarkii Nevling. Daphnopsis subg.
Neivira is characterized by monopodial stems with
axillary inflorescences, male (lowers with petaloid
appendages forming a 1owt ring at the mouth of the
calyx tube, and deeply lobed nectary discs. The
species from the Guayana Highland differ from the
majority of the species in the subgenus, which grow
in the mountains of southeastern Brazil, by the ab¬
sence of petaloid appendages, and probably by hav¬
ing orange to yellow fruits. Further study will likelv
support recognition of a new subgenus for the cau¬
liflorous species from the Guayana Highland.

Daphnopsis granitica Pruski & Barringer, sp.
nov. TYPK: French Guiana. Montagnes de la
Trinite, sommet nord alt. 400 m environ, foret
dense sur crete avec cahos granitiques en sous
bois, 12 Jan. 1984. de Granville, C. Berg,
V/. Jansen-Jacobs & ,/. van Setten 5890 (ho-
lotype, CAY 009959; isotypes, NY, P not seen,
LJ not seen, US 3205040). Figure 1.

Daphnopsi steyermarkii affinis, sed folds brevipetiolatis,
lamina anguste oblanceolata subcordata vel rotundata at-
tenuata, fructibus aurantiacis diversa.

Shrubs to small trees to 4 m tall; stems mono¬
podial, subterete, 2.5—5 cm diam., glabrous, bark
brown, the terminal bud narrowly deltoid, 2—3 mm
long. Leaves simple, alternate, clustered at the stem
apices, estipulale, subsessile; petiole to 2 mm long,
thick; blade narrowly oblaneeolate. 1 1.5—25 X 1.7-
3 cm, stiffly chartaceous or subcoriaceous, gla¬
brous, venation pinnate, reticulodromous, basally
rounded or subcordate, sometimes slightly clasping,
apically attenuate, margins entire. Staminate flow¬
ers not seen. Pistillate inflorescences borne in axil¬
lary bracteate (dusters (braehyblasts) on older, leaf¬
less stems, ea. 20-30 cm below the stem apices,
the primary peduncle, rachis, and secondary pe¬
duncles basically absent, pedicels to 0.5 mm long,
lew- to several-flowered, compact. Pistillate flowers
white; calyx urceolate, the tube 2-2.5 mm long,
longitudinally pleated, glabrous to minutely pubes¬
cent, 4-lobed, the lobes ligulate, 0.8—1 X 0.5— 0.7
mm, obtuse, pubescent toward the apex on the in¬
ner surface, longitudinally ridged on the inner sur¬
face, strongly recurved; petaloid appendages ab¬
sent; staminodia absent; pistil fusiform, sessile, ea.

3 mm long, partially surrounded by a deeply-lobed.

eoroniform nectary disc; ovary superior, style ter¬
minal. stigma exserted, capitate, papillate, white.
Fruit a drupe-like berry, indehiscent, 1 -seeded, ob-
ovoid to subglobose, 8-10 mm diam. when mature,
smooth, glabrous, green when young, orange at ma¬
turity, the style base persistent, the calyx tearing
irregularly, persistent at the base of the fruit.

Distribution and habitat. This species is found
at the base of the granite inselbergs Montagues de
la Trinite and Montagnes des Nouragues, where it
has been collected between 1 10 and 400 m ele¬
vation.

Phenology. Plants flower in January and bear
mature fruits from late February through early
April.

Daphnopsis granitica appears to be related to D.
steyermarkii Nevling. D. guaiquimimae Steyermark,
and D. longipedunculata Gilg ex Domke, each of
which also has monopodial branching and minute
flowers borne in braehyblasts (Nevling, 1967; Stey-
ermark. 1987). Daphnopsis steyermarkii, I). guai¬
quimimae, and D. longipedunculata, which occur
on sandstone substrates in the Guayana Highland,
differ from D. granitica by having broader leaves
and white or green (not orange) fruits. Daphnopsis
granitica is also similar to D. granvillei Barringer,
described below, in having braehyblasts and orange
fruits but differs from it by narrowly oblaneeolate
(vs. oblaneeolate to oblong) leaves, these slightly
(vs. not) clasping.

Specimens very similar to I), granitica have been
seen by Lucia Rossi (pers. comm.), of the Instituto
de Botanica, Sao Paulo. The specimens were col¬
lected in the states of Roraima and Amazonas, Bra¬
zil, and have slightly wider and more oblaneeolate
leaves than D. granitica and yellow' fruits, these
occasionally present near the stem apex. It is pos¬
sible that these represent an additional new spe¬
cies. although this cannot be confirmed without
flowering material.

Paratypes. FRENCH GUIANA. Montagnes de la Trin¬
ite. Bassin de la Mana, G. Cremers 12824 (CAY. MO. NY.
US); Res. Nat. de la Trinite, Bassin de la Trinite, Fourre
de savane-roche, R. DutrSve 50 (CAY); Inselberg grana-
tique Y Fextr&mite NordOuest de Monts de la Trinite, J.-
J. de Granville 4 778 (CAY); Montagnes de la Trinite, Som¬
met Nord, J.-J. de Granville, C. Berg, M. Jansen-Jacobs &
,/. van Setten 5808 (CAY, NY); loret Basse humide en lis-
iere de savane roche, J.-J. de Granville, C. Berg, XI. Jan¬
sen-Jacobs & J. van Setten 5923 (CAY, NY); Montagnes
de la Trinite, Bassin de la Mana, foret Basse sur pente
d’inselberg, pied du versant Sud 1’inselBerg oriental, J.-J.
de Granville 13276 (CAY. LIS); Montagnes de la Trinity,
zone sud, Bassin de la Mana, J.-J. de Granville & F. Gro-
zier 13560 (CAY, NY); Montague des Nouragues, Bassin
de TArataye, D. I Mr pin 504 (GAY ); Montagnes des No¬
uragues, Bassin de TApprouague— Arataye, D. Larpin 849

52

No von

Figure I. Daphnopsis granitica Pruski & Barringer. — A. Habit. — B. I .cal. — C. Brachyblast. Drawn from the holotype
J.-J. de Granville el al. 5890 (CAY).

(CAY, NY, US); I nselberg Trinity, Savane- Roche, J. Villiers
& C. Sarthou 6284 (CAY ).

Daphnopsis granvillei Barringer, sp. nov. IV PIC
French Guiana. Saul, S. <!u Mont Galboa, 13
Dec. 1976, J.-J. de Granville 11.5402 (holo¬
type, CAY 007764). Figure 2.

Daphnopsi steyermarkii affinis, sed foliis magnis oblan-

ceolatis vel obovatis aouminatis, fructibus rubris vel au-
rantiaeis diversa.

Sparsely branched, shrub to 2 m tall; sterns
monopodial, terete, to 2 cm diarn., glabrous, the
young stems red-brown, leaves simple, alternate,
clustered at the stem apices, estipulate, short-pet-
iolate; petiole 4—7 mm long, thick, slightly winged
on the abaxial side; blade oblanceolate to obovate.

Volume 15, Number 1
2005

Barringer & Pruski

New Species of Daphnopsis

53

Figure 2. Daphnopsis granvillei Barringer. — A. Halul. — I?. Leaf. — C. Brachyblast. Drawn from the holotype,
de Granville B.5402 (CAY).

15—30 X 5—9 cm, subcoriaceous, glabrous, vena¬
tion pinnate, reticulate, obscure, basally cuneate,
but abruptly rounded at the junction with the pet¬
iole, apically long-acuminate, margins entire. Sta-
minate inflorescences borne from axillary, bracteate
brachyblasts on older, leafless stettts, tbe primary
peduncle to 1(2) mm long, rachis absent, secondary
peduncles 1—1.5 mm long, pedicels 1.2— 1.5 mm
long, few-flowered, compact. Staminate flowers
while; calyx tubular-campanulate, the tube 2.1— 2.5
mm long, 1.5—2 mm wide at the mouth, 4-angled,
especially in bud, the lobes triangulate to ovate,
0.5— 0.7 mm long, 1 mm wide at the base, spread¬

ing, cymbiform in bud, apically acute to obtuse,
acuminate with a small papillate knob; stamens 8,
in 2 whorls, the antesepalous whorl borne on the
surface of the calyx lobes near the apex, the fila¬
ments 0.5— 0.7 mm long, the alternisepalous whorl
borne at the mouth of the calyx tube, the filaments
0.3— 0.5 mm long; disc free, deeply lobed, the lobes
acute, to ca. 1 mm long; pistillode glabrous,
stalked, 1.2— 1.7 mm long. Pistillate flowers not
seen. Fruit a drupe-like berry, ovoid, to 10 mm
long, to 8 mm wide, reddish or orange, glabrous,
apiculate, the stvle often persistent.

54

Novon

Distribution and habitat. This species grows on
hilltop forests from 500 to 000 in elevation.

Phenology. Plants bloom in December and Jan¬
uary and bear mature fruit in March.

This species is most similar to Daphnopsis gran-
itica, described above. Like that species, the flow¬
ers of I). granvillei are borne in compact, axillary
inflorescences on older, leafless stems and both
have orange fruits, die only species where this is
known. Daphnopsis granvillei grows at higher ele¬
vations and in more localities than D. granitica, but
both arc found on the Montagnes de la Trinite. The
narrowly lanceolate, slightly clasping leaves of I).
granitica are distinctive and make it easy to distin¬
guish from I). granvillei.

Paratypes. FKKNCH GUIANA. Montague Cacao, SF
tie Cayenne, C. Feuillet 549 (CAY); Region de Paul Is-
nanl. Massif du Decou Decou, C. Feuillet 689 (CAY); Ren¬
te NK des Monts Galboa, it It) km SW de Saiil, J.-J. de
Granville 2875 (CAY); Montagnes de la Trinite. sommet
NK, in forest on slopes, J.-J. de Granville et al. 6877
(CAY). SURINAM. Marowijnc Distr., Nassau Mis.. Plateau
C, M. J. Jansen- Jacobs. J. Behari-Ramdas, A. Grant, G.
Ramharrakh, II. ter Steege, 0. Bdnki, F. van Troone A' R.
Ho Tsoi 6877 (MO. U not seen), 6484 (MO. U not seen).

Acknowledgments. We thank Jean-Jacques dc
Granville (CAY) as well as the curators of l\Y, U,
and US for allowing us to study their specimens.

We especially thank Dr. de Granville, Lucia Rossi
(SP), and Scott Mori (NY) for checking for material
of Daphnopsis among their duplicates and recent
collections. We thank Zachary Rogers and Lucia
Rossi for reviewing the manuscript, and Peter Ste¬
vens for helpful discussion. The illustrations were
drawn by Paul Harwood (BKL).

I .iterature Cited

Rarringer, k. & I.. I. Nevling Jr. 2004. Thymelaeaceae.
Pp. 372-374 in N. Smith, S. Mori. A. Henderson, I).
Stevenson & S. Heald (editors). Flowering Plants of the
Neotropics. Princeton Univ. Press, Princeton.

Herber, B. E. 2003 [2002 1. Thymelaeaceae. Pp. 373-396
in k. Kubitzki (editor). The Families and Genera of
Vascular Plants, Vol. 5. Springer, Berlin.

Kawasaki, M. L. & S. A. Mori. 2002. Thymelaeaceae. Pp.
71 1-712 in S. A. Mori, G. Cromers, C. Gracie, J.-J. de
Granville, S. Heald, M. Hoff & J. Mitchell (editors).
Guide to the Vascular Plants of Central French Guiana,
Part 2. Dicotyledons. Mem. New York Rot. Gard. 76(2):
viii 4- 1—776.

Nevling, L. 1., Jr. 1959. A revision of the genus Daphn¬
opsis. Ann. Missouri Bot. Gard. 46: 257—358.

- . 1963. Notes on Daphnopsis. J. Arnold Arbor. 44:

402-110.

- . 1967. Thymelaeaceae. Pp. 452-455 in J. A.

Steyermark, B. Maguire <X collaborators. Botany of the
Chimanta Massif — II. Mem. New York Bot. Gard. 17:
440-464.

Steyermark, J. A. 1987. Flora of the Venezuelan Guayana
III. Ann. Missouri Bot. Gard. 74: 609—658.

Staurogyne rubescens (Acanthaceae), a New Species from

Southeastern Brazil

Denise Monte Braz and Reinaldo Monteiro

Departamento de Botanica, Instituto de Biociencias, Universidade Estadual Paulista-UNESP,
C.P. 199, 13506-900, Rio Claro, SP, Brazil, dmbraz@rc.unesp.hr; reimonte@rc.unesp.br

Abstract. A new species of Acanthaceae, Stau¬
rogyne rubescens, is described and illustrated. It is
endemic to the Brazilian moist forests on the east¬
ern range of mountains named “Sena do Mar" and
is thus far known only from the states of Rio de
Janeiro and Sao Paulo. Staurogyne rubescens can
be distinguished by its lax terminal racemes, with
bracts and bracteoles that are foliaceous, and the
flowers that have a pink calyx and red corolla.
Staurogyne itatiaiae (Wawra) Leonard also has red
flowers and occurs in the highlands of the Atlantic
rain forest. The distinguishing characters for these
two species are presented in tabular format.

RKSUMO. A nova especie Staurogyne rubescens
(Acanthaceae), endemica da Serra do Mar nos Es-
tados do Rio de Janeiro e de Sao Paulo, e descrita
e ilustrada. Staurogyne rubescens e reconhecida
pelo racemo terminal laxo, com hracteas e bracteo-
las foliaceas, e pelas flores com calice rosado e
corola vermel ha. Staurogyne itatiaiae (Wawra) Le¬
onard tambem apresenta flores vermelhas e ocorre
em altitudes elevadas na Horesta Atlantica. As ca-
racterfsticas distintivas entre as duas especies estao
apresentadas em tabela.

key words: Acanthaceae, Brazil, Nelsonioi-
deae. Staurogyne.

Staurogyne Wallich comprises about BO herba¬
ceous or shrubby species (Wasshausen, 1995).
Plants belonging to this genus may be recognized
by flowers with the calyx formed by five unequal
lacinia, the posterior larger than the others, the an¬
terior pair of intermediate size, and the lateral pair
smaller than the others; four stamens, usually di-
dynamous, and with a staminode between the two
posterior stamens; and stigma with usually one of
the two lobes deeply concave or divided into two
segments. Staurogyne is a pantropical genus and in
the Americas, where it comprises ca. 28 species, it
is widely distributed especially in South America;
only one species reaches the subtropical areas of
North America. In Brazil there are ca. 27 taxa, ac¬

cording to an undergoing taxonomic revision of
Staurogyne (Braz & Monteiro, in prep.).

The subfamily Nelsonioideae Pfeiff of Acautha-
eeae, of which Staurogyne is the richest genus, can
be recognized by the imbricate descending cochle¬
ar aestivation, and absence of hooklike retinacula
in the capsules and cystoliths in the leaves (Scot¬
land & Vollesen, 2000). Recent phylogenetic stud¬
ies based on morphological and molecular data
support the basal placement of Nelsonioideae as a
sister group of the other Acanthaceae (Hedren et
ah, 1995; Scotland et al., 1995; McDade & Moody,
1999; McDade et al., 2000; Scotland & Vollesen,
2000; Manktelow et al., 2001; Schwarzbach &
McDade, 2002), showing no reason to treat them
separately. In Brazil, Elytraria Michaux and Nel-
sonia R. Brown are the Nelsonioideae representa¬
tives that differ from Staurogyne in having 2-sta-
minate flowers.

Fhe Neotropical species of Staurogyne have in-
fundibular flowers that are white or lilac-colored,
sometimes purple-red maculate at the limb and
throat, or narrowly tubular corollas that are red or
yellow'. In Brazil. Staurogyne species occur in hu¬
mid and shaded habitats, especially close to water
courses, in the gallery forests, in the cerrado bio¬
geographic domain, but mainly in the understory of
natural forest vegetation, in the Amazon and Atlan¬
tic rain forest domains.

Staurogyne rubescens D. M. Braz & R. Monteiro,
sp. nov. TYPE: Brazil. Rio de Janeiro: Mun.
Parati, subida para o Cuscuzeiro, 500 m.s.m.,
APA-Cairugu, 28 Mar. 1995 (fl), M. G. Bovini,
/.. C. Giordano , ./. Miguel & C. Silva 724
(holotype, RB). Figure 1.

Ah omnibus speciebus generis inflorescentia racemo
terminali, laxo, bracteis et bracteolis viridibus, floribus
calyce roseo et corolla tubulosa rubescenti differt.

Herbs, 50—90 cm tall, stem cylindrical, erect or
decumbent, scarcely branched, with scattered sim¬
ple trichomes. Leaves elliptic, 3-10.3 X 1. 1-3.2
cm, apex acute to somewhat acuminate, base acute
to attenuate, upper surface glabrous or with scat-

Novon 15: 55-58. 2005.

56

Novon

Figure I. Staurogyne rubescens I). M. Braz & R. Monteiro. — a. Flowering stem. — 1>. Open calyx; external view with
the posterior lacinia in tlie middle and the two pairs of lateral and anterior lacinias. — c. Corolla. — d. Open corolla,
stamen, and staminode. — e. Anther. — 1. Staminode. — g. Stigma with the deeply concave posterior lobe iti first plane,
and the anterior lobe at the back. — h. Bract, bracteoles, calyx, and immature fruit. Drawn from Braz & Bessel 77
(HRCB).

Volume 15, Number 1
2005

Braz & Monteiro 57

Staurogyne rubescens from Southeastern Brazil

Table 1. Comparative features of Staurogy

ne rubescens and S. itatiaiae.

Characters

S. rubescens

.S', itatiaiae

Pairs of secondary veins ol the leaf blade

8 to 10

12 to 16

Inflorescence

simple, terminal, lax

compound, terminal and axillary.

Bracts and bracteoles

green, pinnate venation

congested

red to purplish, acrodromous venation

Pedicel length (nun)

3-9

0.8-2

Corolla length (mm)

29-36

16-20

Corolla lobe shape

oblong

orbicular

length (mm)

3-5

1.5-2

Stamen length (mm)

posterior pair

1 7.5-26

5.5-8

anterior pair

18-27

7.5-9

Ovules (number) per locule

14 to 18

20 to 22

Style length (mm)

31-37

12-16

Stigma posterior lobe

deeply concave

truncate

tered simple triehomes al the base of the main vein,
lower surface with scattered simple triehomes, es¬
pecially at the veins; secondary veins 8 to 10 pairs;
petiole 0.6— 2.0 cm long. Inflorescence a raceme,
terminal, leafy-bracteate, lax, simple, rachis with
scattered glandular triehomes; peduncle 21-29 mm
long; bracts foliaceous, 10—12 X 3—3.5 mm, the
upper gradually smaller; bracteoles lanceolate, 4—
12 X 0.8— 2. 1 mm, green, with pinnate venation;
pedicel 3—9 mm long. Calyx pink, slightly purple-
red near the margins, with scattered glandular tri-
chomes, posterior lacinia elliptic to oblong-elliptic,
12—27 X 2—3.5 mm, apex acute, lateral pair su¬
bulate, 10—19 X 0.5— 1.5 mm, anterior pair linear-
lanceolate, sometimes slightly asymmetric, 12—20
X 1.3—2 mm, apex acute; corolla tubular, red to
light pink. 29—36 X 4—7.3 mm. basal tube well
defined, 2.5—6 mm long, throat 23.5—29 mm long,
lobes oblong, anterior slightly orbicular. 3—5 mm
long; stamens subexserted, inserted 3—4.5 mm
above the corolla base, posterior pair slightly short¬
er (17.5—26 mm long) than the anterior (18—27 mm
long), staminode straight. 2—2.8 mm long, apieally
truncate to antheriform. inserted slightly below the
stamens; style 31—37 mm long, posterior lobe of the
stigma deeply concave, with the sides slightly sa¬
lient, anterior lobe longer, oblong, apex obtuse. Im¬
mature capsule white, becoming brown when ma¬
ture, subterete, 14—16.5 X 5—4 mm, sparsely
pilose; seeds 14 to 18 per locule, subglobose, gla¬
brous.

Habitat , distribution, and phenology . Staurogy¬
ne rubescens is endemic to the Serra do Mar, oc¬
curring in the Atlantic rain forest, in the extreme
south of Rio de Janeiro State (Parati municipal dis¬
trict), and in the state of Sao Paulo (Ubatuba mu¬

nicipal district). Plants occur in protected areas of
the conservation units of A PA Cairuyii ( A PA. Area
de Protegao Ambiental ), the ecological station of
Joatinga, and the state park of Serra do Mar. Stau¬
rogyne rubescens is found in shaded and wet habi¬
tats, between 500 and 1080 m elevation. Plants
flower between March and April.

Staurogyne rubescens can be recognized by its
leafy-bracteate inflorescences, with bracts and
bracteoles always green, and by its flowers with a
pink calyx and red tubular corolla. W ith regard to
the red corolla, S. rubescens is similar to S. ita-
tiaiae, but it differs especially by the terminal lax
inflorescence, with green bracts and bracteoles, and
by the notably smaller habit, usually herbaceous;
in S. itatiaiae the racemes are terminal and axillary,
congested, the bracts and bracteoles are red. and
the habit is shrubby (Table 1). Staurogyne rubescens
is also similar to S. minarum (Nees) Kuntze in hav¬
ing a lax terminal raceme and green bracts and
bracteoles. These species may be distinguished by
habit (herbaceous in S. rubescens vs. frutieous in S.
minarum ), flower color (red vs. yellow), and also by
having allopatric and restricted distributions, with
S. minarum being only found in the state of Minas
Gerais.

The name of the new taxon refers to its red co¬
rolla. Staurogyne rubescens and 5. itatiaiae are the
only American species that have tubular red corol¬
las. Both species are almost entirely restricted to
the Atlantic rain forest, above 500 m elevation.
Staurogyne itatiaiae also reaches the “Serra da
Mantiqueira,” where it is widespread in the munic¬
ipal district of Itatiaia, in Rio de Janeiro State,
down to the “Serra da Boeaina,” through which it
ranges to Sao Paulo State, being the first record for

58

Novon

this state. Staurogyne rubescens occurs on coastal¬
facing slopes of the “Serra do Mar."

Raratypes. BRAZIL. Kio de Janeiro: Parati. Praia
Negra. Pico do Cairuyu. Mata de Altitude, 25 Mar. 1992
(II), barney A' /.. Silva , ./. (.. Silva .1123 (RB). Sao
Paulo: Ubatuha. proximo it divisa com Sao Luiz do Par-
aitinga, 5 Abr. 2002 (II), / ). M. lira z A K. Ressel 77
(HRCB); Ubatuba. proximo it divisa com Sao Luiz do Par-
aitinga, 5 Abr. 2002 (II), I). M. firaz A K. Ressel 78
(HRCB, MO).

Acknowledgments. The authors are grateful to
Tarciso l ilgueiras for Latin corrections; to Massimo
Iiovini and Ronaldo Marquete for information about
the locality of the type collection; to FAPKSP (Fun-
dayao de Amparo ft Pesquisa do Estado de Sao Pau¬
lo) for financial support; to the curators of the her¬
baria C, F, HR. MRM. R. RB, and RRR for loan of
specimens; to Jaime Roberto Somera for the illus¬
trations; to the professionals and/or friends who
helped on field trips. Maria M. M. Rraz. Hermann
Schimitt, Alexandra M. S. Pitolli, Rita C. Koch, and
especially to Kaila Ressel.

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lationships in the Aoanthaceae and related families as

suggesteil by cladistio analysis of r6cL nucleotide se-
quences. PI. Syst. Evol. 194: 93—109.

Leonard, E. C. 1937. Notes on the genus Staurogyne. J.

Wash. Acad. Sci. 27(9): 398-402.

Manktelow, M.. L. A. McDade, B. Oxelman, (7 A. Furness
& M. J. Balkwill. 2(K)1. The enigmatic tribe Whitfiel-
dieae (Acanthaceae): Delimitation and phylogenetic re¬
lationships based on molecular and morphological data.
Syst. But. 2b: 104-1 19.

McDade, L. A. A M. I.. Moody. 1 999. Phylogenetic rela¬
tionships among Acanthaceae: Evidence from noncod¬
ing trnh-trnV ehloroplast l)NA sequences. Amer. J. Bot.
86: 70-80.

- . S. E. Masta, M. L. Moody A E. Waters. 2000.

Phylogenetic relationships among Acanthaceae: Evi¬
dence from two genomes. Syst. Bot. 25: 106—121.

Nees, C. G. 1847. Acanthaceae. In: C. E P. von Martius,
A. G. Fielder A 1. Urban (editors). Flora Brasiliensis 9:
1-164.

Schwarzbach, A. E. A L. A. McDade. 2002. Phylogenetic
relationships of the mangrove family Avicenniaceae
based on ehloroplast and nuclear rihosomal DNA se¬
quences. Syst. Bot. 27: 84—98.

Scotland, R. W. A K. Vollesen. 2000. Classification of
Acanthaceae. Kew Bull. 55: 513—589.

- , J. A. Sweere, P. A. Beeves A B. (7 Olmstead.

1995. Higher-level systematics of Acanthaceae deter¬
mined by ehloroplast DNA sequences. Amer. J. Bot. 82:
266-275.

Wasshausen, I). (7 1995. Acanthaceae. Pp. 335—373 in J.
A. Steyermark, P. E. Berry, B. k. Holst A k. Yatskiev-
yeh (editors). Flora ol the Venezuelan Guayana. Vol. 2.
Missouri Botanical Garden, St. Ixvuis.

New Taxa in Lythraceae from Latin America

Tatiana B. Cavalcanti

Embrapa Genetic Resources an<l Biotechnology— Embrapa/Cenargen, Parque Estagao
Ecologica, Final W5 Norte, CP 02372, CEP 70.770-900, Brasilia, RE, Brazil.
taciana@cenargen.embrapa.br

Shirley A. Graham

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

shirley.graham@mobot.org

ABSTRACT. Two new species, two new varieties,
and two new combinations for Lythraceae from Bo¬
livia, Brazil, Cuba, and Mexico are proposed. The
new species are Cuphea aquilana from Mexico and
C. potamophila from Brazil. The new varieties are
C. pulchra var. corollata from Brazil and Dipluso-
don virgatus var. occidentalis from Brazil and Bo¬
livia. Cuphea sessilifolia subsp. bahiensis is elevat¬
ed to the rank of species. Cuphea grisebachiana,
from Cuba, is reduced to varietal status as C. par-
sonsia var. grisebachiana, and the first report of its
chromosome number is given. The range of Cuphea
xanthopetala, previously known only from Mato
Grosso, Brazil, is extended by discovery of the spe¬
cies in Amazonas, Brazil. Additional collections
also expand the range ol the recently described Di-
plusodon bolivianos in Santa Cruz, Bolivia, and
Mato Grosso, Brazil.

Resumo. Duas novas especies, duas novas va-
riedades e duas novas eombinayoes para I .ythra-
ceae da Bolivia, Brasil, Cuba e Mexico sao propos-
tas. As novas especies sao Cuphea aquilana, do
Mexico e C. potamophila, do Brasil. As novas va-
riedades sao C. pulchra var. corollata . do Brasil e
Diplusodon virgatus var. occidentalis , do Brasil e
Bolivia. Cuphea sessilifolia subsp. bahiensis e ele-
vada ao estatus de especie. Cuphea grisebachiana
da Cuba e reduzida ao estatus varietal como C.
parsonsia var. grisebachiana e o primeiro registro
sobre o seu numero cromossomico e fornecido. A
amplitude de distribuiyao de C. xanthopetala . prev-
iamente conhecida apenas para o Mato Grosso,
Brasil, e extendida pela descoberta da especie na
Amazonia, Brasil. Coleyoes adicionais do Mato
Grosso, Brasil, expandem tambem a distribuiyao
conhecida de Diplusodon bolivianos, reeentemente
descrita para Santa Cruz, Bolivia.

Key words: Bolivia, Brazil, chromosome num¬
ber, Cuba, Cuphea, Diplusodon, Mexico.

Collaborative revisionary and monographic stud¬
ies under way in the Lythraceae have led to the
discovery of several novelties in the most speciose
genera. Cuphea P. Browne and Diplusodon Pohl. We
describe two new species, two new varieties, and
make two new combinations for members of floras
from Bolivia, Brazil, Cuba, and Mexico. We also
report range extensions for two recently described
species of Cuphea and Diplusodon.

1. Cuphea aquilana S. A. Graham & T. B. Cav¬
alcanti, sp. nov. TYPE: Mexico. Michoacan:
Aquila, en Las Brisas del Mirador, 24 km al
S de F4 Ranehito, 20 Oct. 1985. ./. C. Soto N.
1 1 194 (holotype, MEXU; isotypes, MEXU,
MO). Figure I.

Herba annua delicatula, 30—35 cm alta. Kolia petiolis
5—27 mm longis insidentia, 20—50 mm longa, 10-35 mm
lata, ovala, cordata, supra sparse strigosa. Raeeini simpli-
ces; pedieelli 2—5 mm longi, filiformes. Flores I in quoque
nodo, 4—5 mm longi, sine calcare, dorsaliter remoti his-
pidi; lobus dorsalis quam eeterus amplior; petala 0; sta¬
mina inclusa, supra medium tubi inserta, tube distincte
breviora; ovula 2; discus nectarifer curvus super ovarium,
crassus, ligulatus. Semina elliptica, 2. 2-2. 4 mm longa,
margine arguta cineta.

Herbaceous annuals, or possibly suffrutescent,
short-lived perennials, 30—35 cm tall, erect, stems
slender to nearly filiform, much branched, the
branches varying in length, some as long as the
primary stem, bearing thin, weak, colorless or pur¬
ple glandular trichomes ol varying lengths to 0.5
mm long; internodes mostly shorter than the sub¬
tending leaves but internodes, leaves, and petioles
highly variable in length. Leaves opposite, petioles
5-27 mm long; blades 20—50 X 10—35 mm, thinly
membranous, ovate, base shallowly to deeply cor¬
date, apex acute to acuminate, margin plane; blade
surfaces sparsely strigose above, glabrous below ex¬
cept for minute trichomes on the veins; leaves grad¬
ually reduced in size toward the stem apex to form

Novon 15: 59-68. 2005.

60

Novon

Figure I. Cuphea aquilana S. A. Graham & T. B. Cavalcanti. — A. Habit. — B. Flower in late fruiting stage, lateral
view. — C. Flower in early fruiting stage, lateral view. — I). Flower opened dorsally. — E. Seed. — F. Leaf, adaxial
surface. (.Soto Nunez 11194, MO.)

the bracts of the inflorescence; bracts paired, un¬
equal. one strongly reduced or absent. Inflores¬
cences leafy racemes; flowers solitary, interpetiolar.
alternate; pedicels 2—5 mm long, filiform, the brac-
teoles (prophylls) represented by 3 to 5 glandular
trichomes. Floral tubes 4—6 X 1.5— 1.7 mm, the
base rounded or the dorsalmost rib extenders 0.2

mm al the base of the lube to form a minute dorsal
spur, the neck ol the floral tube not contracted at
anthesis, the tube greatly distended in fruit becom¬
ing ovoid to suborbicular. the mouth tightly closed;
outer surface completely green or dorsally flushed
red-purple, glabrous or sparsely hispid toward the
base, the trichomes 0.2 mm long, pale or reddish;

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Cavalcanti & Graham
Lythraceae from Latin America

61

inner surface neither bialate nor vesiculate, lightly
pilose above the stamens, glabrous below; calyx
lobes strongly unequal, the dorsal calyx lobe 0.5—
0.75 X 2.0 mm, semicircular, white, deflexed in
fruit closing the mouth of the tube, the other 5 lobes
0.2— 0.3 mm, triangular, white; epiealyx appendages
absent or represented by slight thickenings at each
sinus between adjacent lobes; petals 0; stamens 1 1,
included, all inserted at ca. % the length of the
tube, glabrous, the 2 dorsalmost stamens shortest,
the antesepalous longest; filaments glabrous; an¬
thers not reaching the sinuses of the lobes; pollen
oval-triangular in polar view, tricolporate, syncol-
pate, pores protruding, exilic striate on the equator
between the poles, the striae extending ca. Vi the
distance to the poles, 22.5 /am diam. in lactic acid;
style included, glabrous; stigma capitate to punc-
tiform, at the level ol the anthers; ovules 2; nec¬
tariferous disc I mm long, 0.2 mm wide, ligulate,
curved over the dorsal side of the ovary. Seeds 2.
2.2— 2.4 X 2.0 mm, bilaterally compressed, ellip¬
tical to oval in outline, brown, with a narrow, round¬
ed encircling margin, the margin thinned. 0.1 mm
wide at the base of the seed flanking the hilum.

Phenology. Flowering and fruiting in Septem¬
ber anti October.

Distribution and ecology. On slopes of the Si¬
erra Madre, coastal Michoacan, Mexico, just south
of the Colima— Michoacan border; in low deciduous
woods (selva baja caducifolia); 210 m.

Cuphea aquilarm is one of several species with
flowers 6 mm long or less that combine the key
characters of sections Brachyandra Koehne (deeply
inserted stamens) and Heterodon Koehne (enlarged
dorsal calyx lobe; Koehne, 1903). The sections are
polyphyletic and their defining characters have
evolved multiple times (Graham, 1988, 1998). Cu¬
phea (updlana is morphologically most similar to C.
trochilus S. A. Graham in section Heterodon and for
the present time is placed in that section. It shares
with C. trochilus the same vegetative habit, having
slender filiform stems and thinly membranous cor¬
date leaves with petioles typically ca. 15 mm long.
It differs by the absence of petals and by a rounded
floral tube base that contrasts with the prominent
conical, horizontal to ascending spur of C. trochilus.
Seeds are 2 rather than 3 to 6 as in C. trochilus,
and the nectariferous disc is erect and strap¬
shaped, rather than triangular and deflexed. Pollen
viability of C. aquilana is approximately 100%
based on a cotton-blue/lactic acid test (Kearns &
Inouye, 1993). Position of the anthers and style at
the same level deep in the floral tube suggests the
species reproduces primarily by self-fertilization.

Paratype. MEXICO. Michoacan: Aquila, en El Mir-
ador, a 3 km al W de la desviacidn a Aquila, carr. Teeo-
man, Colima— Playa Azul, Mich., 24 Sep. 1983, E. Mar¬
tinez S. 4488 (MEXU, MO).

2, Cuphea hahiensis (Lourteig) I. 8. Cavalcanti
& S. A. Graham, comb, et stat. nov. Basionym:
Cuphea sessilifolia Martins subsp. hahiensis
Lourteig, Sellowia 39: 34. 1987. T\ PE: Brazil.
Bahia: Mono do Chapeu, Morrao, 15 Jan.
1977, G. Hatschbach 86428 [in the protologue
erroneously cited as 306428] (holotype, P not
seen: isotype, MBM). Figure 2.

Erect subshrubs, 0.60—1 m tall; branches pu¬
bescent. bearing abundant white, short, erect, glan¬
dular trichomes, 0.5—1 mm long; internodes typi¬
cally 2—3 mm long, mostly ca. Vi the length of the
subtending leaves. Leaves opposite or rarely 3-ver-
ticillate, sessile, crowded and overlapping, 5—13 X
3-9 mm, coriaceous, ovate-lanceolate, base cordate
to amplexicaul, apex acute, margin thickened,
plane or subrevolute, ciliate with short glandular
trichomes; blades bearing sparse glandular tri¬
chomes, the midvein and secondary veins promi¬
nent on the adaxial surface; leaves abruptly re¬
duced in size toward the stem apex. Inflorescences
bracteate racemes, 7—15 cm long, internodes of the
inflorescence mostly 10—30 mm long; bracts paired,
hypsophylloid, with glandular trichomes; flowers
solitary, interpetiolar, alternating al the nodes,
sometimes 1 to 3 alternate to nearly opposite on
condensed axillary branehlets; pedicels 3.8— 4.5
mm long, with the same indumentum as the
branches, bibracteolate. Floral tubes 9-11 X ca. 2
mm, including a straight to deflexed, rounded or
truncate spur extending 1.5-2. 1 nun beyond the
pedicel; outer surface wine-red dorsally including
the ribs and spur, green ventrally, hirsute, hearing
erect, minute non-glandular trichomes and erect,
pale glandular trichomes 0.5— 0.7 mm long; inner
surface not bialate or vesiculate, villose above the
stamens, lightly pilose to glabrous below; petals 6,
subequal, 4.5—6 X 2.5—4 mm, rose-violet to lilac,
caducous; stamens 11, the 2 dorsalmost shortest,
included, the others slightly exserted; filaments vil¬
lous or glabrous; pollen suborbicular in polar view,
tricolporate, syncolpate, pores not protuding, colpi
bordered by narrow margo, exine psilate under light
microscope, 25—27.5 yarn diam.; style exserted be¬
yond the anthers at maturity, glabrous, red; stigma
punctiform; ovules 3; nectariferous disc broadly at¬
tached, deflexed, the apex curled under forming a
concavity. Seeds 3, ca. 2X2 mm, bilaterally com¬
pressed, orbicular in outline, the margin thick,
rounded.

62

Novon

Figure 2. Cuphea baldens is (Lourteig) T. B. Cavalcanti & S. A. Graham. — A. Habit. — B. Leaf, abaxial surface. —
C. I .cal. adaxial surface. — D. Flower, lateral view. — E. Flower, opened dorsally, without petals. — F. Seed. ( Cavalcanti
el al. 2480, CEN.)

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Cavalcanti & Graham
Lythraceae from Latin America

63

Phenology. Collected in flower and fruit from
September to March.

Distribution and ecology. Known only from ha¬
ll i a. Brazil, in the vicinity of Mono do Chapeu;
forming extensive populations along the roadsides
in the municipality; in dry sandy soils, campo ru-
pestre; 870—1 190 m.

Cuphea bahiensis is exceptional among the oth¬
erwise very similar species of section Trispermum
for the terminal, highly floriferous inflorescence,
which is well-differentiated from the vegetative
stem. Other diagnostic features of C. bahiensis are
the clasping, crowded overlapping leaves on ex¬
tremely shortened internodes, and the large flowers
(9—1 1 mm long vs. mostly less than 8 mm long in
the other rose-flowered species of the section). In
the most recent treatment of the section (Lourleig,
1987), C. inaequalifolia Koehne and C. gracilis
IIBk are also described with terminal inflorescen¬
ces. but examination of numerous specimens, in¬
cluding those cited by Lourteig, reveals instead
leafy indistinct racemes with flowers partially ob¬
scured by the leaves. Although Cuphea sessilifolia
is vegetalively highly variable, it is unlike C. bah¬
iensis except in the commonly held characters that
define this taxonomically difficult section, i.e., 3-
ovulate ovary, apex of the nectariferous disc curled
under to form a concavity, and seeds orbicular w ith
a rounded margin. The full description of C. bah¬
iensis expands the original brief diagnosis of the
variety.

Selected representative specimens. BRAZIL. Buhia:
Mono do Chapeu. estrada p/o Mono da Torre de trans-
missao, ca. 10 km a partir da sede, Amorim , Carvalho,
Silva A’ Jardim 1049 (MO); Mono do Chapeu— Bonito, km
12. I Mar. 1989, Cavalcanti. Pereira-Silva, Kirkbride Jr.
& Roath 457 (CEN, MO); Mono da antena, entrada pela
estrada para Wagner, Cavalcanti, Pereira-Silva, Kirkbride
Jr. & Roath 460 (CEN, MO); Mono do Chapeu— Piritiba,
km 8, Cavalcanti , Pereira-Silva. Kirkbride Jr. & Roath 466
(CEN, MO); estrada Utinga— Morro do Chapeu, ea. 10 km
de Morro do Chapeu, 12 Jan. 1999, Cavalcanti. Giulietti,
Silva & Harley 2480 (CEN. HUEFS, MO); Telebahia Tow¬
er, ea. 6 km S of Morro do Chapeu, Mori & Room 14445
(MO. NY); 1—2 km S da eidade, na estrada para Utinga,
16 Nov. 1984, Noblick 4478 (CEN. HUEFS); eslaeao re-
transmissora da Telebahia, ea. 6 km A da BA-046, Quei-
roz & Nascimento 4282 (HUEFS, NY); Utinga, 4 km de
Morro do Chapeu. 25 Sep. 1985. Wanderley s.n. (CEN,
SB).

3. Cuphea parsonsia var. grisebachiana (Ko¬
ehne) S. A. Graham, comb, et slat. nov. Bas-
ionym: Cuphea grisebachiana Koehne. in Mar¬
tins, FI. Bras. 13: 225. 1877. Parsonsia
grisebachiana (Koehne) Jennings, Ann. Car¬
negie Mus. II: 199. 1917. TYPE: Cuba. Cuba
occidental, 1863, C. Wright 2546(11 442) (lee-
totype, designated here, GOET; isotypes, BM,
GH, K. MO).

Robust perennial herbs to 40 cm long with semi-
woody stems erect to trailing, sparsely branched.
Leaves with petioles 1—3 mm long, blades 10—25
X 5—10 mm, ovate-oblong, membranous, somewhat
fleshy, glaucous, base rounded, apex acute. Floral
tubes 6—7.5 mm long, glabrous; petals 4 mm long;
stamens 11. Seeds 6, 2 X 1.75 mm, orbicular-ob¬
long. Chromosome number: n — 32 + 1-3 B chro¬
mosomes (Graham 1125, HAG).

Phenology. Flowering throughout the year.

Distribution. Cuba, in Pinar del Rio. Isla de la
Juventud, anil western La Habana provinces; wet
low places in savannas, disturbed grassy fields,
muddy banks; 50-500 m.

Koehne distinguished Cuphea grisebachiana by
quantitative differences that overlap those of the
more widely distributed Antillean C. parsonsia (L.)
R. Brown. Cuphea grisebachiana is judged insuffi¬
ciently distinct morphologically from C. parsonsia
to merit the rank of species in Cuphea and is here
reduced to varietal rank.

Key to the Varieties ok Cuphfa parsossia in Ci ba

la. Floral lubes 4—6 mm long, flowers inconspicuous
at the nodes; leaves elliptic to ovate, thinly mem¬
branous, petioles 0—1 mm long; stamens (1 — )5 —

8( — 1 1) . \iar. parsonsia

lb. Floral tubes 6—7.5 mm long, flowers conspicuous
at tbe nodes; leaves ovate-oblong, somewhat
fleshy, glaucous, petioles 1—3 mm long; stamens

11 . var. grisebachiana

Plants of Cuphea parsonsia var. grisebachiana
are more robust than those of variety parsonsia and
consistently have larger flowers and the full com¬
plement of 1 I stamens. Both varieties are sell-fer¬
tilizing (Graham, 1998, and pers. obs.) and differ
in chromosome number. First chromosome counts
for the Cuban plants are as follows: Cuphea par¬
sonsia var. parsonsia: n — 16. Cuba: Isla de la Ju¬
ventud, betw. Nueva Gerona and La Fe, .S'. Graham
1122 (HAC); Cuphea parsonsia var. grisebachiana :
n = 32 + 1—3 B chromosomes. Cuba: Pinar del
Rio, ca. 3 km NE of Pinar del Rio city, swale E ol
the autopista, S. Graham 1125 (HAC). The counts
suggest that variety grisebachiana originated as an
autopolyploid of variety parsonsia through self-fer¬
tilization of unreduced gametes. None ol the other
species of Cuphea sect. Brachyandra in Cuba is
sufficiently morphologically similar to imply a hy¬
brid origin for variety grisebachiana. The range of
variety grisebachiana is restricted to Cuba and lies
w ithin that of variety parsonsia, which is distributed
ihroughout the Greater and Lesser Antilles (Gra¬
ham. 2003).

Selected representative specimens. CUBA. C. Iluhunu:

64

Novon

Lomas tit* la Lilt*, S. Miguel de Casanova, lira. Iron 71631
(N\ ). Isla tie hi Juventud (Isla tit* I'inos): near Nueva
Gerona, Curtiss 433 (BM, GH, K, MO, NY, l S); savanna
near Nueva Gerona, Roig & Cremata 1754 (NY). La Ha-
liana: cerca tie Artemisa, Wilson 1729 (HAC, NY). Pinar
tlel Kite Rangel, Sierra del Rosario, Bra. Alain 192 ((ill.
NY); vie. of Coloma, Britton , Britton A Gager 7036 ( N N );
vie. of Herradura, Britton & Carle 6567 (NY, US); ea. 3
km NK t>f Pinar tlel Rio city tin the autopista, Graham
1 125 (HAC, MO); not far from Lagoon Maeeriges, S of Los
Palaeios. Bro. Iron 7354 (NY); vie. of Los Palacios. Shafer
1 1666 (A, NY. US).

F. 4 .uplicu polamnpliila T. IL Cavalcanti & S. A.
Graham, sp. nov. TYPK: Brazil. Goias: Caval-
cante, margem direita do rio Macacao. 13 Dec.
2000, G. Pereira-Silva & ./. B. Pereira 4500
(holotype, CEN; isotypes. MO. SPF). Figure 3.

Frutieulus 80 em altus, sint* ramis, caules pil is mal-
pighiaeeis obsitis. Folia subsessilia, elliptiea at! oblonga,
30—100 mm longa, 15—35 mm lata. Racemi simplices,
Irondosi. Flores 19—21 mm longi, virides apiee sanguineo,
ealcari 2—3 mm longo, rlescendenti; lobus dorsalis baud
produetus; appendices lobis breviores; petala 6. subae-
qualia, 4—5 mm longa, sanguinea; ovula 6; discus nectar-
iler horizontal is. erassus.

Perennial subshrubs to 80 cm, steins erect, sev¬
eral from a woody rootstock, unbranched, reddish
brown bark exfoliating in narrow thin strips at the
base, distal ly bearing dense soft curled pubescence
and abundant malpighiaceous, unequally bi-armed
trichomes, the basal-directed arm shortest; inter-
nodes Vi the length of the subtending leaves.
Leaves opposite, petioles 2 mm long; blades 30—
100 X 15—35 mm, thickly membranous, elliptical
to lanceolate or oblong, spreading, base attenuate,
apex acute or short-acuminate, margin plane; blade
surfaces scabrous, the trichomes minute, ap-
pressed, tin* upper surface also with scattered mi¬
nute* bi-armed trichomes; leaves gradually reduced
in size toward the stem apex. Inflorescences ter¬
minal leafy racemes, 6—10 cm long, sparsely flow¬
ered, interpetiolar flowers solitary, alternate, I to 3
additional flowers on axillary branchlets; pedicels
6— 10 mm long, hibracteolate; bracteoles ovate, ea.

I mm long. Floral tubes 19-21 X 4—5 mm. includ¬
ing a descending spur 2—3 mm long, the floral tube
dorsally straight, the neck not contracted in fruit,
the mouth blunt to slightly oblique by extension ol
the ventral side; outer surface green, becoming red
distally and vertically from the level of anther in¬
sertion to the margin of the tube, bearing minute
bi-armed trichomes, the ribs with conspicuous red-
purple setae to 2 mm long; inner surface lightly
pilose above the stamens and on the vein below
each short dorsal stamen; calyx lobes equal, 1 X
1.5 mm, broadly triangular; appendages ea. Vi as

long as the calyx lobes, green, thick, the margin
free, bearing 1 to 3 red-purple setae to 3 mm long;
petals 6, subequal, 4—5 X 2—4 mm, clear bright
red, oblong to slightly obovoid; stamens I I. the 2
dorsalmost shortest, inserted below the ventral 9, 5
antesepalous stamens exserted, the others reaching
the margin of the tube; filaments lightly villous; pol¬
len oval-triangular in outline, tricolporate, syncol-
pate, the exine striate between the poles, 25 gin
diam.; style long-exserted. glabrous; stigma capitate
to punctiform; ovary non-gibbous, glabrous; ovules
6; nectariferous disc 1.5 mm long, 2.5 mm wide,
thick, rounded on the margin, in the same plane as
tin* ovary. Seeds 3 X 2.7 mm. bilaterally com¬
pressed, suborbicular, dark greenish brown, the
rounded margin paler in color.

Phenology. Known to flower and fruit in De¬
cember and February.

Distribution and ecology. Brazil, in west-cen¬
tral Goias; rare in an inundated area at the margin
of rio Macacao in a gallery forest, and in a nearby
humid field on the slope of the serra; 300-380 m.

Cuphea potamophila (Gr., “river-loving") is most
similar to C. grandiflora Koehne in section Melvilla
subsect. Pachycalyx, differing principally by the
green floral tube with red apex and red-purple se¬
tae. In C. grandiflora, the floral tube is red with a
yellow apex and the indumentum is strigose or hir¬
sute. The petals of C. potamophila are larger than
those of C. grandiflora (4—5 mm long vs. 1.5— 2.5
mm long) and clear bright red rather than deep
wine-black. Vegetatively the species are very sim¬
ilar and both grow in riverine habitats in Goias.

Paratypes. BRAZIL. Goias: Cavalcante, estrada Vila
Veneno— Serra Branca, km 3.6 (margem direita do rio Ma¬
cacao), 20 Felt. 2000, G. Pereira-Silva, ./. B. Pereira, ./.
A. Jesus. M. C. Silva & S. <las Santos 5900 (CEN); Cav¬
alcante. margem direita do rio Macacao, 29 Nov. 2001. G.
Pereira-Silva, ./. II. Pereira. J. A. Jesus & J. P. B. Pastore
5765 (GEN, MO).

5. Guplieu polcbra Moricand var. corollutn T.
B. Cavalcanti S. A. Graham, var. nov. T\ PF:
Brazil. Bahia: Piata, estrada Piata— Boninal,
Tiguco. 1 180 m, 12 Nov. 1996, //. P. Bautista
& N. Hind (PCD) 4226 (holotype, HUFF'S;
isotypes. ALCB. CF1N [4|, SPF). Figure 4.

A varietate typica 6 petalis conspicuis, sanguineis dif¬
fer!.

Shrubs 1 — 1.2 m tall, much branched distally,
branches bearing abundant, short glandular tri¬
chomes; internodes 5—20 mm long. Leaves oppo¬
site, sessile, blades 10—23 X 5—18 mm, oval-lan¬
ceolate to ovate, c hartaceous, base subcordate to

Volume 15, Number 1
2005

Cavalcanti & Graham
Lythraceae from Latin America

65

Figure 3. Cuphea potamophila T. 15. Cavalcanti & S. A. Graham. — A. Habit. — H. Flower, opened dorsally. C.

Flower, lateral view. — I). Ovary with basal nectariferous disc (Pereira-Silva & Pereira 4500 , CEN). — F. Seed ( Pereira -
Silva et al. 5900, CEN).

66

Novon

5mm

Figure 4. Cuphea pulchru var. corollata T. 15. Cavalcanti & S. A. Graham. —A. Habit. B. Seed. C. Leaf, abaxial
surface on left, adaxial surface on right. —I). Flower, lateral view. — F. Flower, opened dorsally, without petals. (Bautista
* Hind 4226, CFIY)

Volume 15, Number 1
2005

Cavalcanti & Graham
Lythraceae from Latin America

67

cordate, apex obtuse to acute or sometimes acu¬
minate, margin subrevolute, short filiate with
sparse glandular trichomes; blade surfaces sca¬
brous, non-glandular trichomes minute and ap-
pressed, glandular trichomes short, sparse; leaves
gradually reduced in size toward the stem apex.
Inflorescences terminal leafy racemes of 2 or 3
flowers; pedicels 9—12 mm long, bibracteolate;
bracteoles broadly ovate, 1—1.2 mm long. Floral
tubes 12-20 X 4—5 mm. including a descending
spur 1—4.5 mm long, the tube dorsally straight; out¬
er surface red-orange with dark red veins, becom¬
ing yellow-red apically and ventrally, pubescent;
inner surface glabrous or lightly pubescent espe¬
cially around the base of the ovary, pilose above
the stamens; calyx lobes equal, 0.5 mm long; epi-
calyx appendages 0.5— 0.7 mm long, green, bearing
minute glandular trichomes; petals 6, subequal, 3-
3.5 mm long, 1.5— 2.5 mm wide, deep red, oblong
to obovoid; stamens 11, 5 antesepalous exserted.
the others reaching the margin of the tube; style
lightly pilose; stigma punctiform; ovary glabrous;
ovules 3 to 6; nectariferous disc 1.5 mm long. ea.
2 mm wide, thick, semi-circular in dorsal view.
Seeds not seen.

Phenology. Mowers collected in November and
April, immature fruits present in April.

Distribution and ecology. Brazil, in central Ba¬
hia. in campo rupestres and along margins of cer-
rado woodlands within the range of Cuphea pulchra
Morieand var. pulchra ; 1 180 m.

Ki- 'i to mi: Variktiks ok Cuphea vt i.chr i in Bahia,
Brazil

la. Petals 0; floral tubes pilose to villose within; in¬
florescences terminal racemes typically with ca.

8 to 1 < flowers . var. pulchra

lb. Petals 6, deep red; floral tubes glabrous to pilose

within; inflorescences limited to 2 or 3 terminal
flowers . var. corollata

I he presence of six deep red petals in Cuphea
pulchra var. corollata easily separates it from apet-
alous C. pulchra var. pulchra Morieand. Plasticity
of petal presence and reduction in petal number or
size within a species is found in other species of
Cuphea and in other genera of the Lythraceae, as
for example in Cuphea paucipetala S. A. Graham
(Graham, 1988) and Ammannia latifolia L. (Gra¬
ham. 1985). (riven that tin1 flowers of variety co¬
rollata are otherwise very similar or identical to the
non-petaled ones of variety pulchra and occur on
plants within the larger range of variety pulchra, we
choose to recognize this distinction at the varietal
level.

Paratype. BRAZIL. Italiia; Jacobi na, Sena da Ja-

queira, na subida da Fazenda do Japones, 2 Apr. 1999,
R. M. Harley & A. \1. Giulietti 53695 (CEN, HUEFS).

6. Diplusodon virgatus var. occidentals T. B.

Cavalcanti & S. A. Graham, var. nov. TYPE:
Bolivia. Santa Cruz: Velasco, Par. Nae. Noel
Kempff Mercado, Camp. Huanchaca I. cerra-
do, 650 m. 17 May 1994, L. Arroyo. H. Mos-
tacedo , H. Gonzales, S. Cabrera & ./. Suruhi
684 (holotype. MO; isotype. LJSZ not seen).

Differt haec varietas a habitu robusto frutieulo fnliis
sessilis basi cuneatis, floribus carnosioribus globosiorihus.

Robust shrubs 0.7-2 m tall. Leaves sessile, 20-
48 X 7—13 mm, subcoriaceous, ovate-lanceolate,
the base cuneate, the apex obtuse to ret use, vena¬
tion eucamptodromous. Inflorescences leafy, race¬
mose, the accessory branches ascending; pedicels
2 mm long; bracteoles 5-6 X 2-3 mm, elliptie-
laneeolate, reaching the sinuses of t lie calyx lobes
or slightly below. Moral tubes 5—6 mm long from
base to the sinuses of the calyx lobes, 4.5— 5.5 mm
wide, subglobose in fruit; calyx lobes I ,L> — 2 mm
long, 1—1.3 mm wide; epiealyx segments 2.5—3 X
0.5—1 mm, exceeding the calyx lobes, linear-ob¬
long. thick, partially deflexed in fruit; stamens 12:
style 10 mm long; ovary glabrous; ovules 38.

Phenology. Flowering and fruiting in May.
fruiting in June.

Distribution and ecology. Eastern Bolivia, west¬
ern Mato Grosso, and Mato Grosso do Sul, Brazil,
in woody savanna |cerrado]; 500—650 m.

Kia to thi: Variktiks ok Dii't.i so do \ vircati s in Brazil
and Bolivia

la. Leaves petiolate, long-attenuate; branches of the

inflorescence diverging from the main stem . . .

. var. virgatus

lb. I .eaves sessile, cuneate at the base; branches of

the inflorescence ascending . var. oecidentalis

Among the 60 or more species of Diplusodon. I).
virgatus Pohl is unique in having white-petaled
flowers and a small tree habit (4 m). The rest of the
genus consists of subshrubs and shrubs (mostly
0.5-2.0 m) with rose to purple flowers. This new
variety is also white-petaled but is a robust shrub
with more coriaceous leaves and thicker epiealyx
segments on the flowers than in variety virgatus. Of
greatest diagnostic value are the sessile, narrowly
cuneate leaves and short ascending branches of the
inflorescence, which form a narrow angle with the
stem. Diplusodon virgatus var. virgatus has slender,
long-attenuate, petiolate leaves; the branches of the
inflorescence typically form a wide angle with the
stem.

68

Novon

An intermediate collection (BRAZIL. Mato
Grosso: ca. 9 km INK of Barra do Garzas, 6 Mai.
1973, W. R. Anderson 9795 | MO, NY, U B]), with
subcoriaceous, sessile leaves and widely divergent
inflorescence branches, is known from easternmost
Mato Grosso near the border with Goias in a geo¬
graphically intermediate position between variety
occideritalis and the widespread eastern Brazilian
variety virgatus. Diplusodon virgatus var. occiden¬
tal is is the second Diplusodon to be discovered in
Bolivia. Diplusodon bolivianos T. B. Cavalcanti &
S. A. Graham is also from cerrado in Noel Kempff
M. National Park, in an area south of the I). vir-
gatus var. occidentals locality (Cavalcanti & Gra¬
ham, 1996). It is easily separated from I). virgatus
var. occidentals by the rose-petaled flowers, dis¬
tinctly terminal inflorescences, and by the hirsute
to sericeous yellowish indument that covers all
parts of the plant.

Paratypes. BRAZIL. Mato Grosso: Juruena, Apr.
PXtO, K C. line line 179 6, 1798 (R), May 1909, F. C.
Hoehne 1846 (R); Nova Ubirata, estr. rio Teles Pires. I
May 1997, A. G. Rave 1415 (CEN); Biamantino, Faz. IV-
queno Figueiredo, cerea de 4 km em linha reta de l)ia-
mantino, 22 May 1997, V. C. Souza, A. R.. Duarte, P.
Souza A- V. R. Scalon 16844 (CEN, ESA); Tapurah, estr.
Capixaba, ea. 20 km de Tapurah, 12 June 1997, V. C.
Souza, A. R. Duarte, J. P. Souza & Miyagi 17880 (CEN,
ESA). Mato Grosso do Sul: Seluria, Faz. de Ensino e
Pesquisa da UN ESI’ Campus lllia Solteira, 22 Apr. 1991,
J. Santos 870 (UB), 5 Apr. 1991, 0. Tiritan 584 (UB),
May 1991, J. Santos 418 (UB).

Taxonomic Notks

Guplicu xanlliopet ala S. A. Graham & I. B.
Cavalcanti was described from a single collection
from Mato Grosso, Brazil (Graham Si Cavalcanti,
1999). The following collection extends the range
of the species to the state of Amazonas and con¬
firms it as a species of white sand habitats. BRA¬
ZIL. Amazonas: rodovia do Estanho, rnargem da
rodovia a 150 km de Humaita, eampina, solo ar-
enoso Branco, 25 Sep. 1979, G. Vieira, ./. Zarucchi,
A. Silva, C. Mota Sc 0. Monteiro 148 (I NBA, MO,
NY).

Diplusodon bolivianos I. B. Cavalcanti & S.
A. Graham, previously known only from the type
collection, is further represented by the following.
BOLIVIA. Santa Cruz: Velasco Prov., Par. Nac.

Noel Kempff Mercado colecta general de Huancha-
ca I, en la pampa cerea de la pista de Huanchaca,
A. Soto, Parnfil, Moore Sc Soliz 865 (MO): eolectado
a 2 km de la pista, Soto, Pamfil , Moore & Soliz
427, 468 (MO). BRAZIL. Mato Grosso: BB 364,
Vilhena-Comodoro, 84 km S de Vilhena, 25 June
1997, Cavalcanti, Pereira-Silva, Baltazar Sc Gra¬
ham 2881 (CKN, NY); Tapurah, estr. Capixaba, ca.
20 km NE de Tapurah, 10 June 1997, Souza. Duar¬
te, Souza Sc Miyagi 17617 (CEN, ESA), 12 June
1997, Souza, Duarte, Souza Sc Miyagi 17868 (CEN,
ESA).

Acknowledgments. We thank the curators of
MEXU, NY, and HUEFS for the opportunity to
study their collections; Rogerio da Costa Vieira,
Aeeio Amaral dos Santos, and Jose Geraldo Alves
Vieira (CEN) for specimen and data transfer to MO;
and an anonymous reviewer for helpful comments.
The support of Embrapa and GNPq made possible
field and herbarium studies by T. B. Cavalcanti and
S. Graham with the assistance ol Glocimar Pereira-
Silva. 'fhe National Science Foundation is grate¬
fully acknowledged for partial support to S. Graham
of this research (DEB-9509524). I he illustrations
were drawn and inked by Jeanitto Sebastian Gen-
tilini and Taeiana Cavalcanti.

Literature Cited

Cavalcanti, T. B. & S. A. Graham. 1996. Diplusodon bo¬
livianos sp. nov. (Lythraceae), the first report of the ge¬
nus for Bolivia. Novon 6: 253—255.

Graham, S. A. 1985. A revision of A mmannia (Lythraceae)
in the Western Hemisphere. J. Arnold Arbor. 66: 395—
420.

- . 1988. Revision of Cuphea section Heterodon

(Lythraceae). Syst. Bol. Monogr. 20: l-U>8.

- . 1998. Relacionamentos entre as especies aut6-

gamas de Cuphea I’. Browne seeao Hracliyarulra Koehne
(Lythraceae). Acta Dot. Brasil. 12: 203—214.

- . 2003. Biogeographic patterns of Antillean Lyth¬
raceae. Syst. Bot. 28: 410—420.

- & T. B. Cavalcanti. 1999. The yellow- flowered

species of Cuphea (Lythraceae), including three new
taxa. Brittonia 51: 24—30.

Kearns, C. A. & I). W. Inouye. 1993. Techniques for Pol¬
lination Biologists. IJniv. Press Colorado, Niwol, Colo¬
rado.

Koehne, E. 1903. Lythraceae. In: A. Engler (editor). Das
Pflanzenreich IV. 216: 1—326.
liOurteig, A. 1987. Lythraceae Austroamerieanae. Adden¬
da et corrigenda II. Sellowia 39: 5—48.

Notes on Grasses (Poaceae) for the Flora of China, III:
A New Name in Festuca

Chen Xiang

Herbarium, Institute of Biology, Guizhou Academy of Sciences, Xiaohe, Guiyang,
Guizhou 550009, China, gzcxcn@yahoo.com

Sylvia M. Phillips

Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TWO 3AB, United Kingdom.

s.phillips@kew.org

Abstract. The new name Festuca sinomutica X.
Chen & S. M. Phillips (Poaceae) is proposed for a
species of Festuca from Yunnan Province, China.

Key words: China, Festuca , Poaceae, subgenus
Drymanthele.

During ihe course of revisionary work on Festuca
L. for (lie Flora of China Poaceae account, it was
found that a new name is required for Festuca mu-
tica S. L. Lu, which is a later homonym of F rnutica
Chevall. Lu Sheng-Lian was the author of the ac¬
count of Festuca in the Chinese-language Flora
(2002), used as the basis for the English-language
Flora oj China , now in preparation.

Festuca sinomutica X. Chen & S. M. Phillips,
nom. nov. Replaced name: Festuca rnutica S.
L. Lu, Acta Phytotax. Sin. 30(6): 534, f. 3.
1992, non F. rnutica L. E. Chevallier, El. Gen.
Env. Paris ed. 1: 167. 1827, nee F. rnutica F.
X. Wulfen, El. Noriea Phan.: 145. 1858.
T\ PE: China. Yunnan: Eryuan, in the vicinity
of Nandaping Heishanmen, 2900 m, in mead¬
ows, 31 July 1963, Jinshajiang Expedition 63-
6359 (holotype, PE not seen; isotype, NJU not
seen).

Festuca is a large genus in China, with 58 spe¬
cies recorded as present in the Flora of Chirm ac¬
count, almost half ol them endemic and many of
these based on single specimens. This species, F.
sinomutica, appears to lie- known only from the
type, which has not been seen. However, a full de¬
scription and detailed illustration are provided in
the protologue.

Festuca sinomutica is a member of subgenus
Drymanthele V. Kreczetowiez & Bobrov, which
comprises relatively robust species with the follow¬

ing characteristics: large loose panicles, flat leaf
blades, coriaceous scales at the- base of the shoots,
leaf sclerenchyma as individual bands opposite the
vascular bundles, awnless lemmas keeled only
above the middle, and a pubescent ovary apex.

S. L. Eli (1992) placed this species (as Festuca
rnutica ) in subgenus Drymanthele and made it the
type species ol the new section Muticae S. E. Lu,
including also Festuca changduertsis L. Liou, F
dolichantha P. C. Keng, and F. modesta Steudel.
Festuca modesta occurs in northwestern India and
Nepal as well as China, but tlx* others in section
Muticae are Chinese endemics. Festuca chang-
duensis and F. dolichantha are anomalous in sub¬
genus Drymanthele as they have a glabrous ovary.
It is clear that much more work is needed on Fes¬
tuca in China when more material is available.

A key is provided below to separate the four. spe¬
cies of section Muticae. All are over 60 cm tall,
usually with flat leaf blades and with awnless lem¬
mas.

Key to Species of Festuca Sect. Muticae in China

la. Panicle 7—9 cm; florets 9 to 10 .... F. sinomutica
II). Panicle more than 15 cm; florets 3 to 5.

2a. Spikelets 6—7 mm; first lemma ea. 5 mm;

anthers 1 .5—2 mm . F. changduertsis

2b. Spikelets 7—15 mm; first lemma 7—1 I mm;
anthers 2.5—4 mm.

3a. Leaf sheaths smooth or scabrid; anthers
3—4 mm; ovary apex glabrous ....

. F. dolichantha

3b. Leaf sheaths with retrorse hairs at base;
anthers 2.5— 3.2 mm; ovary apex pubes¬
cent . F. modesta

Literature Cited

Lu, S. 1.. 1992. New taxa of Festuca I,, from China. Acta
Phytotax. Sin. 30; 529—540.

- . 2002. Festuca. In Flora Keipublicae Popularis

Sinica, Vol. 9(2): 40-84.

Novon 15: 69. 2005.

Novae Gesneriaceae Neotropicarum XIV: Four New Species of
Alloplectus from South America

John Littner Clark

Department of Biological Sciences, George Washington University, 2023 G Street, N.W.,
Washington, DC 20052, U.S.A., and Botany Department, MRC-166, Smithsonian Institution,
1*0. Box 37012. National Museum of Natural History, Washington, DC 20013-7012, U.S.A.

clarkjo@si.edu

Laurence E. Skog

Botany Department, MRC-166, Smithsonian Institution, P.O. Box 37012, National Museum
of Natural History, Washington, DC 20013-7012, ll.S.A. skogl@si.edu

Ira y<l a Salinas

Division de Botamea, Museo de Historia Natural, Av. Arenales 1256, Casilla 14-0434,

Lima 14, Peru. Lsa_hi@yahoo.com

Ahsth act. A monographic revision of Alloplectus
(Gesneriaceae) has revealed four species new to
science from Colombia, Ecuador, and Peru. These
species are described here and include A. carpish-
ensis from Peru, A. inflalus from Colombia, A. ser¬
pens from Colombia and Ecuador, and A. oblongi-
ealyx from Colombia and Ecuador.

Key words: Alloplectus , Colombia. Ecuador.
Gesneriaceae, Peru.

The most recent treatment of the genus Alloplec¬
tus Marlins is that of Hanstein (1865). Ilansteins
publication focused on tfie Gesneriaceae at the bo¬
tanical garden in Berlin to provide an overview of
the family, and Alloplectus was only a part of a
monograph of the known Gesneriaceae. Hanstein
recognized 30 Alloplectus species in his treatment.
Recent estimates of the genus range from 65 (Wieh-
ler. 1973) to 75 species (Burtt & Wiehler, 1995).
From a thorough review of the I 19 names listed in
Index Kewensis, only 32 are considered by us to be
currently accepted names in Alloplectus.

Preliminary phylogenetic analyses based on the
nuclear ribosomal internal transcribed spacer re¬
gion (ITS) have shown that Alloplectus is polyphy-
letie (Clark & Zimmer, 2003). A more restricted
monophyletic group containing the type species for
the genus, Alloplectus hispidus (Kunth) Martins, in¬
cludes the new Alloplectus species presented here.
This more restricted definition of Alloplectus is
characterized by the presence of resupinate flowers,
a unique synapomorphy for the clade. The species
described here were frequently collected and rep¬

resented in numerous herbaria, but were annotated
as belonging to other species of Alloplectus. I he
four new species below are derived from a mono¬
graphic study of Alloplectus and allied genera, a
review of herbarium collections (4000+ collections
from 30+ herbaria), and live years of fieldwork by
the first author in Ecuador with recent forays of
fieldwork in Peru and herbarium work in Colombia.

Alloplectus carpisliensis J. E. Clark & I. Salinas,
sp. nov. TYPE: Peru. I luanuco: Chinchao, San
Pedro de Carpish, above Carpish tunnel, ca.
47 km N of Hu&nuco on the road to lingo
Marfa, 09°43'14"S, 76°06'53"W. 2770-2900
m, 1 Nov. 2001, I. Salinas 224 (holotype,
USM; isotype, US). Figure I.

Ad A. serpentem similis sed in foliis elliptieis dense
pubescentibus, lobis calycis apice acuminatis et corollis
uniforiniter flavis differ! .

Subshrubs; terrestrial, stems erect to horizontal
(on steep slopes), rarely branched, to 2 m tall, sub-
woody, subquadrangular to terete, glabreseent be¬
low, densely villous above. Leaves opposite, equal
in a pair; petioles terete, 1-4 cm long, green,
densely villous; blades subcoriaceous when dry, el¬
liptic to slightly falcate, 8—19 X 2—5 cm, base
acute to oblique, apex acute, margin serrate, adax-
ially green, uniformly sericeous, abaxially pale
green, densely sericeous (especially on veins). In¬
florescence epeduneulate, reduced cyme, appearing
fasciculate, with I to 7 flowers per node; bracteoles
ovate, 2—3 X 1-2 cm; pedicels shorter or longer

No von 15; 70-79. 2005.

Volume 15, Number 1
2005

Clark et al.

Novae Gesneriaceae Neotropicarum XIV

71

1cm

Figure 1. Alloplectus carpishemis J. I,. Clark & I. Salinas. — A. Habit. — B. Bract. — C. Calyx. — I). Corolla. — K.
Corolla opened to show stamens. — F. Calyx opened and corolla removed to show valvate nectary glands. — G. Mature
fruit. — H. Seeds. (A— H from holotype, /. Salinas 224, USM.)

than the petiole, 1—4 cm long, densely villous.
Flowers zygomorphic, resupinate; calyx lobes 5,
nearly free, each appressed to adjacent lobe, folded
lengthwise with the margin curved inward, erect,
subequal, ovate, 10—17 X 6-12 mm, base cordate,
apex acute, margin serrate, bright red, outside
sparsely to densely pilose, inside sparsely pilose,
persistent in fruit; corolla horizontal in calyx, to 4
cm long, tubular, tube 3.5 cm long, gibbous pos¬

teriorly on lower surface, spur absent; base 6—8 nun
diam., middle ampliate. becoming apically ventri-
cose on upper surface, throat constricted, appearing
laterally compressed, to I cm wide (at mouth), out¬
side yellow, densely pilose, inside mostly yellow
with red spots and small glandular trichomes, limb
rellexed, to 1 cm wide, lobes subequal, rotund, to
3 nun long, to 3 mm wide, rounded, entire; nectary
deeply bilobed to separate valvate glands, on ven-

72

No von

Iral surface of ovary, glabrous; stamens 4, didyna-
mous, included; filaments coiling after anthesis. 1.9
cm long, connate at base, adnate to base of corolla
tube for 6—8 mm, glabrous; anthers broader than
long, ca. 2 mm long, ea. 8 mm wide, dehiscing by
longitudinal slits; staminode present; ovary ovoid,
3—6 X 3—5 mm. densely pilose, style 1—2.5 cm
long, glabrous, stigma stomatomorphic. Fruit a
(leshy capsule, pendent when ripe, ovoid, bivalved,

1.5 X 1.5 cm, pilose; seeds numerous, fusiform, I —

I. 4 X 0.3 mm, longitudinally striate, dark brown.

Phenology. Flowering from December to May
and August to November; fruiting from June
through November.

Distribution (Fig. 5). Alloplectu. s carpishensis is
known from the eastern Andean slopes ol Peru from
2200 to 3000 m. Sixteen collections have been
made from Carpish in the department of Huanuco.
Two other collections are known from the depart¬
ments of Cuzco and San Martin. No collections ol
A. carpishensis have been made outside of Peru,
thus making this species geographically distinct
from the similar A. serpens of Ecuador and southern
Colombia.

Alloplectus carpishensis is distinguished from the
similar A. serpens and other species of Alloplectus
by the densely pubescent, elliptic- leaves, and uni¬
formly bright yellow corollas. Additional characters
that help differentiate A. carpishensis are the ovate
calyx lobes and acute apices instead of the more
oblong calyx lobes with rounded apic es found in A.
serpens. The nectary in this new species is unique
because it ranges from a single bilobed gland to
two separate valvate glands instead of the more-
common single bilobed gland in most other Allo¬
plectus species. Specimens of A. carpishensis were
annotated in herbaria as A. ichthyoderma Hanstein.
The stems of A. carpishensis lack the scales that are
typical for A. ichthyoderma and have uniformly
bright yellow flowers, in contrast to tlu- yellow flow¬
ers with a dorsal red stripe in A. ichthyoderma.

Para types. PERU. Cuzco: La Cnnvencicm, T. It. Dud¬
ley 10845 (NA. US). Iliianiico: above Acomayo. P. C.
Hutchison, ./. h. Wright & R. M. Slr<uc 5058 (US, USM);
Distrito Chinchao, term Carpish, near Carpish tunnel,
45—50 km N of Huanuco on the road to lingo Marfa. R.
Ferreyru 1220 (US, USM). R. Ferreyra 6 848 (US[2], USM),
C. Davidson & ./. Jones 0100 (LAM, US). R. A. Stein &
C. To, Liu 2208 (US, USM). II. A. Allard 21085 (RH. US).

J. L. Clark. I. Salinas A- II. Beltran 8247 (US. USM). t.
II. Gentry 44888 (US). R. Ferreyra 2880 (MO. US|2|,
USM), I.. F. Shog. ./. F. Skog, G. Dimas M. & J. Selmnke
V. 5140 (MO, US. USM), ./. L Clark. I. Salinas & II.
Beltran 8240 (US. USM). ./. Schunke V. 5225 (F, US). C.
Sandeman 5172 (k); F,1 Mirador, 15 km from Chinchao,
J. Schunke V. 5240 (F, US); trail from San Pedro do Car¬

pish. 3 km E ot tunnel, B. A. Stein. ./. Kallunki A C. Diaz
8851 (US. USM). San Martin: Mariscal Caceres, P. N.
Rio Abiseo, near Gran Pajaten ruins, h. Young 1200 (F).

Alloplectus inflatus J. L. Clark <X E. E. Skog, sp.
tiov. TYPE: Colombia. Cundinamarca: Mu-
chindote Valley, E side of Ouebrada Negra, 13
km NE of Gaehela, 2800 m, 4 July 1044. M.
I,. Grant 959 6 (holotype, COE; isotypes. MO,
N A 1 2 1 . NY, US. WIS). Figure 2.

Ad A. hispidum similis seel pauce ramosus, in lohis ca-
lyois valvatis margine base appressis et in corollis atro-
sanguincis differ!.

Shrubs; terrestrial or epiphytic, stems scandent,
sparingly branched; to 2 m tall, woody, terete,
densely hirsute, leaves opposite, usually equal in a
pair; petioles 0.5—2 cm long, densely hirsute;
blades coriaceous when dry, broadly elliptic, 3—10
X 1—5 cm, base somewhat rounded to acute, apex
obtuse to acute, margin serrate, adaxially dark
green, tomentose, abaxially light green to red.
sparsely to densely hispid especially on the veins.
Flowers solitary; bracteoles absent: pedicels longer
than the petiole, 2—7 cm long, pilose. Flowers sub¬
regular. not resupinate; calyx lobes 5, nearly free,
basally appressed to adjacent lobe, erect, subequal,
broadly ovate, 15—20 X 5—10 mm, base truncate,
apex attenuate, margin laciniate-serrate, red, out¬
side densely pilose, inside sparsely pilose; corolla
oblique in calyx. 4.0— 5.5 cm long, tubular, tube ca.
4.0 cm long, gibbous posteriorly on lower surface,
spur absent; base ca. I cm diam., middle ampliate,
becoming ventricose on upper surface, throat con¬
stricted. not appearing laterally compressed, ca. I
cm wide at mouth, uniformly red. outside densely
pilose, inside glabrous, limb oblique, ca. I cm
wide, lobes nearly equal, rotund, spreading, 2 X

3.5 mm. rounded, entire; nec tary gland bilobed, on
ventral surface of ovary, glabrous; stamens 4, di-
dynarnous, included; filaments coiling after anllie-
sis, ca. 1 .5 cm long, basally connate, adnate to base
of corolla tube for ca. 2 mm, glabrous; anthers
broader than long, I mm long, 2 mm wide, dehisc¬
ing by longitudinal slits: staminode absent; ovary
ovoid, ca. 7X4 mm, densely pilose, style 1 .4 cm
long, glabrous, stigma shallowly bilobed. Fruit a
fleshy capsule, pendent when ripe, ovoid, bivalved,

1.5 X 1.5 cm, pilose; seeds numerous, fusiform, ca.
1.3 X 0.4 mm. longitudinally striate, dark brown.

Phenology. Flowering from February through
December; fruiting from August through November.

Distribution (Fig. 5). Alloplectus inflatus is
known from the- Andes of Colombia. It has been
collected from departments in the Cordillera Cen-

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Clark et al.

Novae Gesneriaceae Neotropicarum XIV

73

1cm

Figure 2. All opted us inflatus ,1. I,. Clark & I,. K. Skog. — A. Habit. — B. Adaxial leal pubescence. — C. Corolla.

D. Corolla opened to show stamens. — E. Calyx. — F. Calyx opened and corolla removed to show bilobed nectary
gland. — C. Immature fruit. — II. Seeds. (A, 15 from L. Uribe II. 2594. US: C-F from G. Huertas & L. A. Canuirgo
5007. COL; C. II from F. W. Fennell 9385. I'll.)

74

Novon

tral (Caldas, Cauca, Tolima) and the Cordillera Ori¬
ental (Boyacci, Cundinamarea, Meta) from 2000 to
3500 m.

A llopt fetus inflatus resembles the type species for
the genus, A. hispidus (Kunth) Martins, but diflers
by the more inflated and remarkable pouched re¬
gion near the apex of the corolla. Both .4. hispidus
and 4. inflatus are scandent herbs, but 4. hispidus
can cover an entire tree trunk for 3-5 m, whereas
4. inflatus is not so prolifieally branched (pers.
obs.). Other characters that are useful for distin¬
guishing 4. inflatus are the dark red flowers and
the nearly valvate calyx lobes with basally ap-
pressed margins.

Paratypes. COLOMBIA. Boyaca: Cordillera Orien¬
tal, NK of Arcabuco, border of Boyacd & Santander del
Sur, II. Garcia Barriga A: R. Jaramillo M. 2026 I (h. (41.
US). Caldas: Cordillera Central, Alaska, above Salento,
F W. Pennell 9383 (GH, I'll); W slopes of the Cordillera
Central, valley of Rfo Otun, Pena Bonita, J. Cuatrecasas
23333 (F, US). Cauea: K slopes Cordillera Central, region
of Moseopan, valley of Bfo San Jos£, Aguabonita, J. Cua¬
trecasas 23484 (F, US); road Popaydn— La Plata, km KM).
Bes. For. Merenberg, C. Murcia 12 (COL). Cundinamar-
ea: bet w. Puebloviejo & Hoerfas, near Boca de Monte, L.
Uribe U. 6387 (COL); Cerros de San Miguel. SW of La
Sabana de Bogota, /.. Uribe U. 2394 (COL, US); Cordillera
Oriental, Moquentiva valley, 14 km NW of Gachetd, M.
/,. Grant 9322 (NA, US); Sibate, FI Pendn, G. Huertas A-
/,. A. Camargo 3007 (COL); Pdramo de Guasca, M.
Schneider 1196 (COL); vie. San Miguel, R. E. Weaver &
//. Kennedy 1367 ((ill). Meta: F slopes of the Cordillera
Central, Sunapds, Quebrada FI Bnque, S. Diaz P. 26/ 7
(MO); main tributary of Bio Grande, S of Cordillera tie
Las Cruces, S slopes of Paramo de Sumapaz, F. R. Fosberg
20862 (US); Quetame, Guayabetal police inspection sta¬
tion, road to Calvario, G. l/ozano C. 4063 (COL), /.. Pen-
uela & J. Clavijo 16 (COL). Tolima: near Quindio hwy.,
along divide. F. P. Fillip A- G. Varela 34612 (COL, US).

Alloplcctus oblongicalyx J. L. Clark & L. E.
Skog, sp. nov. TYPE: Ecuador. Pichincha:
Canton Quito, trail betw. I.loa & Mindo, 30 km
W of Lloa, 00°1 1 'S, 78°41'W, 1800 m, 12 Feb.
1997, J. /.. Clark & S. C. Clark 3903 (holo-
type, QCNE; isotypes, AAU, MO, NY. US).
Figure 3.

Ad 4. grandicalycem similis set! in lobis calycis oblon-
gis et in corollis uniformiter roseis dilfert.

Herbs to subshrubs; terrestrial, stems erect, un¬
branched; to 3.0 m tall, usually woody, succulent
when young, terete to quadrangular, glabreseent or
glabrous, sometimes sparsely pubescent, leaves op¬
posite, usually equal in a pair; petioles 5-12 cm
long, sparsely to densely pilose; blades coriaceous
when dry, elliptic, 12—22 X 8— 15 cm, base acute
to rounded, apex acute to obtuse, margin serrate or
serrulate, adaxially green, sparsely strigose, abaxi-

ally all green to all red, glabrous to sparsely pilose
on veins. Inflorescence epedunculate, reduced
cyme, appearing fasciculate, with 3 to 5 flowers per
node, bracteoles ovate, ca. 20 X 8 mm; pedicels
shorter than the petiole, 3-5 cm long, pilose. Flow¬
ers zygomorphie, resupinate; calyx lobes 5. nearly
free, each appressed to adjacent lobe, folded
lengthwise with the margin curved inward, erect, 4
equal, oblong, 4-5 X 20—30 mm, base cordate,
apex obtuse, margin serrulate or entire, all green,
all red, or mostly red with green-white at base, out¬
side pilose at the base, sparsely pilose above, in¬
side sparsely pilose, fifth (dorsal) lobe smaller and
narrower, oblong, 3—4 X 0.5—1 cm; corolla oblique
in calyx, 5-6.5 cm long, tubular, tube 4.5-6 cm
long, gibbous posteriorly on lower surface, spur ab¬
sent; base 0.5-1 cm diam., middle ampliale, be¬
coming ventricose on upper surface, throat not con¬
stricted. appearing laterally compressed, 13 mm
wide (at mouth), usually uniformly dark red, some¬
times appearing orange to Vermillion, outside
sparsely sericeous becoming densely sericeous dis¬
till ly. inside mostly glabrous, becoming glandular
apically, limb reflexed, to 1.5 cm wide, lobes nearly
equal, rotund, spreading, slightly recurved, 4—5 X
4.5 mm, rounded, entire; nectary gland bilobed, on
ventral surface of ovary, glabrous; stamens 4, di-
dynamous, included; filaments coiling after anthe-
sis, ca. 2.5 cm long, connate at base for 5 mm,
adnate to base of corolla tube for ca. 5 mm, gla¬
brous; anthers broader than long, ea. 1 X 3 mm,
dehiscing by longitudinal slits; staminode absent;
ovary ovoid, ca. 5X4 mm, densely pilose, style
3-4 cm long, glabrous, stigma stomatomorphie.
Fruit a fleshy capsule, pendent when ripe, ovoid,
bivalved, ca. 2 X 1.3 cm, pilose; seeds numerous,
fusiform. 0.9 X 0.3 mm. longitudinally striate,
brownish red.

Phenology. Flowering and fruiting throughout
the year.

Distribution (f ig. 5). Alloplectus oblongicalyx is
known from tin' northwestern Andean slopes of Ec¬
uador and southern Colombia from 1400 to .1200
m. Despite its distribution from the eastern Andean
slopes in Colombia (Cordillera Oriental), it has not
been found on the adjacent eastern slopes in Ec¬
uador. fins could be tht' result of displacement by
the congener Alloplectus grarulicalyx J. L. Clark &
L. E. Skog where the ranges of these two species
would lie expected to overlap.

Alloplectus oblongicalyx is distinguished from
other members of the genus by the combination of
uniformly colored corollas, an erect unbranched
habit, and calyx lobes that are conspicuously ob-

Volume 15, Number 1
2005

Clark et al.

Novae Gesneriaceae Neotropicarum XIV

75

Figure 3. Alloplectus oblongicalyx J. I.. Clark & L. E. Skog. — A. Habit. — B. Abaxial leaf pubescence. — C. Corolla.

I). Corolla opened to show stamens. — E. Calyx. — F. Calyx opened and corolla removed to show bilobed nectary
gland. — (;. Immature fruit. — H. Seeds. (A from l. Zak & J. Jaramillo 2311, US; B from G. L. Webster & II Castro
30230 , US; C, I). C. II from J. I.. Luteyn, ./. J. Pipoly, M. Lebron-Luteyn & ./. A. Kallunki HH 1 9, US; E. E from ./. /,.
Clark & S. G. Nazzaro 4588, US.)

76

Novon

long. Alloplectus oblongicalyx resembles A. gran-
dicalyx, and some ol the paratypes for the latter
(Clark & Skog 2002) are considered here as be¬
longing to A. oblongicalyx. The oblong calyx lobes
in A. oblongicalyx is the most consistent character
for differentiating it from the congener, A. graruli-
calyx. The color ol the calyx lobes in A. oblongi¬
calyx is variable and ranges from greenish-white to
red. The color ol the leaves also varies from green
to red. Many of the herbarium collections of A. ob¬
longicalyx were annotated as ,4. tetragonoides
Mansfeld or A. tetragonus (llanstein) Hanstein. The
calyx lobe margin of A. oblongicalyx is serrulate or
entire in contrast to the deeply serrate margins of
A. tetragonus. The calyx lobes of A. tetragonoides
are smaller and more ovate than A. oblongicalyx.

Paratypes. COLOMBIA. Caqueta: Cordillera Orien¬
tal. road Guadalupe— Florencia, km 14, \. Londono A L.
I’. Kvist 99 (COL, QCA, US); E slopes of the Cordillera
Oriental. Quehrada Rfo Hacha. J. Cuatrecasas 8553 (COE.
US). Gauca: Santa Rosa, Finca Teresa (El Cajdn). 2 km
above Quehrada San Antonio, near Cerro Ea Vieja, L. ./.
Ruhiano & C. Moreno 0. 507 (COE). Iluila: confluence
of Rfo Villalobos & Rfo Cauchos, R. E. Selwltes A' M.
Villarreal 5201 (COL, <411. US); road La Plata-Popayan,
km 101, li. A. Stein 5730 (COL, K. MO. US); route tow.
Agua Bonita, S. Diaz P., C. Lozano A ./. Torres R. 729
(COE); San Jose de lsnos, route tow. El Hornito, G. Ixizano
C. 3301 (COL); W slope of Cordillera Oriental, bel. Ga-
binete, ravine above San Andres, ./. Cuatrecasas 3620
(COE, US). Putumayo: E slopes of Cordillera Oriental,
betw. Sachamates & San Francisco de Sibundoy, ./. Cua¬
trecasas 11460 (COE. US); Sibundoy, betw. San Francisco
& Rfo Blanco, vfa a Moeoa. E. Hernandez. A. Guerrero A
A. Estrada 401 (US); road San Francisco— Mocoa, km 92
from Pasto, sitio Eos Monos, T. Plowman A W. Davis 4332
(COE); Valle de Sibundoy, 1.5 km E Sibundoy, M. I,. Rris-
tol 300 (COE, GH, US). ECUADOR. Carehi: above Mal¬
donado, II. van der Werff A E. Cudino 10308 (MO. OUNC¬
ES); road Tulcdn— Maldonado, ea. 14 km SF of Maldonado,
C. Ilarling A L. Andersson 12330 (GB. SEE, US). Coto¬
paxi: Cantdn Sigchos, Parroquia San Francisco de las
Pampas, Bosque Integral Otonga, ./. L. (dark & A. Munoz
0 I II (QCNE, US). Pichiucha: 42.2 km \V from Colocol-
lao, E near bridge across Rfo Alambi, II. MacRryde 100
(QCA, US); Canton Quito, Bellavista Lodge, next to Mindo
Nambillo Reserve, ea. 6 km S of Tandayapa, ./. L. Clark.
II. Creeney, M. Lysinger A T. Walla 3531 (QCNE. US);
Cordillera de los Yumbos, trail Ea Victoria—1 Chiriboga, ./.
L Clark A S. C. Nazzaro 4533 (AAU, COE. E. MO. QUA.
QCNE, SRP, US); Parroquia Calacalf, Reserva (ieobotan-
ica Pululahua, Sla. Rita, C. E. Cerdn M.. M. Cerdn A .S'.
Chipanlasi 4702 (MO. QCNE, US); Parroquia Nanegal,
Bosque Protector Maquipucuna, above Rfo Picluin, C. I..
Webster A R. Castro 30250 (DAV, QCNE, US): Parrocpiia
Nono, El Pahuma Orchid Reserve,,/. /,. (dark. It. Elcome,
M. Elcome. I\. Harris A M. Mailloux 7054 (QCNE, US);
road Quito— Nono. 12—14 km W NW ol Nono, J. L. Luteyn,
./. ./. Pipoly, M. Lebron-Luteyn A ./. 4. KallunJci 3319
(CAS, NY, QUA. QCNE, US); hwy. Quilo-Chiriboga-Em-
palme, betw. km 75 and 85, V. Zak A ./. Jaramillo 2311
(MO. NY. US).

Alloplectus serpens J. L. Clark & L. E. Skog. sp.
nov. TYPE: Ecuador. Morona-Santiago: Cantdn
Uimon-lndanza. main road betw. Gualaceo &
Plan de Milagro, “Tinajfllas,” cow pasture &
patches of montane forest, 03°00'19”N,
78°36'36"W, 2800 m, 10 Jan. 2001, ./. L.
Clark. F. Sanchez & L. Jost 59 04 (holotype,
QCNE; isotypes, AAU. HA, K. MO. NY. QCA,
US). Figure 4.

Ad A. carpishensem si m i I is sed in lobis calyeis oblon-
gioribus apiee rotundatis differt.

Lianas to subshrubs; epiphytic, stems st andout
or erect, rarely branched; to 2.0 m long, subwoody,
subquadrangular. glabrescent below, densely hir¬
sute above, leaves opposite, equal iu a pair; peti¬
oles 1—5 cm long, densely yellow pilose; blades
membranous when dry, ovate, 4—15 X 3—10 cm,
base acute to rounded, apex acuminate, margin ser¬
rate to crenate, adaxially green, sparsely pilose,
abaxially uniformly pale green to uniformly red.
sparsely to densely pilose, especially on veins. In¬
florescence epedunculate. reduced cyme, appearing
fasciculate, with I to 3 flowers per node; hraeteoles
ovate, 1-2 X 0. 5-1.0 cm; pedicels shorter to longer
than the petiole, 1-3.5 cm long, densely pilose.
Flowers zygomorphic, resupinate; calyx lobes 5,
nearly free, each appressed to adjacent lobe, folded
lengthwise with the margin curved inward, erect,
subeqnal, ovate, 20-25 X 20-25 mm, base sagit¬
tate. apex rounded, margin serrate, red, outside
sparingly pilose with the midvein densely pilose,
inside pilose; corolla horizontal in calyx, 2— 4 cm
long, tubular, tube 2^1 cm long, gibbous posteriorly
on lower surface, spur absent; base 9—1 I mm
diam., middle ampliate, becoming ventrieose on
upper surface, throat slightly constricted, appearing
laterally compressed, to 1.5 cm (at mouth), outside
glabrous at base, otherwise densely pilose, inside
glabrous, limb reflexed, ea. 0.5 cm wide, lobes
nearly equal, rotund, ea. 5 X 3 mm. rounded, en¬
tire; nectary gland bilobed, on ventral surface ol
ovary, glabrous; stamens 4. didynamous. included;
filaments coiling after anthesis, to 1 .9 cm long, con¬
nate at base for 8 mm, adnate to base of corolla
tube for ca. 8 mm, glabrous; anthers broader than
long, 1.3 mm long, 2.0 mm w ide, dehiscing by lon¬
gitudinal slits; staminode absent; ovary ovoid, ca. 5
mm long, ca. 4 mm wide, densely pilose, style to
2.4 cm long, glabrous, stigma stomatomorphic.
Fruits and seeds not observed.

Phenology. f lowering throughout tin* year with
collections of mature flowers made in January, Feb-
ruary, March. May, and December.

Distribution (Fig. 5). Alloplectus serpens is

Volume 15, Number 1
2005

Clark et al.

Novae Gesneriaceae Neotropicarum XIV

77

Figure 4. Alloplectus serpens J. L Clark & L. E. Skog. — A. Habit. — B. Side view of flower. — C. Face view of flower.
— I). Corolla opened to show stamens. — E. Calyx opened and corolla removed to show bilobed nectary gland. (A— 1)
from ./. L. Clark, M. Thurber A I). Ranosa 5627, IJS; E from ./. /.. Clark, K Sanchez A /.. Jost 5900, US.)

78

Novon

85“ 80° 75° 70° 65°

Figure 5. Distribution of Alloplectus carpishensis, A. grandicalyx, A. inflatus, A. oblongicalyx, and A. serpens, bach
symbol represents collections from specific localities cited on herbarium collections. Maps were generated using
Are View CIS 3.2a (ESRI, 2000).

known from the eastern Andean slopes of Ecuador
and tht‘ southern Andes of Colombia from 1500 to
3200 m.

For the distinguishing characters of A. serpens,
see the discussion of A. carpishensis above.

Paratypes. COLOMBIA. Nnriiio: B. I\. La Planada,
7 km from Chueun^s. (). R enmities 8989 (MO. US). Pu¬
tumayo: edge of La Cocha Lagoon. Quebrada de Santa.
,/. Cuatrecasas 11827 (COL, b, US). ECUADOR. Carchi:
17.5 kin E of Santa Barbara, road Tulcan— Alegria via Car-
amelos & Santa Barbara, C. //. Dodson & A. //. Gentry
12124 (MO, QCNE, SLL). Morona-Santiago: Canton Li-
ui6n hidanza. hwy. Gualaeeo— Plan de Milagro. J. /,. Clark,
F Sanchez A' /.. Jost 5900 (HA, OCA. QCNE, US); L
slopes of Matanga Paramo, ca. 30—10 km S of Sigsig, road
tow. Cualaquiza, ./. /,. Luteyn & E. Cotton 1 1 191 (QCNE,
US); road Gualaceo— Limdn, km 35.5, I’. M. Jorgensen, C.
Ulloa X II. 0llgaard 92879 ( A A l . QCNL). IN apo: betw.
Santa Barbara and La Bonita, C. Darling & L. Andersson
12510 ((ill, SLL, US); Canton Arehidona. P. N. Sumaeo
Napo-Galeras, near Voledn Sumaeo crater, J. L. Clark
2264 (QCNL, US); Canton Ouijos, Parroquia Cuyuja, N
border of Res. bool. Antisana, betw. the town of Cuyuja

and the confluence of Rfo Tablon and Rfo Quijos, J. L.
Clark, M. Thurber X’ I ). Ranosa 5627 (QCA, QCNL, US);
Canton Sueumbios, Playbn de San brancisco, road Sta.
Bdrbara-La Bonita, ./. Jaramillo 9328 ((ill); Guagra Ureu,
small hill SL of summit. /.. Holm-Nielsen, ./. Jaramillo, F.
Coello X ' F. Azanza 27633 (AAll, MO). Sueumbios: b
slopes of Voledn Reventador, Res. bcol . Cayambe Coca,
,/. L. Clark 4428 (QCNL, US). Zaniora-Chinchipe: above
Valladolid, on road to Yangana, C. Ilarling X • /,. Andersson
21391 ((ill, US); LI Pangui. Cordillera del Condor, near
military base Cbndor Mirador, 7 Montenegro 154 (QCNL,
US).

Acknowledgments. Support for this project for
the first author was provided by the Elvin Mc¬
Donald Research Endowment Fund of the Ameri¬
can Gloxinia and Gesneriad Society, the National
Science Foundation (DEB 0206512), the Explorers
Club Washington Group, and a Fulbright Graduate
Study and Research Abroad Scholarship. We are
grateful to Cathy Pasquale-Johnson and Alice Tan-
gerini for preparing illustrations of the new species
and Tom Hollowell for help in preparing the dis-

Volume 15, Number 1
2005

Clark et al.

Novae Gesneriaceae Neotropicarum XIV

79

trilmtion map. Hamilton Beltran is also acknowl¬
edged for his guidance in the fieldwork component
in Peru. We are especially grateful to James Smith.
John Boggan. Christian Feuillet, and an anonymous
reviewer for their helpful comments on the manu¬
script and Harold Robinson for assisting with the
Latin diagnoses.

Literature Cited

Burtt, B. L. & H. Wiehler. 1995. Classification of the fam¬
ily Gesneriaceae. Gesneriana 1: 1-4.

Clark. J. L. & L. E. Skog. 2002. Novae Gesneriaceae Neo¬

tropicarum XI: Alloplectus grandicalyx, a new species
of Gesneriaceae from Ecuador. Novon 12: 173—178.

- & E. A. Zimmer. 2003. A preliminary phylogeny

of Alloplectus (Gesneriaceae): Implications for the evo¬
lution of flower resupination. Syst. Bot. 28: 365—375.

ESRI. 2000. ArcView G1S version 3.2a. Environmental
Systems Research Institute, Redlands, California, http:
//www.esri.com.

Hanstein, J. 1865. Die Gesneraceen des Koniglichen Her¬
bariums unrl der Garten zu Berlin, nebst monogra-
phischer Uebersicht der Eaindie im Ganzen, II.
Abschnitt. Gattungen und Arten. Drittes Stuck. Die Eu-
gesnereen, Rhytidophylleen und Beslerieen. Linnaea
34: 225-462.

Wiehler, H. 1973. One hundred transfers from Alloplectus
and Columnea (Gesneriaceae). Phytologia 27: 309-329.

New Species of Araeeae from Bolivia and I he Tropical Andes

Thomas B. Croat

R A. Schulze Curator of Botany, Missouri Botanical Garden, P.0. Box 299, St. Louis,
Missouri 63166-0299, IJ.S.A. thomas.croat@mohot.org

Amparo Acebey

Herbario INacional, La Paz, Bolivia; Universitilt Gottingen, Germany

Ahstract. A total of 16 taxa (15 species and I
variety) of Araeeae are new to science. These in¬
clude Anthurium acebeyae Croat, A. beckii Croat &
Acebey, A. besseae Croat, A. kunthii Poeppig var.
cylindricum Croat, A. ottobuchtienii Croat & Ace-
bey, A. stephanii Croat & Acebey, A. yungasense
Croat & Acebey, Diejfenbachia williamsii Croat,
Monstera kessleri Croat, Philodendron cotapatense
Croat & Acebey, P. kroemeri Croat, Rhodospatha
mukuntakia Croat, Stenospermation dictyoneurum
Croat & Acebey, S. killipii Croat & A. Gomez, Xan-
thosoma puberulum Croat, and X. seidelii Croat.

Kksijmkn. Un total de 16 taxones (15 especies y
I variedades) de Araeeae son nuevas para la cien-
cia. Estas incluyen Anthurium acebeyae Croat, A.
beckii Croat & Acebey, A. besseae Croat, A. kunthii
Poeppig var. cylindricum Croat, A. ottobuchtienii
Croat & Acebey, A. stephanii Croat & Acebey, A.
yungasense Croat & Acebey, Diejfenbachia william¬
sii Croat, Monstera kessleri Croat, Philodendron co¬
tapatense Croat & Acebey, P. kroemeri Croat, Rho¬
dospatha mukuntakia Croat, Stenospermation
dictyoneurum Croat & Acebey, .S', killipii Croat &
A. Gomez, Xanthosoma puberulum Croat, and X.
seidelii Croat.

Key words: Anthurium, Araeeae, Bolivia, Diej¬
fenbachia, floristics, Monstera, Philodendron, Rho¬
dospatha, Stenospermation, Xanthosoma.

In preparation of the Araeeae treatment for the
checklist of Bolivian plants being carried out at the
Missouri Botanical Carden, we have discovered a
number of taxa that arc new to science. A complete
treatment of the Araeeae of Bolivia, including pro¬
visional descriptions of many species believed to
be new but for which there is inadequate material
to describe, will eventually be available on the the
Missouri Botanical Garden web page: http://www.
mobot.org/MOBOT/Besearch/southamericaprojects.
shtml/bolivia. This will include a key to the species
of Diejfenbachia, Monstera, and Stenospermation for

Bolivia as well as full descriptions and complete
exsiccatae for species in other Araeeae genera.

Anthurium

Aiitliiiriiiin acebeyae Croat, sp. nov. TYPE: Bo¬
livia. La Paz: Nor Yungas, 14.3 km NE of
Chuspipata, 16°14'S, 67°47'W, 2000 m, 23
Mar. 1984, ./. Solomon, B. Stein & M. Uehling
12090 (holotype, MO-3213306; isotypes, k,
LPB, US). Figure I.

Planta epiphytiea; internodia 1—2 cm longa, 1.7 cm
diam.; catuphylla 11-15 cm longa, persistentia semi-in-
tacta; pctiolus 30—54 cm fungus, subteres; lamina trian-
gulari-sagillata, 41—54.5 cm longa, 23—28 cm lata; lobuli
postici I 1—16.5 cm longi, tt-10 cm lati; nervis primariis
lateralibus 8—10 utroque; pedunculus 23—39 cm longus;
spatha viridis, 8.5—15 cm longa, 1 .1—1.5 cm lata; spadix
7.5—22.5 cm longus, 4—9 mm diam., stipitatus 3—20 mm,
viridis.

Epiphyte; internodes 1—2 cm long, 1.7 cm diam.;
cataphylls I 1—15 cm long, dark brown persisting
semi-intact with apex remaining intact and pale fi¬
bers showing toward the base, sometimes forming
an almost completely fibrous network. Petioles 30—
54 cm long, about half as long as blades, subterete,
narrowly, obtusely and shallowly silicate, semiglos¬
sy, medium green, drying dark brown to blackened.
I,eaf blades triangular-sagittate, 41—54.5 X 23—28
cm, 1 .7— 2.2 times longer than wide, deeply lolled
at base, acuminate at apex, subcoriaceous, medium
to dark green and semiglossy above, slightly paler
and glossier below, drying dark green to dark brown
above, yellow-green to yellow-brown below; poste¬
rior lobes I 1—16.5 X ft — 10 cm, usually directed
downward, sometimes somewhat outward; anterior
lobes straight to broadly concave; basal veins 6 to
8, 1st pair free to base, 2nd pair sometimes free to
base, 4th and higher order pairs coalesced 5—6 cm;
posterior rib naked for 2.5— 5.5 cm; midrib narrow,
slightly paler and bluntly acute above, paler and
narrowly round-raised below, drying concolorous or
slightly darker than surface, sometimes acute

INovon 15: BO- 103. 2005.

Volume 15, Number 1
2005

Croat & Acebey

Araceae from Bolivia and Tropical Andes

81

Figure I. A-I). Anthurium acebeyae Croat (Croat et al. 84274). — A. Habit. — I?. Leaf blade adaxial surface. — C.
Close-up of inflorescence. — D. Stem with cataphylls and petiole bases.

82

Novon

above, darker or lighter than surfaee below; primary
lateral veins 8 to 10 pairs, arising at 30°-50° angle
to midrib, weakly quilted-sunken and eoncolorous
above, narrowly raised and slightly paler below; ter¬
tiary veins mostly not raised, drying darker than
surface. INFLORESCENCE erect; peduncle 23-39
cm long; spathe green, 8.5—15 X 1. 1—1.5 cm, re-
Ilexed-spreading; spadix 7.5—22.5 cm X 4-9 mm,
stipitate for 3-20 mm. medium to dark green, se-
miglossy. Flowers 1.8— 2.2 mm long, 7 to 9 visible
per spiral; lateral tepals 0. 9-1.0 mm wide, inner
margin rounded, outer margin 2-sided; anthers 0.4
X 0.6 mm; pollen white.

Distribution and habitat. Anthurium acebeyae
is endemic to Bolivia in La Paz (Nor Yungas, Sur
Yungas, Caranavi) and Cochabamba (Carrasco) De¬
partments at 1500—2300 in elevation in a Premon-
tane moist forest (P-mf) life zone (lloldridge et al.,
1971). The new species is a member of section lie-
lolonchium Schott emend. Engler, and is charac¬
terized by its epiphytic habit, short internodes, per¬
sistent. semi-intact cataphylls, triangular-sagittate
leaf blades with the margins usually concave, and
the long-pedunculate inflorescence with a stipitate
green spadix.

Anthurium acebeyae is most likely to be confused
with the widespread and variable A. incurvatum
Engler, which also occurs in the highlands of Bo¬
livia, but it is distinguished from that species by
its concave blade margins and green inflorescences
(the spathe with conspicuously convex blade mar¬
gins and the spadix purple to violet-purple in A.
incurvatum). file new species can also be confused
with A. yungasense Croat, which shares similar per¬
sistent cataphylls and stipitate green spadices, but
differs in having semiglossy to glossy blades that
are convex along the margins of the anterior lobes.
In addition, the spadix of A. acebeyae is usually
much more conspicuously stipitate (frequently by
1-2 cm).

The species is named in honor of Amparo Ace-
bey. Bolivian aroid specialist and coauthor of this
paper, who assisted in collecting the type specimen.

Paratypes. BOLIVIA. Cochabamba: Carrasco, Co¬
chabamba— Villa Tunari, km 141 on old rd., Kessler et al.
7730 (LPB, MO). La Paz: Caranavi, Serranfa Bella Vista,
37 km from Caranavi on the rd. to Sapecho, Kessler et al.
11345 (LPB, MO); Nor Yungas, Caranavi-Yucumo, 37.6
km NE of Caranavi, T. B. Croat et al. 34274 (B, COL,
LPB, MO. NY. USM); NK of La Paz, 5.4 W of rd. to
Coroico, Croat 51459 (LPB, MO); 16.5 km NE of Chus-
pipata, 13.6 km SW of Yolosa, ./. Solomon & Uehling
12166 (LPB, MO); Cotapata NP. T. Kroemer & Aeebey 979
(LPB); Est. Biol. Tunquini, senda del pajonal, atras del
Hfo Cedroni, K. Bach et al. 970 (LPB); Bajo Hornuni. trail
to the camp of Don Pedro, trail to the mine, I). Quintana

et al. 5 (LPB, MO). Bach et al. 970 (LPB, MO). Garcia et
al. 4391 (LPB, MO), 4431 (LPB, MO); Sud Yungas, lluan-
can£, 6.5 km S, on new rd., S. Beck 3058 (LPB. MO).

Antliiiriuiii beckii Croat & Aeebey, sp. nov.
TYPE: Bolivia. La Paz: Abel Iturralde, Tu-
mupasa-San Jos6 de Uchupiamonas, NW of
Tumupasa along slope leading up to Madidi
NP. 5.5— 5.8 km above jet. in main rd. near
Tumupasa, 15°45'N, 67°50'W, 830-850 m, 9
Aug. 2000, T. B. Croat, A. Aeebey & T. Kroemer
B43B7 (holotype, MO-5187101; isotypes, K.
LPB, NY, US). Figure 2A.

Planta epiphytica; internodia brevia, 1 cm diam.; ca-
taphylla 3.5 cm longa, semi-intacta, persistentia, fibris iu-
ferne; petiolus teres, 5-15 cm longus, 1.5 mm diam.; lam¬
ina oblongo-lanceolata, 25—36 cm longa, 1 .2—4.5 cm lata;
nervis primariis lateralibus 5—7 utroque; peduneulus
(13)26-36.5 cm longus, 1. 5-2.5 mm diam.; spatha lan-
ceolata, 4.3-5.5 cm longa, 5-7 mm lata, ex purpureo vir-
idis; spadix atropurpureus, 7—10.5 cm longus.

Epiphyte; internodes short, I cm diam.; cata¬
phylls 3.5 cm long, semi-intact, the fibers parallel,
promptly becoming fibrous with paler brown fibers.
Petioles terete, 5-15 cm long, 1.5 mm diam. on
drying, medium green, weakly glossy, drying light
yellow-brown. Leaf blades oblong-lanceolate, some¬
times weaklv falcate, 25—36 X 1.2— 4. 5 cm, 9—11
limes longer than wide, coriaceous, matte, acute to
very narrowly rounded at the base, narrowly acu¬
minate at apex, moderately bicolorous, dark green
above, moderately paler below, drying dark olive-
green to dark yellow-brown above, moderately paler
and yellow-brown below; midrib narrowly raised
and ± eoncolorous above, bluntly raised and paler
below1; primary lateral veins 5 to 7 pairs, scarcely
visible on fresh leaves, arising at 15°— 20° angle
from midrib, drying flat to deeply sunken above,
scarcely visible on lower surface; collective veins
arising from the base, drying weakly raised and
conspicuously more prominent than primary lateral
veins on lower surface, flattened to sunken on up¬
per surface; tertiary veins obscure. INFLORES¬
CENCE erect; peduncle (13)26—36.5 cm long, 1.5—
2.5 mm diam., terete, tinged purple; spathe spread¬
ing, lanceolate, 4. 3-5.5 cm long, 5-7 mm wide,
green, tinged purple to violet-purple, malic; spadix
dark violet-purple, matte, 7—10.5 cm long. Flowers
3 or 4 per spiral. 2.4— 2.6 X 1.8-2. 2 mm; lateral
tepals 1.6-1. 8 mm wide, the inner margin very
broadly rounded, outer margin 2-sided; pollen pale
orange. Berries orange, subglobose, 6 mm long. 4—
5 mm diam.

Distribution and habitat. Anthurium beckii is
endemic to Bolivia, where it is known only from the

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Croat & Acebey

Araceae from Bolivia and Tropical Andes

83

Figure 2. — A. Anthurium beckii Croul & Acebey (Croat et al. 84387). Habit. 15. C. Anthurium besseae Croat (Croat

71836). — B. Habit. — C. Close-up of leaf and inflorescence. — If. Anthurium kunthii var. cylindricum Croat (Hesse el
al. 1790). Herbarium specimen.

84

No von

mountains of northeastern La Paz Department in
t he provinces of Abel llurrahle. J. Bautista Saave¬
dra. Franz Tamayo, Sud Yungas, and Caranavi at
600-1550 m.

The new species is a member of section Porphy-
rochitonium Schott and is superficially close to An-
thurium friedrichsthalii Schott, which ranges from
Costa Rica to western Colombia, but is not found
in the Amazon basin. Ihe latter species differs in
being a pendent epiphyte with proportionately lon¬
ger leaves (to 14—16 times longer than wide) with
the upper blade surface epunctate and in having a
spadix that is green at anthesis (turning brownish
in age). In contrast, A. beckii is a more or less erect
herb, with leaf blades no more than I I times longer
than wide and glandular-punctate on both surfaces,
and lias a spadix that is dark purple at anthesis.

Paratypes. BOLIVIA. La Paz: Caranavi, Serrama
Bella Vista, Caranavi— Sapecho, km 47. P. Kessler el al.

1 1661 (l.l’B, MO): Saavedra, Pauji-Vuyo, km 3, Apolo-
Charazani, Kessler et al. 9732 (LPB, IMO); Sud Yungas,
Alto Beni, Sapecho, Colonia 16 de Julio, subiendo la sen-
da hacia la antena de ENTEL, Kroemer & Acebey 711
(LPB, MO); Tamayo, Apolo-Charazani, km 52. bey. Cor-
reo, Campamento Calabatea, S. Heck R. hosier IH577
(LPB); Campamento Alto Liinoti, Limdn— Mara, trail San
Jos6— A polo, Helme 556 (LPB. MO).

Aiithiiriiiiii besseae Croat, sp. nov. TY PL: Boliv¬
ia. Cochabamba: Chapare, old rd. from Cocha¬
bamba to Villa Tunari, vie. Villa Tunari, 240
m. cultivated at Selby Botanical Gardens (orig¬
inally collected live by Libby Hesse , 25 Jan.
1986), 29 Sep. 1996, T. B. Croat 79380 (ho-
lotype, MO-3301223; isotypes, B. k, LPB, NY.
SKL. US). Figure 2B. C.

Planta terrestris; caudex fasciculatus, brevis; cataphylla
ad 5.5 cm longa, persistent ia intaeta versus apicem, fibris
inferne; petiolus plus minusve teres, (10)19—33 cm lon-
gus; lamina ovato-cordata vel anguste ovato-cordata, 13—
33 cm longa, 8-23 cm lata; nervis primariis lateralibus
2-4 utroque; pedunculus (4)9-25(36.5) cm longus; spatha
1.3— 5.5 cm longa, 2—8 mm lata, viridis; spadix 1.2— 7.3
cm longus, 5-7 mm diatn., atropurpureus.

Terrestrial; stems densely clustered and short;
internodes short, ca. 1—2.5 cm diam.; cataphylls to
5.5 cm long, drying dark brown, persisting semi¬
intact in part, the remainder fibrous. Petioles
(10)19—33 cm long, erect-spreading, ± terete,
green, tinged red with white spots in lower half,
matte; geniculum 2.5 cm long, 3.5 mm diam.
LEAVES semi-erect with the blades ± pendent
from semi-erect petioles. Leaf blades ovate-cordate
to narrowly ovate-cordate, 13—33 X 8—23 cm, sub-
coriaceous, deeply cordate at base, acuminate at
apex, medium dark green and velvety above, much

paler to moderately paler and matte below, drying
matte on both surfaces, undulate to erenulate along
the margins; anterior lobe 13—26 cm long, the mar¬
gins broadly rounded; posterior lobes 3.5— 8.5 cm
long, rounded; midrib and basal veins convex to
sharply raised to knife-edged and paler above, mar¬
ginally somewhat paler along the major veins
above, drying paler than surface both above and
below; midrib almost Hat and paler to darker below;
primary lateral veins 2 to 4 pairs, arising at 55°—
65° angle, weakly raised to convex and much paler
than surface above, weakly raised and darker be¬
low; tertiary veins weakly visible below, drying un¬
dulate; basal veins 3(4) pairs, all free to the base,
weakly raised above, weakly raised and green be¬
low; sinus narrowly V-shaped to oblong or closed,
3-5 cm deep, lo 8 mm diam. INFLORESCENCE
erect-spreading, usually held somewhat above
leaves; peduncle (4)9-25(36.5) cm long, 2-3 mm
diam., maroon to weakly purplish-spotted; spathe
1.3— 5.5 cm long, 2—8 mm wide, medium green,
tinged maroon to purple on inner surface, minutely
warty, initially erect, becoming reflexed and re¬
curled post-anthesis; spadix 1.2-7. 3 cm long, 5-7
nun diam., curved forward, dark purple to purplish
violet, cylindroid to slightly tapered; tepals matte.
Flowers 2—2.2 mm wide and long, 5 per spiral;
sides straight, parallel to spiral: tepals matte, lat¬
eral tepals 1.3-1. 6 mm w ide, weakly pale-punctate;
inner margins rounded; stamens weakly exserted at
anthesis, closely aggregated; pollen orange; pistils
purple, sometimes exserted. Berries not seen.

Distribution and habitat. Anthurium besseae is
endemic to the type locality in Bolivia in the De¬
partment of Cochabamba, Province of Chapare, at
240 m.

The new species is a member ol Anthurium sect.
Cardiolonchium Schott and is distinguished by its
short stem, short internodes, subterete petioles, and
ovate-cordate, velvety blades with a narrow sinus
or w ith the posterior lobes actually overlapping and
with the major veins raised and much paler on the
upper surface. Also characteristic is the purple spa¬
dix with slightly exserted stamens. It is superficially
similar to A. regale Linden, from central Peru, in
having velvety leaves with pale veins on the upper
surface. That species typically has blades many
times longer, ranging from 35 cm to about 100 cm
long. Anthurium regale also differs in having the
basal veins mostly fused into a posterior rib rather
than free to the base as in A. besseae. In addition,
A. regale has a creamy white rather than dark pur¬
ple spadix.

I he species is named in honor of Libby Besse,

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Croat & Acebey

Araceae from Bolivia and Tropical Andes

85

long-time benefactor. Board member, anti volunteer
staff member of the Marie Selby Botanical Gardens
in Sarasota, Florida, who sponsored many expedi¬
tions to Latin America, especially in Ecuador, and
collected the live plant from which the type spec¬
imens were prepared.

Paratypes. BOLIVIA. Cochabamba: Chapare, along
old rd. from Cochabamba to Villa Tunari, vie. Villa Tunari,
Christenson 1134 (SEL). Ingram 972 (MO, SEL). Ingram
& Atwaixl 1029 (MO. SEL), Croat 71H30 (MO). All para¬
types made from cultivated plant at Selby Botanical Gar¬
dens 82—85, originally collected as Libby Besse el al.

Anlluiriiim kiinthii Poeppig, Nov. Gen. Sp. PI. 3:
84-85. 1845. TYPE: Peru. Loreto: Maynas,
Poeppig s.n. (holotype, W destroyed).
Aiitliiiriinn kiinthii Poeppig var. cylimlricum

Croat, var. nov. TYPE: Bolivia. La Pa/: Lare-
caja, Caranavi— ' Tipuani, km 60, 550 m, 24—26
Jan. 1983, /.. Besse, C. & ./. Luer & R. Vdsquez
1790 (holotype, SEL-04732). Figure 21).

Ab Anthurio kiinthii var. kunthii spadice clavato. lamina
oblanceolata angustius et nervis primariis lateralibus de-
b i 1 1 bus (Illicit.

Hemiepiphytie climber; internodes 6—9 cm long,
1 cm diam., drying moderately smooth, pale yellow-
brown: cataphylls narrowly triangular, pale yellow-
brown. persisting ± intact at upper nodes. Petioles
subterete, 49 cm long, drying to ca. 3 mm diam.,
pale yellow-green. Leaf blades narrowly oblaneeo-
late, 24—27.5 X 4.2— 4.5 cm, gradually long-acu¬
minate at apex, attenuate at base, drying dark yel¬
lowish brown and matte above, moderately paler,
medium yellowish gray-brown and weakly glossy
below; midrib drying narrowly raised to acute and
concolorous above, thicker than broad and slightly
paler than surface below, the midrib often with a
weak medial sulcus; petiolules 2.5— 3.5 cm long, ca.
1.5 mm diam.; primary lateral veins arising at 55°—
65° angle, not apparent on upper surface, some¬
times visible below, ca. 6 pairs, generally scarcely
more apparent than interprimary veins. INFLO¬
RESCENCE erect; peduncle 32 cm long, drying 2.5
mm diam.; spathe green, 4 cm X 7 mm; spadix
green, weakly elavate, 4.5 cm long, 8 mm diam. on
drying, rounded at apex. Flowers 3.0— 3.2 mm wide
and long, 5 to 6 visible per spiral; lateral tepals
1.8 mm wide, the inner margin very broadly round¬
ed, outer margin 2-sided.

Distribution and habitat. Ant Iturinm kunthii
var. cylindricum is endemic to Bolivia, where it is
known only from the type locality in La Baz De¬
partment, Province of Larecaja.

The new variety differs from the typical variety

in having blades that are more narrowly oblanceo-
late (vs. usually more nearly elliptic), with obscure
or very weak primary lateral veins (vs. typically
moderately obvious), and a elavate spadix (vs. lon¬
ger and narrowly tapered).

Anlliurium ottobuchtienii Croat & Acebey, sp.
nov. TYPE: Bolivia. La Paz: Murillo, 44 km
below Lago Zongo dam, vie. of Cahua hydro¬
electric plant, 16°03'S, 68°01'W, 1200 m, 12—
15 Sep. 1983, J. Solomon 10749 (holotype,
MO-3158417; isotypes, B. k. LPB, NY. US,
USM). Figure 3 A, B.

I’lanta terrestris; cataphylla (5.5)10—13.5 cm longa, pro
parte maxima decidua; petiolus (11)11—28 cm longus,
subteres; lamina (12)17—44 cm longa, (3)4— 8.0 cm lata,
plcmmque oblongo-lanceolata vel anguste oblongo-oblan-
ceolata; nervis primariis lateralibus 10—14 utroque; pe-
dunculus (6)17—25 cm longus; spatlia lanceolata,
(2. 3)5. 5— 6.5 cm longa, 7—10 mm lata; spadix atroviolaceo-
atropurpureus, (3)4.5— 8 cm longus.

Terrestrial on steep bank; stem 30—100 cm long;
internodes (0.5) 1.0—1. 5(2.5) X 1.0— 1.8 cm, medium
green and weakly glossy, soon gray-green to gray-
brown. semiglossy, often longitudinally and narrow¬
ly ribbed, sometimes transversely cracked; cata-
phylls (5.5)10—13.5 cm long, medium green, soon
dead except parts of base and thin pale fibers at
upper nodes, sometimes turning brown and per¬
sisting intact at upper nodes, finally weathering to
pale fibers with fragments ol epidermis ol the base
of the cataphylls, finally completely deciduous. Pet¬
ioles (11)14—28 cm long, drying 1—2 mm wide, sub-
terete, oval, narrowly and sharply to obtusely flat¬
tened, sometimes weakly flattened, sometimes
weakly and narrowly sulcate to deeply sulcate,
sometimes sharply flattened with weak medial rib,
medium green, weakly glossy. Leaf blades (12)1 7 —
44 X (3)4— 8.0 cm. (2. 7)3.4— 8 times longer than
wide, oblong-lanceolate to narrowly oblong-oblan-
eeolate, sometimes ovate-elliptic to narrowly ellip¬
tic, 1 .3—2.5 times longer than petioles, narrowly
acuminate at apex, abruptly acute to obtuse al base
(sometimes inequilateral), subcoriaceous, semi¬
glossy, slightly bicolorous, medium to dark green
above, moderately paler below, drying dark green
to dark reddish brown to yellow-brown above, mod¬
erately paler and yellow-green to reddish brown be¬
low; midrib acute to narrowly raised and coneolo-
rous above, often in deep valleys above, narrowly
raised and slightly paler below, drying acute and
concolorous above, paler than surface, round-raised
and brown to reddish brown, sometimes finely cos¬
tate below; primary lateral veins 10 to 14 pairs,
arising at 40°-60°(70°) angle, weakly quilted-sunk-

86

Novon

Figure 3. A, B. Anthurium oltobiu'htienii Croat & Acebey (Croat ct al. 84733). — A. Habit. — B. Close-up showing
inflorescence at anthesis and young infructescence. — C. Anthurium stephanii Croat & Acebey (Beck 3109). Herbarium
specimen. — I). Anthurium yungasen.se Croat & Acebey (Croat et al. 84264). Habit.

Volume 15, Number 1
2005

Croat & Acebey

Araceae from Bolivia and Tropical Andes

87

en ami concolorous above, usually about as prom¬
inent as the collective veins above, narrowly raised
and concolorous below, scarcely more prominent
than the interprimary veins; collective veins arising
from one of the lowermost primary lateral veins
(sometimes from near the middle of the blade), or
from the 1st basal veins, 1—7 mm from margin,
loop-connecting the primary lateral veins. INFLO¬
RESCENCE erect to semi-erect; peduncle (6)17-25
cm long, less than 3 mm diam. on drying; spathe
lanceolate, spreading, (2. 3)5. 5-6.5 cm X 7-10 mm,
light to medium green, margin purple; spadix
(3)4.5— 8 cm long, dark purple- violet, erect to
slightly curved, matte. Flowers ca. 3 visible per spi¬
ral, 2.5—4 mm long; lateral tepals shield-shaped,
1.6— 2.4 mm wide, the inner margin broadly round¬
ed, the outer margins 2-sided; pistils early-emer¬
gent. INFRUCTESCENCE to 14 cm long, nodding;
berries subglobose, ca. 4 mm diam., dark violet-
purple.

Distribution and habitat. Anthurium ottobuch-
tienii is endemic to Bolivia in the departments of
l.a Pa/ (Nor Yungas, Sud Yungas, Caranavi, and
Murillo) and Cochabamba (Prov. Carrasco), at
I 100-2650 m.

The new species is a member of section Xialo-
phyllium Schott, and is characterized by its gen¬
erally terrestrial habit; cataphylls persistent, intact
only at upper nodes; frequently long, narrow leaf
blades; and purple-violet spadix. It is also char¬
acterized by having blades that dry dark green with
tin1 primary lateral veins and the collective veins
deeply sunken in “valleys” on drying. The species
tends to have proportionately broader leaf blades at
higher elevations (2400—2650 m).

The new species is perhaps most easily confused
with A. mindense Sodiro, of Ecuador and northern
Peru, but the latter differs in having longer and
proportionately more slender internodcs and gen¬
erally proportionately broader leal blades. Anthur¬
ium ottobuchtienii is also similar to A. microspadix
Schott, but that species has proportionately wider
blades (less than 6 times longer than wide) that dry
green rather than brown and has a green rather than
purple spadix.

A few collections have leaf blades that are short¬
er than normal, ranging between 2.9 and 3.5 times
longer than wide. These also appear to belong here,
but they are at the extreme in terms of size and
shape and differ from typical A. ottobuchtienii in
the color and venation of the dried leaf blades.

The species is named in honor of Otto Buchtien,
a German who made many collections in Bolivia

during the early 20th century and who made the
first collection of the species.

Paratypes. BOLIVIA. Prov. Unknown: Cordillera
Heal. Nequejahuira, /. Tate 633 (MO. NY). Cochabam¬
ba: Carrasco, Cochabamba-Villa Tunari on old rd., km
I 18. Kessler el al. 7 WO (LPB, MO), km 130. H. Kessler et
<d. 7153 (I, I’ll. MO). Fa Pa/,: Murillo, Valle de Zongo, T.
13. Croat 51426 (LPB. MO); 32.8 km beyond the summit.
300 m S of Planta Hidroeleetrica Chururaqui, Solomon et
(d. 19062 (1.1*8, MO); 30.5 km N |below] dam at Lago
Zongo, trail up Rio Jachcha Cruz, ,/. Solomon 9098 (LPB,
MO); 44 kins despu^s del alira de Zongo. S. Beck 7245
(LPB); Nor Yungas, Polo-polo hacia Coroico, 0. Buchtien
3667 (NY, US); 9.5 km NE below Chuspipata, 20.2 km S
of Yolosa, Solomon 8103 (COL, LPB, MO); Serranfa de
Bella Vista, 16 km N of Carrasco, 37 km N of Caranavi
on rd. to Palos Blancos, Solomon & Nee 12716 (MO);
Unduavi— Caranavi, 34.8 km E of Unduavi, 16.3 km SW
of Yolosa, Croat 51575 (MO): Cotapata NP. Est. Biol., Tun-
quini, vie. of Chairo, NW of Coroico, 23 km W of Yolosa,
Croat et al. 84754 (LPB. MO, W); Senda del Pajonal.
Bach 973 (LPB, MO), E. Garda et al. 4434 (LPB, MO);
trail to Campo de Don Pedro, trail to the mine, Bach 443
(LPB, MO); trail to Homundi, Palahral et <d. 4 (LPB).
Palabral et al. 46 (LPB); Sacramento, Chuspipata— Yolosa,
4. Gentry & Solomon 52045 (F, Oil. LPB. MO); 8 km de
Chuspipata hacia Coroico, M. Kessler et al. 12049 (LPB.
MO); Serranfa Bella Vista, Rurrenabaque— Caranavi, 28.4
km SW of middle of bridge over Rfo Beni near Sapecho,
21.4 km SW of Rio Piquendo, Croat et id. 84733 (LPB,
MO); Caranavi hacia Sapecho, km 37. Kessler et al. 1 1349
(I.BP, MO); Hacienda Simaco sobre el camino a Tipuani,
Buchtien 5345 (GH, NY); Casaniacamino a Tipuani.
Buchtien 7172 ( US).

Aiilliuriiini slcpliunii Croat cN Acebey, sp. nov.

TYPE: Bolivia. La I’az: Nor Yungas, Est. Biol.
Tunquini, 16°12'S, 67°51'W, 1700 m. 3 Apr.

2000, T. Kroemer & A. Acebey 9 94 (holotype,
MO -56502 50; isotypes, B, F, K. LPB. NY. US,
USM, VEN). Figure 3C.

Planta hemiepiphytica; internodia brevia, minus quam
l cm in sicco; cataphylla 5—10 cm longa, persistentia;
petiolus teres, 16.5—24 cm longus; lamina oblongo-lan-
ceolata vel oblanceolata, 28—38.5 cm longa, 7.2-9. 1 cm
lata; nervis primariis lateral i bus 13—15 utroque; pedun-
culus 20 cm longus; spatha lanceolata, viridis, 7.5—12 cm
longa, 9—12 mm lata, spadix 10.3—12 cm longus, 3 mm
diam. in sicco, stipitatus 6 mm.

Hemiepiphytic vine; internodes very short near
the apex, drying less than 1 cm diam., dark reddish
brown; cataphylls 5—10 em long, slender, persistent
along with the petiole bases. Petioles terete, 16.5-
24 em long, sheathed 35%^10% their length, dry¬
ing dark yellow-brown; geniculum 1.5— 2.0 cm long,
darker, terete. Leaf blades oblong-lanceolate to ob-
lanceolate. 28—38.5 X 7.2— 9. 1 cm wide. 3.8— 4.6
times longer than wide, broadest at middle or above
middle, narrowly acuminate at apex, acute at base,
semiglossy on both surfaces, dark green above,

88

Novon

slightly paler below, drying medium to dark green
above, reddish brown to yellow-brown above, nar¬
rowly raised and concolorous to weakly paler
above, drying ± aeute toward the middle of the
blade above, narrowly convex and darker than sur¬
face below; primary lateral veins 13 to 15 pairs,
weak and scarcely visible on both surfaces; collec¬
tive veins arising from the base, 7-10 mm from
margin, scarcely raised and concolorous on upper
surface, weakly raised on lower surface, weakly
loop-connecting the primary lateral veins; tertiary
veins obscure. INFLORESCENCE erect-spreading;
peduncle 20 cm long, drying 2 mm diam.; spathe
lanceolate, green, 7.5-12 X 9-12 mm; spadix
10.3—12 cm long, drying 3 mm diam., stipitate 0
mm. Elowers 2.3— 3.7 mm long, 3 flowers visible per
spiral; lateral tepals 1.7 mm wide, inner margin
rounded, outer margin 2-sided. Berries unknown.

Distribution and lud>itat. Anthurium stephanii
is endemic to Bolivia in Ea Paz Department at
1900—2100 in elevation.

The new species is an unusual member of An¬
thurium sect. Calomystrium Croat, characterized by
its slender stems, persistent, lanceolate, intact cat-
aphylls, and lanceolate dark-brown-drying blades.

This species is different from its close congeners
because there are no other species of section Cal¬
omystrium with lanceolate leaf blades.

file species is named in honor of Stephan G.
Beck, noted German botanist who has spent most
of his professional career, from the 1980s to the
present, in Bolivia.

Paratypes. BOLIVIA. La Paz: Franz Tamayo, Madidi
NP. I’inalito, 30 km E of Apolo on trail to San Jose de
Ucfiupiamonas, Puentes el <d. 4688 (LPB, MO. IIS); Nor
Yungas, Kst. Biol. Tunquini, trail to Campo Don Pedro, P.
Garcia el td. 4888 (LPB. MO), Barb el id. 66 8 (I.PB, MO),
Quintana et al. 8 (LPB, MO); Sud Yungas, Htianeane. 7.5
km to the S over a new rd., S. G. Beck 8109 (LPB).

Aiithiiriimi yungasense Croat & Acebey. sp. nov.
TV PE: Bolivia. La Paz: Nor Yungas, Anmi Co-
tapata NP, Est. Biol. Tunquini, NW ol Coroico,
NNE of La Paz, vie. ol El C.hairo. 23 km W
of Yolosa. 16°12'S, 67°50'W, 1300-1500 m.
21 Aug. 2000, T. B. Croat. A. Acebey & T.
Kroemer B47B2 (holotype, MO-5185105; iso¬
types, B. COL, GOET, K. LPB, NY, QCNE,
LIB. US. USM. VEN). Figures 3D, 4A.

Planta terrestris; calaphylla 10—16 cm longa, pler-
umque intacta; petiofus 51—56 cm longus, suhleres, leni-
tcr eomplanatus; lamina anguste ovata, 49—60(65) cm lon¬
ga. 23—29(35) cm lata, profunde sagiltata; lobi postici
(11)13—16(18) cm longi. (8)10—10.5(12.5) cm lati: nervis
hasalihus 7—10 utroque; nervis lateral i bus 4— 6(7) utroque;

pedunculiis 32—38 cm longus; spatlia 6.5—13 cm longa,
J5— 15 mm diam.; spadix atroviridis vel olivaccus. stipitatus
4—6 mm, 6.5—13.5 cm longus, 12 mm diam.

Terrestrial to ca. 1 m tall; internodes short or lon¬
ger than broad, 1-6 X (1.5)2. 0-2. 7(4. 5) cm, gray-
green and ± scurfy; cataphylls 10—16 cm long,
turning brown, intact or often merely as pale fibers
toward the base of the cataphylls at upper nodes,
these thin and appressed to the stem. Petioles 51 —
56 cm long, subterete to oval in cross section, ob¬
tusely flattened adaxially, medium green, semiglossy,
unmarked or sometimes short-lineate; geniculum

2.5 cm X 7 mm. slightly thicker than petioles. Leaf
blades narrowly ovate, 49—60(65) cm long, 23-
29(35) cm wide. 1.7-2. 1 times longer than wide,
1.0-1. 2 times longer than petioles, subcoriaceous,
dark green and matte-subvelvety, sometimes faintly
pale-speckled above, slightly to moderately paler
and matte below, drying matte, gray-green above,
moderately paler and yellowish green below, deeply
sagittate-lobed at base, briefly short-acuminate at
apex; posterior lobes (11)13—16(18) cm long,
(8)10—10.5(12.5) cm wide, directed downward and
curved inward; anterior lobes (21)40—42(65) cm
long, usually broadly rounded along margins, some¬
times almost straight, rarely weakly concave; sinus
hippocrepiform to obovate (sometimes closed), 10—
12(16) cm deep, 3— 5.5(8) cm wide; basal veins 7
to 10 pairs, 1st pair free to the base, 2nd pair al¬
most free. 5th anil higher order pairs coalesced 5—

5.5 cm; posterior rib naked 2—4.5 cm along the
sinus; midrib acutely triangular to narrowly convex
(more acute toward the apex), and slightly paler to
concolorous, slightly paler to moderately paler and
densely pale-speckled above, narrowly rounded to
bluntly acute and paler to moderately paler below,
drying yellow-brown to dark brown below; primary
Intend veins 4 to 6(7) pairs, scarcely more conspic¬
uous than the interprimary veins, arising at 40°-
65° angle, narrowly raised and bluntly acute, slight¬
ly paler to concolorous above, narrowly raised and
acute to bluntly acute, slightly paler below, usually
drying yellow-brown and darker than surface or yel¬
lowish and paler than surface below, arching prom¬
inently upward along the margin but often not form¬
ing a collective vein throughout most of the length
of the blade; often w ith the lowermost pair of lateral
veins smaller than primary laterals, forming well
above the petiole and spreading at a broad angle
to merge with the uppermost (1st) basal vein; col¬
lective veins lacking or arising from one of the up¬
permost primary lateral veins. INFLORESCENCE
erect; peduncles 32—38 cm long (to 49 cm long in
fruit), terete, semiglossy, medium to dark green;
spathe 6.5—13 cm X 8—15 nun, reflexed-spreading,

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Croat & Acebey

Araceae from Bolivia and Tropical Andes

89

Figure 4. — A. Anthurium yungaseme Croat & Acebey (Croat A- Acebey 84264). Close-up of inflorescence. — B.

Diejfenbachia williamsii Croat (Croat et at. 84485). Side view of whole plant showing stem, leaf, and inflorescence. C,
1). Monstera kessleri Croat ( Kessler 8479). — C. Type specimen with adult leaves, sheet 1. — I). Type specimen with
pre-adult leaves, sheet 2.

90

Novon

under-rolled and sharply folded under twice, cau¬
date-acuminate at apex, the margins rolled under;
spadix dark green to olive-green, matte, stipitate 4-
6 mm (6 mm diam.), 6.5-13.5 cm long, 12 mm
diam. at base, 1 1 mm diam. midway, 8 mm diam.

1 cm from apex (drying 4—6 mm diam.). f lowers
1. 9-2.0 mm long, 2. 0-2. 2 mm wide, 7 to 9 visible
per spiral; tepals 1—1.6 mm wide, broadly rounded
to almost straight on the inner margin, 2-sided on
inside margin; stamens 0.4 mm long, 0.5 mm wide;
pollen pale yellow. INFRUCTESCENCE to 16—36
cm long, 2 cm diam.; berries purple toward apex,
white below middle.

Distribution and habitat. Anthurium yunga-
sense is endemic to Bolivia, where it is known prin¬
cipally from Nor and Slid Yungas (hence the name).
It is also known from the provinces of Caranavi,
Murillo, Muneeas. and Saavedra in La Paz Depart¬
ment and Chapare Province in Cochabamba De¬
partment. at 1200-1700 m elevation.

The new species is characterized by its longer
than wide internodes, cataphylls persistent intact
or fibrous, subterete petioles, and narrowly ovate,
matte-subvelvety, usually grayish-drying leaf
blades with mostly free-ending primary lateral
veins. Also characteristic is the stipitate, green spa¬
dix.

The new species is a member of Anthurium sect.
Cardiolonchium Schott and is most easily confused
w ith A. incurvation, which has leaf blades of similar
size and shape and a stipitate spadix. The latter
species differs in having proportionally shorter and
broader internodes. blades that are semiglossy, not
matte-subvelvety above, typically drying yellow-
green. and (lowering spadices that are purple to
purple-violet, not green. Though dried specimens
of the two species are more difficult to separate,
careful observation at high magnification shows the
upper leaf blade surface of A. incurvation appearing
minutely granular, whereas that of A. yungasense is
alveolate with the center of each epidermal cell
usually sunken.

Paratypes. BOLIVIA. Cochabamba: Chapare, Co¬
chabamba— Villa Tunari, Croat 51258 (LPB, MO); vie. Ho¬
tel Caballeros, km 94, Croat 51268 (LPB, MO). La Paz:
Caranavi, Serranfa Bella Vista, Caranavi-Sapeeho, 44 km
from Caranavi, Kessler el al. 11580 (LPB, MO); Cultural
Unidos, 36 km de Caranavi hacia Sapecho, Kessler et al.
1 1684 (LPB, MO); Murillo. Valle de Zongo, 13.2 km above
the kahua Power Plant, Croat 51444 (LPB, MO); 45.5 km
below lh«> dam at Lago Zongo, vie. of Cahua hydroelectric
plant, Solomon 12655 (MO); Muneeas, Hfo San Cristdbal
near Consata, Hesse et al. 562 (SEL); Nor Yungas, Hach
et al. 1174 (LPB, MO); Cotapata NP, Kroemer & Acebey
680 (LPB. MO); 4.6 km below Yolosa. then 19.1 km on
rd. up Kfo Huarinilla, Solomon 8802 (LPB, MO); 8.7 km

below Chuspipata on rd. to Yolosa, Solomon 6208 (I, I’ll,
MO); 13.7 km on rd. up Rfo Huarinilla, Solomon 6417
(LPB, MO); Coroico- Yolosa, 10 km up Rfo San Juan, Heck
7552 (LPB. MO); Serranfa Bella Vista, Caranavi-Yueumo,
34.3 km NE of Caranavi, Croat et al. 84264 (LPB. MO);
Unduavi— Caranavi, 34.8 km E of Unduavi, 16.3 km SW'
of Yolosa, Croat 51572 (LPB, MO); Sur Yungas, Heck
24880 (LPB, MO); 7-9.5 km NE of (above) Huanean£,
Luteyn & Dorr 12741 (NY), Heck 2056 (LPB, MO). Heck
2122 (LPB, MO); Saavedra, Charazani-Apolo, km 15, Kes¬
sler et al. 10456 (LPB, MO); Cerro Asunta Pata, entre
Apolo y Charazani, Kessler et <d. 10204 (LPB, MO).

DlKFFENBACUIA

Dielleiibaeliiu williamsii Croat, sp. nov. I V PE:
Bolivia. La Paz: Abel Iturralde, Tumupasa-
San Jose Uchupiamonas, NW ol Tumupasa,
along slope leading up to Madidi NP, 5.5— 5.8
km above jet. to San Jos£ near Tumupasa,
15°45'S, 67°50'W, 830-850 m, 9 Aug. 2000,
T. II. Croat , A. Acebey & T. Kroemer H43H5
(holotype, MO-5187105; isotypes, 8. K, LPB,
MO, US). Figure 4B.

Planta 30-90 cm alta; internodia 2-3 cm longa, 1.5-
1.7 cm diam.; petiolus 1 1-17 cm longus; lamina ovata vel
ovato-elliptica, 12—20(26) cm longa, 4.5—10(13) cm lata;
nervis primariis lateralibus 7-1 1 utroque; pedunculus 4—
6.5 cm longus, 6 X 4.5 mm latus; spatha 11 — 16 cm longa,
1-1.7 cm lata; spadix 9.5-11.5 cm longus; pars feminea
5.8-7 cm longa, pars masculina eontingens.

Small herb, 30-90 cm tall: internodes 2-3 cm
long, 1.5— 1.7 cm diam., semiglossy to weakly
glossy, drying matte, usually yellowish green, some¬
times medium to dark orange-brown. Petioles I I —
17 cm long, sheathed 0.3-0. 6 their length, medium
green, semiglossy, smooth, drying matte, medium
yellow-green; sheath acute to narrowly rounded at
apex; free part sharply and broadly silicate, drying
narrowly sulcate. Leaf blades narrowly ovate to
ovate-elliptic, 12—20(26) X 4.5—10(13) cm,
(1 .6)2. 0-3. 0(3.6) times longer than wide. 1.2 times
longer than petioles, weakly inequilateral, one side
5—13 mm wider, inequilateral at base, gradually
acuminate at apex, one side acute to obtuse, the
other rounded or subcordate, dark green and weak¬
ly glossy above, moderately paler and matte below;
midrib flattened-convex and eoncolorous above,
narrowly rounded and slightly paler below, drying
slightly raised and eoncolorous to slightly darker,
often orange-brown below; primary lateral veins t
to I 1 pairs, arising at 40°— 50° near apex. 50°-60°
near base, weakly quilted-sunken above, weakly
convex-pleated below, drying only weakly visible
above, slightly raised and darker than surface be¬
low; minor veins few. moderately obscure. INFLO¬
RESCENCE 1 per axil; peduncle 4—6.5 cm X 6-

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Croat & Acebey

Araceae from Bolivia and Tropical Andes

91

4.5 mm at midpoint, pale green, glossy; spathe 11—
16 X 1 — 1.7 cm, pale yellow-green, matte outside,
slightly paler and glossy inside; spadix 9.5—1 1 .5 cm
long, the pistillate portion 5.8-7 cm X 9 mm at
base, 6 mm diam. at I cm from apex, contiguous
with the staminate portion; pistils 38 to 41. sparsely
arranged with no more than 3 across the axis; sta-
minodia 3 to 4. not fused at base.

Distribution and habitat. Dieffenbachia wil-
liamsii ranges from Peru (Loreto) to Bolivia at 200-
1300 m elevation. In Bolivia it is known from the
departments of Beni (Ballivian, Yacuma), Cocha¬
bamba (Carrasco, Chapare), La Paz (Iturralde,
Franz Tamayo, Nor Yungas), and Santa Cruz (leh-
ilo).

Dieffenbachia williamsii is characterized by its
small, mostly yellow-green-drying, ovate to ovate-
elliptic blades, petioles that dry sharply sulcate,
and especially by the spadix, which lacks a sterile
area between the staminate and pistillate portions.

I lie first collection of the species was made by
R. S. Williams on 28 November 1901 at San Bue¬
naventura in La Paz (across the river from Rurren-
abaque), anil the species is named in his honor.

Paratypes. PERU. Loreto: Coronel Portillo, Arbore¬
tum von Humboldt. Carretera Federico Basadra, km 99.
Diaz et al. 685 (MO). BOLIN IA. Beni: Ballivian. 12 km
by lumber trail. SW of km 12 on Yucumo— Rurrenabaque
Rd.. Kessler et al. 10795 (LPB, MO); Yuc •umo— Rurrena¬
baque, Eseuela Tecniea Agropecuaria Rio Colorado, 35
km N of Yucumo, Killeen et al. 2881 (LPB); Ballivian and
Yacuma, Esl. Biol. Beni, Cerro Ocho, Guareco 85 (LPB,
MO); Yacuma, Chiman 8. Acebey 48 (LPB, MO). Cocha¬
bamba: Chapare, Carrasco NP, Cavernas de Repechon,
Kessler et al. 8315 (LPB, MO); Par. Machia, I km E of
Villa Tunari, Kessler et al. 8472 (LPB. MO); 1.5 km 5 K of
El Palmar on rd. to Avispas, Kessler et id. 8184 (LPB,
MO); El Palmar, km 155 on old Cochabamba— Villa Tunari
Rd., Kessler et al. 8118 (LPB. MO); km 159. Kessler el al.
8224 (LPB, MO); Carrasco NP. Cavernas del Repechon,
Kessler 8253 (LPB. MO). 8290 (LPB, MO); S of Campa-
mento Ichoa. Acebey 563 (LPB, MO). 580 (LPB. MO); km
143 on old rd. from Cochabamba to Villa Tunari, Kessler
<567 (LPB. MO); km 149 on old rd. from Cochabamba to
Villa Tunari, Kessler et al. 7995 (LPB). La Paz: Caranavi.
Serrama Bella Vista, Caranavi— Sapecho, km 42. Kessler et
al. 11516 (LPB); Iturralde, Rio San Antonio, 46 km de
Ixjamas a Alto Madidi, Kessler et al. 11128 (LPB, MO);
San Buenaventura, R. S. Williams 533 (K); Nor Yungas,
Polo-Polo bei Coroieo, Buchtien 3657 (US); 4.5 km below
Yolosa, then 10 km \\ on rd. up Rio Huarinilla, Solomon
8562 (LPB, MO); Franz Tamayo, Madidi NP. Refugio
Chalalan, vie. Campamento Estabon, Kroemer & Acebey
111! (LPB), 1134 (LPB. MO); 42 km W and I km N of
Rurrenabaque, Helme 423 (LPB); Sud Yungas, 5 km de
Chamaca a La Asunta, Kessler et id. 5803 (LPB. MO).
Santa Cruz: Ichilo, 3 km by river above Campamento
Maeunueu, Kessler et al. 8700 (LPB. MO).

Mosstera

Monstera kessleri Croat, sp. nov. TYPE: Bolivia.
Cochabamba: Chapare, Parque Machia, 1 km
E of Villa Tunari, 16°48'S, 65°24'W, 350 m,
14 Sep. 1996, T. Kessler 8479 (holotype, M0-
05042748; iso type, LPB). Figures 4C. I), 5A.

Planta hemiepiphytica; petiolus 15-36 cm longus, va-
ginatus ad geniculum; lamina anguste ovata, ovato-ellip-
tica, 25—41(46.5) cm longa, 13—25 cm lata; pedunculus
22 cm longus, ca. 5 mm diam. in sicco; spadix 1 1.5 cm
longus, 2.5 cm diam.

Hemiepiphyte; stems with pre-adult internodes
1-2 cm long, drying smooth, dark yellow-brown,
densely and minutely granular on magnification;
adult internodes unknown. Petioles 1 5-36 cm long,
sheathed throughout to geniculum, drying matte,
medium-dark yellow-brown, weakly and closely
ridged; sheath margin narrow, sometimes flaking
free. Leaf blades narrowly ovate to ovate-elliptic,
25-41(46.5) X 13-25 cm, (1.2) 1.6-1. 9(2.7) times
longer than wide, 1.1 — 1.7 times longer than peti¬
oles, rounded to acute at apex, acute to weakly at¬
tenuated or rounded at base, drying dark brown and
matte above, dark yellow-brown and semiglossy be¬
low; margins entire and with a single elongated per¬
foration or with only 1 to 2 deep, narrow sinuses
open to the margin (only I deep sinus on pre-adult
blades); midrib drying weakly raised and concolo-
rous above, convex and slightly paler below; pri¬
mary lateral veins 9 to I I pairs, arising at an acute
angle then spreading at 3()°-50° angle, flat and
weakly visible above, convex and slightly darker
than surface below; minor veins weak, fine, paral¬
lel. IN I LORESCENCES with peduncle 22 cm long,
ca. 5 mm wide on drying, drying dark yellow-
brown; spadix 11.5 cm long, 2.5 cm diam. Styles
3-4.5 nun diam. on drying, irregularly angular,
matte, concave; stigma raised, oblong, 1 .8-2.2 mm
long, 0.3— 0.4 mm wide, the margins light brown,
with a deep medial furrow.

Distribution and habitat. Monstera kessleri is
known only from the type, collected near Villa Tun¬
ari in Cochabamba, Bolivia, but juvenile material
collected near Rurrenabaque and an aberrant spec¬
imen from Peru (see below) almost eertainlv belong
to this species, too.

The newr species is characterized by its narrowly
ovate, dark brown-drying leaf blades with only I to
2 deep sinuses per side. They are similar in size,
shape, and color to those of Monstera spruceana
(Schott) Engler, which also has similar pre-adult
blades, but differs in having the peduncle shorter
than the spadix (vs. about twice as long in M. kes-
sleri ). A collection from the department of La Paz.

92

Novon

Figure 5. — A. Monstera kessleri Croat (Kessler 8479). Adult plant with blade lacking perforations [hole in leaf is an

artifact of collecting!. — B. Philodendron colapatense Croat & Acebey (Kroenier & Acebey 1858). lype specimen. (.. I).
Philodendron kroemeri Croat X Acebey (Groat & Acebey 84820). — C. Habit. — D. Close-up of adaxial surface of blade.

NV 2319113

Volume 15, Number 1
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Croat & Acebey

Araceae from Bolivia and Tropical Andes

93

Province Franz Tamayo (Helme 252), collected 42
km west and 1 km north of Rurrenabaque at
14°25'S, 67°55'W, 330 m, is very likely a juvenile
plant of M. kessleri, and it has exserted leaves rath¬
er than the shingle-leaves characteristic of M. spru-
ceana.

The species is named in honor of Michael Kes¬
sler, a German plant ecologist from the University
of Gottingen who has devoted much of his profes¬
sional career to the plants of Bolivia. Kessler has
collected many interesting and new Araceae during
the course of his studies and made an extensive
study of the Araceae in Bolivia (Kessler & Croat,
1999).

Philodendron

Philodendron cotapatense Croat & Acebey, sp.
nov. TYPE: Bolivia. Fa Paz: Nor Yungas, Co-
tapata NP, trail to Est. Biol. Tunquini, toward
Rio Cedroni, 16°12'S, 67°52'W, 1800 m, 12
Sep. 2000, T. Kroemer & A. Acebey / 558 (ho-
lotype, MO-5207999; isotype, LPB). Figure
5B.

l’lanta hemiepiphytica; internodia 1—3 cm longa, 1.7
cm diam.; cataphylla 8 cm longa, acute 1-costata, decidua;
lamina ohlonga, 41.5—4-3 cm longa, 6—6.7 cm lata, anguste
acuta ad basim; nervis primariis non manifestis; inflores-
centiae 2 in quaque axilla; pedunculus 4—6 cm longus;
spat ha 8.5— 9.5 cm longa.

A |)| tressed-( limbing hemiepiphvte; internodes I —
3 cm long, 1.7 cm diam., drying initially blackish
and smooth, becoming minutely, finely and irregu¬
larly ridged, matte, finally light yellow-brown and
smooth with big folds, sometimes with the epider¬
mis loosening; cataphyils 8 cm long, sharply I-
ribbed on drying, drying blackened, deciduous.
Leaf blades oblong, 41.5—43 X 6— 6.7 cm, 6.9 times
longer than wide, 2—2.3 times longer than petioles,
narrowly acute at base, narrowly caudate-acumi¬
nate at apex, semiglossy, slightly bicolored, drying
medium gray-green above, slightly paler and yel¬
lowish gray-green below; midrib broadly convex
and slightly darker than surface above, narrowly
convex and darker than surface below, drying finely
and inconspicuously ridged on both surfaces; pri¬
mary Intend veins not at all apparent; minor veins
numerous, fine, arising at 35°-45° angle, sometimes
drying weaklv undulate. INFLORESCENCES 2 per
axil; peduncle 4—6 cm long, drying black, 3 mm
diam. midway; spathe 8.5— 9.5 cm long, drying 1.2—

1 .5 cm diam.; stipe 4 mm long in front. 5 mm wide;
spadix 8.7 cm long, ca. 0.8 mm shorter than the
spathe; pistillate portion of spadix 2.1 cm long, dry¬
ing 7 X 3 mm wide; staminate portion of spadix

6.5 cm long, drying 5X8 mm wide. Pistils 0.8—
1.0 mm long; 5- to 6-locular; stigmas 0.6— 0.8 mm
diam.; ovules 1 per locule, basal 0.6— 0.7 mm long;
enveloped in a gelatinous envelope 2 mm long; fu-
nicle ca. 2 times as long as ovule.

Distribution and habitat. Philodendron cota¬
patense is endemic to Bolivia, where it is known
only from the type locality in Parque Naeional Co-
tapata (hence the name), iti La Paz Department, at
1800 m elevation.

fhe species is a member of section Calostigma
(Schott) Pfeiffer, subsection Glossophyllum (Schott)
Engler, series Glossophyllum (Schott) Croat. It is
characterized by its somewhat grayish drying, nar¬
rowly oblong-oblaneeolate blades that lack any ob¬
vious primary lateral veins and its moderately
small, narrowly pedunculate, blackish drying inflo¬
rescences. It is closest to P. paxianum K. Krause,
which differs in having leaf blades with distinct
primary lateral veins.

An apparently juvenile collection (Solomon
9549 ) has blades with similar venation, but leaves
that dry light yellow-green. It is uncertain if that
collection belongs here or if it represents another
species.

Paratype. BOLIVIA. La Paz: Nor Yungas, Est. Biol.
Tunquini, Homuni Bajo, senda del campo l)n. Pedro al
pajonal detras del no Cedroni, Bach et al. 1717 (LPB,
MO).

Philodendron kroemeri Croat, sp. nov. T\ PE:
Bolivia. La Paz: Nor Yungas, Serrama Bella
Vista, Caranavi— Yucumo, 34.3 km NE of Car-
anavi, 15°41'30"S. 67°29'37"W, 1567 m, 5
Aug. 2000, T. H. Groat, A. Acebey A T. kroemer
84820 (holotype, MO-5185172—3; isotypes, R.
E, GOET, K. LPB, NY, US. USC. USM). Fig¬
ures 5C, D. 6A.

Planta hemiepiphytica; internodia brevia, 3 cm diam.;
cataphylla 22 cm longa, acute 2-costata, persistentia in-
tacta; petiolus 43 cm longus, subteres; lamina late ovata,

42.5 cm longa, 28.5 cm lata, profunde lolmta ad basim;
lobi postici 12.5—14 cm longi, 10 cm lati; nervis basalibus
6—7 utroque; nervis primariis lateralibus 6 utroque; inllo-
rescentiae 3—7 in quaque axilla; pedunculus (1.5)4— 7.7
cm longus; spatha 4.5— 5.5 cm longa; spadix 4.8— 5.2 cm
longus; pars masculina 1.2— 2.1 cm longa, 6 mm diam.;
pars feminea 3.0 cm longa, 6.5—7 mm diam.; ovarium 4—
5 loculare; ovula I in quoque loculo.

Appressed-climbing hemiepiphtye; internodes
short, 5—6 cm long, to 3 cm diam., drying dark
yellow-brown, narrowly ridged longitudinally; cat¬
aphylls 22 cm long, sharply 2-costate, medium
green, obscurely darker green-1 ineate, marcescent
and persistent semi-intact at least at the upper

94

Novon

Figure 6. — A. Philodendron kroemeri Croal & Acebey ( Croat & Acebey 84820). Close-up of stem, petiole bases, and

inflorescences. If. C. Rhodospatha mukuntakia Croat (Croal el al. 84386). — If. Habit. — C. Close-up of petioles with
fibrous sheath and an unopened inflorescence. — 1). Stenospermation killipii Croat & A. Gomez (Plowman & Schunke
7402). Potted plant showing erect habit with somewhat eernuous inflorescence.

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Araceae from Bolivia and Tropical Andes

95

nodes, drying light yellowish brown, soon decidu¬
ous. Petioles 43 cm long, subterete, medium dark
green and semiglossy, obtusely flattened adaxially,
unmarked, drying dark brown to blackened, 8—10
cm diam., matte. Leaf blades broadly ovate, 42.5
cm long, 28.5 cm wide. 1.5 times longer than wide,
subcoriaceous, dark green and weakly glossy to se¬
miglossy above, slightly paler and semi glossy be¬
low, abruptly short -acuminate at apex, deeply lobed
at base, drying dark blackish brown above, slightly
paler and dark yellow-brown below; anterior lobe
32.5 cm long, the margins broadly convex; posterior
lobes 12.5—14 cm long, 10 cm wide; basal veins 6
to 7 pairs, the 1st pair free or nearly so to base,
3rd and higher order basal veins coalesced 1.5—4
cm; posterior rib scarcely naked, mostly to ca. 1
cm; midrib broadly convex and concolorous to
slightly paler above, convex to narrowly rounded
and slightly paler below, drying slightly darker than
surface above, moderately darker and irregularly
and finely striate below; primary Intend veins 6
pairs, arising at 55° angle near apex, to 65°— 85°
angle in lower half of blade, convex, darker than
surface, sometimes drying blackened and much
darker than surface; minor veins moderately indis¬
tinct with a few visible, drying close and line but
not markedly visible. INFLORESCENCES 3 to 7
per axil; peduncle (1.5)4— 7.7 cm long, 9 mm diam.,
pale green at base, darker green toward apex, dark
brown to blackened on drying; spathe white at an-
thesis, becoming [tale green, greenish white and
glossy inside, 4.5— 5.5 cm long, 6—9 mm diam.,
abruptly pointed at apex, drying blackened to dark
brown; spadix 4.8— 5.2 cm long; staminate portion
1 .2—2.1 cm long, 6 mm diam.; sterile staminate por¬
tion not obvious on live material; pistillate portion
3.0 cm long in front, 2.3— 2.6 cm long in back, 6.5—
7 mm diam. at base, 6.0 mm diam. at apex. Pistils
bottle-shaped, subterete to rhombic in cross sec¬
tion. the upper I mm tapered inward, 1.0— 1.7 mm
diam.; locules 4 to 5 per ovary; ovules I per loeule,
0.6— 0.9 mm long, 0.3— 0.4 mm diam.; placentation
basal; funiele shorter than ovule; stigma 0.5— 0.7
nun diam., 2-3 mm thick; androecium 1.0— 1.7 mm
diam., broadly convex at apex, subrounded to ir¬
regularly angular, 4- to 5(6)-sided, the angles
broadly rounded, f ruits not known.

Distribution and habitat. Philodendron kroe-
meri is known from Bolivia at the Cotapata National
Park in La Paz and Peru at Pasco Province (Oxa-
pampa) at 1650—2400 m elevation.

The new species is characterized by its persis¬
tent, more or less intact cataphylls, terete, dark
brown to black-drying petiole, broadly ovate, dark

blackish brown-drying blades, and cluster of 3 to
7 inflorescences. Similar material has been col¬
lected in Peru, Department of Pasco, in the vicinity
of Oxapampa at 1600—2080 m \Vdsquez et al.
26142, Soukup 2321 (GH); Poster et al. 7736 (F,
MO); I). Smith 2705 (MO, USM)]. This material
may prove nonspecific, but differs in having decid¬
uous cataphylls and blades that dry reddish brown
on the lower surface (rather than dark yellow-
brown).

The species is named in honor of German bota¬
nist Thorsten Kroemer, specialist on the bromeliads
of Bolivia, who has collected many interesting and
new species of Araceae during the course ol inves¬
tigations for his Ph.D. thesis from the University of
Gottingen.

Paratypes. BOLIVIA. La Paz: Nor Yungas, Cotapata
NP. Est. Biol. Tunquini, senda del campo de Don Pedro
al camino de la mina, Garcia 4435 (LPB); Hornuni Bajo,
near Campamento Don Pedro, Kroemer & A. Acebey 1745
(EPB. MO), 1590 (LPB, MO); Hornuni Bajo, senda eafetal
al camino de la mina, Bach et al. 1623 (LPB, MO). PERU.
Pasco: Oxapampa, Chontabamba, carretera camino a La
Sniza antigua (km 10—13), Lingan et al. 373 (B. k, MO.
US, USM); carretera camino a La Suiza nueva, Lingan et
al. 376 (MO. USM).

Riiodospatha

Rhodospatha mukuntakia Croat, sp. nov. T\ PE:
Bolivia. La Paz: Abel Iturralde, Tumupasa—
San Jose de Uchupiamonas, NW of Tumupasa,
Madidi NP. 5.5— 5.8 km above jet. to San .lose
near Tumupasa, 15°45'N, 67°50'W, 830—850
m. 9 Aug. 2000, T. B. Croat, A. Acebey & T.
Kroemer 84386 (holotvpe, MO-5187103; iso-
types. B, F. k. LPB, NY, US. USM, UB). Fig¬
ure 6B, C.

Planta terrestris; intemodia brevia, 2.5—3 cm diam.; pe-
tiolus 35—71 cm longus; lamina plus minusve elliptica,
37.5—63 cm longa, 18—31 cm lata; nervis primariis later-
alibus 25—35 utroque; pedunculus 16-35 cm longus; spa-
tha eremea, plus minusve elliptica; spadix 9.5—16 cm lon¬
gus, 1-1.5 cm diam., stipitatus 5-18 mm, dilulus
aurantiacus vel cremeus.

Terrestrial herb, 50—100 cm tall; internodes
short, 2.5—3 cm diam., the epidermis always cov¬
ered by overlapping petiole sheaths. Petioles 35—
71 cm long (averaging 54 cm), drying light yellow¬
ish brown, closely fissured adaxially, deciduous,
sheathed to within 1—9 cm of the genieulum; sheath
light brown, highly weathered, with pale, ± parallel
strands of fibers and sometimes with fragments ol
brown epidermis persisting; genieulum 1.5—4 cm
long, narrowly silicate, drying sharply sulcate, re¬
mote 2.5— 3.5 cm from the blade. Leaf blades ±
elliptic. 37.5—63 X 18—31 cm, averaging 55 X 26

96

Novon

cm, 0.76—1.2 times longer than petioles, broadest
at the middle, slightly inequilateral (one side ea. I
cm wider), somewhat inequilateral, acute to obtuse
and attenuate at base, acute to obtuse or narrowly
rounded and abruptly acuminate at apex, drying
brownish green above, reddish brown beneath; mid¬
rib sunken and concolorous above, much thicker
than broad below, drying brown, softly and densely
crustose-puberulent; primary lateral veins 25 to 35
per side. (4)7-21 nun apart (mostly 15—20 mm)
arising at an acute angle then spreading at 65°— 85°
angle, weakly curved to the margin, drying dark
brown, smooth; interprimary vein 1, usually much
weaker than the primaries; minor veins usually 2
(sometimes 3). alternating with the interprimary
and primary veins, the interprimary and minor
veins usually sparsely granular; cross-veins con¬
spicuous and close, extending across the whole sur¬
face, the surface greenish and densely reddish
granular punctate on magnification. INFLORES¬
CENCE, erect; peduncle 16—35 cm long, drying 3—

5 mm diam., 0.29—0.79 times as long as the peti¬
oles. averaging about Vi as long as petioles; spathe
cream, turning reddish brown on drying, somewhat
elliptic, acute at apex, ca. 1.5 cm longer than the
spadix; spadix 9.5—16 cm long, 1 — 1.5 cm diam., to
ca. 7 mm diam. ca. I cm below apex, tapered
slightly toward both ends, stipitate 5—18 mm, light
orange to cream (variously described on labels as
B & k yellow-red 8/2.5, 8/5, yellow-red 9/7.5, 8

6 k yellow 8/2.5, and 8 & k yellow-red 9/10).
Pistils often regularly pentagonal or 4-sided, some¬
times irregularly 5- to 6-sided, 1—2 mm diam.
(those at the base of the spadix often larger), fre¬
quently 2.4—3 nun diam., the sides straight to weak¬
ly convex, rarely concave; style faintly purplish
brown with a frost-like matte surface; stigma 0.6—
0.9 mm long, 0.3— 0.4 mm wide, brown, raised, usu¬
ally sulcate medially; anthers narrowly [jointed to¬
ward apex. 1.8 mm long, 0.7 mm diam., promptly
weathering at apex.

Distribution and habitat. Rhodospatha mukun-
takia ranges from southern Colombia (Putumayo) to
central Peru (Amazonas, Cuzco, and Madre de
Dios) and Bolivia (La Paz) from 180 to 930 m. Most
collections are from Nangaritza Canton (at 930 111)
in Zatnora-Chinchipe, Ecuador, and the drainage of
the Kfo Santiago in the Department of Amazonas
at 180-200 m elevation in Peru.

Rhodospatha mukuntakia is recognized by its
terrestrial habit, short internodes obscured by the
petiole bases, reddish brown-drying petioles,
ribbed adaxially and with a deciduous, highly fi¬
brous sheath extending to within 1—9 cm of the

geniculum. The genieulum also dries sharply sili¬
cate, the blades are more or less elliptic, drying
reddish brown and reddish granular- punctate on
the lower surface with prominent, close cross-veins,
the lower midrib densely crustose-puberulent.

The species epithet is based on the common
name “mukuntak” used by the Aguaruna indige¬
nous people of the Rfo Cenepa and Bfo Santiago
drainages in northern Amazonas, Peru, one of the
localities where most collections of this species
have been made.

Paratypes. COLOMBIA. Caqucta: 10 km SW of San
Jose de Fragua, SW of Florencia, Davidse el al. 5704
(COL, MO). Putumayo: Mocoa. Scludles A Cabrera
10045 (US); Villa Gamin. Rfo Cineo. 8 km W of Villa
(Luzon. Plowman 2057 (F. (ill). ECUADOR. Morona-
Santiago: Pumpuentza, W NW of village, Brandbyge &
Asanza .32555 (AAU); Pumpuentza, SSW of village,
Rrandbyge & Asanza 52451 (AAU); Los Tayos, /.. Rodri¬
gue: et al. 5I5A (MO); Taisha, Rrandbyge & Asanza 5IR70
(AAU, MO); Cualaquiza, Mision Bomboiza Salesiana,
Sparre 10150 (S). Napo: lies. Florfstiea “El Chuneho"
Payamino, 5 km al NW de Coca, Kst. Exp. INIAP-Napo,
Cerdn 2407 (M0); vie. Tena, Asplund 8897 (MO); Bfo
Aguarico near confluence with Bfo Pavayacu Bravo, Go¬
mez 256 (MO). Sueumbios: Canton Gonzalo Pizarro, Par-
roquia Pto. Libre (San Pedro de los Cofanes), Cofdn Si-
nangfie, Cerdn et al. 22078 (QAP). Zamora-Chinchipe:
Nangaritza, Cordillera del Condor, Shaimi, frente a des-
taeamento militar, Bfo Nangaritza, Palacios et al. 8711
(MO, QCNE); Shaime, jet. of Bfo Nangaritza and Bfo
Numpatakaime, Palacios 0 011 (MO, OCNE); Nangaritza,
Cordillera del Ctindor, Parroquia Guayzimi, Campamento
militar Miazi, S of Bfo Nangaritza, Cerdn et al. 17870
(DAP). PEBl . Amazonas: I km below La Po/.a, E of Rfo
Santiago, D. Pena 128 (MO); Bfo Cenepa, near Huarn-
pama, near Chavez Valdivia, Kujikat 512 (MO); Bagua, La
Peca, Cordillera Collin, SE of La Peca, 3rd camp. Harbour
4145 (MO. USM): Shimpunts, ehacra al lado de Shim-
punts, Aneuash 1521 (MO); vie. Kusu, Bfo Numpatkin,
Bfo Cenepa drainage system, monte orilla de Chapis, An¬
euash 7410 (MO); ca. 5 km E of Chavez Valdfvia, Aneuash
II 60 (MO); S of Huampami trail to house of Theodora, S
of Bfo Cenepa, Berlin 1 665 (MO); Kusu, Bfo Numpatkin,
vie. Kusu, Kuyap 515 (MO, US); Condorcanqui, Bfo San¬
tiago. 2—3 km behind Caterpiza, Tunqui 004 (MO); 2 km
behind Caterpiza, E of Caterpiza, ca. 65 km N of Pinglo,
Huashikal 505 (MO), 2060 (MO), 2256 (MO), 2287 (MO),
2505 (MO). 2542 (MO); I km behind Caterpiza. E of Ca¬
terpiza, Bfo Santiago. Huashikal 751 (MO). 886 (MO); Bfo
Santiago, otra banda de Caterpiza, 2 km en trocha de me-
tallar, Tunqui 128 (MO). Cuzco: La Conveneidn, Echarati
Cashiriari, 3 well site, 5.0 km S of Camisea Biver. I\uiiez
el al. 25080 (MO. US). Madre de Dios: Manunear Shin-
tuya on rd. to Salvacibn, Plowman & Davis 5076 (Gil).
San Martin: Mariscal Caceres, Santa Cruz, Schunke 7001
(MO). BOLIVIA. Beni: Ballivian. Burrenahaque-Sape-
cho, 21.7 km NW of La Cascada, 38.8 km NW of Las
Delicias, Croat et al. 84666 (LPB, MO). Cochabamba:
Chapare, vie. Villa Tunari, along Bfo Espfritu Santo on
trail to Baja Copacabana, Croat 51281 (LPB, MO). La
Paz: Sud Yungas, Caranavi— Yucumo, 47.4 km NE of Yu-
cumo, 5.1 km from border of Beni, Croat et al. 84550
(LPB, MO).

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Araceae from Bolivia and Tropical Andes

97

S TE NOS PERM A TION

Stenospermation dictyoneurum Croat & Ace¬
bey, s|>. nov. TYPE: Bolivia. La Paz: Nor Yun-
gas, Serranfa Bella Vista. Caranavi— Yucumo,
37.6 km NE of Caranavi, 15°40'37"S,
67°29'34"W, 1508 m, 5 Aug. 2000, T. B.
Croat, A. Acebey & T. Kroemer 84270 (holo-
type, MO-5185298; isotypes, K, LPB, US).
Figures 7C, I). 8A.

Planta epiphytica; internodia 1.5-2 cm longa. Petiolus
(10)13-21 cm longus; lamina 13-31 cm longa, 3-5.3 cm
lata; pedunculus 22—29 cm longus, 1.5— 2.5 mm diam.,
atrobrunneus in sicco; spatha nivea; spadix niveus albus,
3.3— 6.2 cm longus, 5—10 mm diam.

Epiphyte at 2—3.5 m; stems soon brown; inter¬
nodes 1.5-2 cm long, drying 6-7 mm diam., usually
drying dark brown and moderately smooth, some¬
times gray and weakly glossy. Petioles (10)13-21
cm long, sheathed 0.43%-0.71% dieir length, me¬
dium green, matte, dry ing dark brown; sheath erect,
narrowly rounded at apex, ending 2.2-1 I cm below
the geniculum. Leaf blades 13-31 X 3-5.3 cm,
3.5— 6.3 times longer than wide, subcoriaceous,
dark green and semiglossy above, moderately paler
and glossier below, drying dark yellow-brown to
grayish brown and matte above, yellowish brown to
olive-green and semiglossy to weaklv glossy on low¬
er surface; midrib sunken and slightly paler above,
thicker than broad and slightly paler below, drying
weakly raised, sometimes narrowly sunken toward
apex, ± coneolorous above, drying narrowly raised
on lower surface near the base, convex to weakly
ridged and coneolorous to slightly darker below;
primary lateral veins weakly visible above, scarcely
apparent below, drying weakly, narrowly and acute¬
ly raised above, much weaker and somewhat darker
below, scarcely more apparent than the minor veins
below; minor veins also acute and narrowly raised
above with numerous wavy, acutely erect cross¬
veins. these sometimes appearing to be reticulate,
those of the lower surface scarcely apparent; lower
surface weakly and densely granular. INFLORES¬
CENCE erect to weaklv cernuous; peduncle 22—29
cm long, drying dark brown, 1.5-2. 5 mm diam.;
spathe white, soon brown, promptly deciduous; spa¬
dix directed weakly to one side, greenish white,

3. 3-6.2 cm long, 5—10 mm diam., cylindroid,
rounded at both ends. Flowers 4 to 5 visible per
spiral, 1.0— 1.4 mm long, 1.2— 1.4 mm wide, trun¬
cate at apex, drying dark brown; stigma subglobose,
ca. 0.5 mm wide, brush-like, raised above the sur¬
face of the stigma.

Distribution and habitat. Stenospermation dic¬

tyoneurum is known only from Bolivia in La Paz at
1300-1850 m elevation.

I he new species is characterized by its generally
epiphytic habit, rather short, slender internodes,
petioles sheathed usually to %— % their length, and
leaf blades drying dark brown above, moderately
paler and yellow-brown below, with the minor veins
narrowly and acutely raised on the upper surface
with distinct oblique and transverse cross-veins and
sparse, short pale lineations between the minor
veins.

The species epithet is based on the Creek “dic¬
tion" (net) and “neuron" (nerved) alluding to the
prominent net-like reticulum of its veins.

Paratypes. BOLIVIA. La Paz: Caranavi, Serranfa
Bella Vista, Caranavi— Sapecho, km 37, Kessler el at.

1 1848 (LPB, MO); Nor Yungas, Lst. Biol. Tunquini, Bach
969 (LPB, MO), Kroemer & Acebey 1556 (LPB. MO); Saa¬
vedra, Camata to Apolo, km 10, Kessler 10824 (LPB, MO).

Stenospermation killipii Croat & A. Gomez, sp.
nov. I YPE: Peru. Huanuco: Leoncio Prado,
Hermilio Valdizan, La Divisora, along rd. from
Pumahuasi to La Cumbre, 1600-1660 m, orig¬
inally collected live by T. Plowman, cultivated
at MO, 26 June 1978, I B. Croat 78848 (ho-
lotype, MO-05015335; isotypes, B, E, K, NY,
US, USM). Figures 6D, 7A, B.

Planta terrestris vel epiphytica; internodia 0.5— 4.0 cm
longa, 1.2-2.0 cm diam.; petiolus 9-12.5 cm longus; lam¬
ina elliptica vel oblongo-elliptica vel obovato-elliptioa,
15-24 cm longa, 3.5— 6.5 cm lata, superfieie dilute brevi-
lineata; pedunculus 23-33 cm longus; spatha 8-9.5 cm
longa, 2.5 cm lata, persistens, viridis vel virello-alba.

Terrestrial or epiphytic; stems clustered, (0.5)1-
1.5 m tall; internodes 0.5— 4.0 cm long, 1. 2-2.0 cm
diam., dark green, medium green, becoming gray-
green, weakly glossy, drying dark brown, closely
ridged, densely and finely granular. Petioles 9—12.5
cm long, sheathed from % its length or to the ge¬
niculum, matte to weakly glossy, dark olive-green;
sheath decurrent, ineurled hut mostly open; free
portion sub-terete, bluntly sulcate; geniculum 1.8
cm long, sharply sulcate, drying sharply sulcate.
Leaf blades elliptic to oblong-elliptic or obovate-
elliptic, 15-24 cm long, 3.5— 6.5 cm wide, 2.5— 4.6
times longer than wide, 1 .5-1.6 times longer than
petioles, subcoriaceous, acute at base, acuminate
at apex, dark green and weakly glossy to subvelvety
above, moderately paler and semiglossy to weakly
glossy below, drying dark yellow-brown to yellowish
green above, medium yellow-brown below; midrib
obtusely and deeply sunken and coneolorous above,
slightly paler below, drying darker below, drying
finely striate; primary lateral veins not apparent

98

Novon

Figure 7. A, B. Stenospermation killipii Croat & A. Gomez (Plowman & Schunke 7402). —A. Erect inflorescence
showing straight peduncle and elliptic spathe. — B. Close-up of inflorescence with open spathe exposing spadix. C, D.
Stenospermation dictyoneurum Croat & Aeebey (Croat et al. 84270). Habit.

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Araceae from Bolivia and Tropical Andes

99

above, weakly visible below; minor veins drying
weakly raised, with numerous pale, short-pale-l id¬
eations in between the minor veins above, weakly
raised but less dense below with fewer pale cellular
inclusions. INFLORESCENCES erect: peduncle
23-33 cm long, drying dark brown 1-2 mm diam.;
spathe 8—9.5 cm long, 2.5 cm wide at anthesis, 7.5
cm wide when flattened, green to greenish white,
erect, semiglossy outside, weaklv glossy inside,
persisting after anthesis, dark reddish brown; spa¬
dix white, 4.5—7 cm long, 1—1.5 cm diam., with
reduced flowers and bluntly pointed at apex. Pistils
3—3.5 mm diam., 8 to 9 visible per spiral.

Distribution and habitat. Stenospermation kil-
lipii ranges from Ecuador (Zamora-Chinchipe) and
Peru (Amazonas), San Martin (Rioja), Huanuco
(Coronel Portillo, Leoncio Prado), Junin (Oxapam-
pa, Tarma), Ayacucho (La Mar), and Cuzco (Pau-
cartambo) to Bolivia at (700)1600—2100 m eleva¬
tion. In Bolivia it occurs in Cochabamba (Chapare)
and La Paz (Sud Yungas). It may also occur in Bra¬
zil in Bahia (see below).

Stenospermation killipii is characterized by its
erect, frequently terrestrial habit, long stems, often
fully sheathed petioles, weakly glossy to subvelvety,
dark yellow-brown to yellow-green-drying blades,
and especially by its short spadix and persistent
spathe.

The new species is similar to Stenospermation
wallisii Masters in having a persistent spathe, but
differs in having leaf blades that dry yellow-green
to light yellow-brown and lack the dense, pale,
short lineations between the minor veins.

Gentry et al. 21954, from the far north of Peru
in Loreto (Maynas: Rfo Gueppi), is very similar to
S. killipii, but differs in being from oidy 200 m
elevation and in having a persistent spathe and
stems drying a paler rusty brown. Since no other
collection was made below 1000 m, it seems likely
that this represents another species.

The new species is named after Smithsonian bot¬
anist E. P. killip who, along with A. C. Smith, was
the first (1929) to collect it in Peru. This species
was described with A. Gomez, a former student of
lorn Croat, who studied the Stenospermation of
Central and South America lor her master’s thesis
al St. Louis University in St. Louis, Missouri,
U.S.A.

Paratypes. ECUADOR. Morona-Santiago: Sangay
NP, trail to Sardina Yacu, Rfo Sardina-Rfo Volean, Mon¬
talvo A Cerdn 1 1 A (GAP). Zamora-Chinchipe: Nangar-
itza Canton, Rfo Nangaritza, Miazi, confluence of Rfo
Chumbiriatza— Rfo Nangaritza, Palacios A Neill 6 789
(MO, QCNE). PERU. Amazonas: near border with San
Martin, van der Werff el al. 16690 (DUKE, MO. SEE. UR.

USM); Ragua, 12 km E of La Peca, Harbour 2630 (MO);
12-17 km E of Ea Peca, Harbour 2527 (MO, USM);
Chachapoyas, Rodriguez e Mendoza, Cochamal, local idad
*‘Tinas,” Montana de Yanamonte, Diaz et al. 4512 (MO).
Ayacucho: Aina, Huanta-Rfo Apurimac, Killip A- Smith
22550 (NY, US); La Mar, below Huanhuachayo on the
Capprichio— Puncu trail, Rfo Apurimac valley, Madison
10255-70 (US). Cajaniarca: San Ignacio, San Jose de
Lourdes, base of Cerro Picorana, Diaz et al. 10284 (MO).
10777 (K. MO. N\, UR, USM); San Ignacio, San Josf5 de
Lourdes, Camana. Campos A Corrales 3760 (MO, QCNE,
US, USM); base of Cerro Picorana, Vdsquez el al. 26729
(E, MO, USM). Cuzco: Paucartambo, Kosnipata Valley,
km 150, San Pedro, jet. of Rfo Unidn and Rfo Kosnipata,
P. Nunez 1 1992 (MO). Iliinnuro: Coronel Portillo, Padre
Abad, La Divisora, near Rfo Chino, Schunke 9275 (MO);
Leoncio Prado, Hermillo Valdizan, La Divisora. Pumahu-
asi-La Cumbre, Plowman A Schunke 7402 (SEE): 2.8 km
N of divide, Croat A Sizemore 81716 (MO. USM); Cordil¬
lera Azul, on rl. 16, near border with Ucayali Department,
38 km N of lingo Marfa. Davidson A Jones 3457 (LAM);
39.2 km E of Tingo Marfa on rd. to Pucalpa, Davidson A
Jones 9338 (LAM, MO); Tingo Maria— Pucalpa, “Ea Divi¬
sora,” 3.8 km N of Ucayali border, Croat A Sizemore
81694 (MO, USM); Tingo Marfa-Huayna Capac, 10.0 km
A of Rfo Huallaga, Croat A Sizemore 81865 (MO, USM).
Juiun: Chanchamayo Valley, Schunke 475 (E); Tarma, Ea
Merced— Satipo, ea. 5 km E of Ea Merced along Rfo Chan¬
chamayo, Croat A Sizemore 81924 (MO, USM). Pasco:
Oxapampa, Chontabamba, Smith A Hrack 3073 (USM):
Gran Pajonal, vie. Chequitavo, Smith 6866 (MO); W of
Oxapampa, low pass betw. Chontabamba and Suissa. Fos¬
ter A Smith 7572 (F. MO, USM); 20 km \V of Oxapampa,
pass before Ea Suiza, Smith 5381 (MO). San Martin: Rio-
ja-Pedro Ruiz, van der Werff et al. 15464 (MO); 99 km
from Rioja on rd. to Pomacocha, Stein A Todzia 2182
(MO); Elias Soplin Vargas, Centro poblado Porvenir, Yae-
umama, rd. to Rfo Negro, Sanchez E A Dilbm 8066 (E);
Nuevo Cajamarca near Poblado Miraflores, Sanchez V. A
Dillon 8462 (CPUN, MO). BOLIVIA. Cochabamba:
Chapare, Cochabamba— Villa Tunari, vie. of Hotel Cabal¬
leros at km 94 from Cochabamba, Croat 51323 (I. PR.
MO). La Paz: Sud Yungas, de Chulumani, 5 km hacia
Irupana— Apa Apa, Beck 24759 (LPR); Nor Vungas, Cota-
pata NP, Kroemer 732 (LPR, MO); Est. Riol. Tunquini,
entrance to trail to Hornuni, Heck 28554 (EPR).

Cultivated specimens. PERU. Huanuco: 1 .eoncio Pra¬
do, Hermillo Valdizan, Ea Divisora, rd. from Pumahuasi
to La Cumbre, 1600-1660 m, originally collected as Plow¬
man A Schunke 7402), 12 Oct. 1987, Croat 68443 (MO).

Xanthosoma

Xanlhosoma puberuluiii Croat, sp. now TYPE.:
Bolivia. La Paz; Nor Yungas, La Paz— Caranavi,
Chuspipata-Yolosa, 19.0 km SW of Yolosa,
16°15'55"S, 67°47'18"W, 2234 m, 4 Aug.
2000, T. B. Croat, A. Acebey & T. Kroemer
84253 (holotype, MO-5185310; isotypes, K,
LPB). Figures 8B-D, 9A.

Planta terrestris, 1.5— 2.5 m alta; internodia brevia,
(9)13—18 cm diam.: petiolus 63—100 cm longus: lamina
ovata, sagiltata ad basim, 60—85 cm longa, 47—66 cm lata;
lobi postici 26—39 cm longi, 20—30 cm lati, nervis inferne

100

Novon

Figure 8. Stenospermation dictyoneurum Croat & Acebey ( Croat el al. 84270). — A. Close-up of inflorescence. B— D.
Xanlhasoma puberidurn Croat {Croat el al. 84258). — If. I,eaf blade adaxial surface. C. I,eal blade, abaxial surface.
— I). Stem showing inflorescences.

Volume 15, Number 1
2005

Croat & Acebey

Araceae from Bolivia and Tropical Andes

101

Figure 9. — A. Xanthosoma puberulum Croat ( Croat el al. 84253). Inflorescence with lower portion cut away to expose

pistillate portion and sterile staminate portion of spadix. — 15. Xanthosoma seidelii Croat {R. Seidel & E. Richter 1198
MO-3512834). Paratype spec 'iiiii'n (inset of inflorescence is from second sheet of the same collection, MO-5187956).

puherulentis; inflorescentiae 5-6 in quaque axilla; pedun-
eulus 16—24 cm longus; spatha 15-21 cm longa, tubus
5.5— 6.5 cm longus, 2.8— 3.4 cm diam.; pistilla viridia.

Terrestrial. 1.5— 2.5 m tall; stem 0.5— 1.2 m tall;
internodes short, (9)13—18 cm diam. at base, 8-11
cm diam. at apex; basal portion ol stem repent,
brown with pale brown fibers and fragments of epi¬
dermis of petiole bases persisting. Petioles 63—100
cm long, ca. 2 cm diam. midway, obtusely flattened
adaxially with a faint medial rib, slightly spongy,
about as long as or to 1.2 times longer than the
blades, erect-spreading, sheathed to above middle,
weakly glossy, medium green, very weakly striate,
tbe free portion weakly flattened adaxially, moder¬
ately spongy, incurled, slightly paler, bullate and
undulate on margins. Leaf blades semi-erect, ovate
and deeply Inbed at base, 60—85 cm long, 47—66
cm wide, thin, matte-subvelvety above, weakly
glossy, medium to dark green above, slightly paler
and weakly glossy below, drying dark yellow-brown
to medium yellow-green above, moderately paler
and yellowish brown to yellowish green below; pos¬
terior lobes 26—39 cm long, 20—30 cm wide, ob¬
tusely pointed at apex; posterior rib naked for 3—5
cm; basal veins I I to 13(14) pairs, none of them
free to the base, 2nd pair and higher order veins

coalesced into a straight posterior rib with the basal
veins regularly branching off. 5 to 6 of the basal
veins acroscopic, 7 to 8 ol them basiscopic; midrib
sunken and slightly paler above; sinus I 1-20 cm
deep, miter-shaped; major veins round-raised and
slightly paler below, drying slightly paler on both
surfaces; primary lateral veins 5 to 8 pairs, arising
at 45°— 70° angle, sunken and concolorous above,
round-raised, slightly paler and weakly striate be¬
low; tertiary veins weakly sunken above, promi¬
nently raised below; all major and minor veins
prominently puberulous on lower surface, the pu¬
bescence of the midrib most dense along the lower
margin near the surface of the blades, forming a
fringe; collective veins usually 2.3-10 mm from the
margins. INF LORESCENCFiS erect, 5 to 6 per axil;
peduncle 16—24 cm long, obtusely triangular, flat¬
tened on one side, medium green, matte; spathe 15—
21 cm long, matte, the spathe tube 5.5— 6.5 cm
long, 2.8— 3.4 cm diam., medium to dark green out¬
side, sometimes also inside but weakly tinged pur¬
ple inside at base usually in lines, or purple, the
blade 14—16 cm long, 2.0 cm diam., creamy white
on both surfaces, matte outside, weakly glossy in¬
side; spadix 17—20 cm long, equal to or slightly
longer than the spathe pre-anthesis, the pistillate

102

Novon

portion yellowish, 1.8 cm long in front, 1.2 cm long
in back, 1.6 cm diam. at base, 1.2 cm diam. at
apex; sterile staminate portion of spatlix 2.5 cm
long, 1.8 cm diam. at base; constricted portion of
spadix 1.1 cm diam.; fertile portion of staminate
spadix cream-colored, 1.1 cm diam. midway, 6 mm
diam. at 1 cm from apex. Pistils medium to dark
green; stigmas greenish yellow. INFRUCTESC-
ENCES to 8 cm long, 5 cm diam.

Distribution and habitat. Xanthosomn puberu-
lum is endemic to Bolivia, where it is known only
from Ea Paz in the provinces of Nor Yungas, 1. Mu-
necas, and Sur Yungas at 1400—2980 m elevation.
It frequently grows along watercourses on steep
slopes in partial shade.

The new species is recognized by its arborescent
habit with a huge, thick stem partly creeping over
the ground, its huge semi-erect leaves with the pet¬
ioles sheathed to above the middle, and ovate,
prominently lobed blades with sunken major veins
and its spathe lube medium to dark green inside,
weakly tinged purple inside at the base, usually in
lines. This latter character and the puberulent veins
on the lower blade surfaces are the most important
features distinguishing this species. This species is
similar to other arborescent species such as X. un-
dipes (K. Koch & C. I). Bouehe) K. Koch, but that
species differs in having the lower blade surface
glabrous and the spathe tube dark purple on the
inside. Heck 24546 may represent a juvenile plant
of this species; however, it is much less pubescent
than more mature plants.

Paralypes. BOLIVIA. La Paz: Nor Yungas, Unduavi-
Caranavi, 34.8 km E of Unduavi, 16.3 km SW of Yolosa,
('.rout 51575 (LPB, MO); 13.7 km NW ol San Pedro on
rd. through Incahuara— Mejillones and along trail to 12 de
Oct., Solomon 96 19 (LPB, MO); 17 km S\X of Yolosa on
rd. to Chuspipata, 13.2 km NE of Chuspipata, Solomon &
Uehling 12259 (LPB, MO); 14.4 km NE of Chuspipata,
13.}! kin above Yolosa. Solomon 5615 (LPB, MO); Yolosa,
15 km toward Chuspipata, .S’. Heck 15550 (LPB, MO); Yo¬
losa hacia Chuspipata, pasando Sacramento Bajo y el der-
rumbe grande antiguo, .S'. Beck 22477 (LPB); Garanavi—
Yucumo, 32.4 km NE of Caranavi, Croat el al. 84267
(LPB, MO, UB, US); Anmi Cotapata NP. Est. Biol. Tun-
quini, NW of Coroieo, NNE of La Paz, vie. Chairo, 23 km
W of Yolosa, Croat et al. 84770 (LPB, MO); Chulumani—
Irupana, ea. km 5, entering toward Apa Apa, Beck et al.
24782 (LPB. MO); de Chulumani 7 km haeia Irupana,
Apa Apa, .S'. Beck 24546 (LPB, MO); Tarila, Hinojosa &
Aparicio 520 (LPB); I. Muneeas. Llusta, Flores 46 (LPB).

Xantliosoimi seiilclii Croat, sp. nov. TYPE: Boliv¬
ia. Santa Cruz: lehillo, 5 km WSW of El Hon¬
do, “Potrerillo,” W of Quebrada La Concha,
1 7°40'20"S, 63°28'35"W, 400 m, 13 Feb.
1994, M. Nee X I. Vargas 44911) (holotype,
MO-046391 10; isotypes, LPB, NY. USZ). Fig¬
ure 9B.

Planta terrestris, minus quam 1 m alta; tuber depresso-
globosum, 6 cm diam.; petiolus 60—71 cm longus, 18-30
cm vaginatus; lamina ovato-sagittata, 30^10 cm longa,
20—25 cm lata; pedunculus 2.3 cm longus; spalha 13.5—
19 cm longa, tubus 5. 5-7. 5 cm longus, atropurpureus in-
lus.

Terrestrial, medium-sized herb to less than I m
tall; tuber depressed-globose, those seen less than
6 cm diam., densely rooted. Petioles 60—71 cm
long, subterete, drying greenish brown, finely stri¬
ate, sheathed 18—30 cm at base, ca. 40% its length;
sheath acute at apex. Leaf blades ovate-sagittate,
30—40 X 20—25 cm, drying very thin, dark brown
to dark olive-green and matte above, moderately
glossy and grayish yellow-green below; posterior
lobes 1 1-14 cm long, 7-1 1.5 cm wide, rounded to
narrowly rounded at apex; sinus 7—12 cm deep, ±
hippocrepiform; basal veins 4 to 5 pairs, the first
free to the base or nearly so, the remainder vari¬
ously coalesced into a straight posterior rib that
extends straight to the apex; the posterior rib not
at all naked; primary lateral veins 4 pairs, arising
at an acute angle then spreading at 40°-50° angle,
nearly straight to marginal collective veins: minor
veins thin, moderately distinct but not at all raised
on drying. INFLORESCENCES with peduncle to 23
cm long, drying dark brown, 5 mm diam.; spathe
13.5-19 cm long; spathe tube 5. 5-7. 5 cm long,
green to purplish on outside, dark purple on inner
surface; spathe blade 10 cm long, creamy white on
both surfaces; spadix slightly shorter than spathe;
pistillate portion to 2.7 cm long in front, 1.9 cm
long on back, drying 7-8 mm diam., narrowed to¬
ward apex (color not reported but drying grayish
lavender with fine dark purple longitudinal stripes
throughout its length, the stigma drying dark brown
to purplish, 1.0— 1.2 mm diam.); sterile staminate
portion 3.5 cm long, ca. 2.0 cm diam., the lower¬
most row ol flowers 3-4 mm long, 1—2 mm wide;
the other flowers about as broad but much longer,
7—9 mm long, fertile staminate portion not seen.

Distribution and habitat. Xanthosomn seidelii
is apparently endemic to Bolivia, where it is known
in La Paz and Santa Cruz at 400—1400 m.

I he new species is characterized by its medium
size, subglobose tubers, ovate-sagittate blades that
lack a naked posterior rib and dry moderately
glossy on the lower surface, and purple inner
spathe tube and pistils that dr)' with fine purplish
lines on the ovary.

The new species is most similar to X. sagittifol-
ium K. Koch, which also has leaf blades lacking
naked posterior lobes, but differs in its moderately
glossy (rather than matte and bluish green) lower

Volume 15, Number 1
2005

Croat & Acebey

Araceae from Bolivia and Tropical Andes

103

blade surface, and purple (rather than green) inner
spathe tube surface.

A juvenile collection (. M . Moraes & E. Oviedo
2025) from Beni in Yacuma may also be this spe¬
cies. Its blades differ in being proportionately nar¬
rower with posterior lobes narrowed to a blunt
point.

The species is named in honor of German bota¬
nist R. Seidel, who along with E. Richter collected
the species for the first time in 1988.

Paratype. BOLIVIA. La I’az: Nor Yungas: Km Un-
rluavi valley, Santa Rosa— Machacamarc-a, above Lamina
San Silvestre, R. Seidel E. Richter 1 1 OR (LPB. MO).

Acknowledgments. The authors wish to ac¬

knowledge the herbarium staff at LPB. MO, NYr.
US/, and US for making material available to us;
Fred Keusenkothen (MO), who generously digitized
and processed the digital images; and Monica Car-
lsen (MO), responsible for initial preparation, and
Emily Yates (MO), responsible for final preparation
and editing of the manuscript and images. The au¬
thors also thank Mike Grayum for advice on Latin
usage.

Literature Cited

Holdridge. L. IL. W. C. Grenke, W. II. Hatheway, T. Liang
& J. A. Tosi Jr. 1971. Forest Environments in Tropical
Zones. Pergamon Press, Oxford.

Kessler, M. & T. B. Croat. 1999. State of knowledge of
Bolivian Araceae. Selbyana 20: 224—234.

Teratophyllum hainanense (Lomariopsidaceae), a New Species from

Hainan Island, China

Dong Shi -Yon g

South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China,
and Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.

shiyongd@yahoo.com.cn

Zhang Xian- Chun*

Institute of Botany, Chinese Academy ol Sciences, Beijing 100093, China.
xianclumzhang@yahoo.com.cn. * Author for correspondence

A ns m ACT. A new species of Lomariopsidaceae,
Teratophyllum hainanense S. Y. Dong & X. C.
Zhang, from Hainan Island, China, is described
and illustrated. It is related to T. koordersii, but
differs in having 10 to 12 pairs of pinnae in mature
bathyphylls and the scales of sterile pinnae on both
midribs and veinlets as substellate, appressed, and
small, with diameters only to ca. 0.2 nun. The ge¬
nus Teratophyllum itself is reported from China for
the first time.

Key words: China, Lomariopsidaceae, Terato¬
phyllum.

Teratophyllum Kuhn is confined to the Paleo-
tropies and comprises 12 species (Holttum, 1966,
1978). It extends from southern Burma to eastern
Polynesia and Queensland, with most species in
Malesia (Kramer, 1990). 'Phis genus has not been
previously reported from China.

During an expedition of the fern flora of Hainan
Island in May 2002. we collected a climbing fern
without fertile fronds on Mt. Tiedingling and did
not know its identity. We made another trip to Mt.
Tiedingling in April 2003 and observed fronds ol
the unknown species at various stages and became
convinced that it belongs to the genus Teratophyl¬
lum. Compared with T. aculeatum (Blume) Metten-
ius ex Kuhn and T. ludens (F6e) Holttum, two spe¬
cies from neighboring regions of the Indochina
Peninsula, this species from Hainan is very differ¬
ent. Il is distinguished from T. aculeatum by once-
pinnate bathyphylls with crenate pinnae and from
T. ludens by pinnae lanceolate and asymmetric at
base in both bathyphylls and acrophylls. In addi¬
tion to T. ludens , there are three other species in
Teratophyllum with the pinnae of bathyphylls cre-
nate, i.e., T. arthropteroides (H. Christ) Holttum. T.
clemensiae Holttum. and T. koordersii Holttum, all

in section Teratophyllum (Holttum, 1978). The new
species from Hainan is distinct from T. arthropter¬
oides and T. clemensiae by its linear fertile pinnae.
However, it is similar to T. koordersii from Luzon
and the Celebes, differing in having 10 to 12 pairs
of pinnae on mature bathyphylls (vs. 2 to 6 pairs)
and the scales of sterile pinnae on both costae and
veinlets as substellate, appressed, and small (ca.
0.2 mm diam.).

Teratopliy Hum hainanense S. Y. Dong X X. (>.
Zhang, sp. nov. TYPE: China. Hainan Island:
Qiongzhong County, N slope of Mt. Tiedin¬
gling. 18°57'10"N, 109°42'19"E, 720 m, 11
Apr. 2003. S. Y. Dong ttltt (holotype, PE |2
sheets]; isotypes, L, M0, P, PE). Figures 1, 2.

At finis T. koordersii a quo differt bathyphyllorum ina-
turorum pinnis lateralibus 10—12 jugatis, squamis pinna-
rum acrophyllorum ad costam medium et nervulos nas-
oentibus appressis suhstellatis ca. 0.2 mm diametro,
pinnis fertilibus linearibus, 13—18 cm longis et 0.8 mm
latis.

Mature plants. Rhizome , 3—4 X 2 nun. green,
dorsiventral, roots on the ventral surface and 2 rows
of fronds on the dorsal surface, without obvious
spines but with a few remaining spine bases; apex
of rhizome and buds of fronds covered w ith brown
lanceolate scales (ca. 8 mm long). Sterile acrophylls,
usually 50-65 X 18-23 cm, stipe 8-13 cm long,
swollen at base; rachis not winged, stramineous;
pinnae 8 to 11 pairs, alternate, all articulate; pet-
iolules 2-7 mm long, pinnae usually about I 1 X
2.5 cm, largest to 16 X 3.4 cm, broadest near base,
the proximal and basal portions nearly parallel-sid¬
ed, tapering gradually from the middle to an acute
apex, acroscopic base rounded or round-truncate,
basiscopic base cuneate; texture green, papyra¬
ceous; midribs prominent both above and below,
usually pale; veins ca. 1.5—2 mm apart, slightly

Novon 15: 104-108. 2005.

Volume 15, Number 1
2005

Dong & Zhang
Teratophyllum hainanense

105

Figure I Teratophyllum hainanense S. Y. Dong & X. C. Zhang. Based on the holotype, S. Y. Dong 818 (PE). — A.
Bathyphylls, early stage. — B. Dimidiate bathyphy 11s. — C. Mature bathyphylls. — D. Scale from lower surface of sterile
acrophyll. — E. Pinnae of sterile acrophyll. — F. Pinnae of fertile acrophyll. Drawn by Sun Ying-Bao.

prominent on both surfaces, forked once or twice,
ending ca. 0.2 mm from the margin or reaching the
margin; margin nearly entire or slightly crenate
(lig. IE); midribs and veinlets abaxially hearing

numerous minute suhstellate scales (ca. 0.2 mm
diam.), the scales brown, appressed (Fig. ID). Fer¬
tile acrophylls, 3 seen, ca. 30 X 15—20 cm, stipe
5—8 cm long, stipe and rachis pale; pinnae about

106

No von

Fi^un* 2. Teratophyllum hainanense. — A, 15. SKM micrographs of spores. (A: scale bar 23.1 /xm; 15: scale bar 3.8
/im). — C. Adult sterile and fertile leaves. — I). Young and mature hathyphylls. All from S. Y. Dong HIM (PE).

10 pairs, petiolules 6 mm long, pinnae linear, usu¬
ally 13-16 X 0.8 mm, the longest to 18 cm long;
sporangia covering the abaxial surface and appear¬
ing to cover both surfaces when mature, the width
of the pinnae then expanding to ca. 1.5 mm (Fig.

I F). Sporangia containing 32 spores each, spores
large, 50—72 /xm diam., with a folded echinulate
perispore (Fig. 2 A, B).

Early stage. Very young fronds ca. 2 cm long,
with 6 small pinnae on the basiscopic side of the

Volume 15, Number 1
2005

Dong & Zhang
Teratophyllum hainanense

107

Table 1. Features distinguishing Teratophyllum ludens, T. koordersii, and 7. hainanense.

Characters

7. ludens

T. koordersii

/.' hainanense

Pinnae of transition

elliptic or ovate, symmetric

lanceolate, asymmetric at

lanceolate, asymmetric at

bathyphylls

at base, 5 to 9 on basis-

base, 2 to 6 on basiscop-

base, 8 to 14 on basis-

copic side of rachis and

it: side ol rachis and 2 to

copic side of rachis and

0 to 2 on acroscopic side

5 on acroscopic side

4 to 12 on acroscopic
side

Petiolules of pinnae of
sterile acrophylls

0—5 mm long

0—5 mm long

2—7 mm long

Scales on pinnae of

lanceolate, eiliate, ca. 0.6

lanceolate with branches at

substellate, appressed, ca.

acrophylls

mm long on midribs, and

base, curled, ca. 1 mm

0.2 mm diam., on midribs

substellate, ca. 0.2 mm
diam. on veinlets

long, only on midribs

and veinlets (Fig. ID)

Fertile pinnae

14—20 cm long, 2—3 mm

20-24 cm long, 2.5—3 mm

13—16(18) cm long, 0.8 mm

wide

wide

w ide (Fig. 1 F)

rachis and 4 on the acroscopic side, the basal pinna
like those in other stage bathyphylls, reflexed and
overlapping the rhizome at a right angle, the 1 pin¬
nae on the acroscopic side near the apex of fronds;
fronds to develop next ea. 3.5 cm long, with 7 pin¬
nae evenly distributed on the basiscopic side of
rachis and 3 on the acroscopic side: 1 at the base,
1 on the distal portion, and 1 near the apex (Fig.
1 A); rachis winged throughout, both surfaces of
fronds scaly, the scales substellate with branches
less than 0.2 mm long, like those on mature plants
but smaller, with fewer branches and fewer cells in
the center, the scales on the joint of rachis and
rhizome lanceolate, ca. I mm long and 0.5 mm
wide at base.

Dimidiate bathyphylls. Fronds 3—10 X 0.5— 1.5
cm. the terminal part shallowly or deeply lobed,
usually 7 to 9 pinnae presenting on the basiscopic
side ol rachis and no pinnae on the acroscopic side,
rachis narrowly winged throughout; pinnae 0.5—1
X 0.5 cm, ± flabellate or ± rhomboid, crenate on
the margin, rounded at apex, upper base subtrun¬
cate, lower base narrowly euneate (Fig. IB).

The form of the dimidiate fronds is not constant.
The 2 to 6 pinnae near the apex of the fronds are
inserted on the acroscopic side of the rachis in
some cases. This stage bathyphylls in some plants
and in branches above ground absent.

Transition bathyphylls. Fronds 10—15 X 2—4
cm, sessile, asymmetric at base, 2 to 3 acroscopic
basal pinnae usually absent, the terminal pinna
jointed to the rachis; rat his narrowly winged
throughout, with 8 to 14 pinnae on the basiscopic
side of rachis and 4 to 12 pinnae on the acroscopic
side; pinnae 1-2.5 X 0.7—1 cm, crenate, round or
blunt at apex, upper base truncate or broadly eu¬
neate, lower base narrowly euneate, veins usually
forked, the end not reaching the margin (Fig. 1C);

laminar tissue papyraceous or subcoriaeeous, the
adaxial surface glabrous, the abaxial surface dense¬
ly covered with minute substellate scales along
midribs and veinlets.

A population of about 20 plants was observed in
primary rain forest on the north slope in Mt. Tie-
dingling on Hainan Island, China. Early stage and
dimidiate stage bathyphylls were presented only on
stems creeping on the ground. Stems bearing next
stage fronds were observed as climbing on tree
trunks. When the rhizomes have ascended to 2—3
m, the fronds on the middle and lower parts of
stems wither and are shed. Fertile fronds are pro¬
duced near the apex of high-climbing stems. When
fertile fronds are present, except for fertile pinnae
and several sterile fronds, the other lower fronds
are absent.

Of the known species of Teratophyllum. the nor¬
mal bathyphy 11 stage is absent only in T. ludens and
T. koordersii (Holttum, 1932). Teratophyllum hain¬
anense is the third species without the normal bath-
yphyll stage (fronds having pinnae equally on both
sides, and the pinnae being gradually smaller in
size toward apex, the frond apex not being articu¬
late). It is not close to T. ludens. but instead to T.
koordersii. The distinguishing characters between T.
ludens, T. koordersii, and T. hainanense are given
in Table 1 .

Paratype. CHINA. Hainan Island: Qiongzhong
County, N slope of Ml. Tiedingling, similar to holotype, 25
May 2002, ,S. Y. Dong & Z. C. Chen 639 (PE).

Acknowledgments. This study is supported by
Kadoorie Farm & Botanic Garden of Hong Kong
and the Chinese Academy ol Sciences. We are
greatly indebted for the loan of collections from F
and P. We also thank Wang Wen-Tsai for preparing

108

Novon

the Latin diagnosis. Sun Ying-Bao for drawing the
illustrations, and Xiao Yin-Hou for SEM assistance.

I .iterature Cited

Holttum, It. E. 1932. On Stenochlaena, hnnariopsis and
Teratophyllum in the Malayan Region. Card. Bull.
Straits Settleni. 5: 245—313.

- . 1966. The genera Lomariopsis, Teratophyllum,

and Lomagramma in the islands of the Pacific and Aus¬
tralia. Blumea 14(1): 215—219.

- . 1978. Lomariopsis group. Pp. 255—330 in R. E.

Holttum (editor). Flora Malesiana ser. II. Vol. 1, part 4.
Martinus Nijhoff & W. Junk, the Hague.

Kramer, K. U. 1990. I xnnariopsidaeeae. Pp. 164—172 in
k. kubitzki (editor), The Families and Genera of Vas¬
cular Plants, Vol. I. Pteridophytes and Gymnosperms.
Springer- Verlag, Berlin.

A New Combination in Oxypolis Rafinesque (Apiaceae)

,/. R. Edmondson

National Museums Liverpool, Liverpool Museum, William Brown St, Liverpool LB BEN,
United Kingdom, john.edmondson@liverpoolmuseums.org.uk

ABSTRACT. Oxypolis ternata (Nutlall) A. Heller
(Apiaceae) is superseded by the new combination
0. denticulata (Baldwin) J. R. Edmondson, which
is lectotypified herein.

Key words: Apiaceae, lectotypification, North
America, Oxypolis.

While preparing a catalog of a collection of Wil¬
liam Baldwin’s plants from Georgia in the Smithian
herbarium of the Linnean Society of London
(LINN), I became aware that a species of Apiaceae,
Peucedanum ternatum Nuttall, published in July
1818, was synonymous with Siam denticulatum
Baldwin (Baldwin in Elliott, October 1817), which
was represented at LINN by an isotype.

When Rafinesque established the new7 genus Ox¬
ypolis (Rafinesque, 1825) he included Siurn denti¬
culatum under Oxypolis , but failed to make a new
combination for the species Sium denticulatum (ac¬
cording to the Gray Herbarium index, ref. Id:
60428103—2). The genus itself was subsequently
lectotypified by N. L. Britton (in Britton & Brown,
1913, vol. 2: 637) with the type species 0. rigidior
(L.) Rafinesque (Sium rigidius L.) as the generitype.

Oxypolis ternata (Nuttall) A. Heller is the com¬
monly cited modern name (see, for example, Dun¬
can & Kartesz, 1981: 99) for Sium denticulatum,
but as this combination is based on an epithet pub¬
lished later than the one which appeared in Elliott’s
work, it is contrary to Article I 1.4 of the rules of
nomenclature (Greuter et al., 2000). It is therefore
necessary to make a new combination:

Oxypolis denticulata (Baldwin) J. R. Edmondson,
comb. nov. Basionym: Siurn denticulatum Bald¬
win, in Elliott, Sketch Rot. S. Carolina, 1: 354
(Oct. 1817). Peucedanum ternatum Nuttall, Gen.
N. Amer. PI. 1: 182 (July 1818). TYPE: U.S.A.
Georgia: W. Baldwin s.n. (lectotype, designated
here, LINN-Smith no.: 508.5).

I have chosen the Linnean Society specimen as
lectotype as it is the only one I have personally
examined; Elliott did not typify Baldwin’s name as
the type concept had not then been established.

Acknowledgments. This work was based orr the
Linnean S<x:iety’s database of their Smithian Herbar¬
ium, compiled by Claire Sedgwick, to whom I am
greatly indebted for creating this invaluable resource.
I am also grateful to Kanchi Gandhi (Harvard Uni¬
versity) for checking my nomenelatural note on Sium
denticulatum in the manuscript catalog.

Literature Cited

Britton, IN. I,. & A. Brown. 1913. An Illustrated f lora of
the Northern United States, 2nd ed., 3 vols. Charles
Scribner’s Sons, New York.

Duncan, W. H. & J. T. Kartesz. 1981. Vascular Flora of
Georgia: An Annotated Checklist. Univ. Georgia Press,
Athens.

Elliott, S. 1816—1824. A Sketch of the Botany of Soulh-
Carolina and Georgia. 2 vols. J. R. Schenck, Charleston,
Georgia.

Greuter, W., J. McNeill, F. R. Barrie, II. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). H. Nicolson, P. C. Silva,
J. E. Skog, I5. Trehane, N. J. Turland N I). E. Hawks-
worth (editors). 2000. International Code ol Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Nuttall, T. 1818. The Genera of North American Plants,
and a Catalogue of the Species to the Year 1817. 2 vols.
Printed by D. Hearlt, Philadelphia, Pennsylvania.
Rafinesque, C. S. 1825. Neogenylon, or indication of sixty-
six new genera of plants of North America. Published
by the author, Lexington, Kentucky.

Novon 15: 109. 2005.

Cyanotis repens (Commelinaceae): A New Species from

Tropical Africa

Robert R. Fa den

Department of Botany, MRC-166, National Museum of Natural History,
Smithsonian Institution, P.0. Box 37012. Washington, D.C. 20013-7012, U.S.A.

faden.robert@nmnh.si.edu

David M. (Aimer on

Natural Science Division, Pepperdine University, Malibu, California 90263, U.S.A.

David.Cameron@pepperdine.edu. Current address: The University of Iowa,
College of Medicine, Department of Pharmacology, Iowa City, Iowa 52242, U.S.A.

Abstract. Cyanotis repens, a mat-forming peren¬
nial. is described from Kenya in tropical Africa. It
differs from related species by its prostrate habit,
indeterminate flowering shoots, and fusiform swell¬
ing terminating the style. Two subspecies are rec¬
ognized, diploid C. repens subsp. repens from Ken¬
ya. Tanzania, and Uganda, and polyploid C. repens
subsp. robusta from Kenya, Tanzania, Uganda.
Rwanda, Democratic Republic of Congo, and Ga¬
bon. Cyanotis repens belongs to the Cyanotis foe-
eunda species group, the species of which have a
basic chromosome number of x = 13 and often
distinctive karyotypes.

Key words: Commelinaceae, Cyanotis, tropical
Africa.

W ith approximately 25 species (of 50 worldwide)
in Africa and Madagascar, Cyanotis I). Don is the
third largest genus of the Commelinaceae in the
region (after Commelina L. and Aneilema R.
Brown). The genus and several of its species are
widespread outside of forested habitats. There is no
major concentration of species, but endemism is
relatively high in the flora of southern Africa and
flora of West Tropical Africa and low in other parts
of the African continent (Faden, 2001). The new
species described below has been known to the se¬
nior author since the 1960s, and it has sometimes
been cultivated as “Cyanotis somaliensis” perhaps
even longer.

Cyanotis repens Faden & I). M. Cameron, sp. nov.
TYPE: Kenya. Kilifi District: Nwarakaya [ =
Mwarakaya], 3°47.5'S, 39°42'K, 150 m. lime¬
stone rocky outcrops, 21 Nov. 1978, J. P. M.
Rrenan, J. //. Brenan, ./. B. Gillett et al. 14672
(holotype, K; isotypes, EA, US). Figure 1.

Herba perennis base distincta carens plerumque tege-
tem plus minusve prostratam formans. Folia baud decres-
centia apicem versus surculorum florentium; vaginis foli-
aribus 3—13 mm longis, 2.5— 9.0 mm latis. Inflorescentiae
omnes sessiles axillares in vaginis foliaribus inclusae.
Flos stylo glabro in tumorem terminalem fusiformem de-
sinente.

Perennial herb lacking a distinct base, usually
forming a ± prostrate mat; roots thin, fibrous;
shoots sparsely to densely branched basally, un¬
branched distally, repent to decumbent and rooting
at nodes; intemodes to 6 cm long, densely pilose
or sometimes slightly woolly to glabrescent or gla¬
brous, often red to maroon; flowering shoots distal.
Leaves distichous, not decreasing in size distally
on the flowering shoots; sheaths 3—13 X 2.5— 9.0
mm, sparsely to densely pilose or slightly woolly,
ciliate at apex, lamina sessile, succulent, ovate to
ovate-elliptic, narrowly elliptic or lanceolate-ob¬
long, (0.7 — ) 1 — 6(— 9.5) X 0.5— 1.8 cm, the apex
acute, rarely acuminate, sometimes ± mucronate,
the base broadly euneate to rounded or cordate,
often amplexicaul, the margins ciliate. the surfaces
subglabrous to densely pilose, the adaxial lustrous
green, often dotted with maroon, the abaxial often
maroon or suffused with maroon. Inflorescences
sessile, axillary, contracted, 1- to several-flowered
cincinni, enclosed in leaf sheaths, borne at many
nodes; bracteoles scarious, subulate or linear-ob-
lanceolate, 1.5— 8.5 mm long, glabrous or pilose.
Flowers bisexual, 4—8 mm wide, sepals 5.0— 9.4 mm
long, fused basally for 2. 8-5.4 mm, the lobes su¬
bulate, oblanceolate to obovate, elliptic or linear,
densely pilose, the apex acute; corolla 8.0-10.8
mm long, the tube 4.7— 6.7 mm long, the lobes ovate
to obovate, 3.1— 5.0 X 2.5— 4.5 mm, bluish purple
or blue-violet, rarely pink, the apex acute to obtuse;
stamens 6, fertile, equal, filaments 9.8—11.8 mm

Novon 15: 110-116. 2005.

Volume 15, Number 1
2005

Faden & Cameron

Cyanotis repens from Tropical Africa

111

Figure I. Cyanotis repens Faden & D. M. Cameron. A— L. Cyanotis repens subsp. repens. — A. Habit. — H. Inflores¬
cence. — C. Inflorescence, leaf removed. — I). Flower, lateral view. — E. Flower, front hall removed. — F. Stamen. —
G. Anther, showing dehiscence by basal pores. — H. Gynoecium. — I. Capsule. — J. Capsule, sepals removed. — K.
Seed, ventral view. — L. Seed, dorsal view. M. Cyanotis repens subsp robusta. — M. Inflorescence. A— 1. from Cameron
02/02 ex Faden & Faden 74/1174 (cultivated); \1 from Cameron 02/07 ex Faden. Frans A' Lye 69/1067 (cultivated).

112

Novon

long, with an ampulliform or fusiform subapical
swelling, pale hlue-violet, densely bearded distally
for 2.2— 3.3 mm with white-tipped, blue or bluish
purple, moniliform hairs, the anthers oblong-ellip¬
tic to ovate, 0.5-1 .0 X 0.6-0.9 mm, dehiscing by
basal pores, entirely yellow; ovary 1.1 — 1.7 mm
long, densely pubescent at or near the apex, the
style 7.5—11.0 mm long, whitish, glabrous, with a
terminal fusiform swelling 1.2— 2.1 mm long. Cap¬
sules trilooular, trivalved, 2.0— 3.7 X 1.0— 1.5 mm,
the apex hirsute; seeds (1 or)2 per locule, oblong-
ellipsoid to ovoid-ellipsoid, 1. 1-2.0 X 0.75—1.05
mm, the testa brown or reddish black, finely striate
longitudinally, with transverse pits and grooves.

Two subspecies art' recognized.

Key to the Subspecies of Cyanotis kepens

la. Leaf blades mostly 1.0— 2. 5 cm long, rarely to 5.2
cm long; sheaths 3— 5(— 7) X 2. 5—4.0 mm (1.0—

6.5 mm wide when pressed); inflorescences I - or
2(or 3)-flowered (to 5-flowered in cultivation);
bracteoles 1.5— 2.5 mm long; filaments 9.8—10.9
mm long; capsules 2.0— 2. 6(— 3.5) mm long; seeds
1.10— 1.25(— 1.50) mm long . subsp. repens

lb. Leaf blades mostly 2-6 cm long, the longest gen¬
erally at least 4 cm long, sheaths 5—13 X 5—9
mm (3.5— 7.0 mm with* when pressed); inflores¬
cences 5- or more-flowered; bracteoles 4.0— 8.5
mm long; filaments I 1.2—1 1.8 mm long; capsules
2.6— 3.7 mm long; seeds 1.25— 2. (K) mm long . .

. subsp. robusta

la. Cyanotis repens Laden & I). M. Cameron
subsp. repens. Figure I A— L.

l,eaf blades 1 .0-2.5(-5.2) X 0.5-1.0(-1.8) cm;
sheaths 3— 5(— 7) X 2.5-4.0 mm (1.0-6.5 mm wide,
when pressed); inflorescences usually l-or2(or3)-
Howered (to 5-flowered in cultivation); bracteoles
1.5— 2.5 mm long; filaments 9. {{—10.9 mm long; cap¬
sules 2.0— 2. 6(— 3.5) mm long; seeds 1.10— 1 ,25( —

1 .50) mm long, the testa sometimes reddish black.

Ilahitat. Dry or moist forest and forest edges,
thickets, bushland, river banks, woodland, moist
grassland; often associated with rocky outcrops, es¬
pecially limestone, almost always in shade or par¬
tial shade, in sandy soil to clay; just above sea level
to L000(— 1980) m; flowering plants have been col¬
lected in all months, with a slight peak from July
through October.

Chromosome number. 2 n — 20 ( l aden & Suda,
1980, as Cyanotis sp. A; Laden, unpublished).

Distribution. Southeastern Kenya (K7— flora
districts of Polhill, 1988), northeastern Tanzania
(T3, 0), and eastern Uganda (U3). Ligure 2.

Paratypes. KKNYA. Kilifi: Cha Simba, 22 km SW of

Kilili, R. R. Adams 98 (EA, K); Kaloleni— Kilifi rd. (via N
rd.) ca. 14 km, R. li. Faden A A. Evans 70/785 (EA, US);
Sokoke Forest, rd. to Jilore Forest Station, 3.2 km from
turnoff on Kilifi— Malindi rd., R. R. Faden A A. J. Faden
71/715 (EA, US); Chasimba, Mariakani— Kaloleni rd. ca.
km 14, ca. 13 km NE ol Kaloleni, R. R. Faden & A. J.
Faden 71/708 (EA, k. US); Mariakani-kaloleni rd., 8.6
km, R. R. Faden & A. J. Faden 71/884 (EA. k. US); Sala
Gate— Malindi rd., 7 km tow. Sala Gate from jt. w/Bamba—
kakoneni rd., R. R. Faden A A. J. Faden (EA, k. US);
Arabuko, R. M. Graham R070 \ K2061] (EA, k); Sokoke
Forest, G. W. Jeffrey K257 (EA); Sokoke, G. Hi Jeffrey
K886 (EA, k); Jaribuni (Kauma), R. M. Musyoki A 0. J.
Ilansen 906 (EA. k); Marafa, 25 mi. [40 km) NW of Mal¬
indi. R. M. Polhill A ,S. Paido 788 (EA, k); (ilia Simba,
.S'. A. Robertson 5217 (EA. k, MO. US); Mangea Hill. .S'.
A. Robertson 5218 (EA. k. MO, US); Mwarakaya, S. A.
Robertson A' V. C. Gilbert 8700 (EA, k(: Ndzovuni (Jorge
at ford 7 km N of Mtsengo, .S’. A. Robertson A' (J. Luke
0100 (EA, k, US). 0161 (US); ex Dakabuko Hill on Gal¬
ana Hanch, ,S. A. Robertson A (J. Lake 0409 (EA. US).
Rilifi/kwale: Giryama & Tsimba Mountains. 1887, IE
Taylor s.n. (BM). Kwale: Mwachi Biver, P. R. (). Rally
18988 (EA); Buda Mafisini Forest, 8 mi. 1 13 km| WSW
of Gazi, R. R. Drummond A J. II. Hemsley 8945 (K);
Shimba Hills Reserve, beside path to Sheldricks Falls. A.
Faden 8/82 [grown at Smithsonian Institution, pressed as
I). \L Cameron 02/12 (US)]; Mganemi, 2.8 km from Sam-
buru on Voi rd., R. R. Faden A II. Reentje 85/08 (EA, k.
US) [also grown at Smithsonian Institution, pressed as I).
M. Cameron 02/04 (US)]; Lungalunga-Vanga rd. ca. 0.5
km, R. R. Faden A A. J. Faden 77/805 (EA. E. k. MO.
US, WAG); Mwachi Forest, N end, along Mwachi River,
R. R. Faden et al. 77/491 (EA); Mombasa— Nairobi rd.,
2.5 km tow. Mombasa from turnoff to Maji ya Chumvi
Railway Station, R. II. Faden A A. J. Faden 77/575 (EA.
E, US); Nairobi-Mombasa hwy., ca. 8 km E of Makinnon
Stn., P. Kueliar 7827 (EA); Maluganji For. Res. (including
kaya Mtae), S. A. Robertson A' (J. Luke 0017 (EA, US).
Lamu: Garsen— Witu rd.. near Nyangoro bridge, R. R. Fa¬
den A A. J. Faden 74/1154 (EA, k, MO); Lunghi Forest
Reserve (proposed), (J. Luke A” .S’. A. Robertson 1455 (EA).
Tana River: Garsen— Malindi rd., ca. 8 km from Garsen,
R. II. Faden 72/281 (EA. k, l)S); Galole— Garsen rd., 13
km from Galole. R. R. Faden A A. J. Faden 74/1084 (EA.
k, MO); Malindi— Garsen rd., 0.8 km tow. Garsen from
turnoff to Kibusu, R. R. Faden A A. J. Faden 74/1174
(EA, k, MO) [also grown al Smithsonian Institution,
pressed as I). M. Cameron 02/02 (US)]; SE of Garsen on
Garsen— Malindi rd., P. J. Greenway in EA11007 (EA); 2.7
km N of Lango ya Simba, E of new dyke, .S’. .4. Robertson
A Q. Luke 5800 (EA). Leila: Mt. Kasigau, lower slopes
of S side above Bungule, R. R. Faden A A. Evans 70/
900A (EA. k, MO); Voi-Taveta rd.. 18 km tow. Taveta from
Nairobi-Mombasa rd. turnoff, R. R. Faden A A. J. Faden
74/584 (k, EA, MO); 18 km on Voi-Taveta rd., from Nai¬
robi-Mombasa rd., at yellow sign post CH 620, R. R. Fa¬
den A A. J. Faden 77/812 (K, MO. US); Taita Hills, Msau
River Valley, Taita llills Expedition 742 (EA) [also grown
at Smithsonian Institution, pressed as I). M. Cameron 02/
08 (US) |. TANZANIA. I’wani Region. Hagainoyo:
1’ongwe | lores! reserve], R. J. //. Harris , 7. Pdcs A 7. M.
Tadros in R. ./. 77. Harris 4410 (K). Tanga Region. Pan-
gani: Segera Forest, Faulkner 8880 (K). Faulkner 4001
(k). Ilandeni: Kwa Mkono, 30 km E of Handeni, Arcli-
bold 2228 (K); 4.4 km tow. Maramba from Daluni turnoff
on Maramba— Korogwe rd., R. R. Faden et al. 74/829

Volume 15, Number 1
2005

Faden & Cameron

Cyanotis repens from Tropical Africa

113

(MO); bridge over Msangasi stream, ca. 50 km straight S
of Korogwe (rd. to Dar es Salaam), Flock 463 (EA. K).
Lushoto: Mtai Forest Reserve, Mbunta, Kindeketa el al.
114 (MO. US). Liisliolo/llundeni: Korogwe— Handeni rd.,
Faulkner 1329 (K). Tanga: Perani forest, SE Umba
steppe, R. B. Drummond & J. H. Hemsley 3714 (K); Mu-
heza— Amani rd.. 1.5 km tow. Muheza from Kisiwani (near
Sigi), R. B. Faden & A. ./. Faden 74/343 (EA, k, MO);
Usambaras, Diiga, G. Holst 3163 (K); Tanga, E. Organ in
EA149I2 (EA. k). UGANDA. Mhale: Kapchorwa, F.
Lind 345 (K).

lb. ( Cyanotis repens sulrsp. robusta Faden I).
M. Cameron, suhsp. nov. TYPE. Kenya. Narok
District (Masai District of Polhill, 1988): Nger-
endei. near Mara River. 5500 ft. |1676 in],
edge of water course associated with Euclea,
Olea thicket, 13 Apr. 1961, [P. E.\ Glover, | M.
D. \ Gwynne & Samuel [ Kibuwa ] 463 (holotype,
k; isotypc, FA). Figure 1M.

A subspecie typiea foliorum laminis longioribus et va-
ginis longioribus latioribusque, inflorescentiis minimum 5-
floris, bracteolis longioribus, fibs staminum longioribus
atque capsulis seminibusque plerumque longioribus differ!.

Leaf blades ( 1 .5— )2— 6(— ' 9.6) X 0.8— 1.5 cm (the
longest generally at least 4 cm long); sheaths 5—13
X 5—9 mm (3.5— 7.0 mm wide when pressed); inflo¬
rescences 5- or more-flowered; bracteoles 4.0— 8.5
mm long; stamen filaments 11.2—11.8 mm long;
capsules 2.6— 3.7 mm long; serais 1.25—2.00 mm
long, the testa brown.

Habitat. Thickets, thicket edges, forest edges,
stream and river banks, lake edges, sometimes as¬
sociated with rocks, shade or partial shade, sandy
soil to clay loam; 700— 1820(— 2130) in. Flowering
March through December.

Chromosome number. 2 n = 78 (Faden & Suda,
1980. as Cyanotis sp. B); 2 n = 52 (Faden, unpub¬
lished).

Distribution. Northwestern Tanzania (IT),

southwestern Kenya (K5, 6), southern Uganda (U2,
4), Rwanda, Democratic Republic of Congo, and
Gabon? (see discussion). Figure 2.

Paratypes. DEMOCRATIC REPUBLIC OF CONGO.
It. •ni: Parc Nat. Albert. Mutsora riv. Talya ( a f f I. droit
Lumu [= l.ume]), G. F. de Witte 10534 (lilt, US). Riilun-
gu: vers Makamba, L. Pauwels 5562 (BR). I.liofa: Ipamu,
II. Yanderyst 12857 (BR). Katako kombe: Katakokombe
(Maniema), ,/. Lebrun 6116 (BR, US). Kinshasa: Dolo,
Bavicchi 275 bis (BR); I 'res de Menkao. II. Breyne 2368
(BR); Kinshasa, University Lovanium Plateau des Eacul-
tes, C. Evrard 6426 (BR, MO). Kole: Bena Dibele. ./.
Claessens 204 bis (BR). Kutu: Mongonjiale, Flamigni
6273 (BR). Madimha: Mpese, II. Yanderyst 34673 (BR)
& 39007 (BR). Mamhasa: Ituri, ca. 14 km SSW of Epulu
and 2 km S of Tito River, R. E. Gereau et al. 5282 (MO,
US). District Unknown: Kasai. Achten 526b (BR); Pare
Nat. Albert, Marais de la Malebolau. aid. Ilimba. G. F. de
Witte 10789 (BR); Marende inselberg, on Bukavu— Kisan¬
gani rd., S. Porembski & 13. Biedinger 1810 (US) [also
grown at Smithsonian Institution, pressed as I). M. Cam¬
eron 02/05 (US)]: Kashindi, Terashima 2 1 3 (BR). Locality
Unknown: Demeuse 96 (BR), L. Mastrornatteo 1062
[grown at Smithsonian Institution, pressed as I). M. Cam¬
eron 02/11 (US)]. GABON. District Unknown: Libre¬
ville, Baudon 108 (BR). KENYA. kisumn-Londiani:
Kisumu, Opiko in B. R. 0. Bally B664 (EA, K). Masai:
Masai Mara Res., Telek River, P. R. 0. Bally B5269 (K);
Masai Mara Nat. Park, in riverine forest bordering Mara
River at Hippo Lookout, A. Faden 4/79 |grown at Smith¬
sonian Institution, pressed as I). M. Cameron 02/10 (US)];
Masai Mara, near res. st., M. G. Gilbert 5734B (EA, K).
RWANDA. Byuniha: Reg. Du Mutara. env. de Nimuli,
reserve IRSAC, Biumba, G. Troupin 3306 (K). kibungo:
Parc Nat. de 1‘Akagera, plateau de Matinza, G. Bouxin &
M. Radoux 13RS/210 (BR. EA). TANZANIA. Bukoba:
Minziro Forest Res., Kalagara-Bikunyu Subvillage, L. Fes-
to 1807 (US); Rwasina |= Ruasina| to Kiziba. ./. Ford 493
(K). UGANDA. Bunyoro: Murchison Falls Nal. Park.
Murchison Falls, R. B. Faden, A. Evans A’ K. Lye in R.
B. Faden 69/1060 (EA); 2.3 mi. [37 km] N of Rutiaba
turnoff on Butiaba— Murchison Falls Nat. Park rd., 6 mi.

1 10 km| N of Sambive River crossing, near Kisansya, It
B. Faden. A. Evans A' K. Lye in R. B. Faden 69/1067
(EA) [also grown at Smithsonian Institution, pressed as I).
M. Cameron 02/07 (US)], kigczi: Queen Elizabeth Nat.

114

Novon

Park (just outside). South Maramagamho Central Forest
lies., 1 mi. [1.6 km | up rd. Kaizi-Bitereko off the Katun-
guru— Ishasha rd. (Congo Rd.), B. B. Baden, A. Brans &
K. Lye in B. B. Faden 69/1095 (FA), \1asaka: ea. 6 mi.

1 10 km | Masaka-Bukakata rd.. C. 11. >. Kabuye & T. May-
anja 294 (EA, K); Buddu County, Lake Kaganja, E side,
K. Lye 2620 (EA). Mengo: Bavum County, Victoria Is¬
lands, near Kitamiro, II. C. Dawkins 1)406 (EA); Entebbe,
T. I ). Muellands 106 (k): Site 362, 8.29 mi. [13.34 km]
\\ of Lwampanga, I. Langdale- Brown 2170 (k). Miiben-
de: Wattuba, Singo County, A. B. Katende k95 (k); 10 km
NW of katera. K. Lye 2029 (k). Toro: Queen Elizabeth
Nat. Park. J. M. Dick 60/257 (EA); Ruwenzori, F. Scott
Filial 70I0A (k). District Unknown: sin loc., B. Dim¬
mer 2600 (BM, k).

The collections Faden & Faden t 1/B04 and Kin-
deketa el al. 114 are unusually large for Cyanotis
repens subsp. repens. Faden & Faden 71/804 has
the longest and broadest leaves (to 5.2 cm long and
1 .8 cm wide in different leaves) and the longest and
broadest sheaths (to 7 mm long and 6.5 mm wide
pressed) for this subspecies. It also has up to 3-
flowered, instead of the usual 1- or 2-flowered, in¬
florescences. The capsule matches the longest seen
in any other collection of subspecies repens (3.5
mm long), but the seeds (1.15-1.35 mm) fall in the
overlap between those of the two subspecies. If this
plant were from the range of subspecies robusta,
there is no doubt that we would have identified il
as such. Faden cV Fallen 71/834 was collected in
Acacia thorny deciduous bushland on sandy soil (in
sched.), a habitat drier than most other collections
of subspecies repens, but that ecological difference
seems insufficient to account for its size and ro¬
bustness. Unfortunately, no living material was
gathered at the time of the collection, so the ploidy
cannot be determined. Thus the assignment of this
collection to subspecies repens is tentative. Kindek-
eta et al. 114 has leaf sheaths to 6 mm long and 5
mm wide (pressed) and the lamina to 3.3 cm long,
but we think it is just a robust specimen of sub¬
species repens.

The collection Lind 045 from Uganda, which we
have treated as subspecies repens based on its mor¬
phology, is geographically remote from the other
collections of this subspecies (Fig. 2). It also comes
from a much higher elevation (1980 m) than any
other specimen of this subspecies. Lind 045 is out¬
side the range of subspecies robusta, too. although
it is much closer geographically to that subspecies
than to other collections of subspecies repens.

The single collection from Gabon, Baudon 108,
is from the capital Libreville, so perhaps it was
from cultivation. It is far from the nearest collection
of Cyanotis repens but, considering the frequency
of subspecies robusta in western Democratic Re¬
public of Congo (Fig. 2), an extension of its con¬

firmed range further west and north would not be
surprising.

Cyanotis repens has been known as a distinct en¬
tity for a long time and has been variously cited in
the literature. Cyanotis repens subsp. repens has
been called Cyanotis sp. A in Faden and Suda
(1980) and Cyanotis sp. nov. in Robertson and
Luke (1993). Cyanotis repens subsp. robusta has
been cited as Cyanotis sp. B in Faden and Suda
(1980), Cyanotis sp. 1 in Flore du Rwanda Sper-
matophytes (Malaise, 1988), and Cyanotis sp. A in
Upland Kenya Wild Flowers (Faden, 1994).

Cyanotis repens belongs to a group of species that
may be called tin* Cyanotis foecunda species group.
This group is characterized by the presence of only
sessile to subsessile inflorescences, a glabrous style
and. insofar as is known, a basic chromosome num¬
ber of x — 13. The C. foecunda group is mainly
African, ranging from the Arabian Peninsula to
South Africa and Madagascar and extending into
Central and West Africa. The species comprising
this group, in addition to C. repens, are: C. nycti-
tropa Defiers of the Arabian Peninsula, C. somal-
iensis C. B. Clarke of northern Somalia, C. foecunda
Hasskarl, which ranges from Ethiopia west to Cam-
moon and south to Mozambique, Zimbabwe, Bot¬
swana, and Namibia, C. speciosa (L. f.) Hasskarl,
which is distributed from Tanzania and Democratic
Republic of Congo to South Africa and in Mada¬
gascar, C. lapidosa E. Phillips of South Africa, and
probably C. robusta Obermeyer of South Africa and
C. hepperi Brenan of Nigeria and perhaps Came¬
roon. Cyanotis flexuosa C. B. Clarke from Angola,
if il proves distinct from C. foecunda, would also
belong to this group. It is noteworthy that aside
from the three relatively widespread laxa, C. foe¬
cunda, C. repens, and C. speciosa, the species in
this group tend to be local and allopatric.

Cyanotis repens appears to be most closely re¬
lated to C. foecunda. the only species in this group
with whose range il overlaps. The two species have
long been confused, especially from herbarium
specimens, because they both may have flowering
shoots with numerous sessile, axillary cineinni and
both may form mats, at least in Kenya and Tanza¬
nia. In addition, they both have similar karyotypes
(a basic number of x = 13 and all chromosomes
with subterminal centromeres) (Faden & Suda,
1980). The major differences are that C. repens al¬
ways forms mats (whereas C. foecunda only some¬
times forms mats), plants lack a definite base (vs.
having a definite base), plants lack conns (or bulbs)
(vs. sometimes having them), the flowering shoots
have indeterminate growth, i.e., an inflorescence
does not terminate the shoot (vs. determinate

Volume 15, Number 1
2005

Faden & Cameron

Cyanotis repens from T ropical Africa

115

growth, with an inflorescence always terminating
the shoot), the leaves or foliaceous bracts that sub¬
tend the individual cincinni do not decrease in size
distally (vs. strongly decreasing distally), the ter¬
minal style swelling is fusiform with a relatively
broad stigma (vs. obpyriform with a very narrow
stigma), and the plants may be polyploid or diploid
(vs. strictly diploid).

Cyanotis repens has also been confused with C.
somaliensis. Indeed, a plant received from the Mis¬
souri Botanical Garden many years ago as C. so¬
maliensis ( Cameron 02/08 , US) has proven to be C.
repens subsp. robusta. However, other plants culti¬
vated as C. somaliensis, particularly by succulent
enthusiasts, e.g., Brewerton (1977) and Keen
(1982). have basal rosettes and are clearly not C.
repens. Whether they are true C. somaliensis, which
has rarely been collected in the field, or some other
species is yet to be established. True C. somaliensis,
which was reinvestigated by the senior author at
kew in 2008, including a study of liquid-preserved
flowers, differs from C. repens in having the flow¬
ering shoots arising from a sterile basal rosette,
flowering shoots with determinate growth, with up
to 7 (vs. numerous) cincinni per shoot, with the
leaves of the flowering shoot distally decrescent,
and an obpyriform style swelling with a very narrow
stigma. The karyotype of C. somaliensis is also dis¬
tinct from that of C. repens (Faden & Suda, 1980;
Jones & Kukkonen, 1971) (see below).

Cyanotis somaliensis has also been confused with
C. foecunda. Indeed, the plant used to illustrate C.
somaliensis in Faden (1995) proved to be a white-
pubescent plant of C. foecunda from xeric condi¬
tions in eastern Kenya ( Faden & Faden 77/576,
US).

Cyanotis nyctitropa from the Arabian Peninsula
differs from C. repens in the same features as C.
somaliensis. In addition, it has thick, almost tuber¬
ous roots, instead of the fibrous roots found in C.
repens, C. foecunda, and C. somaliensis.

A final plant that has to be contrasted with Cy¬
anotis repens is C. somaliensis var. uda C. B.
Clarke, which was described from western Demo¬
cratic Republic of Congo. It is poorly understood
and is known for certain only from the type ( Cillet
2140 : holotype, BR; photos K, US). The vegetative
shoots closely resemble C. repens subsp. robusta,
but the flowering shoots are determinate (i.e., the
shoots are terminated by inflorescences), the indi¬
vidual cymes are more elongate (up to 2 cm long)
and many-flowered, and the braeteoles are larger
and more falcate. It may be just an odd specimen
of C. repens subsp. robusta, within whose range it
occurs, or a taxon closely related to this subspecies.

file type, Cillet 2140, resembles certain mat-form¬
ing specimens of C. foecunda, e.g., Cameron 02/09
(ex Faden 69/1804) from southwestern Kenya, but
the bracts subtending the individual cincinni in the
former are neither reflexed nor strongly reduced
distally, as they are in C. foecunda, and the elon¬
gate cincinni also are not present in C. foecunda.
I’he determination of specimens such as Cameron
02/09 as C. foecunda is based on their inflores¬
cences, style swelling, and diploid chromosome
count that are identical to other specimens of C.
foecunda. Thus Cyanotis somaliensis var. uda is not
C. foecunda, but its relationship to C. repens re¬
mains unclear. Moreover, Cyanotis somaliensis var.
uda shows no close resemblance to C. somaliensis.
Clarke’s notes on the type of C. somaliensis var. uda
indicate that he thought it differed from the typical
variety only by its longer and weaker stems that
were possibly due to its growing in a moister hab¬
itat. The elongate cymes of variety uda and its lack
u( a basal rosette are important differences from C.
somaliensis, and there is little likelihood of a close
relationship between these taxa.

f aden and Suda (1980) found variation in ploidy
within Cyanotis repens. A population of Cyanotis
repens subsp. repens (as Cyanotis sp. A) from the
Coast Province in Kenya ( Faden & Faden 74/! 174)
was diploid, with a mitotic count of 2 n = 26 (Faden
& Suda, 1980, fig. 18 — voucher slides for chro¬
mosome counts in the Department of Botany, Smith¬
sonian Institution), whereas a population of C. re¬
pens subsp. robusta (as Cyanotis sp. B) from
Uganda (Faden, Evans & Lye in Faden 69/1067)
was hexaploid (2 n = 78) (figs. 2 1 . 22). Subsequent
counts from three further collections of C. repens
subsp. repens, all from Kenya’s Coast Province (A.
Faden 8/82: Fallen & Reentje 85/68: and Tail a
Hills Expedition 742), were also 2n = 26 (Faden,
unpublished). In contrast, a collection of C. repens
subsp. robusta from Masai Mara National Park in
southwestern Kenya (A. Faden 4/79) had 2 n = 52
chromosomes and was tetraploid (Faden, unpub¬
lished). Thus C. repens subsp. repens appears to be
consistently diploid, whereas C. repens subsp. ro¬
busta is polyploid, either tetraploid or hexaploid.

Karyotypes appear to be important in character¬
izing and suggesting relationships among the spe¬
cies of the C. foecunda group. The karyotype of
Cyanotis repens is unusual for the genus and for the
C. foecunda group in lacking metacentric chromo¬
somes. All chromosomes in both subspecies have
subterminal centromeres (Faden & Suda, 1980).
Within the C. foecunda group this karyotype is oth¬
erwise known only from C. foecunda, which is al¬
ways diploid (n = 13, 2n = 26), whereas C. repens

116

Novon

can he diploid, tetraploid, or hexaploid (Faden, un¬
published; Faden & Suda, 1980; Lewis. 1964).

Al least some metacentrie or submetacentric
chromosomes are present in all other species of the
C. foecunda group that have been investigated: five
pairs of metacentrics and eight pairs ol subtermin¬
als in C. somaliensis (Jones & Kukkonen. 1971); at
least two pairs of metacentrics among the 13 c hro¬
mosome pairs in C. nyctitropa (Faden & Suda,
1980); and six pairs of submetacentrics and seven
pairs of subterminals in C. speciosa (Jones & Kuk¬
konen, 1971). All of these species have been re¬
corded only as diploids. Some variation in basic
chromosome number in C. speciosa was reported by
Lewis (1964).

Within Cyanotis, besides the C. foecunda group,
only Cyanotis villosa (Sprengel) Schultes f. and the
closely related C. racemosa Hasskarl, both of India
and Sri Lanka, are known to have a basic chro¬
mosome number of x = 13 (Bai et al., 1995; Faden.
unpublished; Jones & Kukkonen, 1971). Bai et al.
(1995) have shown convincingly that C. villosa has
x = 12, as well as x = 13, in India, and both basic
numbers have been found in C. racemosa from Sri
Lanka (Faden, unpublished). Plants of both species
frequently have some pedunculate inflorescences.
Thus the morphological and cytological characters
of the Indian species are different from those of the
C. foecunda group. A clear relationship between
these two species groups is yet to be demonstrated.

Cyanotis repens subsp. robusta was observed in
cultivation in Kenya for the first time in 2003.
Flowering material was collected at the Naro Moru
River Lodge west of Mt. Kenya (Faden & Faden
2003/001 , US. FA). Additional plantings were ob¬
served but not collected on the grounds of the Na¬
tional Museums of Kenya, Nairobi.

Acknowledgments. The participation of the ju¬
nior author in this project was part of the 2002
Research Training program at the National Museum

of Natural History, Smithsonian Institution. This re¬
search was supported by a grant from the National
Science Foundation Research Fxperienees for Un¬
dergraduates program. Award Number DBI-
9820303. We thank Stanley Yankowski for assis¬
tance in the laboratory, Dan Cole for help with the
distribution map, and Alice Tangerini for the draw¬
ings of the specimen.

Literature Cited

Bai. k. L., P. I. Kuriachan & C. A. Ninan. 1995. Infra¬
specific karyotype evolution in Cyanotis villosa Schultes
f. (Commelinaceae) by centric fusion. J. Cytol. Genet.
30: 109-114.

Brewerton, D. V. 1977. Cyanotis somaliensis Clarke. Kakt.
Sukk. 28(4): 98.

Faden, R. B. 1994. Commelinaceae. Pp. 303—309 in A.
I). (J. Agnew & S. Agnew, Upland Kenya Wild Flowers,
Kd. 2. East Africa Natural History Society. Nairobi.

- . 1995. Commelinaceae. Pp. 79—94 in M. Timlin

(editor). Flora of Somalia. Vol. 4. Royal Botanic Gar¬
dens. kew.

- . 2001. The Commelinaceae of northeast Tropical

Africa (Eritrea. Ethiopia, Djibouti, Somalia, and Ken¬
ya): Diversity and phytogeography. Pp. 213-231 in 1.
Friis & (). Ryding (editors). Biodiversity Research in
the Horn of Africa Region. Proceedings of the Third
International Symposium on the Flora of Ethiopia and
Eritrea at the Carlsberg Academy. Copenhagen, 25-27
August 1999. Biol. Skr. Vol. 54.

- & Y. Suda. 19(50. Cytotaxonomy of Commelina¬
ceae; Chromosome numbers of some African and Asi¬
atic species. Bot. J. Linn. Soc. 81: 301—325.

Jones, k. & I. kukkonen. 1971. The comparative cytology
of some Cyanotis species. J. Indian Bot. Soc. Golden
Jubilee Vol. 50A: 332-339.

keen, W. C. 1982. Cyanotis somaliensis. Natl. (fact. &
Succ. J. 37(2): 40.

Lewis, W. H. 1964. Meiotic chromosomes in African Com¬
melinaceae. Sida 1: 274—293.

Malaise, P. 1988. Commelinaceae. Pp. 119-142 in (5.
Troupin (editor), Flore du Rwanda Spermatophytes, Vol.
4. Mus6e Royal de l'Afrique Centrale, Tervuren.
Polhill, D. 1988. Flora of Tropical East Africa. Index of
Collecting Localities. Royal Botanic Gardens, kew.
Robertson, S. A. & W. R. (J. Luke. 1993. Kenya Coastal
Forests. World Wide Fund for Nature, Nairobi.

Three New Species of Schefflera (Araliaceae) from l he Espinhago
Range in Minas Gerais, Brazil

Pedro Fiasclii and Jose Rubens Pirani

Departamento de Botanica, Institute) de Biociencias, Universidade de Sao Paulo,
Caixa Postal 1 1461, 05422-970, Sao Paulo, SP, Brazil, pedrofiaschi@hotmail.com

Abstract. Three new species of Schefflera from
the Espinhago Range in Minas Gerais, Brazil, are
described and illustrated: Schefflera botumirimensis
Fiaschi & Pirani resembles S. glaziovii (Taubert)
Frodin & Fiaschi, but differs by its elongated inflo¬
rescence and leaflet morphology. Schefflera fruti-
cosa Fiaschi & Pirani appears to be related to S.
vinosa (Chamisso & Schlechtendal) Frodin & Fi¬
aschi, but can be distinguished by its reduced in¬
florescence with up to five partial inflorescences at
each primary branch. Schefflera villosissima F iaschi
& Pirani is similar to S. macrocarpa (Chamisso &
Schlechtendal) Frodin, but differs by its more
dense indumentum and the flowers grouped in
heads.

RKSUMO. Tres especies novas de Schefflera da
Cadeia do Espinhago em Minas Gerais, Brasil, sao
descritas e ilustradas: Schefflera botumirimensis Fi¬
aschi & Pirani assemelha-se a S. glaziovii (Taubert)
Frodin & Fiaschi, no entanto difere desta pela in-
floresceneia alongada e a morfologia dos foliolos.
Schefflera fruticosa Fiaschi & Pirani parece rela-
cionada a S. vinosa (Chamisso & Schlechtendal)
Frodin & Fiaschi, mas pode ser distinta pelas in-
florescencias reduzidas com ate 5 inflorescencias
parciais por ramo primario. Schefflera villosissima
Fiaschi & Pirani assemelha-se a S. macrocarpa
(Chamisso & Schlechtendal) F rodin, diferindo pelo
indumento mais denso e as flores agrupadas em
capftulos.

key words: Araliaceae, Brazil, Espinhago
Range, Minas Gerais, Schefflera.

fhe genus Schefflera J. R. Forster & G. Forster
includes ca. 900 species and is thus the largest in
the family Araliaceae (Frodin, 1995; Frodin & Go-
vaerts, 2003). It is widely distributed in tropical
areas, where it is best represented at high altitudes
in southeastern Asia. New Caledonia, and the
mountain ranges of the Andes.

I he current delimitation of pantropical Schefflera
has been shown to be highly unnatural and to in¬
clude four to five independent lineages; one of

these independent lineages comprises the Neotrop¬
ical species (Wen et ah. 2001; Plunkett et ah.
2004). In Brazil, the genus is represented by ca.
50 species, most of which are restricted to small
areas in the Guayana Shield (near die border with
Venezuela), the Atlantic Coastal forests of Rio de
Janeiro and Espfrito Santo states, western Amazon¬
ia, and the Central Plateau, with the highest per¬
centage of locally endemic species occurring along
the Espinhago Range in Minas Gerais.

During a taxonomic study of Schefflera in south¬
eastern Brazil (Fiaschi, 2002), several new species
have been recognized. As with most Brazilian spe¬
cies of Schefflera that were previously included in
Didymopanax Deeaisne & Plant lion (essentially
distinct by its bicarpellate ovary), the new species
described here belong to this group.

These species are from the Espinhago Range in
Minas Gerais, being “endemic to only one moun¬
tain,” a distributional pattern described by Giulietli
and Pirani (1988) that is characteristic of many
species of the compos rupestres vegetation. Descrip¬
tions of this special type of vegetation are found in
Harley (1995) and Giulietti et al. (1997).

Schefflera botumirimensis Fiaschi N Pirani. sp.
nov. T't PE: Brazil. Minas Gerais: Botumirim,
contrafortes orientais da Serra da Canastra.
trilha do Cruzeiro, 1280 m, 29 Sep. 1997 (fr),
l\. Mello Silva , 17. /.. Kawasaki A A. Rapini
1475 (holotvpe. SPF; isotypes, BHCB. K. MO,
NY). Figure 1.

Species nova a Schefflera glaziovii inflorescentia elon-
gata, pedicellis fructuum longioribus (1—2 cm longo), bas-
ibus foliolorum rotundatis differ!.

Shrubs up to ca. 2 m tall; branehlets 4—8 mm
diam., grayish-sericeous or glabrescent, nigrescent
when dried. Feaves crowded at the terminal portion
of branehlets; stipules 4—6 mm long, apex generally
bifid; petiole 13—23 cm long, glabrescent; leaflets
7 to II, deflexed, conduplicate, papery', glabrous
on adaxial surface, grayish-sericeous or glabrescent
on abaxial surface; venation broehidodromous;
main vein prominent on both surfaces, more so

Nov ON 15: I 17-122. 2005.

118

Novon

Figure 1. Schefflera botumirimensis Fiaschi & Pirani. — A. Fruiting branch. — B. Median leaflet. — C. Adaxial view
of the stipule. — I). Fruits. Drawings based on the type.

abaxially; secondary veins 7 to 9, impressed on
both surfaces; intersecondary veins absent. Median
leaflet: petiolule 1.5—2 cm long, canaliculate on ad¬
axial surface, glabrescent; blade 5.8— 8.6 X 1—2.7

cm, narrowly elliptic or oblong to ovate, apex cau¬
date, base rounded (subcordate), margin entire,
slightly revolute; basal leaflets with petiolule up to
ca. 1 cm long, these blades 5—7 X 1—1.8 cm, sim-

Volume 15, Number 1
2005

Fiaschi & Pirani
Schefflera from Minas Gerais

119

ilar in shape to the median ones, symmetric or
slightly asymmetric. Inflorescence terminal, gla-
brescent; peduncle up to ca. 3.5 cm long, rachis
absent or up to ca. 8.5 cm long; primary branches
3 or 4, 21.5—34 cm long; secondary branches 7 to
9, 4—15 cm long; tertiary branches up to ca. 3.5
cm long; ultimate inflorescence units umbellate;
floral bracts ca. 0.5 mm long. Flowers not seen.
Drupe laterally compressed, 4.5—7 X 8.5—10 mm;
pedicel 1-2 cm long; styles 2(3); pyrenes 2(3), 6—
7.5 X 4.5— 5.0 mm, indurate.

Schefflera hotumirimensis is probably endemic to
the Serra da Canastra and appears to be restricted
to an area northwest of the municipality of Botu-
mirim in the northern portion of the Espinhago
Range. Its restricted geographic range and the
knowledge of just one collection from an unpro¬
tected area suggests that this species should be
considered critically endangered (IUCN, 2001).

Schefflera hotumirimensis resembles S. glaziovii ,
differing clearly by its elongated inflorescence and
pedicels (1-2 cm long in fruit) and by the mor¬
phology of the leaflets, which are characterized by
rounded bases, in sharp contrast with those of S.
glaziovii , which have cuneate to obtuse bases.

Schefflera hotumirimensis, along with S. gardneri
and .S’, glaziovii , comprises a group of shrubby spe¬
cies characteristic of rocky outcrops of campos ru-
pestres vegetation along the Espinhayo Range in the
Brazilian state of Minas Gerais. This species group
is characterized by the presence of leaflets with the
blade slightly to strongly conduplicate and the apex
acute to caudate or cuspidate.

Schefflera fruticosa Fiaschi & Pirani, sp. nov.
TYPE: Brazil. Minas Gerais: Jaboticatubas,
Parque Nacional da Serra do Cipo, mono do
lado esquerdo da Cachoeira da Farofa, 15 June
2000 (11. fr), P. Fiaschi & F. /V. Costa 286 (ho-
lotype, SPF; isotypes, BHCB, K. MBM. MO,
NY, KB, SP). Figure 2.

Species nova a Schefflera vinosa inflorescentia brevi
paueiflora (usque ad 5 ramulis per ramum primarium),
foliolis medianis angustioribus (3.5— f 1 cm longis, 0.7— 2.7
cm latis).

Shrubs 1-1.5 m tall; branchlets 3—5 mm diam.,
densely oehraceous-yellowish to grayish-sericeous
to glabrescent; internodes up to ca. 2.5 cm long.
Stipules reduced (ca. 3 mm long), apex entire or
slightly bifid; petiole 3—13 cm long, ascendent, gla¬
brescent, slightly striate longitudinally; leaflets (1)3
to 8, ascendent, plane, papery, glabrous on adaxial
surface, oehraceous-yellowish to grayish-sericeous
to glabrescent on abaxial surface; venation brochi-

dodromous; main vein prominent on both surfaces,
more so abaxially; secondary veins 6 to 9. promi¬
nent only abaxially; intersecondary veins absent.
Median leaflet with petiolule 1—2.5 cm long, sub¬
winged; blade 3.5—1 1 X 0.7— 2.7 cm, narrowly ob-
lanceolate, apex acute to rounded (truncate), mu-
cronate, base narrowly cuneate to long-attenuate,
margin entire, revolute; basal leaflets with petiolule
up to ca. 0.6 cm long, blade 2.2-7 X 0.5— 1.3 cm,
gradually shorter than the median, symmetric to
clearly asymmetric. Inflorescence terminal, erect,
included within the foliage, oehraceous-yellowish to
grayish-sericeous to glabrescent; peduncle up to ca.
0.5 cm long, rachis absent or up to ca. 5.5 cm long;
primary branches (2)3 to 7, all but 1(2) of which
are terminal, 2.5—8.5(13) cm long; secondary
branches 1 to 5. 0.3— 5.5 cm long, bracts 2—3 mm
long: ultimate inflorescence units umbellate, with 7
to 18 flowers; floral bracts ca. 1 mm long, trian¬
gular. Pedicel 0.4—1 cm long; calyx ochraceous to
yellowish-sericeous, lacinia evident; petal color
cream-greenish, ca. 3 X 1.5 mm, elliptic, sericeous
abaxially, glabrous adaxially; filaments ca. 0.7 mm
long, anthers 1.8—2 X 0.8—1 mm, oblong and apic-
ulate; styles 2, free, ascending in flower, reflexed
in fruit. Drupe laterally compressed, 6-7.2 X 10—
11.5 mm, glabrous, green when immature, turning
dark purple at maturity; pedicel 5—8 mm long; py¬
renes (1)2(3), 5.5—8 X 4.5—6 mm, indurate.

Schefflera fruticosa is presumably endemic to the
Serra do Cipo, in the southern Espinhayo Range,
where it is found between rock outcrops in shallow
soil fields. It has been collected with flowers in
February and June and with fruits in June and Sep¬
tember.

Several features, both vegetative and reproduc¬
tive, can be used to distinguish Schefflera fruticosa
from other members of the genus in eastern Brazil,
the most remarkable being the reduced and few-
flowered inflorescences and the very narrow, ascen¬
dent leaflets. The inflorescences of Schefflera fru¬
ticosa bear both bisexual and male flowers, the first
ones essentially on the distal part of the branches
and the others more proximally, thus characterizing
an andromonoecious sexual system.

The specific epithet refers to this plant's shrubby
habit, a common feature among woody species en¬
demic to the campos rupestres of the Espinhayo
Range.

Paratypes. BRA/IF. Minas Gerais: Jaboticatubas,
Alto <la Serra da Fagoa Dourada, A1. Roque et al. 104
(CTES, F. G. R. SPF, UEC); morro do lado esq. da Cach-
oeira da Farofa, P. Fiaschi et al. 68 (SPF, U); entrada do
Canyon das Bandeirinhas, P. Fiaschi et al. 61 (SPF); San¬
tana do Riacho, estrada vicinal da Rodovia MG 010, trilha

120

Novon

Figure 2. Schefflera fruticosa Fiaschi & I’irani. — A. Fruiting branch. — B. Lateral view of stipule. — C. Floral bud.
— I). Flower (one petal removed). — F. Adaxial view of the stamen. — F. Abaxial view of the stamen. Drawings based
on the type (A, C-F) and A. Roque et al. 104 (B).

para Cachoeira do Gaviao, I’. Fiaschi & F. A. Costa 340
(HUFFS. K. SPF, UB).

Schefflera villnsissimn Fiasc hi & Pirani, sp. nov.
T\ PE: Brazil. Minas (Gerais: Serra do Espin-
ha^o, at Lapinha, ca. 19 km N of Serro, on
road (MG 2) to Diamantina, ca. 1200 in. 24
Fell. 1968 (fl). //. .S'. Irwin, II. Maxwell & I).
C. Wasshausen 20809 (holotype, K; isotypes,
NY, II B, photo SPF). Figure 3.

Species nova a Schefflera macrocarpa indumento denso
villoso, lloribus in capitulis eongestis differt.

Shrubs to 4 m tall; branchlets densely ochra-
ceous to yellowish-villous. Leaves crowded at the
terminal portion of the branchlets; stipules ca. I
cm long, entire; petiole 20—23 cm long, cylindrical,
densely ochraceous to yellowish to grayish-villous;
leaflets 7 to 9, horizontal, plane, leathery, the ad¬
axial surface with trichomes restricted to the prox-

Volume 15, Number 1
2005

Fiaschi & Pirani
Schefflera from Minas Gerais

121

Figure 3. Schefflera villosissima Fiaschi & Pirani. — A. Leaf. — B. Detail of the indument on the abaxial surface of
leaflet. — C. Ultimate inflorescence unil. — I). Floral bud. — E. Longitudinal section of the floral bud. — F. Fruit.
Drawings based on R. Mello-Silva el al. CFCR 7870 (A. B). //. Irwin el al. 20809 (C— E), and U. R. Anderson 8/8 6
(E).

i null portion of the main vein, abaxial surface
densely ochraeeous to yellowish-villous; venation
brochidodromous; main vein prominent on both
surfaces, more so abaxiallv; secondary veins 10 to
12, prominent abaxially, but impressed adaxially;
intersecondary veins absent. Median leaflet with
petiolule 3—5.5 cm long, laterally compressed,
densely villous; blade 12—12.5 X 7—8 cm, elliptic
or oblong, apex retuse to rounded, base obtuse to

rounded, margin entire, revolute; basal leaflets with
petiolule up to 0.5— 1.5 cm long, blade 8.5—9 X
2.5—4 cm, similar to median ones. Inflorescence
terminal, erect, densely ochraeeous to yellowish-
villous; peduncle ca. 1 cm long, raehis reduced;
primary branches 6, 15—43 cm long, secondary
branches 25 to 4J. 2.5-7 cm long, with only one
terminal to 10 racemosely arranged capitulate ul¬
timate inflorescence units; floral bracts ca. 1.5 mm

122

Novon

long, triangular. Flowers sessile, densely oehra-
eeous to yellowish-villous; calyx villous, lacinia ev¬
ident; petals 2.5— 3.7 X 1 .3-2.1 mm, ovate or ellip¬
tic, apex acute, glabrous adaxially, tomentose
abaxially; filaments 0.3 mm long; anthers 2. 2-2. 4
X 1. 1-1.2 mm wide, oblong, apiculate; styles 2(3),
free, ascending in flower, reflexed in fruit. Drupe
laterally compressed, 6— 7 X 9—1 1 mm, persistently
villous at apex, with two pyrenes.

I'he new species is endemic to mountains of the
Diamantina Plateau in the Fspinhayo Range. It has
been collected with (lowers in August and February,
and with fruits in January. February, April, and
July.

As commonly found in most Brazilian Schefflera ,
the inflorescences of S. villosissima have both bi¬
sexual and male flowers, being thus characterized
as having an andromonoecious sexual system.

Schefflera villosissima closely resembles 5. ma-
croearpa, differing from it by having flowers
grouped in heads (vs. ultimate inflorescence units
umbellate) and the dense villous indumentum that
covers the undersurface of leaflets and petiolules,
as well as the flowers and inflorescences (vs. in¬
dumentum not so dense).

Although one collection ( Marlins s.n., M) bears
the name Didymopanax macrocarpum var. villosis-
simutn [inod.], written in E. Marchal’s handwriting,
this name was never published. We have decided
to retain the varietal epithet since it fits well with
the material described here. As the diagnostic fea¬
tures cited above are well marked in populations
from the Diamantina Plateau, we thought it better
to recognize Schefflera villosissima as a distinct spe¬
cies rather than as a variety of .S’. macrocarpa.

Paralypes. BRAZIL. Minas Gerais: without precise
locality, C. F. P. Marinis s.n. (M); Couto de Magalhaes de
Minas, 5 km by road NF of Rio Manso Couto de Ma¬
galhaes, 960-1000 m, W. R. Anderson 8786 (UB); Dia¬
mantina, Serra do MourSo, G. Hatschhach 40869 (MBM);
diamond district, G. Gardner 4706 (BM); Felisberto Cald-
eira [= 53o Contain do Rio Preto], Curtidor. G. Hatsch-
Ixieh & /.. /. Ahumada 81681 (MBM): Rio Vermelho, Ped-

ra Menina, Serra do Ambrosio, Fspigao do Meio, R.
Mello-Silva et at. CFCR 7870 (SPF).

Acknowledgments. The authors are grateful to
Emiko Naruto for the inking of the illustrations, to
the anonymous reviewers, and to EAPESP (Eun-
dagSo de Amparo a Pesquisa do Estado de Sao Pau¬
lo) and the Margaret Mee Botanical Foundation for
financial support to the first author.

Literature Cited

Fiasehi, P. 2002. Fstudo Taxonomico do Genero Schefflera
.1. R. Forst. & G. Foist. (Araliaeeae) na Megiao Sudeste
do Brasil. Unpublished M.S. Dissertation, l niversidade
de Sao Paulo, Sao Paulo, Brazil.

Frodin, D. G. 1995. Neotropical montane Araliaeeae: An
overview. Pp. 421—430 in S. P. Churchill, H. Balslev,
F. Forero A J. L. Luteyn (editors). Biodiversity and
Conservation of Neotropical Montane Forests. New' York
Botanical Garden, New York.

- & If. Govaerts. 2003. World Checklist and Bib¬
liography of Araliaeeae. Royal Botanic Cardens, kew.
Ciuliclti, A. M. & J. R. Pirani. 1980. Patterns of geograph¬
ic distribution of some plant species from the Fspinhayo
Range, Minas Gerais and Bahia, Brazil. Pp. 39-69 in
P. F. Vanzolini & \\. R. Heyer (editors). Proceedings of
a Workshop on Neotropical Distribution Patterns. Aca¬
demia Brasileira de Cieneias, Rio de Janeiro.

- , - A If. M. Harley. 1997. Espinhayo Range

region. Pp. 397—104 in S. 1). Davis, V. II. Heywood, 0.
Herrera-MacBryde, J. Villa-Lobos & A. C. Hamilton
(editors). Centers of Plant Diversity: A Guide and Strat¬
egy for their Conservation, Vol. 3: The Americas. The
World Wild Life Fund for Nature/IUCN, Washington.
D.C.

Harley, R. M. 1995. Introduction. Pp. 1-42 in B. L. Stan¬
dard (editor). Flora of t he Pico das Almas, Chapada
Diamantina, Bahia, Brazil. Royal Botanic Gardens,
Kew.

IUCN. 2001. IUCN Red List Categories and Criteria: Vers.
3.1. IUCN Species Survival Commission. IUCN, (Band,
Switzerland, and Cambridge, United Kingdom.

Plunkett, G. M.. J. Wen & P. P. Lowry II. 2004. Infrafam-
ilial classifications and characters in Araliaeeae: In¬
sights from the phylogenetic analysis of nuclear (ITS)
and plastid (trnl.-IrnV) sequence data. PI. Syst. Fvol.
245: 1-39.

Wen, J., G. M. Plunkett, A. I). Mitchell & S. J. Wagstaff.
2001. The evolution of Araliaeeae: A phylogenetic anal¬
ysis based on ITS sequences of nuclear ribosomal DNA.
Syst. Bot. 26: 144— 167.

Validation of a New Species of Schwartzia (Marcgraviaceae) and
Synopsis of the Genus for Ecuador

Diego Giraldo-Canas

Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogota,

D.C., Colombia, dagiraldoc@unal.edu. co

ABSTRACT. A new species of Schwartzia, S. pter-
osara, is validated. This species is known only from
the vicinity of Baeza, Napo Province, Ecuador. A
key to and a synopsis of the Ecuadorean species ol
Schwartzia are provided. Four Schwartzia species
are presently recognized from Ecuador, with S. cho-
coensis and S. diaz-piedrahitae as first records for
the country.

Bushmen. Se valida una nueva especie de
Schwartzia, S. pterosara. La nueva especie se co-
noce unicamente de los alrededores de Baeza en la
Provincia de Napo (Ecuador). Se presentan una
clave y una sinopsis para las especies ecuatorianas
de Schwartzia. Asf, Schwartzia esta representado en
Ecuador por cuatro especies, siendo registradas por
primera vez para el pais S. chocoensis y S. diaz-
piedrah itae.

Key words: Ecuador, Marcgraviaceae, Noran-
tea, Schwartzia.

Schwartzia Vellozo is a small genus of the Neo¬
tropical family Marcgraviaceae comprising 17 spe¬
cies distributed in wet lowland forests or montane
rain and cloud forests from Costa Rica to southern
Brazil, and also in the Lesser Antilles. This genus
is easily distinguished from Norantea Aublet and
other related genera [Marcgraviastrum (Wittmack
ex Szyszylowicz) de Boon & S. Dressier and Sar-
copera Bedell, which constitute the Norantea com¬
plex] by its relatively short raceme with flowers that
are borne on long pedicels, and by the stalked eym-
biform, cyathiform or helmet-shaped nectaries that
are attached at or below the middle of the pedicel,
or only rarely at the base of the pedicel, e.g., S.
diaz-piedrahitae Giraldo-Canas. The flowers in
Schwartzia are larger than those found in Norantea.
A key to the four genera of the Norantea complex
was provided in Giraldo-Canas (2005).

A thorough taxonomic treatment of the Norantea
complex was the unpublished Ph.D. thesis of Be¬
dell (1985). Additional research on the group has
been carried out by de Boon and Dressier (1997)
and Giraldo-Canas (2001a, 2001b, 2001c, 2002a,

2002b. 2002c, 2003). Schwartzia pterosara, a new
species proposed by de Roon and Bedell (Bedell,
1985) is validated here. The description, Latin di¬
agnosis, and taxonomic history are based on Bedell
(1985), with some changes made herein.

Schwartzia pterosara de Roon & Bedell ex Gir¬
aldo-Canas, sp. nov. TYPE: Ecuador. Napo:
along rd. betw. Quito & Baeza, 3 mi. W of
Baeza, 0°25'S, 77°51'W, 2000 m, 2 Oct. 1980.
T. B. Croat 50284 (holotype, MO; isotypes,
GH, MARY not seen). Figure 1.

Frutex. Folia obovata (5— )8— 12.5 cm longa et (2.2— )3. 8—
6.2 cm lata basibus acutis apicibus oblusis; glandes hy-
pophyllae 50—68 in quoque folio in aequalibus seriebus
]-] 1 mm distantibus ex marginibus foliorum. Axes inflo-
rescentiarum 10—17.2 cm longi; ffores 18—42; pedicelli
3.5— 5.5 cm longi et 2-3 mm lati basin versus 4—5 mm
lati apicem versus; nectaria cyathiformia, 0.6— 1.0 cm lon¬
ga, 0.8-1. 6 cm ex basibus pedicellorum inserta; stamina
12; ovarium 5-loculatum; stigma mammiforme.

Sprawling epiphytic shrubs; branches woody and
subterete with glabrous, grayish yellow bark and
longitudinal lenticels. Leaves petiolate and coria¬
ceous, dull green above, reddish brown below, pro¬
ducing a non-ciliate fracture when broken perpen¬
dicular to the midvein; petioles 3—6 X 2—3 mm;
lamina obovate, (5 — )8 — 12.5 X (2.2— )3. 8— 6.2 cm,
base acute to cuneate, apex obtuse, mucronate or
occasionally refuse through the loss of the mueron
with 50 to 68 small- to medium-sized hypophyllous
glands per lamina in 2 ± uniform rows located 1—
1 1 mm from the margin; midvein obscure or im¬
pressed above, prominulous below, lateral veins ob¬
scure on both surfaces or prominulous below. Inflo¬
rescence a dense, multiflorous, broom-like raceme,
axis 10-17.2 cm long, with 18 to 42 flowers borne
on slender, tapered pedicels 3.5— 5.5 cm long, 2—3
mm wide basally, thickening to 4—5 mm apically,
attached to rachis at angles of 45°— 85°; foliaceous
bracts ovate, 1.4— 2.1 X 0.9-1. 2 cm, each with 3
pairs of hypophyllous glands; nectariferous bracts
leathery and somewhat succulent, greenish red, cy¬
athiform. 0.6-1 X 0. 6-0.8 cm on attenuate stalks

Novon 15: 125-127. 2005.

124

Novon

Figure 1. Schwartzia pterosam de Roon & Bedell ex Oiraldo-Canas. — A. Floriferous branch. — B. Hu<l and nectar-
ii( 'rous hract. — C. Part of corolla with attached immature stamens (from Croat 50284, holotype, MO).

Volume 15, Number 1
2005

Giraldo-Canas
Schwartzia in Ecuador

125

3—4 mm long, attached 0.8— 1.6 cm from t lit* base
of pedicels. Flowers pale green with red at maturity;
buds 0.6— 0.9 cm long; bracteoles elliptic to obovate
with membranaceous margins, (4— )8— 11 X (4— )7—
9 mm, tightly appressed to the calyx; sepals orbic¬
ular with membranaceous margins, (4— )5— 7 mm
long and wide; petals lree, elliptic to oblanceolate,
8—11 X 3—5 mm; stamens 12, 6—8 mm long; fila¬
ments free, flattened and broader apical ly, adnate
basally to the corolla; anthers subsaggitate, pollen
bright yellow; ovary pyriform or turbinate, 4—5 mm
tall. 5-locular; style cylindrical, I mm tall; stigma
smooth, mammiform. Fruit 0.9— 1.1 X 0.8—1 cm,
globose, apiculate, green or brown; seeds reniform
to elliptic, reticulate, shiny, numerous, 1 .2-1.5 X
ea. 0.7 mm.

Distribution and habitat. Schwartzia pterosara
is known only from wet Andean forests at 2000 m
in the vicinity of Baeza (Napo, Ecuador). However,
there may be more specimens at other herbaria
from Ecuador (e.g.. OCA. QCNE, GUAY ). but un¬
fortunately I do not have access to their collections.
It is hoped that increased collecting activity will
yield additional specimens of this species.

Bedell (1985) cited another collection from Co¬
lombia \Cuatrecasas 22618 (F, US)], but I have not
seen it. despite tin' Curators of F and US sending
me all their Maregraviaceae collections (with the
exception of vouchers belonging to Marcgravia) de¬
posited in these herbaria. Stefan Dressier (1 B. pers.
comm.) said there is one specimen from Peru at FR
and MO ( Campos & Nunez 4612), but I have not
seen this collection either. If these collections be¬
long to S. pterosara , the distribution of the species
is broader than stated above.

Phenology. Flowering September to October;
fruiting June to July.

Etymology. flic epithet pterosara is composed
by the Greek words pteron, feather, and saron,
broom, referring to the feathery, broom-like ap¬
pearance of the inflorescence.

Schwartzia pterosara differs from the other spe¬
cies of the genus by its large number of hypophyl-
lous glands, its dense, multiflorous raceme, tapered
pedicels, and the size and shape of the nectaries.
With these new species, four Schwartzia species are
presently recognized for Ecuador (.S’. chocoensis Gir¬
aldo-Canas, S. diaz-piedrahitae, S. lozaniana Gir¬
aldo-Canas, and S. pterosara).

Paratype. FCUADOIC Napo: vie. Baeza, 2000 rti, /..
Besse, II. Kennedy & R. Baker 1504 (MO).

A Synopsis ok tiik Genus Schwartzia in

Eoi A DOR

Schwartzia Vellozo, FI. Flumin. 5: 221. 1825
1 1829]. TYPE: Schwartzia glabra Veil. [=
Schwartzia brasiliensis (Choisy) Bedell ex Gir¬
aldo-Canas].

Sprawling terrestrial or epi- to hemiepiphytic
shrubs, heaves spiraled, subsessile or petiolate,
blades glabrous, oblong, elliptic, or obovate, occa¬
sionally asymmetrical, venation brochidodromous,
obscure or impressed above, obscure to prominent
beneath, coriaceous, with hypophyllous glands. In¬
florescence racemose, terminal, rarely lateral, mul¬
tiflorous or occasionally pauciflorous; flowers pen-
tamerous, borne on elongate pedicels, subtended by
the stalked sacciform, tubular, cymbiform, eyathi-
form, or helmet-shaped nectaries that are attached
at or below the middle of the pedicel, or only rarely
at the base of the pedicel (e.g., S. diaz-piedrahitae );
sepals 5, imbricate in 2 whorls; petals 5, free or
basally connate, reflexed at anthesis; stamens It) to
numerous, rarely 5. in I or several whorls; filaments
linear to broad and somewhat flattened, free or ba¬
sally connate and occasionally adnate to base of
petals; anthers basifixed to subbasifixed, subeor-
dale or subsagittate; ovary conical, pyriform, or tur¬
binate, completely or incompletely 3- to 5-locular;
stigma mammiform, subsessile, lobed or radiate.
Fruit capsular, globose to subglobose, apiculate
with persistent style and stigma, loculieidally and
septifragously dehiscent from the base; seeds hemi¬
spherical or reniform, reticulate, few to numerous,
and with a shiny black testa.

Vellozo (1825) described the genus Schwartzia
based on a single species, S. glabra, a small tree
of the coastal forests of Brazil characterized by hav¬
ing slightly evaginated nectariferous bracts inserted
at or near the middle of the pedicel. This tree
proved to be the previously described Norantea
brasiliensis Choisy (1824). Schwartzia was soon
placed in synonymy, and its species referred to N.
brasiliensis [= S. brasiliensis (Choisy) Bedell ex
Giraldo-Canas], Delpino (1869), however, noted the
distinctive bract position and proposed a monotypic
subgenus, Norantea subg. Cochliophyilum, for N.
brasiliensis. More recently, the genus Schwartzia
was revived by Bedell (1989) and recognized by de
Boon and Dressier (1997) when they accepted the
segregation of the Norantea complex in four genera
( Marcgraviastrum , Norantea , Sarcopera, and
Schwartzia).

126

Novon

Key to the Ecuadorean Species of Schwartzia

la. Nectariferous bracts cyathiform, 0.6—1 cm long;

50 to 68 hypophyllous glands per lamina; brac-
teoles (4— )8-l I nun long; sepals (4—) 5— 7 nun
long; petals 8-11 mm long . S. pterosara

lb. Nectariferous bracts saccate, tubularifom to glo¬
bose, 0.9— 3.3 cm long; 0 to 14 hypophyllous
glands per lamina; bracteoles 1.8—5 mm long;
sepals 2-7 mm long; petals 5-13 mm long.

2a. Nectariferous bracts attached at the base of

the pedicel; stamens 14 ... S. diaz-piedrnhitae
2b. Nectariferous bracts attached 1—1. 6 cm from
the base of the pedicel; stamens 22 to 50.

3a. Stamens 22 to 26; bracteoles ca. 2 mm
long; sepals 2.2— 2.8 mm long; ovary ca.

2.5 mm long; ovary 4-locular; urn of the
nectariferous bracts 0.9— 1.7 cm long

. S. lozaniana

3b. Stamens 50; bracteoles 2.5—5 mm long;
sepals 4—7 mm long; ovary 3-8 mm
long; ovary 5-locular; urn of the nectar¬
iferous bracts 1.5— 3.3 cm long . . .
. S. chocoensis

I . Schwartzia chocoensis Giraldo-Cafias, Revis¬
ta Acad. Colomb. Ci. Exact. 25; 478. 2001.

I Y PE: Colombia. Chocd: carr. Ansermanuevo—
Sail Jose del Palmar, 8.4 km del Alto del Ga-
lapago, 1600 m, 19 Feb. 1977. E. Forero, A.
Gentry, A. Sugden & I). Duly 3000 (holotype,
COL; isotypes. CHOCO, MO).

Description and illustration: Bedell (1985: 202-
206, as “Schwartzia foreroi” de Boon & Bedell, no-
men nudum)-, Giraldo-Cafias (200.3: 10-12).

Schwartzia chocoensis is easily recognized by its
large (lowers on long and thick pedicels, with suc¬
culent, saccate or tubularifom nectaries, and by its
long leaves. This species was known only front the
Chord region of Colombia (Bedell. 1985: 203-206;
Giraldo-Cafias, 2003: 12). Recent studies of the
Norantea complex have revealed its occurrence in
one locality in Pichincha Province, Ecuador. The
study of specimens from Colombia and Ecuador
leaves no doubt about their identity. Schwartzia
chocoensis occurs as a sprawling shrub in different
types of wet forests below 1600 m. “Schwartzia for¬
eroi" de Roon & Bedell is a nomen nudum that
appears in Bedell (1985: 202) and in Forero and
Gentry (1989: 103); it is synonymous of S. cho¬
coensis.

Additional specimens examined. ECUADOR. Pichin¬
cha: km I 10 Quito-Nono-Tandayapa-Mindo-Puerto Quito,
C. Dodson A. Embree 13091 (MO).

2. Schwartzia diaz-picdrahitae Giraldo-Cafias,
Revista Acad. Colomb. Ci. Exact. 25: 480.
2001. TYPE: Colombia. Valle del Cauea: Bajo
Calima, Concesidn Pulpapel/Buenaventura,

3°55'N-77°W. 100 m, 17 Oct. 1984. M. Mon-
salve 434 (holotype, COL; isotypes, JADM,
MO, NY).

Description and illustration: Bedell (1985: 168-
1 72, as “Schwartzia colombiana" de Roon & Be¬
dell. nomen nudum)-, Giraldo-Cafias (2003: 12—14).

This species is unusual by having its nectarifer¬
ous bracts attached at the base of the pedicel, a
feature unique within the genus Schwartzia.
Schwartzia diaz-piedrahilae was known only from
the Choco region in Colombia (Bedell, 1985: 169—
170; Giraldo-Cafias, 2003: 13). However, recent
studies of Colombian and Ecuadorean specimens
have confirmed its presence at one locality in the
province of Carehi, Ecuador. This species is a com¬
mon and sometimes abundant element of wet low¬
land forests at 0-450 m. “Schwartzia colombiana”
de Roon & Bedell is a nomen nudum that appears
in Bedell (1985: 168) and in Forero and Gentry
(1989: 103); it is synonymous with S. diaz-piedra-
hitae.

Additional specimens examined. ECUADOR. Carehi:
trail along plain above Tobar-Donoso & Rfo Guape, W.
Hoover 1254 (MO).

3. Schwartzia lozuniuna Giraldo-Cafias, Calda-
sia 23: 384. 2001. TYPE: Colombia. Narifio:
Junfn-Tumaco rd., 6-1 1 km W of Junin, road¬
side thickets & forest edge, 850—1030 m, 27
Feb. 1979, J. Luteyn & M. Lehrdn-Luteyn 6 330
(holotype, COL; isotypes, MO, NY, IJ not
seen).

Description and illustration: Bedell (1985: 198-
201, as “Schwartzia venusta” de Roon & Bedell,
nomen nudum)-, Giraldo-Cafias (2003: 15-17).

Schwartzia lozaniana is easily recognized by its
flowers borne on long and slender pedicels, its sac¬
cate nectaries, and its elliptic-obovate to oblong
leaves with acuminate to attenuate apices. This
species has been recorded in Narifio (southern Co¬
lombia) and Esmeraldas (Ecuador) (Giraldo-Cafias,
2001b, 2003) and now, too, in the province of Car-
chi (Ecuador). Schwartzia lozaniana is uncommon
in Colombia and Ecuador, and its occurrence is
limited to some wet montane forests at 400-1600 m.

Additional specimens examined. ECUADOR. Carehi:
trail to Rfo Gualpi Chico, along ridge line near Awa en¬
campment, W. Hoover et al. 2533 (MO); border area betw.
Carehi & Esmeraldas, ea. 20 km past Lita on rd. Lita—
Alto Tambo, II. van der Wetff et al. / 1992 (MO). Esmer-
aldas: Quinindf; Gantdri, Rilsa Biol. Slat., Res. Ecol.
Mache-Chindul, 40 km NW of Quinind6, Loma de los
Cuerrilleros, J. Clark et al. 3974 (COL); Lita-San Lorenzo
rd., 18 km W of Rfo Lita Bridge, on old rd. below Lita,
6.6 km W of bridge over Rfo Chuchubf, T. Croat et al.

Volume 15, Number 1
2005

Giraldo-Canas
Schwartzia in Ecuador

127

32631 (MO); Lita— San Lorenzo nl.. 10—20 km N\V of Lila,
A. Gentry el nl. 70033 (MO).

4. Schwartzia pterosara <le Room & Redell ex
Giraldo-Canas.

Acknowledgments. I thank Rosa Ortiz-Gentry
(MO), Victoria Hollowed (MO), Olga Martha Mon¬
tie] (MO), Diana Gunter (MO). Rodrigo Bernal
(COL), Carlos Parra (COL), and Enrique Forero
(COL) for valuable help, and Stefan Dressier (FR)
and Alvaro Idarraga (HUA) for providing important
references. The curators of following herbaria are
acknowledged for the loan of specimens: CHOCO,
COAH, COL. CR, CIJVC. F. HUA. IBGE, JAUM,
MEDEL. MEXU, MO, MPU, NY. PSO. RSA, SI.
SP, L IS. US, and VEN. Visits to MO and RSA were
financed by the Universidad Nacional de Colombia
(Bogota). Missouri Botanical Garden (St. Louis.
Missouri), and Rancho Santa Ana Botanic Garden
(Claremont, California). Duban Canal (COL) pro¬
vided the illustration of S. pterosara. I thank two
anonymous reviewers for their critique of the man¬
uscript. This paper is derived from the project “Es-
tudios sistematicos en el complejo Norantea Aubl.
( Marcgraviaceae)," project No. 803765 of the “Di¬
vision de Investigation (DIB) de la Universidad Na¬
cional de Colombia,” Bogota.

Literature Cited

Bedell, H. G. 1985. A Generic Revision of Marcgravi¬
aceae, 1. Ihe Norantea complex. Ph.D. Dissertation,
University of Maryland, College Park.

- . 1989. Marcgraviaceae. In: R. A. Howard (editor).

Flora of the Lesser Antilles 5: 300—310. Arnold Arbo¬
retum, Harvard University, Cambridge.

Choisv. J. I). 1824. Marcgraviaceae. In: A. P. de Candolle
(editor). Prodromus Systematis Naturalis Regni Vege-
tabilis 1: 565—566. Victoris Masson, Paris.

Delpino, G. G. F. 1869. Rivista monografica della famiglia
delle Marcgraviaceae. Nuovo Giorn. Bot. Ital. 1: 257—
290.

de Room A. C. & S. Dressier. 1997. New taxa of Norantea
Aubl. s.l. (Marcgraviaceae) from Central America and
adjacent South America. Bot. Jahrb. Svst. 119: 327-
335.

f orero, F. & A. H. Gentry. 1989. Lista Anotada de las
Plantas del Departamento del Cliocd, Colombia. Bibli-
oteca Josd Jeronimo Triana 10: 1 — 142. Instituto de
Cieneias Naturales-Museo de Historia Natural-! niver-
sidad Nacional de Colombia, Bogota.

Giraldo-Canas, D. 2001a. Schwartzia brasiliensis (Marc¬
graviaceae), nueva combinacidn. Caldasia 23: 341—342.

- . 2001b. Una nueva especie de Schwartzia (Marc¬
graviaceae) de la vertiente occidental andina de Colom¬
bia y Ecuador. Caldasia 23: 383—388.

- . 2001c. Dos nuevas especies de Schwartzia

(Marcgraviaceae) de Colombia. Revista Acad. Colomb.
Ci. Exact. 25: 477—182.

- . 2002a. Novedades taxondmicas y coroldgicas en

Marcgraviastrum, Sarcopera y Schwartzia (Marcgravi¬
aceae) de Sudamerica. Hickenia 3(32): I 19-123.

- . 2002b. Estudios en el genero Schwartzia Vellozo

(Marcgraviaceae): Una nueva especie de la Cordillera
Central Andina de Colombia. Novon 12: 456—159.

- . 2002c. Los generos Marcgraviastrum y Norantea

(Marcgraviaceae) en Brasil. Revista Acad. Colomb. Ci.
Exact. 26: 469-176.

- . 2003. Revision de las especies colombianas del

genero Schwartzia (Marcgraviaceae). Caldasia 25: 1-21 .

Vellozo, J. M. de C. 1825. Schwartzia. In: Florae Flumi-
nensis 5: 221—222. F. J. Marianus Ed., Rio de Janeiro.

Belamcanda Included in Iris , and the New Combination
I. domestica (Iridaceae: Irideae)

Peter Goldblatt

It. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 209,

St. Louis, Missouri 63166, U.S.A. peter.goldblatt@mobot.org

David ./. Mabberley

Nationaal Herbarium Nederland, University of Leiden, The Netherlands, and Royal Botanic
Gardens Sydney, Australia, david_mabberley@yahoo.co.uk

Abstract. The eastern Asian genus Belamcanda
(Iridaceae: Irideae), with its sole species, B. chi-
nensis, die leopard or blackberry lily, has long been
understood to be most closely related to Iris dicho-
toma (syn. Pardanthopsis dichotoma ), but has nev¬
ertheless been maintained as a separate genus be¬
cause of its distinctive floral, fruit, and seed
morphology. Molecular l)NA sequence evidence
shows B. chinensis and its sister species, /. dicho¬
toma, to be nested within the large Northern Hemi¬
sphere genus Iris (ca. 280 spp.). Not only does con¬
sistent taxonomic treatment of genera of the
Iridaceae require that Belamcanda be transferred
to Iris, but we argue that taxonomy should follow
the principle of monophyly, which requires that Be¬
lamcanda and any other genus nested in Iris be
treated as members of that genus. A new combi¬
nation, /. domestica (basionym Epidendrum domes-
ticum), is made for B. chinensis (based on Ixia chi¬
nensis), because the name Iris chinensis is
preoccupied. The names Belamcanda pampaninii
Leveille and B. chinensis var. taiwanensis S. S. Ying
are here included in the synonymy of /. domestica.

Key words: Belamcanda, Iridaceae. Iridoideae,
Iris, paraphyly. phytogeny.

The genus Belamcanda Adanson, now usually
regarded as comprising a single species, B. chinen¬
sis (L.) DC., the leopard or blackberry lily (Mathew,
1981), is a member of the predominantly Old World
tribe Irideae of subfamily Iridoideae of the Irida¬
ceae (Goldblatt, 1990). The tribe includes the
Northern Hemisphere genus Iris L (ca. 280 spp.)
of Eurasia, North America, and North Africa, the
largely sub-Saharan African Moraea Miller (ca. 196
spp.), the southern African Bohartia L. (15 spp.),
Dietes Salisbury ex Klatl (6 spp.). which is largely
sub-Saharan African but has one species on Lord
Howe Island in the southern Pacific region, and the
tropical and southern African Eerraria Burman ex

Miller (ea. 12 spp.). Other small, formerly recog¬
nized, largely African genera, including Barnar-
diella Goldblatt (1 sp.), Galaxia Thunberg (15
spp.), Gynandriris Parlatore (9 spp.), Hexagloltis
Ventenat (6 spp.), and Homeria Ventenat (32 sp.),
found to be nested in Moraea, have been reduced,
rendering Moraea monophyletic (Goldblatt. 1998;
Goldblatt et ah. 2002).

In contrast, the small genera clearly allied to.
and evidently nested in. Iris, including Belamcan¬
da , Hermodactylus Miller (1 sp.). and Pardanthop¬
sis (Hanee) I.. W. Lenz (1 sp.). are usually recog¬
nized in floristic accounts. Thus Iris, according to
current circumscriptions, is paraphyletie. The be¬
lief that these genera are most closely related to
particular species or species groups within Iris has
now been confirmed by molecular study using chlo-
roplast DNA regions (Tillie et ah, 2001). A consis¬
tent treatment of genera of the Iridaceae requires
that the names of these genera be treated as syn¬
onyms of Iris, and, we argue, a taxonomy that fol¬
lows the phylogenetic principle of taxonomic mono¬
phyly demands such treatment. That Belamcanda
chinensis is nested in Iris is not only amply dem¬
onstrated by molecular analysis using two chloro-
plast DNA sequences (Tillie et al.. 2001), but it is
also the most parsimonious interpretation of evi¬
dence from more classical characters. Its basic
morphology closely resembles that of Iris ( Pardan¬
thopsis ) dichotoma Pallas in its aerial, suberect rhi¬
zome that is in effect a stem, the more or less di-
chotomously branched inflorescence, and short
subequal dark green spathes enclosing the flowers
of each inflorescence unit (a rhipidium) (Mathew,
1981). Both species have indeed been referred to
the genus Pardanthus ker Gawler. In addition, both
species are fully deciduous, unlike most other,
though by no means all. Iris species. These two
species share the same, apparently derived, chro-

Novon 15: 128-132. 2005.

Volume 15, Number 1
2005

Goldblatt & Mabberley
Belamcanda and Iris

129

mosome number, 2 n = 32, and karyotype, and, de¬
spite their apparently grossly different (lowers, can
readily be crossed. Neither species can lx; crossed
with any other species of Iris tested (Simonet,
1934). Morphological similarity combined with the
biosystematic data led Lenz (1972) to segregate /.
dichotoma as the monotypic genus Pardunthopsis,
named lor its similarity to Pardanthus, a nomen-
clatural synonym of Belamcanda (Ker Gawler,
1804). The “intergeneric” hybrids between Belam¬
canda and Pardanthopsis have been named XPar-
dacanda Lenz, for obvious reasons.

I he segregation of Pardanthopsis from Iris over¬
looks its similarities to some species of Iris, in¬
cluding /. japonica Thunberg and other far eastern
Iris species that have an aerial rhizome. Tillie et
al.s (2001) molecular analysis places /. ( Pardan¬
thopsis ) dichotoma sister to Belamcanda chinensis
with strong bootstrap support (BS 98%), but there
is only moderate support for the nesting of these
two species within a well-supported clade that com¬
prises subgenus Iris, a western Asian and Kuropean
assemblage. Nevertheless, both Belamcanda and
Pardanthopsis are deeply nested in Iris, and we see
no reasonable alternative to including both in that
genus.

While the single species each of Hermodactylus
and Pardanthopsis were originally, and are occa¬
sionally still, included in Iris, as /. tuberosa L. and
/. dichotoma, respectively, Belamcanda has never
been so treated. Thus there is no available com¬
bination in Iris for B. chinensis. We remedy this
here, and formally place Belamcanda in the syn¬
onymy of Iris. According to Garay (1997), the ear¬
liest specific epithet available for transfer to Iris is
provided by Epidendrum domesticum I... and the
new combination Iris domestica is therefore provid¬
ed here. I he specific epithet from Belamcanda chi¬
nensis, based on Ixia chinensis I.., cannot be trans¬
ferred to Iris because of the name /. chinensis
Curtis, a synonym of I. japonica Thunberg, another
eastern Asian species.

SVSTKMATICS

Iris L., Sp. PI. 38. 1753. TYPE: Iris Xgerrnanica
L. (pro sp.).

Belamcanda Adanson, Para. PI. 2: 60 (as “ Belam-Canda ”)

& 524 (as “ Belamkanda ”). 1763, nom. cl orth. cons.
Syn. nov. I ^ PE: Belamcanda chinensis (L.) DC., typ.
cons.

Pardanthus Ker Gawler, Koenig & Sims Ann. Pot. 1: 246.
1804, nom. illegit. superfl. pro Belamcanda. TYPE:
Pardanthus chinensis (L.) Kcr Gawler.

Iris domestic:! (L.) Goldblatt & Mabberley, comb,
nov. Basionym: Epidendrum domesticum L.,
Sp. PI. 2: 952. 1753. Vanilla domestica (L.)
Druce, Bot. Kxeh. Club Soc. Brit. Isles 3: 425.
1913. TYPE: Kaempfer, Amoen. Exot. Fasc.
5: t. 869, fig. 1 [Angurek Warna|. 1712, based
on material given to Kaempfer by “Nic. Moel-
lerus” in Jakarta [“Batavia”], Java, Indonesia
(icon, lectotype, designated by Garay (1997)).
EPITYPE: [Europe, cultivated,] E. Duvall in
Herb. J. E. Smith 89.42 (bequeathed to Smith
in 1 798)— LIN N-SM 45, fiehe seen, designated
here).

Ixia chinensis L„ Sp. PI. 36. 1753. Belamcanda punctata
Moeneh, Methodus 529. 1794, nom illegit. superfl.
pro Ixia chinensis L. Moraea chinensis (E.) Thunberg,
H. Jap. 34 1784. Belamcanda chinensis (1,.) DC., in
Bedoute, I. iliac. 3: ad t. 121. 1805. Pardanthus chi¬
nensis (E.) Ker Gawler, Koenig & Sims Ann. Bot. I:
247. 1804, nom. illegit. superfl. pro Belamcanda chi¬
nensis | Pardanthus sinensis Van Houtte, El. Serres
.lard. Eur. 26: t. 1632. 1865—67. orthog. var. |. Gem-
mingia chinensis (L.) Kuntze, Bevis. Gen. PI. 2: 701.
1891. comb, illeg., gen. inval. TYPE [icon]: Bheede,
Port. Malalr. I I : t. 37. 1692 (lectotype, designated
here).

Belamcanda pampaninii Level lie, Bepert. Spec. Nov. Beg-
m Veg. 8: 59. 1910. TYPE: China. Guizhou: Sfiui-
yang Xian, Wangcaoba [28°12'N, I07°26'E] or
Wang-ts ao-pa ea. 107°12,E| (as Kouy-

Tcheou, Choui-mi-tsin, Hoang-Tsao-Pa), flowers yel¬
low, June 1909, Esquirol 1565 (holotype, E).

Belamcanda chinensis var. taiwanensis S. S. Ying, Col.
Illustrated Plants of Taiwan I: 237. 1980. TYPE:
Taiwan Keelung, Hopingtao, Aug. 1979, S. S. Ying
s.n. (HAST not seen).

I he type of Epidendrum domesticum is a some¬
what stylized illustration in Kaempfer (1712).
whose description is apparently based on two com¬
pletely different plants (Garay, 1997), one an or¬
chid, probably a Cymbidium species, and Belam¬
canda chinensis. In Kaempfer’s account of Japanese
plants it is one of the plants collected in 17th cen¬
tury Java by one “Nic. Moellerus,” though, remark¬
ably, unmentioned in the compendium of van
Steenis-Kruseman (1950), and given to Kaempfer
who was on the island in 1689—1690 and 1692—
1693. It was then included in Linnaeus’s Species
Plantarum, where it received its first acceptable
name. The plant was described by Kaempfer as a
scandent parasite with variegated six-petaled flow¬
ers. I he illustration, however, shows no indication
of a climbing habit: only the upper portion of a
branched flowering stem is drawn, with stalked
multi-flowered inflorescences rather crudely shown.
Ihe flowers have six mottled tepals, five of them
subequal and one irregularly twisted into what

130

Novon

could l>c mistaken for an orchid labellum. Stamens
and details of the style are not shown. Although no
confirmatory (“typotype”) specimen could he found
by DJM in Kaempfer's collection in the Sloane Her¬
barium (BM; cf. also Hinz. 2001), we agree with
Garay that the illustration is a flowering stalk of
Helamcanda with the individual rhipidia bearing
two flowers raised above the characteristically short
spathes. In interpreting the mixed illustration thus.
Garay was able to avoid upsetting orchid nomen¬
clature and. at the time when Helamcanda was con¬
sidered distinct from Iris, this action had no effect
on t lit- iridaeeous element. Here we provide an ep-
itype as (ICBN Art. 0.7; Greater et al., 2000) an
“interpretative type ( as | the lectotype . . . associated
with a validly published name, is demonstrably am¬
biguous.’' We have chosen an early cultivated col¬
lection from Europe, as we have been unable to find
a suitable early sheet from Java where Moellerus
gathered his material for Kaempfer.

The genus Helamcanda was named in 1763 by
Adanson, who did not transfer Ixia chinensis, the
single species that lie cited, to the genus. That ac¬
tion was left to I)e Candolle who provided the com¬
bination in 1805. That same year John Ker Gawler
assigned /. chinensis to his new genus Pardanthus,
evidently unaware that this name was a later syn¬
onym of Helamcanda. Gemmingia, a genus listed
in indices of plant names as another synonym of
Helamcanda. is as far as we can determine invalid,
lacking a description. The name was used by Fa-
bricius (1763), who attributed the* name to Heister,
but we have not yet been able to find mention of
the genus in Heisler's publications. Because it is
evidently invalid, we have not listed Gemmingia in
the synonymy above. Two species were listed by
Fabricius as referable to Gemmingia, both listed as
polynomials in the genus Ixia. Currently these are
Iris domestica and Aristea africana (E.) Hoffman-
segg. the basionym of which is Ixia africana (Lin¬
naeus. 1753). Kuntze (1891) provided the combi¬
nation G. chinensis, which is illegitmate because
Gemmingia is invalid. 01 the works containing il¬
lustrations of the plant cited in the protologue of
Ixia chinensis we choose the illustration in
Rheede’s Hortns Malabaricns as the lectotype, as
best representing the1 species. I’he specimen ol the
species in the Linnaean Herbarium is not available
as a lectotype because it was added to the collec¬
tion after Linneaus’s death.

Indexes of plant names list Iris tripedalis Fischer
ex Roemer & Schultes as a synonym of Helamcan¬
da chinensis, but it is an invalid authorless name,
mentioned in discussion only, in the Mantissa to
volume 1 of Roemer & Sehultes’s Systema vegeta-

bilium under the account of /. dichotoma (Roemer
& Schultes, 1822: 306). The unlisted name Par-
danthus tricolor Arruda ex Almeida (1873: 273)
was based on material grown in Brazil but no type
is known, making its identity uncertain, although
this plant is very likely /. domestica. We are in¬
debted to Joseph Kirkbride for drawing our atten¬
tion to this name. We assume that Helamcanda fla-
hellata, described by C. H. Grey in 1934 for
yellow-flowered plants believed to have come from
Japan, is a color variant ol Iris domestica, but we
have been unable to locate type material and do
not include this name in synonymy. The yellow-
flowered variety H. chinensis var. taiwanensis was
included in H. chinensis by Zhao et al. (2000), a
treatment we endorse. Although we have not seen
the type, C.-l. Peng (pers. comm.) considers it a
trivial variant of /. domestica (H. chinensis) with
slightly smaller (lowers than variety chinensis,
which is also native in Taiwan. At least one other
heterotypic synonym is known for the species. He¬
lamcanda pampaninii (LeveilU*. 1910; McKean,
1986), described by Hector Leveille. is based on
(apparently) wild-collected plants from China, also
with predominantly yellow flowers.

All other names in Helamcanda and Pardanthus
(Moeneh, 1794) found in standard indexes of plant
names are combinations in those genera for South
African species now included in Sparaxis Ker Gaw¬
ler or Tritonia Ker Gawler. Pardanthus dichotomus
(Pallas) Ledebour is /. dichotoma, while P. nepa-
lensis Sweet, a name without description, may be
B. chinensis.

Iris domestica is believed to be native to eastern
China, Japan, Korea, Taiwan, and the Ussuri region
of Russia (Mathew, 1981). but the plant has been
in cultivation for so long a time, persists where
planted, and spreads so readily from gardens, that
its original distribution remains somewhat uncer¬
tain. It is treated in the Flora of China as native
there, and is listeil as also occurring in Japan. Ko¬
rea, Myanmar, Vietnam. India, the Philippines, and
Russia, but the authors did not differentiate its na¬
tive from introduced localities (Zhao et al., 2000)
and noted that the plant is “usually cultivated.”
Indeed, the name Helamcanda is perhaps a corrup¬
tion of a southwest Indian vernacular name (Nic-
olson et al.. 1988: 294).

The distinctive features of Iris domestica are the
subequal, spreading tepals. weakly differentiated
into limb and claw (unlike other Iris species), and
the bizarre lepal coloration, a light pink to orange
base with speckles of orange to scarlet pigmenta¬
tion (Fig. 1). Even more singular are the free tepals,
while the style base is not embedded in hypanthi-

Volume 15, Number 1
2005

Goldblatt & Mabberley
Belamcanda and Iris

131

Halin' I. Iris domestica, flowering l>ranch, stem base with leaves, ripe capsule, and detail of the style and style
branches, each with abaxial stigma lobes and short, suberect vestigial paired crests; style branch details much enlarged.
Seale bar I cm. Drawn by John Manning from plants cultivated at the Missouri Botanical Garden.

132

Novon

urn tissue and the style is divided into narrow, al¬
most filiform branches, the latter seemingly quite
different from the broad, tangentially flattened, usu¬
ally petal-like sty It* branches of Iris that terminate
in paired petal-like crests. Examination (with a
I OX hand lens), however, shows that the style
branches are Iris- like in miniature (Fig. I). The
stigma is a small abaxial lobe below the apex of
each style branch, while two small flaps of sterile
tissue form crest-like appendages at the adaxial
apices of lilt; style branch. This structure seems
best interpreted as homologous with the more prom¬
inent style branches of other Iris species. The ap¬
parent reduction of the Iris- like style branches in
/. domestica is paralleled in several species of the
related African genus Moraea, notably species in
section Homeria (Goldblatt, 1986, 1998). In Mo-
raea the reduction of the style branches is associ¬
ated with a shift in pollination system (Goldblatt &
Bernhardt, 1999). A shift in pollination system
therefore seems likely in /. domestica.

Iris domestica also differs from other Iris species
in its globose, smooth, shiny back seeds (fig. I),
evidently an apomorphie character state. The seeds
are quite different from those of other Iris species,
including /. dicholoma, and leave us marveling at
their unusual structure, which we suggest is an ad¬
aptation to avian dispersal, for the seeds collec¬
tively remain attached to the axile placentas after
the capsule walls have dried and curved outward,
the infrueteseenee thereby resembling a blackberry.
This is reflected in one of its common names,
blackberry lily, a name used in North America
where it is widely naturalized (Goldblatt, 2002).

Acknowledgments. We thank Mary Sliffler. li¬
brarian at the Missouri Botanical Garden, for pro¬
viding copies of literature needed for this study,
David Boufford, Brian Mathew, Ching-1 Peng, and
Peter II. Raven for helpful comments on the biology
and geography of Helamcanda, Joseph Kirkbride
for pointing out Almeida Pinto’s publication, Char¬
lie Jarvis for help with Linnaean typification, Stans
Kofman for information on early Javanese collec¬
tions, and Roy Gereau for nomenclatural advice
and the examination of Keister's publications.

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Fabricius, P. C. 1763. Knumeratio Methodica Plantarum Hor-
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Goldblatt, P. 1986. Convergent evolution of tin* *7 tumeric"
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- . 1990. Phylogeny and classification of Iridaeeae.

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- . 1998. Reduction of Barnardiella, Galaxia, Gyn-

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- . 2002. Iridaeeae: Helamcanda. Pp. 395— 396 in

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- & P. Bernhardt. 1999. Pollination of Moraea spe¬
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- , V. Savolainen, (). Porteous, I. Sostarie, M. Powell,

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Greuter, W., J. McNeill, F. R. Barrie, II. M. Burdet, V.
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A New Anthurium sect. Pachyneurium (Araceae) from Minas

Gerais State, Brazil

Eduardo G. Gonsalves

Curso de Ciencias Biologicas, Universidade Catolica de Brasilia, Predio Caspar Bertoni, sala
M-206. OS 7, Lote 1, EPTC, CEP 71030-170, Taguatinga, I)F, Brazil. eduardog@ucl).l)r

Abstract. A new species, Anthuriurn leonii E. G.
Gonsalves from Minas Gerais Slate in southeastern
Brazil, is described, illustrated, and compared with
A. solitarium Schott, the most similar species. An¬
thurium leonii belongs to section Pachyneurium
Schott series Pachyneurium (Schott) Croat and
seems to occur only above I 100 m elevation in tin'
Caparad National Park. This new species is so far
known from the type locality and surrounding areas.

Bksi mo. L ina nova espeeie d e Anthurium {A. leo¬
nii E. G. Gongalves), proveniente do estado de Mi¬
nas Gerais, sudeste do Brasil, e descrita, ilustrada
e comparada com A. solitarium Schott, a espeeie
mais similar. Anthurium leonii pertence a segao Pa¬
chyneurium Schott, serie Pachyneurium (Schott)
Croat e parece apenas ocorrer cm altitudes acima
de 1 100 m. no Parque Nacional do Caparad. Ate
onde se eonhece, esta espeeie e apenas conhecida
para a loealidade tfpica e areas vizinhas.

Key words: Anthurium , Araceae, Brazil, section
Pachyneurium.

The genus Anthurium Schott, comprising approx¬
imately 1000 species (Croat, 1999), is exclusively
Neotropical and is the largest genus in the family
Araceae. Section Pachyneurium is the only section
completely revised in recent times (Croat, 1991),
with at least I 15 species, including A. xanthophyl-
loides G. M. Bai toso, added after the publication of
the revision (Gongalves Salviani, 2001). The di¬
agnostic feature for this section is the presence of
involute vernation, which is supervolute in the oth¬
er sections (Croat, 1991). The species of section
Pachyneurium are usually large herbs with short
internodes.

During mv first visit to the herbarium Guido
Pabst (GJfP) in October 2000, I made note of a
specimen of Anthurium from Caparad National
Park, originally identified as A. solitarium Schott.
Its observed morphology consistently varied from
the typical .4. solitarium that usually occurs in the
eastern Brazilian states of Minas Gerais, Espfrito
Santo, and Rio de Janeiro. Eater, 1 observed living

plants of this species in the field and have con¬
cluded it is a new species, here described. Descrip¬
tive terminology follows Croat and Bunting (1979),
and the term metaphyll for the second cataphyll
follows Grayum (1986).

Anthurium leonii E. G. Gongalves, sp. nov.
TYPE: Brazil. | Minas Gerais:] Alto Caparad,
Parque Nacional do Caparad. 1300 m. 29 Oct.
1996, L S. Leoni 3500 (holotype, GEJP; iso¬
type, UB). Figure I.

Ad sectionem Pachyneurium seriem Pachyneurium per-
linens. Planta epilithica; internodia brevia, 4—7 cm diam.;
prophyllum trigonum, 2-carinatum; metaphyllum laneeo-
latum non carinatum; petiolus 6-13 cm longus, 0.7-1. 5
mm diam., U-formatus, adaxiale sulcatus. marginilms ro-
tundatis; lamina coriacea, obovata, 46—58 cm longa, 21.5-
32 cm lata, nervis primariis lateralibus 10-15 utroque,
arcuatis; pedunculus 30-46 cm longus, 0.5— 0.7 cm diam.;
spatha lanceolata vel ovato-lanceolata, 11-17 cm longa,

1 .8-2.5 cm lata, marginibus ad basem obtusis sed abrupte
acute decurrentibus; spadix castaneus, 9.5—25 cm longus,
inferne 7—1 l mm diam., sursum attenuatus.

Epilithic; stem with internodes short. 4—7 cm
diam.; roots numerous, dense, green, smooth; cat-
aphylls dimorphic, prophyll elongate-triangular. 6—
9 cm long, clearly 2-keeled, metaphyll lanceolate,
12—16 cm long, non-keeled, acute at apex, both
persisting as long brown fibers. Leaves erect to
spre ading; petiole 6—13 cm long, 0.7—1 .5 cm diam.,
U-shaped, narrowly sulcate with rounded margins
adaxially, rounded abaxially, the surface dark
green, pale-speckled; geniculum slightly thicker
than the petiole, pale green, 0.5— 1.5 cm long;
sheath 5.2—9 cm long; blade coriaceous, 46—58 cm
long. 21.5—32 cm wide, obovate, rounded at apex
with a small mucron up to 2 mm long at apex,
obtuse to rounded at base, broadest at tin* middle
or slightly above, the margins very weakly undu¬
late; upper surface matte, medium green, lower sur¬
face matte, slightly paler; both surfaces drying
matte, yellowish green to yellow-brown; midrib
slightly prominent above, obtusely raised below;
primary lateral veins 10 to 15 per side, departing
midrib at 40°— 50° angle, arcuate, slightly raised in

Novon 15: 133-135. 2005.

2cm

134

Novon

Figure I. Anthurium leonii E. G. Gonsalves. —A. Inflorescence. — B. Habit. — C. h lower, longitudinal section. I).
Flower, view from above. — E. Tepal, side view. — F. Pistil, side view. — G. Stamen. — H. Petiole, cross section. — I.
Midrib, cross section. — J. Prophyll. — K. Metaphyll. Drawn from the paralype, Convolves et al. 725 , by the author.

Volume 15, Number 1
2005

Gongalves

Anthurium leonii from Brazil

135

grooves on upper surface, raised below; interpri¬
mary veins poorly developed or absent; tertiary
veins obscure to slightly sunken above in living
material, prominulous in dried leaves, obscure to
slightly raised below in living material, prominu¬
lous in dried leaves; reticulate veins prominulous
when dried; collective vein appearing near the apex
or absent. Inflorescences usually pendent, usually
shorter than the leaves; peduncle 30—46 cm long,
0.5— 0.7 cm diam., 2.3 to 3.6 times longer than the
petiole; spathe spreading, coriaceous, lanceolate to
lanceolate-ovate, rarely lanceolate-elliptic, purplish
tinged with maroon, 11-17 cm long, 1.8— 2.5 cm
wide, broadest slightly above the base, sometimes
almost at middle, inserted at 45°— 50° angle on the
peduncle, acuminate at apex, obtuse at base,
spathe margins almost meeting obtusely then short¬
ly and acutely decurrent to the peduncle; spadix
brown, stipitate by ca. 4 mm, tapered, slightly
curved, 9.5—25 cm long, 7-1 I mm diam. at base,
3-4 mm diam. at apex, broadest near base; (lowers
rhomboidal, 14-2 mm long, 0.8-2 mm wide; 9 to
13 flowers visible in principal spiral, 7 to 10 flow¬
ers visible in alternate spiral; tepals matte; lateral
tepals 0.6—1 mm wide, the inner margins convex,
the outer margins 2-sided; pistils not emergent;
stigma ellipsoid; stamens obscuring the pistil; fila¬
ments not exserted; anthers 0.4— 0.5 X 0.3— 0.8 mm;
thecae elliptic to ovoid, 0.4— 0.5 X 0.2— 0.5 mm.
Infructescenee unknown.

Anthurium leonii is very similar to A. solitarium,
but differs in having a proportionally broader blade
with the length X width ratio about 1.8-2. 3 as com¬
pared to 2.8— 4.8 in A. solitarium. Anthurium leonii
usually has a higher number of primary lateral
nerves (10 to 16, rather than usually 8 to 9(to 10)
in A. solitarium ), and these nerves are completely
arcuate, whereas they are usually found "running
straight from the midrib to the margin, and then
steeply and arching rising along the margin” in A.
solitarium (Croat, 1991: 729). Another important
difference is that the spathe margins meet obtusely
then are shortly and acutely decurrent onto the pe¬
duncle in A. leonii , whereas the spathe margins are
uniformly and acutely decurrent in A. solitarium.
Anthurium solitarium is known to occur in eastern
Minas Gerais State and has been collected in the

vicinity of the Capa rad National Park (e.g., Leoni
2266 and 2260), usually at elevations up to 600 m.
Because of the proportionally broad leaf blade and
long pendent inflorescences, no other species in
section Paehyneurium may be confused with A. leo¬
nii.

Anthurium leonii belongs to section Paehyneu-
rium series Paehyneurium because of the involute
vernation, as defined by Croat (1991).

Anthurium leonii is epilithic in gneissie outcrops,
usually growing together with Philodendron edmun-
doi G. M. Barroso. The plants usually grow in direct
sunlight, but some individuals are also found in
half-shade. Despite the fact that it is locally abun¬
dant. A. leonii is known only from the type locality.
It is possible that it also occurs in neighboring ar¬
eas. above 1000 m.

The epithet was chosen in honor of Lucio S. Leo¬
ni, curator of the herbarium GFJP, one of the most
important plant collectors of the Caparao National
Park and surrounding areas.

Paratype. BRAZIL. Minas Gerais: Alto Caparao,
Barque Nacional do Caparao, Vale Verde, 1100 m s. m.,

I Feb. 2001. E. 6. Gonsalves, E. R. Salviani, L. S. Leoni
& M. Peixoto 725 (GFJP, MO, UB).

Acknowledgments. I thank Lucio Leoni for
showing me living plants of Anthurium leonii, as
well as for the herbarium facilities at GFJP. Field
trips were sponsored by the Institute Plantarum de
Fstudos da Flora Lida. I am also grateful to L. Sal¬
viani for field assistance and Thomas B. Croat (MO)
for valuable comments on the manuscript. The au¬
thor was funded by a grant from FAPESP (99/
02921-7).

Literature Cited

Croat, T. B. 1991. A revision of Anthurium section Pa-
chyneurium (Araceae). Ann. Missouri Bot. Card. 78:
539-855.

- . 1999. The Araceae of Peru — Distribution, spe¬
cies diversity and centers of endemism. Arnaldoa 6(2):
45-80.

- & 6. S. Bunting. 1979. Standardization of An¬
thurium descriptions. Aroideana 2(1): 15—25.

Gonyalves, E. G. & E. R. Salviani. 2001. Anthurium xan-
thophylloides G. M. Barroso (Araceae) re-found in Es-
pfrito Santo state, eastern Brazil. Aroideana 24: 13—17.
Gravum, M. If. 1986. Revision of Philodendron subgenus
Pteromischum (Araceae) for Pacific and Caribbean
Tropical America. Syst. Bot. Monogr. 47: 1-233.

Cuphea nivea (Lythraceae), a New Species from Bolivia

Shirley A. Graham

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

sliirley.graham@mobot.org

Abstract. Cuphea nivea from northwestern Bo¬
livia is described and illustrated. Seed and pollen
characters relate it most closely to C. ciliata, a spe¬
cies occurring from southern Venezuela to southern
I ’em. Ilie small xylopodium and heath-like leaves
of the species suggest it is adapted to grassland
habitats that experience seasonal fires.

llESUMEN. bn el presente trabajo se describe e
i lustra Cuphea nivea de la region noroccidental de
Bolivia. Los caraeteres de las semillas y de los gra-
nos de polen la relacionan muy estrechamente con
C. ciliata, una especie que se distribuye desde el
sur de Venezuela hasta el sur de Peru. El xilopodio
pequeno y las hojas ericoides de la especie sugi-
eren su adaptacion a habitats dominados por pastes
(pie sufren impactos de quemas estacionales.

Key words: Bolivia, Cuphea, grasslands, Lyth¬
raceae.

Hecent collecting efforts in support ol the Cat¬
alogue of the Vascular Plants of Bolivia (Jprgensen
et al., in progress) concentrated on a previously un¬
dercollected area northeast of Apolo, in La Paz
province. A species new to Cuphea 1’. Browne, tin1
largest genus of Lythraceae with over 260 species
(Graham et ah, 1993), is described from those col¬
lections.

Gupliea nivea S. A. Graham, sp. nov. TYPE. Bo¬
livia. La Paz: Franz Tamayo, Senda Apolo-San
Jose de Uchupiamonas, a media hora antes de
llegar al Naranjal, 1798 m, 1 Oct. 2002, 7.
Miranda, C. Maldonado, F. Canqui, R. Alvarez
& J. Tito 141 A (holotype, LPB; isotype, MO).
Figure 1.

Herbae perennes 15—25 cm, rhizomate crasso ad 1.5
cm diam.; eaules 1 -aliquot, pro parte erecti. ascendentes,
strigosi et glanduloso-hispiduli. Folia ovata vel anguste
ovata, 3-6 X 1 .5-3 mm, eoriacea, suberecta, uninervia,
basi rotundata ad subcontata. Racemi simplices, foliati;
pedicelli 1.5-2. 5 mm. Flores 6-meri, 4—5 mm longi, sine
calcari, glanduloso-hispiduli; petala 6, 1.7— 2.3 mm longa,
nivea, vinoso-nervata; stamina II, profunde inserta, tub-
um longe non aequantia; ovula 4; nectarium basi ovarii
0.5 X 0.3 mm, erectum, earinatum. Semina orbicularia,
1.7 X 1.6 mm, exalata, margine obtusa.

Diminutive perennial herbs 15—25 cm; stems 1
to few, unbranched or sparsely, irregularly
branched, from a woody xylopodium ca. 1.5 cm
diam.; stems wine-red, densely strigose and glan¬
dular-hispid, the white, sharp-pointed trichomes
oppressed and antrorsely oriented, the glandular
trichomes erect, 0.5 mm long; internodes slightly
shorter to slightly longer than the subtending
leaves. Leaves opposite, sessile, erect, loosely to
tightly oppressed to the stem, ovate to narrowly
ovate, 3—6 X 1.5—3 mm, coriaceous, appearing uni-
nerved, only the midvein visible, base rounded to
subeordate, apex acute, margin slightly thickened
or inrolled, upper surface glabrous, lower surface
minutely white-strigose; leaves scarcely reduced in
size toward the stem apex. Inflorescences simple
leafy racemes; flowers solitary, alternate, interpetio-
lar; pedicels 1.5— 2.5 mm; bracteoles 0.5 mm, ob¬
long, green. Floral tubes 6-merous, 4—5 X 1 mm
including a rounded base, the base extended 0—0.2
mm beyond the attachment of the pedicel, the neck
of the floral tube not contracted, the mouth blunt;
outer surface wine-red dorsally, green venlrally,
white-strigose and glandular-hispid, the glandular
trichomes costal and intercostal; calyx lobes equal,
0.5 X 0.7 mm, triangular; appendages of the epi-
calyx represented by wine-red thickenings at the
sinuses of the calyx lobes, short-setose, t he margin
not free; inner surface neither bialate nor vesicu¬
late. densely white villous above the stamens, gla¬
brous below; petals 6, subequal, 1.7— 2.3 X 0.8—1
mm, narrowly oblong-elliptic, white with wine-red
midvein, the 2 dorsal petals subtended by a bright
yellow nectar guide; stamens 11, the 2 dorsalmost
shortest and inserted below the other 9, the 9 un-
iseriate, inserted deeply in the floral tube, alter¬
nately unequal, the anthers not reaching the sinus¬
es of the tube; filaments glabrous to sparsely white
villous; anthers light yellow; pollen triangular in
outline, tricolporate, non-syncolpate, pores non¬
protruding; exine psilate at 40X; 15 pun diam.;
style included. 2.5 mm, glabrous; stigma puncti-
form; placental base in the ovary fleshy, semi-cir¬
cular with smooth margin, 2 mm wide at capsule
dehiscence; ovules 4; nectariferous disc 0.5 X 0.3

Novon 15: 136-138. 2005.

5 mm

Volume 15, Number 1
2005

Graham

Cuphea nivea from Bolivia

137

1mm

Figure 1. Cuphea nivea S. A. Graham. — A. Habit. — IF Leaf, abaxial surface. — C. Seed. — I). Flower, lateral view.
— E. Flower opened dorsally, exposing llie nectariferous, unilateral disc at tbe base of the ovary and the included
stamens and stigma. Four petals removed for clarity. Drawn from the isotype, Miranda el al. 141 (MO).

138

No von

mm, erect at the base of the ovary, keeled. Seeds
suborbicular to orbicular, 1.7 X 1.6 mm, light
brown, the margin rounded, paler than the rest of
the seed.

Distribution and phenology. Known only from
the type area; in grassy pasturelands; 1785—1798
m. Collected in flower and fruit October, December,
and April.

Cuphea nivea (sect. Hrachyandra ) is recognized
by the bright white petals with wine-red midvein
and the stems bearing thick, appressed, narrowly
ovate leaves. Pure white petals are uncommon in
the genus, although many species bear pink or pale
purple petals or have four white ventral petals in
combination with two purple dorsal ones. The spe¬
cific epithet refers to the white petal color. The sec¬
tional assignment of the species follows the current
classification of the genus (Koehne, 1903) in which
section Hrachyandra is defined by stamens so
deeply inserted in the floral tube that the anthers
do not reach the sinuses of the calyx lobes. Cla-
distic analyses based on morphology (Graham,
1998) and molecular data (Graham et al., in prep.)
indicate that section Hrachyandra is polyphyletic
and the stamina! character is part of a floral syn¬
drome tied to an autogamous breeding system. The
species that constitute the section represent at least
four independent lineages, and the floral morphol¬
ogy of the primarily self-fertilizing species is con¬
vergent.

In section Hrachyandra, Cuphea nivea shares
seed and pollen features with species of the lineage
that includes: Cuphea calophylla Chamisso &
Schleehtendal, C. melanium (I,.) R. Brown ex Steu-
del, and C. ciliata Ruiz & Pavon. Seeds of these
species are non-winged with round, thickened mar¬
gins, and the pollen is small lor the genus (ea. 15
/rm), nearly psilate, non-syncolpate, and without
protruding pores (Graham, 1998). The habit of C.
nivea approaches that of C. ciliata, an Andean spe¬
cies occurring from southern Venezuela to southern

Peru. Cuphea ciliata differs in having more lax.
much-branched stems, linear leaves that are not so
consistently appressed to the stem, pedicels 7—14
mm, and 6 to 10 ovules, not 4 as in C. nivea. Stems
of C. nivea are less woody and more erect than in
C. ciliata , the leaves are more appressed, and the
xylopodium is better developed.

The thick, heath-like leaves and woody xylopo-
dia of Cuphea nivea are also characteristic of sev¬
eral, mostly Brazilian, species of section Euandra
subsect. Oidemation, and C. nivea could, at first
glance, be considered a member of that subsection.
However, numerous differences in floral and pollen
morphology indicate that the similarity is due to
convergence. Members of both sections occur in
open savannas where coriaceous, closely appressed
leaves and xylopodia are considered advantageous
to survival during seasonal fires. The presence of
C. nivea suggests that the collection area, now pas-
tureland given to cattle-grazing, was once a natural
savanna subject to burning.

Paratypes. BOLIVIA. La Pa/,: Franz Tamayo, Senda
Apolo-San Jos6 de Uehupiamonas, a 15 minutos del ar-
royo Turnia hacia el K de la senda, 2 Dec. 2002, T. Mi¬
randa, C. Maldonado & V. Canqui 388 (LPB not seen,
MO); a media hora antes de llegar al Naranjal, 17 Apr.
2003, T. Miranda. L. Cayola, F. Canqui, K. Alvarez & ./.
Tito 718 (LPB not seen, MO).

Acknowledgments. I thank Yevonn Wilson-
Ramsey for preparing the illustration, and Peter
Jprgensen for information about the Bolivian col¬
lection site.

Literature Cited

Graham, S. 1998. Relacionamentos entre as espO-ies au-
togamas de Cuphea P. Browne segao Hrachyandra Ko¬
ehne (Lythraeeae). Aeta Bot. Brasil. 12: 203—214.

- , J. Crisei & P. C. Hoch. 1993. Cladistie analysis

of the Lythraeeae s.l. based on morphological charac¬
ters. Bot. J. Linn. Soe. 1 13: 1—33.

Koehne, K. 1903. Lythraeeae. In: A. Engler (editor). Das
Pflanzenreich. IV. 216. Heft 17. Wilhelm Englemann,
Leipzig.

Two New Species of Hortia (Rutaceae) from Amazonia

Milton Groppo and Jose R. Pi rani

Instituto tie Biocieneias, Universidade de Sao Paulo, Caixa Postal 1 1461-05422-970,
Sao Paulo. Brazil, groppojr@yahoo.coin.br

Abstract. Hortia nudipetala Groppo and //. van-
delliana Groppo (Rutaceae, Toddalioideae, Todda-
liinae) from Amazonia are described and illustrat¬
ed. The first species is known from the sandy,
periodically inundated margins of tributaries of the
upper Rio Negro, and the second from Brazilian
and Venezuelan Amazonia. Comments on affinities
of the two species, geographic distribution, and
phenology are provided.

Rksumo. Hortia nudipetala Groppo e H. vandel-
liana Groppo (Rutaceae, Toddalioideae, Toddali-
inae) da Amazonia sao deseritas e ilustradas. A pri-
meira especie e conhecida das margens arenosas e
periodicamente inundadas dos tributaries do alto
Rio Negro e a segunda da Amazonia Brasileira e
Venezuelana. Sao apresentados comentarios sobre
as afinidades, distribuiyao geografica e fenologia
das duas especies.

Key words: Amazonia, Brazil, Hortia, Rutaceae.

Hortia Vandelli is a Neotropical genus of Ruta¬
ceae traditionally included iu subtribe Toddaliinae,
subfamily Toddalioideae (see Engler, 1931). The 10
species are distributed from Panama to the state of
Sao Paulo in Brazil, but most occur across Ama¬
zonia. These taxa are trees or shrubs with simple
leaves crowded near the apex of the branches and
have showy, wide corymbose terminal inflorescenc¬
es, red to pink flowers, and a baccate fruit with
abundant oil glands. During a study aimed at a
revision ol the genus, two new species were rec¬
ognized.

The descriptions are based only on collections
from mature individuals. Flowers and fruits of her¬
barium specimens were rehydrated before being
measured and drawn. The terminology for shapes
ol leaves and other organs follows Hickey (1979)
and Radford et al. (1974); lor inflorescences, We-
berling (1989); and for fruit, Spujt (1994).

Ilortia nudipetala Groppo, sp. nov. TYPE: Brazil.
Amazonas: Sao Gabriel da Cachoeira, alto Rio
Negro, Rio LJaupes, Igarape Tubuari, acima do
Seringal Pago, 22 Nov. 1987 (fl), //. C. de
Lima , M. L. Kawasaki, ./. Ramos & R. P. de
Lima 3278 (holotype, SPF; isotypes, IN PA,
NY, RB). Figure T

Ah omnibus speciebus generis petalis glabris distinota.
Species H. brasilianae similis, sed foliis angustioribus dis-
tincte petiolatis atque nervo inframarginali manifesto dif-
ferl.

Shrub 2—3 ni tall; diameter of trunk unknown;
branches erect, glabrous; outer bark scarcely fis¬
sured, the inner bark pale yellow. Leaves eharta-
ceous, patent to ascending, glabrous, petiolate;
blade 15.8—24.1 X 2.9— 3.6 cm, with abundant pel¬
lucid dots, apparently lustrous on adaxial surface,
dull on abaxial surface, narrowly oblong to lance¬
olate, apex rounded, acute or retuse, apiculate or
not, margin entire, revolute, thickened, base atten¬
uate; midrib thick, flat on the adaxial surface,
prominent below, the secondary veins in 18 to 22
pairs diverging from midrib at 55°— 64°. prominent
on both faces, united in an evident inframarginal
vein, the tertiary veins prominent on both faces;
petiole 1.4—3 cm long, clearly distinct from blade,
winged, thickened at base. Inflorescence a pleio-
thyrsoid, 5—7 X 7—8 cm, the branches slightly com-
planate, minutely fissured transversely; bracts and
bracteoles triangular, the proximal ones 1 X 3 mm,
not crowded at base of inflorescence, the distal ones
ca. 1 X 1 mm; thickness of the axis 0.5 cm at base,
0.2 cm after the 4th order of ramification; pedicels
ca. 2 mm long. Flowers 5-merous, ca. 7— 8 mm
diam.; buds ovoid; calyx eotyliform, 5-lobed, the
lobes triangular, ca. 1 .4 mm long, obtuse or acu¬
minate, ciliate; petals 5, pink, oblong to triangular,
ca. 6 X 2.5 mm, apiculate, glabrous; filaments ca.
3—5 X 1 mm, inserted on the apex or side of the
disc; anthers ca. 1.5 mm long, oblong; disc eotyli¬
form, 5-lobed, fleshy, adnate to a very short gyno-
phore; gynoecium ovoid, ca. 5X2 mm. the base
immersed in the disc; ovary 5-lobed, slightly raised
upon the gynophore; style conical, ca. I mm long.
Fruit (immature) a globose berry, ca. 6 X 6 mm,
green; seeds (mature) not seen.

Distribution and habitat. Hortia nudipetala is
known so far by only a few collections made in the
region of the upper Rio Negro, in the vicinity of
Sao Gabriel da Cachoeira, state of Amazonas. Bra¬
zil (Fig. 3). It was found on the periodically inun¬
dated margins of some tributaries of the Rio Negro

Novon 15: 139-143. 2005.

140

Novon

Figure I. Hortia nudipetala Groppo. — A. Habit with detail of the venation and inframarginal vein on the abaxial
surface of the leaf. — B. Flower. — C. Hud. — I). Gynoeeium. — E. Gynoecium in longitudinal section. — F. Petal. —
G. Stamen. — II. Stamen with open thecae. — 1. Same in abaxial view. — .1. Immature fruit; note the remnant of the
style. — K. Immature fruit in longitudinal section. — L. Two ovules, the lower one aborted. A— I and F from Lima el al.
3278; J, k from Schultes A' lApez 96 87.

Volume 15, Number 1
2005

Groppo & Pirani
Hortia from Amazonia

141

(such as llic Uaupes River) and along some igara-
pes (small, black-water rivers) on sandy soils, in
campina (an open vegetation of low trees and
shrubs). Specimens were collected with flowers in
November and January to March, and with imma¬
ture fruits in March.

Hortia nudipetala is the only species in the ge¬
nus where the petals are totally devoid of the char¬
acteristic; trichomes that otherwise occur on the ad-
axial surface of the petals. It is similar to //.
brasiliana Vandelli ex DC. (an arborescent species
of the Atlantic coast of Brazil), in particular be¬
cause of the flowers, i.e., with similar shape of the
ovary and stamens; however, the leaves of H. nu¬
dipetala are narrower (to 3.6 cm vs. 3— 9(— 1 I) cm
in //. brasiliana) with an evident inframarginal
vein, and the petiole is more distinct. In addition,
the two species are largely allopal ric.

No economic uses or common names are report¬
ed for this species.

Paralypes. fill AZ lb. Amazonas: Sao Gabriel da
Cachoeira, Rio Negro, Igarape Toury (Tubuari), R. L Froes
27916 (IAN); Igarape Tuari (Tubuari), lado oposto a ilha
de Aparecida, W. A. Rodrigues 10878 (NY); Rio Uaupes,
Tarac-ua, R. E. Schultes & F. Ldpez 9687 (Cl I, MO).

Hortia vandelliana Groppo, sp. nov. TYPE; Bra¬
zil. Amazonas: Sao Paulo de Olivenya, alto Rio
Solimoes, plato ao sul da cidade, estrada para
a localidade de Bom Kim, 25 Nov. 1986
(buds), C. A. Cid et al. 8538 (holotype, SPF;
isotypes, II AM All, IN PA 2 sheets, NY). Fig-
ure 2.

Species foliis II. coccinae comparabilis, sed petalis
triangularibus, alabastris ovatis atque petalis triehomati-
bus majoribus differt. II. brasilianae etiam sirnilis, sed
alabastris fructibusque mirioribus atque foliis nervis non
manifestis differt.

free or shrub (2— )4— 20 m tall, trunk up to 60
cm diani.; branches erect, glabrous; outer bark
scarcely fissured, the inner bark pale yellow.
Leaves chartaceous to strongly coriaceous, patent
to ascending, glabrous, petiolate; blade 10-22.2 X
4.1— 7.2 cm, with abundant pellucid dots, appar¬
ently lustrous on adaxial surface, dull on the ab-
axial surface, obovate, the apex rounded, seldom
retuse, the margin entire, revolute, thickened, the
base attenuate to decurrent; midrib thick, flat on
the adaxial surface, prominent below, the secondary
veins in 10 to 12 pairs diverging from midrib at
35°— 53°(60°), immersed in the mesophyll on both
faces, usually not visible on the abaxial surface, not
united in an inframarginal vein, the tertiary veins
not visible on both surfaces, immersed in the me¬
sophyll; petiole 1.2— 2(— 3) cm long, sometimes not

clearly distinct from blade, winged, thickened at
base. Inflorescence a pleiothyrsoid, 9—13 X 6— I I
cm, the branches terete, minutely fissured trans¬
versely; bracts and braeteoles triangular, the prox¬
imal ones ca. 3 X 5 mm, not crowded at base of
inflorescence, the distal ones ca. 0.5 X I mm, eil-
iate; thickness of the axis 0.6-0. 7 cm at base, 0.2
cm after the 4th order of ramification; pedicels ca.
1.5 mm long. Flowers 5-merous, ca. 8 mm diam.;
buds ovoid; calyx cotyliform, 5-lobed, the lobes tri¬
angular, ca. 1.4 mm long, obtuse or acuminate, not
ciliate; petals 5, pink, triangular, ca. 4 X 2.5 mm,
apieulate, with a tuft of trichomes on the base of
the adaxial face, the trichomes straight, 0.5— 0.9
mm long; filaments ca. 4 X 0.9 mm, inserted on
the apex of the disc; anthers ca. 2.5 mm long, ob¬
long; disc cotyliform, 5-lobed, fleshy, adnate to a
short gynophore; gynoecium ovoid, ca. 3 X 2 mm,
the base immersed in the disc; ovary slightly 5-
lobed, raised upon the gynophore; style conical, ca.

1 mm long. Fruit an ellipsoid berry, ca. 3.5 X 2.5
cm, green or light yellow when mature; seeds con¬
ical, ca. 8X6 mm, the testa blackened, the hilum
basal; cotyledons 5—6 X 4 mm, oval.

Distribution and habitat. Two of the six speci¬
mens of Hortia vandelliana (the type collec tion and
Silva et al. 532 ) have been collected in eastern
Amazonas state, Brazil, where it occurs in the Bio
Solimoes basin, in Amazonian “caatinga” (arbores¬
cent vegetation on sandy soils), and in “terra fume”
forest (non-inundated tall forest); another two col¬
lections ( Ribeiro 1026 and Froes 28029) arc- from
river margins of the Bio Negro basin, on sandy or
clay soils. There is one record from the department
of Amazonas, Venezuela, and another from Cura-ei-
ha range (600 m of altitude), state of Roraima, Bra¬
zil (Fig. 3). The species was collected with flowers
in June and November, and with fruits in February,
March, and July.

The shape, size, and nervation of the leaf blade
of this species are similar to those of Hortia coc-
cinea Spruce ex Engler. Hortia vandelliana differs
from that species, however, by a petiole that is
sometimes indistinct (vs. c dearly distinct) from the
blade, by smaller (8 mm vs. 14 mm diam.) flowers,
and by the triangular (vs. oblong) petals bearing
longer trichomes (0.5— 0.9 vs. 0.2— 0.3 mm) that are
present just as lull on the base of the- (vs. the- entire)
adaxial surface.

The flowers of Hortia vandelliana arc- smaller
than those of H. brasiliana (8 mm vs. 13 mm
diam.), but similar in all other aspects. In //. van¬
delliana the secondary and tertiary veins are im¬
mersed in the leaf blade, while in II. brasiliana

142

Novon

|-J I fcftok *inWfe

h'i I'll rt* 2. Hortia vandelliana Groppo. — A. Habit, with detail of the fissures in the axis of the inflorescence. — B.
leaves. — G. Bud. — I). Bud with two petals and two stamens removed. — E. Petal, adaxial view. — F. Gynoeeium of
two flowers iri longitudinal section. — G. Fruits. — H. Seed. — I. Seed in longitudinal section, showing embryo (part of
endosperm on the right). A, C-F from Cid et al. 8538; B, left leaf from Silva el al. 532, right leaf from Mur^a-Pires et
al. 16945 ; G— I from Mur^a-Pires at al. 16945.

Volume 15, Number 1
2005

Groppo & Pirani
Hortia from Amazonia

143

they are evident and prominent. In H. vandelliana
(vs. II. brasiliana) t he secondary veins diverge from
the midrib at less obtuse angles [35°— 53°(60°) vs.
60°— 69° | and the fruits are smaller (ca. 3.5 X 2.5
cm vs. 4.5— 5.5 X 4.5 cm in II. brasiliana) and
globose to obovoid (vs. ellipsoid).

The specific epithet honors Domenico (Domingo
or Domingos) Vandelli (1730—1816), Italian physi¬
cian and botanist who described the genus Hortia.

No economic uses or common names are report¬
ed for this species.

Paratypes. BRAZIL Amazonas: Carauari, Riojurua,
Poyo Jurua I. A. S. L. da Silva, A'. A. Rosa &: R. P. Pallia
5.32 (INPA); Sao Gabriel da Cachoeira, rio lyana, perto da
cachoeira Malaeaeheta, R. I- Frdes 2H0.39 (IAN); luarete,
margern do rio Uaupes, P. G. S. Ribeiro 1026 (MG). Ko-
raima: Rio Uraricoera, Serra Cura-ci-lia, J. Mur/pi - Pires,
W. A. Rodrigues, A. A. Rosa & G. Ranzani 16045 (INPA,
MBM, MG). VENEZUKLA. Amazonas: Guainfa, along
road from Maroa to Yabita, ca. 700 m from Yabita, P.
Acevedo- Rodriguez, G. Ayrnard, P. E. Perry & R. Schargel
10285 (NY).

Acknowledgments. The first author thanks the
curators of GH, HAMAB, IAN. INPA, MRM, MG,

MO, NY, and 1111 for the loan of specimens, and
especially the curator of NY. for the gift of impor¬
tant collections of Hortia to SPK; Eduardo Kiek-
hofel for the illustration; and Jacquelyn Kallunki
for both her helpful suggestions and improvements
to the English text and her constant anil enthusi¬
astic help. Financial support from FAPESP (Fun-
da^ao de Amparo a Pesquisa do Estado de Sao Pau¬
lo. grant no. 00/07401-0) is duly acknowledged.

Literature Cited

Engler, A. 1931. Rutaceae. Pp. 187—359 in A. Engler &
K. Prantl (editors), Nat. PHanzenfam. 2nd ed., \ol. 19a.
Wilhelm Engelmann, Leipzig.

Hickey, L. J. 1979. A revised classification of the archi¬
tecture of dicotyledonous leaves. Pp. 25—39 in C. R.
Metcalfe & L. Chalk (editors). Anatomy ol the Dicoty¬
ledons, 2nd ed.. Vol. 2. Clarendon Press, Oxford.
Radford, A. E., W. C. Dickinson, J. R. Massey & C. R.
Bell. 1974. Vascular Plant Systematics. Harper & Row,
New York.

Spujt, R. W. 1994. A systematic treatment of fruit types.

Mem. New York Rot. (lard. 70: 1-181.

Weberling, E 1989. Morphology of Flowers and Inflores¬
cences. Cambridge Univ. Press, Cambridge.

A (New Name in Korean Silene (Caryophy I laeeae)

Suk-Pyo Hong * and Hye-Kyoung Moon 11

Laboratory ol Plant Systematics, Department of Biology & The Basic Sciences Institute,
Kyung I lee University, Seoul 130-701, Korea. usphong@khu.ac.kr; l,lycopus@korea.eom

Abstract. While pre[>aring a treatment of the ge¬
nus Silene L. lor the Generic Flora of Korea, the
following novelty was found and is published in this
contribution. Accepting the current generic concept
of Silene, we noticed that one Korean endemic spe¬

cies, Melandrium umbellalum Nakai. should be
transferred to Silene. However, transfer of the spe¬
cific epithet to Silene would create a later homonym
of Silene umbellata Gilibert (= Silene armeria L.).
Therefore, we here provide the following new name:

F igure I. The holotype of Silene myongcheonensis Suk-Pyo Hong & Hye-Kyoung Moon in I I.

Novon 15: 1 44—145. 2005.

Volume 15, Number 1
2005

Hong & Moon
Korean Silene

145

Silene myongcheonensis Suk-Pyo Hong & Hyc-
Kyoung Moon.

Key words: Caryophyllaceae, Korea, Silene.

The revision of the genus Silene L. (Caryophyl¬
laceae) lor the Generic Flora of Korea is in progress
(Hong, in prep.). The circumscription of Silene has
long been debated, and there has been a tendency
toward expanding the genus during recent decades
(Greuter, 1995). At the infrageneric level, a lump¬
ing trend has been prevalent (Chowdhuri, 1957;
Sou rkova, 1978) and is also supported by nuclear
gene sequence data (Oxelman & Liden, 1995).
Such generic limits have been adopted for the (Ge¬
neric Flora oj Korea treatment.

The Korean endemic species Melandrium iim-
bellatum Nakai was originally described from Mt.
Chilbo-Shan, Myongcheon-Gun, I lanikyungbuk-
Do. This species is similar to Silene capitata Ko¬
marov by having pinkish petals, but is distinct in
its umbel-like cymes and by having glandular tri¬
e-homes on both calyx and peduncles. The prior de¬
limitation of Melandrium from other relatives, in¬
cluding Silene s. str., was strictly on the basis of
carpel and hence style number; this character is
not invaluable among these taxa. This necessitates
a new combination in the genus Silene for Melan¬
drium umbellatum. Unfortunately, the resulting
name of the new combination would be a later hom¬
onym of S. umbellata Gilibert [= Silene armeria L.

I 753: 420). which is illegitimate under Article 53. 1
of the Saint Louis Code (Greuter et al.. 2000).
Therefore, a new name is needed and is here val¬

idated for its use in the upcoming account of the
genus in the Generic Flora oj Korea.

Silene myongcheonensis Suk-Pyo Hong & llve-
Kyoung Moon, nom. nov. Replaced name: Me¬
landrium umbellatum Nakai, Rot. Mag. (To¬
kyo) 33: 48. 1919, not Silene umbellata
Gilibert. FI. Fit. Inch. 2. 170. 1782. TYPE:
“Corea, in montis Hichihozan | Korea, Ha-
mkyungbuk-Do, Myongcheon-Gun, Ml. Chil-
bo-shan],” 4 Aug. 1918. T. Nakai 6985 (ho-
lotype, TI). Figure 1.

Acknowledgments. We are grateful to H. Ohba
(TI) for allowing us to examine material. We are
also thankful to W.-T. Lee and K.-O. 'too for pro¬
viding useful data. The research was supported by
a grant (no. 052-041-026) from the Core Environ¬
mental Technology Development Project for Next
Generation funded by the Ministry of Environment
o! the Korean Government.

I .iterature Cited

Chowdhuri, I*. K. 1957. Studies in the genus Silene. [Notes
Roy. Hot. Card. Edinburgh 22: 221-273.

Greuter, V. 1995. Silene (Caryophyllaceae) iu Greece: A
subgeneric and sectional classification. Taxon 44: 543—
581.

- . J. McNeill, F. If. Rarrie. II. M. Runlet, V. I)e-

moulin, T. S. Filgueiras, I). H. Nieolson. P. (7 Silva. J.
E. Skog, P. Trehane, N. J. Turland & 1). I.. Hawksworth
(editors). 2000. International Code of Botanical Nomen¬
clature (Saint Louis Code). Kegnum Veg. 138.
Oxelman, R. & M. Lidtm. 1995. Generic boundaries in
the tribe Sileneae (Caryophyllaceae) as inferred from
nuclear rl)NA sequences. Taxon 44: 525—542.
Sourkovd, M. 1978. Caryophyllaceae subfam. Dianthoi-
deae— Regrenzung, Charakteristik und Gliederung. Pres-
lia 50: 139-152.

Studies in the Cleomaceae II: Cleome boliviensis, a (New, Spiny,
Large-Flowered Andean Species

Hugh H. litis

Herbarium, Department of Botany, University of Wisconsin-Madison, 430 Lincoln Drive,
Madison, Wisconsin 53706-1381, U.S.A. tscoclira@facstaff.wisc.edu

Abstract. Cleome boliviensis litis, a large-flow¬
ered new species of the speeiose. mostly spiny sec¬
tion Tarenaya (Rafinesque) litis [validated here], is
described, illustrated, and its relationship to other
species discussed. This xerophytic subshrub, wide¬
spread and common in the Andes of southern Peru
and Bolivia (and becoming weedy in adjacent Am¬
azonian lowlands), differs from the allopatric, more
eastern, lowland, annual C. hassleriana by its more
or less perennial habit, whitish to pinkish white
flowers, finely glandular-pubescent fruits, and
smaller seeds with abundant, minute (use 20X
lens), oil-containing, light -refracting, caruncular
bulliform cells. These densely cover the flat seed
faces and often the mouth of t he seed “cleft," and
appear to be related to dispersal by ants. At the
edge of its range, the new species seems to hybrid¬
ize with other species of section Tarenaya, such as
C. parviflora and long-fruited C. spinosa s.l. re¬
cently introduced from northeastern Brazil. Myr-
mecochory in the Cleomaceae is discussed and ap¬
pears to have arisen independently several times.

RKSUMEN. Se describe e ilustra Cleome boliviensis
litis (Cleomaceae), una nueva especie de flores
grandes, perteneciente a la seccidn Tarenaya (Ra¬
finesque) litis, una seccidn con numerosas especies
en sn mayorfa espinosas; se discute su relacidn con
otras especies alines. Es un subarbusto xerofftico,
ampliamente distribuido en los Andes del sur de
Peru y de Bolivia (llega a ser una maleza adventicia
en areas bajas aledanas de la Amazonia), difiere de
la especie alopatrica C. hassleriana por su habito
mas o menos perenne, flores blanquecinas basta
rosado blanquecinas, por sus frutos finamente pu-
bescente-glandu lares y semillas mas pequenas con
abundantes diminutas (use una lupa de 20 X) ee-
lulas carunculares buliformes, que contienen ac-
eites y reflejan la luz. fistas cubren densamente las
caras aplanadas de las semillas y a menudo la boca
de la “escotadura" tie la semilla y sugieren la dis¬
persion por hormigas. En el h'mite de su distribu-
cidn geografica. esta especie parece formar hfbridos
con otras especies tie la sect. Tarenaya eomo C.
parviflora y C. spinosa s.l., una especie de fruto

largo de reciente introduccidn en el nororiente de
Brasil. Se discute la mirmecoeoria en las Cleoma¬
ceae que parece haber evolucionado varias veces,
independientemente.

Key words: Andean biogeography, ant-dispers¬
al, Bolivia, Cleomaceae, Cleome sect. Tarenaya,
myrmecochory.

Throughout extensive montane and submontane
areas of Bolivia and adjoining southern Peru (Kig.
3) there grows a common large-flowered, very spiny,
shrubby Cleome, one of the more than 40 members
of the speeiose, mostly spiny, almost exclusively
New World Cleome sect. Tarenaya (see below and
Jacobs, 1960; litis, 1952. 1967). Though this strik¬
ingly beautiful species is represented by many her¬
barium collections and has long been recognized as
distinct (litis, 1952, 1967), it lacks a properly pub¬
lished binomial.

Cleome sect. Tarenaya (Rafinesque) litis, comb,
et stat. nov. Basionym: Tarenaya Rafinesque,
Sylva Telluriana 111. 1838. TYPE: Tarenaya
spinosa (Jacquin) Rafinesque = Cleome spi¬
nosa Jacquin.

Mostly annual herbs, usually glandular pubes¬
cent, often with epidermis-derived paired spines or
thorns in the stipular position at the base of the
multifoliolate, palmately compound leaves, with
many-flowered bracteate inflorescences, flowers
with 6 stamens, elongate gynophores and many-
seeded siliques, the seeds with a deep hollow cleft
chamber closed on all sides by distinct cleft mem¬
branes from embryo vertex to seed mouth (litis et
ah, in prep.).

A group of ca. 40 New World species and one
West African (litis, 1967).

The above change in status, made previously in¬
formally by Jacobs ( Flora Malesiana / 6(1): 100,
1960). is herewith validated.

The forthcoming enumeration ol the Bolivian flo¬
ra by the Missouri Botanical Garden and tin1 Na¬
tional Herbarium of Bolivia and of the genus Cle-

Novon 15: 146-155. 2005.

Volume 15, Number 1
2005

litis

Cleome boliviensis

147

ome for Bolivia by l.upita Sanchez (Sanchez, in
prep.) necessitates its formal publication as:

Cleome boliviensis litis, sp. nov. TYPE: Bolivia.
La Paz: “Plantae Andium Boliviensium, Prov.
Larecaja, vicinus Sorata. Challapampa, ad riv-
ium, in schistosis, Reg.[ion] temp[lada], 2600
m. Sept. 1857— Jan. 1858,” G. Mandon 938
(holotype, WIS; isotypes, BM, F, G, GH, K,
LE, LPB, NY, P, PR, S, W). Figures 1-4.

Haec species a Cleome hassleriana et C. spinosa habitu
suffruticoso perenni (vel annuo?), capsula glanduloso-
puberula, seminibus minoribus cellulis bulliformibus ea-
runcularibus membranas fissurales omnino obtegentibus
atque floribus albidis vel albido-roseis distinguitur.

Erect, herbaceous to suffruticose, much-
branched, ill-smelling, coarse, perennial, spiny
herbs or subshrubs, (0.5 — ) 1 — 1 .6( — 2.2) m tall, to 2
m across, densely viscid glandular-pilose to -hir¬
sute generally, the hairs rather short and stiff and
frequently intermixed with many scattered spi-
nules, these often also gland-tipped; stems spiny
and detisely glandular-pubescent, with the paired,
stout “stipular” spines 8—6 mm long, 2—6 mm wide
at the base (where often glandular pubescent), di¬
vergent at 90° to ± recurved. Major leaves 5- to 7-
foliolate, on petioles 2—9 cm long, prominently
armed with ascending or divergent prickles and
spines (f ig. 1); leaflets narrowly elliptic to oblan-
ceolate-elliptic, broadly acute to acuminate at the
tip, attenuate to the base, 6— 15(— 30) X 2— 8(— 10)
cm, rarely larger on vigorous young shoots, often
somewhat eonduplicate, entire (but microscopically
serrulate, the denticulations scarcely more than
gland-tipped hairs), glandular-pubescent and some¬
what scabrous, rarely subglabrous above, glandular
hirsute-pilose beneath, particularly along the mid¬
rib and prominent veins; lateral veins on each side
of the midrib 8 to 12. Racemes often very dense,
elongating to 40 cm or more by the time they have
finished blooming, the primary ones each with up
to 200 or more flowers opening acropetally (i.e.,
estimated by counting the persistent floral bracts),
but with only ca. 20 to 40 buds and 3 to 5(to 10)
open flowers at any one time at the apex; floral
bracts 1-foliolate (or the lowest flowers frequently
subtended by 3-foliolate bracts or rarely lull-sized
leaves), narrowly to broadly ovate- to oblong-ellip¬
tic, acute to obtuse at the tip, cuneate to rounded
or subcordate at the base, mostly 6-15 X 4-12
mm. their 1—8 mm long petioles frequently sub¬
tended by a pair of minute prickles. Flowers on
pedicels ( 1 3— )22— 32( — 4 1 ) mm long; sepals narrow¬
ly lanceolate, acute, 6-10 X 2-3 mm, soon decid¬
uous, densely glandular-pubescent throughout, es¬

pecially toward the tip. the glands often mostly
sessile or subsessile; petals spatulate, 24—35 X 4—
10 mm, showy, basically mostly white, but often
variously flushed with lilac or pale pink, or rarely
dark blood red to a rich pinkish purple, glabrous
or rarely with a few scattered glandular hairs on the
back and top when in bud, the blade obovate-ellip-
tic, rounded, gradually attenuate to the slender
claw, the claw' Y3 to % as long; disk (androgyno-
phore) slightly fleshy, conic, 3— 4(— 5) mm long; sta¬
mens 6, the filaments glabrous, 50—70 mm long,
red, reddish violet, purple, or deep maroon, the an¬
thers (7— )9— 13 mm long; ovary cylindric to fusi¬
form, 10—13 X 1 mm, finely and densely glandular-
pubescent; gynophore 4—7 cm long, glabrous, and
together at least initially with pedicel and silique
often strongly arching upward at 45°. Mature si-
liques oblong to cylindric, sometimes somewhat
clavate (i.e., wider toward the tip), broadly acute to
rounded at the apex, obtuse to long-attenuate at the
base, (4— )5— 8(— 9) X 0.5— 0.8 cm, usually stout with
the valves firm to somewhat lignified, and with the
evenly smooth to scaberulous and continuous (nev¬
er torulose) surface ± sparsely and minutely glan-
dular-puberulent (the relatively large glands sessile
or subsessile on short stalks); stigma capitate, ses¬
sile or subsessile; gynophore glabrous, stout,
straight, (35— )50— 95 mm long; androgynophore 3—
6 mm long; pedicel (15— )25 — 40(— 47) mm long, di¬
vergent to stiffly ascending and continuous with the
gynophore and silique, often arching upward to¬
gether; seeds often many, 100 to 400 or more per
silique, reniform-subspherical or -obovoitl, very
small, 1.5-1. 9 X 1.4-1. 7 X 1.1— 1.3 mm, brown to
dark brown or almost black when mature, smooth
or essentially so, finely longitudinally striate, the
internal cleft cavity oblong to obovate in cross sec¬
tion, narrowed toward the vertex, relatively small;
seed cleft-membranes (i.e., on the 2 ± grooved or
flat sides of the seed connecting the two curved
ends, or “claws”) totally covered over by ± densely
aggregated, small (use 20X lens), varnished, light-
reflecting, oil-containing, bulliform cells which may
spread into, and completely occlude, the mouth of
the cleft and spill over onto the surrounding tesla,
and sometimes even onto the back of the seed,
forming a densely caruncular surface (see below
and Fig. 2); claws subequal, the radicular claw con¬
siderably longer. 2 n = 20.

Habitat and distribution. Foothills and moun¬
tains of southern Peru and Bolivia (Fig. 3), at ele¬
vations of (250-) 1 700-2800(-3500) m, on open,
dry, stony slopes among scattered shrubs, in rocky
beds of dried-up mountain streams, and along

148

Novon

Figure I. Cleome boliviensis litis. — A. Habit, with characteristic ascending young siliques. — B. Petiole with spines
mostly at right angles or somewhat antrorse. A, B based on Ugent 5112. Drawing by Kandis Elliot.

shrubby, moist riversides, from “bosques montan-
osos | cloud forests] con muchas epffitas” down to
transitional humid savannas, in open sandy soils
and arid waste places, rocky roadsides and lields.

at lower elevations weedy and adventive in fields
and along roads, rarely cultivated for its showy
flowers in gardens.

Phenology. Flowering and fruiting intermittent-

Volume 15, Number 1
2005

litis

Cleome boliviensis

149

2 mm

Figure 2. Four assorted seeds of Cleome boliviensis showing characteristic oil-filled bulliform cells in seed cleft and
on seed faces, each from a different specimen. Drawing hy the author.

ly throughout the year but apparently mostly from
October to July.

Vernacular names and uses. Peru. Cuzco: “ Qui -
chita ” (. McCarroll 59), as a “remedy for pains of
rheumatism.” BOLIVIA. Cochabamba: “ Khichita ”
( Baer 223), reported simply as “medicinal,” and
“ Kichita ” (Ugent 4593). where “the leaves when
crumpled and stuffed into the ear are said to cure
earaches” |see litis, 1960: 289, for similar reports
from Africa, where the mustard-oil-containing, ac¬
rid leaf juices, seeds, or whole seed pods of the
nearly pantropic weedy Cleome gynandra (L.) L.
are placed into the ear canal to dissolve ear wax,
and where poultices of the leaves or fruits are ap¬
plied for various purposes, including to relieve the
pain of rheumatism],

Cleome boliviensis is a common and widespread,
mostly low-to- -mid-elevation Andean xerophyte,
with a modern range that evidently now heavily re¬
flects human influence. Not only is it often weedy.

but in Bolivia (, Saravia II 156) and in Argentina
(Pederson 177) it is reportedly cultivated as an or¬
namental in gardens for its “fine pink flowers" (Bo¬
livia, Brooke 50IB). E, k. Balls (ex sehed.) made
the following field observations on C. boliviensis in
Bolivia: “Bushy shrubs to 5 ft. tall and 6 ft. or more
through. Leaves sticky, with an unpleasant smell,
stems hairy and woody, usually red. Flowers pink
with deep maroon stamens." The showy flowers
must make this a beautiful species, though the
abundant spines might well discourage garden cul¬
tivation.

Relationships. Cleome boliviensis is one of a
half dozen or more segregates of the old “C. spi-
nosa” s.l. complex to which all of the larger-flow¬
ered, 5- to 9-foliolate, spiny Cleome plants had
been referred previous to my Ph.D. thesis (litis,
1952). Of the South American taxa, it is apparently
most closely related to the commonly cultivated,
sometimes adventive C. hassleriana Chodat, the

150

Novon

Figure 3. Cleome boliviensis: geographic distribution. (The location of Fiebrig 1375, “Alto Paraguay 21°S,” is ques¬
tionable.)

“Pink Queen,” “Spider Flower,” or “C. spinosa" of
American gardens, an allopatric native of swamps
and riversides from northeastern Argentina and
Uruguay to southeastern Brazil (Minas Gerais), and
perhaps even to the extremely rare, giant-flowered,
also allopatric C. lilloi S. A. Gomez (G6mez, 1953),
native to the lowland swamps of the Chaco, front
northern Argentina to Bolivia and Paraguay (litis,
1967, 1998). Very spiny specimens of C. hassleri¬
ana can be nearly indistinguishable from C. boli¬
viensis , but have, as a rule, fewer spines and less
pubescence, 5 to 9 thinner, larger and more point¬
ed, long-acuminate leaflets, deeper-colored pink to
purple flowers (except for an occasional cultivated
albino), and are less-branched, more herbaceous,
annuals. Most significantly, C. hassleriana has to¬
tally glabrous petals, ovaries, and fruits, and larger

seeds (mostly 2—2.3 mm) that always lack the mi¬
nute oil-inflated bulliform cells on the cleft-mem¬
branes (use 20X lens) that are so characteristic of
C. boliviensis. In contrast to the lowland, riverine
C. hassleriana, Cleome boliviensis is a much more
xeromorphic perennial or short-lived shrub ol the
seasonally dry Bolivian and southern Peruvian
mountains, glandular throughout, including the si-
liques, though it, too, may grow on streamsides or
sometimes, at lower elevations, as a weed in wet
ditches.

Some Cleome Problems in Bolivia

Bolivian species with which Cleome boliviensis
may be confused, especially when in flower, include
the following:

Volume 15, Number 1
2005

litis

Cleome boliviensis

151

Cleome

Figure 4. Cleome boliviensis and related Bolivian laxa: mature siliques, each showing (from top) pedicel, receptacle,
gynophore, and silique [all to same scale; siliques from asterisked (*) herbarium]: (deome werderrnannii Ernst. — A.
Steinbach 6 282 (A, ( ,~l H I . K*). Santa Cruz, Cercado. — B. Coro-Rojas 1438 (LPB, WIS*), Tarija, Bermejos. — C. Nee
37880 (CAS, K, ML, MA, MO, NY, W IS*), Santa Cruz, W-side of the city. —I). Werdermann 2397 (B, MO*. S), Santa
Cruz, Misiones Guarayos. — E. Cleome spinosa s.l., Abbott 10139 (BEREA, MO, US/, WIS*), from San Jose de Chi-
quitos, a peculiar population of C. spinosa with strigose, eglandular pubescence (see text). — I . Cleome spinosa s.l..
Beck 17030 (EBP, W'IS*), Pando, Cobija, a weedy recent introduction into Amazonian lowlands (see text). — G, H.
Hybrid or introgressant of Cleome boliviensis with C. spinosa s.l.. Nee & Saldias 37270 (N\. W IS*). I-N. Cleome
boliviensis (see text: for all, W IS*). — 1. J. Mandon 938 (the holotype). — K. (Jgent 3105. — L. Nee 37588. — M. lewis
35099. — N. 11 gent & Cardenas 4895.

I. The rather uncommon white-flowered and
spiny Cleome werderrnannii Ernst is a Bolivian en¬
demic of low and mid elevations (300—1600 m) that
has small, essentially glabrous, smooth and narrow
siliques (Fig. 4A— D), long-attenuate from far below'
the middle, that are sharply deflexed on a quite
short, ± curved, gynophore (only 12-25 mm long).
Its slightly tubercled seeds are somewhat larger
than those of C. boliviensis and lack caruncular.

bulliform cells. Cleome werderrnannii is evidently
related to the widespread, polymorphic C. spinosa
s.l. complex, perhaps to some Central American
types, and does not seem to introgress with C. bo¬
liviensis.

2. White-flowered Cleome spinosa of certain
common, long-fruited, and torulose phenotypes of
northeastern and central Brazil have recently be¬
come introduced in forested Amazonian lowlands

152

Novon

near the Brazilian border (e.g., Bolivia: Pando,
Cobija, Beck 17030 (LPB, WIS), Fernandez-Casas
8053 (MA. MO. N^. photocopies \\ IS)]. Aside from
the enormously long fruits, to 15-17 cm (Fig. 4F),
they differ in their larger seeds devoid of minute
bulliform earuncular cells.

2a. One collection each from the Bolivian De¬
partments of Santa Cruz (Nee & Saldias 37270, NY,
WIS; Fig. 4G, II) and Cochabamba (Holliday 67.
K) suggests interspecific hybridization with C. bo-
liviensis by repeated putative backcrossing to these
long-fruited C. spinosa phenotypes. The first of
these, a glabrous specimen, has the long (13—14
cm), hanging siliques (Fig. 4G, II) typical of the
Brazilian C. spinosa mentioned above, yet has seeds
with abundant earuncular bulliform cells typical of
C. boliviensis.

3. Some sporadic and puzzling specimens from
northwestern Peru and southeastern Ecuador are
problematic |e.g., Peru: Department Tumbes, Zar-
umilla. quebrada Faical. D. II. Simpson & Schunke
559 (F, N\, US); Dept. La Libertad, Trujillo, Bar-
razu, A. Sagastegui 7879 (Gil, F, k, MO. NY). Ec¬
uador: Prov. El Oro, Zaruma, El Calvario, C. Bon-
ifaz X X. Cornejo 3686 (GUAY, WIS); Prov. Loja,
Macara, G. Hurling N /,. Andersson 22539 (GB,
WIS); Zanibi, F. Vivar 2816 (A All, LOJA, photo¬
copy WIS)]. They grow in irrigation ditches and
nearby desert areas, edges of towns and waste plac¬
es, and have in the past been misidentified by me
as Cleome boliviensis. With better recent collections
available, it became clear that most lacked the dis¬
tinctive earuncular bulliform cells on their seeds,
but that others had eglandular hispid pubescence
mixed with shorter, glandular hairs unique in the
polymorphic C. spinosa complex. These may thus
represent a local variant ecotype or mutant rarely
collected and as yet unnamed (litis & Cornejo, un¬
published), while others may yet prove to be C.
boliviensis or weedy C. spinosa or C. hassleriana
Chodat, recently introduced near Guayaquil, Ec¬
uador, or simply the variable offspring of a hybrid
swarm between these species.

4. Two separate collections with very similar pu¬
bescence. cited in detail below (Abbott 16139 and
Jardin & Mamani 3698), have recently turned up
far away in eastern Bolivia, both from San Jose de
Chiquitos, growing in roadside ditches and marshy
areas. This old, small town, a railroad stop in a vast,
arid plain, is, according to Michael Nee (pers.
comm.), characterized by a hot and seasonally dry
climate resembling somewhat that of the forested
coastal plain to the west and north of Guayaquil,
Ecuador. Superficially, these plants look like hun¬
dreds of other C. spinosa collections, were it not for

their unique, dense, closely appressed, strigose and
totally eglandular pubescence, one that somewhat
resembles that of the Pacific coastal plain desert
plants mentioned above, but even more so, espe¬
cially on tilt1 young inflorescences and sepals. In
addition, Abbott’s collection has mature siliques
(Fig. 4E) with seeds 2.3 mm in diameter, which
distinguish themselves by a pronounced scruffiness
of relatively large, dried and collapsed blisters on
their backs, much larger and more pronounced than
those sometimes found on the seeds of Cleome spi¬
nosa, and quite unlike the minute ones on the seeds
of C. boliviensis. Are these distinctive plants indeed
recent weedy immigrants from the Peruvian coast,
or in fact members of a local autochthonous endem¬
ic that, geographically and ecologically isolated,
speciated allopatrically in this ecotonal border town
between the Chaco (mineral-rich, recent alluvial
soils derived from the Andes) and the Cerrado (nu¬
trient-poor soils derived in situ from the pre-Cam¬
brian Brazilian shield; Michael Nee. pers. comm.)?
Whether endemic or not, these are questions that
need, above all, intense fieldwork and many addi¬
tional collections to be answered. | Bolivia: Santa
Cruz, Chiquitos Province: suffrutescent herb to ca.
2 m tall, petals light whitish purple, filaments pur¬
ple (siliques 6— 7.5 cm long, narrowly torulose, on
slender gynophores 5—6 cm long), San Jose de Chi¬
quitos, ca. 3 km W of town square; roadside ditch
and marshy area, I7°49'S, 60°44'W, ca. 500 m, 6
Feb. 1995 (fl, fr), Abbott 16139 (BEREA, MO. USZ,
WIS); hierba de 1.5 m. flores blancas-rosadas claro
con olor dulce, 2 km E de San Jose de Chiquitos,
yendo hacia Bobore del pozo del agua con Ludwig-
ni, Tessaria [ integrifolia ], suelos areillosos, 18C19'S,
60°63 ' W. 200 m, 15 Nov. 1996 (fl), Jardin & Ma¬
mani 3698 (MO, WIS)].

5. Finally, I have seen a half dozen perplexing
collections of Cleome boliviensis from Madre « le
Dios in southern Peru |e.g.. Plowman & Davis 5088
(WIS)], which have narrower, somewhat torulose si-
liques and smaller white flowers, thus resembling
C. parviflora HBK subsp. psoraleifolia (DC.) litis
(based on C. psoraleifolia DC., Prodr. 1: 239. 1824),
yet have very small seeds (1.5 X 1.4 mm) with
well-developed bulliform cells. That most of them
are reported as weedy suggests hybridization with
C. boliviensis , a problem that, like many others
here, needs study.

Biogkogkaphy and Taxonomy

The three large-flowered southern South Ameri¬
can endemics, Cleome hassleriana, C. lilloi, and C.
boliviensis, constitute a series of geographically re-

Volume 15, Number 1
2005

litis

Cleome boliviensis

153

placing ranges from southeastern Brazil. Paraguay,
and northeastern Argentina west to southern Boliv¬
ia and Peru (litis, 1967: 961. fig. 6; litis. 1998:
438, lig. 5) and suggest a clade that evolved from
a single southern South American ancestor by al-
lopatric speciation. Furthermore, this clade seems
to he ancestral to two other, geographically very
distant and highly restricted, local endemics: the
annual swamp-dwelling Cleome chapalaensis litis
(litis, 1998) of the Lago Chapala Basin of south¬
western Mexico ( Cleome “ pringlei ” [ined.] of litis,
1967: 961, fig. 6>); and the extremely rare local en¬
demic of the Dominican Republic, Cleome domin-
gensis litis (a renaming of Cleome erosa Urban,
1912, not Cleoma erosa (Nuttall) Eaton, 1836,
which is Polanisia erosa (Nuttall) litis, 1958; cl.
litis, 1958). Since* both species grow in wet habitats
similar to those of C. hassleriana and C. lilloi and
at geographically equivalent latitudes but north of
the equator, one may assume that they developed
their macro-disjunctions by avian long-distance
dispersal (Carlquist, 1966, 1981; litis. 1967; Ra¬
ven, 1963), a view supported by recent molecular
analysis (Sanchez, 2002).

Whether the West African swamp and riverside
Cleome afrospina litis (litis, 1967), the only indig¬
enous Old World species of section Tarenaya , is
derived from the same New World clade or another,
closely related one (e.g., C. trachycarpa Klotzsch
ex Fielder), again no doubt by avian long-distance
dispersal (litis. 1967), awaits future molecular stud¬
ies. In any case, C. hassleriana , C. boliviensis , and
C. chapalaensis all have the same number of chro¬
mosomes (n = 10, 2 n — 20), as does the polymor¬
phic C. spinosa s.l. of Central America, the West
Indies, and South America north of Bolivia (Ka-
wano & Kowal in litis, 1998; Costa e Silva, 2000;
litis. Kawano, Kowal, Pazy & Pzywara, unpub¬
lished). Yet only C. boliviensis, among all these
taxa, exhibits die unique glistening bulliform cells
on its small seeds (see below), a specialization pu¬
tatively related to dispersal by ants, and one sug¬
gesting autochthonous evolution in situ of this in¬
teresting species.

Aints, Ahii.s, and Skk.d Stri cti kk
IN I lit: Cl.KOM \CKAK

Seed structure in the Cleomaeeae is of great di¬
agnostic value (litis, 1952, 1967; Ruiz-Zapata &
Fscala. 1995: Ruiz-Zapata. 1995; Sanchez. 2002).
and myrmecochory, suggested by the presence of
usually oily appendages as food for ants, is of spe¬
cial interest.

Various specializations for ant dispersal are

found in the seeds of several groups of Cleomaeeae.
Some are related to each other, others not, these
evidently having arisen independently at least sev¬
en times: (I) Small, light-reflecting, oil-containing,
inflated caruncular cells at the mouth of the seed
cleft and on the cleft membranes, one of the main
diagnostic characters of C. boliviensis, suggest that
these seeds are ant dispersed. (2) The cluster of
some five taxa centering on Cleome rosea Vald s.l.
(litis, 1952; Costa e Silva, 2000; litis, in prep.), a
group of ca. seven eastern Brazilian taxa that vary
from spinose to, rarely, unarmed, and form a closely
interrelated complex in section Tarenaya only dis¬
tantly related to C. boliviensis, have bulliform cells
on the strongly transversely rugose seeds, that,
somewhat different from those of C. boliviensis,
most probably arose independently by convergent
evolution. (3) Bulliform cells almost identical to
those that cover the cleft membranes connecting
the two seed claws of C. boliviensis occur only in
one other annual species of Cleome , a member of
a group of three ebracteate unarmed species, prob¬
ably belonging to Tarenaya (litis, 1952). namely the
glabrous, spineless C. paludosa Willdenow ex Fi¬
elder of lowland northern Argentina, Uruguay, and
southern Brazil, this surely again a case of conver¬
gent evolution. To prove the point, anil most curi¬
ously, such cells are totally lacking in its closely
related sister taxon, the common and weedy annual
C. serrata Jacquin of Central America, the West
Indies, and northern South America, a species
which, except (or the total absence of bulliform
cells on its seeds, differs from its southern South
American relative only by the lack of an elongated
gynophore (litis, 1952. 2001). (4) fihiite different
and more prominent are the large, white, spongy
funicular arils, so-called elaiosomes (Bresinsky,
1963), which occur in the spiny, weedy, and wide¬
spread tropical American C. aculeata U. of section
Tarenaya s.l. and its several tropical South Amer¬
ican relatives (litis, 1952; Ruiz-Zapata & Fscala.
1995; Ruiz-Zapata & litis, 1998; Costa e Silva &
litis, in prep.). (5) Independently evolved, although
similar in appearance, are the white arils ol the
unarmed, weedv Old World C. rutidosperma DC. of
section Rutidosperma litis (litis. I960: 287. fig. 1;
Jacobs, I960; Ruiz-Zapata & Fscala, 1995), which
unlike the New World aculeata group has a quite
open seed cleft. (6) Similar to the spongiform fu¬
nicular arils of C. aculeata, but undoubtedly in¬
dependently evolved as well, and for the third time,
are the generically characteristic and prominent fu¬
nicular arils of the 26 species (Cochrane, 1997) of
the annual to perennial, herbaceous to woody New
World genus Podandrogyne Ducke (Woodson,

154

Novon

1948; litis & Cochrane, 1989; Cochrane, 1977.
1978, 1997), which may have had their origin in
(7) the quite inconspicuous small (funicular?) aril
that projects from the very tip of the radicular claw
of the seeds of the widespread, low-elevation an¬
nual C. pilosa Bentham, which, it so happens, is
clearly related, albeit distantly, to Podandrogyne
(litis, 1952; litis ct al„ 2005; litis & Cochrane,
1989; Ruiz-Zapata & litis, 1998: 151) as verified
by morphological assessments and recent molecular
studies (Hall et al., 2002: 1830; Sanchez. 2002).

Paratypes. ARGENTINA. Corrientes: Parque Ou¬
tre., Pedersen 177 (P) [probably cultivated in a city park|.
BOLIVIA. Prov. unknown: Vinto, W. 47. A. Brooke 5018
(BM, I ). Beni: Distr. Ballivian, Serranfa de Pildn l.ajas,
Bosque Montafioso, al borde de camino, San Borja, 75 km
haeia La Paz, Si. G. Berk 6087 (W IS); La Cumbre, Berk
13251 ( W IS); vertiente oriental, 21 km de Yueumo, I). N.
Smith, G. Quintana & V. Garcia 13202 (LPB, MO, \\ IS).
C.hiiquisacu: Distr. Oropeza, rocky roadside, base of cliff,
16 km NE of Sucre on Aiquile Rd., I ). Ugent A' 47. Gdr-
denas 1805 (IBUG, MY, QCA, WIS); Distr. 'lomina, Mu-
suru (16 km de Padilla), rumbo a Monteagudo, C. Saravia
1 1 156 (CTES); Sucre, open, stony, dry slopes, E. K. Balls
6187 (DAB, E, K. NA, UC, US); des ravine de Veleo mayo
[sic! = Pilcomayo?] prbs Chuquisaca, A. D'Orbigny s.n.
(P); Distr. B. Baleto, 700 m N of town square of Nuevo
Mundo, on road to Villa Serano, then 300 m left (downhill)
of road, I). M. Spooner, H. G. van der Berg & 47. /,. Ugarte
6786 (WIS). Cochabamba: Cervecerfa Colbn, Cocha¬
bamba, in a stony bed of a mountain stream. Balls 6205
(E, UC, US); Tunari, K. GraJ 521 (NY); 20 km E of Co¬
chabamba, among stones, thickets on riverbank, IT ./.
Eyerdam 24752 (F, K, NA); 2 km N of Cochabamba, Ug¬
ent 4503 (P. U, US, W IS); Tiquipaya, 10 km N of Cocha¬
bamba, Ugent 5003 (I, II,, UC, WIS); Distr. Carrasco, Co¬
chabamba 142 km from Cochabamba at Santa Cruz, Beck
6830 (WIS); roadside thickets, km 180 on Cochabamba—
Sta. Cruz road between Duraznillo X Challuani, Ugent
5105 (G, K, NY, UC, US. WIS); rock wall crevice, Tuti-
mayo, 25 km E of Cochabamba, Ugent 4643 (Cll. K, SMF,
WIS); Rio Mi/.que, Mizque [riverside thickets|, C. Troll
146-1 (B, M); Tako-Tako near Mizque, Brooke 5016 (BM,
F, NY); stony valley near stream, Cochabamba. T. G. Tulin
1612 (MO); Liriuni, B. Scolnik A.- B. Lull 500 (SI, WIS);
Distr. Ayopaya, euenca Rfo Tambillo, Independencia, B.
Baar 223 (LPB, W IS); San Benito, 44 km de Cochabamba
a Sta. Cruz, J. Ferndndez-Casas 7721 (MA, MO, NY); l.a
Maica, V/. Cardenas 3440 (S, US); Prov. Carrasco al borde
de camino y de las chacras, Cochabamba, km 142 haeia
Sta. Cruz, Prov. Carrasco, Beck 6830 (W IS); valle de Co¬
chabamba. J. Steinbach 8741 (RM, F, G. GIL IM. K. M.
NY, PH, S, U, UC, US, /); Tunari, B. F. Steinbach 117
(DAV, GH, MO. NY. UC, WIS); progeny of seeds collected
in Cochabamba (grown in Washington, D.C.), B. 47. Bing
* L F. Bishop 7716 (B, IBUG, K. MARACAY, MEXU,
QCNE, TEX, UB, US, USM. W IS); Cochabamba, 0. Buch-
lien 2371 (NY. US); Pairumani, Musch 16 (SI). La Paz:
Abel Iturralde Distr., lado de carretera en la handa del
Rfo Tekeje, a 10 km tie Ixiamas, Ixiamas Tacana, G. Bour-
r/v 2771 (111 D. LPB, UMSA, WIS); delta que forrnan el
arroyo Arana y el Rfo Tuichi, pozo Yariapo X-l, B. Kuno
353 (LPB, W IS); Distr. Inquisivi, Inquisivi, 3 km haeia
Cireuata, Beck 4477 (W IS); 4 km SF of Inquisivi, 47. Nee

37588 (NY. W IS). M. Lewis 35000 (LPB. MO. WIS); Rfo
Sara at Rancho Cuesta Sara, Inquisivi, Lewis 882007
(LPB. MO, WIS); Distr. Nord-Yungas, Milliquaya, Buch-
tien 4330 (US); Hacienda “FI Choro,” Coripata, Buchtien
8002 (NY. US); Isapuri, l>. E. Williams 684 (NY, l S); Rfo
Palea valley. Ml. Illimani, B. Julio 110 (C. US); Coroico,
./. Solomon, B. Stein & 47. Uehling 12066 (MO. W IS 2
sheets); forest edge on bank ol Rfo Beni, 20 km upstream
from Rurrenabaque, L. G. Holliday 60, 67 (K); Rurrena-
baque. mossy boulders near river. I). C. Daly et al. 6638
(NY, W IS); Reyes, Rurrenabaque, Rfo Beni, Fleischman
476 (S); hrecha 22. heliport 14, Rfo Tuichi, A. Fournet
480 (W IS); 12 km along Rfo Santa Barbara, below Yolosa,
rd. to Caranavi. Solomon 8808 (MO. W IS); Distr. Sud-
Yungas, Arroyo San Antonio, bajando la Concesion Trop¬
ical (ribera del rfo), Alto Beni, El. Vargas, B. Seidel & L.
Cruz 2230-A (LPB, W IS); Inicua, 0. 47. B.(ERTS) 373
(W IS); rd. to Chulumani, 23 km F of Unduavi, 7.’ B. Groat
51534 (MO. W IS); FI Chaco, Valle de Unduavi, A. Lour-
teig 2625 (P): Cafiamina. II. H. Busby 66 (BKL, NY); Valle
del Rfo Unduavi de la escuela de Sirupaya por abajo B.
Seidal & E. Richter 1250 (W IS); Sirupaya near Yanacachi,
Buchtien 566 (F. G, GH, NY, US, W ); Cotafia, am Illimani,
Buchtien 3173 (NY, US); dry rocky soil on roadside 21
km W of 1 ’uente Villa X 26 km F of Unduavi, G. Davidson
5003 (RSA); Puente Villa 2 km arriba del Rfo Tamanpaya,
Beck 17781 (LPB, WIS); rd. to Puente Villa, 5 km W of
Chulumani, D. 47. Spooner, B. G. van den Berg & W.
Garcia 6734 (LPB, W IS); lowland forest along Rfo Satar-
iapo, 12 km SW' ol Ixiamas, A. Gentry & li. Foster 70883
(W IS); |probably La Paz| Yungas, 1890, 47. Bang 485 (C,
G, LI), I.F, I.IL. M. MO. NDG, NY. PH, R. US, W. WIS,
Z). Santa Cruz: roadside between Santa Cruz— Cochabam¬
ba, km 240, IE ./. Badcock 468 (K [an exceptionally spiny
plant]); Manuel Maria Caballero Distr., ladera eon bosques
ralo, cerea de Comarapa en direccion a Fpizana, Ferndn¬
dez-Casas 7001 (MA, MO, NY); Bachraender [borders of
creek], Samaipata, C. Troll 1063 (B): bosque montafioso
con poeo influencia humana, Prov. Cordillera, Carapari-
cito, 40 km de Camiri. B. de Michel 181 (W IS); 35 km
de Camiri, Michel 65 (W IS); Distr. Vallegrande, low, semi-
deciduous forest. 10 km F of Guadaloupe, valley of Rfo
Piraymiri, “Juntas de Guaricongas,” 47. Nee, P. Bettella &
.47. Saldias 36104 (LPB, NY, W IS); 5 km W NW of Com¬
arapa on road to Cataiinas, I). Ugent & V. Ugent 5112 (B.
UC, W IS). Sara [now Prov. Ichilo]: Cerro de Amborb,
play itas arenosas del Yrama, J. Steinbach 2071 (GH, LIL,
NY, SI); San Ignacio, ./. Steinbach 3537 (G, (ill. K. LIL,
SI); Distr. Ichilo, Parque Nacional Ambord, tropical ev¬
ergreen lores!, sandy edges of Rfo Saguayo, 1 km down¬
stream from Ouebrada Yapoje. Nee 38152 (NY. WIS); ev¬
ergreen tropical lowland forest along Rfo Moile, opposite
Campamento Moile, 6 km SW of Villa San German, 47.
Nee & E. Chavez 51613 (NY, W IS); Rfo Saguayo, Nee &
Saldias 37270 (NY. W IS). PARAGUAY. Chaco: Alto Par¬
aguay, Fiebrig 1375 (Z) [this location is questionable],
PERU. Cuzco: Distr. Cuzco, Camisea, Campamento Ar-
mihuari, P. Acevedo- Bdgz. 0230 (NY, US. WIS); Anta, FI
Chaeean, Brunei 584 (M(), W IS); Cillapuyu, FI Chaccan,
Brunei 350 (MO); Distr. Convencion, Amaibamba, Liuc-
umayo, G. Vargas 4562 (W IS); Distr. Echarate, Armihuari
Well Site, P. Niinez & Beltran 10863 (US, W IS); Distr.
Paucartambo, Atalaya, hillside X riverside, jet. Rfo Car¬
bon w ith Rfo Madre de Dios, B. Foster 3037 (US); abierta
pedregoso, Pileapata, La Raya, Vargas 1 1320 (CUZ, W IS);
arenoso, cerea al rfo, Atalaya-Kosnipata, Vargas 13087
(W IS); Distr. Quispicanchis, Tfo-San Pedro, Vargas 0730

Volume 15, Number 1
2005

litis

Cleome boliviensis

155

(A, CUZ, Vi IS); ruderal, Chili-Chili, Marin 2436 (LIE);
horde camino, en lugares abicotos, Murayaca, Mareapata,
Vargas 3756 (I II., MO); Quince Mil. Vargas 7772 (MO).
Madre de Dios: Distr. Manu, rd. from Shintuya to Sal-
vaeion, T. Plowman & E. 1C Davis 5088 (GH). Puno:
Distr. Sandia, Sandia, /.. Hoogte A' C. Roersch 3447 (MO);
trail from Aricoma Pass to Sto. Domingo, D. McCarroll 59
(CUZ, MICH).

Acknowledgments. 1 wish to thank my former
student, master ethnobotanist and potato specialist
Donald Ugent, Professor Emeritus of Botany, South¬
ern Illinois University— Carbondale, for collecting
the excellent series of Cleome boliviensis popula¬
tions in 1963; Lupita Sanchez for help with the
Bolivian specimen citations; our Botany Depart¬
ment artist, Kandis Elliot, for her careful work on
the illustrations; Euz Marfa Gonzalez for translating
the abstract into Spanish; Barbara NeufHer and an
anonymous botanist for carefully reviewing the
manuscript; Barbara Schaack, Eiz Lipsett, Xavier
Cornejo, and Josh Sulman for dependable and
cheerful help with my manuscript; and especially
Victoria Hollowell, Michael Nee, and Theodore S.
Cochrane for the exemplary care with which they
whetted the final manuscript to perfection.

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nus Cleome (Brassicaceae— Cleomoidae) based on Chlo¬
roplast Sequence Data. Master's Thesis, University of
Missouri— St. Louis.

Woodson, R. E., Jr. 1948. Gynandropsis, Cleome. and Po¬
dandrogyne. Ann. Missouri Bot. Card. 35: 139—146.

Carex hangzhouensis and Section Hangzhouenses, a New Species
and Section of Cyperaceae from Hangzhou, Zhejiang,

Eastern China

Jin Xiao-feng', Ding Bing-yang1-2*, and Zheng Chao-zong'

'College of Life Sciences, Zhejiang University, Hangzhou, Zhejiang 310012,
People’s Republic of China, jinxf583@hotmail.com
2School of Life & Environment Science, Wenzhou Normal College, Wenzhou, Zhejiang,
325027, People’s Republic of China. *eorresponding author, dingby@mail.hz.zj.cn

Abstkact. Carex hangzhouensis C. Z. Zheng, X.
F. Jin & B. Y. Ding, a new species of Cyperaceae
from eastern China, is described and illustrated,
and the distinguishing characters are discussed.
Owing to its peculiar habit, Carex hangzhouensis
does not fit into an existing section. Therefore,
based upon this type, the new Carex sect. Hang¬
zhouenses is identified, and a key to this and the
re lated sections in eastern and southern Asia is
provided.

Key words: Carex, China, Cyperaceae, section
Hangzhouenses , Zhejiang.

Carex L. is the largest genus within the family
Cyperaceae. The genus is cosmopolitan with over
2000 species distributed in various habitats (Nel-
mes, 1951; Ohwi, 1953; Dai et al., 2000). China
has nearly 500 species of Carex (Dai et al., 2000).
During the course of collecting and identifying Car¬
ex specimens collected in Zhejiang, new records,
new species, and new varieties have been discov¬
ered (Zheng, 1989, 1993; Jin et al., 2003, 2004a,
2004b). Among the specimens of Carex collected
in 1978 and 1986 in Hangzhou City, Zhejiang Prov¬
ince, some specimens are peculiar in their spike
arrangement. However, the perigynia of these spec¬
imens are immature. We repeatedly attempted to
find mature specimens at the type locality, Lingyin.
Hangzhou City, and were finally successful in May
2003. Compared with other species of the genus,
specimens with mature perigynia have a subrace-
mose panicle, a terminal staminate spike, the lat¬
eral branched spikes arising from a utriculiform
cladoprophyll with a single pistillate flower (a per-
igynium), and a unique aehene shape.

All tlu *se features distinguish the taxon from oth¬
er sections in Carex subg. Indocarex Baillon
(Clarke, 1904; Kukenthal, 1909; Ohwi, 1936; Nel-
ines. 1951; Dai et al.. 2000). Therefore, we identify
the taxon as a new species and establish a new
section based on it.

Carex sect. Hangzhouenses C. Z. Zheng, X. F.
Jin B. Y. Ding, sect. nov. TYPE: Carex
hangzhouensis C. Z. Zheng, X. F. Jin & B. Y.
Ding.

Haec sectio Carici sect. Mundae affinis, sed ah ea in-
florescentiis suhracemoso-panieulatis, spica terrninali
staminata hasi eramosa, spieis secundariis lateralibus ex
cladoprophyllo saccato floreni pi si i Hat uni solitarium evo-
lutum gerente orlis, perigynio apiee in rostrum curvum ore
hidentatum contractu ah|ue achenio apice in rostrum cur¬
vum hrevem ahrupte contractu differt.

Bracts shortly foliaceous, long vaginate; sheaths
smooth. 1 .5— 2.5 cm long. Panicle subracemose; ter¬
minal spike staminate, long pedunculate, base of
the staminate spike unbranched; lateral secondary
inflorescences 1 to 3, racemose, each with 6 to 10
closely branched spikes, androgynous. Branched
spikes arising from a saeeulate cladoprophyll with
a single developed pistillate flower. Perigynia gla¬
brous, trigonous, multinerved, apex with a genicu¬
late, bidentate beak. Achenes trigonous, concave
on lateral surfaces, apex retuse with short persistent
style base, base with vestigial rachilla.

The new species is distinguished within Carex
by its lateral branched spikes arising from a sae¬
eulate cladoprophyll with a single developed pis¬
tillate flower. The laterally concave aehene shape
uniquely distinguishes the taxon. In particular,
Carex hangzhouensis may be contrasted with other
species in Carex sect. Mundae Kiikenthal and sect.
Japonicae Kukenthal. which are postulated as its
closest putative relatives. Species in Carex section
Mundae differ in their racemose inflorescences ter¬
minated by androgynous spikes as well as sym¬
metrical perigynia with erect beaks that are neither
dentate nor toothed. Species in Carex seel. Japon¬
icae have spieate inflorescences with terminal an¬
drogynous spikes. Perigynia are symmetrical and
have straight beaks, and the beaks are obliquely
truncate. Characterizing the new species, the peri-

Novon 15; 156-159. 2005.

Volume 15, Number 1
2005

Jin et al.

Carex from Hangzhou

157

gynia of Carex hangzhouensis are asymmetrical,
with a geniculate beak to 2 mm long. This beak is
bidentate, with teeth evident, to ca. 5 mm long.

This new section is currently monotypic, distrib¬
uted in Hangzhou, Zhejiang Province, eastern Chi¬
na.

Carex hangzhouensis is the sole species of its
section, but shares its fertile cladoprophyll char¬
acter with five species in eastern and southern
Asia, namely, C. satsumensis Franehet & Savatier,
C. munda Boott, C. fragilis Boott, C. dissitiflora
Franehet, and C. yulongshanensis If C. hi. Carex
satsumensis belongs to section Japonicae and the
other four species belong to section M undue Ku-
kenthal (kiikenthal, 1909; Ohwi, 1936; Nelmes,
1951; hi, 1990; Dai et ah, 2000).

Species of the other sections of Carex subg. In-
docarex differ in that they have lateral spikes ex¬
tending from a sterile (empty) cladoprophyll (Nel¬
mes, 1951). A key to the above-mentioned sections
follows:

Kky to Carex sect. Hang/houenses and Related Sec¬
tions in Eastern and Southern Asia

la. Inflorescence spicate; spikes densely arranged;

bracts unsheathed . sect. Japonicae

lh. Inflorescence racemose or paniculate; spikes
loosely arranged; bracts long sheathed.

2a. Inflorescence racemose, terminal spike an¬
drogynous, base branched; perigynia sym¬
metrical, with a smooth, erect and truncate
beak; aehenes not concave on lateral surfac¬
es . sect. Mundae

2b. Inflorescence paniculate, terminal spike sta-
minate, base unbranched; perigynia asym¬
metrical, with a scabrous, geniculate and bi¬
dentate beak; aehenes concave on lateral
surfaces . sect. Hangzhouenses

Carex lumgzhouensis C. Z. Zheng, X. F. Jin &
B. Y. Ding, sp. nov. TYPF: China. Zhejiang:
Hangzhou City, hingyin, Feilaifeng Mt., on
rocks in forests, 50 m. 30°14,26”N,
120°05'56"E, 13 May 2003, Y. F. Jin & F ./.
Wu 0702 (holotype, HZh; isotype, PE). Fig¬
ure I .

Haec species Carici yulongshanensi affinis, sed ah ea
inflorescentiis secundariis quaque ex spicis secundariis 6
ad 10 constante. squamis staminatis oblongis ca. 0 mm
longis, pistil latis late ovatis ca. 8.5 mm longis 3-nervis
glabris apice in aristam ca. 3 mm longam desinentibus.

Herbs perennial; rhizomes short; culms densely
caespitose, 30—60 cm tall, tenuous, trigonous, gla¬
brous. heaves longer than culms, plane or margins
involute, 2.5—4 mm wide. Bracts shortly foliaceous,
shorter than inflorescences, long vaginate; sheaths
smooth, 1 .5— 2.5 cm long. Panicle subraeemose;

secondary inflorescences 1 to 3, racemose, lateral;
terminal spike and uppermost secondary inflores¬
cence binate, others solitary. Terminal spike sta-
minate, clavate, 4—5.5 cm long, peduncle 3.5—7 cm
long; each secondary inflorescence with 6 to 10
closely branched spikes; branched spikes androg¬
ynous, arising from a sacculate cladoprophyll with
a single developed pistillate flower, terminal sta-
minate spike 1.5—2 cm long, much longer than lat¬
eral spike; lateral spike apex with 4 to 6 staminate
flowers, pistillate spike with a single perigynium.
howermost peduncle to 8 cm long. Staminate scales
oblong, ca. 8 mm long, membranous, stramineous,
apex obtuse, mucronate; pistillate scales broadly
ovate, ca. 8.5 mm long, subcoriaceous, green, 3-
veined, glabrous, apex acuminate, shortly aristate;
arista 3 mm long. Perigynia obovoid, obtusely tri¬
gonous, 6.5—7 mm long, brownish green, glabrous,
multiveined, base attenuate, apex contracted with
a geniculate beak; beak ca. 2 mm long, margin sca¬
brous; teeth ca. 0.5 mm long, bidentate. Aehenes
ellipsoid to obovoid at maturity, subsessile, trigon¬
ous. castaneous, concave on lateral surfaces, 3.5—
4 mm long, apex refuse with a short persistent style
base; style base incrassate; stigmas 3; vestigial
rachilla ca. 2 mm long.

Flowering and fruiting April— May.

This new species is similar to Carex yulongsha¬
nensis P. C. Ei, from which it differs in having the
inflorescence subraeemose paniculiform, the ter¬
minal spike staminate, the base unbranched, each
lateral secondary inflorescence base with 6 to 10
cdosely branched spikes, staminate scales oblong,
ca. 8 mm long, pistillate scales broadly ovate, ca.
8.5 mm long, veins glabrous, apex aristate, arista 3
nun long, perigynia glabrous, beak bidentate,
aehenes concave on lateral surfaces, apex refuse
with a short persistent style base.

Paratypes. CHINA. Zhejiang: Hangzhou City, Lin-
gyin, Feilaifeng Mt., on rocks in forests, 30 m, I May
200.3, X. F. Jin & II. Wang 0657 (HZH 2 sheets), 0658
(MO), 0665 (HZU), 0666 (HZF). 18 Apr. 1080. C. Z.
Zheng 3015 (HZU), I May 2003, II. Wang 0360 (HZU),
0371 (MO). 4 May 200.3. II. Wang 0378 (HZU 3 sheets).
1.3 May 2003, X. F. Jin & F. ,/. Wu 0703 (HZU); Hangzhou
City, Nangaofeng Mt., in forests, 5 Apr. 1978, ('.. /. Zheng
s.n. (HZU).

Acknowledgments. We express our gratitude to
8. Y. Eiang (CAS) and E. k. Dai (CAS) for contin¬
uous support and encouragement, to Victoria C.
Hollow el I (MO) and Amy McPherson (MO) for ed¬
itorial help with the manuscript, lo two anonymous
reviewers for their comments, to E. J. Wu (CAS) for
preparing the illustration, and to E. J. Wu (CAS)
and H. Wang (ZBS) for kind help and collections.

158

Novon

h igure I. Care. x hangzhouemis (.. Z. Zheng, X. F. Jin & B. Y. Ding. Drawn from the holotype and isotype (X. F. Jin
& F. J. Wu 0702). — A. Habit of plant. — B. Pistillate scale, perigyniuin with style branches and staminate (lowers
exserted. — C. Staminate scale. — D. Pistillate scale. — K. Perigyniuin. — K Aehene. — G. Kachilla and perigynium.

Volume 15, Number 1
2005

Jin et al.

Carex from Hangzhou

159

Literature Cited

Clarke. C. B. 1904. List of the Cariees of Malaya. J. Linn.
Soc. Hot. 37: 1-10.

Dai, L. k., S. Y. Liang, P. C. Li X V C. Tang. 2000. Carex
(Cyperaeeae— Carioideae). In: L. k. Dai & S. Y. Liang,
Flora Keipublicae Popularis Sinieae 12: 56—528. Sci¬
ence Press. Beijing.

Jin, X. K, C. /. Zheng & B. \. Ding. 2003. Carex sub-
capitata, a new species of the Cyperaeeae from Zhe¬
jiang, China. Acta Phytotax. Sin. 41: 566-568.

- , - & - . 2004a. New records of Carex

L. in Zhejiang from China. J. Wuhan Bot. lies. 22: 49—
51.

- . - & - . 20041). New laxa of Carex

(Cyperaeeae) from Zhejiang, China. Acta Phytotax. Sin.
42: 541-550

Kiikenthal, G. 1909. Carex (Cyperaeeae— Carioideae). In:

A. Kngler, Das Pflanzenreich, IV, 20 (Heft 38): 67—767.
Engelmann, Leipzig.

Li. P. C. 1990. New taxa of the Cyperaeeae from Heng-
duan Mountains (2). Acta Bot. Yunnan. 12: 137—150.
Nelmes, F. 1951. The genus Carex in Malaysia. Uein-
wardtia 1: 221-450.

Ohwi. J. 1936. Cyperaeeae Japonicae 1. A synopsis of the
Caricoideae of Japan, including the knriles, Saghalin,
korea, and Formosa. Mem. Coll. Sei. kyoto Imp. Univ.
Ser. B 1 1: 229-513.

- . 1953. Carex (Cyperaeeae). Pp. 164-216 in J.

Ohwi, Flora of Japan. Shibundo Press, Tokyo.

Zheng, C. Z. 1989. A study of the Cyperaeeae Family in
Hangzhou. J. Hangzhou Univ. (Nat. Sei.) 16: 460-470.

- . 1993. Carex (Cyperaeeae). In: Q. Lin. Flora of

Zhejiang 7: 286—322. Zhejiang Science and Technology
Publishing House. Hangzhou.

Passiflora xishuangbannaensis (Passifloraceae):
A New Chinese Endemic

Shawn E. Krosnick

Department of Evolution, Ecology and Organismal Biology, Ohio State University,
1315 Kinnear Road, Columbus, Ohio 43212-1157, U.S.A. krosnick.2@osu.edu

Abstract. Passiflora xishuangbannaensis , a new
species endemic to Yunnan Province, China, is de¬
scribed and illustrated. It is distinguished from oth¬
er members ot Passiflora supersection Disemma by
being completely glabrous, having one to two flow¬
ers per node, thickened coronal filaments that are
brown to purple at the base, deeply bilobed leaves
with strong variegation along the three primary
veins, and a v-shaped arrangement of numerous ab-
axial laminar nectaries.

Key words: China, Decaloba, Disemma, Passi¬
flora, Passifloraceae, Xishuangbanna, Yunnan.

Pass I flora I,, is a primarily Neotropical genus
with more than 530 species worldwide. Of these,
approximately 20 species are native to the Old
World, primarily distributed throughout mainland
India, China, Southeast Asia. Australia, and the Pa¬
cific (De Wilde, 1072). All of these species are
placed in subgenus Decaloba, supersection Disem¬
ma (Lahillardiere) J. M. MaeDougal & Keuillet. As
part of a phylogenetic study of Disemma, fieldwork
in Yunnan Province, China, was completed, and
during the course of this research a new species
was found. At the time of collection, all plants were
sterile; cuttings were taken, and the plants have
since flowered continuously in the Plant Biology
Greenhouses at the Ohio State University, Colum¬
bus, Ohio, U.S.A. Descriptions of the flowers and
fruit were made based on the cultivated material.

Passiflora xishuangbannaensis Krosnick, sp.
nov. TYPE: China. Yunnan: Xishuangbanna,
Jinghong County, N side of Da Du Gang Town
near 2765 mile post on llwy. 213, on side of
trail in shady area near stream ca. 1.5 km E
of llwy. 213, mixed secondary forest and ag¬
riculture, 1 2(H) m, 22°19.60'N, 100°58'E, 3
Mar. 2003, S. E. Krosnick & II. Wang 254 (ho-
lotype, OS: isotypes, IIITKC. KUN. MO. NY).
Figure 1.

A Passiflora altebilobata planta glabra omnino, glandi-
bus petioli dispositis prope laminam, foliis minus penitus
bilobatis, nervo medio valde variegato, nectariis numerosis

dispositis ad lineam sub angula 20° abeuntibus, inflores-
centiis 1-2 fioris, floribus grandioribus, filamentis incras-
satis coronarum purpureo-brunneis basi differt.

Weakly tendrillate, clambering vines, 1—3 m
long, glabrous throughout; stem subtriangular, 2.5-
5 mm diam.. new shoot growth straight. Ueaves al¬
ternate; stipules 0.5—1 X 1 mm, lanceolate, sessile,
margins entire, deciduous; petioles 1 .5—2.5 cm
long, often with two short-cylindrical, glandular
nectaries in the distal quarter, tin1 nectaries 1-2
mm diam., short-cylindrical to I mm long, tlx* nec¬
tariferous surface borne terminally; lamina 4. 5-8. 5
X 2.5—5 cm, white- variegated adaxially along the
three major veins, margins entire, the lateral lobes
greatly exceeding the midvein, creating a sinus
about half as long as the leaf, midvein mucronate,
muero 1-2 mm long, abaxial surface light green,
often with purple venation near base of lamina,
laminar nectaries 8 to 12(to 15), discoid ami ses¬
sile, arranged in two lines divaricating at a 20° an¬
gle from the midvein and running partway down the
lateral lobes. Two flowers per node, one flower
sometimes aborting during maturation; pedicels
1.5-2 cm long, bearing 3 floral bracts; floral bracts
linear, 1-2 mm long, sessile, alternate and three-
ranked, 0.5—1 cm beneath the articulation. Flower
buds sub-conic; flowers 3. 2-3.8 cm diam. at anthe-
sis, with 3—5 mm long stipe, orientation of flowers
upward, petals and sepals completely reflexed at
anthesis, smelling strongly of Ive; sepals If)- 18 X
5-7 mm, triangular-ovate, acute and rounded at
apex, abaxial surface light green, strongly veined,
adaxial surface white basally, green at apex, mar¬
gins membranous; petals 10-12 X 2—3 mm, lan¬
ceolate. white; hypanthium 0.8-1 cm wide; coronal
filaments in 2 series, outer series 1 cm long, with
ca. 33 to 35 filaments, thickened, purple-brown in
the basal third, the upper portion bright yellow,
wide spreading at anthesis, inner series 5 mm long,
with ca. 00 to KM) filaments, purple-brown with a
yellow-capitate apex; operculum 3—4 mm high, pli¬
cate, green with purple striations; limen 5—6 mm
diam., annular, green, purple at outermost edge;
nectary yellow, 2 mm deep; staminal filaments con-

Novon 15: 160-163. 2005.

Volume 15, Number 1
2005

Krosnick

Passiflora xishuangbannaensis

161

Figure I. Passiflora xishuangbannaensis krosnick. — A. Habit, shoot tip with buds and flower at anthesis. — B. Cross
section of flower with detail of coronal filaments, limen, and operculum. Drawn by the author, from the holotype,
krosnick & Wang 254 (OS).

nate. 5—7 mm along androgynophore, the free por¬
tions 5—6 mm long, flat; anthers 3—4 mm long, ver¬
satile. elongated with a small basal protrusion;
ovary 4—6 X 2—3 mm, ellipsoid, subtriangular in
transverse section, sessile on the gynophore, gla¬
brous; styles 5—7 mm long including stigmas, stig¬

ma 1—2 mm wide, obliquely expanded at the apex,
stigmatic surface papillate. Fruit 1 .5 X 1 cm, ovoid,
dark blue al maturity; arils yellowish orange, the
distal margin fimbriate. Seeds 7 to 10 per fruit, 5—
6 X 3-4 mm, 2 mm thick, obovate, foveolate-retic-
ulate with 70 to 80 foveae per side.

162

Novon

Fable 1. Characteristics

distinguishing Passiflora xishuangbannaensis

from P. altebilobata.

P. xishuangbann a ens is

P. altebilobata

Pubescence

glabrous

hirsutulous

Stem shape

subtriangular

round

Leaf variegation

present on all major veins of the land- limited mottling present on juvenile

ua in both juvenile and adult leaves leaves, absent in adult leaves

Leal shape

leaves strongly two-lobed with sinus
0.2— 0.3 times the length of the
lateral lobes

leaves strongly two-lobed, mid lobe
truncate, sinus 0.3— 0.0 times the
length ol the lateral lobes

Petiolar nectaries

sometimes absent; if present, paired,
located in the distal quarter of the
petiole

paired, located midway along the
petiole

Abaxial laminar nectaries

8— 12(— 15), discoid, arranged at a 20°
angle from the midvein

2(— 4), paired near the apex of the
midvein

Flowers per node

1-2

10-20

Flower diameter at anthesis

.3.2— 3.8 cm

0.75-1 cm

Coronal filaments

Outer series

thickened to 10 mm, purple-brown at
base, upper 2A yellow

thin to 5 mm. white throughout

Inner series

5 mm long, purple-brown will) yellow-
capitate apex

1 mm long, pale yellow with
subeapitate apex

Fruit size

1 .5 X 1 cm, ovoid

1 X 1 cm, round

Habitat

1200 m, sunny or shady, wet edge of
forest

000 m. wet. shady, in dense forest

Phenology. Specimens not flowering in wild
population at (lie time of collect ion (early March);
flowering continuously in greenhouse conditions in
Columbus, Ohio.

Etymology. The specific epithet is based on the
unique area of endemism, the Xishuangbanna Pre¬
fecture, which encompasses three counties (Jingh¬
ong. Mengla, and Menghai) in Yunnan, China.

Discussion

Passiflora xishuangbannaensis was collected in
Jinghong County in the Xishuangbanna Prefecture
from one of three individual plants observed at the
type locality. Upon later visits in 2003, two of the
larger plants were noted to have disappeared. The
habitat is sandy wet forest openings near streams
in both sunny and shady locations, with agricultural
fields bordering the opposite sides of the streams.
The site of this collection is now under construction
for a new highway and the forest may no longer be
present. Thus, for conservation purposes, its status
should be considered endangered. However, given
the proximity of similar habitats in Mengla and
Menghai Counties, it is possible that P. xishuang¬
bannaensis may still be found there.

Passiflora xish a a ngba nna ens is was not in flower
at the time of collection. Cuttings were made for
cultivation, and there was no observed difference
between the vegetative aspects of the field-collected

and cultivated material. The cultivated plants have
flowered continuously throughout the year in the
greenhouse; however. Passiflora xishuangbannaen-
sis does not appear to be self-compatible. Hand pol¬
lination was performed using pollen from P. xi-
shuangbannensis as well as other closely related
species, including I*. cupiformis Masters and P. pap -
ilio H. L. Li. All self-pollination events failed,
whereas all out-crossed flowers set fruit. Viability
tests of the seeds were not performed. Given the
self-incompatibility observed in this species, the
conservation of more than one haplotype is impor¬
tant.

Passiflora xishuangbannaensis is similar to the
Yunnan native P. altebilobata Hemsley, but is easily
distinguished by several morphological and habitat
features (Table 1). Passiflora xishuangbannaensis is
a weakly tendrillate, clambering herb, while P. al¬
tebilobata is a strong climber. Passiflora xishuang¬
bannaensis is completely glabrous, whereas P. al¬
tebilobata is hirsutulous on the stems, leaves, and
inflorescences. 'File abaxial laminar nectaries are
arranged in a v-pattern in P. xishuangbannaensis ,
while in P. altebilobata there are two (to four) large,
paired nectaries located near the apex of the mid¬
vein. Additionally, P. xishuangbannaensis has one-
to two-flowered inflorescences, while P. altebilobata
has 10 to 20 flowers per inflorescence. Passiflora
xishuangbannaensis has larger and more colorful

Volume 15, Number 1
2005

Krosnick

Passiflora xishuangbannaensis

163

flowers than P. altebilobnta, which has pale green
sepals, petals, and coronal filaments (Table 1).

The habitats of Passiflora xishuangbannaensis
and P. altebilobata appear to differ: P. xi-
shuangbannaensis is found in open, wet, sunny or
shaded locations at 1200 m. whereas P. altebilobata
is found at lower elevations, ca. 900 m, under
dense forest in shaded areas. Passiflora altebilobata
is also found most frequently northwest of Xi-
shuangbanna, throughout Simao and Lin Gang
Counties, whereas P. xishuangbannaensis is only
found within Xishuangbanna.

Paratype. U.S.A. Ohio: Columbus, Ohio Stale Uni¬
versity Plant Biology Greenhouse, in cultivation, krosnick
401 (OS).

Acknowledgments. Thanks to Hong Wang of the

Xishuangbanna Tropical Botanical Garden for ini¬
tially locating this new species and for subsequent
assistance and collaboration during fieldwork to
collect P xishuangbannaensis, John Freudenstein
for the Latin translation, John MacDougal, Peter
Jprgensen, and Mesfin Tadesse for critical com¬
ments on the manuscript, and Joan Leonard for
maintenance of the living collections. This research
was funded by the Ohio State University Alumni
Grants for Graduate Research and Scholarship, and
the American Society of Plant Taxonomists Grad¬
uate Research Award.

Literature Cited

W ilde, W. J. J. 0. de. 1972. The indigenous Old World
Passilloras. Blumea 20: 227—250.

Isoetes orientalis (Isoetaceae), a New Hexaploid Quillwort

from China

Liu Hong and Wang Qing-Feng*

Herbarium, College of Life Sciences, Wuhan University, Wuhan 430072, People’s Republic
of China, supercjt@sohu.com; waugqf97@263.net. * Author for correspondence

W. Car l Taylor

Department of Botany, Milwaukee Public Museum, BOO W Wells Street, Milwaukee,
Wisconsin 53233-1478, U.S.A. ct@mpm.edu

Abstract. Isoetes orientalis is a previously un-
described, rare, and endangered quillwort of south¬
east China with a hexaploid chromosome count of
2/i = 66. Megaspores of /. orientalis have a retic¬
ulate ornamentation consisting of irregularly anas¬
tomosing muri witfi ragged crests. Microspores have
an eehinate-tuberculate ornamentation. Herbarium
specimens of/, orientalis previously have been mis-
identified as /. sinensis. Isoetes orientalis differs
from /. sinensis in spore morphology and chromo¬
some number.

Key words: China, Last Asia, Isoetaceae, Isoe¬
tes, quillwort.

Three basic diploid species and one tetraploid
species of Isoetes have been reported from China
(Liu el ah, 2002; Wang et ah. 2002). These arc /.
hypsophila Handel-Mazzetti (2// = 22), /. taiwa-
nensis DeVol (2 n = 22), /. yunguiensis Wang O. F.
& W. C. Taylor (2// = 22), and /. sinensis Palmer
(2n = 44). Over the last five years, field, herbarium,
and laboratory studies have been conducted in the
assessment of Isoetes populations over a large part
of mainland China. As part of these studies, ex¬
amination of herbarium specimens of Isoetes col¬
lected from Songyang County in southeast China
revealed specimens that had been previously iden¬
tified as /. sinensis. Closer inspection of these spec¬
imens showed that they differed from /. sinensis in
spore ornamentation. Further analysis of these her¬
barium specimens and living specimens from Son¬
gyang County indicate that these collections are
distinct from /. sinensis in spore morphology and
chromosome number. They represent a unique, pre¬
viously undescribed species of Isoetes, which is
named and described as follows:

Isoetes orientalis II. I, in & Q. F. Wang, sp. nov.
TYPE: China. Zhejiang: Songyang County, ca.
0.15 km S of Anming Town, 12 Dec. 2002, Liu
II. & Wang J. Y. WH2002I214 (holotype,
WH). Figures 1, 2.

Planta aquatica ex eaudice trilobo emergens. Folia 20
ad 40, viridia, usque ad 10-20 cm longa. ea. 2.0 mm lata
ad medium longitudinis, hasi dilatata et complanata: fibris
fihrolignosis praesentibus; ligula subtriangulata, 1 .5— 2.0
mm longa, 2.0— 3.0 mm lata. Velum rudimentarium. Spo¬
rangium basale, 5-6 mm fongum; megasporae superfieie
retic ulata ex ponds irregulariter anastomosantibus eristis
laeeratis coronatis constante, 350—450 yam diametro; mi¬
crosporia pallide earns in massam dispositis, elliptieis,
19—29 yam longis, eehinatis. Chromosomatum numerus 2 n
= 66 (Fig. 1 B).

Plant aquatic, emergent. Root stock 3-lobed.
Leaves white basal ly, green above, spirally ar¬
ranged. w ide spreading, 10-20 cm long, ca. 2.0 mm
wide at mid-length, in lulls of 20 to 40, flattened
on adaxial side, rounded on abaxial side, base alate
(Fig. 1 A); peripheral fiber strands present; central
intrastelar canal one; ligule ovate-subtriangular,
1.. 5-2.0 X 2.0— 3.0 mm. Sporangium basal, obovate,
ca. 5—6 X 3.8— 4.5 mm; velum rudimentary, cov¬
ering only distal edge of sporangium. Megaspores
white when dry, gray when wet, ca. 350—450 yam
diam. (mean = 420 yam), with reticulate texture
composed of irregularly anastomosing muri with
ragged crests, girdle obscure (Fig. 2A-D). Micro¬
spores gray in mass, elliptic, ea. 19—29 yum long
(mean = 22 yum), tuberculate-echinate (Fig. 2F-I).
Chromosomes: 2 n = 66.

Distribution. In China, Songyang County of
Zhejiang Province.

Ecology. 1'he marshes containing Isoetes orien¬
talis are in a unique, swampy, loamy meadow be¬
tween two mountains. The* water in these marshes
has a pH of 5.8. The reddish soil in which the
plants are rooted has a pH of 5.4. Common asso¬
ciates include Eriocaulon alpestre Hooker f. &
Thomson, Juntas ejfucus l„, Cyperus haspan L.,
Murdannia triguetra (Wallich) Bruckner, and Oen-
anthe javanica (Illume) DC.

Isoetes orientalis and /. sinensis both occur in
Zhejiang Province, but they are separated by ele-

Novoin 15: 164^167. 2005.

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2005

Liu et al.

Isoetes orientalis from China

165

Figure I. Isoetes orientalis II. I. in & O. F. Viang. — A. Holotvpe: Liu H. & Wang ./. Y. WH20021214 (W II). — 15.
Somatic chromosomes in mitotic root tip squash. — C. Roots with reddish rhizosphere. Scale bar: A = 2 cm; 15 = 10
gm; C = 200 gin. Figures 15 and C from isotype: Liu II. & Wang J. Y. WH20020215 ( W II).

vation. The /. orientalis populations occur at 1200
m, whereas /. sinensis grows between 130 and
300 m.

Leaf features. In cross section, the leaves of Is¬
oetes orientalis are oval and almost completely oc¬
cupied by four, thin-walled, oblong air chambers
with the two adaxial air chambers being slightly
larger than the abaxial ones. In contrast, the air
chambers in the leaves of /. sinensis appear to be
smaller, more circular, and have thicker walls. F ur¬
thermore, Isoetes orientalis has one intrastelar ca¬
nal. whereas /. sinensis has three or more intrastelar
canals (Takamiya et al.. 1997).

Root features. A noteworthy feature of I. orien¬
talis is the reddish, sheathing mantle formed
around its dichotomously branched roots (Fig. 1C).
The reddish color of this mantle or rhizosphere is
reportedly doe to oxidation of iron in the soil ef¬
fected by oxygen released from the roots (Tessenow
& Baynes, 1975, 1978). Certain kinds of bacteria
and fungi grow in the rhizosphere ol Isoetes (Shar-
ma, 1998). This feature has not been observed on
the roots of /. sinensis, and it has not been previ¬
ously reported for any species of Isoetes in Asia.

Spore size and texture. Spore measurements

and photomicrographs were made using a Hitachi
S-800 scanning electron microscope at 15 Kv.
Megaspores of I. orientalis are ea. 350—450 gm
diam. (mean = 420 /xm diam., N = 25), whereas,
according to Huang et al. (1992), megaspores of/.
sinensis are ca. 360—390 /xm diam. Microspores of
/. orientalis are 19—29 /xm long (mean = 22 /xm
long, N = 40), whereas, according to Huang et al.
(1992), the microspores of/, sinensis arc 19—20 pan
long and average 20 /xm long.

Megaspores of /. orientalis have an irregular cris¬
tate-reticulate ornamentation (Fig. 2A— I)), whereas
those of /. sinensis are cristate with fewer anasto¬
mosing ridges (Watanabe et al.. 1996: 290, figs. 39—
43). Megaspore ornamentation of /. orientalis re¬
sembles that of the octaploid /. pseudojaponica M.
Takamiya, Mitsu. Watanabe & K. Ono (Watanabe
et al.. 1996: 288, figs. 15—18), but they differ with
respect to microornamentation. At higher magnifi¬
cation, the fibrils on the megaspores of /. orientalis
(F ig. 2E) are longer and larger than the fibrils on
the megaspores ol /. pseudojaponica (Watanabe et
al.. 1996: 288. fig. 19). Megaspore texture of /. or¬
ientalis is also similar to that of /. japonica A.
Braun, but Isoetes orientalis differs from I. japonica

166

Novon

Figure 2. Isoetes orient alis. — A. Megaspores. — B. Proximal view of megaspore. — C. Distal view of megaspore. —
I). K. Megaspore surface microornamentation. — F. Microspores. — G. Distal view of microspore. — II. Proximal view
of microspore. — I. Microspore surface microornamentation. Scale bar: A = 200 /laid; B & C = 75 gm: I) & F = 15
/Ain; E & I = 2 /Am; G & H = 4 /tin. All from isotype: Liu //. <£ Wang J. Y. WH2002U215 (\\ II).

in microspore texture. Microspores of /. orientalis
have an echinate-tuberculate ornamentation (Fig.
2F-II), whereas /. japonica microspores are nearly
laevigate (Watanabe et al.. 1996: 284. figs. 8—13).
At higher magnification, the short spines of /. or¬
ientalis look more tuberculate and a granulate mi-
croornamentation is apparent (Fig. 21). In contrast,
the microspores of /. sinensis are clearly echinate
(Huang et ah, 1992: 147, fig. 3A— C; Watanabe et
ah. 1996: 290. figs. 46-50).

Although Isoetes orientalis and /. sinensis differ
in spore ornamentation, the plants are similar in
general appearance and growth habit. Both species
are much alike in leaf, sporangium, ligule, and ve¬

lum characters, as well. Early descriptions of Is¬
oetes species were based on general appearance
and habit, and this might be the reason /. orientalis
was originally identified as /. sinensis.

Cytology. Chromosome counts were made from
root tips of plants collected al the type locality and
prepared following the methods of Liu et ah (2002).
Counts from a total of 80 cells found in root tip
squashes of 12 individuals showed that /. orientalis
is a hexaploid w ith a chromosome number of '2n =
66 (F ig. I B). The base number for Isoetes is x =

1 1 (Love et ah, 1977).

Takamiya et ah (1994) recognized 2 n — 44. 65,
66, and 68 cytotypes for /. sinensis. Takamiya et ah

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Liu et al.

Isoetes orientalis from China

167

(1997) revised lliis earlier taxonomy recognizing the
tetraploid cytotype as /. sinensis var. sinensis and
the 2 n = 65, 66, and 68 cytotypes as /. sinensis
var. coreana (Y. H. Chung & H. K. Choi) M. Tak-
amiya. Mitsu. Wantanabe & K. Ono. According to
Takamiya et al. (1997), both of these varieties have
cristate megaspores and echinate microspores.
Their megaspore texture is distinct from /. orien¬
talis. , which has reticulate megaspores and eehi-
nate-tuberculate microspores.

Phenology. Megaspores of Isoetes orientalis ma¬
ture from mid May to early September. Microspores
develop from late June to mid October. By Decem¬
ber, most sporophylls have decayed and only a few
depauperate, green leaves remain, but like other
emergent or amphibious quillworts in China, /. or¬
ientalis stays green, at least to some extent, through
the winter.

I, ike all other quillworts in China, /. orientalis is
a rare and endangered species. To date, it has been
found at only two sites, one of which has been dec¬
imated. The last known extant /. orientalis popu¬
lation covers no more than 200 square meters and
contains fewer than 100 individuals. Pollution and
increasing urbanization continue to raise the risk
of extinction for this species. Protection of the one
remaining population of /. orientalis is crucial, and
searches for additional populations are urgently
needed.

Paratype. CHINA. Zhejiang: Songyang County, An-
min Town. Andaihou, lx>u I.. II. 227 (SY. XJh'C).

Acknowledgments. We are grateful to Wang Jin-
Yuan and Liu Xing for assistance in conducting
field studies. We are particularly indebted to Wang
Yong, Chen Jing-Ming. Dong Yuan-Huo, and Wu
Cui, members of the pteridophyte research group
of the laboratory of Plant Systematic^ and Evolu¬

tionary Biology at Wuhan University, for help and
suggestions in preparing this manuscript. We thank
Roy Gereau for assistance with the Latin diagnosis
for Isoetes orientalis. This study was funded by the
State Key Basic Research and Development Plan
of China (G2000046805).

Literature Cited

Huang, T. C., II. J. Chen & I.. C. Li. 1992. A palynolog-
ical study of Isoetes taiwanensis DeVol. Airier. Fern J.
82: 142-150.

Liu Xing. Y. Wang, O. F. Wang X Y. 11. Guo. 2002. Chro¬
mosome numbers of the Chinese Isoetes and their tax-
onomical significance. Acta Phytotax. Sin. 40(4): 351 —
356. | In Chinese with English abstract.]

Love, A., I). Love & R. E. G. Pichi Sermolli. 1977. Cy-
totaxonomical Atlas of the Pteridophyta. .1. Cramer, Va¬
duz.

Sharma, B. I). 1998. Eungal associations in the root of
three species of Isoetes L. Aquatic Hot. 61: 33—37.
Takamiva, \1., \1. Watanabe & K. Ono. 1994. Biosyste-
matie studies on the genus Isoetes (Isoetaceae) in Japan.
1. Variations of the somatic chromosome number. .1. PI.
Res. 107: 289-297.

- , - & - . 1997. Biosystematic studies

on the genus Isoetes (Isoetaceae) in Japan. IV. Mor¬
phology and anatomy of sporophytes, phytogeographv
and taxonomy. Acta Phytotax. Geohot. 48: 89— 122.
Tessenow, U. & Y. Baynes. 1975. Redox-dependent ac¬
cumulation of Fc and Yin in a littoral sediment sup¬
porting Isoetes lacustris L. Naturw issenschaften 62:
342-343.

- & - . 1978. Experimental effects of Isoetes

lacustris L. on the distribution of Eh, pi I, Fe and Mil
in lake sediment. Verh. Int. Vereinigung Tlieor. Limnol.
20: 2358-2362.

Wang, (J. E, Liu Xing. W. C. Taylor & He Zhao-Rong.
2002. Isoetes yunguiensis (Isoetaceae), a new basic dip¬
loid quillwort from China. Novon 12: 587—591.
Watanabe, M., M. Takamiya, T. Matsusaka X K. Ono.
1996. Biosystematic studies on the genus Isoetes (Is¬
oetaceae) in Japan. III. Variability within qualitative
and quantitative morphology of spores. J. PI. Res. 109:
281-296.

A New Species of Freesia (Iridaceae: Crocoideae) from the
Succulent Karoo, South Africa, and Notes on Nomenclature and

Infrageneric Classification

John C. Manning

National Botanical Institute, P. Bag X7, Claremont 7735, South Africa. Manning@sanbi.org

Peter Goldblatt

B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299,
St. Louis, Missouri 63166-0299, U.S.A. peter.goldblatt@mobot.org

Abstract. Freesia marginata (Iridaceae: Crocoi¬
deae) is a new species from the Breede River valley
near Worcester in Western Cape Province, South
Africa. A local endemic of succulent karoo shrub-
land, it resembles F. caryophyllacea in its flowers
and short, prostrate leaves, but it is distinguished
from all other species of Freesia by the leathery,
somewhat glaucous leaves with a prominent sub¬
marginal vein. The four species previously placed
in the genus Anomatheca arc recognized in Freesia
as the new subgenus Anomatheca. Three of the spe¬
cies, F. grandifiora, F laxa, and F. verrucosa, are
segregated into section Anomatheca, and F. viridis
is placed in the new section Alatae.

Key words: Crocoideae, Freesia, Iridaceae,
South Africa, succulent karoo.

Freesia Klatt is a small tropical and southern
African genus with a marked center of diversity in
the southern part of Western Cape Province, South
Africa, an area of winter rainfall and summer
drought (Goldblatt, 1982; Manning el al., 2002).
The genus is well defined by its conical conns cov¬
ered by netted tunics, strongly inclined to horizon¬
tal spikes with the flowers borne on the upper side,
style branches divided for hall their length, rugose
or papillate capsules, and seeds in which the raphe
is more or less inflated and the surface smooth or
rugose but without secondary sculpturing (Goldblatt
& Manning, 1995). In its earlier circumscription,
Freesia comprised only those species with highly
fragrant flowers in which the funnel-shaped peri¬
anth is divided into a slender basal portion that
widens abruptly into a longer, more or less cylin¬
drical upper portion (Goldblatt, 1982). The genus
was expanded by (Goldblatt and Manning (1995) to
include the four species of the genus Anomatheca
Ker Gawler. Freesia currently comprises 16 spe¬
cies, including the recently described Freesia fu-

cata J. C. Manning & Goldblatt from the Breede
River valley between Worcester and Villiersdorp in
Western Cape Province (Manning & Goldblatt,
2001).

The discovery of Freesia fucata from such a well-
populated area is remarkable, but is matched by
the discovery of yet another species from the same
valley, here described as Freesia marginata. The
discovery of this species provides further evidence
that the drainage basin ol the Breede River, which
forms the region known as the Worcester Karoo, is
a distinct enclave of endemism for Iridaceae. Spe¬
cies of Iridaceae in the southern African winter-
rainfall zone display a high degree of substrate fi¬
delity (Goldblatt & Manning. 1996), and edaphie
and topographic diversity appear to have played a
significant role in speeiation in the flora (Goldblatt
X Manning. 1998, 2000, 2002). The high number
of locally endemic species of Iridaceae in the
Worcester Karoo can probably be ascribed to local
isolation combined with the mosaic of soil and as¬
sociated vegetation types that occur in the basin
(Low & Rebelo, 1996). These range from fynbos on
the sandstones of the fringing mountains, through
a transitional belt of renosterveld on better watered
clay soils to succulent karoo on the drier soils in
valley bottoms and on sunbaked north-facing hill
slopes.

Freesia marginata .1. C. Manning X Goldblatt, sp.
nov. TYPE: South Africa. Western Cape Prov¬
ince: Worcester District, stony washes between
Worcester and Roberston, 25 May 2002, ./. C.
Manning 2734 (holotype, NBG; isotypes, MO.
PRE). Figure I.

Plantae 4-10 cm altae, cormo conico 15—20 mm diam.
tunicis pallide reticulatis, foliis 4—7 plemmque prostratis
lanceolatis 20—60 X 6—15 mm submarginal is prominen-
tibus, caule ram uniramoso dense et minute puberulo, spi-
ca inclinata 4— 10-florum, fforihus 30—37 mm longis fra-

Novon 15: 168—172. 2005.

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2005

Manning & Goldblatt
Freesia from South Africa

169

f igure I. Freesia marginata J. C. Manning & Goldblatt. — A. Whole plant. — 15. Flowering spike. — C. Flower in
front view. — I). Half flower. — K. Inner bract (left) and two different outer bracts to show variation (center and right).
— F. Capsules. — G. Seed. Drawn from ./. C. Manning 2734 (holotype, NI5G).

grantibus albis plerumque externe purpureis suffusis,
tepalis inferioribus aurantiacis notatis. lubo perianthii 1 fi —
25 mm longo infime filiforme 6—10 mm longo, tepalis in-
aequalibus, dorsale grandiore suberecto obovato 12—15 X
7—8 mm, inferioribus recurvatis, filamentis 15—17 mm lon-

gis, antheris 5—5.5 mm longis, stylo diviso prope apieem
antherarum.

Plants small, 4—10 cm high, usually ± prostrate;
conn conic, 15—20 mm diam., the tunics of pale,

170

Novon

medium-textured, reticulate libers, accumulating
with age and forming a long neck around the base
of tin* stem: cataphylls membranous, flushed deep
purple above ground level; stem flexed outward
above the uppermost leaf sheath and then prostrate
or horizontal, densely puberulous, simple or rarely
with one branch. Foliage leaves (4)5 or 6(7). pros¬
trate or rarely erect, oblong to lanceolate, acute,
leathery, ± glaucous, with relatively inconspicuous
pseudomidrib but distinct submarginal vein. 20—
50(60) X 6) — 1 5 mm, slightly shorter than to half as
long as the stem. Spike horizontal, 4- to 10-flow-
ered; bracts solt, herbaceous, green with hyaline
margins, the outer all tricuspidate or the upper ob-
ovate, with a prominent central cusp tipped with
rusty brown, 5—7 mm long, the inner bracts slightly
longer, 7—8 mm long, forked for 2—3 mm, cusps
tipped rusty brown. Flowers (30— )33— 37 mm long,
strongly violet-scented, white, usually lightly
flushed purple on the reverse of the tepals, the base
of the lower tepals yellow-orange, usually with a
dark midline; perianth tube 18—25 mm long with a
basal narrow part 6—10 mm long, widening ±
abruptly into a wider, flared upper portion; tepals
unequal, the dorsal largest, erect, obovate, 12—15
X 7—8 mm, the upper laterals slightly narrower,
oblong-lanceolate, the lower tepals recurved, the
lower laterals oblong-lanceolate, 11—13 X 5.0— 5.5
nun, the lower median oblong, 1 1—12 X 4—5 mm;
filaments arcuate, inserted at the base of the wid¬
ened upper portion of the perianth tube, included,
15—17 mm long; anthers 5.0— 5.5 mm long; ovary
ovoid, 3 mm long; style dividing below or opposite
the anther apices, the branches deeply forked. Cap¬
sules oblong or subglobose, three-lobed, minutely
papillate when fresh, (5— )6— 10 X 5—10 mm; seeds
globose with lightly to moderately inflated chalaza
and raphe, rugose, glossy reddish or purplish
brown, 2.0— 3.0 mm diam.

Distribution. Freesia marginata is currently
known from a single population between Worcester
and Roberston in the valley of the Breede River in
Western Cape Province, South Africa. An uncon¬
firmed sighting (Snijman, pers. comm.) suggests,
however, that further populations occur in the vi¬
cinity. Plants from the type population are restrict¬
ed to washes at the base of a prominent sandstone
knoll where they grow in decaying sandstone with
the conns deeply buried in sandy loam in rock
crevices. The vegetation at the site is open succu¬
lent karoo, dominated by Ruschia caroli (L. Bolus)
Schwantes (Aizoaceae) and Pteronia paniculata
Thunberg (Asteraceae), with scattered bushes of
Wiborgia mueronata (L. f.) Druce (Fabaceae).

Plants of Freesia marginata are typically low-grow¬
ing, with prostrate leaves and more or less prostrate
inflorescences, although individuals growing in the
shade of small bushes develop suberect leaves and
an inflorescence that is borne horizontally above
the ground. The stem usually remains prostrate in
fruit and the capsules are borne on the ground and
often lightly covered by soil.

Phenology. Plants flower in the early winter,
from late May to early June.

Etymology. From the Latin marginata , ‘'mar¬
gined,” referring to the prominent submarginal vein
of the loliage leaves.

The puberulous stem, green bracts, and narrowly
ovate lower tepals of Freesia marginata are char¬
acteristic of section Viridibractea (Goldblatt, 1982)
and indicate its placement in this section. Within
the section it closely resembles F. caryophyllacea
(Burman f.) N. F. Brown in its flowers, prostrate
leaves, and early flowering, but is quite distinct
from that species in the leathery texture ol the
leaves, their more or less glaucous coloring, incon¬
spicuous pseudomidrib, and especially in the prom¬
inent submarginal vein. This is clearly visible in
fresh and dried leaves as a translucent line running
parallel to but ca. 0.5 nun in from the leaf margin.
Anatomically, the leaves of F. marginata are char¬
acterized by ca. 20 layers of mesophyll, and vas¬
cular bundles that are not differentiated into pri¬
mary anil secondary bundles and in which
sclerenehyma phloem caps are rarely developed. A
marked exception are the submarginal bundles,
which are situated very near the marginal vein, and
in which the bundle sheaths are massively devel¬
oped to form a complete sheath extending to the
epidermis (Fig. 2B). The leal of Freesia caryophyl¬
lacea , in contrast, has ca. 10 layers of mesophyll
and a prominent pseudomidrib with the bundles on
either side clearly differentiated into primary and
secondary bundles, each primary bundle with a
well-developed sclerenehyma phloem cap. The
submarginal veins are some distance from the mar¬
ginal vein and have normally developed phloem
caps that do not extend as complete sheaths to the
epidermis (Fig. 2A). The two species also differ in
their ecology: Freesia caryophyllacea is a species
of renosterbos shrubland on stony clay soils. It is
relatively widespread in the Breede River valley
anil in the Overberg from Caledon to Swellendam.
Populations of F. caryophyllacea in the Breede Riv¬
er between Worcester and Roberston appear to
flower in mid to late June, thus later than F. mar¬
ginata. Freesia marginata, in contrast, occurs on
decaying sandstone in succulent karoo and flowers

Volume 15, Number 1
2005

Manning & Goldblatt
Freesia from South Africa

171

sb

mb

Figure 2. Comparison of the anatomy of the leaf margin of Freesia caryophyllacea (A) and V. marginata (B). sl>,
submarginal bundle, mb, marginal bundle. (A) from plants cultivated at Kirstenbosch Gardens, originally collected at
Bredasdorp; (B) from plants from the type population near Worcester.

some weeks earlier than this, in late May or early
June.

The submarginal vein in the leaves of Freesia
marginata is unique in the genus but occurs in
certain species of Tritonia Ker Gawler, including T.
disticha (Klatt) Baker, T. drakensbergensis VI. I de
Vos, and T. lineata (Salisbury) Ker Gawler (De Vos,
1999). The curious, tricuspidate bracts that occur
in F. marginata are found elsewhere in the genus
in F fucata, another recently described local en¬
demic species of the same valley system (Manning
& Goldblatt, 2001 ).

Freesia marginata was first collected in 2000 by
Deon Kotze, horticulturist at the Karoo Botanic
Garden, who showed us living plants in the collec¬
tion in June 2001. Although past flowering, the dis¬
tinctive leaves made if apparent that they repre¬

sented an undescribed species, and flowering
material was collected the following year.

Infragenekic Classification of Freesia

The last revision of the genus Freesia (Goldblatt,
1982) recognized the two sections Freesia sect. Vir-
idibractea Goldblatt and section Freesia, encom¬
passing 11 species. With the transfer into Freesia
of the species previously placed in the genus An-
omatheca Ker Gawler (Goldblatt & Manning,
1995), however, it is now necessary to emend the
classification of the genus to accommodate them.
Until further knowledge is forthcoming concerning
species’ relationships in the genus, it seems best to
recognize the four species previously placed in An-
omatheca as a separate subgenus, circumscribed by

172

Novon

their hypoerateriform flowers with the stamens in-
serted near the month of the corolla tube, which is
slender and cylindrical throughout or widened only
at the mouth. Freesia subg. Freesia is recognized
by the distinctly funnel-shaped flowers in which the
stamens are inserted near the middle or in the low¬
er hall ol the perianth tube, separating the narrow
lower portion from the abruptly widened upper por¬
tion. Within Freesia subg. Anomatheca, F. viridis
(Aiton) Goldblatt & J. C. Manning is immediately
distinguished by its ridged or winged inflorescence
axis and unmarked, green or brownish green flow¬
ers, and is appropriately placed in a separate sec¬
tion, Freesia sect. Alatae. The remaining three spe¬
cies of Freesia sect. Anomatheca ( Freesia
grandiflora (Baker) Klatt. F. verrucosa (Vogel) (iold¬
blatl J. C. Manning, and F. laxa (Thunberg)
(ioldblatl J. (i. Manning) have pink or red (rarely
white to bluish) flowers marked with darker nectar
guides on the lower tepals, and terete flowering
stems.

We therefore suggest the following infrageneric
classification for Freesia :

Freesia Klatt, l.innaea 34: 672. 1866. TY PE: Free¬
sia refracta (Jacquin) Klatt.

I. Freesia subg. Anomatheca (Ker Gawler) J. (i.
Manning & Goldblatt, stat. nov. Basionym: An¬
omatheca Ker (iawler, Koenig & Sims Ann.
Bot. I: 227. 1804. Anomaza Lawson ex Salis¬
bury, Trans. Hurt. Soc. London I: 323. 1812.
Lapeirousia subg. Anomatheca (Ker Gawler)
Baker, Handb. bid.: 173. 1892. TYPE: Anom¬
atheca verrucosa (Vogel) Goldblatt (Ixia verru¬
cosa Vogel).

Freesia [subg. Anomatheca] sect. Anomathe-
ca (K< ■r Gawler) J. ( . . Manning d Goldblatt.
comb, et stat. nov. Type as for subgenus.

1. Freesia grandiflora (Baker) Klatt

2. Freesia verrucosa (Vogel) Goldblatt «Y .1. C.
Manning

3. Freesia laxa (Thunberg) Goldblatt & ,1. (7
Manning

Freesia [subg. Anomatheca] sect. Alatae J. (7
Manning & Goldblatt, sect. nov. TV PE: Freesia
viridis (Aiton) Goldblatt & J. (7 Manning.

Plantae foliis ereetis plan is vel crispis, caule complan-
ato striato vel alalo, hraeteis viridis, floribus hypocrateri-
formibus tube cylindrico ad apicem eurvato, viridis vel
brunneis.

Foliage leaves plane or crisped; stem longitudi¬
nally ridged or winged; bracts green; flowers hy¬

poerateriform with tubular perianth tube sharply
curved at the apex, greenish or brownish.

4. Freesia viridis (Aiton) Goldblatt & .1. G. Man¬
ning

11. Freesia subg. Freesia. TY PE: Freesia refracta
(Jacquin) Klatt.

Freesia [subg. Freesia] sect. Viridibraetea
Goldblatt, J. S. African Bot. 48: 56. 1982.
TY PE: Freesia (dim (G. L. Meyer) Gumbleton.

5. Freesia alba (G. L. Meyer) Gumbleton

6. Freesia caryophyllacea (Burrnan I.) N. E.
Brown

7. Freesia fergusoniae L. Bolus

8. Freesia fucata J. (7 Manning tY Goldblatt

9. Freesia leichtlinii Klatt

10. Freesia marginal a J. (7 Manning & Goldblatt
I I. Freesia sparmannii (Thunberg) N. E. Brown
Freesia |subg. Freesia | sect. Freesia

12. Freesia andersoniae L. Bolus

13. Freesia corymbosa (Burrnan I.) N. E. Brown

14. Freesia occidentalis L. Bolus

15. Freesia refracta (Jacquin) Klatt

16. Freesia speciosa L. Bolus

Acknowledgments. We thank the l.ourens fam¬
ily for permission to collect on their farm, Cornelia
Klak for help with the identification ol the Aizo-
aceae, and Roy Gereau for his infallible assistance
in matters of nomenclature. Material was collected
with a permit from Western Cape Nature Conser¬
vation.

Literature Cited

De Vos. M. P. 1999. Tritonia. In: 0. A. I.eistner (editor).
Flora of Southern Africa 7(2): 89—128. Botanical Re¬
search Institute, Pretoria.

Goldblatt, P. 1982. Systematies of Freesia Klatt (Irida-
ceae). J. S. African Bot. 48: 39-91.

- & J. C. Manning. 1995. Phylogeny of the African

genera Anomatheca and Freesia (Iridaceae: Ixioideae),
and a newr genus Xenoscapa. Syst. Bot. 20: 161—178.
- & - . 1996. Two new edaphic endemic spe¬
cies and taxonomic changes in Gladiolus (Iridaceae) of
southern Africa, and notes on Iridaceae restricted to
unusual substrates. Novon 6: 172—180.

- & - . 1998. Gladiolus in Southern Africa.

Fernwood Press, Cape Town.

- <Y - . 2000. Cape Plants: A conspectus of

the Cape flora of South Africa. Strelitzia 9: 1—743.
- tY - . 2002. Plant diversity of the Cape re¬
gion of southern Africa. Ann. Missouri Bot. Card. 89:
281-302.

Low. A. B. tY A. (7 Rebelo. 1996. Vegetation of South
Africa, Lesotho and Swaziland. Department ol Environ¬
mental Affairs & Tourism, Pretoria.

Manning, J. & P. Goldblatt. 2001. Two new renosterveld
species of Iridaceae: Crocoideae from South Africa.
Bothalia 31: 189-192.

- , - tY I). Snijman. 2002. Cape Bulbs and

their Allies. Timber Press, Oregon.

Aechmea sucreana, a Mew Species of Bromeliaceae from
Espfrito Santo State, Brazil

Gustavo Martinelli and Claudia Magalhaes Vieira

Instituto <le Pesquisas Jardim Bolanico do Rio de Janeiro, Rua Pacheco Leao 915,
Horto Florestal, Rio de Janeiro, RJ, Brasil, CEP 22460-030. gmartine@jbrj. gov.hr;

cnivieira@netrio.com.br

Ahstract. A new species of Bromeliaceae, Aech¬
mea sucreana, found in Espfrito Santo State, Brazil,
is described and illustrated. It is distinguished from
its closest relative, A. blanchetiana, on the basis of
inflorescence features.

Key words: Aechmea, Brazil, Bromeliaceae.

During development of the Biology and Conser¬
vation of Bromeliaceae of the Atlantic Rainforest
Project, a scientific program of the Rio de Janeiro
Botanic Garden Research Institute, I I scientific
expeditions were made between 1998 and 1999.
The project encompassed the largest remaining
area of rain forest, and major fragments. A total of
92.394 km of roads and trails were accessed in 78
conservation units and 59 private areas through 14
Brazilian states. Seven hundred twenty-nine taxa
were obtained and observed, some of them consid¬
ered new taxa to this family, representing ea. 69%
of the total Bromeliaceae recounted in the literature
for the Atlantic Rainforest bionic (Martinelli, un¬
published data). Information on the biology and the
status of conservation of the Bromeliaceae species
and their respective populations were obtained and
stored in a database. In addition to voucher spec¬
imens deposited in RB, one live collection was se¬
cured for taxa considered endemic and/or threat¬
ened with extinction and placed into cultivation. A
total of 1842 specimens were collected and are be¬
ing cultivated in a specially built greenhouse in the
Rio de Janeiro Botanic Garden Research Institute.
The living collections are part of an ex-situ con¬
servation program, including the maintenance of a
germplasm bank and the propagation and later re-
introduction, when necessary, of the species into
their original habitats.

This paper presents a new species of Aechmea
collected by the project during the expedition in
Espfrito Santo State, Brazil.

Aechmea sucreana Martinelli & G. Vieira, sp.
nov. TYPE: Brazil. Espfrito Santo: Einhares,
Imbiriba, Eazenda Sao Jorge, proximo da Re-
serva Florestal de Einhares, Mata Atlantica,
Restinga aberta, 10 m s.m., I9°11'27"S,
39°52'83"W, 5 Aug. 1999 (fl), G. Martinelli
1555V & L. Heron, R. Azoury (holotype, RB
353539). Figure 1 .

Species nova Aechmeae blanchetianae affinis sed inflo-
rescentia breviore, rachidibus intemodiis minoribus, flo-
ribus plus dense dispositis, ae foliis minoribus differt.

Plant flowering to 1.2 m high. Eeaves ca. 25,
erect or suberect, forming a broadly funnelform ro¬
sette; sheaths elliptic, 22—30 X (9.5)14—16 cm, spi-
nose-serrate toward apex, pale brown, densely lep-
idote on both surfaces; blades yellowish green with
reddish nuances on both surfaces, otherwise totally
red. ensiform to narrowly triangular, 34—53 X (4)6—
I I cm, lepidote on both surfaces, mainly on abaxial
surface, spinose-serrate, spines dark brown, 1—2
mm long, apex acute, ending in a rigid, pungent
spine. Inflorescence erect; scape reddish green at
base, red toward apex, often exceeding the leaves,
41—57 X 0.5— 0.7 cm, sparsely white lanate; scape
bracts reddish pink, erect, strongly imbricate, en¬
folding the scape, oblong, entire, with acuminate
apex, 9—17 X 3—4.5 cm, ehartaceous, lepidote on
both surfaces; fertile part of inflorescence 2- to 3-
branched, many-flowered, with central rachis red,
(25)36—48 cm long, sparsely white-lanate to gla-
breseenl; primary bracts reddish pink, shorter than
the branch but greatly exceeding the basal sterile
portion, entire with acuminate apex, 8—15 X 1.5—
3.5 cm, shorter toward inflorescence apex, charta-
ceous, lepidote on both surfaces; primary branches
red-pink, 9 to 13, spreading-ascending, the lowest
(9)12—22 cm long; secondary bracts similar to pri¬
mary bracts, 3.5—5 X 0.9—2 cm; spikelets 3- to 10-
flowered, 4—12 cm long, with internodes 0.8—
1.3(1. 5) cm; floral bracts red-pink, slightly shorter
than to little exceeding the sepals (but often ex¬
ceeding floral buds), 2—3.5 times as long as I lie

Novon 15: 173-175. 2005.

174

Novon

(showing longitudinal callosities and petal appendages) and antepetalous stamen. — F. Stigma. — G. Leaf.

internodes, not covering the raehis, ovate, convex,
multi-nerved, (1.8)2. 4— 3.8 X 1-1.1 cm, entire, with
acuminate apex, sparsely lepidote on abaxial sur¬
face. Flowers polystichous, laxly to densely ar¬
ranged, sessile, ea. 4 cm long; floral buds green-
yellow; sepals green with apex rose and mucro red,
connate at base for ca. 3 mm, ovate or narrowly
ovate, strongly asymmetric, 1.6— 2.1 X 1 cm, mu-
cronate, mucro 1—2 mm long, sparsely white-lanate
to glabreseent on abaxial surface; petals pale yel¬
low. becoming pale brown after anthesis, free, lin-
ear-spatulate, apex suberect at anthesis, acute, 3.2—
3.4 X 0.6 cm, bearing two longitudinal callosities
parallel to staminal filaments, ca. 2.2 cm long; petal
appendages 2 per petal, basal, spatulate-undulate
with apex unevenly toothed, ca. 3 X 2—3 mm; sta¬
mens shorter than petals, included at anthesis, ca.
2.8 cm long; filaments 2.4— 2.5 cm long, the ante¬
petalous ones adnate to the petals for ca. 1.8 cm;
anthers linear-oblong, dorsifixed at the middle, 8—
0X4 mm, apex acute; ovary subtrigonous, 8—10
X 4—5 mm, pale green, sparsely white-lanate, pla¬

centae fixed in the upper half of the ovary, ovules
numerous, caudate; epigynous tube ca. 4 mm long;
style ca. 3.2 cm long; stigma conduplieate-spiral,
ovoid. Fruit ellipsoid, 10—12 X 7—8 mm, red.

I bis species was found growing in open thicket
vegetation (restinga), as terrestrial in white sand,
forming dense groups. Although the occurrence of
Aechmea blanchetiana (baker) L. 8. Smith in res¬
tinga of Linhares was registered by herbarium col¬
lections (KB), no overlap was noticed between these
two species. The local vegetation where A. sucreana
was found is endangered due to cattle activities de¬
veloped in the area.

Aechmea sucreana is similar to A. blanchetiana
in overall appearance, but can be easily distin¬
guished by its shorter inflorescence, spikelets with
shorter internodes, flowers more densely arranged,
and shorter leaves.

The specific epithet is a tribute to the Brazilian
botanist Dimitri Sucre, who made extensive plant
collections throughout the Brazilian state of Espfr-
ito Santo.

Volume 15, Number 1
2005

Martinelli & Vieira
Aechmea sucreana from Brazil

175

Paratypes. BRAZIL. Espirito Sanlo: I. inhares, Im-
biriba. Fazenda Sao Jorge, proximo da Reserva Florestal
de Linhares, Mata Atlantica, Restinga aberta, 10 m s.m.,
1 9° 1 1 '27"S, 39°52'83"W, 5 Aug. 1999 (fl), G. Martinelli
15559- A <£■ L. Heron , R. Azoury (MO), G. Martinelli
15559-B & /.. Hewn, R. Azoury (CEPEC), G. Martinelli
15559-G /.. Heron, R. Azoury (R), G. Martinelli 15559-

D & /.. Heron, R. Azoury (K), G. Martinelli 15559-E & /..
Heron, R. Azoury (SPF) (Inst. Bot.), G. Martinelli 15559-
F & /,. Heron, R. Azoury (CVRD).

Acknowledgments. The authors thank Patricia
Villela for the illustration of the new species.

A New Species of Burmeistera
(Campanulaceae: Lobelioideae) from Ecuador

Nathan Muchhala

Department of Biology, University of Miami, P.0. Box 2491 18, Miami, Florida 33124, IJ.S.A.

n.muehhala@umiami.edu

Thomas (>. hi turners

Department of Biology and Microbiology, University of Wisconsin Oshkosh. Oshkosh.
Wisconsin 54901, U.S.A. lammers@uwosh.edu

Abstract. Burmeistera auriculata is described
from a cloud forest remnant in Pichincha, Kcuador.
Although allied to B. borjensis, this new species is
distinguished from all known congeners by its au-
rieulate calyx.

Rksumkn. Se describe Burmeistera auriculata de
un remanente de bosque nublado en Pichincha, Ec-
uador. Aunque aliado a B. borjensis, esta especie
nueva se distingue de las demas Burmeisteras con-
oeidas por su ealiz aurieulado.

Key words: Burmeistera, Campanulaceae, Kc¬
uador, Lobelioideae, South America.

Burmeistera Triana (Campanulaceae: Lobelioi¬
deae) comprises 102 species ol robust plants (her¬
baceous or suffrutieose perennials, shrubs, and li¬
anas) distributed from Guatemala to Peru
(hammers, 1998, 2002). The genus is distinguished
by a combination of moderate-size (lowers (corollas
averaging 25—35 mm long) usually borne singly in
the axils of the upper leaves on ebracteolate pedi¬
cels; green or yellow corolla often suffused with ma¬
roon or purple; corolla tube neither fenestrate nor
cleft dorsally; dorsal corolla lobes larger than die
ventral lobes and falcate or reflexed; anther tube
wide open at the orifice; baccate, often inflated
fruit; and oblong to fusiform seeds much longer
than broad. With this circumscription, the genus is
most likely monophyletic (Luteyn, 1986; Stein,
1987a, 1987b, 1987c), a hypothesis supported by
preliminary molecular data (Pepper et al., 1997; K.
Knox, pers. comm.).

The last comprehensive account of Burmeistera
was the monograph by Wimmer (1943), who rec¬
ognized 77 species. More recent treatments are
available for the species of Costa Rica (Wilbur,
1975), Ecuador (Jeppesen, 1981), Guatemala
(Nash, 1976), Honduras (hammers & Maas, 1998),
Panama (Wilbur, 1977, 1981), and Peru (Stein.

1987c). Mere and in other publications (MeVaugh,
1943, 1949a. 1949b, 1957, 1965; Wimmer, 1955.
1968; Gomez Gdmez-Laurito, 1986; Lozano &
Galeano, 1986; Luteyn, 1986; Stein, 1987b; ham¬
mers. 1998. 2002), 44 new species were described,
while some taxa recognized by Wimmer (1943)
were reduced to synonymy.

Wimmer (1932, 1943. 1968) divided the genus
into two sections: Burmeistera (Imberbes, nom. in¬
valid), with all five anthers sparsely soft-pubescent
or glabrous at apex; and Barbatae L. Wimmer. with
the two ventral anthers barbate at apex. Though
Wimmer further divided section Burmeistera into
two subsections, Stein (1987b) transferred all mem¬
bers of subsection Aequilatae E. Wimmer to the
genus Siphocampylus Fold. This sectional classifi¬
cation is unsatisfactory, if only for the lack of struc¬
ture afforded so large a genus. It also is not sup¬
ported by preliminary molecular data (E. Knox,
pers. comm.).

The senior author is currently studying the pol¬
lination biology of the genus (Muchhala & Jarrin-
V., 2002). This research has revealed that, while
the closely related genera Centropogon C. Presl and
Siphocampylus are primarily hummingbird-polli¬
nated, Burmeistera has shifted to bat pollination
(Muchhala 2003, unpublished data). Although
hummingbirds visit Burmeistera flowers in the early
morning and at dusk (Feinsinger et al., 1987;
Muchhala, 2003), they rarely transport pollen and
only in small quantities. Meanwhile, a single visit
by a bat will deposit thousands of grains on a re¬
ceptive stigma.

In the course of this research, a new species was
discovered in the Otonga reserve in northern Ec¬
uador. This species, here described and named,
demonstrates a number of the floral adaptations to
bat pollination that are common to the genus, in¬
cluding nocturnal anthesis, dull coloration, rela-

Novon 15: 1 76-179. 2005.

Volume 15, Number 1
2005

Muchhala & Lammers
Burmeistera auriculata from Ecuador

177

tively wide corollas, a distinctive odor, and well-
exposed flowers.

Burmeistera auriculata Muchhala & Lammers,
sp. nov. TYPE: Ecuador. Pichincha: Bosque
Integral Otonga, 00°25.264’S, 79°00.779'\\.
cloud forest. 2233 in, 7 July 2002, A. Much¬
hala 120 (holotype, OLA; isotype, MO). Fig¬
ure 1 .

Ah omnibus caeteris speciebus Burmeisterae calycis
lol)is auriculatis differt; species sect. Burmeisterae at I mis.
B. borjensi, sed ah hac specie lamina lanceolata parviore
7.5—14 cm longa 2.8— 4.4 cm lata, pedieellis pubeseenti-
hus. hypanthio hreviore 5.5—6 mm longo, calycis lohis
brevioribus 9—1 I mm longis, antheris dorsalibns longior-
ihus 9.5—10.5 mm longis, et baccis globosis brevioribus
18—20 mm longis facile distinguenda.

Scandent hemi-epiphytic subshrub, 3 m tall;
stems 2.0— 3.8 mm diam., glabrous, occasionally
branching; latex white. Leaves strictly distichous,
patent; lamina lanceolate, 7.5—14 X 2. 5-4. 4 cm;
adaxial surface minutely scabrous, dull green; ali-

axial surface minutely and more densely pubescent,
dull light green; margin subentire; apex acuminate
to almost caudate; base rounded, occasionally
asymmetric; petiole 1.4— 2.0 cm long, 1.0— 1.5 mm
diam., minutely pubescent, 1/12—1/6 as long as the
lamina. Flowers solitary in upper leaf axils; pedi¬
cels 70—100 mm long, straight and ascending at
anthesis, 100—150 mm long, curved and declined
in fruit, 1—1.8 mm diam., minutely pubescent. 1 1 y-
panthium broadly obeonie, 5.5—6 mm long, 5-6.5
mm diam.. minutely pubescent; calyx lobes broadly
triangular. 9—1 1 X 7—9 mm, with reflexed auricles
4—7 mm long in the sinuses, ascending, overlapping
at base, finely reticulate, minutely scabrous; margin
sinuate; apex obtuse, almost emarginated; corolla
light green. 18—20 mm long, sparsely and minutely
pubescent or almost glabrous; tube slightly tie-
curved. 8—9 mm long, distended at base to 8.5— 9.5
mm diam., dorsiventrally compressed at middle,
3. 0-3. 5 mm tall. 4.0—4. 5 mm wide, abruptly ex¬
panding to 7.5— 8.0 mm at mouth; dorsal lobes lan¬
ceolate. strongly falcate, 23-25 X 5 mm, acuminate

178

Novon

at apex; ventral lobes narrowly triangular, falcate,
1 1-12 X 4 mm, acuminate at apex; staminal col¬
umn long-exserted; filament tube 26-28 mm long,
ca. 1.8 mm diam., sparsely puberulent; anther tube
curved-cylindrical, long-exserted, 5 mm diam., gla¬
brous; dorsal anthers 9.5—10.5 mm long; ventral an¬
thers 5.5— 6.5 mm long; pollen grains trieolpate,
prolate, 72 /ami diam. (equatorial) X 63 /ami diam.
(polar), the surface reticulate. Berries inflated, glo¬
bose, light green, 18—20 mm long, 17—21 mm
diam., crowned by the persistent calyx lobes; seeds
ellipsoid, 0.9 mm long, 0.3 mm diam.

Additional illustration. Muchhala and Jarrfn-V.
(2002), fig. 2A.

Distribution and ecology. Burmeistera auricu-
lata is endemic to the Andes of northwestern Ec¬
uador and known only from the type locality, where
it grows in cloud forest at elevations of 1990 to
2250 m. It flowers throughout the year, with indi¬
vidual flowers opening nocturnally between 1715
and 1800 hr. and senescing after 5 to 6 days. It is
pollinated by glossophagine bats of the genus An-
oura (Muchhala & Jarrfn-V., 2002. as "Burmeistera

sp/’).

Etymology. The name derives from the Latin
adjective auriculatus, ’’with little ears,” given in
reference to the unique calyx appendages.

Relationships. The most distinctive feature of
this species is the large reflexed auricle associated
with each sinus of the calyx. These appear to rep¬
resent basal lobules of adjacent calyx lobes that
have become connate. Though otherwise unknown
in Burmeistera, similar auricles are found else¬
where in the Campanulaceae, characterizing some
species of North American Ijobelia L. sect. Lobelia
(e.g., /.. siphilitica L., I.. appendiculata A. DC., /..
brevifolia Nuttall ex A. DC.; McVaugh, 1943), as
well as certain Eurasian genera of Campanuloideae
(e.g., Favratia Feer, Cryptocodon Fedorov, Mi-
chauxia I "Heritier).

In Jeppesen's (1981) treatment of Ecuadorian
Burmeistera, the new species keys readily to II. bor-
jensis Jeppesen, known from Napo and adjacent Co¬
lombia. The two species shared the following char¬
acters: (1) glabrous seandent stems; (2) abaxially
pubescent lamina with subentire margins; (3) the
length of petioles and pedicels; (4) broad overlap¬
ping reticulately veined calyx lobes with sinuate
margins; (5) the overall length and color ol the co¬
rolla; and (6) the inflated berries. Burmeistera bor-
jensis, however, differs in its larger (15—20 vs. 7.5—
14 cm long, 5—8 vs. 2.8— 4.4 cm wide) elliptic (vs.
lanceolate) lamina, cuneate (vs. rounded) at base;
glabrous (vs. pubescent) pedicels; longer hypanthi-

um (6—8 vs. 5.5-6 mm) and calyx lobes ( 14-16 vs.
9—1 I mm); longer corolla tube (15-18 vs. 8—9 mm);
shorter dorsal corolla lobes (ca. 16 vs. 23—25 mm);
shorter anthers (dorsal pair 8-9 vs. 9.5—10.5 mm);
larger (30—35 vs. 18—20 mm long) ovoid (vs. glo¬
bose) berry; and of course the exauriculate calyx.

In their combination of large inflated berries,
curved-cylindrical anther tube, and overall floral
morphology, Burmeistera auriculata and B. borjensis
would appear to belong to a group of species com¬
prising B. refracta E. Wimmer, II. ignimontis E.
Wimmer, B. cuyujensis Jeppesen, B. truncata Zahl-
bruckner, II. glabrata (Kunth) Bentham & Hooker
(. ex B. 1). Jackson, and II. oyacachensis Jeppesen.
Among these, only B. cuyujensis and B. ignimontis
share the abaxially pubescent lamina of II. auri¬
culata and B. borjensis, an unusual trail in the ge¬
nus. Both differ from the new species in their el¬
liptic or obovate (vs. lanceolate) lamina cuneate (vs.
rounded) at base, glabrous pedicels, longer hypan-
thium (8-12 vs. 5.5—6 mm) with narrowly triangular
or lanceolate (vs. broadly triangular) calyx lobes,
longer corolla (24—35 vs. 18-20 mm) with longer
tube (14—22 vs. 8—9 mm) and shorter dorsal lobes
(18-22 vs. 23—25 mm).

Paratypes. LCUADOR. Cotopaxi: Bosque Integral
Otonga, entre Quito y Sto. Domingo, eerca de San Fran¬
cisco de las Pampas, C. Nowicki & ./. Mutke 1213 (QCA);
Bosque Integral Otonga, N. Muchhala 24 (QCA).

Acknowledgments. The senior author thanks
Hugo Navarrete and the Pontificia Universidad Ca-
tolica de Ecuador for logistical help and access to
the QCA herbarium, and Giovanni Onore for access
to the study site (Bosque Integral Otonga). He also
thanks Paulina Moreno for help in identifying col¬
lections; it was she who first suggested that this
specimen from Otonga might represent an unde¬
scribed species.

Literature Cited

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K. (7 Murray. 1987. Disturbance, pollinator predict¬
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Gomez. F. I). & J. Gbmez-Laurito. 1986. Planlae mesoam-
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- . 2002. Seventeen new species of Lobelioideae

(Campanulaceae) from South America. Novon 12: 206—
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15: 53-56.

Luteyn, J. L. 1986. A new Burmeistera (Campanulaceae:
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McVaugh, R. 1943. Campanulaceae (Lobelioideae). N.
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- . 1949a. Studies in South American Lobelioideae

(Campanulaceae) with special reference to Colombian
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- . 1949b. Seven new species of Lobelioideae (Cam¬
panulaceae). J. Washington Acad. Sci. 39: 157—162.

- . 1957. Two new species of Lobeliaceae from Costa

Rica. Brittonia 9: 30—32.

- . 1965. South American Lobelioideae new to sci¬
ence. Ann. Missouri Rot. Card. 52: 399-409.
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- & I’. Jarri'n-V. 2002. Flower visitation by bats in

cloud forests of western Ecuador. Biotropica 34: 387—
395.

Nash, I). L. 1976. Flora of Guatemala, Campanulaceae.

Field Mus. Nat. Hist.. Rot. Sen 24: 396-431.

Pepper, A. S.-IL, M. H. G. Gustafsson & \. A. Albert.
1997. Molecular systematics of Neotropical Lobelioi¬
deae (Campanulaceae), with emphasis on Burmeistera,
Centropogon. and Siphocampylus, and the utility of fruit

and lloral characters in lobelioid classification. Amer.
J. Rot. 84 (6, Supph): 222 (Abstract).

Stein, R. A. 1987a. Systematics and Evolution of Centro¬
pogon subg. Centropogon (Campanulaceae: Lobelioi¬
deae). Ph.D. Dissertation, Washington University, St.
Louis.

- . 1987b. Siphocampylus oscitans (Campanulaceae:

Lobelioideae), a new name for Burmeistera weberbaueri
from Peru. Ann. Missouri Bot. Card. 74: 491—493.
- . 1987c. Synopsis of the genus Burmeistera (Cam¬
panulaceae: Lobelioideae) in Peru. Ann. Missouri Rot.
Card. 74: 494-496.

Wilbur, R. L. 1975. A synopsis of the Costa Rican species
of Burmeistera (Campanulaceae: Lobelioideae). Bull.
Torrey Bot. Club 102: 225-231.

- . 1977. Flora of Panama, Campanulaceae. Ann.

Missouri Rot. Card. 63: 593—655.

- . 1981. Additional Panamanian species of Bur¬
meistera (Campanulaceae: Lobelioideae). Ann. Missouri
Rot. Card. 68: 161-171.

Wimmer, F. E. 1932. Burmeistera, eirie umstrittene Pflan-
zengattung und ihre Arten. Repert. Spec. Nov. Regni
Veg. 30: 1—52 + pi. cxxiii-cxxvi.

- . 1943. Campanulaceae— Lobelioideae, I. Teil. Pp.

i-vi. 1-260 in R. Mansfeld (editor). Das Pflanzenreich.
IV. Teil, Abteilung 276b. Wilhelm Engelmann, Leipzig.

- . 1955. Lobeliacearum species novae austro-amer-

icanae. Brittonia 8: 107-111.

- . 1968. Campanulaceae— Lobelioideae supplemen-

tum et Campanulaceae— Cyphioideae. Pp. i— x, 815—
1024 in S. Danert (editor), Das Pflanzenreich, IV. Teil.
Abteilung 276c. Akademie-Verlag. Berlin.

Dos INuevas Especies de Gomphrena
(Amaranthaceae; Gomphrenoideae) de
los Valles Secos de Hoi i v i a

Teresa Ortuno

Museo Naeional de llistoria Natural, Herbario Naeional de Bolivia, (iota Cota Calle 27
Campus Universitario, Casilla 10077 Correo Central, La Paz, Bolivia, lpb@acelerate.com,

casstol@yahoo.com

Thomas Borsch

Necs-Institut fur Biodiversitat der Pflanzen, Fried rich-Wilhelms-Universitat Bonn,
Meckenheimer Alice 170, 531 15 Bonn, Germany, borsch@uni-bonn.de

Rksumkn. Se describe la morfologfa, palinologfa
y la distribueidn geografica actualmente eonoeida
de dos especies nuevas de Gomphrena, ambas en-
demicas de los valles secos de Bolivia. Gomphrena
fuscipellita es una especie perenne que se dife-
rencia de otras especies de Gomphrena por lener
lallos decumbentes v divididos dicotomicamente,
por estar densamente cubiertos por pelos fusco-
amarillos y por presentar inflorescencias compues-
tas de 5-6 eapftulos y hojas involucrales grandes.
Gomphrena mizqueensis tiene habito anual v es
morfoldgicamente afm a G. pallida, G. phaeotricha
y G. ferruginea. Esta nueva especie difiere de las
tres ultimas por presentar tallos erectos y ramifi-
cados, tepalos con margen dentado en el apice,
bracteas cimbiformes y dentadas en el apice, y ho¬
jas con un par de nervios laterales evidentes.

ABSTRACT. Morphology, pollen characters, and
geographical distribution as currently known are
described for two new species of Gomphrena. both
endemic to dry valleys of Bolivia. Gomphrena fus¬
cipellita is a perennial, differing from all other spe¬
cies oi Gomphrena by its decumbent habit, gener¬
ally dichotomously branching stems, dense
yellowish brown indumentum, and compound inflo¬
rescences with large involucral bracts. Gomphrena
mizqueensis is an annual, morphologically allied to
G. pallida, G. phaeotricha, and G. ferruginea. From
the three it differs by its erect habit branching in
its upper parts, tepals with dentate apex, cymbi-
form, apically dentate bracts, and ovate leaves with
one conspicuous pair of lateral veins.

Key words: Amaranthaceae, Bolivia, dry val¬
leys, endemics, Gomphrena. pollen.

Gomphrena L. es uno de los generos con mayor

numero lie especies en la familia de Amarantha¬
ceae (Eliasson, 1988; Siqueira, 1992; Townsend,
1993), e inclusive posee mas especies que Alter-
nanthera. La mayor diversidad del genero se pre-
senta en el Nuevo Mundo (80—100 spp.), en parti¬
cular en el Neotrbpico (principalmente en Bolivia.
Brasil) y en Australia (33 spp.; Palmer, 1998). Las
especies de Gomphrena son subarbustos, hierbas
perennes o hierbas anuales. El genero se caracte-
riza por hojas opuestas. flores solitarias y agrupadas
en eapftulos subglobosos o cortamente cilfndricos,
filamentos casi completamente connados formando
un tubo profundamente emarginado o 2-lobado, an-
teras sesiles o subsesiles y alternando con los lobos
qne son tan largos como las anteras o ligeramente
mas eortos, v estigma formando dos ramas cilfndri¬
cos o filiformes. Los linicos tratamientos taxono-
micos sobre especies neotropicales fueron realiza-
dos por Holzhammer (1955, 1956), y por Siqueira
(1992), quien estudio solamente taxa brasileros.
Otras importantes contribuciones para el genero
Gomphrena en America del Sur fueron realizados
por Pedersen (1976, 1990, 1997. 2000), quien (lis¬
ente la taxonomfa y nomenclature de algunas es-
pecies.

Gomphrena forma parte de la subfamilia Gom¬
phrenoideae que se caraeteriza por tener anteras
biloculares (Schinz. 1893. 1934). Besultados ob-
tenidos a traves de investigaeiones palinologicas
(Borsch, 1998) y de la filogenia obtenida sobre la
base de secuencias de ADN (rhe L. Kadereit et ah,
2003; ma/K//rnK, Muller & Borsch, 2005; ndliV,
Pratt. 2003; trn L-F, Sanchez del-Pino et al.. ined.)
sugieren que la subfamilia Gomphrenoideae es
monofiletica. Sin embargo, estos estudios tambien
sugieren que Gomphrena s.l. no es monofiletica ya
que un grupo de especies cercanas a Gomphrena

Novon 15: 180—189. 2005.

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2005

Ortuno & Borsch
Gomphrena de Bolivia

181

elegaris Marti us forma una Ifnea distinta. Las es-
pecies aquf examinadas pertenecen al genero Gom¬
phrena sensu stricto, el que sf es monofiletico de
acuerdo a resultados preliminares de uu analisis
filogenetico del genero Gomphrena (Ortuno &
Borsch, ined.i.

Durante la ejecucion de los proyectos actual-
mente vigentes sobre la diversidad de la familia
Amaranthaceae en Bolivia y sobre las relaciones
filogenetieas del genero Gomphrena , se reeonocen
dos nuevas espeeies singulares y facilmente distin-
guibles. En este trabajo deseribimos las dos espe¬
eies y presentamos datos sobre su distribution,
eeologfa y una evaluation preliminar sobre sus es-
tados de amenaza.

Gomphrena fuseipellita Ortuno N Borsch. >p.
nov. TIPO: Bolivia. Cochabamba: Mizque,
Tintin, camino de Mizque a Raykampampa,
ladera exposicion este, con pendiente de 35°,
estacionalmente humeda, 18°09,95”S,
65°25'98"W, 2870 m. 3 abr. 2003. T. Borsch
A- T. Ortuno 3594 (holotipo, LPB; isotipos.
BOIA. BONN. k. MO). Kiguras 1. 3A, 3C.
4A, 4C.

Herba manifeste perennis radice crassa; caulibus 1 vel
2 (vel 3), decumbentibus, dichotome ramosis, usque ad 25
cm longis, trichomatibus fuscis dense pilosis. Lamina fol-
iaris ovata, 1.7— 2.3 cm longa, 1.3— 1.8 cm lata, dense pi-
losa. Inflorescentia composita ex spicis (5 vel) 6 subglo-
bosis congestis loliis involucratibus 5 (sicut foliis
irifenoribus dense pilosis) quant spicis longioribus sub-
tentis constans; bractea ovata, membranacea, glabra.
2. 0-2. 2 mm longa. brevissime mucronata; braeteolis lan-
ceolatis, membranaceis, (lore aequilongis vel breve lon¬
gioribus, 3. 1-5.0 mm longis, dorsal iter crista munitis. Flos
tepalis dissimilibus. margine piloso, scariosis, externis
lanceolatis 3. 2-5.0 mm longis, 0.8—1. 3 mm lalis, inter-
medio si in i I i. trinervi, sicut externis trichomatibus undu-
latis e base ortis, internis linearibus, 3.0— 4.9 mm longis,
0.2— 0.8 mm latis; filamentorum tubi lobulis aeutis liberis
ca. 0.4 mm longis; antheris 1.0— 1.4 mm longis; stigmate
in ramos filiformes desinente.

Hierba perenne, tallos 1 — 2 ( — 3). decumbentes,
ramificados dicotomicamente, 20—25 cm de largo,
densamente hirsutos, tricomas fuscos, 0.6— 1.2 mm
de largo, uniseriados, con 4—6 celulas, la celula
basal mas amplia, las paredes celulares con gra-
nulos. Hojas opuestas, peciolo corto e inconspicuo,
laminas ovadas o elfpticas, 1.7— 2.3 X 1.3— 1.8 cm,
apice agudo o ligeramente acuminado, base corta-
mente deeurrente, superfieie densamente hirsuta,
pelos ± adpresos y similares a los del tallo. Inflo-
rescencias ascendentes, con pedunculos de 2—7 cm
de largo, eapitulos solitarios de 1 .2— 2.2 cm de dia-
metro, terminales o axilares. compuestos de 6 es-
pigas subglobosas, protegidas por 5 hojas involu-

crales. Hojas involucrales de color verde claro en
la superfieie adaxial y de color verde palido en la
superfieie abaxial, ovadas, sesiles, 0.5— 1.1 X 0.3—
1.0 cm. muy densamente hirsutas; braeteas ovadas.
cimbilormes, membranosas, hialinas y glabras, lle-
gando ± a la mitad de la altura de la llor. 2. 0-2.2
mm de largo X 0.9— 1.0 mm de ancho, con 1 ner-
vadura cortamente exeurrente (< 0.2 mm); brac-
teolas lanceoladas, cimbiformes, membranosas, gla¬
bras. blancas, casi del tamano o tin poco mas largas
que la (lor, 3.1— 5.0 X 0.9— 1.1 mm. en la parte
dorsal presentan una cresta, prolongada hasta cerca
la base, ligeramente encorvada, distalmente den-
tada. Tepalos desiguales, 5. los dos externos lan-
ceolados, 3.2— 5.0 X 0.8— 1.3 mm. 3-nervados, es-
cariosos y algo coriaceos desde la base hasta mas
o menos la mitad, el margen algo dentado con po-
cos tricomas blaneos y simples, dispuestos en un
angulo de 90°; el tepalo intermedio similar a los
externos, 3.1— 5.0 X 0.6— 1.0 mm, los dos interims
lineales, 3.0— 4.9 X 0.2— 0.8 mm. 1-nervados, es-
cariosos, con margen glabro; tricomas blaneos on-
dulados de hasta de 4 mm saliendo de la base de
los tepalos externos e intermedio; filamentos 3.1—
5.0 mm de largo, completamente unidos formando
un tubo, lobulos fibres, 0.4 mm de largo, anteras
de 1.0— 1.4 mm de largo; gineeeo subgloboso, 0.5
mm; estilo 0.4— 0.5 mm; estigma de 0.8 a 1.3 mm
de largo con ramas filiformes.

Polen esferoidal, de 14.4—16.0 pan de diametro
y con 34—44 aperturas. Font 3. 0-3.3 /am; corpus-
culos ectexinosos de la membrana 47—60. espar-
cidamente distribuidos en el centre del poro (el
margen no tiene corptisculos). La estruetura de la
membrana es metareticulada con elementos pen-
tagonales y hexagonales. Mesoporios delgados. 0.3
(Lim de ancho. Tectum prolongado hacia las partes
distales y reducido en las partes verticales. colu-
melas eortas fibres (visibles); la vista transversal
del tectum tiene forma delgada triangular termi-
nando en una punta aguda, 3 veces mas alta que
la columela; sin espfnulas y con conexiones pun-
teagudas.

Afinidades. Las espeeies mas similares a Gom¬
phrena fuseipellita por sus caracteristicas morfolo-
gicas son G. bicolor Martius y G. oligocephala
Remy. Estas dos espeeies son muy semejantes y
segun Pedersen (1976), la linica diferencia es en
el color de las braeteas y bracteolas; mientras (jue
en G. oligocephala son blancas, en G. bicolor son
cafes. Las caracteristicas compart idas por G. bicolor
y G. oligocephala como descritas por Holzhammer
(1955. 1956) y G. fuseipellita son las inflorescen-
cias compuestas de 6—7 eapitulos con un mismo

182

Novon

Hgura 1. Gomphrena fuscipellita Ortuno & Borsch. — A. Habito. — B. Hojas involucrales. — C. Flor con bract£olas y bractea separada. — [). Vista interior del tubo del androeceo
con gineceo y estigmas filiformes, vista exterior (derecha). — E. Tepalos, extemo con 3 venas (izquierda), medio con tres venas, interno con una vena (derecha). (Dibujo del holotino.
Borsch & Ortuno 3594, LPB.)

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Gomphrena de Bolivia

183

numero de hojas involucrales. Ademas, en estas es-
peries las bracteolas lienen erestas dentadas en el
apiee. El tubo estaminal de G. fuscipellita es similar
al de G. bicolor (lobulos entre las anteras unidos y
con apariencia de pseudoestaminodios), aunque los
lobulos son mas cortos: asf en G. fuscipellita miden
ca. 4 mm de largo y en G. bicolor miden hasta 8
mm de largo. Sin embargo, las bracteas de G. fus¬
cipellita son muy parecidas a las de G. oligocephala
ya que son blancas pero presentan una pequena
punta excurrente (< 2 mm).

Un tratamiento de Gomphrena fuscipellita como
una especie distinta es justificada por varias carae-
terfsticas singulares, siendo la mas notable la pre-
sencia de tricomas uniseriados fuscos densamente
dispuestos en sus en sus organos vegetativos. Ade¬
mas, los tallos en G. fuscipellita son ramificados
dicotomieamente y sus hojas se distribuyen homo-
geneamente por toda la planta. Otra caracteristica
es (|ue sus tallos siempre erecen inelinados liacia
la pendiente, mientras <|iie en G. oligocephala y G.
bicolor los tallos erecen siempre reetos y no se ra¬
mi bean . y la mayorfa de las hojas se encuentran
liacia la base de los tallos.

Los lfmites altitudinales donde fueron encontra-
das las poblaciones de Gomphrena fuscipellita
(2280—2870 m) son parecidos a los de G. oligoce¬
phala (2500—3300 m), mientras que G. bicolor se
eneuentra en lugares con mayor altitud (3400—3900
m). Muestras revisadas de las dos primeras espe-
cies son elementos de la vegetacion que erece sobre
suelos arenosos y pedregosos en la region de los
valles secos < lei sur de Bolivia (Cochabamba, Chu-
quisaca, Potosf). Al contrario, G. bicolor aparece
distribuido mas al norte hasta Peru encontrandose
tambien en el bosque hiimedo montano subtropical
(3400 m) y en pastizales de la puna (3900 m).

Distribucion y ecologia. La mayorfa de los es-
pecfmenes revisados de G. fuscipellita proviene de
la provincia de Mizque, departamento de Cocha¬
bamba, en un rango altitudinal de 2280 a 2870 m.
Gomphrena fuscipellita crece en laderas con expo-
sicion este, con pendientes (|ue varfan de 25° a 35°
sobre o cercana a afloramientos rocosos, en suelos
con lutitas y pizarras, arenosos, algo arcillosos y
rojizos. poco profundos, ligeramente aeidos y con
poca materia organiea acumulada, la cual esta cu-
bierta en la estacion humeda. Otras especies (pie
erecen en el habitat de G. fuscipellita son Selagi-
nella spp., Dodonaea viscosa Royen ex Blume (cha-
catea), Eupatorium buniifolium Hooker & Arnott
(tollita), Croton spp.. Salvia spp., h>pech inia spp.,
Stevia samaipatensis B. L. Robinson, Puya spp.,
Cheilantes pruinata Kaulfuss y Pellaea ternifolia
(Cavanilles) Link.

Paratipos. BOLIVIA. Potosf: Cliarcas, Toro Toro,
cerro Marika Paqui entrando la quebrada Mula wacana. ./.
Wood, M. Atahuachi & M. Mercado 19239 1 1 SOI .V. k.
LPB). Cochabamba: Mizque, camino viejo de Raykam-
pampa a Molinero, T. Borsch & T. Ortuno 3608 (BONN,
LPB); Raykampampa, Kollpana, E. Gutierrez 57 (BOLV);
Ramadero, A. Lopez & E. Saravia 381 (BOLV); camino a
herradura de Molineros hacia Botijas, A. Ldpez & E. Sar¬
avia 151 (BOLV); Cantdn Molinero a Ravkampapa Ra¬
madero, M. Sigle 237 (BOLV).

Gomphrena mizqueensis Ortuno & Borsch, sp.
nov. TIRO: Bolivia. Cochabamba: Campero,
Aiquile, camino nuevo de Raykampampa a Ai-
quile, a 1 hora de Aiquile, 18°11'57"S,
65° 17 '28" W, 2906 m, 4 abr. 2003. T. Ilorsch
& T. Ortuno 3614 (holotipo, LPB: isotipos,
BOLV, BONN, K. MO). Figuras 2, 3B, 3D,
4B, 41).

Herba annua radice tenui; can I i Dus erectis, triehotome
ramosis, usque ad 40 cm altis, trichoniatibus uniseriatis
2.0— 3.5 mm longis hirsutis. Lamina foliaris ovata vel ova-
to-elliptica, 1. 8-2.5 cm longa, 1.7-2. 1 cm lata, dense pi-
losa. lnflorescentia terminalis vel axillaris, longe pedun-
culata, composita ex spicis (5 vel) 0 subglobosis congestis
foliis involucralibus 5 (sicut loliis inferioribus dense pi¬
losis) quam spicis longioribus subtentis constans; bractea
elliptica, membranacea, glabra, 2. 0-2. 2 mm longa, den-
tata; bracteolis ovato-lanceolatis, membranaceis, flore ae-
quilongis, 3. 0-4. 2 mm longis, dorsaliter crista apiee den-
ticulata munitis. Flos tepalis dissimilibus, glabris,
scariosis, apiee denticulatis, externis lanceolatis, 2.3— 3.8
mm longis, ca. 0.6 mm latis, intermedin similis, sicut ex¬
tends trichotomatibus undulatis paueis e base ortis. inter-
nis linearibus, 2.1— 3.6 mm longis, ca. 0.4 mm latis; fila-
mentorum tubi lobulis ca. 0.6 mm longis, vicinis connatis
pseudostaminodia formantibus; antheris 1.0-1. 2 mm lon¬
gis; stigmate in ramos filiformis desinente.

Hierba anual, tallos erectos, muy ramificados en
la parte superior y divididos tricotomicamente, 35—
40 cm de largo, el color varfa de verde a rojizo,
laxamente pilosos con tricomas ± adpresos, de
blancos o palido amarillos, uniseriados, 2.0— 3.5
mm de largo, con 8—9 eel u las. la celula basal mas
amplia, las paredes celulares con granulos. Hojas
opuestas, subpecioladas, laminas ovadas, 1.8— 2.5
X 1.7-2. 1 cm, apiee agudo a ligeramente acumi-
nado, base decurrente, superficie densamente pi-
losa, con tricomas adpresos similares a los del tallo.
Inflorescencias erectas, con pedunculos de 1.0-14
cm de largo, capftulos solitarios de 0.6— 2.5 cm de
diametro, terminales o axi lares, compuestos de 6
espigas subglobosas, protegidas por 5 hojas invo¬
lucrales. Hojas involucrales de color verde claro en
la superficie adaxial, y de color verde palido en la
superficie abaxial, ovado-lanceoladas, sesiles. 0.5—
1 .9 X 0.2— 0.6 cm, densamente pilosas con tricomas
similares a los de las hojas. Espiga con (lores api-
eales mas pequenas, pero manteniendo el tamano

184

No von

Figura 2. Gomphrena mizqiutensis Ortuno & Borsch. — A. Habito. — B. Hojas involucrales. — C. Hoja. — 1>. Flor con
braet£olas y brfictea separada. — E. Iepalos, externo (izquierda), medio, interno (derecha) y un acercaniiento del apice
del t£palo externo. — F. Vista interior del tubo del androeeo con gineceo y estigmas (iliformes, vista exterior (derecha).
(Del holotipo. Borsch & Ortuno .if) 14, LPB.)

Volume 15, Number 1
2005

Ortuno & Borsch
Gomphrena de Bolivia

185

4

*00

V00

Figura 3. Fotomiorograffas del polen en MEB, polen limpiado con dimethoxypropano v acetona (secado en el punto critico). — A, C. Gomphrena fuscipellita (isotipo. Borsch &
Ortuno 3594, BONN). — B. I). Gomphrena mizqueensis (isotipo. Borsch A: Ortuno 3614 , BONN). La escala es igual a 1.0 pm.

186

Novon

ligura 4. Fotomicrograffas de pelos en MEB, ernes de las hojas, en grupo (arriha) y un detalle de la union de las e^lulas. — A. C. Gomphrena fuscipellita (isotipo Borsch & Ortuho
3608, BONN). — B, I). (Gomphrena mizqueensis (isotipo. Borsch & Ortuho 3614, BONN). La escala en A y B es igual a 100 pun, en C y D a 10 fim.

Volume 15, Number 1
2005

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187

de sus est met Liras reproductivas (gineceo, anteras);
bracteas elipsoides, cimbiformes, menibranosas,
hialinas, llegando ± a la mi tad de la altura de la
(lor, 2.0— 2. 1 X 1.2 mm, con I nervadura excurrente
(0.1 mm), hordes dentados en la parte apical; brac-
teolas ovadas, cimbiformes, menibranosas, glabras,
blancas, de iguai tamano que la (lor, 3. 0—4. 2 X 1.0
mm, en la parte dorsal presentan una cresta ubi-
cada en el Vi superior, distalmente dentada. Tepa-
los desiguales. 5, los dos externos lanceolados, 2.3—
3.8 X 0.6 mm, escariosos, con los apices dentados,
glabros, no se distingue claramente el numero de
nervios, el tepalo intermedin similar a los externos,
2. 2-3. 7 X 0.5— 0.6 mm, los internos lineales, a ve-
ees encorvados, 2.1— 3.6 X 0.4 mm, 1-nervados, es¬
cariosos, con margen glabro; tricomas blancos on-
dulados de 3 mm, saliendo de la base de los
tepalos, en algunas (lores mas densos (pie en otras;
filamentos 1.5— 4.6 mm tie largo, completamente
unidos (ormando un tubo, Idbulos (lobulos de los
estambres veeinos unidos parecidos a pseudoes-
laminodios) unidos a la mitad, 0.6 mm de largo;
anteras de 1 — 1.2 mm de largo; gineceo subgloboso,
0.2— 0.4 mm; estilo 0.2— 0.4 mm; estigma de 0.6
mm, con ramas (ililormes.

Polen esferoidal, de 13.0—14.0 /xm de diametro
y con 30—37 aperturas. Poro 1.6— 1.8 /xm; corpiis-
culos ectexinosos de la membrana 33—38, con for¬
ma irregular, densamente distribuidos en todo el
poro. La estructura de la membrana es metareti-
culada con elementos penta- y hexagonales. Me-
sosporios delgados, 0.3 /am de ancho. Tectum pro-
longado hacia las partes distales y completo en las
partes vertieales, columelas no visibles; la vista
transversal del tectum tiene forma delgada trian¬
gular terminando en una punta aguda; con espf-
nulas.

Afinidades. Gomphrena mizqueensis, al iguai
que G. ferruginea Pedersen, G. p<dlida (Suessen-
gutli) Pedersen y G. phaeotricha Pedersen es tam-
bien una espeeie anual. Estas especies se distri-
buyen en los valles secos de Bolivia y en el chaco
serrano, la prepuna y en valles secos de monte de
Argentina. Debido a ([ue las cuatro especies son
morfologieamente similares, se presenta una clave
para distinguirlas. La informal' ion sob re los earac-
leres de G. ferruginea, G. pallida, G. phaeotricha
se obtuvo de Pedersen (1976).

Clave paha Kspeciks Anuai.es Cekcanas a G. mi /,
ijt t: t:\sts (paka Fi-OREs en Lstado Maduho)

I . Bracteas cimbiformes, nervio cortamente ex¬
currente, con hordes dentados en la parte api¬
cal; apice de los tepalos dentado . . G. mizqueensis
I . Bracteas ovado-lanceoladas, agudas (nervio no

excurrente), con margen liso; apice de los te¬
palos liso o algo asemdado . 2

2(1). Braeteolas % mils largas (|ue los tepalos; cresta

con dientes notorios . G. ferruginea

2'. Braeteolas iguales o un poco mas largas que los
tepalos; cresta ± entera (dientes solamente cer-

ca del apice) . 3

3(2). Tepalos internos mas cortos, sin tricomas on-

dulados . G. phaeotricha

3'. Tepalos internos iguales en longitud, con tri-

comas ondulados . G. pallida

Suessenguth (1940) describio Gomphrena oligo-
cephala var. pallida Suessenguth sobre la base de
una planta anual colectada por Rodriguez (sub no.
1420, 22 Mar. 1914, LIL, LP. M). Pedersen (1976)
p resen t6 claras diferencias entre var. pallida y G.
oligocephala, y transfirio la variedad a nivel de es-
pecie (G. pallida). Debido a que Gomphrena miz¬
queensis y G. pallida crecen simpatrieamente, es
neeesario puntualizar todas sus diferencias. Gom¬
phrena mizqueensis es una planta alia (liasta 40
cm), erecta, con mas ramas y liojas en la parte su¬
perior. Sus liojas son ovadas y presentan un par de
nervaduras tan evidentes como el nervio central.
Las plantas tienen un indumento denso en los tallos
con tricomas que alcanzan 3.5 mm. Las liojas in-
volucrales presentan poca variacion en el tamafio y
estan regularmente distribuidas. El apice de los te¬
palos es dentado y los filamentos estan completa-
mente (usionados con los lobulos unidos (parecen
ser pseudoestaminodios). Mientras que, G. pallida
es una planta pequena (15 cm), ramificada desde
la base, con liojas oblaneeoladas y varies pares de
nervios tan evidentes como el nervio central. El
indumento presenta tricomas de hasta 6 nun de lon¬
gitud que estan esparsamente distribuidos en los
tallos. Las liojas involucrales son notoriamente va¬
riables en el tamano y el apice de los tepalos es
asemdado. Los filamentos estan fusionados par-
eialmente en un 90% con lobulos laterales y fibres.
Investigaeiones moleeulares preliminares sugieren
que G. mizqueensis es muy distinta a G. pallida,
mientras que es mas afin a G. fuscipellita (Ortuno
& Borsch, ined.).

Existen otras especies anuales que crecen en
esta region como Gomphrena boliviano Mart., G.
martiana (fillies ex Moquin-Tandon, G. platycepha-
la R. E. Fries y G. tarijensis R. E. Fries, pero estas
especies son morfologieamente diferentes y perte-
necen a otras li'neas de Gomphrena. Al contrario de
las anuales incluidas en la clave, G. martiana y
especies affiles presentan pequenos lobulos per-
pendiculares al tubo del androceo. Ademas de es¬
tas cuatro especies, existen dos especies anuales
mas pequenas (G. umbellata Remy y G. radiata
Pedersen), que se diferencian por presentar el an-

188

Novon

droceo sin lobos y las bracteolas sin nervios. De
los sistemas de clasifieaeion realizados por Seubert
(1875), Holzhammer (1955, 1956) y Siqueira
(1992), las (-species aqm nombradas solo estan in-
cluidas en los trabajos de Holzhammer (1955,
1956) ya que estos son mas amplios geogrdfica-
mente. Asf G. oligocephala Renty, G. bicolor Mar-
tius y G. pallida (Suessenguth) Pedersen |sub G.
oligocephala var. pallida Suessenguth] perteneeen
a la seccidn Gomphrena ; G. umbellata a la seccidn
Chnoanthus (Philippi) R. E. Fries; y G. martiana
s.l. con espeeies alines estan en Gomphrenula Seu-
bert. Estas seceiones fueron separadas por algunos
caracteres morfoldgicos (tipo de infloresceneia, pre-
seneia de eresta en bracteolas), pern esta por de-
terminarse si estos caracteres son plesiomdrficos o
apomdrficos en los analisis filogeneticos que estan
siendo efeetuados.

Distribut ion y ecologta. Las muestras analiza-
das de G. mizqueensis provienen de las provineias
Miz(|ue y Campero en un rango altitudinal de 2900
a 3600 m, ereciendo freeuentemente en suelos ped-
regosos de material suelto de pizarra, y junto a es¬
peeies como Kageneckia lanceolate Ruiz <& Pavdn,
Schinopsis haenkeana Engler, Dodonaea viscosa,
Gomphrena pallida y Salvia spp. La mayona de las
plantas fueron eneontrados eti Mizque; el nombre
Gomphrena mizqueensis hace referenda a esta pro-
vincia, que es conocida por sus valles seeos. Se
designd el tipo de esta (‘specie a una coleccidn de
la provineia Campero porque el material es de me-
jor calidad.

Ibiratipos. BOLIVIA, Corhaluunba: Mizque, eainiiio
de Mizque a Aiquile, segundo puente antes de llegar a
Miz(|ue, T. Horsch & T. Ortuno ,i5H4 (BONN, LPB); cum-
lircs entre Rakaypampa y Lagunas, C. Antezana 796
(BOI.V. BONN. LPB).

Gonsk.h vaciOn

La principal amenaza de la espeeie Gomphrena
fuscipellita esta relacionada eon la perdida de habi¬
tat, especialmente cerca de las loealidades de Ray-
kampampa y Aiquile. Esto se debe a la implemen-
tacidn de cultivos extensivos y monocultivo sin
descanso, lo que ademas de eausar una Incite de-
gradaeidn de la vegetaeidn xerdfila natural ha pro-
voeado la introduction de plagas por la ulilizaeidn
de semillas infestadas (Sigle. 1988). Por otro lado,
el sobrepastoreo de animales como eabras, ovejas,
cerdos, vaeas, acrecienta el peligro de la erosion.
Gomphrena mizqueensis fue encontrada en lugares
mas pedregosos, por lo eual, existe menus en areas
eultivadas y al pareeer no esta siendo afeetada por
aetividades humanas. Este trabajo podrfa sets una

base e incentive para (jue en el futuro se comple-
mente con mas informaeidn sobre las dos espeeies.

Vale reealear (jue estas espeeies. Gomphrena fus¬
cipellita y G. mizqueensis, estan distribuidas denim
de los valles seeos de Bolivia (jue constituyen eco-
sistemas diversos en espeeies vegetales, con 1300
espeeies registradas y poteneialmente mas de 2000,
ademas de presentar un alto nivel de endemismos,
eon un 18% de las esjiecies nativas (Ldpez, 2003).
Razon por la eual los trabajos como el de Navarro
et al. (1996), Antezana y Navarro (2002), Ldpez y
Reck (2002), ajumtan a conseguir mecanismos es-
tatales para resguardar estas zonas de importaneia
biologica por el riesgo y deterioro irreversible en
que se encuentran grandes extensiones con este
tipo de vegetaeidn.

Agradecimientos. Este trabajo es jiarte del
proyecto “Sistemdtica y diversidad de las Amar-
autaeeas de Bolivia”, realizado en eolaboracidn con
el Herbaria Nacional de Bolivia. Museo Naeional
de Historia Natural y el Nees-lnstitut fiir Biodi-
versitat der PHanzen de la Universidad de Bonn.
Damns las gracias a la Direccidn General de Bio-
diversidad. La Paz, por los permisos legales; a la
Deutsche Forschungsgemeinschaft (DEG) j>or la
financiaeidn del proyecto (mimero de jiroyecto BO
1815-1/1); al DAAD |>or brindarnos una beca para
T.0. para realizar estudios en Bonn; Darwin Initia¬
tive for the Conservation of Species para el finan-
eiamiento de una jiarte del trabajo de T.O.; a los
euradores del herbario BOEV y los curadores de
los otros herbarios meneionados por prestar mues-
tras, a Ramiro Ldpez, Carola Antezana, Jasivia
Gonzalez, kai Muller, Nelson Loza, John Wood y
Stephan Beck por sus utiles aportaciones; a Ivonne
Sanchez del-Pino y Alina Ereire Fierro por co-
mentarios a este manuscrito, a Roy Gereau por re-
visar el texto en latin, a Carlos Maldonado |>or su
eolaboracidn eon los dibujos y a Valentin Franco
Ramirez (tor su apoyo en el trabajo de campo.

Literatura Citada

Antezana, C. & C. Navarro. 2002. Contribucidn al andlisis
biogeografieo y caUilogo preliminar de la flora de los
valles seeos interandinos del centro de Bolivia. Rev.
Keol. Bol. 12: 3-38.

Borsch, T. 1998. Pollen types in the Amaranthaceae. Mor¬
phology and evolutionary significance. Grana 37: 129—
142.

Kliasson, U. H. 1988. Floral morphology and taxonomic
relations among the genera of Amaranthaceae in the
New World and the Hawaiian Islands. Bol. J. [.inn. Soe.
96: 235-283.

Holzhammer, K. 1955. Die amerikanischen Arten der Gat-
tung Gomphrena L. Teil 1. Milt. Bot. Staatssamml. Miin-
ehen 13: 85-1 14.

- . 1956. Die amerikanischen Arten der Gattung

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2005

Ortuno & Borsch
Gomphrena de Bolivia

189

Gomphrena L. Teil 2. Mill. Hot. Staatssamml. Mtinehen
14-15: 178-257.

Kadereil 0., Th. Borsch. K. Weising & H. Kreitag. 2008.
Phylogeny of Amaranthaceae and Chenopodiaceae and
die evolution of C, photosynthesis. Int. J. PI. Sci. 164:
959-986.

Lopez, R. P. 2003. Diversidad floristica y endemisnio de
los valles secos bolivianos. Ecol. Bolivia 38: 27—60.

- & S. Beck. 2002. Phytogeographical affinities and

life form composition of the Bolivian Prepnna. Candol-
lea 57: 77—96.

Vliiller k. & Borsch, T. 2005. Phylogenetics of Amaran-
thaceae based on malK/trriK sequence data — Evidence
from parsimony, likelihood, and Bayesian analyses.
Ann. Missouri Hot. Card. 92: 66—102.

Navarro, G., S. Arrazola. C. Antezana, E. Saravia & M.
Atahuachi. 1996. Series de vegetacion de los valles in-
ternos de los Andes de Cochabamba (Bolivia). Rev. Bob
Ecol. 1 : 3—20.

Palmer, J. 1998. A taxonomic revision of Gomphrena
(Amaranthaceae) in Australia. Austral. Syst. Bot. II:
73-161.

Pratt. I). B. 2003. Phylogeny and Morphological Evolution
of the Chenopodiaceae— Amaranthaceae Alliance. Ph.D.
Thesis, Iowa State University, Ames.

Pedersen, T. M. 1976. Studies in South American Amar¬
anthaceae II Darwinians 20: 269—302.

- . 1990. Studies in South American Amaranthaceae

111 (including one amphi- Atlantic species). Bull. Mus.
Natl. Ilisl. Nat.. Ser. 4. 12: 69-97.

- . 1997. Studies in South American Amaranthaceae

IV. Adansonia, ser. 3. 19: 217—251.

- . 2000. Studies in South American Amaranthaceae

V. Bonplandia 10: 83-1 12.

Schinz, II. 1893. Amaranthaceae. Pp. 91-118 en A. En-
gler & k. Prantl, Die natllrlichen Pflanzenfamilien, ed.
I. 3 (la), Leipzig.

- . 1934. Amaranthaceae. Pp. 7-85 en A. Engler &

k. Prantl, Die natiirlichen Pflanzenfamilien, ed. 2, 16c,
Leipzig.

Seubert, M. A. 1875. Amaranthaceae. En: C. E. P. von
Martins (editor), Flora Brasiliensis 5(1). Mtinehen.
Sigle, M. 1988. Observacion de malezas en sistemas tra-
dicionales de produccion agrfcola en las regiones de
Aiquile y Rakaypampa, Dpto. Cochabamba. Ecol. Bo¬
livia 11:' 1-23.

Siqueira. .1. C. tie. 1992. 0 genero Gomphrena I.. (Amar¬
anthaceae) no Brasil. Pesquisas, Bot. 43: 5—197.
Suessenguth, k. 1940. Einige neue und seltene Amaran-
taceen, Rhamnaceen und Vitaceen. Repert. Spec. Nov.
Reg. Vcg. 40: 5-15.

Townsend, C. C. 1993. Amaranthaceae. Pp. 70—91 en k.
Kubitzki et al.. Families and Genera of Flowering
Plants, Vol. 2. Springer, Berlin.

New Species and Nomenclatural Notes in Mesoamerican
Ardisia (Myrsinaceae)

John J. Pipoly III

Fairchild Tropical Garden, 1 1935 Old Cutler Road, Coral Cables (Miami),
Florida 33156-4299, U.S.A. jpipoly@fairchildgarden.org

Jon M. Picket son

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

jon.ricketson@mohot.org

Abstract. Taxonomic study of the Myrsinaceae
for the Flora Mesoamericana treatment resulted in
the discovery of three new species from Panama,
which are described and illustrated: Ardisia lierrer-
ana Pipoly cK Ricketson of subgenus Ardisia, and
.4. nana Pipoly & Ricketson and ,4. pseudocuspidata
Pipoly & Ricketson, both of subgenus Icacorea. In
addition, two new combinations within Ardisia
subg. Icacorea are made: Ardisia mexicana subsp.
siltepecana (Lundell) Pipoly & Ricketson anti Ar-
disia furfuracella subsp. veraguasensis (Lundell) Pi¬
poly & Ricketson, the latter with two heterotypic
binomials newly relegated to synonymy (.4. oerste-
diana Lundell and A. caudatifolia Lundell). Final¬
ly, we lectotypily Ardisia compressa Kunth var. mex¬
icana Oersted, a new taxonomic synonym, and
place A. jaliscensis Lundell into synonymy of A.
mexicana Lundell subsp. mexicana.

Rest MEN. El estudio taxonomieo de la familia
Myrsinaceae para el proyecto Flora Mesoamericana
resulto en el descubrimiento tie tres nuevas espe-
cies panamenas las cuales se describen e ilustran:
Ardisia herrerana Pipoly & Ricketson del subge-
nero Ardisia, A. nana Pipoly Ricketson yd. pseu-
docuspidata Pipoly Ricketson, ambas tlel subge-
nero Icacorea. Ademas, se proponen tins
combinaciones nuevas ilentro tie Ardisia suhg. Ica¬
corea-. Ardisia mexicana subsp. siltepecana (Lun¬
dell) Pipoly & Ricketson y Ardisia furfuracella
subsp. veraguasensis (Lundell) Pipoly & Ricketson,
y como novedad se relega a la sinonimia tie la ul¬
tima subespecie a tlos binomios heterotfpicos (.4.
oerstediana Lundell y .4. caudatifolia Lundell). Fin-
almente, se lectotipifica Ardisia compressa Kunth
var. mexicana Oersted, un nuevo sinonimo taxon-
omico, y se ubica a .4. jaliscensis Lundell como si-
ndnimo tie A. mexicana Lundell.

Key words: Ardisia, Ardisia subg. Ardisia, Ar¬

disia subg. Icacorea, conservation assessment. Flo¬
ra Mesoamericana, lectotypilications. Mexico. Myr-
sinaceae, Panama.

The pantropical genus Ardisia Swartz is the larg¬
est in tilt* family Myrsinaceae, containing perhaps
as many as 500 species (Chen & Pipoly, 1996). Its
circumscription has been problematic due to the
lack <>l a comprehensive treatment since that of Mez
(1902). The genus has traditionally been separated
from all others in the Myrsinaceae by what he in¬
terpreted as lree filaments, and by pluriseriate
ovules (Mez. 1902). Pipoly and Ricketson (1998)
found that the stamens in all subgenera are actually
connate basallv by their filaments to form a hyaline,
inconspicuous tube, but that the tube is free from
the corolla. The hyaline staminal tube, free from
the corolla tube, is the only unique characteristic
that distinguishes Ardisia from the other genera of
the family, and thus defines it.

While preparing the taxonomic treatment of Ar¬
disia for inclusion in the Myrsinaceae for Flora Me¬
soamericana, one new species was discovered with¬
in Ardisia subg. Ardisia and two additional ones
from subgenus Icacorea (Aublet) Mez. Also, two
new combinations were found to be needed, and
three binomials are relegated to synonymy. Finally,
while working up the synonymy for Ardisia mexi¬
cana, it was determined that one of its newly pro¬
posed taxonomic synonyms, A. compressa var. mex¬
icana, needed to be lectotypified.

Ardisia Swartz subg. Ardisia

Ardisia subg. Ardisia, as will be circumscribed
in the upcoming Flora Mesoamericana treatment,
comprises 1 1 species in Mesoamerica. Members of
this group are mostly locally common, occur in for¬
est margins and survive moderate levels of distur¬
bance (Pipoly, unpublished), and their ripe black

Novon 15: 190-201. 2005.

Volume 15, Number 1
2005

Pipoly & Ricketson
Mesoamerican Ardisia

191

to purple-black fruits provide a large food source habitats. They are easily propagated by cuttings

for birds, as well as the humans that live in their (Grijalva, pers. comm.).

Lev to Mesoamkrican Ardisia si bo. Arijisia

la. Pedicels covered with minute papillae, or simple multicellular trichomes.

2a. Calyx lobes 2.5 mm or longer . A. herrernna Pipoly & Ricketson

2h. Calyx lobes smaller, less than 2.2 mm.

3a. Pedicels of mature flowers 0—3 mm . A. bracteosa A. DC.

3b. Pedicels of mature flowers longer, slender, 4—7 mm.

4a. Leaf blades coriaceous, generally over 10 cm; pistil 6.1— 6.8 mm, the ovary 1.4— 1.6 mm, the

styles 4.5— 5.2 mm; mature fruits 3—5 mm diam . A. revoluta Kunth

4b. Leaf blades chartaceous, generally under 10 cm; pistil 7-8.4 mm, the ovary 1.6—2 mm. the

styles 6.4— 6.6 mm; mature fruits 5—7 mm diam . . A. escallonoides Schlechtendal & Chamisso

lb. Pedicels glabrous.

5a. Base of peduncle and apex of the branchlet with minute, dense papillae, and/or with simple multicellular
trichomes.

6a. inflorescence as long as or longer than the leaves; pedicels 6.2-12.4 mm . . A. colombiana Lundell

6b. Inflorescence much shorter than the leaves; pedicels 0—4.5 mm . A. granatensis Mez

5b. Base of peduncle and apex of the branchlet glabrous.

7a. Anthers versatile, 1.5— 1.6 X 0.4— 0.5 mm . A. breedlovei Lundell

7b. Anthers basifixed, more than 2 X 0.6 mm.

8a. Pedicels of mature flowers or fruits more than 6 mm . A. paschalis Donnell Smith

8b. Pedicels of mature flowers or fruits 0—1) mm.

9a. Leaf blades membranaceous, 2.8— 3.6 cm; flowers small, calyx lobes 1.3— 1.6 mm, es¬
sentially epunctate; corolla lobes 2.9— 3.1 mm; stamens 2.4— 2.5 mm, anthers 2.3— 2.5

mm . 4. escuintlensis Lundell

9b. Leaf blades chartaceous to coriaceous, 3.8—11.7 cm; flowers larger, calyx lobes 3.2—

7.5 mm, punctate; corolla lobes 6.7—12.8 mm; stamens 6.9—11.2 mm, anthers 2.8— 1). 9
mm.

10a. Calvx lobes 3.2— 6.2 mm; corolla lobes 6.7—10 mm, the corolla tube 2—4.7 mm;

the anthers 2. 8-5. 4 mm X LI -1.9 mm .

. 4. joetida Willdenow ex Boomer & Schultes

10b. Calyx lobes 7—7.5 mm; corolla lobes 12.2-12.8 mm, the corolla tube 1 — 1.2 mm;

the anthers 6.6— 6.9 mm X 1.9— 2.2 mm . A. perinsignis Lundell

I . Ardisia herrerana Pipoly & Ric ketson. sp.
iiov. TY PE: Panama. Darien: Parque Nacional
Darien, trail between Estacion Pine and Cerro
Pine, 08°00'N, 077°45'W. 100-300 m. 11 Pel)
1991 (fl. ft), //. Herrera 945 (holotype, MO;
isotypes, K. FTG, PM A not seen). Figure I.

Haec species quoad folia magna pedicellos papilloso-
tomentosos atque lobulos ealycinos magnos A. cabrerae
arete affinis, sed ab ea larninis foliaribus chartaceis (non
coriaeeis) ad apices acutis (nec acuminatis), inflorescen-
tiarum pinnatarum (non hipinnatarum) ranmlis 20 ad 10
(nec 7 ad 1 l)-floribus, pedicellis 5.3— 7.3 (nec 2.4—5) mm
longis. lobulis calycinis 3.1— 3.4 mm (non 2.7— 2.9) mm
longis ad apices rotundatis (nec obtusis), lobulis coroll inis
ellipticis usque lanceolatis (non ovatis) 5.8— 6.3 X 2.6— 2.9
(non 4.3— 4.5 X 3.4— 3.5) mm, staminibus 6.4— 6.7 (non
3.5— 3.7) mm longis, denique antheris 1.7— 1.9 (non 1.1 —
1.2) mm latis statim separabilis.

Trees to 9 in tall. Branchlets slender, terete, 6—8
mm diam., densely and minutely papillose, with
scattered to dense, prominent lenlieels. leaves with
blades chartaceous, oblong, elliptic, or oblaneeo-
late, 27—35 X 9.5—11.2 cm. apicallv acute, the
acumen 0.5— 1.3 mm long, basally cuneate, dccur-
rent on the petiole, inconspicuously punctate and

punctate-lineate, glabrous, the midrib impressed
above, prominently raised below, llic secondary
veins 52 to 70 pairs, prominulous above and below,
I lie margins entire, flat; petioles slender, marginate,
2.5—3 cm long, glabrous except often densely and
minutely papillose basally. Inflorescence terminal, a
pinnate panicle of racemes, 20-27 X 13—17 cm.
usually shorter than the leaves, densely and mi¬
nutely papillose, the branches bearing 20 to 40
flowers each; the peduncle 1.3— 5.2 cm long, the
lower branches subtended by leaves, densely and
minutely papillose or with minute simple multicel¬
lular trichomes; inflorescence primary bract and
branch bracts unknown (only scars visible); sec¬
ondary branches 0.9— 2.6 cm long; floral bracts ca¬
ducous, membranous, ovate, 9—21 X 7—15 mm,
apically acute, prominently punctate and punctate-
lineate, glabrous within, densely and minutely pa¬
pillose or with minute simple multicellular tri¬
chomes without, the margins entire, hyaline,
sparsely glandular-ciliolate; pedicels slender, te¬
rete, 5.3— 7.3 mm long, inconspicuously punctate
and punctate-lineate, densely and minutely papil¬
lose or with minute simple multicellular trichomes.

192

Novon

Figure I. Ardisia herrerana Pipoly & Ricketson. — A. F lowering branch. — B. Flower. — C. Detail ol stamen and
petal, showing dense yellow papillae. - — 1). Fruit. A— D drawn from the holotype, //. Herrera 945 (MO).

Flowers 5-merous; calyx lobes chartaceous, ovate,
3.1— 3.4 X 3.2— 3.5 mm, apieally rounded, conspic¬
uously punctate and punctate-lineate, glabrous ex¬
cept with sparse minute cupulate scales within, the

margins entire, hyaline, sparsely ciliolate; corolla
white, membranous, 8.4— 8.7 mm long, the tube
2.4— 2.6 mm long, the lobes elliptic to lanceolate,
5.8— 6.3 X 2.6— 2.9 mm, apieally acute to rounded.

Volume 15, Number 1
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Pipoly & Ricketson
Mesoamerican Ardisia

193

conspicuously punctate and punctate-lineate, gla¬
brous except with dense yellow papillae basally
within, the margins entire, hyaline; stamens 6.4—
6.7 mm long, the filaments 4—4.2 mm long, the sta-
minal tube 1.4— 1.7 mm long, the apically free por¬
tion 2.4— 2.6 mm long, inconspicuously punctate
and punctate-lineate, glabrous, the anthers free,
lanceolate, 2.2— 2.5 X 1.7— 1.9 mm, apically apic-
ulate, basally snbcordate, the connective incon¬
spicuously punctate; pistil unknown. Fruit (imma¬
ture) green, globose, 3. 5-5. 5 mm diam.,
conspicuously and prominently punctate and punc¬
tate-lineate.

Distribution. Ardisia her reran a is endemic to
the Parque Naeional Darien, on Cerro Pine in Da¬
rien Province, Panama, growing from 160 to 300 m
elevation.

Ecology and conservation assessment. Ardisia
herrerana is an apparent point endemic. There have
been a number ol large-scale botanical expeditions
to Cerro Pine, and the existence of only one col¬
lection points to the species’ rarity. According to
IUCN (2001) Red List criteria, this species would
be critically endangered (CRB2aD), owing to (R)
area ol occupancy estimated to be less than 10 km2
and (2a) known only from one collection, so (D) the
total population size estimated at less than 50 ma¬
ture individuals. Although A. herrerana occurs in
Parque Nat ional Darien, and its remote location
reduces threat, there remains the possibility of
threat from unforeseen factors (treefalls, storms,
etc.).

Etymology. This taxon is named for Heraclio
Herrera, its collector.

Ardisia herrerana belongs to Ardisia subg. Ardi¬
sia, which is circumscribed by racemose to spicate
inflorescence branches; calyx lobes symmetrical,
longer than wide, and not anriculate basally, glan-
dular-granulose adaxially at the base, and without
a subapieal notch along the margin. Ardisia herrer¬
ana could initially be confused with the common
species A. revoluta by its general leaf shape, size,
and large inflorescence. I lowever. Ardisia herrerana
is easily distinguished from A. revoluta by its mi¬

nutely papillose (not glabrous) branchlets; calyx
lobes chartaceous (not membranaceous), 3.1— 3.4 X

3.2— 3.5 (not 1.9— 2.1 X 1.6— 1.8) mm, apical ly
rounded (not acute); corolla 8.4— 8.7 (not 7.4— 7.8)
mm long, and by its stamens 6.4—6. 7 (not 5. 1-5.3)
mm long.

Ardisia herrerana is most similar to A. cabrerae
Pipoly because of its large leaves, long pedicels
that are densely covered with minute papillae and/
or simple multicellular trichomes, and the large ca¬
lyx lobes. However, A. herrerana differs from A. ca-
hrerae by its chartaceous (not coriaceous) and api¬
cally acute (not acuminate) leaf blades, its pinnate
(not bipinnate) inflorescence, its inflorescence
branches with 20 to 40 flowers (not 7 to I I), its
pedicels 5. 3-7.3 (not 2.4-5) nun long, its calyx
lobes 3.1— 3.4 (not 2.7— 2.9) mm long, and apically
rounded (not obtuse), its corolla lobes elliptic or
lanceolate (not ovate) and 5.8— 6.3 X 2.6— 2.9 (not

4. 3- 4.5 X 3. 4-3. 5) mm, its stamens 6. 4-6.7 (not
3. 5-3. 7) mm long, and its anthers 1.7— 1.9 (not 1.1—
1.2) mm wide.

Ardisia Swartz subg. Icacorea (Aublet) Mez

Ardisia subg. Icacorea, as will be circumscribed
in the upcoming Flora Mesoamericana treatment,
comprises 31 species with 33 taxa in Mesoamerica.
Of these, 12 are widespread, highly variable forest
margin and gap species, w ith extensive quantitative
plasticity in morphological characters that has led
to much taxonomic over-description. Members of
this group are mostly locally common, and because
they produce so many bright red fruits per panicle,
they provide a large food source for birds. They
have broad ecological tolerance, and thus would be
suitable for early introduction in any forest resto¬
ration effort.

The key below includes the new species and pre¬
sents our revised circumscriptions of Ardisia fur-
Jiiracella and A. mexicana. Vegetative or fruiting
calyx characters are used whenever possible to fa¬
cilitate identification, because the majority of her¬
barium specimens are in fruit. Terminology used
here follows that of Ricketson and Pipoly (2003).

Key to Mesoamerican Ardisia subg. Icacorea

la. Inflorescences axillary and terminal (sometimes apparently terminal, but with axillary inflorescence buds
easily visible).

2a. Leaf margins entire.

3a. Leaf apex short-acuminate to subacuminate, the base acute and decurrent on the petiole, but not
to its base; branchlets without interpetiolar ridges or angles.

4a. Branchlets, leaves, and inflorescences glabrous; anthers 3. 2-3.4 mm long; styles 4. 1-4.3

mm long . \. alajuelae (Lundell) Pipoly & Ricketson

4b. Branchlets, leaves, and inflorescences furfuraceous-lepidote; anthers 2.8-3 .2 mm long; styles

4.4— 5.2 mm long . \. hintonii Lundell

194

Novon

3b. Leaf apex long-acuminate to caudate or cuspidate, the base cuneate, rounded or obtuse, and
decurrent on the petiole to its base; branehlets with sharp angles or interpetiolar ridges.

3a. Leaf decurrent to petiole base and on stem; branehlets with interpetiolar ridges.

6a. Leaf blades coriaceous, apically acuminate but not caudate; petioles canaliculate; calyx

lobes 1.2— 2.2 mm wide; corolla lobes 4.5—6 X 2—2.4 mm . A. fendleri Lundell

6b. Leaf blades membranous to chartaeeous, apically caudate; petioles marginate; calyx

lobes 0.5- 1.2 mm wide; corolla lobes 2.9-4.6 X 1.6-2. 1 mm . . A. fiufuracella Standley
7a. Leaf blades membranous, essentially glabrous below, except sparsely to densely
and minutely furfuraeeous-lepidote along the midrib; pedicels glabrous; calyx
lobes 1.4— 1.6 X 1—1.3 mm; corolla lobes 4.4 — 4.6 X 1.7— 2.1 mm; anthers 2.2—

3.1 mm long; styles 4. 7-5.3 mm long .

. 4. furfuracella subsp. veragimsensis (Lundell) Pipoly & Ricketson

7b. Leaf blades chartaeeous, sparsely and minutely furfuraeeous-lepidote below, the
scales denser along the midrib; pedicels with scattered, minute furfuraeeous-
lepidote indument; calyx lobes 0.9— 1.2 X 0.5—1 mm; corolla 2.9— 3.2 X 1.6— 1.8

mm; anthers 1.6— 1.9 mm long; styles 3.6— 3.9 mm long .

. A. furfuracella Standley subsp. furfuracella

5b. Leaf base decurrent to petiole base, but not on stem; branehlets with angles, but without
interpetiolar ridges.

8a. Leaf blades chartaeeous, basally cuneate; calyx lobes 1.3— 1.5 X 1. 1-1.3 mm; styles

1.6- 5.3 mm long . 4- brevis Lundell

8b. Leaf blades membranous, basally obtuse to rounded; calyx lobes 1—1.3 X 0.8— 1.1 mm;

styles 2.9— 3.8 mm long.

9a. Corolla lobes 2.3— 2.6 X 1.1— 1.4 mm; anthers 1.4— 1.6 X 0.5— 0.7 mm; styles 2.9—

3.2 mm long; fruits 3. 5-4. 5 mm diam.; Cocos Island . A. cuspidala Bentbam

9b. Corolla lobes 3.2— 3.5 X 1.7— 1.9 mm; anthers 1.8—2 X 0.7— 0.9 mm; styles 3.5—

3.8 mm long; fruits 5-6 mm diam.; southern Darien/ northern Chocd .

. A. pseudocuspidata Pipoly & Ricketson

2b. Leaf margins undulate, crenulate, dentate or serrate.

10a. Flowers 4- or 5-merous in the same inflorescence . A. guianensis (Aublet) Mez

10b. Flowers strictly 5-merous.

1 la. Corolla lobes f.9-6.2 mm long; anthers 3.7—4. 4 mm long; Mexico to Guatemala .

. A. mexicana Lundell

12a. Calyx lobes 1.2-1. 4 mm long, apically acute; corolla lobes 6—6.2 mm long; anthers

4.1— 4.4 X 0.6-0.8 mm; styles (>-6.2 mm long ... .4. mexicana Lundell subsp. mexicana
12b. Calyx lobes 0. 9-1.2 mm long, apically rounded; corolla lobes 4. 9-5. 2 mm long; anthers

3.7— 3.9 X 1.8-2 mm; styles 4.9— 5.3 mm long .

. A. mexicana subsp. siltepecana (Lundell) Pipoly & Ricketson

I lb. Corolla lobes 3.8-4.3 mm long; anthers 2.2-2.8 mm long; Nicaragua to Panama.

13a. Leaf margins serrate-dentate; calyx lobes 1.3-1. 4 X 1. 2-1.4 mm; anthers 2.2-2. 4 mm

long . .4. longicaudata Lundell

13b. Leaf margins undulate to crenulate; calyx lobes 1-1.3 X 0.6— 0.8 mm; anthers 2.4—

2.8 mm long . A. fruticosa Lundell

lb. Inflorescences strictly terminal, with no axillary buds visible at nodes below.

14a. Inflorescence a compact, ovoid capitulum . A. glomeriflora .1. K Morales

14 b. Inflorescence an open panicle.

15a. Leaf margins undulate, crenulate, serrate or dentate.

16a. Styles 3.5—4. 5 mm long.

17a. Ultimate branches of inflorescence racemose; calyx lobes 1.3— 1.5 mm long; corolla

lobes 4.4— 4.7 X 2—2.2 mm; anthers 2.9— 3.1 mm long . A. geniculata Lundell

17b. Ultimate branches of inflorescence corymbose; calyx lobes 0.9— 1.3 mm long; corolla
lobes 3. 7-4.2 X 1.5— 1.9 mm; anthers 1.9-2. 5 mm long.

18a. Branehlets with conspicuous interpetiolar ridges; styles 4.2— 4.5 mm long ....

. 4. amanuensis Lundell

18b. Branehlets terete or angled, without interpetiolar ridges; styles 3.5— 3.8 mm long

. A. irasuensis Oersted

16b. Styles 5.4— 6.1 mm long.

19a. Inflorescences 2.5— 3.2 X 0.9-1. 5 cm; calyx lobes 0.9-1 mm wide . . . A. cookii Lundell
19b. Inflorescences 4—24 X 3—21 cm; calyx lobes 1 — 1.3 mm wide.

20a. Anthers 2. 9-3.2 mm long; stvles 5.9-6. 1 mm long . 4. subcrenulata Lundell

20b. Anthers 3.1—4 mm long; styles 5.4— 5.9 mm long . 4. compressa Kunth

15b. Leaf margins entire.

21a. Leaf blades membranous to chartaeeous.

22a. Calyx lobes 0,6— 0.8 X 0.0— 0.8 mm; corolla lobes 3—3.4 X 1.7- 1.9 mm; anthers 1.5—

1.6 mm long, oblong . -4. scheryi Lundell

22h. Calyx lobes 1—2.3 X 1-2.2 mm; corolla lobes 4.3—7. 1 X 1.9— 2.7 mm; anthers 2.8—

4.8 mm long, lanceolate.

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21b.

23a. Leal blades apieally obtuse to subaeuminate; calyx lobes 1.7— 2.3 X 1.7— 2.2 mm;

styles 6.8— 8.8 mm long . 4. maxonii Standley

23b. Leaf blades apieally acute to acuminate; calyx lobes 1—1.5 X 1 — 1.3 mm; styles
4.8— 5.9 mm long.

24a. Calyx lobes 1.4— 1.5 mm; corolla lobes 2.4— 2.5 mm long; anthers 2.8—3 mm

long; style 4.8— 4.9 mm long; petioles 0.3— 0.8 cm long . 4. storkii Lundell

24b. Calyx lobes 1 — 1.4 mm long; corolla lobes 3.4—4 mm long; anthers 3.4—4

mm long; style 5.4— 5.9 mm long; petioles 0.6— 1.6 cm long .

. 4. compressa Kunth

Leaf blades subcoriaceous to coriaceous.

25a. Leaf blades decurrent to petiole base and on branchlets, the branchlets with prominent
interpetiolar ridges.

26a. Flowers 4-merous; calyx lobes 1-1.2 X 0.6— 0.7 mm; corolla lobes 3-3.2 mm

long; anthers 0.5— 0.6 nun wide; styles 3.6— 3.7 mm long .

. A. nana Pipoly & Ricketson

26b. Flowers 5-merous; calyx lobes 1.2— 2.9 X 1.1—2 mm; corolla lobes 4.5— 6.5 mm
long; anthers 0.9— 1.2 mm wide; styles 5—6.9 mm long.

27a. Branchlets and inflorescence rachis glabrous; petioles absent or to 0.5 cm
long, slender; calyx lobes 2.5— 2.9 X 1.5—2 mm; corolla lobes 2.3— 2.5 mm

wide; styles 6.3— 6.9 mm long . A. subsessilifolia Lundell

27b. Branchlets and inflorescence rachis with scattered, minute furfuraeeous-
lepidote indument at least apieally, usually glabrescent with age; petioles
absent or to 2.2 cm long, stout; calyx lobes 1.2— 2.2 X 1.1— 1.5 mm; corolla

lobes 1 .5— 2.2 mm wide; styles 5—6.1 mm long . A. copeyana Standley

25b. Leal blades decurrent only to petiole base; branchlets without interpetiolar ridges.

28a. Branchlets terete.

29a. Calyx lobes 1.3— 1.5 mm long; corolla lobes 1.6— 1.8 mm wide; anthers 0.8—

0.9 mm wide; styles 5.8-6. 1 mm long . A. rigidifolia Lundell

29b. Calyx lobes 1.1 — 1.3 mm long; corolla lobes 1.9— 2.1 mm wide; anthers 1 —

1.1 mm wide; styles 4.3— 4.5 mm long . A. jefeana Lundell

28b. Branchlets angulate.

30a. Anthers 2—2.2 X 0.7— 0.8 mm; stamens 3.8—4 mm long; pedicels 3—1.5 mm
long; inflorescences 4—7 cm long; leaf blades short-acuminate, inconspic¬
uously punctate and punetate-lineate . A. guancheana Lundell

30h. Anthers 2.3— 1.4 X 0.8— 1.2 mm: stamens 4.1— 7.3 mm long; pedicels 5—8
mm long; inflorescences 9-14 cm long; leaf blades short-acute to -acumi¬
nate, conspicuously and prominently punctate and punetate-lineate ....
. A. costaricensis Lundell

i. Ardisia nana Pipoly & Ricketson, sp. nov.
T\ PF: Panama. Panama: Cabecera del Rfo Pi-
riadi, 09°11'05"N, 078°16'00"W. 100-150 m,
1.3 June 1994 (11). //. Herrera l(>27 (holotvpe,
MO; isotvpes, F, FTG. MO. PM A not seen.
STR1 not seen). Figure 2.

Haec species oh petiolos ad ramulos deeurrentes, ra-
mulos inter petiolos-porcatos, laminas foliares subeori-
aceas vel eoriaceas A. subsessilifoliae et 4. copeyanae arete
affinis, sed ab eis floribus 4- (non 5-) metis, lobulis ca-
lycinis 1.0— 1.2 X 0.6— 0.7 (non 1 .2— 2.9 X 1.1— 2.0) mm
longis, lobulis eorollinis 3.0— 3.2 (non 4.5— 6.5) mm longis,
antheris 0.5— 0.6 (non 0.9— 1.2) mm latis, denique sty 1 is
3.6— 3.7 (non 5.0— 5.9) mm longis statim separabilis.

Shrubs to I m tall. Branchlets slender, angulate,
2—4 mm diam., densely and minutely furfuraceous-
lepidote at least apieally, usually glabrescent with
age, with prominent interpetiolar ridges. Leaves
with blades subcoriaceous, elliptic or oblong to ob-
laneeolate, 3—10.1 X 0.9— 2. 1 cm, apieally acute to
acuminate, the acumen 8—12 mm long, basally
acute to cuneate, decurrent on the petiole and con¬

tinuous with ihe interpetiolar ridges, inconspicu¬
ously punctate and punetate-lineate, glabrous
above, with scattered, minute furfuraceous-lepidote
indument below, glabrous along the midrib, the sec¬
ondary veins 16 to 24 pairs, the margins entire, flat;
petioles slender, marginate, 0.3— 0.5 cm long, gla¬
brous. Inflorescence terminal, erect, bipinnately to
tripinnately paniculate, 4—5 X 3—3.5 cm. pyrami¬
dal. shorter than the leaves, densely and minutely
furfuraceous-lepidote, the branches loosely con¬
gested into 3- to 9-flowered corymbs; peduncles 2—
5 nun long; inflorescence branch bracts caducous,
membranous, ovate to oblong, 1.5—2 X 0.6— 0.9
mm, apieally acute, prominently punctate and
punetate-lineate, glabrous, the margins minutely
erose, hyaline; floral bracts similar to the inflores¬
cence branch bracts but ovate. 0.6— 0.8 X 0.4— 0.5
mm; pedicels slender, terete, 4—5 mm long, prom¬
inently punctate and punetate-lineate, glabrous.
Flowers 4-merous; calyx 1.3— 1.5 mm long, the tube
0.2— 0.3 mm long, the lobes ovate. 1 — 1.2 X 0.6— 0.7
mm, apieally acute to rounded, conspicuously and

196

Novon

Figure 2. Ardisia nana Pipoly & Ricketson. — A. Flowering branch. — 15. Flower. — C. Fruit. A— C drawn from the
holotype. //. Herrera lf>27 (MO).

prominently punctate and punctate-] ineate, gla¬
brous. the margins minutely erose, hyaline; corolla
white, 4.2— 4.3 mm long, the tube 1 — 1.2 mm long,
the lobes lanceolate, 3—3.2 X 1.5— 1.7 mm, apically

acute, prominently punctate and punctate-lineate,
glabrous, the margins entire; stamens 3.5— 3.7 mm
long, the filaments 1.7— 1.8 mm long, the staminal
tube 0.3— 0.5 mm long, the apically free portion

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Mesoamerican Ardisia

197

1.2— 1.5 mm long, epunctate, glabrous, the anthers
free, lanceolate, 2.1— 2.2 X 0.5— 0.6 mm, apieally
apiculate, basal ly subcordate, the connective in¬
conspicuously punctate; pistil 4.4— 4.6 mm long,
glabrous, the ovary ovoid, 0.6— 0.7 mm long, the
style 3.6— 3.7 mm long, epunctate, the ovules 7 to
9. Fruits unknown.

Distribution. Ardisia nana is endemic to the
Kfo Piriadi area in the province of Panama, Pana¬
ma, from 100 to 150 m elevation.

Ecology and conservation assessment. This spe¬
cies occurs in the unnamed valley that forms the
drainage basin of I .ago Bayamo and Kfo Chepo. It
houses tropical wet forest facing the Pacific, where
the Choco Floristic Province of Panama, Colombia,
and Ecuador has its northernmost extension. The
valley is just east of where the Pan American High¬
way ends, so threats to the area owing to timber
harvest and settlement are much less than they are
farther westward. The species’ extent of occurrence,
based on availability of suitable habitat, is approx¬
imately 4150 kur, and the known area of occupan¬
cy is less than 1 km2. Therefore, we regard this
species as Endangered, ENBlal). sensu the II CN
Bed List criteria (ILCN, 2001). I he Endangered
status was determined based on the fact that the
(111) extent of occurrence is less than 5000 km2, (a)
known from less than five locations, and (I)) the
population size is estimated at fewer than 250 in¬
dividuals. It is significant that the type collection
was gathered from one of the most heavily collected
areas ol Panama, so we can be relatively certain
that it is at least a rare species in terms of number
of individuals per hectare.

Etymology. The epithet comes from the Latin
word "nana," meaning dwarf, referring to the small
leaves and flowers.

The decurrent leaf bases running to interpetiolar
ridges, and subeoriaceous to coriaceous leaf blades
indicate that Ardisia nana is close to A. subsessili-
folia and A. copeyana. However, A. nana is imme¬
diately separable from the latter two species by the
4-merous flowers, shorter and narrower calyx lobes,
smaller corolla lobes, and smaller anthers and
style.

2. Ardisia pscudocuspidatu Pipoly & Ricketson,
sp. nov. TYPE: Panama. Darien: SW ridge
leading to Alturas de Nique on the Colombian
border, 600 m. 28 Dec. 1980 (fl), R. Hartman
12363 (holotype, MO; isotypes, LL. PMA not
seen). Figure 3.

Haec species propter rhachides inflorescentiarum an-
gulatas et longitudinal iter interamulari-poreatas, laminam

foliarem membranaceam (non chartaceam), ad basim ob-
tusam vel rotundatam (nee euneatam), necnon lobulos ca-
lvcinos 1 — 1.3 X 0.9— 1.1 (non 1.3— 1.5 X 1. 1 — 1.3) mm A.
cuspidatae valde arete af fin is, sed ah ea lobulis corollinis
3.2— 3.5 X 1.7— 1.9 (non 2.3— 2.6 X 1. 1 — 1.4) mm, antheris
I 8— 2.0 X 0.7— 0.9 (non 1.4— 1.6 X 0.5— 0.7) mm. stylis
2.9— 3.2 (non 3.5— 3.8) mm longis, fructibus 5.0-6. 0 (non
3. 5-4.5) mm diametris, praeclare distat.

Small trees 3 — 1 m tall, to 9 cm diam. Hranchlets
slender, sharply angulate, 1-4 mm diam., densely
and minutely furfuraceous-lepidote at least apical-
ly. usually glabrescent with age. leaves with blades
membranous, ovate to elliptic, 4. 3-8.7 X 1.8-3. 7
cm. apieally cuspidate, the acumen 7—14 mm long,
basally obtuse to rounded, decurrent to petiole
base, but not onto stem, prominently punctate and
punctate-lineate, glabrous above, with scattered mi¬
nute furfuraceous-lepidote indument below, the
secondary veins 28 to 35 pairs, obscure to promi-
nulous above and below, the margins entire, flat;
petioles slender, marginate, 3—5 mm long, glabrous
above, densely and minutely furfuraceous-lepidote
below. Inflorescences lateral and terminal, erect,
pinnately to bipinnately paniculate, 3—5 X 3—4.5
cm. pyramidal, shorter than the leaves, densely and
minutely furfuraceous-lepidote, the branches loose¬
ly congested into 3- to 7-flowered corymbs; pedun¬
cles 0.3— 0.6 cm long; inflorescence branch bracts
unknown, early caducous; floral bracts caducous,
membranous, ovate, 0.6—1 X 0.4— 0.8 mm, apieally
acute, conspicuously and prominently punctate and
punctate-lineate, glabrous within, with scattered
minute furfuraceous-lepidote indument without, the
margins minutely erose, hyaline; pedicels slender,
terete, 2.9-4. 1 mm long, prominently punctate and
punctate-lineate, densely and minutely furfura¬
ceous-lepidote. Flowers 5-merous; calyx 1 .6-1.8
mm long, the tube 0.4— 0.6 mm long, the lobes
ovate, 1 — 1.2 X 0.9— 1.1 mm, apieally acute, con¬
spicuously and prominently punctate and punctate-
lineate, glabrous within, with scattered minute fur¬
furaceous-lepidote indument without, the margins
minutely erose, hyaline; corolla white, membra¬
nous, 4.4— 4.7 mm long, the tube 1 — 1.2 mm long,
the lobes lanceolate 3.2— 3.5 X 1.7— 1.9 mm. api¬
eally acute, prominently punctate and punctate-li¬
neate, glabrous, the margins entire; stamens 3.4—
3.7 mm long, the filaments 2-2.2 mm long, the
staminal lube 0. 7-0.9 mm long, the apieally free
portion 1.1— 1.3 mm long, prominently punctate and
punctate-lineate, glabrous, the anthers free, oblong
to lanceolate, 1.8—2 X 0.7— 0.9 mm. apieally apic¬
ulate, basally cordate, the connective inconspicu¬
ously punctate and punctate-lineate; pistil LI-4.5
mm long, glabrous, the ovary ovoid, 0.7— 0.9 mm
long, the style 3.5— 3.8 mm long, inconspicuously

198

Novon

5 cm

Figure 3. Ardisia pseudocuspidata Pipoly & Ricketson. — A. Flowering branch. — B. Detail of inflorescence. — C
Fruit. A c!4 B drawn from the holotype. R. Hartman 12363 (MO): C from A. Gentry & A. Clewell 70R2 (MO).

punctate and punctate-lineate, the ovules 38 to 47.
Fruits globose, 5—6 mm diam., prominently punc ¬
tate and punctate-lineate, glabrous.

Distribution. Ardisia pseudocuspidata is en¬
demic to the province of Darien, Panama (belong¬
ing to the Choed Holistic Province), near the* Col¬
ombian border, from 800 to 1400 m elevation.

Ecology and conservation assessment. This spe¬
cie's occurs along ridges in cloud and elfin forests.
The high-altitude areas on the Panantanian-Col-
ombian border are remote and might be safe from
imminent destruction owing to civil unrest. The
area lies within Darien National Park and has Ka-
tios National Park adjoining it on the Colombian

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Mesoamerican Ardisia

199

side, both oi which afford further protection. The
species is restricted to that portion of the Serrania
del Darien that comprises Cerro Pine and Nique,
a mountain chain running from southwest to north¬
east and forming the border between Panama and
Colombia. Therefore, we estimate the possible ex¬
tent of occurrence for the species to be approxi¬
mately 500 km2 and the area of occupancy certainly
less than that. Using the IUCN (2001) Red List
Criteria, Ardisia pseudocuspidata is Endangered,
EN B2aD, because the area of occupancy is esti¬
mated to be less than 500 km2 (B2), the species is
known from less than five locations (a), and the
population size is estimated to be less than 250
mature individuals (D).

Etymology. The specific epithet refers to its
similarity to Ardisia cuspidata.

Paratypes. PANAMA. Darien: Cerro Campamcnlo, 5
of Cerro Pine. 20-22 Mar. 1908 (IV). ./. Duke 1 5676 (MO,
PM A not seen); summit of Cerro Pirre, 29 Dec. 1972 (fr),
,4. Gentry <£ .4. Clewell 7082 (IT. MO. PIMA not seen).

Owing to its angulate inflorescence raehises and
sharply angulate branch lets, membranous leaf
blades obtuse to rounded basally, and calvx lobes
from 1—1.3 X 0.9— 1.1 mm. Ardisia pseudocuspidata
is closely related to ,4. cuspidata. However, Ardisia
pseudocuspidata is clearly separated from A. cus¬
pidata by its large corolla lobes 3.2— 3.5 X 1.7— 1.9
(not 2.3— 2.6 X 1.1— 1.4) mm. larger anthers 1 .8—
2.0 X 0.7— 0.9 (not 1.4— 1.6 X 0.5— 0.7) mm, longer
styles 3.5— 3.8 (not 2.9— 3.2) mm long, and larger
fruits 5.0— 6.0 (not 3.5— 4.5) mm in diameter.

Taxonomic revision of Ardisia subg. Icacorea also
disclosed the necessity of making the following two
combinations, syonymizations and typifieation.

3a. Ardisia furfuracella Standley suhsp. furfur-
acclla, Ann. Missouri Bot. Card. 25: 832.
1938. TV PE: Panama. Chiriqui: valley of up¬
per Rfo Chiriqui Viejo, 1300—1900 m, (II). D.
& 6. White 8 (holotype, E; isotypes A, EE neg.
1971-49. MO).

Distribution. Ardisia furfuracella subsp. furfur¬
acella occurs throughout the Cordillera de Tala-
manea in the provinces of Cartago and Puntarenas,
Costa Rica, to the province of Chiriqui, Panama,
from 1300 to 2200 m elevation.

Ecology and conservation assessment .
Subspecies furfuracella has an extent of occurrence
nearing 24,000 km2 and an area ol occupancy of
approximately 12.000 km2. The species is known
to be locally common, but the populations are very
fragmented as this subspecies specializes in I lie
transition zone from montane wet forest to cloud

forest habitats. That transition zone is the upper
limit for coffee cultivation and a number of other
products, and therefore, the disturbance noted by
Pipoly ( Pipoly 7608) is likely to increase. We there¬
fore consider this species to be Vulnerable,
VUCl,C2ai, where we note that (Cl) the population
sizes are estimated to number fewer than I (),()()()
mature individuals and (C2) a continuing decline
is inferred in numbers of mature individuals and
(a) population structure is in the form of (i) no sub¬
population estimated to contain more than 1000
mature individuals.

3b. Ardisia furfuracella Standley subsp. vera-
guasensis (Lundell) Pipoly & Ricketson.
comb, et stat. nov. Basionym: Ardisia veragu-
asensis Eundell, Wrightia 5: 64. 1974. Icacorea
veraguasensis (Eundell) Eundell, Phytologia
49: 352. 1981. TYPE: Panama. Veraguas: Ca¬
ribbean slope above Rfo Primero Brazo, 5 mi.
NW Santa Fe, 700-1200 m. 18-19 Mar. 1973
(fl). R. Liesner 986 (holotype. EE: isoty pes. GH,
MO. PM A not seen).

Ardisia caudatifolia Lundell, Wrightia 5: 278. 1976. Syn.
nov. Icacorea caudatifolia (Lundell) Lundell. Phvto-
logia 49: 347. 1981. TYPE: Panama. Veraguas: along
stream between Santa Fe & Eseuela Agrfcola Alto
Piedras, without elev., 29 Aug. 1974 (fr). /. Groat
27244 (holotype, IT, F neg. 55648; isotvpes, MO.
PMA not seen).

Ardisia oerstediana Lundell, Wrightia 6: 84. 1979. Syn.
nov. Icacorea oerstediana (Lundell) Lundell, Phyto¬
logia 49: 350. 1981. TYPE: Panama. Veraguas: 0.6
mi. beyond Eseuela Agrfcola Alto Piedra. 730 m. 4
\pr. 1976 (fl), T. Groat & ./. Folsom 84055 (holotype,
IT, F neg. 55678; isotypes, MO. PMA not seen).

Distribution. Ardisia furfuracella subsp. vera¬
guasensis is known from Nicaragua southward to
the province of Veraguas, Panama, from 100 to
1600 m elevation.

Ecology and conservation assessment. Phis sub¬
species occurs in open, windswept forest margins
and has significant tolerance to drying and other
disturbance. The extent of occurrence is 108.900
km2, with an area of occupancy of nearly 12,100
km2 and with 60 collections known from 27 popu¬
lations. There are no data regarding number of in¬
dividuals per population, but all of the environ¬
ments mentioned were disturbed ones. I lierefore
we must assume that the plant is rather resilient,
and we would therefore classify it as of least con¬
cern, EC.

Ardisia furfuracella subsp. veraguasensis may be
separated from subspecies furfuracella by its mem¬
branous (not chartaceous) leaf blades that are es¬
sentially glabrous below, except with scattered to

200

No von

densely and minutely furfuraceous-lepidote scales
along the midrib (not scattered furfuraceous-lepi¬
dote across the leaf below), the glabrous (not lepi-
dote) pedicels, the larger calyx lobes 1.4— 1.6 X
1.1 — 1.3 (not 0.9— 1.2 X ( ),5 — I ) mm, with larger co¬
rolla lobes 4.4— 4.6 X 1.7— 2. 1 (not 2.9— 3.2 X 1 .6) —

1 .8) mm, larger anthers 2.2— 3.1 (not 1.6— 1.9) mm
long, and longer style 4.7— 5.3 (not 3.6— 3.9) mm
long.

The type of Ardisia caudatifolia is qualitatively
identical to the type of A. veraguasensis , differing
only in its longer leaves and inflorescences. Like¬
wise. the type of A. oerstediana is notable only for
its large membranous leaf blades that are extremely
bullate, and the inflorescences that are larger and
much wider than those of the type of A. veragu¬
asensis. In all other aspects the two are identical.
The variation seen in the types of Ardisia cauda¬
tifolia as well as that observed in A. oerstediana fall
well within the range seen across the species. Vie
therefore set* no support for their separation as spe¬
cies, and have thus placed Ardisia veraguasensis
and A. oerstediana in synonymy with A. furfuracella
subsp. caudatifolia.

We recognize Ardisia furfuracella subsp. vera¬
guasensis as distinct from subspecies furfuracella
based on its leaf blades, which are membranous
(not chartaceous), essentially glabrous below, ex¬
cept sparsely to densely and minutely furfuraceous-
lepidote along the midrib (not sparsely and mi¬
nutely furfuraceous-lepidote below, the scales
denser along the midrib), pedicels glabrous (not
with scattered, minute furfuraceous-lepidote indu-
ment), calyx lobes 1.4— 1.6 X 1 — 1.3 (not 0.9— 1.2 X
0.5—1) mm; corolla lobes 4.4— 4.6 X 1.7— 2.1 (not
2.9— 3.2 X 1.6— 1.8) mm; anthers 2.2-3. 1 (not I .(>-

1.9) mm long; and styles 4.7— 5.3 (not 3.6— 3.9) mm
long.

During our studies of a group of entities related
to Ardisia mexicana , the need to leetotypify its syn¬
onym, Ardisia compressa var. mexicana, became ap¬
parent. and is accomplished herewith.

4a. Ardisia mexicana Lundell subsp. mexi-
cana, Wrightia 3: 77. 1963. Gentlea mexicana
(Lundell) Lundell, Wrightia 5: 44. 1974.
TYPE: Mexico. Jalisco: crest of ridge facing
the Pacific, 10 mi. S of Aullan. ea. 5700 ft.
|I737 nij, 20 Aug. 1949 (fl), R. Wilbur A- C.
Wilbur 2460 (holotype, LL; isotype, MICH).

Ardisia compressa Kunth var. mexicana Oersted, Vidensk.
Meddel. Dansk Naturhist. Karen. Kjabenhavti 1861:
125. 1862. TV PE: Mexico. Oaxaca: Prope Quatulco,
without elev., Oct. 1812 (fl). F. Liebmann 21 (lecto-
type, designated here, C).

Icacorea jalisccnsis Lundell, Phytologia 53: 112. 1983.
Syn. nov. Ardisia jalisccnsis (Lundell) Lundell, Phy¬
tologia 61: 65. 1986, nom. inval. Ardisia jaliscensis
(Lundell) Pi poly & Ricketson, Sida 18: 513. 1998.
TYPE: Mexico. Jalisco: along road to Jirosto, ea. 10
km WNW of Purificaeion. ca. 19°44'N, 104°42'W,
400 m. II Jan. 1979 (fr), //. litis & M. Nee 1427
(holotype, LI. -TEX; isotype, MEXll 2 sheets. US.
W IS not seen).

In A. Oersted’s (1862: 125) original description
of Ardisia compressa var. mexicana, three collec¬
tions were cited: “Cl. Liebmann hanc varietateum
prope Quatulco et Mirador a mense Oetobris in jan-
uariurn usque florentem et juxta Colipa Martin fruc-
tificantem legit." Without a clear indication ol a
holotype, a lectotype must be selected. Unfortu¬
nately, t be specimen from Quatulco is the only re¬
maining Liebmann collection of this species at C
where the Oersted types are housed. The Missouri
Botanical Garden houses a duplicate of the Colipa
collection, and no collection from Mirador can be
located. Thus we hereby designate the F. Liebmann
21 collection from Quatulco at C as the lectotype.

Distribution. Ardisia mexicana subsp. mexi¬
cana is restricted to the foothills of the Sierra Ma-
dre del Sur mountain range in the broad sense,
from the state of Nayarit in the north to Oaxaca in
the south, and in (lie foothills of the mountains of
Veracruz, from 100 to 300 m elevation.

Ecology and conservation assessment. The ex¬
tent of occurrence for this subspecies is approxi¬
mately 48,000 km2, with an area of occupancy of
less than 1000 km2, and collections made from only
15 locations. Label data from herbarium specimens
indicate that the plant is rare in these localities,
often at the lower margin of pine-oak forests, where
access roads typically occur. W'e classify this spe¬
cies as Vulnerable, VU B2 bii, biii, biv, where B2
indicates that the area of occupancy is estimated
to he It jss than 2000 km2, and there is a continuing
decline observed and projected in the (bii) area of
occupancy, (biii) quality of habitat, and (biv) num¬
ber of locations or subpopulations. The populations
from Nayarit have not been observed since 1957,
and new collections coming to us from floristic pro¬
jects in Mexico have not included any new mate¬
rial.

41». Ardisia mexicana Lundell subsp. siltepe-
cana (Lundell) Pipoly & Ricketson, comb, et
stat. nov. Basionym: Ardisia siltepecana Lun¬
dell. Wrightia 4: 66. 1968. Icacorea siltepecana
(Lundell) Lundell, Phytologia 49: 351. 1981.
TYPE: Mexico. Chiapas: Cascada near Silte-
pec, 1600 m, 1 Mar. 1945 (fl), E. Matilda .5/6/
(holotype, LL; isotypes, F, LL 2 sheets,
MEXU).

Volume 15, Number 1
2005

Pipoly & Ricketson
Mesoamerican Ardisia

201

Distribution. This subspecies is endemic- to the
southern portion of Chiapas, Mexico, from 200 to
1600 m elevation.

Ecology and conservation assessment. Ardisia
mexicana subsp. siltepecana occurs along river
banks in dry forests. Using the IUCN (2001) cri¬
teria we consider it endangered because its extent
of occurrence is less than 5000 km2, it is known
from fewer than five gatherings at separate loca¬
tions, the area of occupancy is less than 500 km2,
and the population size is estimated to number less
than 2500 mature individuals. KNBla, 2a.

This subspecies differs from Ardisia mexicana
subsp. mexicana by its chartaceous (not membra¬
nous) leaf blades with scattered furfuraceous-lepi-
dote scales over the entire surface (not restricted to
the veins), the smaller calyx lobes 0.9— 1.2 X 0.5—
1 (not 1.4— 1.6 X 1—1.3) mm, the smaller corolla
lobes 2. 9-3.2 X 1. 6-1.8 (not 4.4-T.6 X 1.7-2. 1 )
mm, the smaller anthers 1.6— 1.9 (not 2.2— 3.1) mm
long, and the shorter styles 3.6— 3.9 (not 4.7— 5.3)
mm long. Subspecies siltepecana is also restricted
to dry forests instead of the humid habitats of sub¬
species mexicana.

Acknowledgments. We are grateful to the Mis¬
souri Botanical Garden (MO), to Gerrit Davidse
(MO) as well as the Flora Mesoamericana Project,
and the Fairchild Tropical Garden for their support
of this inter-institutional research effort. Many
thanks are due to Hoy Gereau (MO) for reviewing
the Latin diagnoses. The drawings were prepared
by Jon Kieketson.

I .iterature Cited

Chen, ,1. & J. .1. Pipoly III. 19%. Myrsinaceae. Pp. 1—38
in Wu Xheng-yi & P. H. Raven (editors). Flora of China,
Vol. 15. Science Press, Beijing, and Missouri Botanical
Garden Press, St. Louis.

IUCN. 2001. IUCN Bed List Categories and Criteria, ver¬
sion 3.1. (http://www.iucn.org/themes/ssc/red-lists.htm)
Vic/.. C. 1902. Myrsinaceae. In: A. Lngler (editor). Das
Pflanzenreich I\. 236 (Heft 9): 1—137.

Oersterl, A. S. 1862. Myrsineae Centroamericanae et Mex-
icanae. Vidensk. Meddel. Dansk Naturhist. Fpren.
Kjpbenhavn 1861: 117-142.

Pipoly, J. .)., Ill J. VI. Ricketson. 1998. A revision of
the genus Ardisia subgenus Graphardisia (Myrsinaceae).
Sida 18: 433-472.

Ricketson. J. M. & J. J. Pipoly III. 2003. Revision of
Ardisia subgenus Auriculardisia (Myrsinaceae). Ann.
Missouri Bot. Card. 90: 179—317.

New Species of Siparuna (Siparunaceae) IV. A New Subcanopy
Free from White-Sand Areas in Brazil and Venezuela

Susanne S. Renner

Kakultat fur Biologie, Ludwig-Maximilians Universiliit Miinchen, Munich, Germany,
renner@lrz.uni-muenchen.de; Missouri Botanical Garden, St. Louis, Missouri 63100, U.S.A.

Gerlinde Hausner

Werner-Hilpert-Str. 07, D-65197 Wiesbaden, Germany

ABSTRACT. A new species of Siparuna (Siparun¬
aceae), S. ficoides, is described, illustrated, and
placed in a phylogenetic context based on morpho¬
logical and 1)N A sequence data. The species,
which is a monoecious subcanopy tree, is known
from three collections made near Manaus, Brazil
(two from the same tree), and one in the state of
Bolivar, Venezuela.

KKSUMKN. Sc describe y se iluslra una nueva es-
pecie de Siparuna (Siparunaceae), S. ficoides, ade¬
nitis sc la uhica en un contexto filogenetico basado
cn datos morloldgicos y en sequencias de DNA.
Esta nueva especie es un arbol monoico de sub-
doscl, de la cual se ban registrado tres colecciones
ccrca de Manaos, Brasil (dos del misrno arbol) y
una coleccidn en el Estado de Bolivar, Venezuela.

Key words: Brazil, Central Amazon basin, forest
on sandy soils, Siparuna, Siparunaceae, Venezuela.

The Ducke Forest Reserve comprises some
10,000 ha ol non-floodcd forest on the outskirts of
Manaus, capital of the state of Amazonas, Brazil.
Between 1955 and 1978, about 7000 collections of
vascular plants were made there, and another 7354
were added between 1992 and 1998, as part of a
British-Brazilian technical cooperation that result¬
ed in a “Field Guide to the Vascular Plants” (Ri-
beiro et al., 1999; M. Hopkins, pers. comm.. May
2004). This collection density makes the Ducke
forest the best-documented site in the Amazon ba¬
sin, al least with regard to vascular plants. It is all
the more remarkable that new species of trees con¬
tinue to come to light among collections made in
the reserve during the 1990s that are now available
for study, mainly in the herbaria INPA, k, MG, NY,
RB, and SR. One such species is described here,
based on collections from tin; Ducke reserve and
from the Venezuelan state of Bolfvar. This distri¬
butional range makes sense phytogeograph ically
because the Ducke area harbors many species

adapted to sandy soils, such as are typical of the
Rio Negro basin and the Venezuelan Gran Savanna.

Siparunaceae consist ol just two genera, Sipa¬
runa Aublet (including Bracteanthus Ducke) and
its sister group Glossocalyx Bentham, a monotypic
West African genus very similar to Siparuna (Ren¬
ner, 2004; Renner & Hausner, 2005). Most of the
53 species ol Siparuna are dioecious, but the phy-
logenetically basal species (in a molecular phytog¬
eny; Renner & Won, 2001) are monoecious and
include the large tree species as well as most low¬
land species found in the genus. Besides monoecy,
they share entire-margined leaves, fruitlets that
lack slylar arils, and little-developed lepals. (“Sty-
lar aril” is a term introduced by Corner 1 1976| for
the fleshy appendages present on some Siparuna¬
ceae fruitlets; others, for example, Endress |1980|
and Barroso 1 1999], have referred to these append¬
ages as superarils because they do not derive from
tin; seed or funiculus like a true aril [van der Pijl,
1982: 139, fig. 25], Instead, the appendages in Si¬
paruna develop as asymmetric outgrowths at the
style base. The stylar arils in Siparuna are orange
or red in color and rich in oil, but contain little or
no starch. As far as known, monoecious species
lack stylar arils, and their drupelets’ exo- and ine-
soearp instead often becomes slimy.) The new spe¬
cies described here has been sequenced for one
chloroplast locus and one nuclear locus (Renner &
Won, 2001, under the name S. all. monogyna), and
based on these data it belongs in a clade that also
includes S. cristata (Poeppig & Endlicher) A. DC.,
the species to which it is most similar morpholog¬
ically (see below).

Siparuna ficoides Renner & Hausner, sp. nov.
TYPE: Brazil. Amazonas: Res. f lor. Ducke,
Manaus— Ilacoatiara, km 20, 02°53'S,

59°58'W, permanently tagged tree 3325-74,
29 Sep. 1995, C. A. Sothers, E. da C. Pereira
Si C. E. da Silva 591 (holotype, IN PA; iso¬
types, K, MG. MO, NY, R, SP. U). Figures 1.2.

Novon 15: 202-200. 2005.

Volume 15, Number 1
2005

Renner & Hausner
New Species of Siparuna

203

Figure I. — A. Habit of Si/xiruno ficoides Renner Hausner ( Sot hers el ul. 597, MO, isotype). — B. Distribution of
.S', ficoides.

204

Novon

figure 1. Sifmruna ficoides Renner i\ Hausner. - — A. Inflorescences with male (left) and female flowers (right, some
broken oil); (Sothers el al. 501, MO, isotype). — I). Lower leal surface, showing the loose tertiary venation (Hriceflo 427,
\). paratype). (,. fruiting receptacle (Assun^no el al. 808, N), paratype). — I). Rranehlet with voting fruiting recep¬
tacles (Keserva f lorestal Ducke, tree ,51. id-74; vouchers from this tree are Assun^ao el al. 808 and Vicentini el al.
IOIO). h. Inflorescence with male flowers (Reserva f lorestal Ducke, tree .5.525-71; a voucher from this tree includes
the isotype Sol hers el al. 501, MO). Photos I) and K: M. ,1. (7 Hopkins.

Volume 15, Number 1
2005

Renner & Hausner
New Species of Siparuna

205

A Siparuna crislata foliis brevioribus el angustioribus
(9-15 X 4—8.8 cm) cum venis tertians irregulariter an-
astomosantibus laxe dispersis fructibus subglobosus (ca.
2 X 1.1 cm) pilosibusque differt.

Monoecious subcanopy tree, 13—20 m tall, with a
DBH ol 10—11 cm, the trunk straight, the slash
wound light tan or yellowish and strongly lemon-
scented; young hranchlets terete or flattened at the
nodes, glabrous. Leaves opposite; petioles 1—2.2 cm
long; lamina drying light green , yellowish or grayish
green, shiny, stiff-chartaceous to coriaceous, ob-
ovate or elliptic, 9—15 X 4.4— 8.8 cm, the base ob¬
tuse and shortly attenuate, t he apex obtuse, acu¬
minate, truncate, or emarginate, mature leaves
glabrous except for very few peltate or stellate-lep-
idote hairs on the lower leaf surface or on the mid¬
rib; with 9 to 12 pairs of secondary veins, these flat
above, slightly raised below, anastomosing near the
leaf margin, the tertiary veins irregularly reticulate
and loosely spaced (Fig. 2B), the margin entire.
Cymes in the axils ol extant leaves or on leafless
nodes, about 1.5 cm long, with 6 to 12 flowers, the
peduncle 4— 6 mm long, the pedicels of male flowers
gradually elongating during anthesis and becoming
up to 8 mm long, those of the female flowers and
fruiting receptacles remaining short (Figs. 1. 2);
cymes persistently velvety. Fresh flowers green with
a cream-colored center, dried golden-brownish in
color; male flowers cup-shaped, 3—3.5 mm high, 4—
4.5 mm diam., the indumentum as on the cymes,
the tepals reduced to a circular rim surrounding the
flower center with its harely developed tomentose
floral roof (Fig. 2A, E), stamens 27 to 30. (leshv
and with small anther flaps, dorsally with a few
stellate hairs; female flowers subglobose, 2.5— 3.5
mm high, 4—5 mm diam., the tepals reduced to a
circular rim around the flower center (Fig. 2A); the
floral roof conical and velvety; carpels ca. 8, the
styles basally fused to a short column and hardly
protruding from the floral roof, the free parts of the
styles thin and easily detached. Fruiting receptacle
subglobose, ca. 1.1 cm in height and 2 cm diam.
(Fig. 2C), when fresh greenish yellow and densely
pubescent (fig. 21)), dried material persistently vel¬
vety brown-pubescent, the drupelets protruding
(Fig. 2C), drupelets about 6, apparently lacking a
stylar aril.

Distribution (Fig. 1 B), habitat, and phenolo¬
gy. Siparuna ficoides is known from the Central
Amazon basin and the state of Bolivar in Venezue¬
la, where it grows on sandy soils at altitudes up to
about 410 m. Flowers and fruits have been col¬
lected in August and September.

Etymology. The species is named for its leaves
resembling certain species of Ficus L. (Moraceae).

Common name. flic Venezuelan Yekuana (or
Yekuna) Indians refer to this species as Medebadi
and apply the leaves against snakebites ( Briceho
427).

Siparuna ficoides resembles S. cristata, which
ranges throughout the Amazon basin and into
northern Venezuela, adjacent northernmost Colom¬
bia. and Panama, and which also occurs in the
Dueke reserve. The two species have similar gla¬
brous leathery leaves, but S. cristata has glabrous
fruits that are elongate in shape, reaching 3.5—4 cm
in length, w ith a diameter of (l-)2-3.5 cm, whereas
the fruits of S. ficoides are densely velvety pubes¬
cent and subglobose in shape. In addition, S. cris¬
tata usually has 10 to 18 stamens, while S. ficoides
has 27 to 30 stamens, the leaves of S. cristata dry
dark brown and their blades measure 18— 2 7 ( — 38)
X (7-)9-I4 cm, while leaves of S. ficoides dry light
green or light brown and have blades measuring 9—
15 X 4.4— 8.8 cm. Vet another difference lies in the
leaf venation: in .S', cristata, the tertiary veins cross
between adjacent secondaries in parallel paths
without branching, while in S. ficoides they are ir¬
regularly anastomosing and loosely spaced (Fig.
2B) compared to other species of Siparuna. A few
duplicates of S. ficoides were distributed as S. mon-
ogyna Jangoux, a name that we briefly thought
might pertain to the new entity, but a study of the
type ol S. monogyna revealed that this name does
not apply. Another monoecious species with velvety
fruits is S. gentryana S. S. Renner, occurring in
Ecuador and Colombia. However, S. gentryana has
tubereulate fruits, rather than smooth, subglobose
ones, and dark brown leaves with a venation more
like S. cristata.

Paratypes. VENFZUELA. Bolivar: Foraneo Aripao,
Botadero Norte. 8 May 1994, E. Briceno 427 (NY). BRA¬
ZIL. Amazonas: Res. Flor. Dueke, Manaus— Itacoatiara,
km 26, near observation tower, permanently tagged tree
3133-74, 5 Sep. 1995, A. Vicentini, P. A. C. /,. Assunyao,
C. F. da Silva A E. da C. Pereira 1010 (INPA, k. MG,
MO, NY, R, RB, SP, U), same tree collected again, 6 Mar.
1998, P. A. C. L. Assunyao, E. da C. Pereira & C. F. da
Silva BOB (INPA, k, MG, MO. NY, RB, SP, U. UB).

Acknowledgments. We thank Mike Hopkins
(MG) and Cynthia Sothers (K) for photos of the new
species, and the reviewers David Lorence and Juan
Martfnez-Laborde for their comments.

Literature Cited

Barroso, G. M. 1999. Fmtos e Semeules: Morlologia Apli-

eada a Sistematiea de Dicotiledoneas. Viyosa, Editora

UFV, Universidade Federal de Y’iyosa. Brasil.

Corner, E. J. H. 1976. The Seeds of Dicotyledons. Cam¬
bridge Univ. Press, Cambridge.

Endress, P. K. 1980. Ontogeny, function and evolution of

206

Novon

extreme floral construction in Monimiaceae. PI. Syst.
Evol. 134: 79-120.

Pijl, I,, van dcr. 1902. Principles of Dispersal in Higher
Plants, 3rd ed. Springer, Berlin.

Benner, S. S. 2004. Siparunaceae. Pp. 353—355 in N. P.
Smith, S. A. Mori, A. Henderson, 1). W. Stevenson &
S. V. Heald (editors). Flowering Plants of the Neotrop¬
ics. Princeton Univ. Press, Princeton.

- & C. Hausner. 2005. Siparunaceae. Flora Neotro-

pica 95.

- & II. Won. 2001. Bepeateil evolution of dioeey

from monoecy in Siparunaceae (Faurales). Syst. Biol.
50: 700-712.

Bibeiro, J. E. L. da S., M. J. G. Hopkins, A. Vieentini, C.
A. Sothers, M. A. da S. Costa, J. M. de Brito. M. A. I).
de Souza, L. H. P. Martins. L. G. Lolimann, P. A. C. I,.
Assunyao, F. da C. Pereira, C. F. da Silva, M. B. Mes-
quita & F. G. Proeopio. 1999. Flora of the Ducke Re¬
serve: Field Guide to the Vascular Plants of a Terra
Finna Forest in Central Amazonia. INPA/DFID, Brazil.

A New, Red-Flowered Species of Tripodanthus
(Loranthaceae) from Colombia

Francisco ./. Roldan

Departamento de Biologia, Universidad de Antioquia, Apto. 1226,
Medellin, Antioquia, Colombia

Job Kuijt

Department of Biology, University of Victoria, Victoria, BC V8W 3N5, Canada.

jknijt@uvic.ca

Abstract. A new species of Tripodanthus, T. bel-
mirensis Roldan & Kuijt, is described and illus¬
trated from Antioquia, Colombia. In contrast to the
other two, more southern species of the genus, T.
acutifolius (Ruiz & Pavon) Tieghem and T. flagel-
laris (Chamisso & Schlechtendal) Tieghem, it lias
bright red, much larger flowers. This is the first
report of Tripodanthus from Colombia.

Key words: Colombia, Loranthaceae, Tripodan-
tlius.

A remarkable red-flowered member of the mis¬
tletoe family Loranthaceae was collected some
years ago in the area of Belmira, Colombia, and the
present report documents the species as a member
of the genus Tripodanthus (Kichler) Tieghem. Until
this time, Tripodanthus has consisted of two white-
flowered species, T. flagellaris (Chamisso & Schle¬
chtendal) Tieghem (northern Argentina, adjacent
Uruguay, and southern Brazil) and 7. acutifolius
(Ruiz & Pavon) Tieghem (northern Argentina, Uru¬
guay, and southern Brazil to Venezuela; not yet
known from Colombia).

Tripodaiitlms belinirensis Roldan & Kuijt, sp.
nov. IA PE: Colombia. Antioquia: Belmira,
Einca El Paramo, 3000—3130 m, 6°35'N,
75°32'W, 21 Apr. 1993, R. Fonnegra & I).
Tuberquia 4576 (holotype, HU A; isotype,
LEA), f igures 1 . 2.

Caulis subquadrangulus, folia opposita, laneeolata. In-
florescentia terminalis, paribus triadarum 5 -8. Flores basi
lutei, apice rubri.

Robust, glabrous plants lacking epieortieal roots
on the stem, but base of plant unknown; young
stems somewhat fleshy and quadrangular, older
stems ± terete and bearing numerous small pus¬
tular lenticels; lateral branches subtended by 2 or
more pairs of scale leaves; prophylls subtended by

distal foliage leaves very prominent, acute, 2 mm
long. Leaves paired, when fresh with yellowish to
brown margin and prominently raised, yellow lower
midrib running into apex or nearly so, lanceolate,
to 5 X 2.5 cm, petiole 3—5 mm, canaliculate; apex
of blade acute, base obtuse to nearly truncate, ve¬
nation pinnate. Inflorescences terminating leafy in¬
novations, very compact, with 5 to 8 crowded pairs
(sometimes trios?) of triads; triad peduncle angular,
7 mm long, the primary bract 2-5 mm long, ca¬
ducous; all flowers on pedicels 5 mm long, the two
lateral ones with a caducous bract similar to the
primary one. Mature bud 3 cm long, straight, bright
red above, yellow below, slightly expanded above
the calyculus, and narrowing upward but expanding
again somewhat below the nearly acute apex, ovary

5 mm long, narrow below, widening to 3 mm above,
the calyculus smooth to very shallowly dentate; pet¬
als 2.5 cm long, very slightly dimorphic, becoming
yellow at anthesis; ligule small, ca. 3 mm above
base of petal; filaments 8 mm long, inserted at two
different heights on the petals; anthers dorsifixed
and versatile, 3 mm long, with small, terminal eon-
nectival horn; a few long, unbranched, and very
slender yellow hairs also present, growing out from
the petals near the insertion of the filaments; nec¬
tary small, 6-cornered. Pollen somewhat similar to
that of T. acutifolius (Eeuer & Kuijt, 1980, figs. 23

6 24), polar length ca. 12 gm, diameter slightly
more than 20 p m, isopolar, amb triangular, deeply
concave; tricolpate, syncolpate, the colpal portions
nearest the poles psilate but from about halfway to
the equator distinctly deepened, this portion also
psilate but flanked by small, irregular marginal
lobes; colpal margins raised, psilate, continuous
across the equator where somewhat widening.
Equatorial faces irregularly depressed rugulate-ver-
rucate. Fruit red ( Roldan et al. 2575).

Assignment of the new species to Tripodanthus

Novon 15: 207—209. 2005.

208

Novon

Figure 1. Tripodanthus belmirensis Roldan & Kuijt (based on Fonnegrn el id. 5400, I, FA), —a. Habit of (lowering
shoot. — b. Triad in bud, nearly mature (arrows: caducous bract and bracteole). — c. Style and two petals, and associated
stamens, showing dimorphism. — d. Prophyll (I.: leaf).

Volume 15, Number 1
2005

Roldan & Kuijt

Tripodanthus belmirensis from Colombia

209

Figure 2. Pollen of Tripodanthus belmirensis (based on Fonnegra el al. 5400 , LEA). — a. Polar view. - — 1>. Equatorial
view, showing pore.

would seem to be correct, but its occurrence at high
elevation and its isolation from the other two spe¬
cies of Tripodanthus, neither of which are known
from Colombia, is surprising. Other genera that
might be considered briefly are Gaiadendron C.
Don and Tristerix Martius. The former genus readi¬
es from Bolivia to Venezuela and Ml. Boraima, and
has the same subparamo preference as our new
species; it also has large bracts and bracteoles and
flowers which, however, range from white to golden
yellow and nearly orange. Its pollen structure is
also somewhat similar to that of T. belmirensis , even
though the syncolpus is very narrow. Gaiadendron
shows a series of unique features relating lo ger¬
mination and establishment, which at this lime can¬
not be checked in T. belmirensis. Among these are
its frequently terrestrial habit, lack of primary
haustorium, presence of a tuber in the seedling,
and deeply grooved endosperm (Kuijt. 1963). It is
extremely unlikely, however, that T. belmirensis
shares those features.

The general aspect of the plant is much more
reminiscent of Tristerix than Tripodanthus. Its
leaves and especially the curious, acute prophylls
are similar to those in some Tristerix species (Kuijt.
1988; see fig. 16e). Geographically, also, the range
of Tristerix is suggestive. However, the occurrence
of triads in the inflorescence, dimorphic stamens,
and palynological details of T. belmirensis do not
support assignment to Tristerix and agree with Tri¬
podanthus. Chromosome numbers in Gaiadendron
and Tristerix (n = 12) arc very different from Tri¬
podanthus (n — 8) (Barlow & Wiens, 1973), and
knowing this characteristic in the new species
would provide significant input, as would knowl¬

edge of its haustorial attachment; Gaiadendron
lacks a primary haustorium. and Tristerix lacks ep-
icortical roots and, if epicortical roots are present
al the base of T. belmirensis, this would lend sup¬
port to the present assignment. The curious cadu¬
cous bracts and bracteoles are also present in T.
acutifolius (Abbiatti, 1946, fig. 16A). Palynological
features are strongly suggestive of T. acutifolius and
T. flagellaris (Feuer & Kuijt, 1980), the densely
packed equatorial rods or shorter sculpturing ele¬
ments of those species, respectively, here seemingly
replaced bv a low rugulate-verrucate surface.

Para types. VENEZUELA. Antioquia: Mf rio. Belmira,
cam i no Belmira-San Jose de la Montana, finca El Paramo,
camino a Labores-San Jose, 3130 in, 6°35'N, 75°32'W,
Roldan et al. 2.175 (HU A, LEA); Finca FI Paramo, as the
type. Fonnegra et al. 5400 (HU A, LEA).

Acknowledgments. JK wishes to acknowledge
the continuing financial support of the Natural Sci¬
ences and Engineering Research Council of Can¬
ada. We are obliged to David Eye and Brent Dowen
for assistance with the photographs.

Literature Cited

Abbiatti, I). 1946. Las Lorantaeeas Argentinas. Revista
\1us. La Plata. Secc. Rot.. N.S. 7: 1-1 10.

Barlow, B. A. & I). Wiens. 1973. The classification of the
generic segregates of Phrygilanthus (= A otanthera) of
the Loranthaceae. Brittonia 25: 26—39.

Feuer, S. M. & J. Kuijt. 1980. Fine structure of mistletoe
pollen. III. Large-flowered neotropical Loranthaceae
and their Australian relatives. Amer. J. Bot. 67: 34—50.
Kuijt. J. 1963. On the ecology and parasitism of the Costa
Rican tree mistletoe, Gaidendron punc latum (Ruiz &
Pavrtn) G. Don. Canad. J. Bot. 41: 927—938.

- . 1988. Revision of Tristerix (Loranthaceae). Syst.

Bot. Monogr. 19: 1-61.

A New Species of Tibouchina (Melastornataceae) from
Minas Gerais, Brazil

Rosalia Romero

Inslituto <le Biologia, Universidade Federal de Uberlandia, Caixa Postal 593,
CEP 38400-902, Uberlandia, Minas Gerais, Brazil, rornero@inbio.ufu.br

Paulo Jose Fernandes Guimaraes

Jardim Botanieo do Rio de Janeiro, Rua Pacheco Leao 915, CEP 22460-030,
Rio de Janeiro, Brazil, paulo@jbrj.gov.br

ABSTRACT. The new species Tibouchina rubro-
bracteata belongs lo section Pleroma I). Don. It is
distinguished from related species by the presence
of larger braeteoles than those usually found in sec¬
tion Pleroma, glabrous staminal filaments and style,
the hypanthium not constricted at the apex, and a
globose fruit. This species occurs in highland
“campo rupestre” vegetation in the Serra da Gan-
astra National Park in the state of Minas Gerais, an
important center of endemism in Brazil.

Kk.su MO. A especie nova Tibouchina rubrobrac-
teata pertence a se^ao Pleroma I). Don. fi distinta
das espeeies proximas pela presenga de bracteolas
maiores que o usualmente encontrado na segao
Pleroma, filetes e estilete glabros. hipanto nao con-
strito no apice e fruto globoso. Esta especie ocorre
nos campos rupestres do Parque Nacional da Serra
da Canastra, no estado de Minas Gerais, um im-
portante centro de endemismo.

Key words: Brazil, campo rupestre, endemic,
highland vegetation, Melastornataceae, Minas Ger¬
ais, Tibouchina.

The region of Serra da Ganastra is located in the
Gerrados domain and contains many endemic spe¬
cies in different families (Romero & Nakajima,
1999). The vegetation of the region is composed of
forests, including gallery' forests that occur along
creeks and rivers as well as small patches of cloud
and montane forests, savanna with different physi¬
ognomic types, and campos rupestres, a mainly
shrubby vegetation of rocky outcrops. This new
species of Tibouchina was discovered during the
floristic survey carried out by Herbarium Uberlan-
dense (HUFU) from the Universidade Federal of
Uberlandia in the Serra da Ganastra National Park,
southwestern Minas Gerais state, Brazil.

Tibouchina ruhrobracteata B. Romero & P. .1.
F. Guimaraes, sp. nov. TYPE: Brazil. Minas
Gerais: Sao Roque de Minas, Par. Nae. Serra
da Canastra, estrada para o sftio Joao Domin¬
gos, Serra da Canastra, afloramentos roehosos,
46° 1 5 ' -47°00 ' W, 20°00'-20°30'S, 26 May
1996, ,/. N. Nakajima 6c R. Romero 1R07 (ho-
lotype, HUFU; isotypes, K. MO. BB. UEG,
US). Figure 1A-F.

Frutex 0.4—1 m altus. Laminae ovado-oblongae, apice
acuto, base rotundata, margine integro, supra et suhtus
sericeo, 5—7 nervis. Thyrsus 7—12 cm longus, 5—9 flores;
2-bracteolae concavae, 1 1.0-12.0 X 12.0- 14.0 mm; flores
5-meri, pedicellati; hypanthium campanulatum; calyces
lobis 6—8 X 3-4 mm, in fructu non persistentibus; 10
stamina, filamenta glabra; ovarium 5-loculare, apice seri-
ceo-glanduloso; stylus glaber. Capsula globosa.

Shrub, 0.4—1 m tall, the stem, branches, petioles,
leaves, inflorescence, braeteoles, and hypanthium
densely covered with glandular hairs; branches
quadrangular. Petioles 1.2— 3.3 cm long; leaf blade
6.0-10.0 X 3.0— 5.3 cm. ovate-oblong, apex acute,
base rounded, margin entire, above sericeous, cov¬
ered with eglandular and some glandular hairs, be¬
low sericeous, covered with glandular and eglan-
dular hairs, 5- to 7-nerved, with inconspicuous
lateral veins, confluent in the base. Thyrse termi¬
nal, 5 to 9 flowers, 7.0—12.0 cm long; braeteoles 2,
11.0—12.0 X 12.0-14.0 mm, red, asymmetrical,
concave, outer surface densely covered with glan¬
dular hairs, caducous later. Flowers 5-merous, ped¬
icellate, pedicels 1 .8-3.0 mm long; hypanthium
5.5— 8.5 X 6—8.0 mm, eampanuiate, not constricted
at the apex; calyx lobes 6—8 X 3—4 mm, caducous
in the fruit, triangular, apex acute, outer surface
densely covered with glandular hairs, mainly on the
lobe base, sometimes between the lobes with a
glandular hair longer than the others; petals purple,
15.0—17.0 X 10.0—16.0 mm, obovate, apex round¬
ed, glandular-ciliate, with an apical glandular hair

Novon 15: 210-212. 2005.

5 mm

Volume 15, Number 1
2005

Romero & Guimaraes
Tibouchina from Brazil

211

Figure I. Tibouchina rubrobracteata R. Romero & I’. J. F. Guimaraes. — A. Habit. — R. Fruit. — C. Petal. — I).
Bracteole. E. Lateral view of stamens. — K Ovary anil style. Drawn from the holotype, Nakaiinui & Romero IR07
(HUFU).

212

Novon

longer; stamens 10, subisomorphic; filaments 7.0—
8.5 mm long, glabrous; anthers 7.0— 8.0 mm long,
subulate, apex attenuate, connective prolonged
1.7-2. 2 mm below the thecae, with ventral bilohed
appendage; ovary superior, 5-locular, free apical
portion densely covered with glandular hairs; style
14.0-15.0 mm long, filiform, glabrous, stigma trun¬
cate. Capsule 6.7-7. 8 X 6.3-7.0 mm, globose;
seeds 0.5— 0.7 X ca. 0.5 mm, numerous, cochleate,
tubereulate.

Tibouchina rubrobracteata has 5-merous flowers
in a thyrsoid inflorescence, two conspicuous hrac-
teoles 11-12 X 12—14 mm, caducous, calyx lobes
caducous in the fruit, and subulate anthers with
attenuate apex. These characters refer T. rubro¬
bracteata to section Pleroma, but the sectional lim¬
its in Tibouchina are weak since most sections re¬
flect previously recognized genera, and they are not
monophyletie clades on the basis of morphological
characters.

The 11-12 X 12—14 mm bracteoles, a hypan-
thium not constricted at the apex, the capsule glo¬
bose, as well as the filaments and style glabrous,
distinguish T. rubrobracteata from T. carvalhoi Wur-
dack, to which it is most closely related. The latter
differs in having smaller bracteoles, 2-5 X 0.7-
0.75 mm, a hypanthium constricted at the apex,
which is best observed during the development of
the fruit, the capsule elongated, and the filaments
and style with glandular hairs in the basal portion.
Tibouchina carvalhoi is apparently endemic to Cha-
pada Diamantina, Bahia state, brazil. Wurdack
(1083) related T. carvalhoi with T. clidemioides Cog-
niaux. We have examined tin* syntypes of T. clide¬
mioides (in G and K) and can verify that the brac¬
teoles are elongate (7-12 nun long), but much more
narrow (2—3 mm), and the calyx lobes are also very
narrow and, as in T. carvalhoi, the filaments and
style have glandular hairs toward the base. Accord¬
ing to Cogniaux (1883-1885), the type collection is
from Vitoria, Espfrito Santo state, in eastern Brazil.
These three species are strongly related by the in¬
dumentum of the branches, leaves, and shape of
the inflorescence.

Tibouchina rubrobracteata is endemic to the re¬
gion of Serra da Canastra, Minas Gerais, Brazil, on

rocky slopes. This species flowers from April to Au¬
gust. and fruits from May to July.

The specific epithet refers to the characteristi¬
cally red color of the bracteoles. I his red coloration
frequently can also be seen in the inflorescence,
pedicels, hypanthium, and calyx lobes of Tibouch¬
ina rubrobracteata.

Paratypes. BRAZIL. Minas Gerais: Sao Roque de
Minas, Par. Nac. Serra da Canastra, paredao da Caclmeira
Casca d'Anta, 20 Apr. 1994, R. Romero, J. N. Nakajima
A- F. A. G. Guilherme 959 (HUFU); paredao da Caehoeira
Casca d'Anta, 22 Ago. 1994, R. Romero. J. N. Nakajima
I). G. Simao 1 1 SI (ESA, HUFU, SP); .1 km da sede
administrativa, 10 May 1995,/ N. Nakajima, R. Romero.

M. R. Alcantara & C. A. Prado 1012 (HUFU, k). 15 May

1995, J. N. Nakajima. R. Romero & M. R. Alcantara 1107
(C, HUFU), 15 July 1995,/ N. Nakajima, R. Romero, M.
A. Farinaceio & V. F. (). Miranda 2.195 (HUFU, MBM,
SPF); Caehoeira Casca d'Anta, 12 May 1995, / N. Na-
kajima, R. Romero, M. R. Alcantara A- C. A. Prado 1050
(BHCIL HUFU, R); Caehoeira Casca d'Anta, 10 July

1996, ./. N. Nakajima, K. Ressel & F. A. G. Guimardes
1 946 (HUFU, NY); Garagem de Pedras, I 1 July 1996, J.

N. Nakajima, K. Ressel F. A. G. Guimardes 2002
(HUFU, VIC); Delfintipolis, estrada da represa dos Peix-
otos, 22 May 1996. R. Romero & ./. N. Nakajima 5410
(HUFU, NY.’ RR, UKC, US).

Acknowledgments. The first author is grateful to
the Universidade Federal de Uherlandia for the fa¬
cilities of HUFU, the Instituto Brasileiro do Meio
Ambiente c dos Recursos Naturais Renovaveis
(IBAMA) for permission to collect at Serra da Can¬
astra National Park, Wagner Moreira Lima, former
director of Serra da Canastra National Park, for lo¬
gistical and facilities support, and the Conselho Na-
cional de Desenvolvimento Cientifico e Tecnologico
(CNPq Grant N° 300918/93-3) for a research grant.
Fieldwork for this study had financial support from
the Diretoria de Pesquisa from the Universidade
Federal de Uherlandia. The authors thank Peter
Gibbs for helpful corrections and Eduardo Kick-
hofel for preparing the illustration.

Literature Cited

Cogniaux, A. 1883-1885. Melastomataceae. Tribus Mi-
crolicieae e Tibouchinieae. Pp. 1—510 in C. F. P. de
Martius & A. G. Fielder (editors). Flora Brasiliensis,
Vol. 14, Parte 3. Fried. Fleischer, Lipsiae.

Romero, R. & J. N. Nakajima. 1999. Ksp^eies endemieas
do Parque Nat ional da Serra da Canastra, Minas Ger¬
ais. Revisla Brasil. But. 22 (2-suplemento).

Wurdack, J. J. 1983. Certamen Melastomataceis XXXVI.
Phytologia 53: 121-137.

Heteroflorum : Un Nuevo Genero del Grupo Peltophorum
(Leguminosae: Gaesalpinioideae: Caesalpinieae),
Endemico para Mexico

Mario Sousa S.

Departamento de Botanica, Instiluto de Biologfa, Universidad Nacional Autonoma de
Mexico, Apdo. Postal 70-367, 04510 Mexico, D.F. Mexico, sousa @servidor.unam.mx

Rksumkn. Sc describe e i lustra a un nuevo ge-
nero monotfpico, arboreo y dioico, endemico a Mex¬
ico: Heteroflorum de las Leguminosae, Caesalpi-
nioideae, Caesalpinieae, grupo Peltophorum,
incluyendo la especie nueva II. sclerocarpum de la
Cuenca liaja del Balsas y de la Planicie Costera
del Pacifico Mexicano. Se le relaciona estrecha-
mente eon Conzattia y Peltophorum.

Abstract. A new monotypic arboreal and dioe¬
cious genus, endemic to Mexico, is described and
illustrated: Heteroflorum, within the Leguminosae,
Gaesalpinioideae, Caesalpinieae, Peltophorum spe¬
cies group, including the new species H. sclerocar¬
pum from the Balsas River basin and the Mexican
coastal plains ol the Pacific Ocean. The new taxon
is closely related to Conzattia and Peltophorum.

Key words: Caesalpinieae, Gaesalpinioideae,
endemic, Leguminosae, Mexico, Peltophorum
group.

Mexico es considerado un centro seeundario de
diversification de Leguminosae con 26 tribus, 135
generos y 1724 espeeies (Sousa & Delgado, 1993);
hoy dla tenemos informat ion de 139 generos y
1850 espeeies (Sousa et al., 2001, 2003). Kespecto
al endemismo el 51.9% tie las espeeies lo son, y a
nivel tie genero se cuenta con cinco (3.6%), en las
Ires subfamilias, Gaesalpinioideae, Caesalpinieae:
Conzattia Rose y Heteroflorum ; Mimosoideae, Mi-
moseae: Calliandropsis H. M. Hernandez e lngeae:
Guinetia L. Rico & M. Sousa; Papilionoideae, Mil-
lettieae: Hesperothamnus Brandegee.

Heteroflorum pertenece a la tribu Caesalpinieae
tlentro tie la caracterizacion planteada por Cowan
(1981) y Polhill y Vidal (1981). Dentro de la trilm
Caesalpinieae pertenece al grupo Peltophorum pm-
puesto por Polhill y Vitlal (1981), el cual. se sugie-
re un cambio de circunscripeion por Lewis y Schri-
re (1995) pasando a Conzattia del grupo
Caesalpinia al grupo Peltophorum. Haston el al.
(2003) sugieren emplear los marcadores trn L intron
y /rnL-F del cloroplasto para delimitar al grupo Pel¬

tophorum incluyendo a 16 generos, siendo cl grupo
de tlistribucion pantropical.

Heteroflorum M. Sousa, gen. nov. TIPO: Hetero¬
florum sclerocarpum M. Sousa.

Arhores inermes; plantae per minutionem dioeciae. Fo¬
lia bipinnata, stipulis acicularibus, mox caducis; pinnis
o[)f>ositis, loliolis alternis. Inflorescentiae racemosae, ra-
cemis axillaribus, bractea florali ad l)asem pedicelli. Flo-
res flavi. pedicellati, pedicello articulato, sine bracteolis,
ad basem floralem hypanthium formantes; flores staminati
quam pistil lati majores; calyx gamosepalus, 5-lobatus,
lol)is integris; corolla 5 petalis distinctis, plus minusve
isomorphis, el I i pticis vel obovatis, sine unguibus; stamina
It), libera, lilamentis ad basem pilosis, antheris versatili-
lius; ovarium sessile, multiovulatum; stylo tubulari, apice
lato facto, piloso; stigmate circulari, lato. applanato vel
leviter concavo. Legumen indehiscens, cylindricum, lig-
noso-scleroideum, laeve, dissepimentis fibroso-spongiosis;
semina discoidea vel subsphaerica, h i lo apicali; plantulae
epigaeae, eophyllis alternis, 1-pinnatis, paripinnatis.

Arboles inermes, el tronco con proyecciones co-
nicas suberosas no espinosas. Hojas en disposit ion
helit •oitlal, bipinnadas; estlpulas aciculares, pronto
deciduas; pinnas opuestas motleradamente nume-
rosas, reducidas la basal y la apical; foliolos nu-
merosos, alternos. Inllorescencias racemosas, raci-
mos axilares, bractea floral presente y subyacente
al pedicelo. Plantas dioicas por reduccion; (lores
amarillas. polinizadas por abejas, pediceladas, pe-
dicelo articulado ± de la mitad al tercio superior,
sin bracteolas, flores de 2 tamanos, las estaminadas
de mayor tamano que las pistiladas. la base de la
flor desarrolla hipantio; caliz gamosepalo, 5-lobu-
lado, lobulos enteros; corola polipetala, 5-petalos,
± homomorfos, elipticos a obovados, sin urias; es-
tambres 10 libres, exertos. los lilamentos pilosos en
la base, insertados en el apice del hipantio; anteras
versatiles, longituilinalmente dehiscentes hacia a-
tlentro; ovario sesil, 9- 1 3-ovulado. eslilo tubular,
ensanchado en el apice, piloso; estigma circular,
ancho, aplanatlo a ligeramente concavo. Legumbre
indehiscente, cillndrica, lenoso-esclerotiea, lisa,
glabra, con tabiques fibroso-esponjosos entre las
semi lias formando cavidades seminfferas; semillas

Novon 15: 213-218. 2005.

214

Novon

discoides a subesfericas, eon tegumento duro, hilo
apical. Plcintulas epigeas; eotiledones coriaceos; eo-
filos alternos, 1-pinnados, paripinnados.

Hetcroflorum soleroearpuin M. Sousa, sp. nov.
TIPO: Mexico. Micboaean: l.a lluaeana, a I I
km al 0-S0 de La lluaeana, 300 m, 18 Jill.
1979, Mario Sousa S. 10683, R. Sousa P., i.
Guevara F., I,. Ortiz Arias, A. Morelos Borja &
,/. Ih'az de la Cruz (holotipo, MEXU; isotipos,
PM. MEXU, MO). Figura I.

Arbor 3.5—15 in alia, foliis caducis, caule projecturis
conicis 3-lobatis, suberosis, non spinosis, cortice laevi,
plumbeo, cicatricibus foliaribus in lineis horizontalibus
elevatis disposilis; rami fistulosi, sulcati, pilosiusculi, mox
glabri, lenticellis aurantiacis numerosis, loliis spiraliter
disposilis. Stipulae 7-9 nun longae, aciculares, patentes,
mox caduoae; folia 14-32 cm longa, bipinnata, pinnis (2-)
4-7, (.3.5— )5-6.5(— 12) cm longis, in foliis immaturis op-
positis, postea basin versus oppositis, apicaliter alternis,
basali cl apicali reductis vel parum reductis, irregulariter
imparipinnatis, 21-27 foliolis in quaque pinna; petiolus
4—10 cm longus, eglandulosus, canaliculatus, sulcatus, pi-
losiusculus, mox glaber; rhacbidi foliari 5— 1 5( — 23) cm
longa, canaliculata, pilosa, in appendice f) — H nun longa,
acieulari, mox caduca terininata; foliolis 1—2.4 X 0.7— 1.1
cm, alternis, elliplicis vel leviter oblongis, aliquantum bi-
coloribus, utrinque pilosis, base parum asymmetricis, a p i -
ce obtusis, mucronalis. Inflorcsccntiae racemosae, racemis
axillaribus, 4— 18(— 32) cm longis; bractea Horalis stami-
nata 4—5 nun longa, aciculari-filiformis, mox caduca; brac¬
tea Horalis pistillata 4—7 mm longa, angusle elliptica vel
anguste lanceolata, aliquanto persistens. Flores staminati
12—14 nun longi, bypantbio 2—3 mm longo; calycis loliis
5, 5—6 mm longis, ligulatis; petala Hava, 9-10 X 4—5 mm,
glabra, adaxialiter maculis aurantiacis; stamina 10. 9—10
mm longa, filamentis basi canescenti-pilosis, aliter gla-
bris, antheris 0.8—1 mm longis; gynoecium 2.5—3 mm Ion-
gum, moderate canescenti-pilosum. Flores pistillati 7 — H( —
1 1) mm longi. hypanthio 1.7— 2( — 3) mm longo; calycis lob-
is 5, 3.5— 4.5(-8) mm longis, ligulatis; petala flava. 6— 7(—
10) X 2.7— 3(— 5) mm, glabra; staminodia 10, 5—7 mm lon¬
ga, filamentis basi moderate vel sparse pilosis, aliter
glabris, antheris 0.7— 0.9 mm longis; gynoecium 6— 7(— 12)
mm longum. Legumen (12 — )14 — 18( — 21) cm longum, 2.3—
2.7 cm latum, indehiseens, lignoso-seleroideum, teres vel
leviter applanatum, rectum vel lev iter arcuatum, inter
semina aliquantum constrietum, laeve, glabrum; semina
fi — 1 2 in quoque fructu, 8.3—10.5 X 8. ,3— I I X 6—8 mm.
discoidea vel subsphaerica, laevia, niti<la, atrobrunnea.
duobus annulis pellueidis marginalibus.

Arboles 3.5—15 m alto, eadueifolios, inermes, el
troneo eon proyecciones ednieas 3-lobadas, su-
berosas no espinosas; eorteza lisa, pardo-plomiza.
con cicatrices foliares en lineas horizontales y len-
tieelas ligerarnente realzadas; ranias frecuente-
mente fistulosas, suleadas, pi Insulas, pronto gla-
bras, eon abundantes lentieelas anaranjadas, las
hojas dispuestas helieoidalmente. Estfpulas 7—9
mm largo, aciculares, patentes, pronto eaducas; ho¬
jas 14—32 cm largo, bipinnadas, las pinnas (2— )4—

7, (3.5— )5— 6.5(— 12) cm largo, en hojas inmaduras
opuestas, posteriormente opuestas haeia la base, al-
ternas apiealmente, la basal y la apical reducidas
a ligerarnente reducidas, irregularmente imparipin-
nadas, 21—27 folfolos por pinna; peefolo 4—10 cm
largo, eglandular, canaliculado, sulcado, pildsulo.
pronto glabro; raquis foliar 5— 15(— 23) cm largo,
canaliculado, pildsulo, rematando en tin apendiee

6— 8 mm largo, aeicular, pronto eaduco; folfolos 1 —
2.4 X 0. 7-1.1 cm, alternos, elfpticos a ligerarnente
oblongos, algo bicoloros, pildsulos en el haz y el
enves, la base ligerarnente asimetrica, id apice ob-
tuso, mucronado. Inflorescencias racemosas, los ra-
cimos axilares, 4— 18(— 32) cm largo; brdctea floral
estaminada, 4-5 mm largo, acicular-fili forme, pron¬
to caduca, bractea floral pistilada, 4—7 mm largo,
angostamente elfptica a angostamente lanceolada,
algo persistente. Flores estaminadas, 12—14 mm
largo; hipantio 2—3 mm largo; Idlnilos del caliz. 5,

5— 6 mm largo, ligulados, patentes, esparcidamente
pildsulos; petalos amarillos, 9-10 X 4—5 mm. gla-
bros, id interior (estandarte) con maeulas anaran¬
jadas; estambres 10, 9-10 mm largo, los lilamenlos
canescente-pilosos en la base, sino glabros. las an-
teras 0.8—1 mm largo; gineeeo 2.5—3 mm largo, mo-
deradamente canescente-piloso. Flores pistiladas,

7 — 8( — 11) mm largo; hipantio 1 .7 — 2( — 3) mm largo;
ldbulos del caliz 5. 3.5-4.5(-8) mm largo, ligula¬
dos, patentes, pildsulos; petalos amarillos, 6— 7(— 10)
mm largo, 2.7— 3(— 5) mm ant ho, glabros; estami-
nodios 10, 5—7 mm largo, los lilamenlos moderada
a esparcidamente pilosos en la base, sino glabros.
las anteras 0. 7-0.9 mm largo; gineeeo 6 — 7 ( — 12)
mm largo, densamente piloso en la base, esparci¬
damente en el estilo. Legumbre ( 1 2 — ) 1 4 — 1 8(— 2 1 ) X
2.3— 2.7 cm, indehiscente, lenoso-esc derdtica, rolli-
za a ligerarnente aplanada, recta a ligerarnente ar-
queada, algo constricta entre las semillas. lisa, gla¬
bra; semillas f) — 1 2 por fruto, 8.3—10.5 X 8.3—1 I X

6— 8 mm. discoides a subesfericas. lisas, lustrosas,
castano oscuro, con hordes elaros, formando dos
anillos. hilo apical, tegumento grueso y duro.

Nombres vulgares. “Ojo de venado" (./. L. Con¬
treras ./. 2381. MEXU) [por el parecido superficial
de sus semillas a las de Mucuna ] (Guerrero); “lri-
jolillo" (M. Chazaro 748. MEXU), “ojo de venado"
(,/. C. Soto 3524, MEXU), “teyapo" (,/. C. Soto
1410, I ). Ramos, MEXU), “vainilla" (.S. Rangel-
lAimla 540. MEXU) (Michoacan), “maravilldn del
cerro" (M. Florsa C. 6003, MEXU) (Oaxaca).

Usos. Forrajera (las hojas), la madera no tiene
tisos, dada su poca compactividad.

Distribueidn v habitats. Se trata de una especie
de la cuenca baja del Rio Balsas y de la planicie

Volume 15, Number 1
2005

Sousa

Heteroflorum : Un Nuevo Genero

215

l igura I. Heteroflorum sclerocarpum M. Sousa. — A. Proyecci6n cdnica del tronco. — B. Rama con inflorescencia
estaminada (tornados del holotipo en MEXU, Sousa et al. 10683). — C. Rama con inflorescencias pistiladas. — 1). Flor
pistilada (tornados de Soto et al. 1420). — E. Flor estaminada (tomada de Sousa et al. 10683). — F. Fruto. — G. Semilla.
— 11. Planlula (tornados do Toledo s.n.).

5cm

216

Novon

costera del Pacffico (Kig. 2). En altitudes entre 15-
600 m. Kn vegetacidn primaria de selvas bajas a
tnedianas caducifolias, con Cyrtocarpa procera
Kunth, eactaceas columnares, burseras, legumino-
sas, Ziziphus Miller v Cordia I.. Kiparia en rfos de
temporal y permanentes. Sobre suelos someros eon
grandes roeas sedimentarias, en suelos arenosos,
aluviales y tepetatosos.

Sus {lores son visitadas por abejas, “mayates”
hembras de Xylocopa ( J . C. Solo 1420. I). Homos ,
MKXU). Sus frutos son dispersados por corrientes
de agua.

Fenologfa. Florece de finales de abril a finales
de junio, fructifica, con frutos maduros de septiem-
bre a diciembre.

Dentro del grupo Peltophorum, Haston el al.
(2003) delimitan a un grupo monofiletieo central de
ocho generos, c|ue comprende a Schizolobium Vo¬
gel, Bussea Harms, Peltophorum (Vogel) Benlham.
Parkinsonio L„ Conzottia, Delonix Rafinesque, Col¬
ei Ilea Bojer y Lemuropisum Perrier. I)e ellos He¬
teroflorum tiene mas afinidad con Conzottia y Pel¬
tophorum eon los que morfologicamente constiluye
un subgrupo y comparten los siguientes caracteres:

Arboles inermes. Hojas en disposition helicoi-
dal, bipinnadas; estfpulas aciculares, pronto deci-
duas; pinnas opuestas; foliolos numerosos.

Infloresceneias raeemosas. brdetea floral presente y
subyacentc al pedicelo; (lores amarillas. poliniza-
das por abejas, pedieeladas, el pedicelo articulado,
sin braeleolas. la base de la dor desarrolla hipantio;
caliz gamosepalo, 5-lobulado, los Idbulos enteros;
corola pt)l ipetala, 5-pelalos; estambres 10 libres,
exertos. los (ilamentos pilosos en la base, insertados
en el apice del hipantio; anteras Versailles. longi-
tudinalmente dehiscentes hacia adentro. Plantulas
epigeas; cotiledones coriaeeos; eofilos alternos, 1-
pitmados, paripinnados.

En la invesligacidn en progreso Sotuyo (com.
pers.) ha elaborado una filogenia preliminar de la
tribu Caesalpinieae haciendo enfasis en Coesalpi-
nia E. y generos cereanos, empleando cuatro mar-
eadores trn\.. /ral.-K. trn\\-psbA y Acc D-p.su I del
cloroplasto, y eneontrd <]ue Heteroflorum es el ge-
nero hermano de Conzottia. y que el genero Pel¬
tophorum es cercano.

Comparando a Heteroflorum eon Conzottia y Pel¬
tophorum (vease Tabla 1) se puede apreciar que a
[>esar de los varios caracteres que comparten, He¬
teroflorum es marcadamente diferente sobre todo a
nivel de (lores, frutos y semillas.

Paratipos. MKXKX). Mielioaean: Arteaga, Cerea la
Presa de Kl Infiemillo, M. Chdzaro it. 74H (MKXU); 26
km N of Presa Kl Infiemillo. C. P. Hughes 1 849 (MKXU);

Volume 15, Number 1
2005

Sousa

Heteroflorurrr. Un Nuevo Genero

217

Table 1.

Comparacion entre Heterofiorum, Conzattia

y Peltophorum.

Heterofiorum

Conzattia

Peltophorum

Foliolos

alternos

opuestos

opuestos

Mores de dos tamanos

si

no

no

Lobulos del

eai iz

patentes, aplanados

reflexos, cuculados

reflexos, cuculados a aplana¬
dos

Petalos

homoinorfos

heteromorfos

heteromorfos

Unas de los

petalos

ansentes

presentes

presentes

Estilo

tubular robusto, se ensancha
en el apice

hb forme, no se ensanch:
el apice

a en filiforme, no se ensancha en
el apice

Estigma

ancho, circular aplanado, li-
geramente edneavo

eapitado

ancho, aplanado v peltado

l.egumbre

indehiscente, cilfndrica

dehiscente, aplanada

indehiscente, aplanada

1 iispersion

por corrientes de agua

por viento

por viento y probablemente
por agua

Semillas

discoidales a subesfericas

ovadas a lanceoladas

lanceoladas

Hilo

apical

subapical

subapical

14 km SW de la desv. a K1 Infiernillo, earn La Presa,
Carr. Nueva Italia-Playa Azul, ./. C. Soto V 1086, I. Ro¬
man dr Soto (MEXU. MO); cerros eercanos al poblado do
LI Infiernillo, Soto 1649, G. Silva R. (MEXU); 18 km SW
de la desv. a El Infiernillo, earn Arteaga— Nueva llalia, M.
Sousa 8050, ,/. C Soto A L. Sousa P. (MEXU); 8.2 km by
road NE of Puerto San Salvador, along Mex.. V. W. Stein-
mann 2422. G. Puime A' R. Vrskovy (MEXU); El Infier¬
nillo, solo frutos y semillas fueron colectados, V. M. Toledo
s.n . (MEXU). Churumuco, 92 km de La Huacana, en la
brecha a Uuetamo, E. ./. Lott 2254, 2256, E. Martinez S.
(MLXU); 18 km W de San Jeronimo, camino a Churu-
mueo, E. Martinez S. 1445, ./. C. Soto N. (MEXU); Las
Cliieas, 15 km NW' de San Jeronimo, Soto 1419, I). Ramos
7. (MEXU); El Pitayo, 17 km NW de San Jeronimo, Soto
1420, Ramos (MEXU); 1 km NE de Nuevo Churumuco,
Soto 8524 (MEXU). La Huacana, 3 a 6 km SW de La
Huacana, La Huacana, Soto 1442, 2283, Roman de Soto
(MEXU); El Varillo, 6 km SE de La Huacana, earn a
Inguaran, Soto 3501 (MEXU). Tumbiscatfo, El Caulote, 2
km SE de I .as Cruces, Soto 3610 (MEXU). Guerrero:
Coahuayutla de Guerrero, I km NE de Matamoros, ./. Ca-
Idnico Soto 15173 (ECME, MLXU); San Jose de Anota,

15 km N de Coahuayutla, camino Coahuayutla— Matamo¬
ros, ./. /.. Contreras ./. 2374 (MEXU); 1 km S de Coahua¬
yutla, camino a La Union, Contreras 2379 (MEXU); El
Mezquital, 6 km S de Coahuayutla, por el camino a La
Union, Contreras 2381 (MEXl ); El Infiernillo, 4 km S on
the E bank of the Rio Raisas, close to junction of track
lo Rarranca San Diego, C. E. Hughes 1845 (MEXU). Pe-
tatlan. Petatlan, 2 km SE, C. I). Johnson 1265-79
(MEXU). Oaxaca: Poehutla, Miguel del Puerto, earn ha-
cia Santa Marfa Xadani. 2 km N de Zimatan, M. Elorsa
(.. 238 (MEXU); Rancho La Escalera, 15.6 km N de Pu¬
ente Zimatan. ,/. Rivera II. 2513 (MEXU); Rancho La Es¬
calera. A. Savnes L 2202, 2203. 2204, M. Elorsa, N. le-
Idsquez, E. Martinez S., M. Sousa S. A A. Reyes (MEXU);
Copalita, 15 km NE de Santa Cruz-Huatulco, /. Trejo
2713, I1. Tenorio L. & R. Dirzo (MEXU). Tehuantepec, San
Pedro Huamelula, por San Isidro Chacalapa, 5 km N de
la earn Costera, Elorsa 600 (MEXU); 3 km N de Ayntla,
camino a Guadalupe Victoria, E. Martinez S. 32145, M.
Elosa C. & C. Ferret (MEXU); Playa Grande, 500 m E,

camino Laguna Agua Grande, Rivera 2563 (MEXU); 4 km
S de Chacalapa, .S'. Salas M. 1773, Rivera (MEXU); San
Isidro Chacalapa, 5.5 km S, San Pedro Huamelula, Salas
2922 (MEXU); Camino a Laguna Mazealeo, 500 m S, Sa¬
las 2962 (MEXU); earn a San Isidro Chaealpa, 5 km N
de la cam fed.. A. Sanchez A. 68, .S'. Salas M.. ./. Rivera
H. & M. Elorsa C. (MEXU). Santiago Astata, camino de
la vereda hacia Zimatan a 3 km de Rarra de la Cruz al
NW. Elorsa 263 (MEXU); por Marcalco a 2 km S de la
cam, Elorsa 456 (MEXU); Barra de la Cruz, 4 km E,
Elorsa 3027 (MEXU); 5 km la desv. a Chacalapa. por la
cam de Santiago Astata-San Pedro Poehutla, G. Flores
Franco 3662 (MEXU); Camino a Mazealeo, Saynes 2056,
S. Salas M. A M. Elorsa C. (MEXU); 5 km N de la desv.
a Chacalapa, la desv. 38 km SW de Astata. earn Salina
Cruz-Poclmtla, R. Torres C. 5224, C. Avendano A ./. Mar¬
tinez (MEXU).

Agradecimientos. A Jeny Solange Sotuyo V. por
compartir informat ion inedita y por revisar el ma-
nuscrito; a Fernando Chiang C. por tradueir al La¬
tin las diagnosis; a Gloria Andrade M. por resolver
problemas del texto y recabar informacion; a Silvia
Salas M. y Jose Carmen Soto N. por efectuar nu-
merosas colectas ex profeso tanto en Oaxaca como
en Michoacan respectivamente; a Linda Ellis por
su magmfica lamina botanica y a Gabriela Sanchez
C. por elaborar el mapa.

Literature Citada

Cowan, R. S. 1981. Caesalpinioideae. Pp. 57—64 en R. M.
Polhill & P II. Raven (editores). Advances in Legume
Systematics, Pt. I. Royal Botanic Gardens, Kew.
Hasten. E. M., G. P. Lewis & J. A. Hawkins. 2003. A
phylogenetic investigation of the Peltophorum group
(Caesalpinieae: Leguminosae). Pp. 149—159 en B. B.
Klitgaard & A. Bruneau (editores). Advances in Legume
Systematics, Pi. 10. Royal Botanic Gardens. Kew.
Lewis, G. P. & B. I). Schrire. 1995. A reappraisal ol the
Caesalpinia group (Caesalpinioideae: Caesalpinieae)

218

Novon

using phylogenetic analysis. Pp. 41-52 en M. I). Crisp
& J. J. Doyle (editores), Advances in Legume Systeni-
atics, Pt. 7. Royal Botanic Gardens, Lew.

Polhill. R. M. & J. E. Vidal. 1981. Caesalpinieae. Pp. 81-
95 en R. M. Polhill & P. H. Raven (editores). Advances
in Legume Systematics, Pt. I. Royal Botanic Gardens,
Kew.

Sousa S., M. & A. Delgado S. 1993. Mexican Legumino-
sae: Phytogeography, endemism and origins. Pp. 459-

51 I en T. P. Ramamoorthy, R. Rye, A. Lot & J. Fa
(editores). Biological Diversity of Mexico: Origins and
Distributions. Oxford Univ. Press, New York.

- , M. Ricker & H. M. Hernandez. 2001. Tree spe¬
cies of the family Leguminosae in Mexico. Harvard Pap.
Rot. 6: 339—365.

- , - & - . 2003. An index for the tree

species of the family Leguminosae in Mexico. Harvard
Pap. Rot. 7: 381-398.

Euphorbia derickii (Euphorbiaceae), a New Species from

Central Mexico

Victor W. Steinmann

Instituto de Ecologfa, A.C., Centro Regional del Bajio, Ad’. 386, 61600 Patzcuaro,

Michoacan, Mexico

Abstract. Euphorbia derickii, a new species in
Euphorbia sul>g. Chamaesyce (Euphorbiaceae), is
described and illustrated. It is endemic to central
Mexico and presently known from 1 1 widely sep¬
arated collections in the states of Guanajuato,
Guerrero, Michoacan, and Morelos. Although sim¬
ilar to Euphorbia liirta and E. lineata in most fea¬
tures, it can be distinguished from these and all
other Mexican species of Euphorbia subg. Cha¬
maesyce by its seeds, which possess sharp, well-
defined, uninterrupted, transverse ribs.

Resumen. Se describe e ilustra Euphorbia deri¬
ckii, una especie nueva de Euphorbia subg. Cha¬
maesyce (Euphorbiaceae). La especie nueva es en-
demica del centro de Mexico y conocida de once
eolectas de los estados de Guanajuato, Guerrero,
Michoacan y Morelos. Es semejante a Euphorbia
liirta y E. lineata pero difiere de el las y las otras
especies mexieanas de Euphorbia subg. Chamae¬
syce por tener semillas con costillas transversales
notarias, continuas y con el borde agudo.

Key words: Euphorbia, Euphorbiaceae, Mexico.

Mexico is the center of diversity for Euphorbia
L. subg. Chamaesyce Rafinesque (— Chamaesyce
Gray), and I estimate that it contains ea. 1 10 of the
nearly 300 species of this subgenus. Its members
are well represented throughout the country in al¬
most all types of vegetation from sea level to over
3000 m. During die preparation of the family Eu¬
phorbiaceae for the Mexican Clara del Bajio y de
Regiones Adyacentes, various collections of a dis¬
tinctive taxon were encountered from both within
and outside the boundaries of the Flora. Because
these plants do not appear to belong to any previ¬
ously described species, they are here proposed as
new.

Euphorbia derickii V. W. Steinmann, sp. nov.
TYPE: Mexico. Morelos: mpio. Miacatlan, eji-
do Coateteleo, 10 km al E de Mazatepee, 14
abr. 1986, Vdzquez 45 (holotype, IEB; isotype,
ENCB). Figure I.

Planta annua, ereeta vel ascendens, usque 35 cm alia;
caules teretes; internodia 1 — 14 cm longa; folia opposita,
petioli 1—1 mm longi, laminae ovatae, oblongae vel obov-
atae, 0.5— 2.9 cm longae, 0.4— 1.6 cm latae. apices obtusi,
bases cuneatae vel hemicordatae, saepe ohliquae, margi-
ues serrulati vel crenulati; cyatliia in capitulis densis ter-
minalibus; pedunculi 0.3— 1.8 mm longi, glabri; involucra
infundibuliformia vel campanulata, 1.1 — 1.6 mm longa,
0.8— 1.0 mm diametro; glandulae 4, eireulares vel trans¬
verse oblongae, 0.1— 0.2 mm longae (radialiter), 0.1— 0.3
mm latae (tangentialiter), concavae; appendices flabella-
tae, 0.2— 0.9 mm longae, 0.3— 0.8 mm latae, margines in-
tegri; llores staminati ca. 20—30; ovarium 3-lobatum; styli
3, 0.3 mm longi, bifidi, filiformes vel leviter clavati, ad
apices leviter dilatati; capsula 3-lobata, 1.2— 1.6 mm lon¬
ga, 1.3— 1.8 mm lata; columella 0.9— 1.2 mm longa; semina
ovoideo-quadrangulata, 0.9— 1.0 mm longa, 0.6— 0.7 mm
lata, superficies dorsales costis transversis argutis contin-
uis 6—8.

Plants annual, erect or ascending, to 35 cm tall:
root slender, sometimes diffusely branched; primary
stems single or many arising from the base, little
branched below but much branched distally. terete,
0.8-4 mm thick, internodes 1—14 cm long, strigu-
lose or rarely pilose, sometimes glabrate, trichomes
0.2— 0.4 mm long, almost straight to slightly re¬
curved, white to light brown or yellow. Leaves op¬
posite, stipules interpetiolar, separate, undivided or
laeiniate into 3 to 5 segments, subulate to almost
filiform, 0.8— 2.4 mm long, glabrous to pilose, punc-
tiform glands ca. 0.1 mm diam. sometimes present
at the base of the stipules, petioles 1—4 mm long,
strigulose, blades ovate to oblong or rarely obovate,
0.5— 2.9 cm long, 0.4— 1.6 cm wide, sometimes with
a small lobe toward the base of one side and/or a
red spot in the center, apex obtuse, base euneate
to hemicordate, frequently oblique, with a promi¬
nent midvein and 2 to 4 less-distinct lateral veins
emerging from the base, glabrate above and sparse¬
ly strigulose below, margin irregularly serrulate to
crenulate. Cyathia in dense, terminal, leafless ea-
pitula, peduncles 0.3— 1.8 mm long, glabrous. In¬
volucre funnelform to campanulate, 1. 1-1.6 mm
long excluding the glands, 0.8—1 .0 mm diam. below
the glands, glabrous below, strigulose above, lobes
rectangular to triangular, 0.2— 0.3 mm long, apex
acute to truncate, glands 4, slightly stipilate, cir-

Novon 15; 219-221. 2005.

220

Novon

cular to transversely oblong, 0.1— 0.2 mm long (ra¬
dially), 0. 1-0.3 mm wide (tangentially), concave,
appendages flabellate, sometimes asymmetrical
with one side prolonged, 0.2— 0.9 mm long, 0.3— 0.0
mm wide, entire, white and sometimes becoming

purple-pink in age. Staminate flowers ± 20 to 30.
Gynophore glabrous, exserted 0.5— 1.1 mm, ovary
strigulose, 3-lobed, styles 3, divided to the base,
0.3 mm long, filiform or slightly elavate, subcapi-
late. Capsules broadly ovate and widest below the

Volume 15, Number 1
2005

Steinmann

Euphorbia derickii from Central Mexico

221

middle, 1.2-1. 6 mm long, 1.3-1 .8 mm diam., 3-
lobed, shortly strigulose, shallowly furrowed be¬
tween the carpels; columella 0.9— 1.2 mm long.
Seeds ovoid-quadrangular, 0.9-1. 0 nun long, 0.6-
0.7 mm wide, orange-brown, subacute to obtuse at
the apex, obtuse to truncate at the base, the dorsal
facets with 6 to 8 well-defined, sharp, narrow trans¬
verse ribs that do not pass through the dorsal keel,
these generally uninterrupted but rarely anastom-
izing.

Distribution and habitat. Euphorbia derickii is
endemic to central Mexico and currently known
from 1 I widely separated collections in the states
of Guanajuato, Guerrero, Miehoacan, and Morelos
at elevations of 600 to I 750 m. Four collections are
reported to have been made in cultivated fields.
None of the remaining specimens provide specific
information about the surrounding vegetation ex¬
cept one that is reported from an “open riparian
area.” However, based on the general vegetation of
the localities where it has been encountered, it
probably occurs iu subtropical scrub and tropical
deciduous forest.

Conservation status. At four of the known lo¬
calities, this taxon was reported to be weedy in cul¬
tivated fields, and at two others it was described as
abundant. Due to its tendency to invade disturbed
habitats, it probably is not in danger ol extinction
at present.

Phenology. This species apparently is repro¬
ductive throughout the year under favorable con¬
ditions; it has been collected with flowers and fruits
in July, August, October, January, February, March,
and April.

Etymology. Euphorbia derickii is named in
honor of my father, Derick Otis Steinmann (1943-
2002), to whom I owe not only my existence but
also many invaluable lessons about life.

Discussion. As noted by Wheeler (1911) and
McVaugh (1961), the most useful distinguishing
traits in many groups of Euphorbia subg. Chamae-
syce are those of the capsules, seeds, and pubes¬

cence. For Euphorbia derickii , the seeds are most
characteristic. In other features, this species strong¬
ly resembles both E. hirta L. and E. lincata S. Wat¬
son, and all of the collections were previously iden¬
tified as one of these taxa. However, the sharp,
well-defined, uninterrupted, transverse ribs of the
seeds are unmistakable and serve to distinguish it
from both of these species, whose seeds are rugu-
lose or with subtle, mostly rounded transverse ridg¬
es. In fact, the seeds of E. derickii are so distinctive
that they readily separate it from other North Amer¬
ican and Mexican species of this subgenus.

Like most Mexican members of Euphorbia subg.
Chamaesyce, the common name of Euphorbia der¬
ickii is Golondrina (Rodriguez I did, Torres 414).
Another common name is Verdolaga Cimarrona
(Torres 414).

Paratypes. MEXICO. Guanajuato: mpio. Yuriria,
Yuriria, margenes de la laguna, 3 sep. 1931, Zizumbo Z-
815 (I KB), Z-816 (I KB). Guerrero: mpio. de Coyuea, Co-
yuea, 25 Jan. 1934, Hinton el al. 5544 (K); km 12 car-
retera Chilpancingo, 22 ago. 1974, Rodriguez 1313
(EINCB, MEXU). Miehoaean: Coenqueno, Yurec uaro, 20
feb. 1982, Mores 52 (IBUG). Morelos: Alpuyeca, 18 ago.
1949, Atmar 77 (MEXU); Tlaltizapan, Centro Internacion-
al de Mejoramente de Maiz y Trigo, 3 Sep. 1978, litis et
al. 831c (MICH, WIS); Miaoatlan. Miranda 1282 (MEXU);
mpio. Coatlan del Hfo, Coatlan del Rio, 27 jul. 1976,
Torres 414 (MEXU); Mazatepec, 2 mar. 1968, Vdzquez
1499 (MEXU).

Acknowledgments. For access to their collec¬
tions, I am indebted to the curators and staffs at
IBIJG, IEB, K, MEXU, MICH, and WIS. I thank
Rogelio Cardenas for preparing the illustration.
Economic support was provided by a grant from the
Comision Nat ional para el Conoeimienlo y Uso de
la Biodiversidad to the Institute de Eeologia, A.C.
(account number 902-03).

I .iteralure Cited

McVaugh, R. 1961. Euphorbiaceae Novae Novo-Galici-
anae. Brittonia 13; 145—205.

Wheeler, E. (7 1941. Euphorbia subgenus Chamaesyce in
Canada and the United Slates exclusive of southern
Florida. Rhodora 43: 97-154, 168-205, 223-286;
plates 654—668.

Fourteen New Species of Gonolobus (Apocynaceae,
Asclepiadoideae) from Mexico and Central America

W. I). Stevens

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

stevens@mol)ot.org

Abstract. Fourteen new species of the genus
Gonolobus (Apocynaceae, Asclepiadoideae) from
Mexico and Central America are described and il¬
lustrated. Gonolobus asterias, G. hammelii, G. trun-
catifolius , G. ustulatus, and G. variabilis are endem¬
ic to Costa Rica. Gonolobus grayumii is from Costa
Rica and adjacent Panama and G. hadrostemma is
from Costa Rica and adjacent Nicaragua. Gonolobus
> dbiflorus , G. croceus, and G. sandersii are from
western Mexico. Gonolobus pallidas is from south¬
ern Mexico and G. exannulatus is from southern
Mexico and Guatemala. Gonolobus ancoriferus is
found in southeastern Mexico and Honduras and G.
cuajayote is found in southern Mexico, El Salvador,
and Nicaragua.

Kksiimkn. Se describen e ilustran catorce nuevas
especies <1**1 genero Gonolobus (Apocynaceae, As¬
clepiadoideae) de Mexico y Centroamerica. Gono¬
lobus asterias , G. hammelii, G. truncat if olios, G. us¬
tulatus, y G. variabilis son especies endemieas de
Costa Rica. Gonolobus grayumii es de Costa Rica
y Panama y G. hadrostemma es de Costa Rica y
Nicaragua. Gonolobus albi floras, G. croceus, y G.
sandersii son del oeste de Mexico. Gonolobus pal¬
lidas os del sur de Mexico y G. exannulatus es del
sur de Mexico y Guatemala. Gonolobus ancoriferus
se encuentra en el sureste de Mexico y en Hon¬
duras y G. cuajayote se encuentra en el sur de Mex¬
ico, El Salvador y Nicaragua.

Key words: Apocynaceae, Asclepiadoideae,
Gonolobus, Mesoamerica, Mexico.

Gonolobus is a genus of about 150 species rang¬
ing from the southern U.S.A. to Argentina and the
Caribbean, with one species introduced in western
tropical Africa. The greatest diversity is centered
in southern Mexico and Central America, where a
number of species remain to he described. I his
contribution continues the effort (Stevens & Men¬
del. 2002, 2004) to account for the known but mi-
described species, now about half completed. To

facilitate the description of individual species, a
description of the genus is provided.

Gonolobus Michaux

Twining ( dextrorsum externe visas) herbaceous or
woody vines, corky bark absent from woody stems,
stems nearly glabrous to variously pubescent, hairs
of uniform length or more often a mixture of long
hairs and short and/or short capitate-glandular
hairs, indumentum of internodes uniform or ar¬
ranged in I or 2 lint's beginning between leaf axils
and ending at leaf axils; latex white; roots fibrous.
Leaves opposite, without pseudostipules, petiolate,
with colleters at adaxial union of blade and petiole,
lateral veins pinnate. Inflorescence extra-axillary, 1
per node or rarely with a second inflorescence
borne on an axillary short shoot, racemiform or pan-
ulieuliform; calyx divided nearly to base, normally
with colleters within below the sinuses; corolla ro¬
tate. eampanulate with a rotate limb, or occasion¬
ally tubular, aestivation imbricate and dextrorse,
mostly with a faucal annulus around corona, hairs
of inside of corolla unicellular; androecium and gy-
nostegium entirely fused and forming a gynoste-
gium; corona gynostegial, of 5 largely connate
lobes, adnate to base of gynostegium and to corolla;
anthers indistinct, completely adnate to style apex,
normally with a laminar or fleshy appendage on dis¬
tal-dorsal margin, terminal appendages appressed
to margin of style apex, guide rails linear, often
indistinct, ± vertical or tilted under base of gynos¬
tegium; poll inaria ± horizontal, along anther mar¬
gin. corpusculiim mostly sagittate*, translator prom¬
inently translucent-winged, pollinia symmetrically
or mostly asymmetrically obovoid, rarely cylindri¬
cal, with proximal part sterile and ± excavated;
style apex pentagonal or ± star-shaped, shallowly
concave, smooth or rugose. Follicles 1 or 2, base
asymmetrical, borne at 90°— 180° to pedicel, mostly
with 5 complete w ings, occasionally w ith fewer and/
or incomplete wings or rarely without wings, smooth
to tuberculate; seeds numerous, obovate, flat, with
a prominent margin, eomose.

Nov ON 15: 222-244. 2005.

Volume 15, Number 1
2005

Stevens 223

Gonolobus from Mexico and Central America

figure I. Gonolobus albiflorus W. I). Stevens. — A. Flowering branch. — B. Flower. — C. Gynostegium. — I). I’olli-
narium. — h. Diagnostic longitudinal section of flower. Drawn from Stevens et al. 2554 (MO).

1. Gonolobus albiflorus W. I). Stevens, sp. nov.
TYPh: Mexico. Jalisco: Mpio. Tuxpan, lower F
slope ol Nevada de Colima, ea. I mi. above
Fresnito, 28 July 1976, IF I). Stevens , M. ./.
Dorwgh ue & M. L. Scott 2554 (holotype, MO;
isotypes, KNCB, MSC). Figure I.

G onolobo gonoloboidi affinis sed floribus rninoribus
tubo corollae angustiore gynostegio exserto.

Perennial parts unknown, young stems densely
white-pilose, hairs spreading to somewhat reflexed.
0.3- 1.1 mm long, internodes 10—15 cm. Leal
blades ovate-el liplie. 5.4— 8.5 X 1 .8-3.8 cm, apex
attenuate, base lobate, lobes descending, sinus
0.4— 0.8 em deep, ± densely glassy-pilose, hairs

0.4-1. 0 mm long, lateral veins 4 to 6 pairs, middle
veins 45 —55° to midrib, eolleters 2 to 4; petiole
1.7-4 em, densely white-pilose. Inflorescence con-
gested-racemiform to paniculiform (twice-
branched), 1- to 40-flowered, densely white-pilose,
peduncle 2-1 I mm, axis 0.5-1 1 mm, pedicel 9-
17.5 mm, bracts 1.4-2. 5 X 0. 1-0.2 mm. linear;
calyx tube 0.5-1. 5 mm long, with 1 colleter per
sinus, lobes elliptic to debate with acute to atten¬
uate tips, 4.5— 6.4 X 1 — 1.4 mm, green or tinted
purple, pilose outside, glabrous inside except lip
pilose; corolla tubular, constricted in middle (above
corona), uniformly white or tinted purple at base
outside, lube 6—8.8 mm long, pilose outside with
hairs 0. 3-0.4 mm long except glabrous al base, in-

224

Novon

side white-velutinous with re flexed hairs 0.4— ().8
mm long except base glabrous, faucal annulus ab-
sent, lobes debate to lanceolate with acute to at¬
tenuate tips, 9.5-12.5 X 2. 6-3. 4 mm, reflexed, pi-
lost' outside with hairs 0.3-0.4 mm long, inside
glabrous except with curly, white hairs ca. 0.1 mm
long in a dense fringe along the proximal half or
two-thirds of one side and in a small triangle at
base; gynostegium with stipe 5.5— 6.3 X 0.6— 1.3
mm, prominently 5-winged, each wing with a tooth
near top, corona an erect, fluted, glabrous ring 1.2—
1.5 mm long, adnate for most of length to corolla,
anther appendages ± deflexed, spatlmlate, concave
above, 1-1.6 X 0.9-1. 6 mm in outline, fleshy, pur¬
ple, terminal appendages tightly appressed to style
apex, 0. 1-0.2 X 1.2- 1.4 mm, purple, guide rails
straight, parallel, indistinct, 0.6— 1.5 X 0.1— 0.2
mm; corpusculum 0.3—0.39 X 0.1 1—0.13 mm, sag¬
ittate. dark brown, translators 0.35—0.38 X 0.1 5 —
0. 18 mm, pollinia 1.02—1.08 X 0.3— 0.33 mm; style
apex 2.S-3.4 mm wide, yellow-green. Fruits and
seeds unknown.

This species belongs, with Gonolobas gonoloboi-
des (Greenrnan) Woodson and G. arizonicus (A.
Gray) Woodson, in Gonolobus subg. Pseudolachnos-
loma Woodson, characterized by campanulate to tu¬
bular, white corollas without faucal annuli, rela¬
tively short coronas that do not line the inside of
the corolla tube, and gynostegia with long stipes.
This new species is easly distinguished from the
other two by relatively narrower, reflexed corolla
lobes, a relatively narrower corolla tube, and an
exserted gynostegium. Gonolobus albijloriis seems
to be restricted to a small area of the eastern slopes
of the Nevado de Colima, Mpio. Tuxpan, Jalisco,
Mexico, between 2000 and 2100 m elevation. It is
known to flower in July and August.

Pnratypes. MKXICO. Jalisco: Mpio. Tuxpan, I I km
al 0 de la carr. Atenquique— Colima, brecha al Playon, ./.
Villa C„ S. I ). Korh A' ./. Chavez L 86/ (CHAPA, MO);
antenas de microondas de Cerro Alto, I km al S de Atenq¬
uique, earr. a Colima, luego 14 km al 0 por brecha al
Volcdn de Colima y al Cerro Alto, M. hientes (). 547
(CHAPA, MO).

2. Gonolobus ancoriferus W. I). Stevens, sp.
nov. TYl’K; Honduras. Yoro: 0.9 km F, of San
Jos6 Texiguat by road, then 3.5 km up Rio
Texigua, IN of Cerro Cabeza de Negro,
15°29'N, 87°27'W. 200-300 m, 22 Apr. 1994,
A. E. Grant & I ). Hazlett 2904 (holotype, MO;
isotype, FAR). Figure 2.

Conolobo xanlhotricho affinis, a qua imprimis differt co¬
rona annulo 2— 3plo longiore.

Perennial parts unknown, young stems with
mixed indumentum, denser at the nodes, long hairs
spreading or somewhat reflexed, pale yellow to
brown, 0.5-2. 5 mm long, short and glandular hairs
white to dark brown, 0.05-0.15 mm long, inter¬
nodes 5— 24 cm. Leal blades elliptic' to oblong, 7.1 —
13.6 X 2.fi — 5.7 cm, apex acuminate, base rounded,
truncate or shallowly lobate, lobes divergent to de¬
scending, sinus to 0.4 cm deep, adaxially glabrous
or with sparse, appressed hairs 0.3— 0.4 mm long,
abaxially hispidulous with hairs 0.2— 0.3 mm long,
lateral veins 6 to 9 pairs, middle veins 60°— 70° to
midrib, colleters 2 or 3; petiole 1.1—2./ cm, pilose.
Inflorescence congested-racemiform, 2- to 16-flow-
ered. peduncle 1—9 mm, glabrous or with sparse
mixed indumentum, axis 2—7 mm, pedicel 1 <— 37
mm, with dense mixed indumentum, bracts 1 — 1.5
X 0.2-0. 3 mm, lanceolate or debate; calyx tube
0.4-1 mm long, with I colleter per sinus, lobes
lanceolate with acute tips, 5.3— 6.4 X 2-2.4 mm,
green or with purple or brown margins, abaxially
hispidulose with hairs 0.2-0. 9 mm long, adaxially
glabrous; corolla rotate, green or yellow-green, ap¬
parently sometimes with red or purple markings,
tube 2.2—3 mm long, glabrous, faucal annulus an
erect tube, entire or inconspicuously 5-lobed, 0.5—
l.l mm tall, upper margin with a fringe of erect
white hairs 0.6-1. 5 mm long, corolla lobes ligulate,
with thickened margins, nearly valvate, distally cu-
eullate (tips indexed in bud) and deeply rugose,
apex rounded when flattened but strongly folded
and contorted, glabrous inside, hispidulous outside
with hairs 0.2 mm long or sometimes nearly gla¬
brous, 6-8 X 3.2-4.3 mm, patent; gynostegium
with stipe 1 .5-2. 1 X 0.4— 1.4 mm, smooth or slight¬
ly 5-ribbed, corona fleshy, yellow-orange, bowl¬
shaped, 2.3—3 mm tall, 5-lobed, lobes angular,
slighllv retuse or entire, base adnate to annulus,
anther appendages bifid, stalk patent, 0.9— 1.2 X
0. 6-0.8 mm, arms twisted and arching upward.
1.3-2 X 0.3-0. 5 mm, lips of arms 2.5-4 mm apart,
terminal appendages appressed to style apex, 0.2—
0.3 X I ,Y> — 2 mm, apparently white, guide rails
straight, parallel, indistinct, ea. 0.15 X 0.1 mm;
corpusculum 0.21—0.25 X 0.13—0.15 mm, sagit¬
tate, red-brown, translators 0.28— 0.36 X 0.15—0.19
mm. pollinia 1.04—1.31 X 0.43—0.46 mm, sym¬
metrical or slightly asymmetrical; style apex 2.3-3
mm wide. Fruits and seeds unknown; ovaries 5-
ribbed, glabrous.

Ibis new species is most similar to Gonolobus
xanlhotrichus Brandegee, but most conspicuously is
much less pubescent overall, the corolla lobes are
never barbate within, the dorsal anther appendages

Volume 15 Number 1
2005

Stevens 225

Gonolobus from Mexico and Central America

c

Figure 2. Gonolobus ancoriferus W. I). Stevens. — A. Flowering branch (Hernandez I OHS, MO). — IF Flower (Martinez
& Soto 13643, MO). — O. Gynostegiurn (Martinez & Soto 13643. MO). — I). Pollinarium (Martinez & Soto HU >13, MO).
— F. Diagnostic longitudinal section of Mower.

are longer and thinner, and the corona is two to
three times longer than, rather than shorter than to
equaling, the faueal annulus. This species has a
relatively large distribution but seems to he uncom¬
mon: it has been found from sea level to 700 m
and is known to (lower from March to June.

Paratypes. MFXICO. Veracruz: Mpio. Catemaco,
Playa Escondida, 10 kin N of Sontecomapan, .1/. Nee
26692 (K MO. NA, XAL): Mpio. San Andres Tuxtla, Playa
Balzapote, R. Cedilla T. 3577 (MO). Oaxaca: bpto. Jueh-
itdn, Mpio. Santa Maria de Chimalapa. Juyupac, 2.5 km
al N de Santa Marfa, II. Hernandez G. 1035 (CAS, MO).
Chiapas: Mpio. Ocosingo, El Tumho, camino Chancala—
Monte Efbano, E. Martinez S. M. A. Soto 1. 13643 (F.
MEXU, MO).

3. Gonolobus aslerias W. I). Stevens, sp. nov.
TA PE: Costa Rica. San Jose: basin of El Gen¬
eral, May 1940, A. F. Skutch 49311 (holotype,
US; isotypes, (ill, E, NY). Eigure d.

Amplitudine et forma florum Gonolobo versicolor i simi-
lis. praecipue differt ealyee majore lobis corollae longior-
ibus latioribus attenuatioribus ecarnosis.

Perennial parts unknown, young stems with
mixed indumentum, most densely at the nodes and
in 2 lines on internodes, long hairs ferrugineous,
somewhat reflexed, 0.7— 0.9 mm long, short and
glandular hairs 0.04—0.06 mm long, internodes 4—
8 cm. Leaf blades elliptic, 6.2— 7.2 X d. 9—4.6 cm.

226

Novon

Figure 3. Gonolobus asterias W. I). Stevens. — A. Flowering branch. — B. Flower. — C. Gynostegium from above. —
I). Corona and faucal annulus, from above with gynostegium removed. — E. Gynostegium, corona, and faucal annulus,
from side. — F. Pollinarium. — G. Diagnostic longitudinal section of flower. Drawn from holotype, Skulch 4938 (US).

apex abruptly acuminate, base lobule, lobes de¬
scending, sinus 0.2— 0.4 cm deep, sparsely ap-
pressed-puberulent with hairs 0.1— 0.2 nun long,
lateral veins 5 to 7 pairs, middle veins 55°— 60° to
midrib, colleters 2, black with pale lips; petiole
1 .9— 2.7 cm, with short and glandular hairs, mostly
on adaxial side and distally. Inflorescence congest-
ed-raeemiform, 3- to 4-flowered, glabrous, pedun¬
cle 2—6 mm, axis 3—6 mm, pedicel ea. 45 mm,
bracts 4—1 I X 1.2— 1.8 mm, lanceolate; ealvx tube

ca. 0.5 mm long, with 1 to 2 colleters per sinus,
lobes lanceolate-attenuate with acute tips, 13—16
X 4.5— 5.5 mm, green or tinted purple, glabrous or
appressed-puberulent on distal margin; corolla ro¬
tate, green, glabrous except puberulent on faucal
annulus and in lines from annulus to corolla sinus¬
es, hairs erect, translucent, 0. 1-0.2 mm long, tube
4-4.5 mm long, faucal annulus pentagonal in out¬
line, of 5 erect lobes opposite corolla sinuses con¬
nected bv a thin line, lobes deflate, ca. 0.4 nun tall.

Volume 15, Number 1
2005

Stevens 227

Gonolobus from Mexico and Central America

corolla lobes lanceolate-attenuate with acute tips,
35-40 X 6 — 8 mm, apparently patent; gynostegnuu
essentially sessile, corona a pentagonal disk, 1.3—
1.6 mm wide, densely papillate, deeply rugose ex¬
cel)! 5 smooth patches, each smooth patch with a
raised, fleshy cushion, anther appendages some¬
what deflexed, somewhat bilobed, 0.8— 0.9 X 2—2.2
mm in outline, laminar, terminal appendages tightly
appressed to margin of style apex, ca. 0.3 X 2.7
mm, apparently translucent, guide rails straight,
parallel, indistinct, ca. 0.1 X 0.1 mm, essentially
horizontal under style apex; corpuseulum ca. 0.38
X 0.16 mm, sagittate, pale brown, translators ea.
0.34 X 0.16 mm, pollinia ca. 1.23 X 0.48 mm;
style apex ca. 4.7 mm wide. Fruits and seeds un¬
known.

This is a remarkable new species that in the
years since its only collection, in 1940, has twice
been supplied with manuscript names, but neither
formal description was consummated. In floral mor¬
phology it is closest to Gonolobus versicolor Wood-
son and somewhat less so to G. cteniophorus (S. F.
Blake) Woodson. Williams (1968) associated Gon¬
olobus breedlovei L. 0. Williams and G. longipe-
tiolalus Woodson (= G. salvinii Hemsley) with G.
versicolor but I consider neither to be closely re¬
lated. On the the other hand, there are at least two
other undescribed Mexican species that share the
very long and narrow corolla lobes, more or less
entire dorsal anther appendages, and a discoid,
deeply fimbriate corona.

Gonolobus usterias was found in a well-collected
area of the best-known country in the American
tropics. Since its large flowers would not be easily
overlooked, especially not by the keen Costa Rican
collectors, this species might well be extinct.

4. Gonolobus croceus W. I). Stevens, sp. nov.

TYPE: Mexico. Sinaloa; Colomas. July 1897,

./. N. Rose 1717 (holotype. MO; isotvpes, GH.

US). Figure 4.

Gonolobo jaliscensi prime aspeetu maxime simile, sed
foliis fere glabris floribus 2— 3plo majoribus corolla praeter
annulum hirsutum glabra.

Perennial parts unknown, young stems glabrous
or sparsely tomentose, especially at nodes, hairs
0.1— 0.3 mm long, white or tinted red, internodes
11—16 cm. Leaf blades elliptic, lanceolate or nar¬
rowly obovate, 6.4— 9.4 X 1.7— 2.3 cm, apex acute
to attenuate, base lobate to truncate, lobes descend¬
ing to divergent, sinus to 0.4 cm deep, glabrous or
sparsely pilose on midrib and margin, hairs 0.2—
0.3 nun long, lateral veins indistinct, 5 to 8 pairs,
middle veins 40°— 60° to midrib, colleters 2 to 6;

petiole 1.1— 1.6 cm, glabrous or sparsely pilose. In¬
florescence eongested-racemiform, 1- to 8-flowered,
peduncle 2—1 I mm, glabrous or sparsely pilose in
I line, axis 0.5—3 mm, pedicel fl — 1 6 mm. sparsely
pilose in 1 line, bracts 0.7-1 X 0. 1-0.2 mm, linear
to lanceolate; calyx tube 0.2— 0.5 mm long, with I
to 2 colleters per sinus, lobes elliptic with acute
tips, 2.9—4 X 1.2— 1.4 mm, green or tinted purple,
glabrous or sparsely pilose at base outside and on
margin; corolla tubular in basal half of connate
part, then rotate, yellow (Cowan <£■ Nieves 4785),
drying yellow-green with deep green veins on base
of corolla lobes, tube (connate part of corolla) 2.6—
3.6 mm long (erect, tubular part 1-1.3 mm, reflexed
part 1.6— 2.6 nun), glabrous except on faucal an¬
nulus, faucal annulus erect, somewhat pentagonal
in outline, densely hispid with erect, glassy hairs
0.15—0.4 mm long, lobes elliptic with obtuse tips.
5.5— 7.2 X 3.2— 4.3 mm, reflexed, glabrous outside,
inside glabrous except sparsely white-papillose on
basal half; gynostegium with stipe 0.7— 0.8 X 0.7—
0.8 mm, 5-ribbed, corona an erect, fluted, some¬
what pentagonal, glabrous ring ea. 0.5 mm tall, an¬
ther appendages patent or somewhat deflexed.
broadly spathulate, 0.4-0.7 X 1-1.4 mm in outline,
fleshy, apparently while, terminal appendages tight¬
ly appressed to style apex. 0.2-0. 4 X 1.1-1. 5 mm.
apparently translucent, guide rails straight, paral¬
lel. indistinct, 0.15-0.3 X 0.05-0.13 mm, essen¬
tially horizontal under style apex; corpuseulum
0.17-0.18 X 0.07—0.08 mm, sagittate, dark brown,
translators 0.3—0.35 X 0.17—0.18 mm, pollinia
0.6— 0.7 X 0.22—0.3 mm; style apex 2.3— 2.6 mm
wide, apparently greenish white. Fruits and seeds
unknown.

The known collections of this species are from
tin ■ Pacific foothills of the central coast of Mexico,
at about 900 m elevation; flowers are recorded from
May to July. It is a distinctive species, with rela¬
tively small and narrow, bieolorous, nearly glabrous
leaves and small, nearly glabrous flowers. It shares
the leaf size and shape, small flowers, similar dorsal
anther appendages, and general geographic area
with Gonolobus jalicensis B. L. Robinson & Green-
man. However, that species is found generally fur¬
ther to the south, is densely pubescent throughout,
and has green rather than yellow corollas with the
inner face densely white-hispid.

Paratypes. VI KX ICO. Nayarit: Mpio. San Bias, 3 mi.

K of Jaleacatdn. I). II. Norris N I). ./. Taranto 14148 (MO);
Mpio. Tepic, 17.7 km al 0 de Tepic (ruta 15) sobre ruta
28 (66), 7 km al 0 de Carranza, C. P. Cowan & G. Nieves
II. 4785 (MO. TEX).

228

Novon

c

Figure 4. Gonolobus croceus W. I). Stevens. — A. Flowering branch (Rose 1717 , MO). — B. h lower from above, flattened
(Cowan & Nieves 4783, MO). — C. Flower from side (Norris & Taranto 13138, MO). — I). Gynostegium (Norris
Taranto 13138, MO). — E. Pollinarium (Rose 1717, MO). — F. Diagnostic longitudinal section of flower.

5. Gonolobus cuajayote W. 1). Stevens, sp. nov.
TYPE: Nicaragua. Managua: 2. '3 km from Hwy.
12 on road along ridge of Sierra de Managua
from llwy. 12 at Km 17 to Hwy. 2. 12°04'N,
86°22'W, 450—550 m, 29 July 1977. W. 1). Ste¬
vens 2932 (holotype, MO). Figure 5.

Ex Conolobo nigro corolla viridi pedunculis brevioribus
annulo saepe ciliato distinguenda.

Herbaceous or woody below, young stems with
mixed indumentum, denser at the nodes and in 2
lines on internodes, long hairs absent or restricted

to nodes, spreading or somewhat reflexed, [tale yel¬
low', 0.3— 0.7 mm long, short hairs white. 0.05—0.1
mm long, glandular hairs pale brown, ea. 0.05 mm
long, internodes 2—15 cm. Leaf blades elliptic to
ovate, 3.9—12.1 X 1.9—7 cm, apex acuminate, base
lobate, lobes divergent to convergent, sinus 0.2—2
cm deep, adaxially glabrous to sparsely pilosulose
with long hairs erect, 0.4— 0.6 mm long, and short
and glandular hairs on veins, abaxially with sparse
mixed indumentum, lateral veins 5 to 7 pairs, mid¬
dle veins 30°— 40° to midrib, colleters 2 to 4; petiole
2.8— 9.8 cm, with sparse mixed indumentum. Inflo-

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Stevens 229

Gonolobus from Mexico and Central America

Hgure 5. Gonolobus cuajayote W. I). Stevens. — A. Flowering branch ( Stevens 2932, MO). — B. Flower from side
(Stevens 2932, MO). — C. Pollinarium (Villacorta 537, MO). — I). Diagnostic longitudinal section of flower.

rescence I or rarely 2 per node, racemiform, 12-
to 22-llowered, with mixed indumentum, peduncle
(0— )15-22(— 45) mm, axis 2-17 mm, pedicel 9—24
mm. bracts 1.1— 2.2 X 0.2— 0.4 mm, linear or del-
tate; calyx tube 0.7- 1.7 mm long, with 1 eolleter
per sinus, lobes deflate with acute to rounded lips,
2.3—4 X 1.3— 2.2 mm, bright green, abaxially gla¬
brous or with sparse mixed indumentum on proxi¬
mal half, adaxially glabrous; corolla rotate, deep
green, abaxially densely hispidulose with short and
glandular hairs 0.05—0.1 mm long, adaxially (ex¬
cept annulus) glabrous, tube 1 .2-2.4 mm long, fau-
cal annulus a membranous, erect lip. inconspicu¬
ously 5-lobed, 0.5— 0.7 mm tall, upper margin
glabrous or with a thin fringe of erect white hairs
0.2— 0.3 nun long, corolla lobes lanceolate to ellip¬

tic, apex acute to obtuse or rounded, with conspic¬
uous to inconspicuous auricles at the sinuses, one
margin of each lobe with a thin, dull white callus,
4—7 X 3—4.2 mm, strongly reflexed; gynostegium
with stipe 0.4— 0.7 X 0.7— 1.3 mm, 5-winged, each
wing with a small, fleshy projection near top, co¬
rona lleshy, yellow-orange, bowl-shaped, 5-lobed,
0.4— 0.8 mm tall, 1.3— 1.8 mm wide at lobes, 0.7—
1.4 mm wide at sinuses, anther appendages de-
llexed, reniform to nearly spathulate, somewhat
fleshy, smooth, orange, 1-1.2 X 0.9— 1.2 mm, ter¬
minal appendages appressed to margin of style
apex, 0.2— 0.5 X 1.8— 2.4 mm, white, guide rails
indistinct, ca. 0.2 X 0.1 mm; corpusculum 0.22—
0.28 X 0.07—0.1 mm, sagittate, tip slightly hooked,
red-brown, translators 0.22-0.34 X 0.12-0.15 mm.

230

Novon

pollinia 1.02-1.2 X 0.32-0.44 mm. symmetrical or
slightly asymmetrical; style apex 3—3.6 mm wide.
Follicles ellipsoid, apex obtuse, base abruptly nar¬
rowed, doubled back against follicle body, ca. 10
X 5 cm, with 2 complete wings 3-4) mm wide and
3 distal wing fragments, glabrous; seeds obovate,
7—0 X 4.3—4.55 mm. brown with brown-black ra¬
dial streaks, margin 0.5-0. 6 mm wide, erenulate on
distal half, surface smooth, coma 2—3 cm long,
white.

This new species is most similar to Gonolobus
niger (Cavanilles) 11. brown ex Schultes, ranging
from northeastern Mexico to Guatemala, and 6.
roeanus L. O. Williams, endemic to Guatemala.
Gonolobus roeanus differs from the other two most
prominently by having a thick, fleshy faucal an¬
nulus (vs. membranous), calyx lobes exceeding the
corolla lobes (vs. about half the length), and the
corolla internally papillate (vs. smooth and gla¬
brous). Gonolobus cuajayote is more similar to G.
niger but differs most conspicuously in having
green corollas with a white callus along one side ol
each lobe rather than deep purple-black corollas
with concolorous calluses, shorter peduncles, and
a faucal annulus that is sometimes ciliate at the
top, rather than always glabrous. Gonolobus cuajay¬
ote has winged follicles, while those of G. niger are
unwinged, although the follicles of both are gen¬
erally similar in size and shape; the follicles of G.
roeanus are unknown. Gonolobus niger is most often
found on limestone, while G. cuajayote is appar¬
ently restricted to soils of recent volcanic cinders.
Furthermore, in the southern (Mesoamerican) part
of the range of Gonolobus niger it has been found
at elevations of 750 to 1700 m, while G. cuajayote
occurs up to 000 m, with only two collection above
455 m. The species flowers from July to October
and the single mature fruit was collected in March.
In Flora tie Nicaragua (Stevens et ah, 2001: 251),
this species was described as “ Gonolobus sp. C.”

Most, if not all, species of Gonolobus have fruits
that are eaten when young, either raw or cooked.
Most of the recorded local names are some ortho¬
graphic variant of cuajayote, talayote, or cucham-
per, and one of the common forms is borrowed for
the epithet of this species (erf. Greater et ah, 2000,
Art. 23.2).

Paratypes. MKX1CO. Chiapas: Mpio. Cintalapa, 5
km W of Rizo de Oro. I ). E. Breedlove 36686 (CAS); Mpio.
Ksperanza, Contua, E. Matilda 18342 ( F ). FI. SALVA¬
DOR. Ahuuchupan: I’.N. El Impossible, Finca San Al¬
fonso, R. Villacorta. R. Toledo & R. Ixipez 853 (H, LAGIJ,
MO). La l.ibertud: Volcan Caldera, A. Monro, J. Monter-
ossa & R. Carballo 3786 (MO); Quezaltepeque, Rio Sucio,
FI Platanillo, R. Villacorta 587 (R, I.AGU, MO). NICA¬

RAGUA. Cliiiwindegu: Volcan San Cristobal, Hacienda
Fas Rojas, A. Grijalva. I). Soza II. M. Herndndez 3652
(MO); Los Ralcones, 9 km de Somotillo, carr. a Cineo
Pinos, I1. F. Moreno 11509 (M0); 4-1 km al SK de San
Pedro de Potrero Grande, Los Faureles, P. P. Moreno
11709 (M0); Carr, entre Cineo Pinos y San Francisco, R.
Rueda & R. Dolmas 1180 (HUFF. MO). Leon: S base of
Vole dii Teliea, above Ouimera. //. II. litis. R. h. Benz &
A. Grijalva 30813 (MO, \\ IS). Managua: Reserva FI Clio-
coyero. Sierra de Managua, IF I). Stevens cC 0. M. Montiel
26520 (HNMN, MO).

6. Gonolobus exannulaius W. I). Stevens, sp.
nov. TYPE: Mexico. Chiapas: Mpio. Angel Al¬
bino Corzo, between Finca Cuxtepeque and
Finca Cabanas, 1100 m, 9 May 1983, I). h.
Breedlove & M. Bourell 6 7424 (holotype,
CAS). Figure 6.

Species corolla alba omnino glabra rotata exannulata a
congeneris distinguenda.

Perennial parts unknown, young stems with
mixed indumentum ± concentrated in 2 lines, long
hairs patent to reflexed, pale yellow, 0.4— 1.2 mm
long, short and glandular hairs 0.03-0.15 mm long,
internodes 1-18 cm. Leaf blades elliptic or some¬
what ovate, 6.4— 8.3 X 3. 3-5. 4 cm, apex acuminate,
base lobate, lobes descending to diverging, sinus
0.3-0. 8 cm deep, adaxially glabrous or with sparse
appressed or erect hairs 0.2— 0.4 mm long, abaxially
with erect hairs 0.2-0. 4 mm long, mostly on the
nerves, lateral veins 5 to 6 pairs, middle veins 35°-
40° to midrib, colleters 4; petiole 2.2— 3.2 cm, with
mixed indumentum. Inflorescence congested-race-
miform, 2- to 12-flowered, with mixed indumentum,
long hairs sparse, peduncle 3—12 mm, axis to 5
mm, pedicel 12—19 mm. bracts 1.2— 1.5 X 0.2— 0.4
mm. lanceolate; calyx tube 1-1.5 mm long, with 1
colleter per sinus, lobes elliptic to lanceolate with
obtuse to rounded tips, 5.2— 7.5 X 3.1— 3.5 mm,
green, glabrous or with a few hairs on abaxial base,
patent with recurved tips; corolla very briefly eam-
panulate at base and then rotate, white or pale
green, entirely glabrous, tube 3. 1—3.5 mm long,
faucal annulus absent, lobes ovate with obtuse to
rounded tips, 9.5-12.5 X 5.2-4). 1 mm. patent with
recurved lips, gynostegium with stipe ca. 1 .3 X 2.5
mm, 5-winged, corona an erect, fleshy, 5-lobed
bowl, with 5 radial ridges continuous with wings of
stipe, 1.2— 1.5 mm tall at lobes, 0.8-1 mm tall at
sinuses, anther appendages somewhat deflexed, ob¬
long to rounded-trapezoidal and retuse, fleshy,
somewhat concave, 1.9— 2.2 X 2.8— 3.4 nun in out¬
line, terminal appendages tightly appressed to style
apex, 0. 1-0.2 X 2. 2-2. 4 mm, apparently translu¬
cent, guide rails straight, parallel, 0.8—1 X 0.1— 0.2
mm, nearly vertical; eorpuseulum 0.25—0.26 X

Volume 15, Number 1
2005

Stevens 231

Gonolobus from Mexico and Central America

Figure 6. Gonolobus exunnulatus W. I). Stevens,
up of flowers.

A. Scan of holotype (Breedlove <£ Bourell 67424 , CAS). — It. Close

0.1 1—0.13 mm. sagittate, brown, translators 0.25—
0.34 X 0.18—0.19 mm, pollinia 1.53—1.65 X 0.5—
0.51 mm; style apex 3.7—1 mm wide. Fruits and
seeds unknown, ovaries glabrous and 2-rihbed.

This new species, apparently known from four
specimens, has an unusual set of characters. There
arc only a few species of Gonolobus with white or
nearly white flowers, only a few that entirely lack
a laucal annulus, only a few with an entirely gla¬

brous corolla, and none other than this species with
a combination of all three. The dark-colored anther
appendages and corona contrast strikingly with the*
corolla. It has been collected at between 750 and
1500 m elevation and in flower in March and May.

Paratypes. GUATEMALA. Guatemala: near Finca
La Aurora,./. /. Aguilar 251 (F). San Marcus: Finca FI
Porvenir, ./. A. Steyermarh 52510 (F). Suchitcpequez:
near Pueblo Nuevo, P. C. Standley 66625 (F).

232

Novon

Kigure 7. Gorwlobus gruyumii W. I). Stevens. - — A. Scan of holotype (Hanvut’l & (>rayutn 14096, MO). H. Close-
up of flowers.

7. (ronolobus grayumii W. I). Slev<'iis. sp. nov.
TYPE: ( ]<>sta Rica. Heredia: just S of Vara
Rlanca on rot id to San Miguel, 10°09'N,
84°10'W, 1900 m, 2 July 1985, B. E. Hummel
& M. II. Grayum 141)96 (holotype, MO; iso¬
type, CK). Eigure 7.

Species notahilis appendicibus dorsalibus antherarum
inflatis a speeiebus milii cognitis bene distineta.

Perennial parts unknown, apparently somewhat
woody, older stems glabrescent, lentieellate, young
stems with mixed indumentum, long hairs spread¬
ing to somewhat reflexed, white or pale yellow. 0.5—

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2005

Stevens 233

Gonolobus from Mexico and Central America

1.5 mm long, short hairs 0. 1-0.2 mm long, inter¬
nodes 1—30 cm. Leaf blades ovate or elliptic, 7.1 —
I 1.7 X 3.2— 7.5 cm, apex acuminate, base truncate
to lobate, lobes descending to convergent, sinus to
1.5 cm deep, pilose, hairs 0.5— 1.2 mm long, lateral
veins 5 to 8 [tails, middle veins 40°— 50° to midrib,
colleters 4 to 6; petiole 2.8— 7.5 cm, with mixed
indumentum, short hairs sometimes absent. Inflo¬
rescence congested-paniculiform, nearly umbelli-
form or interrupted, 5- to 13-flowered, with mixed
indumentum, short hairs sometimes absent, pedun¬
cle 4—77 mm, pedicel 6—36 mm, bracts 0.8— 2.5 X
0.5— 0.6 mm, debate; calyx tube ca. 1.2 mm long,
with 0 or 1 colleter per sinus, lobes lanceolate with
attenuate tips, 10—10.5 X 2.5—3 mm, green, his-
pidulose, hairs outside 0.4— 0.6 mm long, hairs in¬
side 0.1— 0.2 mm long; corolla short-campanulate
then rotate, green, hispidulose outside with hairs
0.2— 0.3 nun long, inside hispidulose, hairs brown,
0.2— 0.3 mm long proximally, becoming shorter dis-
tally, tube ca. 3.5 mm long, laucal annulus an erect,
5-lobed lip, with the same indumentum as the ad¬
jacent tube, lobes ca. I mm tall, corolla lobes ovate
to elliptic, eucullate, rugose, with obtuse to asym¬
metrically acuminate tips. 7.5— 7.7 X 4.9— 5.1 mm,
reflexed; gynostegium with stipe ca. I X 1.5 mm,
5-ribbed at tip. 5-winged at base, corona a thin, 5-
lohed disk with 5 radial wings continuing from
stipe wings, ea. 1.5 mm wide at lobes, 1 mm wide
at sinuses, adnate for most ol length to corolla, an¬
ther appendages sharply deflexed. fleshy, spathu-
late when flattened, distal-lateral margins thick¬
ened, concave between, proximal margins thin,
overlapping with proximal margins of adjacent ap¬
pendages and covering corpuseula, 1.5— 1.7 X 2.8—
3 mm in outline, apparently purple, terminal ap¬
pendages loosely overlapping style apex, ca. 0.2 X
1.8 mm, guide rails vertical, straight, parallel, ca.
0.5 X 0.1 mm; corpusculum ca. 0.38 X 0.23 mm,
sagittate, dark brown, translators ca. 0.25 X 0.19
mm. pollinia ca. I X 0.3 mm, sinuately obovate;
style apex ca. 2.8 mm wide. Follicles ovoid, apex
acute, base broad, at a right angle to follicle body,
9.5—13 X 5—7 cm, with 2 complete or nearly com¬
plete wings 0.7—1 cm wide and I or 2 smaller wing
fragments, glabrous, minutely luberculate; seeds
obovate, 8.5 — 8.7 X 4.4—5 mm, pale brown with
darker brown streaks, margin 0.6— 0.8 mm wide,
crenulate on distal two-thirds, surface inconspicu¬
ously verrueose, coma ca. 3 cm long, white.

I he dorsal anther appendages of this species are
unlike any yet recorded (or the genus; they are ap¬
parently inflated in natural condition and dry with
the appearance of crinkled varnish. This is exactly

the appearance of the dorsal appendages of F is¬
churia DC., but iu that genus the dorsal appendages
have grown to die point ol subsuming the terminal
anther appendages, while the terminal appendages
of Gonolobus grayumii are still apparent, although
thicker and more loosely attached than normal,
furthermore, Fischeria lias abundant glandular
hairs and smooth follicles, while Gonolobus gra¬
yumii lacks glandular hairs and has winged folli¬
cles. The first record of this species ( Tonduz 12IB5)
has been in collections for more than a century, but
has only old inflorescences with all flowers fallen
and young inflorescences with immature buds. It is
curious that this original collection has short pe¬
duncles, up to 4 mm long, and more or less truncate
leaf bases, while the two modern collections have
peduncles 21—77 mm long anil deeply lobate leaf
bases, but otherwise the collections match well, in-
eluding the several unique floral characters. The
species has been found between 1390 and 1900 m
elevation and has been collected in flower in May
and July and with mature fruits in January and
March.

far <ily pcs. COSTA RICA. Piintarenas: Buenos Ai¬
res. headwaters of Quebrada Dorora, 9 km NE of Ujarras,
V7. II. Grayum & A. Badilla I (. >268 (MO). Sail Jose: An
bord du Rio Pedregoso au Copey, A. Tonduz 12185 (RR,
P, US). PANAMA. Chiriqm: S side of El llatodel Vulcan,
M. Nee 14141 (MO, WIS).

8. Gonolobus hadrosteiimia W. I). Stevens, sp.
nov. TYPE: Costa Rica. Piintarenas; Monte¬
verdi-, 800 m, 29 Nov. 1980. W. Haber 412
(holotype, MO), f igure 8.

Ex affinilate Gonolobi calycosis et speeierum affinium
corona ampla exserta, ex (',. hammelii floribus minoribus
pilis eorollae brevioribus distinguenda.

Perennial parts unknown, young stems with
sparse mixed indumentum, especially in the nodes,
short hairs spreading, white with brown septa,
0.05—0.1 mm long, glandular hairs 0.05—0.1 mm
long, internodes 10—18 cm. Leaf blades elliptic,
8.5—15.6 X 3—6.4 cm, apex acuminate to attenuate
or occasionally apiculate, base abruptly and incon¬
spicuously cordate, sinus 0.1— 0.3 cm deep, adaxi-
ally glabrous or very sparsely puberulent with hairs
0.1 mm long, abaxially puberulent with hairs 0.2-
0.3 mm long, lateral veins 6 to I I pairs, middle
veins 50°— 60° to midrib, colleters 2 to 4; petiole
LI-3.7 cm, with indumentum of stem. Inflores¬
cence congested-racemiform, 2- to 6-flowered, with
indumentum of stem, peduncle 1—2 mm, axis 0.5—

4 mm, pedicel 7—14 mm, bracts 0.7— 0.9 X 0.3-0. 4
mm, deltaic; calyx tube 0.5— 0.7 mm long, with I
colleter per sinus, lobes narrowly lanceolate to del-

234

Novon

Figure 8. Gonolobus hadrostemma W. I). Stevens. — A. Flowering branch (Moreno 6411, MO). — B. F lower from side
(Moreno 6411, MO). — 0. Details of faucal annulus and corona (Moreno 6411, MO). — I). Pollinariutn (Moreno 6260.
MO).

late, with acute tips. 5.5—10.3 X 0.9— 1.8 mm.
green or tinted purple, puberulent outside, glabrous
inside; corolla rotate, cream or pale purple, distal ly
puberulent outside with hairs 0.1 -0.3 mm long, in¬
side uniformly hispidulose with white hairs 0.05—
0.1 mm long, tube 1-1.5 mm long, faucal annulus
an erect tube, 5-lobed, 0.7-1. 2 mm tall, lobes hor¬
izontal and adnate to the base of the corona, corolla
lobes elliptic, with thickened margins, nearly val-
vate, distally somewhat cucullate (indexed in bud),
apex asymmetrically acute, 7.2— 9.5 X 3.7— 4.7 mm,
reflexed; gynostegium with stipe 2.2— 2.6 X 2.8— 3.2
mm, with 5 wings entirely adnate to corona, corona
lleshy, purple, bowl-shaped, 1.8— 2.5 mm tall, with
5 cucullate lobes alternating with 5 blunt knobs
above stipe wings, base adnate to and largely cov¬

ering the annulus, anther appendages obsolete, re¬
duced to 2 fleshy, marginal ridges, terminal ap¬
pendages loosely overlapping style apex. 0.6— 0.7 X
1.4— 1.6 mm, pale purple, guide rails straight, in¬
distinct; corpuseulum 0.45—0.55 X 0.22—0.35 mm,
sagittate with tip curved outward, red-brown, trans¬
lators 0.3—0.43 X 0.2—0.28 mm. pollinia 1.13-1.4
X 0.55-0.6 mm. nearly symmetrical; style apex 3-
3.7 mm wide. Fruits and seeds unknown; ovaries
5-ribbcd. densely puberulent.

I bis species belongs to the group within Gono-
l obits originally described as Trichostelma Baillon.
The group is characterized by having an erect, tu¬
bular faucal annulus and a corona that is almost
entirely adnate to the inside of the annulus tube

Volume 15 Number 1
2005

Stevens 235

Gonolobus from Mexico and Central America

and adnate to the gynostegium stipe by narrow
wings, creating 5 deep nectaries. Additionally, die
corolla lobes are only slightly overlapping and with
indexed tips in bud, resulting in corolla lobes that
often have thickened margins, more or less sym¬
metrical disposition of the indumentum, and are
distal ly somewhat to distinctly cueullate. The an¬
thers are oriented more to the margin of the gynos¬
tegium than under the style apex and the dorsal
anther appendages that are a unique feature of
Gonolobus are often reduced or even obsolete. This
new species, and the related Gonolobus hammelii
described below, differ from the others in having a
massive corona that is exserted from and largely
obscuring the annulus tube; the annulus is also
more prominently lobed than in the other species
of the group. In Flora tie Nicaragua (Stevens et ah,
2001; 251), this species was described as “ Gono¬
lobus sp. IT"

Paratypes. NICARAGUA. Granada: Volcan Morn-
bacho. arriba y SE de Hacienda Cutirre, P. P. Moreno
6 260 (MO); Volcan Mombacho, Finca San Joaquin, P. P.
Moreno 6 411 (MO). Rivas: Volcan Maderas, P. \lt-
Gillivary 86 (F).

9. Gonolobus liamnielii W. I). Stevens, sp. nov.
TYPE; Costa Rica. Puntarenas: Canton de
Osa, Valle del Terraba, fachada S de Fila Re-
tinta. al N de Palmar Norte. Quebrada Ben¬
jamin, 8°59'N, 83°28'W, 300 m, 13 Dec.
1989, B. E. Hammel & M. //. Grayum 17699
(holotype. MO; isotypes, CR. MO). Figure 9.

Fx affinitate Gonolobi calycosis et specierum alfinium
corona ampla exserta, ex G. hadrostemma floribus inajor-
ibus pilis corollae longioribus distinguenda.

Perennial parts unknown, young stems glabrous
except with very sparse mixed indumentum at the
nodes, short hairs spreading, white with brown sep¬
ta, 0.2 mm long, glandular hairs 0.1 mm long, in¬
ternodes 12-25 cm. Leaf blades elliptic, paler be¬
low, 13.5—16 X 6.3—7 cm. apex acuminate, base
abruptly and inconspicuously cordate, sinus 0.1—
0.2 cm deep, adaxially glabrous, abaxially sparsely
puberulent with hairs 0.1 mm long, lateral veins 10
to 1 1 pairs, middle veins 60°— 70° to midrib, col-
leters 4 to 7, raised on a fleshy pad and somewhat
connate; petiole 2.4— 3.6 cm. glabrous. Inflores¬
cence eongested-racemiform, 2- to 4-flowered, with
mixed indumentum of short and glandular hairs,
peduncle 1-2 mm, axis 1-2 mm, pedicel 18-20
mm, bracts ca. 0.4 X 0.2 mm. deltate; calyx tube
ca. 0.1 mm long, with 2 colleters per sinus, lobes
narrowly lanceolate with acute tips, 7. 5-7. 9 X 1.6-
1.7 mm, green with purple margins, with mixed in¬

dumentum outside, glabrous inside; corolla rotate,
deep purple-black, distally puberulent outside with
appressed hairs 0.1 mm long, inside uniformly his-
pidulose with white hairs 0.5— 0.8 mm long on lube
and annulus and gradually decreasing to 0.2 mm
long at tips of lobes, tube ca. 2.2 mm long, faueal
annulus an erect tube, 5-lobed, 1.4— 2.3 mm tall,
lobes horizontal and adnate to the base of the co¬
rona, corolla lobes elliptic, with thickened margins,
nearly valvate, distally slightly or not at all eueul-
late (slightly indexed in bud), apex acute, 12-12.5
X 6. 1—6.4 mm, reflexed; gynostegium with stipe ca.
0.7 X I mm, 5-ribbed, corona fleshy, purple, bowl¬
shaped. 1.5—3 mm tall, angularly 5-lobed, each
lobe with small tooth directed toward gynostegium.
base adnate to annulus, anther appendages obso¬
lete, reduced to a fleshy pad on base of anther
back, terminal appendages appressed to style apex,
0.3 X 1.8 mm, apparently pale purple, guide rails
indistinct; corpuseulum ca. 0.43 X 0.28 nun. sag¬
ittate with tip curved outward, red-brown, transla¬
tors ca. 0.38 X 0.15 mm, pollinia ca. 1.16 X 0.71
nun. asymmetrical; style apex ca. 3.3 mm wide.
Fruits and seeds unknown; ovaries 5-ribbed, gla¬
brous.

This new species is most closely related and su¬
perficially quite similar to Gonolobus hadrostemma ,
but differs in significant details. The differences in
the size, color, and indumentum of the flowers
could be an artifact of the few known collections,
but the nearly sessile gynostegium and the lack of
the partitions connecting the corona and the gy¬
nostegium stipe are important distinctions.

10, Gonolobus pallidus W. I). Stevens, sp. nov.
TYPE: Mexico. Chiapas: near Laguna Ocotal
Grande, ca. 25—30 km SF of Monte I Abano,
950 m, 12 Aug. 1954, A\ L. Dressier 1655 (ho¬
lotype, NY; isotype, US). Figure 10.

Amplitudine et forma florum Gonolobo versicolori ae-
mulans, differt foliis fere glabris lobis corollae glabrissim-
is ecarnosis.

Perennial parts unknown, young stems with
mixed indumentum, most densely at the nodes and
in 2 lines on internodes, nearly glabrous at mid¬
internode, long hairs pale yellow, sharply reflexed.
0.4— 0.6 mm long, short and glandular hairs 0.05—
0.1 mm long, internodes 3—18 cm. Leaf blades el¬
liptic to obovate, bicolorous, 7.6—12.2 X 2.8—5 cm,
apex acuminate, base acute, obtuse or rounded,
adaxially glabrous or with a few hairs at base of
midrib and margin, abaxially with erect short and
glandular hairs 0.05—0.2 mm long, lateral veins 6
to 10 pairs, middle veins 60°-65° to midrib, col-

236

Novon

Figure; 9. Gonolobus hammelii W. I). Stevens. — A. Flowering branch. — lb Gynostegium, corona, and faucal annulus,
from above. Drawn from holotype ( Hammel & Cray urn I 7699, MO).

leters 2; petiole 0.8—3 cm, with sparse and incon¬
spicuous short and glandular hairs. Inflorescence
racemiform to eongested-racemiform, 3- to 12-flow-
ered, glabrous, peduncle 23—42 nun, axis 0.2— 1.3
mm, pedicel 37—61 mm, bracts 2—3 X 0.2— 0.5 mm,
deltate; calyx tube 0.8-1 mm long, with I colleter
per sinus, lobes narrowly lanceolate with acute tips,
10.7-13.1 X 1.8—2 mm, green, glabrous; corolla
briefly campanulate then rotate, pale green or yel¬
low with a brown or purple tint, glabrous, tube 2.9—
3 nun long, faucal annulus an erect tube, adnatc to

base of corona and only visible as a 5-lobed flange
on corona tube, 1—1.5 mm tall, corolla lobes nar¬
rowly lanceolate, w ith small auricles at the sinuses,
3—40 X 3.3— 4.5 mm, apparently patent; gynoste¬
gium with stipe ohconieal. 1.6— 2.5 X 1—2 mm, 5-
ribhed, corona an erect, 5-lobed, fleshy, obconical
tube, 2.4— 3.5 mm tall from annulus to lobe tips,
1.6—3 mm tall at sinuses, anther appendages sharp¬
ly deflexed, 2-armed, the arms curved outward and
upward to vertical, forming a “W” on back ol an¬
ther, 1.4-1 .5 X 1.4— 1.8 mm in outline in natural

Volume 15. Number 1
2005

Stevens 237

Gonolobus from Mexico and Central America

position, terminal appendages loosely overlapping
margin of style apex, ea. 0.3 X 1.5 mm, apparently
translucent, guide rails straight, parallel. 0.5— 0.6
X 0.2— 0.3 mm: corpusculum 0.34—0.36 X 0.14
mm, sagittate, dark red-brown, translators ca. 0.33

X 0.16 mm, pollinia 0.71—0.73 X 0.34—0.36 mm;
style apex 2.5—3 mm wide. Fruits and seeds un¬
known.

The llower size and shape of this species almost

238

Novon

Figure I I. Gonolobus sandenii W. I). Stevens. — A. Flowering branch. — 15. Flower, from above. — C. Flower, from
side with corolla pulled back. — 1). Pollinarium. — K. Diagnostic longitudinal section of flower. Drawn from Sanders el
al. X522 (MO).

exactly match those of Gonolobus versicolor ; hut oth¬
erwise tin; plants are quite different and the two
species are probably not closely related; G. pallidus
differs in having pale green, bicolored leaves that
lack long hairs, an entirely glabrous corolla that has
flat lobes, and dorsal anther appendages that are 2-
armed rather than entire. The tubular annulus al¬
most entirely adnate to the base of the much longer
tubular corona is a unique character in the genus.

Baratype. MKXICO. Chiapas: Mpio. La Trinilaria, 10
km KINK of Dos Lagos above Santa Klena, D. E. Breedlove
522X2 (CAS).

I I . Gonolobus sandersii W. I). Stevens, sp. nov.
TYPE: Mexico. Colima: Mpio. Comala, Ran¬
cho El Jabalf, around Lago Epazote,
19°26'30"N, 103°40'48"W, 1375 m. 2 Aug.
1991, L. Vdzquez V. V74 (holotype, MO; iso¬
type, UCR). Figure 1 1 .

Species habitu cum Gonolobo jaliscensi optime con-
gruens, sed differt appendieibus dorsalibus antherarum
longioribus gracilioribus.

Perennial parts unknown, young stems with sim¬
ple indumentum, denser at the nodes and ± in 2
lines on internodes, hairs spreading or reflexed.

Volume 15, Number 1
2005

Stevens 239

Gonolobus from Mexico and Central America

white to brown, 0.4— 1.3 mm long, internodes 1—15
cm. Leaf blades elliptic, 5. 7-9.9 X 1 .9^4.4 cm,
apex acuminate to attenuate, base lobate or rarely
truncate, lobes descending to convergent, sinus to
0.6 cm deep, adaxially with sparse erect hairs 0.2—
0.8 mm long, abaxially with brown glandular hairs
ca. 0.1 mm long, especially on the nerves, and oc¬
casionally a few long hairs near base of midrib,
lateral veins 4 to 7 pairs, middle veins 40°— 50° to
midrib, colleters 4 or 6; petiole 2.5— 3.3 cm, with
glandular hairs and often a few long hairs. Inflo¬
rescence congested-racemiform or sometimes inter¬
rupted. 1- to 20-flowered, puberulent with curly,
non-glandular hairs ca. 0.1 mm long, uniformly or
on 1 side, peduncle (2— )9— 52 mm, axis 2—10 mm,
pedicel 15—32 mm, thin and lax, bracts 1.2— 2.2 X
0.2-0. 6 mm. lanceolate or debate; calyx tube 0.4—
0.8 mm long, with 1 colleter per sinus, lobes debate
or ovate with acute to rounded tips, 1.4—2 X 0.9—
1.3 mm, green or purple, abaxially with appressed
hairs 0.1—0.15 mm long, adaxially glabrous; corolla
campanulate, uniformly green, abaxially glabrous,
adaxially pubescent on faucal annulus, tube, and
base of lobes, hairs bright white, 0. 1-0.2 mm long,
hairs of outer lobes of annulus lobes curly, hairs of
inner annulus lobes and corolla lube straight, lube
1.7— 2.4 mm long, faucal annulus of 10 rounded
lobes, lobes opposite corolla lobes erect or ascend¬
ing. outer surface green and sparsely pubescent,
inner surface white and densely pubescent, 1 — 1.6
mm long, alternate lobes sharply indexed, white
and densely pubescent, 0.4— 0.9 mm long, corolla
lobes elliptic with revolute margins and somewhat
twisted, apex acute to rounded, 5.5— 6.3 X 1.8— 2.4
mm, reflexed; gynostegium with stipe 0.4— 0.8 X
0.8— 1.2 mm, 5-ribbed, corona fleshy, white or mar¬
gin sometimes brown, bowl-shaped, 10-lobed, each
lobe with a central ridge, 0.9— 1.2 mm wide at lobes
opposite outer annulus lobes, 0.5— 0.8 mm wide at
alternate lobes, base adnate to annulus, anther ap¬
pendages white, ascending, with a long, thin stalk
and a bifid tip. stalk 1 .5— 3.2 X 0.2— 0.5 mm, arms
divergent to coiled, tip in natural outline 0.4—1 X
0.7— 1.2 mm, terminal appendages appressed to
style apex, 0.15—0.3 X 0.9— 1.3 mm. bright white,
guide rails indistinct, vertical, 0.1— 0.3 X 0.1 mm;
corpusculum 0.14—0.19 X 0.06—0.08 mm, subsa-
giltate, red-brown, translators 0.16—0.24 X 0.09—
0.13 mm, pollinia 0.5—0.64 X 0.18—0.19 mm,
asymmetrical; style apex 1.5—2 mm wide, red-pur¬
ple. Fruits and seeds unknown.

This new species is not apparently allied to any
other. It is superficially most similar to Gonolobus
jaliscensis, especially vegetatively and in the small

flowers with a strongly reflexed green corolla, anti
G. sandersii occurs at similar elevations within the
range of G. jaliscensis, but G. jaliscensis is generally
much more pubescent, including both sides of the
corolla, has shorter and more rigid inflorescences,
green-on-green reticulate corollas, and, most con¬
spicuously, short, bilobed dorsal anther appendag¬
es. The anther appendages of Gonolobus sandersii
are a more prominent part of the flower than any
other species of Gonolobus , and on collectors’ notes
are sometimes called petals.

Paratypes. MEXICO. Colima: Mpio. Comala. Rancho
Jabalf, S and E sides of Lago El Calabozo, A. C. Sunders ,
/. Garcia R., G. Ballmer <V T. Krantz 8522 (MO. UCR), L.
Vazquez V. & B. L. Phillips 880 (MO, UCR), 95/ (MO.
UCR), L. Vazquez V. I0B4 (MO. UCR); Mpio. Comala,
Rancho Jabalf, around l.ago El Epazote, L. Vazquez V. A
B. L. Phillips 725 (MO, UCR), /.. Vazquez V. 1002 (MO,
UCR). 1502 (MO. UCR). Jalisco: Mpio. Teealitlan. 3 km
al E de IJanitos, brecha a Canutillo, J. Villa C. & S. I ).
Koch 777 (CHAPA, MO).

12. Gonolobus truncatifolius W. I). Stevens, sp.
nov. TYPE.: Costa Rica. Limon: Cordillera de
Talamanca, Atlantic slope, Valle de Sileneio,
just N of Cerro Hoffman, 4.5 airline km W of
Panama border, 9°08'N, 82°58'W, 2350-2450
m, 8 Sep. 1984, G. Davidse, G. Herrera G. &
M. II. Grayum 28625 (holotype, MO; isotype,
CR). Figure 12.

Gonolobo chiriquensi proxima forma ct amplitudine f lo¬
in m foliorumque sed differt praesertim indumento prom¬
inent i el nervis secundariis foliorum paucioribus.

Perennial parts unknown, young stems with
dense simple or mixed indumentum, long hairs
0.5—2 mm long, pale yellow, reflexed to nearly pat¬
ent . short hairs, when present, ca. 0.1 mm long,
internodes 1—19 cm. Leal blades elliptic or some¬
what ovate or obovate, 6.3—10 X 2.6—5 cm. apex
acuminate to attenuate, base rounded to truncate,
pilose adaxially with hairs 0.4— 1.5 mm long, midrib
sometimes with short hairs, pilose or pilosulose
abaxially with hairs 0.1— 1.5 mm long, lateral veins
5 to 6 pairs, middle veins 40°— 60° to midrib, col¬
leters 2, long and thin and often with dark bases
and light tips; petiole 1.5— 3.9 cm. pilose, hairs an-
trorse. Inflorescence congested-racemiform, 2- to de¬
flowered, with simple or mixed indumentum, pe¬
duncle 3-17 mm, axis ca. 0.2 mm, pedicel 5-18
mm, bracts 1.2—5 X 0.4—1 mm, lanceolate; calyx
tube 0.5— 1.5 mm long, with 0 to 2 colleters per
sinus, lobes lanceolate to elliptic with acute tips.
6—7.8 X 2.2— 3.3 mm, green, adaxially glabrous,
abaxially pilosulose with hairs erect to appressed,
0. 1-0.5 mm long; corolla rotate, purple or green

240

Novon

i

Figure 12. Gonolobm Iruncatifolius W. I). Stevens. — A. Sean of holotype (. Davidse el al. 28625, MO). — H. Close-up
of flowers.

with brown veins, abaxially pilosulose with hairs
appressed to erect, 0.2— 0.5 mm long, adaxially the
tube, surface of annulus, and one margin of each
lobe with curly white hairs 0.05—0.1 mm long, tube
2—5 mm long, faucal annulus an erect lip, 0.2— 0.3

mm tall, upper margin without distinct hairs, lobes
asymmetrically elliptic with asymmetrically obtuse
lips, 9-12 X 7—8.4 mm. patent; gynostegium with
stipe 0.3— 0.5 X 1—1.3 mm, 5-ribbed, corona a 5-
lobed disk, fleshy and deeply rugose, each lobe

Volume 15, Number 1
2005

Stevens 241

Gonolobus from Mexico and Central America

Figure 13. Gonolobus ustulatus W. I). Stevens. — A. Sean of holotype ( Gomez 11413, F). — B. Close-up of flowers.

somewhat eueullate and retuse at tip, outer margin
deeply striate. 1.4— 2.5 mm wide at lobes. 1.1—2 mm
wide at sinuses, anther appendages deflexed. with
a broad, rounded base and an obdeltate tip, apex
shallowly 3-lobed to shallowly crenulate, 0.7— 0.9

mm long, 0.9— 1.3 mm wide at tip, somewhat fleshy,
terminal appendages tightly oppressed to style
apex, 0.2— 0.5 X 2—2.2 mm, brown, guide rails
straight, parallel, indistinct, nearly vertical, ca. 0.2
X 0.1 mm; corpuseulum 0.26-0.3 X 0.1—0.15 mm.

242

Novon

sagittate, red-brown to nearly blac k, with a hooked
tip, translators 0.24—0.25 X 0.14—0.15 mm. pollin-
ia 0.83—0.94 X 0.31—0.34 mm, sigmoidially ob-
ovate; style apex 3-3.3 mm wide. Immature folli¬
cles fusiform, apex acute, base* narrowed, at a right
angle to follicle body, ea. 9.5 X 2.5 cm. with 5
complete wings ea. 5 mm wide, glabrous, glaucous;
seeds unknown.

This new species is clearly closest to Gonolohus
chiriquensis Woodson, found in similar habitats in
Costa Rica and Panama. The two share generally
similar leaf size and shape, especially the more or
less truncate leaf bases, and generally similar flow¬
er size and shape, especially the short, broad,
strongly asymmetrical corolla lobes that are remi¬
niscent of many species of Matelea Aublet. There
are, however, a number of striking differences. Gon-
olohus chiriquensis is practically glabrous on all
parts while G. truncal iftorus is conspicuously vil¬
lose. notably including the outside of the corolla,
the new species lias 5 to 6 secondary leaf veins (vs.
8 to 14), and the style apex is 3—3.3 mm in di¬
ameter (vs. 1 .6—2 mm). The corona of Gonolohus
t runcal if olius is much more massive and produces
a white mucilage when soaked in water, perhaps
indicating that it is secreting something other than
nectar and a phenomenon not seen in other species.

This is a cloud forest species known from 1500
to 2750 m, among the highest elevations known for
Gonolohus. It is known to flower in July, September,
and December, and the immature fruits were col¬
lected in September.

Paratypes. COSTA RICA. Heredia: I*. N. Braulio
Carillo, ile la Kst. Barva I km hacia Laguna Barva, G.
Rivera 409 (INB. MO). Puntarenas: R. I. Ujarrds. R. B.
Diirika, Q. Jimenez A' A’. Aguilar 1940 (INB, MO). San
Jose: I’. N. Chirripd, camino a Chirripd, E. Alfaro 989
(INB. MO).

13. Gonolohus ustulatus W. I). Stevens, sp. nov.
TYPE: Costa Rica. Alajuela: R. K. San Ramon.
10°I3'N, 84°37'W, 900-1200 m, 12-15 Mar.
1987. J. Gdmez E. / 1413 (holotype, F). Figure
13.

Gonolobo leiantho maxime similis sed caulibus foliis
infloreseentiis ferrugineis pilosis lobis calyces longioribus
angustioribus corollis intra tantum papillatis.

Perennial parts unknown, young stems with
dense mixed indumentum, long hairs 2-2.5 mm
long, brown, short and glandular hairs ea. 0.1 mm
long, internodes 22—25 cm. Leaf blades narrowly
elliptic to ovate, 7—8 X 2.3—3 cm. apex attenuate,
base lobate, lobes descending, sinus 0.3— 0.4 cm
deep, pilose with brown hairs 1 — 1.5 mm long, lat¬

eral veins 10 to 12 pairs, middle veins ea. 60° to
midrib, colleters 2. with dark bases and light tips;
petiole 1.4-2 cm, with mixed indumentum. Inflo¬
rescence congested-racemiform, 2-flowered, with
mixed indumentum, peduncle 22—25 mm, pedicel
(immature) 3—8 mm, bracts 3.6—8 X 0.8-1 mm,
lanceolate; calyx tube ea. 0.8 mm long, with 1 eol-
leter per sinus, lobes lanceolate with acute t i ps,
14—15 X 3.2— 3.4 mm, green, adaxially glabrous,
abaxially with mixed indumentum, long hairs
brown, erect, 1 — 1.5 mm long; corolla rotate, green,
reticulate, abaxially glabrous or with a few hairs at
tips of lobes, adaxially with lube and a broad band
along one margin ol each lobe papillate, lube ea. 5
mm long, faucal annulus a thin, erect lip. ca. 0.3
mm tall at lobes, 0.1 mm tall at sinuses, hispidulose
with white hairs ea. 0. 1 mm long, corolla lobes nar¬
rowly asymmetrically elliptic with asymmetrically
acute tips, with inconspicuous auricles at the si¬
nuses. 20-21 X 9—10 mm, patent; gynostegium
with stipe ca. I X 1.5 mm. 5-winged, corona a 5-
lobed disk, the 5 wings of the stipe continuing as
radii on the corona, ea. 3.5 mm wide, papillate,
outer margin vertically striate, anther appendages
slightly deflexed. asymmetrically reniform, ea. 1.4
X 2.3 mm in outline, fleshy, brown, terminal ap¬
pendages tightly appressed to style apex, ea. 0.3 X
3.2 mm, translucent white, guide rails straight, par¬
allel. indistinct, tilted under style apex. ea. 0.2 X
0.1 mm; eorpuseulum ea. 0.43 X 0.18 mm, sagit¬
tate. dark red-brown, with a hooked tip. translators
ea. 0.34 X 0.24 mm, pollinia ca. 1.66 X 0.48 mm,
± symmetrical, fertile (distal) body with a distinct
longitudinal ridge on one face; style apex ea. 4.5
mm w ide, green. Fruits and seeds unknown.

In technical characters, this imperfectly known
new species is probably closest to Gonolohus leian-
thus Donnell Smith, a species ranging from south¬
ern Mexico to northern Nicaragua, but is quite dif¬
ferent in general appearance because all parts
except the flowers are covered with long brown
hairs. Additionally, the new species can be recog¬
nized by having narrower leaves with more second¬
ary veins, longer and narrower calyx lobes, corolla
lobes merely papillate and lacking a white callus,
and a papillate corona.

14. Gonnlnhus variabilis W. D. Stevens, sp. nov.
TV PE; Costa Rica. Puntarenas: NE slope of
l ila de Cal, along roar! between San Vito de
Coto Rrus and Ciudad Neily, 8°41'()0”N,
82°56'30"W, 620 m, 7 Mar. 1987, M. //. Gra-
yum & C. Fleming HI 18 (holotype, MO). Fig¬
ure 14.

Volume 15, Number 1
2005

Stevens 243

Gonolobus from Mexico and Central America

Figure 14. Gonolobus variabilis W. I). Stevens. — A. Scan of holotype ( Grayum & Fleming 81 18, MO). — B. Close-
up of flowers.

A Gonolobo albomarginato floribus minoribus trichom-
alibus corollae extus intusque brevioribus, a G. eduli co¬
rolla atrovirenti eallis loborum corollarum trichomatibus
corollae extus longioribus trichomatibus corollae intus
brevioribus recedit.

Perennial j>arts unknown, young stems with
dense mixed indumentum, somewhat organized into
2 lines in internodes, long hairs 1—2.5 mm long,
yellow to brown, short and glandular hairs 0.1— 0.2

244

Novon

mm long, internodes 3-18 cm. Leaf blades elliptic
to ovate, 8.5—13.5 X 4.6— 8.4 cm, apex acuminate
to attenuate, base lobate, lobes descending to over¬
lapping. sinus 1—2 cm deep, sparsely pilose adax-
ially with hairs 0.3—1 nun long, midrib also with
short and glandular hairs, abaxially with mixed in¬
dumentum. long hairs 0.2-1 nun long, lateral veins
6 to 8 pairs, middle veins 35°— 50° to midrib, col-
leters 2 to 6, often with dark bases and light tips;
petiole 2.0— 7.7 cm, with mixed indumentum. Inflo¬
rescence congested-racemiform, 2- to 10-flowered,
with mixed indumentum, peduncle 6—30 mm, axis
0.1— 0.6 mm, pedicel 15—45 mm, bracts 0.7— 5.6 X
0.2— 0.7 mm, linear; calyx tube 0.7—1 mm long, with
1 colleter per sinus, lobes lanceolate with acute
tips, 5.3-6 X 1.2—2 mm, green with purple tint,
adaxially glabrous or with a few hairs on distal mid¬
rib, abaxially with mixed indumentum, long hairs
erect, 0.4— 0.8 mm long; corolla briefly eampanulate
and then rotate, green, abaxially with sparse to
dense, simple or mixed indumentum, long hairs
0.2—1 mm long, adaxially with tube and proximal
third to half of lobes hispidulose or papillate with
while hairs to 0.05 mm long and with a band of
curly white hairs 0.2— 0.4 mm long immediately
along white callus ol lobes, tube 1.5—2 mm long,
faucal annulus erect. 0.5—1 mm tall, hispidulose
with white hairs 0.2— 0.4 mm long, lobes narrowly
lanceolate or elliptic with acute lips, with conspic¬
uous auricles at the sinuses, one margin of each
lobe with a thick, bright white callus, 9—14.1 X
.3.1— 4.8 mm, patent; gynostegium with stipe 1-1.5
X 0.7— 1.8 nun. 5-winged, corona a 5-lobed, fluted
bowl, the 5 wings of tin* stipe continuing as radii
on the corona and forming 5 chambers, 0.8—1 .8 mm
tall at lobes, 0.5— 1.4 mm tall at sinuses, anther
appendages deflexed, reniform to nearly spathulate,
with the broad apex shallowly crenulate, 1.2— 2.4 X
1.8— 2.5 mm in outline, fleshy, pale yellow to pale
pink, terminal appendages tightly appressed to
style apex, 0.1— 0.2 X 1 .8— 2.4 mm, white to pah-
brown or purple, guide rails straight, parallel, in¬
distinct to prominent, vertical to tilted under style
apex, 0.1— 0.7 X 0.1— 0.2 mm; corpusculum 0.26—
0.33 X 0.1—0.11 mm, sagittate, brown or red-
brown, with a hooked tip. translators 0.31—0.39 X
0.18—0.19 mm, pollinia 1.05—1.38 X 0.36—0.46
mm. fertile (distal) body distinctly flattened or con¬
cave on one face; style apex 2.6— 4 mm wide, pale
yellow. Fruits and seeds unknown.

While this species is easily characterized by a
series of characters, especially the indumentum of

the corolla, and unlike any other known Gonolobus,
it is at the same time curiously variable. Two of the
collections have long peduncles and two short,
three have the outside of the corolla with a dense
mixed indumentum while one has only scattered
long hairs, three have pale yellow dorsal anther ap¬
pendages and in one they are pale pink, two have
distinctly larger style apices, and three have nor¬
mal, indistinct, 0.1— 0.2 nun long guide rails under
the style apex and one has prominent, nearly ver¬
tical guide rails about 0.7 mm long. One could in¬
vent this species by combining characters from
Gonolobus albomarginatus (Pittier) Woodson, G. ed¬
it l is Hemsley, and G. heterophyilus (Hemsley) W. I).
Stevens, and perhaps hybridization has been in¬
volved, but the four known collections are tempo¬
rally and geographically separated and easily rec¬
ognized. Gonolobus albomarginatus, G. taylorianus
W. 1). Stevens & Montiel, and G. fraternus Schle-
ehtendal. among Mesoamerican species, are at least
equally variable.

This is a cloud forest species known from 620 to
1300 m and is known to flower between November
and March.

Paratypes. COSTA RICA. Alujiiclu: If. B. Montev¬
erdi-. IVfias Blancas. IF lluber A- E. Hello 6535 (MO); l.a
Leiia. 4. Smith P2154 (MO). San Jose: Rfo Claro valley
below La Palma NE of San Jeronimo, It. C. Burger A R.
A. Liesner 6260 (F).

Acknowledgments. Alba Arbelaez prepared the
line drawings and Fred Keusenkothen and Sandra
Arango prepared the specimen scans. Stephen
Koch (CHAPA) and Andrew Sanders (UCR) made
a special effort to provide interesting specimens.
Olga Martha Montiel, Victoria Hollowell. and Neil
Harriman made significant improvements to tin-
original manuscript.

I .iterature Cited

Oreuter. W.. J. McNeill, F. It. Barrie, 11. M. Burdet, V.
Demonlin, T. S. Filgueiras. I). H. Nicolson, P. C. Silva,
.1. F. Skog, P. Trehane, N. J. Turland & 1). I,. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Stevens, W. 1). A ( ). M. Montiel. 2002. A new species ol
Gonolobus (Apocynaceae, Asclepiadoideae) from Me-
soamerica. Novon 12: 551—554.

- & - . 2004. Gonolobus ineerianus (Apocy¬
naceae. Asclepiadoideae), una nneva especie de Me-
soameriea. Novon 14: 350-353.

- , C. Ulloa L.. A. Pool & (). M. Montiel. 2001.

Flora de Nicaragua. Monogr. S\st. Rot. Missouri Hot.
Card. 85: i-xlii. 1-2666.

Williams, L. (). 1968. Tropical American plants, X. Kiel -
diana, Bot. 32: 35—61.

A New Genus and Two New Species of Ferns from Vietnam

Wu Sugong

Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204,
Yunnan, China, sugong@public.km.yn.cn

Phan hr Loc

Institute Ecology and Biological Resources Department of Botany,

Nghia Do-Cau Giay-Ha Noi, Vietnam, pkeloc@yahoo.com

Xiang Jianying

Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204,
4 un nan. China, xiangjianying@mail.kib.ac.cn

Abstract. The new genus Kontumia (Polypodi-
aceae?) and its type species Kontumia heterophylla
from Vietnam are described and illustrated, kon¬
tumia is distinguished from Gymnogrammitis by its
seandent habit, dimorphic leaves, elathrate trans¬
parent scales on the rhizome and petiole bases, anil
aspects of leaf morphology. Also, a new species of
Cyrtomium from Vietnam is described and named
as C. elongation (Dryopteridaceae). Cyrtomium
elongatum is similar to C. wulingense S. F. Wu, but
differs by its longer leaves (to 90 cm) and more
numerous pinnae (more than 60 pairs) with cordate
rather than oblique bases, abaxially with sparse
small brown scales, and with brown punetations
within the areoles.

Key words: Cyrtomium, Dryopteridaceae, fern,
kontumia , Vietnam.

Because of its inaccessibility and steep terrain,
about two-thirds of the montane areas with rich
plant diversity in Vietnam have not been fully ex¬
plored and studied. According to a recent summary
(Phan. 1998), there are 291 families, 2010 genera,
and 9628 plant species recorded for Vietnam. It is
estimated that only 80% of the country’s plants
have been described so far (Averyanov et al., 2000;
T. H. Nguyen et al., 2000; Phan et al., 2002). For
instance. 676 species of pteridophytes were de¬
scribed in the Flore Generate de Flndo-Chine (Tar-
dieu-Blot & Christensen, 1939—1951). However, at
least 776 pteridophyte species have been recorded
more recently (Pham. 1991), and a new genus of
fern, Caobangia A. R. Smith & X. C. Zhang, was
described in 2002 (Smith & Zhang. 2002). In 2002.
during a trip by the first author to northern Viet¬
nam. a number of specimens were examined at 11 N.

these collected by Phan Ke Loc and others. In
2003, the authors jointly visited central and north¬
ern portions of Vietnam again to collect additional
fern specimens. From our survey and examination,
one new genus and two new species are described
below.

Kontuinia heterophylla S. K. Wu & L. k. Phan,
gen. et sp. nov. TYPE: Vietnam. Kon Turn
Prov.: kon Plong Distr., Ngoc Tern Municipal¬
ity, Mang La State Forest, ca. 14°41'4I"N,
108°22'21"F, ca. 1 100 m. 9 Apr. 2000, Plian
ke Loc P-10551 (holotype, HN; isotypes,
kUN, LE, MO). Figures 1. 2.

Deiius novum valde insigne probahi liter affine Gymno¬
grammitis, a qua differt fronde sterili a fertili valde div-
ersa, paleis rhizomatis et stipitis basis translucide elath-
rat is, stipite frondis sterilis anguste alato, lamina crasse
membranacea, venis elevatis; fronde fertili fronde sterili
longiore, stipite tereti, utrinque sine alato, lamina tenuiter
tripinnatifida, segmento ultimo tenuiter lineari vix 1 mm
lato, apice latiore, obtuso, vena singulari ad marginem non
altingenti; soris orbieularibus, apicem nervorum occupan-
libus.

Fern. Stem long seandent, cylindric, with dictyo-
stele, densely covered with scales; scales 1 cm X
1 mm. lanceolate, acuminate at apex, margins
sparsely denticulate, transparent elathrate, base
peltate. Leaves dimorphic, approximately or re¬
motely and spirally arranged, sterile leaves shorter,
ca. Vi the length of fertile ones; petiole terete, stra¬
mineous, inconspicuously articulated. Petiole of
sterile leaves to 10 cm long, ca. 2 mm diam., nar¬
rowly winged on each side, glabrous; lamina ob¬
long-lanceolate, ca. 15 cm long, middle 5—8 cm
wide, obtuse at apex, truncate at base, bipinnati-
partite, crassimembranous; veins pinnate, simple in

Novon 15: 245-249. 2005.

246

Novon

Figure 1. Kontumia helerophylla S. K.Wu & L. K. Phan. — A. Plant with fertile leaf. — B. Seale of rhizome. — C.
Part of fertile pinna. — 1). Part of sterile pinna. Drawn from the holotype and the KLIN isotype.

each pinnule, not reaching the margin; primary pin¬
nae 10 to 15 pairs, spreading, alternate, oblong-
lanceolate, ca. 4 X 1.5 cm, obtuse at apex, cuneate
at base, short-stalked, secondary pinnules lorate, 1

cm long, ca. 2 mm wide, obtuse at apex, margin
lobed, primary and secondary rachises glabrous,
winged. Petiole of fertile leaves to 26 cm long, 2
mm diam., both sides wingless, glabrous; lamina to

Volume 15, Number 1
2005

Wu et al.

Ferns from Vietnam

247

Figure 2. Scanning electron micrograph of a spore of
Kontumia heterophylla (taken from Wu et al. WP-135).

18 cm long, 5 cm wide, tripinnatifid, segments lin¬
ear, less than 1 mm wide, ultimate pinnae apex
broader, obtuse, primary and secondary raehis
wingless. Sori orbicular, terminal on the veinlets,
exindusiate, with 15 to 18 sporangia, lacking pa-
raphyses. Spores 64 per sporangium, 45—65 pun in
longest dimension, ellipsoidal, the thin perispore
with irregular spines, rods, and globules.

Habitat and ecology. This species is found in
the primary closed evergreen tropical seasonal
broad-leaved submontane forest on upper slopes of
sandstone and gneiss mountains, sometimes in sub¬
montane coniferous forest on ridges dominated by
Fokienia hodginsii (Dunn) A. Henry & II. II.
Thomas or Dacrydium elatum (Roxburgh) Wallich
ex Hooker. The forest ground frequently has deep
soil rich in humus; rock outcrops were rare.

fhe habitat of kontumia heterophylla is found on
the summits of the southern Truong Son range. Due
to its unique geographic location, the rains from
both southwestern and northeastern monsoons pre¬
vail in this area, which result in a locally humid
climate. By referring to the climatological data of
neighboring stations of the southern Truong Son
range, such as Ba To and Gia Vuc in Quang Ngai
Province on the eastern slope and Kon Turn and
Dac To in Kon Turn Province on the western slope
(K. V. Nguyen et al., 2000), we estimate t Hal the
mean annual precipitation al the type locality is at
least 2500 mm, with at most two diy months. Mean
annual temperature is about 21°C; the normal tem¬
perature is above 17°C.

kontumia is an unusual genus. It seems closely
related to Gymnogrammitis Griffith, which has tra¬
ditionally been accepted as a member of Davalli-
aceae (Kramer, 1990). However, Ching (1966)
placed Gymnogrammitis in an independent family,
Gymnogrammitidaceae, and suggested its possible
affinities to Polypodiaceae. According to analysis of

the combined data sets of the chloroplast genes
rbcL and r/«4, Schneider et al. (2002) demonstrated
that Gymnogrammitis belongs to the polygrammoid
clade of Polypodiaceae, not Davalliaceae. The mon-
olete spores (Fig. 2) of kontumia with the thin per¬
ispore beset irregularly with globules and spines
and/or rods (some degraded) are quite similar to
those of some species of Polypodiaceae and those
of Gymnogrammitis (as illustrated by Tryon & Lu-
gardon, 1991), but different from those found in
Davalliaceae.

The new genus we describe hen1 has a scandent
habit, dimorphic leaves, the scales of the rhizomes
and petiole bases clathrate, the petiole of sterile
leaves with narrow wings, lamina crassimembran-
ous, with elevated veins. The fertile leaves are lon¬
ger than the sterile ones, with a terete petiole, wing¬
less tripinnate lamina, linear ultimate pinnules less
than 1 mm wide, apex broader, obtuse, and a single
veinlet in each pinnule not reaching the margin.
The sori are orbicular, terminal on the veinlets but
not dorsal, and thus are very different from those
of Gymnogrammitis. More detailed studies on tin*
systematic position of these ferns are needed in the
future.

Paratypes. VIETNAM. Kon Turn Prov.: Kon Plong
Distr., Hieu Municipality, Mang La forest enterprise,
14°39'N, 108°25'E, 1000—1200 m, primary evergreen
broad-leaved wet forest on steep mountain slopes on sand¬
stone and gneiss, 15 Apr. 2000, Averyanov et al. 17 1-5129
(IIN, I,E, MO); Kon Plong Distr., 1 1 ieu Municipality,
14°41'48"N, 108°22'23"E, ca. 1250 m, 2(1 Nov. 2003, Wu
et al. WP- 136 (HN, KUN. MO); Kon Plong Distr., Po E
Municipality, first village, 14°43'I6"N, 108°38'18"E, ca.
1200 m, logged primary closed evergreen seasonal tropical
broad-leaved submontane forests on slope of mountains,
22 Nov. 2003, Wu et al. WP-201 (HN. KUN, MO).

Gy rtnmiiim elongation S. K. Wu L. K. Phan,
sp. nov. TV PE: Vietnam. Can Bang Prov.:
Nguyen Binh Distr., Yen Lac Municipality, ca.
22°44'N, 105°50'E, ca. 1.300 m, Phan ke Loc
el al. CBL- 1 586 (holotype, I IN; isotypes, KUN,
MO). Figure 3.

Species in genere Cyrtomio valde insignis, aff in is C.
wulingensi, sed plantis ad 90 cm aids, pinnis plus quam
60-jugis, pinna basi eordata non obliqua, supra glabra
subtus paleis brunneis sparse obtecta, in areola punctis
brunneis dilfert.

Plant to 90 cm tall. Petiole 30 cm long, base 4
cm diam., brown or stramineous, adaxially grooved,
abaxially densely covered in brown linear-subulate
denticulate scales. Leaves lanceolate, ca. 60 cm
long, ca. 4.5 cm broad at mid-point, acuminate at
apex, base gradually narrow, simply pinnate, cori¬
aceous; pinnae ca. 62 pairs, alternate or suboppos-

248

Novon

F igure 3. Cyrtomium elongatum S. k. Wu & L. k. I’liau. — A. Plant with fertile leaf. — B. Part of pinna. — C. Seale
from lower portion of stipe. Drawn from the holotype and the KUN isotype.

ite, approximate to each other; upper middle pinnae
spreading-lanceolate, 2.2 X 1.2 cm, the basal pairs
1 .5 X 1 cm, smaller than others, deflexed from ra-
cltis, obtuse at apex, cordate tit base, entire on the

margin, abaxially sparsely with small brown scales,
brown punctations within areoles. Veins pinnate¬
forked, inconspicuous adaxially, slightly elevated
abaxially; costa adaxially immersed, abaxially ele-

Volume 15 Number 1
2005

Wu et al.

Ferns from Vietnam

249

vated; lateral veins joining on each side of costa
into 2 serial areoles. Kachis terete, adaxially
grooved, sparsely covered with linear small brown¬
ish scales. Sori orbicular, attached along a line on
each side of costa; indusium caducous, with ea. 35
sporangia. Spores ellipsoidal with prominent tuber¬
cles.

Cyrtomium elongatum is similar to the Chinese
species C. wulingense S. F. Wu, which was de¬
scribed from western Hunan Province, but differs
by the plant to 90 cm tall with more than 60 pinna
pairs, the pinnae cordate at the base (not oblique),
the sparse small brown abaxial pinna scales, and
the brown punctations in the areoles of the veins.

Habitat and ecology. This species occurs on
vertical cliffs of a north-facing, shaded limestone
ridge. This karst region is dominated by relatively
dry, species-rich, primary evergreen coniferous for¬
est on peaks and ridge tops or less commonly by
rather open forests, file floor of these forests is a
tangle of rocks, crevices, small cliffs, and plants.

Acknowledgments. We are grateful to C. Yat-
skievyeh ol the Missouri botanical Garden for the
SKM photo of spores and his valuable suggestions
on the manuscript. We are also indebted to M. Veith
and the electron microscopy facility at Washington
University, St. Louis. The field survey and research
were funded by the U.S. National Geographic So¬
ciety, grant numbers 6300-98 and 7512-03. We ex¬
press our sincerest gratitude for these grants. Spe¬
cial thanks are extended to Nguyen Tien Hiep, S.
T. Averyanov, and Yang Yongping for their great
support and help. We thank J. K. Yang for the il¬
lustrations (T igs. 1 and 3).

literature Cited

Averyanov, h. V., T. II. Nguyen, I.. k. Phan & I). II.

Dzuong. 1996. Fndangered Vietnamese Paphiopedil-

ums. Part I. Paphiopedilum helenae. Orchids (West
Palm Beach) 65: 1062—1068.

- . - . - & A. L. Averyanova. 2000. Pre¬
liminary orchid checklist of Can Bang Province. Lin-
dleyana 15: 130—164.

Ching, R. C. 1966. Gymnogrammitidaeeae Ching, a new
fern family. Acta. Phytotax. Sin. 9: I 1-1 (>.

Kramer, k. U. 1990. Davalliaceae. Pp. 74— 80 in k. Ku-
bitziki (general editor). The Families and Genera of
Vascular Plants, Vol. 1. Pteridophytes and Gymno-
sperms. Springer- Verlag, Berlin.

Nguyen, k. V. (general editor), T. H. Nguyen, I.. k. Phan
& T. H. Nguyen. 2000. Bioclimatic Diagrams of Viet¬
nam. Vietnam National University Publishing House,
Hanoi [in Vietnamese],

Nguyen, T. II.. L k. Phan N L. V. Averyanov. 2000. Some
new species for the flora of Vietnam collected from the
limestone mountains of Can Bang province. ,|. Biol. (Ha¬
noi) 22(4): 1 — 11.

Pham, II. II. 1991. Cayco Vietnam [An Illustrated Flora
of Vietnam], Vol. I, Pt. 1. Mekong Printing, Hanoi.

Phan. F. k. 1998. On the systematic structure of the Viet¬
namese flora. Pp. 120—129 in Zhang A. & Wu S. (edi¬
tors), Floristic Characteristics and Diversitv of East
Asian Plants. Proceedings of the First International
Symposium on Floristic Characteristics and Diversity of
East Asian Plants, Kunming, 1996. China Higher Ed.
Press, Beijing, and Springer- Verlag, Berlin.

- . T. II. Nguyen & L Averyanov. 2002. What is

new to the llora ol Cao Bang. Pp. 37—45 in Sustainable
Development and Protection of the Forest and Biodi¬
versity on Limestone Mountains of Vietnam. Forest In¬
ventory and Planning Institute, Hanoi [in Vietnamese].

Schneider. II.. A. K. Smith, B. Cranfill, C. II. Haulier, T.
A. Ranker & I. Hildebrand. 2002. Cymnogrammitis
dnreiformis is a polygrammmoid fern (Polypodiaceae) —
Resolving an apparent conflict between morphological
and molecular data. PI. Syst. Evol. 234: 121—136.

Smith. A. R. & X. C. Zhang. 2002. Caobangia, a new
genus and species of Polypodiaceae from Vietnam. No-
von 12: 546—550.

fardieu-Blot, [M. I..| & C. Christensen. 1939—1951. Crvp-
togamies vaseulaires. Pp. 1—596 in II. I. ecomte (editor),
Flore Generale de I'lndo-Chine, Vol. 7(2). Masson &
Cie, Paris.

Tryon, A. F. & B. Fugardon. 1991. Spores of the Pteri-
dophyta. Springer- Verlag, Berlin.

Lectotypifications in the Genus Andropogon (Poaceae)

Ana Zanin

Departamento de Botaniea, Universidade Federal de Santa Catarina, Trindade, Floriandpolis,
Santa Catarina, 88040-900, Brazil, anazanin@terra.eom.br

Hilda Maria hmgh i- Wagner

Departamento de Botaniea, Universidade Federal do Rio Grande do Sul, Avenida Bento
Gonsalves, 9500, CEP 91501-970, Porto Alegre, Rio Grande do Sul. Brazil.

hmlw@plugin.eom.br

Abstract. During a taxonomic revision ot the
Brazilian species of Andropogon, the need for lec-
totypification of the following names became ap¬
parent: Andropogon arenarius Hackel f. snbcornple-
tus Hackel, Andropogon bicornis L. var. gracUlimus
Hackel, Andropogon carinatus Nees, Andropogon
carinatus var. leiophyllus Hackel, Andropogon hy-
pogynus Hackel, Andropogon ingrains Hackel, An¬
dropogon palustris Pilger, and Andropogon selloan-
ns (Hackel) Hackel.

Key words: Andropogon, lectotypification, Po¬
aceae.

Of approximately 100 species of Andropogon L.
(Clayton & Renvoize, 1986), 28 occur in Brazil
(Zanin, 2001). Lectotypes are designated here for
eight names related to some of these taxa. The lec-
totypifications were necessary because: the holo-
types were destroyed, several syntypes were men¬
tioned in the protologue, no holotype was selected
from the isotypes of the only collection number cit¬
ed by the author, or only the locality of collection
was cited in the protologue.

For the designation of the lectotypes the best
preserved specimens that fit the original description
were chosen. Further, specimens that had clear in¬
dication to have been studied by the author of the
species, for instance with notes in the author’s
handwriting, or preserved in the herbaria where the
author worked or where his main collections are
deposited, were considered.

Andropogon arenarius Hackel, Flora 68(8): 184.
1885. TYPE: Uruguay. Montevideo: arenales
da Costa, s.d., Arechavaleta 204 (holotype, W;
isotvpes, ( i. K).

Andropogon arenarius Hackel f. subcompletus Hackel, in
Findman, Kongl. Svenska Vetenskapsakad. Handl.
846: 6. PMK). TYPE: “Brasiliae austr., Rio Grande
do Sul, I Ilia dos Marinheiros prope oppidium Rio

Grande, in collibus arenae profunda,” s.d., C. A. M.
Lindman, Exp. I. Regnell. ,4. 699 (lectotype, desig¬
nated here, S; isotypes, S, W).

Hac kel (1900: 6) mentioned “C. A. M. Lindman,
Exp. Regnell. A. 699” in the protologue. One spec¬
imen at W, with label data slightly different from
the protologue, and two specimens at S, where the
Regnellian collection is preserved, were found. The
three specimens are well preserved and are iden¬
tified by Hackel’s own handwriting. The lectotype
was chosen from the two S specimens whose label
data are the same as that cited in the protologue.

Andropogon bicornis L., Sp. PI. 1046. 1758.
TV PE: “Porto Rico, Mayagiiez, between Monte
Mesa and the sea,” 27 Oct., A. Chase 247 (typ.
cons. MO, not seen).

Andropogon bicornis L. var. graeillimus Hackel. in Martins
& Kidder. FI. Bras. 2(8): 285. 1888. TYPE: “In Par¬
aguay, l,ambor6 pr. Assumpcion, Balansa n. 271, a.
1874” (lectotype, designated here, W; isotypes, 0

1 3], K).

Two collections were mentioned in the proto¬
logue, Mosen, from Brazil, and Balansa 271, from
Paraguay. Five specimens of Balansa 271 were
found, one at W, three at G, and one at K. Of these,
the one from W, where most of the specimens Hack¬
el studied are deposited, is the best preserved and
the only one identified by him. This specimen is
chosen as the lectotype. One specimen of Mosdn
was found at W and was also identified by Hackel,
but it is not as well preserved.

Andropogon carinatus Nees, Agrostologia Bras-
iliensis, in Mart., FI. Bras. Enum. PI. 2(1):
330. 1829. TYPE: “Brasilia, Sellow" (lecto¬
type, designated here, K).

Andropogon carinatus var. leiophyllus Hackel, in A. DC.
& DC., Monogr. Phan. 6: 434. 1889. TYPE: “Bra-

INovon 15: 250—252. 2005.

Volume 15, Number 1
2005

Zanin & Longhi-Wagner
Lectotypifications in Andropogon

251

silia, Prov. Minas Gerais,” Glaziou 1 7381 (lectotype,
designated here, W; isotype, K).

Nees (1829) mentioned “Habitat in Brasilia aus-
traliore — Sellow vidi in herb. Reg. Berol.” in the
protologue of Andropogon carinatus. An extensive
search by the curatorial staff at B and by the first
author failed to locate the cited specimen or any
collection of A. carinatus by Sellow. The material
was probably destroyed during World War II. Two
specimens annotated “Brasilia, Sellow;' identified
as A. carinatus by Nees, one at K and one at W.
were found. Kven though the W specimen has bet¬
ter inflorescences, the K specimen is being desig¬
nated as the lectotype because it is the only one
with an original label from B attached to its sheet.
According to Filgueiras (2003), there is one isotype
at LE, but this specimen was not checked for this
work.

Two collections of Andropogon carinatus var.
leiophyllus, Glaziou 17381 and Glaziou 17382.
were mentioned in the protologue. Both were found
at W. file first one has a duplicate at K and the
second at BR. The specimen Glaziou 17381 de¬
posited at W is the more complete and best pre¬
served. and agrees with the morphology described
in the original description. Therefore, it is chosen
as the lectotype.

Andropogon liypogynns Hackel, in Martins &
Eichler. FI. Bras. 2(3): 290. 1883. TYPE:
“Brasilia, Riedel 1655" (lectotype, designated
here, G; isotype, K).

Hackel ( 1 883) described Andropogon h ypogvnus
var. genuinus, variety anatherus, and variety con-
jugens. For A. hypogynus var. genuinus the author
mentioned Weddell 1858, Sellow s.n.. Warming s.n.,
and Riedel s.n.: the first two were not located during
this research. The specimen of Warming deposited
at W bears an inflorescence of the variety conjugens
mixed with a complete specimen of the variety gen¬
uinus, as mentioned by Hackel (1883). Further, the
same sheet contains one envelope with a fragment
of Riedel 1655, corresponding also to A. hypogynus
var. genuinus. probably the same as Riedel s.n. cit¬
ed by Hackel (1883). All this mixed material was
identified by Hackel. One specimen of Riedel 1655
was found at G. It was chosen as the lectotype of
A. hypogynus because it is more complete and bet¬
ter preserved than the Warming collection at W.
There is no other material mixed on the sheet.
There is no doubt that it corresponds to Riedel men¬
tioned by Hackel (1883) in the protologue of the
species because it is identified in the author’s hand¬
writing as “A. hypogynus Hackel in Mart. F. Br.”

I here is also one duplicate of Riedel 1655 at K. for
which there is no indication of having been seen
by Hackel.

Andropogon ingratus Hackel. Oesterr. Bot. Z.
51(5): 151. 1901. TYPE: “Brasilia, Provincia
Minarum, 1891, Glaziou n. 18681" (lectotype.
designated here, W; isotypes, US, \\ (.

Hackel (1901) indicated Glaziou 18681 in the
protologue of Andropogon ingratus, of which there
are two specimens at W annotated by him. There
is one specimen at US also annotated by Hackel.
fhe two specimens of W are better preserved: the
more complete one is being designated as the lec¬
totype and was annotated as such.

Andropogon palustris Pilger, Bot. Jahrb. Syst.
30(1): 137. 1901. TYPE: “Brasil, Mato ( j rosso,
Jul. 1899, Herrmann Meyer 732" (lectotype,
designated here, US).

d’he holotype deposited at B was destroyed dur¬
ing World War II. The US specimen seems to be
the only sheet extant for the typic collection Meyer
732 and is designated as the lectotype.

Andropogon selloanus (Hackel) Hackel, Bull.
Herb. Boissier. 2(4): 266. 1904. Andropogon
leucostachyus Kunth subsp. selloanus Hackel,
in A. DC. & DC.. Monogr. Phan. 6: 420. 1889.
TYPE: “Brasilia Sello ” (lectotype, designated
here, W; isotypes, K, W).

In the protologue of Andropogon leucostachyus
subsp. selloanus, Hackel (1889) cited two collec¬
tions from Brazil, Sellow s.n. and Weddel 2711. be¬
sides Balansa 279a from Paraguay. Two unnum¬
bered specimens of Sellow and one of Balansa
279a were found at W, all of them with original
notes made by Hackel. The morphology of the two
Sellow specimens agrees better with the original
description, and the more complete one was se¬
lected as the lectotype. Another unnumbered spec¬
imen of Sellow was found at K, also annotated by
Hackel.

Other Sellow specimens of A. leucostachyus
subsp. selloanus were examined at h. and W. How¬
ever, although their morphology agrees with the
original description, they were not considered for
lectotypification because it appears that Hackel did
not see them.

Renvoize (1998) cited Balansa 279 as an iso-
synlype of A. selloanus, but this specimen, also
from Paraguay, was collected in a different locality
from that of Balansa 279a and was not considered
for lectotypification.

252

Novon

Acknowledgments. The authors are grateful to
the curators of the herbaria cited and to the Mar¬
garet Mee Foundation for financial support to the
first author. The second author thanks the Brazilian
Research Council (CNPq) for a fellowship.

Literature Cited

Clayton, W. D. & S. A. Renvoize. 1986. Genera Gramiu-
uin: Grasses of the World. Lew Bull. Addit. Ser.. 18.
Filgueiras, T. S. 2008. Andropogon. In: F. (). Zuloaga, 0.
Morrone, G. Davidse, T. S. Filgueiras. P. J. Peterson. U.
J. Soreng & E. Judziewicz, Catalogue of New World
Grasses (Poaceae): 111. Subfamilies Panicoideae, Aris-
tidoideae. Arundinoideae and Danthonoideae. Contr.
ll.S. Natl. Herb. 46: 1-662.

Hackel, E. 1883. Gramineae— Andropogoneae. In: C. F. P.

von Martius A A. W. Eichler (editors), FI. Bras. 2(3):
245-326.

- . 1889. Andropogoneae. In: A. I)e Candolle & C.

l)e Candolle (editors), Monogr. Phan. 6: 1-716.

- . 1900. Andropogon. In: C. A. M. I.indman, Bei-

triige zur Gramineenflora SUdamerikas. Kongl. Svenska
Vetenskapsakad. Hand!. 346: 6.

- . 1901. Neue (baser. Tribus Andropogoneae. Oes-

terr. Rot. Z. 51(5): 150—152.

Nees. (7 G. 1829. Agrostologia brasiliensis. In: C. F. P.
von Martius (editor), FI. Bras. Enum. PI. 2(1): 320—331.

Renvoize, S. 1998. Gramineas de Bolivia. Royal Botanic
Gardens, kew.

Zanin, A. 2001. RevisSo d e Andropogon L. (Poaceae— Pan-
icoideae— Andropogoneae) no Brasil. Tese de Doutorado,
Instituto de Bioeiencias, Universidade de Sao Paulo,
Sao Paulo, Brasil.

Volume 15, Number I. pp. 1-252 of NOVON was published on 29 April 2005.

Volume 15 NO VON

Number 2

2005

Three New Species of Bomarea (Alstroemeriaceae) from the
Andean Region of Colombia

Fernando Abate Guarin

Posgrado de Biologfa, Universiclad de Antioquia. Apartado Aereo 1226, Medellin,
Antioquia, Colombia. fernando_alzate@hotmail.eoni

Arstract. Three new species of Bomarea (Al¬
stroemeriaceae), B. caucana, B. euryantha, and B.
straminea. found during a revision of the genus for
the Flora of Colombia, are described and illustrat¬
ed and their relationships discussed.

Rksumen. Se clcscriben e ilustran ties nuevas es-
pecies de Bomarea (Alstroemeriaceae), B. caucana,
B. euryantha y B. straminea, encontradas durante
la revision del genero para la Flora de Colombia,
y se diseuten sus relaciones.

Key words: Alstroemeriaceae, Bomarea, Co¬
lombia.

The genus Bomarea (Mirbel, 1804; Alstroemer¬
iaceae) mostly inhabits Andean forests between
1500 and 3000 m. with a few species occurring
elsewhere both at lower elevations anti at higher
elevations on forest edges and in paramo forma¬
tions. Bomarea is a monophyletic group closely re¬
lated to Alstroemeria L. (Aagesen & Sanso. 2003;
Sanso & Xifreda, 2001); Dahlgren et al. (1985)
placed both genera in the family Alstroemeriaceae.

In the last taxonomic revision ol Bomarea. Baker
(1888) recognized 75 species; since then nearly
150 species have been described, especially from
the northern Andean region of Colombia, Peru, and
Keuador (Sodiro. 1908; killip. 1936a).

Within Bomarea three subgenera are currently
recognized (Baker, 1878, 1882, 1.888; Pax & Hoff¬
mann. 1930; killip. 1936b; Smith X Gereau.
1991): Bomarea subg. Bomarea, distinguished by
flowers with completely inferior ovaries, dehiscent
(capsular) fruits, resupinate leaves, and a usually

elongate, climbing habit; Bomarea subg. Sphaerine
(Herbert) Baker, characterized by a completely in¬
ferior ovary, indehiscent (berry-like) fruits, usually
resupinate leaves often crowded on the distal por¬
tion of the stem, and erect to decumbent (never
climbing) stems; and Bomarea subg. Wiehuraea (M.
Roemer) Baker, characterized by a partly inferior
ovary, dehiscent (capsular) fruits, non-resupinate
linear leaves with revolute margins, and stiffly erect
stems with decurved stem apices. Notes regarding
subgeneric placement are included in the discus¬
sion of each new species.

During preparation of a revision of Bomarea for
the Flora of Colombia, I found several collections
belonging to three undeseribed species.

1. Bomarea caucana F. Alzate. sp. nov. 15 PK:
Colombia. Cauca: Mun. El Tambo, Reserva
Natural El Tambito, selva de niebla, 1600 m.
I Dec. 1998, H. A. Serna, N. Gomez. O. Cas-
anas, /,. Ordonez, M. Burbano & C. Gonzales
TBS (holotype, CA11P). Figure 1.

Herba sarmentosa; caule glabro glabrescenleve. Lamina
ovata usque ovato-lanceolata, — 1 4 X 2.2— 8.0 cm, infera
folia glabra, nervis pilosis. InHorescentia composita; brac-
teis foliaceis, 1.5— 5.5 X 0.3— 0.8 cm; bracteolis lanceo-
lato-linearibus; radiis 2 ad 4. 2- ad 4-ramosis; pedicellis
glabris pubescentibusve. Flores penduli; sepalis rubris,
2.0— 2.0 cm longis; petalis rubris, 12—20 mm longis.

Sarmentose herb; roots not seen: stem 2—4 mm
diam., glabrous to glabrescent when young; inter-
nodes 5—10 cm long. Leaves pseudopetiolate, re¬
supinate; pseudopetiole 9—17 mm long, pilose;
blade 6—14 X 2.2— 8.0 cm, ovate to ovate-lanceo-

Novon 15: 253-258. 2005.

254

Novon

figure I. Bomarea caucana l\ Al/.ate. — A. Plant. — It. Leaf pubescence, abaxial surface. — C. Ovary, external view.
Drawn from R. A. Serna el al. 785 (CAIJP).

late, glabrous above, nerves pilose beneath, the in-
ternerves pustulate or reticulate, thinly papyra¬
ceous to chartaeeous when dry, base acute, apex
acuminate to cuspidate, margin revolute. Inflores¬
cence compound, pendulous; bracts leaflike, pseu-
dopeliolate to sessile, 1.5— 5.5 X 0.3— 0.8 cm, linear
to lanceolate, acute al base, acuminate at apex, the

margin revolute; braeteoles sessile, 1.0-2. 5 X 0.3-
0.9 cm, linear to lanceolate, lanulose or pustulate
beneath, acuminate at apex; rays 2 to 4. 10—30 cm
long, 2- to 4-branched, pilose to glabrescent; ped¬
icels 1.0— 2.5 cm long, glabrous to pubescent. Flow¬
ers pendulous; inner perianth segments longer than
the outer; outer tepals 2.0— 2.6 cm long, ovate, gla-

Volume 15, Number 2
2005

Alzate

Bomarea from Colombia

255

1 nous, red, obtuse at apex, the claw 1 mm long;
inner lepals 1 .2— 2.0 cm long, oblanceolate, gla¬
brous, red with yellow spots and purple costa, ob¬
tuse at apex; filaments 14—18 mm long, glabrous;
anthers 3—4 nun long; ovary inferior, pubescent to
glabrous, rugose to pustulate; style 22—24 mm long,
glabrous; stigma trifurcate, decurrent, strigose.
Capsule turbinate, 1.5— 2.0 cm diam.; seeds 3—4
mm diam., spheroid to obloid, the sarcotesta yel¬
low-red.

Distribution and habitat. Bomarea caucana is
known from Cauea and Putumayo Departments in
Colombia, where it is found in disturbed areas of
premontane and montane wet forests (Holdridge,
1978) at altitudes of 1600—2910 m.

Phenology. This species presents two flowering
and fruiting periods during the year; tin* first is
February to April and the second July to Septem¬
ber.

Etymology. The specific epithet makes refer¬
ence to the department of Cauea, Colombia, where
the largest population of this species is found.

Taxonomic relationships. This species is related
to Bomarea edulis (Tussac) Herbert, sharing such
features in common as a compound inflorescence
and foliar indurnent and habit; Bomarea caucana
is distinguished by its papyraceous leal blades,
which are pilose beneath only in the nerves, and
its linear to lanceolate bracts. Because of its sar-
mentose stems, resupinate leaves, inferior ovary,
and capsular fruit, B. caucana belongs to Bomarea
subg. Bomarea.

Paratypes. COLOMBIA. Cauea: Man. K1 Tambn, Par.
Nac. Nat. Munchique, T. Croat & ./. Baskin 80040 ( AFP.
CAUP, MO); Corn 20 de Julio, earn abandonada cerca do

2 km de la cabana del Inderena, It. Ruiz el al. 1180
(CAUP, COL, MO); vertiente occidental tlel no Munc¬
hique, //. Garc(a- Barriga et al. 12947 (COL); Mun. Tim-
bio, Km 54 of road from Timbio to 20 de Julio, It Maas
& T. Plowman 2144 (COL, F). Putumayo: Mun. San
Francisco, earn bacia FI Mirador, It. Guarin 792 (I’SO).

2. Bomarea euryantha F. Alzate, sp. nov. TYPE:
Colombia. Cundinamarca: Mun. Alban, 2300
m, 17 Apr. 1950, J. Idrobo BOO (holotype,
COL). Figure 2.

Herba sarmentosa; caule glabro. Lamina ovata, glabra.
Inflorescentia simplex; bracteis foliaceis, 5.5—10.0 X 2.0—
4.5 cm; bracteolis foliaceis, 5—7 X 2—4 mm. Flores cam-
panuli, segmentis perianthii aequalibus; sepalis 5.0— 5.5
cm, brunneo-guttalis; petalis viridibus, roseo-maculatis;
ovario glabro.

Sarmentose herb; roots not seen; stem 3—5 mm
diam., terete, glabrous, pendent at apex; internodes
5.0— 6.5 cm. Leaves pseudopetiolate, resupinate;

pseudopetiole 8-15 mm long, glabrous; blade
10.5—15.0 X 4.5— 6.5 cm, ovate, glabrous, granular
above, pustulate beneath between major veins, car¬
tilaginous when dry, the base obtuse to rounded,
the apex acuminate, the margin distinctly revolute.
Inflorescence simple; bracts leaflike, petiolate, 5.5—
10.0 X 2.0— 4.5 cm, ovate, glabrous, obtuse at base,
acuminate al apex, the margin entire; bracleoles
leaflike, sessile, 5—7 X 2—4 mm, linear to lanceo¬
late, glabrous, acute to acuminate at apex, the mar¬
gin entire; rays 3, 7.0— 8.5 cm long. Flowers with
perianth widely campanulate, the segments equal
in length; outer lepals 5.0— 5.5 cm long, ovate, pink
with brown spots, glabrous, obtuse at apex, the claw
3—4 mm long; inner lepals 5.0— 5.5 cm long, oblan¬
ceolate, green with pink spots and purple costa,
glabrous, rounded to obtuse at apex, tbe claw 2—3
mm long; filaments 30—44 mm long, glabrous; an¬
thers 5—8 mm long; ovary fully inferior, turbinate,
rugose; stvle 40—48 mm long, glabrous. Fruit and
seeds not seen.

Distribution and habitat. Known only from two
collections, in the central region of the Cordillera
Oriental of Colombia, at elevations between 2000
and 2400 m. The habitat of the type collection was
not described by tin* collector, but to judge by the
geographical data, it inhabits wet montane forests
(Holdridge, 1978).

Phenology. Flowering in April and August.

Etymology. The epithet makes reference to the
perianthal amplitude, observed in the large flowers
of this species.

Taxonomic relationships. I bis species is related
to Bomarea andreana Baker in having large flowers
(5.0— 5.5 cm long) and a simple inflorescence type.
However, it differs by its low number of flowers (3)
with a very wide perianth, which is a diagnostic
character of this species. Bomarea euryantha is in¬
cluded in Bomarea subg. Bomarea because of its
elongated, sarmentose stems, resupinate leaves,
and fully inferior ovary. Definite subgeneric place¬
ment is not possible without knowledge of the fruit.

Paralype. COLOMBIA. Cundinamarca: Mun. Al¬
ban. vereda La Maria, Oranjas del Padre Luna, M. Amaya
& ./. Granados 737 (COL).

3. Bomarea straiuinea Fillip ex F. Alzate, sp.
nov. TYPE: Colombia. Valle del Cauea: Mun.
Cali, Los Farallones. extreme N, vertiente ori¬
ental, entre Alto del Buey y Las Cascadas,
bosques, 3250— 3050 m, 13 Oct. 1944, J. Cua-
trecasas IBI02 (holotype, VALLE; isotype,
LIS). Figure 3.

Herba suberecta; caule glabro. Lamina lanceolata, 5.5—

256

Novon

Figure 2. Homarea euryantha F. Alzate. — A. Plant. — lb Lower surface of leaf. — C. Ovary, external view. Drawn
from ./. ldrobo 300 (COL).

8.0 X 1.0- 1.6 cm. infera (olia glabra, nervis sparse pu-
bescentibus. Inllorcscenlia composita; bracteis foliaceis;
bracteolis foliaceis. pubescentibus; radiis 3 ad 5. 2- vel
3-ramosis, glabris, glabrescentibusve. Flores penduli,
campanulati: sepalis aurantiaco-rubris, apice v i ride; pc-
talis aurantiaco-rubris, purpureo-maculatis; ovario pubes-
ecnte.

Suberect herb; roots not seen; stem 3—5 mm
diam., glabrous; internodes 0.5— 2.2 cm long.

Leaves pseudopetiolate, resupinate; pseudopetiole
3—5 mm long, sparsely pubescent; blade 5.5— 8.0 X
1.0-1. 6 cm, lanceolate, glabrous above, the nerves
glabrous to sparsely pubescent beneath, tin1 inter¬
nerves papillose or reticulate, coriaceous when dry,
the base acute to attenuate, the apex acute to acu¬
minate, the margin entire, revolute. Inflorescence
compound; bracts leaflike, pseudopetiolate, 3.5— 7.5

Volume 15, Number 2
2005

Alzate

Bomarea from Colombia

257

Figure 3. Bomarea straminea Fillip ex K Alzate. — A. Plant. — B. Lower surface of leaf. C. Ovary, external view.
Drawn from ./. Cuatrecasas 18102 (VALLE).

X 0.6— 1.2 cm, lanceolate, glabrous to glabrescenl.
acute at base and apex, the margin moderately rev¬
olute; bracteoles leaflike, sessile, 3.0— 4.0 X 0.3—
0.8 cm, lanceolate, pubescent, acuminate at apex,
the margin revolute; rays 3 to 5. 7—12 cm long, 2-
or 3-branched, glabrous to glabrescent; pedicels
1 .5-3.5 cm, glabrous. Flowers pendulous campan¬
il late; outer tepals 1.6— 2.0 cm long, ovate, orange-
red, green at apex, acute at apex, strigose to gla¬
brous. the claw 1 mm long; inner tepals 1 .6-2.0 cm
long, orange-red with purple spots and pink costa.

oblanceolate to spatulate. mucronate at apex, stri¬
gose; filaments 15-22 mm long, glabrous; anthers
3—4 mm long; ovary inferior, papillose, rugose to
strigose; style 18—24 mm long, pubescent. Capsule
turbinate, 1.8— 2.5 cm diam.; seeds 2—3 mm diam.,
obloid. the sareotesta bright orange.

Distribution and habitat. Bomarea straminea is
known from two disjunct localities in Colombia, one
from the Eastern Cordillera in the department of
Norte de Santander, and a second from the Western

258

Novon

Cordillera in Valle del Cauca where die species
grows in montane forests and paramo borders at
elevations of 2800—3250 m.

Phenology. This species has been observed in
flower and fruit during February, July, and October.

Etymology. The specific epithet is a reference
to the purple stems, observed principally in the
young apex. The name of this species was suggest¬
ed by Ellsworth I*, killip in several specimens in
major Colombian herbaria, but it was never pub¬
lished.

Taxonomic relationships. Bomarea straminea
seems to be related to B. lehmanni Maker because
ol its similar habit, compound umbels, and coria¬
ceous leaf texture. However, Bomarea straminea is
easily distinguished from this species by the num¬
ber of flowers per inflorescence ray (2 or 3) and its
larger flowers (1. 6-2.0 cm long).

Paratypes. COLOMBIA. Norte de Santander: Mini.
Pamplona, vertiente oriental, cerro al NK, paramo, J. Cua-
trecasas & II. Garc4a-Barriga 1020.1 (COL). Valle del
Cauca: Man. Cali. Cordillera Occidental, Los Karallones,
vertiente NW, quebrada de Kamos. ./. Cuatrecasas 21840
(VALLE).

Acknowledgments. My thanks in particular to
Ricardo Callejas ( 1 1 U A ) for his teaching and help
in my studies, R. E. Gereau (MO) for useful com¬
ments on tin' manuscript, and Francisco Morales
(I IN Rio) for reading the manuscript. 1 am thankful
to the staff of CAUP, COL, CUVC, F, IIUA, MO.
PSO, NY, and VALLE for tin- loan of herbarium
specimens and collaboration during visits. The ex¬

cellent drawings were made by Gloria Mora (HUA).

Financial support was received from the Universi-

dad de Anlioquia and the Missouri Botanical Gar¬
den.

Literature Cited

Aagesen. L. & A. Sanso. 2003. The phylogeny of the Al-
stroemeriaceae, based on morphology, rps 1 0 intron, and
rhe L sequence data. Syst. Bot. 28: 47-69.

Baker, J. G. 1878. A new key to the genera of Amarylli-
daceae. J. Bot. 16: 161—169.

- . 1882. On a collection of Bomares made by M. E.

Andre in New Granada and Ecuador. J. Bot. 20: 201-
206.

- . 1888. Handbook of the Amaryllideae. George

Bell & Sons, London.

Dalhgren, IT. II. Clifford & I’. Yeo. 1985. I'lie Families of
the1 Monocotyledons. Springer- Verlag, Berlin.

Holdridge, L. K. 1 97ft. Kcologfa Basada en Zonas de Vida.
Instituto fnteramericano de Ciencias Agrfcolas, San
Jos6, Costa Rica.

Killip. E. P. 1936a. Bomarea Midi. In: J. E. Macbride,
Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser.
13(1): 640-660.

- . 1936b. Bomarea, a genus of showy Andean

plants. Natl. Hort. Mag. 15: 115-128.

Mirbel. C. E. It. 1804. Bomarea. llistoire Naturelle. Gf1-
nerale et Particulifcre, des Plantes 9: 71.

I’ax, E k. Hoffmann. 1930. Amaryllidaceae. In: A. En-
gler, Nat. Pflanzenfam. (Aufl. 2) 15a: 391-430.

Sanso, A. & C. Xifreda. 2001. Generic delimitation be¬
tween Alstroemeria and Bomarea (Alstroemeriaeeae).
Ann. Bot. (London) 88: 1057-1069.

Smith, I). N. & It. f.. Gereau. 1991. Bomarea alhimontana
(Alstroemeriaeeae), a new species from high Andean
Peru. Candollea 46: 503—508.

Sodiro, A. I *408. Sertulae Florae Ecuadorensis 2. Quito,
Ecuador.

Dubautia kalalauensis , a New Species of the Hawaiian Silversword
Alliance (Compositae, Madiinae) from Northwestern Kaua'i, U.S.A.

Bruce G. Baldwin

Jepson Herbarium and Department <>1 Integrative Biology, 1001 Valley Life Sciences
Building, University of California, Berkeley, California 94720-2465, U.S. A.

bbaldwin@berkeley.edu

Gerald I). Carr

Department of Botany, University of Hawaii, 3190 Made Way, Honolulu. Hawaii 96822.

U.S.A. gerry@hawaii.edu

ABSTRACT. Dubautia kalalauensis B. G. Baldwin
& C. I). Carr is a newly described species o( the
Hawaiian silversword alliance (Compositae, Madi¬
inae) from wet forests in the vicinity of the rim of
Kalalau Valley. Kana'i. The taxon has been treated
within the circumscription of D. laxa Hooker &
Arnott subsp. hirsuta (Hillebrand) G. I). Carr, but
differs by having a more openly branching habit,
softer leaf surfaces, greenish cream (rather than
vellow) corollas, less glandular corolla tubes and
throats, and longer pappus elements. Based on mo¬
lecular phylogenetic data, D. kalalauensis is less
closely related to the widespread D. la.xa than to
other Kaua‘i endemics, including D. imbricata H.
St. John & C. I). Carr, I). kenwoodii G. I). Carr, D.
laevigata A. Gray, D. syndetica G. I). Carr A Uor-
ence, and I). waialealae Rock. Unlike all previ¬
ously described laxa of Dubautia Gaudiehaud-
Beaupre, D. kalalauensis combines opposite leaves,
eglandular peduncles, receptacular bracts through¬
out the heads (one per floret), greenish cream co¬
rollas, and pappi of setiform scales ca. 4 mm long.

Key words: Asteraceae, Compositae, Dubautia,
Hawaii, Kauai. Madiinae, silverswords, tarweeds.

The Hawaiian silversword alliance (Argyroxi-
phium DC., Dubautia Gaud ie baud -Beaupre, and
Wilkes in A. Gray) is most diverse on kana'i (Carr,
1985, 1999), the oldest high island of the archipel¬
ago (5.1 Ma; Clague & Dalrymple, 1987), where the
number of endemic angiosperm taxa in general is
higher than on any other island of the chain (Sakai
el al.. 2002). Both species of Wilkesia and all but 2
of 13 species of Dubautia on Kana'i are known only
from Kana'i (Carr, 1985, 1999). Based on results of
molecular phylogenetic analyses, the 15 indigenous
species on Kaua'i represent two of four major line¬
ages of the silversword alliance and may have de¬

scended from a common ancestor there (Baldwin &
Robiehaux, 1995; Baldwin, 1997. unpublished).

One major lineage of the silversword alliance from
Kaua‘i resolved with molecular data (nuclear ribo-
somal l)N A transcribed spacer sequences) compris¬
es all but three species of Dubautia known from the
island (i.e., all species except I), latifolia (A. Gray)
1). I). Keck, D. paleata A. Gray, and I), rail 'lard 'ioides
Hillebrand) and all taxa with nuclear genomic ar¬
rangements designated by Carr and Kyhos (1986) as
Dubautia Genome (DC) 1 and DG 3 (Baldwin. 1997.
unpublished). Members of the DG I + DG 3 lineage
are found in mesie. wet, or boggy habitats and in¬
clude trees, subscandent to erect shrubs, and cush¬
ion plants (Carr, 1985, 1999; Baldwin & Robie haux,
1995). Expanded sampling of taxa and DNA se¬
quences has led to resolution of a well -supported,
finer-scale lineage within the DG I + DG 3 group
that includes taxa of Dubautia with DG 3 or un¬
known genomic arrangements, including three of
four species of the silversword alliance described
since 1980: D. imbricata H. St. John & G. I). Carr.
D. kenwoodii G. D. Carr, and D. syndetica G. 1). Carr
& Lorence (Baldwin, unpublished), each of highly
limited geographic distribution (Carr. 1999). Here,
we describe yet another geographically restricted
species, from northwestern Kaua'i. that evidently be¬
longs to this same, recently resolved lineage (Bald¬
win, unpublished).

Dubautia kalalauensis B. G. Baldwin & G. I).
Carr. sp. nov. TYPE: U.S.A. Hawaii: Kaua'i.
Hanalei, Koke'e State Park, rim of Kalalau Val¬
ley, ca. 100 m N of Hwy. 550 (22°08'56. 1 "N.
1 59°38'10.4”W), along SW tributary of Kalalau
Stream, E slope, wet forest, 1205—1235 m. 6
Nov. 2004. B. (i. Baldwin & S. ./. Bain bridge
1441 (holotvpe. UC; isotypes. BISII. K HAW.
MO, NY. PTBG, US). Figure I.

Novon 15: 259-263. 2005.

260

Novon

Figure I. A— I',. Dulxiutia ktilnla uensis B. G. Baldwin iK G. I). Carr. — A. Flowering Branch. — B. Head wilh peduncle.
— Floret and associated receptacular bract. — I). Cypsela with pappus. — F,. Habit (central plant in foreground) and
wet-forest habitat. Drawn from the type.

Volume 15, Number 2
2005

Baldwin & Carr

Dubautia kalalauensis from Hawaii

261

A speeiebus ceteris Dubautiae differt characteribus
conjuncte: foliis oppositis, pedunculis eglandulosis, capi-
tulis bracteis receptaculorum ubique (1 bractea in qtioque
flosculo), eorollis viridulis-eburneis, squamis papporum
setiformibus ca. 1 mm longis.

Openly branching shrubs or trees 1.8— 3.5 in tall,
vegetative sterns ± strigose (trichomes often arcu¬
ate' or flexuous) and dark purple or black dislally,
glabrescent and pale brown proximally, leaf scars
glabrous or ± strigose; flowering stems ± strigose
or hispid-hirsute. Leaves opposite, sessile, com¬
monly 8—14 X 1.6— 3.2 cm (up to 20 X 5.6 cm in
juvenile plants), blades narrowly elliptic to oblan-
eeolate, lime green and moderately ± strigose
adaxially, yellow-green and somewhat more densely
and coarsely ± strigose abaxially; margins shallow¬
ly toothed along distal % to 34 of length, sparsely
ciliolate or eciliate; apices acuminate; bases atten¬
uate. petioloid; venation acrodromous, with 7 basal
to suprabasal veins. Inflorescences as heads, dis¬
coid, (6 to) 15 to 55. in dense, cymo-corymbiform
(Hat-topped or rounded) arrays, 4—10 cm long, 4—
10 cm wide, the stalks and peduncles densely his¬
pid-hirsute, eglandular, peduncles 0— 5(— 7) mm
long; receptacles glabrous, flat or convex, false in¬
volucres (of outermost reeeptacular bracts) cylin¬
drical to narrowly campanulate, reeeptacular bracts
8 to 16, one associated with (and partially envel¬
oping) each floret, linear to linear-lanceolate, free
or coaleseent to Vs of length proximally, the outer¬
most 4.5—6 mm long, usually green, sometimes pur¬
plish. ± strigose, eglandular; florets 8 to 16. corol¬
las exserted. greenish cream, aging pinkish to
reddish brown (often drying dark purple), 3—5.5
mm long, usually slightly exceeding pappi, tubes/
throats arcuate or recurved, glabrous or sparsely
glandular proximally, moderately glandular distally,
throats abruptly dilated distally, lobes narrowly tri¬
angular. usually reflexed. 0.8—1 mm long, sparsely
setulose abaxially; anthers reddish purple or ma¬
roon; cypselae black, straight or arcuate, ca. 2—3
nun long, ± strigose; pappi of 19 to 24, stramineous
to brown or reddish purple, mostly subequal, fim¬
briate, setiform scales ca. 4 mm long.

Distribution. The species is known only from
the vicinity of the southern rim of Kalalau Valley,
north of koke'e Road (Highway 550), from near
Kalalau Lookout east to slopes bordering a south¬
western tributary of Kalalau Stream, west of Ru’u o
kila. known collections span an elevational range
of 1205—1235 m.

Habitat. Dubautia kalalauensis occurs on
densely vegetated, windward slopes and at stream-
sides in wet forests. Along the tributary of kalalau
Stream west of Pu‘u o kila. the first author observed

26 plants (mostly juveniles) during a single traverse
from the upper eastern slope to the streamhed.
Woody associates include species of Metrosideros
Banks ex Gaertner, Scaevola L., Broussaisia Gau-
diehaud-Beaupre, Cheirodendron Nuttall ex See-
mann, Pittosporum Banks ex Gaertner, and Hedy-
otis L. Associated ferns include species oi
Elaphoglossum Schott ex J. Smith, Sadleria kaul-
fuss. Cibotium Kaulfuss, and Pneumatopteris Nakai.
Alien Rubus argutus Link (Florida blackberry ), He-
dychium gardnerianum ker Gawler (Kahili ginger),
and Passiflora mollissima (Kunth) L. H. Bailey (ba¬
nana poka) are well established. Passiflora mollis¬
sima appears to pose a particularly serious threat:
most large, flowering individuals of I), kalalauensis
observed by the first author at the type locality were
almost entirely obscured by a dense covering of
banana poka. Taxa of Dubautia observed in prox¬
imity to D. kalalauensis are D. knudsenii llille-
brand subsp. knudsenii , I). laevigata A. Gray, and
I), microcephala Skottsberg. Natural hybrids be¬
tween I). kalalauensis and D. laevigata ( Carr l i 70.
BISH. HAW, NY. UC, WS; Flynn & Lowrey I B 75.
PTBG), reported earlier by Carr (1999), have been
collected on and just below the rim of Kalalau Val¬
ley.

Phenology. Flowering specimens have been
collected from August to November.

Relationships. Members of Dubautia kala¬
lauensis were previously treated provisionally with¬
in the circumscription of the morphologically sim¬
ilar D. laxa Hooker & Arnott subsp. hirsuta
(Hillebrand) G. I). Carr (Carr, 1985, 1990. 1999).
Field observations in the 1980s by Tim Flynn, Cu¬
rator of the Herbarium at the National Tropical Bo¬
tanical Garden, first brought attention to these
plants from the vicinity of Kalalau rim that ap¬
peared to be members of I), laxa subsp. hirsuta
with white, rather than yellow, corollas. Based on
examination of specimens from the field and her¬
baria, Carr (1990: 301) indicated that the Kalalau
plants differed from other populations of I), laxa
subsp. hirsuta (including the type) in leaf vestiturc
(“finer pubescence”) as well as corolla color and
“may represent a new taxon." He later noted that
the Kalalau plants also differ from typical I). laxa
subsp. hirsuta by having longer pappus elements
and less glandular corolla tubes (Carr. 1999). Com¬
parison of 1 8S-26S ribosomal l)NA sequences of
the external and internal transcribed spacers (ETS
and ITS) with those of all other recognized taxa of
the silversword alliance revealed that the Kalalau
plants represent a unique lineage within a well-
supported elade that also includes D. imbricata, D.

262

Novon

kenwoodii, I). laevigata , I), syndetica, and D. wai-
alealae Rock (Baldwin, unpublished).

The relationship between D. kalalauensis and D.
laxa is sufficiently remote based on rl)NA data to
lead us to conclude that similarities between the
two taxa are likely the result of evolutionary con¬
vergence and shared primitive (plesiomorphic)
characteristics. In hindsight, the presence of only
moderate, abaxial glandulosity on the corolla tidies
and throats of I). kalalauensis is perhaps the best
morphological evidence for a distant relationship
with / ). laxa ; densely and coarsely glandular co¬
rolla tubes arc* diagnostic and universal for the1 lin¬
e-age comprising I). laxa. I ). knudsenii , and I). pau-
cijlorula H. St. John <A G. I). Carr (Carr, 1985;
Baldwin, 1997). Ancient gene flow across lineages
cannot be ruled out as a potential explanation for
shared morphological c haracteristics of D. kala¬
lauensis and I). laxa subsp. hirsuta, although rDNA
sequences provide no evidence consistent with that
possibility. Cytogenetic evidence from a natural hy¬
brid between I). kalalauensis and I). laevigata
(Carr, 1999) and from artificial hybrids involving
I). laxa. I). laevigata, and other species of Duhau-
tia (Carr, 1985; Carr & Kyhos, 1986) demonstrates
that I). kalalauensis and D. laxa are, at most, dif¬
ferentiated genomicalb bv two reciprocal translo¬
cations, a minor barrier to introgression involving
non-recombined regions of the genomes (see Carr,
1995). Opportunities for modern gene exchange* ap¬
pear to be limited extrinsic-ally; the two taxa arc1 not
known to co-oceur and may be ecologically distinct,
with I). laxa subsp. hirsuta occurring largely in
boggy situations and on foggy, wind-swept ridges,
rather than in wet forests (Carr, 1985, 1999). Di¬
vergence in ecological and physiological features
associated with occupancy of different habitats has
been a major feature of diversification of the si I -
versword alliance (Robichaux et ah, 1990; Baldwin
<A Robichaux, 1995) and may be sufficient under
natural conditions to prevent introgressive hybrid¬
ization between interfertile taxa, e.g., through post¬
dispersal selec tion against hybrids (Kyhos et ah,
1981; Carr, 1995).

Etymology. The species epithet refers to the
rim of Kalalau Valley, Kaua‘i, where all known pop¬
ulations of the taxon occur.

Conservation status. Duhautia kalalauensis ap¬
pears to be in danger of extinction based in part on
its rarity and small geographic: range. As noted
above (under “ Habitat ”), invasive plants threaten
the* only known, sizable population, at the type lo¬
cality, where measures to control Passijlora mollis-
sitna (especially). Ruhus argutus, and Hedyehium
gardnerianum may be necessary to prevent extir¬

pation of I). kalalauensis. Efforts to relocate histor¬
ical occurrences and discover additional popula¬
tions along the rim of Kalalau Valley are warranted.

Paratypes. I .S.A. Hawaii: Kaua'i, Hanalei, KokeV
State Park, E rim of Kalalau Valley, 0. Degener 23854
(lilSH, UC); Kalalau Pali, C. A'. Forbes I034.K (HISII),
Forbes 1697. k ( It I S 1 1 ); just below lookout, S. Carlquisl
1112 (ItlSIl. UC); Hwy. 550 near mi. 18.3, at back of Ka¬
lalau Valley, 7: lh Flynn 1171 (BISH, HAW), Flynn 1352
(PTBG); rim of Kalalau Valley near Kaluapulii Trailhead
betw. Kalalau <A Pu'u o Kila lookouts, G. I). Carr & Flynn
1330 (ItlSIl): just below trail along Kalalau Valiev rim,
Flynn & /.’ K. hnorey 1871 (ItlSIl. ITBG).

Acknowledgments. Research support was pro¬
vided in part by a grant from the National Science
f oundation (DEB-9458237, to BGB) and an award
from the U.C. Berkeley Chancellor's f und (to BGB).
We thank Timothy W. Flynn for originally bringing
our attention to the uniqueness of the Kalalau
plants described here. Susan J. Bainbridge for field
assistance, Bridget E. Wessa for lab assistance,
John L. Strother and Alan R. Smith for assisting
with the Latin diagnosis and for reviewing the man¬
uscript, Lesley Randall (or preparing the illustra¬
tions, and Wayne Souza (Hawaii Division of State
Parks) for permission to collect Duhautia in Koke'e
State Park.

I .iterature Cited

Baldwin, B. G. 1997. Adaptive radiation of the Hawaiian
silversword alliance: Congruence and conflict of phy¬
logenetic evidence from molecular and non-molecular
investigations. Pp. 103—128 in T. J. Givnish <A K. J.
Sytsma (editors). Molecular Evolution and Adaptive Ra¬
diation. Cambridge Univ. Press, Cambridge.

- <A R. 11. Robichaux. 1995. Historical biogeogra¬
phy and ecology of the Hawaiian silversword alliance
(Asteraceae). Pp. 259-287 in W. L Wagner & V. A.
Funk (editors), Hawaiian Biogeography: Evolution on a
Hot Spot Archipelago. Smithsonian Institution Press.
Washington. P.C.

Carr, G. I). 1985. Monograph of the Hawaiian Madiinae
(Asteraceae): Argyroxiphium, Dubautia. and Wilkesia.
Allerlonia 4: 1-123.

- . 1990. Dubautia Gaud. Pp. 292-308 in W. L.

Wagner. I). R. Ilerbst & S. H. Solimer, Manual of the
f lowering Plants of Hawai'i, Vol. I. Univ. Hawaii Press
and Bishop Museum Press, Honolulu.

- . 1995. A fully fertile intergeneric hybrid deriva¬
tive from \rgyroxiphium sandwicen.se subsp. macroce-
phalum X Dubautia tnenzie.sH (Asteraceae) and its rel¬
evance to plant evolution in the Hawaiian Islands.
Amer. J. Bot. 82: 1574-1581.

- . 1999. A reassessment of Dubautia (Asteraceae:

Heliantheae— Madiinae) on kaua'i. Pacific Sei. 53: I 14—
1 58.

- <A I). \\. Kyhos. 1986. Adaptive radiation in the

Hawaiian silversword alliance (Compositae— Madiinae).
II. Cytogenetics of artificial and natural hybrids. Evo¬
lution 40: 959-976.

Clague, I). A. tA G. B. Dalrymple. 1987. The Hawaiian-

Volume 15, Number 2
2005

Baldwin & Carr

Dubautia kalalauensis from Hawaii

263

Kmperor volcanic chain. Pp. L— 54 in R. W. Decker, T.
I.. Wright & P. H. Stauffer (editors), Volcanism in Ha¬
waii. I .S. Geological Service Professional Paper 1350.
LJ.S. Government Printing Office, Washington. D.C.
Kyhos, I). W., C. Clark & \\. C. Thompson. 1981. The
hybrid nature of Enceliu laciniata (Compositae: Helian-
theae) and the control of population composition by
post-dispersal selection. Syst. Hot. 6: 399—41 I.

Robiehaux, R. H., G. I). Carr. M. Ciebman & R. W. Pear-
cy. 1990. Adaptive radiation of the Hawaiian silver-
sword alliance (Compositae-Madiinae): Ecological,
morphological, and physiological diversity. Ann. Mis¬
souri Rot. Gard. 77: 61—72.

Sakai. A. K., W. E. Wagner & E. A. Mehrhoff. 2002. Pat¬
terns of endangerment in the Hawaiian flora. Syst. Biol.
51: 276-302.

INew Combinations in Chinese Polyspora (Theaceae)

Bruce Bartholomew

Botany Department, California Academy of Sciences, 875 Howard Street, San Francisco,
California 94103-3009, U.S.A. hbartholomew@calacademy.org

Ming Tienlu

Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming,
Yunnan 650204, People’s Republic of China

Ahsthact. For the Theaceae treatment in the
Flora of China, volume 12, it was found that sev¬
eral specific epithets have not yet been validly
transferred from Gordonia Ellis to Polyspora Sweet
ex G. Don. In this article the following five species
are transferred from Gordonia to Polyspora : Polys¬
pora chrysandra (Cowan) I In ex B. Bartholomew &
T. L. Ming, P. hainanensis (I lung T. Chang) C. X.
Ye ex B. Bartholomew & T. h. Ming, P. longicarpa
(Hung T. Chang) C. X. Ye ex B. Bartholomew & I.
h. Ming. P. speciosa (Kochs) B. Bartholomew <X T.
L Ming, and P. tonkinensis (Pitard) B. Bartholomew
& T. L. Ming. Three species. Gordonia axillaris
Roxburgh ex Ker Gawler, Gordonia tonkinensis Pi¬
tard, and Tliea speciosa Kochs, as well as the one
variety Gordonia axillaris var. acuminata E. Pritzel,
are lectotypified.

Key words: China, Gordonia, Polyspora, Thea¬
ceae.

Recent molecular evidence has shown that the
Chinese species often treated as Gordonut Ellis are
not closely related lo the North American species
G. lasianthus (I,.) Ellis, the type species of the ge¬
nus, and should he treated as a different genus
(Prince & Parks, 2001; Yang ('I ah. 2004). On the
basis of priority, the generic' name that must he
used for these species is Polyspora Sweet ex G.
Don. Although several of the Chinese species have
been treated in the literature as Polyspora (Hu,
1963; Prince & Parks, 2001; Yang cl ah, 2004; Ye,
1990), for the recognized laxa only the combination
Polyspora axillaris (Roxburgh ex Ker Gawler)
Sweet ex G. Don has been validly made. In order
to treat the Chinese species of Polyspora in the
Flora of China, volume 12, five combinations must
be validly made. The six species of Polyspora that
occur in China include the following:

I . Polyspora axillaris (Roxburgh ex Ker Gawler)
Sweet ex G. Don, Gen. Hist. I: 574. 1831.

Basionym: Camellia axillaris Roxburgh ex Ker
Gawler. Rot. Reg. 4: t. 349. 1818. Gordonia
axillaris (Roxburgh ex Ker Gawler) I). Die¬
trich. Syn. PI. 4: 863. 1847. TYPE: plate in
the original protologue, Bot. Reg. 4: t. 349.
1831 (lectotype, designated here).

This species occurs in forests and thickets from
100 to 800(— 2300) m in Guangdong, Guangxi, Hai¬
nan, and Taiwan, China (Chang, 1998), and Viet¬
nam (Ho, 1991).

There is no type specimen for this species, so
the plate in the original protologue is chosen as the
lectotype.

2. Polyspora chrysandra (Cowan) Hu ex B. Bar¬

tholomew & T. E. Ming, comb. nov. Basionym:
Gordonia chrysandra Cowan, Notes Roy. Bot.
Card. Edinburgh 10: 184. 1931 . TYPE: China.
Yunnan: lava bed W of Tengyueh. 25°N. 5000
ft., Nov. 1912, G. Forrest 9234 (holotype, E;
isotype, A).

This species occurs in forests and thickets from
1100 to 2400 m in Guizhou, Sichuan, and north¬
western Yunnan, China, and northern Myanmar
(Chang, 1998; Ming. 1997).

H. II. Hu proposed the combination Polyspora
chrysandra (Hu, 1972) but did not include the bas¬
ionym reference citation.

3. Polyspora hainanensis (Hung T. Chang) C. X.

Ye ex B. Bartholomew & T. E. Ming, comb,
nov. Basionym: Gordonia hainanensis Hung T.
Chang, Acta Sei. Nat. Univ. Sunyatseni 22(2):
111. 1983. TYPE: China. Hainan: Jianfeng-
ling, T. (J. Yu 11206 (holotype, SYS).

This species occurs in forests from 300 to 1500
m in Hainan, China (Chang, 1998).

(7 X. Ye proposed the combination Polyspora
hainanensis (Ye, 1990) but did not include the bas-
ionyin reference citation.

IMovon 15: 264—266. 2005.

Volume 15, Number 2
2005

Bartholomew & Ming
Chinese Polyspora

265

4. Polyspora lougirarpu (Hung T. Chang) C. X.

Ye ex B. Bartholomew & T. L. Ming, comb,
nov. Basionym: Gordonia longicarpa Hung I.
Chang, Acta Sei. Nat. Univ. Sunyatseni 22(2):
111. 1983. TYPE: China. Yunnan: Tenge hong.
./. Chen 2-77 (holotype, KUN).

This species occurs in forests from (1000— )1 700
to 2500 m in southeastern to southwestern Yunnan
China, northern Myanmar, northeastern Thailand,
and northern Vietnam (Chang. 1998; Keng, 1972;
Ming, 1997). Keng (1972) recorded Gordonia ax¬
illaris as occurring in northeastern Thailand, which
was before G. longicarpa had been named, but from
his description the plant is clearly Polyspora lon¬
gicarpa.

C. X. Ye proposed the combination Polyspora
longicarpa (Ye, 1990) but did not include the bas¬
ionym reference citation.

5. Polyspora speciosa (Kochs) B. Bartholomew

& T. L. Ming, comb. nov. Basionym: Thea spe¬
ciosa Kochs, Bot. Jahrb. Syst. 27: 597. 1900.
Camellia speciosa (Kochs) Cohen-Stuart, Med-
ed. Proefstat. Thee 60: 67. 1916. TYPE: Chi¬
na. Sichuan: Maup’o shan, im Urwald, 3 Sep.
1891, C. Hock ' & A. von Rost horn 754 |as
“454" | (holotype, B destroyed; leetotype, des¬
ignated here. ( iZU).

Gordonia axillaris (Roxburgh ex Ker Gawler) Sweet ex G.
Don var. acuminata K. Pritzel, Bot. Jahrb. Syst. 29:
473. 1900. Gordonia acuminata (E. Pritzel) flung T.
Chang, Acta Sei. Nat. Univ. Sunyatseni 22(2): I 12.
1983. TYPE: China. Sichuan: Nan ch'uan. fa ho
kou. 31 Julv 1891. C. Rock & A. von Rost horn 147
(syntvpe, B destroyed; leetotype, designated here, <);
isotypes, GZU. 0). China. Sichuan: Mao p’o shan.
Sep. 1891. C. Rock & A. von Rost horn 754 (syntvpe,
B destroyed; isosyntype, GZU).

Gordonia kwangsiensis Hung T. Chang, Acta Sei. Nat.
Univ. Sunyatseni 22: 112(2). 1983. UY PI*,: China.
Guangxi: Damiao Mountain, S. II. Glum 16467 (ho¬
lotype, I BSC).

This species occurs in forests from 1200 to 2000
m in northern Guangxi, Guizhou, southern to south¬
western Sichuan (including the Chongqing Munic¬
ipality, formerly part of Sichuan), and Yunnan. Chi¬
na (Chang, 1998).

The only specimen cited by Kochs (1900) for
Then speciosa was “Bock & von Rosthorn 454.
Although the holotype at B is lost, there is com¬
pelling evidence that 454 is an error, and the num¬
ber in the protologue should have been 754. By
checking tin* dates and collection numbers cited in
Diels (1900—1901) it is evident that Hock & von
Rosthorn numbers between at least 192 and 707
were collected in August 1891. In the protologue

for Thea speciosa, Kochs stated that the collection
was made in September, whereas Bock cY von Ros¬
thorn numbers for September 1891 started with
708. The holotype of Thea speciosa was not only
collected on the same date, but also at the same
Chinese collection locality (with minor transcrip¬
tion differences in spelling) as Bock & Rosthorn
754. Furthermore, in the treatment for Simaruba-
ceae by Pritzel in Diels (1900—1901), Bock & von
Rosthern 454 is identified as Picrasma ailanthoides
(Bunge) Planchon. Unfortunately, no archival infor¬
mation that might throw additional light on this
matter is available at B (Robert Vogt, pers. comm.).
GZU (Anton Drescher, pers. comm.), or 0 (Per
Sunding, pers. comm.). Quite likely collection num¬
ber 754 was misread as 454 either when the hand¬
written label on the specimen was transcribed or
possibly when a handwritten manuscript was set in
type.

In flower Polyspora and Camellia (Thea) are very
similar, and this no doubt was why Kochs originally
placed the flowering specimen of Bock cY von Ros-
llwrn 754 in Thea. However, because the fruit of
Polyspora and Camellia (Thea) are very different,
once it was realized that the fruiting specimen of
Bock cY von Rosthorn 147 was the same species, it
would have been clear that Thea speciosa was in
the wrong genus. In the treatment of Theaceae in
Die Flora von Central-China (Diels. 1900—1901),
Pritzel and Kochs realized that T. speciosa should
be treated as a Gordonia. This treatment in Diels’s
article was by both Kochs (who authored Thea) and
Pritzel (who authored the rest of the family). Pritzel
did not reduce Thea speciosa to a variety of Gor¬
donia axillaris, but rather selected a different in¬
fraspecific epithet, based his Gordonia axillaris var.
acuminata E. Pritzel on two syntypes (one of w hich
was the- holotype of T. speciosa), and gave T. speciosa
Kochs as a synonym for his new variety. However,
at the1 specific level T. speciosa has priority, so Pritz-
el’s specific epithet speciosa must be used when
treating this taxon as a species. Because the holo¬
type of T. speciosa and both syntypes of G. axillaris
var. acuminata have been destroyed, it is appro¬
priate to lectotypify T. speciosa on the only known
isotype and to lectotypify G. axillaris var. acumi¬
nata on one of the extant fruiting isosyntypes.

G. X. Ye proposed the combination Polyspora
kwangsiensis (Ye, 1990) but did not include the
basionym reference citation and misspelled the
specific epithet as “ gwangsiensis .” However, this
combination is not being validated here because
the species is now reduced to a synonym. II. I.
Chang proposed the species Gordonia szechiianensis
(Chang, 1983) for what is the same species as Po-

266

Novon

lyspora speciosa, but he did not supply Latin de¬
scription or diagnosis.

6. Polyspora tnnkineusis (Pitard) 1L Bartholo¬
mew & T. L. Ming, comb. nov. Basionym: Gor-
donia tonkinemis Pitard, in Lecomte, FI. Gen.
Indo-Chine 1: 348. 1910. TYPE: Vietnam.
Tonkin environ de Ninh-binh. 1883—1891,
Abbe R. P. Ron 60 (lectotype, designated liere,

P).

This species occurs in forests and thickets from
1200 to 2100 m in southern Guangxi, southeastern
Yunnan, China, and northern Vietnam (Ming,
1997).

km to the Species oe Podspor i in China

la. I.cal blade apex rounded, obtuse, or emarginate.

2a. Petiole 10—15 mm, glabrous; leaf blade mar¬
gin entire or apieally remotely erenate . . .

. I . P. axil laris

2b. Petiole 3—5 mm, oppressed pubescent; leaf

blade margin serrate . 2. P. chrysandra

lb. Leaf blade apex bluntly acute, acuminate, or
shortly acuminate.

3a. Current year branchlets, petioles, leaf
blades abaxially. and filaments basal ly all
with trichomes; capsule 4-5 cm long ....

. 4. P. longicarpa

3b. Current year branchlets, petioles, leaf
blades, and filaments glabrous (branchlets
subglabrous in P. tonkinensis); capsule 1—3.5
cm long.

4a. Young branches slender; leaf blade nar¬
rowly oblong to oblanceolate, 8—13 X

2—3 cm; flowers ca. 4 cm diam .

. 3. P. hainanesis

lb. Young branches stout; leaf blade elliptic
to oblong-elliptic, 9—18 X 3—6 cm;
flowers 5 cm or more diam.

5a. Terminal buds large, purplish red,
glabrous; petiole 1.5—2 cm: flowers
8-10 cm diam.; style ca. 2 cm . .

. 5. P. speciosa

5b. Terminal buds small, white seri¬
ceous; petiole 0.8— 1.5 cm; flowers
5—6 cm diam.; style ca. 1 .5 cm . .
. 6. P. tonkinensis

Acknowledgments. We thank Thomas F. Daniel
(GAS), Anton Drescher (GZU), Peter W. Fritsch
(CAS), Per Sunding (0), and Robert Vogt (B) for
assistance during preparation of this article.

Literature Cited

Chang, H. T. 1983. Gordonia. P. 469 in Anonymous, Icon-
ographia Cormophytorum Sinicorum Supplementum II.
Science Press, Beijing.

- . 1998. Gordonia. FI. Keipubl. Popularis Sin.

19(3): 206-2 1 I. Science Press, Beijing.

Diels, L. 1900-1901. Die flora von Central-China. Bot.
Jahrb. Syst. 29; 169-659.

Ho, P. H. 1991. Gordonia axillaris. III. FI. Vietnam 1(1):

537. Pham Ho&ng Hy, Montreal.

Hu, H. H. 1963. Theaceae. P. 345 in W. 5. Chun. Mate¬
rials for the Flora of Hainan. Acta Phytotax. Sin. 8:
341-357.

Keng. II. 1972. Gordonia axillaris. FI. Thailand 2(2): 144.

Applied Scientific Research Corporation, Bangkok.
Kochs, J. 1900. fiber die Gattung Tliea und der ehine-
sisehen Thee. Bot. Jahrb. Syst. 27: 577—635.

Ming, T. L. 1997. Gordonia. FI. Yunnan. 8: 315-319.
Prince, L. M. & C. R. Parks. 2001. Phylogenetic relation-
shi, >s of Theaceae inferred from chloroplast DNA se¬
quence data. Amer. .1. Bot. 88: 2309—2320.

Yang, S. X.. J. B. Yang. L. (7 Lei, I). Z. I.i, II. Yoshino
X I. Ikeda. 2004. Reassessing the relationships be¬
tween Gordonia and Polysftora (Theaceae) based on the
combined analysis of molecular data from the nuclear,
plastic! and mitochondrial genomes. PI. Svst. Kvol. 248:
45-55.

Ye, C. X. 1990. The range of Gordonieae (Theaceae) and
limitation of genera in the tribe. Guihaia 10: 99-10.3.

Pennellia brachycarpa (Brassicaceae), a New Species from

Jujuy, Argentina

Mark Beilstein

Department of Biology, University of Missouri— St. Louis. 8001 Natural Bridge Road,
St. Louis, Missouri 63121, U.S.A. inab347@studentniail.umsl.edu

Ihsan A. Al-Shehbaz

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

ihsan.al-shehbaz@mobot.org

ABSTRACT. Pennellia brachycarpa (Brassicaceae),
a new species from the province of Jujuy in north¬
ern Argentina, is described and illustrated. The
new species is distinguished from other members
of die genus by having corymbose rather than lax
racemes and forked and Y-shaped instead of den¬
dritic triehomes.

key words: Argentina, Brassicaceae, Jujuy,
Pennellia.

Pennellia Nieuwland (Brassicaceae) is a genus
of 7 to 1 1 species distributed in die southern Unit¬
ed States, Mexico, Central America, Colombia, and
disjunctly into Bolivia and northern Argentina
(Rollins, 1980; Appel A Al-Shehbaz, 2003; Fuen-
tes-Soriano. 2004). The genus is easily distin¬
guished from its nearest relatives in the Halimolo-
bine clade (Bailey et al., 2002), which consists of
Halimolobos Tauscb, Mancoa Weddell, Pennellia,
and Sphaerocardamum. S. Sehauer, by its cup¬
shaped flowers, purple to purple-tipped sepals, and
petals sub-equaling or slightly longer than sepals.

the novelty described herein, Pennellia brachy¬
carpa, was collected during fieldwork in northern
Argentina in connection with a broad, ongoing phy¬
logenetic study of the family Brassicaceae. Phylo¬
genetic results from the chloroplast gene ndhY in¬
dicate that this species is closely related to die
North American P. longijolia (Bentham) Rollins, a
species distributed in Arizona, New Mexico, Texas,
and southward throughout Mexico to Guatemala
(Fuentes-Soriano, 2004). Pennellia was previously
known from Argentina and Bolivia only by P. bo-
liviensis (Muschler) Al-Shehbaz (Al-Shehbaz. 1990;
Fuentes-Soriano, 2004). The inclusion of I1, bra¬
chycarpa in Pennellia reinforces the southern dis¬
tribution of the genus.

IVuiicIliu brachycarpa Beilstein N Al-Shehbaz.
sp. nov. TYPE: Argentina. Jujuy: Abra Pampa,
S of Abra Pampa City off route 9, among rocks
on isolated hilltop, 3650 m, 22°49.352'S,
65°41 .324 AY. 10 Feb. 2003. Mark Beilstein.
Noah Whiteman & Donna Eakman 03-148
(holotype, M0). Figure I.

Media perennis 7.5—27 cm alta, pilis furcatis brevi-stip-
ilatis et simplicihus praedita. Kolia basal ia oblaneeolato-
spathulata, 1—3 X 0.3—1 cm, margine subintegra vel ser-
rufata; folia caulina sessilia, non auriculata, 0.7— 2.6 cm
X 1—5 mm. Racemi 7— 30-ffori, ebracteati: pedicelli frue-
tiferi tenues, recti, 7—10 mm longi. Sepala oblonga, sparse
pilosa. ea. 2 X I mm; petala alba, anguste spathulata, 2
2.5 X 0.6— 0.8 mm; ovula 50—64; stylo 0.1— 0.2 mm longo.
Fruclus lineares, 1.3— 1.7 cm X ea. I mm, teretes, glabri;
semina ovata, ca. 0.25 X 1 mm, subbiseriata.

Plants perennial, 7.5—27 cm tall: triehomes of
stems, leaves, and sepals short- stalked and forked,
these mixed with simple ones along stem and leaf
midvein, to 0.3 mm long, rarely a few dendritic
ones on leaf margin; stems erect, single, few-
branehed, and glabrous above. Basal leaves sub-
sessile, oblaneeolate-spatulate, 1—3 X 0.3—1 cm,
base attenuate, margin subentire to serrulate, apex
obtuse; cauline leaves sessile, not auriculate at
base, oblong-linear, 0.7— 2.6 cm X 1—5 mm, margin
entire, apex subacute. Raceme ebracteate, corym¬
bose, 7- to 30-flowered, rachis straight; fruiting
pedicels very sparsely hairy, ascending, straight,
slender, 7—10 nun long. Sepals oblong, green with
purple tips, ca. 2 X 1 mm, not saccate, sparsely
pubescent below apex; petals white, narrowly spat-
ulate, not clawed, 2—2.5 X 0.6— 0.8 nun: filaments
1 — 1.2 mm long; anthers ovate, ca. 0.6 mm long;
ovules 50 to 64; style 0.1— 0.2 mm. Fruit linear,
1 .3— 1.7 cm X ca. I mm, terete, slightly curved;
valves glabrous, smooth; midvein distinct basally,
obscure distally; stigma entire; seeds subbiseriate,
brown, ovate, ca. 0.25 X 1 mm.

Novon 15: 267-269. 2005.

268

Novon

Figure 1. Pennell'ui brachycarpa Beilstein & Al-Sliehbaz. — A. Plant. — B. Portion ol infructeseence. — (i. Triehome.s.
— I). Sepal. — E. Petal. — F. Fruit. Seale: A, B. 1). E = I mm; C = 0.1 mm; I' = 5 mm. Drawn by Al-Shehbaz from
the holotype (Beilstein, Whiteman & Eakman 03-148, MO).

Volume 15, Number 2
2005

Beilstein & Al-Shehbaz

Pennellia brachycarpa from Argentina

269

Pennellia brachycarpa, which is known only from
the holotype specimen, is easily distinguished from
the other species of Pennellia hv having corymbose
instead of lax racemes and forked and Y-shaped
instead of dendritic trichomes. It is related to a
group of four species (the South American P. boli-
viensis and the North American P. patens ((). E.
Schulz) Hollins. P. micrantha (A. Gray) Nieuwland.
P. lasiocalycina (O. E. Schulz) Rollins) with terete
to subterete. ascending to erect fruits (Fuentes-So-
riano, 2004). From these. P. brachycarpa is also
distinguished by its shorter (1.3— 1.7 cm) instead of
longer (more than 2 cm) fruits.

Ongoing studies on the South American genera
of Brassicaceae should clarify generic boundaries
and establish relationships among and within gen¬
era.

Acknowledgments. We are grateful to Victoria

C. Hollowell and two anonymous reviewers for their
comments on the manuscript. We thank Sandra Al-
iscioni for help in the field.

Literature Cited

Al-Shehbaz. I. A. 1990. New or noteworthy species in the
South American genera Mancoa, Pennellia, and Sis¬
ymbrium (Brassicaceae). Harvard Pap. Bot. 2: 11-16.
Appel. 0. & 1. A. Al-Shehbaz. 2003. Brassicaceae. Pp.
75—174 in k. Kubitzki (editor). The Families and Gen¬
era of Vascular Plants. Vol. 5. Springer- Verlag. Berlin.
Heidelberg & New York.

Bailey. C. I).. B. A. Price & J. J. Doyle. 2002. Systematic*
of the halimolobine Brassicaceae: Evidence from three
loei and morphology. Syst. Bot. 27: 318—332.
Fuentes-Soriano, S. 2004. A taxonomic revision of Pen¬
nellia (Brassicaceae). Harvard Pap. Bot. 8: 173—202.
Rollins. R. G. 1980. The genus Pennellia (Cruciferae) in
North America. Contr. Gray Herb. 210: 5-21.

Inga colonchensis (Fabaceae, Mimosoideae), una Nueva Endemica
del Bosque Seco Tropical en Ecuador

Xavier Cornejo y Carmen Bonifaz

Herbario GUAY, Facultad de Ciencias IV at males, Universidad de Guayaquil.

Casilla 09-01-10634, ( mayaquil. Keuador. xeornejoguay@hotmail.com

Rest MKN. Se describe Inga colonchensis romo un
nuevo taxon del fraginentado bos(|ue seco tropical
de la cordillera Chongon-Colonche (Prov. Guayas),
en la region occidental del Keuador. Por su inflo-
rcsccncia capituliformc y estructuras floralcs dc
muy pequenas dimensiones, subglabras, csta es-
pecic sc ooloca cn la seccidn Leptinga. La nueva
(■specie cs similar a /. allenii de Costa Rica y Pana¬
ma, dc la que difiere principalmente por tener nec-
tarios foliarcs doliiformes vs. clavilormcs, fruto mas
pequefio, con apicc obtuso vs. rostrado y foliolos
con mayor mimero venas secundarias por lado. 14—
19 vs. 6-8, rcspccti\ amenle.

Austkact. Inga colonchensis is described from
fragmented tropical dry forest in the cordillera
Chongon-Colonche, Guayas Province, western Ke¬
uador. It is placed in section Leptinga on account
of the capituliform inflorescence and very small
subglabrous flowers. I his new species is similar to
/. allenii from Costa Rica and Panama but differs
in its barrel-shaped foliar nectaries, smaller fruits
with obtuse apex, and higher number of lateral
veins per side.

Key words: tropical dry forest (bosque seco
tropical). Keuador, endemic, Inga colonchensis, sec¬
tion Leptinga.

Inga Miller (Fabaceae— Mi mosoideae) es un ge-
nero que comprende aproximadamente 300 espe-
eies, ampliamente distribuidas en America, desde
24°N en Mejieo hasta 34°S en Uruguay, incluyendo
las Antillas (Pennington, 1997). Kn Keuador se ban
registrado 75 especies, distribuidas en bosques se-
eos y luimedos, desde las /onas bajas a ambos lados
de los Andes hasta el bosque montano andino (Pen¬
nington & Revelo, 1997; Cornejo, obs. pers.). Kste
genero ha sido dividido en 14 seeeiones. entre las
cuales leptinga. se caracteriza por presentar: (I)
pubescencia finamente puberula o subglabra, (2)
neetarios foliarcs casi siempre sesiles, (3) inflores-
eencias capitadas o umbeladas, (4) flores glabras o
puberulas de pequenas dimensiones, (5) estambres
de 1—2 cm de largo, en mimero de 20—50, (6) ovario

I -carpelado y (7) legumbre plana a convexa eon los
margenes angostos, no expandidos (Pennington,
1997).

Durante la lase de eampo para la elaboraeidn de
la Flora de! Bosque de Garda de la comuna hnna
\lt a (Bonifaz X Cornejo, 2004). en las margenes
de los nos localizados en las partes bajas de la
cordillera Chongon-Colonche, eneontramos una es-
peeie que presenta las caraeterfstieas de leptinga.
la misrna que es distinta de las 30 especies de esta
seccidn que se encuentran en la region occidental
de America del Sur v de las 1 1 especies registradas
en America Central. Ksta es descrita a conlinua-
cidn.

Inga colonchensis X. Cornejo X C. Bonifaz, sp.
nov. I IPO: Keuador. Guayas: Cordillera Chon-
gdn-Colonche, Comuna l.oma Alta, Rfo seco,
l°46'S. 80°40'0, 200 m, 29 May 1999 (II), X.
Cornejo & C. Bonifaz 6B58 (holotipo, GUAY;
isotipos, AAU, COL, GB. L0JA, MO, OCA.
QCNK). Figura I.

Race species Ingae allenii aflinis, a qua differt glan-
dulis extra rhacliiales. nectariis interfoliolarihus doliifor-
inihus, ven is lateral i bus majorihus, legumine ajiiee obtu¬
so.

Arbol hasta 6 m alto; ramas abundantemente
lenticeladas, las terminales cilfndricas hasta algo
anguladas, longitudinalmente canalieuladas; brotes
jdvenes (sub)eilfndrieos hasta subangulados, a vo¬
ces algo eomplanados. adpreso-puberulos; estfpulas
(—9 mm, oblongas hasta oblanceoladas, glabras o
adpreso-puberulas. caducas. Glandulas extra-ra-
(|uiales dispuestas antes de las bases de los peefo-
los y de los pedunculos de las inflorescencias. Pe-
efolo 1.1—4 cm (sub)eilfndrieo, glabro; raquis 2.5—
12.3 cm, alado hasta ampliamente alado (2 cm), en
la mitad o hasta dos tercios superiores del entrenu-
do. glabro; el apendiee ausente; neetarios foliares
doliiformes con pequena apertura circular. 0.8— 1.3
mm; pecidlulo 2—3 mm; foliolos (2-)3 pares; par
terminal 10—28 X 3.2—1 LI cm, elfptico a angosto
obovado, el apice acuminado hasta 1.5 cm, base
atenuada; par basal 4.7-12.5 X 1.8-6 cm, elfptico.

Novoim 13: 270-273. 2005.

Volume 15, Number 2
2005

Cornejo & Bonifaz

Inga colonchensls de Ecuador

271

2cm

rigura 1. Inga colonchensis X. Cornejo & C. Bonifaz. — A. Hama florecida. — B. Nectario foliar y parte superior
adyacente del raquis alado. — C. Infforesceneias terminales. — I). Hor. — E. Legumbre rnadura. — I'. Seecibn dr* la
legumbre. (A— I): l)ibujo del holotipo, Cornejo & Bonifaz 6858\ E. I*: Cornejo <ft Bonifaz 5801.)

272

Novon

el apice aeuminado, la base atenuada; raramente
un poco obtusa; liaz opaco, glabro; enves con muy
cortos pelos ad preso-pube rules o glabro; venaeidn
eucamptddroma; 14—19 pares de venas seeundarias
arcuadas. venaeidn tereiaria perpendicular o debil-
mente retieulada. Inflorescencias eapituliformes,
solitarias o geminadas. terminales, axilares o rami-
floras; pedunculo 2—3.3 cm, adpreso-puberulo: ra-
(piis floral 1.5-2. 5 X 1-1.3 nun. clavado; bracteas
1 — 1.7 mm. espatuladas, eadueas; (lores sesiles; ca-
liz abierto en prefloraeidn. el tubo 1.5—2 mm. los
Idbulos 0.2— 0.3 mm. adpreso-puberulo; el tubo de
la eorola 4—5.5 mm, los Idbulos 1—1.5 mm, adpre¬
so-puberulo; estambres ea. 20—35, tubo estaminal
3—5 X 0.4 mm, incluido o apenas exserto, filamen-
los lilues. 5—6 mm; ovario de 1 earpelo, glabro, el
estilo 9—10 mm. Legumbre madura (5— )l 1-16 X
2—2.8 X 0.8— 1.5 cm, recta, el apice y la base ob-
tusos, las superficies ligeramente convexas, apenas
hinchadas sobre las semillas, con debil venaeidn
transversa, margen 4-6 mm, blanquecino, con
abundantes lenticelas de color bianco, glabra.

Dentro de la seccidn Leptinga, Inga colonchensis
es affn a la centroamericana /. allenii J. Leon de
Costa Rica y Panama (Leon, 1966; Pennington,
1997), la oral presenta similar estructura de la in-
floresceneia, las dimensiones florales, la abertura
de sirs neetarios foliares. el numero y forma de los
folfolos; pero difiere de esta por tener neetarios fo¬
liares doliiformes vs. claviformes, fruto con apice
obtuso vs. rostrado, de menor longitud, (5 — ) 11 — 16
vs. 19—24 cm, folfolos con 14—19 vs. 6—8 venas
laterales por lado, con apices acuminados vs. cau-
dado-acuminados; en vivo, los margenes del fruto
y las lenticelas son blancas vs. cafd y se observan
distintivas glandulas extra-raquiales dispuestas an¬
tes de las bases de los pecfolos y de los pedrinculos
de las inflorescencias, siendo diffcil de apreciar
esta ultima caracterfstica en material seen. Tanibien
es parecida a Inga jinicuil G. Don, distribuida des-
de Mexico hasta cl bosque muy humedo del noroc-
cidente de Ecuador, de la cual difiere por tener
raquis alado vs. no alailo, apendice ausente vs. pre¬
sente, de 4— 7 mm de largo y legumbre mas angosta,
2—2.8 vs. 6-7 cm de ancho, con base simetriea vs.
fuertemente asimetrica. Entre las especies de Lep-
tinga de America del Sur, Inga colonchensis es pa-
recida a /. umbratica Poeppig & Endlieher, am-
pliamente distribuida en el bosque muy humedo de
la amazonfa, desde el sur de Venezuela hasta Bo¬
livia, de la eual difiere en cuanto a su fitogeograffa,
zona de vida y morfoldgicamente por presentar nee¬
tarios foliares doliiformes nuis angostos, de 0.8—1 .3
mm de diametro vs. pianos o pateliformes, 2—3 nun

de diametro; folfolos eon base atenuada vs. cordada
hasta truncada; raquis de la hoja glabro vs. den-
samente puberulo; raquis floral clavado vs. globoso;
flores con pedicelos ausentes vs. presentes. de
0.75—5 mm de largo y el tubo estaminal 3—5 mm
vs. 9-1 1 mm de longitud.

Ecologfa v distribucidn. Inga colonchensis se
encuentra en las margenes de los nos de la cordi¬
llera costera Chongdn-Colonche. entre 130-250 m
s.n.m. La base de la mencionada cordillera esta
constituida por rocas basicas perteneeientes a un
complejo volcano-sedimentario del perfodo ereta-
cico. Su zona de vida corresponde a la formacidn
vegetal de bosque seco tropical con vegctacidn ca-
ducifolia, adaptada a perfodos seeos de aproxima-
damente seis meses de duracion.

Fenologi'a. Esta especie florece en febrero,
mayo y junio, fructifica en julio hasta lines de oc-
tubre.

Interacciones bioldgicas. Las flores son visita-
das por hormigas y abejas durante el dfa.

Etimologia. El nombre propuesto para esta
nueva especie se debe a la cordillera costera Chon¬
gdn-Colonche v a su genie, la misma (]ue rescald
vivo al segundo autor en el accidente aereo de 1993
en el que fallecio A. H. Gentry.

Uso. De las poblaciones silvestres existentes,
los habitantes de Loma Alla ocasionalmente comen
sus frutos. Se dice (pie los monos tambien lo co¬
men, a lo (pie debe uno de sus nombres locales
(“Guaba de mono"). La suave pulpa blanca (jue
recubre las semillas tiene un sabor dulce y agrada-
1 )le al madurar, sin embargo esta especie no ha sido
domesticada.

Nombres locales. “Guaba". “Guaba de mono".

Paratipos. ECUADOR. Guayas: Cordillera Chongdn-
Colonche, La Bramona, bosque seco tropical, 130 m, 9
Jtil. 1995 (fr), X. Cornejo & C. Ronifaz 4205 (GB, GUAY,
K); Cordillera Chongdn-Colonche, Comuna Loma Alta, le-
eho dc rfo California, bosque seco tropical, formacidn dc
lccho dc rfo. 150 m. 27 Sep. 1997 (fr), ,Y. Cornejo C.
Ronifaz 5801 (GB, (ill AY, k); miembabital humedo, 7
Feb. 1 999 (fl). Y. Cornejo S: C. Ronifaz 0756 (GUAY).

Agradecimientos. El cabildo de la Comuna
Loma Alta nos otorgd el permiso de coleccidn res¬
pective; el eomunero Celso Tomala nos brindd el
hospedaje y los comuneros Mauricio, Eelix v Wil¬
son Tomala fueron nuestros gufas de campo. T. I).
Pennington (K) revisd el paratipo X. Cornejo <£• C.
Ronifaz 4205. Los herbarios visitados son: AAU,
k. MO, NY. OCNE. La fundacidn GAIA finaneid
parcialmente los gastos de la fase de campo.

Literatura Citada

Bonifaz, C. & X. Cornejo. 2004. Flora del Bosque dc Ga-

rua dc la Comuna Loma Alta, Provincia del Guayas,

Volume 15, Number 2
2005

Cornejo & Bonifaz

Inga colonchensis de Ecuador

273

Ecuador. Universidad de Guayaquil/Missouri Botanical
Garden/ Fundacion GAIA. Guayaquil, Ecuador.

I .eon. J. 1966. Central American and West Indian species
of Inga. Ann. Missouri Hot. Gard. 53: 265—359.

Pennington, T. 1). 1997. The Genus Inga — Botany. The
Royal Botanic Gardens, Kew.

- & N. Revelo. 1997. El Genero Inga en el Ecua¬
dor. The Royal Botanic Gardens. Kevv.

A New Species of Hypericum (Clusiaceae) from
Shaanxi Province, China

Du Xi-chun and Ren Yi*

College of Life Seienees, Shaanxi Normal University, Xi'an 710062,
People’s Republic of China

Author lor correspondence: renyi@snnu.edu.cn; nMivi@nwu.edu.cn

Austuact. The new species Hypericum qinlin-
gense (Clusiaceae) is described and illustrated from
the southern slope ol Mount Qinling in Shaanxi
Province, China. The new species belongs to Hy¬
pericum sect. Adenosepalum by its glabrous, peren¬
nial, herbaceous habit, persistent perianth and an-
droecium, sepals with large stalked black glands,
three bundles of stamens, and three free styles, as
well as by the alveolate seeds. The new species is
similar to Hypericum elodeoides Choisy in its scor-
pioidal diehasium, but differs in its bracts and flow¬
ers. A key to the Chinese species of Hypericum
sect. Adenosepalum is included.

Key words: China, Clusiaceae, Hypericum sect.
Adenosepalum, Shaanxi Province.

The genus Hypericum L. (Clusiaceae) consists of
more than 400 species and is cosmopolitan except
for the Polar Region. There are 8 sections, 55 spe¬
cies, and 8 subspecies of Hypericum in China,
mainly in the southwest region (l.i et ah. 1990). The
new species belongs to Hypericum sect. Adenose¬
palum Spaeh, which consists of 34 species (Li et
ah, 1990); 7 occur in China: H.filicaule (Dyer) N.
Robson, II. monanthernum Hooker f. & Thomson
ex Dyer, II. wight ianum Wallich ex Wight & Arnott,
II. himidaicum l\. Robson, H. elodeoides Choisy, II.
hengshanense W. T. Wang, and the new species.

Hypericum qiulingense X. C. Du & Y. Ren, sp.
nov. TYPE: China. Shaanxi: Yangxian. Huay-
ang, Xinglongling, 33°41'73"N, 107°35'94"E,
2450 m, 3 July 2001, ETNWU (Expedition
Team of Northwest University) 40053 (holo-
type, SANU). Eigure 1.

Have species Hyperico elodeoidi similis, sed ah eo bruc-
Icis base glandulis nigris stipitatis praeditis, sepalis pe-
lalisque minoribus, stain i n i bus in quoque fasciculo pau-
cioribus, filamentis brevioribus atque stylo longitudine ca.
tertiam partem ovarii aequante differ!.

Herbs perennial, 30—40 cm high; stems 1 to sev¬
eral from basal rosette, cylindrical, glabrate, with
inconspicuous glandular spots and vertical long

Novon 15: 274-276. 2005.

glandular stripes, internodes 1 — 4.5 t-ni long. Leaves
stiffly papyraceous, elliptic to obovate, 1.3-2. 9 X
0.5—1 cm on main stems, 6 — 10 X 2-3 mm on
branches, with small light glands on the surface
and larger black glands near the edge, slightly re-
tuse or obtuse at apex, the margin entire and rev¬
olute, cuneate at the base and tapering into a pet¬
iole about I mm long, green on upper surface and
light green below. Inflorescence a terminal scor-
pioidal diehasium, bracts lanceolate, 3-5 mm long,
ea. 1 mm wide, with stalked black glands at the
base of the back and the edge. Flowers yellow, ca.
8 mm diam.; perianth persistent; sepals 5, lanceo¬
late, 3— 4 mm long, ca. 1 mm wide, with yellow-
brown vertical glandular stripes and occasional
black stalked glands at the edge; petals 5. linear.
4—6 mm long, ca. 1 mm wide, with black glands at
the apex and yellow-brown vertical glandular
stripes; androeeium persistent, with three stamen
groups ol 10 to 12 each, filaments ca. 4 mm long,
anthers yellow, ca. 0.1 mm long, with longitudinal
slit, with black glands at the connective; carpels 3,
united, ovary ovoid, ea. 4 X 3 mm. with axial pla-
centation, styles 3, 1 — 1.8 mm long, ca. xh length of
the ovary, with occasional black glands on stigma.
Capsules ovoid, with septicidal dehiscence, ca. 5
mm diam., with yellow-brown glands, with ca. 180
seeds inside. Seeds ca. 0.6 mm long, alveolate.

Habitat and distribution. The new' species oc¬
curs on exposed slopes and in secondary forest, as
well as roadsides, ranging from 1320 to 2450 m
elevation. The primary vegetation of the area is de¬
ciduous broadleaved forest consisting of species of
Quercus L. or temperate coniferous and broadleav¬
ed mixed forest composed of species of Betula L.
and Abies Miller.

The new species is known only from Yangxian
County of Shaanxi Province, China. Yangxian
County is located in the middle of the southern
slope of Ml. Qinling.

The new species belongs to Hypericum by its
herbaceous habit, yellow flowers without glands be-

Volume 15, Number 2
2005

Du & Ren

Hypericum qinlingense (Clusiaceae)

275

Figure 1. Hypericum qinlingense X. C. Du <N, 1. Ken. — Plant. — II. Dorsal view of leaf. C, D. Bracts. (.. Dorsal
view. — I). Ventral view. — K. Flower. — F. Petal. — G. Stamens. — FI. Capsule with persistent sepals and styles and
with petals and stamens removed. — I. Seed. Scale bars: \ = 2 cm; I? = 5 mm; C. I) — 1 mm; K = •> mm: F — 1
mm; G = I mm; 11=2 mm; I = 0.2 mm. Drawn by Guo Mu-seng from the holotype, ETNWU 40053.

276

Novon

tween l lie stamen groups, stamens basally united
into three groups, septieidal c apsules, and seeds
without wings.

There are eight sections of Hypericum in China
(hi el al., I960). Hypericum qinlingense is readily
distinguished from the related Chinese species in
section Ade nosepal um by its stalked black glands
at the* base of the back of the bracts, sepals ca. 3—
4 X 1 mm, petals 4—6 X ca. 1 mm, stamens basally
adnate in 3 groups of 10 to 12 each, filaments ca.
4 mm long, and d carpels with d styles 1-1.8 mm
long, ca. Vs of ovary length.

Hypericum (/inlingense is similar to //. elodeoides
in its scorpioidal dichasium, but differs from the
latter in its bracts with black stalked glands at the
base, smaller sepals and petals, fewer stamens per
bundle, shorter filaments, and much shorter styles
that are only about Vs ol the length ol the ovaries,
file new species may be contrasted with six other
species of Hypericum known to occur in China in
the following key.

k.K'l TO HYPKHICI \l SKCT. AUKXOSKPAIJ 1/ FROM CHINA

la. Inflorescence a scorpioidal dichasium.

2a. Bracts w ith stalked black glands at the base
of the back and the edge; sepals ca. 3-4 X
I mm; petals 4—6 X ca. I mm; stamens ba¬
sally adnate in 3 groups of 10 to 12 each,
filaments ca. 4 mm long; 3 carpels with 3
styles, styles 1 — 1.8 mm long ca. Vs of ovary

length . II. qinlingense

2h. Bracts lacking glands; sepals ca. 6 X 3 mm;
petals ca. 15x4 mm; stamens basally ad¬
nate in 3 groups of 20 each, filaments ca.
0.8-14 cm long; 3 carpels with 3 styles,

styles at least twice ovary length .

. II. elodeoides

lb. Inflort ■scence a single flower or aggregate inflo¬
rescence, not scorpioidal diehasia.

3a. Sepals 4 . H. filicaule

3b. Sepals 5.

4a. Style 1.4—7 times longer than ovary, ex-

serted from flower . II. hengshnnense

41). Style shorter than or not exceeding the
ovary, inserted within flower.

5a. Flowers to 2 cm diam .

. II. monanthemum

5h. Flowers less than 1.5 cm diam.

6a. Annual herbs, with black
stalked glands on leaf margins

. II. wighlianum

6h. Perennial herbs, lacking mar¬
ginal glands .... II. himalaicum

Parntypes. CHINA. Shaanxi: Yangxian, Huayang,
Cang'eryan, 26 June 2001, ETNWU 40055 (SANU), 25
Aug. 2000, ETNWU WA2-6-25 (SANU); Dongping, 26
Aug. 2000. ETNWU EE-025 (SANU); Maoping, Pochayu,
2 i Aug. 2000. ETNWl EA-004 (SANU); same locality as
type, ETNWU 1 0544 (MO).

Acknowledgments. The reviewer, John J. Pipoly
III (f TG), and scientific editor, Victoria C. Hollow-
ell (MO), were extremely helpful in clarifying the
text; we also thank Roy Gereau (MO) for the Latin
diagnosis. This report was supported by State Key
Basic Research and Development Plan of China
(No. G20000468()0() I ).

I iteralure Cited

Fi Hsiwen, Li Yanhui, Long Shaoquan. Tao Gouda, Zhang
Pengyun & Zhang Yaojia. I W0. Hypericum. Pp. 2-72
in Flora Reipublieae Popularis Sinicae 50(2). Science
Press, Beijing. 1 1 n Chinese.]

Nomenclatural Notes on rhi *ee Species of Cactaceae from

South America

Urs Eggli

Sukkulenten-Sammlung Zurich, Mythenquai 88, CH-8002 Zurich, Switzerland.

urs. eggli@gsz.stzh.ch

Abstract. The names Cumulopuntia crassicylin-
drica (from Peru), Espostoa guentheri (from Bolivia),
and Yungasocereus inquisivensis (from Bolivia) are
validated here. All had been previously published,
but were invalid under Article 33.3 of the Inter¬
national Code of Botanical Nomenclature.

key words: Cactaceae, Cumulopuntia. Espos¬
toa, nomenclature. South America, Yungasocereus.

In the course of preparing a checklist of accept¬
ed names of Cactaceae and their synonyms for use
in the forthcoming German translation of Anderson
(3001), a number of names have been found to be
in “current use" despite the fact that they are not
validly published under Article 33.3 of the current
International Code of Botanical Nomenclature
(Greuter et al.. 2000).

Failures to meet the requirements for valid pub¬
lication under Article 33.3 of the Code in the con¬
text of publishing a new combination are sometimes
difficult to establish, and this has been the case for
the two names in question, whose combinations
were published by F. Ritter: yungasocereus inqui¬
sivensis (Ritter, 1980) and Cumulopuntia crassicy-
lirulrica (Ritter, 1981). Both these names are in¬
valid because of missing page numbers in the
basionym reference, but were listed as valid by Eg-
gli and Taylor (1991), who used information from
Rowley et al. (1982. 1983). In the case of the at¬
tempted combination Espostoa guentheri by Bux-
baum (1959), its invalid status (because of tbe
missing reference to the basionym) was somewhat
cryptically stated (as “comb, subnud.”) bv Rowley
i 1961: 16), but was fullv indicated by Kggli and
Taylor (1991).

In order to validate these names to be available
for future use. tin* necessary combinations are pub¬
lished as follows:

Ciiinulopiintia crassiev lindrica (Ranh N Baek-
eberg) F. Ritter ex Fggli. comb. nov. Basionym:
Tephrocactus crassicylindricus Ranh &
Backeberg, in Baekeberg, Descr. Cact. Nov.:
8. 1956 | 1957]. TYPE: Peru. Rept. Arequipa:
1956, W. Ranh k 152 (holotvpe, IIEII) not
seen).

Cumulopuntia crassicylindrica is a poorly known
taxon, presently only reported from Dept. Arequipa,
Peru, and seems to be known from the type collec¬
tion only (Iliff. 2002).

Espostoa guentheri (Kupper) Buxbaum ex Eggli,
comb. nov. Basionym: Cephalocereus guentheri
Kupper, Monatsschr. Deutsche Kakt.-Ges. 3:
161—162. 1931. TYPE: Bolivia. Dept. Clm-
quisaca: 1927. C. Troll s.n. (holotype, M not
seen).

Espostoa guentheri is reported from the Bolivian
departments of Chuquisaea, Cochabamba, and San¬
ta Cruz (Navarro, 1996). It was described on the
basis of photographs and a living stem tip sent to
Munich by the collector. It is assumed that the liv¬
ing material was preserved at Munich after having
died in cultivation, but this has not yet been veri¬
fied. In the absence of the specimen, the illustration
in the protologue (p. 160) might serve as lectotype.

Yungasocereus inquisivensis (Cardenas) F. Ritter
ex Eggli, comb. nov. Basionym: Samaipaticer-
eus inquisivensis Cardenas, Cactus (Paris)
12(57): 246-247. 1957. TYPE: Bolivia. Dept.
La Paz: 1956, M. Cardenas 5 III I (holotype,
BOI.V not seen; isotype, US not seen).

Yungasocereus inquisivensis is only known from
the Bolivian Department La Paz (Navarro, 1996).
The holotype specimen supposedly is at BOI.V, as
a recent survey (by the author and B. E. Eeuen-
berger) of the Cardenas Cactaceae specimens at
LIE did not locate any material of this collection
number. A “cotype” (i.e.. isotype) is cited for US
in the protologue, but the on-line version of the type
specimen list at US does not include an entry for
diis number.

Literature Cited

Anderson, E. F. 2001. The Cactus Family. Timber Press,
Portland.

Buxbaum, F. 1959. Die behaartbliitigen Cephalientrager
Sudamerikas. Osterr. Bot. /. 106: 138—158.

Kggli. U. & N. P. Taylor 1991. IDS Index of Names of

Novon 15: 277-278. 2005.

278

No von

Cactaceae published 1950—1990 from Repertorium
Plantarum Succulentarum. Royal Botanic Gardens,
Kew/Stiidtische Sukkulenten-Sammlung, Zurich.

Greuter, W.. J. McNeill, F. II. Barrie, II. M. Burdel, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
J. K. Skog, P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.

Ilifl, .1. 2002. The Andean Opuntias: An annotated check¬
list of the indigenous nou-platyopuntioid opuntias (Cac-
taceae— Opunlioideae) ol South America. Slice. PI. lies.
6: 133-244.

Navarro, G. 1996. Catdlogo ecoldgico preliminar dc las
cacldceas dc Bolivia. I.azaroa 17: 33—84.

Bitter, F. 1980. Kakteen in Sildamerika. Band 2. Argen-
tinien/Bolivien. Selhstverlag, Spangenhcrg.

- . 1981. Kakteen in Sildamerika. Band 4. Peru.

Selhstverlag, Spangenhcrg.

Rowley, G. I). (editor). 1961. Repertorium Plantarum Suc-
culentarum. X. International Association of Plant Tax¬
onomists, Utrecht. Regnum Veg. 21.

- , I.. K. Newton X N. P. Taylor (editors). 1982. Re-

pertorium Plantarum Succulentarum. XXXI. Interna¬
tional Organization for Succulent Plant Study, Addle-
stone.

- , - & - (editors). 1983. Repertorium

Plantarum Succulentarum. XXXII. International Orga¬
nization for Succulent Plant Study, Addlestone.

Caloch art us mendozae (Calochortaceae), una Nueva Especie de

San Luis Potosi, Mexico

Adoljo Espejo Serna , Ana Rosa Lopez- Ferrari y Jacqueline Ceja Romero
Departamento de Biologfa, Division de C. 15. S., Universidad Autonoma Metropolitana
Unidad Iztapalapa, Apartado Postal 55-535, 09340 Mexico, D.F. aes@xanum.uam. nix;

jcr@xanum.iiam.mx

RESUMEN. Se describe e ilustra a Calochortus
mendozae Espejo, Lopez-Ferrari X Ceja, especie
eonocida hasta ahora del munieipio de San Nicolas
Tolentino, en el estado de San Luis Potosi, Mexico,
en donde crece sobre taludes yesosos, en vegeta-
eidn de matorrral xerdlito. I.a nueva especie se ubi-
ea dentro de la subseccion Ghiesbreghtiani de la
seeeidn Cyclobothra del genero Calochortus y la es¬
pecie mas cereana al nuevo taxon es C. hintonii
Bullock ex Ovvnbey, eon la eual se compara.

ABSTRACT. Calochortus mendozae Espejo, Lopez-
I' erraii & Ceja, a new species from the municipality
ol San Nicolas Tolentino in the state of San l.uis
Potosi, Mexico, is described and illustrated. The
new species grows on gypseous soils in xeroplivtie
scrub. Calochortus mendozae is placed in section
Cyclobothra subsection Ghiesbreghtiani and is
closely related to C. hintonii Bullock ex Ownbey,
with which it is compared.

Key words: Calochortaceae, Calochortus, San
Luis Potosi, Mexico.

Al identificar material de monoeotiledoneas
proveniente de San Luis Potosi, nos encontramos
con un ejemplar del genero Calochortus Pursh que
no pudimos asignar a ninguna de las espeeies re-
portadas para Mexico. La revision euidadosa de la
literature pertinente y del material, nos permit id
estableeer que se trata de una especie no descrita
por lo que proponemos:

Calochortus mendozae Espejo. Lopez-Ferrari X
Ceja, sp. nov. TIPO: Mexico. San l.uis Potosi:
munieipio de San Nicolas Tolentino, 3 km des¬
pues del Polrero de Santa Gertrudis, rumbo a
Buenavista, 22°1 1'30"N, l()0°20'58"W, 1320
m s.n.m., matorral xerdfilo, 21 Nov. 2002. A.
Mendoza Ruiz 6 56 (holotipo, UAMIZ; isotipo.
MO). Figure 1.

Herba ad 54 cm alta. Caules llexuosi, ramosi, glauei.
Kolia hasalia 1-2. linearia; folia caulina plantae apieem
versus gradatim decreseentia. Inflorescentiae axillares, di-

chotome ramificantes, arcuatae. t lores camparnilati, er-
ecti, fla\i, tepalis exterioribus ellipticis, apiee obtusis,
sine glandula, tepalis intends obovatis, hasi cuneatis, ap-
ice integris vel parum undulatis rotundatisque, hasi tri-
ehomatibus papillosis, glandula hippocrepiformi in tepali
dimidio ba.sali per membranam laciniatam obtecta; an-
tlieris oblongis, apice apieulatis; ovario longe elipsoideo,
ramis stigmaticis flavis.

Hierbas de hasta 54 cm de alto incluyendo el
bulbo, los bulbos globosos a ovoides, de 1.9 cm de
largo, de 1.6 cm de diametro, eubiertos por vainas
fibroso-reticuladas, pardas. Tallos erectos, flexuo-
sos, ramifieados, glabros, glaucos. Hojas basalt's I —
2, linear-lanceoladas, de 20—25 cm de largo, de ea.
1.2 cm de ancho, glabras; hojas eaulinares si mi la¬
res a las basales en forma, de hasta 18 cm de largo,
de 5—10 mm de ancho, reduciendose gradualmente
hacia la poreidn apical de la planta. Infloresceneias
axilares, dicotomicamente ramificadas, arqueadas,
las bracteas 3, una de ellas generalmente mas pe-
quena, linear-lanceoladas, largamente aeuminadas,
de 1.2— 3.3 cm de largo por 0.1— 0.5 cm de ancho
en la base; los pedunculos lineares, arqueados, es-
triados, glabros, de 4.5—13 cm de largo: los pedi-
celos lineares, arqueados, estriados, glabros, eon
tonalidades rosadas a purpuras, de 5—7 cm de largo
en (lor, de 8.5—16 cm en fruto; (lores campanuladas,
aparentemente nutantes por los pedunculos y pedi-
celos algo arqueados pero en realidad erectas; te-
palos externos amarillos, elipticos, de 12—16 mm
de largo, de 4—5 mm de ancho, enteros, obtusos en
el apice, glabros excepto por linos escasos trieomas
presentes en la porcion media basal, sin glandulas;
tepalos internos amarillos, obovados, euneados en
la base, enteros a ligeramente ondulados y redon-
deados a laciniados en la porcion apical, de 12—14
mm de largo, de 7—10 mm de ancho, glabros ex¬
cepto en la porcion basal en donde presentan tri¬
eomas a los lados de la glandula; la glandula en
forma de herradura situada en la mitad basal del
tepalo, cubierta por una membrana laciniada y eon
trieomas papilosos; filamentos subulados, amarillos.
de 4 mm de largo, de ca. 0.6 mm de ancho en la
base, las anteras oblongas, amarillas, de 4—6 mm

Novon 15: 279-281. 2005.

280

Novon

I cm

Figura 1. Calochortm mendozae Espejo, L6pez-Ferrari & Ceja. — A. Habito. —IF Flor. — C. Flor disecada. — 1).
Detalle de la glarulula. — E. Gineeeo. — F. Estambre. Basado en el holotipo.

de largo, apieuladas en el dpiee, el apfculo bianco;
ovario largamente elipsoide, triangular, de ca. 1 cm
de largo, de ca. 2.5 mm de diametro, glauco, las
ramas estigmatieas 5. amarillas, de 2 mm de largo,
recurvadas, agudas. Fruto una capsula largamente
elipsoide. triangulada, de 3 cm de largo, lie 0.5 cm
de diametro; semillas alargadas, triangulares en

corte transversal, de 5 mm de largo, de I nun de
ancho, con la testa finamente reticulada.

La especie se eonoce unieamente de la localidad
tipo, en donde erece sobre taludes yesosos, en ve-
getacidn de material xeiofito. Calochortus mendozae
se ubica dentro del subgenero Cyclobothra (Painter,
1911) o la seecidn Cyclobothra (Ownbey, 1940), ya

Volume 15, Number 2
2005

Espejo et al.

Nueva Especie de Mexico

281

'labia 1. Diferencias entre Calochortus mendozae y C. hintonii.

Calochortus mendozae

Calochortus hintonii

llores

erectas, campanuladas, tie 24 a 32 mm tie
diametro, tie color amarillo

erectas, ampliamente campanuladas, tie 18 a
27 mm tie diamentro, tie color rojo oscuro,
ocasionalmenle amarillas

Tepalos externos

elfpticos, 12-16 X 4—5 mm, obtusos en cl
apice, glabros excepto por unos escasos tri¬
comas presentes en la porcion media basal

elfptico-lanceolados, 9—1 1 X 3.5—5.55 mm,
agudos a obtusos en el apice, barbados cer-
ca de la base

Tepalos internos

obovados, 12—14 X 7-10 mm, redondeatlos a
laciniados en el apice. glabros excepto en
la porcion basal a los lados de la glandula;
tricomas multiseriados

elfpticos a angostamente obovados. 12—13.5

X 5-7.5 mm, obtusos a agutlos en el apice,
densamente barbados cerca tie la base; tri-
comas uniseriados

Glandula

en forma de herradura, cubierta por una
membrana laciniada y tricomas papilosos

en forma de media luna, densamente cubierta
por tricomas no papilosos

Kilamentos

subulados, amarillos, tie 4 mm tie largo

lineares, rojos, de 5 mm tie largo

Anteras

oblongas, de 6 nun de largo, apieuladas en (“1
apice

elfpticas, tie 1.7 a 3 mm tie largo, obtusas a
agudas en el apice

( Ivario

ca. 10 inm de largo, vertle, glauco; ramas es-
tigmaticas amarillas

6.5—7 mm tie largo, vertle, glauco; ramas es¬
timations rojas a purpuras

Habitat

matorral xerdlito. solire suelos yesosos

bosque de encino o tie pino-encino, sobre
suelos rieos en materia organiea

1 listribucion

conocida solo tie San Luis Potosf

endemica tlel estatlo tie Mexico

< I tit- presenta bul bos cubiertos [tor calafilas fibroso-
reticulailas y capsulas alargadas y trianguladas y
pertenece a la subseecion Ghiesbreghliani, que
agrupa especies con los tallos flexuosos. raramente
bulbfferos en las axilas de las hojas caulinares y
llores pequenas y erectas con los tepalos inconspi-
cuamente barbados (Ownbey, 1940). hi nuevo ta¬
xon presenta (lores aparentementc similares a las
de C. hintonii Bullock ex Ownbey, especie endemi-
ca del Estado de Mexico que tambien se ubica en
la subseecion Ghiesbreghtiani (Apendice 1). de la
que adenitis se conoce una variedad de flores ama-
ri I las. Sin embargo, existen diferencias claras entre
ambus especies, como puede apreciarse en la
labia 1.

El nombre de la especie esta dedicado al M. en
C. Aniceto Mendoza Ruiz, colega y amigo de la
Universidad Autdnoma Metropolitana l/.tapalapa.
companero de numerosos viajes de coleeta, quien
liizo el liallazgo del nuevo taxon.

Agradecimientos. Los autores agradecen a los
Ores. Fernando Chiang Cabrera y Abisai Garcia
Mendoza, la revision crftica del manuscrito y al pri-
mero de ellos la elaboracidn de la diagnosis latina
de la nueva especie. La figura que acompana al

trabajo fue realizada por Rolando Jimenez Macho-
rro.

l.iteratura Citada

Ownbey, M. 1940. A monograph of the genus Calochortus.

Ann. Missouri Hot. Gartl. 27: 371—561.

Painter, J. H. 1911. A revision of the subgenus Cyclo-
bothra of the genus Calochortus. Conti. U.S. Natl. Herb.
13: 343-350.

ApTnimck 1. Material revisado de Calochortus hinto¬
nii bullock ex Ownbey. MEXICO. Estado tie Mexico:
Municipio de Tejupilco de Hidalgo, 5 km antes de la des-
viaoion a Almoloya tie las Granadas, por el camino Te-
inascaltepec— Tejupilco, A. Espejo. A. R. hipez- Ferrari &
t. Flores C. 3396 (UAMIZ); Municipio tie Tejupilco de
Hidalgo, alrededores de Rincon tie Ugarte, Tejupilco, A.
Fspejo. A. R. Ldpez- Ferrari A' /. Reyes ./. 5537 (UAMIZ);
Municipio de Tejupilco de Hidalgo, I km sobre el camino
a Almoloya tie las Granadas, a partir tit1 la carretera Te-
masealtepec— Tejupilco, A. Flores C„ A. I\. hi jtez- Ferrari

6 A. Fspejo 985 (l AMIZ), 986 (UAMIZ); Municipio tie
Pejupilco tie Hidalgo, Cerro Muneca, G. Ilinton 1383
(MO. GH, NY, US): Municipio de Tejupilco de Hidalgo,

7 km despues de Pejupilco, rumbo a Temascal tepee, A.
R. Ldpez- Ferrari. A. Fspejo & I. Reyes ./• 2335 (UAMIZ);
Municipio tie Tejupilco de Hidalgo, “Rancho Quieto”,
Rincon de Ugarte, Tejupilco, /. Reyes ./. 48 (UAMIZ);
Municipio de Tejupilco de Hidalgo, 7 km adelante tie Te-
mascaltepec, rumbo a Tejupilco, 500 in antes de la des-
viacion a San Simon. ./. Santana C. 113 (UAMIZ).

Ditassa obscura (Apocynaceae: Asclepiadoideae, Asclepiadeae),
a New Combination from Minas Gerais State, Brazil

Maria Ana Farinaccio

Departamento de Botanica, Instituto de Bioci6ncias, Universidade de Sao Paulo,

Caixa Postal 1 1461, CEP 05422-970, Sao Paulo, SP, Brazil, mafarinaccio@hotmail.com

Tatiana U. F. Konno

Instituto do Botanica de Sao Paulo, Divisao de Fitotaxonomia, Segao Curadoria <lo Herbario,
Av. Miguel Estefano 3687, Caixa Postal 4005, CEP 04301-012, Sao Paulo, Brazil.

tkonno@uol.com.br

Abstract. Ditassa obscura (E. Fournier) Farinac¬
cio & T. 1). P. Konno (Apocynaceae: Asclepiadoi¬
deae, Asclepiadeae), a new combination from Mi¬
nas Gerais State, Brazil, is proposed, and its
relationships are discussed.

Key words: Apocynaceae, Asclepiadeae, Ascle¬
piadoideae. Brazil. Ditassa , Minas Gerais, Neotrop¬
ics.

Ditassa R. Brown is a Neotropical genus of the
large tribe Asclepiadeae (Asclepiadoideae, Apocy¬
naceae), containing approximately 100 species
(Konno & Fontella-Pereira, 2004). It is one of the
most species-rich genera in Brazil, with a center of
diversity in the states of Bahia and Minas Gerais
(Fontella-Pereira et al., 1987, 1989; Rapini et ah,
2001; Farinaccio & Mel lo-Silva, 2004a).

The germs has a problematic delimitation (Fari¬
naccio & Mello-Silva, 2004a, 2004b), but can usu¬
ally be recognized by its subaxillary umbelliform
cymes and flowers with a double corona, in which
the outer segments are united at least at the base.

During the preparation of a monograph of the
subfamily Asclepiadoideae for the Serra da Can-
astra National Park, in the state of Minas Gerais,
southeastern Brazil (Farinaccio & Mello-Silva,
2004a), a specimen came to light that exhibited the
same set of characters as the type collection of Me-
tastelma obscurum E. Fournier. Its clear affinity
with Ditassa tomentosa (Decaisne) Fontella, espe¬
cially in relation to floral morphology and also veg¬
etative features such as indument and leaf venation,
justify its transfer to Ditassa (Fontella-Pereira,
1979). Therefore, we propose the new combination
Ditassa obscura (E. Fournier) Farinaccio & T. U. P.
Konno.

Ditassa obscura (F. Fournier) Farinaccio & T. U.
P. Konno, comb. nov. Basionym: Metastelma
obscurum F. Fournier, in Marlins, FI. Bras.
6(4): 208. 1885. TYPE: Brazil. “In prov. Min-
arum in Serra de Araxa," s.d., St. Hilaire Cl-
449 (holotype, P). Figure I .

Vines, densely tomentose; leaves opposite, pat¬
ent; petiole 2-7 mm long, somewhat sulcate, dense¬
ly tomentose; leaf blade 2-4.7 X 0.6— 1.5 cm, el¬
liptic to obovate, apex acute, mueronate, base
cuneate, discolorous, opaque, chartaceous, adaxial
surface tomentose, abaxially pubescent, margins
strongly revolute, venation brochidodromous, mid¬
rib abaxially prominent, secondary veins conspic¬
uously parallel, approximately perpendicular to the
midrib, 2 to 4 colleters at the base on the adaxial
side; inflorescence subaxillary, glomerulate, alter¬
nate, 5- to 10-flowered, erect; peduncles 0.3-1 nun
long, densely tomentose, bracts 1.1 — 1.5 X 0.2— 0.6
mm, lanceolate, abaxial surface tomentose, adaxial
surface glabrous. Pedicel 0.3-0. 5 nun long, tomen¬
tose. Calyx light green to green, lobes 1.7—2 X
0.55—0.6 mm, lanceolate to ovale, apex acuminate,
abaxial surface tomentose, adaxial surface gla¬
brous, with 1 colleter present below each sinus;
corolla white to cream, subcampanulate, tube 0.7-
0.75 mm long, abaxial surface glabrous, adaxial
surface barbellate, glabrous only in the proximal
region; lobes 1.5-2 X 0.7—0.95 mm, ovate, erect,
recurved from about the middle, apex acute, ab¬
axial surface glabrous, adaxial surface incanous-se-
riceous, barbellate in the proximal region, margins
hyaline; corona white to cream, in 2 series: outer
corona with 5 lobes joined at the base or free, not
attached to the corolla tube, each lobe 0.64—0.8 X
0.35—0.4 mm, oblong to ovate, asymmetric, apex
cuspidate, apiculate to unguiculate, shorter than
the gynostegium; inner segments strongly reduced

Novon 15: 282-285. 2005.

Volume 15, Number 2
2005

Farinaccio & Konno

Ditassa obscura from Minas Gerais

283

Figure 1. A— K. Ditassa obscura (E. Fournier) Farinaccio & T. U. P. Konno. — A. Flowering branch. — B. Flower with
one lobe removed showing corona and gynostegium. — C. Abaxial surface of anther. — I). Pollinarium. — E. Fruit. (A
from Romero , Nakajima, Farinaccio & Roque 4770, HUFU, SPF; B— E from Farinaccio & Campos Filho 273, BHCB,
F. (i. HKCB. K. MBM. MO, NY, BB. SPF.) Drawn by Bogerio l.upo.

284

No von

to al>sent; gynostegium cream, 045—1.1 mm long,
().(>— 0.64 mm diam., sessile, cylindrical, apex cap¬
itate; anthers 0.4—0.52 X 0.32—0.36 mm, rectan¬
gular, gibbous, terminal appendage 0.35—0.4 X
0.16—0.24 mm, ovate, strongly acuminate, wings
straight, longer than the dorsum; corpusculum
0.07—0.12 X 0.05—0.06 mm, ovate to conical, apex
rounded, translator arms 0.03—0.05 mm long, bat¬
tened. broad, translucent, pollinia 0.14—0.16 X
0.05—0.07 mm, oblong to elliptic; fruit a single fol¬
licle. immature green to dark purple, 3.8—4 X 0.3—
0.4 cm, oblong-lanceolate, hirsute; seeds 5—5.5 X
2—2.5 mm, ovate, eomose at the micropylar end.

Habitat and distribution. Ditassa obscura oc¬
curs at the borders of small forest patches (“ca-
pdes"), in gallery forests, and in grasslands with
stony sandy soil.

Phenology. Flowering from October to Novem¬
ber and from April to May; fruiting from March to
May.

Ditassa obscura forms a complex with D. tomen-
tosa. I). cipoensis (Fontella) Rapini, I). itambensis
Rapini, D. longicaulLs (F. Fournier) Rapini. and I).
longisepala (llua) Fontella & F. A. Schwarz (Table
I). It has bullate leaves like D. longisepala ; nev¬
ertheless, its Moral morphology and the shape and
color of the leaves are distinctive. Also. I). obscura
is very similiar to I). tomentosa, but the corolla
lobes of I), tomentosa are twice as long as those of
I). obscura (Table I). Moreover, I). obscura has bul¬
late leaves and much shorter peduncles and pedi¬
cels (Table 1) so that the inflorescences are almost
sessile, densely clustered, and glomerulate.

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Specimens examined. BRAZIL. Minas Gerais: Sao
Roque de Minas, Par. Nac. da Sena da Canastra. Rio do
Peixe, proximo ii area de desenvolvimento, Farinaccio it'
Campos Filho 273 (RHCR. F. (',. HRCB, k. MBM. MO,
NY. RB, SPF). Farinaccio , Campos Filho A' Vecchi 231
(CKSJ. C I KS, KSA, HUFFS, SPF); 20°l 5'27.B"S.
46°24'55.9"W, Farinaccio , Pontes. Magenta A Magenta
514 (B. BHCB, F!SA. G. SP. SPF); proximo £* trilha para
a Casca d'Anta, parte de eirna, Farinaccio A Pontes 516
(BR, FUEL, IN PA, M. P. S. SP. SPF. U, UEC); base do
mono proximo id sede admin. Nakajima. Romero. Zanini
A Simdo 1433 (HUFl . SPF); margens do Rio Sao Fran¬
cisco, Romero. Nakajima A Guilherme 167 (IIUFU, SPF);
Corrego dos Passageiros, Romero. Nakajima A Farinaccio
4179 (IIUFU. SPF); Chapadao da Zagaia de frente para a
Serra das Sete Ybltas, apos torre de observa^ao, Romero,
Nakajima. Farinaccio A Roque 4770 (IIUFU. SPF).

Acknowledgments. This work is part of a mas¬
ter's thesis by the first author, prepared under the
supervision of Renato Mello-Silva at the Universi-
didade de Sao Paulo. We are grateful to Rosana
Romero and Jimi Nakajima. coordinators of the
Flora of Serra da Canastra Project. Roth authors

cd

J,

cd

o s5 O

Volume 15, Number 2
2005

Farinaccio & Konno

Ditassa obscura from Minas Gerais

285

have been supported by FAPESP and CNI\] (MAh ).
Special thanks to the Margaret Mee Foundation,
whic h sponsored the visit of TUPK to the Museum
National d'Histoire Naturelle. in Paris. The line
drawings were prepared by Rogerio Lupo.

Literature Cited

Farinaccio, M. A. & It. Mello-Silva. 2004a. Aselepiadoi-
deae (Apocynaceae) do Parque Nacional da Sena da
Canastra, Minas Gerais, Brasil. Bol. Bot. Univ. Sao Pau¬
lo 22(1): 53-92.

- & - . 2004b. Ditassa insignis (Apocynaceae,

Asclepiadoideae), a new species from Serra da Canas¬
tra, Minas Gerais, Brazil. Few Bull. 59: 145-148.
Fontella-Pereira, ,1. 1979. Contribuigao ao estudo das As-
clepiadaceae Brasileiras, XIII. Ditassa tamentosa (De-

caisne) Fontella, uma nova combinagao. Bol. Mus. Bot.
Mimic. 39: 1-4.

- , M. da C. Valente & N. M. F. da Silva. 1987.

Asclepiadaceae. In: A. M. Giulietli et al.. Flora da Serra
do Cipo: Caracterizagao e li.sta das especies. Bol. Bot.
Univ. Sao Paulo 9: 22-24.

- . - , B. M. Harley & N. F. S. Marquete.

1989. Contribuigao ao estudo das Asclepiadaceae Bras¬
ileiras — XXIV. Checklist premilinar do Estado da Ba¬
hia. Kodriguesia 41(67): 81-1 15.

Konno, T. U. P. & J. Fontella-Pereira. 2004. Some no-
menclatural and taxonomic notes on Brazilian Ditassa
(Apocynaceae: Asclepiadoideae). Kew Bull. 59: 297—
300.

Rapini, A., R. Mello-Silva & M. L. Kawasaki. 2001. As¬
clepiadoideae (Apocynaceae) da Cadeia do Espinhago
de Minas Gerais, Brasil. Bol. Bot. Univ. Sao Paulo 19:
55-169.

Crotalaria mwangulangoi (Fabaceae, Faboideae), a New Species
from the Udzungwa Mountains, Tanzania

Roy E. Gereau and Sharon Bodine

Missouri Botanical Garden, P.0. Box 209, St. Louis, Missouri 63166-0299, U.S.A.
roy.gereau@niobot.org; sharon.bodine@mobot.org

Abstract. We describe and illustrate the new
species Crotalaria mwangulangoi (Fabaceae, Fa¬
boideae) from the Udzungwa Mountains of south¬
central Tanzania and compare it with the morpho¬
logically similar species C. hemsleyi from the Nguru
and Uluguru Mountains of eastern Tanzania.

hey words: Crotalaria, Fabaceae, Faboideae,
Tanzania, Udzungwa Mountains.

Crotalaria L. (Fabaceae, Faboideae) is a genus
of ea. 600 species found throughout the tropics and
subtropics (Polhill, 1982), with the greatest con¬
centration of species in eastern and southern trop¬
ical Africa. Furthermore, it is the largest genus of
vascular plants in tropical Africa (Polhill, 1982).
W ith more than 200 species now known from Tan¬
zania. Kenya, and Uganda, it is also the largest
genus in the area covered by the Flora of Tropical
Fast Africa (Polhill. 1971; Beentje, pt ars. comm.).

Together with other digitatcly 3(to I 1 )-foliolate
genera of Fabaceae subfamily Faboideae, Crotalar¬
ia has variously been treated as part of the largely
north temperate tribe Genisteae (Adanson) Ben-
tham (e.g.. Wilezek, 1953; Polhill, 1971) or the pri¬
marily tropical tribe Crotalarieae Hutchinson (e.g..
Polhill. 1976, 1982; Van Wyk & Schutte, 1995; Du
Puy & l.abat. 2002). Crotalaria may be distin¬
guished from the other genera of Faboideae1 in trop¬
ical East Africa by the following combination of
characters (Gillett et ah. 1971): leaves usually dig-
i lately 3(to 7)-foliolate, sometimes simple or uni-
foliolate; stipels lacking; leaflets entire; calyx not
spalhaeeous; keel beaked, not helically coiled: sta¬
mens united; filaments not expanded beneath an¬
ther; anthers alternately long and short; legume
usually inflated.

I he Udzungwa Mountains National Park, gazett¬
ed in 1992, protects approximately 20% of the
whole Udzungwa Mountains, which form the south¬
ern end of the Eastern Are Mountains of Kenya and
Tanzania (Luke & Beentje, 2003). Recent botanical
inventories in I dzungwa Mountains National Park
reveal the presence of a new species of Crotalaria

apparently allied to another species from adjacent
parts of the Eastern Arc Mountains.

Crotalaria mwangulangoi Gereau Bodine, sp.
nov. TYPE: T anzania. Iringa Region: Iringa
Rural Distr.. Udzungwa Mountains National
Park, grassland, 7°37'47"S, 36°36'58"E. 1600
m, 10 Sep. 1999, N. A. Mwangulango, A. //.
Seki & C. Leliyo 402 (holotype, Nil T; isotypes,
GRA. K. MO. S). Figure 1.

Haec species quoad stipulas non foliaceas, folia trifo-
liolata, foliola utrinque puhescentia, braeteas elongatas
persistentes. calycem contra pedicellum deflexion diini-
diuni longitudinis carinae non attingentem, carinam prope
medium rotundatam, rostrum non tortum atque legumen
subsessile triebornatibus patentibus vestitum ad Crotalar-
iam hemsleyi maxime accedit, sed ab ea stipulis longior-
ibus, foliolis late elliptieis, bracteolis prope medium ped-
ieelli insertis, alis earina brevioribus, rostro longiore atque
legumine majore facile distinguitur.

Shrub to I m tall; branches very finely ribbed,
densely pubescent with fine spreading tawny Iri-
chomes. Stipules subulate to setaceous, 2-4 mm
long, largely obscured by indumentum: petiole 0.5-
1.5 cm long, with fine spreading tawny trichomes;
leaves 3-foliolate; leaflets broadly elliptic, obtuse to
rounded and often apieulate at apex, broadly eu-
neate to obtuse at base, the terminal 1.2-2. 7 X
0.7-1. 7 cm, the laterals 0.7— 2.3 X 0.5— 1.6 cm,
coarsely and somewhat spreading-pubescent on up¬
per surface, densely pubescent on lower surface,
with trichomes more spreading and pronounced
along nerves. Racemes terminal, 5—12 cm long,
with ca. 15 to 30 closely arranged flowers; bracts
subulate-caudate, expanded at base, 3—6 mm long,
long persistent; pedicels 3—4 mm long at anthesis;
bracteoles in middle to upper half of pedicel, fili¬
form. 2—3 mm long. Calyx becoming basally trun¬
cate and moderately deflexed against pedicel, the
tube 3-4' mm long, pubescent with irregularly ar¬
ranged mostly spreading brown or reddish brown
trichomes, the lobes acuminately triangular, 4—5
mm long; standard subcircular, ca. 1.5 cm long,
yellow inside, with maroon lines outside, glabrous
outside or with very' few white trichomes along mid-

Novon 15: 286-289. 2005.

Volume 15, Number 2
2005

Gereau & Bodine

Crotalaria mwangulangoi from Tanzania

287

Figure I. Crotalaria mwangulangoi Gereau & Bodine. — A. Flowering and fruiting branch apex, habit. — B. Flower
and pedicel, lateral view. — C. Attached fruit with infructescence node. — I). Seed and funicle. — F. Leaf, abaxial
surface. — F. Leaf, adaxial surface. — G. Ovary, style, and stigma. — H. Long anther, lateral view, and short anthers,
lateral and ventral views. — I. Keel, lateral view. — J. Wing petal, abaxial surface. — K. Standard petal, abaxial surface.
Drawn from Mwangulango et al. 402 (GRA, K. MO, and S isotypes).

288

Novon

ril> near apex; wings slightly shorter than keel; keel
rather strongly rounded a little below middle, with
a slightly incurved, untwisted beak, 1.5— 1.6 em
long, lime green with maroon lines. Legume sub-
sessile, cylindrical, rounded at base, 2.7— 3.5 X
0.7— 1.0 em. with spreading tawny triehomes ea. I
mm long outside, glabrous inside, ea. 30-seeded;
seeds oblique-cordiform, 2.2— 2.6 nun long, smooth,
shiny, reddish brown.

Habitat and distribution. Known only from the
type collection, in grassland at 1600 m. The re¬
cently described Vernonia luhomeroensis Q. Luke
& Beentje (Asteraceae) was collected within 3 km
of Cralalaria mwangulangoi and at similar eleva¬
tions (Luke & Beentje, 2003); however, V. luhom-
eroensis is a species of submontane to montane for¬
est. The recent discovery of two new species in this
area demonstrates the need for much more inten¬
sive botanical inventory of Ud/ungwa National
Park.

Provisional conservation status. With an Extent
of Occurrence (EO) of less than 5000 km2 and oc¬
curring only in the Udzungwa Mountains National
Park, Tanzania, and already listed as potentially
threatened (Gereau & Luke, 2003): Endangered
(Blab), using IUCN (2001) criteria.

In the infrageneric classification presented by
Polhill (1982), Crotalaria mwangulangoi clearly
falls within Crotxdaria seel. Hedriocarpae Wight «X
Arnolt subsect. Macrostachyae (Bentham) F. A. Bis-
by & Polhill by the following combination of char¬
acters: hypanthium not prominent; calyx less than
half as long as keel; standard with maroon lines
outside; keel rounded near middle, with untwisted
beak; style not dilated nor bifurcate at apex, not
geniculate; legume subsessile, rounded at base,
glabrous inside, cylindrical. Within this subsection,
it is morphologically most similar to C. hemsleyi
Milne-Redhead, from which it may be distin¬
guished by the following key:

Kkv Charactkus <u Ckotaiaria HKMLSLE) i and C. l/ll t\

GVLANGOI

la. Stipules ea. I mm long; leaflets elliptic. 2.5— 7.0

cm long; bracteoles at base of calyx; calyx
st rough deflexed against pedicel; wings a little
longer than keel; legume 2.5— 2.7 X ea. 0.6 cm;
seeds 2.5— 3.0 mm long . C. hemsleyi

lb. Stipules 2—4 mm long; leaflets broadly elliptic.

9.7— 2.7 cm long; bracteoles in middle to upper
half ol pedicel; calyx moderately deflexed against
pedicel; wings slightly shorter than keel; legume

2.7— 3.5 X 0.7— 1.0 cm; seeds 2.2— 2. 0 mm long
. O’. mwangulangoi

The morphologically similar Crotxdaria hemsleyi
is known only from upland grasslands between

1200 and 2000 m in tlx* Nguru and Uluguru Moun¬
tains of Tanzania, while C. mwangulangoi is known
only from the type collection in montane grassland
at 1600 m in the Udzungwa Mountains. Lovett
(1998) subdivided the Eastern Arc Mountains of
Kenya and Tanzania into three subregions: North-
mil, comprising the Taita Hills, North and South
Pare Mountains, East and West Usambaras; Cen¬
tral, comprising the Nguu, Nguru, Ukaguru, Ulu-
guru, and Rubeho Mountains; and Southern, com¬
prising the Udzungwa and Mahenge Mountains.
Gereau et al. (in press) document a limited degree
of species-level endemism in the Central subregion,
to which Crotalaria hemsleyi contributes; C. mwan¬
gulangoi is apparently a sister species to C. hem¬
sleyi, endemic to similar habitats in the adjacent
Southern subregion.

Acknowledgments. We sincerely thank the
WWF Tanzania Programme Office for support of
Nathan Mwangulango during his work for the Tan¬
zania Botanical Training Programme, Ouentin Luke
for alerting us to the publication of Vernonia lu¬
homeroensis, John Flanagan of the library at the
Royal Botanic Gardens, Kew, for expediting a re¬
print of that article, and John Myers for another fine
illustration.

I .iterature Cited

Du Pay, I). ,|. & J.-N. I .a bat. 2902. Crotalarieae. Pp. 009—
798 in I). .1. Ihi Pay, J.-N. I.abat. If. Rabevohitra, J.-F.
Villiers, ,|. Bosser & J. Moat. The Leguminosae of Mad¬
agascar. Hoyal Botanic Gardens, Kew.

Gereau, B. 1C & W. If. D. Lake. 2993. List of Potentially
Threatened Plants in the Eastern Are Mountains and
Coastal Forests of Tanzania & Kenya Biodiversity Hot¬
spot. Unpublished Report, Critical Ecosystem Partner¬
ship Fund & Conservation International, Washington,
DC.

- , C. M. Taylor & W. If. (J. Luke. In press. Endemic

plant species of the Eastern Arc Mountains of Kenya
and Tanzania: Analysis and refinement of distributional
patterns. Proceedings of the XVII A E l i A T Congress,
21-20 September 2003. Addis Ababa.

Gillett, J. B.. If. M. Polhill & B. Verdeourt. 1971. Legu¬
minosae subfamily Papilionoideae. In E. Milne-Red¬
head & If. M. Polhill (editors). Flora ol Tropical East
Africa. Crown Agents for Oversea Governments and Ad¬
ministrations. London.

IUCN. 2991. II CN Red List Categories and Criteria Ver¬
sion 3.1. Prepared by the 11 CN Species Survival Com¬
mission. IUCN, Gland, Switzerland.

Lovett. J. C. 1998. Importance of the Eastern Are Moun¬
tains for vascular plants. J. E. African Nat. Hist. 87: 1-
20.

Luke, (J. & II. Beentje. 2993. A new Vernonia (Coinpos-
itae: Vernonieae) from the Udzungwa Mountains. Tan¬
zania. kew Bull. 58: 977—980.

Polhill, If. M. 1971. Crotalaria. Pp. 817—994 in J. B.
Gillett, If. M. Polhill & If. Verdeourt, Leguminosae sub¬
family Papilionoideae. In: E. Milne-Redhead & R. M.

Volume 15, Number 2
2005

Gereau & Bodine

Crotalaria mwangulangoi from Tanzania

289

Polhill (editors), Flora of Tropical Kast Africa. Crown
Agents for Oversea Governments and Administrations,
I .ondon.

- . 1976. Genisteae (Adans.) Benth. and related

tribes (Leguminosae). Bot. Syst. 1: 143—368.

- . 1982. Crotalaria in Africa and Madagascar. A.

A. Balkema, Rotterdam.

Van Wyk, B.-K. & A. I.. Schutte. 1995. Phylogenetic re¬

lationships in the tribes Podalyrieae, Liparieae and Cro-
talarieae. In: M. Crisp J. .1. Doyle (editors), Advances
in Legume Systematics 7 (Phylogeny): 283—308. Royal
Botanic Gardens, Kcvv.

Wilczek, R. 1953. Genisteae. In: Comile ex6cutif de la
Flore do Congo beige el le Jardin Botanique de 1' ft tat,
Flore do Congo beige et du Ruanda-Urundi 4: 46—
288.

A New Synthetic Allopolyploid Naranjilla,
Solarium indianense (Solanaceae)

Charles Heiser

Department of Biology, Indiana University, Bloomington, Indiana 47405, U.S.A.

ebheiser@bio. indiana.edu

Jorge Soria

Avenida La Bepiibliea 700, Quito, Ecuador, jasv@intertelec.net

Carlos Miller

Department of Biology, Indiana University, Bloomington, Indiana 47405, U.S.A.

millerc@indiana.edu

Gregory Anderson

Department of Ecology and Evolution, University of Connecticut, U-43, Storrs,
Connecticut 06269, U.S.A. ander@uconnvm.uconn.edu

Abstract. The chromosomes of two nearly sterile
hybrids (Solarium t/uitoense X .S'. sessUijloriim) have
been doubled by colchicine treatment and the re¬
sulting tetraploids are described as a new species,
.S’, indianense. I lie parents, now grown in Ecuador
under the names Puyo and Palora, are dignified
with cultivar status.

Bl-si MKN. Los cromosomas de dos hfbridos easi
esterdes (Solanum quitoense X .S', sessilifiorum) han
sido duplicados mediante tratamiento eon colchi-
cina y los tetraploides resultantes s<‘ describen
como nueva especie. S. indianense. Los padres que
se cultivan actuahnente en Ecuador, bajo los nom-
bres de Puyo y Palora, son reeonocidos bajo la ca-
tegorfa de cultivares.

Key words: corona, Ecuador, hybrid, naranjilla,
pol y p loi dy, Sol anu m .

Over 90% of the naranjilla, much esteemed for
its fruit juice, grown in Ecuador today comes from
the f j hybrids Solatium quitoense Lamarck X S.
sessilifiorum Dunal var. sessilifiorum and S. </ui-
toense X .S', sessilifiorum var. georgicum (R. E.
Schultes) \\ halen. The former is commonly known
under the name Palora and the latter as Puyo from
the places where they were first grown in Ecuador.

I hese nearly sterile diploid hybrids are propagated
vegetatively. A discussion of their origins may be
found in Heiser (1993) and Heiser and Anderson
(1999). Both ol these hybrids had their chi omo-
somes doubled by colchicine in the greenhouses of

Novon 15: 290-292. 2005.

Indiana University. I he tetraploids, propagated by
seed, are described as a new species.

Solatium indianense C. Heiser & J. Soria, sp. nov.
[Solarium quitoense Lamarck X S', sessilifiorum
Dunal]. I YPE: U.S.A. Indiana: Bloomington,
cultivated in the greenhouse of Indiana Uni¬
versity (tetraploid Palora), 15 Eeb. 2004, C.
Heiser CHI -l (holotype, MO; isotypes, INI),
QCNE).

Plantae hybrido Solano quitoensi X S. sessilifloro simi¬
les sed prope omnibus partibus majoribus; pollinis grana
diametro 26—28 yam; 60% vel plus tingibiliu; frutus 60—
seminalis vel plus, seminibus plus quam 3.0 mm longis;
chromosomatum numerus tetraploideus (n = 24).

Lignescent perennials 1 .5-2.8 m high, unarmed,
vegetative parts green, rarely with red or purple
tinges; stems woolly, with sessile to stalked stellate
trichomes (stellae), stalks 1—6 mm, lateral rays 5 to
9, 0.4-11.7 mm long, midpoints shorter than or
nearly equaling rays. Leaves felly-pubescent, very
large, blades to nearly I m long, broadly ovate,
repand, cordate at base with 5 to 7 major veins on
either side; the repand margins with 5 to 6 deltoid,
acute- or obtuse-tipped lobes at lateral vein ter¬
minations; interlobal sinuses often with 1 to 3
smaller, tooth-like lobes; adaxial leaf surface with
mostly sessile stellae with elongate midpoints 1-2
mm long and reduced lateral rays 0.05—0.3 mm
long, a distinct class of smaller stellae present or
lacking; major veins with both stalked and sessile

Volume 15, Number 2
2005

Heiser et al.

Synthetic Allopolyploid Naranjilla

291

stellae, stalks to 1 mm; abaxial leaf surfaces dense¬
ly tomentose with interwoven stalked and sessile
stellae, usually lacking midpoints and with well-
developed rays of varying length, sometimes over I
mm long; petioles typically 14 to Vi the length of
the blade, inflorescence 3- to 20-flowered; the low¬
ermost 4 to 6 flowers perfect; subsequent ones fe¬
male-sterile; pseudoaxis 0.7—22 mm long; pedicels
4—12 mm long in flower; ealvx broadly campanu-
late, green, stellate-pubescent; calyx lobes broadly
ovate, acute tipped, 14-19 mm long; corolla white,
4.8— 6.2 cm across, divided to near base into ovate
or ovate-lanceolate lobes, externally stellate-pubes¬
cent; anthers 10-14 X 3—5 mm; style glabrous, 8—

I I X 1. 5-2.0 mm. berries I to 5 per inflorescence,
globose, 4. 2-7.0 cm diam., 4-locular, orange when
ripe with light green to orange flesh, densely stel¬
late-pubescent at maturity, hairs soon deciduous,
pericarp stellae sessile with elongate midpoints,
0.6— 1.2 mm long with 9 to 15 lateral rays near base
about 0.2 mm long; seeds from ea. 60 to 150 or
more, light tan. lenticular, broadly ovate, 3.0— 3.8
mm long. Chromosome number, n — 24.

The following key should serve to distinguish the
new species from the other large-fruited members
of Solarium sect. Lasiocarpa cultivated in the
Americas.

K i.x to Lak<;k-Fiu ited Cli.iivatki) American Species
ok Soiam u Sect. Iasiocari’a

la. I. eaves and stems prickly . 1.5. quitoense

Ih. Leaves and stems unarmed.

2a. Berries deep red or brown to orange- red or
orange, usually over 5.0 cm diam. with pu¬
bescence of very readily deciduous, soft
stellate hairs with midpoints rarely longer
than 1 mm; fruit flesh orange; young leaves
and stems green; mostly grown below 500 m

. 2. .S', sessiliflorum var. sessiliflorum

21). Berries orange, small to large, with pubes¬
cence of deciduous, slightly harsh stellate
hairs with midpoints often longer than 1
mm; fruit flesh orange to green; young leaves
and stems purplish, reddish or green; mostly
grown above 900 m.

3a. Fruit flesh usually deep green; fruit
diam. 4.5— 5.5 cm; midpoints of stellae
of most fruit hairs over 3.5 mm long;
young leaves and stems w ith strong red¬
dish or purplish cast; pollen 19—21 gun,
over 90% of pollen taking stain; seeds
several hundred, usually less than 3.0

mm long . 1 • .S', quiloense

3b. Fruit flesh orange or light green; fruit
small to large; midpoints of stellae of
fruits less than 2 mm long; young leaves
and stems green or slightly reddish or
purplish; pollen of various sizes, fre¬
quently less than 90% stainable; seeds

usually 100 or less, usually over 3.0
mm long.

4a. Fruit (lesh very light green or or¬
ange; berries small to large.

5a. Berries naturally less than 3.8
cm diam. (nearly to 6.0 cm
when sprayed with 2,4-1 )); pol¬
len stainability less than 40%;
pollen around 21 gin, tricol-
pate; seeds few or none ....

. 3. S. quitoense X

S. sessiliflorum “Puyo’
51). Berries 4.0 to 5.4 cm diam.;
pollen stainability seldom less
than 70%; pollen 26—28 gun;
some grains tetraeolpate; seeds

60 or more . 4. S. indianense

(tetraploid Puyo)

4b. Fruit flesh orange; fruits usually
over 5.5 cm diam.

6a. Seeds few1 or none; pollen
stainability 40%' or less; pollen
around 21 gun, tricolpate . . .

. 3. .S. quitoense X

S. sessiliflorum “Palora"
6b. Seeds 60 or more; pollen stain¬
ability usually over 70%, pol¬
len 26-28 gun, some grains te-

tracolpate . 4. S. indianense

(tetraploid Palora)

1. Solarium quitoense. The prickly naranjilla
dominates the area of cultivation from Colombia to
Guatemala. Unarmed plants are grown in Ecuador
and Colombia. This species is also grown as an
ornamental in the United States. In Latin America
the naranjilla may escape front cultivation and es¬
tablish weedy populations (Whalen et al.. 1981).

Two autotetraploids of a naranjilla have been se¬
cured by colchicine treatment. The plants have pro¬
duced few fruits and still fewer seeds. I luce nearly
sterile hybrids have been secured in a cross with
S. indianense as the male parent.

2. Solatium sessiliflorum. The typical variety,
usually known as eoeona, is grown primarily in the
eastern lowlands of the northern Andean countries
extending into Brazil (Whalen et al.. 1981). Unlike
S. quitoense it is quite variable, particularly in char¬
acters of the fruit. Escapes from cultivation are not
uncommon (Whalen et al., 1981).

3. Solatium quitoense X .S’, sessiliflorum var.
georgicum “Puyo" and S. quitoense X .S’, sessiliflo¬
rum var. sessiliflorum “Palora.” These F, hybrids,
which are propagated by cuttings, are mostly inter¬
mediate between their parents, although some char¬
acters are slightly larger than in either of the par¬
ents because of hybrid vigor. I he Puyo hybrid
normally produces a rather small fruit (3.0— 3.8 cm
diam.), but spraying the (lowers with a dilute so¬
lution of 2,4-D gives a much larger fruit. This prac-

292

Novon

lice Ims become a regular part of its culture in Ec¬
uador.

4. Solarium indianense. As can be seen from
the key, the two tetraploids that constitute ibis new
species are propagated by seeds and are very sim¬
ilar to their diploid progenitors, which are dis¬
cussed in #4 above. Their very thick stems usually
readily distinguish the tetraploids from the diploids
but it is difficult to give precise measurements be¬
cause stem size is influenced by soil fertility. The
tetraploids are more variable than their diploid pro¬
genitors in height, presence and amount of red pig¬
ment in the leaves and stems, fruit size, flesh color,
fertility, and time of flowering. The great variability
can probably be explained by chromosome behav¬
ior at meiosis. Thirty-five cells were examined in
one of each of the two tetraploids. From 12 to 24
bivalents were found with the other chromosomes
associating in up to six quadrivalents (rarely as tri-
valents and univalents). If regular pairing (24 pairs)
occurred with normal segregation of the chromo¬
somes, little or no variation would be expected
(Stebbins, 1971). Other causes of variation are also
now known in neopolyploids (Sehranz & Osborn,
2001).

It is too early to propose eultivar names for the
new species, both of the tetraploids, as well as hy¬
brids between them, are now being grown in Ec-
uador. From these plants, selections will be made
with more desirable combinations of characters that
will bee •ome more widely grown and deserving of
names. \\ ith selection we can also anticipate better
chromosome pairing and an increase in fertility
(Ramsey & Schcmske, 2002).

Discussion

Nearly a hundred years ago when it was discov¬
ered that colchicine could induce polyploidy, there
was great hope that newly created polyploids would
add greatly to our cultivated crops. Only a few have
proven successful and these usually only alter
many years of work (Simmonds & Smartt. 1999).
I here is some reason to believe that Solarium in-
dianense may be successful, for it should be able
to outperform its chief competition, the diploid hy¬
brids, neither of which can be readily improved as
they rarely reproduce sexually. In addition to be¬
coming established as cultivated plants, it is pos¬
sible that plants of S. indianense will become weedy
escapes.

Acknowledgments. We thank Marti Crouch, El¬
len Dean, Gerald Gastony, Loren Rieseberg, and
Tyson Roberts.

Literature Cited

Heiser, C. 1993. The naranj ilia (Solarium quitoense), the
cocona (Solarium sessiliflorum ) and their hybrid. Pp.
29—34 in J. Gustafason, If. Appels & P. Raven, Gene
Conservation and Exploitation. Plenum Press, New
York.

- & G. Anderson. 1999. “New” solanums. Pp. 379—

384 in J. Janick, Perspectives on New Crops and New
Uses. ASUS Press, Alexandria, Virginia.

Ramsey, J. & I). Schcmske. 2002. Neopolyploidy in flow¬
ering plants. Annual Rev. Ecol. Syst. 33: 589-639.
Sehranz, N. & I. Osborn. 2001. I)e novo variation in life-
history traits and responses to growth conditions of re¬
synthesized polyploid Brassica no pus (Brassicaceae).
Amer. J. Rot. 91: 174-183.

Siinonds, N. & J. Smartt. 1999. Principles of Crop Im¬
provement. Blackwell, London.

Stebbins, G. 1971. Chromosomal Evolution in Higher
Plants. Addison- Wesley, Menlo Park, California.
Whalen, M.. I). Costieh & C. Heiser. 1981. Taxonomy of
Solarium sect. Ixisiocarpa. ( rentes Herb. 12: 41—129.

New Combinations and a New Species in Morelia (Myricaceae)

Jane Herbert

School of Biology, Sir Harold Mitchell Building, l niversity of St Andrews, k\ 16 9TH,

United kingdom, jh53@st-and.ac.uk

ABSTRACT. I he molecular support for current ge¬
neric concepts in Myricaceae lias made it necessary
to make new combinations in Morelia. Morelia com¬
prises approximately 47 species found in the Old
and New World tropics, three new combinations
are made here: Morelia adenophora, M. nan a , and
M. punctata. Morelia nana is leetotypified, and a
new species, M. rivas-martinezii, is described.

Key words: Canary Islands. China, Cuba, Mo¬
relia, Myriea, Myricaceae, Taiwan.

With tin* exception of the monotypic genera
Comptonia E’Heritier ex Aiton (northeastern
U.S.A.) and Canacomyrica Guillaumin (New Cale¬
donia), the species of Myricaceae have traditionally
been referred to the Uinnaean genus Myriea (Cron-
quist, 1981: kuhitzki. 1993; Zomlefer, 1994; Takh-
tajan, 1997). There are, however, significant differ¬
ences between the two deciduous species with dry
fruits of the temperate Northern Hemisphere (Myr¬
iea gale L. and M. hartwegii S. Watson) and the
remaining ca. 47 evergreen species with papillose,
fleshy fruits that are disjunctly distributed in the
Old and New World tropics. Some authors have rec¬
ognized these two groups at the subgeneric level
(Engler, 1888; Melchior, 1964; Elias, 1971), while
others have given them generic status (Spach,
1841: Chevalier, 1901; Baird, 1969, unpublished
thesis, University of North Carolina, Chapel Hill;
Wilbur, 1994; Judd et ah, 1999; Pol hill & Verd-
court, 2000). Previous authors have used both Myr¬
iea E. and Morelia Loureiro to refer to the evergreen
species, and a long history of nomenclatural diffi¬
culties is associated with the typification of these
two genera.

In the only monograph of the family, Chevalier
(1901) placed the dry-fruited species in the genus
Gale Duhamel. It can be assumed that he consid¬
ered one of the fleshy-fruited species to typify the
genus My rica, in which he placed the majority of
species. However, he did not cite types for these
genera. Subsequent authors (e.g.. Baird, 1969, un¬
published; Wilbur, 1994) have retained Myriea for
the dry-fruited species, adopting Morelia for the re¬
maining taxa.

Recently, Verdeourt and Polhill (1997) made a

proposal to conserve the name Myriea with the con¬
served type of M. cerifera E. (one of the fleshy-
fruited species) in an attempt to minimize the num¬
ber of necessary name changes. However, the
Committee for Spermatophyta (Brummill, 1999) re¬
jected the proposal. Thus Myriea gale is the lee-
totype of Myriea (Herb. Einn. No. 373 1 1 list.
France]; lectotype designated by Jonsell & Jarvis
(in Jarvis et ah, 1993)), and the majority of species
previously known as Myriea must be transferred to
Morelia. The following key separates the two gen¬
era:

ki \ to Diffkr kntiatk Myrica and Morelia

la. Shrubs deciduous; leaves with sunken stomata;

inflorescences borne on the previous year's
growth; fruits dry with adherent spongiose brac-
teoles, water-dispersed . 1 1 erica

lb. Shrubs or trees evergreen; leaves with stomata
not sunken; inflorescences borne on the present
year’s growth; fruits papillose, fleshy, sometimes
with a waxy covering, bird-dispersed .... Morelia

New combinations in Morelia have been made
for species in North America (Wilbur, 1994). Africa
(Killick et ah, 1999). Malesia (Turner, 2001 ), South
America (Parra-O., 2002), and some in Central
America and the Caribbean (Wilbur, 2001; Knapp,
2002). In the course of the preparation of a Ph.I).
thesis on Myricaceae, in whic h new molecular data
support the recognition of Morelia (Herbert, 2005),
it became apparent that some new combinations in
Morelia were necessary. A new species is described
and three new combinations are made here to pro¬
vide valid names for species recognized in a forth¬
coming molecular phylogenetic paper.

Morelia adenophora (Hance) J. Herbert, comb,
nov. Basionym: Myrica adenophora Hance, J.
Bot. 21: 357. 1883. TYPE: [China.) “In di-
cione Ting-on ins. Hai-nan, m. Nov. 1882,” II.
C. Henry 22159 (holotype, BM).

This species occurs in Hainan, Guangdong, and
Guangxi in China (Lu & Bornstein, 1999); variety
kusanoi Hayata is recognized in the* Flora of Tai¬
wan (Yang & Lu, 1996). Morelia adenophora is a
shrub (up to 3 m in height) and is closely related

Novon 15: 293-295. 2005.

294

Novon

to Morelia liana , which occurs in the neighboring
provinces of Yunnan and Guizhou (see below). Mo¬
relia adenophora is also closely related to Morelia
rubra Loureiro, which is widely cultivated in China
and also occurs in Japan. The fruits of M. rubra
(up to 3 cm diam.) are significantly larger than
those of M. adenophora (up to 1 cm diam.). All
three species share the character of a single fruit
developing at the apex ol tlx- infrueteseence, thus
differing from the other Asian species, such as M.
javanica (Illume) I. M. Turner, in which more than
one fruit per infrueteseence develops.

Morelia nana (A. Chevalier) J. Herbert, comb,
nov. Basionym: Myrica nana A. Chevalier,
Mem. Soc. Sei. Mat. Cherbourg 32: 202. 1901.
TYPE: China. Yunnan Province: Bois de Mao-
Kou-Tehang, an dessus du Tapintz, P. ./. M.
Delaray 148 (lectotype, designated here, P).

This species is found in central and northern
Yunnan and western Guizhou in China. At up to 3
m in height, it is much smaller than the more wide¬
spread M. rubra (up to 15 m). When Chevalier
(1901) described this species he had not seen spec¬
imens of M. adenophora ; the ranges of the two spe¬
cies are contiguous, but detailed population-level
studies of morphology and/or genetics would be re¬
quired to confirm their status. Morelia nana is rec¬
ognized in the Flora of China (Eu & Bornstein,
1 999).

\lorella punctata (Grisebach) J. Herbert, comb,
nov. Basionym: Myrica punctata Grisebach,
Mem. Acad. Amer. Sci. Art N. S. 8 [Plantae
Wright ianae, [it. I|: 177. 1860. TYPE: [Cuba.]
“In saxosis prope Monte Verde, Feb,” C.
Wright 1460 (holotype, (ill).

This species is endemic to Cuba. It has entire
oblong-ohlanceolate leaves (3—5 cm long) and ovoid
fruits (Leon & Alain, 1951). distinguishing it from
the widespread Morelia cerifera (L.) Small, which
is found throughout the Caribbean (also in the Unit¬
ed States and Central America) and which has lon¬
ger leaves (to 8 cm) and globose fruits.

Morelia rivas-martinezii A. Santos & ,|. Herbert,
sp. nov. TYPE: Spain. Canary Islands: Isla tie
Hierro, La Dehesa, 22 Feb. 1976, A. Santos
24690 (holotype, OB I ).

A. Santos (1980) intended to publish “ Myrica
rivas-martinez ” as a new species. Because different
gatherings were cited as “llolotypi atque isotypi”
(holotypes and isotypes), the name was not validly

published (Greuter et al., 2000: Art. 37.1 & 37.2,
with Art. 8.1 & 8.2). Dr. Santos has accepted my
invitation to publish the taxon here as a new spe¬
cies of Morelia (on the basis of his 1980 descrip¬
tion), and this we do above.

This species is sympatric with Morelia faya (Ai-
ton) Wilbur in the laurel forests of La Gomera, El
Hierro, and La Palma in the Canary Islands. The
mature leaves, which are small (up to 20 mm) and
spathulate, are distinct from the much larger and
oblanceolate leaves of M. faya.

Acknowledgments. I thank Sandra Knapp,
Thomas Lammers, and one anonymous reviewer for
comments on an earlier draft of this manuscript.
Ian Hedge (E) is thanked for comments on the man¬
uscript. John McNeill (E) is thanked for nomencla-
lural guidance, Peter Phillipson (MO, at P) and Ar-
noldo Santos (OB I ) are thanked for checking types.
The assistance of the following is gratefully ac¬
knowledged: the curator of the herbarium ol the
Natural History Museum, London (BM) for loan of
material, the curator and staff ol the Boyal Botanic
Gardens Kewr (K) for use ol the herbarium, and the
staff at the Harvard University Herbaria (GH) and
The New York Botanical Garden (NY) for supplying
digital images of types. My studies on the generic
concepts in Myrieaeeae were supported by an
NEBC studentship (NER/S/A/2(X)0/03638).

Literature Cited

Baird, J. If. 1969. A Taxonomic Revision of the Plant
Family Myrieaeeae of North America, North of Mexico.
Ph.D. Thesis, University of North Carolina at Chapel
Hill. | Unpublished.]

Rrummitt, If. K. 1999. Proposals to conserve or reject.
Report on the Committee on Spermatophyta (1291—
1292). Taxon 48: 367-368.

Chevalier, A. 1901. Monographic des Myricac^es. Mem.

Soc. Natl. Sci. N Nat. Math. Cherbourg 32: 85—340.
Cronquist, A. 1981. Myrieaeeae. Pp. 214—217 in An In¬
tegrated System of Classification of Flowering Plants.
Columbia Univ. Press, New York.

Elias, T. S. 1971. The genera of Myrieaeeae in the south¬
eastern United States. J. Arnold Arbor. 52: 305—318.
Kngler, A. 1888. Myrieaeeae. Pp. 26-28 in A. Engler &
k. Prantl (editors), Die natiirliehen Pflanzenfamilien III.
t. Engelmann, Leipzig.

Greuter, W., J. McNeill, F. If. Barrie. II. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). If. Nicolson, P C. Silva,
J. E. Skog, P. Trehane, N. J. Turland I). I.. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Herbert, J. 2005. Systematies and Biogeography of Mvri-
caceae. Ph.D. Thesis, University of St Andrews. (Un¬
published. |

Jarvis, C. E., F. R. Barrie, I). M. Allan & J. 1,. Reveal.
1993. A List of Linnaean Generic Names and their
Types. Regnum Veg. 127.

Judd, W. S.. C. S. Campbell, E. A. Kellogg & I’. F. Stevens.

Volume 15, Number 2
2005

Herbert

Myricaceae

295

1999. Myricaceae. Pp. 314—315 in Plant Systematics,
a Phylogenetic Approach. Sinauer, Sunderland, Mas¬
sachusetts.

K i II ii k. D. J. IP. R. M. Polhill & B. Verdcourt. 1999. New
combinations in African Myricaceae. Kew Bull. 53:
993-995.

Knapp, S. 2002. A new combination in Morelia (Myrica¬
ceae) in Mesoamerica. Novon 12: 200.

Kubitzki, K. 1993. Myricaceae. Pp. 453-457 in K. ku-
bitzki, J. G. Rohwer & V. Bittrich (editors). The Fami¬
lies and Genera of Vascular Plants, Vol. 2. Flowering
Plants: Dicotyledons. Springer- Verlag, Berlin.

Leon, H. & H. Alain. 1951. Myricaceae. Pp. 36-37 in
Flora de Cuba. Dicotiledoneas: Casual inaccae a Mcli-
aceas. Contr. Ocas. Mus. Hist. Nat. Colegio “De La
Salle." 10. Havana.

Lu, A. M. & A. J. Bomstein. 1999. Myricaceae. Pp. 275—
276 in W. Zheng-yi & P. II. Raven (editors). Flora of
China, Vol. 4. Science Press, Beijing, and Missouri Bo¬
tanical Garden Press, St. Louis.

Melchior. 11. 1964. Myricaceae. P. 40 in Fngler’s Syllabus
der Pflanzenfamilien. Borntraeger. Berlin.

Parra-O., C. 2002. New' combinations in South American
Myricaceae. Brittonia 54: 322—326.

Polhill. R. M. & B. Verdcourt. 2000. Myricaceae. Pp. I —
12 in H. J. Beentje & S. A. L. Smith (editors). Flora of
Tropical East Africa. Balkema, Rotterdam.

Santos, A. 1980. Contribucion al conocimiento de la flora
y vegetaeion de la isla de Hierro. (I. Canarias). Fund.
Juan March, Ser. Univ., Cicnc. Agrar. I 14: 45—16.

Spach, E. 1841. Myricaceae. Pp. 256—266 in Histoire Na-
turelle des Vegetaux. Phanerogames. Vol. II. Paris.

Takhtajan, A. 1997. Myricaceae. Pp. 153-154 in Diversity
and Classification of Flowering Plants. Columbia Univ.
Press, New York.

Turner, I. M. 2001. New combinations in Malesian Myri-
caceae. Card. Bull. Singapore 53: 323—325.

Verdcourt, B. vK R. M. Polhill. 1997. Proposal to conserve
the names Myrica and Gale (Myricaceae) with con¬
served types. Taxon 46: 347—348.

Wilbur. R. L. 1994. The Myricaceae of the United States
and Canada: Genera, subgenera, and scries. Sida 16:
93-107.

- . 2001. f ive new combinations in the genus Mo¬
relia (Myricaceae) for Neotropical species. Rhodora
103: 120-122.

Yang. Y. P. & S. Y. Lu. 1996. Myricaceae. Pp. 19—22 in
T. C. Huang (editor). Flora of Taiwan, Vol. 2. Angio-
spermae. Editorial Committee of the Flora ol Taiwan,
2nd edition. Taipei.

Zomlefcr, W. B. 1994. Myricaceae. Pp. 174— 176 in Guide
to Flowering Plant Families. Univ. North Carolina Press,
Chapel Hill & London.

A New Combination in Myrcia (Myrtaceae)

Bruce K. Holst

Marie Selby Botanical Gardens, 81 1 South Palm Ave., Sarasota, Florida 34236-7726, l .S.A.

bholst@selby.org

Abstract. In preparation for 1 he treatment of
Myrcia (Myrtaceae) for Flora Mesoamericana, the
new combination Myrcia zetekiana (Standley) B.
Holst is required.

Key words: Flora Mesoamericana, Myrcia, Myr¬
taceae.

My rein zetekiana (Standley) B. Holst, comb. nov.
Basionym: Eugenia zetekiana Standley, J.
Wash. Acad. Sci. 15: 286. 1925. Aulomyrcia
zetekiana (Standley) Amshoff, Ann. Missouri
Bot. Gard. 45: 170. 1958. TYPE: Panama. P.
C. Standley 2750.1 (holotype, US).

Myrcia zetekiana is one of the most unusual spe¬
cies in the subtribe Myreiinae, with dense purplish
indument and large flowers. Based on the shape of

the embryo. Amshoff (1958) correctly recognized its
placement in the Myreiinae rather than Eugeniinae,
making the transfer into Aulomyrcia. Aulomyrcia,
however, has been considered a synonym of Myrcia
for more than KM) years (kiaerskou. 1893), and so
it is a bit surprising that this striking, though in¬
frequently collected, species has not been trans¬
ferred until now.

I .iterature Cited

Amshoff, C. J. II. 1958. Myrtaceae. In It. K. Woodson, It.
W. Sehery & Collaborators, Flora of Panama, Part VII,
Fasc. 2. Ann. Missouri Bot. Gard. 45: 165—201.
Kiaerskou, It. 1893. Fnumeratio Myrtacearum brasilien-
sium. In: K. Warming (editor), Symbolarum ad Floram
Brasiliae Centralis Cognoscendam 39: 1—200. Copen¬
hagen.

Novon 15: 266. 2005.

Prockia krusei (Salicaceae), una Especie Nueva del Estado de

Guerrero, Mexico

Jaime Jimenez Ramirez v Ramiro Cruz Duran
Herbario de la Faeultad de Cieneias (FCME). UNAM, Apartado Postal 70-399,
Delegacion Coyoacan 04510, Mexico, I).K. jjr@hp.fciencias.unarn.mx

RESUMEN. Se describe e ilustra Prockia krusei
(Salicaceae). una especie nueva del estado de
Guerrero, Mexico. Se discute su ubicacibn en los
generos Batumi, Hasseltia, Hasseltiopsis, Macrohas-
seltia, Neosprucea, Olmediella, Pleuranthodendron,
Pineda y Prockia.

Abstract. Prockia krusei (Salicaceae), a new
species from the state of Guerrero, Mexico, is de¬
scribed and illustrated. Its position relative to the
genera Banara, Hasseltia, Hasseltiopsis, Macrohas-
seltia, Neosprucea, Olmediella. Pleuranthodendron,
Pineda, and Prockia is discussed.

Key words: Banara, Guerrero. Hasseltia, Has¬
seltiopsis, Macrohasseltia, Mexico, Neosprucea. Ol¬
mediella, Pineda. Pleuranthodendron, Prockia, Sal¬
icaceae.

Kn la revision de la coleccibn KRUSE en el Her¬
baria Nat ional de Mexico (MEXU), se encontrd una
especie nueva de Salicaceae (sensu Chase et ah.
2002) y se la ubieo en el genero Prockia P. Ihowne
ex I,, debido a que los sepalos alados en el baton
floral y la inflorescencia del taxon nueva (ver fig.
1C y D) son indudablemente semejantes al genero
referido. Sin embargo, la especie nueva [Prockia
krusei) tiene ovario unilocular con tres placentas
intrusivas (ver Fig. 1G). en lugar de tener ovario
eon 3—6 loculos como ocurre en Prockia, por lo que
su ubicacibn esta lejos de ser indisputable.

Los generos Banara Aublet, Neosprucea Sleumer
v Olmediella Baillon tienen ovarios uniloculares
con placentas fuertemente intrusivas, como en la
especie nueva. pero los dos primeros siempre po-
seen petalos bien desarrollados y los sepalos en el
baton floral no son alados como en el nueva taxon,
ni en Prockia. En el caso de Olmediella, la dislin-
cibn es mas sencilla pues sus flares apetalas son
unisexuales y su fruto es capsular, mientras que
Prockia krusei y todos los generos mencionados
basta este momento poseen llores bisexuales y ba-
yas.

Los generos Hasseltiopsis Sleumer. Macrohassel¬
tia L. O. Williams y Pleuranthodendron L. 0. Wil¬

liams presentan ovarios uniloculares con placentas
muy ligeramente intrusivas, el primero tiene peta¬
los, flares unisexuales y bayas, mientras que cl se-
guntlo y tercero tienen fruto capsular, aunque Ma¬
crohasseltia presenta (lores unisexuales y carece de
petalos, y Pleuranthodendron posee petalos. Es ob-
vio q uc no es posible ubicar con certeza al taxon
nueva, por sus flores bisexuales y su baya. en nin-
guno de estos generos.

Hasseltia Kunth y Pineda Ruiz tN I’avon tambien
pertenecen a la t ri bn Prockieae (sensu Chase et ah,
2002) como easi todos los generos mencionados, a
except ion tie Olmediella que esta ubieada en la
tribu Flacourtieae (sensu Chase et ah, 2002), por
ello es conveniente examinarlos. Hasseltia tiene
ovario 2(-3)-locular, petalos y baya, y Pineda pre-
senta ovario unilocular, petalos y capsula, carae-
tenslieas que impiden la insert ion del nuevo taxon
en alguno de el los (sobre todo en Pineda).

A la luz de la eomparacibn hecha hasta el mo¬
mento, la ubicacibn generica tiene varias posibili-
tlades, si se adopta una posit ion muy estricta sobre
los 1 unites morfolbgicos de los taxones, se justifica
la creacibn tie un genero nuevo, ya t|ue no puede
insertarse en ninguno de los descritos basta la fe-
cha, [)ues ninguno tiene flores apetalas, ovario uni¬
locular con placentas fuertemente intrusivas y
bayas simultaneamente; sin descartar esta posibi-
lidad, no existen en este momento suficientes es-
tutlios filogeneticos y tie desarrollo floral de los ge¬
neros de la familia para optar por esa solucibn. Si
se incluye en Prockia (como se hace en el presente
trabajo) es necesario hacer una modificacibn tie su
circunscripcibn, donde se admita la posibilidatl de
tener ovario unilocular con placentas intrusivas y
no solo ovarios 3—6 loculares. La ausencia de pe-
talos en Prockia krusei ya esta ineluida en su con-
cepcibn actual, pues Prockia crucis P Browne ex
L. presenta ocasionalmente flores apetalas. La ter-
cera alternativa es tratar de ubicar a la especie
nueva en los generos con ovarios uniloculares con
placentas fuertemente intrusivas, flores petalfferas
V bayas, como ocurre en Banara y Neosprucea o en
Hasseltia ya que comparte con estos generos los dos

No von 15: 297-300. 2005.

298

Novon

ultirnos caracteres, aunque posee ovario 2(— 3)-locu-
lar. Htinara y Neosprucea parecen ofrecer opciones
muy interesantes para insertar al nuevo taxon, pues
comparten el mismo tipo tie plaeentaeion y de fruto,
(‘1 ilnieo obstaeulo aparente es que siempre presen-
tan petalos, sin embargo, la balanza se inelina lige-
ramente a favor del genero Batumi, pues tiene fi-
lamentos alargados y anteras snbglobosas romo la
especie nueva (ver Fig. 1C y I)); en eambio. Neo-
sprucea presenta (ilamentos muy cortos y anteras
lineares, pero ambos generos earecen de sepalos
alados en el boldn, eomo oeurre en Prockia y P.
krusei, por lo que no fueron las opeiones elegidas.
Hnalmente, queda Hasseltia, pero puede ser faeil-
mente excluida por la preseneia de petalos, ovario
2(-3)-locular y porque tiene disco con glandulas.
mientras (|ue P. krusei, Banara y Neosprucea care-
cen de disco.

Prockia krusei J. Jimenez Ramfrez & R. Cruz, sp.
uov. TIPO: Mexico. Guerrero: Mpio. Chilpan-
eingo, Salto Valadez, 17°27'20"N, 99°28' 10"0,
1400 m, 2 ago. 1070 (fl), Hubert Kruse I9B5
|d| (Imlotipo. MEXU; isotipos, FCMK. MEXU,
MO). Figura I.

Differt haee species ah afftnibus floribus apetalis, 94—

I 12 staminibus, ovario unilooulari, 3 placentis lamellifor-
mibus, baeea 9 mm longa et lata atque seminibus 3.5—4'
X ea. 2.5 mm.

Arboles o arbustos hasta 4 m de alto; tallo
ligeramente estriado, cafe elaro, con abundantes
lenticelas; ramas jdvenes bianco pubescentes, con
abundantes lenticelas amarillentas. Ilojas ovado-
elfpticas; estfpulas lanceoladas a subuladas 2—2.5
mm, pubescentes en ambas earas, terminando en
una glandula, en oeasiones otra lateral cerea de la
base; peefolo (0.5— )0. 8— 1.8 cm, pubescente; lamina
(6— )6.5— 12 X (4— )4. 5— 8(8.5) cm; apice agudo; base
cordada a redondeada, con 5 nervios basales, 3^4
venas laterales; margen glandular-erenado, una
vena terminando en una glandula circular en el
apice de cada diente; dientes 13— 29(— 31), 4—7 mm
de largo; haz pubescente, con apariencia rugosa de-
bida a la venacidn; enves visiblemente reticulado,
pubescente, trieomas rectos, pubescencia mas
abundante en la venacidn. Inflorescencias cimosas
(un dieasio terminal y una flor lateral; la antesis
inicia en la flor terminal y la lateral basal), termina-
les o axilares (aparecen eomo axilares cuando (4
fruto ha madurado, y una vez que el apical se ha
renovado), (3.3-)4— 4.5 cm de largo, eon (l-)3— 4
(-5) (lores, eje de la inflorescencia 1.2-1. 6 cm de
largo, pubescente; peduneulos 1-3 mm de largo.
Flores amarillas-verdes-rojas, apdtalas; pedicelos

1—1.6 cm de largo, ligeramente pubescentes, con
una bractea basal de 3 mm de largo, laneeolada,
pubescente en ambas earas; caliz con 3(— 4) sdpalos
reduplicado-valvados (fuertemente alados en el bo-
tdn), erectos, (8— )9— 10 X 8—10 mm, ovados, pu¬
bescentes en ambas earas, connados hasta 5 nun.
ligeramente lobulados en la base (en oeasiones
cada lobulo con una glandula), dpiee agudo. con
una glandula; estambres 94—1 12. 3 mm de largo,
nunea exsertos; anteras oblongo-elipsoides; pislilo
7 mm de largo; ovario esferieo, 4 mm didmetro,
unilocular, glabro; plaeentaeion parietal, 3 placen¬
tas lameliformes; estilo 3 mm de largo, recto, gla¬
bro; estigma brevemente trilobado. Fruto una baya
globosa, 9 X 9 mm, con el caliz persistente de 1.5—
1.8 X 1.2— 1.8 cm (se desarrollan 3—4 frutos por
inflorescencia); semillas ovoides ca. 3.5—4 X ca.
2.5 mm, (1— )4— 8 fertiles y generalmente 4 aborta-
das.

Habitat y distribucidn. Se encuentra en bosque
tropical eaducifolio, en altitudes de 1400-1420 m,
asoeiada a Casearia corymbosa Kunth, Nopalea sp.,
Opuntia sp.. Agave sp.. Ipomoea sp., Bursera bi-
pirmata (DC.) Engler, Bursera glabrifolia (Kunth)
Engler, Eysenhardtia sp. y Tiliaceae, en roeas cali-
zas. Hasta el momento solo se ha eolectado en el
paraje conocido eomo Salto Valadez en el estado de
Guerrero y la poblacidn es sumamente eseasa.

Fenologta. Florece de julio a agosto y fructifica
en septiembre.

hpiteto especifico. El nombre de la especie hon-
ra al Ing. Hubert Kruse, importante coleetor para
el estado de Guerrero durante los anos 1959 a
1968(— 1973), en su coleecidn. con cerca de 3500
ejenqilares botanieos. destaean numerosas es|ieeies
nuevas y endemicas para el estado de Guerrero,
Mexico.

A Contini aciOn Sk Pkksknta una Clavk (Modikicada
en Parte he Sleumer, 1980) para Distinguir a la Es¬
pecie Nl E\ \ DEI. Re.STO l)E LAS EsPECIKS DEL GENERO

Prockia

la. Estfpulas fuertemente inequildteras o semicor-
dadas o semireniformes, 5 — 10(— 1 7) mm tie largo.

2a. Estfpulas fuertemente inequildteras o semi¬
reniformes; sepalos eon margen entero o easi
enlero, 4—7 mm de largo.

3a. Hojas hasta 2 veces mds largas que an-
( has o menos. Se distribuye de Mexico

hasta Sudamdriea .

. P. crucis P. Browne ex b.

3h. Hojas generalmente 2.5 mds largas que
anchas o mds. Conocida de Costa Riea

. P. costaricencis Standley

2h. Estfpulas eon hase semieordada; sepalos eon
<4 margen serrado-glandular (al menos en la

Volume 15, Number 2
2005

Jimenez & Cruz
Prockia krusei de Mexico

299

Figure 1. Prockia krusei J. Jimenez Ramirez & R. Cruz. — A. Rama con (lores. — B. Rama con frulos. — C. Nor con
cuatro sepalos, uno removido. — I). Flor con ties sepalos — K. S^palo. - — F. Margen y enves de la lioja. — G. Placen-
tacidu. — H. Seniillas, vista lateral y ventral. — I. Kstfpula. (A, C, I), K, Gel, Kruse 1985 [d]; B, Kruse 1985 [a]; F y
H, Kruse 1985 [c].) Ilustrado por Ramiro Cruz Duran.

300

Novon

base), 7—10 mm de largo. Conocida de Ve¬
nezuela . P. flava Karsten

lb. Estfpulas con la base sim£trica, lineares, laneeo-
ladas a subuladas, 1—2.5 mm de largo,
la. Estfpulas lineares, I mm de largo o menus;
(lores con 5 sepalos y 5 petalos. Sc distri-

buye en Ecuador y Peru .

. P. pentamera A. Gentry

4b. Estfpulas laneeoladas a subuladas, 2—2.5
mm de largo; (lores con 3( — 4) sepalos. ape-
talas. Distribucidn restringida a Guerrero,

Mexico .

.... P. krusei .1. Jimenez Hamfrez & R. Cruz,

sp. nov.

Prockia krusei se puede diferenciar lacilmente
de todas las especies del genero, por tener ovario
uniloeular, eon Ires placentas intrusivas, en lugar
lie tener ovario triloenlar y plaeenlaeidn axilar. A
priniera vista, las especies eon sepalos grandes
eomo P. pentamera (10 X 3—8 nun) y P. flava (7—
10 X 5—8 mm) se pareeen a la especie nueva (8—
10 X 8—10 mm), sin embargo, es sencillo distin-
guirla de la priniera porque esta tiene 5 scpalos y
5 petalos, mientras que P. krusei tiene 3-4 sepalos
y eareee de petalos. I.a separaeidn de la segunda
se debe a que esta especie posee 50—70 estambres,
baya de 1—8 mm de diametro y semillas de 1 mm
de longitud, en eontraste. la especie nueva tiene
94-1 12 estambres, baya de 0 mm de diametro y
semillas de 3.5-4 mm de longitud.

Prockia costaricencis (una de las dos especies
con sepalos de basta 7 mm de largo) y la especie
nueva pueden separarse porque P. krusei presenta
estfpulas de laneeoladas a subuladas. de 2-2.5 mm
de largo y fruto de 9 mm de largo, frente a estfpulas
semireniformes. de 5—9 mm de largo y los frutos de
7 mm de largo de P. costaricensis.

Prockia cruris (la segunda especie con sepalos
de 7 mm de longitud o menus) se puede distinguir
de P. krusei por el tamano de las estfpulas (5—8 mm
de largo en la priniera y 2—2.5 mm de largo en la
segunda especie), y por el tamano del fruto y las
semillas (baya de 6—8 mm de diametro y semilla
de 1 — 1.5 mm de largo en P. cruris y baya de 9 mm
de diametro y semilla de 3.5—4 mm de largo en P.
krusei).

Paratipos. MEXICO. Guerrero: Mpio. Cliilpancingo,
Salto Valadez, Hubert Kruse 1 985 |a| (KCME, MEXU),
Hubert Kruse 1985 |l>| (I'CME. MEXU), Hubert Kruse 1985
[c| (I’CME, MEXU), R. Cruz Durdn 5974 (ECME).

Agradecimientos. A1 Dr. Mario Sousa del Her¬
baria Nat ional (MEXU) por las facilidades otorga-
das para la consulta del material de la coleceibn
KRUSE, a la P. de Biol. Gloria Andrade por los
date is biograficos referentes al Ing. Hubert Kruse, a
la M. en (7 Martha Martinez Gordillo la diagnosis
en lathi y la revision erftica al manuserito, al M.
en C. Carlos Ruiz Jimenez el apoyo tecnico en el
manejo de la Figura 1, asf eomo al Biol. Jose An¬
tonio Hernandez Gomez de Mierocine, Faeultad de
Ciencias, UNAM, la digitalizacidn de la misma. En
particular, a la l)ra. Sue Zmartzy por la cuidadosa
revision y atinadas sugerencias al manuserito y a
los revisores anonimos sus valiosas observaciones.

bitcratura Citada

Chase, M.. S. Zmartzy, M. Lledd, J. Wurdack, S. Swensen
& M. Eay. 2002. When in doubt, put it in Elacourtia-
ceae: A molecular phylogenetic analysis based on plas-
lid rbe\. I)NA sequences. Kew Bull. 57: 141—181.
Sleumer, II. 1980. Elacourtiaceae. Pp. 1-499 en C. T. Ro-
gerson (editor), Elora Neotrop. Monogr. 22. The New
York Botanical Garden, Bronx, New York.

A New Species of Gymnosporia (Celastraceae) from
Southern Africa

Marie Jordaan

South African National Biodiversity Institute, Private Bag X101, Pretoria, 0001, and
Department of Botany, University of Pretoria, Pretoria, 0002, Republic of South Africa.

jordaan@sanbi.org

Abraham E. van Wyk

H.G.W.J. Schweickerdt Herbarium, Department of Botany, University of Pretoria, Pretoria,
0002, Republic of South Africa. braam.vanwyk@up.ac.za

Abstract. Gymnosporia gariepensis Jordaan (Ce¬
lastraceae), here described as a new species, is
known only from the Gariep Center, an arid region
of high iloristie endemism in the northwestern cor¬
ner of the Northern Cape (South Africa) and adja¬
cent parts of southern Namibia. An illustration and
a distribution map are provided. Diagnostic char¬
acters include: the robust thorns that terminate the
short, abbreviated lateral branches on older shoots,
and the capsules that are reddish, smooth, and ob-
conic-trigonous. The new species is related to G.
tenuispina (Sonder) Szyszylowiez, a species with
slender thorns with which it shares arils that are
reduced to a fleshy rim at the base of the seeds.

Key words: Celastraceae. Celastroideae, Gariep
Center of Endemism, Gymnosporia, southern Afri¬
ca. taxonomy.

Gymnosporia (Wight & Arnott) Hooker I.. an Old
World woody genus of about 108 species and sub¬
species in subfamily Celastroideae of the Celastra¬
ceae, has recently been reinstated (Jordaan. 1995;
Jordaan & Van Wyk, 1999; Archer X Jordaan,
2000). Celastroideae are characterized by dry de¬
hiscent capsules and arillate seeds. Included in
Gymnosporia are all the thorn-bearing members
previously classified under Maytenus Molina s.l.
(Marais, I960: Robson, 1965, 1966, 1994).

During a study of Gymnosporia. a number of un¬
described species have been discovered. The pur¬
pose of this paper is to describe one of the new
species; it has a restricted geographical distribution
centered in the Richtersveld. an arid mountainous
region in the northwestern corner of the Northern
Cape, South Africa, as well as in the adjacent
southernmost part of Namibia.

'l'he new species shows affinity with Gymnosporia
tenuispina (Sonder) Szyszylowiez, a species with

slender thorns of the summer-rainfall savanna re¬
gion m northeastern South Africa with which it
shares arils that are reduced to a fleshy rim at the
base of the seeds. A similar reduced aril is also
encountered among species of Gymnosporia in
northeastern tropical Africa, a region considered,
in addition to southern Africa, as one of the centers
of diversity for the genus (Sebsebe. 1985). Hol istic
similarities between the arid parts of southern Af¬
rica and the Horn ol Africa are well established
and strongly support the proposed former existence
of periodic biogeographical connections between
these two regions (Van Wyk X Smith, 2001).

Gymnosporia gariepensis Jordaan. sp. nov.
TYPE: South Africa. Northern Cape: Richters¬
veld Nat. Park. Die Toon, E of Tatasberg, 1
July 1995. ./. E. Rourke 2085 (holotype, NBC;
isotypes, MO. PRE). Figure 1.

Gymnosporiae tenuispinae similis, sed omnino glabro;
spinis semper ramos secundarios terminantibus; ramulis
juvenibus rubrobrunneis; lamina coriaeea, margine sem¬
per integra; tloribus fructibusque in pedieelfis longis pen-
dulis, semper in brachyblastis in ramulis lateralibus se-
cundariis; floribus majoribus, capsulis rubris, viride
suffusis, dilfert.

Lax, few- to multistemmed shrubs, up to 2 m
high, thorny, glabrous, dioecious; branches terete;
young branchlets smooth, purplish red to red-
brown, becoming gray with small, round slits in
bark, each containing a sunken lenticel, flaking in
uneven white pieces; thorns slender and axillary on
young shoots, becoming robust and terminating the
abbreviated lateral branches on older shoots, up to
15 mm long. Leaves fasciculate in axils ol braehy-
blasts on abbreviated lateral branches, alternate on
young branchlets, coriaceous, pale green, subses-
sile, glabrous; lamina oblong or obovate-oblong,
12—45 X 2—5 mm. apex rounded or subemarginate,

Novon 15: 301-304. 2005.

302

Novon

Figure I. Gymrwsporia gariepensis Jordaan. — A. Flowering branch. — 15. Young branchlet. — C. Fruiting branch.

I). (.apsule. F. Seed with aril reduced to a rim at base. — I*. Female flower. A and F drawn from the holotype, Rourke
20R5 (INH(»); R from Drijfhout 2969 (PRE); C, I), and E from Jurgens 28856 (PRE).

base cuneate, margin entire, venation obscure (at
least in dried material); petiole very short. ± 0.5
mm long; stipules very short, subulate, ± 0.5 mm
long. Inflorescence cymose. axillary or on short ab¬
breviated lateral branchlets; peduncle 1—5 mm

long; pedicels 5—13 mnt long. Flowers functionally
unisexual, pentamerous, 7-10 mm diam.; sep<ils tri¬
angular-ovate, 1—2 mm long, apex acute, margin
ciliolate; petals oblong, yellowish green, 2.5— 4.0
mm long, margin subentire; disc flat, entire or shal-

Volume 15, Number 2
2005

Jordaan & van Wyk 303

Gymnosporia gariepensis from Southern Africa

Figure 2. Known geographical distribution of Gymnosporia gariepensis (•).

lowly 5-lobed: male flowers: stamens shorter than
petals; filaments ± 2 mm long, arising at lower out¬
side edge of disc; pistillode small; style short, un-
branched: female flowers: staminodes slightly short¬
er than stamens in male flowers; ovary 3-locular,
subglobose; style very short, terete; stigma 3-
b ranched. Capsules 3-valved, obconic-trigonous,
smooth, chartaceous, pendulous, 9-12 mm long,
upper part red. lower part green; seeds reddish
brown, rugose, ± 5 mm long; aril yellow, reduced
to a fleshy rim at base of seed.

This species is a laxly branched shrub with pur¬
plish red to red-brown stems, becoming gray with
age. On older branches the robust thorns terminate
short, abbreviated lateral branches. The leaves are
coriaceous, long, and narrow, and the venation is
obscure. Inflorescences and capsules are borne
mainly on short abbreviated lateral branchlets and
are rarely axillary. The inflorescences are few-flow¬
ered, or flowers are sometimes solitary. I he cap¬
sules are pendulous, obconic-trigonous, red, and
smooth, with the veins not visible. Ibis new species
is allied to Gymnosporia tenuispina, which has slen¬
der thorns, usually in the axils of the leaves or
brachyblasts, and small, yellow', smooth, somewhat

inflated and obconic-trigonous capsules. In G. ten¬
uispina, the leaves are chartaceous, tin* venation is
visible, and the margins are shallowly glandular
denticulate to subentire, whereas in G. gariepensis
the leaves are leathery, the venation is inconspic¬
uous. and the margins are entire or with only a lew
remnants of teeth.

Distribution and habitat. Plants usually grow
among large rocks in sparse, semidesert vegetation
in the extremely arid northern Richtersveld and ad¬
jacent southernmost part of Namibia, where it is
currently known from only a few populations (Fig.
2). The geographical distribution ol Gymnosporia
gariepensis and its closest ally is markedly disjunct.
Gymnosporia tenuispina occurs in eastern Botswa¬
na. southern Zimbabwe, and in South Africa in the
Limpopo Province, North-West, Gauteng, Mpuma¬
langa, the northern Free State, and northern
KwaZulu-Natal.

Etymology. The specific epithet refers to the
Orange River, which cuts through the distribution
area of the species — Gariep is the Khoekhoe name
of the Orange River and means “Great River.”
Known as the Gariep Center of Endemism, this re¬
gion along the lower reaches ol the Orange River

304

Novon

is well known lor its high levels of florislic ende¬
mism and forms part of the larger Succulent Karoo
Region (Van Wyk & Smith. 2001). Within the range
of tin- new species the mainly winter rainfall is usu¬
ally less than 50 mm per annum, and extremely hot
temperatures (> 40°C) are frequently experienced
in summer (Hi Iton-Taylor, 1996; Van Wyk & Smith.
2001).

Conservation status. As most individuals of
Gymnosporia gariepensis are known from the Rieh-
tersveld National Park, the species enjoys at least
some protection. Unfortunately, domestic livestock
is allowed inside the park and the resultant over¬
grazing by goats and sheep is causing serious dam¬
age to the vegetation.

Haratypes. NAMIBIA. Mara, Craven 2644 (WIND);
Knersberg, I km N\Y of Sambokrivier, 20 Apr. 1988. I an
Jaarsveld & Leitch 6726 (NBG, PRE). SOUTH AEBICA.
Northern Cape; Gannakouriep, Hardy 261)2 (PRE);
Richtersveld. Die loon, E of Tatasberg, 17 Sep. 1989.
Jurgens 28856 (PRE), Eeb. 1995, G. & F. Williamson
5576 (NBG); Richtersveld, S slopes of Rosynt jieherge, I I
June 1980. Drijjhout 26 66 (NBG. PRE).

Acknowledgments. We are most grateful to the
curators of NRG and WIND lor the loan of speci¬
mens; John Rourke for collecting the type material;
Marietjie Steyn for doing the line drawings; Hugh
(den lor providing the Latin diagnosis; Kmsie du
Pit \ssis for critically reading and improving the
manuscript; and Hester Steyn for preparing the
map.

I .iterature Cited

Archer, R. H. & M. Jordaan. 2000. Celastraeeae. Pp. 2 14 —
220 in 0. A. Eeistner (editor). Seed Plants of Southern
Africa: Families and Genera. Strelitzia 10. National Bo¬
tanical Institute, Pretoria.

Hilton- Taylor, C. 1996. Patterns and characteristics of the
flora of the Succulent Karoo Biome, southern Africa. Pp.
58—72 in L. .1. G. van der Maesen ct al. (editors). The
Biodiversity of African Plants. Kluwer Academic Pub¬
lishers, Dordrecht.

Jordaan, M. 1995. A I axonomic Revision of the Spiny
Members of Subfamily Celastroideae (Celastraeeae) in
Southern Africa. M.Se. Thesis, University of Pretoria,
Pretoria. [Unpublished. |

& A. E. Van Wyk. 1999. Systematic studies in
subfamily Celastroideae (Celastraeeae) in southern Af¬
rica: Reinstatement of the genus Gymnosporia. S. Afri¬
can J. Bot. 65: 177-181.

Marais, W. 1960. An enumeration of the Maytenus species
of southern Africa. Bothalia 7: 381-386.

Robson. N. K. B. 1965. New and little known species from
the Flora Zambesiaea area XVI. Bob Soe. Brot., s£r. 2,
39: 6-2.5.

- . 1966. Celastraeeae. Pp. 355—418 in A. W. Exell.

A. Fernandes & H. W ild (editors). Flora Zambesiaea 2.
Crown Agents for Oversea Government and Administra¬
tions, London.

- . 1994. Celastraeeae. Maytenus. Pp. 1-21 in R. M.

Polhill (editor), Flora of Tropical East Africa, Celastra¬
eeae. A. A. Balkema, Rotterdam.

Sebsebe Demissew. 1985. The genus Maytenus (Celastra¬
eeae) in NE tropical Africa and tropical Arabia. Symb.
Bot. Upsal. 25: 1-101.

Van Wyk, A. E. & G. E Smith. 2001. Regions al Floristic
Endemism in Southern Africa: A Review with Emphasis
on Succulents. Umdaus Press, Pretoria.

Dupuya , a New Genus of Malagasy Legumes (Fabaeeae)

Joseph H. Kirkbride Jr.

USDA, Agricultural Research Service, Systematic Botany and Mycology Laboratory, Bldg.
01 1 A, Rm. 304. BARC-West, Beltsville, Maryland 20705-2350, U.S.A.joe@nt.ars-grin.gov

ABSTRACT. The new endemic Malagasy genus
Dupuya is [imposed based on morphological char¬
acters. It has two species, D. haraka, comb, nov.,
and D. rriadagascariensis, comb, nov., and is distin¬
guished from related genera in having no petals,
numerous stamens, a whorl of staminodes inside
the stamens, and a number of distinctive seed char¬
acters. A new combination is also made for Dupuya
madagascariensis subsp. tamarindoides. Dupuya is
a member of the aldinoid elade with the genera
Aldina , Amburana , Cordyla, Dussia, Mildbraedio-
dendron , and Myrospermum.

key words: aldinoid clade, Cordyla, Dupuya.
Dupuya haraka. Dupuya rriadagascariensis, Faba-
ceae, heguminosae, Madagascar.

In 2002. the outstanding Leguminosae of Mad¬
agascar was published (Du Buy et al.. 2002). 1 his
comprehensive treatment fills a critical lacuna in
our knowledge of the world’s legumes; 34% of the
100 native Malagasy genera and 80% of the 573
native species are endemic. All taxa were described
in detail, and the fruit and seed descriptions are
some of the most complete descriptions found in
any Holistic work. 1 compared all their descriptions
to those in our database of legume genera (kirk¬
bride et ah, 2003). For all the Malagasy genera,
only minor revisions were needed to encompass the
variations found in Madagascar, with one notable
exception, Cordyla Loureiro, for which our seed
data were completely at odds with Du Puy and ha-
bat's (2002) descriptions and illustrations of the ge¬
nus and its two constituent species in Madagascar.
|)u Puv and Labat (2002: 294) had noted, “the Mal¬
agasy species are distinctive as the inner stamens
are sterile.” Kxamination of the USDA, ARS, U.S.
National Seed Herbarium (BARG) material re¬
vealed that our seed data were based only on Af¬
rican specimens of Cordyla, and corresponded with
that of Corner (1976) for C. africana Loureiro. After
examining specimens of all five African Cordyla
species (see Appendix 1), as w'ell as the two in
Madagascar, it was apparent that Du Puy and habat
(2002) had correctly described the latter and that
our data for the African members were also correct.
Based on the presence of staminodes in the Mala¬

gasy species and their absence in the African spe¬
cies. along with the many differences in the seeds
of the two groups (see the key to genera below), the
two Malagasy species are here described as Dupuya
J. H. Kirkbride, a new genus endemic to Madagas¬
car.

Systematic History ok Cordyla s.i..

Loureiro (1790) described the genus Cordyla
with a single species, C. africana. Milne-Redhead
(1937) speculated that Loureiro collected the type
specimen in Mozambique because the common
name that he used for it, “mutonda.” was still in
use for it there. Milne-Redhead (1937) revised the
four African species known to him. and later Gillett
(I960) added a fifth. C. sornalensis J. B. Gillett.
V iguier ( 1948) described the first Malagasy species,
C. madagascariensis R. Viguier. and Capuron
(1968) published the second, C. haraka Capuron,
in his revision of Malagasy Cordyla. Following Ca¬
puron. all other Cordyla treatments have been Ho¬
listic in nature (Aubreville, 1970; Berhaut, 1975;
Brenan, 1967; Brunei et al., 1984; Cowan, 1981;
Du Puy & Labat, 2002; Gilbert & Boutique. 1952;
Gillett. I960. 1962; Hedburg & Kdwards, 1989;
Hutchinson, 1964; Keay, 1958, 1989; Keay et al.,
1964; Lebrun & Stork, 1992: Ross, 1977; Timlin.
1993) without attempting to compare or re-evaluate
its generic delimitation based on an analysis of all
its members.

Herendeen (1995) carried out extensive phylo¬
genetic analyses of the 1 I genera in tribe Swart-
zieae sensu Cowan (1981). He used 29 morpholog¬
ical characters, including 3 from seeds and I from
l he fruit. In addition, 19 Sophoreae and 3 Caesal-
pinieae genera were scored. The Sophoreae genera
were included because Herendeen (1995: 123)
considered them to be “the likely sister group to
Swartzieae” and some genera, such as Ateleia (DC.)
Bentham, Cyathostegia (Bentham) Schery, and Hol-
ocalyx Micheli, were difficult to place in the correct
tribe. The Caesalpinieae genera served as out¬
groups. The Swartzieae genera fell into three dis¬
tinct clades in association with various genera of
Sophoreae and Orphanodendron Barneby & J. W.

Novon 15: 305-314. 2005.

306

Novon

Grimes. Swartzieae is obviously a polyphyletic
group, and Herendeen (1995: 130) proposed that
“the concept o( Swartzieae . . . should be aban¬
doned and all members of Swartzieae should be
referred to Sophoreae. ""

In Herendeen’s analyses, Cordyla formed a clade
with Mildbraediodendron Harms, well supported by
three nonhomoplasious forward changes: absence of
a corolla, stamens united basally, and fruit globose
to elongate with a fleshy endocarp and two homo-
plasious forward changes: punctate leaflets and ab-
axial stomata with a variable number of subsidiary
cells, more than two. Our seed observations (kirk-
bride et al., 2003) strongly support this relationship
(figs. 1—3). Cordyla and Mildbraediodendron seeds
are large, lack a testa, have straight embryos par¬
alleling the seed length, the' radicle tip straight, and
the* plumule well developed, and are centered rel¬
ative to the cotyleelons.

Pennington et al. (2001) studied the evolution of
the basal faboid legumes to clarify the broad-scale
relationships within subfamily Faboideae. They
carried out phylogenetic analyses using trn\. intron
sequences for 122 taxa, representing 14 of 15
Swartzieae sensu Polhill (1994a, 1994b), 26 of 41
Sophoreae, the 20 Dipterygeae and Dalbergieae,
and various placeholders for derived groups; 4 Cae-
salpinioideae genera were used as outgroups. Using
both the trnL intron and spacer. Ireland et al.
(2000) conducted a phylogenetic analysis focusing
on tribe Swartzieae. They sequenced 14 of 15
Swartzieae genera, tin* genus Robgunnia .1. H. kirk-
bride & Wiersema, 19 Sophoreae genera, and 20
Dipterygeae and Dalbergieae, using 3 Caesalpi-
nioideae genera as outgroups. Neither study in¬
cluded the Swartzieae genus Candolleodendron R.
S. Cowan. Roth studies used sequences from Cor¬
dyla madagascariensis, which is lien* treated as
Dupuya madagascariensis (R. Viguier) J. 11. kirk-
bride, and no African Cordyla s. str. was included
in either study. Their results were similar, differing
mainly in scope. Roth Swartzieae and Sophoreae
are polyphyletic groups whose genera belong in
various other tribes. I he sister group to the re¬
mainder of the faboid legumes is the swartzioid
clade (Pennington et al., 2001), consisting of Ro¬
bgunnia, Swartzia Schreber, Rocoa Aublet, Cy-
athostegia, and Ateleia, which might be recognized
as either a redefined tribe Swartzieae or a fourth
legume subfamily. Also in both studies, the aldi-
noid clade (Pennington el al.. 2001) consisted of
the genera Myrospermum Jacquin, Russia kmg &
Urban ex Taubert. Amburana Schwacke & Taubert,
Mildbraediodendron, Dupuya, and Aldina Endlieh-

er. Neither of these two chides had been previously
recognized in any other study.

I he swartzioid and aldinoid clades (except Al¬
dina) shari1 absence of the 50-kilobase inversion in
the chloroplast genome (Doyle et al.. 1996; Ireland
et al., 2000), which is also absent from subfamilies
Mimosoideae and Caesalpinioideae and present in
most of subfamily Faboideae. Because Aldina has
the CP inversion and the remainder of the aldinoid
genera do not, its relationships should be re-ex¬
amined (Ireland cl al., 2000). and more species be¬
longing to the aldinoid clade should be screened
for the CP inversion confirming its presence in oth¬
er genera of this clade, clarifying the circumscrip¬
tion of the clade, and validating the membership of
Dupuya in it.

Aside from the presence of staminodes and the
differences in seed characteristics, Cordyla and
Dupuya are so similar morphologically that they
probably are sister genera (sensu Wiley, 1981), and
therefore Cordyla s. str. also is probably a member
of the aldinoid clade. The morphological differenc¬
es separating genera of the aldinoid clade are pre¬
sented in the following key.

ki;v to Genkra oi the Aldinoid Clade

I. Flowers with I or (3 to)4 to 7 petals; fruits de-
liisoenl or indehiseent (Myrospermum): Neo¬
tropics . 2

1 . f lowers without petals; fruits indehiseent; trop¬
ical Africa and Madagascar . 5

2(1). flowers with (3 to)4 to 7 petals; cotyledons with
surface grooved, ridged, or rugose, unlobed;
embryonic axis oblique or perpendicular to

seed length . 3

2'. Flowers with 1 petal (the standard); cotyledons
with surface smooth, lobed; embryonic axis par¬
allel to seed length; southeastern Peru to north¬
eastern Mrazil and south to northern Argentina

. Amburana

3(2). Leaflets pellucid-glandular; fruit winged, inde-
hiscent; epiearp veined; seed 1(>—I7 X 4-7.2
X 4.5— 5.5 mm; Mexico, Central America, and

northern South America . Myrospermum

3'. Leaflets not glandular; fruit not winged, dehis¬
cent; epiearp not veined; seed 20—60 X 10—50

X 10-40 mm . 4

4(3). Corolla papilionaceous; fruit elliptic or fusi¬
form, slipitate, with values enrolling; seed 20-
45 X 10-20 X 10-20 mm. not overgrown; testa
present; radicle bulbose, straight at tip; Antil¬
les, Mexico, Central America, northwestern
South America, and the western Amazon basin

. Russia

4'. Corolla not papilionaceous, with petals ±
equal; fruit obovate, not stipitate, with values
revolute; seed 35-60 X 30-50 X 22—10 mm,
overgrown (1 seed completely filling the fruit
cavity); testa absent; radicle triangular, curved
at tip; Venezuela and the upper Amazon basin
. Aldina

Volume 15, Number 2
2005

Kirkbride

New Malagasy Genus

307

Figure 1. Seeds of Aldina, Amburana, Mildbraediodendron, Cordyla, and Dupuya. — A. Seed of Aldina latifolia
Bentham (Ducke 24049. BARC). — B. Seeds of Amburana eearensis (Allemao) A. C. Smith (PI 109210. BARG). — C.
Seed with endoearp of M. excelsum Harms (BARG). — D. Seeds with and without endocarp of C. africana Goureiro
(List #95, Shipment #41, BARC). — E. Seeds with testa of / ). madagascariensis R. Viguier suhs(). madagascariensis
(l)u Puy. Camlet & Labat 2463. P).

5(1). Flower hods subglohose, calyx splitting into (2
to)3 unequal lobes down as far as the disc; sta¬
mens 12 to 18, basal ly united into a single
whorl at the edge of the disc; tropical Africa

. Mildbraediodendron

5'. Flower buds turbinate, calyx splitting into 2 to
4 unequal to subequal lobes down as far as the
apex of the hypanthium; stamens numerous (20
to 130), basally united into approximately 3 or

4 whorls at apex of hypanthium . 6

6(5). Staminodes in a single whorl inside stamens,
y6 to 14 as long as stamens; testa present, en¬
dosperm present, cotyledons folded, partially
concealing radicle, one inner cotyledon face
with a central ridge and the other with a central
groove, embryonic axis oblique to the seed and
cotyledon lengths, radicle bulbose; Madagascar

. Dupuya

6'. Staminodes absent; testa absent, endosperm ab¬
sent, cotyledons not folded, fully concealing the
radicle, inner cotyledon faces flat, embryonic
axis parallel to the seed and cotyledon lengths,
radicle triangular; tropical Africa . Cordyla

Dupuya .1. II. Kirkbride, gen. nov. TV PE: Dupuya
madagascariensis (K. Viguier) J. II. Kirkbride.
Figures I E, 2E, and 3E.

A Cordyla el Mildbraediodendro staminibus mollis,
staminodiorum verticillo intra stamina, staminodiis stam¬
inibus circa quater brevioribus, seminibus brevioribus, ar¬
il lo, testa, endospermioque praesentibus, cotyledonibus

ambabus plicatis, cotyledonis unius superficie interiore
crista et cotyledonis alterius superficie interiore sulco, em-
bryonis axe obliquo ad seminis et cotyledonum axem, rad-
it ula bulbosa et aliquantum occulta cotyledonibus differt.

Trees, 5-35 in tall, deciduous, the branches with
large, corky, cream-colored lenticels, bilobed with
the lobes elliptic, parallel to the axis ol the branch,
with a sulcus between them; stipules small, cadu¬
cous. Leaves alternate, imparipinnate, with alter¬
nate or subopposite leaflets, without slipels, the ra-
chis with 2 narrow adaxial ridges, the closest one
of the pair interrupted at each leaflet insertion and
the other entire, sometimes extended into stipel-
like points, the leaflets emarginate, with glandular
slreaks oblique to the midrib, sometimes just glan¬
dular dots, smaller at the base of the leaf and be¬
coming gradually larger toward the apex of the leaf,
“light green above and beneath (drying much dark¬
er above)” (Du Puy & Labat, 2002: 294). Inflores¬
cences terminal or rarely axillary, racemose, some¬
times flowers solitary in the upper leaf axils, with
a caducous bract subtending each pedicel and a
pair of bracteoles (sometimes caducous) on ihe ped¬
icel at ca. the middle or above it. Flowers with a
conspicuous hypanthium, the calyx entire, with a
tuft of hairs at the apex, splitting into 2 to 4 un¬
equal to subequal lobes, apetalous, the stamens nu-

308

Novon

Figure 2. hmbryos and cotyledons of Aldina, Amburana , Mildbraediodendron, Cordyla, and Dupuya. — A. Aldina
lalifolia ( Cid Ferreira et id. 71 1H, BARC). — If. Amburana eearensis (Glaziou 20882, BARG). — C. M. excelsitm (BARG).

I). C. q/rieana (l.isl #9o, Sliipnient #11, BARC), — K. I). madugascariensis suhsp. madagascariemis ( R, Caouron
s.n., BARC).

melons, approximately arranged in three concentric
whorls at the hypanthinm apex, with a fourth, inner
whorl of staminodes, the filaments basally connate,
die anthers elliptic, dorsifixed, longitudinally de¬
hiscent, the staminodes filament-like, Vu to XA as
long as the stamens, basally adnate to the stamens,
the ovary stipitate, multiovulate, glabrous, exserted
above the anthers. Fruits unilocular, straight, ellip¬
tic to oblong, terete, with or without a straight beak
(5—8 mm long), the same color and texture as fruit,
with the apex and base tapered or rounded and
aligned with longitudinal axis of fruit; drupaceous
when Iresh (I)u Puy & Lahat, 2002) or ligneous
when dry, with the margin not constricted or slight¬
ly constricted along both margins (when 2 or more
seeds in fruit), stipitate, indehiscent; epicarp dull
or glossy, light to dark brown or reddish to purplish
brown, glabrous, longitudinally ribbed with 20 to
40 ribs; mesocarp thick, 2-layered, ligneous; en-
docarp dull, white when fresh (Du Puy & Lahat,
2002) or pale brown when dry, spongy when dry
and disintegrating at the touch, with seeds I to 3(4)
with their length parallel with fruit length; seeds
not overgrown (sensu Corner, 1951), elliptic to ob¬
long or slightly ovate, compressed, with surface
smooth, with a dry rim-aril, entire, brown; testa

completely adhering to endocarp, dull, with fre¬
quent streaks, reddish to dark brown or red (Du
Puy & Labat, 2002), with brown overlay, faintly
grooved or striate, coriaceous; raphe from liilum
through lens to base of seed and terminating, ligh¬
ter than testa, reddish brown, recessed; liilum vis¬
ible, with the lips of the faboid split the same color
as the rest of the liilum, elliptic, recessed, within a
halo lighter in color than testa; lens not discernible;
endosperm thick, covering at least Vi but not the
entire embryo, adnate to testa; cotyledons the same
thickness and ± of equal length, with I to 6 lon¬
gitudinal grooves on outer faces, with I cotyledon
partially folded with flaps unequal and not touching
and outer face flat, the other fully folded with flaps
equal and touching and outer face concave, par¬
tially concealing radicle, notched at radicle with
lobes touching (auriculate) forming a basal groin,
white, inner face ol fully folded cotyledon with cen¬
tral ridge and central groove on partially folded
one; embryonic axis straight, oblique to seed
length; radicle bulbose with lobe lip straight; plu¬
mule moderately developed.

The genus is named in honor of David J. Du Puy,
senior author of The Leguminosae of Madagascar

Volume 15, Number 2
2005

Kirkbride

New Malagasy Genus

309

Figure 3. Embryos, with cotyledons removed, of Aldina, Amburana, Mildbraediodendron, Cord via, and Dupuya. — A.
Aldina latifolia ( Cid Ferreira el al. 7118, BARC). — B. Amburana cearensis (Glaziou 20882, BARC). — C. M. excelsum
(BARG). — I). (.. africana (List #95, Shipment #41, BARC). — E. I). madaga.scarien.sis suhsp. madagascariensis (R.
Capuron s.n., BARC).

(Du Pn v et al., 2002). I)u Puy and Cabal's (2002)
treatment of tribe Swartzieae and the genus Cordyla
brought this new genus to my attention and sup¬
plied essential data enabling its publication.

Key to Dirrw Species in Madagascar

I. Stipules straight, obovate to narrowly so; leaves
with 6 to 13 leaflets, the petiolules 1—2.5 mm
long, the leaflets acute to obtuse at the base, I —

4.5 cm wide: inflorescence bracts elliptic, 2.5—3
X 1—2.7 mm; flower bracteoles elliptic to narrow¬
ly so, 3.5— 6 X 1.5— 2.5 mm; calyx 7— B mm long;
staminodes 3—3.5 mm long; ovary stipe 15—22

mm long; seeds 25 X 14—15 13—15 mm .

. I). haraka

I '. Stipules curved, falcate; leaves w ith 15 to 44 leaf¬
lets, the petiolules 0.5—1 mm long, the leaflets
oblique at die base, (0.3— )0.5— 1 cm wide; inflo¬
rescence bracts narrowly obovate, 1.5— 2.5 X
0.5— 0.7 mm; flower bracteoles narrowly obovate,

1—2 X 0.4— 0.6 mm; calyx 4.5—5 mm long; stam¬
inodes 1.5— 2.5 mm long; ovary stipe 9—13 mm
long; seeds 15—21 X 12—13 X 9—11 mm ....
. I), madagascariensis

1. Dupuya haraka (Capuron) J. H. Kirkbride,
comb. nov. Basionym: Cordyla haraka Capu¬
ron. Adansonia, ser. 2, 8: 220. 1968. TYPE:
E Madagascar. Massif de Earankaraina, a l'Est
d’Andranofotsy (Maroantsetra), 24 Nov. 1953,
Sen ice Forestier (R. Capuron) 86 58 (holotype,
I* not seen; isotypes, B not seen, BB not seen,
G not seen, K not seen. I, not seen. MO. P.
WAG not seen).

A large tree 10—35 m tall, I)BH 25—100 cm, the
"trunk tall, often unbranched except near the
crown, flaring basally into rounded buttresses, the
bark deep chestnut-brown, flaking off in large ir¬
regular pieces, some areas with large corky cream
lenticels" (Du Puy N Labat, 2002: 295). youngest
branches strigillose with older branches gradually
becoming glabrate to glabrous. Stipules straight,
obovate to narrowly so, acute at the apex, 1 .2— 4.5
X 0.4— 1.8 mm, strigillose to glabrate. Leaves 8—19
cm long, with 6 to 13 leaflets, the raehis 6—13 cm

310

Novon

long, strigillose to glabrate, the petiolules 1—2. .3
mm long, strigillose to glabrate, with the blade ob-
ovate to rarely broadly so or elliptic to rarely broad¬
ly so, acute to obtuse at the base, emarginate to
sometimes obtuse to acute at the apex, 1 .3— 8.5 X
1—4.5 cm, strigillose to glabrate with hairs concen¬
trated over the midrib and veins, “becoming deep
green and glossy above, pale green beneath" (Du
Puy cK Labat, 2002: 295), coriaceous to charta-
ceous. Hacemes terminal or rarely axillary, (1- to)9-
to 22-flowered, (3— )6— 14 cm long, the bracts ellip¬
tic. obtuse to subacute at the apex, 2.5—3 X 1-2.7
mm, strigillose, the pedicels 4.5—10 mm long, stri¬
gillose to glabrate, with the bracteoles elliptic to
narrowly so, acute to obtuse at the apex, 3.5—6 X
1.5— 2.5 mm. glabrate to sparsely strigillose, the hy-
panthium expanding upward, 2.5-4 mm long, 2-
3.5 mm diam. at the base and 3.2— 4.5 mm diam.
at the apex, strigillose to glabrate outside and gla¬
brous inside, the calyx 7-8 mm long, glabrous to
glabrate outside and sericeous inside, the filaments
unequal. 11-19 mm long, glabrous, the anthers
0.7—1 mm long, glabrous, the staminodes 3—3.5 mm
long, til** stipe 15—22 mm long, 0.7— 0.8 mm diam.,
glabrous, the ovary 4.5—6 X 1.5-2 mm, glabrous,
tin- style 3—5 mm long, glabrous. Fruits ellipsoid to
clavate. with a beak 2-4 mm long, 4—7 X 2—3.5
cm, glabrous, with the stipe 2—3 cm long. Seeds 25
X 14-15 X 13-15 mm.

Illustrations. Capuron (1968: fig. 5. p. 221); Du
Puy & Labat (2002: fig. 55, G-J, p. 296. pi. 18.
A).

Habitat and distribution. From Toamasina to
Vohemar, southeast of Antsiranana, and also on the
island of Nosy B6 and the Ankarana Massif in Mad¬
agascar. Found up to 700 m, in tall evergreen rain¬
forest and also in deciduous forest in the north (Du
Puy & Labat. 2002).

Common names. Haraka (Service Forestier (R.
Capuron) R65R. P); Harakafotsy (Service Forestier
1571 1. P): Vahonda (Du Puy & Labat. 2002).

Most individuals of this species arc found in the
eastern, evergreen, low-altitude forests, without a
distinct dry season and very high annual rainfall,
from Toamasina to Vohemar. At the northern end of
its range, the species stretches across Madagascar
at approximately 13°S latitude into the evergreen,
Sambirano, low-altitude forests, with a distinct dry
season of 4 to 6 months and high annual rainfall,
from the vicinity of Vohemar to the island of Nosy
lie. The morphology ol specimens from (lit1 eastern
forests differs slightly from those collected in the
Sambirano forests. The eastern-forest specimens
have, in general, slightly larger structures, for ex¬

ample, stems bigger in diameter, larger leaflets and
flowers, longer inflorescences, as well as more co¬
riaceous leaflets than the Sambirano-forest speci¬
mens. The Sambirano-forest specimens are inter¬
mediate in some characteristics between the
eastern-forest specimens and I). madagascariensis.

Specimens examined. MADAGASCAR. K Madagas¬
car. Masoala Peninsula, western side, NF of Antalavia, I
liours on foot along Antalavia River (ca. 4—5 km ?),
1 5°46'S, 50°2'F. 285 m. 16 Nov. 1989, I). ./. Du Puy, li.
P. I)u Puy, G. Rafamantanantsoa A' G. Sehatz Mi 75 (P);
Toamasina, Diego Suarez region, SF of town and SF of
Ambilobe, near Daraina on road to Vohemar (Iharana), I).
Meyer’s lemur study site. I f 70 S. 10 lo t,. 200 m. G.
McPherson 14755 (K, MO); District Fenerive. dans le foret
< It* Bemahogo, 2 km 500 tl Test du village de Andapibe
el d'Ambodranpana, 30 Mar. 1952, Rakoloniaina H (MO);
Toamasina, Masoala Peninsula, just F of Ambanizana in
hills above village. 15°38'S, 49°58'F, 10-200 in, 24 Nov.
1989, G. E. Sehatz & P. P. hurry II 2836 (MO); Farau-
karaina, it Test d’Andranofotsy (Maroantsetra), 24 Nov.
1953, Service Forestier (R. Capuron) 8658 (P); foret orien-
tale, foret d’Antandrokaolo, entre Morafeno, sur la Ran-
tabe et Amboditavolo, basin de la Fananehana, 400 m,
Feb. 1954, Service Forestier (R. Capuron) 9029 (k. P);
Ouest (Nord). Plateau de TAnkarana entre les P.k. 84-85
(entre Anivorano-Nord et Ambondromilehy), 14 Nov.
1958, Service Forestier ( R . Capuron) 20001 (MO. P); Ouest
(Nord), vestige de foret entre Belinta el Ambetratra, it
quelques kilometers au NW de Vohemer. 14 Oct. 1966,
Service Forestier (R. Capuron) 24853 (k, MO, P); Ouest
(Nord), vestige de foret entre Belinta et Ambetratra, au
NW de Vohemer. au Sud de Maintialaka, 10—18 Dee.
1966, Service Forestier (R. Capuron) 27337 (MO. P); foret
d'Amboangisay, Canton d’Ampahano-Antalaha, 20 June
1952, Service Forestier I33R.I40 (MO): Tampolo, l eneri-
ve. 13 Nov. 1954, Service Forestier 12470 (MO. P). 2 Mar.
1955, Service Forestier 13100 (BARC, P): Farankaraina.
Maroahitra, 5 Mar. 1956, Service Forestier 1571 1 (BARC,
k. P).

Dupiiyu madagascariensis (R. Viguier) J. II.
Kirkbride, comb. nov. Basionym: Cordyla
madagascariensis R. Viguier, Notul. Syst. (Par¬
is) 13: 355. 1949. TYPE: W Madagascar. Ba¬
sin du Fiherenana, 400 m, Oct. 1926. //. Per¬
rier de la Bdthie 16617 (Icetotypc. designated
by Du Puy & Labat (2002: 295). P not seen:
isotypes, K not seen, MO not seen, P). Figures
1 E, 2E, and 3E.

A tree 5-25 m tall. DBH 15—70 cm, “the bark
with shallow fissures forming a net pattern, even¬
tually flaking off in large pieces, pale grey-brown”
(Du Puy N Labat, 2002: 295), youngest branches
strigillose with older branches gradually becoming
glabrate to glabrous. Stipules curved, falcate, nar¬
rowly acute at the apex, 3—4 X 0.2— 0.7 mm, stri¬
gillose to densely so. Leaves (7—) 14— 22 cm long,
with 15 to 44 leaflets, the rachis (6-)12-18.5 cm
long, strigillose to glabrous, the petiolules 0.5—1

Volume 15, Number 2
2005

Kirkbride

New Malagasy Genus

311

mm long, strigillose to glabrate, willi l he blade nar¬
rowly obovate to narrowly elliptic, oblique at tbe
base, emarginated at tbe apex, with entire to cre-
nate margins, (0.6— ) 1.4—3. 6 X (0.3— )0.5-l cm, gla¬
brate to glabrous with hairs concentrated over the
midrib and veins, “pale green (often drying darker
above)” (Du Puy & Labat, 2002: 295), chartaceous.
Racemes terminal, rarely axillary, or sometimes a
solitary flower in tbe upper leaf axils, I- or (10-
to)12- to 38-flowered, 6—18 cm long, the bracts,
pedicels, anil bracteoles strigillose to glabrate, the
bracts narrowly obovate, narrowly acute at the apex,

1.5— 2.5 X 0.5— 0.7 mm, the pedicels 3—6 mm long,
with the bracteoles narrowly obovate, narrowly
acute at the apex, 1-2 X 0.4— 0.6 mm, the hypan-
thium expanding upward, 1.5—2 mm long, 1.1—2
mm diam. at the base and 1 .5-2.5 mm diam. at tbe
apex, glabrous to glabrate outside and glabrous in¬
side, the calyx 4.5—5 mm long, glabrous outside
anil sericeous to sparsely so inside, the stamens
glabrous; the filaments unequal, 9-15.5 mm long,
the anthers 0.5— 0.8 mm long, the staminodes 1.5—
2.5 mm long, the gynoeeium glabrous, the stipe 9—
13 mm long, 0.5— 0.7 mm diam., the ovary 3—4.5 X

1.5— 2 mm, the style 1.5— 4.5 mm long. Fruits ellip¬
soid to cylindrical to moniliform, with a beak 2-8
mm long, 3—6.5 X 2—3 cm, glabrous, with the stipe
2-3.5 cm long; seeds 15—21 X 12— 13 X 9-1 I mm.

Illustration. Du Puy & Labat (2002: pi. 18. 8).

Common names. Anakaraka ( XI. F. C. Ixmvel
160. P); Hazomena (Service Forest ier (R. Capuron )
3031. P); Karabo ( Service Forestier 2838, P); Lan-
drazo (Du Puy & Labat, 2002); Lazaza (Du Puy A
Labat, 2002); Madiroala (Du Puy et al. M75S, P);
Madivoala (Du Puy & Labat, 2002); Maimbohazo
(Du Puy & Labat, 2002); Sikilihazo (Du Puy & La¬
bat. 2002); Vaivay (Du Puy & Labat. 2002).

Key to Subspecies of Dupuya m. wa casc wkxsis in Mad¬
agascar

1. Stipules 3.5-4 X 0.2— 0.3 mm; leaflets 17 to 30(to 40);
staminodes ca. 1.5 mm long; fruit beak 5—8 mm long . .

. I). madagascariensis subsp. madagascariensis

I '. Stipules 3—3.5 X 0.5— 0.7 mm; leaflets (10 to)38
to 44; staminodes 2—2.5 mm long; fruit beak 2—

5 mm long .

. I). madagascariensis subsp. tamarindoides

There arc two subspecies recognized for Dupuya
madagascariensis in Madagascar, one of which, the
non-autonym, needs a new combination.

Dupuya madagascariensis subsp. madagasear-
iensis. Cordyla madagascariensis var. mada¬
gascariensis, Adansonia, ser. 2, 8: 220. 1968.
Cordyla madagascariensis subsp. madagascar¬

iensis, in Du Puy et ah, Leg. Madagascar 297.
2002. TY PE: W Madagascar. Basin du Fih-
erenana, 400 m, Oct. 1926. //. Perrier de la
Bdthie 16617 (lectotype, designated by Du
Puy & Labat (2002: 295), P not seen; isotypes,
K not seen, MO not seen. P). Figures 1 E, 2E,
and 3E.

Illustration. Du Puy & Labat (2002: fig. 55, A—
F, p. 296).

Specimens examined. MADAGASCAR. 40 km
d’Ankazoabo, route Sakaraha, Feb. 1970. ,/. Ilosser 19956
(P); A Madagascar, Province of Toliara (Tulear), INF of
Morondava, ca. 4 km INF of Andranomena, route to Mar-
ofandila, 20°9'S, 44°27'F, 30 m, 20 Oct. 1989, I). Du
Puy, R. P. I)n Puy A ,/. Andriantiana M361 (K. P): W
Madagascar, Toliara Province, Forest of Kirindy, ca. 60
km IN INF of Morondava (Swiss Forestry Concession, “Piste
Conoco"), 20°5'S, 44°37'F, ca. 30 m, 18 July 1989, I). ./.
Du Puy. R. P. Du Puy A I). Raum M297 (k, P); Tulear,
Tulear, Manombo, Foret de Mikea, env. 2 km au N de
Beroroha, 22°52'54"S, 43°33'25"E, 60 m, 0 Dec. 1993,
./. -/V. Labat, I). ./. Du Puy & .S’. Comtet 2463 (PARC, MO.
P); Tsinembo, 21 Oct. 1932, J. Ijeandri 444 (P); Trangaliy,
31 Oct. 1932. ./. Leandri 481 (MO. P), ./. leandri s.ri. (P);
foret a feuilles caduques sur calcaires de l’Antsingy, vers
Ambodiriana (F d’Antsalova), 100—150 m, 21—27 Jan.
I960, ./. Leandri & P. Saboureau 2751 (MO); foret a feuil¬
les caduques sur calcaires de I’Antsingy, vers Andobo (F
d’Antsalova), 100-150 m, 7 Feb., J. Ijeandri A P. Sabou¬
reau 3037 (MO, P); forets de I’Ouest, Anakarake, Moron¬
dava, Feb. 1923, M. I.oiwel 160 (P; syntype of Cordyla
madagascariensis)', Toliara, 55 km NF of Morondava via
route 8 at the forestry concession of the Centre de For¬
mation Professional Forestiere (CFPF). kirindy Forest, 5
km F of route 8 along principal concession road (Piste
CONOCO), along S boundary of forestry block, 20°4'S,
44°40'F, 35 m, 21 Mar. 1992, R. D. Noyes, I). K. Harder.
F. A. Rakotobe, T. G. Razafindrabeaza A J. P. Abraham
1053 (MO); Toliara, N of Toliara, in Foret de Mikea, ca.

I ft road-km W of Vorehe, 22°15'S, 43°25'F, 50 m, G.
McPherson, S. Razafimandimbison, M. Olsen A R. Alongi
17354 (MO); Hois de Bona (Ankazoabo) sud, Feb. 1970,
P. Marat 3574 (MO); Ankaladiny, sur la Bebibokn, July
1904, //. Perrier de la Rdthie 1414 (P: syntype of Cordyla
madagascariensis)', sur le Manambato, Oct. 1913, II. Per¬
rier de la Rdthie 4240 (P; syntype of Cordyla madagas¬
cariensis); environs de Majunga, & Ankatsepe, Aug. 1911,
II. Perrier de la Rdthie 4749 (P; syntype of Cordyla mad¬
agascariensis); Toliara, Zombitsy Forest, along Route Na¬
tional 7, 10 km from Sakaraha, 22°54'S, 44°38'E, 650 m,
27 Feb. 1990, P. R. Phillipson A J. R. Milijaona 3639
(MO); Toliara, Prefecture Morondava, approximately 60
km (by road) NF of Morondava, in the kirindy Swiss for¬
estry concession, 20°5'S, 44°38'F, 45 m, 25—31 Oct.
1990, C. F. Schatz 2982 (MO); Ainpandra, Tulear.
22°56'S, 44°32'F, 6 Feb. 1951, Sen ice Porestier 2838 (P);
Belo/Tsiribihina, foret d’Andranofotsy, 8 Jan. 1955, Service
Forestier 13154 (MO); T. Ankatoky, Beroroha, 16 May
1955, Service Forestier 13295 (MO).

312

Novon

l)ii|»uya nuuhiguscurieiisis sul>sp. tamaiimloi-
<les (Capuron) J. II. Kirkbride, comb. nov.
Basionym: Cordylu madagascariensis var. ta-
marindoides Capuron, Adansonia, s£r. 2. 8:
220. 1 968. Cordylu madagascariens is subsp.
tamarindoides (Capuron) I). ,1. Du Puy & Ca¬
bal, in Du Puy et al.. Log. Madagascar 207.
2002. TYPE: N Madagascar. Plateau calcaire
de I’Ankarana, environs d’Ambondromifehy,
Service Forestier (R. Capuron) 3031 (holotype,
P not seen; isotypes, K not seen, MO not seen,

P).

Specimens examined. MADAGASCAR. District
d'Ambilobe, montagne d’Andavakafanihy, calcaires de
PAnkarana, km 105 de la route d’Ambilobe, 200 m, 3
Feb. I960, ('.ours X II. Humbert 5620 (P); IN Madagas¬
car, Antsiranana Province, SE of Antsiranana (Diego Sua¬
rez), NV\ of Irodo village, Analafandrika forest, 12°38,2,,S.
49°3 1 '40"F, 60 m, 24 Mar. 1994, IK ./. Du Puy, J. An-
driantiana X 0. Andrianuntoanina M755 (k, P); N Mad¬
agascar, Antsiranana Province, “Windsor Castle” peak
near llomotro, NW of Antsiranana (Diego Suarez),
I2°I3'N, 49°I0'E, 2(H) in, 10 June 1989, I). ./. Du Puy,
II. R. l)u Pw\ X G. Rajamantanantsoa M267 (k. I’): An¬
tsiranana, Vovo Village Distr., ca. 6 km from Diego Suarez
oil Diego Suarez— Vovo Village road, and ca. 1.5 km S of
beach on Indian Ocean, 12°19'5"S, 49°23'7"F,. 85 m. 23
Apr. 1993, I). K. Harder. V/. C. Merello, S. G. Razafiman-
dimisan X T. G. Razajinfrabaeza 1693 (MO); plateaux cal¬
caires de PAnkarana du nord entre Ambilobe et Anivor-
ano, 1 2°57'S, 49°8'E, 200-350 m, 4-9 Mar. 1951. //.
Humbert & R. Capuron 25485 (P); Province de Diego-
Suarez. collines et plateaux calcaires de PAnkarana du
nord, vers grotte dn Eaniliv. 30—350 m, 24 Jan.— 29 Feb.
1960. //. Humbert & G. Goars 32728 (P); Antsiranana.
Ambilobe, Matsaborimanga, Reserve Spdciale
gd’Ankarana, Tsingy an dessus du lac Vert, 12°55'26”S.
49°5' 10"K, 300 m, 20 INov. 1996, ./.-A'. Labal , F. Ran-
drianjohany X (). Andrianuntoanina 2790 (I*); Antsiran¬
ana, Ambilobe, Matsaborimanga, Reserve Sp^eiale (PAn¬
karana, |)isle du lac Vert, debut de la zone calcaire, avant
Amposatelo, 12°55'7”S. I9°5,46”E. 180 m, 21 Nov. 1996,
./.-A', Dibat. E. Randrianjohany (V 0. Andrianantoanina
2803 (P); Antsiranana, Reserve Speeiale (PAnkarana.
piste vers le lac Vert, plaque du Tsingy, I2°54'43"S,
49°6'39"K. 180 in, 18 Feb. 1994, R. ./• Rakotomalaza 142
(MO); NW Madagascar. Antsiranana province, Ankarana
Reserve, 3.4 km from Ambondromifehy, 12°52'S,
49°14'E, ca. 220 m, II. IK Sell ire. IK J. Du Huy & G. R.
leu: is 2542 (MO. P); Marolandilia. Bevoav-Morondava, 14
Nov. 1952, Service Forestier 6062 (P); Andranolava- Pulsar,
Foret de Vohibasia, 15 Jan. 1954, Service Forestier 8296
(P); Montague des Franyais-Diego Suarez, Service Forestier
12731 (k, P); District Morondava, Canton de Rcfasy, Ren¬
te moyenne, Anadabo, 8 Oct. 1956, Service Forestier
16555 (MO. P).

Du Puy and Labat (2002: 297) gave the distri¬
bution ol Dupuya ma dagascarie n s is from south¬
western to northern Madagascar in seasonally dry
forest with a dry season of 6 to 8 months, found up
to ca. 500 m elevation, from the Forest ol Sakaraha
to Antisiranana. Dupuya madagascariemis subsp.

madagascariensis lias the largest distribution of the
two subspecies throughout southern and southwest¬
ern Madagascar, and I). madagascariensis subsp.
tamarindoides occurs at the northern tip of Mada¬
gascar. mainly to the north ol the Ankarana and
Analamerana Massifs (Du Puy & Labat, 2002:
297). Dupuya haraka and IK madagascariensis are
allopatric, except for their populations on the An¬
karana Massif. According to Du Puy and Moat’s
(2002: map 4) vegetation zones, the Ankarana Mas¬
sif has both humid Sambirano forests and season¬
ally dry forests. The herbarium labels are not sul-
ficient to determine the exact habitat ol each
species on the Ankarana Massif, but they are sym-
patric.

Acknowledgments. 1 thank the following insti¬
tutions for loan ol their Cordylu— Dupuya specimens
that made this study possible: K, MO. P. and US.
1 also thank Amy Y. Rossnian ( H PI). John II. Wier-
sema (BARG), and two anonymous reviewers for
constructive criticisms of the manuscript.

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Appendix 1. Cordyla Specimens Examined.

Cordyla africana Loureiro

KENYA. I ,amu District, W'itu Forest near Mambosasa
Game Forest Post, 2°23'S. 40°32'E, 10—20 m. 20 Feb.
1977. R. B. Faden Ac A. ,/. Faden 77/567 (BARC, MO.
P): kilili District, Pangani, crossing of Lwandani Stream
on Chonyi— Ribe road, 3°5US, 39°4UE, 160 m. If) Feb.
1977. R. B. Faden, A. ./. Faden. J. B. Gillett A A'. Gaeh-
athi 77/447 (MO. US); Lamu District. Witu Forest. N of
road, forest at northern point of reserve and up to 1 km
S. If) Nov. 1988, .S. A. Robertson Ac (J. Luke 54BB (k. US);
kwalc District, Mrima Hill. 10 Apr. 1978. B. Verdcourt
5274 (k). MALAWI. Mangochi District. Monke\ Bay/
Mangochi road, roadside, ./. /.. Balaka, A. ,/. Salubeni Ac
R. II. Rwatha I2B2 (MO); Machinga District. Liwonde,
roadside, 14 Aug. 1986, A. ./. Salubeni, ./. L. Balaka Ac
C. A. Musa 4699 (MO); Zomha District, Chigwere Village.
Chisi Island, Lake Chilwe, Chisi Hill slopes, 10 Nov.
1986. A. ./. Salubeni Ac R. B. Kivatha 4850 (MO): Zomha
District, Zomha Botanic Garden, edge of Mulunguzi river.
21 Oct. 1987. F. ./. Tawakali Ac M. S. Thera 1152 (MO).
MOZAMBIQUE. Gaza, entre Javanhana e Caniyado,
proximo do rio Limpopo, 9 Oct. 1958. /.. 4. Grandvaux
Barbosa 8548 (k); Province du Gorongosa. Village di Sa-
binga, Rivier du Vunduzi, 23 Sep. 1906. G. \asse 418 (P).
SOUTH AFRICA. Zululand. St. Lucia, western shores.
Hell’s Gate Peninsula along side road, 30.5 in, 12 Mar.
1982, A. Nicholas Ac I). MacDevette 1276 (MO). TAN¬
ZANIA. Mtvvara District, Tangazo Village, 30 km S of
Mtwara on road to Maharunga and Ruvuma R. mouth, 6
Nov. 1978. F. C. Magogo & R. R. lanes RRI476 (k): Moshi
District, Moshi— Arusha road, 19 Nov. 1951, McCoy-Hill
19 (k): Bezirk Lindi, Lutamba-See, 40 km westlieh Lindi,
200—250 m, 7 Sep. 1934. //. ./. Sehlieben 5267 (Pi: East¬
ern Province, kilosa District, Old Boma, 18 Oct. 1961. S.
R. Semsei 3820 (k). ZAMBIA. Eastern Province, foothills
E of Machinje Hills, 1000 m, 12 Oct. 1958, A. K. B.
Robson 66 ( is. ). ZIMBABWE. District Mrewa. I zumbi
Tribal Trust Land. Chiore School Road. 5 Oct. 1969, II.
I). I.. Corby 2 1 88 (k); District Mrungwe, Mana Pools, Sep.
1969. T. A. S. Gordon 44 (k); Sabi valley, above 549 m,
23 Mar. 1905, W. M. hnigden 18182 (US); Causeway,
1975. \ational Botanic Garden s.n. List Foil Shipment Fill
(BARC); Weller s.n. List #95 Shipment #41 (BARC).

Cordyla densillora Milne-Redhead

TANZANIA. Mpwapwa. 1067-1219 m. 18 June 1933, B.
I). Barit 4755 ( k ); Iringa Province, Iringa, 914-1219 m,
18 Sep. 1936. II. F. Fmson 618 (k); Iringa District,
Msembe, 823 m, Apr. 1970. F. ./. Greenway 4420 (K);
Msembe, near the Rest House, 762 m, 5 Aug. 1969. B. J.
Greenway Ac Kanuri 18690 (k); Morogoro Region, Moro-
goro District, Ruaha gorge on Morogoro-Iringa road,
36°40'S. 7°30'E. 26 June 1986, ,/. ! Arnett A1 ./. hwett 852
(k); Iringa District, Nvangolo. 16 July 1956, F. Milne-
Redhead Ac F. Taylor 11228 (k); Iringa District. Ruaha
National Park, kimiramatonge Hill. 823 m, 23 Oct. 1970,
//. M. Richards Ac S. Arasululu 26828 (BARC, k).

Cordyla pinnata (A. Richard) Milne-Redhead

GAMBIA. Near Georgetown, 10 Jan. 1927../. M. Dalziel
8029 (k): hords de la Gambia. Mar. 1829. G. S. Ferrottet
s.n. (P). GUINEA. Region de Gaoual. pres koumbia. 14
Jan. 1979. S. Lisowski 51176 (k). MALL Diondiou. If)
June 1899. A. Chevalier 8845 (P); kakoulou (kayos), I 1

314

No von

Jan. 1937, R. Dubois 248 (P); & 18 km de S6gou en di-
rection de Mopli, 14 May 1989, D. Gautier- B/guin & L.
Gautier-B/guin 1414 ( MO). NIGEKIA. Province Niger,
District Mokwa, 29 Dec. 1964, I). I‘. Stanfield 55965 (k).
SENEGAL. Thies-Diourbel. Kaolack, 23 Mar. 1954, R.
P. Berhaut 4945 (P); S&lhiou (Casamancc), Feb. 1900, A.
Chevalier 8587 (P), Heudelot 867 (P); Boundou, Woulli,
1836, Heudelot s.n. (P); Belidjibara, 99 km apr£s kous-
saus, Saiisatic. 15 Feb. 1983, P, Lurie 1720 (P), G. S.
Perroltel 22 (P), s.n. (P); km 220 cntrc Diourbel et Gas-
sane, II Dec. 19.30, J. Trochain 1 1 14 (P); Tambacounda-
Diabougou-Vtdingara, 21 June 1931.,/. Trochain 1258 (P).

Conlylu richardii Milne-Bedbead

SUDAN. Waw, Bahr Al Ghazal, 28 Feb. 1926, G. Aylmer
27/22 (k); Waw, A. II. Braun s.n. (k); Imatong Mountains,
just S of Ngarama along the road to Molongori. near the
northernmost point of the continuous chain, 700 m, 13
Mar. 1982, /. Friis & K. Vollesen 1186 (k). UGANDA.

West Madi district, at Metuli, 1219 m, Dec. 1931, /V. V.
Brasnett 209 (k); Metu, Madi. ca. I mi. SW of Metu rest-
house, on S side of Dzele hill, I 158 m, 5 Dec. 1947, II.
C. Dawkins 202 (k).

Cordyla soiualiensis J. 8. Gillett subsp. littoralis
J. B. Gillett

SOMALIA. Merca, 29 Jan. 1954, Bally 9527 (k).

Gordyla soiualiensis J. B. Gillett subsp. soiualiensis

ETHIOPIA. Oagden: W of Warder, 6°50'N, 44°10'E,
914 m, 16 Oct. 1953, E. Ellis 126 (k): 10 mi. SW Warder,
16 Oct. 1954, C. F. Henning 271 ( k); 4 mi. E of Mersin.
15 July 1958, C. F. Henning 1557 (k). SOMALIA. 17
km S of Gellinsoor, 520 m, 7 Feb. 1973, P. R. 0. Bally
& R. Melville 16282 ( k); ca. 10 km from Bula Burti to
Ceel Bum; 4°0'N, 45°30'E, 15 May 1979, l. I. Finn &
(). J. Hanson 4019 (k).

A New Species of Rytigynia (Rubiaceae, Vanguerieae) from
Northeastern Tanzania, with Notes on I lie
Circumscription of the Genus

//. Lantz

Department of Systematic Botany, Evolutionary Biology Centre, Uppsala University, Norbyv.
I8D, SE-752 36 Uppsala, Sweden, henrik.lantz@ebc.uu.se

Roy E. Gereau

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

roy.gereau@mobot.org

Abstract. We discuss the history and circum¬
scription of the genus Rytigynia (Rubiaceae, Van¬
guerieae) and present the morphological characters
used for the delimitation of t lie genus in a new.
restricted sense. We describe Rytigynia dichasialis,
a new species from northeastern Tanzania that is
unique in the genus by having long-pedunculate
and dichasially cymose inflorescences with well-de¬
veloped secondary axes. We compare the new spe¬
cies to other, morphologically similar species of Ry¬
tigynia and present the characters used to
distinguish among them.

Key words: Rubiaceae, Rytigynia, Tanzania,
Usambara Mountains, Vanguerieae.

The Vanguerieae are a tribe of ca. 600 species
in the family Rubiaceae with a distribution in Af¬
rica, Madagascar, South and Southeast Asia, the
Pacific, and Australia. The center of diversity is in
East Africa, and these are also the species that
have been studied most extensively. In the treat¬
ment of this tribe for the Flora oj Tropical East
Africa (Verdcourt & Bridson, 1.991) and in the pub¬
lications that laid the foundation for this work
(Bridson, 1985, 1986, 1987a, 1987b; Verdcourt,
1981, 1987), the current circumscriptions of most
of the genera were established and a large number
of new species were described. Both specific and
generic delimitations in the tribe are notoriously
difficult. Lantz et al. (2002) and Lantz and Bremer
(2004) have shown that 5 of the 27 currently rec¬
ognized genera are para- or polyphyletic, and sev¬
eral other genera lack readily observable diagnostic
characters.

Rytigynia Blume (1850), a genus of about 60
species with a widespread distribution in Africa
and Madagascar, is one of these problematic genera
in Vanguerieae. Phylogenies based on molecular

data (Lantz et al., 2002; Lantz & Bremer. 2004)
have shown that Rytigynia s.l. (e.g.. Verdcourt,
1987) consists of a core group of species, here re¬
ferred to as Rytigynia s. sir., that are closely allied
to Fadogia Schweinfurth, Fadogiella Robyns, and
Hutchinsonia Robyns. However, Rytigynia s.l. also
includes some species more closely related to other
genera (as noted by Verdcourt & Bridson, 1991:
803). Lor example, Rytigynia bugoyensis (K. Krau¬
se) Verdcourt anil R. mrimaensis Verdcourt are
more closely related to the type species of Can¬
thium Lamarck. C. coromandelicum (Burman f.)
Alston, than to other species of Rytigynia. Both
species have supra-axillary spines, often in com¬
bination with brachyblasts, a diagnostic character
of Canthium s. str., to which genus both have re¬
cently been transferred (Lantz & Bremer, 2004).
Rytigynia fuscosetulosa Verdcourt, a species with
linear calyx lobes that is currently placed in Ryti¬
gynia subg. Sali Verdcourt, is closely related to
Vangueria Jussieu (Lantz & Bremer, 2004) anil
should not remain in Rytigynia s. str. (Lantz & Bre¬
mer, in prep.).

Traditionally, Rytigynia s.l. has been delimited
by a lack of characters present in other genera of
Vanguerieae rather than by unique diagnostic char¬
acters of its own. In practice, Rytigynia s.l. has
included shrubby species with few-flowered inflo¬
rescences lacking the most obvious characters of
the plurilocular genera related to and including
Vangueria. The vaguely defined combination of
characters used in its circumscription has evolved
multiple times in the tribe, and Rytigynia s.l. is
thus polyphyletic. Also complicating ihe circum¬
scription of Rytigynia is the equally vague circum¬
scription of Canthium s.l. (shown to be polyphyletic
by Lantz & Bremer. 2004), with focus only on the
bilocular ovaries, which has made it difficult to dis-

Novon 15: 315-318. 2005.

316

Novon

tinguish between this genus and Rytigynia. It is
sometimes unclear on which basis the species are
referred to one genus or the other, as in the above-
mentioned Rytigynia bugoyensis and R. mrimaensis.

W ith the knowledge of the phylogeny of the tribe
based on molecular data (Lantz et ah, 2002; Lantz
& Bremer, 2004). Rytigynia s. sir. is morphologi¬
cally delimited by the following combination of
characters: shrubs or small trees; leaves persistent:
domatia often present (on abaxial leaf surface in
axils of secondary nerves, the normal position in
Vanguerieae); inflorescences usually shortly pedun¬
culate 5 mm) and with poorly developed sec¬
ondary axes; calyx truncate or very shortly toothed;
and ovaries hi- or plurilocular. The truncate or very
shortly toothed calyx is of special importance to
distinguish Rytigynia s. str. from Rytigynia s.l.
Some species of Afrocanthium (Bridson) Lantz &
Bremer (e.g.. Afrocanthium pseudoverticillatum (S.
Moore) Lantz; see Lantz & Bremer, 2001) match
much of this description, but these are without ex¬
ception deciduous. Thus any species currently
placed in Rytigynia s.l. but deviating from this de¬
scription are likely candidates for exclusion from
the genus, though we will await molecular data be¬
fore any more transfers are made. However, only a
small number of the remaining Rytigynia species
will be excluded from Rytigynia s. str. and further
transfers will not change the geographic distribu¬
tion of Rytigynia ; Rytigynia s. sir. is widespread in
Africa and Madagscar.

During the identification of a large number of
Vanguerieae specimens collected during the Tan¬
zania Botanical Training Programme, the first au¬
thor discovered an undescribed species that could
only be identified to genus with difficulty. The com¬
bination of 3-locular ovaries, domatia, and a trun¬
cate calyx clearly indicate a position in Rytigynia
s. str.. but the many-flowered inflorescences with
well-developed evmose branching are not known
from other species in the genus. The specimen rep¬
resents a new species of Rytigynia s. str. exhibiting
hitherto unknown features in the genus in this re¬
stricted sense.

Rytigynia diehasialis Lantz & Gereau, sp. nov.
T\ PE: Tanzania. Tanga Region: Muheza Dist.,
Amani East Forest Reserve, near edge of road.
5°05'36"S, 38°36'01"E, 500 m, 29 Jan. 1999.
A. Ntemi Sallu 247 (holotype, NH L: isotypes,
BR. K. MO. UPS, eucfp (reference collection
of East Usambara Catchment Forest Project)).
Figure 1.

Haec species quoad stipulas longe appendieulatas

bracteas connatas cupuliformes etiam fforium aspectum ad
Rytigyniam monantham niaxime aecedit, sed ab ea foliis
majoribus atque infforescentiis 5- ad 14- (nee I - vel 2-)
flnris, a eongeneris ad Rytigyniam sensu strieto pertinen-
tibus infforescentia dichasiali pluriflora (nee fasciculata
subumbellata umbellatave plerumque paueiffora) perfacile
distinguitur.

Shrub ca. 2.5 m tall; twigs light brown, lenticel-
late. densely short-pubescent with straight to some¬
what curved or wavy trichornes when young, gla-
brescent with age; bark smooth. Stipules with basal
parts connate into a sheath 2.0— 2.5 mm long with
trichornes present within, lower part of sheath pu¬
bescent externally, upper part glabrous and shiny,
each stipule with free apical filiform appendage 5—
7 mm long, soon deciduous; petiole 2— A mm long;
leaf blade ovate to ovate-oblong, 4.7—10.5 cm long,
1.5— 4.4 cm wide, L/W = 2.3— 3.1, long-acuminate
and minutely apiculate at apex, acute to obtuse and
sometimes asymmetric at base, with short adpres-
sed to somewhat spreading trichornes with papil¬
lose-swollen bases above, with slightly longer (‘pa¬
pillose trichornes beneath, denser than above but
surface not obscured, the costa and secondary
nerves more densely pubescent; secondary nerves
4 to 6 on each side of costa, eucamptodromous at
base to weakly brochidodromous at apex; domatia
present as tufts of white trichornes in axils of sec¬
ondary nerves. Inflorescence axillary, 5- to 14-flow-
ered. evmose (dichasial), all parts shortly pubes¬
cent; peduncle slender, 7-13 mm long; bracts and
bracteoles present as connate sheaths or sometimes
free, then ovate to lanceolate; dichasium with 2 or
3 orders of branching, the central solitary flower
apparently always present, the lateral secondary
and/or tertiary axes often reduced and somewhat
monochasial; pedicels 4—6 mm long in flower, elon¬
gating to 17 mm in fruit. Flower buds acuminate,
glabrous or with a few short trichornes on outside
of corolla lobes, slightly constricted below lobes;
calyx tube obovoid, 1.0— 1.8 mm long, short-pubes¬
cent, limb tube 0. 2-0.3 mm long, glabrous, trun¬
cate or somewhat undulate-crisped and thus ap¬
pearing very slightly toothed; corolla 5-tnerous,
cream, glabrous outside at anthesis, the tube ea. 2
mm long with deflexed trichornes attached at throat
inside, these rather straight but usually somewhat
curled toward rounded apices, the throat with a
smooth rather cartilaginous ring, the lobes 4—5 mm
long, narrowly triangular, the apex acute with api-
culum ca. I mm long; anthers 5, inserted in corolla
throat with upper half visible, dorsal side without
darkened connective; ovary 3-locular; sty It* exsert-
ed 1.0— 1.2 mm from corolla throat: stigmatic club
green, cylindrical, 1.0— 1.2 mm long, with pro-

Volume 15, Number 2
2005

Lantz & Gereau

Rytigynia dichasialis from Tanzania

317

|'j ir(] rc I. Rytigynia dichnsiulis I,ant/ & Gereau. — A. f lowering plant, habit. B. Flower, with corolla opened to
show internal triehomes. — C. Flower laid. — 1). E. Nolle of twig with leaf bases and stipule with appendage. — I*. Node
of twig with stipule, appendage fallen. — O. Node with stipule and one inflorescence (of two). — II. Immature fruit. A-
F and H based on isotypes at BIT k, and MO; inflorescence detail G based on isotype at UPS.

nounceil ridges, 3-lobetl at apex. Immature fruit
subglobose, to 6 mm long and 7 mm wide, glabrous,
with 3 pyrenes.

This species is unique in the genus in its rela¬
tively long-pedunculate and several-flowered inflo¬
rescences w ith well-developed branching. I he most
mature flowers and fruits consistently appear in a
terminal position, indicating a determinate and es¬
sentially cymose inflorescence type that is best de¬
scribed as a dichasium, though with some of the

secondary and/or tertiary branches often reduced
anti monochasial. This inflorescence condition is
unique in Rytigynia s. str., which characteristically
has inflorescences that are fasciculate or shortly pe¬
dunculate and umbellate or subumbellate. and
rarely have more than four flowers. The mostly con¬
nate bracts and bracteoles of this new species are
also distinctive. Similar bracts and bracteoles are
known from several other species in the genus, al¬
though the larger number of flowers in R. dichasi-

318

Novon

alis makes them more noticeable. Sterile specimens
of R. dichasialis could be confused with R. mon-
antha (K. Schumann) Kobyns, which is quite sim¬
ilar to R. dichasialis in characters of leaf shape,
long-appendaged stipules, bracts, braeteoles, and
flowers. However, R. dichasialis lias larger leaves
(to 10.5 X 4.4 cm in R. dichasialis vs. to 0X4
cm in R. monantha) and is only known from the
type locality at an elevation of 500 m in northeast¬
ern lanzania, while R. monantha is known from
elevations of 775— 1940 m and is widespread in Bu¬
rundi, Rwanda, Malawi, Tanzania, and Zambia. A
poorly known species from Malawi, R. pawekiae
Verdeourt, shares a similar number of flowers per
inflorescence with R. dichasialis , but is differenti¬
ated by smaller, more densely pubescent leaves,
peduncles only up to 4 mm in length, inflorescenc¬
es congested and subumhcllate, and external tri¬
e-homes present on the corolla lobes at anthesis.
Rytigynia duhiosa (l)e Wildeman) Robyns, known
from Cameroon, the Democratic Republic of the
Congo, and Uganda, has inflorescences with up to
If) (lowers (Verdeourt & Rridson, 1991: 810). but
has subumbellate inflorescences with short pedun¬
cles (up to 6 mm) and glabrous leaves.

Rytigynia dichasialis is known only from the type
collection. Amani East Forest Reserve was origi¬
nally gazetted in 1955 (Iversen, 1991: 30), and was
tlx- smallest ol the six forest reserves that were
combined in 1997 to create the Amani Nature Re¬
serve (Doody et ah, 2001: 0), the largest bloc of
forest in the East Usambara Mountains. Of the 264
species of trees and shrubs recorded from vegeta¬
tion plots in the Amani Nature Reserve, 19(7.2%)
are endemic to the Usambara Mountains, and 1 I
(4.2%) are endemic to tin- East Usambara Moun¬
tains; tin- Reserve thus contains a significant pro¬
portion of regionally restricted species within an
area of only 83.6 km- (Doody et ah, 2001: 62). hike
most continuous forest blocs ranging from lowland
to submontane forest in the Eastern Are Mountains,
the Amani Nature Reserve is particularly rich in
Rubiaceae. The East Usambara Mountains have
had a history of moderately intensive botanical re¬
search and collection dating back at least to 1892

(see Iversen, 1991: 1-11), but nevertheless there
remain new taxa to be discovered among existing
and future collections.

Acknowledgments. We are greatly indebted to
Charlotte Taylor for the illustration and insightful
comments on earlier versions of the manuscript. We
sincerely thank Albert Ntemi Sallu for having col¬
lected this probably rare species in the field, and
Nike Doggart of the Tanzania Forest Conservation
Group for valuable information about tlx* Amani
Nature Reserve.

Literature Cited

Illume, C. L. 1850. Museum botanicum Lugduno-Batavum
1(12): 177-192. K. J. Brill, Leiden.

Bridson. I). M. 1985. The reinstatement of Rsydrax (Ru¬
biaceae, subfam. Cinohonoideae tribe Vanguerieae) and
a revision of the African species. Kew Bull. 40: 687-
725.

- . 1986. The reinstatement of the African genus

Keetia (Rubiaceae subfam. Cinehonoideae, tribe Van¬
guerieae). Kew Bull. 41: 965—994.

- . 1987a. The recognition and recircumscription of

the African genus Multidentia (Rubiaceae— Vanguer¬
ieae). Kew Bull. 42: 641-654.

- . 1987b. Studies in African Rubiaceae— Vanguer¬
ieae: A new circumscription of Pyrostria and a new sub¬
genus, Canthium subgen. Bullockia. Kew Bull. 42:
61 1-639.

Doody, K. /.. K. M. Howell & E. Fanning (editors). 2(H) I .
Amani Nature Reserve: A biodiversity study. Fast
Usambara Conservation Area Management Programme
technical Paper 52: i— xiii, 1 — 123. Frontier Tanzania,
Ministry of Natural Resources and Tourism, and Met-
sahallitus Consulting; Dar es Salaam, Tanzania, and
Vantaa, Finland.

Iversen. S. I. 1991. Tlx- Usambara Mountains, NF Tan¬
zania: Phytogeography of the vascular plant flora. Symb.
Rot. Upsal. 29(3): i-vi, 1-234.

Lantz, H. & B. Bremer. 2004. Phylogeny inferred from
morphology and l)NA data: Characterizing well-sup¬
ported groups in Vanguerieae (Rubiaceae). Rot. .1. Finn.
Soc. 146: 257-283.

- . K. Andreasen & B. Bremer. 2002. Nuclear rl)NA

ITS data used to construct the first phylogeny of Van¬
guerieae (Rubiaceae). PI. Syst. Evol. 230: 173-187.
Verdeourt, B. 1981. Notes on African Rubiaceae. Kew
Bull. 36: 493-557.

- . 1987. Notes on African Rubiaceae— Vanguerieae.

Kew Bull. 42: 123-199.

- & I). Bridson. 1991. Rubiaceae (Part 3). In: R.

M. Polhill (editor). Flora of Tropical East Africa. A. A.
Balkema, Rotterdam/Brookfield.

Validation of Mimulicalyx paludigenus (Scrophulariaceae),
a Species Endemic to China

Li De-Zhu and Cai Jie

Kunming Institute of Botany, Chinese Academy of Sciences,

Kunming, Yunnan 650204, China, dzl@mail.kib.ac.cn; j.tsai@mail.kib.ac.cn

Ahstract. In (he course of writing the account of
Scrophulariaceae for the Flora Yunnanica, it was
noticed that one species of Mimulicalyx, M. palu¬
digenus, described from Yunnan Province, south¬
western China, in 1979 was invalid because two
type specimens (one fruiting, one in flower) were
designated. The name is validated here, with the
fruiting specimen designated as the holotype.

Key words: China, Mimulicalyx, Scrophulari¬
aceae, validation.

Mimulicalyx P. C. Tsoong has been described as
a genus endemic to China by Tsoong (1979) in his
account of Scrophulariaceae in the Chinese-lan-
guage Flora Reipublicae Popularis Sinicae, Tomas
67. Up to now only two species, M. rosulalus P. C.
Tsoong and M. paludigenus, were recorded in this
genus. While preparing the account of Scrophular¬
iaceae for the Flora Yunnanica, we found that a
nomenelatural mistake existed in M. paludigenus.
Two specimens, representing (lowering and fruiting
stages, were designated as types simultaneously
(Tsoong, 1979). The name, therefore, did not accord
with Article 37 of the International Code of Botan¬
ical Nomenclature (Zhu, 2001) and made the name
Mimulicalyx paludigenus Tsoong invalid. Unfortu¬
nately, this problem was not discovered in the Ln-
glish ami updated edition of the Flora of China
(Hong et al., 1998). The specimen that represented
flowering material could not be found during re¬
examination, so Mimulicalyx paludigenus is vali¬
dated here by reference to the previously published
diagnosis by Tsoong (1979) and by designating the
fruiting specimen, C. W. Wang HI 476, as the ho¬
lotype.

Mimulicalyx puludigenus P. C. Tsoong ex I). /.
Li & J. Cai. sp. nov. TYPE: China. Yunnan:
Mong-tze Hsien, Shau-shin-jay, 1400 m, 5
Sep. 1935 (fr), C. W. Wang BI476 (holotype,
PE).

hitin diagnosis: Tsoong, in Flora Reipublicae
Popularis Sinicae 67(2): 400. 1979.

Although the holotype of Mimulicalyx paludigen¬
us, which represents merely the fruiting stage, does
not exhibit floral characters, it is easy to distinguish
the pair of species by their vegetative characters.
In M. paludigenus, leaves are 3-4 cm long, borne
on the stems oppositely and are mostly amplexi-
caulous except the basal one or two pairs with 0.4
cm long petioles; the margin is entire or with a few
small teeth: the pedicel is longer than the bract,
and to 4 cm long or longer at l lit* fruiting stage. In
M. rosulatus, basal leaves are in a cluster and more
than 4 cm long with a 1—1.5 cm long petiole; can-
line leaves are 0.8-2. 5 cm long and amplexieau-
lous; bracts are conspicuously shorter than pedi¬
cels, but the pedicel is usually not more than 3 cm
long al the fruiting stage.

Literature Cited

Hong. I). Y., H. I?. Yang, C. L. Jin & N. H. Holmgren.
1998. Scrophulariaceae — Mimulicalyx. Pp. 52—53 in Z.
Y. Wu & P. H. Raven (editors). Flora of China, Vol. 115.
Science Press, Beijing, anti Missouri Botanical Garden
Press, St. Louis.

Tsoong, P. C. 1979. Scrophulariaceae — Mimulicalyx. Pp.
209-21 I. 400 in S. S. Chien & W. Y. Chun (editors).
Flora Reipublicae Popularis Sinicae, Vol. 67, No. 2.
Science Press, Beijing.

Zhu, G. H. 2001. International Code oi Botanical Nomen¬
clature (St. Louis Code), Chinese Ed. Science Press,
Beijing, and Missouri Botanical Garden Press, St. Lou¬
is.

Novon 15: 319. 2005.

Notes on Succulent Cynanchum (Apocynaceae, Asclepiadoideae)

in East Africa

Sigrid Liede-Schumann and Ulrich Move

Department <>l Plant Systematics, University ol Bayreuth. 95444 Bayreuth, Germany,
sigrid. I iede@uni-bayreuth.de; ulrieh.meve@uni-bayreuth.de

Austhact. Changes in sucenlent Cynanclwm U.
species are needed for the Flora of Somalia ac¬
count of Apocynaceae subfamily Asclepiadoideae.
1 he following changes are proposed within the tribe
Asdepiadeae, subtribe Cynanchinae. Cynanchum
viminale subsp. crassicaule, another subspecific
taxon in the C. viminale complex, is described as
new, based on morphological, ecological, and mo¬
lecular evidence. Its chromosome number is spec¬
ified with 2 n = 22. The new combination Cynan¬
chum viminale subsp. stipitaceum is proposed, and
Sarcoslernma andongense Hiern is recognized as a
new synonym of C. viminale subsp. stipitaceum.

Key words: Africa, Apocynaceae, Asclepi-
adeae, Asclepiadoideae, Cynanchinae, Cynan¬
chum, Kenya, Sarcoslernma , Somalia.

Becently it has been shown that the African
members of Sarcoslernma R. Brown are nested in
the succulent Cynanchum clade, which further
nests within/among the Madagascan Cynanchum
species (Uiede kunze. 2002). Meve and l.iede
(2002) therefor*1 concluded that Madagascar is the
* enter of origin for taxa previously summarized un¬
der Sarcostemma and that the taxonomic difficulties
in the ( lynanchum viminale complex reflect the am¬
biguity associated with a comparatively young, rap¬
idly evolved and still evolving taxon. Nevertheless,
tin* traditional concept that florally distinctive taxa
are recognized at the specific level, while florally
similar but vegetatively distinctive taxa are consid¬
ered subspecies under Cynanchum viminale (l.iede
& Meve, 1995; Meve & Uiede, 1996), is still a valid
approach to handle diversity in this group. A re¬
cently conducted RAPI) analysis (l.iede & Meve.
unpublished) supports an infraspecific concept for
C. viminale. W hile the unraveling of the Cynan¬
chum complex as a whole has to wait for the con¬
clusion of this analysis, the upcoming publication
of the Flora oj Somalia makes it necessary to pub¬
lish here a new subspecies within the Cynanchum
viminale complex.

Cynanclium viminale I,, subsp. crassicaule
Uiede & Meve. subsp. nov. T\ PE: Kenya. Tai-
ta Oistr.: SVi side of Ml. Kasigau. 1 150 m, 12
Feb. 1996, I). ./. Coyder. F. S. Masinde, U.
Meve & C. Whitehonse 4029 (holotype, K; iso¬
types, EA, UBT in alcohol). Figure 1.

Cynancho viminali typico af finis se<l habitu non-tortuo-
so arcuato, caulibus crassis, inflorescentihus sessilibus ex-
tra-axillaribus e-t terminalilnis, gynostegio conspicue lain
differt.

Plants erect to decumbent, sarmentose (arching
over the ground or scandent in bushes), non-twin¬
ing. richly irregularly branched; subterranean or¬
gans consisting of fibrous roots; shoots perennial,
succulent, prominently glaucous, sparsely glabres-
eent over the entire surface with flexuous tri-
cliomes, 600—800 gm long; internodes cylindrical.
1—20 cm long. 6—1 1 mm diam.; latex white, caustic
(fide P. Powys. Kenya, pers. comm.), heaves reduced
to scales, not caducous, sessile, blades fleshy when
young, papery when old, 1.4—2 mm long, ca. 1.5
mm wide, triangular-deltate. Inflorescences always
one per node, extra-axillary and terminal, growth
continued beyond inflorescence, 5- to 10-flowered,
all flowers open synchronously, sciadioidal. sessile.
Flowers faintly unpleasantly scented; floral bracts
ca. 0.8 mm long, ca. 0.6 mm wide at the base,
triangular, with trichomes; pedicels 5—7 mm long,
densely covered over the entire surface with flex-
uous trichomes, 600—800 gun long; florid buds 5-4)
mm long, 5 mm diam., conical; calyx basally fused,
abaxially with trichomes, lobes 1.2— 1.5 X 1-1.2
mm. triangular-deltate, apically acute; corolla (>— 8
mm long, cream to bright yellow; lobes 2.5-3 mm
wide, patent, oblong, apically acute, with revolute
margins. Gynostegial corona ivory in color, of free
staminal lobes (Cs) connate to a ring of staminal
and interstaminal parts [C(is)]; staminal corona (Cs)
1.5— 1.8 mm high, equaling the gynostegium, lobes
unguiculate, filled with loose parenchyma; ring of
staminal and interstaminal parts [C(is)] 1-1.2 mm
high, shorter than the gynostegium, cyathiform. Gy¬
nostegium 2.6—3 mm high, 4-4.5 mm diam., ses¬
sile; stamens with filament of 800 gun length, an-

Novon 15: 320-323. 2005.

Volume 15, Number 2
2005

Liede-Schumann & Meve
Succulent Cynanchum

321

figure I. Cynanchum viminale I,, subsp. crassicaule Fiede & Meve. — A. Habit, with flowering (left) and fruiting
stems (right). — lb Budding extra-axillary inflorescence. — C. Flower. — I), (iynostegium with corona in lateral view.

_ F. Pollinarium. _ F. Style-heail with two pollinaria attached. (Drawn hy U. Meve from the type. Cmyder cl al. 4029.

A drawn from a photograph of the type plant taken at the type locality; B-F drawn from alcohol material of the isotype
in UBT.)

322

Novon

thers about as long as broad, trapezoidal, abaxially
planar; anther wings 700 /ini long, convergent, con¬
sisting of distal ridge alone; anther wings of adja¬
cent anthers parallel to each other, basally wid¬
ened. centrifugal; connective appendages 0.5— 0.6 X
0. 1-0.5 nun, triangular, narrower than the stamen,
slightly indexed. Pollinarium : eorpusculum 220-
240 grn long, elliptic, margins of the corpuscular
cleft parallel; caudieles 200 gm long, (sub-)basally
inserted at the eorpusculum, cylindrical, genicu¬
late; pollinia apieally attached to the caudieles,
pendulous, 300 gm long, 100 gin wide, elliptical
in cross section, elavate. Style-head white, 1.2-1. 3
mm diam., 0.8—1 mm long, upper part 0.4— 0.5 mm
long, equaling the lower part, bifurcate. Follicles
always one per (lower, 60—100 mm long, 4—5 mm
diam.. narrowly oblong, round in cross section, api -
cally acute, but not beaked, with sparse indumen¬
tum; seeds 6—7 mm long, 2.5—3 mm wide, ovate,
light brown; seta side smooth, aseta side with
patchily arranged trichomes, 50 gun long; margin¬
ally with 0.5— 0.7 mm wide wing with entire margin;
coma 20—25 mm long.

Chromosome number. 2 n — 22 (vouchers: Goy-
der et at. 4029, Liede & Newton 2172).

Phenology. Flowering and fruiting from Feb¬
ruary to November.

Habitat and distribution. Cynanchum viminale
subsp. crassicaule occurs at altitudes of around sea
level close to shore up to the higher foot of Ml.
Kilimanjaro at ca. 1400 m a.s.l. Typically it grows
in Acacia savanna and scrub of semi-arid to arid
habitats in lauzania. Kenya, Uganda, and Somalia.

Conservation status. Cynanchum viminale
subsp. crassicaule cannot be regarded as threatened
because it is comparatively widespread. However,
where it occurs, it is usually not as frequent as most
ol the other subspecies of C. viminale (including
subsp. stipitaceum).

Cynanchum viminale subsp. crassicaule is not
only morphologically distinctive, but also the three
representatives included in a recently undertaken,
but still unpublished, HAIM) analysis form a rea¬
sonably well supported group. The habit of C. vi¬
minale subsp. crassicaule is quite conspicuous
since it is non-twining, arching over the ground for
up to 2 m or scandent in and over bushes. With
glaucous and strictly cylindrical stems that reach
up to I I mm in diameter, its stem morphology dif¬
fers from all other subspecies of C. viminale ; the
stems are the thickest known in all former Sarcos-
temma. I he flowers are also comparatively large for
forme r Sarcostemm a, reaching 15 mm in diameter;
however, most significant are the dimensions of the

Figure 2. Hofotype (Welwitsch 4257. K) ol Sarcostemma
andongense Hiern, referred herein to Cynanchum viminale
subsp. stipitaceum ,

gynostegium (including corona). The enormous
width with respect to Cynanchum or Sarcostemma
of up to 4.5 mm in diameter contrasts to the modest
length, which is ca. 2.7 mm (Fig. ID); this results
in a length/width quotient of around 0.6. In no other
taxon of former Sarcostemma is such a low length/
width quotient, meaning such a flat gynostegium,
known. In 6. viminale subsp. brunonianum (Wight
& Arnott) P. I. Forster the gynostegium is only a
little less flat (length/width quotient around 0.7),
but this is a non-African taxon and its flowers never
reach more than 10 mm in diameter. In all African
taxa of the C. viminale complex, the length/width
quotient lies between 0.77 and 1.0 (Meve & Liede,
unpublished data). In Rumuruti (Laikipia Plateau,
Kenya), where C. viminale subsp. crassicaule oc¬
curs among other former “ Sarcostemma " taxa (C.
viminale subsp. suberosum Meve & Liede, 5. slo-
loniferum Adams & Holland), grazing animals dis¬
dain subspecies crassicaule because of its caustic
latex, while the other taxa are eaten.

The name Sarcostemma andongense Hiern has
been frequently misapplied to C. viminale subsp.

Volume 15, Number 2
2005

Liede-Schumann & Meve
Succulent Cynanchum

323

cross icaule (e.g.. Blundell, 1987: 149, fig. 225;
Schultz & Powys, 1998). However, the name .S', an-
dongense refers to a taxon originally described from
Angola (Fig. 2), which we recognize as a synonym
of C. viminale subsp. stipitaceum (Forsskal) Meve
& Liede (see below). Most obvious, the inflores¬
cences are typically stipitate in subspecies stipita¬
ceum (of. Fig. 2). whereas they are sessile in sub¬
species crassicaule (Fig. 1 B).

Paratypes. KENYA, kitui Dislr.: Mutoma Hills. I).
Camming 393 (UBT, in cult.). Laikipia Distr.: Powys
farm nr. Rumuruti, .S'. Liede & L. hi. Newton 3173 (PPM);
()| Doinyo Losses, /.. E. Newton s.n. (K). Macliakos
Distr.: Lukenya, P. R. 0. Roily 119 (K). N Frontier
Distr.: 01 Lolokwe, M. G. Gilbert 5371 (K). Turkana
Distr.: Lokitaung Gorge, S. Carter & B. Stannard 125 (k).
SOMALIA. Juhhadu House Distr.: Dick's Head (Has
Chiamboni), ,/. Adamson sub P. R. (>. Rally 5933 (K).
TANZANIA. Moshi Distr.: Kilimanjaro, Nanuai f arm , J.
M. Grimshaw 9413 (K). UGANDA. Karamoja Distr.:
Dodoth county, ca. 5 mi. West Loyoro on Koputh and Rom
rd.. II. C. Dawkins 647 (FHO).

Cynanchum viminale L. subsp. stipitaceum

(Forsskal) Meve & Liede, comb. nov. Basion-
ym: Asclepias stipitacea ForsskSl, I I. Aegypt.:
50. 1775. Sarcostemma stipitaceum (Forsskal)
Schult., Syst. Veg. (ed. 1| 6: 116. 1820. Sar¬
costemma viminale (L.) R. Br. subsp. stipita¬
ceum (Forsskhl) Meve & Liede, Bot. J. Linn.
Soc. 120: 32. 1996. TYPE: Yemen. Million, P.
Forsskal s.n. (holotype, C).

Sarcostemma andongense Hiern, Cat. Afr. PL: 689. 1898.
Syn. nov. TYPE: Angola. Pungo Andongo, more el¬
evated rocky places in the praesidium, Dec. 1856,
h. M. ./• Welwitsch 4257 (holotype, K). Figure 2.

Acknowledgments. We thank David Goyder (K)
and Siro Masinde (FA) for help with specimen in¬
formation. SL-S thanks Leu Newton, UM thanks
David Goyder, Siro Masinde, and Chris Whitehouse
for pleasant companionship in the field. For loan of
specimens, we thank the directors of LA. f HO,
and K.

Literature Cited

Blundell, M. 1987. Collins Guide to the Wild Flowers of
East Africa. Collins, London.

Liede, 5. & H. Kunze 2002. Cynanchum and the Cynan-
ch inae (Apocynaceae— Asclepiadoideae) — A molecular,
anatomical and latex triterpenoid study. Org. Divers.
Evol. 2: 239-269.

- & U. Meve 1995. The genus Sarcostemma R. Br.

(Asclepiadaceae) in Madagascar. Bot. J. Linn. Soc. 1 18:
37-5 1 .

Meve, U. & S. Liede 1996. Sarcostemma R. Br. (Ascle¬
piadaceae) in East Africa and Arabia. Bot. J. Linn. Soc.
120: 21-38.

- & - . 2002. Floristic exchange between

mainland Africa and Madagascar: A case study of Apo-
cynaceae- Asclepiadoideae. J. Biogeogr. 29: 865-873.
Schultz, R. & A. Powys 1998. Pinguone, Kenya: Succu¬
lents and their Environment. Schultz Publishing, lees-
dale. VIC, Australia.

Leucocoryne talinensis (Alliaceae), a New Species from Chile

Levi M. Mansur and Mauricio A. Cisternas

Facultad de Agronornfa, Pontificia Universidad Catdlica <le Valparaiso, Camilla 4-d. Quillota,

V region, Chile. levi@entelehile.net

Abstract. A new species from Chile, Leucoco¬
ryne talinensis, is described and illustrated. It is
known from Tali nay and Fray Jorge in Filmin' Prov¬
ince'. from roadsides and adjacent fields. Its chro¬
mosome number and relationship to the most close¬
ly related species are discussed.

Resumen. Se describe e ilustra una nueva espe-
cie de Chile, Leucocoryne talinensis, conocida de
las localidades de Talinay y Fray Jorge (Provincia
de l.imarf. .JO’S), donde crece a orillas de* la carre-
tera y en cam[>os adyacentes. Se da a conocer sn
numero crnmnsomico y se discute su relacion con
la especie mas cercana.

Key words: Alliaceae, Chile, chromosome num¬
ber, Leucocoryne.

The genus leucocoryne Lint I ley (Alliaceae) is en¬
demic to Chile and is comprised of about 15 to 20
laxa (Kahn. 1098). It grows from the Atacama des¬
ert in Region I of Turapaea near the city of Iquique
(20°S) in northern Chile (Pinto, 1999), to the humid
Lake District Region X de Los Lagos of southern
Chile (39°S) (Munoz, 2000).

It appears that the center of diversity for the ge¬
nus lies between tin* Regions of Coquimbo and Val¬
paraiso, specifically between the cities of Valparai¬
so (33°S) to north ol La Serena (29°S) from sea level
to 2800 m a.s.l. In this area approximately 75% of
the described species are found. In addition, this
area has some taxonomically undescribed popula¬
tions (Mansur, 2004; Zoellner. 1072).

I lie genus presents great morphological vari¬
ability, shown especially in floral structure, which
causes difficulties in its taxonomic classification
(Mansur, 2004; Munoz, 2000).

Leucocoryne talinensis L. Mansur & M. Cister¬
nas, sp. nov. TYPE: Chile, l.imarf: Talinay,
cultivated at Ouillota from material collected
I 7 Nov. 1097, along Highway 5, km 355,
30°49'S, 71°33'W, type prepared 10 Aug.
20(H), L. Mansur / (holotype, CONC; isotype,
SI). Figure 1.

Lx affinitate /,. odoratae, fforihus magnis differt. Peili-

Novon 15: 324-326. 2005.

eelli inaequales, 1.3— 3.7 cm longi; perianthium tubo cy-
lindrico et hasi globosa; tc|iala ovali-lanceolata ad apicem
leviler acuminata, 10—16 mm longa et leviter reflexo-coch-
leariformia; staminodia patentia, luteola el cylindrica ad
apicem obtusa 3.4—7 mm longa; ovarium tricarpelare, sty¬
lus brevis et persistens. Capsula oblonga et leviter rostra-
ta.

Herbac ■eous scape 16—30 cm tall: bulbous geo-
phyte, perennial; bulbs tunicate, with strong garlic
odor, creamy-white, globular, 12—21 mm high. 11 —
10 mm diam. covered with grayish white cataphylls.
Leaves 4 to 5, all basal, linear, slightly canalicu¬
late. 12—18 X 1.2— 2.5 cm. Spathes 2. membranous
and dry at anthesis, 1. 6-2.5 X 2. 0-4.0 cm, with a
prominent central vein and 8 secondary veins. In¬
florescence a pseudo-umbel, with (3)4 to 8(0) white
flowers; pedicels unequal, 1.3-3. 7 cm long. Peri¬
anth with a cylindrical tube, dilated at the base,
6.9—1 I X 2.0— 4.5 mm, white, with 6 brown-violet
veins running longitudinally; tepals 6 arranged in
2 series, ovate-lanceolate, acute, unequal. 3.7—8 X
10-16 mm, entire to erose, acute apex, with a
greenish central vein; tepals slightly spatulate and
reflexed; stamens 3, fertile; anthers sessile, 1.50-
1.55 mm long, yellow, adnate to the tube of the
perianth above the pistil; staminodes 3, patent,
fleshy, broadly acute, 3.4-7.0 X 1.0-1. 7 mm.
creamy yellow, at the base of the interior tepals;
tricarpellate ovary, with axial placentation; stigma
capitate-papillate; style persistent and short with 3
septal nectaries at the base ol the ovary. Capsule
oblong, slightly rostrate, 9.5-14.8 X 4. 0-4.8 mm;
seeds black and small. 36.6 ± 1.1 mg 50 ', cren-
ulate, irregularly rounded and append aged.

Habitat and distribution. The new species is
known from Talinay, where it grows along Highway
5 and in adjacent fields with populations of Leu¬
cocoryne purpurea Gay at 250-300 m a.s.l., in al¬
luvial, sandy-loam soils (Rodriguez, 1000). leuco-
coryne talinensis is also occasionally found in the
upper part ol tin* Fray Jorge National Park in Chile.
leucocoryne talinensis was originally collected in
1007 and cultivated in Ouillota, where the type col¬
lection was prepared in 2000.

I he new species is exposed to damaging human
activity, such as plantings of exotic Atriplex species.

Volume 15, Number 2
2005

Mansur & Cisternas
Leucocoryne talinensis from Chile

325

Figure I . Leucocoryne talinensis L Mansur & M. Cisternas. — A. Habit and inflorescence. — B. Flower. — C. Opened
flower showing the stamens. — I). Capsule. — h. Seeds. Drawn from (resh material.

as well as grazing, especially by goats. Its conser¬
vation status may be considered vulnerable.

Etymology. The specific epithet refers to the
geographic origin of the new species.

Leucocoryne talinensis presents phenotypic char¬
acteristics that are stable in regard to the color of
its flowers and the cataphylls of the bulb. However,
it is frequently possible to find individuals within

326

Novon

Talinay populations that vary in the form of the
tepals, some having entire to erose tepals. This
plant was referred to previously as Leucocoryne sp.,
“Talinay” (Araneda et al., 2004; Mansur et al.,
2004), and as L. sp ecotipo Talinay (de la Cuadra
& Mansur, 2004). Its diploid chromosome number
was determined to be 2 n = 18, with an idiogra-
matic formula of 14 metacentric + 2 sub-telocen¬
tric + 2 telocentric (Araneda et ah. 2004).

The new species is somewhat similar to Leiico-
coryne odorata Lindley. However, unlike L. talinen-
sis, L. odorata has a chromosome count of 2// = 10
(Crosa, 1988), filiform staminodes, an inflorescence
with only 2 to 4 flowers, and tepals ovate with an
acute apex. Its bulb is covered with brownish cat-
aphylls versus grayish white in L. lalinensis. Fur¬
ther, /,. odorata is distributed south of 32°S in
Chile, whereas L. lalinensis is found only in the
type locality in Talinay at 30°49'S.

Paratype. CHILE. Coquimbo: Limarf, Fray Jorge.
450 m, Sep. 1972. E. Navas 132389 (CONIC).

Acknowledgments. The authors are grateful to
Rosa Cuaglianone and Roberto kiesling. Institute
de Botanica Darwinion, Argentina, for valuable
comments and suggestions on the manuscript, and
to Roberto Rodriguez, Departamento de Botanica,
Universidad de Concepcion, for providing herbari¬
um material. We also thank Ms. Marlen Olivares
for drawing the figures of Leucocoryne lalinensis and

Mr. Luis Arriagada for assistance in the collection

of specimens.

Literature Cited

Araneda, I... P Salas & I,. Mansur. 2004. Chromosome
numbers in the Chilean endemic genus leucocoryne
(Huilli). J. Amer. Sue. Hurt. Sci. 129: 77— 80.

Crosa, (). 1908. Los eromosomas de nueve especies del
gdnero leucocoryne Lindley (Allieae— Alliaceae). Boh
Invest. N° 17, Fac. Agron. Uruguay.

de la t iuad ra, C. & L. Mansur. 2004. Descripeion de las
primeras etapas del ciclo de vida de Ires genotipos de
leucocoryne sp.: Semilla a bulbo. Agric. T<V. 64(2):
205-212.

Mansur, L. 2004. Life cycle, self-incompatability and ge¬
netic plasticity in the design and color of its (lowers.
Pp. 9-15 in leucocoryne , a Chilean Native Genus and
its Use as a Garden Plant, I" ed. Reimco, Vina del Mar,
Chile.

- , M. Gonzalez, I. Rojas & P. Salas. 2004. Self-

incompatibility in the Chilean endemic genus leuco¬
coryne Lindley. .1. Amer. Soe. Hurl. Sci. 129(6): 836-
838.

Munoz, M. 2000. Consideraciones sobre los gdneros en-
demicos de monocot ileddneas de Chile. Boh Mus. Nac.
Hist. Nat. 343: 16-27.

Pinto, R. 1999. Oasis de Niebla. Una expedicidn botanica
a los cerros costeros de lquique durante “El Nino”
1997. Ograma. Chile.

Rahn, k. 1998. Alliaceae, Pp. 70—78 in k. kuhitzki (ed¬
itor), Flowering Plants, III. Monocotyledons. Lilianae
(except Orchidaceae). Springer- Verlag, Berlin.

Rodriguez, M. 1990. Ceograffa Agricola de Chile, 2“ ed.
Ed. Universitaria, Santiago.

Zoellner, 0. 1972. El genero leucocoryne. Analcs Mus.
Hist. Nat. Valparaiso 5: 9-83.

Nuevas Especies de Weinmannia (Cunoniaceae) para
Costa Rica y Colombia

./. Francisco Morales

Institute) Nacional de Biodiversidad ( I IN Rio), Apto. 22-3100, Santo Domingo de Heredia,

Costa Rica, fmorales@inbio.ac.cr

RESUMEN. Se described Weinmannia cogolloi y W.
vulcanicola (Cunoniaceae), dos nuevas especies de
Colombia y Costa Rica respectivamente. Weinman¬
nia cogolloi es un taxon bastante singular dentro
del genero, por la presencia de inflorescencias pa-
niculadas con ejes secundarios evidentes; auii(|iie
se encuentra algo relacionada con W. latifolia C.
I 'its I y W. macrophylla Kunth, estas ultimas se se-
paran por sus inflorescencias racemosas. Weinman¬
nia vulcanicola se encuentra relacionada con W.
trianaea Weddell, de la que se separa por sus liojas
con mayor numero de folfolos, con la lamina mas
pequena y el raquis alado, asf como por el conspi-
cuo indumento de sus ramitas jovenes.

Abstract. Weinmannia cogolloi and W. vulcani¬
cola (Cunoniaceae), two new species from Colombia
and Costa Rica, respectively, are described. Wein¬
mannia cogolloi is distinct within the genus on ac¬
count of its paniculate inflorescences, with conspic¬
uous secondary axes; it is somewhat related to 1C
latifolia C. Presl and W. macrophylla Kunth, but
these species are easily separated by their racemose
inflorescences. Weinmannia vulcanicola is somewhat
related to W. trianaea Weddell, but it is distin¬
guished by its leaves with more and smaller leaflets,
winged rachis, and the indument of the young stems.

Key words: Colombia, Costa Rica, Cunoni¬
aceae, Rosales, Weinmannia.

Weinmannia b. (Cunoniaceae) es un genero pan-
tropical con cerca de 150 especies, mayormente
distribuidas en el hemisferio sur (Bernardi, 1961,
1963; Bradford, 1998; Stein, 2001). En forma gene-
rid, este genero se distingue facilmente por sus ho-
jas opuestas, simples a pinnadamente compuestas,
con esti'pulas foliaceas y conspicuas, inflorescen¬
cias racemosas, con las (lores dispuestas en fasci¬
cules o subfascfculos (en el neotropico) y frutos
capsulares. Las ultimas revisiones de las especies
neotropicales fueron hechas por Bernardi (1961,
1963). Sin embargo, la ulilidad de estas monogra-
fi'as es relativamente baja, sobre todo por la caren-
cia de descripciones morfologicas completas y pa-

ralelas, asf como la ausencia de una lista completa
de espeefmenes examinados. Por otro lado, varias
secciones que el propuso fueron problematicas,
rlado que algunas fueron invdlidamente pnblicadas
|)or la no designation de un tipo (e.g., sect. Sim-
plicifoliae) o que no han sido apoyadas por analisis
cladfsticos recientes (e.g., Bradford. 1998. 2002).
Por lo tanto, a t raves de este trabajo se siguen los
resultados de Bradford (1998. 2002) que incluyen
a la section Simplicifoliae (sensu Bernardi, 1963),
dentro de la section Weinmannia. Mas reciente-
mente, Marling (1999) realize el tratamiento de Cu¬
noniaceae para la Flora of Ecuador ; trabajo de re-
lativa alta importancia, por el hecho de proveer
descripciones extensas, citacion de espeefmenes
examinados e ilustraciones de todos los taxones, asf
como por clarificar en parte, la confusion taxonomi-
ca presente en el genero. Durante el proceso de
elaboration de las Cunoniaceae para la Flora Me-
soomericana y el Manual de las Plantas de Costa
Rica, dos nuevas especies fueron encontradas. Una
de ellas con hojas simples, es conocida solo del
Departamento de Antioquia, Colombia, siendo bas¬
tante notable dentro del genero por sus inflorescen¬
cias conspicuamente paniculadas, con los ramos la-
terales secundarios conspicuos, mientras que la
segunda, con hojas pinnadas, ha sido conocida por
numerosas coleeciones desde hace mas de 40 aiios
en Costa Rica como Weinmannia trianaea Weddell,
una especie que de otro modo es restringida al SE
de Colombia y Ecuador. Por lo tanto. ambas son
descritas a continuation.

Weinmannia cogolloi J. E Morales, sp. nov.
T1P0: Colombia. Antioquia: Par. Nac. Nat. Las
Orqufdeas, sector Cruces, margen derecha del
Rio Calles. camino de Cruces hacia Venados,
23 Feb. 1989 (fl). A. Cogollo, I). Cardenas &
E. Alvarez 4196 (holotipo, JAUM; isotipo, MO).
Figura 1 .

Haec species Weinmanniae latifoliae affmis seel foliis
majoribus et inflorescentiis paniculatis differt. 14. macro-
phyllae aflinis sed (oliis coriaceis, supra glabris et inflo¬
rescentiis paniculatis differt.

Novon 15: 327-331. 2005.

328

Novon

Figura 1. Kotugraffa. ramita (lorffera de Weinmarmia cogolloi J. F. Morales (/l. Cogollo , I). Cdrdenas & E. Alvarez
419 6, MO).

* . 1 1 1 n till il ii m i i|ii 1 1 1 1 ii ill m ) i ntai ii ii 1 1 nil i n . « 1 1 1| ( 1 1 1 pii II

11 21 21 41 5l 6l 7l

Arboles 10—18 m, los tallos conspicuamente an-
gulados y aplanados en tallos jovenes, subteretes
con la eda<l. densamente tomentosos o tomentulo-
sos. cl indumento cafe. Hojas simples, la lamina
12-20 X 0.5—14 cm. anchamente eh'ptica, carta-
cea. agudas y mucronadas apicalmente, anclia-
mente agudas basalmente, serradas marginalmente.

esparcidamente puberulentas adaxialmente, den-
samente puberulentas abaxialmente, no revolutas
marginalmente, pecfolos 19—22 mm. tomentosos;
estipulas 10—12 mm, anchamente ovadas. eonspi-
cuamente cordadas basalmente, caducas. Inflores-
ceneias panfculas pareadas, conspicuamente rami-
ficadas, 9—17 cm de longitud, los ejes laterales de

Volume 15, Number 2
2005

Morales

Nuevas Especies de Weinmannia

329

1—3.1 cm dc largo, el raquis esparcidamente hir-
suto, el indumento amarillento, diminuto, ca. 0.5
mm de largo. Flores ea. 1.5 mm de longitud, los
pedicelos 1.5— 2.5 mm de largo; Idbulos del caliz
0.6— 0.8 mm de largo, cortamente acuminados api-
calmente, glabrescentes; petalos ca. 1.5 mm de lar¬
go; ovario 1.5—2 mm de largo, glabro. Frutos in-
maduros ca. 4 mm, glabros.

Distribution, ecologia y habitat. Endemica al
noroeste del Departamento de Antioquia, Colombia,
donde crece en bosques muy hiimedos entre los
800 y 1400 m. Flores y frutos inmaduros hail sido
recolectados entre febrero y marzo.

Weinmannia cogolloi se encuentra relacionada
con W. latifolia C. Presl y W. macrophylla Kunth,
compartiendo caracteres tales como hojas simples,
con laminas grandes y desarrolladas, asf como in-
llorescencias usualmente pareadas. Sin embargo,
W. cogolloi se separa facilmente de ambas por sus
inflorescencias paniculadas, con conspicuas ramas
laterales (vs. inflorescencias racemosas, sin rami-
ficaciones). Ademas, 11. latifolia tiene hojas con las
laminas muelio mas pequenas ((5— )6— 8(— I 1 ) cm vs.
12—20 cm) y W. macrophylla tiene laminas foliares
coriaeeas y usualmente glabras abaxialmente (vs.
laminas cartaceas y puberulentas en W. cogolloi ),
con las margenes muy esparcidamente crenado-se-
rratlas (vs. conspicua y densamente serradas).

Fs importante liacer notar (|iie la presencia de
inflorescencias paniculadas dentro del genero, con
ramificaciones o ejes secundarios conspicuos es
algo inusual, va que en forma general, practica-
mente todas las especies tienen inflorescencias ra¬
cemosas. Bradford (1998) realizd un analisis deta-
llado de la arquitectura tie la inflorescencia en el
genero. determinando que debido a la incapacidad
de aplicar en forma satisfactoria algunas de las ter-
rninologfas de lipos de inflorescencia anteriormente
propuestos (e.g., Briggs & Johnson, 1979; Weber-
ling. 1989), las inflorescencias en Weinmannia de-
bfan catalogarse como racimos, aunque si bien, va-
rios tipos de variaciones podi'an estar presentes. De
esta forma, la descripcion de Weinmannia cogolloi,
con inflorescencias claramente paniculadas, es algo
bastante raro dentro del genero y que sale fuera de
contexto de la clasifieacion propuesta j>or Bradford
(1998).

El epfteto de esta especie esta dedicado a Alvaro
Cogollo, director del Herbario Joaquin Antonio Uri¬
be (JAIIM), Medellin, Coloml )ia, en reconocimiento
a su trabajo en pro de la Flora de Colombia y por
agradecimiento a su apoyo personal v facilidades
brindadas en mis visitas a Medellin en los aims
2001, 2002 y 2003.

Fa ra tipos. COLOMBIA. Antioquia: Crrao, Bar. Nac.
Nat. I.as Orquideas, sect. Cabana de Calles, Quebrada La
Agudelo, 31 Mar. 1992. I). Cardenas & E. Alvarez 3173
(JAUM); Bio Calles, 9 Die. 1993, A. Cogollo el al. 7H62
(INB. JAUM, MO); seel. Calles, Quebrada l.a Agudelo, 5
Jill. 1991, E. Ramirez 4134 (JAUM), 8 Jul. 1991, E. Ra¬
mirez 4307 (JAUM).

Weininannia vuleanicola J. E. Morales, sp. nov.
TIPO: Costa Rica. Alajuela: Greeia, Par. Nac.
Volcan Poas, sendero a La Laguna Botos, 28
Sep. 1999 (fl). ,/. F. Morales 7 1 R3 (Imlotipo.
INB; isotipos, K, MO). Figura 2.

Haee species Weinmanniae trianaeae afiinis sed rhaehi-
dibus alatis et foliolis minoribus differt. W. polyphyllae
affinis sed rhachidibus angustioribus, inflorescentiis ag-
glomeratis et bracteis florigeris minoribus differt.

Arbustos 2—8 m, los tallos conspicuamente an-
gulados y aplanados en tallos jovenes, subteretes
con la edad. densamente tomentosos o tomentulo-
sos. el indumento ferrugfneo a matron ferrugfneo.
algo glabrescentes con la edad. Hojas compuestas,
lamina de los folfolos (0.6)0. 8— 1.5 X 0.3— 0.8 cm.
elfptica, los folfolos (11)15 a 25(29), subcoriaceos.
la mayorfa obtusos a redondeados apicalmente, mas
raramente agudos, obtusos a redondeados basal-
mente, crenado-serrados, usualmente glabros, solo
con pelos a lo largo del nervio central abaxial¬
mente, variadamente eoncavos y revolutos margi-
nalmente, entrenudos del raquis alados 1.5— 2.5(3)
mm de ancho, el raquis densamente tomentoso o
tomentuloso abaxialmente, el indumento ferrugfneo
a matron ferrugfneo; peciolos 4—6 mm. densamente
tomentosos, usualmente no alados; estfpulas 6— 7 X
8—9 mm, anchamente ovadas. Inflorescencias ra¬
cemosas, usualmente pareadas, 6—12 cm de longi¬
tud, el raquis y los pedicelos esparcidamente
puberulentos, el indumento ferrugfneo o marron-
ferrugfneo, diminuto. Flores 2.5—3 mm de longitud.
los pedicelos 2—4 mm de largo; lobulos del caliz
0.7-1 mm de largo, cortamente acuminados api¬
calmente, glabros o glabrescentes; petalos ca. 2 mm
de largo; ovario 1 .5—2 mm de largo, glabro. Frutos
4—5 mm, glabros.

Distribucidn, ecologia y habitat. Esta especie
esta restringida por el momento a Costa Rica, don¬
de crece en bosques nubosos, areas de sucesion
volcanica, robledales y vegetacion secundaria re¬
lacionada, entre 2300 y 3000 m. Es probable que
este taxdn se encuentre en el norte de Panama,
especfficamente en las estribaciones de la Cordille¬
ra de Talamanca. Especfmenes con flores ban sido
recolectados entre febrero y octubre. Especfmenes
con frutos se recolectaron entre agosto v diciembre.

Paratipos. COSTA BICA. Alajuela: Volcan I’oas. I I

330

Novon

Figura 2. Fotograffa, ramita florffera de Weinmannia vulcanicola J. F. Morales (J. F. Morales 7183, INB).

Sep. 1965, I,. Bernardi 10535 (K (j, MO), 21 Abr. 1968,
C. Cordoba 018 (CR), 5 Mayo 1991, F. Dobbeler & G.
Dauphin 5488 (CR), 3 Mar. 1999, M. Gargiullo 81 5 (CR).
18 Mayo 1965, W. Hat he way 1330 (CR), 19 Ago. 1961,
A. Jimdnez 103 (CR, F), 25 Oct. 1964, F. Little G.
Budowski 20020 (CR), 20 Ago. 1965, T. McDaniel 6 718
(MO), 11 Nov. 1967, C. Orozco 13 (CM), 18 Jun. 1971, G.

Proctor 32468 (F. MO), 19 Jul. 1976, ./. Solomon 5320
(MO), 13 Jul. 1962, G. Webster, K. Miller & L Miller
12280 (F. MO), I Sep. 1968. K. Wilbur & D. Stone 10650
(DUKK. F. MO); Monteverde, 30 Nov. 1976, F Dryer 1046
(F); Fila Volcdrt Viejo, San Carlos, Feb. 1986, ./. Gdmez-
L. 11032 (CR, F, USJ); O de Vara Blanca, 21 Fne. 1968,
II. Wilbur A I). Stone 0837 (DUKK. F. MO). Heredia:

Volume 15, Number 2
2005

Morales

Nuevas Especies de Weinmannia

331

Vara Blanca, 31 Die. 1973, F. Almeda, M. Flowers & R.
Wyatt 2206 (DUKE. E); Par. Nac. Braulio Carrillo, Vulcan
Baeva. 13 Ago. 1993, II. Iloyle, ,/. Bradford & K. Sparks
2360 (MO), 14 Ago. 1993, II. Hoyle, A. Iloyle, ./. Bradford
<& K. Sparks 2382 (CR, MO); entre Sacramento y Laguna
<le Barva, Volcan Barva, 12 Abr. 1975,./. Utley & K. Utley
2027 (F). Limon: Talamanca, Sabanas de Durika, Rio
Kuk, 1 km aguas debajo de la conlluencia de los Rios Uk
y Kuk. 20 Oct. 1989, A. Chacon 584 (CR, F, [Ml. MO);
Cordillera de Talamanca, Cerros Tararia. 10 Sep. 1984. C.
Davidse, G. Herrera & M. Grayum 28848 (INB. MO); Par.
Int. La Amistad, Cerros Tararia, Valle del Silencio, 1 Abr.
1997, F. Quesada et al. 1924 (INB).

Weinmannia vulcanicola ha sido tradicional-
mente conocida en Costa Rica como W. trianaea,
pero el analisis de las poblaciones del sur de Co¬
lombia y Ecuador con respecto a las de Costa Rica,
asf como el estudio del ejemplar tipo de W. trianaea
(Triana s.n., RM, K, P, W) determino que el grupo
presente en esle ultimo pais tiene sufieientes carac-
teres distintivos, como para ser tratada como un
taxon aparte. Weinmannia vulcanicola diliere de IE
trianaea por sus hojas con el raquis alado a lo largo
de su extension (vs. alado solo en el tercio distal)
y con muchos nuts folfolos ((11)15 a 25(29) vs.
(9)11 — 17(20)), estos ultimos con la lamina mucho
mas pequena y angosta ((6)8—15 X 3—8 mm vs.
(17— )22— 30 X (7— )9— 14 mm) y la presencia de un
indumento ferrugfneo o cafe-ferrugmeo en los tallos
jovenes. Weinmannia vulcanicola se encuentra re-
laeionada con W. polyphylla Moricand ex Seringe.
olra especie restringida al noroeste de Sur America
eon quien comparte hojas con similar numero y di¬
mension de folfolos; sin embargo, W. vulcanicola
diliere por sus hojas con las alas del raquis mas
angostas, infloresceneias densamente aglomeradas
(vs. laxas), y fasciculus con las brae leas lineares y
rapidamente deciduas, ca. 1 X 0.5 mm (vs. brae-
teas persistentes anchamente ovadas o elfpticas,
1.5-2 X 1-1.5 mm).

Estas tres especies pueden ser separadas por la
siguiente clave:

Clave para las Especies Relacionadas con Weimiax-

MA VULCANICOLA EN COSTA RlCA, Cot.OMUIA Y ECUADOR

la. Raquis de las hojas no alarlo o solamente alado
en el tercio terminal; lamina de los folfolos la-
terales (17)22—30 X (7)9—14 mm . . . . 1L trianaea

lb. Raquis de las hojas alado en toda su extension;

lamina de los folfolos laterales (6)8—15 X 3—8
mm.

2a. Tallos jovenes esparcidamente hirsutulos,
glabrescentes con la edad, el indumento
amarillento, no ferrugfneo; infloresceneias
con los fascfculos laxamente distribuidos;
bracteas persistentes aun en fructificacidn,
anchamente ovadas o elfpticas, 1.5—2 X l—

1 .5 mm; S de Colombia y Ecuador .

. W. polyphylla

2b. Tallos j6venes tomentosos o tomentulosos,
nunca glabrescentes, el indumento ferrugf¬
neo o eale-lcrrugfneo; infloresceneias con
los fascfculos densamente aglomerados;
bracteas rapidamente deciduas, lineares, ca.

1 X 0.5 mm; Costa Rica . IT vulcanicola

Agradecimientos. Quiero agradecer a los cura-
dores y personal de los herbarios CR, DUKE, F, G.
HUA, JAUM, MO, NY, MEDEL, US, USJ, por per-
mitirme el uso de sus colecciones, asf como el per¬
sonal del Herbario de la Universidad de Antioquia
(1IUA) y del Jardfn Botanico Joaquin Antonio Uribe
(JAUM), por las facilidades logfsticas ofrecidas en
mis visitas a Medellin, Colombia. Finalmente, se
agradece la revision crftica del manuserito por par¬
te de un revisor anonimo y de Jason Bradford; esle
ultimo suministro ademas refereneias bibliogralieas
e importantes comentarios sobre la clasificacion
seccional en Weinmannia.

Literatura Citada

Bernardi, L. 1961. Revisio generis Weinmanniae. Pars I:

Sectio Weinmanniae. Candollea 17: 123—189.

- . 1963. Revisio generis Weinmanniae. Pars II: Sec¬
tio Simplicifoliae. Candollea 18: 285—334.

Bradford, .). C. 1998. A cladistie analysis of species
groups in Weinmannia (Cunoniaceae) based on mor¬
phology and inflorescence architecture. Ann. Missouri
Bot. Card. 85: 565—593.

- . 2002. Molecular phylogenetics and morphologi¬
cal evolution in Cunonieae (Cunoniaceae). Ann. Mis¬
souri Bot. Card. 89: 491-503.

Briggs, B. & L. Johnson. 1979. Evolution in the Myrta-
eeae. Evidence from inflorescence structure. Proc. Linn.
Soc. New South Wales 102: 157-272.

Hailing, W. 1999. Cunoniaceae. En: G. darling & L. An-
dersson, FI. Ecuador 61: 1—74.

Stein, B. 2001. Cunoniaceae. En: W. I). Stevens, C. Ulloa
Ulloa, A. Pool & O. M. Montiel (editores). Flora de
Nicaragua. Monogr. Syst. Bot. Missouri Bot. Card. 85:
718.

Weberling, F. 1989. Morphology of Flowers and Inflores¬
cences. Cambridge Uuiv. Press, Cambridge.

Una [Nueva Especie de Werauhia (Bromeliaceae) para Costa Rica

./. Francisco Monties

Instituto Nacional do Biodiversidad (IN Bio), Apto. 22-3100, Santo Domingo do Heredia,

Costa Kira. fmorales@inl)io.ae.er

Resiimkn. Sr describe e ilustra Werauhia anitana
(Bromeliaceae), una nueva especie de Costa Rica,
con infloresceneias dfsticas y brdcteas punteado-
verrucosas, densamente imbricadas aun despues de
la antesis. Se discuten las relaciones con los taxo-
nes relacionados (i.e., W. osaensis, W. viridiflora).

Ahsthact. Werauhia anitana (Bromeliaceae), a
new species from Costa Rica with distichous inflo¬
rescences, and punctate-verrucose and densely im¬
bricate bracts (after anthesis), is described and il¬
lustrated. Relationships with related taxa (i.e., W.
osaensis , W. viridiflora) are given.

Key words: Bromeliaceae, Central America,
Costa Rica, Neotropics, Werauhia.

Cas Bromeliaceae estan representadas en Costa
Rica por aproximadamenle 210 especies y 16 ge-
neros (Morales, 2003c). El genero Werauhia, segre-
gado de Vriesea por Grant (1995), es el mas nu-
meroso con cerca de 80 especies, muchas de el las
restringidas a Costa Rica y Panama, tin intensive
trabajo de campo en los ultimos 15 anos, Uevado a
calm principalmente por el Instituto Nacional de
Biodiversidad (INB) e investigadores asociados del
Missouri Botanical Garden (MO) y del Museo Na¬
cional de Costa Rica (CR), ha incrementado en for¬
ma significativa el numero de especies de brome-
lias conoeidas en forma previa para el pais, con la
subseeuente deseripcion de varias novedades
(Grant iK Morales, 1996; Luther & Kress, 1996;
Morales, 1999; Spencer & Smith, 1991). Cuantlo la
exploracidn Imtanica de Costa Rica empezo a fi¬
nales del siglo IXX y principios del siglo XX, la
region del drea del Paso de la Palma, loealizada al
norte de San Jose entre los volcanes Irazu y Barba,
lue una de las localidades mas colectadas, sobre
todo por su facilidad de acceso desde San Jose y
la presencia de extensas zonas boscosas. Muchas
Bromeliaceae (ueron colectadas por Carlos Werckle
de esta area, especialmente rica en bromelias y epi-
fitas en general (e.g.. Ericaceae, Orchidaceae) y al-
gunas de ellas atin son conoeidas por muv pocas
colecciones (e.g., 1L singulijlora (Mez vN Werckle)
J. R. Grant).

Durante varies viajes de campo a esta area entre

1999 y 2003, eon el objetivo de estudiar y local izar
algunas de estas especies de bromelias pobremente
conoeidas, una nueva especie de Werauhia sect.
Werauhia fue eneontrada, no colectada previamente
por Werckle. En forma general, las especies de
Werauhia de Costa Rica pertenecientes a esta sec-
cion y envueltas en el eomplejo de W. gladioliflora
y W. viridiflora (e.g., U. acuminata (Mez & Werckle)
J. R. Grant, IE bicolor (L. B. Smith) J. R. Grant)
son numerosas y su taxonomfa ha sido poco estu-
diada. |uies las colecciones disponibles son en mu-
chos casos muy escasas y fragmentarias. El estudio
extensivo de las poblaciones en las localidades de
los tipos en los ultimos anos ha llevado al reeono-
cimiento de especies no aceptadas por Grant
(1995), ya que presentaban caracteres bien distin-
tivos (e.g., forma y textura de las brdeteas, preferen-
cias ecoldgicas y altitudinales) que no habfan sido
estudiados previamente en detalle (Morales,
2003b).

Esta nueva especie muestra cercanas afinidades
con Werauhia osaensis (J. F. Morales) J. F. Morales
y W. viridiflora (Regel) J. R. Grant, ya que esle
grupo de especies comparten infloresceneias sim¬
ples, no ramificadas, con bracteas florales de lon¬
gitudes similares. Por lo tanto, es descrita a con¬
tinuation y se discuten las relaciones con estas
especies.

Werauhia anitana J. F. Morales, sp. nov. I IPO:
Costa Rica. San Jose: Paso de la Palma, cami-
no a Bajo Ea Hondura, cerca de las lecherfas
en ruinas. 1300—14001) m. 2 Nov. 2002 (fl), ,/.
F. Morales & R. Abarca 9074 (holotipo, INB).
Figura I .

A Werauhia osaensi, cui similis, foliis angustioribus,
scapo decurvato. et bracteis florigeris panctalis verrucosis
differt; a W. viridiflora. cui uf finis. bracteis florigeris im-
bricatis punctatis verrucosis et [>etalis majoribus differt.

Plantas acaulescentes, con una altura en flora-
cion de 30 a 40 cm. Hojas rosuladas, ± erectas
pero algo reflexas apicalmente, 1 1—38 cm de lon-
gitud, densa a moderadamente punteado-lepidotas,
eon las puntuaciones negras, en especial abaxial-
mente, delgadas, verdes, verde-moradas o total -

Nov ON 15: 332-334. 2005.

Volume 15, Number 2
2005

Morales

Werauhia anitana para Costa Rica

333

mente moradas por ambas superficies; base de la
lamina anchamente eliptica, (4)9-1 1 X (2.2)3 cm,
palida, usualmente concolora con las laminas, den-
samente negro punteado lepidota, en especial
almxialmente; lamina de las hojas ligulada, atenu-
ada o aguda, 7-27 X 1-1.3 cm. Escapo decurvado,
38-41 X 0. 3-0.4 cm, glabro, diminutamente cafe
papilado; bracteas del escapo erectas, 2.5— 3.5 cm
de longitud, mucho mas largas ([ue los entrenudos,
densamente imbricadas, cafe oscuras basalmente,

algo mas claras distalmente, densamente punteado-
verrucosas. Inflorescencias erectas, simples, 1 1 — 13
cm de longitud; bracteas Morales anchamente ova-
das, agudas, 4. 1-4.5 X 2. 2-2. 5 cm, relativamente
delgadas, densamente imbricadas y eonspicua-
mente excediendo los sepalos. de color cafe canela
en la antesis, los hordes ligeramente mas clams,
ecarinadas, conspicuamente punteado-verrucosas.
Flores 5 a 6, con los pedicelos gruesos, 2—5 mm
de longitud, erectos, esparcidamente papilados; se-

334

Novon

palos 1.5— 1.7 cm <lc longitud, credos, angosta-
mente obovados a obovado-elfpticos, coriaeeos,
caf£ claros. Corola blanco-erema o crema, los pe¬
talos obovado-elfpticos, 35—40 mm de longitud, ob-
tusos, con las dos escamas basales enteras; estigma
bianco. Cdpsula desconocida.

Distribut ion y habitat. Werauhia anitana es en-
demica a Costa Rica, donde se encuentra restrin-
gida al Paso La Palma, en bosques montanos nu-
bosos o bosques ventosos premontanos y vegetat ion
perturbada asociada en elevaeiones entre 1350—
1500 m. I* loracidn ocurre en junio y noviembre.

Etimologla. Esta ('specie esta nombrada en
honor a Anita Cooper, en reconocimiento de su ex-
celente y extensive trabajo en ilustraciones de
acuarela para las gufas de campo Bromelias de
Costa Rica (Morales, 2000) y OrquCdeas, Cactus y
Bromelias del Bosque Seco de Costa Rica (Morales,
2001), que las realize mientras estuvo en Costa
Riea en 1996.

Siguiendo la clasifieacihn infragenerica propues-
ta por Grant (1995), Werauhia anitana debe ser
incluida en la st*ct. Werauhia, la que se caracteriza
por sus inflorescencias simples o compuestas, pero
entonees eonspicuamente bipinnadas <> tripinnadas.
Werauhia anitana esta cercanamente relacionada
con W. osaensis (J. K. Morales) J. F. Morales, com-
partiendo caracteres tales como inflorescencias
simples, dfsticas, no ramificadas, con las bracteas
Morales densamente imbrieadas. Sin embargo, W.
anitana es facilmente distinguida por sus hojas eon
la lamina mas angosta (1 — 1.3 cm), escape decur-
vado y bracteas densamente punteado-verrucosas,
las euales son de un cafe canela en la antesis, en
contraposit ion con W. osaensis, que tiene hojas con
la lamina mds ancha ((1.4-)2. 5-3.8 cm), escape
erecto y brdeteas verdes o verde-canela en la an¬
tesis, lisas a levemente rugulosas. Atlemas, ambas
espeeies tienen prefereneias ccologicas diferentes:
mientras W. anitana crece en bosques montanos
nubosos o bosques ventosos premontanos a eleva-
ciones de 1350—1500 m, W. osaensis se encuentra
en bosques muy humedos, entre 0—700 m. Werau¬
hia osaensis fue originalmente descrita en el genero
Vriesea (Morales, 1999), pero dado que las Mores
de esta especie tienen las escamas basales de los
petalos enteras y t'n forma de lengua, asf como una
coloracidn blanca o crema, fue recientemente trans-
ferida a Werauhia (Morales, 2003a).

Werauhia riridiflora (Regel) .1. R. Grant se puede
eonfundir con esta nueva especie, ya tjue ambas
tienen Idminas foliares de dimensiones y formas
similares, inflorescencias simples, no ramificadas y

prefereneias ccologicas similares. Sin embargo, la
principal diferencia se encuentra cn la disposit ion
y textura tie las bracteas Morales, ya que mientras
en W. viridiflora estas son levemente imbrieadas y
usualmente lisas, en W. anitana estan densamente
imbrieadas, con la superficie externa punteado-ve-
rrucosa. Atlemas, his (hires tie W. anitana tienen
petalos mas grandes en relat ion a W. riridiflora
(3.5—4 cm vs. 2.4— 2.6 cm).

Paratipos. COSTA RICA. San Jose: Paso de la Pal¬
ma, ruinas de las leclierfas, 12 Die. 2002, Morales 8919
(INB), 10 Jim. 2003, Morales & Iddrraga 9182 (INB).

Agradecimientos. Ouiero agradecer a Michael
Grayum por sus eomentarios de una version previa
en ingles tie algunas partes de este manuscrito.
Esta publicat ion fue finaneiada por la Asistencia
Holandesa para el Desarrollo (NEI)A) a (raves del
proyecto “Desarrollo del Conocimiento de la Bio-
diversidad y Uso Sostenible en Costa Rica” con-
ducido por el Institute Naeional de Biodiversitlad
(IN Rit>). Atlemas, la investigacion fue posible gra-
cias al eonvenio de cooperat ion entre cl Ministerio
de Ambiente y Energfa (M1NAE) e IN Rio con el
fin tie completar cl Invcntario Nat ional tit' Biodi-
versidad en Costa Rica.

Literatura Citada

Grant, J. R. 1995. The resurrection of Alcanlarea and Wer¬
auhia, a new genus. Trap. Subtrop. Pflanzenwelt 91: 7-
57.

- <X J. I. Morales. 1996. Bitcairnia calcicola (Bro-

ineliaceae), a new species from the tropical dry forest
of Costa Rica. Novon 6: 366—369.

Luther, H. & J. Kress. 1996. Two overlooked species of
Guzmania (Bromeliaceae) of the species-complex Mas-
sangea from Central America. Brittonia 48: 91-95.
Morales, J. E 1999. Scis nuevas espeeies de Vriesea sect.
Xiphion (Bromeliaceae: Tillandsoideae) para Costa
Rica. Novon 9: 401-406.

- . 2000. Bromelias de Costa Rica. Instituto Nacio-

nal dc Biodiversitlad, Santo Domingo de Heredia, Costa
Rica.

- . 2001. Orqufdeas, Cactus y Bromelias del Bosque

Seco de Costa Rica. Instituto Naeional de Biodiversi-
dad, INBio, Santo Domingo de Hcretlia. Costa Rica.

- . 2003a. Nuevas combinaciones y un nuevt) nom-

bre en las Bromeliaceas th' Costa Rica. Polibot&nica 15:
100-103.

- . 2003b. New combinations in Werauhia (Brome¬
liaceae) from Costa Rica. Lundiana 4: 65.

- . 2003c. Bromeliaceae. Pp. 297—375 en B. E.

Hammel, M. H. Grayum, G. Herrera Nr N. Xamora (eth-
tores). Manual de Plantas de Costa Rica, Vol. II: Gim-
nospermas y Monocotileddncas. Monogr. Syst. Bot. Mis¬
souri Bot. Gard. 92.

Spencer, M. & L. B. Smith. 1991. Bitcairnia funhiae, a
new species from Costa Rica. .1. Bromeliad Soc. 41:
214.

Una Nueva Especie tie Sphyrospermum (Ericaceae) para

Costa Rica

./. Francisco Morales y Alexander Rodriguez

Instituto National cle Biodiversidad (INBio), Apto. 22-3100, Santo Domingo de Heredia,
Costa Rica. fmorales@inhio.ac.cr; arodrig@inbio.ac.cr

R HSU MEN. Se describe e ilustra Sphyrospermum
linearifolium (Ericaceae), mi taxon endemieo a
Costa Rica, bastante singular dentro del genero por
sus liojas lineares y hdbito colgante y se comenta
su relation con otras especies dentro riel genero.

ABSTRACT. Sphyrospermum linearifolium (Erica¬
ceae), a new endemic species from Costa Rica, is
described and illustrated, and its relationships are
discussed. The taxon is distinctive within the genus
because of its linear leaves and scandent habit.

Key words: Costa Rica, Ericaceae, Sphyrosper¬
mum.

El genero Sphyrospermum Poeppig & Endlicher
esta representado por cerca de 22 especies. distri-
buidas desde Mexico a Suramerica y las Islas del
Caribe (Luteyn, 1996). Se puede reconocer por su
habito arbustivo, liojas con la venacion usualmente
inconspicua. flores con el pedicelo no articulado al
hipanto, el pedicelo en la mayoria de los casos re-
lativamente largo y filiforme, con dos bracteolas,
hipanto terete, corola cortamente tubular (general-
mente menos de 10 mm de longitud), estambres de
similar tamano entre si, anteras dehiscentes a
traves de aberturas ovaladas e introrsas, ovario fn-
fero y frutos abayados (Luteyn. 1991). En Costa
Rica, algunas especies pueden llegar a ser muy
comunes en eiertas areas, sobre todo en indigenes
y paredones alterados de caminos v carreteras. En
estas condiciones began a ser bastante dominantes,
junto con otras Ericaceas, tales como Disterigma
humboldtii (Klotzsch) Niedenzu y Gaultheria erecta
Ventenat.

En el proximo tratamiento del genero Sphyro¬
spermum para el pais (Luteyn et al., en prep., Ma-
nual de Plantas de Costa Rica), se reconocen cinco
especies, una de ellas endemica y restringida a las
Cordilleras de Tilaran y Talamanca. La exploracibn
de eiertas zonas remotas e inexploradas en la ver-
liente Atlantica del Volcari Turrialba (Cordillera
Central), produjo como resultado el hallazgo de una
notable especie de Sphyrospermum. Esta planta se
caracteriza por su habito colgante y liojas estricta-

mente lineares, este ultimo caracter no reportado
dentro del genero liasta el momento, lo cual de
cualquier manera, no es algo comun en otros ge-
neros de la familia (e.g., Anthopterus Hooker, Cav¬
endish ia Lindley).

Por lo tanto, se brinda una clave para todas las
especies del genero en Costa Rica, asf como la des-
cripcidn de este nuevo taxon.

Clave para las Especies de Sphyrosi’Ermi i/ en Costa
Rica

la. Hojas con la lamina linear .

. . . S. linearifolium Al. Rodriguez & J. F. Morales

lb. Hojas con la lamina de diversas formas, pero
nunca linear.

2a. Pedicelos 1-3 mm; corola rojiza a rojizo-ro-
sada, 7-10 mm de longitud, campanulada

. S. dissimile (S. F. Blake) Luteyn

2b. Pedicelos 5—25 mm; corola blanca a blanco-
rosada, 2-9 mm de longitud, urceolado-ci-
lfndrica, subcilfndrica a cilfndrica.

3a. Flores extendi^ndose mas al l«S de las

hojas S. buxifolium Poeppig & Fndlicher
3b. Flores no extendiendose mas alia de las
hojas.

4a. Corola (4— )5— 7(— 9) mm; pedicelos

15—25 mm . . . S. cordi folium Bentham
4b. Corola 2-3.5 mm; pedicelos 7-15
mm.

5a. Lamina foliar 1.5—4 cm de lar¬
go; corola 3-3.5 mm; zonas ba-
jas en la Peninsula de Osa en
el Pacffico Sur. 100—500 m . .

. S. ellipticum Sleumer

5b. Lamina foliar 4.5-6 cm de lar¬
go; corola 2—2.3 mm; eleva-
ciones medias en las cordille¬
ras de Guanacaste, Tilardn y
Talamanca, 850-1500 m . .
. S. standleyi A. C. Smith

Sphyrospermum linearifolium Al. Rodriguez A
J. F. Morales, sp. nov. TIPO: Costa Rica. Li-
mon: Altos de Pascua, cabeceras Quebrada
Linda, 1 Abr. 2001 (II, fr). J. F. Morales 7823
(holotipo, INIL isotipos, CR, k, MY, MO). Li-
gura 1 .

Ab aliis speciebus Sphyrospermi foliis linearibus distin-
guitur.

Novon 15: 335-337. 2005.

336

Novon

Figure 1. Sphyrospermum linearifolium Al. Rodrfguez iK J. F. Morales (./. F. Morales 7823, holotipo). — A. Ilabito. —
IF I .amina de la lioja. vista adaxial. — C. Lamina de la hoja, vista abaxial. — I). Caliz y pedicelo, mostrando el detalle
de las braetbolas. K. (ailiz v corola. — F. (.aliz, coo la corola removida. — (». Corola abierta, mostrando la posieibn
de las anteras. — II. Anteras, vista lateral, dorsal v ventral, mostrando las aperturas de I os tubulos.

[Manias epffitas y eolgantes; runias delgadas, gla-
brescentes, longitudinalmente estriadas; ramitas de
eoloracidn canela a canela-grisacea, diminuta v es-
pareidamente puberulentas. Pecfolos 1-2 mm de
largo; hojas alternas. la lamina linear. 3.5-14.4 X
0.2— 0.8 cm, el dpiee angostamente agudo, la base
angostamente obtusa, el margen entero, usualmente
revoluto en las liojas viejas, coriaeeas a subeori-
aeeas. glabreseentes adaxialmente, la vena central
muy esparcidamente pildsula, la lamina diminuta v
esparcidamente estrigulosa abaxialmenle. ineonspi-
cuamente 2— 3-plinervadas. Flores axilares. solita-
rias; pedilnculo delgado, eontinuo con el hipanto.

1. 1-3.3 cm de largo, glabro; bracteas 2. ca. 0.65
mm de largo, lineales. deeiduas; hipanto subglo-
boso. terete a subterete, ca. 1.2 mm en diamelro,
glabro; limbo tie I caliz suberecto, 0.75—1 mm (in-
cluyendo los lobulos) de largo, los lobulos 4, de
0.2— 0.6 mm de largo, deltados, agudos, glabros;
corola urceolada. rosada a blaneo-rosada, el tubo
ca. 5 mm de largo, ca. 3 mm en diametro, glabro,
los lobulos de la corola triangulares, ca. 0.65 mm
de largo, agudos; estambres 5, iguales. levemente
mas cortos que el tubo de la corola, los filamentos
ca. 1.75 mm tie largo, glabros a esparcidamente
vilosos, las anteras lisas, ca. 2.3 mm de largo, los

Volume 15, Number 2
2005

Morales & Rodriguez 337

Sphyrospermum linearifolium de Costa Rica

tubulos abriendose por aberturas ovaladas e intror-
sas; ovario inferior, el estilo filiforme, casi tan largo
como la corola, el estigma truncado. Hava globosa
a subglobosa, 5—8 mm en diametro. glabra, blan-
quecina o morado-blanqueeina.

Distribution. Endemiea a Costa Itica. donde ha
sido reportada de la vertiente Atlantiea de la Cor¬
dillera Central, en las faldas de los voleanes Barva
y Turrialba, entre los 700—1500 m.

Fenologia. Especimenes eon flores ban sido re-
colectados en marzo y abril. Colecciones eon frutos
fueron hechas en marzo, abril y julio.

Sphyrospermum linearifolium |)nede ser reeono-
eida fdeilmente dentro del genero por sns hojas li-
neares, usualmente revolutas marginalmente, asi
eorno por su habito eolgante. En forma general las
demas especies en el pais tienen hojas de formas
variadas, pero nunca tan angostas para dar nna sen-
saeidn de ser lineares. Asimismo, el habito eolgante
es nn earaeter bastante irregular, ya que es eomun
que el resto de los taxones en el pais tengan habito
arbustivo o rastrero.

I.a vertiente atlantiea del Volean Turrialba, as i
roimi muehas zonas aledanas (Bonilla de Turrialba,
El Destierro) en las provineias de Cartago y Union,
han sido areas hasta el presente inexploradas desde
el punto de vista botanieo. Exploraeiones mas in-

tensas en un futuro pueden arrojar a la luz otros
hallazgos igual de interesantes y novedosos.

Paratipos. COSTA RICA. Heredia: Par. Nae. Braulio
Carrillo, falda atlantiea del volean Rarva. 28 Mar. 2002,
./. F. Morales 9772 (CR. INB, MO). Union: Altos de Pas-
cua, eabeeeras Quebrada Linda, 1 1 Jul. 2003, ./. /'. Mo¬
rales & II. Callejas 9426 (INB. MO).

Agradecimientos. Deseamos agradecer a James
Luteyn (NY) por ayudarnos a clarificar en forma
previa el status de esta espeeie, asf eomo a Claudia
Aragon por la ilustraeion. Tambien se agradeee la
revision crftiea del manuserito por parte de un re-
visor andnimo. Esta publieaeion fue financiada por
la Asisteneia Holandesa para el Desarrollo (NEI)A)
a traves del proyeeto “Desarrollo del Conoeimiento
de la Biodiversidad y Uso Sostenible en Costa
Kiea eondueido por el Instituto Naeional de Bio¬
diversidad ( I N Bio). Ademas. la investigaeion fue
posible graeias al eonvenio de eooperaeion entre el
Ministerio de Ambiente y Knergia (MINAE) e IN-
Bio eon el fin de conipletar en Inventario Naeional
de Biodiversidad.

Literatura Citada

Luteyn. J. L. 1991. key to the subfamilies anil genera of
Neotropical Ericaceae. Nordic .). But. 11: 623-627.

- . 1996. Ericaceae. In: C. Marling & L. Andersson

(editors), Flora of Ecuador 54: 1—104. Univ. Giiteborg/
Riksmuseum.

Dos Especies Nuevas del Genero Melica L. (Poaceae: Melieeae)

del Uruguay

Jaime Mujica Salles

Laboratorio de Ecofisiologia, Departamento tie Solos, Universidade Federal Rural do Rio de
Janeiro, Rodovia BR465 Km 7 s.n., Serop6dica, Rio de Janeiro, Brasil, CEP 23890-000.

jaime.mujica@uss.br

Marene Machado Marchi

Heihario do Centro de Ciencias Exatas, leenologicas e da Natureza, Universidade Sevenno
Somhra, Av. Exp. Osvvaldo de Almeida Ramos, 280— Bloco Professors Therezinha Coelho de
Souza-Conjunto Universitario, Centro, Vassouras, RJ, Brasil. CEP 27700-000.

marene.marchi@uss.hr

Rksumkn. Se describen e ilustran dos nuevas es¬
pecies de Melica I.. (Poaceae: Melieeae), M. ani-
marum y M. serrana, enddrnicas de la Sierra de
Animas (Maldonado, Uruguay). Melica animarum y
M. serrana son alines a M. hyalina Doll. Melica
animarum se diferencia de M. Iiyalina por la gluma
inferior elfptiea. de apice dentado y la lema de api¬
ce bidentado de hordes irregulares. Melica serrana
se dih "rencia de M. Iiyalina por la gluma inferior
obovada. de apice emarginado e irregular y la lema
de apice irregular eroso.

A US i’ll ACT. Two new species of Melica h. (Po¬
aceae: Melieeae), M. animarum and M. serrana, en¬
demic to the Sierra de Animas (Maldonado, Uru¬
guay), are described and illustrated. Melica
animarum and M. serrana are related to M. hyalina
Doll. Melica animarum is differentiated from M.
hyalina by the elliptic lower glume, with a dentate
apex and lemma with bidentate apex and irregular
margins. Melica serrana is differentiated from M.
hyalina Doll by the ohovate lower glume, with an
emarginated and irregular apex and lemma with an
irregular and erose apex.

Key words: Melica, Melieeae, Poaceae, Uru¬
guay.

El genero Melica U. (Poaceae— Melieeae) presenla
entre 96 (Hempel, com. pers. 2003) y 80 especies
(Clayton & Renvoize, 1989), distribuidas princi-
palmente en las regiones templadas, con except ion
del eontinente australiano. En Uruguay se compor-
tan como especies de ciclo invernal, floreciendo
durante la primavera-verano. habitando principal-
mente en las praderas.

El mayor numero de especies en America del

Novon 15: 338-342. 2005.

Sur, aparece en Argentina, Rrasil y Uruguay. En
Argentina se encuentran 17 especies (Zuloaga et
al., 1994), presentando la mayor diversidad en el
centre y este (Torres, 1980). La mayor concentra-
cidn de especies en Brasil, ocurre en Rio Grande
do Sul, con 12 especies (Longhi-Wagner, 1987),
disminuyendo en numero y expresidn en direceidn
norte. Para Uruguay, son citadas 10 especies por
Rosengurtt et al. (1970), distribuidas por todo el
pafs. Para Chile, son referidas 7 especies, distri-
buidas principalmente en la region central (Muftoz-
Schick, 1984).

Durante el levantamiento de la familia Poaceae
en la Sierra de Animas, en la region Este del Uru¬
guay, se constataron dos especies nuevas de Melica
para la ciencia, Melica animarum y Melica serrana,
endemieas del lugar.

I, a Sierra de Animas, ubicada en el Departa¬
mento de Maldonado, esla constituida por una su-
cesidu tie t erms retlondeados, que se extienden en
sentido norte-sur, donde se local iza el Gerro de las
Animas, lugar tie colecta tie las especies aquf tles-
criptas. El Cerro tit* las Animas es la maxima ele-
vacitin de la format ion, con una altura de 501 m
s.m. Estt>s cerros se encuentran separados por va-
lles profundos, con las laderas cubiertas por vege-
tacidn serrana arbustiva y arborea, sujetos a una
accidn constante del viento proveniente tlel mar. La
combinacidn tie una serie tie faetores, tales como
altitud, viento. radiaeidn luminosa, humedad, suelt)
y vegetacion, determinan la ocurrencia tie mi-
croambientes particulars que propician la existen-
cia tie especies entlemicas tie gramfneas y tie olras
familias, como la Mimosa Imrkartii descripta por
Marchesi (1987).

Volume 15, Number 2
2005

Mujica & Marchi
Melica del Uruguay

339

Melica animarum J. Mujica & M. Marchi, sp. nov.
TIPO: Uruguay. Maldonado: “. . . Sierra de
Animas, Cerro de las Animas, ladera Oeste,
en la cercama de la cima”, 2 Nov. 1991, J.
Mujica 240 (holotipo, MVFA). Figura 1.

Melicae hyalinae affinis sed glurria inferiore el 1 i plica,
apice dentato et lemmate apice bidentado praecipue dif-
fert.

Planta perenne, cespitosa, con rizoma horizontal,
de 25 cm de altura; canas erectas, simples, con
asperezas retrorsas esparcidas; vainas foliares gla-
bras, retroescabrosas; lfgula membranacea, hialina,
glabra, dilacerada, desde muy reducida hasta 0.8
mm de long.; laminas foliares lanceoladas, planas,
tie apice agudo no punzante. dfsticas, de (5— )9— 10
X 0.2—0.28 cm, de faz dorsal estriada, faz ventral
tie nervaduras marcadas y con asperezas antrorsas
abundantes, margenes con pestanas esparcidas. In-
florescencia en panoja lineal, contrafda, ± densa,
de 7-10 X 1—1.5 cm, tie ramas vertieiladas, las
mayores tie 5 cm de long., lisas; ratjuis con escasas
asperezas retrorsas; petlicelos curvos, pilosos. Es-
piguillas comprimidas lateralmente, violaceas, de
6-7.7 mm de long., con 2, raramente I antecio(s)
inferior(es) hermafrodita(s) y 1 a 3 neutros termina-
les reducidos, formando un cuerpo apical; raquilla
articulada por debajo de las glumas, estas caducas
con el antecio maduro; glumas tlesiguales en forma
y tamano, papiraceas en la region de los nervios,
hialinas en los margenes y con una amplia zona
hialina apical; gluina inferior elfptica, de apice
dentado, dorso list), violaceo, a veces con debiles y
escasas asperezas antrorsas en la mitad superior,
glabra, de 6-7.7 X 4—4.8 mm, 7-nervada; gluma
superior oblanceolada, de apice emarginado, de
dorso escabriusculo, glabra, violacea, de nervadu¬
ras muy poco notorias, con una amplia zona hialina
apical, 5-nervada, de 6—7 X 1.8— 2.3 mm. Primer
antecio hermafrodita, con lema elfptica, lanceolada,
papiracea, de apice bidentado, de hordes irregu-
lares, tie 4.8-6 mm (incluyendo el apice hialino tie
1-1.2 mm de long.) X 1.8-2. 4 mm de ancho total,
dorso con nervaduras poco engrosadas, escabrosas,
con pelos submarginales, hialinos, retorcidos, de
hasta 1 .2 mm de long.; palea elfptica, de base es-
treeha, hialina, de apice bidenticulado, 2-nervada,
nervaduras cilioladas en su mitad o cuarto superior,
dorso glabro y liso entre las nervaduras, de 4.8—5
X 1 . 1-1 .2 mm; segundo antecio hermafrodita, simi¬
lar al basal, pero de menor tamano y generalmente
glabro y con escasos pelos submarginales, a veces
neutro y sin palea; antecios superiores neutros, re-
ducidos a las lemas rudimentarias, incluidas una
deutro tie otra, constituyendo un cuerpo apical ob-

cdnico; estambres 3, con anteras lineales de 1.1-
1 .2 mm de long.; ovario glabro, con 2 estigmas plu-
mosos. Cariopsis no observado.

Melica animarum es endemica de la Sierra de
Animas, habiendo sido colectada unicamente en un
campo tie la ladera oeste, proximo a la cima del
Cerro de las Animas. El lugar tie colecta esta
aproximadamente a 500 in s.m., presentando un
suelo oscuro, rico en materia organica, con eober-
tura pedregosa. En el area referida, la especie es
rara, no habiendo sido colectados mas ejemplares
de la misma, a pesar de que los autores Inin retor¬
nado al Cerro de las Animas en diversas ocasiones
con tal finalidad.

Melica animarum fue colectada floreeida, no
fructificada a inicios de noviembre. Esta especie es
affn a Melica hyalina Doll, esta ultima tambien
existente en la region, diferenciandose principal-
mente por la gluma inferior elfptica y de apice den-
tado, y por la lema de apice bidentado tie hordes
irregulares.

Melica serrana J. Mujica & M. Marchi, sp. nov.
TIPO: Uruguay. Maldonado: “. . . Sierra de
Animas, Cerro de las Animas, en horde de
bosque”, 27 Nov. 1995, M. Marchi , J. Mujica
& R. Brescia 1464 (holotipo, MVFA). Figura 2.

Melicae hyalinae affinis sed gluma inferiore obovata
apice emarginato et irregulari, lemmate apice irregulari
eroso differl.

Planta perenne, cespitosa, con rizoma horizontal,
de 25-65 cm tie altura; canas subereetas, simples,
escabras o escabriusculas; vaina foliar eseabrius-
cula, con pelos cortos adpresos sobre las nervadu¬
ras, o glabra y lisa; lfgula membranacea, hialina,
frecuentemente dilacerada, o entera tie apice ob-
tuso, de 0.5—3 mm de long., glabra o con algunos
pelos esparcidos; laminas foliares linear-lanceola-
das, involutas, de apice agudo no punzante, de 5-
15 X 0. 1 -0.3(-0.4) cm, de faz dorsal glabra, es-
cabriuscula, con margenes pestanosos, faz ventral
cortamente pilosa. Inflorescencia en panoja lanceo¬
lada estrecha o lineal, generalmente pauciflora, de
10-22 X 1.5(-4) cm, de ramas vertieiladas aproxi-
madas al raquis, las basales mayores de hasta 6 cm
tie long., glabras, escabriusculas; raquis glabro, es¬
cabriusculo; petlicelos curvos, pilosos. Espiguillas
comprimidas dorsiventralmente, pajizas, tie 5—8
mm tie long., con 2, raramente 1 antecio(s) inferi¬
or's) hermafrodita(s) y 1 a 3 neutros terminales
reducidos a las lemas rudimentarias, formando un
cuerpo apical; raquilla articulada por debajo tie las
glumas, estas caducas con el antecio maduro; glu¬
mas dife rentes en forma y tamano, papiraceas, hia-

340

Novon

I' igura I. Melica unimarum J. Mujioa & M. Marchi. — A. Habito. — B. Espiguilla, vista ventral. — C. Gluma inferior,
vista dorsal. — I). (.luma superior, vista dorsal. — k. Lema I. vista dorsal. — F. Antecio basal, vista ventral. — -G. lYilea
I, vista dorsal. — H. Antecio II, vista dorsal. — I. Antecio II. vista ventral y cuerpo apical. Dibujos basados en el
bolotipo. ./. Mujica 240.

Volume 15, Number 2
2005

Mujica & Marchi
Melica del Uruguay

341

I- ig ma 2. Melica serrana .1. Mujica & M. Marchi. —A. Hdbito. — B. Espiguilla, vista ventral. — C. Gluma inferior,
vista dorsal. — D. Gluma superior, vista dorsal. — E. Lema I, vista dorsal. — F. Palea I. vista dorsal. — G. Antecio II.
vista dorsal. — II. Antecio II y cuerpo apical. — I. Cariopse, vista ventral. — J. Cariopse, vista dorsal, bibujos basados
en los paratipos M. Marchi et at. I 16 / (A— III y M. Marclii el at. 1465 (I. J).

linas en los indigenes y en el apice; gluma inferior
obovada, de apice emarginado de hordes irregula-
res. pajiza, glabra, lisa, con una amplia zona hialina
apical, de 6—8 X 4—5.4 nun. 7— 9-nervada; gluma
superior oblanceolada, de apice obtuso, eroso, irre¬
gular, dorso glabro y liso, o con escasos pelos cortos

sobre el extremo superior de las nervaduras o en
los dos tercios superiores del dorso, a veces con
escasas asperezas antrorsas superiormente. con una
amplia zona hialina apical de 1.1 — 1.4 mm de long..
5-nervada, de (6. 1— )6.5— 7.2 X 1.2-2 mm. Primer
antecio hermafrodita con lema elfptica, de apice

342

Novon

irregular eroso, dorso con nervaduras engrosadas,
con asperezas rctrorsas. pelos hialinos, retorcidos
de liasta 0.7 mm de long, sobre las nervaduras la-
teralcs, de 4.6— 5.5 mm (ineluyendo el dpice hialino
de ().(>— 0.7 mm de long.) X 1.2— 1.7 mm; palea eli'p-
tica, oblanceolada, de apice bidenticulado. 2-ner-
vada, nervaduras cilioladas superiormente, dorso
glabro y liso o con micropapilas puntiformes ('litre
carenas, de 4—5 X 0.9— 1.6 mm; segundo anteeio
hermafrodita, similar al basal, pero algo menor y
glabro; antccios superiores neutros, reducidos a las
lemas rudimentarius, incluidas una dentro de la
otra, constituyendo mi cuerpo apical oblongo; es-
tambres 3, con anteras cleistogamas de 0.6 mm de
long.; ovario glabro, con 2 estigmas plumosos. Ca-
riopsis elipsoideo. de liilo linear, que sc extiende
de extremo a extremo; glabro, liso. castano lustroso,
de 2.7—3 X 0.9 mm.

Melica serrana es una especie endemiea del Ce-
rro de las Animas, siendo observada en campo ar-
bustivo localizado a media altura (aproximada-
mente a 250 m s.m.). No se observo su oeurrencia
en otros lugares de la Sierra. Ksta especie fue co-
leetada a lines de noviembre, presentando algunos
ejemplares con cariopsis maduros. Melica serrana
es all'll a M. hyalina, diferenciandose principal-
mente por su gluma inferior obovada, de apice
emarginado c irregular, y por su lema de apice irre¬
gular eroso.

Paratipos. URUGUAY. Maldonado: Sierra de Ani¬
mas, Cerro de las Animas, 27 Nov. 1995, M. Marchi, J.

Mujica & R. Brescia 1461. 1464a, 1465. 1 466 (MVFA),
1467 (MO).

Agradecimientos. Agradecemos a la lug. Pri-
mavera Izaguirre de Artucio por todo el apoyo re-
cibido en el Uruguay; al lug. Haul Brescia por su
colaboracidn en los trabajos de campo; al Dr. Ro¬
berto Rossiello e Dra. Ariane l.una Peixoto, por su
importante estinmlo; al Sr. Pierino Zerboni por las
ilustraciones; y al Dr. Tarciso Kilgueiras por las di¬
agnosis latinas. Agradecemos tambien a la Comi-
sidn Sectorial de Investigacion Cientffica (CS1C), de
la Universidad de la Republica, por el subsidio
otorgado; y a la Catedra de Botaniea de la Facultad
de Agronomfa de Montevideo, por las facilidades
brindadas.

I.iteratura Citada

Clayton, W. 1). & S. A. Renvoize. 1989. Genera Grami-
num. Kew Bull. Addit. Ser. XIII, 2 ed.

I. ongh i- Wagner, II. M. 1987. Tribo Poeae (Gramineae).
Bol. Inst. Bine. Univ. Fed. Rio Grande do Sul 41: I —
191.

Marchesi, E. 1987. Dos espeeies nuevas de la (lora uru-
guaya. Bol. Invest. Far. Agron. Montevideo 5: 1—8.
Muftoz-Schick, M. 1984. Revision de las espeeies del g 6-
nero Melica L. (Gramineae) en Chile. Bol. Mus. Nae.
Hist. Nat. 40: 41-89.

Rosengurtt, lb. B. R. Arrillaga de Malei & P. Izaguirre de
Artucio. 1970. Gramfneas Uruguayas. Universidad de
la Republica, Montevideo.

Torres, M. A. 1980. Revisidn de las espeeies argentinas
del g^nero Melica L. (Gramineae). Opera Uilloana 29:
1-1 15.

Zuloaga, F. ()., E. G. Nicora, Z. E. Rugolo de Agrasar, 0.
Morrone, J. Pensiero A. M. Cialdella. 1994. Cati'dogo
de la Familia Poaceae en la Republica Argentina. Mo-
nogr. Syst. Bot. Missouri Bot. Card. 47.

A New Species of Beilschmiedia (Lauraceae) from Borneo

Sack iko Nish i da

The Nagoya University Museum, Furo-cho, Chikusa-ku, Nagoya 464-8601, Japan.

nishida@num.nagoya-u.ac.jp

Abstract. A new species of Beilschmiedia , pre¬
viously recognized by Kostermans hut never pub¬
lished, is described as B. oligantha and illustrated.
This new species differs from the other Bornean
Beilschmiedia species in the combination of the fol¬
lowing characters: long erect trichomes on twigs,
six to ten pairs of secondary leaf veins, relatively
large [tale red flowers, and stamens with short pu¬
bescent filaments.

Key words: Beilschmiedia, Borneo, Lauraceae.

Beilschmiedia Nees is one of the larger genera
of Lauraceae and includes about 250 species dis¬
tributed mainly in the paleotropies (Nishida, 2001).
It is distinguished from the other laurel genera by
its paniculate or racemose inflorescences that are
not strictly opposite at the terminal division, bisex¬
ual and trimerous flowers with six equal to sube¬
qual tepals, nine (rarely six) fertile stamens with
two-celled anthers, and fruits lacking cupules. One
of the centers of species diversity is Southeast Asia,
but the taxonomy of the genus in this region is still
confused because of the lack of a comprehensive
revision.

While revising the specimens from Borneo, one
of the larger islands in this region, 1 found several
groups of specimens that were informally recog¬
nized as new species by Kostermans, but never
published. These specimens were annotated by him
with unpublished names, and some of them were
even placed in type folders at a few herbaria, e.g..
BO, K. Since these names have caused taxonomic
confusion. 1 clarified the published names for the
Bornean Beilschmiedia species and described one
new species in my last publication (Nishida, 2001).
Recently, I found another group of specimens that
is different from any known Bornean Beilschmiedia
species. It is distinct from other species in the com¬
bination of the following characters: long erect tri¬
chomes on twigs, six to ten pairs of secondary leaf
veins, and large (ea. 5 mm) pale red flowers, as
well as stamens with short pubescent filaments. 1
conclude that it represents a new species and de¬
scribe it here.

Bcilsclimicdiu oligantha Sachiko Nishida, sp.
nov. TYPE: Malaysia. Sarawak: Kapit. Pela-
gus. I 1 July 1979, B. Lee S40647 (holotype,
L; isotypes, K not seen, KEP, SAN not seen.
SAR). Figure 1.

Ah aliis speciebus borneensibus, ramulis patenter pi¬
losis, nervis secondariis 6—10 utrinque, fioribus rubris 5
mm longis, filamentis pubescentibus et brevibus differt.

Small tree, up to 10 m tall; terminal buds and
twigs densely pubescent with erect, long (up to 1.5
mm), straight trichomes. Leaves spirally and evenly
arranged on branches, (14)17—20(24) X (4)5—10
cm, obovate or sometimes elliptic, chartaceous,
base acute, apex acuminate or cuspidate, margin
flat, upper leaf surface pubescent only along mid¬
rib, lower surface sparsely pubescent with erect,
long, straight trichomes, lower surface glaucous;
midrib almost immersed above, raised below, sec¬
ondary veins 6 to 9(10) pairs, almost immersed
above, raised below, tertiaries not percurrent, minor
venation relatively fine, smallest reticulations ca. 2
mm diam., slightly raised (the smallest veins im¬
mersed) above, slightly raised below; petioles 1 .5—
2 cm long, flat to slightly canaliculate above, dense¬
ly pubescent throughout. Inflorescences axillary,
paniculate, 1—5 cm long, each bearing a few to 30
flowers, rachis reddish tinged, densely pubescent;
floral pedicels of lateral divisions ca. 1 mm long,
pedicels of central flowers ca. 2—3 mm long; bracts
as large as flowers. Flowers pale red. ca. 5 X 5
mm; tepals 6, almost equal, ovate, ca. 3.5 X 2.5
mm, glabrate outside, sparsely pubescent inside;
stamens 9, ca. 2 mm long, 2-celled, the outer 6
with the cells opening introrsely, inner 3 opening
extrorsely or laterally; filaments pubescent, shorter
than anthers, the inner 3 filaments with 2 globose
glands near the base; anthers glabrous with ovate
apex; staminodes 3, ca. 1.5 mm long, sagittate; pis¬
til glabrous, ca. 3 mm long, ovary gradually nar¬
rowed in the style; receptacle relatively deep, pu¬
bescent inside. Fruits ellipsoid, ca. 2.8 X 1 .5 cm,
drying reddish black, surface smooth; fruit pedicels
ca. 2 mm diam., apieally constricted.

Habitat and distribution. In forests dominated

Novon 15: 343-345. 2005.

344

Novon

Figure I. Heilschmiedia oligantha Sachiko Nishida. — A. Habit. — B. Venation pattern (abaxial view). — C. Inflores¬
cence. — I). Flower (front tepals removed). — E. First whorl stamen (adaxial view). — F. Second whorl stamen (adaxial
view). — (i. Third whorl stamen with glands (abaxial view). — H. Staminode (abaxial view). — I. Pistil. — J. Fruit. A,
B, D— I. Drawn from the holotype {H. Tee S40647). C. Reconstructed from the holotype and isotypes. J. Drawn from a
paratvpe (Sibal <ik Luang S22I58, SAR).

Volume 15, Number 2
2005

Nishida

Beilschmiedia oligantha from Borneo

345

Table 1. Differences

between Beilschmiedia oligantha and Bornean

species morphologic;

illy close to 11. oligantha.

Features

B. oligantha

11. maingayi

B. pilosa

B. wieringae

Leaf length (cm)

( 1 4)1 7-20(2 4)

1 6-25(30)

(24)28-30(34)

(16)18-24

Secondary veins (pairs)

6 to 9(10)

8 to 10

(13)14 to 17

8 to 1 1

Leaf shape

obovate to elliptic

elliptic

lanceolate, rarely

elliptic

narrowly elliptic

Petiole length (cm)

1.5-2

1 .8-3

1 .5-2.3

2-3.5

Flower color

pale red

yellowish green

red

white

Stamen filament length

shorter than anthers

'boiler than anthers

as long as anthers

longer than anthers

Stamen filament

pubescent

pubescent

glaluate

glabrale

Fruit pedicels

apically constricted

not constricted

apically constricted

apically constricted

by Dipterocarpaceae (“mixed dipterocarp forests”),
15—1400 m; known (tidy from northwestern Borneo.

This is one of the species informally recognized
as new, but never published, by Kostermans. He
wrote tbe name ‘'Beilschmiedia pauciflora' on spec¬
imens belonging to this species, but this name was
already given to a different species (Beilschmiedia
pauciflora H. W. Li). Thus, 1 translated the epithet
into Greek as an alternative, Beilschmiedia oligan¬
tha, crediting Kostermans's earlier determination.

Beilschmiedia has about 30 known species in
Horneo (Nishida, 2001). Among these. B. oligantha
is most closely related to B. maingayi Hooker 1.
both share a similar leaf shape and inflorescence
structure. However, B. oligantha differs from B.
maingayi in its pilose twigs, pale red flowers, and
fruit pedicels that are apically constricted (Table
I). Collections of B. oligantha have occasionally
been misidentified as B. pilosa Kostermans and B.
wieringae Kostermans, due to their similar indu-
ment type. However, Beilschmiedia oligantha can
be distinguished from B. pilosa by its shorter leaves
(blade length (14)17—20(24) cm vs. (24)28—30(34)
cm in B. pilosa) with fewer pairs of secondary veins
(6 to 9(10) vs. (13)14 to 17) and pubescent fila¬
ments (vs. glabrale). The new species also differs
from B. wieringae by its shorter petioles (1.5—2 cm
vs. 2—3.5 cm in B. wieringae). pale red flowers (vs.
white), and stamens with short pubescent filaments
(vs. stamens with glabrale filaments longer than an¬
thers) (Table 1).

Paratypes. MALAYSIA. Sarawak: Bintula, liukit
Mersing, Sihat ak Luang S2I877 (I., SAR), Sibat ak

Luang S21886 (1,, SAR), Sibat ak Luang S22I58 (L.
SAR), ,/. 11. Purseglove 5276 (k. 1.); Sungai kakns. ca. 4
km K from Minah, M. Hirano & 47. Holla 205 (SAR); Miri,
Lambir Hills. I lias P. S16615 (L, SAR). Rena G. S40280
(L. SAR). Sabah: Beaufort, Beaufort Hill, J. k. l/ijanga h
14514 (K. L, SAN); Keningau, Sg. Siliawan Sepulut For.
Res., Pidilis k. SAN 112019 (K, L); Keningau, Kerumatoi
Liu I Bata 7 Vi Sook, Saikeh L. SAN73254 (L, SAN); ken-
ingau, Ulu Sg. Matud, Sumbing ./. SAN 1 19569 (SAN);
Kinabalu. Kampung Malangkap Tamis, Sungai Kinotoki 1
km dari Malangkap Tamis, L. Lugas 712 (k); Nabawan,
KM 55 Jalan Nabawan/Sepulut, Ulu Sg. Nabawan, Sumb¬
ing SAN 1 28374 (L). BRUNEI. Brunei-Muara: Lumapas.
Bukit Sa'di. Ogata. Ariffin k A Han S. ()g-B105 (I,). Jof-
fre Hj. Ali Ahmad. Niga N., Sharbini 47.. Ibrahim A. &
Jangarun E. I1RUN15389 (l„ SAN). Joffre. Niga. Ariffin.
Hussain //. ()., Ibrahim & Han. BRUN 16741 (L); Bukit
Lumapas, on banks along stream at base of liukil Sa'eli,
A.P. Davis, Bygrave PC., kirkup D., Niga. Joffre. Han.
Ibrahim 512 (K). Belait: new road to Merankin from km
13 Ualii Road, ea. 3 km from junction. ./. Dransfield 6847
(I,); Sukang, P S. Ashton BRUN 116 (BRUN, L); Sukang,
Sungei Kuat, Ulu Belait, Salleh /.. Ibrahim A., Melinau
G. Emran k., Arfin k. & Clayton ./. BRl V 15342 (L).
Tenibiirong: Kuala Belalong, ridge bounded In Belalong
and Temburong rivers, S of Belalong, k. 47. Wong 1234
(L). INDONESIA. Kalinianlan Timur: between bong Ha¬
waii and Panado, R. Geesink 8990 (L).

Acknowledgments. I am grateful to Mutsuko
Nakajima for providing tbe excellent illustration,
and to Hideloshi Nagamasu for improving the Latin
diagnosis. I also thank tin* curators of A, BO, K.
KKP, L. SAN, and SAR for loans.

I .iterature Cit(‘d

Nishida, S. 2001. Notes on Beilschmiedia (I.auraceae) of
Southeast Asia I: Checklist of the Bornean species with
the description of a new species. Acta Phytotax. Geobot.
52(2): 103-113.

Lupinus reitzii (Fabaceae— Faboideae), a New Species of the
Lupinus lanatus Complex from Southern Brazil

Mardiore Pinheiro and Silvia Teresinha Sfoggia Miotto
Depto. Botanica, IB-UFRGS, Campus do Vale, Porto Alegre, RS, Cep. 91.501-970, Brazil.

mardiorepinheiro@brturbo.eom

A as m ACT. Lupinus reitzii , a new species from
southern Brazil, is described and illustrated. The
new species belongs to t lie Lupinus lanatus com¬
plex and is closely related to L. lanatus and L.
magnistipulatus, from which it is set apart princi¬
pally by the size and shape of the stipules and by
the shape and pubescence of the leaflets.

Rksumo. F descrita e ilustrada Lupinus reitzii,
tuna nova especie do Sul do Brasil. Lupinus reitzii
£ relacionada com os taxons do complexo Lupinus
lanatus, especialmente L. lanatus e L. magnisti-
pulatus, diferenciando-se destas prineipalmente
pelo tamanho e forma das estfpulas, forma e pilos-
idade dos folfolos.

Key wards: Fabaceae— Faboideae, Lupinus lan¬
atus complex, Lupinus reitzii, southern Brazil.

During a revision of the genus Lupinus I., in the
Brazilian state of Rio Grande do Sul (Pinheiro &
Miotto, 2001), we had the opportunity to study the
type material of L. magnistipulatus Burkart ex
Planchuelo & Dunn and verify that some of the
paratypos present morphological differences from
the isotype ( Reitz 2377, HBR). In 1961, Burkart,
while analyzing the collections Smith & Reitz 9155
(HBR) and 10135 (IIS, HBR), wrote on these sheets
t hat they represented a new species, which he
dubbed L. reitzii, but never validated the name
(Pinheiro & Miotto, 2001). Cater, Planchuelo and
Dunn (1989) considered these collections to rep¬
resent young individuals of L. magnistipulatus and
referred them to this species. Our analysis of spec¬
imens determined as L. magnistipulatus, with ad¬
ditional collections from southern Brazil, indicates
that a new species is represented among these col¬
lections and is herein described.

Lupinus reitzii Burkart ex M. Pinheiro & Miotto,
sp. nov. I V PC: Brazil. Santa Catarina: Var-
geao, BB 282. km 476, .41 Oc t. 1999 (fl), M.
Pinheiro 207 (holotype, ICN; isotypes, k,
MBM. MO). Figure 1.

Herbae probabiliter biennes, erectae, secundum caules

ramosae. Folia basalia unifoliolata et trifoliolata, altera 5-
foliolata, interdum folia trifoliata in ramis lateralibus; fo-
liola late oblanceolata ad angusta oblanceolata, apiee ro-
tundato, marginibus undulatis, sparse utrinque lanosis,
pili flaveseentes, undulati et mixti; stipulae petiolo adna-
lae, parte distali libera, lanceolata vel laneeolato-caudata,
apice acuniinato, interdum acuto, dorsaliter lanosae, ven-
traliter glabrae. Racemi terminales, densiflori; bracteae
caducae, lanceolato-eaudatae, dorsaliter lanosae, ventral-
iter glabrae. Flores nisei, roseo-purpureive vel colore in-
dico tincti; calyx labio infero trilido, labio supero profunde
bilido; alae oblongae, apice rotundato. I .('gumma lanosa;
semina elliplica.

Plants probably biennial, herbaceous, erect,
branched along the stem, (12)18.5—70 cm tall,
sometimes in cushions more than 50 cm diam.;
stem finely or densely lanate. Basal leaves unifo-
liolate and trifoliolate, evident only in juvenile
plants; adult leaves sometimes trifoliolate in lateral
branches, mostly 5- to 7(8)-foliolate; leaflets 3—
8(9.5) X 1—3 cm, widely to narrowly oblanceolate,
apically rounded, margins undulate, finely lanate
on both sides, with sparse intertwined trichomes
yellow-tinged, about 5-7 mm long; petioles 2.3-
12.5(15) cm; stipules (1 1)18—70 mm, partly aduate
to petioles, the free portion 8—38(49) X 3-9(13)
mm, lanceolate to lanceolate-caudate, the apex
acuminate, sometimes acute, glabrous adaxially, la¬
nate abaxially. Terminal racemes, densiflorous, 25-
to 50-flowered, 4.6—20.0 cm; peduncles (1.3)3—
6. 6(8.9) cm; rachis densely lanate; pedicels 1.5—
4.7(6) mm, pilose as the rachis; bracts 10.7—16 X
2.1— 4.2 mm, caducous, lanceolate-attenuate to lan¬
ceolate-caudate, lanate abaxially. Flowers 10—15.2
mm, pink, purple-pink, or indigo; bracteoles 2.5
mm long, lanceolate, rarely oblanceolate, glabrous
within, finely to densely lanate outside; calyx gla¬
brous within, finely to densely lanate outside; lower
lip 1 1 — 14 X 2.7— 4.5 mm, the apex trifid with cen¬
tral tooth 1.8-3. 8 mm and lateral teeth 1.2— 3.4 mm;
upper lip deeply bifid. 6.3—9 X 4. 1-6 mm; laciniae

1 .8— 2.8 mm wide, w ith a notch between them 4.1—
6.9 mm deep; corolla with standard 13—16.8 X

7.8- 12.3 mm, ovate, sometimes w idely ovate or ob¬
long-ovate; wings 13.5—18.4 X 4.7— 7.6 mm, ob¬
long, apex rounded, the claw 2.4— 3.8 mm long;

Novon 15: 346-349. 2005.

Volume 15, Number 2
2005

Pinheiro & Miotto
Lupinus reitzii from Brazil

347

Figure 1. Lupimis reitzii Burkart ex M. Pinheiro & Miotto. — A. Flowering branch. — B. Stipules. — C. Bract, ventral
view. — I). Calyx, inside surface. — E. Banner petal, ventral view. — F. Wing petal. — G, H. Keel petals. Seale bars =
4 cm in A; 2 cm in B; 5 mm in C-H. (A— G: drawn from the type, M. Pinheiro 207: H: M. Pinheiro 318.)

348

Novon

keels 12.6—16.6 X 3.8— 5.4 mm, the apex falcate,
straight or twisted upward, rarely abruptly falcate-
arcuate, the claw 3-4.5 mm long; ovary with 7
ovules. Legumes (33)43—64(74) X 7—9 mm. dense¬
ly lanate; seeds 4.2— 4.6 X 3—3.4 mm, elliptic.

Habitat and distribution. Lupinus reitzii was
collected in sunny places on roadsides and hare
rocky soils in the southern Brazilian slates of Rio
Grande do Sul, Santa Catarina, and Parana.

Phenology. Flowering and fruiting plants were
collected from October to January.

Etymology. The epithet “reitzii" is an homage
to Brazilian botanist Raulino Reitz.

Pinheiro and Miotto (2001). after a morphologi¬
cal analysis of the isotype of L. magnistipulatus (R.
Reitz 2377, HBR) and the paratypes (/,. B. Smith.
R. Reitz S: R. M. Klein 7703 . HBR, US; L. B. Smith
& R. Reitz 10320 , HBR, R, US; R. Reitz & R. \L
Klein 7117. HBR). verified that these were the only
specimens examined by Planchuelo and Dunn
(1989) which belong to L. magnistipidatus. while
the other paratypes examined belong to L. reitzii.
Moreover, Pinheiro and Miotto (2001), during the
revision ol Lupinus in Rio Grande do Sul, based on
the morphological analysis (stipules, leaflets, hau¬
lier, and calyx) of L. magnistipulatus and /.. seti-
folius Planelmelo & Dunn, concluded that both
species are the same taxon, justifying the synony-
mization of L. setifolias. It should be emphasized
that in the holotype photo of h. magnistipidatus (R.
Reitz 2377, SI), there are many notes written by
Burkart. who indicated that this specimen is iden¬
tical to the specimen B. Rambo 3620B (isotype,
PAG A), designated by Planchuelo and Dunn as the
holotype of I,, setifolius.

Planchuelo and Dunn (1989) considered the
presence ol Iriehomes on the leaflets a distinguish¬
ing characteristic to separate L. magnistipidatus
(hairv leaflets) from L. setifolius (glabrous leaflets).
However, the analysis of L. magnistipidatus popu¬
lations demonstrated an intraspecific variation con¬
cerning their pubescence (Pinheiro & Miotto,
2001). Moreover, it can be emphasized that the de¬
scription of /,. magnistipulatus sensu Planchuelo &
Dunn also includes vegetative and reproductive
morphological characteristics of L. reitzii, which has
hairy leaflets.

The analysis ol the L. magnistipulatus and L. se¬
tifolius types, as well as of a great number of spec¬
imens. in addition to field observations, allowed a
better understanding ol these taxon circumscrip¬
tions and the adoption of L. reitzii as a new species.

Lupinus reitzii is closely related to L. magnisti¬
pulatus and L. lanatus Bentham. From /,. magnis¬

tipulatus it is set apart mainly through the mor¬
phology of the stipules and leaflets, as well as the
flower color. The stipules in L. reitzii are lanceolate
to lanceolate-caudate with an acuminate to acute
apex, with the free part 8—38(49) X 3-9(13) mm,
while in L. magnistipulatus they are lanceolate to
ovate-lanceolate with an acute apex, the free part
1 1-56.3 X 4.4—22.0(25.2) mm; the leaflets in L.
reitzii are widely to narrowly oblanceolate, both (ac¬
es lanate with sparse intertwined yellow-tinged tri-
chomes, while in L. magnistipidatus the leaflets are
w idely oblanceolate to obovate, glabrous or only oc¬
casionally lanose on both sides with sparse inter¬
twined white trichomes. Flowers are pink to purple-
pink. rarely indigo, in L. reitzii, while they are
consistently indigo in L. magnistipulatus.

Lupinus lanatus differs from the new species by
the densely white or whitened lanose, narrowly el¬
liptic to elliptic-oblaneeolate leaflets with plane
margins and an obtuse or rounded apex, as well as
the apices of the wings, which are rounded in L.
reitzii, but have an obtuse beak in L. lanatus. The
flowers in L. lanatus are blue or rarely while, con¬
trasting with the pink, purple-pink, to deeper in¬
digo ol L. reitzii.

The geographical distribution of these Lupinus
species is also distinct: Lupinus magnistipulatus is
known from isolated scattered populations restrict¬
ed to the montane fields from Rio Grande do Sul
and Santa Catarina, in the highest and coldest en¬
vironments of southern Brazil (Pinheiro & Miotto,
2001), while L. reitzii is widespread in southern
Brazil; L. lanatus occurs in Santa Catarina and Rio
Grande do Sul. where it is also widespread.

Paratypes. BRAZIL. Parana: Guarapuava, Guard, 5
Dec. 1968, G. Hatschbach 20102 (MBM, l S|. II. Brtlcher
3003 (K): Sao Jose dos Pinhais. 18 Oet. 1980, 0. Hatsvh-
bach 43213 (K). Rio Grande do Sul: Bom Jesus, estrada
para Sao Jos# dos Ausentes. 16 Nov. 1997, ./. A. Jarenkow
A' E. N. Garcia 3635a (PEL); Cambara do Sul. de Osvaldo
Kroeff, para Sao Jos# dos Ausentes. 5 Jan. 2000. S. T. S.
Miotto & M. Pinheiro 1844 (ICN); Itaimbezinho pr6x. de
Sao Francisco de Paula, lit Nov. 1953, Rambo 54507 (B,
PACA); Gaxias do Sul, Vila Oliva. 29 Oet. 1985. M. I..
Abruzzi 1027 (MAS); Sao Francisco do Paula, 30 Oet.
1995. .S’. T. .S'. Miotto et al. 1456 (ICN); estrada para Serra
do Umbu, 23 Oet. 1998, A. Flores 239 (ICN); ca. I I km
do rio Pinto, 23 Nov. 1998. M. Pinheiro 128 (ICN); Sao
Jos# dos Ausentes, em direyao a Silveira, 6 Jan. 2000. M.
Pinheiro 310 (ICN); Silveira, ea. 2 km em direyao a Bom
Jardim da Serra, 6 Jan. 2000. M. Pinheiro 313 (ICN).
Santa Catarina: Agua Doce, margeni do rio Cliapeoo. 4
Dee. 1964, /.. II. Smith & Klein 13552 (B); pr6x. 4 Ca-
yador. 2 Dec. 1964. I.. B. Smith & Klein 13393 (B. R);

I I km E de Bom Jardim da Serra, 25 Nov. 1980. A. Kra-
poviekas c C R. Vanni 36933 (CETS); Bom Jardim da Serra,
6 Jan. 2000, 1/. Pinheiro 318 (ICN); Cayador, Campos das
Palmas, 52 km 0 de Cayador Joayaba, 24 Dee. 1956, L.
R. Smith cC R. Reitz 9 155 (HBR); Capao Alto. SC 458,

Volume 15, Number 2
2005

Pinheiro & Miotto
Lupinus reitzii from Brazil

349

km 181, a 12 km do trevo da BR 116, 30 Oct. 1999, .S'.
T. S. Miotto 176.3 (ICN); Campo Belo do Sul. km 163, ea.
12 km da divisa com o mun. Campos Novos, 30 Oct. 1999,
1 VI. Pinheiro 198a (ICN); Cages, km 247, perfmetro urba-
no, 2 Nov. 1999, M. Pinheiro 225 (ICN); Ponte Serrada,
BR 282, km 450, 31 Oct. 1999. M. Pinheiro 205 (ICN);
para Faxinal dos Guedes, 13 Oct. 1964, /,. H. Smith &
Reitz 12480 (FLOR, R); para Xanxere, 8 Nov. 1964, L 13.
Smith <K Klein 1.3054 (US); Ponte Alta do Norte, BR 1 16,
km 165, 02 Nov. 1999, M. Pinheiro 221 (ICN); Sao Cris-
tovao do Sul, BR 1 16. km 198, 2 Nov. 1999, )l. Pinheiro
22.3 (ICN); Sao Joaquim, nascente do rio Capivaras, serra
do Oratdrio, 10 km E de Bom Jardim da Serra, 16 Jan.
1957, L P. Smith & Reitz 10135 (HBR, US), 22 Oct.
1961. (',. Pabst 6220 <t' Pereira 6393 (HR. PEC); Var-
geao, BR 282. km 476, em diregao a Faxinal dos Guedes,
31 Oct. 1999, M. Pinheiro 206 (ICN).

Acknowledgments. We are grateful to Marcos
Sobral lor his help, incentive, and suggestions; to
Tarciso Filgueiras for the Latin description; to Mar¬
cia Vignoli da Silva for the illustrations; and to
Capes, CNPq, and Pronex for financial support of
our studies.

literature Cited

Pinheiro, M. & S. T. S. Miotto. 2001. Flora Ilustrada do
Rio Grande do Sul. Ease. 27. Ceguminosae— Fahoideae:
Genero Lupinus C. Bol. Instil. Bioci. Univ. Fed. Rio
Grande do Sul 60: 1-102.

Planchuelo, A. M. & I). B. Dunn. 1989. Two new species
of the Lupinus lanatus complex. Ann. Missouri But.
Card. 76: 303-309.

A New Species of Amyris (Rutaceae) from
Baja California Sur, Mexico

Jon P. Rebman

San Diego Natural History Museum, P.0. Box 121390,

San Diego, California 921 12-1390, U.S.A. jrebman@sdnhm.org

Fernando Chiang

Departamento de Botaniea, Instituto de Biologfa, UNAM,
Apartado Postal 70-233, 04510 Mexico, D.F., Mexico

ABSTRACT. Field exploration and Holistic re¬
search have revealed another plant species endem¬
ic to the peninsula ol Baja California, Mexico. A
new shrubby species (Amyris carterae) from the Si¬
erra Guadalupe and Sierra de la Giganta, Baja Gal-
ifomia Sur. is described here. Distribution, asso¬
ciated vegetation, rarity, affinity to another species
in the genus, and a botanical illustration are pre¬
sented.

RKSUMKN. Trabajo de eampo e investigation flor-
fstiea han puesto de manifiesto una especie de
planta endemica de la peninsula de Baja Califor¬
nia, Mexico. Lin arbusto (Amyris carterae) de la Si¬
erra Guadalupe y Sierra de la Giganta, del estado
de Baja California Sur se describe aquf como es-
pecie nueva para la ciencia. Se presentan la dis-
tribucion. vegetacion asociada, rareza, afinidad con
otra especie en el genero, asi como una ilustracion
del nuevo taxon.

Key words: Amyris . Baja California Sur, Mexi¬
co, Rutaceae.

Lower California is comprised of two Mexican
states (Baja California and Baja California Sur) that
are politically divided at the 28th parallel. These
states compose the Baja California peninsula and
its adjacent islands, which are located in both the
Gulf ol California (Sea of Cortes) and the Pacific
Ocean. This region supports a wealth of species.
W iggins (1980) estimated that a total ol 2958 taxa.
of which 686 are endemic, occur in Lower Califor¬
nia, but recent discoveries and a more extensive
overview of the literature suggests that the flora
probably consists of more than 4000 plant taxa with
a rale ol endemism around 30% (Rebman, 2001).

The Rutaceae are represented in Lower Califor¬
nia by eight native species in six genera (Amyris P.
Browne, Cneoridium Hooker f., Esenbeckia HBk,
Ptelea I... Thamnosma Torrey & Fremont, and Zan-

thoxylum L.). At least four of these species (the new
species Amyris carterae, as well as Thamnosma tri-
foliata I. M. Johnston, Zanthoxylum arborescens
Rose, and Z. fagara (L.) Sargent) are rare or re¬
stricted in their distribution on the peninsula. The
genus Amyris has approximately 40 species distrib¬
uted from Florida, Texas, and the Antilles, south to
Venezuela and Peru (Gereau, 1991). At present,
only this species of Amyris is known to occur on
the peninsula ol Baja California; tin* member of the
genus with the closest geographical proximity is A.
balsamifera I.. in southern Sonora and Sinaloa in
Mexico. A collection (M. Dominguez 235, SD) made
in the Sierra de la Laguna ol the Cape Region may
represent another species ol this genus, but more
research and better collections are needed to sub¬
stantiate this occurrence. The new shrubby species
described here adds to the impressive diversity and
endemism of the flora of Lower California.

Amyris carterae J. Rebman & F. Chiang, sp. nov.
TYPE: Mexico. Baja California Sur: Sierra
Guadalupe, W of Mulege, Guadalupe Canyon
ea. 14 mi. SW ol San Jose de Magdalena, 500
m, 26°56'54"N, I I2°22'15''W, 25 Sep. 1998,
./. P. Rebman 5634 & ./. L. Zuniga (holotype,
SI) 142880; isotypes. BCMEX, HCIB, MEXll,
MO. RSA). Figure 1.

A. madrensi similis, sc<) differt foliolis supra non nitidis,
venis lateralibus non prominente elevatis, inflorcscontia
eonstanter pubescenti, et crescit in solo volcanico.

Rounded to spreading shrub, to 7 m tall, most
branches ascending to erect, the stems densely pu¬
bescent with spreading hairs, canescent especially
on newer growth. Leaves opposite to subopposite,
4-ranked, odd-pinnately compound; petioles gray,
tomentose, 5-1 I mm long; lateral petiolnles 1-3
mm long, the terminal one 2—5 mm long; leaflets 3
to 9, 7—21 mm long, 5—20 mm wide, the leaflet

Novoim 15: 350-353. 2005.

Volume 15, Number 2
2005

Rebman & Chiang
Amyris carterae from Mexico

351

Figure I. Amyris carterae J. Rebman & F. Chiang. — a. Stem segment with leaves, inflorescence, and variable leaflets,
scale bar = 1 cm. — b. Leaf, bar = 1 cm. — e, d. f lower, bar = I mm. — e. Part of paniculate inflorescence, bar =
1 cm. — f. Pubescence on petiole and raeliis, bar = 2 mm. — g. Mature fruit, bar = 1 mm. Drawn from the type
material (Rebman 5634) except for the fruit (g), which was drawn from Rebman 6 038.

352

Novon

shape variable, mostly ovate, but also broadly el¬
liptic or orbicular, the margin cremate or rarely en-
t i re. the apex acute to obtuse, rarely emarginate,
the base cuneate to rounded, sometimes slightly
cordate; upper leaflet surface dark green, densely
brown punctate, puberulent but glabrescent with
age, the lower surface gray to light green, sparsely
punctate, densely pubescent and velvety. Inflores¬
cence a terminal or axillary panicle, densely pu¬
bescent and velvety. Flowers small, perfect; calyx
light green, densely pubescent, with 4 triangular
lobes to 1 mm long; petals 4, Iree, white, obovate,
2—3 mm long, with a central patch of oil-gland dots
and simple hairs on the abaxial surface of each
petal; stamens 8, of which 4 are shorter in length
and alternate with longer ones, separate; anthers
with longitudinal dehiscence. Fruit a 1 -seeded
drupe, ellipsoid to ovoid, black at maturity, 9—13
X 8—1 I mm. puberulent with simple, spreading
hairs.

Phenology. Flowering September to November.

Distribution and habitat. Endemic to the state
of Baja California Sur; elevations 390—1000 m; oc¬
curring in canyon bottoms and lower slopes on vol¬
canic substrates usually associated with vegetation
dominated by Lysiloma divaricatum Bentham, hints
palmeri S. Watson, Celtis reticulata Torrey, Sebas-
tiania hilondaris S. Watson, Stenocereus thurberi
(Engelmann) Buxbaum, and Fouquicria diguetii
(Tieghem) I. M. Johnston. Other plant species pre¬
sent in the canyon community with Amyris carterae
include: Acacia kelloggiana A. M. Carter & Rudd,
Aculypha subviscida S. Watson, Bursera laxifiora S.
Watson. Carlowrightia pectinata Brandegee, Car-
minatia tenuiflora DC., Croton sonorae Torrey, Di¬
em urns alternifolius (S. Watson) Cline & Bray,
Eupatorium pcninsularc Brandegee, Ferocactus pen-
insulae (Engelmann ex Weber) Britton & Rose var.
peninsulae, Galium rnoranii Dempster subsp. acu-
leolatum Dempster, Hedyotis brevipes (Rose) W. II.
Lewis, Jatropha cinerea (Ortega) Muller Argovien-
sis, ./. vernicosa Brandegee, Justicia candicans
(Nees) L. I). Benson, Priva lappulacea (L.) I’ersoon.
Pseudabutilon thurberi (A. (fray) Fryxell, Solatium
americanum Miller. Tithonia thurberi A. Gray, Tour-
nefortia volubilis L., and Vallesia laciniata Bran¬
degee.

Etymology. This new taxon honors the late An-
netta Carter, an enthusiastic botanist who contrib¬
uted greatly to our Holistic knowledge of Baja Cal¬
ifornia Sur. Carter collected intensively (including
paratypes of this new species) in the Sierra de la
Giganta, where together with the Sierra Guadalupe
are the only known localities for this new taxon.

The common name of this species that is used by
the ranchers at Rancho Pie de la Cuesta near the
type locality is “gobernadora." This common name
is normally used, however, for Larrea tridentata
(Sesse & Moyifto) Coville in most of Mexico in¬
cluding parts of the Baja California peninsula.

Amyris carterae is most similar to Amyris mad-
rensis S. Watson, which it superficially resembles
in most aspects, but it does not have an upper leaf
surface that is glossy or lustrous and the lateral
veins are not prominent or obviously raised. The
populations of A. carterae occur on volcanic sub¬
strates in northwestern Mexico, quite disjunct from
those of A. madrensis, which are restricted to lime¬
stone substrates in central and northeastern Mexico
and adjacent Texas (Standley, 1923; Wilson, 191 1).
There is, in addition, a report of A. cf. madrensis
from the state of Jalisco (Lott. 1993). on the Pacific
coast. A consistent feature of Amyris carterae is the
pubescence, always present in the inflorescence
and young leaves, the leaves becoming glabrescent.
whereas in A. madrensis the pubescence may be
present or absent.

Paratypes. MEXICO. Baja California Sur: Sierra de
la Giganta W of Loreto, betw. Arroyo de las Parras &
Arroyo de Santo Domingo, 30 Nov. 1950, Carter & Kellogg
2844 (MEXU); GatVm del Cayuco, E base of Cerro de la
Giganta, 7 Oct. 1951, Carter & Kellogg 2124 (MEXU);
betw. La Victoria & Rancho Santa Marfa, W of Notrf. 22
Mar. I960, Carter & Ferris 2960 (MEXU); S side of Valle
de los Encinos. ft Nov. I960, Carter 4129 (MEXU); cafi-
ada S of Rancho de Ixis Encinos, Valle de los Encinos, 7
June 1963. Carter A' Reese 4558 (MEXU); vicinity of Ojo
de Agua del Carrizal, N from Rancho Viejo, 22 Sep. 1965,
Carter 5005 (MEXU); Sierra Guadalupe, near “Los Enci¬
nos,'" SW of Rancho San Sebastian, 29 Nov. 1953, Carter
& Kellogg 2272 (MEXU); Arroyo de los Gueribos, 25 Oct.
1997, J. ,/. Ptsrez Navarro 1054 (HCIB, SI)); W of Muleg£
at San Borjitas archeological area, 20 Sep. 1997, Rebman
4251. IV. Roberts & IV. Villavicencio (BCMEX, HCIB,
MEXU, SD); Canyon Guadalupe, just SW of Rancho Pie
de la Cuesta, betw. San Jos£ de Magdalena & ex-Mision
Guadalupe, 29 Oct. 1997, Rebman 4674 (MEXU, SI)); W
of MulegG W side of Cumbra de San Pedro betw. ex-
Mision Guadalupe & San Juan do las Pilas, 29 Apr. 1998,
Rebman 5241 A' J. L. Zuniga (SI)); Guadalupe Canyon,
E of the ex-Mision Guadalupe & near Rancho Pie de la
Cuesta, 13 Oct. 1999, Rebman 6028. I ). Silverman, M.
Wellik, & C. Graf (BCMEX, SD).

Acknowledgments. This work was supported in
part by funds from the National Geographic Society
(Grant # 6022-97) and from Sven Lindblad and
Special Expeditions, who funded the 1997 Lind¬
blad Binational, Multidisciplinary Expedition to
central Lower California organized by the San Di¬
ego Natural History Museum. Our gratitude is ex¬
tended to the SD Herbarium staff (especially Judy
Gibson) and volunteers lor their continued assis-

Volume 15, Number 2
2005

Rebman & Chiang
Amyris carterae from Mexico

353

tance with florist ic research projects. A special
thanks goes to Free I Roberts for supplying the bo¬
tanical illustration. The first author is indebted to
the many people who have joined and helped him
with fieldwork on the1 flora of Lower California, es¬
pecially Jose Luis Leon de la Luz at HOB and .lose
Delgadillo at BCMEX, both invaluable friends and
colleagues. A special thanks to the appropriate
Mexican authorities for granting permits and to Ex-
equiel Ezcurra and Patricia Beller for their efforts
in obtaining the papers and permits needed to con¬
duct field research and to collect plant species in
the region.

Literature Cited

Gereau. It. 1991. El genera Amyris (Rutaeeae) en America
del Sur, con dos espeeies nuevas de la Amazonia oc¬
cidental. Candollea 46: 227—235.

Lott, E. J. 1993. Annotated checklist of the vascular flora
of the Chamela Bay region, Jalisco, Mexico. Occas. Pap.
Calif. Acad. Sci. 148: 1-60.

Rebman. J. 2001. Succulent diversity in Lower California,
Mexico. Cact. Succ. J. (U.S.) 73: 131-138.

Standley, P. C. 1923. Rutaeeae. In: P. C. Standley. Trees
and shrubs of Mexico. Contr. U.S. Natl. Herb. 23: 524—
538.

Wiggins. I. 1980. flora of Baja California. Stanford Univ.
Press, Stanford, California.

Wilson, P. 1911. Rutaeeae. N. Amer. I I. 25: 174-224.

Tigridia suarezii (Iridaceae, I igridieae), a New Species from

Jalisco, Mexico

Aaron Rodriguez and Luis Ortiz-Catedral

Departamento <le Botdnica y Zoologfa, Universidad de Guadalajara, Apartado Postal 1-139,
45101, Zapopan, Jalisco, Mexico. rca08742@eueba.udg.mx

Abstract. Tigridia suarezii (Iridaceae, Tigri¬
dieae) is a new species from Mexico. Ms distin¬
guishing features are l lit- dark purple, almost Mack
color of the spreading tepal limbs and the white to
cream floral cup with dark purple spots and stripes.
It is morphologically similar to T. alpestris subsp.
obtusa and T. venusla, from which it differs in the
more robust habit of the plant, the shape, length,
and width of the tepals, and the length of the an¬
ther, filament column, and style branches. Tigridia
suarezii and T. venusta inhabit the understory in
pine-oak forests in an altitudinal range from 2080
to 2800 in, whereas /.' alpestris subsp. obtusa grows
on exposed rocky slopes above 3000 m. Tigridia
suarezii is known only from the state of Jalisco.

Best m KIN. Tigridia suarezii (Iridaceae, Tigridieae)
es una especie nueva para Mexico. Se caracteriza
por el color purpura, easi negro de la parte distal
de los Mpalos y el color bianco o crema con man-
chas o rayas purpuras de la copa floral. Su morfol-
ogia es similar a 7. alpestris subsp. obtusa y T. ven¬
usta de las que se distingue por su habito mas
robusto y en la forma, longitud y aneho de los t£-
palos. lambien son diferentes en la longitud de las
anteras, la columna estaminal y las ramas del es-
tilo. Tigridia suarezii y T. venusta crecen en el so-
tobosque del bosque de pino y encino en mi range
altitudinal de 2080 a 2800 m. En contraste, T. al¬
pestris subsp. obtusa crece en acantilados roeosos
por encima de los 3000 in de altitud. Tigridia suar¬
ezii se eonoce unicamente del estado de Jalisco.

Keywords: Iridaceae, Jalisco, Mexico, Tigridia.

The iris family, Iridaceae, was divided into four
subfamilies, Isophysidoideae, Nivenioideae, Ixo-
ideae, and Iridoideae, by Goldblatt (1990). Simi¬
larly, the subfamily Iridoideae was divided into four
tribes: Tigridieae, Sisyrinehieae, Irideae, and Mar-
ieeae. The tribe Tigridieae comprises the bulbous
and plicate-leafed genera of Iridaceae and is found
exclusively in the New World. It lias radiated ex¬
tensively m temperate and Andean South America

Novon 15: 354-357. 2005.

and in Mexico. I igridieae include 19 genera and
approximately 140 species.

Tigridia Jussieu is the largest genus of Tigri¬
dieae, with 50 species. The group has a notable
disjunction with 37 species in Mexico and Guate¬
mala and 13 species in the South American Andes.
I hirly species are endemic to Mexico. Tigridia is
taxonomieally difficult and, since Molseed (1970),
no other monographic study for the genus has been
completed. In addition, this group is underrepre¬
sented in Mexican herbaria, and three species have
been described based on single collections. Floral
characters useful to define species mav be poorly
preserved when dried, making further taxonomic/
morphological analysis problematic.

In the course of systematic studies of the tribe
I igridieae. Rodriguez et al. (1990) reported a new
locality for Tigridia alpestris Molseed subsp. obtusa
Molseed in the state of Jalisco. Phis species was
previously known only from its type locality in the
Cerro de Taneftaro in the state of Michoacan (Mol-
seed. 1970; Espejo & Lopez-Ferrari, 1996); how¬
ever. a direct comparison with fresh material was
not possible at that time. A collecting trip in August
2002 gave us the opportunity to compare fresh ma¬
terial of T. alpestris subsp. obtusa from the type
locality and these plants from Jalisco. An analysis
of floral characters led us to conclude that the
plants from Jalisco belong to an unrecognized spe¬
cies, which is described below.

Pigruliu suarezii Aaron Rodriguez & L. Ortiz-Ca¬
tedral, sp. nov. TYPE: Mexico. Jalisco: Mpio.
de Tapalpa, camino de Juanacatlan a Tepee,
2330 m, 20 Aug. 1987, A. Rodriguez & ./. C.
Suarez- Jaramillo 972 (holotype, IBUG; iso¬
types. ENCB, MEXU, MO, IJAMIZ). Figure I .

folia husalia nulla, folia caulina hinaria; flores plures,
erecti, crateriformes; cupula alba atropurpureo maculata;
tepala externa ovato-elliptica, parte Itasali eburnea, parte
supra cupulam patente atropurpurea; tepala interna ovata,
concava, unguiculata, obtusa, alba vel eburnea, maculis
atro-purpureis; glandula alba vel cinerascens ad tepalo-
rum partem basalem posita, cordata; antherae liberae ob-
longae ascendentes, per totam longitudinem fertile.s; styli

Volume 15, Number 2
2005

Rodriguez & Ortiz-Catedral
Tigridia suarezii from Mexico

355

f igure 1. Tigridia suarezii Aaron Rodriguez & I.. Ortiz-Catedral. — A. Habit. B. Mower, cross-view. (.. Mower,
lateral view. — D. Fruits. — K. Seeds. A. I). E, drawn from the type, Rodriguez & Suarez '>72 (IBUG); B, C. drawn
from Rodriguez el al. 2058 (IBUG).

356

Novon

I able I. Morphological comparison of Tigridia suarezii and related taxa.

Character

Stern length (cm)
f lower diameter (cm)
f lower color

f lower position
Outer tepal length (mm)
Outer tepal width (mm)
Inner tepal length (mm)
Inner tepal width (nun)
Column length (mm)

Anther length (mm)

Style branches length (mm)
Nectary bands length (mm)
Nectary bands width (mm)

T. suarezii

58-80
2.6-3. 5

dark purple stripes and
spots on a creamy white
background
erect
20-23
8-10
14—16
9-10
7-7.5
7—7.5

5— 5.5
9-9.5

6- 8

T. a I pest ns
subsp. oblusa

30-47

2

uniformly dark purple with
some pale yellow stripes

erect

12.5-18
6-7.5
1 1-13

4— 6.5

5.4— 6

5- 6
3-4

5.5— 7
4.4-5

T. venusta

33-60

4

dark purple stripes and
spots on a creamy white
background
erect or secund
22-25
10-1 1
12-15
6-10
4-5
12-13

9.5-1 1.5
4
3

ramuli bipartiti. ad apicem deelinati, eburnei; mucro in
sinubus styli ramorum instruetum; capsula oblongo-cla-
vata; semina pyriformia eastanea.

Krect. bulbous perennial herb, stein 58-80 ein
high, with 2 or 3 branches, glabrous; bulb ovoid,
d— 7 X 2-3 cm. the outer tunics thin, pale brown.
Masai leaves none; cauline leaves 2. linear, the low¬
er 50- 1 1 3 X 1 .6-0.2 cm. upper one shorter, 10—
25 X 0.f>— 2.4 cm. Inflorescence in a rhi pidium with
boat-shaped subequal spathes, 5.5— 6.5 X 0.7 cm.
Flowers 5 to 7 per rhipidium, erect, bowl-shaped,
2. 6-3. 5 cm diam.; outer tepals ovate-elliptic,
clawed, basallv cream to white, distally dark pur¬
ple, 20-23 X 8-10 mm; inner tepals ovate, con¬
cave, clawed, obtuse, with dark spots and stripes
over a white to creamy background, 14—16 X 9—
10 mm; nectary at base of inner tepals, cordate,
while to grayish, 9—9.5 X 6—8 mm; filaments con¬
nate in a column, 7—7.5 mm long; anthers oblong,
ascending, <—7.5 nun long; style branches 5—5.5
mm long, deeply bifid into 2 style arms slightly
declined at apex, ivory-colored; an obvious mucro
inserted between style arms; ovary 7—8 mm long,
fruit a capsule, oblong-clavate, 25—35 mm long;
seeds brown pyriform with a chalazal crest.

Tigridia suarezii is most closely related to T. al-
pestris subsp. obtusa. I hey share the dark purple,
nearly black color of the spreading tepal limbs.
Moth species have erect flowers, exposed nectaries,
and a mucro between the style arms. However, T.
suarezii differs from 7. alpestris subsp. obtusa m
having a more robust habit and larger and broader
leaves. In addition, the (lowers ol T. suarezii are
larger than those of T. alpestris subsp. obtusa. The
tepals of /. alpestris subsp. obtusa are dark purple

to wine-colored with pale yellow stripes around the
floral cup, whereas /.’ suarezii presents white to
cream tepals with dark spots and stripes at their
bases. The inner tepals of T. suarezii are ovate while
those of I alpestris subsp. obtusa are elliptic to
lanceolate. The filament column is longer in T.
suarezii than in T. alpestris subsp. obtusa. Finally,
there are differences in their habitat preferences in
Mexico. Tigridia alpestris subsp. obtusa grows on
exposed rocky slopes above 3000 m surrounded by
fir-pine forest in the state of Michoaean in west-
central Mexico, whereas /.' suarezii inhabits the un¬
derstory in pine-oak forests in an altitudinal range
from 2080 to 2800 m in the state of Jalisco in west-
mil Mexico.

The outer tepals of Tigridia suarezii are distally
dark purple. This color pattern resembles that of T.
venusta ; however, the outer tepals of T. suarezii art*
ovate-elliptic, while those of T. venusta are oblong.
Moreover, the anthers of T. venusta are fertile only
in their apical portion, whereas those of T. suarezii
are fertile along their full length. Lastly, the style
branches for T. venusta are about twice as long as
the style branches for T. suarezii. I he distinguish¬
ing characteristics of T. suarezii , T. alpestris subsp.
obtusa , and T. venusta are presented in Table I.

Distribution and habitat. Tigridia suarezii is
known only from the Sierra de Tapalpa and Cerro
Viejo in the state of Jalisco in western Mexico.
Common associated elements are Arbutus L. and
Rh us ( lournefort) L. The soil is predominantly silty,
black and rich in organic matter. The blooming pe¬
riod extends from August to October.

Etymology. I be specific epithet honors Jose C.
Suarez-Jaramdlo, first collector of the species and

Volume 15, Number 2
2005

Rodriguez & Ortiz-Catedral
Tigridia suarezii from Mexico

357

a noted enthusiastic collector and photographer of
Mexican Tigridia.

Paratypes. MEXICO. Jalisco: Amacueca, cam i no en-
tre Juanaeatlan y Tepee, 20°3'0"N, I03°30'0"W, A. Rod¬
riguez 2800 (ENCB, IBUC, IEB. MEXU, MO, UAMIZ,
WIS, XAL); Chiquilistlan, brecha Tapalpa-Chiquilistlan a
1 km de la Ifnea municipal, .4. Rodriguez & ,/. Suarez 532
(ENCB, IBUG, MEXU); brecha Tapalpa-Chiquilistlan,
Rodriguez & Suarez 073 (CHAPA, ENCB, IBUG, IEB,
MEXU, MO. NY. UAMIZ, WIS, XAL); laderas del cerro
El Chichimeco, brecha Tapalpa-Chiquilistlan. 1.5 km N\\
de la torre de estacion de microondas Chiquilistlan,
20°0'0"N, 103°47'0"W, Rodriguez 2801 (ENCB. IBUG.
IEB, MEXU, MO. NY. UAMIZ, WIS, XAL); Jocotepec,
Cerro Viejo vere da de la Ventanilla a el Kilo, 20°21 '42"N,
1 03°26'27"W, ./. 4. Machuca 8983 (ENCB, IBUG, IEB.
MEXU); Tapalpa, brecha entre Juanacatlan y Tepee, Rod¬
riguez & Suarez 585 (IBUG), 1577 (ENCB, IEB, IBUG,
MEXU); I 1.5 km a partir de Tapalpa sobre la brecha a
Chiquilistlan, 20°0'06"N, 103°49'05"W, A. Rodriguez et
al. 3058 (IBUG).

Acknowledgments. We thank Jorge Alberto
Perez dr* la Rosa from the Universidad de Guada¬

lajara and Ignacio Garcia Ruiz from the Centro In-
terdisciplinario de Investigacion para el Desarrollo
Integral Regional, Unidad Michoacan, for their
support in the field. We also thank Antonio Ma¬
chuca who gave us the opportunity to analyze ma¬
terial collected bv him. The illustration was made
by Luis Ortiz-Catedral.

Literature Cited

Espejo-Serna, A. & A. K. Lnpez-Ferrari. 1996. Las Mon-
oeotiledoneas Mexicanas, una Sinopsis Elorfstica I. I As¬
ia de Referenda Parte VI. Consejo Naeional de la Flora
de Mexico A. G., Universidad Autonoma Metropolitana-
Iztapalapa. Comision Naeional para el Conocimiento y
Uso de la Biodiversidad, Mexico, I).F.

Goldhlatt, P. 1990. Phylogeny and classification ol Irida-
eeae. Ann. Missouri Bot. Gard. 77: 607—627.

Molseed, E. 1970. The genus Tigridia (Iridaeeae) of Mex¬
ico and Central America. Univ. Calif. Publ. Bot. 54: I —
I 13.

Rodriguez, A.. (). Vargas, E. Villegas & k. J. Sytsma.
1996. Nuevos informes de iridaceas (Tigridieae) en Jal¬
isco. Bob Inst. Rot. Univ. Guadalajara 4: 39—17.

A New Species of Microlicia (Melastomataceae) from
Minas Gerais, Brazil

Rosa n a Romero

Instituto de Biologia, Universidade Federal de Uberlandia, Caixa Postal 593,
(d'd 38400-902, Uberlandia, Minas Gerais, Brazil, romero@inbio.ufu.br

Absthact. A new species in Microlicia (Melas-
tomataeeae) is described and illustrated. Microlicia
tenuifolia is known from I lie rocky slopes of Dia-
mantina, Minas Gerais state, an important center of
endemism (or Melastomataceae species. It resem¬
bles morphologically M. isophylla DC., but it can
be distinguished by the smaller leaves and pedicels
and calyx lobes with an acute uon-apieulate apex.

B l- si mo. L ina nova especie de Microlicia (Melas¬
tomataceae) e dcserita e ilustrada. Microlicia ten-
uijolia ocorre nos campos rupestres dos arredores
de Diamantina, no eslado de Minas Gerais, urn mi-
portante centro de endemismo para especies de
Melastomataceae. Assemelha-se moHologieamente
a M. isophylla DC., mas pode ser distinta pelas
folhas e pedicelos menores e lacmias do ealiee com
apice agndo, nao apiculado.

hey words: Brazil, “eampo rupestre," highland
vegetation, Melastomataceae, Microlicia tenuifolia,
Minas Gerais.

During the course of a taxonomic study of the
Neotropical species of Microlicia I). Don, a new
species from Diamantina, Minas Gerais state, Bra¬
zil. was discovered. I lie genus consists of approx¬
imately 100 endemic species in the Brazilian states
ol Minas Gerais, Goins, and Bahia (Romero, 2003)
with a few outlying taxa in Bern (Brako & Zarucchi,
1993), Bolivia and Guyana (Wurdack & Renner,
1993), and Venezuela and Colombia (Berry et ah,
2001). Species of Microlicia are mostly shrubs or
subshrubs, with 5-merous, pedicellate to subses-
sile, solitary flowers, and capsular fruits with de¬
hiscence longitudinal from the apex to the base.

Microlicia tenuifolia R. Romero, sp. nov. TYPE:
Brazil. Minas Gerais: estrada para Conselheiro
Mata, a 14 km da rodovia, 13 Dee. 1980, /.
Cordeiro, l\. L. Menezes, ./. R. Pirani & A. Par¬
ian 591 (holotype, HUFU; isotypes, MO. RB,
SPF, UEC). Figure 1.

Sullrutex 0.4—1 in alias, caulis teres glaber; ramuli
quadrangnlali punctato-glanclulosi. laminae sessiles er-

Novoin 15: 358-360. 2005.

('ctae elliptieo-laneeolatae 1—3 X 0.2— 0.5 mm margine in-
tegerrima utrinque densiuscule impresso-punctata, 1-3-
nervae. Mores solitarii; pelala magenta apiee acuminata;
ovarium 3-loculare.

M. isophyllae DC. maxime aflinis sed praecipue lamina
et pedicello brevioribus diversa.

Subshrub. 0.4—1 m tall, branching dichotomous-
ly or trichotomously, young branches fastigiate;
stein terete, glabrous; young branches quadrangu¬
lar, glutinous, covered with sessile glands, older
branches brown, quadrangular, leafless proximally,
leaf scars conspicuous, glabreseent. Leaves sessile,
blade erect, adpressed, rarely slightly spreading, I —
3 X 0.2— 0.5 mm, lanceolate to elliptic, apex acute
or obtuse, base truncate-obtuse, margin entire, cov¬
ered with sessile glands on both surfaces, glutinous
or not, I- to 3-nerved, with inconspicuous lateral
veins, flowers 5-merous, solitary, terminal; pedi¬
cels 0.5—1 mm long; hypanthium 2-3 X 1.5-2. 5
mm, campanulate, 5- to 10-costate, rugose, gluti¬
nous. covered with sessile glands, calyx lobes 1.5-
2.5 mm long, triangular-lanceolate, apex acute, not
apieulate, covered with sessile glands; petals 6—10
X 4-6(-8) mm, magenta, obovate, apex acuminate,
margin entire; stamens 10. dimorphic, larger sta¬
mens 5. the filaments 1—2 mm long, purple, the
thecae 1.0- 1.5 mm long, purple, ovate to oblong,
not corrugate, with rostrum ca. 0.2 mm long, white,
the connective prolonged 1.5-2 mm. purple, with
ventral appendage ca. 1.0 mm long, yellow, ex¬
panded and slightly bilobed; smaller stamens 5. the
filaments 1.5-2 mm long, purple, the thecae ca. I
mm long, yellow, ovate, not corrugate, with rostrum
ca. 0.2 mm long, the connective prolonged ca. 0.6
mm long, yellow, without ventral appendage; ovary
superior, 3-loeular, glabrous; style ca. 3 mm long,
filiform, glabrous, stigma punctiform. Capsule 3—4
X 2—3 mm, greenish brow n, globose; seeds 0.4— 0.5
X ca. 0.3 mm, numerous, slightly curved, foveolate.

Distribution and ecology. Microlicia tenuifolia
is characterized by being a subshrub with slender,
fastigiate branches, very small leaves, costate hy¬
panthium. long calyx lobes, and dimorphic sta¬
mens. Endemic to the region around Diamantina,

Volume 15, Number 2
2005

Romero

Microlicia tenuifolia from Brazil

359

Figure 1. Microlicia tenuifolia R. Romero. —A. Habit. —IF Petal. — C. Flower, hypanthium, and calyx lobes. — D
Detail of the branch and insertion of the leaves. — K. Leaf. — F. Lateral view of small stamen. — G. Lateral view of
large stamen. Drawn from the paratype Splett 622 (HR).

360

Novon

Minas Gerais state, where it is locally common in
eampo rupestre.

Phenology. Flowering specimens have been
collected from June !<• December; fruiting speci¬
mens have been collected in January, April, May.
August, September, and October.

I he fust gatherings of this species were made by
Auguste Gla/iou in 1892 from localities near Dia-
mantina, Minas Gerais state (Glaziou 19200.
19201). Gla/iou (1908) subsequently listed both
gatherings under Microlicia denudata Cogniaux.
However, this identification, although perpetuated
by most subsequent collectors/authors, is errone¬
ous. Microlicia denudata Cogniaux seems to be
known only from the type gatherings from Goids
( Gardner d7.il). Isotypes of Microlicia denudata
seen by the author at 8M and k have a hypanthium
covered with dense stalked glands mainly on the
apex, eventually with caducous glands, triangular-
lanceolate calyx lobes setose with acute apex, fil¬
iate margin, and lanceolate leaves with entire mar¬
gin and acute, apiculate apex, and are thus quite
different from the Gla ziou 19200 and 19201 gath¬
erings. rhese two latter gatherings represent a hith¬
erto undescribed species, which I describe herein
as Microlicia tenuifolia. Gatherings of Glaziou

19200 at C and k were determined as "M. denu¬
data var. brevifolia Cogniaux, and those of Glaziou

19201 at (,, k, and OUPR were determined as "M.
denudata var. fastigiata Cogniaux,” although nei¬
ther varietal name was validly published by Alfred
Cogniaux or indeed by anyone else.

Microlicia tenuifolia resembles M. isophylla DC.,
which occurs in Sao Paulo, Minas Gerais, and Rio
de Janeiro states. Roth have erect, lanceolate to
elliptic leaves with entire margins and both surfac¬
es covered with sessile glands. The leaves 1-3 X
0. 2-0.5 mm, pedicels 0.5-1 mm long, and calyx
lobes with an acute non-apiculate apex distinguish
M. tenuifolia from M. isophylla. Microlicia isophylla
has larger (3-7 X 1-2 mm) leaves, pedicels 1.5-2
nun long, and calyx lobes with an acute, apiculate
apex.

Puratypes. BRAZIF. Minus Gerais: Diamuiitina, s.d.,

A. Glaziou 19200 (C, k. P — 3 sheets), 9 Apr. 1892. A.
Glaziou 19201 (C, F, k. OUPR, P, l{, US); Conselheiro
Mala, Lama Preta, June 1934, Rrade 12717 (US); ca. 12
kin SVt ol Diamantina, 18 Jan. 1969, II. S. Irwin. R. R.

Santos. R. Souza <£- F. Fonseca 22190 (k, NY, KB, US);
Cuinda, 7 Sep. 1971. G. Hatscbach 27204 (HUFU, MBM.
US); Planalto Diamantina, 7 Oct. 1972, A. P. Duarte
14026 (US); eslrada Diamantina-Corinto, 1 Dee. 1976, G.
J. Shepherd. J. R. Andrade. I.. S. Kinoshita & ./. K Vani¬
sh iro 2290 (NY. UFC); 24 km na eslrada Diamantina—
Conselheiro Mata, 30 Aug. 1981,4. VI. Giulietti, N. M. L
Menezes, ./. Semir & A. ./. Mattos 1821 (HUFU, SPF, UFC,
US); km 66 da eslrada Curvelo— Diamantina, Serra do Bre-
jo Grande, 9 Apr. 1982, A. Furlan, L. Rossi, N. M. L.
Menezes, N. Ilensold. II. /,. Wagner & E. M. Isejima 2252
(SPF US); eslrada Diamantina-Conselheiro Mata, km 174,
10 Sep. 1986, N. L. Menezes, T. li. Cavalcanti A' ./. C. C.
Gonsgalves 10298 (HUFU, SPF. US); eslrada Diamantina-
Conselheiro Mata, 23 Sep. 1994, 20.3 km depois do as-
falto, Splett 622 (SPF. UB. US); eslrada para Gouveia, 1 1
Oct. 1996, W. Marcondes- Ferreira, A. 0. Simoes & R. Re-
linello 1247 (UFC); eslrada para Gouveia ca. 10 km de
Diamantina, 3 Oct. 1997, M. L Kawasaki A- A. Rapini
1055 (IlllfU, SP); eslrada Diamantina— Gouveia, ca. 12
km sul de Diamantina, 30 Mar. 2001. II. Ronu l TO & J. A.
Nakajima 6065b (HUFU, NY, UFC); eslrada Diamantina-
Conselheiro Mala (km 162), ea. 29 km do trevo, 1 Apr.
2001, R. Romero & J. N. Nakajima 6095 (HUFU).

Acknowledgments. I am grateful to the Univer-
sidade Federal de Uberlandia for the facilities of
the Herbarium Uberlandense (HUFU), the curators
of R. RHCB, C, F. HR. HUFU, MRM, l\Y, OUPR,
R, RB, SP, SPF, UR, UFC, and US for loans of
specimens, Conselho Nacional de Desenvolvimento
Cientffico e lecnologico for a research grant, the
Margaret Mee Foundation for a fellowship to visit
BM, k. P, and W, and Maria Cecilia Tomasi for
preparing the illustration. I thank Peter Gibbs for
correcting the Fmglish and for helplul suggestions,
and John Pruski for useful suggestions and for cor¬
rections to the draft.

I.ilerature Cited

Berry, P., k. Yalskievych & B. k. Holst. 2(H) I . Flora of
the Venezuelan Guayana, Vol. 6, Liliaceae-Myrsina-
eeae. Missouri Botanical Garden Press, Si. Fouis.

Brako, F. & J. F. Zarucehi. 1993. Catalogue of the F low¬
ering Plants and Gymnosperms of Peru. Monogr. Syst.
But. Missouri Bot. Card. 45.

Glaziou, A. F. M. 1908. Liste des plantes du Bresil Cen¬
tral reeueillies en 1861-1895. Melaslomatae^es. Bull.
Soc. Bot. France 54, Mem. 3; 240-298.

Romero, R. 2003. A newly described species ol Microlicia
(Melastomataeeae) from Minas Gerais, Brazil. Novoir
1 16-1 18.

Wurdaek. J. J. N S. S. Renner. 1993. Melastomataeeae.
In: A. B. A. (flirts- Van Rijn (editor). Flora das Guianas,
Fascicle 13. Koeltz Scientific Books, Germany.

Plectranthus igniarioides (Lamiaceae), a New
Species from Somalia

( Hof Ryding

The Botanical Garden and Museum, Gothersgade 130, DK-I 123 Copenhagen k. Denmark.

olofr@snm.ku.dk

ABSTRACT. Plectranthus igniarioides Ryding, a
new species from northern Somalia, is described
and illustrated. Comparison is made with P. argen-
tifolius Ryding, P bariensis Ryding. P. cuneatus (E.
O. Baker) Ryding, P gillettii J. k. Morton, I’, ig-
niarius (Scliweinfurth) Agnew, and P. puberulentus
J. k. Morton. The circumscription of Plectranthus
gillettii, which has been reported to occur in So¬
malia. is refined to include only the material known
from Ethiopia and Kenya.

Key words: Labiatae, Lamiaceae, Plectranthus.
Somalia.

The genus Plectranthus L’Heritier (including Co¬
leus Loureiro and Capitanya Giirke) belongs to
Lamiaceae subfamily Nepetoideae tribe Oeimeae.
It contains about 300 species and is most frequent
iti tropical and southern Africa and Madagascar,
but also occurs in tropical Asia and Australia.
There are 17 species known from Somalia. The So¬
mali material, identified as P igniarias (Schvvein-
furth) Agnew by the collectors, includes two anom¬
alous specimens. When Morton (1998) described P.
gillettii as a new species, he cited these two more
or less leafless Somali specimens as paratypes,
along with material from Ethiopia and Kenya. The
Somali material is, however, geographically some¬
what isolated from the rest of that species. Mats
Timlin recently collected leafy, but otherwise very
similar, plants from the same region in Somalia,
flie characteristic shape of the leaves, as well as
other features of the plants, strongly suggests that
these specimens are distinct from both P. igniarius
and P gillettii and belong to a different hitherto
undescribed taxon. It is described here as a new
species.

With the exclusion of the Somali collections in
question, P. gillettii is more clearly delimited and
is restricted to a more coherent distribution area in
Ethiopia and Kenya. However, the modification of
the circumscription hardly affects Morton's (1998)
description of this species.

Plectranthus igniarioides Ryding, sp. nov.
TYPE: Somalia. Sanaag Region: 43 km from
Ceel Afweyn on road to Ceerigaabo, l()°16'N,
47°12'E, ca. 1480 m. 30 Jan. 2002, M. Thulin
I076H (holotype, EPS, photos at C, MO: iso¬
type, K). Figure I.

Differt ah aliis speciebus cognatis Plectranthis combi-
natione constanti ex foliis inlegris subsessilibus utrinque
ineanis plerumque decicluis sub antbesi, lobo supero ra-
lveis fructicantis cupulato, lobis centralibus labii superi
corollarum fimbriis.

Shruhlet. 0.4-2. 5 m tall, strongly aromatic; young
stems woody-fleshy, hoary; eglandular hairs 0.1-
0.15 mm long, ± recurved, stalked glands slightly
longer, sessile glands orange or colorless; old stems
woody, bark gray. Leaves decussate, mostly decid¬
uous before anthesis, elliptic, up to 33 X 15 mm,
1.7-2. 8 times as long as broad, grayish, hoary es¬
pecially below, with orange sessile glands on both
surfaces; apex obtuse to subacute; base with de-
current margins, petiole up to 2 mm long: margin
entire. Inflorescences thyrsoid, lax, verticillasters 2
to 6, simple, lateral branches short or apparently
gathered into a branched thyrse; bracts ovate, acu¬
minate to acute, cupular, 1-2 mm long, strongly
orange-glandular, quickly deciduous; cymes 1- to
5-flowered. peduncle 0-1.5 mm long; pedicels 6-
13 mm long, spreading, densely glandular with
shortly stalked and larger orange to colorless sub-
sessile glands. Calyx at anthesis 2—3.5 mm long,
upper lobe violet and rather sparsely glandular,
elsewhere violet or green, very densely glandular
with shortly stalked and larger orange or colorless
subsessile glands; corolla 2-lipped, 11—16 mm
long, dark blue to purple; indumentum sparsely
glandular and eglandular; tube base sigmoid,
slightly widened toward the throat, prolonged at the
insertion of the lower lip, 7.5—9 mm long; upper lip
4-lobed, mid-lobes much broader than the lateral
lobes, subtruncate, minutely erose; lower lip much
longer than the upper lip, 5—7 mm long, deeply
boat-shaped, slightly darker than the rest of the co¬
rolla; stamens 4, shorter than the lower lip of the
corolla during early anthesis, becoming longer than

Novon 15: 361-363. 2005.

362

Novon

figure 1. Fleet rant hits igniarioides Ryding. — A. Branch will) leaves (the n pper part of the specimen). — B. Branch
with inflorescences. — C. Calyx at the anlhesis. — 1). Corolla at early anthesis with the stamens short. — E. Young
gynoeeium of a flower bud; s = style; o = ovary lobe; d = disc. — E Anther of an anterior stamen seen from above.

C. I lie same anther seen from below. — H. Fruiting calyx. — I. Map of Somalia showing the known distribution of
I • igniarioides. A— G drawn from Tliulin 10768 (l l*S); II drawn from (Hover A' Gilliland 6 42 (K).

Volume 15, Number 2
2005

Ryding

Piectranthus igniarioides from Somalia

363

the lower lip of the corolla, connate along 1-2 mm,
anterior pair formation slightly longer; anthers be¬
coming depressed at the back by the insertion of
the filament, ± incurved at the opening; anthers of
the posterior pair 0.6— 0.8 mm diam.; anthers of an¬
terior pair 0.7— 0.9 mm diam.; pollen mass white;
style exserted from the lower lip of the corolla, bifid;
disc abaxial lobe slightly enlarged. Fruiting calyx
enlarging to 7—8 mm long, less densely glandular
than (lower calyx, glabrous inside or sparsely glan¬
dular; pedicel insertion dorsal; upper lobe subor-
bicular to slightly oblate, apex acute, slightly mu-
cronate, becoming cupular, margins shortly
decurrent on the lube; lateral lobes 2, obliquely
subtriangular, slightly shorter than the ventral
lobes; ventral lobes 2, narrowly triangular, becom¬
ing 2—2.5 mm long; nutlets (immature?) obovoid, ca.
1.7 X 1.3 mm, smooth, grayish brown.

Distribution and habitat. Gypsum or limestone
hills or on hillsides over volcanic rock; altitude
1450—1850 m. Sanaag Region in Somalia (Fig. II).

The connate stamens (Fig. 1 1)), deciduous
bracts, and the absence of a ring of eglandular hairs
in the calyx throat indicate that P. igniarioides be¬
longs to the subgenus Coleus as defined by Codd
( 1975). The species is perhaps most closely related
to P. igniarius, which it resembles in its entire de¬
ciduous leaves and minutely erose mid-lobes of the
upper lip of the corolla. However, whereas P. ig¬
niarius has a unique calyx shape with a hooded
upper lip and rather short and broad ventral lobes,
P. igniarioides (fig. Ill) resembles most other Piec¬
tranthus species in having the upper calyx lobe cu¬
pular and shortly decurrent along the tube and the
two ventral lobes rather narrow. The species also
differs from P. igniarius in having densely hoary,
subsessile leaves. Piectranthus igniarius occurs in
eastern and northeastern Africa including Somalia
and on the Arabian Peninsula. Piectranthus igni¬
arioides is perhaps also closely related to the other
northeastern tropical African species that have a
similar shrubby habit and deciduous leaves, as dis¬
cussed above. In addition to lacking the character¬
istic leaf shape of P. igniarioides, P. gillettii differs
in being less hairy with slightly longer hairs, in

having the posterior stamens virtually free, and
probably having a paler and partly white corolla
with the tube more strongly widened toward the
throat. Piectranthus cuneatus (E. G. Baker) Ryding
and P. bariensis Ryding both differ from P. igni¬
arioides in having shorter pedicels (2—7 mm long)
and characteristic apically truncate or very broadly
rounded leaves. Piectranthus puberulentus .1. K.
Morton differs from P. igniarioides in having the
fruiting calyx slightly but distinctly hairy inside
and the pedicels densely pubescent with short
eglandular hairs.

Piectranthus igniarioides can be distinguished
from most other Piectranthus in having entire
leaves (Fig. 1 A). Minutely erose mid-lobes of the
upper corolla lip seem also to be a rare feature in
the genus (Fig. ID). Entire leaves, however, also
occur in the Somali and Yemeni P. argentifolius
Ryding. Piectranthus igniarioides differs from P. ar¬
gentifolius and other species of Piectranthus subg.
Calceolanthus in having the fruiting calyx almost
glabrous inside, while the species of Calceolanthus
have the calyx characteristically villous in the
mouth. Piectranthus igniarioides also differs from
P. argentifolius in being taller, woodier, and having
longer pedicels.

Paratypes. SOMALIA. Sanaag Region: Mas Altai,
1 1°00'N, 48°20'E, 7 Aug. 1957, Neubould 822A (k with
P. igniarius on the same sheet); at “Erigavo” (Ceerigaabo),
24 Jan. 1945, Clover & Gilliland 642 (K).

Acknowledgments. Material was loaned from
UPS and K, and material of the presumably related
species was examined during visits to ETH, UPS,
and K. I thank the directors and curators of these
herbaria. I am also indebted to Mats Thulin at the
University of Uppsala (Sweden) for informing me
about the interesting plants collected during his
journey in Somalia in 2002, and to the two review¬
ers, Peter Phillipson and Trevor Edwards (both
South African), for valuable comments on the man¬
uscript.

Literature Cited

Codd, L. E. 1975. Piectranthus (Labiatae) and allied gen¬
era in southern Africa. Bothalia 11,4: 371—442.
Morton, J. K. 1998. Two new species of Piectranthus from

northeast Africa. Kew Bull. 53: 997—999.

Waltheria berteroi (Sterculiaceae, Hermannieae),1 a New
Combination from Colombia and Venezuela

Janice G. Saunders

Instituto de Botdnica Darwinion, C.C.#22, B1642HYI) San Isidro, Pcia. Buenos Aires,
Argentina. jsaunders@darwin.edu.ar

Abstract. A new combination in Sterculiaceae,
Hermannieae, Waltheria berteroi (Sprengel) ,). C.
Saunders, is created and lectotypified. Waltheria
suheordata Standley from Colombia and Venezuela
is newly synonymized to W. berteroi on the basis of
shared bract, leaf, and inflorescence characters.
Additional isotypes of W. subcordata are newly cit¬
ed for Smith 493, and further clarification of non¬
type Smith 493 specimens is given. Waltheria ber¬
teroi has been confused with W. lophanthus J. B. &
C. Forster from the South Pacific Ocean but differs
in its leaf vestiture, leaf apices, stipules, bracts,
thrum stamen tube, and stigma, as well as char¬
acters of the fruit and seed. Apparent morphologi¬
cal intermediates of W. berteroi with W. glomerata
Presl or W. involacrata Bentham are noted to occur
outside the Colombian Sierra Nevada de Santa Mar¬
ta, western Eastern Cordillera, and Sahagun vicin¬
ities.

Key words: Colombia, Hermannieae, Malachra,
Malvaceae, Marquesas Islands, Society Islands,
Sterculiaceae. Venezuela, Waltheria.

A new combination is necessary for a primarily
Colombian species now called Waltheria suheordata
Standley (Sterculiaceae, Hermannieae). Sprengel
earlier described a Bertero collection of the same
species from Colombia as Malachra berteroi Spren¬
gel [as "herterii"\ in Malvaceae (1826: 94): "M.
cattle Jrut ieoso aspero tomentoso, foliis ovato-oblon-
gis (luplicato-dentatis supra stellato pubescentibus,
subtus venoso- retie ulat is tomentosis, racemis axillar-
ibus. involucri foliolis cuneatis coloratis nervosis. Ad.
Jl. Magdalen., Bertero." According to Lanjouw and
Stafleu (1954), Bertero collected in northern Co¬
lombia during 1820—1821; duplicates of these
specimens can be found at B. HAL, L, MEDEL,
MO, and S.

Garcke (1863), a curator at Halle (Stafleu &
Cowan. 1976), apparently examined the type of
Malachra berteroi (at that time at Halle, later trans¬

ferred to B in 1890 (Stafleu & Cowan, 1985), now
presumably lost) and concluded it was in complete
agreement with Sprengel’s original diagnosis. More¬
over. he concurred with the determination by Balbis
for a Bertero collection of the same plant (Berlin
Herbarium specimen E photo 9568 and Garcke,
1863) that Bertero’s collection was not a Malachra
but a Waltheria. Wheareas Balbis had designated
the plant to be W. lophanthus J. K. & G. Forster
and. non Willdenow (Willdenow, 1800) [nom. il-
legit., = W. tomentosa (J. R. & G. Forster) II. St.
John; see ./. K. & G. Forster s.n., K, F Berlin Her¬
barium specimen-photo 9568 and MO Bernhardi
Herbarium specimen 3261414], Garcke differed in
the species determination, stating that W. lophan¬
thus was a different species, but a valid combina¬
tion was not made. Giirke (1892), following Garcke,
excluded M. berteroi from Malachra but treated it
as a synonym of W. lophanthus J. R. & G. Forster.

As Garcke (1863) pointed out, Balbis and later
Giirke misidentified the Bertero specimens at the
species level. Waltheria tomentosa is endemic to
the Marquesas Islands and Society Islands of the
southeastern Pacific Ocean and differs from Spren-
gel’s species of Colombia. Waltheria berteroi is dis¬
tinct from W. tomentosa by its overall vestiture,
which is finely stellate-pubescent and shorter rath¬
er than woolly-pannose, stellate-tomentose; by its
leaf blades, which are ovate, with acute to acumi¬
nate-acute apices, with rounded to narrowly cordate
bases, and adaxiallv pubescent, abaxially tomen-
tose. rather than widely to very widely ovate, with
mostly obtuse-angled to obtuse-rounded apices,
with cordate to deeply cordate bases, and canescent
on both faces; by its stipules, which are narrowly
triangular-subulate and compressed, rather than
triangular and planar; by its less compact inflores¬
cences; by its narrowly oblanceolate or obovate-
spathulate bracts rather than narrowly triangular-
subulate bracts; by its longer thrum stamen
filament tube to 1 .5(2.0) mm rather than 0. 7(0.9)

1 Hermannieae including Waltheria. formerly in Sterculiaceae, is now placed in the Byttnerioideae basal to Grewioi-
deae, formerly in Tiliaceae, in an expanded Malvaceae sd. (Bayer & Kubitzki, 2003).

Novon 15: 364—367. 2005.

Volume 15, Number 2
2005

Saunders
Waltheria berteroi

365

mm long; by its more distended-plumose stigma; by
its pubescent rather than sericeous capsule; and by
its longer seed to 2.5 mm rather than to 2.2 mm
long (cf. Saunders, 1995: 330—337, 780—793). The
Colombian species was validly described as Wal¬
theria subcordata by Standley (1916), who was un¬
aware of the earlier name, based upon another type.
However, both types are conspecific, sharing the
distinctive loosely condensed inflorescences, showy
mostly distinct obovate bracts with obtuse apices
and cuneate bases, and leaf shapes.

Garcke (1863) noted that Sprengel erroneously
cited the provenance of the Bertero collection as
coming from the Rfo Magdalena. The labels of both
Sprengel’s sheet ( Bertero s.n.) and the duplicate an¬
notated by Balbis at B indicated “S. Marthe” (Santa
Marta, Colombia) with labels in Balbis's handwriting
(verified by reference samples in Burdet, 1972). It
is easy to understand the confusion about specific
locality data. The Bertero collections appear to have
been distributed without attached original label data,
as the only early labels on the F isotype photo and
MO specimen are in Balbis’s own handwriting bear¬
ing “S. Marthe,” written between 1820 and 1821
(probable collection dates of Bertero’s Colombian
collections listed in Lanjouw & Stafleu, 1954) and
1831. the date of Balbis’s and Bertero’s deaths (Sta¬
fleu & Cowan, 1976). Balbis moved to I, yon in 1814
from Torino (Stafleu & Mennega, 1992). and appar¬
ently visited the Berlin herbarium during the period
between 1820 and 1831, probably 1824, as noted,
but illegibly, by Kunth on the Berlin duplicate (see
F 9568, photo of B isotype [destroyed]). The source
for Sprengel’s locality information or even Balbis's
locality notations is elusive. Presumably, one or the
other was in correspondence with Bertero, or had
access to knowledge obtained about the localities.
An asterisk in Sprengel’s (1826) original description
indicates living material was studied, so perhaps
separate horticultural records account for the dis¬
crepancy in localities. The Magdalena River empties
into the ocean about 120 km southwest of the pres¬
ent city limits of Santa Marta in the Dpto. Magdalena
in Colombia, but is near to the general locality that
appears to have been broadly designated as Santa
Marta by early collectors.

Waltheria berteroi (Sprengel) J. C. Saunders,
comb. nov. Basionym: Malachra berteroi
Sprengel |as “berterii”], Syst. Veg. 3: 94. 1826.
TYPE: Colombia. Santa Marta, 1820—1821.
[C. G.j Bertero s.n. (lectotype, designated here,
MO 3261414; isotypes, F 9568, photo of B
isotype destroyed).

Waltheria subcordata Standley, Contr. U.S. Natl. Herb. 18:

I 17. 1916. Syn. nov. TYPE: Colombia. Masinga, 500
ft., 25 Oct. 1898-1901, II. H. Smith 49.3 (holotype,
US 533011; isotypes. A, E, F, MO, MPLJ, NY, S).

Distribution, habitat, and phenology: Northern
and eastern Colombia (Atlantico, Magdalena, Bo¬
livar, Cesar, Santander, La Guajira, Vichada) and
northwestern and central Venezuela (Falcon, Zulia,
Apure, Amazonas, Bolivar). It is mostly and best
represented (Hi berteroi s. str. ) from Colombia in
and below the western Sierra Nevada de Santa Mar¬
ta, up to 1600 m, along the western Eastern Cor¬
dillera. and near Sahagun. Elsewhere in Colombia
(Magdalena River mouth to Baranoa; El Cerrejon)
and Venezuela it is less representative ( IT. berteroi
s.l.) with intermediate forms (see below) particular¬
ly near the middle Orinoco River from Samariapo
to San Vicente. Shrubs to 3 m in tropical savannas
in openings or borders of dry forest, along rivers or
streams, roadsides, or in foothills, occurring from
10 to 1613 m elevation. Flowering October through
March, peaking in December and January, with
flowers presumably diurnal.

Additional isotypes of Waltheria berteroi have
been sought at L (absent), MEDEL, and S. because
Bertero’s Colombian collections are there (Lanjouw
& Stafleu. 1954). They were not sought at HAL, as
Sprengels’s herbarium at HAL was transferred to
Berlin (Stafleu & Cowan, 1985).

Additional conspecific collections given the
same number ( Smith 493, LL, MICH, NY, U. YV IS)
but from a different locality in Magdalena at Bonda,
250 ft., on a different date, 13 May, are excluded
from the type of Waltheria subcordata and noted
below in specimens examined. Ayers and Boufford
(1988) discussed the common assignment of the
same collection number from different localities
and dates for presumably conspecific plants in
Smith collections and established that his collec¬
tion data exist on NY material in most cases. Iso-
tvpes for W. subcordata from E, MPIJ, and S sup¬
plement those listed there. The non-type collections
of Smith 493 can be readily identified upon com¬
parison with the NY non-type specimen that bears
a handwritten collection label stating "Bonda." The
non-type specimens also differ by their smaller
leaves, thrum rather than pin flowers, and aspect
of more arid growing conditions. As Standley
(1916) noted, Smith 493 duplicates were distrib¬
uted from NY as W. involucrata Bentham, “narrow
bracted form.”

Riedlea berteriana Balbis ex DC., a synonym of
Waltheria glabra Poiret, is a similar epithet to ber¬
teroi, but would not be considered a homonym
(ICBN, Art. 53.3 ex. 10. Greuter et ah, 2000).

366

No von

Specimens that in my opinion are representative
of Waltheria berteroi s. str. are cited below. Other
Colombian populations of Waltheria berteroi s.l.
from the departments of Atlantic!) (e.g.. D ugand it'
Jaramillo 4034, US; Elias 195, F), and Guajira
(e.g., Bunch et at. s.n., MO), and Venezuelan pop-
n lat ions from Falcon (e.g.. Wingfield 6 009, CTES)
and Zulia (e.g., Nucette 164, CTES) appear to be
morphologically intermediate with W. glomerata
Presl by their compact, glomerate inflorescences
and smaller bracts. Waltheria glomerata, from
Meta, Guaviare, Boyacd, and Uribe in Colombia,
and Panama to Mexico, otherwise differs by its
white petals, its leaves often elliptical or rhombic,
later rugose, and its differently shaped bracteoles
and bracts, with more fusion. In contrast, the mid¬
dle Orinocan River-associated Venezuelan popula¬
tions of W. berteroi s.l. from Amazonas (e.g., Grdger
& Rare raft 295, MO; Stergios 3170. CTES), Apure,
and Bolfvar (e.g.. Amyard & Stergios 3340, MO;
Trujillo 1 1785. MY), and the Colombian ones from
Vichada appear to be morphologically intermediate
with W. involucrata (Saunders, 1995: 49, 335—337,
575—589). All intermediate populations associated
with the lower Orinoco River are placed with W.
involucrata s.l., occurring only east of the Caronf
River but not known between the Chaviripa River
and west of the Caronf River, where non-interme¬
diate W. involucrata s. str. occurs. Their bracts are
wider, more elliptically or rhombicly but variably
shaped, sometimes with the middle two or three
slightly to mostly fused rather than mostly distinct,
with margins irregular, serrulate, undulate, or partly
entire rather than entire. When bracts are together
eupuliform as in IE involucrata, but 4-shallowlv
lobed and I -parted, as occurs near the lower rather
than middle Orinoco River in the Venezuelan de¬
partments of Bolfvar, east of Rio Caronf, and in
Delta Amacuro, the intermediates are unequivocal¬
ly placed with W. involucrata, Waltheria berteroi s.
str. differs from W. involucrata s. str. in non-inter¬
mediate populations by its essentially distinct
bracts that are spathulate, with margins generally
entire, rather than entirely fused bracts that are ur-
ceolate, with lobes that are 1 — 1.7 mm long, and
dentate; by its calyx lobes that are (1.2)1 .6—1 .9 mm
wide rather than 1.1-1. 5(1.6) mm wide; and by its
anthers that are 0.8— 1 .()( 1 .2) mm long rather than
(1.0)1. 1—1.5 mm long. Also, usually the calyx tube
of IE berteroi is shorter, 3.5— 4.2 mm long, rather
than (3.0)4. 0—6.0 mm long as in W. involucrata.

Selected specimens. COLOMBIA. Cordoba: Sahagun,
27 Jan. 1918, Pennell 4085 (NY). Cesar: Mpio. Valle-
dupar, lnspecci6n a Atanquez, entre Atanquez y Guata-
purG 18 Nov. 1968, Jones A Pinto 2856 (NY); Poponte,

Magdalena Valley, 8 Dec. 1924, Allen 826 (MO). Mag-
dab Mia: Minca, 1941. Pro. Apolinar 528 (US); Cincinati,
1932, Giacometto III (US); Ocana, 11 Dec. 1879, Kal-
breyer 1266 (K); Sta. Marta, Uranieta, Valle de Alpar, Kar-
sten s.n. (W); Sta. Marta, Purdie s.n, (K); Camino de La
Gran Vfa a San Pedro, Romero-Castaneda 10644 (F, MO.
NY); Tucurinca, Oct. 1947, Romero-Castaneda 614 (F,
US); La Bonda, 250 ft.. 13 May 1898-1901, 11. II. Smith
493 locale & dale differ from type for IL subeordata (LL,
MICH, NY, U, WIS). Santander: Bucaramanga & vie.,
29 Dec. 1926, Killip & Smith 16251 (A. F, (ill. NY).

Acknowledgments. I thank Carmen Cristobal
and Antonio Krapoviekas for bringing the nomen-
elatural problem to my attention, providing an anal¬
ysis, bibliographic materials, and for reading the
manuscript; Lindsay Woodruff for sending a copy
of Gtirke's article; and lx. Ilerbst, Mahinda Marti¬
nez. and Marguerite Elliot for the translation of
Garckes article. 1 am grateful to Fred Barrie for
nomenclatural advice, lor review, and for rewriting
portions of the later versions of the paper, and also
to Beryl Simpson, Billie Turner, Guy Nesom, Javier
Fuertes, Fernando Zuloaga, Roberto Kiesling, Os-
valdo Morrone, and Victoria Hollowed lor nomen¬
clatural or other helpful comments and review of
earlier or later drafts. Partial financial support for
a larger monographic revision that included this
study was provided by a National Science Foun-
dation-DIS grant (BSR -8800899) and various fel¬
lowship awards from the University of Texas at Aus¬
tin. I am grateful to the curators and directors of
A. CTES, E, F. GH. K, LL, MEDEL, MICH. MO,
MPU, MY, NY, S, US, W, and WIS for the loan of
specimens cited here, and to staff at L, MEDEL,
and S for searching for additional isotypes.

Literature Cited

Ayers, T. J. & D. K. Boufford. 1988. Index to the vascular
plant types collected by H. II. Smith near Santa Marta.
Colombia. Brittonia 40: 400-432.

Bayer, C. & K. Kubitzki. 2003. Malvaceae. Pp. 225-311
in K. Kubitzki & C. Bayer (editors). The Families and
Genera of Vascular Plants, Vol. 5. Flowering Plants Di¬
cotyledons: Malvales, Capparales, and Non-betalain
Caryophyllales. Springer- Verlag, Berlin.

Burdet, H. 1972. Carlulae adbotanicorum graphicem.
Candollea 27: 315-316.

Garcke, A. 1863. Ueber einige ganz unbekannte Malva-
ceen. Bot. Zeitung (Berlin) 21: 275.

Greuter, W., J. McNeill, F. It. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
J. E. Skog. P. Trehane, N. J. Turland & 1). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Giirke, M. 1892. Beitriige zur Systematik dcr Malvaceen.

1. Die gaining Malachra. Bot. Jahrb. Syst. 16: 360.
Lanjouw, J. & F. A. Stafleu. 1954. Index Herbariorum.

Part 11(1). Collectors A-I). Regnum Veg. 2: 51. 71.
Saunders, J. G. 1995. Systematics and Evolution of Walth¬
eria (Stereuliaceae: Hermannieae). Ph.l). Dissertation.

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2005

Saunders
Waltheria berteroi

367

The University of Texas at Austin. [Unpublished. Avail¬
able as Reprint, Dissertation Services, Ann Arbor,
Michigan.]

Sprengel, C. P. J. 1826. Systems Vegetabilum 3: 94. Got¬
tingen.

Slafleu, F. A. & K. S. Cowan. 1976. Taxonomic Literature,
2nd ed.. Vol. 1: A— G. Bohn, Scheltema Holkema,
Utrecht. Regnum Veg. 94: 107—108, 201—202, 912.

- & - . 1985. Taxonomic Literature, 2nd ed..

Vol. 5: Sal— Ste. Bohn, Scheltema & Holkema, Utrecht/
Antwerpen. Regnum Veg. 1 12: 806. 813.

- & E. A. Mennega. 1992. Taxonomic Literature,

Supplement 1: A-Ba. Koeltz Scientific Books, Konig-
stein. Regnum Veg. 125: 287.

Standley, P. C. 1916. A new Waltheria from Colombia.

Contr. U.S. Natl. Herb. 18: I 17-1 18.

Willdenow, C. L. 1800. Sp. Plantarum, 4th ed. 3(1): 587—
588. Berlin.

New Species of Ocotea (Lauraceae) from Northern
Peru and Ecuador

Henk van dvr We iff

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

henk. vanderwerff@inobot.org

ABSTRACT. Seven species ot Ocotea , collected
during fieldwork in the drainage of the Rio Cenepa
in northern Peru, art* described. They will be in¬
cluded in the forthcoming Flora del Rio Cenepa.
The new species are: Ocotea badia van tier Werff,
Ocotea hirtandra van tier Werff, Ocotea imazensis
van tier Werff. Ocotea laevijolia van tier Werff, Oco¬
tea lenitae van tier Werff, Ocotea leptophylla van
tier Werff, and Ocotea vasquezii van tier Werff.

Key words: Ecuador, Lauraceae, Ocotea , Peru,
Rfo Cenepa.

During preparation of a treatment of the Laura¬
ceae for the Flora del Rio Cenepa (Vasquez et al.,
in prep.) a number of collections were encountered
that tlefied attempts at identification. Three species,
belonging to the better known genera Licaria Au-
blet, Nectandra Rottboell, and Pleurothyrium Nees,
were described in an earlier publication (van tier
Werff, 2003), but those belonging to the large and
difficult genus Ocotea Aublet had not been dealt
with, based on these collections from northern Peru
and adjacent Ecuador, seven new species art* de¬
scribed here. Ocotea is the largest genus of Laur¬
aceae in iht* Neotropics, with an estimated 300 +
species (pers. obs.). It was last revised by Mez
(1889), who included 199 species in the genus.
Since then, a steady stream of new species has
been published (van der Werff, 1994, 1999, 2001.
2003; van tier Werff cY Vicentini, 2000). bohvver
(1980) published an overview of the genus, largely
based tin a study of type specimens, lit* recognized
a number of species groups and provided keys to
his species groups and to species within each spe¬
cies group. His contribution has greatly helped our
understanding of the genus. Traditionally, Ocotea
has been characterized by having flowers with nine
4-celletl locelli. the locelli arranged in two super¬
posed paim, and includes all species that do not fit
in other, better defined genera with nine 4-celletl
stamens. A DNA-based phytogeny of the genera of
Lauraceae (Chanderbali et al., 2001) suggests
strongly that Ocotea as currently accepted is not a
monophyletic genus, but is formed by several not

closely related groups of species. These newr in¬
sights have not yet been incorporated in a new clas¬
sification, largely because morphologic al characters
that would support the groups recognized in the
DNA-based phylogeny have not yet been found. A
first step toward a new classification of Ocotea will
be a revision of the known species and the descrip¬
tion of the assorted undescribed species. Such a
revision is planned by the author. Until such a re¬
vision has been completed, I will use the traditional
concept for Ocotea , even though it is very likely too
broad.

The new species discovered in northern Peru are
the following:

I. Ocotea badia van der Werff, sp. nov. TYPE:
Peru. Depto. Amazonas: Bagua Prov., Distr.
Imaza. Yamayakat, 04°55'S, 78°19'W. 400 m
alt.. 24 May 1996 (fl), R. Vasquez <& A. Vasquez
20957 (holotype. HUT; isotypes, HBG, K. MO.
NY, USM). Eigure I.

Ex grege Oeoteae minarum, seel ramulis tomentellis,
venatieme partim brochidodroma et folds suhlus pilis cr¬
eeds praeditis differt.

Trees, to 25 in; young twigs angular, solid or old¬
er twigs narrowly fistulose, densely reddish brown
tomentellous, the indument covering the surface
completely, becoming glabrous with age; terminal
buds densely reddish brown tomentellous. Leaves
10—18 X 3—5 cm, elliptic to slightly obovate, char-
taceous, alternate, the base acute, the margin flat,
the apex acuminate, acumen to 1.5 cm long; ve¬
nation immersed on upper surface, raised on lower
surface; upper surface glabrous, lower surface ini¬
tially densely pubescent with short, erect hairs, the
indument becoming sparser with age, midrib and
lateral veins tomentellous on the lower surface;
domatia lacking; lateral veins 4 to 7 pairs, often
somew hat loop-connected in the distal half; petioles
flat above. Inflorescences to 12 cm long, panieu-
late-cymose, reddish brown tomentellous, in the ax¬
ils of leaves or cataphylls. I' lowers hermaphrodite,
yellow, 2.5— 3.5 mm diam., the tepals erect at an-
thesis, pedicels 2—3 mm long; tepals 6, 1.5 mm

Novon 15: 368-378. 2005.

Volume 15, Number 2
2005

van der Werff

New Species of Ocotea

369

long, ovate-elliptic, moderately pubescent outside,
sparsely pubescent inside; stamens 9, all 4-cel led,
the cells in 2 superposed pairs, the outer 6 ea. 1.2
mm long, tin* glabrous anther ca. 1 mm long, the
short filament pubescent; inner stamens same
length, also with a very short filament, glands pre¬
sent at the base of the inner stamens, staminodia
not seen, pistil glabrous, ca. 1.6 mm long, the style
I mm long, the ovary globose, clearly swollen.
Fruits ellipsoid to ovoid-ellipsoid, 2X1 cm, eu-
pule small, platelike, 6 mm diam., gradually nar¬
rowed into the pedicel.

Ocotea badia, named after the color of the in-
dument on the young twigs, is close to the 0. min-
arum group or part of it, largely because of its el¬
lipsoid or ovoid-ellipsoid fruits and small, platelike
cupules. Kohwer (1986) found two types of flowers
in this group: pistillate and hermaphrodite. The
type of 0. badia has hermaphrodite flowers: the pis¬
til has a clearly thickened ovary, while the locelli
of the stamens open and show some pollen grains
on their flaps. However, the terminal buds are not
as long and slender as is usual in this group. I he
only species in the 0. minarum group or close to
it that have an erect indument are O.fendleri (Meis-
ner) Rohwer, O. ovalifolia (Kuiz & Pavon) Mez. ().
scalariforrnis van der Werff, and 0. odorata (Meis-
ner) Mez. but the first three of these species have
attenuate leaf bases that are usually inrolled and
0. odorata differs in having the lower leaf surface
glaucous. Some of the flowers and buds of the type
collection are larger than average for the species,
and have a denser indument and longer pedicels.
These buds or flowers are diseased and contain
only whorls of gradually reduced tepals, but no re¬
productive organs. The tertiary venation on the low¬
er leaf surface of O. badia is noticably less raised
on the fruiting specimens than on the flowering
ones, but the leaf shape, indument. and the part l\
loop-connected venation are similar in fruiting and
flowering specimens. The Ecuadorian paratype is
placed here provisionally: it differs in having small¬
er leaves, shorter petioles, more pairs of lateral
veins and lacks the loop-connected venation in the
distal half of the leaves.

Phenology. Flowering May— June; fruiting Feb¬
ruary.

Paratypcs. ECUADOR. Carchi: Tulcan Canton, Re-
serva Ethnica Awa-Camumbi (fr). C. Quelal, C. Aulestia
A E Nastacuaz 255 (VIO). PERU. Amazonas: Km Ce-
nepa, S of Huampami trail (fr), B. Berlin 1662 (MO); One-
hrada Huampami (fr), R. Kayap 363 (MO); Condoreanqui
Province, El Cenepa, comunidad de San Antonio (tl). R-
Vasquez, A. Rena, E. Chavez & E. Quiaco 24088 (MO).

2. Ocotea hirtandra van der Werff, sp. nov.
TYPE: Peru. Depto. Amazonas: Bagua Prov.,
Distr. Imaza. camino Putuim-Shimutaz,
<)5°03'S, 78°20'W, 380 m. 22 June 1996 (fl).
lx. Vasquez , ,4. Vasquez, L. Debent a i & M.
Chuimtan 21335 (holotype, HUT; isotypes,
HBG, MO, NY). Figure 2.

Ocoteae multiglandulosae similis, sed staminibus dense
pubescentibus et pistillodio glabro recedit.

Trees, to 30 m; twigs angular or ridged, glabrous
or near the tips minutely appressed pubescent, sol¬
id. terminal buds densely appressed pubescent.
Leaves alternate, 10-21 X 4-8 cm, chartaceous,
obovate or obovate-elliptic. the base acute, margin
flat or slightly inrolled, apex obtuse to bluntly
acute, glabrous, except for the tufts of hairs in the
axils of the basal lateral veins, venation immersed
on the upper surface, midrib and lateral veins
raised, tertiary venation immersed on the lower sur¬
face, petioles 10-15 mm, ridged; lateral veins * to
10 pairs. Inflorescences 8-15 cm, paniculate-cy-
mose, in the axils ol cataphylls near the tips of the
branches, basally nearly glabrous, becoming dis-
tally moderately to densely puberulous with very
short, erect hairs. Staminate flowers 3.5—4 mm
diam., yellowish, pedicels densely puberulous, half
as long to about as long as the flowers; tepals 1.5-
2 mm long, densely puberulous on both surfaces,
half-erect at anthesis, equal; stamens 9, all 4-
celled, the cells in 2 superposed pairs, the outer 6
stamens with the cells introrse, 1.5 mm long, the
filament half as long as the anther, filaments dense¬
ly puberulous, the anther abaxially densely puber¬
ulous or the indument dense along the center ol the
anther and less dense toward the margins; inner 3
with the lower pair of cells extrorse. the upper pair
latrorse, about as long as the outer 6 stamens, fil¬
aments densely puberulous and with 2 conspicuous
glands at tile base; staminodia present, ca. 0.5 mm
long, stipitiform or with a capitate tip. puberulous,
sometimes 2 locelli present on the staminodia; pis-
tillode spindle-shaped, I mm long, glabrous; recep¬
tacle cup-shaped, glabrous inside; pistillate flowers
and fruits not known.

Ocotea hirtandra is characterized by its densely
puberulous flowers and stamens and its obovate or
obovate-elliptic leaves with axillary tufts of hairs.
The position of the inflorescences, in the axils of
cataphylls and not in the axils of leaves, is unusual.
In one of the five known collections (Palacios & Ig-
uago 4639) the inflorescences are not subapical;
here the terminal buds have produced a young shoot
with immature leaves and the inflorescences are low¬
er on the twig, but still not in the axils ol leaves.

370

No von

Figure I.

n.V4«QU#I, fi.Vjsqiin ?0937

fll'JSUUni IJOIONHAt tiAHlM M tftRDAttturi (nil

Ocotea badia van der Werff, I lie MO isotype, Vasquez & Vasquez 20957.

Relationships of the new species are not clear. It
resembles species such as (). multiglandulosa (Ruiz
& Pavdn) Mez and (). cuneifolia (Ruiz & Pavdn)
Mez. I he position of the inflorescences is unusual
in the genus, as are the densely puberulous stamens.
Mature cupules would help in understanding its po¬
sition in the genus. In one collection ( Kayap 972 )
the pistillode is swollen at the base, suggesting a
viable pistil, but the* anthers are normally developed,
possess a few pollen grains, and the thickened pis¬
tillode does not possess a stigma and is clearly short¬
er than the inner 3 stamens. It is possible that this
species is androdioecious, having staminate and her¬
maphrodite (lowers, but more collections are needed
to confirm this. Gynodioecious flowers have been re¬
ported in Ocotea , but androdioecious flowers are not
definitely known in Lauraceae; Chanderbali (2004)
suggested tbe presence of androdioecious flowers in
Endlicheria gracilis Kostermans.

Phenology. Flowering June (Peruvian collec¬
tions), October (Ecuadorian collections).

Paratypes. PERU. Depto. Amazonas: Prov. Condor-
cunqui. comunidad de Tutino (ff). It Vasquez. A. Pena, E.
Chavez & h. Quiaco 24177 (MO); Lugar Huampaini (fl).
It Kayap 972 (MO). ECUADOR. INupo: Canton Archi-
dona, Carretera Hollfn— Loreto, entre Huamani y el Rfo
Pucuno (fl). ft. Palacios & C. Iguago 4619 (MO); km 50
Carretera Hollfn— Loreto (fl). W. Palacios & C. Iguago
4621 (MO).

3. Ocotea imazensis van der Werff, sp. nov.
TYPE: I Vru. Depto. Amazonas: Ragua Prov.,
Distr. Imaza, Comunidad de Kampaenza, Ri¬
bera de la Quebrada Shimutaz, 05°55'S,
78°19'W, 320 m. 23 Sep. 1994 (fl. immature
fr). A'. Jaramillo, It Apanu & S. Katip 477
(holotype, HUT; isotypes, HBG, k. MO, NY,
USM). Figure 3.

Volume 15, Number 2
2005

van der Werff

New Species of Ocotea

371

MISSOURI
BOTANICAL GARDEN
HERBARIUM

R. VA'.que/, 0. VJSQu*j,

MISSOURI BOTANICAL CARDIN HfRRAR IUM <MU >

Figure 2. Ocotea hirtandra van der Werff, the MO isotype, Vasquez et al. 21335.

A congeneris raniulis inflorescentiisque dense tomen-
tellis, basibus foliorum parum decurrentibus, petiolis su¬
pra complanatis et floribus hermaphroditis recedit.

Trees, to 25 m; twigs terete or angular, densely
ferrugineous-tomentellous, the surface completely
covered, solid; terminal buds densely tomentellous.
Leaves 7—10 X 3-7 cm, firmly chartaceous, evenly
distributed along the twigs, green above, with small
wax deposits on the lower surface, glabrous on up¬
per surface, with scattered curled hairs, these op¬
pressed or half-erect, on the lower surface the hairs
denser and becoming tomentellous on the major
veins; domatia lacking; apex acute or shortly acu¬
minate. the base acute, the lamina slightly decur¬
rent on the petioles; lateral veins 5 to 7 pairs; pet¬
ioles brown-tomentellous, 8-14 mm long, flat
above. Inflorescences 2.5-5 cm long, densely to¬
mentellous, in the axils of leaves or bracts, panic¬

ulate-cymose, the lateral branches ending in a
cyme, or racemose. Flowers yellow or cream-col¬
ored, hermaphrodite, ca. 3 mm diam., the pedicels
less than the diameter ol the (lowers; tepals 1.3 mm
long, spreading at anthesis, elliptic-oblong, densely
pubescent on the outer surface, slightly less so on
the inner surface; stamens 9, all 4-eelled, glabrous,
the outer 6 ca. 0.5 mm long, the lilament shorter
than the anther, opening introrse, the cells arranged
in 2 pairs, inner 3 stamens of same size, the lower
2 anther cells extrorse, the upper 2 lateral, glands
present at the base of the inner 3 stamens, stami-
nodia present, columnar, with a slightly thickened
apex, glabrous, about as long as the filaments of the
inner stamens; pistil glabrous, style distinct, ca. 0.2
mm long. Immature fruit enclosed in the hypanthi-
um; I older cupule 7 mm diam., 5 mm high, weakly
double-margined, the outer margin erect and sur-

372

No von

MISSOURI
BOTANICAL GARDEN
HERBARIUM

C«thn nit- T. Arthur I oufiiljl i<m
niasttui nmoNtcot onuot n hi hhai< mu ,run

Figure 3. Ocotea imazensis van der Werff, the MO isotype, Jaramillo et al. 477.

passing the inner margin; fruit (possibly immature)
9 X < 1 mm; tepals deciduous in fruit.

Ocotea imazensis is best recognized by the com¬
bination ol densely pubescent twigs and inflores¬
cences. the slightly decurrent leaf base, the flat pet¬
ioles, and perfect flowers. The new' species does not
seem to be closely related to any of the other Ocotea
species with perfect flowers in South America. Veg-
etatively it resembles a collection from the Arbo¬
retum Jenaro Herrera in the Departamento de Lor¬
eto (Cliota s.n , collected from tree 3/96, MO), but
that specimen has larger flowers with pubescent
stamens; it has not yet been identified.

Phenology. f lowering September.

Paratype. PERU. Dcpto. Amazonas: Bagua Prov.,
Distr. Imaza, Comunidad Aguaruna Pituim (fl). C. Diaz. A.
Pena A- P. Atamain 7208 (MO).

4. Ocotea laevifolia van der Werff. sp. nov.
TV PE: Peru. Depto. Amazonas: Prov. Bagua.
Distr. Imaza, Tayu Mujaji, 05°15'56"S,
78°22'07"W, 1200 m, 21 Oct. 1997 (fl), R. Vas-
quez, ,/. Lirio & (',. Pitug 24602 (holotype,
HUT; isotypes, HBG, MO). Figure 4.

Ex grege Ocoteae aeiphvllae, sed foliis laevibus, inflo-
rescentiis laxis et hahitu fruticoso recedit.

Shrub, 2 m tall; twigs terete or ridged, near the
tip minutely appressed pubescent, soon becoming
glabrous, solid; terminal buds small, densely ap¬
pressed pubescent. Leaves 9-14 X 3.5— 5.5 cm, el¬
liptic or ovate-elliptic, alternate, evenly distributed
along the twigs, ehartaeeous, brittle, glabrous on
both surfaces or minutely appressed pubescent on
young leaves, the base obtuse, slightly inrolled, the
apex acuminate or acute, venation on upper surface

Volume 15, Number 2
2005

van der Werff

New Species of Ocotea

373

MISSOURI
BOTANICAL GARDFN
HtRBARIUM

H. v«tqu«2, J. Lino l G. Pi tug. i?**603

niBSOUNl WIIANICW liAHDfN »« HBAIiMffl (MU )

Figure 4. Ocotea laevifolia van der Werff, the MO isotype, Vasquez el <il. 24603.

immersed, midrib slightly raised, otherwise vena¬
tion immersed on lower surface, lateral veins dif¬
ficult to discern, 6 to 8 pairs; petioles 7—10 mm
long, canaliculate, minutely appressed pubescent
or glabrous. Inflorescences 5—7 cm long, in the ax¬
ils of leaves, laxly flowered, paniculate near the
base, racemose more distally, moderately appressed
pubescent, the indument denser toward the flowers.
Flowers hermaphrodite. 3—3.5 nun diam.. yellow,
appressed pubescent, pedicels 3—5 nun long,
densely appressed pubescent; tepals ea. 1 .5 mm
long, ovate-elliptic, appressed pubescent on both
surfaces, half-erect at anthesis; stamens 9, all 4-
celled. the cells in 2 superposed pairs, 1.2 mm
long, the filaments a little shorter than the anthers,
filaments pubescent, outer 0 stamens with tin* cells
introrse, inner 3 with the cells lateral-extrorse.
glands present at the base of the inner stamens,
staminodia not seen, pistil 1.5 mm. glabrous, style

slightly shorter than the ovary; receptacle cup¬
shaped, densely pubescent inside. Fruits and cu-
pules not seen.

Ocotea laevifolia is only known from the type
collection, made in the summit area of Germ Tayu,
at an altitude of about 1200 m. Cerro Iayu is an
isolated sandstone hill with forested slopes and a
summit area covered with 2 m tall scrub. Several
other collections from Cerro Tayu are identified as
undeseribed species (representing two species of
Cyathea in Cyatheaceae, one other species of Oco-
tea , and one species of Phyllanthns , Kuphorbi-
aeeae) or are new records for Peru ( Aratitiyopea
lopezii (L. B. Smith) Steyermark & Berry. Xyrida-
eeae, and Euceraea nitida Martins, Flacourtiaceae).
It is likely that more novelties will be found with
further collecting on this rather small hill.

Ocotea species with hermaphrodite flowers in

374

Novon

MISSOURI BOTANIC**

/ l J . Can pc« I63<«6

tlAWKN ID UhAttlllM mi)

Figure 5. Ocotea lenitae van der Werfl, the MO isotype, van der Werff et al. 16546.

South America tend to belong to a few species
groups, such as the (). insularis group, the 0. aei-
phylla group, the 0. sodiroana group, and the 0.
indecora group. The strongly canaliculate petioles
point to the (). aciphylla group, while distinctive
characters of the other groups are lacking. Its clos¬
est relative may be an undescribed species from
the Cordillera del Condor in Kcuador, which differs
in having thinner, subopposite leaves with an in-
rolled and decurrent leaf base. The two species
share laxly flowered inflorescences and a pubescent
inner surface of the receptacles.

3. Ocotea lenitae van der Werff, sp. nov. TYPE:
Peru. Depto. de San Martin: Kioja Prov., along
road Yorongos-La Florida near Rioja, 1000 m.
3 1 Mar. 2001 (hermaphrodite fl), //. van dvr
Werff' R. Vasquez, R. Gray & J. Campos 16546
(holotype. Mill’; isotypes, CANR. [IRC, K.
MO, NY. US, USM). Figure 3.

Ex grege Ocoteae minarum, sed foliis obtusis, tepalis
inlus dense pubescentibus et fructibus globosis bene dis-
tincta.

Irees, to 45 m; twigs angled, moderately to
sparsely appressed pubescent, becoming glabrous
with age; terminal buds slender, densely appressed
pubescent. Leaves 10-18 X 3-7 cm, alternate,
evenly distributed along twigs, firmly chartaeeous,
obovate, the base narrowly cuneate, tin* margin flat
or slightly inrolled. the apex obtuse, the upper sur¬
face glabrous, lower surface glabrous or with some
appressed hairs, a few pit-domatia sometimes, but
not always, present in the axils of lateral veins and
visible as bumps on the upper surface, the margins
of the pits glabrous; venation immersed on the up¬
per surface, midrib and lateral veins raised on the
lower surface; lateral veins 7 to 9 pairs; petioles 1 —
2 cm long, shallowly canaliculate. Inflorescences
.5-15 cm long, pan i culate-cy mose, sparsely to mod-

Volume 15, Number 2
2005

van der Werff

New Species of Ocotea

375

N? 5796961

LAURACEAE

Figure 6. Ocotea leptophylla van tier Werff, the MO isotype, Diaz et al. 8076.

erately appressed pubescent, in the axils of leaves
or cataphylls; hermaphrodite flowers ca. 5 mm
diam., pistillate (lowers ca. 4 mm diam., yellow or
green, pedicels ca. 2 mm long. Hermaphrodite flow¬
ers: tepals ca. 1.7 mm long, elliptic, spreading at
anthesis, the outer three sparsely appressed pubes¬
cent on the outer surface, the inner more densely
so; inner surface of tepals densely pubescent; sta¬
mens 0. all 4-eelled, the cells in 2 superposed
[»airs, the outer 6 stamens 1.3 mm long, the fila¬
ments ca. 0.3 mm long, pubescent, the anthers I
mm long, glabrous, opening introrse; inner 3 sta¬
mens 1.5 mm long, the filaments pubescent, ca. 0.5
mm long, the anthers glabrous; globose glands pre¬
sent at the base of the inner stamens; staminodia
lacking; pistil 1.5 mm long, glabrous, the style ca.
I mm long, the stigma discoid, receptacle shallow,
glabrous inside. Pistillate flowers: tepals 1.5 mm
long, staminodia 9, ca. 0.6 mm long, the filaments

very short, pubescent, glands present al the base
of the inner staminodia, pistil 1.6 mm long, gla¬
brous, the style a little longer than the ovary; re¬
ceptacle shallowly cup-shaped, with some ap¬
pressed hairs inside. Infructescences branching
under almost 90°. Fruit globose, 1 cm diam., cupule
small, platelike, 7 mm diam., pedicel and cupule
lenticellate.

Ocotea lenitae belongs to the Ocotea minarum
group based on its hermaphrodite and pistillate
(lowers and its platelike, lenticellate cupule. It is
distinct within this group by the combination of ob-
ovate leaves with obtuse apex, presence of (few) pit-
domatia, and the spreading tepals that are densely
pubescent on the inner surface. Most species in the
(). minarum group have leaves with an acute apex;
in Peru only 0. obovata (Ruiz & I ’a von) Muz is
reported to have leaves with a rounded or obtuse

376

No von

apex, but that species does not have domatia. Oco¬
tea tillettsiana Allen is (lie species most similar to
O. lenitae ; it has leaves with a rounded apex,
spreading tepals at anthesis, and pit-domatia; it dif¬
fers in its sparser indument on twigs and flowers,
in having shorter petioles (to I cm long), more dom¬
atia, and raised tertiary venation. Ocotea tillettsiana
is an incompletely known species reported from
Guyana and adjacent Venezuela.

I he pit-domatia on the leaves of Ocotea lenitae
are not numerous. Many leaves do not have these
domatia and one has to search carefully for their
presence. Several collections placed here in 0. len¬
itae had been previously identified as (). oblonga
(Meisner) Mez, and duplicates may have been dis¬
tributed under that name.

It is a pleasure to dedicate this species to my
wife, Lenita, in recognition of her interest in and
steadfast support of my work.

Phenology. Flowering April-June; fruiting Jan¬
uary.

Paratypes. PERU. Depto. San Martin: Rioja Prov.,
along road Rioja- Yorongos-I ,a Florida (pistillate fl), II.
van tier Her//, It. Vasquez, H. Cray & J. Campos 1 65 It)
(MO); along road Yorongos-La Florida (young fr), //. van
der Her//, R. Vasquez, R. Cray cY ,/. Campos 16545 (MO).
Depto. Amazonas: Quebrada Kayamas, l.ngar Cenepa
(berm, fls), hi Anriiash 170 (MO); Quebrada yutui entsa
(berm, fls), hi. Aneuash 217 (MO). ECUADOR. Pastaza
Prov.: Via Aura, I 15 km S of Coca (fr), P. Hurtado. W.
Palacios & C. Iguago 1575 (MO); 35 kin SE of Curaray
(buds), h.spinoza 500 (\U). QCINE). INapo Prov.: Rio I’ax-
amino (buds), /,. Holm-Nielsen & S. Jeppesen HOI (A A LI,
MO); V ia Coca-Auca. km 17 (fr), HI Palacios 3514 (MO);
PN Yasuni (fr). t. Crijaha 765 (MO, QCNE), (si). M. Au-
lestia & T. Ahue 2766 (MO. QGN E). (berm. fls). M. Au-
lestia 2272 (MO. QCNE), (buds), W. Palacios 2565 (MO.
QCINE); Ueserva F.tniea Huaorani (herm. fls). M. Auleslia
5642 (MO, QCINE), (buds), M. Aulestia 5616 (MO.
QCINE). (berm. fls). M. Aulestia 5655 (MO. QCINE).

6. Ocotea Icplophylla van der Werff, sp. nov.
TYPE: Pt Tii. Depto. Amazonas: Bagua Prov.,
Distr. Imaza, Comunidad Aguaruna de Wanas
(km 92 Carretera Bagua— Imacita), Germs
Chinim, 700-800 m, 29 Aug. 1996 (male II),
C. Diaz, A. Pena, /.. Tsamajain & M. Roca
8076 (holotype. Ill I; isotypes. HBG, K. MO,
NY, US, USM). Figure 6.

A eongeneris loliis subtus glaucis, infloreseentiis foli-
isque ad ramulos hornotinos limitatis distincta.

Trees, to 25 m; young twigs angular, densely
brown tomentellous, the surface completely cov¬
ered, older twigs glabreseent. dark, with scars from
fallen leaves; terminal buds densely brown tomen¬
tellous. Leaves 1—1 I X 2—5 cm, membranaceous
to chartaceous, elliptic to obovate-elliptic, clus¬

tered on llu‘ recent flush, older twigs leafless; base
obtuse to acute, the apex acute, the margin flat;
upper surface glabrous, except for the pubescent
midrib, lower surface glaucous, with scattered erect
hairs or almost glabrous, the indument denser on
the major veins; domatia lacking: midrib and lateral
veins immersed, tertiary venation slightly raised on
the upper surface, midrib raised, lateral veins
weakly raised and tertiary venation immersed on
lower surface; lateral veins 5 to 7 on each side;
petioles 10—15 mm long, densely pubescent, flat
above. Inflorescences to 7 cm long, in the axils of
bracts, rarely in the axils of leaves, along the young
twigs; densely tomentellous, the surface entirely or
largely covered, paniculate-cymose. Staminate
(lowers green or cream-colored, 2.5—3 mm diam.,
tepals ea. 1 mm long, ovale, half-erect at anthesis,
densely pubescent on the outer surface, sparsely
pubescent on the inner surface, but more densely
so at the base, stamens 9. all 4-eelled, in 2 super¬
posed pairs, outer 6 stamens 0.6 mm long, the cells
opening introrse or the lower two lateral, the fila¬
ment very short, 0.2 mm, pubescent; inner 3 sta¬
mens 0.8 mm long, the cells opening extrorse,
glands present at the base of the short filaments,
staminodia not seen, pistillode lacking or small,
glabrous; receptacle shallow, pubescent inside; pis¬
tillate (lowers, fruits, and cupules not known.

Ocotea leptophylla is easily recognized by its
glaucous, thin leaves and its short, densely tomen¬
tellous inflorescences, both of which are confined
to the recent growth. Rarely do a few older leaves
persist, and those have a darker lower leaf surface.
The indument on the lower leaf surface varies from
sparse to almost lacking. The stamens have very
short filaments: normally Ocotea stamens have dis¬
tinct. narrow filaments, while the closely related ge¬
nus Rliodostemonoclaphne Rohwer & Kubitzki has
sessile or almost sessile anthers. Most species of
Rhodostemonodaphne have clearly larger flowers
than those of O. leptophylla, but flowers of R. par-
vifolia Madrinan are almost as small as those of O.
leptophylla. Still, anthers of R. parvifolia are ses¬
sile, while those of 0. leptophylla have a distinct,
short filament; this is the main reason to describe
the new species in Ocotea rather than in Rhodos¬
temonodaphne.

Phenology. f lowering August— September.

Paratype. PERU. Depto. Amazonas: Ragua Prov.,
Distr. Imaza, Comunidad de Kampaenza, N. Jaramillo, R.
Apanu & S. Katip 4H7 (HUT, MO).

Volume 15, Number 2
2005

van der Werff

New Species of Ocotea

377

MISSOURI

BOTANICAL GARDEN
HERBARIUM

N? 5796958

7. Ocotea vasquezii van der Werff, sp. nov.
TYPE: P eru. Depto. Amazonas: Prov. Bagua,
Distr. Imaza. Yamayakat. 05°03'S, 78°20'W,
480 m, 9 Nov. 1996 (fl). R. Vasquez, P. Stern,
R. Rojas <£• R. Aguilar 21739 (holotype, HUT;
isotypes, HBG, K. MO. NY, US). Figure 7.

Ocoteae atirrensi affinis, sed ramulis solidis, staminibus
verticelli 1 1 1 subglabris vel glabris et cupulis parvis, pi an is
recedi t.

Ireelets or small trees, to 10 m; twigs angular to
terete, glabrous or with a few small, appressed
hairs, solid, terminal buds rather densely appressed
pubescent. Leaves 15—35 X 7.5—15 cm, elliptic to
broadly elliptic, alternate, evenly distributed along
the twigs, papyraceous, the base acute to obtuse,
the margin flat, apex acute or acuminate, the acu¬
men I cm long, glabrous on both surfaces, domatia
absent, venation immersed on the upper surface.

midrib prominently raised, lateral veins and tertiary
venation raised on lower surface; lateral veins 7 to
10 pairs; petioles 10—16 mm long, glabrous, ridged
above. Inflorescences 7-20 cm long, paniculate cy-
mose, in the axils of leaves or cataphylls near the
tips of the branches, moderately to sparsely puber-
ulous. Flowers hermaphrodite, 2.5—3 mm diam..
white or yellow, pedicels ea. 2 mm long; tepals
erect or half-erect at anthesis, ca. 1.5 mm long,
ovate, moderately to densely pubescent outside,
when moderately pubescent, the inner tepals with
a denser indument than the outer ones; stamens 9,
all 4-celled, the' cells in 2 superposed pairs, the
outer stamens 6 with introrse cells, ea. 1.3 mm
long, the filament as long as the anther, glabrous or
w ith a few hairs along the filament, the inner 3 with
extrorse cells, ca. 1 mm long, the anther slightly
longer than the filament, glabrous or with a few
hairs along the filament; globose glands present at

378

Novon

the base of the inner stamens; staminodia not seen:
pistil glabrous, 1.5 mm long, the style a little longer
than the ovary, stigma platelike; receptacle shallow¬
ly cup-shaped, glabrous inside. Fruits ellipsoid, 3.5
X 1.7 cm, cupule consisting ol a small plate, 6 mm
diam.; tepals deciduous in fruit.

Ocotea vasquezii is closely related to or belongs
in the Ocotea insularis group. Its appearance is
similar to (). atirrensis Me/ & Donnell Smith from
Central America; large, dark green drying, papy¬
raceous leaves, copiously branched inflorescences,
more or less erect tepals at anthesis, and the habit,
a small tree. It differs from O. atirrensis in having
solid stems, a small, platelike cupule (0. atirrensis
has a small cup-shaped cupule) and in the absence
of the tuft of hairs at the junction of filaments and
anthers of the inner stamens. Ocotea insularis is
less similar; it has domatia, decurrent leaf bases,
firmer leaves drying a brighter green color in ad¬
dition to having a cup-shaped cupule. Ocotea atir¬
rensis is not known from South America, while 0.
insularis s.l. is not uncommon in Colombia, Ecua¬
dor. and to a lesser degree in Peru.

It is a pleasure to dedicate this species to Ro¬
dolfo Vasquez, the collector of the type, who
through his publications and collections has greatly
expanded our knowledge of the flora of Peru.

Phenology. Flowering October— November;

fruiting June-August.

Paralypes. PERU. Depto. Amazonas: Prov. Bagua,
Distr. Ima/.a, Tuyankuwas, comunidad de Yanat (fl), E.
Chare: 77 (MO); Distr. Ima/.a, Yamayakat (fr), R. V usque:.

R. Orliz-Genlry , l\. Jaramillo & R. Apanu 18700 (MO),
(buds), R. Vasquez & /V. Jaramillo 2020-i (MO), (fl), R.
Vasquez. P. Stern . R. Rojas & R. Aguilar 2 1 582 (MO), (fr),
R. Vasquez , A. Pena & E. Chaves 28945 (MO), (fl), R.
Vasquez. A. Pena A' E. Chavez 24902 (MO); Distr. Ima/.a,
Quebrada Kuzu (fr), C. Diaz. S. Katip & A. Pena 0958
(MO); Distr. El Cenepa, Mamayaque (fr), R. Vasquez. R.
Rojas, A. Pena. E. Chavez & E. Quiaco 24509 (MO), (fr),
R. Rojas. A. Pena & E. Chavez 225 (MO).

Acknowledgments. Fieldwork in Peru was sup¬
ported by the Taylor Fund for Ecological Research.
Fred Keusenkothen made the scanned images.
East, and most importantly, 1 thank my colleagues
in Peru for their excellent collections.

Literature Cited

Chanderbali. A. S. 2004. Endlicheria. El. Neotrop. 91: I-
141.

- . H. van der Werff & S. Renner. 2001. Phytogeny

and historical biogeography of Lauraeeae: Evidence
from chloroplast and nuclear genomes. Ann. Missouri
Bot. Card. 88: 104-134.

Mez, C. 1889. Lauraeeae Americanae. Jahrb. Konigl. Bot.
Cart. Berlin 5: 1—556.

Rohwer, J. G. 1986. Prodromus einer Monographic der
Gattung Ocotea Aubl. (Lauraeeae) sensu lato. Mitt. Inst.
Allg. Bot. Hamburg 20: 1-278.

Van der Werff, H. 1994. Novelties in Neotropical Laura¬
eeae. Novon 4: 58—76.

- . 1999. New taxa and combinations in the Ocotea

helieterifolia (Lauraeeae) species group. Novon 9: 571 —
583.

- . 2001. New taxa and combinations in Ocotea

(Lauraeeae) from Central America. Novon 1 I: 501-51 1.
- . 2003. New taxa of Lauraeeae from South Amer¬
ica. Novon 13: 337-357.

- & A. Vicentini. 2000. New species of Lauraeeae

from Central Amazonia, Brazil. Novon 10: 264—297.

New Combinations and Synonyms in Chinese
Pyrenaria s.l. (Theaceae)

Yang Shi- Xiong

Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan, 650204,
People’s Republic of China, shxyang@mail.kib.ac.cn

Abstract. Six new combinations in Pyrenaria
(Theaceae) are made. These are: Pyrenaria specta-
bilis (Champion) (5 Y. Wu & S. X. Yang, P. spec-
tahilis var. greeniae (Chun) S. X. Yang, P. macu-
latoclada (Y. K. Li) S. X. Yang, P. wuiana (Hung
T. Chang) S. X. Yang, P. jongquieriana Pierre ex
Lanessan subsp. multisepala (Merrill & Chun) S.

X. Yang, and P diospyricarpa Kurz var. camelliiflo-
ra (Kurz) S. X. Yang. Pyrenaria burmanica T. K.
Paul & Nayar newly synonym izes to Pyrenaria dios¬
pyricarpa var. diospyricarpa. Pyrenaria tibetana
Hung T. Chang newly synonymizes to Pyrenaria
khasiana R. N. Paul. Then shinkoensis Hayata and
Tutcheria tenuifolia Hung T. Chang newly synony-
mize to Pyrenaria microcarpa (Dunn) II. Keng var.
microcarpa. Camellia buisanensis Sasaki, Tutcheria
symplocifolia Merrill & F. P. Metcalf, and Tutcheria
taiwanica Hung T. Chang & S. X. Ren newly syn-
onymize to Pyrenaria microcarpa (Dunn) H. Keng
var. ovalifolia (H. L. Li) T. L. Ming & S. X. Yang.
Tutcheria brachycarpa Hung T. Chang newly syn¬
onymizes to Pyrenaria spectabilis (Champion) C. Y.
Wu & S. X. Y ang var. spectabilis. Tutcheria austro-
sinica Hung T. Chang, Tutcheria rostrata Hung T.
Chang, and Pyrenaria turbinata S. X. Yang newly
synonymize to Pyrenaria spectabilis (Champion) C.

Y. Wu & S. X. Yang var. greeniae (Chun) S. X.
Yang. In addition, Pyrenaria microcarpa (Dunn) H.
Keng is leetotypified.

key words: China, Pyrenaria, Theaceae.

Keng (1980), Yang (1997), and Chang (1998)
have published regional taxonomic revisions of the
Malesian and Chinese Pyrenaria Blume and Tutch¬
eria Dunn (Theaceae). Keng (1972) first reduced
Tutcheria into Pyrenaria. Yang and Ming (1995a,
1995b) and Yang (1997) supported Keng’s (1972)
opinion and further reduced Parapyrenaria Hung
T. Chang into Pyrenaria. A worldwide taxonomic
revision of Pyrenaria s.l. has been carried out as a
Ph.D. dissertation by the author (Yang, 2000) in
which the more than 90 specific and infraspecific
epithets are reduced to 25 species, including two
subspecies and ten varieties. A detailed mono¬

graphic account of this genus will he forthcoming.
Seven of the 13 Pyrenaria species that occur in
China require one or more new combinations or
other taxonomic treatment and are here published
in order to make the names to available for the
Flora oj China Theaceae treatment.

1. Pyrenaria diospyricarpa Kurz, J. Asia). Soe.
Bengal. 42(2): 60. 1873. TYPE: Myanmar.
Martaban, 15 Aug. 1872. 5. Kurz 1023 (holo¬
type, K).

Pyrenaria burmanica T. K. Paul & Nayar, J. Jap. But.
64(1): 8. 1989. Syn. nov. TYPE: Myanmar. Tenas-
seriu, Humatee, 5000 ft., 10 Apr. 1877. Geo Gallatly
695 (holotype, CAL not seen; isotype, CAL).

Except for the non-foliaceous bractlets and se¬
pals, Pyrenaria burmanica is morphologically in¬
distinguishable from P. diospyricarpa var. diospyri¬
carpa. With the observation of gradual transitions
from non-foliaceous bractlets and sepals to folia-
ceous ones in my field investigations, the gap be¬
tween both taxa is bridged.

la, Pyrenaria diospyricarpa Kurz var. dios¬
pyricarpa

Distribution and habitat. Southern Yunnan,
China; Myanmar; Thailand; Vietnam. Forests on
hills, slopes, or valleys between 1000 and 2000 m
(Yang, 1997).

I b. Pyrenaria diospyricarpa Kurz var. eainel-
liiflora (Kurz) S. X. Yang, comb, et stat. nov.
Basionym: Pyrenaria camelliiflora Kurz, J.
Asiat. Soc. Bengal. 40(2): 46. 1871. TYPE:
Myanmar. Martaban, Doyokee pass, 4000 ft.,
I). Brandis s.n, (holotype, CAL not seen).

This taxon is different from P. diospyricarpa only
by its branchlets and leaves with less pubescence.
It is more acceptable to treat it as a variety.

Distribution and habitat. Myanmar; Thailand;
Vietnam. Forests between 900 and 1500 m (Yang,
1997).

Novon 15: 379-382. 2005.

380

Novon

2. Pyrenaria jonquieriana Pierre ex Lanessan,

PI. Util. Colon. Franc., 295. 1886. TYPE:
l.aos. Attapeu Basin, western bank of Mekong,
Harmaiul 4005 (holotype. P).

2a. Pyrenaria jonquieriana Pierre ex Lanessan
suhsp. jonquieriana

Distribution and habitat. Laos. Forests between
400 and 1 700 in.

2I>. Pyrenaria jonquieriana Pierre ex Lanessan
subsp. miiltisepnln (Merrill cK Chun) S. X.
Yang, comb, et stat. nov. Basionym: Tutcheria
multisepala Merrill & Chun, Sunyatsenia 2(1):
41. 1934. Parapyrenaria multisepala (Merrill
& Chun) Hung T. Chang, Acta Sei. Nat. Univ.
Sunyatseni 18(3): 74. 1979. Pyrenaria multi¬
sepala (Merrill & Chun) H. K.eng, Card. Bull.
Singapore 26(1): 134. 1072. TYPE: China.
Hainan: Wanning (Manyun), Tungshanling, 5
Apr. 1932, II. V. Liang 61515 (holotype, IBSC:
isotype, PE).

Except that it has more sepals (more than 20),
Tutcheria multisepala is almost the same as Pyr¬
enaria . jonquieriana. Considering their disjunct dis¬
tribution, it is suitable for us to treat them as dif¬
ferent subspecies.

Distribution and habitat. Hainan, China. For¬
ests between 800 and 1000 m.

3. Pyrenaria khasiana R. N. Paul, Bull. Bot.

Soc. Bengal. 33: 115. 1979. TYPE: India.
Meghalaya, kliasi lulls, 675—1000 m, S. Kur:
161 (holotype, CAL not seen; isotype, CAL).

Pyrenaria tibetana Hung T. Chang, Acta Sei. Nat. Univ.
Sunyatseni 22(3): 64. 1983. Syn. nov. TV PE: China.
SF, Tibet, Medog, NW of Didongzhan, in evergreen
forest, 3 Aug. 1974. Qinghai-Xizang (Tibet) Exped.
Acad. Sin. 74-1040 (holotype, PK; isotype, klJN).

No natural differences can be found between the
type specimens of Pyrenaria khasiana and P. tibe¬
tana. It is thus inappropriate to treat them as dif¬
ferent species.

Distribution and habitat. Southeastern Xizang
(Tibet). China; northeastern India. Evergreen forest
between 675 and 2100 m.

4. Pyrenaria niaculatoclada (Y. k. Li) S. X.

Yang, comb. nov. Basionym: Tutcheria macu-
latoclada Y. k. Li, Guihaia 4(1): 35. 1984.
TYPE: China. Guizhou: Sandu, Yaorenshan,
alt. 910 m. 8 Sep. 1982. C. /. Dang 2100
(holotype, I IG AS).

Tutcheria maculatoclada lias been reduced into
Pyrenaria greeniae (Chun) keng (Yang. 1997) or
Tutcheria greeniae Chun (Chang, 1998). Its recog¬
nition as an independent species in Pyrenaria is
based on its succulent pericarp.

Distribution and habitat. Northern Cuangxi
and southern Guizhou, China. Forests between 700
and 1000 m (Yang. 1997).

5. Pyrenaria niicrocarpa (Dunn) II. keng,
Gard. Bull. Singapore 26(1): 134. 1972. Bas¬
ionym: Tutcheria niicrocarpa Dunn, .). Bot. 47:
197. 1909. TYPE: China. Guangdong: Boluo,
Lo Fou Mountains, Ford 610 (lectotype, des¬
ignated here, IBSC).

Then shinkoensis Hayata. J. Coll. Sei. Imp. Univ. Tokyo
30(1): 45. 1911. Syn. nov. Tutcheria shinkoensis
(Ilayata) Nakai, J. Jap. Bot. 16(12): 708. 1940.
TYPE: China. Taiwan: Taipeh, Shinko, Remogansha,
8 June 1906. 7. Kawakami & II. Mori 1424 (holo¬
type, 'll).

Tutcheria tenuifolia Hung I. Chang, Acta Sei. Nat. Univ.
Sunyatseni 22(2): 108. 1983. Svn. nov. 1YPE: Chi¬
na. Cuangxi: Uongzhou, Daqing Mt., near Nailian
Village, 27 June 1957, S. II. Chun 12044 (holotype,
IBSC; isotype, kl N).

Dunn (1909) listed two specimens under Tutch¬
eria niicrocarpa Dunn: Ford 6 10 and Hongkong
llerh. 2050. I checked them in IBSC. Both of them
correspond to the protologue, but the first collection
is in better condition, so it is selected as the lec¬
totype here.

Then shinkoensis was treated as a variety of Pyr¬
enaria niicrocarpa based only on its description
(Yang, 1997). I studied the type specimen of T.
shinkoensis at Tl and found that it shares almost all
features with P niicrocarpa except that a lew leaves
of T. shinkoensis are subacuminate.

Tutcheria tenuifolia is different from P. niicro¬
carpa in its obvious pedicel (7—8 nun long). In fact,
the pedicel of P niicrocarpa is variable, from I to
10 mm in length.

5a. Pyrenaria niicrocarpa (Dunn) II. keng var.
niicrocarpa

Distribution and habitat. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hainan, Jiangxi,
Taiwan, and Zhejiang, China; Ryukyu Islands, Ja¬
pan; Vietnam. Forests in mountain valleys or along
streams between 300 and 1000 m (Yang, 1997).

5b. Pyrenaria niicrocarpa (Dunn) II. keng var.
ovalifolia (H. L. Li) T. L. Ming & S. X. Yang,
Chin. J. Appl. Environm. Biol. 3(3): 287.
1997. Basionym: Tutcheria ovalifolia II. L. Li,

Volume 15, Number 2
2005

Yang

New Combinations in Pyrenaria

381

J. Arnold Arbor. 25(2): 209. 1944. TYPE: Chi¬
na. Hainan: Baoting, in forest, 16 Nov. 1936,

S. A. Lau 28218 (holotype. A; isotype, I BSC).

Pyrenaria buisanensis (Sasaki) C. F. Hsieh, S. Z. Yang &
M. H. Su, Taiwania 49(3): 202. 2004. Syn. nov. Bas-
ionym: Camellia buisanensis Sasaki, Trans. Nat.
Hist. Soc. Form. 21: 222. 1931. Camelliastrum buis¬
anensis (Sasaki) Nakai, J. Jap. Hot. 16(12): 700.
1940. TYPE: China. Taiwan: Pingtung |Pingtlong]
Co., Mt. Jenlishan, 30 May 2004, K. T. lee Y. k.
Wu s.n. (neotvpe designated by Su et al. (2004: 202),
TAI).

Tutcheria symplocifolia Merrill & F. P. Metcalf, Eingnan
Sci. J. 16(2): 172. 1937. Syn. nov. TYPE: China.
Guangdong: Ta-pu, Tai Mo Shan, If) July 1932. It!

T. Tsang 21196 (holotype, IBSC; isotype, PE).
Tutcheria taiwanica Hung T. Chang & S. X. Ren, Acta

Sci. Nat. Univ. Sunyatseni 30(1): 71. 1991. Syn. nov.
TYPE: China. Taiwan: Pingtung [Pingdong] Co.,
Bankinsing, tree 20 ft., 25 Feb. 1893—1894,4. Hen¬
ry 123 (holotype, US).

Through a wide range of herbarium and field in¬
vestigations. two variable foliar trends in Py renaria
microcarpa have been determined: from oblong to
obovate in shape and from acuminate to acute to
obtuse to blunt at the apex, which resulted in the
establishment of a few ambiguous species, includ¬
ing Tutcheria ovalifolia, T. symplocifolia, T. taiwan¬
ica, and C. buisanensis. Hereby, those species with
leaves obovate in shape and obtuse to blunt at the
apex are treated as the variety ovalifolia.

Distribution and habitat. Fujian. Guangdong,
Hainan, and northern Taiwan, China. Forests below
100 to 1000 m (Yang, 1997).

6, Pyrenaria speetabilis (Champion) C. Y Wu X
S. X. Yang, comb. nov. Basionym: Camellia
speetabilis Champion, Hooker’s J. Bot. Kew
Card. Misc. 3: 310. 1851. Thea speetabilis
(Champion) Kochs, Bot. Jahrb. Syst. 27: 595.
1900. Tutcheria speetabilis (Champion) Dunn,
J. Bot. 46: 324. 1908. TYPE: China. Hong
Kong: Hong Kong Island, 1847, Champion s.n.
(holotype. K).

Tutcheria brachycarpa Hung T. Chang, Acta Sci. Nat.
Univ. Sunyatseni 22(2): 109. 1983. Syn. nov. TYPE:
China. Guangxi: Jinxiu (Yaoshan), S. S. Sin 328 (ho¬
lotype, IBSC).

The type specimen of Tutcheria brachycarpa
shows no essential difference from P. speetabilis. It
is unreasonable to retain it as a species.

6a. Pyrenaria speetabilis (Champion) C. Y. Wu
X S. X. Yang var. speetabilis

Distribution and habitat. Fujian, Guangdong,
Guangxi, and Hong Kong, China; northern Viet¬

nam. Broadleaf evergreen forests between 300 and
1500 m (Yang, 1997).

6b. Pyrenaria speetabilis (Champion) C. Y. Wu
& S. X. Yang var. greeniae (Chun) S. X. Yang,
comb, et stat. nov. Basionym: Tutcheria green-
iae Chun. J. Arnold Arbor 9(2—3): 129. 1928.
T\ PE: China. Guangdong: North Biver region.
Pan lin tsze, in open woods, 25 Dec. 1927, IF
Y. Chun 5937 (holotype, IBSC; isotypes. A,
PE).

Tu t ch eri a a usl rosin ica Hung T. C hang | as ''austro-sinica" |.
Acta Sci. Nat. Univ. Sunyatseni 22(2): 167. 1983.
Syn. nov. TYPE: China. Guangdong: Yangohun,
Heweishan, Mei/.i Region, 7 June 1957, lab. Geobot.
Inst. Hot. Austro-Sin. 4258 (holotype. IBSC).
Tutcheria rostrata Hung T. Chang, Acta Sci. Nat. Univ.
Sunyatseni 22(2): 106. 1983. Syn. nov. 14 PE: Chi¬
na. Guangdong: Yangchun, Heweishan to Sanehahe.
19 Oct. 1957, IF T. Tin 30962 (holotype, CANT).
Pyrenaria turbinata S. X. Yang, Chin. J. Appl. Environm.
Biol. 3(3): 281. 1997. Syn. nov. TYPE: China. Gu¬
angxi: Nanning, cultivated in Guangxi Forest Insti¬
tute (introduced from Hexian), 24 Oct. 1991. .S'. A.
Yang & IF ,/. Zhang 91041 (holotype, KIN).

The fruit of this taxon is < ] n i te variable in peri¬
carp thickness (from 1 to 2. even to 4 mm), locale
numbers (ranging from 3, 4, to 5), apex shape (from
acute to obtuse to blunt), which bridge these mor¬
phological gaps between Tutcheria austrosinica, T.
rostrata, Pyrenaria turbinata, and 7. greeniae.

Distribution and habitat. Southern Fujian,
Guangdong, Guangxi. southern Hunan, and south¬
ern Jiangxi, China. Broadleaf evergreen forests be¬
tween 300 and 1200 m (Yang, 1997).

7. Pyrenaria wuiana (Hung T. Chang) S. X.
Yang, comb. nov. Basionym: Tutcheria wuiana
Hung T. Chang, Acta Sci. Nat. Univ. Suny¬
atseni 6(1): 29. 1961. TYPE: China. Guang¬
dong: Funding, Jiayi, Yadaodashan Hill, 6 Sep.
1959, B. S. Wang & L. C. Xu 0014 (holotype,
SYS).

Based on the leaf-like bractlets, large and oblong
sepals, and acuminate fruit, the previous taxonomic
treatment reducing this species into Pyrenaria
grandiflora (Wu) Ming & S. X. Yang (Yang, 1997)
should be corrected.

Distribution anti habitat. Western Guangdong
and eastern Guangxi, China. Forests between 800
and 900 m (Yang. 1997).

Acknowledgments. The study was partially sup¬
ported by funds from the National Natural Sciences
Foundation of China (Grant no. 30470136), a pro¬
ject of the Provincial Natural Sciences Foundation

382

Novon

Committee of Yunnan, P. R. China (97C039Q), and
the special support grants from The Chinese Acad¬
emy of Sciences for biotaxonomy and floristics. I
am most grateful to A, CAL, CANT, HGAS. IBSC,
k., P, PE, SYS, Tl, and US for kindly providing
specimens. I also express my deep gratitude to V.
C. Hollowed (MO). N. Turland (MO), and R. Bar¬
tholomew (CAS) for critical reviews ol the manu¬
script.

I literature Cited

Chang, II. T. 1998. Tutcheria. Pp. 195-206 in Flora Rei-
publieae Popularis Sinieae, Vol. 49(3). Science Press,
Beijing.

Dunn. S. T. 1909. New Chinese plants. J. Hot. 47: 197.
Keng, H. 1972. Two new theaoeous plants from Malaysia

and a proposal to reduce Tutcheria to a synonym of
Pyrenaria. Card. Hull. Singapore 26: 127—135.

- . 1980. The genus Pyrenaria (Theaeeae) in Malay¬
sia. Card. Hull. Singapore 33(2): 262-289.

Su, M.-H., S.-Z. Yang iK C.-F. Ilsieh. 2004. The identity
of Camellia buisanensis Sasaki (Theaeeae). Taiwania
49(3): 201-208.

Yang S. X. 1997. A preliminary revision ol Pyrenaria from
China and its adjacent regions. Chin. J. Appl. Envi-
ronm. Biol. 3(3): 276—288.

- . 2000. Systematics, Diversification and Geograph¬
ical Distribution ol Pyrenaria sensu lato (Theaeeae).
Ph.D. Dissertation, Kunming Institute of Botany ol the
Chinese Academy of Sciences, Kunming. | Unpub¬
lished.!

- & T. L. Ming. 1995a. Fmbryologieal studies on

genera Pyrenaria and Tutcheria of family Theaeeae.
Aeta Hot. Yunnan. 17(1): 67-71.

- X - . 1995b. Studies on tin* systematic po¬
sition of genera Pyrenaria, Tutcheria and Parapyrenaria
of family Theaeeae. Aeta Hot. Yunnan. 17(2): 192—196.

Lectotypifieation of Machilus platycarpa and
Machilus gracillima (Lauraceae)

Yang Yang

Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, 20 Nanxincun, Xiangshan, Beijing, China, ephedra@ibcas.ac.cn

I Irnk van der Werff

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

henk.vanderwerff@mobot.org

Abstract. In the description ot two species of
Machilus , Chun cited two type collections. In each
case, one of the type collections does not belong in
the I .auraeeae. Lectotypifieation of these two names
is proposed.

Key words: China, Lauraceae, lectotypifieation,
Machilus.

During a project aimed at cataloging names
and types of Chinese Lauraceae, problems were
encountered with the types of two species of the
genus Machilus Nees described by Chun (1953).
In Chun’s article, species were first described in
Chinese, including citation of a type and para-
types when available, followed by a translation
in Latin, again with citation of a type and para-
types. In two cases the type collection cited in
the Ch inese description was different from the
type collection cited in the Latin description.
Both names have been used in subsequent pub¬
lications (Lee, 1982; Wei et al.. 1991). In order
to avoid confusion about the correct tvpe speci¬
men for these two names in Machilus, we lecto-
typify both names and explain the reasons for our
lectotypifieation in this note.

1. LECTOTY PI FIXATION OK MACHILUS I’lATYCARPA
CtIUN

Machilus platycarpa was published in Chun
(1953), with the Chinese description on page 163
and the Latin description on page 164. Chun des¬
ignated S. I*. Ko 51351 as the type collection in the
Chinese description and C. Wang 33722 in the Lat¬
in description. Chun worked at IBSC. 1 1 is original
material should be expecteil in IBSC, and indeed
both collections are present in that institution. The
first author had the opportunity to study both col¬
lections and found that C. Wang 33722 belongs to
the Lauraceae and fits the description of Machilus

platycarpa Chun. Further, the collection .S'. P. Ko
51351 does not belong to the Lauraceae, but rep¬
resents Oreocnide frutescens (Thunberg) Miquel
(Urtieaceae). Clearly, a lectotype is needed in this
situation, which we designate below.

Machilus platycarpa Chun, Acta Lhytotax. Sin. 2:
163, pi. 29. 1953. TYPE: China. Guangdong:
Yangehun County, Mount Liping, 13 Nov.
1935, C. Wang 33722 (lectotype, designated
here, IBSC; isotypes, IBK. MO).

2. Lectotypikication of Machii.i s <;hacii.uu \
Chun

The case of Machilus gracillima is similar to the
preceding one. Chun designated C. Wang 41 072 as
the type in the Chinese description and C. Wang
41022 as the type in the Latin description. A
search in IBSC yielded the collection C. Wang
41072, which indeed belongs to the Lauraceae and
fits the description of M. gracillima. However, the
collection C. Wang 41022 could not be located in
IBSC; records kept at IBSC indicated that C. Wang
41022 belonged to Stachyurus chinensis Kranchet
(Stachyuraceae). Subsequently, the first author
found a duplicate of C. Wang 41022 in PE. This
specimen represented Stachyurus chinensis, as in¬
dicated by the records at IBSC. Again, lectotypifi-
cation is required to resolve this taxonomic/nomen-
clatural confusion.

Machilus gracillima Chun, Acta Phytotax. Sin. 2:
170, pi. 35. 1953. TYPE: China. Guangxi:
Nandan County, Mangchang, Mengguo, 10
July 1937, C. Wang 41072 (lectotype, desig¬
nated here, IBSC).

Novon 15: 383-384. 2005.

384

Novon

Literature Cited

Chun, W. Y. 1953. Materials for Flora of South China, new
species of Machilus of Lauraceae. Acta Phytotax. Sin.
2(3): 163-171, pi. 29-35.

Lee, S. k. 1932. Machilus. In: H. W. Li (editor), Flora

Reipublicae Popularis Sinicae, Vol. 31. Science Press,
Beijing.

Wei, Y. T., F. N. Wei & G. Z. Li. 1991. Lauraceae. In: S.

k. Lee & C. F. Liang (editors). Flora ol Guangxi, Vol.

l. Guangxi Science and Technology Publishing House,
Nanning.

Volume 15. Number 2. pp. 253—384 of NOVON was published on 13 July 2005.

Volume 15
Number 3
2005

NOVON

Transfer of the South African Genera Brachycarpaea, Cycloptychis,
Schlechteria, Silicularia, and Thlaspeocarpa to
Heliophila (Brassicaceae)

lh son A. Al-Shehbaz

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

ihsan.al-shehbaz@mobot.org

Klaus Mummenhoff

Department of Botany, Faculty of Biology/Chemistry, University of Osnabriiek, Barbaras! r.

I I, D-49069 Osnabriiek, Germany, rnummenhoff@biologie.uni-osnabrueck.de

ABSTRACT. The genera Brachycarpaea , Cyclop¬
tychis, Schlechteria, Silicularia, and Thlaspeocarpa
are united with Heliophila (Brassicaceae). Except
for Brachycarpaea, the genera are treated as new
synonyms of Heliophila. The characters previously
used to distinguish these genera are critically eval¬
uated. file retention of Brachycarpaea and Sihcu-
laria in Heliophila would not necessitate any no-
menclatural novelties, but t lie transfer of species of
the remaining three genera to Heliophila requires a
new' combination and four new names. The new
combination H. namaquensis and the new names II.
monosperma, H. maraisiana, II. hurkana, and II.
suborbicularis are proposed.

Key words: Brachycarpaea, Brassicaceae, Cy¬
cloptychis, Heliophila, Namibia, Schlechteria, Sili¬
cularia, South Africa, Thlaspeocarpa.

As delimited by Appel and Al-Shehbaz (1997).
the southern African tribe Heliophileae (Brassica¬
ceae) consists of 80 species in the genera Helio¬
phila L. (73 spp.), Brachycarpaea DC. (1 sp.), Cy¬
cloptychis E. Meyer ex Sender (2 spp.), Schlechteria
Bolus (I sp.), Silicularia Compton (I sp.). and
Thlaspeocarpa C. A. Smith (2 spp.). Of these. 78
species, including those of the 5 smaller genera
above, are distributed in the Cape provinces of
South Africa and 8 grow in Namibia, 3 in Lesotho,

and I in Swaziland (Marais, 1970; Pienaar & Nich¬
olas. 1988; Bean. 1990). With the exception of Bra¬
chycarpaea, which has spirally coiled cotyledons,
the taxa of the tribe have diplecolobal cotyledons
(twice transversely folded). The latter type of cot¬
yledons also occurs in 3 species of h-pidiuin L.
subsect. Diploploca Hewson (Hewson, 1982), but
molecular studies (Mummenhoff, unpublished)
clearly show this feature evolved independently in
I cpidi urn and the Heliophileae.

Molecular studies bv Miihlhausen (2002) and
Mummenhoff et al. (2005), using internal tran¬
scribed spacers of nuclear ribosomal DNA (ITS)
and the trn I , intron of cpDNA, reveal that all six
genera of the Heliophileae form a monophyletic
clade, with Brachycarpaea, Cycloptychis, Schlech¬
teria, Silicularia, and Thlaspeocarpa nested within
the morphologically far more diversified Heliophila.
It is concluded that the Heliophileae should consist
of one inclusive Heliophila, and the nomenelatural
changes needed to unite the smaller genera with
Heliophila are proposed below. In our opinion,
these smaller genera are based on one-character
fruit differences and. in the absence of fruits, their
species cannot be distinguished from those of He¬
liophila, For a detailed comparison of all six "gen¬
era." see Table 2 in Appel and Al-Shehbaz I 1997).
Koch et al. (2003) summarized many similar mo-

Novon 15: 385-389. Published on 21 September 2005.

386

Novon

leeular studies in the Brassicaceae that leil to the
union of monotypic or oligotypie genera witli their
larger, paraphyletie, more inclusive relatives.

Brachycarpaea

Hayek (191 1) placed Brachycarpaea in tlie tribe
Heliophileae, and Schulz (1936) assigned it to the
Lepidieae. In their tribal assignment of this genus,
Schulz emphasized the presence of angustiseptate
fruits (flattened at a right angle to the septum),
whereas llayek relied on the overall similarities in
habit, flowers, distribution, and the presence of lin¬
ear cotyledons. Except for fruit compression and
cotyledonary coiling, B. juncea (P. .). Bergius) Mar¬
ais is indistinguishable morphologically from spe¬
cies of Cycloptychis and many Heliophila. The pres¬
ence of angustiseptate fruits in Brachycarpaea is
unique in the Heliophileae, but as shown by Appel
and Al-Shehbaz (2003), this and other types of fruit
compression occur within several individual genera
of the Brassicaceae (e.g., Graellsia Boissier, les-
querella S. Watson (now Physaria (Nuttall) A.
Gray), Smelowskia C. A. Meyer).

Indeed, the fruits of B. juncea are only slight I v
angustiseptate, but they have woody, one-seeded,
indehiscent valves borne on a tiny carpophore and,
therefore, they are schizoearpie much like those of
Cycloptychis. The latter genus is easily distin¬
guished from Brachycarpaea by having a longer
carpophore and woody, rugose (vs. smooth) latisep-
tate (flattened parallel to the septum) instead of an¬
gustiseptate fruits. In tin* molecular study (Mum-
menhoff et ah. 2005), the two genera form a
well-supported elade nested within Heliophila.

The single species of Brachycarpaea was origi¬
nally described by Bergius (1767) as Cleome juncea
P. J. Bergius. Druee (1017) transferred it to Helio¬
phila and Marais (1966) to Brachycarpaea, but
Marais (1966, 1970) overlooked I truce's transfer.
Therefore, the species should be known in Helio¬
phila as //. juncea (P. J. Bergius) Druee, and no
nomenclatural changes are needed from the union
of Brachycarpaea and Heliophila.

Cycloptychis

In the absence of fruits, both species of Cyclop¬
tychis are morphologically indistinguishable from
Brachycarpaea juncea and several species of Helio¬
phila. The genus is based solely on having sehiz-
ocarpic. latiseptate fruits with a distinct carpophore
and rugose, woody valves. All species of Heliophila
have dehiscent, smooth, non-woody fruits without a
carpophore. As indicated above, molecular data
(Mummenhoff et ah. 2005) reveal that Cycloptychis

forms with Brachycarpaea a monophyletic subelade
deeply nested within Heliophila. Although the pres¬
ence of typically schizoearpie fruits with a distinct
carpophore in Cycloptychis is unique among all of
th<‘ Old World genera of Brassicaceae, the feature
evolved independently in the South American Men-
onvillea DC. (Rollins, 1955; Appel & Al-Shehbaz,
2003). As shown below, the transfer to Heliophila
of the basionyms of C. virgata (Thunberg) E. Meyer
ex Sender and C. marlothii 0. E. Schulz would cre¬
ate two later homonyms, and the new' names H.
maraisiana and //. hurkana, respectively, are pro¬
posed to facilitate those transfers and the union of
Cycloptychis with Heliophila.

SCHLECHTERIA AND THIASPEOCARPA

Hayek (1911) assigned Schlechteria to the tribe
Alysseae and Schulz (1936) placed it in the Lu-
narieae, but both authors overlooked the dipleco-
lobal cotyledons in S. capensis Bolus. However,
Compton (1953) was the first to consider the genus
to be closely related to Heliophila , a position ac¬
cepted by subsequent authors (e.g.. Marais, 1970;
Appel & Al-Shehbaz, 1997).

Thlaspeocarpa was proposed by Smith (1931) as
a new name replacing the later homonym Palms-
truckia Sonder. The genus remained monotypic for
nearly 100 years, and Marais (1966) added a sec¬
ond species to the one first discovered by Sonder
(1860). Thlaspeocarpa has long been considered to
be closely related to Heliophila, and their place¬
ment in the same tribe was not questioned.

Both Schlechteria and Thlaspeocarpa differ from
Heliophila primarily by having indehiscent. esep-
tate fruits. These differences in fruit characters are
minor, and one finds similar or more extreme sit¬
uations within several genera of Brassicaceae. For
example, Lepidium (including Cardaria Desvaux,
Coronopus Zinn, and Stroganowia Karelin & Kiri¬
lov). Physaria (including Lesquerella), Tropidocar-
pum Hooker (including Twisselmannia Al-Sheh¬
baz), and Vella L. (including Boleum Desvaux and
Euzomodendron Cosson) have species with dehis¬
cent or indehiscent and septate or eseptate fruits.

Appel and Al-Shehbaz (1997) separated Thlas¬
peocarpa from Schlechteria primarily by their being
annual herbs instead of shrubs with appendaged
versus unappendaged filaments and winged versus
w ingless seeds. However, all of these characters oc¬
cur in a mosaic pattern within Heliophila. Based on
our examination of all 73 species of Heliophila . 44
species are annuals, 7 are perennial herbs, and 22
are shrubs or subshrubs. Of the annuals, 9 have
staminal appendages and seed wings, 6 have nei-

Volume 15, Number 3
2005

Al-Shehbaz & Mummenhoff
Transfers of South African Genera

387

iher, 5 have seed wings but not appendages, and
19 have staminal appendages but not seed wings.
Seed or filament information is not recorded for 5
species. Of the 22 shrubs and subshrubs, 2 have
die seed wing and filament appendage, 14 have
neither, I has only the appendage, and 3 have only
the seed wing. Therefore, the differences between
Schlechteria and Thlaspeocarpa are not supported,
and the union of these genera with Heliophila is
justified on morphological and molecular grounds.

The transfer of Schlechteria capensis Bolus and
the basionym of Thlaspeocarpa capensis (Sonder) C.
A. Smith to Heliophila would create later hom¬
onyms, and new names are proposed below to ac¬
commodate such transfers.

SlUCULARIA

Silicularia basically mirrors the morphological
situation of Schlechteria and Thlaspeocarpa in re¬
lation to Heliophila. It, too, has indehiscent fruits
without a septum (sensu Appel & Al-Shehbaz,
1997) or with a fenestrated septum (Compton,
1.953), but Silicularia (fillers from these three gen¬
era by having woody, rugose fruits. In all other aspects
of the plant, Silicularia polygaloides (Schlechter)
Compton, which is a shrub with unappendaged fil¬
aments and wingless seeds, is remarkably similar
to numerous species of Heliophila and to Schle¬
chteria capensis. In the absence of fruits, Compton
(1953) indicated that the species (as Silicularia sig-
illata Compton) is indistinguishable from those of
Brachycarpaea and Schlechteria. In fact, the spe¬
cies was originally described by Schlechter (1899)
as II. polygaloides Schlechter and, therefore, its re¬
tention in Heliophila would not necessitate any no-
menclatural changes, though Compton (1953), see
below, mishandled the nomenclature of this and
other closely related species of Heliophila.

Heliophila L., Sp. PI., ed. 2, 2: 926. 1763. TYPE:
Heliophila integrifolia L, Sp. PI., ed. 2, 2:
926. 1763, nom. i 1 1 eg. [Art. 52.1] = Heliophila
africana (L.) Marais, in L. E. Codd et ah, FI.
S. Africa 13: 55. 1970; based on Cheiranthus
africanus I,., PI. Rar. Afr. 13. 1760.

Brachycarpaea DC.. Mem. Mas. Hist. Nat. 7: 247. 1821.
TYPE: Brachycarpaea various DC., Syst. Nat. 2:
699. 1821. = Brachycarpaea j unce a (P. J. Bergius)
Marais, Bothalia 9: 112. 1966. Heliophila juncea (P.
J. Bergius) Druee, Bot. Exch. Club Soc. Brit. Isles
1916: 627. 19t7; based on Cleome juncea P. J. Ber¬
gius, Descr. PI. Cap. 164. 1767.

Cycloptychis E. Meyer ex Sonder, in Harvey & Sonder, FI.
Cap. 1: 34. I860. Syn. nov. TYPE: Cycloptychis vir-
gata (Thunberg) E. Meyer ex Sonder, in Harvey &
Sonder, FI. Cap. I: 34. I860 (lectotype, designated

by Marais, 1970: 80); based on Cleome virgata
Thunberg, Prodr. 109. 1800.

Schlechteria Bolus, Bot. Jahrb. Syst. 24: 455. 1987. Syn.
nov. TYPE: Schlechteria capensis Bolus, Bot. Jahrb.
Syst. 24: 455. 1897.

Silicularia Compton, S. African J. Bot. 19: 147. 1953.
Syn. nov. TYPE: Silicularia sigillata Compton, S. Af¬
rican ,1. Bot. 19: 147. 1953. = Heliophila polygalo¬
ides Schlechter, Bot. Jahrb. Syst. 27: 137. 1899, non
Heliophila polygaloides (Sonder) Compton, S. Afri¬
can .1. Bot. 19: 152. 1953.

Thlaspeocarpa C. A. Smith, Bull. Misc. Inform. Kew 1931:

155. 1931. Syn. nov. TYPE: Thlaspeocarpa capensis
(Sonder) C. A. Smith, Bull. Misc. Inform. Kew 1931:

156. 1931; based on Palmstruckia capensis Sonder,
in Harvey & Sonder, FI. Cap. 1: 35. 1860. based on
Peltaria capensis Thunberg, Prodr. 107. 1800, non
F. f., Suppl. PI. 296. 1782.

Heliophila monosperma Al-Shehbaz & Mum¬
menhoff, nom. nov. Replaced name: Schlech-
terio capensis Bolus, Rot. Jahrb. Syst. 24: 455.
1897, non Heliophila capensis Kuntze, Revis.
Gen. PI. 3(2): 5. 1893, nec Heliophila capensis
(L. f.) C. A. Smith, Bull. Misc. Inform. Kew
1931: 155. 1931. TYPE: South Africa. Cape:
Clanwilliam, Koudeberg, 1300 m, 30 Aug.
1896. R. Schlechter B765 (holotype, not seen;
isotypes. RM, K, MO).

Thi s new name should be used when Schlechteria
is united with Heliophila.

Heliophila juncea (P. J. Bergius) Druce, Rot.
Exch. Club Soc. Rrit. Isles 1916: 627. 1917.
Basionym: Cleome juncea P. ,1. Bergius, Descr.
PI. Cap. 164. 1767. TYPE: South Africa.
Grubb s.n. (holotype, SBT not seen).

This species name should be used for the trans-
fer of Brachycarpaea to Heliophila. flic name was
not listed in Marais (1970). I)e Candolle (1821)
listed the nomen nudum H. juncea Vald in the syn¬
onymy of //. suavissima Burch ex DC., but that
name has no relevance to the species recognized
herein.

Heliophila inaraisiana Al-Shehbaz N Mummen¬
hoff, nom. nov. Replaced name: Cleome virgata
Thunberg, Prodr. 109. 1800, non Cleome vir¬
gata Steven, in Candolle, Prodr. I: 240. 1824,
nec Heliophila virgata Burch ex DC., Syst.
Nat. 2: 693. 1821. Cycloptychis virgata (Thun¬
berg) E. Meyer ex Sonder, in Harvey & Sonder,
FI. Cap. 1: 34. 1860. TYPE: South Africa. C.
P. Thunberg s.n. (holotype, UPS not seen).

Heliophila maraisiana is named in honor ofWes-
sel Marais, whose accounts of the South African
Brassicaceae (Marais, 1966, 1970) remain classic.

388

Novon

The species is most closely related to //. hurkana ;
both species were previously placed in the genus
Cycloptychis. The characters distinguishing these
species are discussed under //. luirkana below.

Ilcliophila iiama<|iiensis (Marais) Al-Shehbaz &
Mummenhoff, comb. nov. Basionyrn: Thlaspeo-
carpa namaquensis Marais, Rothalia 9: 1 10.
1960. TYPE: South Africa. Cape: Namaqua-
land. Steinkopf, ./. II. I*. Acock s 19536 (holo-
type, PRE).

This specific epithet should not be confused with
Heliophila namaquana Rolus, an entirely different
species with linear, dehiscent siliques 1.5— 3.7 cm
X 1—1.5 mm, instead of suborbicular, indehiscent
silieles 1—1.3 X 0.8— I cm.

Ilidiophila hurkana Al-Shehbaz & Mummenhoff,
iioni. nov. Replaced name: Cycloptychis mar-
lothii O. E. Selmlz. Rot. Jahrb. Sysl. 66: 97.
1933, non Heliophila marlothii (). E. Schulz,
Rot. Archiv. 31: 533. 1931. TYPE: South Af¬
rica. Cape: Ceres, [Swartruggens], 1000 m,
Oct. 1903. R. Marloth 3362 (holotype, PRE).

The species is named after Herbert Hurka (Uni¬
versity of Osnabrtiek) for his life-long research on
the Rrassieaeeae and on the occasion of bis 65lh
birthday. Erom the other species of Ilcliophila. II.
hurkana is easily distinguished by being a shrub
with schizocarpic silieles, thick-walled valves, and
a slender carpophore. From the closely related II.
maraisiana, it easily distinguished by having fruits
8.5—12 mm long (excluding the style), styles 1.5—2
mm long, and leathery, finely reticulate-ridged fruit
valves with an obscure midvein. Ry contrast, II.
maraisiana has fruits 5-6 mm long (excluding the
style), stvles 4—6 mm long, and woody valves with
a prominent midvein and four ridges on each side.

Heliopliilu polygaloides Schlechter, Rot. Jahrb.
Syst. 27: 1.37. 1899. TYPE: South Africa.
Cape: Koude Rokkeveld, 5000 ft., 8 Sep.
1896, R. Schlechter 3900 (holotype, R).

As indicated above, this species was transferred
by Compton (1953) to the monotypic Silicularia.
However, Compton mishandled the nomenclature or
taxonomy of this species in two ways. First, he
transferred Cycloptychis polygaloides Sender (Sun¬
der. 1860) to Heliophila and made a later hom¬
onym, II. polygaloides (Sunder) Compton. Second, he
failed to retain t he name II. polygaloides Schlechter
in Heliophila and replaced it with the superfluous
and therefore illegitimate II. nuhigenoides Comp¬

ton. Taxonomieally, he described the new genus
and species Silicularia sigillata without realizing it
is nonspecific with Schlechter’s H. polygaloides.
Therefore, the sole species of Silicularia should be
known in Heliophila as II. polygaloides Schlechter.

Ilcliophila suborbieularis Al-Shehbaz & Mum¬
menhoff, nom. nov. Replaced name: Reltaria
capensis I hunberg, Prodr. 107. 1800. non Rel¬
taria cape ns is L. f„ Suppl. PI. 296. 1782. nee
Heliophila capensis Kuntze, Revis. Gen. PI.
3(2): 5. 1893. nee Heliophila capensis (L. f.)
C. A. Smith, Hull. Misc. Inform. Kew 1931:
155. 1931. Palmstruckia capensis Sunder, in
Harvey & Sunder, El. Cap. 1: 35. 1860. Tldas-
peocarpa capensis (Sunder) C. A. Smith, R nil.
Misc. Inform. Kew 1931: 155. 1931. TYPE:
South Africa. Cape: C. R. Thunberg s.n. (ho¬
lotype. UPS not seen).

The species name is descriptive of the fruit
shape. The name should be used for the transfer of
Thlaspeocarpa capensis to Heliophila.

Acknowledgments. We are grateful to Victoria
C. Hollowed, Peter Goldblatt. Nicholas J. Turland,
and an anonymous reviewer for their review and
valuable comments on the manuscript.

I .iterature Cited

Appel, O. & I. A. Al-Shehbaz. 1997. Re-evaluation of the
tribe Heliophileae (Brassicaeeae). Mitt. Inst. Allg. Bot.
Hamburg 27: 85-92.

- & - . 2003. Cruciferae. In K. knbitzki & C.

Bayer (editors). Families and Genera of Vascular Plants
5: 75—174. Springer- Verlag, Berlin, Heidelberg, and
New York.

Bean, P. A. 1990. The unusual newly discovered Helio¬
phila ephemera from the southern Cape, and its position
in the family Brassicaeeae. S. African J. Bot. 56: 670—
674.

Bergius, P. J. 1767. Descriptiones Plantarum ex Capite
Bonae Spei. I.aur. Salvius, Stockholm.

Compton, R. II. 1953. Silicularia. a new genus of Cruci¬
ferae, with notes on related genera. S. African J. Bot.
19: 147-155.

I)e Candolle, A. I’. 1821. Regni Vegetabilis Systema Na-
turale. Treuttel & Wiirtz, Paris.

I truce. G. C. 1917. Nomenelatorial notes: Chiefly African
and Australian. Rep. Bot. Exch. Club Soc. Brit. Isles
1916: 601—653.

Ilayek. A. von. 1911. Entwurf eines Crucifcren-Systems
auf phylogenetischer Grundlage. Beih. Bot. Centralbl.
27: 127-335.

Hewson, H. J. 1982. The genus Lepidium L (Brassica-
ceae) in Australia. Brunonia 4: 217—308.

Koch, M., I. A. Al-Shehbaz & k. Mummenhoff. 2003.
Molecular systematies, evolution, and population biol¬
ogy in the mustard family (Brassicaeeae). Ann. Missouri
Bot. Gard. 90: 151-17U

Volume 15, Number 3
2005

Al-Shehbaz & Mummenhoff
Transfers of South African Genera

389

Marais, W. 1966. Notes on South African Cruciferae.
Bothalia 9: 97—1 12.

- . 1970. Cruciferae. In L E. Codd, B. I). Winter,

I). J. B. killick & H. B. Bycroft (editors). Flora of
Southern Africa 13: 1-1 IB.

Muhl hausen. A. 2002. Molekulare Analysen zur Syste-
matik der Tribus Heliophileae (Brassicaceae) in Slid Af-
rika. M.S. Thesis, University of Osnabrlick. [Unpub¬
lished.]

Mummenhoff, K., I. A. Al-Shehbaz, f. I. Bakker, 11. P.
Linder & A. Miihlhausen. 2005. Phylogeny, morpholog¬
ical evolution, and speciation of endemic Brassicaceae
genera in the Cape flora of southern Africa. Ann. Mis-
sou ri Bot. Card.: 92: 399-123.

Pineaar, B. J. & A. Nicholas. 1988. Heliophila cornells-

bergia, a new species from the Richtersveld. Bothalia
18: 183-188.

Rollins, B. C. 1955. A revisionary study ol the genus Men-
onvillea (Cruciferae). Contr. Gray Herb. 1,7: 3-57.
Schlechter, B. 1899. Plantae Schlechterianae novae vel
minus cognitae describuntur. II. Bot. Jahrb. Syst. 27:
86-220.

Schulz, 0. E. 1936. Cruciferae. In A. Engler & H. Harms
(editors). Die naturlichen Pflanzenfamilien, Ed. 2. 1 7B:
227-658. Verlag von Wilhelm Engelmann, Leipzig.
Smith, C. A. 1931. Palmstruckia of the Flora Capensis.

Bull. Misc. Inform, kew 1931: 154—156.

Sonder, 0. W. 1860. Cruciferae. In Vi. 11. Harvey & 0. W.
Sonder (editors), Flora Capensis I: 19—54. Hodges,
Smith, Dublin.

Tripogon debilis (Poaceae), a New Species from Western China

Cai Lian-bing

Northwest Plateau Institute of Biology, The Chinese Academy of Sciences, Xining,
Qinghai 810001, China, cailh@nwipb.ac.cn

ABSTRACT. A new species of Tripogon from west¬
ern China (Sichuan Province), T. debilis L B. Cai,
is described and illustrated. This species is similar
to both T. chinensis (Franchet) Hackel and T. si-
chuanicm S. M. Phillips & S. I.. Chen, but distin¬
guished from these two species by its pendent
spikes, relatively long glumes and lemma awns,
denticulate upper glumes, and its paleas strikingly
shorter than the lemmas.

Key words: China, Poaceae, Tripogon.

The genus Tripogon Boomer cX Schultes consists
of approximately 30 species. One species is dis¬
tributed in Australia and two in America; the re¬
mainder are in Asia and Africa. The greater num¬
ber are concentrated in southern Asia, especially
in India, for which Bor (1960) reported Id species.
China has 10 previously recognized species, but
only I are endemic, and most of them occur in the
mountains of western China (Keng, 1959; Keng <X
Liou, I960; J. L Yang, 1983; X. L Yang, 1983,

1 990; Phillips <X Chen. 2002). liecent research on
the genus in this region revealed two collections
representing an unknown taxon. Thorough, imme¬
diate investigation and collection in the localities
of these specimens in the summer of 2003 revealed
that this taxon is different from all other species of
thi s genus in its external morphology. Hence, it is
reported as a new species here.

Tripogon debilis L B. Cai, sp. nov. TYPE: China.
Sichuan: Jiulong County, Tanggu, 3210 m,
29°6'N, 101°36'E, 23 July 2003, L. B. Cai
0318 (holotype, HNWP; isotype, CDBI). Fig¬
ure 1.

Culmi tenues, 25—34 cm alii. Spieae laxae, graciles, 8—
15 cm longae. Spiculae 6-8 mm longae (aristis exclusis),
6- ail 8-flores gerentes; glumis lanceolatis, 1- ad 3-ner-
vibus, infera integra, 2.5— 3.5 mm longa, supera 4.0— 5.0
mm longa, infra apicem hidenticulata; lemmate prime
3.5— 4.5 mm longo (arista exelusa). nervo medio in aristam
3—4 mm longam extenso; paleis lemmatibus ea. 1.1 mm
breviorihus.

Perennial herb, with thick fibrous roots. Culms
erect or slightly geniculate below, smooth, eaespi-
lose. 25—34 cm tall, ca. 0.8 mm diam., 2- or 3-

noded. Leaf sheaths shorter than the internodes,
white-villose at abaxial sheath summit; ligules very
short or nearly absent; leaf blades involute, 4-1 1
cm long, lower surface glabrous, upper surface sca¬
brous or sometimes pubescent toward the base.
Spikes lax. slender, pendent, 8-15 X ea. 0.25 cm;
inflorescence rachis internodes glabrous, generally
5—13 mm long. Spikelets sessile, brownish green,
oppressed along the rachis, usually 6- to 8-flow-
ered. 6-8 mm long (excluding awns); glumes lan¬
ceolate. glabrous, slightly membranous along the
margins; lower glume 1 -nerved, entire, pungent at
the apex, 2.5— 3.5 mm long; upper glume obscurely
3-nerved. 2 denticulate below the apex, 4—5 mm
long; lemmas glabrous, lanceolate, 3-nerved; first
lemma 3. 5^4.5 mm long, its median vein extended
into a straight awn 3—4 mm long, lateral veins ex¬
tended into macros 0. 3-0.5 mm long, toothed be¬
tween the macros and the median awn; callus ob¬
tuse, with ca. 0.3 mm long hairs; paleas
semi-membranous, 2-nerved, shorter than the lem¬
mas by ea. 1.1 mm. densely ciliate along upper
keels; anther I. rarely 2 or 3, yellow, 1.3— 1.5 mm
long. Caryopses unknown.

Ktymology. The epithet “debilis" refers to the
slender spikes and culms of the new species.

Distribution and habitat. Tripogon debilis is
known only from the Hengduanshan district of
Sichuan Province, western China, where it grows
on roadsides, wasteland, and stony slopes from
3100 to 3800 m. According to field observations,
this species is locally rare.

Relationships. This new species is similar to
Tripogon chinensis (Franchet) Hackel in its slender
culms to 34 cm tall, lax and graceful spikes, short
spikelets appressed to the rachis, and its lemmas
with only central awns. It differs from T. chinensis
in having pendent spikes and spikelets with 6 to 8
florets; lower glumes 2. 5-3.5 mm long; upper
glumes 4-5 mm long, with 2 teeth below tin1 apex;
lemma awns 3-4 mm long, and paleas being strik¬
ingly shorter than the lemmas. In addition, the gen¬
eral appearance of spikes in Tripogon sichuanicus
S. M. Phillif is & S. I„ Chen suggests some similar-

Novon 15: 390—392. I’iibi.ished on 21 September 2005.

Volume 15, Number 3
2005

Cai

Tripogon debilis (Poaceae)

391

Figure 1. Tripogon debilis L. B. Cai. — A. Plant. — B. Spikelet. — C. Lower glume. — I). Upper glume. —
view of the first lemma. — F. Ventral view of the first lemma. — G. Palea. — H. Anthers. Scale bar: A = 12
0.17 mm; C. I) = 0.13 mm; F.— H = 0.11 mm. A— II drawn from the holotype (/.. B. Cai Oiiltt. HINW P).

K. Lateral
nun: B =

392

Novon

ilies with this species, but T. sichuanicus can be
distinguished from this species by its erect or
slightly curved spikes, lower glumes 2.1— 2.7 mm
long; upper glumes entire 3.5— 4.3 mm long, lem¬
mas oblong-ovate 2.8— 3.0 mm long, with blunt
teeth and short central awns 1.8— 3.3 mm long, and
its paleas being subequal to the lemmas.

Morphologically, Tripogon debilis, T. chinensis, T.
sichuanicus, and some other speeies, such as T. hu-
milis X. L. Yang and T. purpurascens Duthie, form
a distinct group within Tripogon. Members of the
group have slender spikes, entire lower glumes, the
lemmas without the lateral awns (at most muero-
nate), and central awns that are shorter than or sub¬
equal to the lemmas. Such speeies as 7’ bromoides
Hoemer & Schultes, T. yunnanensis J. L. Yang ex
S. M. Phillips cK S. L. Chen, T. longiaristatus Hack-
el ex Honda, and T. filiform is Nees ex Steudel have
broader spikes, lobed lower glumes, and lemmas
with lateral awns as well as central awns that are
longer than the lemmas. Tripogon nanus Keng not
only has lateral and central awns longer than the
lemmas, but also distinct awns on the lobes be¬
tween lateral and median awns. Further research is
needed to determine whether these speeies groups
should be recognized as sections.

Paratypes. CHINA. Sichuan: Jiulong County, Tang-
gu, 1. H. Yang 2947 8 (HNWP); Daocheng County, Haizlii.
/. G. Lion 29030 (HNWP).

Acknowledgments. I thank Ma Bi-hua, Qiao
Zhi-xin, and Zhang Chang-quan for assistance with
fieldwork. Song Yu-zhu for preparing the excellent
illustration, and Chen Chun-fang for making the
holotype collection. This work was financially sup¬
ported by the Systematic and Evolutionary Biology
Foundation of the Chinese Academy of Sciences.

Literature Cited

Bor, N. L. I960. Grasses of Burma, Ceylon, India and
Pakistan. Pergamon Press, Oxford.

Keng, P. C. & I.. I.iou. I960. A study on the tribe Fra-
grostideae and its two genera new to China. Acta Bot.
Sin. 9: 48-75.

Keng, V L. (Editor). 1959. Pp. 458-462 in f lora lllus-
tralis Plantarum Primarum Sinicarum Cramineae. Sri.
Publ. House, Beijing.

Phillips, S. M. & S. L. Chen 2002. The genus Tripogon
(Poaceae) in China. Kew Bull. 57: 91 1-924.

Yang. J. L. 1983. New speeies of Cramineae from Sichuan.

Acta Bot. Yunnan. 5: 47—53.

Yang, X. L. 1983. New taxa of Tripogon and Aeluropus.
Acta Bot. Yunnan. 5: 72—74.

- . 1990. Tripogon. Pp. 58—63 in S. L. Chen (editor).

Flora Reipublieae Popularis Sinicae, Tomus 10(1). Sci¬
ence Press, Beijing.

Studies in I lie Capparaceae XXVI. Capparis bonifaziana, a New
Species and Western Ecuadorian Sister to the
Mostly Amazonian C. macrophylla

Xavier Cornejo

Herbario GUAY, Universidad de Guayaquil, Av. 25 de Julio via al Pto. mantinio, Casilla
09-01-10634, Guayaquil, Ecuador, xcornejoguay@hotmail.com. Present address: Department
of Botany, University of Wisconsin, 430 Lincoln Drive, Madison, Wisconsin 53706, U.S.A.

Hugh II. litis

Department of Botany, University of Wisconsin— Madison, 430 Lincoln Drive. Madison,
Wisconsin 53706, U.S.A. tscochra@wisc.edu. (To whom request for reprints should be sent.)

Ahstkact. From subgenus Capparidastrum s.l.
(Capparaceae), Capparis bonifaziana, a small to
medium-sized tree endemic to the western. Pacific
slope of Ecuador, is newly described as an allo-
patrie segregate of the much more widespread but
rare C. macrophylla s. str., of the mountains and
valleys of the northern Andes to the tropical low¬
lands of western and southwestern Amazonia. I he
new species differs by its more conspicuous and
larger episepalous glands, smaller anthers, ovaries,
and erect petals; smaller number of stamens; ellip¬
tic to oblong, rather than broadly ovate leaves; and
longer and narrower baccate fruits. Both taxa are
described, compared, illustrated, mapped, and with
all their known specimens listed.

Bksljmkn. Capparis bonifaziana. del subg. Cap¬
paridastrum, s.l., Capparaceae, es una nueva es-
pecie alopatrica, de arbolillo o arbol principal-
mente endemico del occidente de Ecuador,
segregada de la infrecuente C. macrophylla, s. str.,
ampliamente distribuida desde las montanas y val-
les al norte de los Andes hasta las tierras bajas
tropicales adyacenles de la Amazonia occidental y
suroccidental. Esta nueva especie diliere por sus
glandulas episepalas grandes v conspicuas; anteras,
ovarios y petalos erectos de menores dimensiones;
menor mimero de estambres; bojas mayormente
eh'pticas a oblongas antes (fue amplio ovadas; y fru-
tos abayados mas largos y angostos. Ambos taxones
son descritos, comparados, ilustrados y mapeados,
con todos los especimenes conocidos cilados.

Key words: allopatrie speciation, Capparaceae,
Capparidastrum. Capparis, edible fruits, endemic,
western Ecuador.

In the forested tropical lowlands on either side

Novon 15: c

of the Andes, from Venezuela to Bolivia, there grow
several taxa of Capparis subg. Capparidastrum
(DC.) Eiehler s.l.. Capparaceae (Eichler, 1865),
which are characterized by (1) pepo-like or bac¬
cate, spherical to cylindric, pendent fruits with an
exocarp that is indehiscent, thick and decomposing
lo tardily and partially dehiscent, by 3 to 1 thinly
coriaceous valves (only in C. macrophylla), with the
many seeds immersed in fleshy pulp and/or sareo-
testa; (2) racemose flowers on stiff, long, basally
naked peduncles; (3) massive, fleshy calyx glands,
which, sitting diagonally on top ol the inside basal
half of each of the sepals, strongly force the latter
to reflex and so partially cover and even obscure
them to a greater or lesser degree; and by (4) usu¬
ally glabrous leaves of often uneven sizes, on pul-
vinate petioles ol variable length.

Involved here are: (1) the widespread, northern
Peruvian- Ecuadorian, spherical -fruited Capparis
petiolaris Kunth, including its xeromorpbic. often
pubescent, narrow-leaved Peruvian ecotype, C.
prisca J. F. Maebride, with which it thoroughly in¬
tergrades; (2) C. coimbrana X. Cornejo & litis, an
allopatrie Bolivian sister species of C. petiolaris,
which mainly differs from tin* former by its oblon-
goid, edible fruits and higher number of lateral
veins (Cornejo & litis, 2005); (3) the Peruvian C.
sprucei Eichler, locally endemic to the arid low¬
lands along the Rio Huallaga from Tarapotd south
to the small village of Cuzco in the province ol San
Martin, this an extreme xerophyte with strongly co¬
riaceous, orbicular to broadly ovate leaves; (4) a
new, highly local endemic from the Rio Nechi
headwaters in Antioquia, northwestern Colombia
(Cornejo & litis, in prep.); (5) the rare local C. cua-
trecasana Dugand from Bogota in the Colombian
Andes; (6) the uncommon C. macrophylla Kunth s.

»3_404. Published on 21 September 2005.

394

Novon

I' igure I. Distribution ol (.< ipparis bonifaziana X. Comejo & litis (solid riots) and C. macrophylla (hollow dots), their
ranges separated by the Andes (the isolated dot ol C. bonifaziana in southern Peru is discussed at end of its specimen
citations).

sir. (Figs. I, 3), a complex species that in one form
or another ranges sporadically along the Andean
arc, mostly in tropical inter- Andean valleys in the
north and western Amazonia to the south in a nar¬
row belt just east ol the Andes, from northern Ve¬
nezuela (inch C. iltisiana T. Kufz) and Colombia
(C. macrophylla s. str.) to Acre in southwestern bra¬

zil (inch C. magnifica Cilg), and is characterized
by often giant, broadly ovate to ovate-lanceolate
leaves and oblongoid fruits; and finally, (7) the new
species C. bonifaziana (Figs. 1. 2) an allopatric,
Ecuadorian segregate of C. macrophylla, the sub¬
ject of this paper.

Capparis macrophylla s.l. has been problematic

Volume 15, Number 3
2005

Cornejo & litis

Studies in Capparaceae XXVI

395

for a long time, partly because of the confusion
caused by the fact that its name had been some¬
times misapplied to tbe widespread but distantly
related C. frondosa Jacquin (C. baducca L.; Du-
gand, 1955: 108; Km'z-Zapata & litis, 1998: 140),

and, steadily since the 1890s, to the common, more
closely related, smaller-flowered and much smaller-
fruited C. osmantha Diels (inch C. guaguaensis
Steyermark) of mostly the tropical Amazonian for¬
ests east of the Andes. This species, in turn, is

396

Novon

more closely related to a Mexican— Mesoamerican
alliance (litis. 2001; Rufz-Zapata <!C litis. 1998)
composed of C. discolor J. I). Smith (inch C. tuer-
kheimii J. I). Smith and C. uniflora Woodson), C.
(fuirigucnsis Stand ley, and C. mollicella Standley

(inch C. lankasterii Standley), and, in Colombia, C.
cuatrecasana Dugand.

The type specimen of Capparis macrophylla
(. Humboldt & Bonpland 1568, P), consisting of
three leaves (cf. Fig. 3) and one flower, is not very

Volume 15, Number 3
2005

Cornejo & litis

Studies in Capparaceae XXVI

397

revealing, since these resemble leaves and (lowers
of some of the other taxa listed above, while tbc
original description, based in part on Bonpland's
field notes (Kunth. 1321: 91), is more helpful. In
any ease, since material of C. macrophylld from
Colombia is often incomplete, the need for more
ample collections, especially from the southwest of
the country, is critical. Unfortunately, the fully ma¬
ture, soft fruits of most of these species, but espe-
cially of C. macrophylld. are uncommonly collected
by botanists, attractive as they are to animals and
to humans who also like to eat them, and they are
furthermore, much as bananas, difficult to dry and
preserve. Finally, fruit size varies enormously with¬
in most species, both clinally, depending on the
regional or local climate, and individually, depend¬
ing on the vigor of the plant (Rufz-Zapata X litis.
1998; litis. 2001).

Nevertheless, in preparing a treatment of Cap¬
paraceae for the Flora of Ecuador, and utilizing the
large number of splendid specimens recently col¬
lected there, our study of all Capparis macrophylld
collections made it abundantly clear that these
could be separated on leaf shape alone into two
allopatrie taxa divisible by the Andes (figs. 2. 3):
narrowly oblong-lanceolate to oblong-elliptic, and
acute to obtuse, from the relatively drier western
Pacific lowlands, and broadly elliptic to ovate, and
acuminate to short-acuminate, from the much wet¬
ter Amazonian lowlands just east of the Andes, with
the latter populations matching closely in their leaf-
outlines the leaves of the C. macrophylld type spec¬
imen from the Colombian Andes (Fig. 3). Several
other characters of flower and fruit were eventually
discovered, which correlated well with the season¬
ally dry to moist forest ecology of western Feuador.

ki.v to Capparis bo\u tz/i vi v\n C. \iu:Hoi‘inu.\

la. Kpisepalous glands a— 8 mm wide, showy and ex¬

posed by divergent to reflexed sepals; stamens
35 to 71; anthers 1.5— 5.3 mm; ovary 5—8 mm;
petals 19—29 mm, divergent to sabered at an-
thesis; fruit indehiscent. Leaf blades oblong-el¬
liptic to oblong-lanceolate, rarely lanceolate or
oblanceolate; endemic (except for one isolated
collection from central Peru) to the southern half
of Ecuador on the western side of the Andes, in
seasonally drv to moist or gariia (fog) forests,
from the lowlands to 850 rn . C. bonifaziana

lb. Kpisepalous glands 2. 5—4 mm wide, inconspicu¬
ous and hidden bv the divergent to ascending se¬
pals; stamens 80 to 130; anthers 6-7 mm; ovary
8-1 1 mm; petals 27—10 mm, reflexed at anthesis;
fruit tardily dehiscent by 3 to 4, thin, flexible co¬
riaceous exoearp segments. Leaf blades widely el¬
liptic to ovate-elliptic or sometimes suborbieular;
in moist to wet lorests of the tropical valleys to
the submontane zone of the Venezuelan and Col¬

ombian Andes, and in the low-elevation, wet, ev¬
ergreen, tropical Amazonian rain forests, from
southern Colombia and eastern Ecuador and Peru
to Amazonas and Acre states of western Brazil,
just east of the Andes . C. macrophylld

With Capparis macrophylld s. str., now more nar¬
rowly defined, the newly segregated Ecuadorian en¬
demic is in need of a name. Because its best and
most complete specimens were eo-collected by Dra.
Carmen Bonifaz (b. 1956), Universidad de ( utaya-
({ii i I Herbarium director and a long-time sophisti¬
cated student of the western Ecuadorian flora, we
are most pleased to name this species in her honor.

I. Capparis bonifaziana X. Cornejo <X litis, sp.
nov. TYPE: Ecuador. Los Rios Prov.: Heda.
Clementina, Puerta Negra. en “Tecal contig-
uo. bosque humedo tropical, 79°21 \\. 1°40'S.
80 m. 9 Oct. 2003 (11. IV), X. Cornejo <C C.
Bonifaz 7BI2 (holotype, GUAY; isotvpes,
A A U . RM. COL, K. LOJA, MO, NY. OCA,
QCNE [2], US [2], USM. WIS [7]). Figures I.
2. 4. 5.

Harr species Cappari macrophyllae af finis, a qua differ!
foliis plerumque oblongo-ellipticis vel oblongo-lanceolatis;
petalis 19-29 X 9-17 mm, erectis vel divergentibus;
staminibus 35—71: glandulis nectariis crassis, 5—8 mm la-
tis; antheris et ovariis minoribus.

Treelet or tree 3—25 m tall, to 25 cm or more
l)BH. unbuttressed, glabrous throughout; new
branches with orbicular to linear lentieels 0.2—4
mm; stipules narrowly triangular. 1-1.8 X 0.5— 0.8
mm. Leaves evergreen, spirally disposed, ea. 5 to
10 spread out near the end of current year's and/
or last year’s growth, with petioles and leal blades
of different lengths and sizes intermixed; petioles
0.3— 1 5( — 24) cm long; pulvinus (2— )4— 12 mm. dark
brown (dry); leaf blades ± coriaceous, oblong-el¬
liptic to oblong-lanceolate, rarely lanceolate or ob¬
lanceolate. 6-45 X 3-15(-20) cm. the apex acu¬
minate to acute or rounded and apieulate, the base
rounded or obtuse to euneate, dark green above,
pale green beneath, with (7— )9— 12(— 1 5) prominent
main lateral veins on each side of the midrib: leaf
margin not retuse at insertion above the pulvinus,
and only rarely slightly deeurrent (Fig. 2). Inflores¬
cences terminal or subterminal, erect, ebracteate
racemes; peduncles ± naked, up to 21 cm long and
8 mm thick, the terminal flower-bearing portion of
the axis ea. 3—12 cm long, 9- to 25-flowered; ped¬
icels (2— )3 — 4( — 5) cm. each subtended by a minute,
caducous, triangular to linear bract and two persis¬
tent minute triangular stipules; calyx 4-parted. to
15 nun across, lip to lip: sepals ascending to some¬
what reflexed at anthesis, broadly deltoid to se-

398

Novon

figure 4. Capparis bonifaziana. — A. Carmen Hon i fa/. holding an immature fruiting branch of the type specimen. —
If. 1* lowering branch, with buds and post-anthesis flowers wit li filaments abscissed, showing the strongly ascendent
petals, large glands and ovaries on elongated gynophores. — C. The same branch as in B, enlarged. — I). Flower bud
just preceding anthesis.

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Studies in Capparaceae XXVI

399

Figure 5. Capparis bonifaziana. — A. Nearly mature fruit with characteristic longitudinal stripes, reflecting the inner,
parietal placentae. — B. Fruit in longitudinal section, showing the many seeds imbedded in a white, sweet, edible pulp.
— C. Branch with two fruits and subtending leaf. A— C: type collection.

miorbicular, 4—6 X 5—9 mm, white to greenish
white with white-hyaline, shallowly erose-ciliolate
margins, each subtending and bearing on top of
their inside base a conspicuous, relatively large (ca.

2—3 X 5—8 mm), white to pink or purple (when
fresh, but drying black), episepalous gland that is
fleshy, rounded, fat and cushion-shaped; flower
buds just preceding anthesis obovoid, 15—25 X 18—

400

Novon

20 nun; petals still and erect to ascending or di¬
vergent, somewhat lleshy, forming a teacup-shaped
corolla at anthesis, broadly ovate to elliptic, 19—29
X 9-17 mm. the apex broadly rounded and cueul-
late, the base subsessile by a minute peg inserted
between, and inside of. the glands, greenish white
without, cream to white within, the margin ± entire,
minutely and irregularly erose-dentieulate; stamens
35 to 71, filaments 5.5—8 cm long, anthers 4.5— 5.3
mm, dorsifixed in the basal third; gynophore 6—8
cm, white; ovary 5—8 X ea. 2—3 mm, green, oblon-
goid to subeylindrieal. sometimes longitudinally
striate; stigma truncate to hemispherical, sessile.
Fruits solitary or up to 6, clustered at the end of
branches, pendent indehiscent and baccate, fleshy
within, eylindric or ohlongoid, rarely ovoid, straight
to somewhat curved (banana-shaped), 7—15 X 2—4
cm, when immature the exoearp green with four
dark green stripes (“resembling a squash fruit,"
Steyermark 52766], in immature herbarium mate¬
rial dark brown or purple-spotted with four black
longitudinal stripes, at maturity yellow to orange,
with 2 major and 2 minor longitudinal, grooved
markings [Fig. 5; “bright yellow with faint red lon¬
gitudinal lines," Neill 11446], eventually turning
soft, or thin and coriaceous. Idled with a white pulp;
gynophore 7-1 I X 0.3— 0.6 cm, bright yellow; ped¬
icels 3— 5(— 6) cm; seeds ea. 25 to 60 per fruit, each
surrounded by a white aril, irregularly beveled,
somewhat flattened. 10-14 X 7-8 mm (but see be¬
low); testa brown or black, the cleft invagination
narrow and very short (ea. 2 mm); embryo white,
the cotyledons convoluted one into the other and
around the radicle, with a thin layer of endosperm
visible (in cross section) on the rounded back of
the seed.

Habitat and distribution. Lowlands to the pre-
montane, fog-enshrouded mountains of western Ec¬
uador, from Esmeraldas south to Azuay, from dry
to moist tropical and (rarely wet) premontane for¬
ests. often in secondary vegetation (usually all that
is now lelt in western Ecuador), from near sea level
(50 m) to 850 m; flowering and fruiting throughout
the year.

I /teal names. Barniz (Spanish), Qnelal et al.
472 (Esmeraldas); Cacho de Venado (Spanish), Cor¬
nejo & Bonifaz 5667 (Los Bios); Guineo de montafta
(Valverde et al., 1991); Maduro (Spanish), Cornejo
N Bonifaz 5884 (Maimin'), Cornejo & Bonifaz I BOB
(Guavas, Bonifaz cK Cornejo, 2004).

Uses. Fruits edible, Cornejo & Bonifaz 5884,
4885, 5667. 7765; Bonifaz & Cornejo 50B5 (Boni¬

faz & Cornejo, 2004). Cultivated as living fences,
Alvarez & Tirado 1545.

Biological interactions. The flowers, open from
early evening until early next morning, are proba¬
bly bat-pollinated, but this has as yet not been ver¬
ified. During the day, flowers are visited by black
bees (Cornejo & Bonifaz 5507) and at least five
species of ants, each observed and collected on a
different tree at the type locality ( Cornejo & Bonifaz
7BI2).

Discussion. The vegetative differences between
Capparis bonifaziana (f ig. 2) and C. macrophylla
(Fig. 3) may be subtle. In addition to the overall
leaf shape. C. bonifaziana has the blade margins
inserted generally at right angles above the pulvi-
nus, and only rarely dee ur rent onto the pulvinus,
whereas C. macrophylla has leaves that are more
often rounded or suheordate at the base, but then
more abruptly decurrent onto the pulvinus.

Ecology. While label data tell us that C. boni¬
faziana grows in a wide range of forest types, from
dry (but not very dry) to moist or even wet. asso¬
ciated species have never been mentioned. Here we
now report two woody Capparaceae collected to¬
gether with C. bonifaziana ( Cornejo & Bonifaz
7115, in sehed.) in remnant moist forests near Cho-
ue: Capparis ecuadorica litis (litis. 1978), a locally
not uncommon Ecuadorian endemic shrub or tree-
let; and. in addition, wonder of wonders, tin* showy
Steriphoma urbani Eggers ( Cornejo & Bonifaz 7550.
in sched.), a super-rare liana endemic to western
Ecuador, here now at its only fourth known station,
and the only white-flowered species in this small,
highly specialized, otherwise orange- to red-flow¬
ered, hummingbird-pollinated genus. Its pollinating
bats are yet to be discovered.

Detailed descriptions of the vegetation associat¬
ed with Capparis bonifaziana may be found in Flo¬
ra de! Bosque de Garua de la Comuna l Anna Alta.
Brovincia del Guavas, Ecuador (Bonifaz Cornejo,
2004), located in the Chongbn-Colonehe mountains
northeast of Guayaquil. Here, the belt of dense,
seasonally moist to wet, low-elevation (400—600 m)
cloud forest is. like the hot super-dry coastal forests
and deserts, the product of a peculiar climatic phe¬
nomenon characteristic of the west coast of Peru
and Ecuador (as it is of the west coasts of South
Africa and Australia), namely oceanic fog. Locally
known as the garua. and drifting inland from the
frigid Pacific Ocean to the west, this fog of fine
water droplets is the result of the chilling effects of
the upwelled, cold coastal waters of the Antarctic
I luml joldt (or Peru) Current on the warm, moisture-

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Cornejo & litis

Studies in Capparaceae XXVI

401

laden tropical westerlies. With its 20—30 in tall
trees “milking” the clouds, such wet forests are
characterized by abundant epiphytes, especially
Araceae (Anthurium Schott, Philodendron Schott),
Orehidaceae, Bromeliaceae, pteridophytes and
mosses, and an occasional treelet of C. bonifaziana.

Although widespread, Capparis bonifaziana is
relatively uncommon, the often solitary treelet or
rare tree occurring in open, seasonally dry (hut not
very dry) to moist or even wet forests, and most
often in secondary vegetation. I he only exception
to this rule was found at the type locality, where
some 30 or more treelets were scattered in the un¬
derstory of a teak or “teca” ( Tectona grandis L. f.)
plantation (a lead in Spanish). This is perhaps not
surprising, since most collections were made in dis¬
turbed forests or fields or intervening arroyo rem¬
nants. In fact, such treelets do well when planted
as living fences [Alvarez & Tirado 1545).

Even though Los Rios, wherein lies the type lo¬
cality'. is the province with both the most disturbed
vegetation in all of Ecuador, with pastures and
fields abutting giant plantations of bananas, and the
most thoroughly studied flora of the country, with
its Flora of the Rio Palenque Science Center (Dod¬
son & Gentry, 1978) and La Flora de Jauneche
(Dodson et al., 1985), neither of which report a
Capparis equivalent to C. bonifaziana , nevertheless
its flora continues to yield botanical novelties. In
these treasured forest remnants, much field explo¬
ration remains to be done before all the wild will
be gone.

Finally, on the basis of geography alone, Cap¬
paris bonifaziana must he the remarkable species
quoted below by the legendary, now much lamented
taxonomist-ecologist, Alwyn II. Gentry (1945—
1993), one of the second author’s former students,
who at age 48. together with his friend and fellow
bioexplorer, the famed ornithologist Theodore A.
Parker III. met a most untimely death on the gari'm-
enshrouded slopes of the Chongon-Colonche moun¬
tains in a tragic airplane accident (Miller, 1996).
In their thorough, book-length survey of Conser¬
vation International's Rapid Assesment Program
(RAP) on the Status of Forest Remnants of the Cor¬
dillera de la Costa, Gentry's (1992: 38) discussion
of the "Dry Forest" highlights, first, the remarkable
widespread abundance and diversity of Cappara¬
ceae in coastal Ecuador, and second, the discovery
of an astonishingly 25 m tall tree Capparis that he
had never seen before. This is of interest, for de¬
spite the fact that some species of the C. macro-
phylla alliance are widespread and yet not com¬
monly collected, and Gentry had as wide a Holistic
experience in northwestern South America (Gentry,

1993) as almost any botanist who ever worked
there, this was, he thought, the first time that he
ever encountered that Capparis, this surely a re¬
flection in part of the enormous, complex, and lo¬
calized biodiversity so characteristic of tropical flo¬
ras. While Gentry evidently had both leaves and
the "ripe banana-sized, striped fruits” in his hand,
he did not collect that tree, strangely enough, per¬
haps awed by the difficulty of cutting it down, the
magnificence of its stature, or its presence in a pro¬
tected area, i.e., in a National Park. Nevertheless,
apparently on the morning of that same day ( 1 8 Jan.
1991 ). lie did collect (but did not recognize) a small
treelet in both flower and fruit of this very same
species [Gentry & Josse 72386 ) southwest of the
small village of Guale on the Rio Ayampe, and only
ea. 3 km from that striking tall tree at Estero Manta
Blanca:

“Although there are some local peculiarities [in
Parque Nacional Machahlla]. the dominant families
in our dry forest sample from Estero Perro Muerto
are Leguminosae and Bignoniaceae (7 spp. each),
just as they are in essentially all Neotropical dry
forest. A noteworthy family in this region is Gap-
paridaceae, which mostly occurs in the very driest
forest and may be more dominant here than any¬
where else in the Neotropics. One striking Capparis
(of which we found only a single tree) al Estero
Manta Blanca is a large tree ca. 25 m tall, among
the largest for this genus. Another interesting Cap¬
paris discovery is that the sometimes confused C.
heterophylla and C. ecuadorica show strong ecolog¬
ical differentiation in areas where they are sym-
patric [this remains to be verified, for these two
species often do occur together]. 1 have not previ¬
ously seen the large-leaved Capparis with edible
banana-sized, striped fruits that we found al Mach-
alilla and it may be an undescribed species (Gen¬
try, 1992: 38).

How right he was!

Paratypes. ECUADOR. Esnieraldas: Chaflu, bosque
humedo tropical, X. Cornejo & C. Bonifaz 5407 (OIL
GUAY, k, WIS); Vi of San Mateo, lies. For. Jardin Trop¬
ical. Univ. Tec. Luis Vargas Torres, A. Gentry A A. Lajones
73056 (MO, QCNE, WIS photocopy); Paraiso de Papaga-
yos. Fund. Rescate de Animales Silvestres, km 2 v ia Ls-
meraldas-Quininde. ./. Clark. /.. Chatrou, P. Maas A (..
Repet ur 3790 (MO. QCNE. WIS [2]); Ancliayacu, Floy Al¬
faro, Mayronga, T. I). Pennington. R. Calero, M. Guevara,
L. Zambrano. G. Rodriguez A' /,. Veloz 14552 (k, W IS |2|):
Parroquia Alto Tarnbo, l inca del Sr. Calderon, a 2 km del
sector HI Cristal, en earretera, C. Quelal 472 (MO. Vi IS
photocopy); Rilsa Biol. Stat .. Mache mtns., 35 km W of
Quininde 5 km W of Santa Isabel, premontane wet forest.
Monkey Bone trail. ./. Clark A B. Adnepos 76 (MO. QCNE,
WIS [2]), ./. Clark. /.. Chatrou. C. Repetur A P. Maas 3B05
(MO. QCNE, WIS |2|). Manabi: Rfo Plata, ca. 50 km

402

Novon

NNK dt* Chone. reinanente de bosque litimedo tropical, X.
Cornejo & C. Bonifaz 71 15 (GUAY, W IS); km 12 vfa Cho¬
ne— Sto. Domingo, I km atrds de la Universidad Catblica,
host | ue humedo tropical, X. Cornejo X C. Bonifaz 7350
(GUAY, QCNE,, \\ IS |2|); 31 kin N of Pedernales along
new coastal hwy., ea. 2 km S of Rfo Maehe, I). Neill X
QCNE botany interns 1 1 BOB (QCNE), \\ IS photocopy); 30
km S of Pedernales on coastal highway, primary forest on
hills, 2 km from seashore, lb Neill X QCNE botany interns
11293 (GUAY, MO, QCNE, WIS); 34 km N of Pedernales
along new coastal hwy., near village of Chindul, I). Neill
X QCNE botany interns 11446 (GUAY, MO, QCNE, WIS);
Parqne Nae. Machalilla, San Sebastian, X. Cornejo X C.
Bonifaz 3004 (GUAY, K, WIS), C. Hernandez 260 (QCA);
Sector entre la Bola de Oro y La Moeora, C. Hernandez
191 (QCA). Guayas: (iuale, bosque seeo tropical, X. Cor¬
nejo X C. Bonifaz 1300 (GUAY, WIS); valley of Rfo Av-
ampe, border w/Manabf, SW of Guale, A. Gentry & C.
Josse 72306 (MO. QCNE, WIS); Cordillera Chongon-Co-
lonche, comuna Oldn, X. Cornejo X C. Bonifaz 2021
((ill AY, WIS); Rfo California, bosque seco, C. Bonifaz &
\. Cornejo 3005 ((>UAY, W IS). Los Rios; Represa Daule
Peripa, El Barro, Boca de Pescadillo, C. Bonifaz 516
(til AY, MO. WIS); La Puntilla, X. Cornejo A' C. Bonifaz
5667 (GUAY, WIS); llcda. Clementina, en tecal, bosque
humedo tropical, X. Cornejo A- C. Bonifaz 4005 (GUAY,
K, WIS), Y. Cornejo & C. Bonifaz 7793 (( ill AY , WIS);
eerro Samama, near Puerta Negra, mainly disturbed sec¬
ondary forest (very common), B. Stahl X X. Cornejo 5020
(GB, GUAY, W IS |2|); above Rfo Mombe, ca. 38 km NE
of Babahoyo, B. Stahl X J. Knudsen 1179 (GB. W IS pho¬
tocopy). Caiiar/Azuuy: llcda. Yubay, at Sanaguin, on S
side of Rfo Patul, ./. Steyerinark 52706 (NY); comuna de
Manta Real, al sur de la carretera Ea Troncal— Zliud, cam-
ino entre Xlmcay y el Rfo Patul. margen izquierda del Rfo
Patul, bosque humedo premontano, A. Alvarez X M. Tirado
1545 (MO, QCNE, W IS). Azuay: 3 km SE of Ea Troncal
on road to Zlmd, beginning of track to Zhucay, G. P. Lewis
X P. hizano 2097 (QCA, QCNE,, W IS); Ea Conferencia y
Santa Cruz, Parroquia Molleturo, /,. Ortiz 132 (QCA.
QCNE). PERU. San IVfartui-Loreto: high area above Rfo
Aguaytia, W bank 10 km below Aguaytia, small tree with
paw-paw | Asimina triloba , Annonaceae, an edible-fruited
common treelet of eastern North America]-like fruits, 4
July 1959 (fr), M. Mathias X I). Taylor 3500 (MO, WIS
photocopy).

I his solitary collection from southern Amazonian
Pern, at the far southern end of San Martin prov¬
ince and close to lingo Marfa, has large elliptic
leaves and fruits exactly like those of Capparis bon-
ijaziana, and may represent that species otherwise
endemic to western Ecuador. Since this station is
disjunct by fully (>()() km and close to the stations
of C. macrophylla [Froehner 407), we included this
collection here somewhat reluctantly. Of course,
such disjunction may be attributed to avian long¬
distance dispersal, a situation not unknown consid¬
ering that these fleshy fruits evolved for animal con¬
sumption and dispersal. On die other hand, with
the fruits of this species eaten by people even now,
and undoubtedly by “indfgenas” some several thou¬
sands of years ago, at the time of the Incas and

before, there is always the possibility that some
horticulturally-inclined persons took seeds or cut¬
tings and transported them to their new home to
enjoy these tasty fruits yet another day. Though the
great distance and different climatic regime would
seem to preclude such a scenario, as would an un¬
likely long-distance avian dispersal, we surely can¬
not discount these possibilities completely.

2. Capparis mucrophylla Kunth, in I IRK. Nov.
Gen. Sp. 5: 91. 1821. Capparidastrum macro-
phyllum (Kunth) Hutchinson, Gen. FI. PI. 2:
310. 1967. TY PE: Colombia. |Bolfvarj: “Cres-
cit ad Rfo Viejo in ripa Magdalenae lluminis.
inter Badillas [Badillo] et Bojorque | Bohdr-
quez]. May 1802 (fl), Humboldt X Bonpland
I SOB (holotype |one leaf and one flower], P |F
neg. 34627, WIS photo]; isotype [two leaves],
P, WIS photocopy). Figures I, 3.

According to Dugand (1941: 40; 1955: 108), a
pioneer scholar of tropical American Capparis , “El
sitio llamado ‘Rfo Viejo por Humboldt y Bonpland
se Italia cerca de San Pablo, abajo de Puerto \\ ilch-
es, a 7°30' de latitud norte. Badillo se encuentra a
7°57', y Bohdrquez a 7° 18' preeisamente f rente a
Puerto Wilches.”

Capparis iltisiana T. Rufz. Ernslia 3(>: 1-2. 1986. TY I’E:
Venezuela. Edo. falcon; l)tto. Acosta, Man. Jacura,
Cerro Ea Mina de Riecito, bosque submontane hu¬
medo, selva siempreverde con afloramientos mooses.
500 m, T. Ru(z X E Ronddn 3709 (holotype, MY;
isotypes, E |2], MY, VEN, W IS).

Shrub “with scandent branches” (“ Frutex ramis
scandentibus. . | Kunth, 1821: 91], this character

probably based on a mistaken perception) to tree
15 m or more high and 15 cm or more DBH, with
terminal branches glabrous, the lenlicels promi¬
nent, orbicular to linear, 0.2—2 mm (of. Dugand.
1955: 109), stipules ca. 1.7 mm long, narrowly tri¬
angular. Leaves evergreen, spiral, with petioles and
blades of different lengths and sizes clustered near
the ends of the branches; petioles (4— )8-30 X 0.1-
0.3 cm; pulvini 3—24 mm long, dark brown, usually
glabrous; leaf blades widely elliptic to ovate-elliptic
or sometimes suborbicular, (2— )10— 59 X (5— )10— 34
cm, with apex acute to usually abruptly acuminate
or rounded and apiculate, at base broadly obtuse
to rounded and frequently shortly decurrent onto
the petiole, (sub)eoriaceous, dark green above with
midvein and secondaries impressed to prominent,
glabrous, paler green beneath with all veins prom¬
inent. with 9 to l3(to 15) main lateral veins on each
side of the midrib. Inflorescences erect, terminal,
ebracteate racemes, the stout peduncles 7—24 cm

Volume 15, Number 3
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Cornejo & litis

Studies in Capparaceae XXVI

403

long, with the flower-bearing portion 6—18 cm long,
corymbiform, with ca. 7 to 14 flowers, each sub¬
tended by a minute, subulate, caducous bract ea.
1.5 X 0.7 min, and laterally by 2 persistent minute
triangular stipules, glandular with short (less than
0.1 mm), dense, colorless, uniseriate trichomes, the
stout pedicels 2— 6(— 10) cm. Flowers greenish or
white, with flower buds just before anthesis ovoid
to oblongoid, 2.5—4 cm long; calyx 4-parted, in two
inconspicuously imbricate series; sepals broadly
deltoid, ea. 5—1 1 X 6— 10(— 15) mm, erect to some¬
what divergent at anthesis, with rounded apex,
greenish, with shallowly and irregularly denticulate
to erose glandular margins, externally glabrous,
subtending and hiding a ring of four inconspicuous
episepalous, fleshy, glabrous glands ea. 1 X 2.5— 4
mm; petals white, elliptic, 27—40 X 16— 18(— 25)
mm, inserted between and inside of the glands, di¬
vergent to reflexed at anthesis, white or cream lo
greenish without, fleshy, longitudinally inconspic¬
uously nerved, with margins entire to denticulate;
stamens 80 to 130, filaments 5.5—10 cm, white, the
anthers 6—7 mm, linear, dorsilixed in basal one
third; ovary 8—11 X 2 mm, oblong to elliptic, gla¬
brous, 3- to 4-angled, unilocular but appearing
falsely 3- to 4-locular by the intrusion of 3 to 4
parietal placentae (Ruiz-Zapata, 1986), the stigma
hemispherical, sessile; gynophore 7—10 cm long.
Fruits pendent, fleshy, baccate [bacca corticata ;
Jackson, 1928 (reprinted 1949)], ellipsoid to oblon¬
goid, (4-)6-14 X 2-4(-6) cm, externally yellow,
glabrous, with (3 to)4 external, longitudinal, vas¬
cular markings or grooves (2 strong, 2 weak, re¬
flecting the 4 parietal placentae inside), at maturity
often tardily partway dehiscent from the lip, with
the 3 to 4 thin, flexible coriaceous exocarp seg¬
ments (i.e., valves) separating and recurving at full
maturity (Ruiz-Zapata, 1985; Schunke & Graham
S 14806) to expose the fleshy interior; gynophore in
fruit (3— )8— 14 X 0.2— 0.4(— 0.6) cm; pedicels 4— 7(—
I 1) cm long; seeds ca. 25 to 65 per fruit, ca. 10—
12 X 6—9 mm, immersed in a white to ivory fleshy
pulp/aril, i.e., “Frutos. . .dehiscentes en 4 valvas,
mostrando las semillas embebidas en masa cremosa
ile naturaleza arilar” (Ruiz-Zapata, 1985; 104—
III); embryo white to cream, the cotelydons folded
one into the other and around the radicle.

Habitat and distribution. In moist to wet, ev¬
ergreen lowland forests of tropical inter-Andean
valleys lo premontane slopes of the northern Andes
ol Venezuela (150—1000 m) and Colombia (50—400
m), and in western Amazonia in a narrow zone on
the wet, eastern side ol the Andes, from southern
Colombia (30 m) through eastern Ecuador (200—

300 m) and Peru (200 m) to Amazonas and Acre
states (100—300 m) of southwestern Brazil; (lower¬
ing and fruiting more or less throughout the year.

Local name. Ecuador: “Bus su bara” (Siona;
fide Reinaldo Lucitande), Vickers 82 (Sucumbios).

Discussion. The location of the ty pe was spec¬
ified by Dugand (1941; 1955: 108-110). who in
addition gave a detailed Latin description of the two
Romero-Cast aiieda (2054, 2055) collections, which
we have not seen, yet which were not only collected
close to the type locality but also according to their
detailed morphological description undoubtedly re¬
fer to Capparis macrophylla. The taxonomic prob¬
lems with C. macrophylla were partially discussed
in our introduction. The type description was based
in part on Bonpland’s field notes, as mentioned, and
only three leaves and one dissected flower survive
in the Paris Herbarium. Its claim that it came from
a “shrub with scandent branches" seems to be an
error. Careful descriptions of C. macrophylla are
found in Dugand (1955) and Ruiz-Zapata (1986).
A beautiful illustration is found among the unpub¬
lished Mutis plates (No. 9 ol Capparis) in the Ma¬
drid Herbarium and another with Rufz-Zapata’s ex¬
ceptionally careful description of C. iltisiana
(Ruiz-Zapata, 1986).

Except for Colombia (and even here), this is a
rarely collected species, widely dispersed in highly
isolated populations, some of which perhaps will
eventually deserve taxonomic recognition once
more material is available.

Additional specimens examined. VENEZUELA. Edo.
Falcon: Dtto. Silva, hacienda El Carmen, 20 km al 0 de
Tucacas, A. Cardoso & R. Wingfield 188 (CORO); Dtto.
Acosta, Cerro de La Mina, c. Riecito, Mini. Jacura, bosque
submontano humedo, L. Ruiz 641 (MER, MY); ca. 2 km
al ENE del P. IN. Cueva de la quebrada El Toro, cerca del
origen del Rfo Paray, R. Wingfield it' R. Smith 8808
(CORO); Dtto. Eederacion, carr. La Ta/.a a la est. del P.
N. Cueva de la quebrada El Toro, T. Ruiz &' M. Ruiz 4158
(MY); Par. Nac. La Cueva de la quebrada El Toro, a la
entrada de la cueva, T. Ruiz , R. Parra & A'. Marino 4058
(MY). COLOMBIA. Antioquia: Segovia, /’.. Renteria 2805
(HUA, JAUM, NY, Vi IS photocopy); Santa Catalina, G.
Basse 8674 (LE, MO. WIS photocopy). Cundinamarca:
Tocayma [Tocaima], M. Goudot s.n. [9?] (P, WIS photo¬
copy). Narino: Costa del Pacffico, corr. de Herrera, cuen-
ca del Rfo Mira, c. Candelillas, Idrovo <£ II. We her
1420e (COL, WIS). Putumayo: Rio San Miguel o Suc-
umbfos, Santa Rosa y los alrededores, R. Schultes 3618
(GH, WIS photocopy). Santander: 10 leguas al S.E. de
Barranca Bermeja, a 3 km de la margen izquierda del no
Opon, terrenos bajos inundables, Romero-Castaneda
5054, 5055 (COL, HUA not seen, fide Dugand, 1955).
ECUADOR. Sucumbios: Rfo Aguarieo, Shushufindi, W.
Vickers 82 (E, W IS photocopy); San Pablo de Los Secoyas,
path W-SW, L. Brandbyge, E. Asanza , /,. Werling, S. leth-
Nissen & F. Larrea 33310 (AAU. OCA. WIS); Est. exp.

404

Novon

Payamino INIAP. Reserva F'lorfstica Kl Chunchu, ./■ Jar-
amilh) 8363 (QCA). PKRU. Amazonas: Huambisa, valle
del Rfo Santiago, ea. 65 kin IN de Pinglo, quebrada Ca-
terpiza, 2—3 km atrds de la coni. Caterpiza, V. Huashikat
46 4, 1283 (MO, WIS). .S. Tunqui 91 (MO, W IS photocopy).
Ucayali: Prov. Coronel Portillo, Bosque Nac. Alexander
von llumholdt. Pucallpa— Tingo Marfa rd, carr. de extrac-
eidn. Ch. Froehner 407 (MO. WIS photocopy); Prov. Pu¬
rus, Rfo la Novia, margen derecha del caserfo de la com.
San Jose, ./. Sehunke A' ./. Graham SI 4806 (WIS [3]).
Mud re de Dios: Prov. Mann. Par. Nac. Mann. Rfo Manu,
Rfo Cumerjali, II. Foster & 8. d'Achille 12002 (F. WIS
1 2 1). BRAZIL Acre: Rfo Acre, Seringal Auristella, F. Fie
9409 (G, k. MG. W IS fragm.); B[au]m, im Seringal S.
Francisco, F. Fie 9410 (G, k, MG, Vi IS photo); near
mouth of Rfo Macauhan (tributary of Rfo Vaco), on terra
firme, 8. Krukoff 3788 (A. GH, NY. W IS).

Acknowledgments. We thank first of all Carmen
Ronifaz, fellow botanist and to the first author long¬
time field companion in the forests of western Ec¬
uador, who helped initiate the taxonomic concept
of Capparis bonifaziana as a separate species; Al¬
icia Lourteig, the former herbarium curator of the
Museum National d’Histoire Naturelle in Paris,
France, for photocopies of the type of C. macro-
jihylla ; Rosa Rankin for helpful comments; Ted
Cochrane for the very careful reading of the man¬
uscript; Lupita Sanchez for whetting the text to per¬
fection; and finally, the ever helpful Kandis Elliot,
artist of the UW Botany Department, for the expert
and inspired illustrations.

Literature Cited

Bonifaz, G. A X. Cornejo. 2004. Flora del Bosque de Gar-
ua de la Coinuna Lorna Alta, Provineia del Guayas, Ec¬
uador. Universidad de Guayaquil/Missouri Botanical
Garden/Fundacion GAIA. Guayaquil, Ecuador.

Cornejo, X. A H. H. Illis. 2005. Studies in the Cappara-
eeae XXIII: Capparis coimbrana, a new species from
Bolivia. Brittonia 57: 53—59.

Dodson. C. II. A A. H. Gentry. 197B. Flora of the Rfo
Palenque Science Center, Eos Rios Province, Ecuador.
Selhvana 4: 1-628.

- . - A F. M. Valverde. 1985. La Flora dejau-

neche, Los Rios, Ecuador. Banco Central del Ecuador,
Quito, Ecuador.

Dugand, A. 1941. FI geuero Capparis en Colombia. Cal-
dasia 2: 29-54.

- . 1955. Capparis macrophylla Kunth. In: A. Du¬
gand. Noticias Botanicas Colombianas XL Caldasia
VI 1(32): 108-1 10.

Fielder, A. W. 1865. Capparideae. In: C. F. P. von Marlins
(editor), Flora Brasiliensis 13(1): 237—292, tab. 51—65.

Gentry, A. H. 1992. Dry forest. In: T. A. Parker III A J.
L. Carr (editors), Status of the Forest Remnants in the
Cordillera de la Costa and Adjacent Areas of South¬
western Ecuador, Rapid Assessment Program (RAP).
Conservation International. Washington. D.C.

- . 1993. Field Guide to the Families and Genera of

Woody Plants of Northwest South America (Colombia,
Ecuador, Peru), with supplementary notes on herba¬
ceous taxa. Illustrations by Rodolfo Vdsquez. Conser¬
vation International, Washington, D.C. [Capparidaeeae,
pp. 315-319.)

litis, II. 11. 1978. Studies in Capparidaeeae XIV. Capparis
ecuadorica, a new species. Selbyana 2: 303—307.

- . 2001. Capparaceae. In: W. I). Stevens, C. Ulloa

Ulloa, A. Pool X (). M. Montiel (editors). Flora de Nic¬
aragua. Monogr. Syst. Rot. Misssouri But. Garth 85(1):
566—584.

Jackson, B. I). 1928 [reprint 1949]. A Glossary of Botan¬
ical Terms. Duckworth, London.

kunth, C. S. 1821. Capparaceae. In: F. W. II. A. von
Humboldt, A. J. A. Bonpland A C. S. kunth. Nova
Genera et Species Plantarum [quarto ed.], 5. Paris.

Miller. J. S. (editor). 1996. Alwyn Howard Gentry 1945—
1993: A tribute. Ann. Missouri But. Garth 83: 433—460.

Rufz-Zapata, T. 1985. Estudio Taxondmieo de la Eamilia
Capparidaeeae en el Estado Falcon: Contribucion al
Conocimiento de la Flora del Estado Falcon. Trabajo
presentado para optar a la categorfa tie Profesor Asis-
tente. Departamento tie Botaniea, Facultad de Agro-
nonua, Universidad Central de Venezuela, Maracay.

- . 1986. Notas sobre Capparidaeeae tie Venezuela

[Capparis iltisiana sp. nov.]. Ernstia 36:1—4.

- X II. II. Illis. 1998. Capparaceae. Pp. 132-157

in ,1. Steyermark, P. E. Berry A B. k. Holst (editors).
Flora of the Venezuelan Guayana, Vol. 4. Missouri Bo¬
tanical Garden Press, St. Louis.

Valverde, F'., G. Rodriguez X C. Garcia. 1991. Estado ac¬
tual de la vegetacion natural tie la cordillera Chougdn-
Colonche. Instituto de Investigaciones tie Recursos Na-
turales, F acultad de Ciencias Naturales, Universidad de
Guayaquil. Ecuador.

Eysenhardtia officinalis (Leguminosae, Papilionoideae),
una Especie Nueva de Mexico

Ramiro Cruz Duran

Herbario de la Facultad de Ciencias (FCME), UNAM, Apdo. Postal 70-399,
04510 Mexico, P.F. (‘dr@minervaux2.fciencias.unani.mx

Mario Sousa S.

Herbario Nacional (MEXU), Institute de Biologia, UNAM, Apdo. Postal 70-367,
04510 Mexico, D.F. sousa@scrvidor.unam. rax

Rksumen. Se describe e ilustra Eysenhardtia of¬
ficinalis (Leguminosae, Papilionoideae), una espe¬
cie nueva de Mexico. Eysenhardtia officinalis guar-
da similitud con E. polystachya (Ortega) Sargent v
E. texana Scheele, pero difiere en el ruimero y mor-
fologfa de los folfolos, inflorescencias mas largas,
la morfologfa de la flor y del fruto, asf como una
distribucion altitudinal menor. Se ineluye un cua-
dro eomparativo para distinguir las espeeies cer-
canas a esle nuevo taxon.

A Its i n \cr. Eysenhardtia officinalis (Leguminosae,
Papilionoideae), a new species from Mexico, is de¬
scribed and illustrated. This species appears sim¬
ilar to E. polystachya (Ortega) Sargent and E. tex-
ana Scheele, but differs in the number and
morphology of llie leaflets, the longer inflorescenc¬
es, the flower and fruit morphology, and in altitu¬
dinal distribution. A table for comparing related
taxa is also provided.

Key words: Eysenhardtia. Hidalgo. Legumino¬
sae. Mexico. Papilionoideae, San Luis Potosi, Ve¬
racruz.

LI genero Eysenhardtia kunlli (Leguminosae,
Papilionoideae) ha estado conformado por 14 es¬
peeies (Pennell, 1919) y mas recienternente por 11
espeeies y 2 variedades (Lang, 1972; Lang & fsely,
1982). Barneby (1982) ineluye al genero en la Tri-
bu Amorpheae, y considera alrededor de 10 espe¬
eies. Se trata de un genero restringido principal-
mente a Mexico, con excepcidn de E. texana
Scheele, E. orthocarpa (A. Gray) S. Watson y E.
spinosa A. Gray (pie se distribuyen hasta el sur de
E.U.. asf como de E. adenostyhs llaillon (|ue se
extiende a Guatemala y El Salvador. A algunas es-
pecies se les ban atribuido propiedades diuretieas
como oeurre con: E. texana, E. polystachya (Ortega)
Sargent y E. adenostylis.

En los tratamientos taxondmicos del genero se
han considerado como caracteres importantes la si-
metrfa del caliz (tomando en cuenta la forma y la-
inani) de los dientes), el tamano d<‘ las hojas. el
numero de folfolos, el tamano de las glandulas en
los folfolos (visibles en el enves), la relacidn de
longitud entre petalos y caliz, el tamano de las es-
tipelas (en funeion de la longitud del [lecfolo), el
tamano, forma y posicion de los frutos y la presen-
cia de glandulas en el estilo (Lang, 1972: Lang &
Iselv. 1982; Pennell, 1919).

Durante la elaboracidn del tratamiento laxo-
ndmieo <jue se realiza para el genero Eysenhardtia
kunlli en el Herbario Nacional de Mexico (MEXU),
se detecto una entidad que difiere de las ya cono-
cidas para este genero, por lo que se propone como
una especie nueva, que se describe a eontiuuacion:

Eysenhardtia officinalis R. Cruz & M. Sousa, sp.
nov. TIPO: Mexico. Veracruz: Mpio. Dos Rfos,
Pachuquilla, 14.9 km al SO de Conejos, If). <
km al SO de Puente Nacional, camino a Las
Bajadas, 19°14'8.1"N, 96°37'5.3"(). 366
m.s.n.m., 4 Nov. 2001 (fl, fr), Ramiro Cruz
Duran , Esteban Martinez Salas cV Clara Hilda
Ramos Alvarez 5459 (holotipo, MEXU; isoti-
pos, FCME, 1EB, MEXU, MO). Figura I.

Arbor vel frutex (1.5-)2-4 in alius; folia (5.5— )7.4 — 1 3(—
14.5) cm longa; foliola (43-)45-65, margine revoluta, sub-
tus duobus tipis glandularum uniformiter obsita; racemi
(7-)l()-19 cm longi; petala ealyeem superantia ad duplo;
legumen 6.5-7 mm longum, (1.5— )1. 7-2 lalum, a.scen-
dens, falcatum, turgidum.

Arboles o arbustos de (1.5— )2— 4 m de alto; eor-
teza lisa, amarillo pajizo, ramas jdvenes pubescen-
tes y con glandulas conicas. Hojas (5.5 — )7 .4 — 1 3( —
14.5) cm de largo, estfpulas (3.5— )4. 5— 10 nun de
largo, subuladas, pubescentes, glandulares, articu-
ladas hasta (0.8-) I (-1 .2) mm, con una glandula
ovada en la base; peefolo 0.4— 0.9(— 1 .2) cm de lar-

Novon 15: 405-409. Published on 21 September 2005.

406

Novon

Figura 1. Eysenhardtia officinalis R. Cruz & M. Sousa, sp. uov. —A. Rama con inflorescencias. — B. Folfolo, haz y
env^s. — C. Raquis, estipelas y gldnclulas. — [). Estfpulas v base del pecfolo. — K. Flor. — F. CAliz. — G. Estandarte.
— H. Ala. —I. Ouilla. — J. Androceo. — K. Pistilo. — L. F rut os. — M. Semillas. (A-K, holotipo, R. Cruz I)., E. Martinez
S. & C. //. Ramos A. 5459 (MEXU); I.. M, M. Riba s.n . (FCME, MEXU)).

Volume 15, Number 3
2005

Cruz & Sousa

Eysenhardtia officinalis (Leguminosae)

407

go, con pubescencia blanca, con numerosas glan-
« In las conicas; rai| n i s acanalado, pubeseente con
glandulas conicas a ainbos lados; estipelas subu-
ladas, 0.1— 0.2 mm de largo, pubescentes. menores
en longitud al peciolulo; peciolulo 0.5—1 mm de
largo, pubeseente; folfolos (43— )45— 65, (7— )9— 14 X
2.4— 3.8(— 4) mm, opuestos a subopuestos, oblongos,
base oblicua, ligeramente cordada, apiee truncado,
margen revoluto, baz pubeseente con pelos cortos,
vena media hendida (dandole apariencia carinal al
folfolo), venas secundarias prominentes, enves pu-
bescente con pelos largos, mas evidentes en la vena
media, glandulas de dos tamanos, distribuidas de
manera uniforme. Racimos (7—) 10— 19 cm de largo,
axilares solitarios o terminales a modo de panfcula
(en ocasiones un solo racimo terminal), pedunculo
pubeseente. Flores 5—6 mm de largo; bracteas 1.5—
2 mm de largo, lanceoladas, pubescentes en ambas
caras, con 3—4 glandulas en el enves, justo abajo
del punto de insercion con el raquis se presenta un
conjunto de pelos largos a modo de rnechon y en
ocasiones tambien se presentan dos glandulas la-
terales prominentes; pedicelo 0.5—1 mm de largo,
eilmdrico, pubeseente; tubo del caliz 2.1— 2. 5(— 2.6)
mm de largo, campanulado, pubeseente, eon glan¬
dulas prominentes, generalmente hacia el lado ca¬
rinal, base del caliz engrosada, a manera de audio
con costillas prominentes en la articulacion con el
pedicelo, los lobulos del caliz verdosos, 0.4— 0.5
mm de largo, con margenes ciliados, interior del
caliz pubeseente, al menus en la mitad superior;
corola 4.5—6 mm de largo, blanca, petalos mem-
branaceos (menos del doble de la longitud del ca¬
liz), el estandarte emarginado, 5.5—6 X 1.6—2 mm,
la una (1.2— ) 1 .4—1 .7 mm de largo; ala 5.2— 5.8 X
1.3— 1.6 mm, la una (1 .3— ) 1 .6— 1 .7 mm de largo;
quilla 4.8— 5.6 X 1.2- 1.4 mm, la una 1.4—2 mm de
largo; androceo 5—5.5 mm de largo, parte connata
(1.1 — ) 1 . 5 — 2 mm en longitud, filamentos verdosos;
el pistilo 5—5.3 mm de largo, ovario 1—1 .2 mm de
largo, glabro, verde, el estilo 3.8—4 mm de largo,
exerto, recto, pubeseente, con una glandula trans-
lucida en el doblez de 0.5— 0.7 mm, (4 estigma li¬
geramente capitado, verde. Legumbres 6.5—7 X
(1.5— ) 1.7— 2 mm, pediceladas, ascendentes, falca-
das. turgentes, con glandulas prominentes y algu-
nos tricomas hacia la parte dorsal, con caliz per-
sistente (este sin abertura dorsal), densamente
aglomeradas y aplicadas al raquis. Semilla una por
fruto, pendula, 3.8—4 X 1.1 X 0.9 nun, oblongo-
falcada, testa cafe-negra, gruesa, brillante, punc-
tulada, el micropilo prominente.

A 'ombre comun. “Taray” (Veracruz, Villanueva

/); “Chiilab” (Huasteco, San Luis Potosi, Alcorn
2147 , 2037).

Fe nolog ui. Florece en (septiembre) octubre-
noviembre (diciembre), fructifica en (octubre) no-
viembre— diciembre.

Altilud. (1 50-)200-477(-550) m.

Distribucion. En Veracruz las poblaciones co-
nocidas de Eysenhardtia officinalis son abundantes
y se |iresentan de manera continua en suelo negro
sobre sustrato calizo, y oeupan una extension de
aproximadamente 20 km2. La especie se distribuye
en selvas bajas caducifolias, se le encuentra aso-
ciada a especies de Mucuna Adanson, Porophyilum
Guettard. Caesalpinia cacalaco Bonpland, Rasselia
Jacquin. fndigofera I ... Acacia Miller, Ijeucaena
Bentham, Acacia cornigera (L.) Willdenow, Des-
rnodium Desvaux, Aeschynomene L., Centrosema
(DC.) Bentham y Graminae. Se presentan tambien
individuos aislados en pastizales o zonas pertur-
badas, en remanentes de selva mediana, asociada
con gramfneas. En el estado de San Luis Potosi se
le ha eneontrado en selva alta perennifolia, asocia¬
da con Sabal mexicana Martius y en el estado de
Hidalgo, con Calliandra houstoniana (Miller)
Standley y Leucaena pulverulenta (Schlechtendal)
Bentham.

EpCteto especifco. El nombre de esta especie
hace alusion al uso medicinal que se le ha dado
prineipalmente en la zona de Plan del Rio, Vera¬
cruz, Mexico.

Usos. Porciones del tallo, junto con la eorteza,
son colocados en un recipiente con agua, asf sa-
turan el lfquido de un color amarillo palido que al
contacto con la luz se torna azul fluorescente. Esta
infusion se ingiere como agua de tiempo. En Plan
del Rio, Veracruz se sabe del uso y comercializa-
cion de la madera y eorteza de esta especie contra
afecciones del rindn y diabetes, al menos por los
ultimos 20 anos, tambien se usa como cerca viva.

En la etnobotanica huasteca las hojas se utilizan
en afecciones de rindn. la eorteza en tratamientos
veterinarios y la madera en la obtencidn de colo-
rante, asf como en la elaboracion de utensilios de
labranza y fabricaeidn de muebles (Alcorn. 1984).

Eysenhardtia officinalis difiere claramente de E.
polystachya y E. texana, por una mayor longitud de
sus hojas, un mayor numero de folfolos, estfpulas
mas largas, estipelas mas cortas (casi nulas), raci¬
mos mas largos, y los petalos menos del doble de
la longitud del caliz. Por otro lado. la distribucion
altitudinal de esta nueva entidad es menos de 600
m, a diferencia de las otras dos que prosperan por

408

Novon

Cuadro 1. Comparaci<5n

entre Eysenhardtia officinalis

y espeeies alines.

F. polystachya

E. texana

F. officinalis

Hojas, longitud (cm)

2-11

3-10

(5.5-)7.4-13(-14.5)

No. folfolos

29-61

1 5-47

(43-)45-65

Folfolos, tamano (mm)

4-14 x 2-4

4-13 X 2-3.8

(7-)9-14 X (2.4— )2.5—

3.8(4)

Estfpulas, longitud (mm)

1 .2-5.2

1 ,5-5.5(-6.5)

(3.5-)4.5-l0

Estipelas. longitud (mm)

(t. 3-1.4, subigual o mayor
al peciblulo

0.2— 0.8, menor o igual al
peeidlulo

0.1— 0.2, menor al peeidlulo

Raeiinos, longitud (cm)

4-14

3-1 1

(7-) 10- 19

Pdtalos, longitud (mm)

4-9, mas del doble de la

5—8, el doble de la longitud

4.5— 6.0, menus del doble

longitud del ealiz

del eiiliz

de la longitud del etiliz

Tubo del ealiz. longitud
(mm)

2-4.2

2-4

2. 1-2.6

Estilo, longitud (mm)

3-4

2.5-3

3.8-4

Legumbre, tamano (mm)

9.5-15.5 X 2-4

6-13 X 2.3-2. 6

6.5-7 X (I.5-) 1.7-2

Legumbre, forma y posieidn

aplanada, recta, retleja, ca-

turgida, aseendente, caliz

turgida, faleada, ascenden-

liz persistente con aber-

persistente con abertura

te. ealiz persistente sin

tura dorsal

dorsal breve

abertura dorsal

Semilla, tamano (mm)

4.5-5. 5 X 2 X 0.6

3. 2-3.7 X 1.4- 1.6 X 0.9

3.8-4 X l.l X 0.9

Testa de la semilla

cafe, lisa

cafe, lisa

cafe-negra, punetulada

Florae ion (perfodo)

jun.— oet.

abr.— nov.

(sep.)oet.— nov.(die.)

Fruclilicacibn (perfodo)

ago.— ene.

may. -die.

(oet.)nov.— die.

Vegetaeibil

pino— encino. chaparral, ma-

matorral halbfilo, bosque de

selvas baja a mediana ea-

torral de mezquite

Juniperus, bosque de
Quercus, matorral sub-
monlano

ducifolias y alia perenni-
folia

Altitud (m. s.n. m.)

( 1 000-)2000-2600(-3200)

1 000 — 1 600(— 2500)

( 1 50-)200-47 7 (-550)

arriba de los 1000 in; asimismo, la espeuie nueva
cavcn en diferentes tipos dr vegetacion (Cuadro 1).

Si bien pareciera c|iie E. officinalis guarda mia
mayor relaeidn eon E. texana, resulta evidente que
la disposition de las glandulas en el enves de los
folfolos no es igual, pues, aunque en ambas enti-
dades se presentan glandulas de dos tamanos, en
E. texana las glandulas de mayor tamano confor-
man una lfnea a ambos lados de la vena media, asi
como en el margen, mientras cpie en E. officinalis
adoptan una distribution irregular o dispersa.
Aunque en ambas esperies se presentan frutos as-
eendentes, en E. officinalis estos son considerable-
mente faltados y ttirgidos, eon caliz persistente,
pero sin abertura dorsal, aunado a la preseneia de
semi lias cale-negras y visiblemente falcadas.

Lang (1972: 204) menciona cpie: “Like most oth¬
er species ol Eyscnhardtia , it | E. texana] hybridizes
with E. polystachya. My primary evidence for this
viewpoint, as stated elsewhere, is that intermediate
forms are found in areas where the two come to¬
gether, and only in these areas. However, two spec¬
imens [one ol these probably Purpus 5638] from
northern Veracruz (beyond the range of E. poly¬
stachya), intermediate in fruit shape and leaflet
glands between these species, may provide minor

exception to this generalization.” Considerando
esto, en nuestras observaeiones de campo, princi-
pahnente en las cercanias de Cerro Gordo en el
estado de Veracruz (si t io donde prospera una po-
blacion bastante grande de E. officinalis), no se en-
contro ninguna de las otras espeeies (E. polystachya
o E. texana), por Io que se desecha la posibilidad
de cpie se trate de una poblaeion hfbrida.

Paratipos. MEXICO. Veracruz: Mpio. Dos Rfos,
29.3 km al SF de Xalapa, cercanfas de Cerro Gordo, R.
Cruz I).. E. Martinez S. tk C. II. Ramos A. 5103 (KCME,
ML XU); 3.48 km al SE de Plan del Rfo, desviaridn a
Actopan, earr. Xalapa— Veracruz, R. Cruz I). et al. 5434
(FCME, MEXll); entre Rancho El Diamante y Rancho El
Capuh'n, camino a Misantla, R. Cruz I ). et al. 5437. 5437
his (FCME, MEXU): km 28 earr. Fed. Xalapa-Veraeruz,
30 km al E de Xalapa, cercanfas de Cerro Gordo, R. Cruz
I). et al. 5359, 5364 (FCME, MEXU); Plan del Rfo, M.
Riba s.n. (FCME, MEXU), R. Villanueva G. I (MEXU); El
Carrizal, F. Ventura A. 2823 (1)S, MICH, NY); FI Aguaje.
F. Ventura A. 9098 (MO); Cerro Gordo, F. Ventura A. 2606
(US, NY, MICH); Mpio. Emiliano Zapata, earr. Carrizal—
Los Banos, km 4.5, G. Castilleja 7 (F); earr. Xalapa-Ve¬
raeruz, desviacion para Los Banos del Carrizal, ,/. I. Cal-
zada 2056 (NY, UC); Barranca de Santa Marfa Zacuapan,
C. A. Purpus 5638 (UC); Halo de La Higuera, C. A. Purpus
s.n. (US); Mpio. Actopan, Idolos, F. Ventura 19231
(MEXU); sin Mpio. 10.1 mi. W of El Tamarindo on Hwy.
140, Seigler, Nickrent & Herlocher I)S 12227 (NY). Hi-

Volume 15, Number 3
2005

Cruz & Sousa

Eysenhardtia officinalis (Leguminosae)

409

dulgo: Huejutla v Maeuxtepetla, camino a Xaltoean, II.
E. Moore 2243 (CH): Huejutla de Los Keyes, 6 kin al SO,
carr. 105 a Paehuea. C. //. Hughes I860 (MEXU, MO.
NY, TEX). San Luis Potosi: Mpio. San Antonio. Tanjas-
nee, ./. B. Alcorn 2147 (MEXU. TEX); Lejem, ./. B. Alcorn
2037 (TEX); El Salto, 2 km al S, R. (Wether & H. < Juero
750 (CAS. MEXU).

Agradecimientos. A los Biologos Esteban Mar¬
tinez Salas y Clara Hilda Ramos Alvarez del Her-
bario Nacional de Mexico (MEXU), Instituto de
Biologia, UNAM. por llamar la atencidn sobre la
existencia de este taxon. A la M. en C. Martha Mar¬
tinez Gordillo por la diagnosis en Latin. Al personal
tecnico de Herbaria Nacional (MEXU) por permit ir
la consulta del material de herbario. A los revisores

del manuscrito por sus acertadas sugerencias. A la
Dra. Martha Riba sus valiosas eolectas.

I .iteratura Citada

Alcorn, J. B. 1984. Huastec Mayan Ethnobotany. Univ.
Texas Press, Austin.

Barneby, R. C. 1982. Daleae imagines. Mem. New York
Bot. Card. 27: 1-892.

Lang, J. M. 1972. Eysenhardtia (Leguminosae); Taxonomic
Revision and Relationships. Ph.D. Dissertation, Iowa
State University, Ames. Diss. Abstr. Ini. B. Sci. Eng.
33(10): 4092. 1973.

- & D. Isely. 1982. Eysenhardtia (Leguminosae:

Papilionoideae). Iowa State J. Res. 56(4): 393—41 7.
Pennell, E. W. 1919. Eysenhardtia. N. Amer. El. 24: 34—
40.

Protium aidanianum, a New Species from Western Amazonia.
Studies in Neotropical Burseraceae XII

Douglas C. Daly

Institute ot Systematic Botany, The New York Botanical Garden, Bronx,

New' York 10458-5126, U.S.A. ddaly@nybg.org

Abstract. Protium aidanianum (Burseraceae), a
distinctive new species from western Amazonian
Ecuador and Peru, is described, illustrated, and
discussed. It resembles !\ nodulosum Swart be¬
cause of its large, coriaceous leaflets and large,
brownish, substipitate, lenticellate fruits, but the
new species has leaves 5- or 6-jugate (vs. 2- to 4-
jugate). lateral leaflet base truncate to slightly cor¬
date (vs. usually acute), inflorescence shorter than
the petiole of the subtending leal, flowers 4-merous
(not 5-merous), petals glabrous and not persistent
in fruit (vs. pubescent and often persistent and re¬
flexed in young fruits), pistillode glabrous, and fruit
larger and with rounded (vs. acuminate) apex. It
resembles P. glabrescens Swart due to its robust
branchlets. the inflorescence shorter than the pet¬
iole of the subtending leaf, and the petiole often
turning black and exfoliating, but the new species
differs by its deeply canaliculate petiole, higher-
order venation impressed adaxially, shorter pedicel,
petal margin long-papillate, and fruit brown or cof¬
fee-colored and much larger. The fact that only 1
of the 16 collections known to date has flowers
highlights both the difficulty of finding flowering
material and the need to actively seek it, particu¬
larly for dioecious tropical trees with small green
flowers.

Key words: Amazonia, Burseraceae, Ecuador,
Neotropics, Peru. Protium.

Documentation of the Neotropical flora has im¬
proved significantly over the past 30 years, but the
increase in collections of newT taxa and progress
toward more realistic distribution records has not
been matched by an increase in the proportion of
systematically useful collections. Particularly in
light of the recent surge in developmental studies
for phylogenetic work (e.g., Costello & Motley,
2001), adequate representation of a species ulti¬
mately requires collections for every developmental
stage of inflorescences, flowers, and fruits, as well
as seeds, germinating seeds, and seedlings.

As in many tropical tree groups, the flowers of
Burseraceae are small, green (usually), and unisex¬

ual (most), the latter implying the need for flowering
material from both staminate and pistillate individ¬
uals. Moreover, for many taxa the flowers persist
only a short time. The net result is that tropical
systematists have cabinets full of new taxa repre¬
sented by material insufficient for lull description
and therefore publication. Two strategies can com¬
pensate for this: first, botanists collecting tropical
trees should always use binoculars to assist in lo¬
cation of flowering trees; and second, phenological
studies often he! p to catch trees that have short
flowering phases. Meanwhile, the following new
species from Amazonian Ecuador and Peru is rep¬
resented by 16 collections, but 3 are sterile, 12
have fruits (most with immature fruits), and 1 has
buds and a few staminate flowers only.

Protium uhlaiiiuiiiuti Daly, sp. nov. TY PE: Ec¬
uador. Napo: La Joya de los Sachas, Pompeya,
S side of Rfo Napo, Rio Jivino, Maxus Camp,
km 5-7 Maxus Road, 00°25’S, 76°37’W. 220
m, 22-24 Aug. 1992 (fl), A. Grijalva & E. Gu-
dino 2 (holotype, NY |2|; isotypes, MO.
k)CNE). f igure 1.

Arbor mediocris vel grandis foliola grandia coriacea
fructus grandis bruneolus lenticellatus a P. noduloso fo-
liolis 5— 6-jugatis (non 2— 4-jugatis) inflorescentia petiolo
breviore floribus 4-partitis (nee 5-partitis) petalis glabris
(ruetu grandiore apice rotundo differt. A P. glabrescenti
nervis tertiariis adaxialibus itnpressis pedicellis breviori-
bus petalis longo-papillatis fructu grandiore bruneolo (nee
rubello) differt.

Tree, reproductive height (6)10—30 m X 12—50
cm; buttresses present at least sometimes (“high”
in Pinkley 478 in sched., at base in SEE 10867);
bark gray-green, smooth, outer bark 2 mm thic k on
small trees (SEE 10367), inner bark yellow-brown;
branchlets robust, to 1.5 cm diam. proximal to apex;
trichomes of four types: malpighiaceous hairs with
lateral length ea. 0.2 mm; fine erect “bristles” to
0.05 mm long (to 0.1 mm on adaxial midvein); cap¬
itate glands; and thick golden appressed hairs to
0.1 mm long, these mostly at inflorescence branch¬
ing points. Leaves 5- to 6-jugate, 43—90 cm long;
petiole 7—31 cm X 6-9 mm, toward the base deeply

Novon 15; 410-413. Published on 21 September 2005.

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2005

Daly

Neotropical Burseraceae XII

411

Figure 1. Protium aidanianum Daly. — A. Portion of leaf, with inset showing detail of petiolule. — B. Young inflo¬
rescence arising from leaf axil. — C. Staminate bud. — I). Staininate flower with part of calyx, one petal, and four
stamens removed (right), and longisection of disk and pistillode (left). — E. Adaxial view of petal. — F. Dorsal, ventral,
and lateral view of stamen. — G. Fruiting branchlet with leal removed. — H. Lateral and ventral views of pyrene. (Based
on A. Cerori 3464, Grijalva & Gudina 2, Tunqui 251.)

canaliculate and broad (nearly semi-clasping), often
lenticellate; interjuga 2—7 cm long, petiole and ra-
chis provided with sparse to scattered malpighia-
ceous hairs, usually with sparse to scattered bris¬
tles, and sometimes with sparse glands, in fruit the
petiole and rachis sometimes partly glabrescent,
when dry the epidermis black and exfoliating in
many spots; lateral petiolules robust, 0.8— 3.2 cm
long, with a conspicuous pulvinulus at distal end,
terminal petiolule 3.5— 6.3 cm long; leaflets coria¬
ceous, drying yellowish brown, the laterals 13.5—

38.3 X 5.5—12 cm, the basal pair (broadly) elliptic
to oblong-elliptic, other laterals oblong to oblong-

elliptic; terminal leaflet 17.5-31 X 8— 1 1 cm, ob¬
long-elliptic to slightly oblanceolate, leaflet apex
abruptly and narrowly acuminate, the acumen
(0.2)0. 6— 1.5 cm long, leaflet base subequal, on lat¬
erals truncate to slightly cordate, on the terminal
one obtuse to slightly cordate, the midvein acutely
prominent abaxially, the remaining vein orders
prominulous, the midvein and secondary veins pro¬
vided abaxially with scattered to sparse bristles
(sometimes to 0.1 mm long) and sometimes scat¬
tered thick appressed hairs; the midvein prominu¬
lous but sunk in a groove adaxially, remaining vein
orders impressed (sometimes the tertiary and higher

412

Novon

orders Hat), the midvein provided with scattered
bristles to 0.1 mm long as well as capitate glands.
Inflorescences (Fig. IB) (3.3)5—15.5 cm X 0.3— 0.4
mm (X 3.5—4 mm in fruit), shorter than petiole of
subtending leaf, secondary axes to 4.5 cm long,
axes provided with sparse to scattered malpighia-
ceous hairs and sparse bristles, sometimes also with
sparse glands; bracts on primary inflorescence axes
(1)1.3— 2 mm long, subulate to ovate, hraeteoles
subtending flowers 0.3— 0.5 mm long, subulate, the
bracts densely provided with thick oppressed hairs;
(lowering pedicel 0.7— 1.5 X 0. 3—0.4 mm, provided
with scattered malpighiaceous hairs and (at the
base) sparse bristles and capitate glands. Staminate
flowers (Fig. 1C— F) 4-merous, ca. 3.3 mm long over¬
all at anthesis; calyx ca. 1—1. 2(1.4) X 2.1— 2.4 mm
overall, deeply cupular and slightly ureeolate, ex¬
ceeding the disk, the lobes (0)0.2— 0.3 mm long,
rounded perdepressed-deltate or sometimes re¬
duced to an apiculum, calyx glabrous except the
lobe apices long-papillate; petals greenish yellow,
ca. 2.7 X 1.2 mm, lanceolate, slightly fleshy, with
an indexed but inconspicuous apiculum 0.3—0.35
mm long, petals glabrous but margin long-papillate;
antcsepalous stamens 1.45—1.5 mm long, filaments
strap-shaped, the anthers ca. 1 mm long and ob¬
long, antepetalous stamens 1.25—1.3 mm, the an¬
thers ca. 0.85—0.9 mm long and slightly ovate; disk
0.2—0.35 X 0.2 mm thick; pistillode 0.65—0.75 X
0.35-0.7 mm overall, ovary ovoid, the style 0.2 mm
high with 4-lobulate apex; pistillate flowers un¬
known. Fruiting pedicel apparently variable in
length, 1.5— 5.5 X 2.5—4 mm, terete, fruit (Fig. IG)
brown or coffee-colored (reported as blackish green
in Gerdn 3464), 3.2— 4.5 X 2.6— 3.5 cm (to 3.8+ cm
broad when more than one pyrene develops), broad¬
ly ovoid, only slightly oblique, tbe apex rounded,
the base substipitate, obtuse to truncate above the
st i pe. the surface lenticellate and sometimes pro¬
vided with scattered bristles and glands; pseudaril
while, pulpy, covering all of pyrene, the pyrene
(Fig. Ill) 2.4— 3.1 X 1.7— 2.3 cm, bony, in dorsi-
ventral view the pyrene broadly ovate with acute to
sharply acuminate apex, in lateral view the poste¬
rior side semi-ovate, the anterior side semi-obovate
but deeply cleft at site of the funicular scar, which
is located 6697— 80% the distance to the pyrene
apex, the funicular scar small, narrowly inverted-
crescent shaped; pyrene surface glabrous; seed with
testa irregularly thickened and cotyledons lobed
and contortuplieate; germination and seedlings un¬
known.

Distribution and ecology. Protium aidanianum
shows a western Amazonian distribution, in Ecua¬

dor and Peru. Unlike most Protium , the new spe¬
cies tends to occur in waterlogged soils, such as in
periodically inundated areas (SEP 10367) and river
island forests (Berlin 3538a), on soils reported as
lateritic (Gentry & Diaz 28254) or red clay dystro-
pept soils (Zak 3831) and at elevations between
180 and 400 m. It is known to flower in August
and to fruit between August and January.

Common names and uses. Shakap (Huambisa,
Berlin 3538a: Huashikat 185, 973. 1038): tofake-
kopa (Kofan, Pinkley 478): chipa (Huambisa, Tun-
qui 251): patu muyu (Quichua, Grijalva A Gudina
2): shirquillo (Quichua, Revelo 94). The Huambisa
use the fruits to make necklaces (Huashikat 1038)
and the resin to paint large earthen jars (Tunqui
251), while among tin1 Kofan the resin has been
applied to the ankles and touched with charcoal to
make a black ring (Kofan, Pinkley 478), and the
lowland Quichua use the resin to make candles
(Revelo 94).

Etymology. It gives me great pleasure to name
this species for my son Aidan. who helped me with
the measurements, and who commented that it is
quite remarkable; the same goes for him.

The new species most closely resembles another
western Amazonian species, P. nodulosum Swart,
because they both show long lateral petiolules,
large coriaceous leaflets that are sometimes oblong
and sometimes glossy adaxially, and large, brown¬
ish. substipitate, lenticellate fruits. Protium aidan¬
ianum differs by its leaves 5- or 6-jugate (vs. 2- to
4-jugate), lateral leaflet base truncate to slightly
cordate (vs. usually acute), higher-order venation
impressed or rarely flat adaxially, inflorescences
shorter than the petioles, bracts on primary inflo¬
rescence axes (1)1.3— 2 (vs. 0.6— 0.9) mm long, flow¬
ering pedicel shorter (0. 7-1.8 vs. ca. 1.5—6 mm),
flowers 4-merous (not 5-merous), calyx more deeply
cupular with lobes either obsolete or short and
broad (0—0.4 vs. ca. 0.4— 0.6 mm long), petals gla¬
brous and not persistent in fruit (vs. pubescent, and
often persistent and reflexed in young fruits), pis¬
tillode glabrous, fruit larger (3.2— 4.5 X 2.6— 3.5 vs.
ca. 4 X 2.4 cm) and with rounded (vs. acuminate)
apex, and funicular scar on the pyrene located
66%— 80% the distance to the pyrene apex (not
subapical).

Protium aidanianum also resembles the south¬
western Amazonian species P. glabreseens Swart
because of their robust branehlets, petiole often
turning black and exfoliating, leaflets coriaceous
and often oblong, secondary veins impressed adax¬
ially (only sometimes in P. glabreseens), adaxial sur¬
face sometimes glossy, and inflorescence shorter

Volume 15, Number 3
2005

Daly

Neotropical Burseraceae XII

413

than the petiole of the subtending leaf. The new
species differs by its deeply (vs. shallowly) cana¬
liculate petiole, higher-order venation impressed
adaxially, pedicel shorter (ca. 0.7— 1.5 vs. 1.5—2
mm), consistently 4-merous flowers (vs. 4- to 5-
merous), deeply cupular calyx, petal margin long-
papillate (not short -papillate), and the fruit brown
or coffee-colored (vs. green, pink, or red-brown),
much larger (3.2-4.5 X 2. 6-3. 5 cm with one py¬
rene, vs. ca. 1. 8-2.1 X 1.1-2 cm), and less oblique.

Paratypes. KCUADOR. Napo: I*. N. Yasunf, near
Amo Sur Heliport, 0()°52'S, 76°05'W. 16—19 Jan. 1983
(fr), Ceron 3464 (NY); La Joya de los Sachas, Pompeya,
S side of Rio Napo, Rfo Jivino, Maxus Camp, km 5-7
Maxus Road, 00°25'S, 76°37'W, 23-29 Nov. 1992 (fr),
Grijalva el al. 234 (NY); Canton Francisco de Orellana.
V ia de los Zorros, Pozo Jaguar I. 00°44,S, i 7°05’W, 23
Oct. 1988 (fr), Palacios 3205 (NY); Dureno, on Rio
A gua rico, 4 Oct. 1996 (st), Pinkley 478 (NY); lena. Fst.
Biol. Jatun Sacha, OHM'S, 77°36'W, 24 Nov. 1994 (fr),
Revelo 94 (NY); P. N. Yasunf, Rfo Anangu near confluence
w/Rfo Napo, 0°3 1 2'S. 76°24'W, 30 June-9 July 1982 (st),
SEP 10367 (NY). I'astaza: Carretera de Petrn-Canada,
Vfa Auca, 1 15 km S of Coca, 4 km S of Rfo Tuguino,
01°15'S, 76°5.5'W, 22-28 Feb. 1989 (st). Zak 3831 (NY).

Sueumbios: Lago Agrio Canton, Res. Cuyabeno, Iara-
poa-Tipischca, Cruce del Rfo Cuyabeno, ()()°01'S,
76°15'W, 14 Nov. 1991 (fr), Palacios el al. 8957 (NY).
PERU. Amazonas: I km S of La Poza, Santiago River. 9
Aug. 1979 (fr), Berlin 3538a (MO. NY); I km from Poza,
K margin ol Rfo Santiago, 24 Aug. 1979 (fr), Huashikal
185 (MO, NY); Prov. Condorcanqui. I km beh. Caterpiza,
E of (Juebrada Caterpiza, Rfo Santiago, 17 Oct. 1979 (fr),
Huashikal 973 (MO), 24 Oct. 1979 (fr), lluashikal 1038
(MO); Rfo Santiago. Quebrada Caterpiza, 2—3 km beh. Ca¬
terpiza. 7 Dec. 1979 (fr), Tunqui 251 (MO. NY). Iliiaii-
uco: Pachitea, Codo de Pozuzo, trail W of settlement to
lower mountain slopes, 9°40'S, 75°28\\. li Oct. 1982
(fr). Poster 9239 (NY). Loreto: Prov. Alto Amazonas, An-
doas, Rfo Pastaza. near Ecuador border, 1 7 Nov. 1979 (Ir),
Gentry & Diaz 28254 (NY).

Acknowledgments. Once again, I thank Bobbi
Angell for a revealing and attractive illustration,
also James Solomon of MO for his timely help with
tracking down duplicates of the type collection.

I .iterature Cited

Costello, A. & T. J. Motley. 2001. Molecular systematics
of Tetraplasandra, Munroidendron, and Reynoldsia
sandwicensis (Araliaceae) and the evolution of superior
ovaries in Tetraplasandra. Edinburgh J. Rot. 58(2):
229-242.

A New Species of Ruyschia (Marcgraviaceae) from llie
South American Andes

Adrian C. de Roon

Nationaal Herbarium Nederland, Utrecht University branch, Heidelberglaan 2,
3584 CS Utrecht, Netherlands, aderoon@ncrvnet.nl

Abstract. Ruyschia andina, a new species found
on the eastern slopes of the Andes in Ecuador,
Peru, and Bolivia at altitudes of around 2000 m, is
described and illustrated. The species is distin¬
guished from the other species of the genus by the
leaves with two conspicuous rows of numerous hy-
pophyllous glands and the 3(or 4)-loeuled ovary.
The relationship) between Ruyschia and Souroubea
is discussed, and a generic key is given.

Key words: Andes, Bolivia, Ecuador, Marcgra¬
viaceae, Peru, Ruyschia . South America.

The genera Ruyschia Jacquin and Souroubea Au-
blet are commonly considered to be closely related
(de Boon, 1975). Swartz (1788) united the two gen¬
era, then still monotypie, under Ruyschia. This was
followed by Martins (1832) and T Viana and Plan-
chon (1802), who described a number of new' spe¬
cies of Ruyschia that currently are placed in Sou¬
roubea. Delpino (1869), the first monographer of the
Marcgraviaceae, separated the two genera again.

1 his was adopted by all subsequent students of the
Marcgraviaceae, namely Wittmack (1878), Szyszy-
lowiez (1893), Gilg and Werdermann (1925), de
Boon (1975), Bedell (1985), and Dressier (2004).
The two most important characters to distinguish
Ruyschia and Souroubea are in the nectariferous
bract and in the pistil. Ruyschia has small, folia-
ceous, flat or gibbous to slightly concave nectari¬
ferous bracts and a (sub)globular, mostly 2-loculed
ovary with a minute or distinct style and a small
stigma, while Souroubea has hollow, cup- or spur¬
shaped and auriculate nectariferous bracts and an
often angular, (3- to)5-loculed ovary with a usually
large, sessile, often radiate stigma. The placement
ol a given species in either one or the other genus
has sometimes been arbitrary. Ruyschia pilophora
Iriana N Planchon was placed by Delpino (1869)
in Souroubea on account of the small, slightly hol¬
low, somewhat Souroubea-Uke nectariferous bracts,
but was placed again in Ruyschia by de Boon
(1975) on account of the 2-loculed ovary. On the
contrary, Ruyschia platyadenia Gilg, with small,
flat, (oliaceous, nectariferous bracts, was placed in

Souroubea because ol the 5-loeuled ovary (de Boon.
1975). The separation ol Ruyschia and Souroubea
remains disputed and possibly needs reconsidera¬
tion.

The current generic subdivision of the Marcgra¬
viaceae— Noranteoideae was based on morphologi¬
cal features (Bedell, 1985; see also Dressier, 2004)
and is not supported by the molecular data pres¬
ently available. Ward and Price (2002) presented
molecular data that suggest a closer relationship
between a species of Ruyschia and a species of
Schwartzia than to investigated species of Sourou¬
bea. In a recent study of wood anatomy and pollen
morphology of the Marcgraviaceae, Van Evelgl iem
(2003) found no support for an amalgamation of
Ruyschia and Souroubea. However, both studies
(Van Evelghem and Ward N Price) included only
one species of Ruyschia and three of Souroubea,
respectively. Decisions regarding possible taxonom¬
ic changes in the Marcgraviaceae have to wait until
sufficient molecular and other data are available.

Up to now, seven species of Ruyschia have been
accepted, all from mountainous areas: R. clusiifolia
Jacquin is confined to the Lesser Antilles (St. Kitts,
Guadeloupe, Martinique, and Dominica); R. enervia
1. undell ranges from southern Mexico to Belize,
Guatemala, Honduras, and Costa Rica; R. phylla-
denia Sandwith is rather common in Costa Rica and
Panama; R. valerii Standley, closely related to R.
phylladenia, and differing in leaf shape only, is a
rare endemic in Costa Rica; R. tremadena (Ernst)
Lundell is common and widespread in Panama (Da-
ri£n), Colombia, and Venezuela; R. pilophora Iriana
& Planchon is endemic to Colombia; and R. pavonii
G. Don is known from Colombia and Peru.

During identification work another hitherto un¬
described species was found. This new species is
distinguished from the other species of the genus
by its relatively large leaves with a conspicuous row
of numerous hypophyllous glands running the
length of the leaf blade on either side of the mid¬
vein, approximately halfway between the midvein
and the leaf margins, a 15—30 cm long raceme w ith
60 to 130 flowers, and flowers with a 3(or 4)-loculed

Novon 15: 414-417. Published on 21 Septembeh 2005.

Volume 15, Number 3
2005

de Room

Ruyschia andina (Marcgraviaceae)

415

ovary. The Iasi character especially is remarkable,
as all other species of the genus have a 2-loculed
ovary.

Ruyschia andina de Roon, sp. nov. TYPE: Ec¬
uador. I’rov. Zamora-Chinchipe: Road Loja-
Zamora, km 28.3, secondary scrub above
storm drain, 3°57’S, 79°3‘V\. 2000 in. 27 Dec.
1997 (fl). O'. P. Lewis, P. Lozano & C. C. Berg
3733 (holotype, U; isotypes, AAU not seen, E
not seen, k no! seen, LOJA not seen, MO not
seen, QCA not seen. QCNA not seen). Figure I.

Frutex scandens saepe hemiepiphyticus. Folia petiolis
ad 10 mm longis; lamina coriacea. obovata, 8—15 cm lon-
ga, 3.5— 7.5 cm lata, apice obtusa vel rotundata, basi acuta
vel cuneata; glandulae hypophyllae numerosae, biseriatac.
Racemi 15—30 cm longi. (lores 60—130; nectaria leviter
cymbiformia, ovarium 3( — 4) loculare.

Shrub, hemiepiphytic shrub, or woody climber,
stem 10 cm diam., strongly branched. I .eaves pet-
iolate, petioles .5-10 mm long, 2—3 mm diam., flat¬
tened to canaliculate above; blades obovate, 8—15
cm long, 3.5— 7.5 cm wide, apex obtuse to rounded
and mucronate or minutely retuse after the mucro
has fallen, base acute to cuneate, margin slightly
revolute, coriaceous, midrib slightly prominent at
base to slightly canaliculate above, prominent be¬
neath, secondary veins obscure to slightly im¬
pressed above, obscure to subprominent beneath;
hypophyllous glands poriform, usually in a con¬
spicuous row of 10 to 25 ca. halfway between the
margin and midrib on either side. Inflorescence a
raceme, 15—30 cm long, with 60 to 130 flowers.
Flowers pedicellate, pedicels 5—8 mm long; nec¬
tariferous bracts slightly eymbiform, 3-4 mm long,
inserted just below the flowers; braeteoles directly
subtending the calyx, sepaloid, oval. 1.5—2 mm
long, ca. 2 mm wide, persistent; sepals suborbicu-
lar, ca. 2 mm long, 2.5—3 mm wide, persistent; pet¬
als oblong, 6—7 mm long, 2—3 mm wide; stamens
5, filaments flattened. 1-1.5 mm long, ca 0.5 mm
wide, abruptly narrowed toward the anthers; an¬
thers 2-3 mm long, 1 mm wide; ovary 1-1.25 mm
long, 3(or 4)-loculed, style conical, 0.75—1 mm
long. Fruit depressed globose, apiculate by the per¬
sisting style and stigma, ca. 5 mm diam., fleshy,
green tinged red.

Phenology. Flowering specimens are collected
in December. January, March, and August, and
fruiting specimens in November, December, Janu¬
ary, March, April, and May.

Distribution. Ruyschia andina is found in pri¬
mary and secondary tropical montane rain forests
and cloud forests at altitudes of 1800—2300 m on

the eastern Andean slopes in Ecuador, Peru, and
Bol i v ia.

Besides the new species, three other species are
known from South America: Ruyschia tremadena in
Venezuela, Colombia, and Panama (Darien), char¬
acterized by gibbous to semiglobose nectaries in¬
serted at the base of the calyx and by 3 stamens;
Ruyschia pilophora in Colombia, characterized by
small, excavated, hat-shaped nectaries inserted at
the base of the calyx; and Ruyschia pavonii in Co¬
lombia and Peru, characterized by (sub)globose
nectaries inserted at about the middle of the ped¬
icel.

ki.'i to the Neotropical Species ok Ri yschia
I a. Stamens 3.

2a. Leaf blade 3—5 cm wide; nectaries gibbous

to semiglobose . R. tremadena

2b. Leaf blade 1.5—3 cm wide; nectaries Lilia¬
ceous, flattened or slightly gibbous.

3a. Leaf blade acuminate; pedicels 3—5 mm

long . R. valerii

3b. Leaf blade obtuse to subacuminate;

pedicels 5—9 mm long . . . R. phylladenia
I b. Stamens 5.

4a. Nectaries inserted near the middle of the

pedicel . R. pavonii

4b. Nectaries inserted at the base of the flowers.

5a. Nectaries hollowed out, hat-shaped . .

. R. pilophora

5b. Nectaries solid, gibbous to semiglobose.

6a. Hy, lophyllous glands numerous in
a distinct row of 10 to 25 ca. half¬
way between the margin and mid¬
vein on either side; ovary 3-celled

. R. andina

6b. Hypophyllous glands few, scattered
or in a row of 2 to 7 between the
margin and midvein on either side,
sometimes absent; ovary 2-celled.

7a. Hypophyllous glands small or
minute; nectaries eymbiform,
concave beneath; pedicels
slender, 4—10 mm long, ca. 1

mm thick . R. clusiifolia

7b. Hypophyllous glands large,
with a light-colored circumval-
lation, nectaries gibbous to se¬
miglobose, flattened beneath;
pedicels stout, 3—5 rnm long,
1-1.5 mm thick .... R. enervia

Paratypes. BOLIVIA. La Paz: Prov. Sud Yungas, Si-
rubaya near Yanacachi, 0. Buchtien 520 (US, W); 7 km
de Huancane en carr. a San Isidro, I). I\. Smith, V. Garcia,
L. Dorr & L. Barnett 13915 (B, LPB not seen. MO not
seen, U). ECUADOR. Prov. /ainora-Cliineliipe: Road
Loja— Zamora, G. P. Lewis, J. Luteyn, R. Fuller & E. M.
Guerrdn 3249 (AAU not seen, R, K not seen, LOJA not
seen, MO not seen, QCA not seen, OCNE not seen); area
of Estacion Cientffica San Francisco, ca. 30 km away from
Loja on hwy. tow. Zamora, S. Matezki 227 (FR, MO not
seen), S. Matezki 453 (FR. UBT not seen); E Andean

416

Novon

Figure 1. Ruyschia andina de Roon. — A. Flowering branch. — B. Flower bud with nectariferous bract. — -C. Flower.
— I). Infructescence. — E. Fruit. — F. Fruit, transverse section. A. B from I a- iris el <d. 3733 (U) ; C from Matezki 433
(FR): I) from Lewis el <tl. 3249 (B); F from Matezki 227 (FR).

slopes. Quebrada El Milagro watershed. I). Neill & QCNE
botany interns 12612 (FR, MO not seen. OC N F not seen);
El Bangui. Cordillera del C6ndor, dost, militar Crtndor
Mirador en la Frontera Ecuador— Peru, G. Pabon, ./. Car-
a ni/ u i tC Grupo Post-Grado MO-QCNE 333 (MO not seen.

U); sandstone escarpment above & \\ of Tikimints, valley
of Rid Coangas. P. hi. Perry & I). Neill 7635 (FR, MO not
seen, QCNF, not seen). Prov. Morona Santiago: Canton
San Miguel de los Cuyes, main trail betw. the villages of
San Miguel de los Cuyes Canazhuma (via Rfo San Mi-

Volume 15, Number 3
2005

de Roon

Ruyschia andina (Marcgraviaceae)

417

gu el de I os Cuves), J. L (dark & C. Morocho 5859 (U.
US not seen). PERU. Cajamarca: Prov. San Ignacio. San
Jose de Lourdes, Camino al Cerro Picorana, ./. Campos ,
L. Campos & L. Zurita 5514 (FR. MO not seen).

Acknowledgments. I thank the curator of the
Senckenberg herbarium (FR) for sending me a
comprehensive collection of Ruyschia and Sourou-
hea for identification, among which the type collec¬
tion and many paratypes of the new species were
found. I am much indebted to Stefan Dressier and
to Gwilym Lewis for valuable suggestions, which
improved the manuscript. I thank Lubber! Wostra
for his assistance with the Latin diagnosis and the
first draft and Hendrik Rypkema for the illustration.

literature Cited

Redell, II. G. 1985. A Generic Revision of Marcgravi¬
aceae I. Phe Norantea Complex. Ph.D. Dissertation,
University of Maryland, College Park. [Unpublished.]
de Roon, A. C. 1975. Contributions Towards a Monograph
of the Marcgraviaceae. Ph.l). Thesis, Rijksuniversiteit,
Utrecht.

Delpino, G. C. F. 1869. Ulteriori osservationi e consider-
azioni sulla Dicogarnia nel regno vegetale. Revista mon-
ografica della famiglia delle Marcgraviaceae precipua-

mente sotto I apetto della biologia ossia delle relazioni
di vita esteriore. Atti Soe. I tal. Sci. Nat 12: 179—213.

Dressier, S. 2004. Marcgraviaceae. Pp. 258—265 in k. ku-
bilzki (editor). The Families and Genera of Vascular
Plants, Vol. 6. Flowering Plants. Dicotyledons: Celas-
trales, Oxalidafes, Rosales, Comales, Frieales. Spring¬
er. Berlin, Heidelberg, New York.

Gilg, F. & E. Werdermann. 1925. Marcgraviaceae. In: A.
Fngler & k. Grant I (editors), Die natirrliehen Pflanzen-
familien, ed. 2, 21: 94—106. Wilhelm Frigelmann. Leip¬
zig.

Martius, C. F. P. 1832. Nova Genera et Species Plantarum
3(3): 174—179. Miinchen.

Swartz, 0. P. 1788. Nova Genera & Species Plantarum
sen Prodromus etc. Stockholm, Uppsala, Abo.

Szvszylowicz, I. 1893. Marcgraviaceae. In: A. Fngler & k.
Prantl (editors). Die natiirlichen Pffanzenfamilien 3(6):
157—164. Wilhelm Frigelmann. Leipzig.

Triana. J. & J. E. Planehon. 1862. Ruyschia. In: Prodro¬
mus Florae Novo-Granatensis. Ann. Sci. Nat. Ser. 4,
tome 17: 376—379.

Van Evelghem, L. 2003. Houtanatomie en Pollenmorfolo-
gie van de Marcgraviaceae. Undergraduate Thesis, In¬
stitute of Botany and Microbiology, k.U. Leuven. [Un¬
published.]

Ward. N. M. & R. A. Price. 2002. Phylogenetic relation¬
ships of Marcgraviaceae: Insights from three chloroplast
genes. Svst. Rot. 27: 149—160.

Wittmack, L. 1878. Marcgraviaceae. In: C. E P. von Mar¬
tius, Flora Brasiliensis 12(1): 213—258. Miinchen,
\\ ien, Leipzig.

A New Species of Crotalaria (Leguminosae, Papilionoideae)
from Southeastern Brazil

Andreiu Silva Flores and Ana M. Goulart do Azevedo Tozzi
Departamento de Botaniea, Institute de Biologia, Universidade Estadual de Campinas,
Caixa Postal 6109, Cani|)inas 13083-970, SP. Brazil. andreiasflores@itelefonica.com.br;

anatozzi@unicamp.br

Abstract. Crotalaria miottoae, a new species of
Leguminosae from southeastern Brazil, is described
and illustrated. The new species is close to Crota¬
laria vitellina Ker Gawler, but differs in having
larger wing petals, fruits, and seeds, in having ter¬
minal inflorescences, rarely leaf-opposed, and the
leaflets usually rhombic or narrowly elliptic.

Rksumo. Crotalaria miottoae, uma nova especie
de Leguminosae no sudeste do Brasil e descrita e
ilustrada. Ksta especie e relacionada h Crotalaria
vitellina Ker Gawler, diferindo desta por possuir
alas, frutos e sementes maiores. inflorescencias ter-
minais, raramente opositifdlias e folfolos rombieos
ou estreitamente elfptieos.

Key words: Brazil, Crotalaria, Leguminosae,
Papilionoideae.

Crotalaria L. is a large legume genus of ca. 600
species distributed in the tropical and subtropical
regions of the world mostly in Africa and Asia (Pol-
hill. 1981, 1982). while 42 species have been re¬
corded lor Brazil. During a taxonomic revision of
the Brazilian species of Crotalaria, an undescribed
species came to our attention and is described
herein as new.

Crotalaria miottoae A. S. Flores & A. M. G.
Azevedo, sp. nov. TYPE: Brazil. Minas Gerais:
Camanducaia, Monte Verde, estrada para Ped-
ra Partida, 2 Jul. 2002, A. S. Flores 1036 <£
R. Schiitz Rodrigues (holotype, UEC: isotypes,
F, HBG, MO, SPF). Figure 1.

Haec species C. vitellinae rarnis puberulis et braeteis
persistentibus similis sed alis longioribus, leguminibus et
seminibus majoribus. foliolis saepe rhombieis vel anguste
ellipticis et infloreseentiis terminalibus raro oppositifoliis
differt.

Erect shrub or subshrub, 1.5—2 rn high; branches
puberulous to finely pubescent with short, yellow
or white triehomes. Internodal stem wings absent.
Leaves digitately 3- to 4- or 5-foliolate, petiole 2.5—
5.4 cm long, puberulous; leaflets usually rhombic

or narrowly elliptic, the apex acuminate or acute,
base cuneate, 3-5. 8(7.4) X I -1.5(2. 5) cm, disco-
lorous, puberulous above, glabrous beneath; stipule
1—1.5 mm long, linear, usually persistent. Inflores¬
cence a terminal or rarely leaf-opposed raceme, 7—
15 cm long, 8- to 25(35)-flowered; peduncle 1.5—
2.5 cm long; pedicel 7-9 mm long; bract 2-4 mm
long, filiform, persistent; braeteole 1-2 mm long,
filiform, inserted at the middle of tin* pedicel, per¬
sistent. Calyx 7-10 mm long, shorter than the co¬
rolla, the tube campanulate; lobes 5, twice as long
as the tube, triangular to lanceolate, finely pubes¬
cent or glabrescent; corolla yellow; standard petal

1.2— 1.8 X 1.1 — 1.7 cm, suborbicular; wing petal

1.3— 1.8 cm long, obovate to oblong; keel 1.1-1. 6
cm long, almost as long as the wing petals, dorsal
margin rounded, the non-twisted beak short, the
margins filiate or glabrous. Stamens dimorphic
with anthers alternately 5 long, basifixed and 5
smaller, dorsifixed; ovary 4—5 mm long, oblong, pu¬
berulous along the margin and glabrous at the cen¬
ter, stipitate; style curved, stigma pubescent. Le¬
gume 3.2— 4.3 X 1.1— 1.4 cm. broadly oblong or
obovoid. puberulous or glabrous, stipitate, black at
maturity, ca. 20-seeded; seeds 6—7 X 5—6 mm.
oblique-cordiform, seed coat black, opaque.

Distribution and ecology. Southeastern Brazil,
in Rio de Janeiro, Espfrilo Santo, and Minas Gerais
states. Crotalaria miottoae grows in campo rupes-
tre, along the margins of wet forest and inside mon¬
tane forests at elevations of 1000— 1900 m.

Phenology. Collected in flower and in fruit dur¬
ing most of the year.

The species is named in honor of Silvia T. Sfog-
gia Miotto, Brazilian legume specialist, who has
stimulated this work since the beginning.

According to the sectional classification pro¬
posed by Bisby and Polhill (1973), Crotalaria miot¬
toae should be assigned to section Chrysocalycinae
(Bentham) Baker f. due to its calyx subequally 5-
lobed and 0.7 as long as the keel. It is closely
related to Crotalaria vitellina Ker Gawler, both spe-

Novon 15; 418-420. Published on 21 September 2005.

Volume 15, Number 3
2005

Flores & Azevedo

Crotalaria miottoae (Leguminosae)

419

Figure 1. Crotalaria miottoae A. S. Flores & A. M. G. Azevedo. — A. Fertile branches. — B. Flower. — C. Calyx. —
D. Details of corolla: standard petal, wings, and keel. — E. Alternative view ol standard petal. — F. Part of infructescence
with mature fruits. (Drawn from the holotype.)

cies having puberulous to finely pubescent branch¬
es and persistent bracts. However, C. vitellina has
smaller wing petals (0.9— 1.3 cm long), legumes
1 .6-3.2 X O.f)— 0.9 cm, subcylindrical or oblong-
ellipsoid, brown at maturity, seeds 4X4 mm,
brown and elliptic:, oblong, or obovate leaflets. Cro¬
talaria miottoae also differs in having terminal in¬
florescences, which are absent in C. vitellina. Cro¬
talaria laeta Martius ex Bentham, proposed as a

variety of C. vitellina by Windier and Skinner
(1982), is considered a distinct species by Flores
(unpublished thesis). Crotalaria laeta differs from
both species in having the wing petal smaller than
the keel. In addition, it differs from C. miottoae in
having a smaller wing petal (7—8 mm long), keel
(9—11 mm long), fruits (1.2— 1.8 X 0.4— 0.7 cm), and
seeds (3X3 mm). Crotalaria miottoae resembles
vegetatively C. m irons Link and has similar fruits.

420

Novon

I tut ('. micans has a keel that is lanate toward the
upper margin witli a slightly incurved beak and ca¬
ducous bracts and braeteoles.

Some specimens ol Crotalaria miottoae that oc¬
cur iu Rio dc Janeiro and Kspirito Santo states have
leaves digitately 3-, 4- or 5-foliolate and more
dense pubescence in branches, ovaries, and le¬
gumes.

Fa ratypes. BRAZIL. Espirito Santo: Domingos Mar¬
lins. I’edra Azul, MR 262 km 86, A. S. Flores 1074, R.
Sehiitz Rodrigues & t. F. Mansano (DEC). Minas Gerais:
Camanducaia, Monte Verde, A. S. Flores 6/9. R. Sehiitz
Rodrigues. L. I). Meireles &• R. Relinello (UEC); Sena do
Cip6, km 135, ea. 150 km N of Belo Horizonte, II. S.
Irwin. II. Maxwell & I). C. Wasshutisen 20534 (F, 145).
Rio de Janeiro: Nova Krihurgo, Pico Nova Caledonia, II.
C. de l.ima 2527, F. F. Rerrx. ./. F. Baumgratz & ./. C.
domes (E, IN PA, RB); Sena dos Organs. I). R. Haul 0471
(SP. LB); Teres6polis, Sena dos Organs, ea. 5 km S\\ of
city of Teresopolis. d. Fiteri <Sz L. T. Film 7174 (SP. LB).

Acknowledgments. We are grateful to Rodrigo

Sehiitz Rodrigues and Leonardo Dias Meireles for
kind help collecting Crotalaria species, Rodrigo
Sehiitz Rodrigues for assistance with the Latin di¬
agnosis. kavna Agostini (or improving the English,
and Rogerio Lupo for preparing the illustration. We
appreciate the helpful review offered by G. 1’. Lew¬
is. This work was supported by the Funda^ao de
Amparo a Pesquisa do Estado de Sao Paulo (FA-
PESP) proc. 00/1 1074-2.

I iterature Cited

Bishy, F. A. & R. M. Polliill. 1973. The role of taximetries
in angiosperm taxonomy II. Parallel taximetrie and or¬
thodox studies in Crotalaria. L. New Phytol. 72: 727—
742.

Polhill, R. M. 1981. Crotalarieae. Pp. 399-402 in R. M.
Polhill (X P. H. Raven (editors). Advances in Legume
Svstematics. Part 1. Royal Botanic Gardens, Kew.
- . 1982. Crotalaria in Africa and Madagascar. Roy¬
al Botanic Gardens, Kew.

Windier, I). R. & S. G. Skinner. 1982. New taxa and new
combinations in the American Crotalarias (Fabaceae).
Phytologia 50: 185—206.

A New Species of Styrax (Styracaceae) from Southern Mexico

Peter It' Fritsch

Department of Botany, California Academy of Sciences, 875 Howard Street, San Francisco,
California 94103, U.S.A. pfritsch@calacademy.org

ABSTRACT. Styrax uxpanapensis, a new species of
Styrax series Valvatae (Styracaceae) from the
Uxpanapa-Chimalapa region in the states of Oaxaca
and Veracruz, Mexico, is described and illustrated.
The stamen morphology of this lowland humid trop¬
ical forest species appears to ally it with two An¬
dean cloud forest species. A section of a key to
Mexican and Mesoameriean species of Styrax, as
modified to incorporate S. uxpanapensis, is includ¬
ed.

Key words: Mexico, Oaxaca, Styracaceae, Sty¬
rax, Veracruz.

Several specimens examined for the revision of
Styrax L. (Styracaceae) for western Texas, Mexico,
and Central America (Fritsch. 1997) were, for var¬
ious reasons, of uncertain placement. Among these
was a lowland collection in fruit from Municipio de
Hidalgotitlan, Uxpanapa region of Veracruz, Mexico
(2 Dec. 1981, T. Wendt 3576). I tentatively identi¬
fied this specimen as .S', panamensis Standley
(Fritsch, 1997). Because S. panamensis is known
only from a single collection in flower hud (Panama,
191 I. II. Pit tier 4242). however, the taxonomic sta¬
tus of tfie Veracruz collection remained unclear.

I have recently examined more material of Styrax
from the Uxpanapa region of Veracruz and adjacent
Chi ma la pa region in eastern Oaxaca and have iden¬
tified two more collections with clear similarity to
Wendt 3576. The combination of floral and fruit fea¬
tures present in these three collections clearly dis¬
tinguishes them from S. panamensis and all other
described Styrax species, such that the description
of a new species is warranted. Here I describe this
new species and incorporate it into a section of a
previously published (Fritsch, 1997) key to Mexi¬
can and Mesoameriean species of Styrax to aid
identification.

Slvrax uxpanapensis P. W. Fritsch, sp. nov.
TYPE: Mexico. Veracruz: Uxpanapa, Esfuerzo
Nuevo, al E, 1 7°10'15"N, 94°21'18"W, 225 m.
3 May 1996, ./. Rivera //. A S. Escobedo 73
(holotype, MEXU; isotypes, CAS, TEX). Fig¬
ure I .

Haec species S. argenteo similis, sed calyce 2-2.5 mm
longo, lobis corollae 7-9 X 1.2 — 2.0 nun, auriculis fila-
menti conspicuis, antheris 4—5 mm longis differt.

Evergreen tree to 23 m tall; young twigs grayish
brown-stellate-tomentose. Petiole 9—1 i mm long:
lamina 13-21.5 X 5. 4-8.2 cm, 2. 1-2.9 times as
long as wide, chartaceous, elliptic, secondary veins
7 to 9 on each side of midvein; apex acute to short-
acuminate; base cuneate to rounded, adaxially gla¬
brous except on midrib and secondary veins, these
slightly impressed; abaxiallv greenish gray to the
naked eye, with a dense, thin base tomentum of
minute greenish white stellate trichomes, and larger
scattered stiff tawny stellate trichomes (especially
prevalent on the veins) with arms to 0.3 mm long,
the base tomentum absent from the primary through
tertiary veins, the tawny to light orange surface of
the primary through tertiary veins easily visible
through the pubescence, the tertiary veins slightly
raised; margin entire. Inflorescences axillary, ter¬
minal, or produced above leal scars proximally
along the branch, racemose or paniculate, 1 to 3
per node, 3-4 cm long, 5- to 23-flowered. greenish
gray-stellate-tomentose with scattered orange to fer-
rugineous stellate trichomes, the lax to somewhat
stiff trichomes to 0.3 mm long; lower pedicels 3-4
mm long. Flowers hermaphroditic, 10-12 mm long;
calyx 2-2.5 X 3-4 mm, shallowly eupuliform,
densely gray ish green-stellate-tomentose. trichomes
at mid-calyx with arms to 0.2 mm long, margin con¬
cave to truncate between the minute teeth: corolla
8-10 mm long, white; tube 0.8-1. 2 mm long, ex¬
tending up to the calyx margin; lobes 5. 7-9 X 1.2-
2.0 mm, spreading, thick; stamens 10; free portion
of stamen tube 0.5 mm long: distinct portion ol fil¬
aments 0.8-1 mm long, ventrally with prominent
auricles bearing a dense mass of tawny-yellow stel¬
late trichomes with arms to 0.8 mm long, trichomes
nearest the proximal end of the auricle with arms
predominantly pointing upward, also often stellate-
pubescent in sinuses of filaments, dorsally gla¬
brous; anthers 4—5 mm long, I lit1 glabrous connec¬
tives slightly exceeding the linear, apically tapered,
short stellate-pubescent thecae; ovary densely gold¬
en orange-stellate-tomentose; style filiform, gla¬
brous; ovules many. Drupe 1.6— 2.3 X 0.9— 1.4 mm.

Novon 15: 421-424. Published on 21 September 2005.

422

Novon

Figure 1. Sty rax nxpanapensis P. W. Fritsch, new species. —A. Branch with distal inflorescences. — B. Leaf surface,
abaxial view. — C. Still stellate trichomes from the abaxial side of the leaf. — I). Branch with proximal inflorescences.
K. flower. f. flower, median long-section. — G. Calyx, long-section, and gynoecium. — H. Stamen, lateral view.
I. Stamen, ventral view. J. fruit. A— f based on Rivera & Escobedo 78 (CAS isotype); J based on Wendt et <d 3578
(MKXU).

Volume 15, Number 3
2005

Fritsch

Styrax uxpanapensis from Mexico

423

grayish green, ellipsoid, apex rounded; wall irreg¬
ularly and very coarsely rugose on herbarium spec¬
imens; fruiting calyx shallowly cleft-cupuliform to
discoid. 5-7 X 1-2 mm. 0.06-0.12 times the
length of the drupe proper.

Etymology. The specific epithet refers to the
Uxpanapa region in southeastern Veracruz and
eastern Oaxaca, from which the type collection of
the species was made.

Habitat , distribution, and phenology. Styrax
uxpanapensis is endemic to the Uxpanapa-Chima-
lapa region in extreme southeastern Veracruz and
eastern Oaxaca, Mexico, where it has been found
in secondary vegetation along arroyos and in eha-
parrera. a type of secondary rain forest (T. Wendt,
pers. comm.; information from specimen labels) at
225—250 m. It was collected in flower in May. and
in fruit in September.

The Uxpanapa-Chimalapa region comprises part
of an area of the Mexican Atlantic slope rain forest
that is rich in endemic species (Wendt. 1987,
1993). The geographic distribution of the available
collections of Styrax uxpanapensis suggests that it
is endemic to this region. Species of Styrax (e.g..
S. glabrescens Bentham, S. warscewiczii I’erk ins)
have been collected from surrounding areas in
Mexico and Central America, but S. uxpanapensis
has not been among these.

Styrax uxpanapensis belongs to Styrax sect. Val-
ratae Giirke series Valvatae Perkins, a clade of
about 78 species of Styrax endemic to the Neotrop¬
ics (Fritsch, 1999). Species of this series are easily
distinguished from other members of Styrax by the
combination of persistent leaves and a fleshy to
juicy mesocarp (Fritsch, 1999). Within this group,
the species is part of a clade that possesses robust
trichomes on the stamen filaments (generally 0.8—
2 mm vs. up to 0.7 mm long) and anther connec¬
tives that are prolonged beyond the anther sacs (vs.
not or only slightly exceeding the anther sacs; see
key below and Fritsch. 1999).

Other than Styrax peruvianus Zahlbruckner, S.
uxpanapensis is the only species of Styrax in Mex¬
ico and Mesoameriea that possesses prominent sta¬
men filament auricles. Other species of Styrax with
this feature are endemic to South America, mostly
in Andean cloud forests. Styrax peruvianus is not
likely to be the closest relative of S. uxpanapensis
because it has an indument of peltate scales instead
of stellate trichomes and linear-triangular (vs. lin¬
ear) anther sacs. Thus, unless prominent auricles
have arisen independently in S. uxpanapensis, the
phylogenetic affinity of this species appears to be
with the Andean cloud forest species S. subargen-

teus Sleumer or S. tornentosus Bonpland, or both.
Styrax uxpanapensis is similar to these species in
its stellate trichomes and linear anther sacs, but is
distinguishable from them by its larger leaves (13-
21.5 cm vs. 5.5—12 cm long) and smaller flowers
(10—12 mm vs. 13—20 mm long; Perkins. 1907;
Sleumer, 1936; P. Fritsch. pers. obs.). Furthermore,
S. uxpanapensis is a lowland species, whereas „S.
subargenteus and S. tornentosus are montane, f lo-
ristic affinities between the Uxpanapa region and
areas of South America have been noted by Wendt
(1993).

Paratypes. MEXICO. Oaxaca: Juchitan. Santa Maria
Chimalapa, 3 km W of San Antonio Nuevo Parafso, Plan
de la Ceiba, 21 Sep. 1997. E. Torres H.. J. Rivera II.. T.
Alvarez R. & I). Acuea V. 1347 (CAS, MENU). Veracruz:
llidalgotillan. hills SE of Poblado 6, 2 Dec. 1981, T.
Wendt. A Villalobos C. & I. Nava r ret e 3576 (CHAPA,
MEXU[2]).

Key to Mkxican and Mesoamkrican Species ok Styhax
with Anthkr Connectives Prolonged reyond hie
Anther Sacs

la. Calyx vestiture of stellate hairs.

2a. Calyx vestiture uniformly golden brown; lam¬
ina 1.5 times as long as wide . . . S. panamensis
2b. Calyx vestiture predominantly or completely
grayish green, occasionally with additional
scattered yellow or orange-brown trichomes;
lamina 2.1— 3.8(— 5.2) times as long as wide.

3a. Calyx 2—2.5 mm long; corolla lobes 7—

9 X 1.2— 2.0 mm; filament auricles con¬
spicuous; anthers 4—5 mm long . . .

. S. uxpanapensis

3b. Calyx 2.5—5 mm long; corolla lobes 8—

14 X 1 .7-4 mm; filament auricles in¬
conspicuous; anthers 5—8 mm long.

4a. Upward-pointing trichomes on the
filament auricles with at least some
arms 1—2 mm long; abaxial laminar
surface and outer calyx grayish
green-stellate-pubescent, without
scattered orange-brown stellate tri¬
chomes; larger drupes 13—18 mm
long (excluding beak, if present),
ellipsoid or ovoid-ellipsoid; corolla
lobes 1.7— 2.3 mm wide, widely

spreading or recurved .

. ,S. argenteus C. Presl

4b. Upward-pointing trichomes on the
filament auricles with arms to I mm
long; abaxial laminar surface and
outer calyx with scattered, orange-
brown, stellate trichomes in addi¬
tion to grayish green pubescence;
larger drupes (15— ) 17— 28 mm long,
ovoid or obovoid; corolla lobes 2.5—

4 mm wide, recurved or reflexed
. S. niearaguensis P. V . Fritsch

lb. Calyx vestiture of radiate, peltate, or laeerate-
margined scales.

5a. Calyx with vestiture of peltate or lacerate-

424

Novon

margined scales; scales at mid-calyx 0.27-

0.44 mm diam . .S', periwianus

5b. Calyx with vestiture of radiate scales; scales
at mid-calyx 0.05—0.43 mm diam.

6a. Abaxial laminar surface glabrous or
nearly so; calyx 4—7 turn long, in fruit
funnelform, 4-8 mm long; anthers 5-8
mm long; corolla lo!>es linear-deltoid . .

. .S’, glabratus Schott

6b. Abaxial laminar surface densely pubes¬
cent; calyx 2.5—3 mm long, in fruit shal¬
lowly eupuliform, to 1.5 mm long; an¬
thers to 4.5 mm long; corolla lobes
linear . .S', steyermarkii P. \V. Fritsch

Acknowledgments. I thank Tom Wendt for
bringing his collection of this species to my atten¬
tion and for insight into the habitat of the species;
Mario Sousa and the curators of MEXU for facili¬
tating examination of MEXU Styrax material; liana
Pazdirkova for the illustration; and Arne Antler-

berg, Jon Kicketson, and one anonymous reviewer

for helpful comments on the manuscript.

I .iterature Cited

Fritsch, P. W. 1997. A revision of Styrax (Styracaceae) for
western Texas, Mexico, and Mesoamerica. Ann. Missou¬
ri Hot. Gard. 84: 705—761.

- . 1999. Phylogeny ot Styrax based on morpholog¬
ical characters, with implications for biogeography and
infrageneric classification. Syst. Hot. 24: 356—378.

Perkins, J. 1907. Styracaceae. In A. Engler (editor). Das
Pflanzenreich 241 (IV.30Hd): 1-1 1 I.

Sleumer, H. 19.36. Vermischte Diagnosen IV. Repert.
Spec. Nov. Hegni Veg. 39: 117-128.

Wendt, T. 1987. I.as Selvas de Uxpanapa, Veracruz-Oa-
xaca, Mexico: Evidencia de refugios florfsticos cenozo-
icos. Anales Inst. Biol. UNAM 58 (1987). Ser. Hot.
(Num. unieo): 29—54.

- . 1993. Composition, floristic affinities, and origins

of the canopy tree flora of the Mexican Atlantic Slope
rain forests. Pp. 595—680 in T. P. Ramamoorthy, R. Rye,
A. l.ol & J. Fa (editors). Biological Diversity of Mexico:
Origins and Distribution. Oxford Univ. Press. New York.

A New Species of Bourreria (Ehretiaceae, Boraginales)

from Costa Rica

Marc Gottschling and James S. Miller

Missouri Botanical Garden. P.0. Box 299. St. Louis, Missouri 63166-0299, U.S.A.
caix@zedat.fu-berlin.de; james.miller@mobot.org

Abstract. We describe Bourreria grayumii (Eh¬
retiaceae) as new from Costa Rica, near the Nica¬
raguan border. It is illustrated, and a distribution
map is provided. The new species may be closely
related to Bourreria huanita, as both share schiz-
ocarpous fruits, an unusual and specialized char¬
acter in Bourreria.

Key words: Boraginales, Bourreria, Costa Rica,
Ehretiaceae.

Bourreria P. Browne comprises approximately 30
New World species and belongs to the Ehretiaceae
of a more broadly circumscribed Boraginales (Bor-
aginaceae s.l .), additionally including the subordi¬
nate taxa Boraginaceae s. sir., Heliotropiaceae,
Cordiaceae, and also Hvdrophyllaceae and Eennoa-
ceae (Thorne, 1983; Gottschling et ah, 2001). In
the course of examining specimens and preparing
the taxonomic treatments of the borages for the
Maraud de Plantas de Costa Brea and for the Horn
Mesoamericana, collections representing a previ¬
ously undescribed taxon were discovered. Former
authors assigned these collections to Bourreria mol¬
lis Standley (= Bourreria oxyphylla Standley),
which is common in Mexico. Belize, and Guatemala
(Nowicke. 1969; Gentry cN Janos, 1974; Miller.
1988) and also occurs in Honduras, Panama, and
Colombia. The present study shows the Nicaraguan
and Costa Rican collections that have been treated
as B. mollis (= B. oxyphylla) differ consistently in
several morphological features and are best consid¬
ered as the distinct new species below.

Bourreria grayumii Gottschling & J. S. Mill., sp.
nov. TYPE: Costa Rica. Canton de Upala,
along Rio Chimurria, vie. Colonia Puntarenas,
ca. 11 km (bv rd.) SE ol Upala, 80—100 m,
10°49'N, 84°53'W, 17 Nov. 1988 (II. fr), M. II.
Grayum, G. Herrera & H. Evans 904B (holo-
type, MO 4265358; isotypes, CR, E 21 17346,
MEXU, MO 4265359). Figures 1, 2.

Arbor vel frutex usque ad 12(-20) tu altus. Lamina gla¬
bra. elliptica, 1.5—10.9 cm longa, (0.4— )0.8-4.3 cm lata,
apice acuminata, hasi acuta; folia petiolo ca. 7 mm lougo,

Novon 15: 42

glabro. luflorescentia plerumque termiualis, thyrsoidea.
Flos fragrans; calyce (4 — )5 — 7 mm lougo. intus tomentoso;
corolla alba, iufundibuliformi, (10— ) 1 2 — I . mm longa, 5-
loba, tubo 7-10 mm lougo, lobulis ovatis, 3-6 mm longis;
staminibus 5, filamentis glabris, ad tubum corollae adna-
lis; stylo simplici, (4— )6— 8 mm lougo. Fructus immaturus
succulcntus, ca. 10 mm diam., maturitate schizocarpus.

Trees or shrubs, to 1 2( — 20 ) m tall; bark light
brown, rarely gray; branches glabrous; lenticels ab¬
sent. Leaves alternate, the blade 1.5—10.9 X
(0.4-)0.8-4.3 cm. elliptic to obovate. occasionally
asymmetric, the apex usually acuminate or apicu-
late. sometimes acute, retuse, or emarginate, the
base acute or cuneate, frequently asymmetric,
sometimes decurrent, the texture coriaceous to
membranaceous, the primary vein ± prominent, the
secondary veins (4 to)5 to 7(to 9). the tertiary veins
reticulate or rarely percurrent, occasionally incon¬
spicuous, the upper surface glabrous, rarely with
multicellular incrustrations, the lower surface gla¬
brous or with lew trichomes on the veins; petiole
0.2— 0.8(— 1 .2) cm long, slender, canaliculate on ad-
axial surface, glabrous or with few trichomes. luilo-
rescence (2— )3— 5(— 7) cm, terminal or sometimes
axillary, comprising up to 10(to 15) flowers, but
usually less, the branching sympodial, the branches
slender or rarely robust, glabrous to varying tomen-
tose, the bracts frequent, leafy, the braeteoles rare,
if present small, lanceolate, velutinous. Mowers fra¬
grant, the pedicels up to 3 mm, the buds widely
obovoid; calyx (4— )5 — 7 mm long, eampanulate or
sometimes tubular, coriaceous, indurate and slight¬
ly striate in fruit, varying puberulent outside, vary¬
ing tomentose at least at margin inside, the lobes
5, 1—3 mm long, aeute or acuminate, usually reg¬
ularly divided, but rarely irregularly (into 3 + 2 or
2 + 2+ I ) in fruit; corolla ( 1 0— ) 1 2— I 7 mm long, sal-
verform, membranaceous, white, the tube 7—10 mm
long, tubular or rarely funnel-shaped, usually gla¬
brous on both surfaces, the lobes 5. 3—6 mm long,
rounded or obtuse, papillose or with few strigose
trichomes outside; stamens 5. included, the fila¬
ments (6 — )7 — 9 mm long, completely adnate to the
corolla tube, glabrous, the anthers I (—2) mm long;
ovary glabrous, the style (4— )6— 8 mm long, includ-

5-428. PuttLiSHFd) on 21 Seitkmufr 2005.

426

Novon

Figure 1. Bourreria grayumii Gottschling & J. S. Mill. — A. Flowering branch. — B. Flower (note the papillose outer
surface of the corolla lobes and the strigose hairs on the outer surface of the corolla). — C. Open corolla with stamens
(note that the filaments are entirely adnate to the corolla tube). — 1). Open calyx with gynoecium. — K. Fruiting branch.
— F, G. External surface of pyrenes showing overlapping lamellae. A and F, G from Espinoza cl al. 213 (MO); I? from
Martinez 93 (MO); C, I) from Flores cl al. 46 (MO); F from Grayum cl al. 904H (MO).

ed. undivided, the stigma capitate, sometimes dis¬
tinctively broader than the style; disc present. Im¬
mature fruits 9-14 X 10-15 X 9-14 mm,
drupaceous, widely to depressed ovoid, ridged, the
exocarp thin, the mesocarp scanty, the mature fruit
schizocarpous; pyrenes 8-9 X 4 X 4—5 mm. each
one quarter sphere in shape, the abaxial surface
with 5 to 6 conspicuous lamellae.

Etymology. This species is named in honor of
Michael H. Grayum (MO), who collected the type
as well as many valuable plant specimens in Costa
Rica and has contributed greatly to our understand¬
ing of the Costa Rican flora.

Distribution. Bourreria grayumii occurs in
southern Nicaragua and northern Costa Rica, usu-

Volume 15, Number 3
2005

Gottschling & Miller

Bourreria grayumii from Costa Rica

427

Figure 2. Distribution of Bourreria grayumii at the N icaraguan— Costa Rican border.

ally at elevations of 40—100 m, but occasionally to
1 100 m, and is sympatric with B. costaricensis
(Standley) A. H. Gentry.

Notes. Bourreria grayumii has been considered
closely related to B. mollis from the Bourreria bac-
cala Rafinesque species complex comprising ap¬
proximately four difficult to distinguish species \B.
baccata, B. mollis, B. polyneura O. E. Schulz, and
B. virgata (Swartz) G. Don). They are similar in
general morphological appearance and have some
overlapping morphological features that are also
shared by B. grayumii at first glance. Fruiting col¬
lections of B. grayumii show that the mature fruit
is schizocarpous, and even its immature fruits have
both a ridged surface and a somewhat four-angled
shape. Schizocarpous fruits in Bourreria break into
four separate mericarpids at maturity, and each of
them has a filament connecting the gynohase with
its apex. This character appears to circumscribe a
monophyletic species group (Gottschling, 2004)
that occurs from Mexico to northern South America
and was recognized as Crematomia by Miers
(1869). Bourreria grayumii would be unique among
those species in exhibiting stamen filaments com¬
pletely adnate to the corolla tube.

Paralypes. NICARAGUA. Ilio San Juan: S;tbalu,
San Antonio, I km Vt of Sdhalo, B. !\ Moreno 26240 (MO.
US). COSTA RICA. Alajuela: G uatuso. Cabana, near Rio
Buena Vista, Q. Jimenez, R. Quesada & R. Castro 1 133
(CR, MO): Uos Chiles, Llanura de Guatuso, R.N.V.S. Cano
Negro, San Antonio, K. Flores el at. 46 (CR. FNCR, MO),
K. Flores A K. Flores 114 (FNCR. MO), K. Martinez HR
(CR. MO), K. Martinez 93 (CR. FNCR); Upala, Canalele,
Jimenez-Saa 125 (F). Guanacaste: Rarque National
Guanacaste, Cerro El Hacha. R. Espinoza 131 (CR, MO);
Fstacibn Maritza, Espinoza et al. 213 (CR. FNCR, MO).

Acknowledgments. We thank Roy Gereau (MO),
who reviewed the manuscript and corrected the
Latin description of Bourreria grayumii, the editor
Victoria Hollowell (MO) for improving a first draft
of the manuscript, and Barbara Alongi (MO) for the
excellent drawings. A portion of this work was sup¬
ported by a fellowship within the Postdoc-Pro¬
gramme of the German Academic Exchange Ser¬
vice (DAAD).

Literature Cited

Gentry, ,1. L., Jr. & D. Janos. 1974. A preliminary generic
key and geographic checklist of the Boraginaceae in
Central America and Panama. Phytologia 27: 445—455.
Gottschling, M. 2004. Floral ontogeny in Bourreria (Fh-
retiaceae, Boraginales). Flora 199: 409-423.

428

Novon

- , H. H. Hilger, M. Wolf <X N. Diane. 2001. Sec¬
ondary structure of the ITS] transcript and its appli¬
cation in a reconstruction of the phytogeny of Boragin-
ales. PI. Biol, 3: 629-636.

Miers, J. 1869. XVII. — On the Ehretiaceae: Crematomia.
Ann. Mag. Nat. Hist. (Ser. 4) 3: 300—313.

Miller. J. S. 1988. A revised treatment of Boragina-

ceae for Panama. Ann. Missouri Hot. (lard. 75: 456—
521.

Nowicke. ,|. W. 1969. Part IX. Family 1 07. Boraginaeeae.
In: K. E. Woodoson & B. W. Scliery, Flora of Panama.
Ann. Missouri Bot. Hard. 56: 33—69.

Thorne, K. F. 1983. Proposed new realignments in the
angiosperms. Nordic J. Bot. 3: 85-1 17.

Studies in Capparaceae XXII. Capparis sclerophylla , a Novelty
from Arid Coastal Peru and Ecuador

Hugh II. litis

Department of Botany, University <>l Wisconsin, 430 Lincoln Drive, Madison,
Wisconsin 53706, U.S.A. tscochra@facstaff.wisc.edu

Xavier Cornejo

Herbario, Universidad de Guayaquil, Av. 25 dc Julio via al Pto. Mantimo. Casilla 09-01-
10634, Guayaquil, Ecuador, xcornejoguay@hotmail.com. Present address: Department of
Botany, University of Wisconsin, Birge Hall. 430 Lincoln Drive, Madison,
Wisconsin 53706-1381. U.S.A.

Alls’ll! ACT. Capparis sclerophylla (subg. Cynophal-
la). Capparaceae, a sharply distinct new species of
shrub or small tree, differs from other, often domi¬
nant. related, but often difficult to distinguish spe¬
cies of subgenus Cynophalla by thick, hard, still,
xerophyllous leaves, these subsessile on short (1—4
mm), dark-brown to black, rigid and fragile petioles:
by large white flowers spirally disposed with reflexed
petals usually borne on thick, congested, short, and
rigid raceme axes: by long-gynophored. usually slen¬
der siliques with white pulp, white seed arils and
green embryos; and by its restriction to arid decid¬
uous thornscrub near the Pacific Ocean from north¬
western-most Peru to west-central Ecuador. Its clos¬
est relative appears to be C. verrucosa of Mexico to
Venezuela. A cytotaxonomic survey of subgenus Cy¬
nophalla proves it to be a polyploid series, with base
number of n = 14, 2 n = 28 (x = 14. 28. 56, 84),
represented by C.flexuosa and other species; several
of these chromosome numbers are reported here for
the first time, but the number for C. sclerophllya is
as yet unknown.

Rksi MKN. Capparis sclerophylla (subg. Cynophal¬
la), Capparaceae, es una nueva y claramente dis-
tinta especie de arbusto o arbolillo que difiere de
las otras especies del subg. Cynophalla, freeuen-
temente dominantes, relacionadas, pero usualmente
dificiles de distinguir. por poseer hojas xerdfilas.
rfgidas, gruesas, subsesiles, con pecfolos muy cur¬
ios 1 1—4 mm), n'gidos. fragiles, de color cafe-obscu-
ro a negro; grandes (lores blancas con petalos re-
flexos que nacen en los ejes gruesos. cortos. rigidos
y sesiles de los racimos congestos; silicuas delga-
das con ginoforos largos, pulpa y arilo de la semilla
blancos v embriones verdes; y por estar restringida
al monte espinoso y bosque muy seco deciduo, cer-
ca del Oceano Pacffico, en la costa del norocciden-

te de Peru hasta el oeste-central de Ecuador. Su
pariente mas cercano parece ser C. verrucosa, dis-
tribuida desde Mexico hasta Venezuela. I n estudio
citotaxondmico del subg. Cynophalla demuestra
<|ue es una serie poliploitle, con un numero base
den = 14. 2 n = 28 (x = 14, 28. 56, 84), repre-
sentada por C. flexuosa y otras especies; se presenta
algunos de estos numeros cromosdmicos aqui por
primera vez. pero el numero de C. sclerophylla es
aun desconoeido.

Key words: Capparaceae, Capparis. chromo¬
some number, coastal Peru and Ecuador, endemic,
extra-floral nectaries, myrmecotrophic. subgenus
Cynophalla, xerophyte.

As a consequence of the dramatic rainshadow
created by the upwelling cold waters of the Peru
(or Humboldt) Current precipitating the humid. Pa¬
cific “westerlies," the coastal regions of western
Peru and Ecuador south of the Equator are covered
by a widespread arid belt of xerophytic vegetation:
mostly deciduous thornscrub with columnar ceroid
cacti, semideciduous woodlands and seasonally dry
tropical forests, in which various species of Cap¬
paris are not only common, but characteristic and
often dominant. Here one finds such highly distinct,
stellate- or lepidote-pubescent species as C. sca-
brida Kuntli (C. angulata Ruiz A Pavdn ex DC.),
C. crotonoides kunth (C. cordata Ruiz A Pavdn ex
DC.), and C. avicennifolia Kunth (C. ovalifolia Ruiz
A Pavdn ex DC.), as well as many species of the
glabrous or barelv pilose taxonomically difficult
Capparis subg. Cynophalla (DC.) Eiehler (Eichler,
1865), this nevertheless a well-defined group char¬
acterized by: (1) distichous and divergent, evenly
spaced out, subsessile leaves; (2) each leaf sub¬
tending an ascending series of one to three, char-

Novon 15: 429-437. Published on 21 September 2005.

430

Novon

acteristic axillary, nectar-producing glands (Pool-
sen, 1879—1880; l)i Sapio et al., 2001) visited by
ants; (3) large, round, cup-shaped sepals in two un¬
equal decussate series, with the inner the larger;
(4) elongate, torulose, dehiscent siliques, each with
several to many seeds embedded in a fleshy orange,
red, or white pulp (endocarp); (5) seeds covered by
a paper-thin testa, and this in turn enclosed by a
white aril, with both together readily slipping off
the naked green embryo; and (6) plants that are
mostly glabrous or at best only minutely puberulent
or pilose with simple short, soft trichomes. Included
here are two closely related, rather thin-leaved spe¬
cies, C. heterophylla Ruiz & Pavdn ex DC. and C.
ecuadorica litis (litis, 1978). In addition there are
several taxa ol a notoriously variable complex of
intergrading taxa from western Ecuador (the poly¬
ploid C. flexuosa (I,.) I,., s.l. complex; cf. litis, be¬
low and 1978; Madsen et al., 2001). the nomencla¬
ture ol which in many regions has yet to be worked
out: the infrequent C. didymobotrys Ruiz & Pavdn
ex DC.; the ubiquitous and enormously variable C.
guayaquilensis kunth and the usually more south¬
ern. less common variant of the latter; C. mollis
Kunth; and their diverse segregating progenies of
some putative, presumably ancient, hybridization
events.

Intermixed and hidden in all this impossible di¬
versity within Capparis subg. Cynophalla was an
unknown, usually a sterile member of Cynophalla ,
superficially resembling in leaf shape Capparis gua-
yaquilensis ( = Capparis lanceolata Ruiz & Pavdn
ex DC., and so cited by Madsen et al., 2001: 153—
155), one that heretofore had escaped notice and is
herewith newly described as Capparis sclerophylla.
A closer examination revealed that its leaves are
remarkably hard, stiff, and evergreen, and are sub-
sessile, when dry on dark brown or black, short,
thick, rigid, fragile petioles. As a consequence, its
leaf blades do not hang loosely down as in some
other, sympatrie species in this subgenus, but di¬
verge laterally and break off when dry at the slight¬
est touch. Most significantly, its large, pure white
flowers with reflexed petals are usually borne on
thick, congested, stubby short shoots or spurs (i.e.,
peduncles) mostly no more than a centimeter long,
these produced at the end and sides of older
branches and unique in subgenus Cynophalla. Oc¬
casionally, flowers are spaced out on young leafless
long shoots more typical of the subgenus. Finally,
the scarce pulpy endocarp is said to be white rather
than red, a condition duplicated only in the wide¬
spread C. verrucosa Jacquin ol Mexico to Venezue¬
la, for this and other reasons the closest relative of
C. sclerophylla. Abundantly distinct, it is thus grat¬

ifying to be able to segregate still another species
out of the polyploid taxonomic jungle that is the
Capparis subg. Cynophalla (litis, 1978; Rufz-Za-
pata & litis, 1998: 139; litis, unpublished).

Capparis sclerophylla is relatively uncommon,
flowers infrequently, and only rarely sets fruit.
When sterile specimens were collected by the first
author near Bah fa de Caraquez, these were initially
taken to be aged, xeromorphie extremes of a vari¬
able population of the similar C. guayaquilensis, a
much more common species with which it grew in¬
termixed. But once the taxon is recognized, the pet-
iolar characters alone readily distinguish it. Finally,
the recent splendid fertile collections of Ecuado¬
rean capparids, sent to the University of Wisconsin
Herbarium (W1S) from the rejuvenated Universidad
de Guayaquil Herbarium (GUAY) by Carmen Bon-
ilaz and the second author, have made it obvious
that this taxon deserves a binomial of its own. Fo¬
cusing on its distinctive sclerophyllous leaves, we
name it:

Capparis sclerophylla litis & X. Cornejo, sp. nov.
TYPE: Ecuador. Manabf: Bahia de Caraquez a
8 km, alrededores de Univ. Catdlica, 80°32'W,
00°45'S, monte espinoso/bosque muy seco
tropical deciduo, fuertemente intervenido, 30
m, 27 Oct. 1997 (fl, fr), X. Cornejo & C. Bon-
ifaz 5822 (holotype, WIS; isotypes, AAU. B.
COL, DAV, F, GB, (ill, GUAY, K*, LOJA, MA,
MEXU, MO*, MY, NY*, P. QCA, QCNE*,
RSA*. SBT, UC, US*, USM, VEN, WIS [3].
[Some specimens incomplete, but all supple¬
mented by a suite of xerox reproductions of
(lowering and/or fruiting isotypes; specimens
with an asterisk (*) include a wood sample.]
Figures 1-3.

Fruticosa 0.5—5 m alta, C. guayaquilensi similis, sed
foliis insigniter diiris, petiofis atrofuscis vel nigris, bre-
viorihus (2-3 mm longis) et fragilibus in sicco, floribus
grandibus (20—30 mm longis) cum petalis reflexis et axi-
bus floriferis brevibus, densis et condensatis differt.

Xeromorphie shrubs to small slender trees, 0.5
to 5 m tall and 10 cm DBH, with many leaves
evenly spaced on the current year and several pre¬
vious years’ growth, somewhat distichous to mostly
irregularly stiffly diverging in all directions but
never all drooping together, the branchlets lent i-
eellate; plants glabrous when mature, but young
branches, petioles, and leaves sometimes with a
fine pubescence of very short, simple trichomes.
Leaves evergreen, thick, hard, stiff, entire, narrowly
oblong to elliptic, less often linear-lanceolate to ob-
lanceolate or obovate, highly variable in size and
shape (e.g., in one extreme case, 8 X 1 .4 cm leaves

Volume 15, Number 3
2005

litis & Cornejo

Studies in Capparaceae XXII

431

W 80° 78° 76°

Figure 1. Capparis sclerophylla. Geographic distribution from northwesternmost Peru and western Ecuador.

adjoining some 23 X 5.5 cm), those on sprout-
shoots often narrowly to broadly linear, (5— )7— 9(—
15) X (1— )2— 3(— 4) cm or even 20 X 1 cm, dark
olive-green above, very light |>ale green beneath,
glabrous when mature, or occasionally with a soft
short dense puberulence on midrib and blade be¬
neath but barely so above, the apex acute to obtuse,
rarely rounded or emarginate, apieulate, the base
subsessile, cuneate to rounded or subcordate (i.e.,
viewed from above, subpeltate, with leaf lobe mar¬
gins overlapping and almost confluent below the
petiole insertion); lateral veins 7 to 12 on each side
of the midrib, somewhat elevated on both sides, the
midrib beneath strongly elevated, yellow, changing
abruptly into the dark brown to black petiole (rarely

green, at least in the herbarium); petioles very
short, (1— )2— 3(— 4) mm, when fresh with 20 or more
deep transverse fissures, when dry extremely brittle
and readily breaking from the branch at the slight¬
est touch (hence many if not most herbarium spec¬
imens leafless), subtended by paired, thin, trian¬
gular, apieulate, ± caducous stipules ca. 1-2 mm
wide; glands above the leaf axils on very young
branches l(to 3), increasing in size upward in a
vertical row, sessile, subspherical, 1—2 mm diam.,
composed of 2 pairs ol decussate, fleshy bracts (the
outer larger) enclosing a brown, nectar-producing
center, subtended by a pair of round-apiculate, pale
and marcescent stipules, on the older, pale gray to
pale brown branches often solitary and soon becom-

432

Novon

figure 2. Capparis sclerophylla litis & X. Cornejo. — A. Branch with flower buds on short leafless terminal long shoot.
— B. Detail, with flowers on condensed short-shoot raceme axis or peduncle. — C. Condensed short-shoot peduncle
showing .1 terminal pedicels and many old spirally disposed pedicel bases with associated stipules and bracts. — ]).
Open (lower w i 1 1 1 stamens fallen off one day after anthesis. — E. Leafy, elongate, long-shoot branch bearing four f ruits
(only two shown). — F. Leaves showing short black petioles: three drawings (left) and photos of two paired leaves (right),
left of each pair showing underside, right of each pair, upper surface with subpeltate insertion. — G. Leaf bases, with
underside (right), and upper surface (left) with subpeltate insertion. — LI. Close-up photo of 2 leaf nodes of very young
shoot, each with a stipulate petiole, its leaf base, axillary bud, and extra-floral nectary. (A— H. from the type, Cornejo
& Bonifaz SH22.)

Volume 15, Number 3
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litis & Cornejo

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Figure 3. Capparis sclerophylla. — A, C. Habitat, ea. 5 km ESE of Bahfa de Caraquez, at Silio Mauricio, close to the
type locality in Ecuador. View of vegetation with Pilosocereus tweedianus (Britton & Hose) Bytes & G. I). Bowley
(Caetaceae), the smooth hark of Cochlospermam vitifolium (Willdenow) Sprengel (Bixaceae), Ceiba trichistandra (A.
Gray) Bakhuizen (Bomhacaceae, shown in center of A), and leafy branch of C. sclerophylla (lower center of E). — B.
Flowers on short-shoot raceme axis. (Photos: A, C, litis; B, Cornejo.)

434

Novon

ing obsolete and barely recognizable as small, erod¬
ed axillary pits. Racemes terminal, subterminal
and/or lateral, mostly on old branches, their axes
(peduncles) of two sorts: most commonly (I) lateral
or subterminal short shoots or spurs on old wood of
leafy branches, ± highly condensed and congested,
thick, stubby and sessile* (never slenderly zig-zag),
5-1 I (-22) mm long, 2—4 mm thick, green when
young or, if old, grayish and scruffy or scaly and
covered with paired, round to triangular, apiculate
stipules and bract surrounding (3 to)6 to 12 or more
old pedicel scars or aborted (lower buds; when pe¬
duncles lateral, then solitary or in twos or threes,
when terminal or subterminal, solitary or in twos to
fives, these often then tightly clustered on the old
wood, each bearing up to 3 to 6 or more flower
buds, the larger peduncle clusters at anthesis with
as many as 25 flower buds arranged in a compound
umbelliform false “panicle,” but with oidy a single
flower of each individual peduncle open at one
time; or finally, less commonly and sometimes to¬
gether w ith, but exceeding in length, the above con¬
dition: (2) slender and uncongested, solitary, leaf¬
less new-growtli (long shoots) terminating new leafy
branches or old leafy branches, in either ease the
peduncles (raceme axes) slender and smooth, un¬
condensed and mostly 20-50 mm long, ca. 2 mm
thick, and ending in ca. 8 to 14 flower buds or
flowers and/or pedicel scars arranged distiehously
as in a fan, and in addition, each with 3 to 0 ax¬
illary nectariferous glands (see above) clustered at
or near the peduncle base, these not subtended by
pedicels but only by minute, paired, pale, scarious,
round to triangular stipules. Flowers showy, white,
spirally disposed on ascending pedicels (14— )20—
30(— 44) mm long; calyx of 4 sepals in 2 unequal
decussate-imbricate series, the smaller, outer ones

4— 6 X 6—8 mm, the larger, inner 7-8 X 9-10 mm,
green on tin* back, with thin, hyaline-papery, trans¬
lucent margins, strongly copulate, in bud nearly
completely covering corolla, but just before anthe¬
sis tightly covering only V2 to Vr, corolla buds at
anthesis obovoid, ca. 15 mm diam., glabrous; re¬
ceptacle obconie, lopped by a circular fleshy disk

5— 6 mm diam., its flat top with I nectar-producing
depressions, the nectar very sweet. Corolla cruci¬
form; petals pure white within, light green at base
without, ovate to oblong, 23—29 X 9-17 111m, with
rounded apex, the base broadly euneate to rounded,
subsessile and inserted into the disk by a short peg.
the open flower ca. 15-20 mm across, with the up¬
per half of the petals at nightfall becoming sharply
reflexed over, and almost to the base of, the sepals;
androgynophore absent, the androecium base form¬
ing a ring around the base of the gynophore in the

center of the disk: stamens 40 to 70, filaments pure
white, glabrous to the very base, subequal, 45—70
mm long, in bud spirally disposed around tin* style,
at anthesis in a pincushion funnel with ca. 90° di¬
vergence; anthers 2—3 mm long; gynophore white,
becoming dark purple with maturity, 45—65 mm
long; ovary green, narrowly oblongoid, 5-7 mm
long, with a hemispherical stigma, the gynophore
and ovary exceeding the stamens. Siliques I to 4
(rarely to 6) per raceme, usually densely clustered
on the short shoots (or more distant on the more
uncommon slender terminal peduncles), linear-cy-
lindric, 6— 14(— 20) X 0.8— 1.2(— 1.5) cm, smooth and
glabrous, usually ± tondose (constricted around
seeds, but see below), yellow-brown to purple, or
green streaked with dark red to purple, when ma¬
ture each continuous with its slightly curved, slen¬
der to stout, 6—9 cm long gynophore and its (1.3—
)2-3(-4) cm long pedicel, with the 2-4 mm long,
6— 8 mm wide receptacle in between; seeds mostly
(2 — )6 — 1 2( — 1 4). usually single-rowed or occasionally
2- or 3- or rarely 4-rowed and square in cross sec¬
tion, gigantic to 20 X 1.5 cm, ca. 60 [Cornejo &
Bonifaz 7154 (WIS)|; seeds (5 — 1 5 X ca. 5—7 mm,
the testa paper thin, dark brown, surrounded by a
white, oily aril, both together (unlike seeds of all
other species in subg. Cynophalla, except C. ver¬
rucosa) embedded in the apparently white pulpy
endocarp, with aril and testa together presumably
slipping oil easily under pressure exposing the na¬
ked, green, ready to germinate embryo; seeds some¬
times hanging on for a short period by a slender
thread from the thin, valve-embedded replum; em¬
bryo slightly curved, but essentially straight.

Distribution and habitat. Northwesternmost
Peru ( lu mbes) to northwestern coastal Ecuador
(Provinces of Loja, El Oro, Guayas, and Manabf as
far north as “El Recreo [(just south of Canoa and
north of Bahia de Caraquez), a beach house labeled
“El Recreo” on the second floor and still standing
in 1977, made famous by the Danish plant explorer
Baron von Eggers (Eggers, 1894: litis. 1978)], in a
variety of xerophytic, super-dry coastal vegetation
types, from deciduous thornscrub or monte espi-
noso, to semi -evergreen dry forest (bosque muy
seco— seco), savanna or bushwood, to tropical de¬
ciduous forest, from sea level to 600 m.

“The habitat of Cornejo & Bonifaz 5822 (the type
collection) is a very disturbed, open, sunny, very
dry tropical forest, with dense scrubby shrubs less
than 2 m high and very few scattered trees, in
which C. sclerophylla is a frequent shrub associated
with Cordia lutea [Lamarck], Mimosa debilis var.
aequatoriana [(Rudd) Barneby], Capparis crotono-

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litis & Cornejo

Studies in Capparaceae XXII

435

ides, C. scabrida, C. heterophylla, C. guayaquilen-
sis, C. didymobotrys, Cucurbita ecuadorensis [Cutler
& Whitaker], Ipomoea nil [(L.) Roth], /. hederifolia
| L. ], and Jacquemontia [Choisy] sp. In other small
remnant forests nearby, mature individuals of C.
sclerophylla ranged from shrubs to 4—5 rn tall and
10 cm dbh treelets, that are associated with Tabe-
buia chrysantha | Nicholson |. Caesalpinia glabrata
|Kunth], Achatocarpus pubescens [C. 11. Wright],
Bursera graveolens [Triana & Planchon], Eriotheca
ruizii |(K. Schumann) A. Robyns], Geoffroea spinosa
[Jacquin], ami Bonellia sprucei [(Mez) Stahl & Kal-
lersjo]” (letter to litis from Cornejo & Bonifaz, 5
Nov. 1997).

Phenology. Flowering from October into March
(May), fruiting mostly from (October) December to
April.

Local names. Fresno (Spanish), Vivar et <d. 883
(Loja); Patacon (Spanish), litis 168 (Manabf); Se¬
bastian (Spanish), fit is 168 (Manabf), Madsen
6)3781. 63782 (Guayas); Vaina morada (Spanish),
Hernandez et al. 222 (Manabf).

local uses. For construction, firewood, and
charcoal (Madsen et al.. 2001).

Discussion. Capparis sclerophylla is vegetative-
ly easily recognized by the thick, stiff, variously
divergent, dark olive-green leaves on short (1—4
mm) dark brown to black petioles. In other respects
it is rather similar to various forms of C. guaya-
quilensis, but the latter have relatively thinner and
more flexible leaves that on drying become dark
reddish brown or mahogany, and are borne on more
flexible, slender petioles 6—10 mm long, from which
the leaves usually hang down loosely, on either side
of the branch. When in flower or fruit, or leafless,
the densely congested, short-shoot peduncles, 2—4
mm thick and usually less than 20 mm long, im¬
mediately identify C. sclerophylla, while most oth¬
ers in subgenus Cynophalla have slender, flexible,
often pronounced flexuous (zig-zag) raceme axes on
which the flowers or flower buds are well-spaced
out and 5—10 mm apart. Like all species of Cyno¬
phalla, C. sclerophylla is characterized by 1 to 3
cauline “glands” or extra-floral nectaries (Poulsen,
1879—1880) in the axil of leaves and/or at the base
of the leafless terminal raceme axis (see above),
which together with the young leaves are visited by
vast numbers of ants of at least three taxa (Cornejo,
pers. obs.).

Among its close relatives, Capparis sclerophylla
is morphologically the most highly specialized xe-
rophyte, judging from the thick, evergreen leaves,
the congested short-shoot peduncles and the short

petioles, the latter shared only hy another mostly
coastal species, Capparis verrucosa, widespread
from western Mexico to coastal Venezuela. This
species not only shares with C. sclerophylla the
strongly recurved petals of the open flower, but also
the color of the fruit pulp, which in both species is
apparently white rather than the typical bright or¬
ange, red, or purple of all the other Cynophalla
species. Whether this white pulp reflects an adap¬
tation to bat seed dispersal during the night is not
known, but deserves investigation.

The unique relationship of these two clear-cut
species, Capparis sclerophylla and verrucosa, is
strengthened by an easily overlooked floral char¬
acter, the presence or absence of pubescence al the
base of the staminal filaments. While most of the
taxa of the Capparis flexuosa s.l. complex (e.g., C.
flexuosa s. str., C. polyantha Triana & Planchon, C.
declinata Vellozo, C. mollis, C. guayaquilensis, C.
didymobotrys, C. tarapotensis Eichler, C. retusa G rise-
bach, etc.) have a dense, short, glandular pubes¬
cence covering the basal */- or so of the staminal
filaments, C. sclerophylla and C. verrucosa lack this
peculiarity, what with their filaments being com¬
pletely glabrous to the very base.

There are two exceptions within the Capparis
flexuosa s.l. complex. First of all, the condition is
variable in C. hastata Jacquin f. coccolobifolia
(Martius) Dugand & litis, a relative of C. flexuosa
s. str. that appears to hybridize with it (litis, 1991).
with staminal filaments pubescent in ca. one-fifth
of the specimens and totally glabrous in the rest.
Capparis hastata is apparently the only species in
the C. flexuosa group that regularly has glandular
pubescent ovaries and gynophores. Second, and
more interesting, is the lack of filament pubescence
in a closely related, but quite distinct, species pair
of subgenus Cynophalla: C. heterophylla and C.
ecuadorica (litis, 1978). Though related to subge¬
nus Cynophalla, they do not belong with the C.
flexuosa s. str. alliance, and their relationship to C.
sclerophylla and C. verrucosa is rather distant. It
may be worth mentioning, however, that much like
C. sclerophylla, both C. heterophylla and C. ecua¬
dorica are also restricted to the lowlands on the
western side of the Andes and nearly endemic to
Ecuador.

Finally, similar in leaf shape is Capparis flexuosa
s. sir., a shrub widespread from Florida, the Carib¬
bean and Central America to Venezuela and the
Brazilian Islas de Noronha in the Atlantic Ocean.
While the leaves of C. flexuosa are generally quite
flexible, the fruit shape resembles that of C. scler¬
ophylla. It is of interest that C. flexuosa is also char¬
acterized by a few large flowers at the end of its

436

No von

branches and weak, lianoid "leaning” stems, which
may also occur in C. sclerophylla, with one of its
collections {Plowman el al. 4610) even called a
“vine.” Considering that the habitat of C. sclero¬
phylla is often heavily disturbed by the gathering
of firewood, animal grazing, or forest clearing
through burning, we also have often seen its flexible
slender branches sprawling like a vine over nearby
vegetation (Cornejo, pers. obs.).

The relative rarity of Capparis sclerophylla, and
the infrequency ol its flowering or fruiting, may be
related to such disturbance. Furthermore, the sec¬
ond author has often observed its firewood-trimmed
shrubs in secondary vegetation overwhelmed by
dense layers of vines, often of the endemic Cucur-
hita ecuadorensis, which, whenever such nest-like
tangles were lifted off from these bushes, revealed
tin- presence ol usually sterile specimens ol our
new species underneath. Evidently, the treelet
growth form is more to its reproductive optimum.

Biological interactions. The flowers ol Capparis
sclerophylla are visited by bees and ants in the ear¬
ly evening (Cornejo, pers. obs., 6:05 p.m.— 6:30
pan.) and also by bumblebee nectar thieves that do
not penetrate the flower bill bite a hole al the base
ol tbe flower to get al the nectar. The flowers pro¬
duce a very sweet nectar by 7:30 p.m.. becoming
fragrant around 8:00 pan., and stay open all night.
Next day. once the sepals, petals, and stamens have
fallen, the nectaries, as well as the axillary glands,
are extensively visited by ants (Cornejo, pers. obs.).
The pollinators are unknown, but probably include
bats.

Cytotaxonorny of Capparis subg. Cynophal-
la. Clear evidence for hybridization or introgres-
sion with other species is lacking, even though C.
sclerophylla quite frequently occurs together with
one or more species ol tin- subgenus. Some species
groups within subgenus Cynophalla appear to be
part ol a polyploid series, and others are homoploid.
Herein may lie the cause ol both the distinctness
of C. sclerophylla. whose chromosome number is as
yet unknown, and the taxonomic problems due to
putative hybridization between the closely related
C. guayatjuilensis and C. mollis of the Pacific slope
ol Ecuador and Peru. Despite Capparis chromo¬
somes being very small and difficult to study,
counts by Batia Pazy (in litis. 1678. vouchered in
A IS) for C. ecuadorica indicated ea. n — 25—26
(probably 28. i.e.. a tetraploid 2n = 56, based on
litis 106) and for the related C. heterophylla, ea. 2 n
= 70—80? (perhaps a hexaploid 2n = 84. based on
litis 207). these two an isolated species pair essen¬
tially endemic to Ecuador.

A similar species pair of Capparis subg. Cyno¬
phalla from Pernambuco in eastern Brazil, both
part of the Capparis flexuosa complex, revealed a
plant with fruits similar to those of C. hastata f.
coccolobifolia (listed in Costa e Silva, 1995, as "C.
cynophallophora") with a chromosome count of 2 n
= 28, while another sympatric species within the
C. flexuosa complex from eastern Brazil, C. decli-
nata (listed as “C. flexuosa"), had a tetraploid count
of 2 n = 56 chromosomes (Costa e Silva, 1995: 58-
61). Finally, n — 14 has been reported for C. flex¬
uosa s. str. from Dominica (Ernst in litis, 1991).
Previously unreported are the following additional
n — 14 counts by Batia Pazy for species of sub¬
genus Cynophalla (litis et al.. 1985. unpublished,
all vouchered in W IS): C. flexuosa s. sir., Jalisco,
Mexico ( litis 20105): C. declinata (C. flexuosa s.l.).
Bahia. Brazil (Mori 1 1241): C. guayaquilensis,
Guayas, Ecuador (litis E-10 ); and C. verrucosa,
Chamela, Jalisco, Mexico (Bullock s.n.. counted by
S. Bullock, pers. comm, to litis). These chromosome
numbers suggest genetic isolation of a putative dip¬
loid C. sclerophylla as supported by the morpholog¬
ical uniformity of C. sclerophylla, in contrast to the
incredible intergrading variability observed among
other sympatric taxa.

Paratypes. PEKl . Tuiubes: Klistenhalbwuste und
Trockenwald-region, Tonnenbaum Mischwald \Ceiba tri-
chistandra mixed forest], Cienego, just along coast S of
Tumbes. II. Ellenberg 1408 (U, GUA\ photo, \\ IS); Zar-
umilla, pampas de Hospital, entre El Caueho y Condor
Flores, C. Diaz <& R. Vdzquez 5044 (MO, A IS fragm.);
Zarumilla, region below El Caueho, T. Plowman 5489
(GH, WIS photocopy). ECUADOR. Loja: In sector algo-
donal, entre Maeard y Zapotillo, P. Uizano & 0. Davis 702
(LOJA); Zapotillo. La Ceiba, P. Vivar 888 (LOJA). El
Oro: near Santa Rosa [fide catalogue], l\ Ishrnann 7649
(k. \\ IS); 18 km S\\ of Santa Rosa, C. Johnson 1288 (MO,
WIS); rd. from Arenillas to Huaquillas, T. Plowman. L.
Jacobs & E. Davis 4610 (GH, S. WIS); 1 km NW of Ar¬
enillas on rd. to Huaquillas, L Albert de Escobar 705
((,)CA. TEX, W IS); 2 km al sur de Chacras eerea a la
entrada a Ralsalilo en la earretera de la Reserva Militar
Arenillas, //. Vargas, C. Canaday & R. Miranda 1171
(MO, QCNE, WIS). Guayas: Capeira. km 21 on rd. Gua¬
yaquil to Panic. Dodson & t. Gentry 12619 (MO, SEL,
A IS); Finca La Gloria, km 53 earr. Guayaquil— Playas, X.
Cornejo A’ II. Grochowski 7797 (GUAY, A IS); Guayaquil—
Salinas rd., 1-5 km E of El Progreso, G. Ilarling & L
Andersson 21 1 18. 21129 (GB. W IS photocopy); G. Barling
& L. Andersson 21105 (tilt. (JCA); km 20 Guayaquil— Sa¬
linas rd. G. Ilarling A’ A. Andersson 21041 (GR, QUA,
A IS); Playas, Guayaquil, G. Sanchez 2. 8 (A IS); km 17
de la vfa Playas— Posorja, G. Bonifaz 8689 (GUAY, A IS);
Chanduy prop** Guayaquil, [Feb.— Aug. I8(>2|. R. Spruce
s.n. (k. GUAY photo, A IS); Isla Puna, trail from Puna
Nueva to Concordia. ./. Madsen. E. Madsen & E. t lantern
85867 (AAU. OCA. A IS photo); vie. Puna Nueva tow.
Zapote. ,/. Madsen 64106 (AAU, A IS photo); El Placer.
0—5 km tow. Rfo Hondo, ./. Madsen 68942 (AAU. A IS

Volume 15, Number 3
2005

litis & Cornejo

Studies in Capparaceae XXII

437

photo); path from Campo Alegre to Midway Rio Hondo../.
Madsen 63781 , 63782 (AAll. WIS photo). Maiialn: Cuen¬
ca del Rfo Ayampe, Cerro Cabeza de Y'aca. X. Cornejo &
C. Bonifaz 818 (GUAY, W IS); Barbasmonte, X. Cornejo X
C. Bonifaz 1027 (GUAY. QCNE, WIS); Portoviejo-Jipijapa
rd„ I km S of La Pila, G. Hurling X /.. Andersson 21821
(GB. WIS photocopy); id. from Colimes de Pajan to Jipi-
japa. C. Dodson X I,. Tliien 1603 (WIS); Cerro Monte-
cristi. 2(H) m, X. Cornejo X (.. Bonifaz 5917 (GUAY, W IS);
N of Manta, 2 km from Monteeristi, C. Dodson X L. I liien
1750 (UC, WIS); Quevedo— Manta rd., 50.9 km W of Kl
Empalrne, R. Sauleda. M. Ragan. II. Luther. R. Wunderlin.
B. Ilansen. L. Davenport X ./. Wiersem a 4023 (LJSf. W IS
photocopy); Par. Nac. Machalilla, N de Puerto Lopez. V.
Cornejo X C. Bonifaz 2322 (GUAY. QCNE, W IS); hstero
Manta Blanca, S of Agua Blanca, A. Gentry X G. Josse
72718 (MO, QCNE, W IS); Las Goteras, C. Hernandez 222
(OCA); Bahia de Caraquez, predios de la Univ. Catolica,
A. Cornejo X C. Bonifaz 7154 (GUAY, WIS |2J); bills
above Rfo Chone, just in back of Sitio Mauricio along
main n I. 4 km SSE of Bahia de Caraquez, II. II. litis X
1/. litis hi- 1 68 (QCA p.p., vv/ C. lanceolata ); 9 km N of
San Vicente, //. H. litis X M. litis K-208 (WIS |3|): llcda.
Ed Recreo, ca. 2 km along coast from Canoa, IB km N of
Bahia de Caraquez. II. II. litis X M. litis E-216 (W IS |2|).

Acknowledgments. This paper has been fully 28
years in gestation. lb the Nave Bequest for Latin
American Studies, administered by the Dean of
Letters and Science, University of Wisconsin, for
partially supporting fieldwork in Ecuador during
the summer of 1977. our thanks. Without fieldwork,
no amount of dried herbarium material will substi¬
tute for viewing living (lowers, fleshy, decomposing
fruits, or the complexities of the ecosystems. Ones
obligations cannot be expressed deeply enough to
that hardy soul who, helpful and uncomplaining,
trudged along as field companion to the senior au¬
thor (and once, literally, as a life-saver) in extreme¬
ly hot, humid, or arid tropical habitats. So. special
thanks to my assistant and son, Michael G. litis,
who braved two months in Ecuador chasing Cyno-
phalla's, and saved that expedition in more ways
than one. To Carmen Bonifaz. Director of the her¬
barium of the Universidad de Guayaquil. Ecuador
(GUAY), and to the GAIA Foundation of Guayaquil,
for supporting the collections of C. sclerophyUa ; to
Flor Valverde and Florinda Trivino, professors of
Botany emeritus at t he Universidad de Guayaquil,
for being helpful in many ways, botanically and
otherwise; to Galaway Dodson and Aleides Miirmol

of the Rfo Palenque Biological Station, for their
generous hospitality; to Craig McFarland, former
Director of the “Fundacion Charles Darwin para las
Islas Galapagos." and his wife, for rescuing in 197 i
two sick travelers and their huge, full plant presses;
and to FUNDACYT of Ecuador, for support of the
second author’s air travel to Madison. Wisconsin;
to all our sincerest gratitude. Finally, we are much
obliged to Balia Pazy, superb cytogenetist of Jeru¬
salem University, Israel, for several diffieult-to-ob-
tain chromosome counts; to Bernardette Costa e Sil¬
va. of the Universidade Federal Rural de
Pernambuco, Recife, Brazil, for allowing us to pub¬
lish her significant, previously unpublished chro¬
mosome counts; and to the unexcelled artist ol the
Botany Department, University of Wisconsin, Kan-
dis Elliot for preparation of the splendid illustra¬
tions.

Literature Cited

Costa e Silva, M. B. 1995. Estudos Taxonomieos sobre <>
Genero Capparis L. (Capparaceae Juss.) em Pernam-
buco-Brasil. Dissertacao apresentada ao Prograrna de
posl-Graduacao em Botanica da Universidade Federal
Rural de Pernambuco. Recife, Brasil. | Unpublished.]

I)i Sapio, O. A.. M. A. Gatusso & I). E. Prado. 2001.
Structure and development of the axillary complex and
extrafloral nectaries in Capparis retusa Griseb. PI. Biol.
(New York) 3: 598-606.

Eggers, H. 1894. Das Ktistengebiet von Ecuador. Deut¬
sche Geographisehe Blatter, Geogr. Gesellsehaft in Bre¬
men 17(4): 265—289, tab. 4.

Fielder, A. W. 1865. Capparideae. In: C. E. P. von Martins
(editor). Flora Brasiliensis 1.3(1): 237—292, tab. 51 65.
Illis, H. H. 1978. Studies in the Capparidaceae \l\: Cap¬
paris ecuadoriea, a new species. Selbyana II: 303—307.

- . 1991. Capparaceae. In: I). H. Nicholson (editor).

Flora of Dominica, Part 2: Dicotyledonae. Smithsonian
Contr. Rot. 77: 60-63.

- . R. Kowal. It. Pazy & L. Pzywara. 1985. Miscel¬
laneous new chromosome counts for Capparaceae and
Cleomaceae. | Unpublished.]

Madsen, J. E„ R. Mix & H. Balslev. 2001. Flora of Puna
Island, Plant Resources on a Neotropical Island. Aarhus
Univ. Press, Denmark.

Poulsen, V. A. 1879-1880. Del ekstrallorale nektarium
hos Capparis cynophallophora |i.e., = C. flexuosa (L.)
L.]. Vidensk. Meddel. Naturhist. Foren. Kjpbenhavn,
31/32: 35-44, tab. II.

Ruiz-Zapata, T. & II. H. litis. 1998. Capparaceae. Pp.
132—157 in P. E. Berry, B. K. Holst N K. Yatskievych
(editors). Flora of the Venezuelan Guayana, Vol. 4. Mis¬
souri Botanical Garden Press, St. Louis.

Magnolia krusei (Magnoliaceae), una Especie Nueva de

Guerrero, Mexico

Jaime Jimenez Ramirez y Ramiro Cruz Duran

Herbario de la haeultad de Ciencias, UN AM, Apartado Postal 70-399, Delegaeidn Coyoacdn
04510, Mexico, D.F. jjr@hp.fciencias.unam.mx

RKSUMEN. Sc iluslra y describe a Magnolia krusei
(Magnoliaceae), una especie nueva del estado de
Guerrero, Mexico. Magnolia krusei es affn a M. il-
tisiana A. Vbzquez y a M. tamaulipana A. Vaz¬
quez. pero difiere de bstas por presentar llores y
folfculos de menor tamano. Tambien se ineluye una
clave dicotbmica para distinguir las especies afines
a este nuevo taxbn.

ABSTRACT. Magnolia krusei (Magnoliaceae), a new
species from the slate of Guerrero, Mexico, is il¬
lustrated and described. It is related to M. iltisiana
A. Vazquez and M. tamaulipana A. Vazquez, but
differs from these by its smaller (lowers and folli¬
cles. A dichotomous key for distinguishing all re¬
lated taxa to this new taxon is also included.

Key words: Guerrero, Magnolia, Magnoliaceae,
Mexico.

En (‘I proceso de curaeibn de la coleeeion KRU-
SE del Herbario Nacional de Mexico (MEXU), Ins-
tituto de Biologfa, UN AM. se detectb una especie
tie Magnolia que difiere en caraeterfstieas de las
hasta ahora conocidas, no incluida en el trabajo de
Vbzquez (1994), ni en trabajos posteriores de este
u otros autores.

A1 examinar las primeras colectas (Kruse
[ 1 9630503—238] y [19750326-239], MEXU), exis-
tfan algunas dudas sobre la madurez de las (lores,
pues cl reducido tamano de los sepalos y pbtalos
podrfa atribuirse a un desarrollo incompleto, por
ello, (lie necesario colectar especfmenes con (lores
indudablemente maduras. Ademas, fueron colec-
tados frutos maduros e inmaduros para describir
sus caraeterfstieas de manera fidedigna.

Magnolia krusei J. Jimenez Ramirez & R. Cruz,
sp. nov. TIPO: Mexico. Guerrero: Mochitlan.
San Roque, 1 7°2 1 '34"N, 99°25'20"W, 1150 m,
26 Abr. 2003, R. Cruz Duran & R. Fragoso
Ramirez 5586 R (holotipo, MEXU; isotipos,
EC ME, MO). Eigura 1.

Arbor 6-10(-25) m alia; sepala 3, (3.2-)3.7-4.2 X 2

Novon 15: 438-441. Published on 21 Sep

cm; petala 6, 3.4— 4.3 X (1 .4— )1 .8-2. 5(— 2. ft) cm; stamina
(99-) 120- It) 1 (-1 70); polyfolliculus globoso-oblongus cum
38—50 folliculis; folliculus 1.4— 1.5 cm longus; semina (7—
)8-9 X 5-6 mm.

Arbol de 6— 1()(— 25) m de alto; tronco 50—100
cm d.a.p., ramificaeibn desde los 1—2 m de altura;
ramas con abuudantes lenticelas blancas; ramillas
lanoso-pubescentes, pubescencia blanca (verde
palido en seen), en ramas maduras la pubescencia
persiste, pero adquiere una coloracibn negra; cor-
teza gris-negra, lisa con algunas fisuras. Ilojas al-
ternas, coriaceas, elfpticas a oblongo lanceoladas,
hasta con 16 pares de venas secundarias, (I I — ) 1 2 —
1 9.5 X (3.5— )4— 8(— 8.2) cm, bpice aeuminado, lig-
eramente cuspidado (o muy rara vez agudo), base
cuneada; peefolo 1.5— 2.5 cm de largo, canalicula-
do, pubescente; haz verdoso, glabro; enves lanoso-
pubescente. pubescencia glauca con pelos largos y
eortos; estfpulas (2.4— )4.2-5.6 X 1.4 cm, blanco-
pubescentes en la cara externa, (rojizas a eafb
cuando secas), los pelos del bpice mbs largos. Elo-
res terminales, solitarias (rara vez en un patron ra¬
cemose). blanco-cremosas (amarillas segun Ixtzada
1810 o amarillas-erema segiin Kruse |238|), 6.5—7
cm de dibmetro; pedunculo ( I— ) 1 .2— 1.6(— 2.5) X
0.5-1. 2 cm, lanoso-pubeseente, pubescencia blan¬
ca (verde palido cuando seco); hot bn floral 3-4 X
2-2.4 cm, ovado-elfptico, lanoso-pubeseente, pu¬
bescencia blanca, brbeteas 2, lanoso-pubescentes
en la cara externa, glabras en la cara interna, la
externa 3.7—4 X 3—4.4 cm; sbpalos 3, oblongo-elfp-
ticos, blanco-cremosos en el haz (verde claro en el
enves en llores jbvenes), consistencia carnosa,
(3.2— )3. 7—4.2 X 2 cm, haz glabro, pubescencia
blanca en el envbs (en algunos casos sblo pubes-
centes cerca de la base), margen ligeramente ci-
liado, no dedexos, mbs largos que los petalos; pe-
talos 6, blanco-cremosos, consistencia carnosa,
glabros, dispuestos dos series, los externos obova-
dos, alternos a los sepalos, 3.4 — 4.3 X ( 1 .4 — ) 1 .8 —
2.5( — 2.8) cm. los internes lanceolado-obovados.
opuestos a los sepalos, (3-)3. 4—3.6 X 1.4-1. 5 cm;
estambres lineares, (99-) 120- 161 (-170), 1.3— 1.4
X 0.15 cm, bpice agudo, bianco (amarillo palido

I’EMBER 2005.

Volume 15, Number 3
2005

Jimenez & Cruz
Magnolia krusei de Mexico

439

Eigura 1. Magnolia krusei J. Jimenez Ramirez & R. Cruz. — A. Rama eon flor. — B. Boton floral. — C. Cineceo. —
I). Fruto inmaturo y estfpula. — E. Polifolfculo maduro. — F. Semillas, con y sin sarcotesta. — G. Estambres, vista
ventral v lateral. A, F v G, A’. Cruz Duran &: R. Fragoso Ramirez 5586 R (FCME, MEXLI, MO); R y F R- (*ruz Duran
& R. Fragoso Ramirez 5585 B (FCME, MEXU); C y I). R. Cruz Duran & A. Gonzalez Zanwra 5571 (FCME, MEXU).
Ilustrado por Ramiro Cruz Duran.

cuando seco); gineceo oblongo, 2 X 1 .2 cm, lanoso-
|)iibescente, pubescencia blanca (verde palido en
seco), con eslilos crectos, verde claro (cafe a negros
cuando secos), estigmas amarillentos, ligeramente
curvos. Polifolfculo 5.2— 5.7 X 3.4— 3.8 cm, globoso-
oblongo; folfculos 38—50, 1.4— 1.5 X 0.6— 0.8 cm,
elipsoides a oblongos, lanoso-pubescentes, pubes¬
cencia verde palido, con estilos persistentes, erec-
tos. Semillas triangular-deltadas, (7— )8— 9 X 5—6

mm, cafe claro, 1 6 2 por folfculo (predomina ge-
neralmente una), con sarcotesta anaranjada-roja en
fresco, con aroma a flor de Annona.

Fruit ificacion . Noviem bre-ab ri 1 .

Floracion. Abril— mayo (Julio, ejemplares cul-

tivados).

Distribucion y habitat. Magnolia krusei se lo-

440

Novon

caliza en altitudes de 1 140—1600 m (2000 m en un
ejemplar cultivado: A. Vazquez el al. 4901 , WIS),
en restos de bosque dr Quercus y hosque mesdfilo
de montafia.

Kn San Roque la poblacion de Magnolia krusei
es eseasa. los individuos prosperan entre roeas de
esquistos donde se retiene abundante humus y se
asocia a Quercus sp., Xilosma velutina (Tulasne)
Triana <X Planehon, Melastomataceae, Rubiaeeae,
Peperomia sp.. Compositae y Rhamnaceae. En
Ximileotitlan se pueden observar arboles de hasta
IS in de altura y mi troneo de hasta 100 cm d.a.p.,
<•011 algunas epffitas como Anthurium sp.. Vittaria
sp.. y asoeiados a Slyrax sp. y Verbenaeeae. Aproxi-
madamente 1 km al 0 de Ximileotitlan se obser-
varon numerosas pldntulas de Magnolia krusei cre-
eiendo en la hojarasca de un bosque mixto de Pinas
sp. y Quercus candicans Nee.

Usos. Segun informes de los habitantes de la
region, una infusion de las Mores frescas se utili/.a
contra los colicos y dolores de estdrnago. y por su
agradahle aroma, tienen uso ritual y ornamental. Kn
particular, los habitantes del poblado de San Roque
meneionan que se esta iniciando la explotaeidn de
esta espeeie eon fines combustibles y senalan <|ii<“
las semillas son eonsumidas y dispersadas por aves
como "tenzapatos", “ cardenales ”, “ adrilenos ” y "]il-
gueros". Eos artesanos del poblado de Aeahui/.olla.
Guerrero (donde no hahita la espeeie nueva) ntili-
zan su madera para la elaboration de utensilios
domestieos como platos, saleros, eueharas, etc. Su
nombre comun es '‘Iolosuehil”, " Yoloxochitl".

Epfteto especi'fico. El nombre de esta espeeie
honra al lug. Hubert Kruse Robinett (1910-1984),
por su mctodica e invaluable labor botaniea en la
reeoleeta de ejemplares botdnicos para el estado de
( hierrero. Mexico.

En los ejemplares de Magnolia krusei eoleetados
por uno de nosotros (A*. Cruz Duran el at. 5.505 P.
550 / It. 5500 H. 5900) el color de los petalos es
bianco eremoso. pero, en los ejemplares de Kruse
\ 1 9600500— 200] la etiqueta dice amarilla-erema v
en Ijozada 1010. amarilla, lo que puede indiear que
el color de la flor varfa eon la edad o que la per-
eepeidn del color es distinta entre los eoleetores.

Magnolia krusei pertenece a la seceidn Therho-
<lon, caraeterizada por tener hojas perennes, estf-
pulas fibres, 2 braeteas. sepalos y petalos subsi-
mi lares, estambres con <4 conectivo agudo a
aeuminado y anteras con dehiseeneia introrsa. Es
pareeida a M. iltisiana A. Vdzquez y M. tamauli-
pana A. Vazquez y puede distinguirse de ellas en
la clave que se present a a continuacion.

Clavk para Distinguir i.as Espkciks Afinks a M. km

SKI SI’. NOV.

la. Mores eon didmetro de I 1 — 15 cm; sepalos ob-
longo elfpticos u oblongo obovados. 5—9.6 cm de
largo; petalos obovados, 5.5—9 cm de largo; fo-
lfeulo 2.4— 2.7 cm de largo.

2a. Ernes de las hojas pubescente; sepalos 5—

6 cm de largo; petalos eon el apice agudo,
todos con tamano similar, 7—8 cm de largo;
estambres 90—100, bianco eremosos; poli¬
folfculos subgloboso-oblongoides; folfculo

2.4 cm de largo . M. iltisiana

2b. Ernes de las hojas pubernlo; sepalos 6.5—

9.6 cm de largo; petalos con el apice redon-
deado, los externos de 5.5—9 cm de largo,
los internos de 4.5— 6.5 cm de largo; estam¬
bres 140—145, con la base y el apice pur¬
pura; polifolfculos ovoide-oblongoides; folf-

culo 2.7 cm de largo . M. tamaidipana

I b. 1 lores con diametro de 6.5—7 cm; sepalos oblon¬
go elfpticos, 3. 7-4.2 cm de largo; petalos exte-
riores obovados y los interiores lanceolados
obovados, 3—4.3 cm de largo; folfcnlo 1 .4—1 .5 cm
de largo . M. krusei

Magnolia krusei difiere de M. iltisiana ]>orque
liene las Mores mas petjuenas (6.5—7 cm, frente a
I 1 — 14 cm de diametro), los petalos exteriores e in¬
teriores de diferente forma y tamano, todos con el
apice redondeado (frente a los petalos totalmente
similares en forma y tamano. eon el apice agudo,
de la segunda espeeie) y los bolones Morales ovado-
elfptieos (mientras que M. iltisiana los tiene ovado-
oblongos), ademas. M. krusei presenta polifolfculos
globoso-oblongos y folfeulos de 1.4-1. 5 cm de lon-
gitud. en eambio. M. iltisiana posee polifolfculos
subglobosos a oblongos y folfeulos de 2.4 cm de
largo.

Magnolia krusei difiere de M. tarnaulipana
porque tiene las Mores mas pequenas (6.5—7 cm,
frente a 12-15 cm), los petalos mas pequenos (3.4—
4.3 cm, en contraste, con los 4.5—9 cm de la se-
gunda espeeie) y los petalos interiores lanceolado-
obovados mientras que en M. tarnaulipana, son
obovados. Eos estambres de la espeeie nueva son
color bianco eremoso. asf como la parte estamiuada
del reeeptaeulo. en eambio. M. tarnaulipana tiene
el apice y la base de los estambres purpura, mismo
color que presenta la poreion estamiuada del re¬
eeptaeulo. Eos polifolfculos de M. krusei son glo-
boso-oblongoides y los folfeulos miden 1.4— 1.5 cm
de largo, en contraste, en M. tarnaulipana los po-
lifolfeulos son ovoide-oblongoides y los folfeulos
miden 2.7 cm.

El examen de las i nidge nes digitales de dos pa-
ratipos de M. iltisiana (Miller & Campos 2949,
WIS, colectado a 5 km de El Parafso, Guerrero y
Vdzquez et al. 4901 , W IS. colectado en Puerto del
Gallo, de un arbol cultivado), permitio reubicarlos

Volume 15, Number 3
2005

Jimenez & Cruz
Magnolia krusei de Mexico

441

en la especie nueva. Esta circunstancia, unida al
hecho de que el resto de los paratipos de Guerrero
de M. iltisiana (no vistos por nosotros) hallan sido
reeolectados en un area geografica restringida (a I 1
kin o menos del poblado de Paraiso, Guerrero),
hace suponer que todos pudieran perteneeer en
realidad a M. krusei. Adenitis, el ejemplar Ixtzada
IRIO (FCME), no visto por Vazquez (1994). desig-
nado ahora eomo paratipo de M. krusei , tambien lue
colectado en la misma poreidn geografica.

Paratipos. MEXICO. Guerrero: Moehitlan. San
Roque, 5 km ESE de Acahuizotla, camino a Ojo de Agua
de San Francisco [XimilcotitlAn], 10 Nov. 2002, R. Cruz
Duran & A. Gonzalez Zamora 5568 (ECME, M EX L . \\ IS),
5571 (ECME. MEXU), 5572 (FCME); San Roque, 20 Abr.
2003, R. Cruz Duran & R. Fragoso Ramirez 5585 R
(ECME, MEXU), 5587 R (ECME, MEXU), 5588 R
(ECME, MEXU), 12 Ene. 2005, l\. Cruz Duran 5688
(FCME); Ojo do Agua de San Francisco [Ximilcolitlan],
Sierra Frente a Acahuizotla, 3 Mayo 1 1962 6 1963|, segun
diurio de la epoea, fcchado el 15 de Mayo de 1963, Hubert
Kruse [ 19630508-288 ] (ECME, MEXU); Ximilcolitlan, 26
Mar. 1975, Hubert Kruse \ 19750826— 239] (MEXU), 12
Ene. 2005, R. Cruz Duran 5965 (FCME); 1 km al 0 de
Ximilcotitlan, 12 Ene. 2005, R. Cruz Duran 5959 R
(ECME); Atoyae de Alvarez, Nueva Deli, 4 km al S. 20
Mayo 1993, /.. Ijozada Perez 1810 (ECME); between Rio
Verde and El Paraiso, a short dirt road (ea. 5 km off the
road between El Paraiso and Atoyae), 23 Mavo 1987, ./.

S. Miller & G. Campos 2949 (\\ IS); breclia Puerto del
Gallo a Cruz de Ocote, I I Jul. 1988, A. Vazquez et al.
4901 (A IS).

Agradecimientos. A I l)r. Mario Sousa del Her-
bario Nat ional (MEXU) por las faeilidades otorga-
das para la consulta del material de la COLEC-
CION KRUSE, a la Biol. Gloria Andrade por la
informacion biografica referente al lug. Hubert
Kruse Robinett, a la Biol. Ma. del Rosario Garcia
IVna, tecnico academico del MEXU, por permitir
la revision de material en prestamo del genero
Magnolia, al M. en G. Lucio Lozada Perez, quien
proporciono sus colectas de Magnolia del estado
de Guerrero; a los Biol. Alberto Gonzalez Zamora
y Rogelio Eragoso Ramirez por la ayuda en el tra-
bajo de campo, a la M. en G. Martha Martinez Gor-
dillo la diagnosis en latin y al Biol. Jose Antonio
Hernandez Gomez de Microcine, Facultad de Cien-
cias, LJNAM. la digitalizacion de la Figura I. Y de
manera especial al l)r. T. S. Cochrane por el envfo
de literatura e imagenes digitalizadas de material
de herbario de Magnolia depositado en W IS.

Literatura Citada

VAzquez-G., J. A. 1994. Magnolia (Magnoliaceae) in Mex¬
ico and Central America: A sinopsis. Rrittonia 46: I —

23.

Two New Species of Myrtaceae from South America

Leslie f\. hindrum

School ol Life Sciences, Arizona State University, Tempe, Arizona 85287-4501, U.S.A.

les. landrum@asu.edu

Abstract. Two new species of Myrtaceae are de¬
scribed and illustrated, Calycolpus aequatorialis of
Ecuador and Psidium huanueoense of Peru. Caly¬
colpus aequatorialis appears to have a close relative
in C. surinamensis, but differs in having a closed
calyx, relatively little indumentum, and strongly 4-
vvinged twigs. Psidium huanueoense is compared
with the similar species P. acutangulum from which
it differs in having velvety indumentum of short,
yellowish, erect hairs and larger leaves. Compari¬
son with P. guineense is also made, from which it
differs in venation, number of ovules, anther length,
and hair length.

Key words: Calycolpus, Ecuador, Myrtaceae,
Peru, Psidium.

Calycolpus 0. Berg is a genus of ea. 12 species
ranging from Central America to Minas Gerais,
Brazil, with the greatest diversity in northeastern
South America in the Guayana Highlands (Mc-
Vaugh, 1969). It is closely related to Psidium L.
and distinguished from it by a combination of in¬
florescence, floral, and seed coat characters (Lan¬
drum & Sharp. 1989). For a few years I have been
aware of an unusual species from northeastern Ec¬
uador, known from a single collection, that I as¬
sumed belonged to Psidium because of its closed
calyx and strongly 4-winged tw igs, characters com¬
mon in Psidium. These same characters were until
now unknown and rare, respectively, in Calycolpus.
The best characters for distinguishing Psidium and
Calycolpus are in the seeds, which are still un¬
known lor this species, so some doubt remains, but
on consideration of other characters, I believe this
species is best placed in Calycolpus. It has ovate,
persistent bracteoles, which are unknown in Psi¬
dium, but found in some species of Calycolpus. It
has bracteate shoot inflorescences with a short axis
and leaves that dry dark reddish brown, both of
which are common characters in Calycolpus and
rare in Psidium. The prominent brochidodromous
venation with the marginal vein closely following
the margin is unusual in Psidium, but essentially
identical to that found in C. surinamensis. This is
the first speeies of Calycolpus known that has a

closed calyx, a character that until now was useful
in separating the genera.

I. Calycolpus aequatorialis Landrum, sp. nov.
TYPE: Ecuador. Sucumbios: Sendero La Hor-
miga, 0°5'25"N, 76°12'50"W, 26 Apr. 1986
(fl), ./. Jaramillo H4H7 (holotype, QCA. ASU
photo; isotypes, NY, QCA, ASL photos). Fig-
u re 1 .

Haec species Calycolpo surinamensi prime aspectu sim-
ilis, sed ealyce ante anthesin elauso, sub anthesi irregu-
lariter lisso, el ramunculis juvenibus 4-alatis differt.

Arbor usque 12-metralis; folia lanceolata, elliptica, vel
oblonga, 8.7—18 X 3.6-6. 7 cm, 2—3.4 plo longiora quam
latiora, apice aeuminato, plerumque falcato; stamina ea.
325, ea. I cm tonga, antherae ea. 3 mm longae, connec-
tivum multi-glanduliferum.

free ea. 12 m high, glabrous to minutely puber-
ulent, sometimes densely so on hypanthium and ca¬
lyx apex within; hairs whitish, to ca. 0.25 mm long,
mainly appressed; young twigs with 4 prominent
wings (to ca. 1 mm w ide), dark reddish brown when
dry. glabrous to moderately appressed puberulent.
Leaves lanceolate, elliptic or oblong, 8.7—18 cm
long, 3.6-6. 7 cm wide, 2-3.4 times as long as wide,
glabrous or nearly so; apex acuminate, usually fal¬
cate; base obtuse to acute; petiole 5—6 mm long,
1.5—2 mm wide, channeled, sparsely appressed pu¬
berulent; midvein prominent below, impressed
above, the venation brochidodromous, with 15 to
20 pairs of lateral veins, these prominent below,
impressed above, leaving the midvein at an angle
of 60—80°, a clear marginal vein connecting with
laterals and ± paralleling the margin, running ca.
2—4 mm within the margin, the tertiary veins con¬
necting the larger veins in a ± dendritic pattern;
blades coriaceous, drying dark reddish brown.
Flower buds pyriform, ca. 18 mm long; peduncles
uniflorous, 12-26 mm long. ca. 2 mm wide, terete
to somewhat flattened, borne in axillary clusters
(short bracteate shoots), obscurely and sparsely pu¬
berulent; bracteoles ovate to suborbicular, ca. 3.5
mm long and wide, often wider than long, clasping
hypanthium, persisting at least until anthesis; calyx
closed in bud. prolonged in an apiculale tip. tearing
irregularly at anthesis, sparsely puberulent without

Novon 15: 442-446. Publishf.d on 21 September 2005,

Volume 15, Number 3
2005

Landrum

Myrtaceae from South America

443

figure 1. Calycolpus aequatorialis Landrum. — A. Branchlet with flower buds and old flowers. — B. Magnified portion
of young 4-winged twig and short bracteate shoot bearing a flower bud. — C. Longitudinal section of a flower bud with
two views of the placenta with numerous ovules, and a cross section of a floral ovary. — D. Magnified view of anthers
showing the glands in the connectives. A— D, Jaramillo 8487 (QCA).

and within, more densely so near apex within, the
remnants ca. 0.2 mm thick when dry; petals gla¬
brous except for a ciliate margin, strongly glandu¬
lar; hypanthium ca. 8 mm long, densely puberulent;
disk subglabrous, ca. 10 mm across; stamens ca.
325, to ca. I cm long; anthers oblong, to ea. 3 mm
long, with 8 to 22 glands in connective; style ca.
12 mm long, glabrous; ovary 3-locular in I flower

dissected; ovules ea. 75 in 1 loeule observed, ca.
10-seriate, the placenta a nearly sessile, peltate
pad. Fruit unknown.

This remarkably distinct species is known from
a single gathering made by J. Jaramillo, who col¬
lected several duplicates, surely realizing that it
was something unusual. Habitat information is

444

Novon

lacking from the label, hut judging from other col¬
lections by Jaramillo from the general vicinity (in¬
formation available through Tropicos at MO, <hltp:
//mobot. mobot.org/W3T/Search/vast.html>), it is a
lowland wet tropical forest at about 250 m that is
rich in Araceae, Bignoniaeeae, Gesneriaceac, Laur-
aceae, etc. This single collection was flowering in
April.

In describing Calyeolpus surinamemis, McVaugh
(1969) drew attention to the strong venation and
large persistent bracteoles as being unusual char¬
acters iu Calyeolpus, both characters that it shares
with C. aequatorialis. The lateral veins in these
species leave the midvein at an angle of about 60°
or more. This type of venation is also found in C.
bolivarensis Landrum and similar bracteoles are
possessed by C. legrandii Mattos, species unknown
to McVaugh. Closed calyces appear to have evolved
numerous times iu Myrtaceae. so it is not surprising
to find them also in Calyeolpus. All the specimens
known to me are of a single collection and arc held
at QC A and NY .

Psidium L. (Myrtaceae) is a genus of at least 50
and perhaps as many as 100 species ranging from
Mexico and the Caribbean to Argentina and Uru¬
guay. The distinguishing characters of Psidium are
discussed in Landrum (2003) and in Landrum and
Sharp (1989).

2. Psidium huaiiucocnse Landrum, sp. nov.
TYPE: Peru. Iluanuco: Puerto Inca, Dtto. Yu-
yapichis, Unidad Modelo de Manejo y Pro¬
duction Forestal DANTAS, 9°40'S, 75°02'W.
270 m, 16—30 Apr. 1990, Telia 47 (holotype,
USM. ASU photo; isotypes, C, ASH photo, NY.
ASU photo). Figure 2.

Haec species Psidio aeutangulo prime aspectu similis.
sed alabastro el fructu pi I is ereetis densis ochraceis dif-
fert.

Arbor usque 15-metralis; ramunculi juvenes 4-alati; fo¬
lia elliptica-ovalia, ovata vel lanceolata, 8.5—21 X 5.5—9
cm. 1.3— 2.5 plo longiora cpiam laliora; calyx ante anlhesin
clauses, sub antbesi irregulariter fissus; stamina ca. 600,
ea. 12 mm longa, antherae ca. 1.5 mm longae; loculi ea.
265-ovulati.

Tree to ca. 15 m high, sparsely to densely pu-
berulent on most surfaces; hairs light tan to whitish,
usually erect, to ca. 0.25 mm long; young twigs with
1 prominent wings, light brown to reddish brown
when dry, moderately puberulent, glabreseent with
age. Leaves elliptic, ovate, or lanceolate, 8.5— 2 I
cm long. 5.5—9 cm wide, 1.3— 2.5 times as long as
wide, moderately puberulent below (especially
along veins), sparsely puberulent to glabreseent
above; apex broadly rounded to acute or acuminate

(sometimes abruptly so); base euneate to rounded;
petiole 5—12 mm long, 1.5—3 mm wide, channeled,
puberulent to glabreseent; midvein prominent be¬
low1, impressed above, the venation brochidodrom-
ous or partially eucamptodromous, with 7 to 10
pairs of lateral veins, these prominent below, im¬
pressed to about flat above, leaving midvein at
about 45°. a clear marginal vein evident at least
distally, arching slightly between laterals, ± equal¬
ing laterals in prominence, running 3-12 mm from
margin, a second weaker marginal vein running
along margin itself sometimes evident, the tertiary
veins connecting larger veins in a dendritic (rarely
ladder-like) pattern; blades coriaceous, drying dark
reddish brown above, olive green below. Flower
buds pyriform, 15—17 mm long, densely puberu¬
lent; peduncles uniflorous or triflorous, 10—22 mm
long, to 2.5 mm wide, flattened, the lateral arms of
the dichasia to ca. 8 mm long; bracteoles lanceo¬
late-linear, 6—10 mm long, 1—2 mm w ide, puberu¬
lent, caducous before anthesis; calyx closed in bud.
terminating in an acute to acuminate tip. tearing
irregularly at anthesis, the remnants 0.5— 0.7 mm
thick when dry; petals obovate to suborbicular,
probably ca. 1.5 cm long when expanded, strongly
glandular, densely appressed puberulent centrally
without; hvpanthium campanulate, about 6— 7 mm
long; disk ca. 6 mm across; stamens ca. 600, ca.
12 mm long; anthers oblong, ca. 1.5 mm long (per-
ha, is longer when fully expanded), with 5 to 10
glands in the connective; style glabrous, to ca. 1.5
cm long, the stigma peltate; ovary 5-locular in 1
ovary examined; ovules ea. 265 iu 1 locale exam¬
ined. Fruit ca. 5 cm diam.; seeds probably more
than 200. 4—5 mm long.

Phenology. Flowering in April: fruiting in No¬
vember.

Habitat and distribution. Based on label data,
the habitat is wet tropical forest at ca. 280 m, with
ca. 2.5 m of annual precipitation and an average
temperature of 23— 25°C; so far known from a re¬
stricted region of Iluanuco. Peru .

Specimens of Psidium huanucoense have pre¬
viously been identified as P. acutangulum DC.,
and it resembles that species except for the vel¬
vety indumentum of fine, yellowish, erect hairs,
this condition being especially distinct on the
flower bud and fruit and quite unlike P. acutan¬
gulum. The leaves of P. huanucoense are generally
much larger (8.5—21 X 5.5—9 cm) than in P. acu¬
tangulum (3.5—13 X 1.5— 5.6 cm). The peltate pla¬
centa with a broad smooth central area is unlike
P. acutangulum and more similar to P. guineense
Swartz. It differs from P. guineense in having a

Volume 15, Number 3
2005

Landrum

Myrtaceae from South America

445

strong marginal vein. The number of ovules is few¬
er in / \ guineense (generally 50 to 100), the an¬
thers usually longer (1—3 mm), and the hairs lon¬
ger (0.3— 0.5 mm). The acute to acuminate tip of
the closed c alyx of P. huanucoen.se is rare in P.
guineense.

Paratypes. I'KIUJ. Huaiiuco: Puerto Inca, Yuyapi-
eliis, Unidad Modelo do Manejo y Produccion Porestal
DANTAS, 9°40'S, 75°02'W, 270 m, 1-15 Apr. 1991

(hud), Telia 1756 (G, MO. USM), I Nov. 1989 (fr). IS. hriill
Saldana 6H4 (ASU).

Acknowledgments. I thank mv colleagues J. Jar-
amillo, k. Romoleroux, B. Leon, and my wife, Sonia
Landrum, whose help made this paper possible. I
am also grateful to the curators ol ASU, 0. MO.
NY. QCA, and USM, whose collections were fun¬
damental to this study. Three anonymous reviewers

446

Novon

and V. Hollowed offered many helpful suggestions.
Figures I and 2 were drawn by Bobbi Angel],

Literature Cited

Landrum, L. R. 2003. A revision of the Psidium salutare
complex (Myrtaceae). Sida 20: 1449—1469.

- & W. P. Sharp. 1989. Seed coat characters of some

American Myrtinae (Myrtaceae): Psidium and related
genera. Syst. Hot. 14: 370-376.

McVaugh, K. 1969. Myrtaceae. In The Botany of the Gua-
yana Highland — Pt. 8. Mem. New York Hot. Card.
18(2): 55-286.

Rubiacearum Americanarum Magna Hama, Pars XVII.
A New Species and Four New Combinations in
Mesoameriean Rondeletieae (Rubiaceae)

David //. Licence

National Tropical Botanical Garden, 3530 Papalina Road, Kalaheo, Hawaii 96765, U.S.A.

lorence@ntbg.org

ABSTRACT. Studies for a forthcoming treatment of
Rubiaceae for the Flora Mesoamericana project re¬
vealed an undescribed species from Honduras and
also the necessity for four new combinations in
Rondeletieae. The new species Arachnothryx nel¬
son ii Lorenee is described from Honduras. It is re¬
lated to the glabrous Mexican species A. purpurea
(Lorenee) Both idi from which it differs in having
leaves sparsely strigillose on the costa, veins, and
margins, flowers with a strigillose hypanthium and
calyx, white externally strigillose corollas, and
subglobose capsules. In addition, the following new
combinations are proposed in Arachnothryx and
Rogiera : Rondeletia myriantha Standley & Steyer-
mark var. armentalis L. 0. Williams is elevated to
species level as Arachnothryx armentalis (L. 0.
Williams) Lorenee; Rondeletia hondurensis Donnell
Smith is transferred to Arachnothryx as Arachnoth¬
ryx hondurensis (Donnell Smith) Lorenee; Ronde¬
letia megalantha is transferred to Arachnothryx as
Arachnothryx megalantha (Lorenee) Lorenee; Ron¬
deletia standleyana A. R. Molina is transferred to
Rogiera as Rogiera standleyana (A. R. Molina) Lor-
ence; and Rondeletia ovandensis Lundell is reduced
to synonymy under Arachnothryx armentalis.

Key words: Arachnothryx, Javorkaea, Mesoam-
eriea, Rogiera. Rondeletia. Rondeletieae, Rubi¬
aceae.

As interpreted in the broad or traditional sense,
Rondeletia s.l. comprises approximately 230 to 250
species of shrubs and small trees restricted to the
tropical regions of Mexico. Central and South
America, and the Antilles. However, the genus dis¬
plays considerable morphological variation and has
been split into segregate genera by various authors,
most often utilizing morphological characters (for
further discussion see Borhidi, 1982, 1987, 2001:
Borhidi A Fernandez Zequeira, 1981; and Lorenee,
1991). I he three principal segregate genera are:
Rondeletia s. str., comprising ca. 140 species main¬
ly in the Antilles (four species in Mesoamerica);
Arachnothryx Planchon with ca. 80 species ranging

from Mexico to South America; and Rogiera Plan¬
chon, with 18 species ranging from Mexico to Pan¬
ama (Mabberley, 1997; Lorenee & Taylor, in prep.).
Initial cytological evidence based on investigations
of two species per genus revealed base numbers of
n = 9 for Arachnothryx , n = 10, 11 for Rogiera.
and n — I I, 14 for Rondeletia s. str. (Kiehn, 1995).

More recently, Borhidi and collaborators have
described several small segregate genera in tbis al¬
liance in Mexico and Mesoamerica based on mor¬
phological criteria, including Javorkaea Borhidi A
Jarai-Komlodi, Rennistipula Borhidi, and Rovean-
thus Borhidi (Borhidi & Jarai-Komlodi. 1983; Bor¬
hidi, 2003, 2004). Rennistipula was described to
accommodate three species with reniform stipules
that Borhidi had previously transferred to Arach¬
nothryx: A. costaricensis (Standley) Borhidi, A. g<d-
eottii (Standley) Borhidi, and A. izabalensis (Stand-
ley & Steyermark) Borhidi. In addition, Borhidi
(2004) described Roveanthus to accommodate two
species he had formerly included in Rogiera: R.
strigosus (Bentham) Borhidi and R. suffrutescens
(Brandegee) Borhidi. Pending further molecular
and phylogenetic studies covering a broad range of
Rondeletieae species, I believe it is premature to
recognize these new genera and prefer to retain
these species in Arachnothryx and Rogiera, respec¬
tively.

Recent molecular-phylogenetic data from /rrcL-F
cpDNA gene sequencing suggest that Rondeletia
s.l. is polyphyletic, and that Arachnothryx and Ja¬
vorkaea are closely related to each other and to
Gonzalagunia Ruiz & Pavbn (also Rubiaceae: Ron¬
deletieae), but less closely so to Rogiera and Ron¬
deletia s. str. (Rova et al., 2002). However, only a
small number of species in the Rondeletia alliance
have been sampled thus far for molecular studies;
more importantly, virtually none of the critical spe¬
cies of Arachnothryx from Mexico and Central
America with unusual floral and seed morphologi¬
cal characters that overlap the generic boundaries
proposed by Borhidi and others have been sampled.

Novon 15: 447-451. Published on 21 September 2005.

448

No von

These include species such as Arachnothryx delimit
(Donnell Smith) Borhidi and A. hondurensis, whose
deviating characters are noted in the key below.
Species comprising the segregate genera Rennisti-
pula and Roveanthus also remain unsampled. Fur¬
ther cytological, morphological, phylogenetic, and
molecular studies are clearly needed to satisfacto¬
rily resolve the systematics of this complex.

In light of available evidence it was decided to
recognize three segregate genera in the broader
Rondeletia alliance (i.e., Arachnothryx, Rogiera,
and Rondeletia s. sir.) lor the Flora Mesoamericana
treatment. During the course of preparing accounts
of these genera, collections representing an unde¬
scribed species from Honduras were encountered.
In addition, four new combinations in Arachnothryx
and one in Rogiera are required. A full treatment
ol the Mesoameriean Rubiaceae will be given in
the forthcoming treatment for Flora Mesoamerieana
by David Lorence and Charlotte M. Taylor (Lorence
& Taylor, in prep.).

Kky to thk Three Genera in the Rondeletia s.e. Ai -
i.iance in the Mesoamerican Recion

la. Lobes ol calyx and corolla usually 4 (corolla
lobes 5 and calyx lobes 5 to 7 in A. hondurensis,
calyx lobes rarely 5 or 6 in A. deamii); corolla
throat glabrous or white papillose-puberulent in
and around throat, not yellow or white barbate
(except in A. hondurensis), lacking a conspicuous
fleshy ring around throat, tube internally pubes¬
cent in basal portion; capsule dehiscence locu-
licidal and often with some septieidal splitting;
seeds usually rhomboid, square, or triangular, not
winged or caudate (except fusiform and winged
in 1. deamii. narrowly winged in A. costaricensis,

A. galeottii, and A. izabalensis), the testa usually
deeply foveolate-reticulate . Arachnothryx

lb. Lobes ol calyx and corolla usually 5 (rarely 4 or
6 in some flowers); corolla densely yellow or
white barbate in and around mouth of tube, or
with a thickened fleshy ring around mouth of
tube, tube glabrous within; capsule dehiscence
strictly loculicidal; seeds fusiform, caudate and
winged at one or both ends ( Rondeletia ), or an¬
gular to rounded or suborbicular (Rogiera), the
testa usually shallowly foveolate-reticulate.

2a. Corolla with a thickened, fleshy ring around
mouth ol tube (also papillose or puberulent
in R. belizensis (Donnell Smith) Lorence);
seeds fusiform, caudate and winged at one
or both ends (discoid with broad wing in R.

belizensis) . Rondeletia s. str.

2b. Corolla densely yellow or white barbate in
and around mouth ol tube; seeds angular
and rounded to suborbicular, unwinged . . .

. Rogiera

I. Arachnothryx iielsnuii Lorence, sp. nov.
TV PE: Honduras. Copan: Montana Los Zapo-
tes, mountain W of Los Zapotes, 18 km N ol

Copan ruinas on road to Agua Caliente from
Copan ruinas and 2-3 km W of Los Zapotes
school, 950-1050 m, 14°58'N, 89°10'W, 30
Apr. 1996 (11. fr), T. Hutchins 993 (holotype,
MO; isotypes, BM. LAP. MEXU, PTBG). Fig¬
ure I.

Haec species Arachnotrychi purpureae affinis, sed ab ea
lamina foliari secns costam venas marginemque parcestri-
gillosa. hypanthio calyceque strigillosis, corolla alba extus
strigillosa, atque eapsula subglobosa differt.

Small Irees 3—8 m tall, the leafy twigs slender,

1- 1.8 mm diam., terete, glabrous, drying dark
brown, wrinkled, the internodes 0.7—10 cm long.
Leaves opposite, those of a pair at a node aniso-
phyllous or subequal, one up to 50% larger than
the other; lamina ovate-elliptic, elliptic, oblong-el¬
liptic or oblaneeolate-elliptic, 4-1 1 X 1.5— 3.5 cm,
the apex abruptly acuminate or caudate, the acu¬
men 1-1.5 cm long, often falcate, the base euneate
to acute, often unequal-sided, drying membrana¬
ceous to thinly chartaceous, greenish brown, adax-
ially sparsely strigillose on costa and secondary
veins, abaxially glabrous or sparsely strigillose on
costa and veins, the secondary veins 3 to 7 pairs,
arcuate, prominulous on both surfaces, eucampto-
dromous, the axils barbate and domatiate, the ve¬
nation visible to quaternary order on both surfaces,
the margins sparsely strigillose-eiliolate; petioles

2— 10 mm long, 0.5— 0.8 mm diam., glabrous or
adaxially strigillose; stipules broadly to narrowly
triangular, acuminate, 0.8—2 mm long, ea. I mm
wide, externally sparsely strigillose, internally pi-
losulous. Inflorescence terminal, corymbiform-cy-
mose, triehotomous, 3-7 X 2.5—5 cm (including the
corollas), sessile or the peduncle to 3 cm long, 12-
to 27-flowered, the primary branches 3, ascending,
0.8— 3.4 cm long, subequal or the central one lon¬
gest, unbranched or branching once again, ending
in 2- to 6-flowered cymules. the axes glabrous, flat¬
tened, bracteoles 0.8-3 mm long, linear-subulate,
strigillose; flowers apparently distylous, subsessile
or on pedicels to 1.2 mm long, minutely bracteo-
late, the hypanthium ellipsoid, 1-1.2 X 0.6— 0.8
mm, strigillose, the calyx limb lobed nearly to base,
calyx cup 0.3 mm deep, calyx lobes 4. strigillose,
unequal. I large and oblong or oblong-elliptic, 2—

4.5 X 0.7— 0.9 mm, the other 3 lobes much smaller,
linear or subulate, 1—2.5 X 0.1 mm; corolla sal-
verform. while, the tube ca. 8 mm long, 0.6— 0.8
mm diam. medially, externally strigillose with as¬
cending hairs, internally hirtellous in basal 1/3,
distally glabrous, the lobes 4, oblong-elliptic, 1.2—

1.5 X 1 — 1.2 mm. externally strigillose-setose, in¬
ternally glabrous; short-styled flowers with stamens

Volume 15, Number 3
2005

Lorence

Mesoamerican Rondeletieae

449

Figure 1. Arachnothryx nelsonii Lorence. — a. Habit with inset showing abaxial leal surface details. — b. Flower, with
hypanthium bearing calyx lobes and style, and corolla tube opened to reveal stamens. — c. Two inflorescence evmules
with flowers in bud. — d. Disintegrating capsules showing vascular bundles. — e. Seed. — f. Node with stipule. All
drawn from the type, Han kins 993. Seale bar 5 cm in a; I cm in b, c, d; 0.5 mm in e; 1 mm in f.

included, sessile, attached ca. 1 mm below apex of
tube, the anthers ca. I mm long w ith style glabrous,
ca. 3.5 mm long with 2 linear stigma lobes ca. 1.5
mm long; long-styled flowers not seen. Capsules
subglobose, 3—4.5 mm diam., glabrate, slightly 1 »i -
silicate, weakly 6- to 8-ribbed, disintegrating with
age to reveal vascular bundles; seeds subglobose-
angulate, 0.4— 0.6 mm diam., the testa deeply retic¬
ulate, dark brown.

Distribution and habitat. Known only from the
departments of Copan and Cortes in Honduras
where it occurs in montane evergreen wet foresl
with Liquidambar. 950—1500 m in elevation. Col¬
lected in flower in February and April, anil in fruit
in April.

Etymology. This species is named for Hondur¬
an botanist Cirilo II. Nelson in recognition of his
contributions to the botany of Honduras.

450

Novon

Arachnothryx nelsonii seems most c losely related
to the nearly glabrous Mexican species A. purpurea
(Lorence) Borhidi from which it differs in having
leaves sparsely strigillose on the costa, veins, and
margins, flowers with a strigillose hypanthium and
calyx, corolla white with an externally strigillose
lube and externally strigillose-setose lobes, and
subglobose capsules.

Raralype. HONDl HAS. (lories: Montana de I.a Ni-
eve, lado SO del Caserfo El Tapiquilar, 20 km S de San
Antonio de (lories. 2d Kel>. 1982 (fl), C. Nelson. J. Escobar
& R. An, lino HOI) I (MO 4324692, PTBG, UN A II).

2. Arachnothryx armentalis (I.. (). Williams)
Lorence, comb, et slat. nov. Basionym: Ran-
deletia myrianlha Standley & Steyermark var.
armentalis I.. 0. Williams, Phytologia 26: 127.
1973. Arachnothryx myrianlha (Standley &
Steyermark) Borhidi var. armentalis (I.. ().
Williams) Borhidi, Acta Bot. Hung. 35: 310.
1989. TYPE: Guatemala. (v)uiche: Nebaj, 5600
It.. 22 Nov. 1 934, A. F. Skutch I 776 (holotype,
K 81581 I, photos at MEXU, PTBG).

Rondeletia ovandensis l.undell, Wrightia 5: 326. 1976.
Syn. nov. Arachnothryx ovandensis (l.undell) Borhidi,
Acta Bot. Hung. 35: 310. 1989. TYPE: Mexico,
(lliiapas: Ml. Ovando near Eseuintla, 24 Oct. 1911.
E. Matilda 0007 (holotype, I.I., photos at MEXL .
I*TBG; isotypes, E. 1.1. not seen).

V\ illiams (1973) described Rondeletia myrianlha
Standley & Steyermark var. armentalis (meaning
“one ol a crowd") based on a single flowering spec¬
imen. lie believed it represented a minor variant
of Arachnothryx myrianlha (Standley X Steyer¬
mark) Borhidi, from which it differed in having
leaves densely arachnoid-tomentose abaxially and
flowers with a sparsely arachnoid-tomentose hypan¬
thium and calyx limb and subequal calyx lobes.
Further study of additional collections has revealed
that A. myrianlha is characterized by having leaves
that are uniformly arachnoid-tomentose on both
surfaces when young and soon glabrescent and its
corollas externally strigillose-pilosulous, whereas
variety armentalis differs in having leaves adaxially
persistently hirtellous or seabrid and corollas ex¬
ternally tomentulous on the tube ami pilose-setose
on the lobes. In view of these fundamentally dif¬
ferent pubescent types and to maintain consistency
with taxonomic concepts in Mexican and Central
American Arachnothryx, Williams's variety is here
elevated to the species level as A. armentalis, the
earliest available epithet. Furthermore, A. ovanden¬
sis is here proposed as a synonym of A. armentalis.
This species is closely related to A. buddleioides
(Bentham) Planchon, which differs in having leaves

adaxially smooth and glabrous or at first sparsely
arachnoid-tomentose and soon glabrescent, inflo¬
rescences mostly longer than the subtending leaves,
and corolla lobes externally arachnoid-tomentose
hut lacking pilose-setose triehomes. Arachnothryx
armentalis occurs in Mexico (Oaxaca, Chiapas) and
Guatemala in mesic and cloud forests at 1 100—
I 700 m in elevation.

Representative specimens examined. MEXICO. Oaxa¬
ca: Sierra Mazateea. Mun. Huautla de Jimenez, alrede-
i lores del Puente de Fierro en direccirtn a Sta. Marfa Chil-
chotla (7 km de Huautla de Jimenez por la carretera a
lent il Ian— Mex 1 32), Munn-Estrada & Mendoza 1204
(I’TIIG). Chiapas: from El Triunfo to El Paval, S slopes
of Sierra de Soconuseo, Xolocotzi A Sharp X. 489 (MO).
GUATEMALA. Sail Marcos: above Finca el Porvenir on
“Todos Santos Chiquitos”, lower S-faeing slopes of Volcan
Tajumulco, Steyermark 87096 (F).

3. Arachnothryx hondurcnsis (Donnell Smith)
Lorence, comb. nov. Basionym: Rondeletia
hondurcnsis Donnell Smith, Bot. Caz. (Craw-
fordsville) 27: 335. 1899. Javorkaea honduren-
sis (Donnell Smith) Borhidi & J. komlodi. Acta
Bot. Hung. 29: 16. 1983. TYPE: Honduras.
Santa Barbara: Rfo Chameleeon, 300 m. Dee.
1888, C. Thieme 5267 (holotype, US 943502;
isotypes, F. US [2 1).

The segregate genus Javorkaea (Borhidi & Jarai-
komlodi. 1983) was described to accommodate
Rondeletia hondurcnsis Donnell Smith, a species
endemic to Honduras and recognized by Standley
(1918) as comprising the monospecific but never
validly published section Hondurenses Standley of
Rondeletia. Javorkaea was characterized similarly
to Standley ’s Hondurenses by 1- to 3-lobed stipules
with well-developed sheaths, I - to 3-storied inflo¬
rescences, a 4- to 7-merous calyx, a large, slightly
zygomorphic 5-merous corolla with lobes puberu-
lent basallv within, a subeapitate and bilobed stig¬
ma. a glabrous nectary disc, 4-eolporate pollen, ob¬
long hypanthium and capsule, capsule dehiscence
at first loculicidal then septicidal and separating at
base, and seeds as in Arachnothryx (Borhidi & Jar-
ai-komlodi, 1983). The subeapitate, shortly bilobed
stigma and 4-colporate pollen grains are distinctive
at the species level but fail to provide sufficient
basis for recognition at the generic level (Lorence,
1991).

Subsequently, Borhidi (1997a, 1997b) trans¬
ferred to Javorkaea six Mesoamerican species, two
from Rondeletia and four that he had previously
placed in Arachnothryx: A. chaeonii (Lorence) Bor¬
hidi |as chaconis], A. macrocalyx (Standley & Stey¬
ermark) Borhidi, R. megalantha Lorence, R. scabra

Volume 15, Number 3
2005

Lorence

Mesoamerican Rondeletieae

451

Hemsley, A. torre.si/ (Standley) Borhidi, and A.
itxpanapensis (Lorence & Castillo-Campos) Borhidi.
These species all display relatively large white or
yellow corollas, puberulent around the mouth of the
tube and Basal ly on the lobes within, but are not
otherwise closely related to Arachnothryx lumdu-
rensis except lor their large flowers, which may rep¬
resent an adaptation for moth pollination. Recently,
Borhidi (2003) described a new species, Javorkaea
pulcherrima Borhidi, from Chiapas, Mexico. Based
on an examination of the isotype (Heath & hmg
881, PTBG) this last species name is clearly syn¬
onymous with Arachnothryx jurgensenii (Hemsley)
Borhidi. The addition of these seven species radi¬
cally changes the original circumscription of Javorkaea
sensu Standley’s section Hondurenses, rendering it
a heterogeneous assemblage of large-flowered
shrubs or small trees differing in no consistent
character or suite of characters from the majority
of species otherwise placed in Arachnothryx.

The primary characters used to differentiate Ja¬
vorkaea, i.e.. tin* relatively long stipular sheath and
calyx cup, 1- to 3-aristate stipules, zygomorphie ca¬
lyx, large corolla with basally puberulent lobes
within, and oblong hypanthium and capsule (Bor¬
hidi, 2003), occur in various combinations in other
species of Arachnothryx, thus representing ex¬
tremes in a continuum of variation rather than
unique discrete character states. For these reasons,
Javorkaea is here considered as a synonym of Ar¬
achnothryx and the necessary combination is made
below for R. megalantha. Borhidi (2004: 101) sim¬
ilarly conceded that it was not possible to maintain
Javorkaea and transferred Rondeletia scabra Hems¬
ley ( Javorkaea scabra (Hemsley) Borhidi) to Arach¬
nothryx.

f. Arachnothryx megalantha (Lorence) Lor¬
ence, comb. nov. Basionym: Rondeletia me¬
galantha Lorence, Novon 4: 132. 1994. Ja¬
vorkaea megalantha (Lorence) Borhidi, Acta
Bot. Hung. 40: 16. 1996-1997. TYPE: Hon¬
duras. Cuyamel. 23 Aug. 1924, M. A. Carleton
653 (holotype, US 1208409; isotype, US).

See the discussion under Arachnothryx hondu-
rensis, above.

5. Rogiera standleyana (A. R. Molina) Lorence,
comb. nov. Basionym: Rondeletia standleyana
A. R. Molina, Ceiba 1: 262. 1951. Arachnoth¬
ryx standleyana (A. R. Molina) Borhidi, Acta
Bot. Hung. 33: 302. 1987. TYPE: Honduras.
Morazan, in pine forest above Zambrano, 1200
m, 20 July 1949, L 0. Williams & .4. Molina
R. 14417 (holotype, EAP not seen; isotypes, F,
US [2]).

Based on its pentamerous flowers and corolla with
internally densely yellow-barbate lobes and throat
Rondeletia standleyana clearly belongs to Rogiera
(see generic key above), not Arachnothryx to which
Borhidi (1987) incorrectly assigned it. Rogiera stan¬
dleyana is closely related to R. nicaraguertsis (Oer¬
sted) Borhidi, which differs by its leaves with the
lamina adaxially hirtellous-pilosulous and abaxially
strigillo.se and stipules 4—5 mm long, in contrast to
being villous on both surfaces and w ith stipules only
1—3 mm long in R. standleyana.

Acknowledgments. Thanks are extended to Anna
Stone for skillfully drawing the botanical illustration.
Roy Gereau (MO) for kindly supplying the Latin di¬
agnosis. Charlotte Taylor (MO) for constructive com¬
ments on the manuscript, and the curators of tin*
herbaria cited herein for access to their collections.

Literature Cited

Borhidi, A. 1982. Studies in Rondeletieae (Rubiaceae) III.
The genera Rogiera and Arachnothryx. Acta Bot. Hung.
28: 65-7 1 .

- . 1987. Studies in Rondeletieae (Rubiaceae) X.

New combinations ol Central American taxa. Acta Bot.
Hung. 33: 301-303.

- . 1997a. Studies in Rondeletieae (Rubiaceae),

XIII. New combinations of Mexican and Central Amer¬
ican plants. Acta Bot. Hung. 40(1—4): 15—16.

- . 1997b. Two new1 species of the genus Javorkaea

Borhidi et J. Komlodi (Rubiaceae). Acta Bot. Hung
40(1-4): 17-20.

- . 2001. Additions and corrections In the "Nomen-

clator of Mexican and Central American Rubiaceae” of
I). H. Lorence. Acta Bot. Hung. 43: 37—78.

- . 2003. Kspeeies nuevas de los generos Arach¬
nothryx y Javorkaea (Rubiaceae, Rondeletieae) en la
flora de Mexico. Acta Bot. Hung. 45: 1 — 11.

- . 2004. El Rondeletia complejo en Mexico. Acta

Bot. Hung. 46: 91—135.

- & M. Fernandez Zequeira. 1981. Studies in Ron¬
deletieae (Rubiaceae) 1. A new genus: Roige.Ua. Acta
Bot. Hung. 27: 309-312.

- & M. Jarai-komlodi. 1983. Studies in Rondele¬
tieae (Rubiaceae). IV. A new' genus: Javorkaea. Acta
Bot. Hung. 29: 13-27.

Kiehn, M. 1995. Chromosome survey ol the Rubiaceae.

Ann. Missouri Bot. Card. 82: 398—108.

Lorence, D. H. 1991. New species and combinations in
Mexican and Central American Rondeletia (Rubiaceae).
Novon 1: 135—157.

Mabberley, I). J. 1997. The Plant-Book: A Portable Dic¬
tionary of the Vascular Plants, 2nd ed. Cambridge Univ.
Press, Cambridge.

Rova, J. E., P. Delprete, L. Andersson & V. A. Albert.
2002. A irnL-F cpDNA sequence study on the Conda-
mineeae— Rondeletieae— Si paneeae complex with impli¬
cations on the phylogeny of the Rubiaceae. Amer. J.
Bot. 89: 145-159.

Standley, P. C. 1918. Rubiaceae. In North American Flora
32: 1-86.

Williams, L. 0. 1973. Tropical American Plants. X 1 \ . Ron-
deletiae Centro-americanae. Phytologia 26: 127—130.

Estudios en las Apocynaceae Neotropicales XVI:

Una Nueva Combinacion en Mandevilla para Sur America

./. Francisco Morales

Institute* Naeional de Biodiversidad (IINlIio), Apartado 22-3100, Santo Domingo, Heredia,

Costa Rica. fmorales@inhio.ae.er

Resi MEN. El estudio de las colecciones tipo dc
los diferentes taxones del genero Mandevilla I ind-
ley (Apocynaceae), ha determinado que Echites spe-
ciosum Kunth, debe ser excluido de la sinonimia
de M. snbsagittata (Rui/ Pavdn) Woodson. Por
lo tanto, sc establece una nueva combinacion, Man¬
devilla speciosa (Apocynaceae, Apocynoideae, Me-
sechiteae), y M. fendleri se reduce a su sinonimia.
Adicionalniente, se designa un neotipo para Mito-
ztis discolor Miers.

Absih \ci . t he study ol die type collections from
the different taxa in the genus Mandevilla Lindley
(Apocynaceae) has established that Eddies specio-
sum must be excluded from the synonymy ol M.
snbsagittata (Ruiz & Pavdn) Woodson. Therefore, a
new combination, Mandevilla speciosa, is estab¬
lished. and M. fendleri is included in synonymy. A
neotype is designated for Mitozns discolor Miers.

Key words: Amblyanthera, Apocynaceae, Apo¬
cynoideae, Echites, Mandevilla. Mesechiteae, Neo-
tropics.

Mandevilla Lindley (Apocynaceae, Apocyno¬
ideae), es el genero mas grande de la tribu Mese¬
chiteae, con eerca de 145 especies distribuidas
desde Mexico hasla el Norte de Argentina y las
Antillas (Morales, datos sin publ.). La ultima mono-
graffa lue liecha por Woodson (1933), qiiien reeon-
oeib 1 OJi taxones. organizados en dos subgeneros y
cineo seeciones. Sin embargo, este trabajo se de-
sactualizd en forma relativamente rapida por la
subsiguiente deseripcidn de varies taxones en las
dos decadas posteriores (e.g., Woodson, 1936,
1939, 1948). Durante el proceso de elaboracidn de
una nueva monograffa de Mandevilla, ha sido ne-
cesaria la revision de la mayorfa de los tipos de los
nombres hasta ahora publicados, con el fin de re-
afirmar la validez de las especies aceptadas por la
ultima monograffa del genero (Woodson, 1933). Tra-
bajos recientes (Morales. 1998; Sales, 1993, tesis
sin publ., Univ. Est. Campinas, Sao Paulo), ban
aclarado en forma pareial, la numerosa sinonimia

prevaleciente dentro del genero para algunas re-
giones geograficas en Mesoameriea y Brasil respec-
tivamente. Como parte de la revision de espeef-
menes tipo en varies herbarios europeos, fueron
revisados la totalidad tie las colecciones deposita-
das en el herbario de Humboldt y Bonpland en
Pans (P-HB). Como resultado, una nueva combi¬
nacion y nueva sinonimia son establecidas aquf.
Los datos de distribucidn geografiea y altitudinal
aca citados fueron tornados de la nueva monograffa
de Mandevilla (Morales, datos sin publ.), por lo que
pueden diferir notablemente de los publicados an-
teriormente por Woodson (1933). Asimismo, las
partes de la corola infundibulilormes siguen lo pro-
puesto por Morales y Fuentes (2004a).

MuiHlcvillu speciosa (Kunth) J. F. Morales, comb,
nov. Basonimo: Echites speciosum Kunth, en
IIBK. Nov. Gen. Sp. 3: 219. 1818 1 1819]. Exo-
thostemon speciosum (Kunth) G. Don. (fen.
Syst. 4: 82. 1838. TIPO: Venezuela. Datos per-
didos (fl. fr). A. Humboldt s.n. (holotipo, P-HB,
fotograffas. INB).

Mandevilla fendleri (Muller Argoviensis) Woodson. Ann.
Missouri Hot. Card. 19(1): 7(1. 1932. Syn. nov. Am-
blvanthera fendleri Miiller Argoviensis. Linnaea 30:
117. I860. TIPO: Venezuela. Merida: eerca de Co-
lonia Tovar, 1857 (fl). A. Eendler 1032 (holotipo, C-
1)C, fotograffa F neg. 26875 en INB \ MO; isotipos,
HK. F, K [en parte], MO, NY).

Mitozus discolor Miers, Apoeyn. S. Amer. 224. 1 878.
TIPO: Venezuela. Local idad y fecha perdida, (II). ,/.
Moritz 1209 (holotipo, BM perdido); Venezuela. l,o-
ealidad y feeha perditla, (II). ,/. Moritz 1392 (neotipo,
seleceion; ido aquf. BM, fotograffa en INB).

Distribucidn. Colombia, Venezuela, Guyana y
el Norte de Brasil, en elevaciones de 800—2250 m.

El basonimo tie Mandevilla speciosa fue incluido
por Woodson (1933, 1936) tie manera incorrecta
bajo la sinonimia de M. snbsagittata (Ruiz & Pa¬
vdn) Woodson, ya que esta ultima especie se car-
acteriza por sus cttrolas estrictamente hipocrateri-
formes, mientras que M. speciosa tiene eorolas
claramente infuntlibuliformes, con la parte superior
del tubo conspicuamente eampanulada. Es impor-

Novon 15: 152—454. Pi bushed on 21 Skitkmblh 2005.

Volume 15, Number 3
2005

Morales

Mandevilla para Sur America

453

tante aclarar que dentro del subgenero Exothoste-
mon, raenos de 10 taxones tienen corolas estricta-
mente hipocraterdormes. En todo caso, basado en
los espeefmenes examinados citados por Woodson
(1933), se infiere que el no examino el tipo de E.
speciosum y es probable (jiie su propuesta de si-
nonimizar este taxon fue basada solamente en la
descripcion del prologo original.

Ahora bien, el estudio de los espeefmenes tipo
de Echites speciosum Kunth y Mandevilla fendleri
(Muller Argoviensis) Woodson ha determinado que
ambas colecciones son similares. Estos taxones
pertenecen al subgenero Exothostemon (Woodson,
1933), grupo que se caracteriza por sus laminas
foliares cou eoleteres a lo largo del nervio central
adaxialmente, asf como corolas con el tubo algo
giboso basalmente. Eas bracteas florales de ambos
taxones son escariosas, compartiendo ademas el
mismo tipo (en forma y dimensiones) de corola,
siendo practicamente identicas en el resto de ear-
acteres morfologicos. Aunque la coleccion tipo de
M. fendleri tiene hojas con un indumento mas den¬
se en la superficie abaxial en relacidn al tipo de E.
speciosum. es importante recordar la alta variabili-
dad de ese caracter dentro del genero Mandevilla ,
lo cual lue comentado en forma previa por Morales
y Fuentes (2004b), por lo cual, no se justifica el
mantener como separados ambos taxones ni siqui-
era a nivel de variedad.

Woodson (1933) relego Mitozus discolor Miers y
Mandevilla linearis N. E. Brown bajo la sinonimia
de M. fendleri. Sin embargo, de estos dos taxones,
unicamente el primero es coespecffieo con los tipos
de Echites speciosum y M. fendleri. Miers (1878)
dcscribid Mitozus discolor y designo a Moritz 1299
como el tipo. Sin embargo, este especimen no pudo
ser localizado en el herbario de Miers en el Museo
Britanico (BM), ni tampoco duplicados adicionales
pudieron ser localizados en los principales lierba-
rios europeos. El unico especimen anotado por
Miers como Mitozus discolor lleva el numero 1892
y se encuentra depositado en el Museo Britanico
(BM), siendo concordante con la descripcion ori¬
ginal dada por el. Este especimen no difiere de los
espeefmenes tipo tie Echites speciosum v M. fend¬
leri. . compartiendo bojas con caracteres similares,
tanto morfologicamente como en la presencia y tipo
de indumento, asf como inflorescencias con brac¬
teas escariosas y corolas infundibuliformes. Ante la
perdida del bolotipo y la ausencia de duplicados
adicionales, la coleccion Moritz 1R92 es designada
como el neotipo.

Eos dos sintipos (F. McConnell & ./. Ouelch 122
y 194) de Mandevilla linearis N. E. Br. citados por
Brown (1901), representan un taxon con hojas li-

neares, (|ue no es affn con el concepto de 17. spe-
ciosa. For su parte, esos sintipos se encuentran re-
lacionados con un complejo de taxones (algunos de
ellos aim sin describir) del subgenero Exothoste¬
mon, que comparten en comun bojas lineares. Dado
que dicbo complejo se encuentra en estudio en este
momento, el estatus final de M. linearis no sera
definido por ahora.

EspecXmenes seleccionados examinados. COLOMBIA.
Antioquia: Ituango, carretera La Granja-Sanla Rita de
Ituango, 12 May. 1988 (st), Zarucchi & Betancur 6 472
(HLA. MO. USF). Boyaca: Alto Cliap6n, NO de Bogota,
3 Jun. 1932 (fl), Lawrence 143 (K MO. US). Cane a:
parque nacional Munchique, El Tambo, vereda La Ro-
melia, 24 Jul. 1993 (fl, fr), Velayos et al. 7010 (COL. MA).
Cundinamarca: estacidn Santana, cerca de Sasaima, Jul.
1945 (fl), Dugand A' Jaramillo 3810 (COL. US). Huila:
La Argentina, 25 Sep. 1984 (fl, fr). lozann et al. 4124
(COL. W IS). Magdalena: entre Pueblo Viejo y San Mi¬
guel. Jul. 1932 (II). Seifriz 369a (MO, US). Nariiio: car¬
retera Narino— Sonsdn, 7 Mar. 1987 (fr, fr), Zarucchi &
Redoya 4542 (COL, HUA. MO, USF). Quindfo: Armenia.
30 Oct. 1983 (fl), Arheldez et al. 228 (HUA, HUO). San¬
tander: entre Pie de Cuesta y Las Vegas. 19-24 Die.
1926 (fl). Killip & Smith 16161 (US). Tolinia: San Agus-
tfn, 28 Mar. 1899 (fl), Sprague 325 (k. US). Valle: Cor¬
dillera Occidental, Hoya de Alban. 26 Oct. 1946 (fl). Cua-
t recast/ s 22576 (COL, F, MO. VALLE). VENEZUELA.
Aragua: entre Guamitas y La Mesa, parque nacional
Henry Pittier, 20 May. 1981 (fr), Rodrigue: et al. 1276 (E.
MY). Bolivar: Gran Sabana, cerca de no Yuruani, 31
Mar. 1988 (fl), Hernandez et al. 4 (M. I’, VEN). Lara:
Moran, carretera Humocaro Alto a Buenos Aires, 19 Jul.
1984 (fl, fr), Agostini et al. 2805 (MO). GUYANA. Region
Potaro-Siaruni, Paramakatoi, 13 Mar. 1989 (fl. fr), Hahn
el al. 5652 (CAY. k. US). BRASIL. Amazonas: Surumu.
Sep. 1909 (fl), Ule 8450 (G [2 larninas|, k).

Agradecimientos. Quiero agradecer a los cura-
dorcs <lc los herbarios BM, BR, E. G, k, MO, NY
y P por el envfo de material en prestamo, asf como
por permitirme el uso de sus colecciones. Tambien
quiero reconocer la ayuda de Michael Grayum
(MO) en la interpretacion del codigo de Nomeni da¬
tura Botanica. Se agradece la revision crftica del
manuscrito de Bruce Hansen y un revisor andnimo.

Literatura Citada

Brown, N. E. 1901. Report on two botanical collections
made by Messrs. F. V. McConnell and .1. ,1. Ouelch at
Mount Roraima in British Guiana. Trans. Linn. Soc.
London, Bot. Ser. 2. 6: 1 — 107.

Miers, J. 1878. On the Apocynaceae of South America.

Williams & Norgatc, London, Edinburgh.

Morales, ,1. F. 1998. A synopsis of the genus Mandevilla
(Apocynaceae) in Mexico and Central America. Britton-
ia 50: 214-233.

- & A. Fuentes. 2004a. Estudios en las Apocyna¬
ceae Neotropicales VIII: Nuevas especies de Mande¬
villa para Peru y Bolivia, con notas sobre la morfologfa
floral en corolas infundibuliformes. Candollea 59(1):
167-174.

454

Novon

- & - . 20()4b. Estudios en las Apocynaceae

Neotropieales VI: Una nueva espeeie, nuevos reporles
y nueva sinonimia en las Apocynaceae de Bolivia. Sida
21: 165-174.

Woodson. B. E. 1953. Studies in the Apocynaceae. IV.
The American genera of Echitoideae. Ann. Missouri
Bot. Card. 20: 605—790.

- . 1936. Studies in the Apocynaceae. IV. The

American genera of Echitoideae. Ann. Missouri Bot.
Card. 23: 169-438.

- . 1939. New or otherwise noteworthy Apocynaceae

of tropical America. VI. Ann. Missouri Bot. (lard. 26:
95-98.

- . 1948. Miscellaneous new Apocynaceae and

Aselepiadaceae. Ann. Missouri Bot. (lard. 35: 233—
238.

Alsmithia Subsumed in Heterospathe (Arecaceae, Arecoideae)

Maria Vi hr Norup

Department of Systematic Botany, University of Aarhus, Build. 540. Ny Munkegade,
8000 Aarhus C., Denmark, and Herbarium, Royal Botanic Gardens, Kew,
Richmond TW9 3AE, United Kingdom, m.norup@kew.org

Abstract. The palm genus Alsmithia H. E.
Moore (Arecaceae, Arecoideae), with a single rep¬
resentative from Fiji, is included in synonymy with
Heterospathe Scheffer as //. longipes (H. E. Moore)
Norup. The species is distinctive within Hetero¬
spathe in having an elaborately sculptured endo-
earp. Alsmithia and Heterospathe share several
characters, most notably the lack of a erownshaft.
persistent peduncular bracts and prophyll. a com¬
pletely encircling prophyll. interfoliar inflorescenc¬
es, and an elongate peduncle. However, the char¬
acters are hornoplasious and are only diagnostic in
combination. The taxonomic transfer is well sup¬
ported by molecular phylogenetic studies based on
two low-copy nuclear DNA markers, PRK and
RPH2.

Key words: Alsmithia, Arecaceae. Areceae,
Arecoideae, endocarp sculpture, Heterospathe,
Indo-Pacific region.

The Fijian Alsmithia was described by Moore et
al. (1982) with the single species A. longipes H. E.
Moore. The monotypic genus is defined by the com¬
bination of long petioles, a highly extended pedun¬
cle, absence of a erownshaft, a completely encir¬
cling base of the prophyll at insertion, and an
irregularly shaped seed enclosed in a highly sculp¬
tured endocarp formed by thick fibers. The com¬
pletely encircling base of the prophyll is a feature
of particular diagnostic importance since it distin¬
guishes Alsmithia from Cyphosperma II. Wendland,
another Fijian genus with which Alsmithia might
be easily confused.

Heterospathe was erected by Scheffer ( 1876), and
although the taxonomic history of Heterospathe is
intertwined with several other genera, it is main¬
tained as a distinct genus in the latest formal clas¬
sification of the palms (Uhl & Dransfield. 1987). In
its present circumscription it accommodates 38
species, with species diversity centered in Newr
Guinea, but the genus extends into the western part
of the Pacific, particularly the Solomon and Phil¬
ippine Islands, and the Moluccas (Govacrts N
Dransfield. 2005; Uhl & Dransfield. 1987). The fol¬

lowing characters define the genus: (1) an interfol¬
iar inflorescence w ith a prominent peduncle, (2) ab¬
sence of a erownshaft, (3) typically the presence of
a single (two in H. trispatha Fernando) prominent
and persistent peduncular bract longer than the
persistent prophyll (but see also Norup et al., in
prep.).

The complex taxonomic history of Heterospathe
and its perceived lack of noteworthy morphological
characters have raised concerns that morphology is
not sufficient to resolve relationships within tribe
Areceae to which both Alsmithia and Heterospathe
belong. Therefore recent efforts have been concen¬
trated on a molecular systematic approach. By com¬
bining data from a number ol nuclear DNA mark¬
ers, it has been possible to obtain well-supported
phylogenetic hypotheses for the tribe (Baker & Eoo.
2004; Eewis Doyle, 2002; Norup et al., 2003).
The results show that in order for the classification
to reflect phylogenetic relationships a number of
nomenclatural changes in tribe Areceae are nec¬
essary.

The latest molecular systematic study of the
Indo-Pacific Areceae members (Norup et al., in
prep.) uses two low-copy nuclear loci, nitron 4 of
the gene encoding the Calvin cycle enzyme phos-
phoribulokinase (PRK) and intron 23 of the gene
for RNA-polymerase II. subunit B f RPR2). The re¬
sults obtained from both the analyses of the genes
separately and in combination show that Alsmithia
is embedded within a strongly supported Hetero¬
spathe (more than 95% bootstrap support; Norup et
al., in prep.). Two accessions of Alsmithia longipes
{ Raker IIR0 , FTG; Zona 10.19, FTG) were se¬
quenced independently to verify the DN A sequenc¬
es and ensure that this remarkable relationship was
not an artifact or a technical error.

The close relationship with Heterospathe is sur¬
prising because Moore et al. (1982) thought Alsmi¬
thia to be quite distinct from other members of the
Clinostigma alliance (named after Clinostigma H.
Wendland), an informal group erected by Moore
(1973). Based on the unusual seed shape and
sculpturing of the endocarp, Moore et al. (1982)

Novon 15: 455-457. Published on 21 September 2005.

456

Novon

suggested a possible relationship with Hurretio-
kentia Piehi Sermolli, Veillonia II. E. Moore, Cy¬
phosperma, and Physokentia Beccari, all of which
are distributed in the western Pacific region (New
Caledonia and Fiji). In their discussion of putative
relationships, Moore et al. (I *452: 124) particularly
noted “a striking habitual resemblance” between
Alsmithia and Cyphosperma, but no reference to
Heterospathe was made. It is, however, worth noting
that when Alsmithia was erected in 1982, the east¬
ernmost record of Heterospathe was in the Solomon
Islands (//. salomonensis Beccari), and it was not
until 15 years later that a new species of Heteros¬
pathe was described from Fiji as //. phillipsii I).
Fuller <X Dowe, in Fuller et al. (1997). The earliest
record of other West Pacific species of Heterospathe
was made just one year earlier, when H. uniformis
Dowe was described from Vanuatu (Dowe & Ca-
balion, 1996).

Both Alsmithia and Heterospathe show morpho¬
logical affinities with many of the other Indo-Paeifie
arecoid genera, thus the molecular data have been
essential in pointing to a putative relationship be¬
tween the two genera. The new well-supported mo¬
lecular phylogeny makes it possible to evaluate
critically the morphology of these two genera (No-
rup et al., in prep.). Alsmithia shares a number of
features with Heterospathe, notably (1) habit small
to moderate, solitary |or sometimes clustering in
Heterospathe], (2) lack of a crownshaft (leaf sheath
splitting to the base), (3) persistent peduncular
bracts and prophyll, (1) completely encircling pro¬
phyll, (5) interfoliar inflorescences (at least at an-
thesis), (6) an elongate peduncle, (7) androecium of
six stamens (stamens ranging from six to many in
Heterospathe), (8) awl-shaped filaments, and (9) ob¬
long. dorsifixed anthers that are indexed in bud
(Moore et al.. 1982; Uhl & Dransfield, 1987). Most
notably, the peduncular bract of Alsmithia greatly
exceeds the persistent prophyll in length just as in
Heterospathe, where this is considered a defining
character of the genus. Many of the characters men¬
tioned above are shared with a great number of
arecoid palms, and Heterospathe, including Alsmi¬
thia, is shown to be distinguished by a combination
of homoplasious characters rather than by unique
synapomorphies (Norup et al., in prep.).

The striking endocarp sculpturing and seed
shape of Alsmithia are quite unknown in Hetero¬
spathe and are undeniably reminiscent of the gen¬
era liurretiokentia, Veillonia, Cyphosperma, and
Physokentia. However, morphological studies have
noted some similarity between the pericarp com¬
position in Alsmithia and Heterospathe, in that they
both possess an expanded series of vascular bun¬

dles that is uncommon in other closely related gen¬
era (Essig et al., 1999). Optimization of ornate en-
docarps on one of the most recently assembled
molecular trees (Norup et al.. in prep.) strongly sug¬
gests that such endocarps have developed repeat¬
edly within the tribe Areceae. In the present ex¬
ample. the placement of a species (//. longipes) with
ornate endocarps in a genus with otherwise smooth
endocarps is indeed interesting. The sculptured en¬
docarp is probably an ecological adaptation, but
until more field observations are available we can
only speculate as to the underlying evolutionary
driving forces. It would also be valuable to estab¬
lish through developmental studies ii the origin of
the endocarp sculpturing is the same in all species
that display this particular feature. A second no¬
table character that differs between Alsmithia and
Heterospathe (sensu Uhl Dransfield. 1987) is the
endosperm condition, which is homogenous in the
first and ruminate in the latter. The homogenous
condition, however, was recently described in II.
uniformis (Dowe Cabalion, 1996). Furthermore,
variation in endosperm condition, both infrageneric
and infraspecific, is known to occur in several palm
genera (e.g.. Zona, 2003) and is a highly labile con¬
dition (Norup et al., in prep.).

Based on the strength of the molecular evidence
and the morphological similarities described above,
Alsmithia is placed in synonymy in favor of a mono-
phyletic Heterospathe. Here the necessary new
combination is made, which brings the number of
species of Heterospathe known from Fiji to two.

Heterospathe Scheffer, Ann. Jard. Bot. Buitenzorg
I: 141. 1876. TYPE: Heterospathe elata Schef¬
fer.

Alsmithia H. K. Moore, Principes 26: 122. 1982. Syn. nov.
TYPE: Alsmithia longipes II. E. Moore | = Hetero¬
spathe longipes (H. E. Moore) Norup].

Harkerwebbia Beccari, Webbia I: 281. 1905. TYPE: Har-
kerwebbia elegans Beccari | = Heterospathe elegans
(Beccari) Beccari |.

Ptyehandra Scheffer, Ann. Jard. Bot. Buitenzorg 1: 140.
1876. TYPE: Ptyehandra glauca Scheffer | = Het-
erospathe glauca (Scheffer) H. E. Moore|.

Heterospathe longipes (H. E. Moore) Norup.
comb. nov. Basionym: Alsmithia longipes H. E.
Moore, Principes 26: 124. 1982. TYPE: Fiji.
Taveuni: wet forest on trail along ridge from
Wainibau River to mountain crest on E side* of
Taveuni beyond Lavena village, 325 m. 25
Mar. 1980, H. E. Moore Jr„ H. //. Phillips &
S. Vodonaivalu 10545 (holotype, BH; isotype.
K).

Specimens examined. FIJI. Collected directly from

Volume 15, Number 3
2005

Norup

Alsmithia Subsumed

457

wild, origin unknown, cultivated at FTG, Baker 1180
(FTG); collected directly from wild, origin unknown, cul¬
tivated at Floribunda Palms and Exotics nursery, Hawaii,
Zona 1080 (FTG); Taveuni, II. K. Moore Jr.. R. II. Phillips
& S. Vodonaivalu 10545 (K).

Acknowledgments. I am most grateful to Jeff
Marcus (Floribunda Palms and Exotics nursery,
Hawaii) for swiftly providing a specimen of die sec¬
ond Alsmithia accession, and lo Carl Lewis (T IG)
for generating an extra set of sequences lo verify
the Alsmithia sequences produced in Kew. My
studies at the Royal Botanic Gardens, Kew. were
enabled thanks lo economic support from the ( Mi-
con Foundation and the Faculty of Science, Uni¬
versity of Aarhus. ON A sequencing was, in part,
funded by grants from the Royal Society, the Fev-
erhulme Trust, and the British American Tobacco
Biodiversity Partnership through the Palms of New
Guinea project. Sincere thanks go to Anders Barfod
(AAU) for invaluable support, supervision, and crit¬
ical comments on the manuscript; and to my su¬
pervisors at the Royal Botanic Gardens, Kew. John
I Mansfield and Bill Baker, who introduced me to
palms and provided great support and many useful
comments. Also thanks to Scott Zona (FTG) and
Victoria G. Hollowell (MO) for thorough and con¬
structive reviews of the submitted manuscript.

Literature Cited

Baker, W. J. & A. Loo. 2004. A synopsis of the genus
Hydriastele (Arecaeeae). Kew Bull. 59: 61—68.

l)owe, .1. L. & P. Cabalion. 1996. A taxonomic account ol
Arecaeeae in Vanuatu, with descriptions of three new
species. Austral. Syst. But. 9: I -60.

Kssig, F. B.. T. J. Manka & L. Bussard. 1999. A systematic
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Fuller, I)., J. I). Powe & M. F. Doyle. 1997. A new species
of Heterospathe from Fiji. Principes 41: 65—69.

Govaerts, R. & ,|. I Mansfield. 2005. World Checklist ol
Palms. Royal Botanic Gardens, Kew.

Lewis, C. F. & J. ,1. Doyle. 2002. Phylogenetics of tribe
Areceae (Arecaeeae) using low-copy nuclear genes. PI.
Syst. Fvol. 236: 1-17.

Moore, H. E., Jr. 1973. The major groups of palms and
their distribution. Gentes Herb. 1 1(2): 27—140.

- . R. II. Phillips & S. Vodonaivalu. 1982. Additions

to the palms of Fiji. Principes 26: 122—125.

Norup. M. V., W. Baker, C. Lewis, A. Loo & .1. Dransfield.
2003. Phylogenetic and biogeograph ieal studies of the
ludo-Pacific clade of tribe Areceae (Palmae), based on
two low-copy nuclear genes. Poster presented at Mono-
cots 111. Rancho Santa Ana Botanical Gardens, Califor¬
nia.

Scheffer, R. H. C. C. 1876. Sur quelques palmiers du
groupe des Arecinees. Deuxieme parte. Ann. Jard. But.
Buitenzorg 1: 103—164.

Uhl, N. W. & J. Dransfield. 1987. Genera Palmarum: A
Classification of Palms Based on the Work of II. E.
Moore Jr. International Palm Society and L. II. Bailey
Hortorium, Lawrence, Kansas.

Zona, S. 2003. Endosperm condition and the paradox of
Ptychococcus paradoxus. Telopea 10: 1-7.

Two New Species of Hypericum (Clusiaceae) from Korea

Seon-Joo Hark

Department of Biology, Yeungnam University, Kyeungsan. Kyeungbuk 712-749, Korea.

sjpark01@yu.ac.kr

Ki-Joong Kim *

Division of Life Sciences and Biotechnology. Korea University, Seoul 136-701, Korea.

kiinkj@korea.ac.kr
^Corresponding author

Abstract. Two undescrihed species of Hyperi¬
cum (Clusiaceae) have been found during florist ic
research in Korea. Hypericum chejuense S.-J. Park
<A K.-J. Kim and //. jeongjocksanen.se S.-J. Park »A
K.-J. Kim are described and illustrated from Cheju
Island in southern Korea and Jeongjock Mountain
in southeastern Korea, respectively. Hypericum che¬
juense and II. jeongjocksanense are easily distin¬
guished from the closely related II. erect um Tliun-
herg ex Murray and //. laxum (Illume) Koidzumi,
respectively, by their habit, petal and sepal size,
inflorescence type, style length, stylar hair distri¬
bution. leal base shape, stem branching pattern,
sepal size, and DNA sequence from the nuclear ITS
regions.

Key words: Clusiaceae. Hypericum, Korea.

Hypericum L. is the largest genus of the family
Clusiaceae (Guttiferae). The cosmopolitan genus
contains about 370 species distributed globally ex¬
cept for arctic and desert areas and the lowland
tropics (Mabberley, 1997). Some Hypericum species
are valuable for ornamentals (e.g., //. inodorum
Miller) and medicinal uses (e.g., //. perforatum L
and II. androsaemum L), and they have become
the subjects of extensive investigation (Hennessy et
al., 2002; Pepping, 1999). Hypericum is classified
into 30 sections based on the character combina¬
tions of habit, stem shape, glandular trichome type,
petal duration, style number, stamen fascicle num¬
ber, plaeentation type, and basic chromosome num¬
ber (Hobson, 1977; Robson <A Adams, 1981).
Twelve species of Hypericum belong to three sec¬
tions and are distributed nearly throughout the Ko¬
rean peninsula, with these distribution ranges ex¬
tending into Japan and China. However, the two
new species, II. chejuense S.-J. Park cA K.-J. Kim
and H. jongjocksanen.se S.-J. Park cA K.-J. Kim. are
only known from limited areas in southern Korea.
Hypericum chejuense and //. jongjocksanen.se are

Novon 15: 458-463. Published on 21 Sep

similar to //. erection Thunberg ex Murray and II.
laxum (Blume) Koidzumi. respectively, both in their
morphology and DNA sequence similarity. The new
species are easily distinguished by their habit, pet¬
al size, inflorescence type, style length, style hair,
leaf base shape, stem type and branching pattern,
sepal size (cf. Table I, Figs. I. 2). and DNA se¬
quences (Park <A Kim. 2004).

Hypericum chejuense S.-J. Park «A K.-J. Kim, sp.
nov. I 'l PK: Korea. Chejudo Prov.— Cheju Is¬
land, Seogwipo Natural Forest, 33°23'N.
126°35'E, 5 Sep. 2000 (fl), S.-J. Hark & K.-J.
Kim 21)011 (holotype, YNUH; isotype, KUS).
Figure 1.

Ilaee species inter eongeneros koreanos ad Hypericum
sectionem Hypericum pertinentes quoad eaulem glanduhs
atratis destitum et capsulam late usque anguste ovoideam
ad Hypericum erection inaxime aecedil, sed ah eo habitu
suffrutieoso, eaule ascendente 10—20 cm alto, foliis basi
acutis atque petalis 4.2-5. 1 mm longis distinguitur.

Subshrub, 10—20 cm tall, decumbent from
rooting base; 5- to 8-branehed; stems rounded,
eglandular. Leaves sessile; blades narrowly tri¬
angular-ovate to narrowly elliptic, 1.52—2.01 X
0.87—1.21 cm; laminar glands black, numerous,
small; intramarginal glands black; base acute;
margin entire, plane; apex obtuse to rounded;
main lateral veins 3- or 4-paired; tertiary veins
densely reticulated. Inflorescences 1- to 3-flow-
ered from 1 to 3 nodes, the whole flat-topped to
cylindric: bracts and bracteoles reduced, leaf¬
like. margin entire. Flower ca. 0.7— 1.0 cm diam.,
substellate; buds narrowly ellipsoid, apex acute.
Sepals free, erect, ovate-lanceolate, subequal to
unequal, ca. 3.2— 5.0 X 1.2— 1.0 mm; laminar
glands black, streaks and dots; intramarginal
glands black, sparse or absent; margin entire to
irregularly glandular-ciliate, apex obtuse to
acute; veins 5; petals bright yellow7, obovate to

IEMUER 2005.

Volume 15, Number 3
2005

Park & Kim

Hypericum (Clusiaceae)

459

1 mm

Figure ]. Hypericum chejuense S.-J. Park & K.-J. Kim. — A. Habit. — IF Flower. — C. Leal blade. — I). Petal. — K
Sepal. - — F. Fruit. Drawn from the holotype, S.-J. Hark &• K.-J. Kim 20011.

oblong, 4.2— 5.1 X 1.2— 1.6 mm; margins densely
blaek-punctate, laminar glands black, streaks,
distal; marginal glands black, inframarginal to
prominent, irregular; margins entire; stamens ca.

2.2— 3.5 mm, 3-fascieled. Ovary ovoid; styles 3.
2.5— 3.2 mm, widely spreading. Capsule ovoid,
ca. 5.2— 6.3 X 2.9— 3.7 mm; seeds yellow-brown,
0.4— 0.6 mm; testa finely reticulate.

460

No von

Table 1. Comparison of diagnostic charade
II. laxum, and II. jeangjocksanense.

rs among the closely

related species Hypericum

erection. II. ehejuense.

< diameter

II. erect um

//. ehejuense

II. laxum

II. jeangjocksanense

Habit

herbs

subshrubs

herbs

herbs

Height (cm)

30-70

10-20

7-25

3-5

Petal length (mm)

7.1-15.2

4.2-5. 1

4.4-5. 2

1 .9-3. 1

Petal w idtli (mm)

1 .8-2.6

1 .2-1.6

1.1-1 .5

0.5-0. 7

Petal color

golden yellow

bright yellow

bright yellow or
orange

bright yellow

Inflorescence type

svmpodial

svmpodial

svmpodial

monop* »dial

f lower number

1 to 3 from apical

1 to 3 from apical

1 to 3 from apical

1 from apical node

or axillary nodes

or axillarv nodes

or axillary nodes

only

Style length (mm)

3.5— 4.0

2. 5-3. 2

2. 1-2.5

LI-1.5

Stylar hair

absent

absent

absent

present

l.eaf base

eordate-amplexicaul

acute

obtuse

obtuse

Leaf veins

4

4

1 to 7

1 to 2

Sepal \eins

5

5

3 to 5

3

Stem tvpe

erect

decumbent

erect

erect

Stem branches

1 to 2

5 to 8

1 to 2

1

Habitat and distribution. Open sunny area of
subtropical evergreen forests; growing on semi-wet
and sand-elay mixed soils. The new species is rare
and only known from the type locality of Cheju Is¬
land. Korea.

Phenology, flowering: June to September
(pers. obs.); fruiting: July to October.

Hypericum ehejuense belongs to section Hyperi¬
cum and appears to be closely related to II. erec-
tum. Hypericum ehejuense. however, can be differ¬
entiated easily from II. erection by its subshrubby
habit, basally decumbent stems, the usually shorter
petal lengths (el. Table I) and smaller and broader
capsules, more acute leaf bases, and shorter
heights. Hypericum erection is widespread through
northeastern Asia (Lee, 1979; Li el ah. 1990; Ohwi,
1965) and a more variable species assigned to sec¬
tion Hypericum subsect. Erecta N. Robson (Robson,
2001). Hypericum ehejuense also belongs to the
same subsection as //. erection on the basis of com¬
mon features.

Etymology. The new species is named for the
type locality, Cheju Island of Cheju Province. Ko¬
rea.

Hypericum jcoiigjocksaiicn.se S.-J. Park & K.-
J. Kim, sp. nov. TYPE: Korea. Kyeong-sang-
nam-do Prov.: the Upo swamp reserve of
Jeong-jock-shan, 550—650 m, 35°27'N,
I29°8'E. 27 Aug. 2000 (fl), S.-J. Hark & k-J.
kirn 20027 (holotype, YNLIH; isotypes. KLS.
MO), f igure 2.

Haec species inter congeneros koreanos ad Hypericum
seclionem Trigynobrathydem pertinentes quoad caulem er-
ectum decumbentemve, folia inferiora elliplica ovatave
patentia superiora ovata foliacea et sepala apice obtusa ad
Hypericum laxum maxime accedit, sed ab eo caule sim-
plici 3—5 cm alto, inflorcscentiae ramis monopodialibus,
pctalis 1.9— 3.1 mm longis atque stylo 1 .1 — 1.5 mm longo
basim versus trichomata gerente distinguitur.

Herbs, annual, 3—5 cm tall, erect; stem solitary,
unbranched below inflorescence. Stem 4-lined,
with scattered glandular dots. Leaves sessile; blade
usually ovate or ovate-trianglar to oblong, 0.22—
0.31 X 0.15—0.23 cm; papery, abaxially paler and
sometimes glaucous; laminar gland dots dense; in¬
tramarginal glands dense; base obtuse, margin
plane, apex obtuse; main lateral veins 1 to 2, basal;
tertiary reticulation not visible. Inflorescence I. ter¬
minal. Flower 0.15—0.3 cm diam.; bud cylindric-
ellipsoid, apex obtuse. Sepals free, erect, narrowly
oblong or rarely lanceolate to elliptic, subequal to
unequal. 0.7— 1.5 X 1.0— 1.2 mm; laminar glands
lined, distally dots; marginal glands absent; apex
obtuse to rounded; veins 3. Petals pale to bright
yellow, obovate to oblong, 1.9— 3.1 X 0.5— 0.7 mm;
laminar glands absent; margins entire, eglandular.
Stamens 5 to 30, irregular or in 5 obscure groups
when few. Ovary broadly ovoid to subglobose; styles
3, 1.1— 1.5 mm, spreading; stylar hairs present on
the lower portions. Capsule cylindric to globose,
1.1— 2.4 X 0. 7-1.2 mm, usually slightly shorter or
slightly longer than sepals. Seeds straw-yellow, ca.
0.2— 0.3 mm; testa finely linear-scalariform.

Habitat and distribution. The

new species

Volume 15, Number 3
2005

Park & Kim

Hypericum (Clusiaceae)

461

Figure 2. Hypericum jeongjocksanense S.-J. Park & K.-J. Kim. — A. Habit. — B. Flower. — C. Leaf blade. — I). Petal.
— Fi. Sepal. — F. Fruit. Drawn from tbe holotype, S.-J. Park & k.-J. Kim 2002/.

grows on the wet clay soil of the Mujaeehi swamp
reserve area of the Jeong-jock-shan of southeastern
Korea. The swamp area is maintained by spring
water and is a unique ecosystem in Korea. The

swamp ecosystem is designated as a protected area
by the Ministry of Environment, Korean Govern¬
ment. The new species is rare and endemic to the
swamp of Jeon-jock-slum.

462

Novon

Phenology. Flowering: Marc h to September
(pers. obs.); fruiting: May to October.

Hypericum jeongjocksariense belongs to section
Trigynohrathys (Y. Kimura) N. Hobson with //. lax-
um and //. japonicum. It is closely related to II.
laxum, differing from it essentially by the mono-
podial inflorescences. Fhe sizes of the* plants, pet¬
als. sepals, and seeds are smaller than those of//.
laxum (Table 1). Also, trichomes exist on the lower
portions of the style1 in H. jeongjocksariense. Hyper¬
icum laxum grows from lowlands to mountains up
to 2000 m. and its morphology is variable according
to its habitat. Hypericum jeongjocksariense has been
collected only in the limited lowland swamp areas
in Korea.

Etymology. The new species is named for llu*
type locality, Jeong-jock-shan of Kyeong-sang-nam-
do Province, Korea.

I he distribution of the two new species is lim¬
ited. Hypericum chejuense is distributed on Cheju
Island, in southernmost Korea. Hypericum jeong-
jocksanense is distributed in the* Upo swamp reserve
of tilt- Jeong-jock-shan of southeastern Korea. Both
species grow on wet soils. In addition to the exter¬
nal morphology, the nuclear ITS sequence data
(Park & Kim, 2004) differentiate the two new spe¬
cies from the closely allied species pairs. I’lie two
new species of Hypericum from Korea can be dis¬
tinguished from the known species by the following
key.

K I X TO I MF KoHK.VN Sl'KClKS (IK HWERta 1/

la. Styles 5-branched, rarely 4-l)ranchecl; stamen
fascicles 5; dark glands absent on floral parts

. H. ascyron L.

lh. Styles 3-branched, rarely 2- or 4-branehed; sta¬
men fascicles 3; dark glands usually present on
floral parts.

2a. Stems 4-ridged; leaves 0.2— 1.0 cm long; an¬
nual herbs; flowers 0.15—0.7 cm across; an¬
ther apex without black glandular dots; cap¬
sules without glands.

3a. Stems erect; lower leaves ovate to
broadly ovate, ascending; upper
leaves lanceolate to linear, often
bract-like; sepals truncate at apex . .

. //. japonicum Thunberg

3b. Stems erect or decumbent; lower leaves
elliptic or ovate, spreading; upper
leaves ovate, leaf-like; sepals obtuse at
apex.

4a. Stems usually branched, 7—25 cm
tall; inflorescence branches sympo-
dial; petals 4.4— 5.2 mm long; sty les
2. 1-2.5 mm long, stvlar hairs ab¬
sent ... II. laxum (Illume) Koidzumi
4b. Stems unbranched, 3—5 cm tall;
inflorescence branches monopodi-
al; petals 1.0— 3.1 mm long; styles

1.1-1. 5 mm long, stylar hairs pre¬
sent on the lower portions . . .

. H. jeongjocksariense

S.-J. Park & K.-J. Kim
2b. Stems terete or with 2 striations; leaves
1.5— 5.0 cm long; perennial herbs; flow¬
ers 0.7— 2.5 cm across; anther apex with
black glandular dots; capsules with
glands.

5a. Styles shorter than ovary .

. H. oliganthum

Franchet A Savatier
5h. Styles longer than or equal to ovary.

6a. Stems erect, with dark glands;
capsules broadly ovoid to nar¬
rowly conic .

. //. attenuation Choisy

6b. Stems erect to ascending, with¬
out dark glands; capsules
broadly to narrowly ovoid.

7a. Subshrubs; stems 10—20
cm tall, basally decum¬
bent; petals 4.2— 5.1 mm
long; leaf bases acute . . .

. II. chejuense

S.-J. Park & K.-J. Kim
7b. Perennial herbs; stems
30—70 cm tall, erect; pet¬
als 7.1-15.2 mm long; leal
bases obtuse .... II. erection
Thunberg ex Murray

Acknowledgments. The authors thank Bov Ger-
eau for the Latin diagnosis, Victoria Hollowed for
critical comments on the manuscript, two anony¬
mous reviewers for the critical readings of the man¬
uscript, and Hae-I.im Lee for die fine illustrations.

I ll is work was supported by grants (nos. 052-041-
026, 052-052-040) from the Core Environmental
Technology Development Project (or Next Genera¬
tion funded by the Ministry ol Environment of the
Korean Government.

I .iterature Cited

Hennessy, M., I). Kellecher, .1. P. Spears. M. Barry, P.
Kavanagh, 0. Back, F. Mulcahy & J. Feely. 2002. St.
John's Wort increases expression of P-glycoprotein: Im¬
plications for drug interactions. Brit. J. Clin. Pharmacol.
53; 75-82.

Lee, l.-B. 1979. Guttiferae. Pp. 544—546 in Illustrated
Flora ol Korea. Hyangmunsa, Seoul. |ln Korean.)

Li, H.-W., 5. -II. Li, S.-(,). Tong & < 7. - 1 ). Tao. 1990.
Hypericum. In Flora Reipublicae Popularis Sinicae,
Tomus 50(2): 1-72. Science Press, Beijing. [In Chi¬
nese.)

Mabberley, I). J. 1997. The Plant-Book: A portable dic¬
tionary of the vascular plants, 2nd ed. Cambridge l niv.
Press, New York.

Ohwi, J. 1965. Guttiferae. Pp. 630—632 in C. G. Meyer &
F. II. Walker (editors), Flora of Japan. Smithsonian In¬
stitution, Washington, D.C.

Park. S.-J. A K.-J. Kim. 2004. Molecular phytogeny of the
genus Hypericum (Clusiaeeae) from Korea and Japan:

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Park & Kim

Hypericum (Clusiaceae)

463

An evidence from nuclear rDNA ITS sequence data. .).
PI. Biol. 47: 366-374.

Pepping, J. 1999. St. John’s Wort: Hypericum perforatum.

Amer. J. Health.-Syst. Pharm. 56: 329-330.

Robson, N. k.. B. 1977. Studies in the genus Hypericum
L. (Guttiferae). I. Infrageneric Classification. Bull. Brit.
Mus. (Nat. Hist.) But. 5: 291-355.

- . 2001. Studies in the genus Hypericum L. (Gut-

liferae). 4(1) Sections 7. Roscyna to 9. Hypericum sensu
lato (Part 1). Bull. Brit. Mus. (Nat. Hist.) Bot. 31(2):
37-88.

- & P. Adams. 1981. Chromosome numbers in Hy¬
pericum and related genera. Brittonia 20: 95—106.

Notes on Grasses (Poaceae) for the Flora of China, IV

Sylvia M. Phillips

Herbarium. Boyal Botanic Gardens, Kew, Surrey TW9 3AB, United Kingdom.

s.|)hillips@kew.org

Chen Shouliang

Herbarium. Jiangsu Institute of Botany, Academia Siniea, I Qianghuhuocun, Nanjing,
Jiangsu 210014, China, herb_nas@yaboo.eom.en

Abstract. The following changes are made in
Poaceae, necessary for the upcoming Flora of Chi¬
na. Kengia chinensis is leetotypified on a specimen
of K. hackelii, and hence becomes a synonym of the
latter. Kengia gracilis is synonymized under K. mu-
vronata. Isachne clarkei is leetotypified. Isachne be¬
tter kei (also leetotypified) and /. tennis are placed
as synonyms of /. clarkei. Isachne alhens var. hir-
suta is leetotypified and placed as a synonym under
/. sylvestris. Isachne sernitalis is leetotypified and
placed in synonymy of /. schmidtii. The distinctions
between the closely related species /. kunthiana, I.
schmidtii, and I. repens are discussed. The new spe¬
cies Arundinella suniana , from Yunnan province
and related It) A. selosa, is described. Arundinella
setosa var. esetosa is validated. The name Arundi¬
nella liupeiensis is rejected as a homonym due to
its similarity to the name A. hubeiensis. Eccoilopus
batnbusoides is transferred to Spoil iopogon, neces¬
sary with the inclusion of Eccoilopus within Spo-
diopogon. The new species Imperata flavida is de-
sc ribed from Hainan, distinguished from /.
cyUndrica mainly by its much shorter spikelet
hairs. E riant has gri/fithii var. trichophyllus is trans¬
ferred to Saccharum arundinaceum as S. arundi-
naceum var. triehophyllum, as Erianthus is includ¬
ed within Saccharum. Polytrias amaura var. nano
is transferred to P. indica, the correct name for P.
amaura. Coelorachis striata var. pubescens is trans¬
ferred to Mnesithea. a necessary transfer when Coe¬
lorachis is regarded as a synonym of Mnesithea.
Discussion of the changes is provided throughout.

Key words: Arundinella. China, Cleistogenes,
Coelorachis , Eccoilopus , Erianthus, Eulalia. Imper¬
ata, Isachne, Kengia, Mnesithea, Polytrias, Sac¬
charum. Spodiopogon.

This paper continues the series for Poaceae in
Novon lor the publication of taxonomic novelties
and nomenelatural changes that have arisen during

the preparation of the grass family account for the
Flora of China.

Kl{ AGROSTIDK \l\

Kengia hackelii (Honda) Packer, Bot. Not. 113:
291. 1960. Basionym: Diplachne hackelii Hon¬
da. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 3:
I 12. 1930. Cleistogenes hackelii (Honda) Hon¬
da. Bot. Mag. (Tokyo) 50: 437. 1936. TYPE:
Japan. Musashi Prov., Shirako. 1880, Matsu-
mura 4 (holotype, Tl).

Kengia chinensis (Maximowicz) Packer, Bot. Not. 1 13:
291. I960. Basionym: Diplachne serotina var. chi¬
nensis Maximowicz, Bull. Soc. Imp. Naturalistes
Moscou 54: 70. 1B79. Cleistogenes chinensis (Maxi¬
mowicz) Y. I., keng, Sinensia 5: 152. 1934. Cleis¬
togenes serotina var. chinensis (Maximowicz) Handel-
Mazzetti, Mazz. Symb. Sinicae 7(5): 12B0. 1936.
Cleistogenes hackelii var. chinensis (Maximowicz)
Oliwi. Bot. Mag. (Tokyo) 55: 309. 1941. TYPE,: Ja¬
pan. Yokohama, “fine Septembris fl.." C. Maximow¬
icz (lectotypc, designated here, LE).

When Maximowicz described his variety chinen¬
sis of Diplachne serotina (now Kengia serotina (L.)
Packer), he based the variety on five syntypes.
These comprised three Bretschneider collections
from near Peking (Takio-sze, Hei-lung-tau, Wan-
shou-, China), and two of his own collections, one
from Mongolia and the other from Yokohama, Ja¬
pan. The Japanese collection is a specimen of Ken -
gia hackelii (Honda) Packer, the only species of
Kengia present in Japan.

There has been confusion over the identity of the
remaining syntypes, mainly comprising Bretschnei-
der's plants from China. Confusion first resulted
from mistakes in the paper by Keng (1934) erecting
the genus Cleistogenes for a group of Eurasian
grasses that had previously been included in Di¬
plachne. Keng made the combination Cleistogenes
chinensis (Maximowicz) Keng based on Maximow-
icz's variety, but his description, illustration, and

No\on 15: 464—470. Pi bushed on 21 September 2005.

Volume 15, Number 3
2005

Phillips & Chen

Notes on Grasses (Poaceae)

465

cited specimens all belong to a different species
now known as Kengia songorica (Roshevitz) Packer.
Keng later (1938: 299) realized his mistake and
described the specimens he had used in his 1934
paper as C. mutica (a synonym of K. songorica).
However, his combination Cleistogenes chinensis re¬
mains validly published.

In 1934 keng had cited several specimens from
China under Cleistogenes serotina. Two years later
Handel-Mazzetti (1936) transferred Maximow iez's
variety to C. serotina as C. serotina var. chinensis.
These are the basis for reports of the distribution
of Kengia serotina extending to China (e.g.. Kit fan,
1985: 579). Tvpical K. serotina occurs from south¬
ern and eastern Europe only as far east as western
Turkmenistan and is not known from eastern Russia
or China (Tsvelev, 1983: 949). Ohwi (1941: 309)
recognized that the eastern Asian populations differ
from European Kengia serotina and assigned both
the Chinese and Japanese elements of Diplachne
serotina var. chinensis to Cleistogenes hackelii , sep¬
arating the Chinese plants as C. hackelii var. chi¬
nensis.

In recent Chinese literature Kengia chinensis has
been recognized as a separate species distinct from
Kengia hackelii (C. P. Wang, 1990: 47). Conert
( 1959: 232) put forward the suggestion that the Chi¬
nese element of Diplachne serotina var. chinensis
(1879), based on Rretschneider’s collections, may
be eonspecifie with Kengia kitagawae (Honda)
Packer (1936: 99).

Clearly a lectotypification is necessary to fix the
use of the name Diplachne serotina var. chinensis
and hence Kengia chinensis. A search was under¬
taken in the St. Petersburg herbarium (EE) to locate
the original syntypes. Three Bretschneider speci¬
mens were found from the type localities near Pe¬
king, but labeled with the manuscript name ''Di¬
plachne rigidula n sp." in Maximowicz’s hand. One
is Kengia kitagawae , and the other a mixed collec¬
tion. No specimen from Mongolia was located.
However, there is a specimen collected by Maxi-
mowicz in Yokohama, and clearly labeled by him
as Diplachne serotina var. chinensis. This is select¬
ed here as the lectotype. Hence C. chinensis be¬
comes a synonym of K. hackelii. and the current
use of the later name K. kitagawae for the Chinese
taxon is preserved.

Kengia inucronata (Y. E. keng ex P. C. keng &
E. Eiou) Packer, Hot. Not. 113: 293. I960.
Cleistogenes mucronata Y. E. keng ex P. C.
Keng & L. Eiou, Acta Hot. Sin. 9: 70. I960.
TYPE: China. Gansu: without locality or date,
T. D. Wang s.n. (holotype, NAS).

kengia gracilis (Y. I,. Keng ex P. C. keng & I,. Eiou)
Packer, But. Not. 113: 293. I960. Cleistogenes gra¬
cilis Y. L. Keng ex 1’. C. Keng & E. Eiou, Acta Bot.
Sin. 9: 69. 1960. Svn. nov. TYPE: China. Shaanxi:
Wu-kung, Chang-kia-kang, 1940. K. S. Tien s.n. (ho¬
lotype, NAS).

Additional specimens. CHINA. Shanxi: Sihsien. 7
Sep. 1935, I V. Wang .1568 (K). Shaanxi: Si Kuan
Is'. . .(illegible), 16 Oct. 1916, E. Licent 8021) (k).

Kengia mucronata and k. gracilis were described
at tin' same time in 1960. Y. C. Keng distinguished
them only by the number ol florets in the spikelet
(4 to 6 in K. mucronata: 5 to 8 in K. gracilis ), and
by the number of veins in the upper glume (often
3 in K. mucronata: I in K. gracilis). Two other spec¬
imens at Kew have 3 to 5 florets and 1 -veined
glumes.

The glumes in all Kengia species are variable.
The cleistogamous spikelets found in the upper leaf
sheaths generally have fewer florets and smaller hy¬
aline glumes than the chasmogamous spikelets of
the terminal panicle. The lower spikelets of the ter¬
minal panicle tend to have smaller, fewer-veined
glumes than those above. When the glumes are pre¬
dominantly 1 -veined, small lateral veins can usu¬
ally be found in some spikelets, sometimes only on
one side of the glume. Hence glume nervation is a
very unreliable character for distinguishing species
in Kengia. When the spikelets have a larger num¬
ber of florets, the spikelets overlap, giving the pan¬
icle branches a denser appearance.

Kengia mucronata can be characterized as fol¬
lows: habit densely tufted, wiry, with basal clumps
of old sheaths, lacking basal scaly buds; leaf blades
numerous, stiff, patent, inrolled, uppermost blades
very short; panicle exserted from uppermost leaf
sheath, open; spikelets with inconspicuously mu-
cronate lemmas.

Kengia mucronata is selected here to have pri¬
ority over K. gracilis because the type specimen is
better. The type of K. gracilis lacks a base.

IsACHNEAE

Isachne clarkei Hooker f.. FI. Brit. India r. 24.
1897 [1896]. TYPE: NE India, kohima, 1520
m, “5000 ft” [1520 m], 20 Oct. 1885. C. II
Clarke s.n. (lectotype, designated here. K).

Isachne heneckei llackel. Oesterr. Bot. Z. 51: 459. 1901.
Syn. nov. TYPE: Indonesia. Java, Prigen, 23 Mar.
1891. F. F. Benecke 22 (lectotype, designated here,
W).

Isachne tenuis Y. E. keng ex P. C. Keng, Acta Phytotax.
Sin. 10: 15. 1965. TYPE: China. Yunnan: Ping-hien
Hsien, Shi-tuen. 29 Sep. 1939, C. IE Wang 82102
(type, PE [at NAS, 2004]).

466

Novon

I liis is a delicate annual species with narrowly
lanceolate leaf blades, and an open panicle of small
(1—1.5 mm) scattered spikelets on slender glandu¬
lar pedicels. The specimen selected as the lecto-
type for Isachne clarkei is one of two sheets at Kew
collected by Clarke at Kohima in the Naga Hills,
and bears the name “clarkei” on the label in Hook¬
ers handwriting. The syntype specimens from Sik¬
kim included by Hooker in the protologue are /.
sikkimensis Bor, a species with longer (2—2.4 mm)
elliptic spikelets and an eglandular panicle.

When Hackel described /. beneckei he compared
it with the protologue ol /. clarkei and apparently
had difficulty finding real differences, mentioning
little more than the thin texture of the leaf blades
ol /. clarkei. In fact, the type specimens of the two
taxa agree very well. Hackel based his species on
two syntype specimens from Java, Henecke 22 and
20. The better specimen, Henecke 22, is selected
here as the lectotype.

Isachne tennis was first established by V. I,. Keng
(1957: 117. 218) and described by him two years
later (1959: 642, t. 580), but both these publica¬
tions lack a Latin diagnosis. In the 1965 protologue
/. tenuis is said to dilfer from /. beneckei by its
smaller spikelets, fewer-nerved glumes, and glan¬
dular pedicels. However, the type of /. beneckei
does also have glandular pedicels, and the other
characters (all within the normal range for /. clar¬
kei.

Isachne sylvestris Ridley, J. Straits Branch Roy.
Asia!. Soc. 44: 206. 1905. Isachne albens var.
sylvestris (Ridley) Jansen. Reinwardlia 2: 280.
1953. TYPE: Malaysia. Perak, Telok Sera, 17
Mar. 1896, II. N. Ridley 7205 (holotype, k).

Isachne albens var. hirsute Hooker f., FI. Brit. India 7: 23.
1897 1 1896]. Syn. nov. Isachne hirsute (Hooker f.) I*.
C. keng. Acta Phytotax. Sin. 10: II. 1965. TYPK:
NE India. Cachar, R. L. Keenan (lectotype, desig¬
nated here, K).

I his is one ol the more vigorous species of Is¬
achne, with culms up to 65 cm tall and a much-
branched panicle about 20 cm long. It is similar to
the widespread southeast Asian species /. albens
Irinius and was first described from northeastern
India as a variety ol this. However, it is clearly
specifically distinct, as set out in the following key
couplet.

KkY TO DlSTINGl ISI! ISACH.XK \I.HE\S AND /. SYLVESTRIS

la. Leaf sheaths glabrous; leaf blades 0.8- 1.8 cm

wide; culm nodes and panicle eglandular; spike¬
lets 1-1.5 mm, whitish green . /. albens

lb. Leaf sheaths densely hirsute; leaf blades 1.3— 2.4

<•111 wide; culm nodes and panicle branches glan¬
dular; spikelets 1.3— 1.9 mm, green or purplish
green . /. sylvestris

A separate specific status for Isachne albens var.
hirsuta was recognized by P. C. keng. who raised
it to specific rank as /. hirsuta. However, this grass
had already been described at specific rank as /.
sylvestris by Ridley, based on a specimen from the
Malay peninsula. The types ol both are at kew, and
are clearly conspecific.

Isachne albens var. hirsuta was described by J.
I). Hooker based on two syntype specimens from
northeast India (Silhet. <le Silva in Wall. Cat. R057
and Cachar, Keenan s.n.). The more complete col¬
lection from Cachar collected by Keenan is select¬
ed here as the lectotype. The other cited collection,
from Silhet by l)e Silva, consists only ol the top of
a culm with one leal and a panicle.

I’iii-; Circumscription ok Isachne ki \thi\i\a
(Wight <!i Arnoit kx Stkudkl) Miqukl.

/. SCHMIDTII HACKKL, AND /. RERUNS RKNC

I his group of three species has been much con¬
fused in the literature. They are low -growing spe¬
cies with lanceolate leaf blades, eglandular pani¬
cles, spikelets over 2 mm long, and similar florets.
The differ* ■nees between them are set out here in
order to clear up the confusion.

I he oldest name is Isachne kunthiana, originally
described from India by Steudcl as Panicum kun-
thianum. I his grass is apparently confined to India
and Sri Lanka and has spikelets with the glumes
clearly longer than the florets. The confusion arose
when Miquel transferred the name from Panicum
to Isachne, while misapplying it to a similar species
in southeast Asia with long glumes. Sinee then /.
kunthiana has been widely described as occurring
in southeast Asia (e.g., Gilliland, 1971; Davidse,
1994; Lazarides, 1980; Iskandar & Veldkamp,
2004). The correct name for this southeast Asian
species is /. schmidtii.

In 1899 Haekel (1899: 721) assigned a grass
from the Ryukyu islands (Japan) to a new variety
nudiglumis of Isachne m vasal is. Later Koyama
(1962) correctly recognized its affinity to the south¬
east Asian species known as /. kunthiana and
transferred it as /. kunthiana var. nudiglumis,
hence compounding the misapplication of the spe¬
cies name /. kunthiana. Meanwhile Y. L. Keng
(1933: 129) had described this grass at tin* species
level as /. repens, based on specimens from south¬
ern mainland China. Koyama knew this, and cited
/. repens as a synonym of /. kunthiana var. nudi¬
glumis.

Volume 15, Number 3
2005

Phillips & Chen

Notes on Grasses (Poaceae)

467

Brief descriptions follow, setting out the most im¬
portant differences, with relevant synonymy. A key
to distinguish these three taxa is given below.

Kky to Distinguish Isachne ki sthiana, I. sunn mil. and

I. REPENS

la. Plant with underground rhizomes; leaf blade base

cordate; glumes w ith a few long setae or glabrous,
longer than florets . /. kunthiana

lb. Plant with surface stolons; leaf blade base obtuse
to rounded; glumes densely scabrid-hispidulous,
longer than or equaling florets.

2a. Glumes longer than florets; leaf blades 2.6—

4.5 cm; panicle contracted, 0.5-1 cm wide,

branches erect . /. schmidtii

2b. Glumes equaling florets; leaf blades 5—7.5
cm; panicle open, 1 .4—2.8 cm w ide, branch¬
es ascending or spreading . /. repens

Isacline kunthiana (Wight & Arnold ex Steudel)
M ique I. FI. Ned. Ind. .4: 460. 1857. Panicum
kunthianum Wight & Arnold ex Steudel, Svn.
PI. Glumac. I: 96. 1854. TYPE: India. "Pen-
ins. Ind. or.,” Herb. Wight 1659 (isotype, k).

Plant w ith underground rhizomes. Leaf blades I —
I I cm, hispid, base cordate, basal margins pecti¬
nate. Panicle contracted to open, branches op¬
pressed to widely spreading, seabrid, pilose at least
in the axils. Glumes chartaceous, longer than flo¬
rets. contracted above florets, acuminate-cuspidate,
with a few long tubercle-based setae or glabrous;
florets elliptic, the upper slightly shorter, more con¬
vex and often pubescent.

Miquel transferred the epithet kunthiana to Is-
achne in 1857. on the basis of a specimen from
Java that had been tentatively placed under Pani¬
cum kunthianum by Steudel. .1. I). Hooker (1896:
21) cited a Ridley collection from Singapore that is
a syntype of/, semitalis (syn. of /. schmidtii). Jansen
(1953: 285) placed I. schmidtii in synonymy under
/. kunthiana. but he does not appear to have ex¬
amined specimens from India.

In fact, although /. kunthiana and /. schmidtii
are similar in facies and in possessing long glumes,
they are clearly distinguishable by the characters
given here in the key and descriptions.

Isachne schmidtii Haekel, Hot. Tidsskr. 24: 97.
1901. IA PE: Thailand. Koh Chang island. ,/.
Schmidt s.n. (holotype,W).

Isachne semitalis Ridley, H. Malay. I Vain. 5: 237. 1 625.
TYPE: Malaysia. Selangor, Rantau Panjang, C. It.
Kloss 78 (lectotype, designated here, K).

Plant with surface stolons. Leaf blades 2.6— 4.5
cm. glabrous or with thinly scattered hairs, base

obtuse to rounded, margins not pectinate. Panicle
contracted, 0.5—1 cm wide, branches erect, almost
smooth, slightly scaberulous, glabrous or with oc¬
casional scattered setae. Glumes herbaceous, lon¬
ger than florets, evenly narrowing above florets,
densely scabrid-hispidulous; florets ovate to sub¬
rotund. monomorphic, slightly glossy, pubescent
only on inrolled margins.

Ridley cited five syntype specimens in the pro-
tologue of Isachne semitalis. All are in the kew her¬
barium and are a good match with the type of /.
schmidtii. The best example is selected here as lec¬
totype.

Glume length is variable in this species. I he
glumes typically exceed the florets by 0.5—1 mm.
However, some spikelets may have glumes scarcely
longer than the florets in panicles where most
spikelets have clearly longer glumes. Possibly /.
repens may better placed as a subspecies under /.
schmidtii, as done by koyama (1962, 1987 as /.
kunthiana subsp. riudiglumis), but we do not wish
to do this here as we have seen insufficient speci¬
mens to be certain.

Isachne repens Y. L. keng. Sunyatsenia 1: 129.
t. 33. 1933. TYPE: China. Guangdong: Ping
Wu Shan. 18 Oct. 1929, C. 6. Tso 21292 (ho-
lotype, IBSC).

Isachne myosotis var. riudiglumis Haekel, Bull. Herb.
Boiss. 7: 721. 1899. Isachne firmula Base var. riu¬
diglumis (Haekel) Rendle, J. Linn. Sue. Bot. 36:
322. 1904. Isachne kunthiana var. riudiglumis
(Haekel) Koyama, .1. Jap. Bot. 37: 236. 1962. Isach¬
ne kunthiana subsp. nudiglumis (Haekel) Koyama,
Grass. Jap. Neighb. Reg.: 511. 1987. TY BE: Japan.
Ryukyu Is., Liu-kiu I., Mt. Yonahadake, Toshiro ex
coiled. Matsumura s.n. (holotype, W).

Plant with surface stolons. Leaf blades 5—7.5 cm.
glabrous or pilose, base obtuse to rounded, margins
not pectinate-hispidulous. Panicle open, 1.4— 2.8
cm wide, branches ascending or spreading, smooth,
glabrous. Glumes herbaceous, equal to florets or
upper very slightly shorter, densely seabrid: florets
ovate to subrotund, monomorphic, slightly glossy,
pubescent only on inrolled margins.

Isachne repens is a slightly larger grass than /.
schmidtii, with bigger leaf blades and a less con¬
tracted panicle. However, the spikelets are similar
except for the shorter glumes.

file taxon is incorrectly named as /. kunthiana
in the recently published Flora of Taiwan (Hsu.
2000: 449). where /. schmidtii is cited in synonymy.
The text therein refers to the glumes exceeding the
florets, but the accompanying illustration clearly

468

Novon

shows thorn subequal and is a good representation
of I. repens.

For distribution within China, see Chen (1990:
178).

Am ndinki 1 1: \i

Ai iiiidinella siiniana S. \1. Phillips & S. L. Chen,
sp. nov. TV PE: China. Yunnan: "leg. bor.,
Yunnanfu.” 19 Sep. 1922. ,/. Cavalerie s.n.
(holotype, BM).

I laec species Arundinellae setosae affinis, sed nodis
barhatis, infloreseentia contraeta, ramis spiculis congestis,
spieulis longioribus (6.8— 7.8 mm non 5—6.5 mm), lemmate
inferiore spieula aequilongo, lemmate superiore hilohato,
lobis triangularibus setosis differt.

Base absent: culm ea. 85 cm tall. 8 mm diam.,
3-noded, nodes lanate. Leaf sheaths smooth, upper
margin and mouth villous, otherwise glabrous; lig-
ule ea. 0.5 mm. membranous, eiliolate; leaf blades
narrowly linear, ea. 22 X 2.5— 3.5 mm. glabrous,
scabrous, apex finely acuminate. Panicle lanceo¬
late. contracted, ea. 10 X 2.5 cm. branches densely
spiculate, lowest branch ca. 3.5 cm; main axis and
branches scabrous puberulous on edges; pedicels
very unequal, the longer 2.5-3 mm, the shorter
0.5—1 mm. not setose at apex. Spikelets 6.8— 7.8
mm, dark gray; glumes lanceolate, conspicuously
setose on veins with stilf tubercle-based trichomes.
apex acuminate-caudate; lower glume ea. 6 mm. 3-
veined, upper glume ea. 7.3 mm. 5-veined; lower
floret staminate or sterile, ca. 6.5 mm. equaling up¬
per glume, lemma smooth, glabrous, 3-veined; up¬
per floret bisexual, elliptic-oblong, ea. 3.8 mm,
apex clearly 2-dentate, shortly pilose on upper
flanks and apical lobes, awned from sinus, lobes
triangular, each tipped by 0.7— 0.9 mm awnlet; awn
geniculate, column ca. 2 mm, limb 4—4.3 mm; an¬
thers 3, 2.5— 2.7 mm. Caryopsis not seen.

I bis species is named after B. S. Sun, the emi¬
nent agrostologist from Yunnan province, and is at
present known only from the type. It is clearly re¬
lated to the widespread and variable Arundinella
setosa Frinius by the possession of lateral bristles
on the upper lemma, but differs by the combination
of characters set out in the diagnosis above. The
contracted panicle of dark -colored, setose spikelets
is particularly noteworthy and reminiscent of that
found in Arundinella hookeri. However, that species
has broader (up to 1.2 cm), densely villous leaf
blades, and an emarginate upper lemma lacking
lateral bristles.

In Arundinella setosa the lower floret is shorter
than the glumes, or at least clearly shorter than tin1
upper glume, and the upper lemma is scabrous, the

lateral bristles arising more or less directly from
the awn base. In contrast, in Arundinella suniana
the lower floret is as long as the upper glume, and
the lateral bristles arise from definite membranous
apical lobes.

Arundinella setosa Trinius var. eselosa Bor ex S.
M. Phillips & S. I,. Chen, var. nov. TYPE: In¬
dia. “Hah. Himal. Bor. Occ., regio trop.." 31
Aug. 1849, T. Thomson s.n. (holotype, K).

Bor (1960: 425) provided a Latin description
when establishing this variety and cited 1 1 i ret* spec¬
imens (T. Thomson s.n.: Stainton, Sykes & Williams
4255. 4461). However, lie neglected to indicate a
type, a necessary requirement of valid publication
after 1958. Flic Thomson specimen is selected here
as type because it clearly shows the setose pedicels
of A. setosa, and also lacks any trace of apical lat¬
eral bristles on the upper lemma, the defining char¬
acter for this variety.

Note on Ari sdineija tit peiensis Kkng & X. P.
Lion, Et.. Huhkiknsis 4: 312. 2001.

The species epithet “ hupeiensis " is confusingly
similar to the epithet huheiensis already in use
in Arundinella for the previously published A. hu¬
heiensis I). M. Chen (1983). Both are named after
the Chinese province of Hubei. According to Arti¬
cle 53.3 of the International Code of Botanical No¬
menclature (Greuter et al.. 2000) such names are
to be treated as homonyms, and are therefore ille¬
gitimate. Arundinella hupeiensis is based on the col¬
lection T. Y. Tai & C. //. Chien 465. stated to be
in the herbarium of Nanjing University. However,
it could not be found there on enquiry by the sec¬
ond author. It seems likely that it is a specimen of
Arundinella setosa var. esetosa,

Andkopogonk \h

Spodiopogon humhiisoidcs (P. C. keng) S. M.
Phillips X S. L. Chen, comb. nov. Basionym:
Eccoilopus hambusoides P. C. Keng, in B. J.
Ceng X C. (t). Song, Cuihaia 13: 320. 1993.
TV PE: China. Guangxi: Guilin. Qixingyan, 27
Oct. 1953, Guangxi Expedition 3820 (holo¬
type. PE | PB in error. 1993], not seen).

This species was originally published as Spodi¬
opogon hambusoides by V. L. Keng (1957: 139,
235) with a brief Chinese description as part of a
key, but without a Latin diagnosis. Two years later
lie provided a full description (1959: 768), but
again only in Chinese. Fhe name was therefore not
validly published. His son P. C. Keng later provid-

Volume 15, Number 3
2005

Phillips & Chen

Notes on Grasses (Poaceae)

469

ed t he necessary Latin description and type infor¬
mation, hut placed it in the genus Eccoilopus , as
noted above.

The genus Eccoilopus Steudel comprises a small
group of species in Asia distinguished from Spo-
diopogon s. str. by a tough raceme rachis (articu¬
lations are present but do not actually fracture),
with both spikelets of a pair pedicellate, rather than
one sessile and the other pedicelled. The genus
Eccoilopus will be included within Spodiopogon in
the upcoming treatment for Poaceae in Flora of
China.

Imperata flavida Y. L. Keng ex S. M. Phillips A
S. L. Chen, sp. nov. TYPE: China. Hainan: Ton
Fao Kacheh river, sandy river shore. 91 m
|“300 ft"). 14 Jan. 1923, Eryl Smith s.n. (ho-
lotype, K).

I lace species Imperatae cylindricae affinis sed pilis ral-
li spirula aequilonga non 3-plo longiore, stigmatihus flav-
ido-brunneis non atropurpureis differt.

Khi/omatous perennial: rhizomes spreading,
vvoodv, internodes closely packed, 3—5 mm; old leaf
sheaths fibrous; culms solitary, erect, 70—125 cm
tall, 3—7 mm diam.; nodes glabrous. Leaf sheaths
crowded near base, longer than internodes, smooth,
glabrous except for silky triehomes at mouth, up¬
permost sheath spathelike with vestigial blade and
enclosing base of panicle; ligule ca. 1 mm. brown,
margin densely eiliolate; leaf blades linear, flat. 20—
60 X 0.5— 1.0 cm (upper culm blades much short¬
er), smooth, glabrous, base narrowed to prominent
midrib, apex acuminate. Panicle cylindrical, 12—17
X 2.5 cm; branches short, erect; pedicels of a pair
unequal, long-pilose, expanded upward. Sp i ke let s
narrowly elliptic oblong, 3—4 mm. membranous;
callus triehomes equal to spikelet or slightly short¬
er; glumes pilose on back in lower third, triehomes
equaling spikelet. lower glume 5-veined, upper
glume 7-veined, upper margins eiliate, apex obtuse
or erose; lower lemma broadly oblong, ca. 2 mm,
eiliate, irregularly denticulate; upper lemma and
palea ca. 1.5 mm, eiliate, irregularly lobed; anthers
2. 2.5— 2.8 mm; stigmas yellowish brown. Caryopsis
not seen.

Distribution. China, Hainan island, in mixed
forest along rivers and in valley bottoms (L. Liu,
1997: 35).

This species was originally published by Y. L.
Keng (1957: 138, 234) with a brief Chinese de¬
scription as part of a key, but without a Latin di¬
agnosis. Two years later he provided a full descrip¬
tion with an illustration (1959: 756, fig. 701) based
on the specimen McClure 7791 , but again only in

Chinese. The name was therefore not validly pub¬
lished. L. Liu (1997: 35) included the specie's un¬
der Imperata , with the reference “Keng ex L. Liu,
LI. Res. Gram. II: 31. 1989.” However, this work
has in fact never been published. The name is val¬
idated here, and the opportunity is taken to provide
a full description in English. It has not been pos¬
sible to trace McClure 7791 , so the species is typ¬
ified on another collection at Kew.

As far as is known, Imperata flavida is confined
to the island of Hainan. It can be readily distin¬
guished from the much commoner /. cylindrica (L.)
Raeuschel by its shorter spikelet triehomes. The
species difference in stigma color is also notewor¬
thy.

Faratypes. CHINA. Hainan: Ton Fao Kacheh river,
sandy river shore, 300 ft., 14 Jan. 1923, Eryl Smith s.n.

( k ): sine loo., 18 Nov. 1932. N. K. Clam 442H2 (NAS);
sine loc., sine dat., F. A. McClure 7791 (whereabouts un¬
known).

Saerharum arundinaceum Retzius var. trielio-
phy limn (Handel-Mazzetti) S. M. Phillips &
S. L. Chen, comb. nov. Basionym: Erianthus
griffithii J. I). Hooker var. trichophyllus Han¬
del-Mazzetti, Akad. Wiss. Wien, Math.-Natur-
wiss. Kl.. Anz. 58: 154. 1921. Erianthus tri-
choph yllus (I landel-Mazzetti) I landel-Ma/.zetli.
Akad. Wiss. W ien. Math.-Naturwiss. Kl.. Anz.
62: 254. 1926. TYPE: China. Yunnan: 19 Mar.
1914, H. Handel-Mazzetti 745 (leetotype. des¬
ignated by Handel-Mazzetti (1936: 1308),
WU).

Distribution. China (Yunnan), Sikkim (Sikkim
Himalaya, 21 May 1874, collector illegible 128 ( K)).

Erianthus is included within Saccharum in the
upcoming treatment for Poaceae in the Flora of
China. Handel-Mazzetti’s variety is best placed un¬
der Saccharum arundinaceum. The species Sac¬
charum griffithii is a different taxon with much lon¬
ger, yellowish callus hairs. It occurs in Pakistan
and Afghanistan and is not known from China.

Handel-Mazzetti cited two of his own collections
in the 1921 protologne, numbers 745 and I 1016.
However, later in another publication (1936: 1308)
he cited 745 as “typus,” which therefore can be
accepted as the leetotype. Handel-Mazzetti 11016
(WU) is a specimen ol Saccharum procerum Rox¬
burgh. Variety trichophyllum appears to be a rather
small, locally occurring form of S. arundinaceum
distinguished mainly by the pilose upper glume of
the sessile spikelet. It can be distinguished from
the normal form as follows:

470

Novon

Kkv to Distinguish Two Varieties ok
Saccharvm AM I MIX ACM 1/

la. Culms up to () rn; upper glume of sessile spikelet

glabrous .

. S. (irundinaceum var. arundinaceum

lb. Culms up to 1.5 m; upper glume of sessile spikelet

thinly pilose . . ,S. arundinaceum var. trichophyllum

1'olytrias milieu (Houttuyn) Veltlkamp var. nana
(keng S. E. Chen) S. M. Phillips & S. L.
Chen, comb. nov. Basionym: Eulalia nana
Kcng & S. L. Chen, FI. Main. 4: 454. 539. f.
1243. 1977. Polytria s amaura var. nana (kcng
& S. L. Chen) S. L. Chen, FI. Reipubl. Po-
pularis Sin. 10(2): 101. 1997. TYPE: China.
Hainan: Dong Fang, along bank of Chang Hua
river. 7 Jan. 1956, Hainan Exped. 531 (leo
totype, designated here. I BSC).

This variety, apparently confined to Hainan Is¬
land, differs from the typical variety in Polytrias
indica by the shorter pubescence on the lower
glume of the sessile spikelet, as shown in the il¬
lustration accompanying the protologuc of Eulalia
nana in Flora Hainanica (1997). It was later re¬
duced to varietal level under the species name Po¬
lytrias amaura , the widely used name for this grass
until recently. However, the correct name for the
species is P. indica, P. amaura being a superfluous
name, as explained by Veldkamp (1991: 180).

I here is a type specimen in the Guangzhou her¬
barium (I BSC), but because it is uncertain whether
this is the holotype, it is selected here as lectotype.

Mncsithcn striata (Steudel) Koning Sosef var.
pukescens (Hackel) S. M. Phillips & S. L.
Chen, comb. nov. Basionym: Rottboellia striata
var. pubescens Hackel, in A. & C. de Candolle,
Monogr. Phan. 6: 302. 1889. Coelorachis stri¬
ata var. puhescens (Hackel) Bor, (basses Bur¬
ma. Ceyl. India Pak.: 121. I960. TYPE: India,
khasia. J. D. Hooker & T. Thomson s.n. (type,

k).

Veldkamp ft al. (1986) reduced Coelorachis to a
synonym ol Mnesithea. Hairy forms of M. striata are
separated at the varietal level in the upcoming
treatment for Poaceae in the Flora of China , re¬
quiring this new combination.

Acknowledgment. We thank the director of the
herbarium at the Vienna Museum (W) for sending
the type ol Isachne benekei on loan to kew.

Literature Cited

Bor, N. L. I960. The Grasses of Burma, Ceylon, India,

and Pakistan. Pergamon Press, Oxford, London, New

York, Paris.

Chen, I). M. 1983. A new species of Arundinella from
Hubei province. Acta Phytotax. Sin. 21: 207-208.

Chen, S. L. 1990. Isachne. In Flora Reipublicae Popularis
Sinicae. 10(1): 176—191. Science Press, Beijing.

Conert. 11. ,1. 1959. Beitrage zur Monographic der Gattun-
gen Cleistogenes und Neyraudin. Bot. Jahrb. 78: 208—
245.

Davidse, G. 1994. Isacline. In; Dassanayake, F. IT Fosberg
& W. 1). Clayton (editors). Flora of Ceylon, Vol. 8. Am¬
erind, New Delhi.

Gilliland, 11. B. 1971. A revised Flora ol Malaya, Vol. 3.
Grasses of Malaya. Government Printer, Singapore.

Greuter, \\.. J. McNeill, F. B. Barrie, IF M. Burdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
J. F. Skog, P. Trehane, N. ,1. Turland & I). L. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.

Hackel, E. 1899. Enumeratio Graminum Japoniae. Bull.
Herb. Boiss. 7: 637—726.

Handel-Mazzetti, II. 1936. Symbolae Sinicae, Vol. 7.
Springer, Vienna.

Hooker, J. I). 1896. Flora of British India, Vol. 7. 1897.
Reeve, Ashford.

Hsu, (7 (7 2000. Flora of Taiwan, Ed. 2, Vol. 5. Grami-
neae. Department of Botany, National Taiwan Univer¬
sity. Taipai.

Iskandar, E. A. P. & J. 1. Veldkamp. 2001. A revision ol
Malesian Isachne sect. Isachne (Gramineae, Panicoi-
deae, Isachneae). Reinwardtia 12: 159-179.

Jansen, P. 1953. Notes on Malaysian grasses I. Reinward¬
tia 2: 225—350.

Keng, V. L. 1933. Two new' grasses from Kwangtung. Sun-
yatsenia I: 128-130.

- . 1934. A newr generic name, Cleistogenes, in the

grasses of Eurasia. Sinensia 5: 147—157.

- . 1938. New grasses from Peiling Miao, Suiyuan

Province, China. J. Wash. Acad. Sei. 28: 298-308.

- . 1957. Claves Generum et Specierurn Graminear-

um Primarum Siniearum. Scientific Books Publishers,
Shanghai.

- . 1959. Flora lllustralis Plantarum Primarum Sin¬
iearum. Science Publishing House, Peking.

kit Tan. 1985. Cleistogenes. In Flora of Turkey, Vol. 9.
Univ. Press, Edinburgh.

koyama, T. 1962. Nomenelatural remarks on some grass¬
es. J. Jap. Bot. 37: 231-241.

- . 1987. Grasses ol Japan and its Neighboring Re¬
gions. Kodansha, Tokyo.

Lazarides. M. 1980. The Tropical Grasses of Southeast
Asia. Cramer, Vaduz.

Liu. I.. 1997. Imperata. In Flora Reipublicae Popularis
Sinicae 10(2): 30—35. Science Press. Beijing.

Ohwi, J. 1941. An account of the tribe Eragrosteae of
Japan II. Bot. Mag. (Tokyo) 55: 309-313.

Lsvelev, N. N. 1983. Grasses ol the Soviet Union. Oxonian
Press, New Delhi, Calcutta.

Veldkamp, J. F. 1991. Miscellaneous notes on southeast
Asian Gramineae. VI. Blumea 36: 179-181.

- . R. l)e Koning & M. S. M. Sosef. 1986. Generic

delimitation of Rottboellia and related genera (Grami¬
neae). Blumea 31: 281—307.

Wang, (7 P. 1990. Cleistogenes. In Flora Reipublicae Po¬
pularis Sinicae 10(1): 41-54. Science Press, Beijing.

Notes on Grasses (Poaceae) for the Flora of China, V. New species

in Cymbopogon

Sylvia XL Phillips

Herbarium, Royal Botanic Gardens, Kew, Surrey TWO 3AB, United Kingdom.

s.ph i 1 1 i ps@kew.org

Peng Una

Herbarium, Institute of Botany, Academia Sinica, Heilongtan, Kunming, Yunnan 650204,

China, hpeng@mail.kib.ac.cn

Abstract. Two new species of Cymbopogon are
described from China, C. liangshanensis and C. mi¬
nor. Both belong to Cymbopogon series Proceri, pre¬
viously mainly confined to Australia. A third spe¬
cies in this series, Cymbopogon quinhonensis from
Vietnam, is transferred from Andropogon.

Key words: China, Cymbopogon, Vietnam.

This paper describes two new species in Cym¬
bopogon Sprengel from China. Both were recog¬
nized as new by Liu Liang, who died in 2001, but
neither has been validly published. They represent
the first records from continental Asia of Cymbo¬
pogon series Proceri Soenarko, the most primitive
series in the genus, which until now was thought
to be confined to Australia and parts of Indonesia
and New Guinea. The apparent disjunct distribu¬
tion is bridged by one species in Vietnam, here
transferred from Andropogon L. to Cymbopogon.

Cymbopogon liangshanensis L. Liu ex S. M.
Phillips & H. Peng, sp. nov. TYPE: China.
Sichuan: Liangshan, 21 Aug. 1964, /J\u Tai-
ping 41 1 (holotype, PE).

Haec species omnibus sinensis racemis non deflexis
cum spiculis homogamis vestigialibus vel nullis, spiculis
pedicellatis redactis manifeste distinguenda; C. procero
culmis 50-110 cm (non 100-200 cm) allis, internodiis
rachidis pedicellisque ciliatis pilis 0.5— 1.5 mm (non 6
mm) differt.

Perennial; culms solitary or in small tufts, slen¬
der. 50-1 10 cm tall, ca. 1.5 mm diam., 4- to 6-
noded. Leaf blades linear, gray green, 20—30 cm X
3—5 mm, slightly scaberulous, lower blades nar¬
rowed almost to midrib toward base; ligule scarious,
1.5— 2.5 mm. Inflorescence a spathate compound
panicle, 30—40 cm long; branches laxly ascending,
bearing several tiers of spathes. Spatheoles green¬
ish brown, narrowly lanceolate, 2—2.5 cm. Racemes
paired, 1.5— 1.8 cm, 3- to 5-noded; peduncle in¬

cluded within spatheole, glabrous. Raceme bases
subequal, linear, not deflexitig. apices cupuliform,
scarious; homogamous spikelets often absent, or
one reduced or vestigial pair present on lower ra¬
ceme base; rachis internodes and pedicels sub¬
equal. slenderly linear, ca. 3/4 length ol sessile
spikelel, ciliate with ca. 0.5 mm long white tri-
chomes increasing to ca. 1.5 mm at apex, apices ol
rachis internodes and pedicels expanded, cupuli¬
form, scarious, toothed. Sessile spikelet 3.7-4 mm:
callus ca. 0.5 mm, shortly and sparsely pilose; low¬
er glume narrowly elliptic oblong, papyraceous, lat¬
erally keeled with indexed margins, glabrous, back
shallowly concave below middle, keels scabrid,
unwinged, 2- to 4-veined between keels above mid¬
dle, narrowed to 2-denticulate apex; upper glume
acute; lower floret reduced to an oblong hyaline
lemma; upper lemma 2-lobed. awned from sinus;
awn geniculate, glabrous, 1.3— 1.5 cm; pedicelled
spikelet narrow, reduced to the glumes, lower
glume 2. 5-3. 5 mm, upper glume 1-2 mm. Cary-
opsis not seen.

Distribution. China: southwest Sichuan.

Liu Liang (1997: 197, fig. 48, 1—4) included this
species in her account of the genus Cymbopogon
for volume 10(2) of the Chinese-language Flora of
China, with a reference to her publication “Plant
Resources of Gramineae II: 14. 1989." In fact, this
work has never been published, and her description
in 1997 was without a validating Latin description.
The species is validated here, and the opportunity
is taken to provide a full description in English.

Cymbopogon liangshanensis differs from all other
members of the genus in China in possessing a
rather lax spathate panicle, and racemes with slen¬
der. non-deflexing raceme bases. The homogamous
spikelet pair normally present on the lower raceme
base in Cymbopogon is reduced or vestigial, or even
completely absent, and the pedicelled spikelets are

Novon 15: 471-473. Published on 21 September 2005.

472

No von

2 mm

Figure I. Cymbopogon minor IF S. Sun & R. Zhang ex S. VI. Phillips iK H. Peng. — A. Hahit. — R. Spatheole and
raceme bases. — C. Spikelet pair. — I). Pedicel with pedicelled spikelet and raehis internode. Drawn from the holotype.
Team oj Northeastern Yunnan 1466 (YUNU), by Margaret Tebbs.

similar, being narrow and reduced to the glumes.
In these characters the taxon approaches the close¬
ly related genus Arulropogon. However, the sub¬
equal raceme bases, and the presence of a homog-

amous jiair (albeit often very small), which is
normally completely absent in Arulropogon, are in¬
dicative of Cymbopogon.

Cymbopogon liongshonensis appears to be a first

Volume 15, Number 3
2005

Phillips & Peng

New species in Cymbopogon

473

record lor continental Asia of Cymbopogon series
Proceri, whose members are otherwise mainly con¬
fined to Australia. This is the most primitive series
in the genus, including species with equal, fre¬
quently non-deflexing raceme bases and reduced
pedicelled spikelets. It is distinguished from the
Australian C. procerus (R. Brown) Domin by its
smaller habit and much shorter trichomes in the
racemes.

Cymbopogon minor B. S. Sun & R. Zhang ex S.
M. Phillips & II. Peng. sp. nov. TV PK: China.
Yunnan: Qiaojia County, Xiaohe, bank of Niu-
lan Jiang [river], roadside in grassland. 900 m,
2B Oct. 1964, Team of Northeastern ) unnun
1466 (holotype, YUNU). Figure 1.

Haee species C. liangshanensi af finis, sed lam inis fo-
liorum angustioribus (1—2 mm non 3—5 mm lalis), spa-
iheolihus angustioribus brevioribus (1.2-1. 5 cm non 2-
2.5 cm longis), praccipuc basibus racemorum dcflcxis
di fieri.

Perennial, tufted: culms wiry, 60-70 cm tall, 1
mm diam.. ea. 4-noded. Leaf blades narrowly lin¬
ear. pale green, ea. 20 cm X 1—2 mm. smooth,
glabrous, narrowed toward base; ligule soarious.
1.3—2 mm. Inflorescence a narrow spathate panicle,
moderately dense, up to 30 cm long, spathes and
spatheoles in clusters; spatheoles red, narrowly lin¬
ear. 1.2— 1.5 cm; racemes paired. 4- or more-noded
(no complete racemes on type); peduncle slender,
glabrous, ea. 1/2 length of spatheole, shortly ex-
serted at maturity; raceme bases subequal, short,
flattened, deflexed at maturity, pubescent, bearded
at apex with ca. 2.5 mm long hairs, apices cupu-
liform, scarious. Homogamous spikelets reduced,
one pair present on lower raceme base, lower spike-
let of pair resembling pedicelled spikelets, upper
spikelet vestigial; rachis internodes and pedicels
subequal, slenderly linear, ea. % length of sessile
spikelet, densely eiliate with ca. 0.7 mm long white
trichomes increasing to 2—3 mm at apex, apex of
internode expanded, eupuliform, scarious, toothed,
apex of pedicel similar but smaller; sessile spikelet
3.8—4 mm; callus ca. 0.3 mm. shortly bearded, a
Irichome tuft projecting into base of lower glume
concavity; lower glume narrowly oblong, herba¬
ceous, margins indexed, lightly keeled below mid¬
dle. sharply keeled above, glabrous, back shallowly
concave below middle, keels seabrid, unwinged, 2-
to 4-veined between keels above middle, narrowed
to emarginate apex; upper glume acute; lower floret
reduced to an oblong hyaline lemma; upper lemma
2-lobed, avvned from sinus; awn geniculate, gla¬
brous, 1.2—1 3 cm; pedicelled spikelet narrow, 0.5—

2.5 mm. reduced to empty lower glume. Caryopsis
not seen.

Distribution. China: northeast Yunnan.

This species is known only from the type spec¬
imen. which bears two determination labels, ”//y-
parrhenia gracilis L. Liu. sp. nov.” and
“ Cymbopogon minor B. S. Sun et R. Zhang, sp.
nov.” Neither of these names has been publis lied.
The taxon is clearly excluded from Hyparrhenia by
the short callus of the sessile spikelet, which is
inserted into the expanded apex of the rachis in¬
ternode below. It was correctly placed in Cymbo¬
pogon bv B. S. Sun and R. Zhang, whose specific
epithet is taken up here.

Cymbopogon minor resembles C. hangshanensis,
as described above, in its reduced homogamous
and pedicelled spikelets. I he small sessile spike¬
lets are also similar, but differ slightly in shape,
texture, and a more strongly bearded callus with a
trichome tuft. However, it differs markedly by its
strongly deflexing. flattened, bearded raceme bases.
This makes it rather anomalous in Cymbopogon se¬
ries Proceri, into which it would otherwise fit com¬
fortably.

A species described in Andropogon from Viet¬
nam by A. Camus also has slender, equal, non-
deflexing raceme bases with absent or reduced ho¬
mogamous pairs, and reduced pedicelled spikelets.
It also is best placed in Cymbopogon series Proceri,
and is transferred to the genus here.

Cymbopogon quinhonensis (A. Camus) 5. M.
Phillips & H. Peng, comb. nov. Basionym: An¬
dropogon quinhonensis A. Camus, Bull. VI us.
Hist. Nat. (Paris) 25: 201. 1 919. TYPE: Viet¬
nam. Annam: Qui-nhon, Balansa 4H72 (holo¬
type, P not seen, K photo).

A. Camus (1919) remarked on the similarity of
this grass to Cymbopogon in the protologue. It has
more conspicuously white-villous racemes than C.
Hangshanensis, and the lower glume ol the sessile
spikelet lacks intercarinal veins.

Additional specimen seen. VIETNAM. Annam: lour-
ane [Da Nang], border of thickets, May-July 1927, ./. &
At. S. Clemens 4454 (K).

Acknowledgment. We thank the directors of the
herbaria at PE and YUNU for sending types on loan
to Kew.

Literature Cited

Camus, A. 1919. Craminees nouvelles de l'Asie orientale.

Bull. Mus. Hist. Nat. (Paris) 2.5: 262-204.

I, iu. L. 1997. Cymbopogon. I’p. IBB— 209 in flora Rei-

publicae Popularis Sinicae, Vol. 10(2). Science Press,

Beijing.

Notes on Grasses (Poaceae) for the Flora of China, VI.
New Combinations in Stipeae and Anthoxanthum

Sylvia M. Phillips

Herb arium, Royal Botanic Gardens, Kew, Surrey TW9 3AB, United Kingdom.

s.phillips@kew.org

Wu /lien- 1 jui

Herbarium, Northwest Plateau Institute of Biology, Academia Sinica, 78 Xiguan Street,

Xining, Oinghai 810001, China

Absik \< a . W lien Oryzopsis is confined to the type
species, Chinese species are placed in Achnathe-
rum and Piptatherum. This necessitates the follow¬
ing new combinations: Piptatherum grandispicu-
lum, Piptatherum aequiglume var. ligulatum,
Piptatherum tibeticum var. psilolepis, Piptatherum
munroi var. parviflorum, Achnatherum henryi, and
Achnatherum henryi var. aciUum. Achnatherum
henryi is lectotypified. The new name Piptatherum
filial replaces the illegitimate name Piptatherum ob¬
tusion, and this species is lectotypified. Hierochloe
potaninii is transferred to Anthoxanthum as A. po-
taninii, as Hierochloe is regarded as a synonym of
Anthoxanthum. All the grass taxa (Poaceae) listed
are endemic to China.

Key words: Achnatherum, Anthoxanthum, Chi¬
na. Iherocldoe. Oryzopsis, Piptatherum, Poaceae.

When preparing the tribe Stipeae, Poaceae, for
the Flora of China, it was necessary to consider
generic concepts in the light ol modern research,
as discussed below. I bis has necessitated a few
new combinations. The opportunity is also taken in
this paper to publish a new combination in An¬
thoxanthum in the tribe Aveneae.

Stick m

Delimitation of genera in Stipeae is still in a state
ol flux, but recent work has led to a realignment of
some familiar genera, based in part on molecular
studies. Opinion has been divided in the Old World
on whether to take a broad view of Stipa itself,
including Achnatherum and some smaller genera,
or whether to recognize Achnatherum separately.
Freitag (1985) took a broad view, as did Cope
(1982) and Noltie (2000). However, Russian and
Chinese authors have generally kept the two genera
separate, for example Tzvelev (1968, 1976) and Lu
and Kuo (1987). liarkworth (1993) has published a

generic reassessment for North America and, with
collaborators, followed this with a report on molec¬
ular studies (Jacobs et al., 2000). The two genera
are now considered separate in America (Soreng,
2003) and will be also in the treatment ol Poaceae
for tht‘ Flora of China.

A further problem involves the genus Oryzopsis
Miehaux. This name lias been widely used in both
hemispheres, with Piptatherum regarded as a syn¬
onym. a view reinforced by Prei tag's paper (1975)
revising the species from southwest Asia. However,
the type species of Oryzopsis, O. asperifolia Mi-
chaux (1803) from North America, has many un¬
usual features and is not closely related to other
species placed in Oryzopsis (Barkworth, 1993). Ory¬
zopsis is therefore considered as monotypic in mod¬
ern treatments (Jacobs et ah. 2000; Soreng, 2003),
with the other species placed in Piptatherum and
Achnatherum.

Most Chinese species placed in Oryzopsis in the
Chinese flora (Lu & Kuo, 1987) already have com¬
binations in either Piptatherum or Achnatherum.
However, a few new combinations are required, as
set out below. All the taxa listed are endemic to
China, or almost so ( Piptatherum kuoi extends to
the Ryukyu islands in Japan and Taiwan).

Piptatherum kuoi S. M. Phillips & /. L. Wu,

nom. nov. Replaced name: Piptatherum obtu¬
sion (Stapf) Roshevitz, in Bot. Mater. Gerb.
Bot. Inst. Komarova Acad. Nauk SSSH 14:
102. 1951, non Piptatherum obtusum Nees &
Meyen. 1841. Oryzopsis obtusa Staph in Hook¬
er's Icon. PI. 24(4): t. 2393. 1895. TYPE: Chi¬
na. Hubei: Ichang and immediate neighbor¬
hood, A. Henry 3507 (lectotype, designated
here, K).

This species differs markedly from other Pip¬
tatherum species in China by its plump, ellipsoid.

Novon 15: 174-476. Published on 21 Shite mb eh 2005.

Volume 15, Number 3
2005

Phillips & Wu

New Combinations in Stipeae

475

glabrous floret and large callus with circular scar.
It also has glabrous anthers. The other eight Chi¬
nese species (e.g., P. munroi (Stapf) Mez) have lan¬
ceolate. usually pubescent florets and a very shal¬
low' callus with a transversely elliptic scar. The
anthers are normally bearded.

The species Piptatherum aequiglume (Dutliie ex
Hooker f.) Roshevitz, P. tibeticum Roshevitz. and P.
munroi (Stapf) Mez belong to a group ol closely
related species in high parts ol western China and
the western Himalaya. The following taxa described
within this group require transfer from Oryzopsis.

Piptatherum graiidispiciilum (P. C. Kuo X X. L

Mil) S. M. Phillips X Z. L. Wu, comb. nov.
Basionym: Oryzopsis grandispicula I*. C. Kuo
X Z. I.. M'ii, Acta Phytotax. Sin. 10: 435. t. 1.
1981. TYPE: China. Xizang: Xigaze, Daiduka,
3700 m. 25 Aug. 1963. Zhi-ye Gin 10210 (ho-
lotype, HNWP).

This is a variant with large 8—9 mm spikelets,
close to P. munroi (Stapf) Mez. but distinguished
from that species and others ol the group by its
smooth leaves and panicle branches.

Piptatherum aequiglume (Duthie ex Hooker 1.)
Roshevitz var. ligulnlum (P. C. Kuo & /. E.
Mu) S. M. Phillips & Z. I,. Wu, comb. nov.
Basionym: Oryzopsis aequiglumis Duthie ex
Hooker f. var. ligulata P. C. Kuo & Z. I,. Wu,
Acta Phytotax. Sin. 19: 435. 1981. TYPE: Chi¬
na. Yunnan: Eijiang— Heqing, 2800 m. 10 Sep.
1962, Exped. Zhongdian 2270 (holotype,
KUN).

This taxon appears to be a local element from
the Piptatherum aequiglumis— P. munroi species
group. The 8—13 mm ligule is remarkably long, and
the 0.4 mm lemma pubescence is longer than in
either aforementioned species. It may prove to mer¬
it separate specific status when better known.

Piptatheriun tibeticum Roshevitz var. psilolcpis

(P. C. Kuo & Z. L. Mu) S. M. Phillips & Z. I,.
M'ii. comb. nov. Basionym: Oryzopsis tibetica
(Roshevitz) P. C. Kuo var. psilolepis P. C. Kuo
& Z. E. M'u, Acta Phytotax. Sin. 19: 435. 1981.
Oryzopsis psilolepis (P. C. Kuo X Z. E. Wu) L.
Eiu, Vase. PI. Hengduan Mountains 2: 2257.
1994. TYPE: China. Sichuan: Aiming, 2400
m. 4 July 1959. Xin Li 70020 (holotype, PE).

This taxon differs from the typical variety by its
smooth glabrous lemmas.

Piptatherum niuiiroi (Stapf) Mez var. parviflo-
runi (Z. E. Wu) S. M. Phillips X Z. E. Wu,
comb. nov. Basionym: Oryzopsis munroi Stapf
var. parviflora Z. E. Wu, Acta Phytotax. Sin.
30(2): 174. 1992. TYPE: China. Qinghai:
Menyuan, Zhugu, Xuelongtan, 2700 m, 7 July
1962, Geobotanical Group 1651 (holotype,
HNM P).

This taxon differs only slightly from P. tibeticum
Roshevitz, but that species is reported to have an
ovate floret and 3 to 5 branches at the panicle
nodes, whereas P. munroi var. parviflorum has a
lanceolate floret and usually paired panicle branch¬
es.

Two species within Oryzopsis in the Chinese flora
(Eu X Kuo, 1987) (it better into Achnatherum than
Piptatherum, namely 0. chinensis and 0. henryi.
These had previously been recognized as diiferent
from other Chinese species in Oryzopsis (Wu &
Zhou, 1981), when a new section Sinoryzopsis was
erected to accommodate them. They are best placed
in Achnatherum on account of their more or less
terete florets with a pubescent callus. The first has
already been transferred as Achnatherum chinense
(Hitchcock) Tzvelev. The second, with its variety,
is transferred below.

Achnatherum henryi (Rendle) S. M. Phillips X
Z. E. M'u, comb. nov. Basionym: Stipa henryi
Rendle, J. Einn. Soc., Bot. 36: 382. 1904. Ory¬
zopsis henryi (Rendle) Keng ex P. C. Kuo, El.
Tsinlingensis 1: 145. 1976. TAPE: China.
Sichuan: S. Wushan, A. Henry 5530 (lectotype,
designated here, K).

The specimen selected as lectotype from the
three syntypes listed in the protologue already
bears a lectotype label “selected by II. Freitag.”
However, this lectotypifieation has never been pub¬
lished. The selected specimen is the best of the
three original syntypes, the other two being poor
specimens consisting of only one culm.

Achnatherum henryi var. acutuin (E. Eiu ex Z.
L. Mu) S. M. Phillips X Z. E. Mu. comb. nov.
Basionym: Oryzopsis henryi var. acuta E. Eiu
ex Z. E. Wu, Acta Phytotax. Sin. 19(4): 435.
1981. Oryzopsis acuta (E. Eiu ex Z. E. Mu) E.
Liu, Vase. PE Hengduan Mountains 2: 2257.
1994. TYPE: China. Yunnan: Meixi, 2500 in,
8 Mar. 1940. K. M. Feng 3729 (holotype, PE).

Anthoxanthlim

The genus Hierochloe B. Brown was included
within Anthoxanthum by Schouten and Veldkamp

476

Novon

(1985) because of the morphological intermediates
between them. This decision was followed by Noltie
(2000) and has been borne out by work on the
group for the Flora of China, where several inter¬
mediates also occur. By unifying these two genera,
their close relationship is emphasized, as evi¬
denced by their unusual 3-flowered spikelet struc¬
ture, with the two lower florets sterile or staminate
and modified, and all three falling together from the
glumes as one unit. The possession ol coumarin,
found in this group, does not occur elsewhere in
the grasses.

Schoulen and Veldkamp made new combinations
in Anthoxanthum for those species of Hierochloe
where combinations did not already exist, but omit¬
ted one species from China.

Anthoxaiithuin potuninii (Tzvelev) S. M. Phillips
<£ Z. L. Wu, comb. nov. Basionym: Hierochloe
potaninii Tzvelev, Bast. Centr. Azii, Mater.
Bot. Inst. Komarov 4: 35. 1968. TYPE: China.
W Gansu: Li-Dsha-Pu, 20 June 1885, G. Po¬
tanin s.n. (holotype, LE not seen).

fhis was reduced to synonymy under Hierochloe
laxa B. Brown ex Hooker f. (Anthoxanthum laxum
(B. Brown ex Hooker f.) Veldkamp) by Y. II. Sun
and P. C. Kuo (1987). However, the two taxa can
be distinguished as follows:

km to PisriNoi isii Axthoxaviw i tnm.xi.xti and A. iam i /

la. Leaf blades scabrous; glumes unequal; bisexual

floret seabrid near apex . 4. potaninii

I b. I.eaf blades smooth; glumes subequal; bisexual

floret pilose or hirsute toward apex . 4. laxum

Anthoxanthum laxum occurs on high mountain
ledges and in alpine pasture over 3000 m in eastern
Afghanistan, northern Pakistan. Kashmir, and Ne¬
pal (Cope, 1982), and A. potaninii is known only

lrom southern Gansu and western Sichuan provinc¬
es ol China.

Acknowledgments. Vie thank tin* curators at
KUN and HNWP for sending types on loan to kew.

I .iterature Cited

bark worth, VI. E. 1993. North American Stipeae (Grami-
neae): Taxonomic changes and other comments. Phyto-
logia 74: 1—25.

Cope, T. A. 1982. Flora ol Pakistan. No. 143. Poaceae.

Department of Botany, l Diversity of Karachi.

Freitag, II. 1975. The genus Piptatherum (Gramineae) in
Southwest Asia. Notes Boy. bot. Gard. Edinburgh 33:
341-408.

- . 1985. l’he genus Stipa (Gramineae) in Southwest

and South Asia. Notes Boy. bot. Gard. Edinburgh 42:
355-489.

Jacobs. S. W. I... .1. Everett. M. E. bark worth & C. Hsiao.
2000. Belationships within the stipoid grasses (Grami¬
neae). Pp. 75-82 in S. W. E. Jacobs & .1. Everett. Grass¬
es: Systematics and Evolution. CSIB0, Collingwood.
Lu. S. E. & Kuo. P. C. 1987. Achnathernm. Pp. 319—329
in P. C. Kuo, Flora Beipublicae Popularis Sinicae, Vol.
9(3).

Michaux, A. 1803. Flora boreali-Amerieana I. Paris,
Strasbourg.

Noltie. II. .1. 2000. Flora of Bhutan. Vol. 3(2): 883. The
Grasses ol Bhutan. Royal Botanic Garden Edinburgh X
Royal Government ol Bhutan.

Schouten, V. & J. F. Veldkamp. 1985. A revision of An¬
thoxanthum including Hierochloe (Gramineae) in Ma-
Icsia and Thailand, blumea 30: 319-351.

Soreng, B. J. 2003. P. 730 in Catalogue of New World
Grasses (Poaceae): IV. Pooideae. National Museum of
Natural History. Washington. I).C.

Sun Y. II. & P. C. Kuo. 1987. Hierochloe. Pp. 17:5-179
in P. C. Kuo. Flora Beipublicae Popularis Sinicae. Vol.
9(3).

Tzvelev, N. N. 1968. Basteniya Central'nov Asii. Vol. 4.
Poaceae. V. 1. Grubov (editor). Nauka Publishers, Len¬
ingrad.

- . 1976. Zlaki SSSB. Nauka Publishers, Leningrad.

Wu, Z. E. & Zhou, X. M. 1981. New supplement and eco-
geographic distribution of the genus Oryzopsis in China.
Acta Phytotax. Sin. 19: 434—4.39.

Kive New Combinations in Gossia (Myrtaceae) from Melanesia

Neil Snow

Department ol Biological Sciences, University of Northern Colorado, Greeley, Colorado 80634,
U.S.A. Current address: H. T. Harvey & Associates, 423 W. h allbrook Ave.. Suite 202,
Fresno, California 9371 1-6138, U.S.A. neil.snow@unco.edu

ABSTRACT. Based on characters of the fruit, seed,
and embryo, live new combinations in Gossia N.
Snow & Guymer (Myrtaceae) are proposed for spe¬
cies occurring in New Guinea, Vanuatu, or ihe Sol¬
omon Islands. These species previously all had
been placed in Myrtus L. or Austromyrtus Burret.
A leetotype is selected for Gossia aneityensis. A key
is provided for the Melanesian species of Gossia.

Key words: Austromyrtus, Gossia. Melanesia,
Myrtaceae, Myrtus, Solomon Islands, Vanuatu.

Recent revisionary work in Myrtaceae has led to
the splitting of Austromyrtus s.l. into Austromyrtus
s. sir., Lenwebbia N. Snow & Guymer. and Gossia
N. Snow & Guymer (Snow et ah, 2003). Gossia is
most species-rich in Australia and New Caledonia,
where together approximately 42 species occur
(Snow, 2000; Snow et ah. 2003). The author is pres¬
ently revising species of Gossia for the Flore de la
Nouvelle- Ga ledon ie and has had the opportunity to
reassess boundaries of all known species ol the ge¬
nus. The following species currently placed in Myr¬
tus or Austromyrtus are all easily attributable to
Gossia and arc here transferred to that genus based
on the amended descriptions of Austromyrtus. Myr¬
tus, and Gossia (Snow et ah. 2003). Gossia floribun-
da (A. J. Scott) N. Snow & Guymer, which is most
common in Australia, also occurs in New Guinea
(Snow et ah. 2003).

Kin to the Species of Gossia ekom New Gi inea, the
Solomon Islands, and Vanuatu

1. Flowers 5-merous . 2

I Flowers 4-merous . 4

2(1). Branchlets rounded . G. eugenioides

2' . Branchlets 4-angled . 1

3(2). Stipular “hairs” prominent, very elongated api-
eallv; leaves broadly elliptic, I2-2B mm long;
hypanthium densely appressed-pubeseent . . .

. G. versteegbii

3'. Stipular “hairs” somewhat obscure, only some¬
what elongated apically; leaves narrowly ellip¬
tic to ovate, 15—75 mm long; hypanthium gla¬
brous to sparsely sericeous . G. floribunda

4(1). Axis of inflorescence with short, spine-like

bracts . G. salomonensis

F. Axis of inflorescence lacking spine-like bracts

5(4). Branchlets 4-angled; leaves very narrowly

ovate, apex apieulate . G. randiana

5'. Branchlets round; leaves narrowly ovate to

ovate, apex acute . 6

6(5). Stipular “hairs” obscure (or if present then not
swollen at base); style hairy at least near base

. G. aneityensis

6'. Stipular “hairs” prominent and greatly swollen

at base; style glabrous . G. floribunda

1. Gossia randiana (Merrill & Perry) N. Snow.

comb. nov. Basionym: Myrtus randiana Merrill
& Perry, .1. Arnold Arbor. XXIII: 237. 1942.
TYPE: Papua New Guinea. Central Division.
Kubuna, Dec. 1933, /,. ./. Brass 5642 (holo-
type. A; isotype, BRI).

The holotype specimen has two illustrations in
pencil, presumably by Merrill or Perry, that illus¬
trate two characters which in part diagnose Gossia
from related genera. The first is the circulate em¬
bryo, the second is the gelatinous endosperm seen
associated with the embryo (Snow et ah. 2003).

2. Gossia eugenioides (A. .1. Scott) N. Snow.

comb. nov. Basionym: Myrtus eugenioides A. ,|.
Scott, Kew Bull. 33: 514. 1979. TYPE: Indo¬
nesia. Manokwari, Vogelkop, Arlak Mis., Angi
gila lake, 2800 m, A. Kostermans 2598 (holo¬
type, k [photos at BRI. GREE]; isotype. E
| photos at BRI, GREE]).

The original description by Scott (1979) indi¬
cates only two collections, both from Irian Java. In
the key to fertile specimens (Snow et ah. 2003: 32),
Gossia eugenioides would key out most closely to
Gossia shepherdii (E. Mueller) N. Snow & Guymer,
but it differs most readily from G. shepherdii by its
moderate to densely sericeous hypanthium and ev¬
ident lack of smooth bark. Further information is
needed about the biology and distribution of this
species.

3. Gossia versteegliii (Merrill N Perry) N. Snow.

comb. nov. Basionym: Myrtus versteeghii Mer¬
rill & Perry, J. Arnold Arbor. XXIII: 238.
1942. TYPE: Indonesia. Irian Java. 9 km NE
of hake llabbema, 2800 m camp. Oct. 1938,
L. ,/. Brass 10618 (holotype. A: isotypes. BRI.
k. h).

Novon 15: 477-478. Ft bushed on 21 September 2005.

478

Novon

The stipular “hairs"' of Gossia versteeghii are
unique in the genus. As with the “type B" stipules
discussed in Snow et al. (2003: 7), they have a
prominently swollen base, but they differ in having
a greatly attenuated apex and can be up to 2 mm
long.

4. Gossia salomonetisis (A. J. Scott) N. Snow.

comb. nov. Basionym: Myrtus salomonensis A.
J. Scott, Kew Bull. 35: 412. 1980. TYPE: Sol¬
omon Islands. Gizon Island, R. Mauriasi et al.
BSIR 11740 (holotype, k | seen as digital im¬
age!).

The species is distinct in Cassia by virtue of the
number of short spinose bracts occurring in the in¬
florescence, which are typically aggregated near the
base of the pedicel; other species in the* genus typ¬
ically have none or only a few such bracts.

5. Gossia aneityensis (Guillaumin) IN. Snow,

comb. nov. Basionym: Myrtus aneityensis Guil-
laumin, J. Arnold Arbor. XII: 254. 1931. Aus-
tromyrtus aneityensis (Guillaumin) Burret, No-
ti/.bl. Bot. Gait. Berlin-Dahlem 15: 506. 1941.
TYPE: Vanuatu ( = New Hebrides). Anelgau-
hat Bay, 23 Feb. 1929, S. J. Kajewski RIO (lec-
totype, elesignated here, P; photos at BBI,
GHEE; isotypes, BBI, k [photo], MEL).

Guillaumin (1931: 254) indicated that the com¬
mon name of this species is “Nivie.” The selected
lectotype is the* first of two specimens mentioned in
the protologue, which I have seen and which is
representative of the species.

Acknowledgments. Thanks to Eve Lucas at the
Royal Botanic Gardens, kew (k), for sending a dig¬
ital image of the type specimen of G. salomonensis ;
Christiane Tirel and Ceeile Aupic at the Museum
National d’Histnire Naturelle (P) for assistance in
the herbarium; and the curators of A, BR1, k, L,
MEL, and P for permission to borrow or view spec¬
imens. The comments of two anonymous reviewers
and Victoria llollowell clarified the manuscript.

Literature Cited

Guillaumin, A. 1931. Contributions to the flora of the New
Hebrides: Plants collected by S. F. Kajewski in 1928
and 1929. J. Arnold Arbor. 12: 220-264.

Scott, A. J. 1979. New species and combinations in Myr-
taceae from Malesia and Australia. Kew Bull. 33: 51 1 —
5 1 5.

Snow, N. 2000. A conspectus of Australasian Myrtinae
(Myrtaceae). kew Bull. 55: 647—654.

- , G. P. Guymer & G. Sawvel. 2003. Systematies of

Austromyrtus, !r nwebbia , and the Australian species of
Gossia (Myrtaceae). Syst. Bot. Monogr. 65: 1—95.

Two New Species of Rhynchospora sect. Tenues (Cyperaceae)
from the Guianas, South America

Mark 7. Strong

Department of Botany, MRC/166, Smithsonian Institution. P.0. Box 37012, Washington, D.C.

20013-7012, U.S.A. strongm@si.edu

ABSTRACT. Two new species are described (R.
cordatachenia and R. saxisavannicola ) in Rhyn¬
chospora sect. Tenues kiikenthal from northeastern
South America (Guianas). A key to the species of
R. sect. Tenues in the Guianas is given, and illus¬
trations are provided for the new species.

Key words: Cyperaceae, French Guiana, Guian¬
as, Rhynchospora sect. Tenues, South America, Su¬
rinam.

Rhynchospora Vahl nom. cons, is a genus of ap¬
proximately 270 species (Strong, 2004). It is most
diverse in the Western Hemisphere, particularly
warm-temperate North America and the Neotropics.
Eighty taxa (including infraspecific taxa and the
two novelties described here) are currently known
from the Guianas (Guyana, Surinam, and French
Guiana) (Strong, 2001, 2004).

Of the 18 sections of Rhynchospora occurring in
the Guianas (Strong, 2004), R. sect. Tenues kiiken¬
thal is the best represented with 14 species and I
infraspecific taxon. This high number of taxa re¬
flects the center of diversity and speciation of
Rhynchospora sect. Tenues in savanna ecosystems
of northern South America, where there are likely
many more species still awaiting discovery. Rhyn¬
chospora sect. Tenues is primarily Neotropical in
distribution, but has representatives in tropical Af¬
rica, Madagascar, and one species in southeast
Asia, R. gracillima Thwaites. In the Neotropics,
species range from the West Indies and Mexico,
south through Central America and northern South
America to southern Brazil and northern Argentina.
Thomas (1998) treated 16 taxa (including several
unknowns) for the Venezuelan highland region of
South America. Eleven taxa were treated (including
2 newly described) by Rocha and Luceno (2002) in
a treatment of R. sect. Tenues for Brazil.

In habit, species of Rhynchospora sect. Tenues
can be characterized as low annuals or perennials,
less than 0.5 m tall, primarily of sandy savanna and
grassland habitats. They typically have linear or
subterete, often involute leaf blades; an inflores¬
cence composed of a terminal anti 1 or 2 corym¬

bose. open or contracted, partial panicles from the
upper bracts; a 2-branched style; and an aehene
that is transversely rugose or rugulose with verti¬
cally oriented linear cells that have finely sinuous
walls, a 2-lobed style base, and bristles wanting.
Achenes of some species have cells with strongly
convex or swollen periclinal walls at their base
characteristic of Rhynchospora sect. Spermodontes
kiikenthal, which can be characterized by the two
lateral margins of the aehene that are generally pro¬
longed and cusp-like at their apices. Two species
that Kiikenthal (1949, 1950) placed in section
Spermodontes (R. fallax Uittien and R. tenerrima
Nees ex Sprengel) are particularly characteristic of
species in section Tenues. They have the linear, ver¬
tically oriented epidermal cells and transversely ru¬
gulose surface typical of that section. However, the
two species exhibit the cusp-like margins of the
aehene apex characteristic of section Spermodontes
and the style base is low and often obtusely trian¬
gular, sometimes decurrent on the upturned cusp¬
like margins. Style bases of Rhynchospora sect.
Tenues are generally 2-lobed at base with lobes that
extend along the shoulders of the aehene apex.
However, in some species the lobes are indistinct,
e.g., R. spruceana C. B. Clarke and R. albida (Nees)
Bockeler, or unlobed altogether as in R. caracasana
(kunth) Bockeler, R. hirsuta (Vahl) Vahl. and R.
tenella (Nees) Bockeler.

The glossy and transversely rugulose aehene sur¬
face seen in Rhynchospora sect. Tenues and other
Rhynchospora species may help protect the embryo
and surrounding tissues from lethal temperatures
brought by full sunlight in open habitats such as
tropical savannas and grasslands. Like lighter sur¬
faces of objects that reflect sunlight and are inter¬
nally cooler, glossy epicuticular waxes on the aehene
surface increase reflection of incoming radiation and
aid in regulating internal temperature (Eller, 1979).
The surface sculpturing of a seed increases its sur¬
face area up to 10 times. The surface irregularity
may increase energy exchange with the surrounding
cooler air and may increase thermodynamic ex-

Novon 15: 479-483. Pubeishkd on 21 September 2005.

480

Novon

change l>y causing turbulence in laminar air flow
around the seed surface (Rarthlott, 1981).

Kky to the Species or RtnxcHosnoiu sect. Tkm ks in

THE GlIIANAS

I. Aeliene body appearing essentially smooth,

at most faintly transversely rugulose . 2

I'. Achene body distinctly transversely rugu¬
lose, rugose, or pitted-reticulate . 3

2(1). Achene body thickly biconvex, tumid, broadly
obovate to subrounded, abruptly narrowed to
short-stipitate base; style base triangular,
blunt at apex; sheaths becoming fibrillose, the
base of plant clothed with rusty brown fibers;
plant of upland savanna or shrub savanna

habitats . R. caracasana (Kunth) Bockeler

2'. Achene body thinly biconvex, not tumid,
narrowly obovate, rounded-cuneate at base,
estipitate; style base triangular with acute to
short-acuminate apex; sheaths not fibrillose;
plant of high-elevation tepui habitats ....

. R. roraimae Kiikenthal

3(1). Achene body with two oval to rounded pro¬
tuberances on either side of the base, ap¬
pearing as though there is a style base on

both ends . R. riparia (Nees) Bockeler

3'. Achene body lacking protuberances at base 4
4(3). heal blades and sheaths uniformly pubescent

. R. Iiirsiiln (Vahl) Vahl

4 . Leaf blades and sheaths glabrous, at most leaf

blades ciliate on margins and midveins ... 5

5(4). Spikelets 1.5-2 mm long: style base forming
2 strap-like lobes proximally, which are de-
current on shoulders of achene bodv to about
the middle, abruptly contracted distally into a
short triangular-lanceolate tip .... R. contractu

(Nees) Ravnal

5'. Spikelets 2.5—1 1 mm long; style base not
strap-like proximallv, trigonous, depressed-
trigonous, cap-like, or subdiscoid, not
abruptly contracted distally except for R.

saxisavannicola . 6

6(5). Spikelets — I 1 mm long . 7

O'. Spikelets 2.5— 5.5 mm long . B

7(6). Achene asymmetrically biconvex, thicker
near apex, obovate to narrowly obovate, wiri¬
est at apex, estipitate at base; style base de¬
pressed-triangular, entire or scarcely 2-lobed

. R. spruceana C. B. Clarke

7' . Achene biconvex, turgid medially, transverse¬

ly oblong or quadrate, widest at middle, with
short-stipitate base; style base triangular, 2-

lobed or shallowly 2-lobed . R. emaciata

(Nees) Bockeler

8(6). Spikelet scales dark brown on sides; aehenes
broadly biconvex to nearly globular; style
base depressed-trigonous or narrowly dis¬
coid. not lobed basally, elliptic as seen from

above . R. tenella (Nees) Bockeler

8'. Spi kelet scales light brown to stramineous or
whitened; aehenes biconvex, obovate, obpyr-
iform. subrounded, oblate, or widely trans¬
versely oblong (squarish); style base often 2-

lobed basally . 9

9(8). Achene cordate or rounded-cordate at base;

sl\ le base narrow l\ triangular-lanceolate w ith

a flaring, 2-lobed base (shaped like a witch's

hat) . R. cordatachenia M. T. Strong

9'. Aehenes narrowed or subrounded at base; style

base triangular or depressed-triangular .... 10

10(9). Aehenes subrounded, oblate, widely trans¬
versely oblong, or quadrate, narrowed
abruptly to a slipitate base, often with a lon¬

gitudinal dark gray band or patch medially

on each side . II

10'. Aehenes obovate, obpyriform, or subrouml-
ed. gradually narrowed to base, brown, pale

brown, or flecked with gray . 12

11(10). Aehenes rounded to oblate; style base tri¬

angular, shallowly and shortly 2-lobed, 0.1 —

0.2 mm wide at base . . R. junciformis (Kunth)

Bockeler

I I '. Aehenes transversely oblong or quadrate; style
base depressed-trigonous, broadly 2-lobed.
0.6-0. 9 mm wide at base . . R. emaciata (Nees)

Bockeler

12(10). Achene narrowly obovate, coarsely rugose

with 5 to 6 rugae per lace .

. R. sanariapensis Steyermark

12'. Achene obovate or elliptic, rugulose with 7

to 12 rugae per face . 13

13(12). Roots often golden yellow; scales strami¬
neous to whitened; achene obpyriform, eu-
neate at base, short-stipitate; stvle base cap¬
like, indistinctly 2-lobed, the lobes not
extending along shoulders of achene body

. R. ulbida (Nees) Bockeler

13'. Roots brown; scales light brown; achene ob¬
ovate. broadly obovate, broadly elliptic, or sub-
rounded, estipitate; style base distinctly 2-
lobed, the lobes extending along shoulders of

achene body . 14

14(13). Spikelet rach ilia strongly flexuose at matu¬
rity; aehenes 0.5— 0.7 mm w ide; base of style
base 0.2— 0.3 mm wide; only known from
French Guiana and Surinam in rock savanna
and seeps on granitic outcrops (inselbergs)

. R. saxisavannicola M. T. Strong

14'. Spikelet rachilla not strongly flexuose; aehenes
0.8-1 mm w ide; base of style base 0., 5-0.7 mm

wide; wide-ranging savanna species . 15

15(14). Corymb rays elongate: spikelets straight or
nearly so, 1—5 mm long; anthers 2—3 mm long;
aehenes obovate, 1—1.4 mm long .... R. tenuis
Willdenow ex Link subsp. tenuis
15'. Corymb rays short; spikelets falcate, 2—1 min
long; anthers 0.8-1 mm long; aehenes sub-
rounded-obovate, 0.6-0.9 mm long . . . R. tenuis
subsp. austrobrasil iensis T. kovnmn

In preparing Rhynchospora for the Flora of the
Guianas, 1 noted two novelties, which are here de¬
scribed and validated.

Rhynchospora cordatachenia M. T. Strong, sp.
nov. TYPE: French Guiana. Environs de Cay¬
enne, Savane du Gal I ion. 9 Sep. 1979, A.
Raynal-Roques 21566A (holotype, US-
3464701) [originally mixed with Rhynchospora
tenerrima Nlees ex Sprengel; distributed as A.
Raynal-Roques 21566]. Figure I.

Volume 15, Number 3
2005

Strong

Rhynchospora sect. Tenues

481

figure 1. Rhynchospora cordatachenia M. T. Strong. — A. Habit. — B. Detail of junction of sliealli and leal blade. —
C. Cross section of leaf blade. — 1). Detail of inflorescence. — E. Spikelet. — F. Spikelet scale. — G. Achene. Drawn
from the holotype, A. Raynal- Roques 21566A (US).

A ceteris speciebus seetionis Tenuium, aehenio ad bas-
im cordato, rostro simili petaso venificae ad basim dilatato
di fieri.

Caespitose perennial. 7—13 cm tall; rhizome
short; roots fine, light brown; culms spreading-as¬
cending, 0.3— 0.6 mm wide, filiform, obtusely tri¬
gonous to terete or nearly so. soft. Hexuous, finely
ribbed, pale green to stramineous, glabrous. Leaves
I to 3 per culm, ascending, basal and lower cau-
line, 0.1—10 cm long; sheaths short, closely clasp¬
ing culm, ligulate with a narrow band of tissue present
at adaxial junction of sheath and blade, herbaceous,
finely ribbed, light brown to stramineous distally,
glabrous, the inner band membranous, truncate at
apex, brown, tinged with red; blades filiform, 0.2-

0.4 mm wide, involute to crescentiform-capillary,
herbaceous, cellular-reticulate adaxially, finely
veined abaxially, green, glabrous, margins smooth
proximally, antrorsely scabrous toward apex, atten¬
uate to subflattened tip. Inflorescence a terminal and
1 or 2 lateral corymbose partial panicles from the
upper bracts, the terminal panicle 4—9 X 2-5 mm,
with 3 to 6 spikelets; bracts leaf-like, 1—7 cm long,
the axillary one elongate and often exceeding sub¬
tending panicle; branches short, very slender and
filiform, flattened-trigonous or subterete in cross sec¬
tion; spikelets slenderly ellipsoid-lanceoloid, 3—1 X
0.6—1 mm, straight, acuminate to apex, narrowly cu-
neate at base; raehilla strongly flexuose at maturity;
scales dorsally obtuse to rounded, herbaceous, distal

482

Novon

scales of spikelet thinly herbaceous to submembran-
ous, brown, proximal and medial scales thickly her¬
baceous, smooth and glossy, uniformly pale brown,
glabrous, margins scarious, midcosta very fine, in¬
distinct proximally, prolonged beyond the obtuse to
acute apex as a short antrorsely scabrous awn, which
is slightly recurved, lateral nerves indistinct; lertile
scales 5 to 0, ovate-elliptic to ovate-lanceolate or
lanceolate, 2-2.7 X 0.9— 1.2 ram; sterile scales 2 at
base of spikelet, ovate-elliptic, 1.2— 1.5 X 0.5— 0.8
ram. Flowers bisexual; stamens 1, the anthers 0.6—
0.8 mm long, apieulate at apex, minutely papillose
at base, basifixed. thecae parallel, longitudinally de¬
hiscent; style 2-branched, equaling to 2/3 length of
unbranched portion, long-exserted from subtending
scale. Achene unequally biconvex, ovate-rounded or
somewhat ovate-quadrate. 0.8— 0.9 X 0.7-0.8 mm.
obtuse to subtruncate at apex, cordate to slightly so
at base, narrowly stipitate, transversely rugulose with
6 to 7 rugae per face, narrowly cellular-reticulate
along margins and at base, shiny with a crystalline
appearance, whitish tinged with brown; epidermal
cells linear, vertically oriented; style base narrowly
triangular-lanceolate with a flaring, 2-lobed base,
flattened, 0.4 X 0.3-0.4 mm, attenuate to apex,
brown to dark brown; bristles absent.

Distribution and habitat. Fndemic to French
Guiana in coastal plain savanna. Known only from
the type collection (see citation above).

Distinguishing features. Among the species of
Rhynchospora sect. Tenues , R ■ cordatachenia is
unique in having an achene with a cordate base
and a style base that is narrowly triangular-lance¬
olate with a flaring. 2-lobed base (shaped like a
witch’s hat).

l{liyiiclios|>oru saxisavannicola M. T. Strong, sp.
nov. TYPE: F reneh Guiana. Mont Saint-Mar¬
cel. zone sud-est du massif, mares gravillon-
naires de savane-roche, 300 m, 02Q23'00"N,
53Q00'20"W. 18 July 2002. J. J. DeGranville,
L. Aliker A: C. Sarthou 15283 (holotype, US-
3451946; isotypes, CAY, NY, P). Figure 2.

A ceteris speciebus sectionis Tenuium, achenio late el-
liptieo, rostra non profunde triangulari ad apicem abrupte
brevi-attenuato diflert.

Caespitose perennial, 3—10 cm long; rhizome
short; roots fine, light brown; culms erect, 0.2— 0.5
mm wide, filiform, flattened-trigonous or subterete,
soft, flexuous, finely ribbed, pale green, glabrous.
Leaves 1 to 3, ascending, primarily basal, 2—5 cm
long; sheaths short, closely clasping culm, ligulate
with a narrow band of tissue present at adaxial

junction of sheath and blade, herbaceous, finely
ribbed, pale brown to stramineous proximally, gla¬
brous, the inner band membranous distally on basal
sheaths, herbaceous on upper cauline sheaths ex¬
cept for membranous orifice, truncate at apex;
blades compressed-filiform, 0.2— 0.3 mm wide, her¬
baceous, finely veined, glabrous, the apex often
curving. Inflorescence 1 to 3 corymbose, lax. partial
panicles from the upper bracts, the terminal pani¬
cle 5—10 X 3—15 mm, with 1 to 5 spikelets; bracts
leaf-like, to 3 cm long; branches very slender and
filiform, flattened-trigonous or subterete in cross
section; spikelets slenderly ovoid-lanceoloid, 3—4.5
X 0.5— 0.9 mm. straight, acuminate to attenuate at
apex, cuneate at base; rachilla strongly flexuose at
maturity; scales dorsally obtuse to rounded, her¬
baceous, distal scales of spikelet thinly herbaceous
to submembranous, minutely cellular-striate, senii-
glossy. reddish brown with slightly darker brown
lineations, glabrous, margins narrowly scarious,
midcosta very fine, indistinct except at apex, pro¬
longed beyond the narrowly acute to acuminate
apex as a short antrorsely scabrous awn that is
slightly recurved, lateral nerves indistinct; fertile
scales 3 to 4, ovate-lanceolate to lanceolate, 2—3 X
0.9— 1.3 mm: sterile scales 2 or 3 at base of spike¬
let, ovate-elliptic, 1-2 X 0.3— 1.2 nun. Flowers bi¬
sexual; stamens 3, the anthers ca. 1 mm long, apieulate
at apex, basifixed, thecae parallel, longitudinally
dehiscent; style 2-branched, equaling to 2/3 length
of unhranehed portion, long-exserted from subtend¬
ing scale. Achene biconvex, elliptic to broadly so
or subrounded, 0.6-0.8 X 0.5— 0.7 mm, rounded at
apex, obtuse at base, transversely rugulose with 5 or
6 rugae per face, narrowly cellular-reticulate along
margins and at base, shining, glossy, stramineous to
brown, often with a longitudinal gray stripe medially;
epidermal cells linear, vertically oriented; style base
shallowly triangular, 0.1 X 0.2-0.3 mm. abruptly
short-attenuate at apex, shallowly 2-lobed at base,
or indistinctly so, the ends of the lobes often slightly
upturned, brown or blackish; bristles absent.

Distribution and habitat. French Guiana and
Surinam. Seeps and pools in savane-roche (rock sa¬
vanna) and granitic outcrops, 280—450 m.

Distinguishing features. Among the species of
Rhynchospora sect. Tenues, R. saxisavannicola can
be distinguished by the combination of a strongly
flexuose spikelet rachilla, elliptic to broadly elliptic
achene body, and shallowly triangular, abruptly
short-attenuate style base.

Paratypes. SURINAM. Sipaliwini Savanna area on
Brazilian frontier, in valley on saddle betw. two “4-Ge-
broeders” mtn. tops, in trickling zone at edge of sloping

Volume 15, Number 3
2005

Strong

Rhynchospora sect. Tenues

483

Figure 2. Rhynchospora saxisavannicola M. T. Strong. — A. Habit. — B. Detail of junction of sheath and leaf blade.
— C. Cross section of leaf blade. — 1). Detail of inflorescence. — E, F. Spikelets. — G. Spikelet scale. — H. Achene.
Drawn from the holotype, J. J. DeGranville, /.. Aliker & C. Sarthou 15283 (US).

granitic outcrop, 450 m. 10 Oct. 1968, F. //. F. Olden-
burger, R. Norde & ./. R. Schulz 254 (U); Sipaliwini Sa¬
vanna area on Brazilian frontier, 280 m, 24 Jan. 1969. F.
11. F. Oldenburger, R. Norde & J. F. Schulz 986 (U).

Acknowledgments. This paper is number 97 in
the Smithsonian Institution’s Biological Diversity of
the Guiana Shield (BI)GS) Program publication se¬
ries. A florist ic treatment of the Gyperaeeae (sedg¬
es) for llie Flora oj the Guianas is in preparation
by the author and collaborators. I thank Dan Nic-
olson (US) and Gerry' Moore (BKL) for reviewing
the manuscript; an anonymous reviewer for cor¬
recting t he Latin; and Alice Tangerini (US) for ren¬
dering the superb illustrations.

Literature Cited

Barthlott, W. 1981. Epidermal and seed surface characters
of plants: Systematic applicability and some evolution¬
ary aspects. Nordic J. Bot. 1: 345—355.

Eller, B. M. 1979. Die strahlungsokologische Bedeutung
von Epidermisauflagen. Flora 168: 146—192.

Kiikenthal, G. 1949. Vorarbeiten zn einer Monographic
der Rhynchosporideae. Bot. Jahrb. 74(1): 375—509.

- . 1950. Vorarbeiten zu einer Monographic der

Rhynchosporideae. Bot. Jahrb. 75(1): 90—126, (2): 127—
195.

Rocha, E. A. & M. Luceno. 2002. Estudo taxonomico de
Rhynchospora Vahl Segao Tenues (Cyperaceae) no Bras¬
il. Hoehnea 29(3): 189-214.

Strong, M. T. 2001. Novelties in Rhynchospora (Cypera¬
ceae) lrom the Guianas. Novon 1 1: 261—273.

- . 2004. An Electron Microscopy Study of the Outer

Pericarp Surface of Achenes (Fruits) of Rhynchospora
(Cyperaceae) in the Guianas, South America, Bearing
on the Delimitation of Species and Sections, and lo
Clarify the Taxonomy and Distribution of Species Oc¬
curring in the Guianas. Ph.l). Dissertation, George Ma¬
son University, Fairfax, Virginia.

Thomas, W. W. 1998. Rhynchospora. Pp. 610-639 in P.
E. Berry, B. K. Holst cN K. Yatskievych (editors). Flora
of the Venezuelan Guay ana, Vol. 4. Missouri Botanical
Garden Press, St. I.ouis.

Pinellia koreana (Araceae), a New Species from Korea

Kyoung-Hwan Tae and Joo-Hwan Kim

Division of Life Science, Daejeon University, Daejeon 300-716, Korea. sysbot@dju.ac.kr;

kimjh@clju.ac.kr

Abstract. A new species in Araceae, Pinellia
koreana, is described from South Korea. Pinellia
koreana differs from P. yaoluopingensis, P. pedati-
secta, and P. ternata by having 3-foliolate leaves
with bilobed lateral leaflets when mature and only
petioles bearing bulbils. A polyploid chromosome
number is reported for the new species (2/i = 104),
representing an octoploid series from a base num¬
ber x — 13.

Key words: Araceae, chromosome number, Ko¬
rea. Pinellia.

The genus Pinellia Tenure (Araceae) consists of
about 10 species restricted to warm temperate and
temperate East Asia (Mayo et al., 1997). All spe¬
cies are small plants, and some are popular as hor¬
ticultural subjects. In Korea and China Pinellia is
used for herbal medicine, particularly for chest
complaints.

Taxonomic studies on the genus have concen¬
trated on morphology (Li et ah. 1977; Wu & Li,
1979; l.i et ah, 1997a) and cytological studies (Ku-
rakubo. 1940; Ito, 1942; Suzuka, 1953; Jones,
1957; Marehant, 1972; Li et ah, 1997b).

Pinellia in Korea was reported by Nakai (191 1),
Chung et ah (1937), and Chung (1956) as consist¬
ing of a single species, Pinellia lernata (Thunberg)
Breitenbach. Yang (1958) reported that the genus
Pinellia in Korea consisted of two taxa, Pinellia
lernata and Pinellia lernata var. viridis Makino, but
P. lernata var. viridis was a synonym of P. lernata.
fob (1978) undertook a taxonomic study to inves¬
tigate differences of morphological features, ana¬
tomical structures, and histochemical and cytolog¬
ical characteristics between P. lernata and P.
tripartita Schott, the latter not recorded from Korea.

While studying the genus in Korea, we found a
population of Pinellia that does not belong to any
taxon hitherto described. The plants grow along the
Hwa-um valley of the southern slope of Mt. Chi ri .
The population of P. koreana in the type locality
occupies a small area of ca. 100 m2 and consists
of about 25 individuals. We herein describe and
illustrate this new species of Pinellia from Korea.

Pinellia koreana K. II. Tae & J. II. Kim. sp. nov.
TYPE: Korea. Jeollanam-do Province, Mt. Chi¬
rk grassy slopes in Hwa-um valley, 480 in. 10
June 1999, K. II. Tae 99-001 (holotype, TUT).
Korean name: Ji-Ri-Ban-Ha.

Herba perennis usque ad 40 cm alia; tuber globosum
1 .0-2.0 cm diametro. Folia 2, ovata, 3-foliolata, iterum
scissa in extrinsecus 2-foliolata, 10—12 cm longa, 7-1 1
cm lata; petiolis 13-20 cm longis. Pedunculi 23-30 cm
longi; spatha ereeta. Numerus ehromosomatum 2 n = 101
(8x).

Perennial herb, to 40 cm tall. Tuber globose,
1.0-2. 0 em diam. Leaves 2. Petiole greenish, 13—
20 cm long, base sheathing. Bulbils present in
sheath at lower part of petiole, 7-8 mm diam. Blade
ovate in outline, green, trifoliolate, lateral leaflets
bilobed when mature, green, 10-12 cm long, 7-1 1
cm wide. Peduncle green, solitary, longer than the
petiole, 23-30 em. Spathe green outside, typically
light purplish green within, tube narrowly cylindric,
6—7 em long. Spadix with pistillate portion basal,
1.5— 1.8 cm long, staminate portion above, 0.9— 1.2
cm long, and appendix violet to green (Fig. 1). Ber¬
ries ovoid, greenish white, stigma persistent. So¬
matic chromosome number 2 n — 104.

Flowering May— June.

fhe new species (Fig. I) appears to be closely
related to Pinellia ternata. The characteristic fea¬
tures and differences among related species are
summarized in Table 1. The new species differs
from the three other species in the leaf shape and
the position of the bulbils. Somatic chromosome
numbers of Pinellia koreana were established as 2n
— 104. This represents the first cytological report
for the new species, which is seen as octoploid,
from an x = 13 series. The plants examined cyto-
logically were collected from the type population
on Mt. Cliiri on 10 June 1999 (K. //. Tae 99-001,
TUT).

Pinellia koreana is distinguished from P. yaolu¬
opingensis X. II. Guo & X. L. Liu, P. pedatiseeta
Schott, and P. ternata in having trifoliolate leaves
with bilobed lateral leaflets when mature, only pet¬
ioles bearing bulbils (Fig. 1), and somatic chro¬
mosome numbers (2 n = 104, Table 1, Fig. 2). Re-

INovon 15: 484-487. Published on 21 Seitembeb 2005.

Volume 15, Number 3
2005

Tae & Kim

Pinellia koreana (Araceae)

485

lcm

Figure 1. Pinellia koreana k. H. Tae & J.-H. Kim. — A. Habit. — B. Spathe and spadix. — C. Monoecious flowers,
showing staminate flowers on the upper portion, and pistillate flowers on the lower portion of the spadix. — I). Young
leaf. — E. Mature leaf. Drawn from the holotype, Tae 99—001 (TUT).

486

Novon

fable 1. Diagnostic morphological and cytologieal characters of the new species and closely related taxa. The
chromosome numbers of P. yaoluopingensis, P. pedatisecta, and P. ternata are based on Kurakubo (1940); llo (1942);
Malvesin-1 abre (1945); Huttleslon (1953); Suzuka (1953); Jones (1957); Marchant (1972); Guo and Liu (1986); and l.i
el al. (1997b).

Characters

P. yaoluopingensis

P. pedatiseeta

P. ternata

P. koreana

Distribution

China

China

China, Korea, Japan

3

Korea

Leaflet number (blade)

3(to 5)

6 to 1 1

3 (5-parted)

Petiole

deep green with
purple spots

green

green

green

Bulbils

present only in
tuber-top

absent

present in petiole
and leaf

present only in
petiole

Spadix

green

yellow

green

green

Flowering period

May— July

June— July

May-July

May— June

Somatic chromosome #

'In = 26

CM

II

2 n = 28, 54, 72. 99.

2 n = 104

(basic #)

(x = 13)

(x = 13)

108. 1 16, 128
(x = 7, 8. 9, 29)

(x = 13)

cently, Li et al. (1997) disc •ussed the taxonomic
relationship among Pinellia species in China based
on bulbil patterns and chromosome numbers. They
suggested that the basic chromosomes x = 7, 8, 9
were derived from x = 13. Therefore, R koreana
may be1 interpreted as one of the primitive chro¬
mosomal type x = 13. from an octoploid series to
2 n = 104.

Acknowledgments. This research was supported
by a grant (PMM)I301— 01) from the Plant Diversity
Research Center of the 21st Century Frontier Re¬

search Program funded by the Ministry of Science
and Technology of the Korean government. We
thank Miss Hye-ryon Kim for the illustration of the
type specimen.

1 .iterature Cited

Chung, T. H. 1956. Rep. FI. Isl. Jin-Do. J. Sungkyunkwan
Univ. 8: 69—95. 1 1 n Korean.]

- . B. S. Do, I). B. Lee & VI. J. l,ee. 1937. Nomina

Plantarum Koreanum. Chosen Nat. History Inst. Seoul.

| In Korean.]

1 0/jm

I

figure 2. A chromosome photograph of Pinellia koreana (2 n = 104).

Volume 15, Number 3
2005

Tae & Kim

Pinellia koreana (Araceae)

487

Guo, X. H. & X. L. Liu. 1986. A new species of Pinellia
from Anhui Province. Acta Bot. Yunnan. 8: 223—224.

Huttleston, I). G. 1953. A Taxonomic Study of the Tem¬
perate North American Araceae. Ph.D. Thesis. Cornell
University. [Unpublished.]

Ito, T. 1942. Chromosomen und Sexualitat der Araceae. I.
Somatische Chromosomenzahlen einiger Arten. Cytolo-
gia 12: 313-325.

Jones, G. E. 1957. Chromosome numbers and phyloge¬
netic relationships in the Araceae. Diss. Abstr. 17:
2394.

Kurakubo, Y. 1940. Uber die Chromosomenzahlen von
Araceae-Arten. Bot. & Zool. 8: 1492.

Li, H., Y. Shiao & S. L. Tseng. 1977. Claves diagnosticae
et taxa nova aracearum sinicarum. Acta Phytotax. Sin.
15(2): 87-109.

l i, M. W., 1). Gu & Y. Liu. 1997a. Several variation pat¬
terns and their evolution of Pinellia (Araceae). J. Wu¬
han Bot. Res. 15(4): 317—322. [In Chinese.]

- . I). Gu, Y. Liu & P. Hsu. 1997b. Relationship

between occurrence of bulbils and chromosome number

and ploidy in Pinellia (Araceae). Acta Phytotax. Sin.
35(3): 208—214. 1 1 n Chinese.]

Malvesin-Fabre, G. 1945. Contribution it la Caryologie des
Aracees. These. Sciences. Univ. Bordeaux, France. [Un¬
published.]

Marchant, C. J. 1972. Chromosome variation in Araceae:
IV. Kew Bull. 26: 395-404.

Mayo, S. J.. J. Bogner & P. C. Boyce. 1997. The Genera
of Araceae. Royal Botanic Gardens, Kew.

Nakai, T. 1911. Flora Koreana. Pars Secunda. J. Coll. Sci.
Imp. Univ. Tokyo. [In Japanese.]

Suzuka, O. 1953. Chromosome numbers in pharmaceuti¬
cal plants II. Seikeu Ziho. 6: 79. |ln Japanese.]

Toh, C. A. 1978. A comparative morphological study on
Gen. Pinellia. Korean Life Sci. Inst. 22: 85—102. |ln
Korean.]

Wu, C. Y. & H. Li. 1979. Pp. 200-206 in Flora Reipub-
licae Popularis Sinicae. Tomus 13 (2). Science Press,
Beijing.

Yang, I. S. 1958. Inv. Rep. PL in Jin-do Isl. J. Kyungpook
Univ. 2: 323—352. | In Korean. |

A New Combination in Chinese Aesculus (Hippocastanaceae)

Nicholas ./. Tu rl an (l

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

nicholas.turland@mol)ot.org

Nianhe Xia

South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510520,
People's Republic of China, nhxia@scih.ac.cn

Austkact. In tlie context of the present authors’
revision of the Hippocastanaceae for tfie forthcom¬
ing Flora of China , Volume 12, two Chinese en¬
demic taxa, Aesculus chinensis Bunge and A. wil-
sonii Rehder, are treated as nonspecific. The latter
taxon, represented hy wild plants, is here recom¬
bined as A. chinensis var. wilsonii (Rehder) 'Fin land

N. II. Xia. comb, et slat. nov. Aesculus chinensis
var. chinensis, with A. chekiangensis Hu & W. P.
Kang (A. chinensis var. chekiangensis (Hn & W. P.
Fang) W. P. Fang) in synonymy, is represented by
cultivated plants and is unknown with certainty in
a wild stale.

Key words: Aesculus, China, Hippocastanaceae.

The genus Aesculus F. (Hippocastanaceae) com¬
prises ea. 12 species of deciduous trees with pa I -
mate leaves, showy flowers borne in thyrses, and
large seeds (buckeyes or horsechestnuts). They are
disjunctly distributed in North America (including
Raja California), southeastern Furope, and Asia
from the Himalayas to Japan.

In the context of the present authors’ revision of
the Hippocastanaceae for the forthcoming Flora of
China, Volume 12. four species are here recognized
as occurring in China. One is endemic: Aesculus
chinensis Bunge, under discussion here; one is rath¬
er widely distributed from Sikkim to northern Viet¬
nam and southern China: .4. assaniica Griffith; and
two are introduced: A. hippocastanum L., from the
Balkan peninsula in Europe, and A. turbinata Illu¬
me. from Japan.

Aesculus chinensis (Bunge, 1833: 10) was origi¬
nally described from cultivated material from the
Beijing area. It is characterized by having petio-
lules 0.5— 2.5(— 3) cm long, leaflet blades 8— 25(— 30)
cm long; thyrses cylindric, 15—35 cm long; petals
white with yellow spots, subequal, oblong-obovate
to oblong-oblanceolate, 8—14 X 3—5 mm; anthers
1 — 1.5 mm long; and capsules dotted but smooth.

Cultivated plants from southern Jiangsu and

northern Zhejiang provinces were described as Aes-
culus chekiangensis Hu & W. P. Fang (Fang, 1962:
86; repeated by Fang & Hu, 1965: 218), which was
later reduced to varietal rank (Fang, 1981: 277), as
,4. chinensis var. chekiangensis (Hu & W. P. Fang)
W. P. Fang. However, they are scarcely distinguish¬
able from plants in the Beijing region and in fact
have some features that were thought to distinguish
A. wilsonii Rehder (see below), namely a thinner
pericarp and a smaller liilum (Rehder, 1913: 499;
Hardin. 1960: 35).

Aesculus wilsonii (Rehder, 1913: 498) was de¬
scribed from wild plants in Chongqing, western Hu¬
bei. and Sichuan provinces, and in its typical form
differs from ,4. chinensis in the above-mentioned
features and in having leaf blades abaxially pubes¬
cent (vs. more or less glabrous) with a more round¬
ed base (vs. cuneate). Within the range of A. wil¬
sonii occur numerous intermediates with A.
chinensis, as was noted by Hardin (1960: 34), who
hypothesized that the latter species might only be
a cultivated form of the former. File comment by
Fang (1981: 276. in Chinese) about native plants
of A. chinensis existing in the (Jin Ling mountain
range (southern Shaanxi province) was possibly
based on a comment by Rehder (1913: 500). which
was questioned by Hardin (1960: 34), who noted
“the specimens which I examined from Shensi [i.e.,
Shaanxi] represent a form of A. wilsonii Indeed,
among the intermediates mentioned above are the
same gathering seen by Hardin (F. N. Meyer 1725,
A, MO) and another from Mianxian County in
southwestern Shaanxi (K. ,/• Fu 5650, MO).

While the extreme forms can easily be assigned
to either Aesculus chinensis or .4. wilsonii, the ex¬
istence of so many intermediates makes it impos¬
sible to delimit species and difficult even to justify
infraspecific taxa. We have therefore decided to
recognize one variable species, A. chinensis, but
nevertheless to retain a separate status for the wild
plants, as A. chinensis var. wilsonii (Rehder) Tur-

Novon 15: 488-489. Published on 21 September 2005.

Volume 15, Number 3
2005

Turland & Xia
Chinese Aesculus

489

land & N. II. Xia. Such a treatment would not be
at odds with two recent analyses. Forest et al.
(2001) published a cladistic analysis of Aesculus
based on morphology oidy. The two most parsimo¬
nious trees obtained showed no resolution between
,4. chinensis and A. wilsonii, or for that matter be¬
tween those two taxa and the only other native Chi¬
nese species: A. assamica. Xiang et al. (1998) pub¬
lished a molecular phylogenetic analysis in which
the single most parsimonious tree based on internal
transcribed spacer (ITS) sequences showed A. chi¬
nensis and A. wilsonii as sister taxa, the former sup¬
ported by eight nucleotide changes and the latter
by six. Aesculus assamica (therein as “A. wangii' ’)
was shown as sister to the A. chinensis— A. wilsonii
clade, supported by 30 nucleotide changes. “Aes¬
culus wangii Hu” was not validly published by ei¬
ther Fang (1962: 99) or Fang and Hu (1965: 227,
as “ wangi ’) because in both cases two gatherings
were indicated as types ( Sino-Ross . Exped. 580, PK,
and K. 17. Feng 12943 , PE), both of which are
clearly referable to A. assamica. flic analysis of
Xiang et al. (1998) did not include other accessions
of A. assamica in addition to that given as “4. wan¬
gii."

The two varieties of Aesculus chinensis may be
summarized as follows. The provincial distributions
are based on the herbarium specimens examined
by the presenl authors (A, GH, IBSC, MO, PE) and
the data given by Rehder (1913), Hardin (I960),
and Fang ( 1 962. 1 98 1 ).

Aesculus chinensis var. chinensis (including 4.
chekiangensis ) has leaflet blades abaxiallv glabrous,
or grayish tomentose on the veins when young, and
the leaflet blade base cuneate to broadly so. It is
cultivated, especially in temple grounds and by
houses, at elevations below 800 m, and is not
known with certainty in a wild state. Il has been
recorded from the following provinces in China:
Hebei, northern Henan, southern Jiangsu, southern
Shaanxi, southern Shanxi, and northern Zhejiang.

Aesculus chinensis var. wilsonii has leaflet blades
abaxially more or less uniformly grayish tomentose
or villous, or grayish tomentose on the veins (some¬
times only when young), and the leaflet blade base
cuneate to rounded or slightly cordate. It grows in
broad-leaved forests, near streams in tall shady for¬

ests, woods, thickets, mountain and hill slopes, ra¬
vines, and roadsides, at elevations of 600— 2000(—
2300) m, and is also sometimes cultivated. It has
been recorded from the following provinces in Chi¬
na: Chongqing, southern Gansu (Kangxian: E S.
Lian 96405 , MO), northern Guangdong, Guizhou,
southwestern Henan, western Hubei, Hunan, west¬
ern Jiangxi, southern Shaanxi, Sichuan, and north¬
eastern Yunnan.

Aesculus chinensis Bunge var. wilsonii (Rehder)
Turland & N. H. Xia, comb, et stat. nov. Bas-
ionym: Aesculus wilsonii Rehder, in Sargent.
PI. Wilson. 1: 498. 1913. TYPE: China. Si¬
chuan: Wenchuan County, “Hsin Wen Ping,
3400 ft.,” I June 1908 (fl), E. //. Wilson 200
(lectotype, designated by Hardin (1960: 35),
A, sheet no. 50582).

Acknowledgments. We thank David E. Boufford
and Anthony R. Brach (both A, GH) for making the
first author’s visit to Harvard University Herbaria
in February 2005 such a pleasant one and for pro¬
viding a copy from the Harvard library of Fang’s
1962 paper. We also thank Guanghua Zhu (MO) for
obtaining digital images of Aesculus types at PE.

Literature Cited

Ifunge, A. A. von. 1833. Enumeratio Plantarum, quas in
China Boreal i Collegit. St. Petersburg.

Fang, W. P. 1962 [“I960”]. Additamenta ad floram sinen-
sem I. Hippocastanaceae. Acta Sci. Nat. Univ. Sze¬
chuan. 1960(3): 77—125.

- . 1981. Hippocastanaceae. Pp. 274-289 in W. P.

Fang (editor). Flora Reipublicae Popularis Sinicae, Vol.
46. Science Press, Beijing.

- & H. H. Hu. 1965. New species of Aesculus from

China. Acta Sci. Nat. Schol. Super. Sin.. Biol. 1965(3):
2 1 7-237.

f orest, F, J. N. Drouin, R. Charest, L. Brouillet & A.
Bruneau. 2001. A morphological phylogenetic analysis
of Aesculus L. and Billia Peyr. (Sapindaceae). Canad. J.
But. 79: 154-169.

Hardin, J. W. 1960. Studies in the Hippocastanaceae, V.

Species of the Old World. Brittonia 12: 26—38.

Rehder, A. 1913. Hippocastanaceae. Pp. 498—500 in C.
S. Sargent (editor), Plantae W'ilsonianae, Vol. 1 . Cam¬
bridge Univ. Press, Cambridge, Massachusetts.

Xiang, Q. Y.. 1). J. Crawford, A. I). Wolfe, Y. C. Tang &
C. W'. DePamphilis. 1998. Origin and biogeography of
Aesculus L. (Hippocastanaceae): A molecular phyloge¬
netic perspective. Evolution 52: 988—997.

Eurya luchunensis, a INew Species of Tlieaceae from

Yunnan, China

Jinhua Wang, Hong Wang, and Dezhu Li

Kunming Institute of Botany, the Chinese Academy of Sciences, Kunming 650204, People's
Republic of China, wangjh@mail.kib.ac.cn; wanghong@mail.kib.ac.cn; dzli@mail.kib.ac.cn

Abstkact. Eurya luchunensis J. H. Wang & H.
Wang, a new species of Tlieaceae from Luchun
County in southern Yunnan, China, is described
and illustrated. It was collected in a tropical broad¬
leaved secondary forest or montane rain forest on
Mt. Huanglian-shan. This species is very similar to
Eurya pseudocerasifera Kobuski, from which it dif¬
fers by having lanceolate to oblong-lanceolate
leaves, 20- to 24-locellate anthers, and the first-
year branches, as well as the pedicels, bracteoles,
and sepals, being glabrous or subglabrous.

Key words: China, Eurya luchunensis, Thea-
ceae, Yunnan.

established in 1785 by Thunberg, Eurya is a
genus of evergreen shrubs or small trees, rarely
larger trees to 2-5 m tall. There are over 130 spe¬
cies distributed primarily in eastern and southeast¬
ern Asia. Indo-Malesia. and islands in the south¬
western Pacific Ocean. There are 81 known species
for China, among which approximately 41 occur in
Yunnan (Ming & Bartholomew, in press; Ling,
1998).

During our study of all specimens of the genus
Eurya deposited in the KUN herbarium, we found
collections from Luchun County, southern Yunnan,
that were rather similar to E. pseudocerasifera Ko¬
buski, which is distributed in southwestern to
northwestern Yunnan and southeastern Xizang in
China. These specimens have 20- to 24-locellate
anthers, are glabrous or subglabrous in the first-
year branches, pedicels, bracteoles, and sepals,
and have lanceolate to oblong-lanceolate leaves.
However, the anthers are 5- to 10-locellate in E.
pseudocerasifera, which further differs by being pu¬
bescent in the first-year branches, pedicels, brac¬
teoles, and sepals, as well as having leaves that are
oblong-elliptic. The Luchun specimens represent a
new species.

Eurya lurliiineiisis J. 11. Wang & II. Wang, sp.
nov. TYPE: China. Yunnan: Luchun County,
Mt. Huanglian-shan, 1700 m, 19 Oct. 1973.
I). I ). Tao 1101 (holotype, KUN). Figure 1.

Euryae pseudocerasiferae valde affinis, sed foliis angus-
te oblongo-lanceofatis vel lanceolatis, (9— )11— 18 cm lon-
gis. 2^1 cm latis. apice falcato- vel caudato-acuminatis 2-
3 cm longis, subtus secus costam adpresso-pilosis,
postremo saepe glabrescentibus, nervis lateralibus reti-
eulatisque utrinque prominentibus, petiolis 1-1.3 cm lon¬
gis, bracteolis sepalisque glabris, tbecis antheraruni 20 to
24 locufatis. ovariis glabris differt.

Shrubs or small trees, 2—5 m tall; first-year
branches brown, terete, glabrous, or sparsely pu¬
bescent at the apex; terminal buds pubescent. Leaf
blades leathery, narrowly oblong to lanceolate, (9—
)1 1 — 18 X 2^4 cm, apex long-acuminate with an
acumen 2—3 cm long, base widely cuneate, margin
entire or sparsely serrate near apex, dark green
above, shiny, glabrous, light green below, midribs
impressed above and prominent below, sparsely ap-
pressed pubescent along the midribs, becoming
glabrescent later, drying purple, secondary veins 1 1
to 13 on each side of midrib, prominent on both
sides together with the reticulate veins; petioles 1 —
1.3 cm long, deeply impressed above, sparsely pu¬
bescent or glabrescent below. Inflorescence of sol¬
itary axillary flowers or 2 to 4 in a cluster; pedicels
4—6 mm, sparsely pubescent or glabrescent; sta-
minate flowers: with bracteoles 2 or 3, broadly
ovate or orbicular, 1.5—2 nun. glabrous, margin eil-
iolate; calyx with 5 sepals, orbicular, ca. 4 mm
diam., apex rounded and with a reluse tip, outside
glabrous, sparsely pubesecent, margin ciliolate; co¬
rolla with 5 petals, not fully open; stamens ca. 16
to 18, anthers ca. 2.5 mm long, 20- to 24-locellate;
pistilloid ovaries ca. 1.5 mm long, glabrous; pistil¬
late flowers: with bracteoles 1 — 1.5 mm long, calyx
with 5 sepals, sepals ca. 2.5 mm diam. and corolla
with 5 petals, only immature seen; ovary globose,
ca. 2 mm diam., glabrous, style 3-parted. Fruit
ovoid, 5.5— 6.5 mm long, 5—6 mm diam., glabrous,
with persistent styles ca. 4 mm long, apex 3-parted.

Distribution and habitat. Eurya luchunensis oc¬
curs in a tropical broad-leaved secondary forest or
montane rain forest, from 840 to 1900 m. So far
the new species is only known from its type locality,
on Mt. Huanglian-shan, Luchun County, southern

Novon 15: 490-492. Published on 21 September 2005.

Volume 15, Number 3
2005

Wang et al.

Eurya luchunensis (Theaceae)

491

Figure I. Eurya luchunensis J. II. Wang & H. Wang. — A. Flowering branch. — B. Pistillate or slanunate flower,
showing braeteoles, calyx, and corolla. — C. Detail of pistillate flower showing longitudinal section. — D. Pistillate or
staminate flower, showing braeteoles and calyx. — E. Detail of degraded pistil and single stamen and filaments of other
stamens with anthers removed. — E Stamen. — E. fruit. Drawn from the holotype, I). I )• Tno 1101 (KEN).

492

Novon

Yunnan. China. According to the collector’s note, it
is easy to find this species just above this area.

Phenology. Flowering middle October to early
December, fruiting probably in later May (of the
following year).

Paratypes. CHINA, Yunnan Prov.: Luchun County,
Mt. Huanglian-shan, I 750 m, 5 Nov. 1974, Luchun Exp.
690 (K UN); Mt. Huanglian-shan, 1700-1900 m. 15 Oct.
I *195, S. K. Wu el al. 049 (KLIN); Mt. Huanglian-shan,
1700-1900 m, 19 Oct. 1995, S. K. Wu el al. 548 (KlIN);
Mt. Huanglian-shan, 1700-1900 m, 23 Oct. 1995, S. K.
10/ el al. M8 (MO); Mt. Huanglian-shan, 840-1300 in,
30 Oct. 1995, S. K. Wu el al. 2101 (KUN); Mt. Huanglian-
shan, 1700-1900 m. 22 Oct. 1995, S. K. Wu el al. 8208
(KlIN).

Acknowledgments. We are indebted to Ming

Tien-lu and Shu Zhi-yun (KlIN) and Robert R. Mill
(E) for their critical reading of the manuscript, and
to th<‘ curator of the KUN herbarium, Kunming In¬
stitute of Rotany, the Chinese Academy of Sciences,
for enabling the study. We also thank Wu Xi-lin for
preparing the illustration. This research was sup¬
ported by a grant from the National Program for R
& D Infrastructure and Facility Development
(20041 DKA30430).

literature Cited

Ling, L. K. 1998. Ternstroemoideae. Flora Reipublicae
Popu laris Sinieae, Vol. 50(1). Science Press, Beijing,
China.

Ming, T. L. & B. Bartholomew. In press. Theaeeae. In:

V. Wu & P. H. Raven (editors), f lora of China, Vol. 12.
Science Press, Beijing, and Missouri Botanical Garden
Press, St. Louis.

Validation of l he Name Hex Xaquipernyi
J. B. Gable (Aquifoliaceae)

Alan T. Whittemore

US National Arboretum, 3501 New York Ave. NE, Washington, DC 20002-1958 U.S.A.

whittema@ars.usda.gov

Abstract. Ilex Xaquipernyi J. B. Gable ex \\ liit-
temore (Ilex aquifolium X I. pernyi), a name that
lias been commonly used in the horticultural lit¬
erature anti the nursery trade for some 60 years, is
validated here. It is known only from artificial
crosses.

Key words: Aquifoliaceae, Ilex.

The genus Ilex L. (holly) consists of some 400
species of shrubs, native to all continents except
Antarctica. Many species bear attractive red berries
in fall and winter, tolerate a wide variety of soils,
and are easy to propagate and grow, so hollies are
cultivated frequently almost worldwide. Temperate
evergreen species are particularly popular since
some of them are very cold-tolerant; they can pro¬
vide winter greenery in climates where few other
evergreen plants will survive.

In the course of checking data for hollies at the
National Arboretum, it was noted that a well-known
name in widespread use in the holly literature, in
general horticultural literature, and in the nursery
trade. Ilex Xaquipernyi, has never been validly
published. This name is validated and typified
here.

Ilex Xaquipernyi J. B. Gable ex V\ hittemore, sp.
nov. TYPE: U.S.A. District of Columbia: be¬
side the parking lot, Holly-Magnolia Collec¬
tion, U. S. National Arboretum, NA 17762. 21
Apr. 2003, A. T. Whittemore 03-005 (holotype.
NA; isotypes. A, K. MOB. MO, TUS).

Hybrid formula: Ilex aquifolium L. X Ilex pernyi
Franchet.

Frutex vel arbor sempervirens ad 6 m alta; ranuili hor-
notini glabri vel puberuli. Lamina lanceolata usque ovala
vel elliptica, uudulata, 31—63 mm longa, 14—35 mm lata,
spinis grandibus, 2—1 in quoque margine. Flores 1-meri,
fasciculati, pedicellis 2—3 mm longis.

Shrubs or small trees to 6 m tall; bark light
gray-brown, rough but not fissured; twigs green or
brown, 2—3.5 mm thick, glabrous or puberulent.
Leaves evergreen; petiole 3—6 mm long, glabrous

or puberulent; blade thick and rigid, glabrous (or
base of midrib puberulent), lanceolate to ovate or
elliptical, weakly to strongly undulate, 31—63 X
14—35 mm (excluding spines); upper surface
shiny; base rounded to broadly obtuse or almost
truncate; apex acuminate and spine-tipped; mar¬
gins narrowly recurved, each side of leaf with 2 to
4 large triangular spine-tipped teeth up to 5 nun
long. Flowers 4-merous, densely clustered in leaf
axils on previous years’ growth, pedicels 2—3 mm
long, glabrous or puberulent, petals white. Berries
red, spherical, 7—9 mm long, each with four py¬
renes.

Illustrations. Figure I: Galle (1952: plate

III).

The name Ilex Xaquipernyi has a long history
in the American nursery trade. Horticultural ma¬
terial grown under this name is descended from
artificial crosses between /. aquifolium and /.
pernyi made before 1940 by Joseph Benson Ga¬
ble (1886—1972), a nurseryman of Stewartstown,
Pennsylvania. Gable apparently distributed plant
material labeled with the name "Ilex Xaquiper¬
nyi'’ for trial in the mid-1940s. The name has
been in common use since that time, and appears
in many standard horticultural references (Ev¬
erett. 1981; Huxley et ah. 1992; Meyer et ah.
1994; Cullen et al„ 1997; Galle, 1997; Dirr,
1998). Several cultivars are available in the trade
(Dirr. 1998. lists six), and /. Xaquipernyi has
been used as one parent in several crosses
(Galle, 1997; Dirr, 1998). Despite this long his¬
tory. the name has never been validly published,
and has been in use as a nomen nudum for some
60 years.

The type specimen was taken from a plant of Ilex
Xaquipernyi 'Aquipem'. a cultivar Gable selected
from seed that he obtained in 1933 from the cross
Ilex aquifolium ‘Pyramidalis’ X I. pernyi (Anony¬
mous, 1953). The cross is not known to occur spon¬
taneously, though it might be expected where the
two species are cultivated together.

Novoin 15: 493-494. Published on 21 September 2005.

494

Novon

Figure 1. Ilex Xaquipernyi .1. B. Gable ex Whittemore.
— Flowering stem (A. T. Whittemore 03-005).

Acknowledgments. I thank Kevin Conrad for
help locating information on these plants, John
Wiersema and Dan Nicolson for comments on the
manuscript, and Don Voss and Mark Koh for help¬
ful discussions.

I .iterature Cited

Anonymous. 1953. Preliminary holly checklist. Holly Soc.
Amer. Bull. 6: 1—56.

Cullen, J. et al. (editors). 1997. The European Garden
Flora: A Manual for the Identification of Plants Culti¬
vated in Europe, Both Out-of-Doors and Under Glass,
Vol. 5. Cambridge Univ. Press, Cambridge.

Dirr. M. A. 1998. Manual of Woody Landscape Plants, ed.
5. Stipes Publishing, Champaign, Illinois.

Everett. T. II. 1981. The New York Botanical Garden Il¬
lustrated Encyclopedia of Horticulture. Vol. (>, Id— Ma.
Garland, New York.

Gallo, E C. 1952. Evergreen hollies for the south. Holly
Soc. Amer. Bull. 5: 1-12.

- . 1997. Hollies: The Genus Hex. Timber Press,

Portland.

Huxley, A., M. Griffiths & M. I^vy. 1992. The New Royal
Horticultural Society Dictionary of Gardening, Vol. 2, I)
to K. Macmillan, London.

Meyer, F. G.. P. M. Mazzeo & I). II. Voss. 1994. A Catalog
of Cultivated Woody Plants of the Southeast United
States. United States National Arboretum Contribution
no. 7. U.S. Dept, of Agriculture, Washington.

Epigeneium gaoligongense (Orchidaceae), a New Species from

Yunnan, China

Hong Yu

Laboratory of Ecogenetics, Yunnan University, Kunming 650091, China, hongyu@ynu.edu. en

Shi -(rang Zhang

Inmol Laboratory of Biotechnology, Kunming 650212, China, fisher@public.km.yn.cn

ABSTRACT. Epigeneium gaoligongense Hong Yu
& Shi-Gang Zhang, a new species of Orchidaceae
from Lushui County in western Yunnan, is de¬
scribed and illustrated. It is related to Epigeneium
rot undatum (Lindley) Summerhayes. hut differs
from it by having smaller leaves 2.5-7 cm long,
narrower petals 1 .5—2 mm wide, a broadly ovate-
orbicular lip with much larger lateral lobes, three
indistinct keels and a callus on the disc, and a
shorter column 1-5 mm long.

Key words: China, Epigeneium , Orchidaceae,
Yunnan.

Epigeneium Gagnepain is an orchid genus of ea.
55 species, distributed in tropical and subtropical
Asia. Seven species of this genus have been re¬
ported from China (Tsi, 1999). During a botanical
trip by the authors to the Gaoligong Mountains in
western Yunnan in autumn 2000, several plants of
Epigeneium were collected from Lushui County at
an altitude of nearly 2500 m, where no less than
20 individuals were found growing on rocks or tree
trunks on the edge of a forest. 1 hey were brought
back and cultivated in the greenhouse of the Inmol
Laboratory of Biotechnology in Kunming and flow¬
ered early in August 2001. The species is similar
in habit and to a lesser extent in floral structure to
Epigeneium rotunda! um (Lindley) Summerhayes. a
species widely distributed from southwest China to
India, Myanmar, and the Himalaya regions, from
the latter it differs by having rather small leaves
2.5—7 cm long, narrower petals 1.5—2 mm wide, a
broadly ovate-orbicular lip with much larger lateral
lobes, three indistinct keels and a callus on the
disc, and shorter column 4—5 mm long. We treat it
here as a new species belonging to Epigeneium
sect. Katherinea (A. I). Hawkes) Garay A G. A.
Komero-Gonzalez, which is characterized by having
two terminal leaves on each pseudobulb.

Epigeneium gaoligongense Hong Yu & S. G.
Zhang, sp. nov. TYPE: China. Yunnan: Gao¬
ligong IMt., Lushui County, alt. 2500 m, 26
Aug. 2001, //. Yu A S'. Zhang 101 (holo-
type, PE; isotype, Herbarium, Inmol Labora¬
tory of Biotechnology, Kunming, Yunnan), fig¬
ure I .

Pseudobulbi 3—9 cm distantes, anguste ovoidei vel cy-
lindrico-ovoidei, 1.5-4 cm longi, 5-0 mm diam., apice
bifolii. Folia ovato-elliptica vel elliptica, 2.5-7 X 1. 2-2.0
cm. Scapus 2—3 cm longus, apice 1-florus; ffos ca. 3 cm
diam.; petala lineari-lanceolata, 15-21 X 1.5-2 mm; la-
bellum ambitu late ovato-orbiculare. 1-1.5 X 1.1—2 cm:
lobis lateral ilms erecto-incurvis ovatis 11X0 mm. lobo
intermedio late ovato 0-10 mm longo et lato, disco ob¬
scure 3-carinato et apice carinarum I -calloso; columna 4—
5 mm longa.

Epiphytic plants with an elongate, creeping rhi¬
zome 2-3 mm thick; rhizome usually branched,
densely maroon-sheathed when young. Pseudo¬
bulbs 3-9 cm apart on the rhizome, enclosed by
maroon sheaths when young, narrowly ovoid to cy-
lindrie-ovoid, 1.5-4 cm long, 5-8 mm thick. 2-
leaved at apex. Leaves coriaceous, rigid, deep
green above, [vale green beneath, ovate-elliptic to
elliptic. 2.5-7 X 1.2-2. 8 cm. bifid at apex; petiole
5-1 I nun long. Scape arising from the top of a rath¬
er young pseudobulb, 2—3 cm long, glabrous, with
2 or 3 maroon, lanceolate bracts at base, terminat¬
ing in a single llower; flower ca. 3 cm across; sepals
and petals pale green, usually with obscurely red¬
dish purple markings dorsally; lip greenish color,
densely red-purple-spotted; dorsal sepal lanceolate,
18—23 X 5-6 mm. obscurely 5-nerved; lateral se¬
pals obliquely lanceolate-ovate, 15—19 X 9—10
mm, obscurely 7-nerved; petals linear-lanceolate,
15—21 X 1.5—2 mm, 1-nerved; lip adnate at base
to the top of the column foot, broadly ovate-orbic¬
ular in outline, trilobed, 1 — 1.5 X 1.4—2 cm; lateral
lobes erect-incurved, ovate, ca. 1 1 6 mm: mid-lobe
broadly ovate, 8-10 mm long and wide, acute at
apex; disc with 3 indistinct keels and at their top

Novon 15: 495-497. Publishf.d on 21 Sfitkmbf.h 2005.

496

No von

Figure 1. Epigeneium gaoligongense Hong Yu & S. G. Zhang. —A. Plant. —It. Dorsal sepal, lateral sepal, and petal,
Imnt view. — C. Lip, front view. —I). Column and column-foot, side view. Drawn from the type specimen, Yu & / hung

4 mm

Volume 15, Number 3
2005

Yu & Zhang 497

Epigeneium gaoligongense (Orchidaceae)

a red-purple callus; column stout, 4—5 mm long and
wide, base with a foot 5—7 mm long.

Artificial Key to Chinese Species of Ei>tci:\h:n \i

la. Pseudobulbs with 1 leaf at apex.

2a. Lip rounded, apiculate or slightly emarginate

at apex . E. nakuharaei (Schlechter)

Summerhayes

2b. Lip conspicuously bilobed at apex.

3a. Leaves ovate or broadly ovate, 1—2.3 cm
long; lip oblong-obovate, with its mid-
lobe deeply bilobed at apex . . E. fargesii
(Finet) Gagnepain

3b. Leaves often obovate or obovate-elliptic,

2.5— 4.7 cm long, lip pandurate, with its
mid-lobe emarginated at apex ....
. E. clemensiae Gagnepain

lb. Pseudobulbs with 2 leaves at apex.

4a. Lip with three indistinct keels and a callus

on the disc . E. gaoligongense

Hong 'll! & S. G. /hang

4b. Lip with 2 or 3 fleshy lamellae on the disc.

5a. Petals narrowly lanceolate or narrowly
oblong, less than 4 mm wide.

6a. Labellum without a claw at base;
mid-lobe hooked-aristate at apex
. . E. fuscescens (Griffith) Summerhayes
6b. Labellum with a distinct claw at

base; mid-lobe apiculate at apex

. E. yunnanense

T. Tang & Z. H. Tsi

5b. Petals lanceolate or oblong-lanceolate,
more than 5 mm wide.

7a. Sepals ca. 4.5 cm long; mid-lobe of

labellum sub rhombic ... E. amplum
(Lind lev) Summerhayes
7b. Sepals 2.2— 2.5 cm long; mid-lobe
of labellum subreniform or nearly
orbicular ... E. rotundatum (Lindley)
Summerhayes

Distribution. This new species is known only
from the type locality in Yunnan, China, where it
is rather common growing on rocks or tree trunks
on the edges of forests.

Acknowledgments. The authors express their
sincere thanks to S. C. Chen for his kindly cor¬
recting the manuscript and to S. Q. Cai and Q. H.
Huang for preparing the drawing ol this new spe¬
cies.

Literature Cited

Tsi. Z. H. 1999. Epigeneium. In Z. H. Tsi (editor). Flora
Reipublicae Popularis Sinicae 19: 156—163. Science
Press, Beijing.

Carex capilliculmis (Cyperaceae), a New Name for
Ca rex fi lame n l o s a

Shu- Ren /hang

Institute of Botany, The Chinese Academy of Sciences, Beijing 100003,
People’s Republic of (China, srzhang@ibcas.ac.cn

Abstract. The new name Carex capilliculmis S.
It. /hang is proposed to replace the later homonym
Carex Jilamentosa K. T. Fu. Carex capilliculmis be¬
longs to section Digitatae and is found in western
China.

Key words: Carex capilliculmis, Carex Jilamen¬
tosa, China, Cyperaceae.

While preparing the taxonomic treatment ol the
genus Carex (Cyperaceae) for the Flora of China,
the author found that Carex Jilamentosa K. T. Fu
was an illegitimate later homonym, and a new name
was necessary for the species. Carex Jilamentosa k.
T. Fu was published in volume 1(1) of the Flora
Tsinlingensis (Anonymous, 1976) and has been ac¬
cepted by other Chinese botanists (Dai et al., 2003;
hi. 1 994, 2000). According to the protologue, this
species is closely related to C. lanceolata Boott. It
differs, however, in its more slender culms (ca. 0.5
mm wide), very distant spikes, lealy bracts with
short blades (to 5 cm) and long sheaths (to 3 cm),
a long and filiform peduncle ol tin* lowest spike,
and smaller glumes (3—4 mm), and should be rec¬
ognized as a distinct species. In C. lanceolata, tin1
spikes are relatively congested, the bracts are
spathe-like with or without setaceous blades al the
top and with short sheaths (ca. 1 cm), and longer
glumes (5—6 mm). Carex Jilamentosa k. T. Fu is
distributed in western China from Gansu, Qinghai,
Shaanxi. Sichuan, and Yunnan, whereas C. lanceo¬
lata is found in Anhui, Gansu, Guizhou, Hebei,
Heilongjiang. Henan. Jiangsu, Jiangxi, Jilin. Liao¬
ning. Nei Mongol, Shaanxi, Shandong, Shanxi.
Sichuan, Yunnan, and Zhejiang (hi, 2000). The dis¬
tributions of the two species overlap in Gansu,
Shaanxi, Sichuan, and Yunnan in China.

Unfortunately, the name Carex Jilamentosa was
previously published by Petrie (1916). The hom¬
onym Carex Jilamentosa k. T. Fu (1976) belongs to
section Digitatae (Fries) Christ (Bull. Soc. But.
Belg. 24(2): 13. 1885), which is distributed in Eu¬
rope, temperate Asia, and temperate North America
(Egorova, 1999), while the earlier name C. Jilamen¬
tosa Petrie belongs to a rather different section

Echinochlaenae T. Holm (Amer. .1. Sci. 16: 462.
1903), which is distributed in Australia, New Zea¬
land. and South America (Kiikenthal, 1909). The
species Carex Jilamentosa Petrie is endemic to New
Zealand and recognized in the Flora of New Zea¬
land. volume 3 (Healy & Edgar, 1980). Thus, the
name Carex Jilamentosa k. T. Fu is an illegitimate
later homonym, and should be replaced by a new
name. Here, Carex capilliculmis S. B. Zhang is pro¬
posed to replace C. Jilamentosa k. I. Fu.

Carex capilliculmis S. B. Zhang, nom. nov. Re¬
placed name: Carex Jilamentosa k. T. Fu, FI.
Tsinling. 1(1): 254, fig. 221. 1976, non Petrie,
Trans. & Proe. New Zealand Inst. 1915 (48):
191. 1916. TYPE: China. Shaanxi: Ping-an
Temple, Mt. Taibai, on slopes, 2850 m, 12
June 1959, ./. X. Yang 907 (holotype, Wllk
not seen; isotype, PE).

Distribution and habitat. China (Gansu, Qing-
liai, Shaanxi. Sichuan, and Yunnan; the distribution
in Yunnan was recorded by hi. 1994, 2000). In
forests, among thickets, on rocky cliffs, slopes, and
alpine meadows; I 100—4350 m.

Fin-Yin name. Si gan tai eao.

Specimens examined. CHINA. Gansu: June, Tao River
Expedition 3256 (PE), Qinghai: Haiyan, P. C. Tsoong
8352, 8301 (I’E). Shaanxi: Ml. Taibai, Ping-an Temple.
./. X. Yang 907 (PE). Sichuan: Dawn. .S'. Jiang 9652 (PE);
Jiulong, C. T. Kuan 270211 (PE); kangding. K. I,. Chit
7032 (PE); Yajiang, Anonymous 5453 (PE), C. T. Kuan
510158 (PE).

Because its terminal spike is staminate, its lat¬
eral spikes are pistillate, and its utricles are pu¬
bescent with tapered bases and short beaks, Carex
capilliculmis belongs to section Digitatae (Fries)
Christ. The species also resembles C. cardiolepis
Nees and C. crebra V. Kreczetowicz of the same
section. Carex cardiolepis is distributed in Qinghai,
Sichuan, Tibet, and Yunnan in China, and also in
the Himalayas, kashmir, and Afghanistan, while C.
crebra is found ordy in Gansu, Qinghai, Sichuan,
Tibet, and Yunnan in China (hi. 2000). These three

NIoxon 15: 498-499. Published on 21 September 2005.

Volume 15, Number 3
2005

Zhang

Carex capilliculmis

499

species together with C. lanceolata can he distin¬
guished with the following key.

Ki-'i to Cakex capilliculmis and Belated Species
in China

la. Bracts spathe-like, sheaths short and loosely em¬
bracing culms, without blades or only the lowest
one with setaceous blade.

2a. Leaf blades flat, margins not involute; pistil¬
late glumes 5—6 mm; utricles much shorter

than glumes, ca. 3 mm . Carex lanceolata

Boott

2b. Leaf blades filiform, margins involute; pis¬
tillate glumes 4.5—5 mm; utricles slightly

shorter than glumes, 3.5— 4.5 mm .

. Carex crebra V. kreezetowiez

lb. Bracts leafy, sheaths long and tightly embracing
culms, with short leafy blades.

3a. Culms ca. 1 mm wide; spikes larger, staminate
one 15—20 X 2—2.5 mm, pistillate ones 10—20
X 3 mm, with many pistillate (lowers: pistillate
glumes more than 4 mm; utricles not longer
than glumes, with 2 lateral keels and several

fine veins . Carex cardiolepis Nees

3b. Culms ca. 0.5 mm wide; spikes smaller, sta¬
minate one 7-10 X 1.5—2 mm, pistillate ones
7—10 X 2-3 mm, with few pistillate flowers;
pistillate glumes less than 4 mm; utricles
slightly longer than glumes, only with 2 lat¬
eral keels and no fine venation .

. Carex capilliculmis S. B. Zhang

Acknowledgments. The author appreciates the
valuable comments on the manuscript hy anony¬
mous reviewers. This study was supported hy a
grant (grant number 30370104) from the National
Natural Science Foundation of China.

Literature Cited

Anonymous. 1976. Cyperaceae. In: H. Tsinling. 1(1):

197—275. Science Press, Beijing.

Dai, L. k., S. Y. Liang & S. IL Zhang. 2003. Cyperaceae.
In: B. Z. Fang (editor), FI. Yunnan. 15: 567—803. Sci¬
ence Press, Beijing.

Egorova, T. V. 1999. The Sedges (Carex L.) of Bussia and
Adj acent States (within the limits of the former
U.S.S.B.). St. Petersburg State Chemical-Pharmaceuti¬
cal Academy, St. Petersburg, and Missouri Botanical
Garden Press, St. Louis.

Healy. A. J. & E. Edgar. 1980. Cyperaceae. In: FI. New
Zealand 3: 144—200. P. I). Hasselberg, Government
Printer. Wellington.

kiikenthal, G. 1909. Cyperaceae— Caricoideae. In: A. En-
gler (editor). Das Pflanzenreich (IV) 20 (Heft 38). W.
Engelmann, Leipzig.

Li, P. C. 1994. Cyperaceae. In: W. T. Wang (editor). Vase.
PL Hengduan Mount. 2: 2321—2386. Science Press,
Beijing.

- . 2000. Carex section Digitatae. In: L. k. Dai &

S. Y. Liang (editors), FI. Beipubl. Popularis Sin. 12:
182-209. Science Press, Beijing.

Petrie, D. 1916. Descriptions of new native phanerogams.
Trans. Proc. New Zealand Inst. 48: 186—192.

Lectotypification ol l lie Chinese Species Stachyurus yunnanensis
and I he Identity of S. callosus (Stachyuraceae)

Yuping Zhu & Zhiyun Zhang *

Laboratory of Systematic and Evolutionary Botany, Institute of Botany,
Chinese Academy of Sciences, 20 Nanxineun, Xiangshan, Beijing 100093, China.
zhangzy@ibcas.ac.cn (*author for correspondence); zhuyp@ibcas.ac.cn

Abstract. The Chinese endemic Stachyurus yun¬
nanensis Franchet is leetotypified. Stachyurus cal¬
losus C. Y. Wu ex S. k. Chen, also endemic to
China, is referred to synonymy of .S', yunnanensis
based on the two entities having inseparable leaf
characters and a sympatric distribution. Stachyurus
esquirolii II. Leveille is regarded as a synonym of
5. yunnanensis, as in Li's and Eauener’s previous
treatments of this taxon.

I\ey words: China, Stachyuraceae, Stachyurus,
Yunnan.

Stachyurus yunnanensis Franchet was first de¬
scribed by Franchet (1898) from three collections
from Yunnan and Sichuan provinces of China. In
the protologue, the characters of the flowers and
especially the leaves of this species were described;
leaf characters, e.g., leaf shape and margin serra¬
tion. were emphasized to distinguish S. yunnanensis
from ,S. praecox Siebold & Zuecarini. No holotype
was designated by Franchet, but he did cite three
specimens, which are therefore syntypes (ICBN
Art. 9.4, Greuter et ah. 2000). J. M. Delavay 322,
J. M. Delavay 2234, and I*. Forges I IS 6 are pre¬
served at P. Up to now no lectotype has been des¬
ignated. as far as we could trace. According to Ar¬
ticles 9.9 and 9.10 ol the ICHN. a lectotype may
be designated lor S. yunnanensis and, under the
circumstances, it must be one of the syntypes. The
specimen ./. 47. Delavay H22 (P) is the most com¬
plete, a beautiful specimen with ovate-oblong to
ovate-lanceolate, leathery leaves, with serration at
the margins, and racemes with dense, subsessile,
yellow flowers. It best corresponds with the diag¬
nosis of S. yunnanensis, and it agrees with the spe¬
cies as currently understood.

Stachyurus yunnanensis Franchet. J. Hot. (Morot)
12: 253. 1898. TYPE: China. Yunnan: Mos-
uoying, 23 Apr. 1883, Delavay 322 (lectotype,
designated her**. P)

Stachyurus esquirolii H. l/vciHc, I I. kouy-Tcheou. 416.

1915. TYPE: China. Guizhou: 1700 m, 20 Mar.
1912, J. Esquirol 3517 (holotype, K).

Stachyurus callosus C. Y. Wu ex S. k. Chen, Acta Hot.
Yunnan. 3: 128. 1981. Syn. nov. TYPE: China. Yun¬
nan: Malipo, 20 Jan. 1940, (.. 11. W ang 33275 (ho¬
lotype, kUN; isotvpe. PE).

Specimens examined. CHINA. Yunnan: Pingbian, 26
Oct. 1954, K. M. Feng 5175 (KLIN, PE).

Habitat and distribution. Stachyurus yunna¬
nensis is an evergreen shrub that is narrowly dis¬
tributed in China from northeastern and southeast¬
ern Yunnan to Guangdong provinces, via Sichuan,
Guizhou, Hubei, and Hunan provinces; it usually
grows in calcareous forests between 650 and 1950 m.

Stachyurus esquirolii II. Leveille was first pub¬
lished by H. I ,cveill£ (1914) based on one collec¬
tion, ,/. Esquirol 3517 (E), but it was not recognized
by later workers. I.i (1943) and Lauener (1966)
treated it as a synonym of S. yunnanensis. After
examining J. Esquirol 3517, we found its morpho¬
logical features, such as persistent, ovate-oblong to
ovate-lanceolate, leathery leaves, distinct serration,
and .5—9 cm long inflorescences with yellow flowers,
are consistent with those of S. yunnanensis. We
therefore agree with Li’s (1943) and Eauener’s
(1966) treatments of it.

Stachyurus callosus G. Y. Wu ex S. k. Chen was
first described based on the holotype collection C.
U. Wang 36275 (Chen, 1981). The protologue in¬
dicated that this species was close to S. yunnanen¬
sis but differed only in having elliptic or oblong-
elliptic leaves with irregular and straight teeth.
Stachyurus callosus has been traditionally recog¬
nized as a species by taxonomists since its 1981
publication (Chen, 1983; Tang et ah, 1983; Shan,
1999). After examining numerous specimens of
Stachyurus and conducting a field expedition to
Sichuan and Yunnan provinces of China, we found
a continuous variation in the leaf blades of S. yun¬
nanensis. For example, in this species, leaves are
leathery or thickly papery, ovate-oblong, oblong-el¬
liptic, or ovate-lanceolate, with either regular or ir¬
regular serration at the margins. Even different

Novon 15: 500-501. Published on 21 September 2005.

Volume 15, Number 3
2005

Zhu & Zhang

Lectotypification in Stachyuraceae

501

leaves on (he same plant of S. yunnanensis may
have variable teetb. The characters of S. callosus
are within the range of that of S. yunnanensis.
Therefore, it is effectively impossible to distinguish
between S. callosus and S. yunnanensis based solely
on morphological characters.

Slachyurus callosus is distributed only in Malipo
and Pingbian Counties of southeastern Yunnan
(based on two specimens), at an altitude of 900—
I 100 m, falling just within the geographic distri¬
bution of S. yunnanensis. In addition, both species
grow in calcareous forests (Chen, 1981). On the
basis of these data, we believe that Stachyurus cal¬
losus should be treated as a synonym of ,S. yunna¬
nensis.

Acknowledgments. We are most grateful to the
curators of BM, E. k, KLIN, P, and PE for the loan
of or permission to examine specimens, and to Tang
Yancheng (PE), John McNeill (E). and Zlui Guan-
ghua (MO) for valuable information and suggestions
on the manuscript. This project is supported by a
grant from the National Natural Science Foundation
of China (NSFC No. 30130030).

I .iterature Cited

Chen, S. K. 1981. A study on the Stachyuraceae from
China. Acta Bot. Yunnan. 3(2): 125—137.

- . 1983. Stachyuraceae. In: Kunming Institute of

Botany, Chinese Academy ol Sciences, (editor). Flora
Yunnanica, Vol. 3: 339-348.

Franehet, A. 1898. Plantarum sinensiurn ecloge seeunda.
J. Bot. 12: 253-264.

Greuter, W.. J. McNeill. F. B. Barrie, II. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). fl. Nicolson, P. C. Silva,
.1. K. Skog, P Trehane, N. J. Turland & l). I.. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Kegnum Veg. 138.

Lauener, L. A. 1966. Catalogue ol the names published
by Hector Leveille. Notes Roy. Bot. Card. Edinburgh
27(1): 1-7.

Leveille, II. 1914—1915. Flore du Kouy-Tcheou. Le Mans.

Li, II. L. 1943. The genus Stachyurus. Bull. Toney Bot.
Club 70(6): 615-628.

Shan. II. B. 1999. Stachyuraceae. In: Delectis f lorae Rei-
puhlieae Popularis Sinicae Agendae Academiae Sinicae
Edita, (editors). Flora Reipublicae Popularis Siniea,
Vol. 52(1): 81-96. Science Press, Beijing.

Tang, Y. C., Y. L. Cao, Y. Z. Xi & J. lie. 1983. Systematic
studies on Chinese Stachyuraceae (1) — Phytogeograph-
ical. cytological, palynologieal. Acta Phytotax. Sin.
21(3): 250.

Volume 15, Number 3, pp. 585—502 of NOV ON was published on 21 September 2005.

Volume 15
Number 4
2005

NOVON

INevv Taxa and New Combinations in Some Mexican and Central
American Melastomataeeae

Frank Almeda

Department of Botany, California Academy of Sciences, 875 Howard St., San Francisco,
California 94103-3098, ll.S.A. falmeda@calacademy.org

Abstract. Diagnoses, descriptions, and illustra¬
tions are presented for four new laxa of Miconia
(M. commutata from Costa Rica and Panama, M.
danielii and M. tonduzii var. hirtella from Costa
Rica, and M. dissita from Panama). Three new com¬
binations (Miconia chinantlana (Naudin) Almeda
for Staphidium chinantlanum Naudin; Miconia Inn-
dellii (Wurdack) Almeda for Clidemia lundellii
Wurdack; and Tibouchina tortuosa (Bonpland) Al¬
meda for Rhexia tortuosa Bonpland) are proposed.
Miconia tonduzii Cogniaux is neotypified and its
infraspecific taxonomy is revised to include three
varieties. Miconia sparrei Wurdack. which was de¬
scribed from Amazonian Ecuador and heretofore
known only from the holotype, is redescribed, il¬
lustrated, and reported from the Caribbean low¬
lands of Costa Rica and Irom Peru. Citations of
representative specimens, distributional and pho¬
nological notes, and comparisons with probable rel¬
atives are provided for each taxon.

Key words: Central America, Melastomataeeae,
Mexico, Miconia , Neotropics, Tibouchina.

This paper presents miscellaneous descriptions
of new taxa, new combinations, a neotypification,
and new distributional records for selected species
in the genera Miconia Ruiz & Pavon and Tibouch¬
ina Aublet. It represents what I hope will be one
of the last precursors to the accounts of the Melas¬
tomataeeae for Flora Mesoamericana and the Man¬
ual de Plautus de Costa Rica. Work on a family
like the Melastomataeeae, which has radiated ex¬
tensively in the wet tropics, is a challenging and
unending synthesis. The Holistic accounts men-

Novon 15: !

tinned above will be the first comprehensive ac¬
counts of the family for the region covered based
on extensive fieldwork by a specialist and collab¬
orators. I see these treatments as progress reports
that will provide the crucial basis for identifying
collections and as compendia that will stimulate
additional exploration and research to deepen our
understanding ol one of the more commonly en¬
countered families of vascular plants in the Neo¬
tropics.

Miconia chinantlana (Naudin) Almeda. comb,
nov. Basionym: Staphidium chinantlanum
Naudin, Ann. Sci. Nat. Bot. 3. 17: 318. 1851.
Clidemia chinantlana (Naudin) Iriana. I’rans.
Linn. Soc. London 28: 135. 1871. TYPE: Mex¬
ico. Oaxaca: Chinantla, Jocotepeque, 1000 m,
1840, II. G. Galeotti 2945 (holotype, P; iso¬
type. BR).

Miconia echinoidea Stand Icy & Steyennark, field Mas.
Nat. Hist., Bot. Ser. 23: 180. 1944. Syn. nov. TV PE:
Guatemala. Huehuetenango: vicinity ol Maxbal,
about 17 miles N of Barrillas, Sierra de los Cucliu-
matanes, ./. Steyennark 4883 1 (holotype. F).

Section Octomeris. Shrub 1.5—4 m tall, the up¬
permost terete cauline internodes, petioles, abaxial
leaf surfaces, inflorescence rachis, hypanthia. calyx
lobes, and exterior calyx teeth copiously covered
with yellowish spreading stalked-slellate hairs.
Leaves of a pair ± equal to unequal in size; peti¬
oles 0.6-4. 8 cm long; blades 9.7—20 X 4—9.5 cm,
ovate to lance-ovate, 5- to 7-plinerved with the in¬
nermost pair of primary veins diverging from the
midvein in opposite or subalternate fashion 0.5—1 .7

503-518. IM bushed on 12 December 2005.

504

Novon

cm above the blade base, apex long acuminate,
base obtuse to broadly rounded and commonly
oblique, the margin inconspicuously ci Mate- serru¬
late to nearly entire, the adaxial surface copiously
covered with smooth spreading apically curved
hairs 0.25—0.5 mm long. Inflorescence a terminal
paniculiform dichasium 4—9 cm long with ultimate
branchlets terminating in simple 3-flowered dicha-
sia, flowers 5-merous, essentially sessile at anthesis
with pedicels to 0.5—1 mm in fruit; bracts and brac-
teoles 1-2 X 0.25—0.4 mm. deciduous, subulate to
elliptic, adaxial surface glabrous, abaxial surface
beset with stalked-stellate hairs, the margin ciliate
and terminating in a smooth hair. Hypanthial in-
dumenl typically not concealing the actual surface
at anthesis; calyx tube 0.25 mm long, the calyx
lobes 2—3 mm long, and conspicuously exceeding
the calyx lobes in length, ovate-rounded to ovate-
oblong, the exterior calyx teeth stoutly subulate;
petals 3—5 X 1.75—2 mm, glabrous, oblong-obovate
and reflexed at anthesis; stamens 10, isomorphic;
filaments 2—3.5 mm long, glabrous; anthers 3—3.5
mm long, yellow, linear-oblong with a ventrally in¬
clined apical pore; connective thickened dorsally
but neither prolonged nor appendaged; ovary 5-loc-
ular, 2A inferior, apex of fruiting hypanthia truncate
and glabrous around the stylar scar but glandular-
puberulent in a wide ring around the style; style 8
mm long, straight, sparsely beset with minute stel¬
late and glandular hairs; stigma punctiform. Berry
6—7 X 6-7 mm when dry, purple-black when ma¬
ture; seeds 0.75—1 mm long, ovoid, the testa vague¬
ly rugulate.

Phenology, f lowering collections have been
made from May through July; fruiting from August
through November.

Distribution. Local and uncommon in rain for¬
ests and \ nne-odk- Liquidambar L. forests from
southeastern Mexico (Chiapas and Oaxaca) to west¬
ern Guatemala (Huehuetenango) at 1250-1900 m.

Miconia chinantlana is here assigned to section
Octomeris Triana based on its elongate subulate an¬
thers and regularly lobed calyx. Most recently col¬
lected material of this species has been identified
as Miconia echinoidea Standley & Steyermark, the
type of which is identical to the type of Staph idium
chinantlanum, the oldest available specific epithet
lor this species. Miconia chinantlana is very close
to and readily confused with Miconia globulifera
Naudin, which also occurs in southeastern Mexico
(Chiapas, Oaxaca, and Veracruz) and Guatemala.
These two species are similar in habit, foliar shape,
venation and indument, and in having 5-merous

flowers and a 5-locular ovary. Miconia globulifera
differs in having the (lowers aggregated in capitate
clusters of more than three flowers at the ends of
the inflorescence branchlets, the calyx teeth are
0.5—1 mm long and barely exceeding the calyx
lobes in length, the hypanthia are beset with simple
and apically bifid hairs 1—1.5 mm long that conceal
the actual surface at anthesis, and the style is es¬
sentially glabrous.

Representative specimens examined. MEXICO. Chia¬
pas: Mun. Cintalapa, SE of Cerro Haul on the border w /
Oaxaca, 16 km NW of Rizo de Oro along a logging road
to Colonia Figaroa, Breedlove 27619 (OS); Mun. Jitotol, 5
kin SE of Jitotol along road to Bochil, Breedlove 52655
(CAS); Mun. Yajaldn, camino la punta del Cerro Venlana,
Mdndez Ton 66 87 (CAS); Rancho Concepcion (Adelante
Cate Simojobel), Miranda 6694 (CAS). Oaxaca: Mun. To-
tontepec, Distr. Mixe, Rancho La Garrapata, 6 km al E de
Totontepec, Ramirez & Ramirez 404 (CAS); Mun. Toton-
tepec, Distr. Mixe, Totontepec. Rivera 1490 (CAS); Distr.
of Choapam, Cerro Lalana, Schultes & Reko B56 (CAS,
MEXU).

Miconia commutnla Almeda. sp. nov. T\ FE: Cos¬
ta Rica. Heredia: Finca La Selva. OTS Field
Station, Rfo Puerto Viejo, just E of jet. w/Rfo
Sarapiquf, along Q. La Selva. 2800 m S X 700
in E, 22 Feb. 1982 (fl), B. Hammel 1 1 237 (ho-
lotype, CAS; isotype, DUKE). Figure 1.

Section Miconia. Arbor 4—15 in; ramuli primum quad-
rangulati vel sulcato-quadrangulati demum teretes sicut
petioli inflorescentia venae primariae subtus hypanthiaque
indumento appresso aniorpho-squamato induti. Lamina
15—31 X 6.3—12.8 cm oblongo-elliptica 3- ad 5-nervata,
supra glabra. Panicula 10—28 cm multiffora; llores 5-meri,
cafycis tubus 0.5 nun altus, lobis interioribus 0.5 mm al-
t is, dentibus exterioribus subobsoletis vel obscurissimis;
fructum calyces decidui; stamina paullulo dimorphica gla-
bra; antherarum thecae 2.5 mm anguste oblongae poro
ventraliter inclinato, connectivo vix (0.5 mm) prolongato
dorsaliter vel antherarum thecae 2.25 mm anguste oblon¬
gae poro truncato, connectivo vix (0.25 nun) prolongato
dorsaliter ad basim bilobulato ornatis; ovarium 3-loculare
et 2A inferum apice glafiro.

Tree 4—15 m tall, the quadrate to quadrisulcate
uppermost cauline internodes, young petioles, in¬
florescence branches, pedicels, hypanthia, and ca¬
lyx teeth moderately to densely covered with an
appressed lepidote or bran-like indument. Leaves
of a pair equal to subequal; petioles (2.5— )4— 7.5
cm long, the blade 15—31 X 6.3—12.8 cm, 3- to 5-
nerved with an additional inconspicuous intramar¬
ginal pair, oblong-elliptic, the apex caudate-acu¬
minate, the base obtuse to rounded, the margin
entire, the adaxial surface glabrous, the abaxial
surface with a sparse deciduous lepidote or bran¬
like indument on the elevated primary veins but
essentially glabrous on the actual surface. Inflores-

Volume 15, Number 4
2005

Almeda

New Taxa in Melastomataceae

505

Figure 1. Miconia commutata Almeda. — A. Habit. — B. Representative leaf (abaxial surface). — C. Petal (adaxial
surface). — I). Antepetalous stamen (profile view). — E. Style and stigma. — F. Antesepalous stamen (ventral view). —
G. Mature berry. — H. Seeds. (A-F from the holotype Hummel 11237 ; G-H from Folsom 9355.)

cence a multiflowered panicle 10—28 cm long, the
sessile flowers 5-merous, the bracteoles apparently
very early deciduous and typically not seen. Hy-
panthium conspicuously 10-ribbed at maturity
when dry; calyx tube 0.5 mm long, the calyx lobes
0.5 X 0.75 nun, broadly triangular, and ultimately
deciduous on fruiting hypanthia to leave a truncate
circumscissile scar, the exterior calyx teeth trian¬
gular. 0.1 mm long and not exceeding the calyx
lobes but typically inconspicuous and appearing

obsolete; petals 2.5—3 X 1.5—1.75 mm, white but
drying yellow7, glabrous, oblong-obovate; anthers
somewhat dimorphic, white, linear oblong, the the¬
cae alternately 2.5 mm long with an emarginate
ventrally inclined apical pore and die connective
prolonged dorso-basally into a deflexed sheath-like
collar 0.5 mm long or thecae 2.25 mm long with a
± truncate pore and the connective prolonged dor¬
so-basally into a blunt tooth-like spur 0.25 mm long
and a bilobed ventro-basal appendage; style 5—7

506

Novon

mm long, glabrous; stigma clavate-truncate; ovary
3-locular, 2A inferior, the apex glabrous and round¬
ed. Berry 5—6 X 5—6 mm when dry, 10-ribbed,
turning from green through yellow, red, and ulti¬
mately purple-blaek at maturity; seeds 1—1,5 mm
long, ± triangular in outline with a raphe that is
expanded into a somewhat compressed elevated
border, the glossy testa with minute but distinct
verruculose angles on the convex face.

Phenology, f lowering collections have been
made in February and March; fruiting specimens
have been collected from February through May.

Distribution. Local and uncommon in lowland
rain forest and swamp forest (0—120 m elevation)
at the OTS Field Station along the Bio Puerto Viejo
on the Caribbean slope of Costa Rica and in the
vicinity of Changuinola on the Caribbean versant
in northwestern Panama.

Miconia commutata is assigned to section Mi-
con in based on its slightly attenuate anthers and
dorsally appendaged anther connective. It is char¬
acterized by its large entire leaf blades, long peti¬
oles, circumseissile calyx that falls away as a ring,
anther connectives that are modified dorso-basally
into a spur or sheath-like collar, 10-ribbed berries,
and more or less triangular seeds with a raphe that
forms a somewhat compressed elevated border. Mi¬
conia commutata is most similar to M. snuiragdina
Naudin in indument details and in the eircumseis-
sile nature of the1 calyx. The latter differs in having
narrower leal blades (2.7— 8.5 cm) that are some¬
what decurrent on the shorter (0.5—1 .9 cm) petioles,
petals that are minutely granulose- papillose (espe¬
cially on the adaxial surface), anther connectives
that are prolonged ventro-basally into a bilobed ap¬
pendage with each lobe appressed to the base of
each anther sac, and a capitate stigma. These two
species also exhibit distinctive distributional and
elevational differences. Miconia commutata ap¬
pears to be restricted to low elevations (0—120 m)
in Costa Rica and Panama, whereas M. smaragdina
occurs at higher elevations (450)850—1500 m from
southern Mexico disjunetly through Central Amer¬
ica to Colombia, Venezuela, and Ecuador (Almeda,
unpubl.).

Among South American species, M. commutata
is superficially like M. rivalis Wurdack of Ecuador,
which also has large leaf blades and a multiflow-
ered inflorescence. The leaf blades of M. rivalis,
however, are undulate-serrulate, the tardily decid¬
uous indument on the inflorescence and hypanthia
is appressed subamorphous-stellulate, the calyx
lobes are persistent, and the anther pores are uni-
formh ventrally inclined.

The flowers ol this species reportedly (fide Ham-
mel 1 1237) have a very strong cinnamon-like or
rose-cinnamon fragrance.

Etymology. The specific epithet is derived from
the Latin word commutata, meaning changing or
changed, in reference to the changing color of ber¬
ries from green to yellow, then red, and ultimately
purple-black at maturity.

Paratypes. COSTA RICA. Heredia: Finca La Selva,
OTS Field Station on the Rfo Puerto Viejo just F of jet.
w/Rfo Sarapiqm Central Trail, 2400 m line, Folsom 9467
(CAS, DUKE), Southwest Trail, 1400—1600 in line, Fol¬
som 9697 (CAS. DUKE), El Swampo Trail. Folsom 9767
(CAS. Dl KE). Central Trail, 2600 in line/V Boundary,
3000 in line, Folsom 98.55 (CAS, DUKE); OTS f ield Sta¬
tion, S Boundary, along (,). El Salto, Hammel 7940 (CAS,
DUKE); OTS Field Station, \X Boundary 1800 in S.
swampv area. Hammel 12566 (CAS, DUKE). Puntaren-
as: IVnfnsula de Osa, Bosque communal La Gloria, a or-
illas de Rfo Claro. Cordero 277 (tilt, INB. MO). PANAMA.
Bo. •as del I oro: a orillas del Changuinola y bosque al-
rededor de la eseuela de Corriente Grande, Correa el al,
4014 (CAS, I’M A).

Miconia danielii Almeda, sp. nov. TYPE: Costa
Rica. Cartago: 4—6 km N of Trinidad on S-
facing slopes of Vulcan Turrialba, 4 July 1977,
F. Almeda, R. /.. Wilbur <£■ 7. F. Daniel 2R76
(holotype, CAS; isotype, CR). Figure 2.

Section Cremanium. Frutex 1.5— 3.5 in; ratnuli primum
obtuse sulcato-quadrangulati demum terries sieut petioli
foliorum sublus venae prituariae et seeundariae hypanth-
iaque modice vel dense pi I is dendritieis el lepidotis iu-
duti. Lamina I 1.5—20 X 6—10.5 cm elliptieo-ovata. 5-ner-
vata. supra glabra. Panicula 8-15 cm multiflora, flores

5- meri, calycis tubus 0.25 mm altus, loliis interioribus
0.5—0.75 mm altis, dentibus exterioribus crass is appressis
inframarginalibus. Stamina isomorphica glabra, anthera-
rum tfieeae 1 — 1.5 mm oblongae latae 4-porosae, poro ven-
traliter inclinato; connectivum ad basim ventraliter paulo
(0.1 nun) bilobulato-prolongatum dorsaliter dente 0.5 mm
longo armatum. Ovarium 3-loculare et Vi inferum apice
glabro.

Shrub 1.5— 3.5 m tall, the rounded-quadrate to
quatlrisulcate upper internodes, young petioles, pri¬
mary and higher order veins on abaxial foliar sur¬
faces, and inflorescence branches moderately to
densely covered with lepidote and ± flattened den¬
dritic hairs. Leaves ol a pair essentially equal in
size; petioles 3.3— 4.5 cm long, blades 11.5—20 X

6— 10.5 cm, 5-nerved, elliptic-ovate, the apex acu¬
minate to caudate-acuminate, the base obtuse to
rounded, the margin conspicuously serrulate, the
adaxial surface essentially glabrous at maturity. In¬
florescence a multiflowered panicle 8—15 cm long,
the 5-merous flowers on pedicels 1 — 1.5 mm long,
the bracteoles 2.5— 4.5 X 0.75—1 mm, oblong to
elliptic-oblong, early-deciduous. Hypanthium spar-

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507

Figure 2. Miconia danielii Almeda. — A. Habit. — B. Representative leaf (abaxial surface). — C. Petal (adaxial sur¬
face). — 1). Stamens, ventral view (left) and profile view (right). — E. Style and stigma. — F. Berry. — <i. Seeds. (A. B.
F from Almeda et al. 288.3; C-F from Utley & Utley 2466; (', from the holotype Almeda el al. 2876.)

inglv scurfy -puberu lent at t lie base to essentially
glabrous throughout; calyx tube 0.25 mm long, the
calyx lobes 0.5—0.75 X 1 mm, broadly rounded;
the exterior calyx teeth bluntly conic, 0.25 mm
long, shorter than the calyx lobes; petals 1.5 X 1.5—

2 mm, white, glabrous, obovate to suborbieular; an¬
thers isomorphic, 1 — 1.5 mm long, white, obovoid-
oblong, 4-celled with 4 pores that are somewhat
ventrally inclined; connective thickened dorsj dly
and prolonged dorso-basally into a deflexed eau-

508

No von

ililorm appendage 0.5 mm long and also prolonged
ventro-basally at die base of each anther sae into
a deflt ^xed lobule 0.1 mm long; style 3—3.5 mm
long, glabrous; stigma peltate; ovary 3-loeular, Vi
inferior, the apex glabrous. Berry 3 X 4-5 mm
when dry; seeds 0.5 mm long, narrowly ovoid, the
testa vaguely granulate.

Phenology. Collected in flower from March
through May and July; in fruit in July and in De¬
cember.

Distribution. Endemic to Costa Rica, where it
is local and uncommon in cloud forests, forest mar¬
gins, stream banks, and montane pastures on the
Cordillera Central (Volean Barva and Volean Tur-
rialba), Valle de Reventazon, and the Cordillera de
Talamanca at 1300—2400 m.

Miconia danielii is here assigned to section Cre-
tnaniurn (1). Don) Naudin based on its obovoid an¬
thers that are 4-eelled and 4-pored. It is defined by
a consistent combination of vegetative and floral
features. It has an indument ol bran-like and ±
flattened dendritic hairs on upper internodes, pet¬
ioles, abaxial foliar veins, and inflorescence
branches; 5-nerved, elliptic-ovate leaf blades with
an acuminate to caudate-acuminate apex and con¬
spicuously serrulate margins; white anther thecae
that are 4-eelled and 4-pored; an anther connective
that is prolonged dorso-basally into a caudiform ap¬
pendage and prolonged ventrally at the base of
each anther sac into a deflexed lobule; a peltate
stigma; and a vaguely granulate seed coat.

Among species of Miconia sect. Cremanium (I).
Don) Naudin, M. danielii resembles M. coriacea
(Swartz) DC. and M. cremadena Gleason in various
details. The former differs from M. danielii in being
dioecious and in having a deciduous-furfuraceous
or matted furfuraceous indument, 2-celled and 2-
pored anthers, a distally fistulose style and capitate
stigma with a erateriform apex, and a smooth seed
coat. Miconia cremadena differs most notably from
M. danielii in its elliptic-oblong leaves with incon¬
spicuously crenulate to subentire margins, and a
glandular puberulence on the filaments and basally
prolonged anther connective.

Etymology. This species is named lor my col¬
league Thomas K. Daniel (b. 1954), co-collector of
the type, student of Neotropical Acanthaceae, and
field companion on several expeditions to Central
America.

Paratypes. COSTA RICA. Carlugo: t! km N of Trin¬
idad on S-fueing slopes of Volean Turrialba, Almeda et al.
2555 (CAS); Canton de Paraiso, I’. N. Tapantf, Valle de
Reventazon, RfoCrande de Orosf, sector la Represa, Mora

527 (CAS, CR. INB, MO); 1-2 km N of El Kmpalme,
along trail N of old Pan-American Highway, Utley & Utley
2455 (CAS, CR). Heredia: headwaters of Rfo Santo Do¬
mingo, ea. 3 km NE of San Rafael de Varablanea, N slope
ol Volean Barva, Grayum 7251 (CAS. CR. MO). S an Jose:
I .as Nubes, Standley 55475 (US); near Finca La Cima,
above Los Lotes, N of El Copey, Standley 42685 (US).

IVlicoiiiu dissila Almeda. sp. nov. TV PE: Panama.
Bocas del Toro: Edwin Fabrega Dam & Reserve
in Fortuna, along Continental Divide trail bor¬
dering Chiriquf, 08°45'04"N, 82°15'04"W,
1200-1300 m. 10 Jan. 1989 (fl), F Almeda. O’.
de Nercrs A G. McPherson 6246 (holotype.
CAS; isotypes, MO. PM A). Figure 3.

Section Cremanium. Frutex vel arbuscula 1.3—5 m;
ramuli obtuse quadrangulati sicut inflorescentia hypanth-
iaque pi I is asperis el pilis stellalis induti. Lamina 5.1-18
X 2.4— 9.8 cm elliptica vel elliptico-ovata 3- ad 5-nervata
vel 3- ad 5-plinervata, supra et subtus glabra. Panicula
4—14 cm multiflora, flores 5-meri, calycis tubus 0.2—0.25
mm altus, lobis interioribus 0.5 X 1 mm, dentibus exter-
ioribus acutis 0.25 mm lobos interiores aequantibus vel
panic brevioribus; stamina isomorphica glabra, anthera-
rum thecae 1.2 mm oblongae 2-porosae, poro ventraliter
inclinato; connectivum nec prolongatum nec appendicu-
latum; ovarium 3-loculare et Vi inferum apice glabro.

Shrub or small tree 1.3-5 m tall with somewhat
swollen uppermost cauline nodes, the rounded-
quadrate uppermost cauline internodes, inflores¬
cence branches, and hypanthia moderately covered
with scurfy asperous-headed and/or stellulate hairs.
Leaves of a pair subequal to somewhat unequal in
size; petioles 0.7-8 cm long, the blades 5.1—18 X
2.4— 9.8 cm, 3- to 5-nerved or if 3- to 5-plinerved
then the innermost pair of primary veins diverging
from the midvein 0.5— 1.1 cm above the blade base,
the blades elliptic to elliptic-ovate, the apex acu¬
minate, the base acute to obtuse or rounded, the
margin entire, glabrous on both surfaces. Inflores¬
cence a multiflowered panicle 4-14 cm long with
lateral branches diverging from the central axis at
essentially right angles; flowers 5-merous, subses-
sile or on pedicels to 0.5 mm long; the bracteoles
0.2 X 0.1 nun. triangular, inconspicuous and early-
deciduous. Calyx tube 0.2-0.25 mm long, the calyx
lobes 0.5 X 1 mm. rounded-triangular, the exterior
calyx teeth 0.25 mm long, triangular, ± equaling
the calyx lobes but barely or not protruding beyond
the calyx lobes; petals 1 — 1.5 X I mm. white, gla¬
brous abaxially but often warty-papillate adaxially
when dry, oblong-obovate to obovate; anthers 1.2
mm long, isomorphic, white, oblong, 4-eelled with
2 ventrally inclined pores; connective thickened
and ± elevated dorso-basally but neither prolonged
nor appendaged; style 4—6 mm long, glabrous; stig¬
ma capitate; ovary 3-loeular, Vi inferior, the apex

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New Taxa in Melastomataceae

509

Figure 3. Miconia dissita Almeda. — A. Habit. — B. Enlargement of cauline indument. — C. Petal (adaxial surface).
— D. Stamens, ventral view (left) and profile view (right). — E. Style and stigma. — F. Fruits. — G. Seeds. (A. B from
McPherson 1 20 1 3; C— E from Almeda et al. 6226 ; F from Antonio 4057; G from Croat 66632.)

510

Novon

glabrous. Berry 3.5—4 X 3.5—4 mm when dry. blue-
green to blue-white at maturity; seeds 0.5 nun long.
± ovoid, the testa murieate.

Phenology. Flowering specimens have been
collected from January through March; fruiting ma¬
terial has been collected from January through
April. June, July, and September.

Distribution. Locally common to uncommon in
cloud forest understory from Costa Rica’s Cordillera
Central (1*. IN. Braulio Carrillo) and Cordillera de
Talamanca (vicinity of Cerro Duan) disjunelly west
to Panama from the Edwin Fabrega Dam area in
Fortuna to Cerro Colorado at 800—1500 m.

Miconia dissita is here placed in section Cre-
manium because of its 4-celled anthers with two
apical pores. Its anther thcceae are more elongate
and less obovoid than many members of section
Cremanium, but its affinities are clearly with mem¬
bers of this section. I initially identified many spec¬
imens of M. dissita as M. tonduzii Cogniaux var.
latifolia Cogniaux (here reduced to the synonymy
of M. tonduzii var. tonduzii) because of its compar¬
atively wide leaf blades, 4-celled anthers, and cap¬
itate stigma. Closer inspection in the field and her¬
barium revealed a number of characters that cannot
be accommodated in the range of variability here
attributed to M. tonduzii. Miconia dissita can be
recognized by its somewhat swollen distal nodes
and lateral inflorescence branches that diverge
from the main axis at angles of 90° or more. It also
has a distinctive induinent on upper internodes, in¬
florescence' branches, and hypanthia that consists
of scurfy asperous-headed and/or stellulate hairs
and anther thecae' that are widest at the' middle with
ne» evielence of a septum protruding beyond the'
apex e>f the' anther thee-ae. The le'af blades e>f M.
dissita are consistently entire, the adaxial petal sur-
fae'es frequently appear warty-papillate (in part)
upon drying, anel the seeel coat is muriculate.

Among species e»f sectiem Cremanium , Miconia
dissita appe'ars to be* me>st closely relateel te> M. ton¬
duzii var. tonduzii. In the latter, the1 elistal imeles are*
neit notie'eably sweillen, the lateral inflorescence
branches eliverge' from the (central axis at angles e>f
abemt 45°. the' induinent is amorphous-furfuraceous
puberulent or lacking, anel the anther thecae are
widest elistally with an evielent se'plum slightly pro-
trueling beyonel the' perimeter of the* anther pore.
Other elifferences that se'l M. tonduzii var. tonduzii
apart from M. dissita are the* remotely anel obscure¬
ly denticulate to subentire leaf margins, completed
smooth pe'tal surfae'es, anel the* obscurely granulate
se'eel coal.

Etymology. The- specific epithet is derived from
the Latin wore!, dissitns, meaning lying apart, re-
meile'. or well-spaced, in referene'e to the spacing eif
lateral divergent branches on the inflorescence.

Paratypes. COSTA 8ICA. San Jose: Bajo Honelura,
Parque Nacional Braulio Carrillo, Davidse el id. 23199
(CAS. CB. MO). Cartago: ca. 2 kin K of Cachf along road
paralleling Bfo Naranjo toward Cerro Duan. Utley 3887
(CAS, IH KK). PANAMA. Hocas ele'l Tore*: along road
betw. Fortuna Dam & Chiriquf Cranele along gravel road
which eleparts main hwy. near Continental Diviefe (4.5 mi.
N of hrielge over Fortuna lake), (coat 08032 (CAS. MO);
forest near headwaters of Kfo Culebra e-a. 5 km ENE of
Cerro Pate' Macho. Hamniel 011 0 (CAS. MO, PM A); vie.
of Cerro Colorado mine above San Felix, along trails N of
road along Continental Divide. McPherson 12013 (CAS.
MO). Chiriqiii: Edwin Fabrega Dam cN Reserve in For¬
tuna along trail to Ufo Hornito above Los Planes. Almeda
et id. 0318 (CAS, MO. PM A); along road betw. Oualaca
(A fortuna Dam site at 10.1 mi. NW of Los Planes de
Hornito. Antonio 1037 (C AS. MO. PM A); Fortuna Dam
area along Quebrada Bonito to E of road. Churchill et al.
1700 (CAS, MO).

Miconia limdcllii (Wurdack) Almeda, comb. nov.
Basionym: Clidemia lundeUii Wurdack, Wrigh-
tia 5: 227. 1976. TV PE: Guatemala. Baja Ver-
apaz: Union Barrios, W of km 159. 13 Aug.
1975, C. /,. I Ain dell & E. Contreras 19626 (ho-
lotype, US; isotypes, F, LL).

Section Oetomeris. Shrub 1—2 m tall, the round¬
ed-quadrate upper internodes, petioles, inflores¬
cence axes, pedicels, hypanthia and calyx teeth
densely covered with rusty-brown straight and/or
curly tangled hairs 1-4 mm long underlain with a
very sparse scattering of minute glands, the peliolar
hairs often inconspicuously bifid apically. Leaves
of a pair ± equal to somewhat unequal in size;
petiole 1-4 cm long, blade 6—1 1.3 X 2.9— 5.8 cm,
elliptic-ovate, 5- to 7-pli nerved with the innermost
pair of primary veins diverging from the midvein
0.5— 1.5 cm above the blade base, apex short-acu¬
minate, base rounded to obtuse, the margin entire
to obscurely crenulate, copiously ciliate, the adax¬
ial surface moderately covered with somewhat
spreading smooth hairs 1—2 mm long, the abaxial
surface (including primary and higher order veins
and actual surface) moderately covered with
spreading rusty-brown smooth hairs ca. I mm long
and sparsely underlain with ± sessile glands. In¬
florescence a terminal paniculiform diehasium 2.5—
5 cm long; flowers 5-merous on pedicels 2—5 mm
long; the bracts and bracteoles 2—9 X 0.75—2 mm,
narrow ly oblong, tardily deciduous, ciliate. sparing¬
ly setose to glabrous on both surfaces. Hypanthium
with an induinent like the cauline internodes, but
the hairs prevailingly straight and not curly; calyx

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New Taxa in Melastomataceae

511

lobes 0.75 X 1 mm, broadly triangular, commonly
concealed by the linear-oblong exterior teedi that
are 1.5—2 mm long; petals 5— 6 X 3—3.5 mm, gla¬
brous, oblong-obovate; filaments 4—5 mm long, an¬
thers 2.7— 3.5 mm long, yellow, the apical pore cen¬
trally inclined; connective thickened dorso-basally
and prolonged below the thecae ca. 0.25 mm, oth¬
erwise exappendieulate; ovary 5-locular, %-inferior.
apex elevated into a depressed ± crateriform dome
that probably flattens out on maturing fruits. Mature
berries and seeds are unknown.

Phenology. This species is known from only the
type and one other collection; the August collection
is in flower, the February collection is in young fruit.

Distribution. Apparently endemic to Guatemala
(Baja Verapaz), where it occurs in montane ever¬
green forests. No elevational data are given on
known collections, but judging from the collection
localities this species appears to occur at elevations
of about 2000-2100 m.

The elongate anthers and regularly lobed calyx
of Miconia lundellii dictates placement in section
Octomeris Triana. Wurdaek describes the inflores¬
cence of this species as seemingly terminal. Ex¬
amination of the two known collections verifies his
observation. There is no indication on either spec¬
imen that lateral shoots eventually overtop the in¬
florescence so that it becomes pseudolateral like a
typical member of the genus Clidemia. In view' of
the apical dominance exhibited by the inflores¬
cence of this species, I see no justification lor main¬
taining it in Clidemia and here transfer it to Mi¬
conia. Fruiting material ol this rare species is
needed before its intrageneric affinities can be as¬
sessed with confidence. Miconia lundellii is readily
recognized by its 5-merous flowers, 5-locular ovary,
copious rusty brown indument of straight and/or
curly tangled hairs, petiolar hairs that are often in¬
conspicuously bifid apically, linear-oblong exterior
calyx teeth that conceal the broadly triangular calyx
lobes, and the unappendaged anther connectives.

Additional specimen examined. GUATEMALA. Baja
Verapaz: Union Barrios on Salama/Coban road, V. 3 km,
on top of hill, Lundell A' Contreras 18958 (CAS. I.L).

Miconia sparrei Wurdaek, Fhytologia 35: 245.
1977. TYPE: Ecuador. Morona-Santiago: Mi-
sidn Salesiana, Gualaquiza, 700—800 m. 3 Oct.
1967, B. Sparre 19290 (holotype, S; isotype,
US). Figure 4.

Section Tamonea. Shrub or small tree 4—8 m tall,
the rounded -quadrate uppermost cauline inter¬
nodes (densely), petioles (moderately), and inflores¬

cence branches (densely) setose with spreading ±
flattened barbellate Irichomes 2-9 nun long. Leaves
of a pair somewhat unequal in size; petioles 3—5.5
cm long; the blades oblong-elliptic to elliptic-ob-
ovate, 30—55.5 X 13—28 cm. 5-plinerved (often
with an additional inframarginal pair of veins ex¬
tending half to % up the length of the blade), the
innermost pair of primary veins diverging 1.5— 3.3
cm above the blade base, the apex short-acuminate,
the base obtuse, the margin denticulate, the abaxial
blade surface essentially glabrous, the adaxial sur¬
face sparsely to moderately covered with spreading
roughened or barbellate hairs on the primary and
secondary veins but glabrous on the venules and
actual surface. Inflorescence a multiflowered pani¬
cle 15—30 cm long with 5-merous sessile flowers
borne in glomerules at the ends of the branehlets;
the persistent narrowly triangular bracteoles 2—5 X
0.75—2 mm with stramineous crisped hairs at or
near the base. Hypanthium sparsely stellulate-pu-
bendent; calyx tube 0.5 mm long, the calyx lobes
0.25 X 1 mm. deltoid; the exterior calyx teeth small
thickened callosities ca. 0.5 mm long that essen¬
tially obscure the calyx lobes; petals 2.5-4 X I —
1.8 mm, white, distinctly but minutely papillose
abaxially and along the distal half adaxially; an¬
thers somewhat dimorphic, linear-oblong; the the¬
cae alternately 2 and 2.3— 2.4 mm long with a ven-
t rally inclined apical pore; the connective not
prolonged abaxially but enlarged at the base of
each anther sac and fringed with conspicuous
glands; style 6 mm long, glabrous; stigma ± trun¬
cate to barely expanded; ovary 3-locular, % inferior,
apex elevated into a low stellate-puberulenl collar.
Berry 4X5 mm when dry, reportedly yellow turn¬
ing black at maturity; seeds 1.5 mm long, ± tri¬
angular with an ungulate convex face, the testa
smooth and glossy.

Phenology. Flowering collections have been
made in June and September; fruiting collections
in February through June and in September.

Distribution. Bare and uncommon in primary
rainforest at 30—800 m. It is presently known in a
region extending from La Virgen to the OTS La
Selva Biological Station in the Caribbean lowlands
of Costa Rica, in Amazonian Ecuador, where it is
known only from the type locality, and from Peru.

Based on its elongate subulate anthers and api¬
cally retuse petals, Miconia sparrei is assigned to
section Tamonea Cogn. I here provide an expanded
description and illustration (Fig. 4) for this species,
which has heretofore been known only from lowland
Amazonian Ecuador. Its discovery in the Caribbean
lowlands of Costa Rica may seem surprising but it

512

Novon

Figure 4. Miconia sparrei Wurdaek.
Petal (udaxial surface). — F. Stamens.
2451.)

— A. Habit. — 1?. Enlargement of eauline indument. — C. Mature berries. — I).
ventral view (left) and profile view (right). — F. Seeds. (A-F from Zamora el al.

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Almeda

New Taxa in Melastomataceae

513

is not unusual in view of distribution patterns seen
elsewhere in the family. Wurdack (1980: 4) notes a
recurrent distributional pattern among Ecuadorian
Melastomataceae that includes Costa Hiea-Pana-
ma, western Colombia, and the lower eastern slopes
of the Andes.

Miconia sparrei is distinguished by its large leaf
blades, upper cauline internodes that are densely
covered with spreading, roughened or barbed ±
flattened trichomes, sessile flowers, anther connec¬
tives that are enlarged at the base and edged with
several large glands, and ± triangular seeds with a
smooth glossy testa and an angulate convex face.

Representative specimens examined. COSTA RICA.
Heredia: Finca La Selva, the OTS f ield Station, Kid
Puerto Viejo at Pasos Perdides entre los posies ,'5000 a
.'5200. Cluudn 613 (l)llkF), in forest along S edge of Var¬
gas property, Hummel 11224 (CAS. DUKE), primary for¬
est. SF section of new property, Hummel / 1659 (CAS,
1)1 KE); Est. Biol. La Selva, Sendero Rihereno a Rfo Peje.
Hummel 1 7439 ( CAS. DUKE): Canton de Sarapii pit. Llan-
ura de San Carlos, La Virgen, Finca La Tirimbina, Zamora
el <d. 2151 (CAS, CR. F, l\B. MO). PERI . Iluaiiuro:
Morawetz A' Wallnofer 13—22333 (US). Sail Martin:
Schunke-Vigo 6 949 (US).

Miconia tonduzii Cogniaux var. tonduzii Cog-
niaux, in A. DC. & C. DC., Monogr. Phan. 7:
1191. 1891. Miconia tonduzii var. furjuracea
Cogniaux, in A. DC. & C. DC., Monogr. Phan.
7: 1191. 1891. Syn. nov. TYPE: Costa Rica.
In sylvis ad Rancho Flores, alt. 2043 m, A.
Tondtiz 2115 (neotype, here designated, HR;
isotype. US).

Miconia tonduzii var. cuneata Cogniaux, in A. DC. & C.
DC., Monogr. Phan. 7: I 101. 1891. Syn. nov. TYPE:
Costa Rica. In sylvis ad Rancho f lores, all. 2500—
2700 in. II. F. Pittier 1913 (holotype, HR; isotype,
US).

Miconia. tonduzii var. latifolia Cogniaux, in A. I Hi. & C.
DC., Monogr. Phan. 7: 1 191. 1891. Syn. nov. TYPE:
Costa Rica. Ad Rancho Flores, mont. Barba, .4. Ton-
duz 2112 (holotype, RR: isotype, US).

Section Cremanium. Shrubs or small trees 2—6
m tall, the young branchlets, elevated primary leaf
veins beneath, and inflorescences sparsely furfu-
raceous-puberulent to glabrate. Leaves of a pair
equal to somewhat unequal in size; petioles 1-4.5
cm long. Leaf blades 7.5—16 X 3—5 cm, 3- to 5-
nerved. elliptic to elliptic-lanceolate, adaxially gla¬
brous, abaxially sparingly furfuraceous-puberulent
to glabrate, apex gradually acuminate, base acute
lo rounded, the margin remotely and obscurely den¬
ticulate to subentire. Panicle 5—8 cm long; flowers
5-merous on pedicels 1—2 mm long, the bracteoles
0.5— 1.5 mm long, subulate and early-deciduous.
Hypanthia obscurely and deciduously puncticulate-

lepidote, but often appearing glabrous; calyx tube
0.5 mm long, the oblong apically rounded lobes
0.5— 0.7 mm long, the bluntly tubereulate teetli 0.5
mm long, oppressed to but not exceeding the lobes;
petals 2—2.5 X 2-2.5 mm, obovate, glabrous; sta¬
mens isomorphic; anthers 1.5 mm long, euneate, 4-
eelled. white or white flushed with [link, the pore
widest distally, somewhat retuse and ventrally in¬
clined with the septum slightly protruding beyond
the perimeter of the anther; connective thickened
dorso-basally into a bluntly lobulate spur and pro¬
longed ventro-basally into a bilobulate appendage;
style 4—5 mm long, glabrous; stigma capitate; ovary
3-locular, 2h inferior, apex elevated into a lobulate
glabrous collar. Berry 4-5 X 4—5 mm, blue-green
then turning purple at maturity; seeds 0.5 mm long,
ovoid, obscurely granulate.

Phenology. Flowering specimens have been
collected from January through May; fruiting col¬
lections have been made in January and from
March through October.

Distribution. Locally common in cloud forests
and elfin forests from Nicaragua (Cerro Pataste, east
to the Cordillera Dariense and north to the Cordil¬
lera Isabella and Cerro Saslaya), south lo Costa
Rica, where il occurs from the Cordillera Central
south to the Cordillera de Talamanea at 1300—
3000 m.

Miconia tonduzii was assigned to section Cre¬
manium by Cogniaux (1891) based on its cunei¬
form. 4-eelled anthers. \\ ben lie described Miconia
tonduzii he neglected to cite any specimens under
the nominate variety. Following the description of
the species he provided diagnoses for six varieties
and also listed a type specimen for each. I here
recognize only two of I he six varieties circum¬
scribed by Cogniaux and describe a third one as
new. Because Cogniaux did not specifically name a
M. tonduzii var. tonduzii , 1 here neotypify flic nom¬
inate variety. In doing so. I have selected the type
of M. tonduzii var. furjuracea to become I he type of
the species. What has passed as M. tonduzii var.
furfuracea is the most commonly encountered va¬
riety in this species complex. The nominate variety
is characterized by its furfuraceous puberulence on
upper cauline internodes, inflorescence branchlets,
and primary veins on abaxial leaf surfaces.

Miconia tonduzii Cogniaux, a variable and puz¬
zling species complex of southern Central America,
is recognized by its 4-celled euneate, broad-pored
anthers with filaments that conspicuously exceed
the petals in length and capitate stigmas. Cog-
niaux’s six varieties were based largely on minor
differences in the size and dentition of leaf blades

514

Novon

and details ol the indument. All of his varieties,
except M. tonduzii var. oblongifolia Cogniaux,
whieh I am relegating to the synonymy ol Miconia
camea Cogniaux, are readily accommodated in
what I consider to he two infraspeeific laxa worthy
of recognition.

In addition to the three varieties treated here,
another seemingly distinct population of Miconia
tonduzii is known from Volcan Mombacho, Nica¬
ragua ( Neill 1396, CAS), and other mountaintops
in Costa Rica and Panama. Plants from these pop¬
ulations are glabrous and the obscurely serrulate
subentire leaf blades are somewhat shorter (4.5— 8.3
cm) than typical for the species. Additional study
is needed to properly evaluate the taxonomic status
of these and other related populations and to de¬
termine whether M. tonduzii s.l. should be merged
with the closely related M. cornea.

All populations of M. tonduzii s.l. that I have
examined have hermaphroditic flowers, but one
specimen ( Almeda & Nakai 3329. CAS) has some
stamens on some flowers that appear to be abortive
and non-polleniferous.

Representative specimens examined. NICARAGUA.
Atlunlicn Norte: near summit of Cerro Saslaya, Neill
3349 (CAS, MO). Jinolega: km 155 de la earr. entre Ma-
lagalpa y Jinolega. en las Pilas. Moreno oil) (CAS). Mad-
riz: Cerro Pataste, ea 20 km SO de Ciudad Somoto, Gri-
jalva 912 ( CAS, MO), \1alagalpa: Cerro Id Pieacho above
Motel Santa Maria de Osliuna, Nee 27634 (CAS, MO).
COSTA RICA. Alajiicla: ea. 15—20 km N of Sareln on
road to Rujos del loro Amarillo, Almeda et id. 7366 (CAS,
CR). Carlago: ea 11 km SL of I'd l .mpalme, Cordillera de
Palamanea, Almeda el al. 3111) (CAS). Heredia: SVY
Hanks ol Volerin Rarva above Sacramento, vie. of Laguna
Rarva, Almeda el al. 3275 (CAS, CR); road betw. San
Rafael »Y Rfo Las Vueltas, N of Cerro Chompipe, Stevens
14037 (CAS, MO). San Jose: Canton de Perez Zeledon,
Cordillera de Talamanea. Cainino a la Auxiliadora, 4 km
K de Villa Mills, Gamboa cV Picado 393 (CAS. INR).

Miconia tonduzii Cogniaux var. birlella Almeda,
var. nov. TYPE: Costa Riea. Cordillera de Tal¬
amanea, on right-hand turn onto a gravel road
oil the Interamerican Hwy. ea. I km beyond
La Georgina en route to San Isidro de El Gen¬
eral. 9700 ft. |2939 m], 9 Mar. 1981 (fl), F.
Almeda it- K. Nakai 4809 (holotype, CAS).
Eigure 5.

Section Cremanium. Frutex vel arbuseula 1—5 m; ram-
uli suleato-quadrangulati sieut petioli loliorum subtus ve¬
nae primariae et seeundariae sparsiuseule setosis pi 1 is
plerumque 0.5—3 mm longis laevibus et lepidotis induli.
Lamina 4.3—13 X I .f> — H cm elliptica v<d elliptico-ovata
5-plinervata, supra glabra vcl sparse setulosa pilis laevi¬
bus ea. 0.5—1 mm longis. Panieula 8—15 cm multiflora,
florcs 5-mcri; ealycis tubus 0.25 mm alius, lobis intcrior-
ibus 0.5 X I mm. dentilms exterioribus acutis 0.3 mm

lobos interiores brevioribus; stamina isomorphiea glabra,
antherarum thecae 1.5 mm oblongo-cuneatae 2-porosae,
poro ventraliter inclinato; connectivum ad has im ventral-
iter paulo bilobulato-prolongatum; ovarium 3-loeulare et
Vi inferum apice glabro.

Shrub or small tree 1—5 m tall, the rounded-
c]uadrate to quadrisuleate uppermost eauline inter¬
nodes and petioles sparsely covered with smooth
spreading hairs 0.5—3 mm long underlain with a
sparse brow n-lepidote indument. Leaves of a pair
equal to subequal in size; petioles 1—3 cm long;
blades 4.3—13 X 1.6— 5(— 8) cm, 5-plinerved, the
innermost pair of primary veins diverging from the
midvein 1 .5 — 3( — 9) mm above the blade base, the
blade elliptic to elliptic-ovate rarely varying to al¬
most obovate, the apex acuminate, the base obtuse
to almost rounded, the margin ciliate-serrulate, the
adaxial surface essentially glabrous or sometimes
with a few scattered smooth spreading hairs 0.5—1
mm long, the abaxial surface sparsely covered with
spreading smooth hairs 0.5—2 mm long on primary
and higher order veins and sparsely brown lepidote
on the actual surface. Inflorescence a multiflowered
panicle 8-15 cm long, the main branches and ped¬
icels sparsely brown-lepidote; flower 5-merous on
pedicels 1-2.5 mm long, the early deciduous brac-
teoles 0.5—0.75 mm long, linear-subulate and fili-
form. Calyx tube 0.25 mm long, the calyx lobes 0.5
X I mm, rounded-triangular, calyx teeth 0.3 mm
long, bluntly triangular, appressed to and shorter
than the calyx lobes; petals 1.5— 2.5 X 1.5—2 mm,
white, glabrous, obovate; anthers 1.5 mm long, iso¬
morphic, white, cuneate and widest apieally, 4-
eelled with two ventrally inclined pores and a sep¬
tum protruding somewhat beyond the perimeter of
the pores; connective thickened and somewhat di¬
lated dorso-basally and prolonged ventro-basally
into a bilobulate appendage; style 4—5 mm long,
glabrous; stigma capitate; ovary 3-locular, %-infe-
rior. the apex glabrous and ± rounded on mature
fruits. Berry 3.5—4 X 3.5—4 mm when dry, pale
blue-green at maturity; seeds 0.5 mm long, ovoid,
the testa granulate.

Phenology. Flowering from January through
May; fruiting from March through June.

Distribution. Endemic to Costa Rica where it is
locally common in cloud forests and montane pas¬
tures on the Cordillera Central (Vol. Rarva, Vol. Ira¬
zu, and Vol. Turrialba), Cerros de Escazu, and the
Cordillera de Talamanea at 2300— 3000 m.

This variety is readily distinguished from the
other infraspecific taxa of Miconia tonduzii by the
ciliate-serrulate leaf margins and the spreading
blackish indument of smooth trichomes on upper

Volume 15, Number 4
2005

Almeda

New Taxa in Melastomataceae

515

Figure 5. Miconia tondiizii Cogniaux var. hirlella Almeda. — A. Habit. — B. Representative leal (abaxial surface).

C. Petal (adaxial surface). — 1>. Stamen (% profile view). — E. Berry. — F. Seed. (A— R from the holotype Almeda &
Nakai 4809; F from Almeda & Nakai 4799.)

516

Novon

cauline intemodes and petioles. These spreading
trieliomes are also found on infloreseenee branches
ol some specimens and the abaxial surfaces of some
leaves. This is the only variety ol M. tonduzii that
is restricted to Costa Rica. Populations from Cerro
Baser differ in having modally larger leaf blades
(up to 13 cm long and 8 cm wide) that are ± ob-
ovate. In all other features this foliar variant is
identical to typical populations ol this variety.

Etymology. I lie varietal epithet is derived from
the Latin word hirtus , hairy or rough, in reference
to lht‘ spreading hairs on distal internodes and pet¬
ioles.

Paratypes. COSTA RICA. Gartugo: Voleun Irazu in
pasture along CR rte. #l() from Cartage to the crater. llar-
ringer et al. 3239 (CAS. F); Villa Mills area near Cerro
de la IMuerte, Haft et al. 2155 (CAS. MO); vie. La I’ieada
A finca Retiro on K slope ol Volean Turrialba, Wilbur
21676 (C \S. I ) l KF,). Gartago/Sau Jose border: 15 km
SSf, of Kl Km pa I me intersection on the Cordillera de Tal-
amanca. Almeila et al. 3375 (CAS); Cordillera de Tala-
manca, vie. of Ojo de Agua, Almeila & Nakai 4799 (CAS).
Heredia: Cantrin de Rarva. Parque \ac. Rraulio Carrillo,
Kstacidn Barva, Fernandez 146 6 (CAS. INB. MO); Volcdn
Rarva approaching La Laguna on Pacific side. Lamer 1177
(CAS. CR. NY). San .1 ose: Cerros de Fseazu, Cerro Baser
from ca. 2(K) m below the crest to the top, Utley & Utley
2285 (CAS, CR); ca. 5 km VY-SW of Aserrf along upper
slopes & summit of Cerro Baser, Almeila & Nakai 3663
(CAS).

M. tonduzii Cogniaux var. serrulata Cogniaux, in
A. DC. cK C. DC., Monogr. Phan. 7: 1191.
1891. 1 Y PL: Costa Rica, secus Rio Pods ad
mont. Iscazn. alt. 2100 m. II. F. Pit tier 2393
(holotype. BR; isotype, US).

Mieonia tonduzii var. parr if alia Cogniaux, in A. DC. & C.
DC.. Monogr. Phan. 7: I P>1. 1891. Syn. nov. TY PE:
Costa Rica, in sylvis ad Rancho Flores, alt. 2043 m.
A. Tonduz 2116 (holotype. BR).

Section Cremanium. Shrubs or small trees 2-5
(—9) m tall, juvenile leaves and vegetative buds
sparsely and deciduously furluraccous-puberulent
to glabrate. Leaves of a pair equal to somewhat un¬
equal in size; petioles 1—3 cm long; blades 6— 12 X
2-4 cm, 3-nerved or 3-plinerved, often with a de¬
pressed pair of intramarginal veins, elliptic, adax-
ially glabrous, ahaxially sparsely and inconspicu¬
ously lepidote, apex gradually acuminate to
caudate-acuminate, base acute, the margin con¬
spicuously serrulate. Panicle 4-10 cm long; (lowers
5(to 6)-merous on pedicels 0.25—0.5 mm long, the
deciduous elliptic bracteoles 2—3.5 mm long. 1 1 v -
panthia obscurely and deciduously puncticulate-
lepidote, but appearing glabrous; calyx tube 0.25
mm long, the oblong apically rounded lobes 0.5—

0. T ( — I ) mm long, the tuberculate teeth 0.5— 0.7 mm
long, appressed to and not exceeding the lobes; pet¬
als 1.5— 2.5 X 1.5—2 mm, obovate, glabrous; sta¬
mens isomorphic; anthers 1-1.5 mm long, 4-celled,
cuneate, white, the broad pore ventrally inclined;
connective thickened dorso-basally into a blunt
lobe or spur, but ventro-basally adnate to the thecae
to form a bilobulate appendage; style 5 mm long,
glabrous; stigma capitate; ovary 3-loeular, Vi infe¬
rior. apex shallowly fluted and sparingly glandular-
puberulent to glabrate. Berry 5-6 X 5—6 mm. blue-
gray or blue-violet al maturity; seeds 0.5 mm long,
ovoid, obscurely rugulate or granulate.

Phenology. Flowering collections have been
made from January through May and in September
and October; fruiting material has been collected
from February through May and in August and Sep¬
tember.

Distribution. Locally common in elfin forests and
cloud forests from Nicaragua at Montana (inspire
(Depto. Jinotega) and I s I a de Ometepe (Depto. Ri¬
vas) to Costa Rica's Cordillera de Tilaran, Cordil¬
lera Central, and Cerros de Fseazu at 400—2200 m.

This variety is distinguished by its conspicuously
serrulate leaf blades that are 3-nerved or 3-pliner¬
ved. Nicaraguan populations ( Pipoly 5955 and
6042, both at CAS) from Cerro La Pimienta (Ze-
laya) and Pipoly 5164 ((7 AS) from Cerro FI Hor-
miguero (Zelaya) superficially resemble this variety
in having elliptic leaf blades that are sparsely lep¬
idote ahaxially, but the foliar margins are obscurely
serrulate and the anthers on the meager material
available for study appear to be 2-celled. Study of
the few flowers on Pipoly 6042 are also suggestive
ol dioecism. II examination ol better material con¬
firms this observation, these puzzling collections
should also be compared with M. glaberrima
(Schlechtendal) Naudin of southern Mexico and
northern Central America.

Representative specimens examined. NICARAGUA.
Jinotega: Montana Cuspire; Moreno 8028 (CAS). Itivas:
ladera N del Vulcan Concepcion. Isla de Ometepe. Mar¬
tinez et al. 1522 (CAS, MEXU); Isla de Ometepe, Lago de
Nicaragua, N slope of Volcdn Maderas, Neill A- Vineelli
3304 (CAS). COSTA RICA. Alajuela: Palmira, Smith
NY 52 7 (CAS, NY): Monteverde, road from Cerro Plano to
Cerro Amigos. Penneys 1490 (CAS, FLAS). Heredia:
headwaters of Rio Santo Domingo, ca. 3 km F of San
Rafael de Vara Blanca, N slope of Volcdn Rarva, Grayum
et al. 1 1 50 (CAS. CR. MO); Reserva Cerro Chompipe,
l manii & Chacdn 546 (CAS, CR); ca. 2 km SF of jet. CR
12(1 & CR 9 on CR 9, ca. 3 km south of Varablanea. File \
<£■ l tley 4392 (CAS. Bl KF). Punlareiias: Cordillera de
Tilaran. Monteverde Cloud Forest Res., Almeila & Daniel
4864 (CAS, CR). San Jose: 6 km NF of Caseajal, Almeila
<V Nakai 4864- (CAS, CR); ca. 5 km WSW of Aserrf along

Volume 15, Number 4
2005

Almeda

New Taxa in Melastomataceae

517

upper slopes and summit of Cerro Daser, Almeda & Nakai

3678 (CAS, CR).

l iheeue liina tortuosa (Bonpland) Almeda, comb,
nov. Basionym: Rhexia tortuosa Bonpland,
Monogr. Melast. 2: 17, pi. 7. 1806-1808. Os-
beckia tortuosa (Bonpland) Sprengel, Syst. Veg.
2: 312. 1825. Chaetogastra tortuosa (Bonp¬
land) DC.. Prodr. 3: 132. 1828. Oreocosmus
tortuosus (Bonpland) Naudin, Ann. Sci. Nat.
Bot. 3, 13: 38. 1850. TY PE: Mexico. Guerrero:
Taxco (as Tasco), A. Bonpland s.n. (holotype,
P; isotype, P |2|).

Pleroma mexicanum DC., Prodr. 3: 152. 1828. TYPE:
M, ;xico. without exact locality, M. Sessd & ./. Alocino
s.n. (holotype, OXK, photos, CAS, F, US).

Suffruteseent herb or slender shrub 0.5— 1.5 m
tall; distal branches quadrate, rounded with age,
the internodes and petioles moderately covered
with appressed simple trichomes 1-4 mm long.
I, eaves of a pair somewhat unequal in size, petioles
2-14 mm long; blades 2.9-9 X 1—3.4 cm, elliptic
to elliptic-lanceolate, apex acuminate, base acute,
margin inconspicuously ciliate-serrulate, 3- to 5-
nerved, adaxially moderately strigose, abaxially
moderately covered with appressed indument of
smooth trichomes on and between the primary
veins. Inflorescence a simple or once-compound
cyme but commonly reduced to paired or solitary
flowers on pedicels 2-4 mm long; braeteoles 3—6
(-10) X 0.5—3 mm. narrowly elliptic to oblong,
sparingly strigose on both surfaces, entire, ciliate,
and early deciduous. Calyx lobes 5—8 X 0.75—1
mm. ciliate, sparingly beset with a few simple tri¬
chomes on both surfaces; petals 10—15 X 8—11
mm, lavender, the margin ciliate, the cilia often
gland-tipped; large stamens, with filaments 5.5-6
mm long, anthers 3.5—6 mm long, subulate, red¬
dish-purple; connective prolonged 1.5— 3.5 mm be¬
low anther sacs, expanded dorsally near filament
insertion into a minute lobule and ventrally into a
yellow bilobed appendage 0.5—1 mm long; small
stamens, with filaments 5—7 mm long, anthers 2—
5.5 mm long, linear-oblong, yellow, connective pro¬
longed 0.25—0.5 mm below anther sacs, the yellow
shallowly bilobed ventral appendage 0.25—0.5 mm
long: style 8—9 mm long. Fruiting hypanthia 6—7 X
4—5 mm, subcylindric to suburceolate, moderately
covered with appressed to antrorsely spreading
smooth hairs 0.5-2 mm long; seeds 0.25-0.5 mm
long, coehleate, the testa tuberenlate.

Phenology. Flowering specimens have been
collected from January through April and from July
through December; fruiting collections have been

made from January through April, July, and from
August through October.

Distribution. Locally common on slopes with
mesophytic montane forests with Fraxinus L.. Firms
L., and Quercus L. in southern Mexico from Mi-
choacan, Queretaro, and Hidalgo disjunctly south
to Chiapas at 950—1950 m.

The name Tibouchina mexicana Cogniaux has
long been attributed to this species (Cogniaux,
1891; Standley, 1924). Because it is based on the
same type as T. tortuosa, it is illegitimate and must
be rejected since it was nomenclaturally superflu¬
ous when published. There has been no previous
attempt to identify the earliest epithet for this spe¬
cies. Cogniaux did not give a date of publication
for Rhexia tortuosa in his familial monograph and
Standley erroneously lists the publication date as
1823, the same year that D. Don described this
entity as Melastorna mexicanum, a later homonym
that is also illegitimate. According to Slafleu and
Cowan (1979: 369), Rhexia tortuosa was published
sometime between 1806 and 1808, which gives it
undisputed priority. I have examined the types of
R. tortuosa at P and these together with the plate
that accompanies the protologue leave no doubt
that what have passed as Pleroma mexicanum and
Tibouchina mexicana are clearly identical.

Tibouchina tortuosa and T. rufipilis (Schlechten-
dal) Cogniaux form a closely related species pair
characterized by narrowly oblanceolate or linear-
spatulate calyx lobes and dimorphic stamens. Typ¬
ical T. tortuosa has a sparse covering of appressed
trichomes on upper eauline internodes, whereas T.
rufipilis usually has a denser cover of spreading
internodal trichomes. Both species appear to be re¬
stricted to southern Mexico, but judging from avail¬
able collections, the latter is less common and
seems to have a range that is more eastern and
centered in the states of Queretaro, Hidalgo, and
Veracruz (Almeda, 1993). Posture and density of
eauline pubescence is known to be* an unreliable
taxonomic character among specie's cemiplexes
elsewhere in the family. I have refrained from com¬
bining these twee species, however, until the varia¬
tion can be evaluateel as part of a ceemprehensive
revisionary study. II these two spee ies prove tee be
conspecific, T. tortuosa is the epithet that has pri¬
ority.

Representative specimens examined. MEXICO. Chia¬
pas: man. Tenejapa in I lie paraje Oshe Wits, Breedlove
77644 (l)S, ENCB). Hidalgo: mun. Meelango. (> km S\\
of Molango, Breedlove & Almeda 59416 (CAS); along Hwy.
105 betw. Huejutla & Molango. 1 1.5 mi. N of Ajaeaya S.
Cristobal & 12.6 mi. X of jet. road to Olongo. Daniel 395
(CAS, MICH). Mexico: mun. Tejupilco, I km al V) de

518

Novon

Nanehilitla, Rzedowski 22104 (CAS, ENCR). Michoufuu:
entre Las Trineheras y El Puerto tie Laureles, aprox. 21
kin al SVV de Zilacuai'o, Soto et <il. 2058 (CAS, MEXU).
Morelos: by streams near Cuernavaca, Pringle 8021
(CAS, MEXU). Puebla: mun. Necaxa, along lake shore
al Nuevo Necaxa, Breedlove & Almeda 50592 (CAS).
Quereturo: mun. Lamia, 2 km al Noroeste ile Neblinas,
Rubio 1 175 (C AS. I EM); mun. Jalpan, 4-5 km al Poniente
tie La Paratla, Servt'n 224 (CAS, IER).

Acknowledgments. Field studies were support¬
ed, in part, by U.S. National Science Foundation
Grants DEB 76-8.4040, DEB 78-25620, BSR 86-
14800 (Flora Mesoamericana), and I lie G. Eindsay
Eield Beseareli Fund of the California Academy of
Sciences. Logistical support was provided by the
Universidad Autdnonia de Mexico. Museo Nacional
de Costa Bica, Instituto Nacional de Biodiversidad,
Costa Bica, Smithsonian Tropical Research Insti¬
tute, and the University ol Panama. The assistance
of several colleagues has advanced my research on
Mesoamerican Melastomataceae in many ways. I
am especially grateful to the following: D. E. Breed¬
love, M. Correa, T. Daniel, G. Davidse, G. de Ne-
vers. M. Grayum, B. Hammel, R. Kriebel, G. Mc-

Pherson, J. Rzedowski, I). Stevens, and N. Zamora.
Ihe illustrations were prepared by Meg Stalcup
d igs. I. 2, 5) and liana Pazdfrkova (Figs. 3, 4). I
also thank the curators and staffs of the following
herbaria for loans, gifts of specimens, or assistance
during study visits: BM, BB. C, CAS. CR, DS,
DUKE. ENCB, F. FLAS, G, IEB, INB. K. EE, MA.
MEXU. MICH, MO. NY. OXE. P. PM A. S, SCZ.
TEX, and US.

Literature Cited

Almeda, F. 1993. Melastomataceae. In: J. Rzedowski &
C. Calderon de Rzedowski (editors). Flora del Rajhi y
de regiones adyacentes 1(1: 1—38. Instituto de Ecologfa
A.C., Centro Regional del Rajhi. Patzcuaro, Miehoaedn,
Mexico.

Cogniaux, A. 1891. Melastomac^es. In: A. & C. de Can¬
dolle (editors), Monographiae phanerogamarum 7: 1—
1251). G. Masson, Paris.

Stafleu, F. A. & R. S. Cowan. 1979. Taxonomic literature,
Vol. II: H-Le. Regnum Veg. 98: 1-991.

Standley, I’. C. 1924. Melastomaeeae. In Trees and shrubs
of Mexico. Contr. U.S. Natl. Herb. 23: 1046-1074.
Wurdaek, J. J. 1980. Melastomataceae. In: C. Marling &
R. Sparre (editors). Flora of Ecuador 13: 1—403. Uni¬
versity of Goteborg; Riksmuseum, Stockholm.

Nomenclatural Notes on Eurasian Arabis (Brassieaceae)

Ihsan A. Al-Shehbaz

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, LJ.S.A.

ihsan.al-shelil)az@mobot.org

ABSTRACT: Two new generic names {Catolobus
and Pseiidoturritis ) and four new combinations
( Boechera falcata, C. pendula, I '■ turrita, and
Rhammatophyllum fruticulosum) are proposed.
Morphological characters supporting tin1 assign¬
ment of seven Eurasian species ol Arabis to Boech¬
era, Catolobus, Fourraea, Pseudoturrit is, Rham-
rnatophylluni, and Turritis are discussed in
connection with molecular or cytological data. A
key to these Eurasian segregates ol Arabis is pre¬
sented.

Key words: Ar<d)is , Boechera, Brassieaceae, Ca¬
tolobus, Fourraea, Pseiidoturritis, Rhammatophyl-
lum, Turrita. Turritis.

Until recently, Arabis I., was considered to in¬
clude as many as I BO species distributed in the
temperate areas of the northern hemisphere (Al-
Shehbaz. 1988). No comprehensive account of the
entire genus exists, though regional Holistic and
monographic works are numerous (see under Lit¬
erature Cited). Schulz (1936) synthesized earlier
works and presented a detailed classification of Ar¬
abis that included 16 sections, of which 10 were
either monotypic or ditypie. Schulz s sectional clas¬
sification was highly artificial because it included
species now assigned to Arabidopsis Heynhold,
Boechera A. Love & I). Love, Catolobus (C. A. Mey¬
er) Al-Shehbaz, Crucihimalaya Al-Shehbaz et ah,
Draba E.. Fourraea Greuter & Burdet, Pseudotur-
ritis Al-Shehbaz, Rhammatophyllum 0. E. Schulz,
Sibara Greene, and Streptanthus Nuttall.

Previous botanists who worked on Arabis delim¬
ited the genus primarily on having linear, latisep-
tate fruits (flattened parallel to the septum),
branched trichomes, and aecumbent cotyledons.
However, this character combination evolved in¬
dependently more than 25 tunes in the Brassica-
eeae (Al-Shehbaz, 2003). Molecular data (Koch,
2003; Koch et ah, 1999, 2000, 2001; kropf, 2002;
O'Kane & Al-Shehbaz, 2003; Price et ah, 2001;
Beilstein, pers. comm.) and morphological studies
(Al-Shehbaz, 2003) have c learly demonstrated that
all recent North American, European, and Asian
authors (see below) have recognized Arabis as a
polyphyletic genus consisting of many unrelated

Noyon 15:

taxa. For example, the delimitation of Arabis by
North American authors (e.g., Hopkins, 1937; Rol¬
lins, 1941, 1993; Mulligan, 1996) included species
presently placed in Arabidopsis, Boechera, Pennel-
lia Nieuwland, Streptanthus. and Turritis E. By con¬
trast. Arabis sensu European authors (e.g.. Jones
Akeroyd, 1993; Talavera, 199.3; Jalas & Suominen.
1994; Tan. 2002) inc luded species of Arabidopsis,
Diplotaxis DC., Draba. Fourraea, Turritis. and
Pseiidoturritis, whereas that of Asian authors (e.g..
Busch, 1939; Zhou et ah. 2001) included species
of Arabidopsis, Boechera. Catolobus, Crucihima¬
laya, Parryodes Jafri, Pseiidoturritis. Rhammato-
phyllum, Transberingia Al-Shehbaz & 0 Kane, and
Turritis.

Arabis s. sir. consists of species (all with x — 8)
that have while to lavender or rarely purplish flow¬
ers, ebracteate racemes, erect to ascending or di¬
varicate (never reflexed or secund) fruits, auriculate
or rarely short-pet iolate cauline leaves, and basal
and lower cauline leaves with stalked stellate tri-
chomes sometimes mixed with fewer, simple or
forked trichomes (see key).

The present paper deals with the generic place¬
ment of seven Eurasian species previously assigned
to Arabis (4. brassica, A. fruticulosa, A. ghtbra, A.
laxa, A. pendula, A. turezaninowii, A. turrita: sea1
below for synonymy and author attribution) and
proposes generic placements based on molecular
and cytological data combined with a critical eval¬
uation of morphology.

Boechera

As delimited by Al-Shehbaz (2003). Boechera is
a well-defined North American genus of more than
65 species (present estimate approaches 100) all
with a base chromosome number of x — 7, fruits
that are falcate, secund. pendent, or strongly re¬
flexed, and trichomes that are dendritic, irregularly
forked, sessile, malpighiaceous, simple or totally
lacking. All Boechera species were previously
placed in Arabis (Rollins, 1993; Mulligan. 1996),
but the latter genus has a base chromosome number
of x = 8, fruits that are straight, ascending to di¬
varicate but never secund, and trichomes that are

519-524. Published on 12 December 2005.

520

Novon

primarily stalked and stellate. Species presently as¬
signed to Arabis that do not have this character
combination are anomalous in the genus and should
be transferred to other genera. In fact. Arabis s. sir.
is most closely related to Draba, whereas Boechera
belongs to an unrelated North American lineage
(including Anelsonia J. F. Macbride & Payson, Cus-
ickiella Hollins. Phoenicaulis Nuttall. Polyctenium
Greene) that diverged from Arabis s. sir. about 19-
25 million years ago (Koch et al., 2001: Mitchell-
Olds et ah, 2005). Except for the following species,
the remainder of Boechera is restricted to North
America.

Boechera falcata (Turczaninow) Al-Shehbaz,
comb. nov. Basionym: Turritis falcata Turczan¬
inow, Bull. Soc. Imp. Naturalistes Moseou 13:
63. 1 010. Arabis Jalcata (Turczaninow) Ber-
kutenko, Bot. /burn. (Moscow & Leningrad)
61: 1597. 1976, non A. falcata Michaux, Fh
Bor. Amer. 2: 31. 1803, nec A. falcata A.
Bichard, Tent. Fh Abyss. 1:17. 1847. A. tur-
czaninowii Ledebour, Fh Rossiea 1: 123.

1811. I 't PE: Russia. Kamchatka: Ochoten-
sem, between Czernoliesk and Allach-Jun,
1835, Ilia Kuznetzojff s.n. (holotype. LE).

Boechera falcata, a species restricted to the Rus¬
sian Far Fast and adjacent Siberia, was correctly
recognized by Yurtsev & Zhukova (1972), Berku-
tenko (1988), and Doronkin (1994) as Arabis tur¬
ezaninowii. Lcdehour (1841) proposed A. turezani-
nowii as a new name to avoid creating a later
homonym by the transfer of Turritis falcata to Ar¬
abis (see synonymy above). Chromosome counts for
B. falcata. all agreeing on 2 n = 14. were listed as
A. falcata by Berkutenko and Gurzenkov (1976)
and as A. turezaninowii Ledebour bv Yurtsev and
Zhukova (1972) and Zhukova and Petrovsky
(1984). flit' species has falcate ly pendent fruits and
indumentum ol primarily dendritic trichomes.
Therefore, both cytological and morphological data
clearly support its placement in Boechera, a dis¬
position representing the first report of the genus
from Asia. It is interesting to note that Schulz
(1936) placed B. falcata (as A. turezaninowii) and
the North American B, holboellii (Hornemann) A.
Love & I). I .five (as A. holboellii Hornemann) in the
ditypic section Arabis sect. Campylocarpus C. A.
Meyer, but such a very close relationship was not
acknowledged by subsequent authors.

Catolohi s

I his name was originally proposed by Meyer (in
Ledebour, 1831. p. 20) as a monotypic section of

Arabis and was maintained as such by Schulz
(1936). Subsequent workers, however, did not rec¬
ognize this section. Meyer (in Ledebour. 1831) dis¬
tinguished section Catolobus and its type. A. pen-
dula I... from section Arabis (as sect. Euarabis) by
having sepals equal and non-saccate instead of un¬
equal and saccate, fruits pendulous instead of erect
or ascending, absence instead of presence of me¬
dian nectar glands, and funieles adnate to septum
instead of free from it. A closer examination of Ar-
alns s. str. worldwide shows that the differences in
fruit orientation and the adnation of funieles to sep¬
tum are reliable and consistent to distinguish Ar¬
abis s. str. and segregate genera, but those of the
nectar glands and sepals are not. f urthermore, the
abaxial leaf surface of 4. pendula has sessile, ap-
pressed, 3- to 6-rayed stellate trichomes with one
ray perpendicular to the surface. To my knowledge,
this type of trichomes is not found in any species
previously placed in Arabis. The indumentum in
Arabis s. str. consists of stalked, stellate trichomes
rarely mixed with forked or simple ones.

Molecular data (O'Kane & Al-Shehbaz, 2003)
show that Arabis s. sir. is remotely related to A.
pendula, a species nested within a well-supported
clade that includes Capsella bursa-past oris (L.) Me-
dikus, Neslia paniculata (L.) Desvaux, Camelitui
microcarpa Andrzejowki ex DC., and Pseudoarabi-
dopsis toxophylla (Bieberstein) Al-Shehbaz, O'Kane
& R- A. Price. Furthermore, chromosome counts
(Berkutenko & Gurzenkov, 1976) show that A. pen-
dula is a triploid with 2 n — 21. All species of
Arabis s. str. are diploid (or rarely tetraploid) based
on x = H (Al-Shehbaz, 2003). Berkutenko et al.
(1984) also reported 2 n = 16 and 2 n = 21 for the
species from the same population, but such dis¬
crepancy in chromosome number needs confirma¬
tion. and further cytological studies on A. pendula
are certainly needed before a meaningful conclu¬
sion is reached. I herefore, molecular and morpho¬
logical data support the removal of A. pendula to a
genus ol its own, and Arabis sect. Catolobus is el¬
evated herein to the generic rank to accommodate
that disposition.

Catolobus (C. A. Meyer) Al-Shehbaz, stal. et
comb. nov. Basionym: Arabis sect. Catolobus
C. A. Meyer, in Ledebour, FI. Altaica 3: 20.
1831. 'TV PE: Catolobus pendula (L.) Al-Sheh¬
baz.

Herbs biennial; trichomes simple and setose,
mixed with smaller, sessile, appressed. stellate, of¬
ten 3- to 6-rayed ones usually with 1 ray perpen¬
dicular to the surface; stems erect to ascending.

Volume 15, Number 4
2005

Al-Shehbaz
Eurasian Arabis

521

often much branc hed above. Basal leaves petiolate,
often withered by anthesis; cauline leaves sessile,
upper ones aurieulate, entire or serrulate to den¬
tate. Racemes ebracteate, often in panicles. elon¬
gated considerably and sometimes second in fruit;
fruiting pedicels slender, ascending to reflexed. Se¬
pals oblong, deciduous, erect, equal; petals white
or rarely pink, oblong to oblanceolate. not differ¬
entiated into blade and claw; stamens 6, slightly
tetradynamous, unappendaged; anthers oblong;
nectar glands confluent, subtending or surrounding
bases of all stamens, rarely median nectaries ab¬
sent; ovules 70 to I 10 per ovary. Fruit dehiscent,
linear, strongly latiseptate, straight, widely spread¬
ing to reflexed; valves often with a distinct midvein,
glabrous or rarely sparsely pubescent, smooth; gy-
nophore obsolete; replum rounded; septum com¬
plete; style obsolete or less than 1 mm long; stigma
capitate, entire; seeds uniseriate or biscriate,
winged all around or only distallv, oblong, ovate, or
suborbieular, flattened: funicles adnate at base to
septum; cotyledons aceumbent.

Catolobus pemlula (L.) Al-Shehbaz, comb. nov.

Basionym: Arabis pemlula L., Sp. PI. 2: 665.

1753. TYPE: Siberia, no collector name given

(lectotype designated by Boczantzeva (1997: p.
1 16). Herb. Finn. No. 842.13. FINN).

Zhou et al. (2001) indicated that the species (as
Arabis pemlula) is highly variable in the develop¬
ment of seed wing, arrangement of seeds per loeule,
and density of indumentum. Because of the dis¬
crepancy in its chromosome number (Berkutenko
et al.. 1984) and its tremendous morphological,
ecological, and altitudinal variation (Zhou et al.,
2001 ), the species is much in need ol detailed stud¬
ies throughout its range. Catolobus pemlula is wide¬
ly distributed in European Russia eastward into
central Asia. China, the Russian Far East, and Ja¬
pan (Zhou et al.. 2001 ).

Fot RR I/', t

Although this monotypic genus was established
two decades ago by Greuter and Raus (1984), it
has not yet received the recognition it deserves. It
differs from Arabis s. str. by being completely gla¬
brous and strongly glaucous and by having some¬
what fleshy leaves, long-petiolate basal leaves, sub-
terete fruits, and a chromosome number of 2 n =
14 (Burdet, 1967). All other species ol Arabis are
non-glaucous and otherwise have stellate tri-
chomes, short-petiolate to subsessile and nonfleshy
basal leaves, flattened fruits, and a base chromo¬
some number x = 8 (Al-Shehbaz, 2003). The type

species. Fourraea alpina (F.) Greuter & Burdet, is
known in Arabis as .4. brassica (Feers) Rauschert
and also, erroneously, under the illegitimate 4. pau-
ciflora (Grimm) Garcke (see Greuter et al.. 1986).
flie species is distributed in southern Europe from
the Iberian Peninsula eastward into Yugoslavia
(Jones & Akeroyd. 1993). Talavera and Yelayos
(1992) reduced Fourraea to a monotypic section of
Arabis, whereas Schulz (1936) placed thi> single
species (as A, paueijiora) in the invalid (nomen nu¬
dum) Arabis seel, lirassicoturritis O. E. Schulz. Mo¬
lecular data (Koch et al.. 1999, 2000; Koch. 2003),
as well as cytological and morphological data
above, strongly support the maintenance of Four¬
raea and amply demonstrate that its single species
is unrelated to Arabis s. str.

PSEl DOTl RRITIS

Arabis turrita F.. another anomalous element
widely recognized in Arabis, is distributed in Eu¬
rope (Jones & Akeroyd, 1993: Jalas & Suominen.
1994), Turkey, Caucasus, and adjacent Russia
(Cullen. 1965), and Algeria, Israel. Febanon. and
Syria (Greuter et al., 1986). It is readily distin¬
guished from all Eurasian and North American Ar-
abis by its yellow flowers, fully bracteate racemes
(rarely bracteate only along lower half), and seeund.
strongly falcate, deflexed fruits (8—) 10— 15 cm long
and with prominently thickened margins. By con¬
trast, Arabis has white or pink to purple flowers,
ebracteate racemes, and non-secund. straight, erect
to divaricate fruits l-4(— 5) cm long and with tm-
thickened margins.

Molecular data (Koch et al.. 1999. 2000; Koch,
2003; Kropf, 2002) demonstrate that Arabis turrita
forms a clade well separated from that including
Draba, Aubrieta Adanson, and Arabis s. str. I hose
findings, along with the significant morphological
differences above and those used by Al-Shehbaz
(2003) to delimit Arabis s. str.. strongly support the
placement of A. turrita in a genus of its own.

Wallroth (1822) [imposed the generic name Tur¬
rita to include two species originally described by
Finnaeus (1753) in Arabis, A. pemlula and A. tur¬
rita. He renamed the latter species as T. major
Wallroth, a name that should be used il Turrita
were to be maintained. However, this generic name
has recently been rejected as a later homonym ol
Turritis (see Brummilt, 2005). Although Bubani
(1901) expanded Turrita to include 12 additional
species from Arabis , his broader delimitation was
not followed by subsequent botanists, and at best
Turrita has since been recognized as a monotypic
section of Arabis (Schulz. 1936; Talavera cY Yelay-

522

No von

os, 1992). Based on the recommendation by the
Committee on Spermatophytes (Brninmilt, 2005:
p[). 535—536), a new monotypie genus Pseudotur-
ritis is proposed to accommodate A. turrita.

Psiiedoturritis Al-Shehbaz. gen. nov. TV PE: P.
lurrild (L.) Al-Shehbaz.

Herba perennis; pili stipitati. stellati; folia caulina ses-
silia. eordato-aurieulata vel amplexicaula; raeemi bractea-
li. valde elongati, petala luteola, spathulata; stamina 6,
tetradynama; ovula 50 ad 80; fructus lineares. seeundi,
eompressi, laleati. (8-)10— 10 cm longi; septum comple¬
tion; semina uniseriata, oblonga, alata, compressa; coty-
ledones accumbentes.

Short-lived perennial herbs, rarely biennial, pu¬
bescent with stalked stellate trichomes mixed with
fewer forked ones; stems erect, simple or few
branched above. Basal leaves rosulate, petiolate,
dentate, often withered by anthesis; cauline leaves
sessile, cordate-auriculate to amplexicaul at base,
denticulate. Racemes bracteate throughout or rare¬
ly along lower half, elongated considerably in fruit;
rachis straight; fruiting pedicels erect and ap-
pressed or ascending. Sepals oblong, deciduous,
erect, not saccate; petals pale yellow, spatulate,
clawed; stamens 0. tetradynamous. unappendaged;
anthers oblong; nectar glands confluent, lateral
ones strongly developed, cup-shaped, median
glands confluent with lateral ones; ovules 50 to 80
per ovary. Fruit dehiscent, linear, strongly latisep-
tate. secund, strongly falcate, deflcxed, (8-)10-16
cm long; valves with a distinct midvein and prom¬
inently thickened marginal veins, glabrous or pu¬
bescent; replum rounded, straight or constricted
between seeds; septum complete; style distinct, to
2 mm long; stigma capitate entire; seeds uniseriate,
winged all around, oblong, flattened; funieles ba-
sally adnate to septum; cotyledons aceumbent.

rseudotiirritis turrita (I,.) Al-Shehbaz, comb,
nov. Basionym: Arabis turrita 1,., Sp. IM. 2:
665. 1753. TYPE: “Brassica sylv. albido flore
nutante siliqua” in Boecone. Museo di Piante:
tab. 72. 1697 (lectotype, designated by Al-
Shehbaz & Turland, in Cafferty & Jarvis,
2002: p. 531).

Epitype. | Italy]. Toscana, La Verna. 27 May
1956, Corradi s.n. (FI).

RllAMMATOl'ttYI.U \l

Rhammatophyilum consists of nine species dis¬
tributed in Afghanistan, Kazakhstan, Kyrgyzstan,
lajikistan, Turkmenistan, and the Mongolian Altai
(Al-Shehbaz & Appel. 2002: Kamelin. 2002). All

species are shrubs with malpighiaceous, submal-
pighiaceous, or rarely subdendritic trichomes, en¬
tire leaves, apiculate anthers, flattened and linear
fruits, torulose valves with a distinct midvein, often
sessile and coarsely capitate stigmas, and oblong,
flattened, and usually winged seeds (Al-Shehbaz &
Appel, 2002). Species of Arabis are always herbs
with stalked stellate trichomes, distinct styles, and
entire stigmas often less in diameter than the style.

A tenth species of Rhammatophyilum, R. fruti¬
culosum, was originally described by Meyer (in
Ledebour, 1831) as a species of the monotypie sec¬
tion Arabis sect. Dendroarabis C. A. Meyer. Schulz
(1936) maintained that section, and the species re¬
mained there to the present. For reasons discussed
below, the species is transferred herein to Rham-
niatoph yllum.

RhainmatophyUum friilictdosuni (C. A. Meyer)
Al-Shehbaz, comb. nov. Basionym: Arabis fru-
ticulosa C. A. Meyer, in Ledebour, FI. Altaic.
3: 19. 1831. TYPE: “In rupestribus montium
Kurtsehum pr. Nabaty,” 10 June 1826. C. A.
Meyer s.n. (lectotype, designated by Bot-
schantzev, 1968: 144. LE).

Rhammatophyilum fruticulosum is widely dis¬
tributed in China (Xinjiang), Kazakhstan. Kyrgyzst-
an. Mongolia. Pakistan, Russia, and Tajikistan
(Zhou et ah, 2001). Although the species has been
maintained in Arabis to the present, it is quite
anomalous in this genus. It consists of shrubs to 50
cm tall and has dendritic trichomes, petiolate
leaves, pubescent fruits, and large stigmas nearly
as wide as the fruit. None of these characters occurs
in Arabis s. str. as delimited by Al-Shehbaz (2003),
and except for trichome morphology, the species fits
exceptionally well in Rhammatophyilum. Although
the trichomes in Rhammatophyilum are malpigh¬
iaceous or submalpighiaceous, in R. afghanieum
(K. H. Reehinger) Al-Shehbaz & (). Appel they are
a mixture of subdendritic and fewer submalpigh¬
iaceous ones, and in the related R. fruticulosum
they are exclusively dendritic. These two species
can also be separated by their fruits and seeds. In
R. fruticulosum the fruits are 1—2 nun wide and the
seeds are narrowly winged (wing to 0.1 mm wide),
whereas in R. afghanieum the fruits are (2.5— )3—
4.5 mm wide and the seeds are broadly winged
(wing 0.5—1 mm wide). Such a slight difference in
trichome morphology should not be alarming be¬
cause many genera in the Brassicaceae (e.g.. ,4/y.s-
sum I,., Draba L., Boechera) exhibit tremendous tri¬
chome diversity (Al-Shehbaz, 2002; Appel &
Al-Shehbaz. 2003).

Volume 15, Number 4
2005

Al-Shehbaz
Eurasian Arabis

523

77 RRITIS

The status of Turritis has fluc tuated in the vari¬
ous Holistic and monographic accounts between ac¬
ceptance and reduction to synonymy of Arabis. I he
literature dealing with the genus and its type spe¬
cies. T. glabra L. ( = A. glabra (L.) Bernhardi). is
too voluminous to cite, and the interested reader
should consult the references on Arabis for leads
(e.g., Al-Shehbaz, 2003). The genus includes two
species, of which the type, T. glabra , is widely dis¬
tributed in Eurasia and North America (Hollins,
1993). file second species. T. laxa (Smith) Hayek
( = A. laxa Smith), is confined to the eastern Med¬
iterranean (Iraq, Israel, Lebanon, Syria, lurkey),
Caucasus, Yugoslavia, and Greece (Townsend &
Guest, 1980: Jones & Akeroyd, 1993: Tan, 2002).

Every molecular marker used in tlu1 study of Ar-
abis s.l. (e.g.. Koch. 2003: Koch et ah. 1999, 2000.
2001; Kropf. 2002; O'Kane & Al-Shehbaz. 2003)
clearly demonstrates that Turritis glabra has no
close affinity to Arabis except for being a remotely
related mustard. The placement of this species in
Arabis would enforce the recognition of a polyphy-
letic genus, a position with which I disagree.

Kr.'i to Gknkha or Eurasian Sereins Purvioisn
Placed in Arahis

la. Racemes bracteate throughout or at least along
lower half; flowers pale yellow; fruits (8— ) 1 0 — 16
cm long, margin conspicuously thickened ....

. Pseudoturrit is

lb. Racemes ebracteale; flowers while, pink, or pur¬
ple (pale yellow in Turritis glabra): Iruits less
than 8 cm long (8-12 cm in 71 laxa), margin nol
thickened.

2a. Shrubs: fruits densely pubescent; stigma dis¬
tinctly wider than style; leaves linear to oh-
lanceolate, petiolate, not auriculate at base

. RhammatophyUum

2b. Herbs; fruits glabrous, rarely sparsely pu¬
bescent; stigma narrower than style; leaves
of various other shapes, sessile, base auric¬
ulate or amplexicaul. rarely cuneate.

3a. Plants glaucous; racemes and upper
cauline leaves glabrous.

4a. Perennials; seeds wingless; basal
leaves long petiolate, persistent; tri-
chonies exclusivelv simple, restrict¬
ed lo petioles of basal leaves, petals

white, rarely pink . hourraea

4b. Biennials; seeds winged; basal
leaves short petiolate lo suhsessile,
withered at anthesis; al least some
trichomes stalked and forked; pet¬
als pale yellow or pink . Turritis

3b. Plants not glaucous; racemes and upper
leaves variously pubescent.

5a. Trichomes stalked stellate, rarely
malpighiaceous; fruits erect to as¬
cending, rarely divaricate .... Arabis

5b. Trichomes dendritic, forked, sessile
stellate, or setose; fruits deflexed or
rarely divaricate (Eurasia).

6a. Trichomes noil-setose, dendrit¬
ic, sometimes also stalked
forked (simple in some North
American species); fruits
(Asia) strongly falcate, not se-
cund; lower cauline leaves ses¬
sile . Haerhera

6b. Trichomes setose, simple,
those on leaves mixed with
sessile, appressed stellate ones
with one ray perpendicular to
leaf surface; fruits straight, se-
eund; lower . . . cauline leaves
petiolate . Catolobus

Acknowledgments. I am grateful to Richard K.
Brummilt. Genii Davidse, and Nicholas J. Iiirlatul
for advice on nomenclatural matters, as well as to
Charlie If. Jarvis and Steve Cafferty for tvpification
of some Linnaean names in Arabis. I am indebted
to Olga Cherneva, Vladimir Dorofeyev. and Alex¬
ander Sennikov for their help in locating the types
of some species at EE, and to Tatyana Shu I k i na for
translating some of the Russian literature. East, but
not least. I thank Victoria G. Ilollowell for editorial
advice and critical checking of the manuscript.

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Agave valenciana (Agavaceae), a Gigantic New Species
from Jalisco, Mexico

M iguel Chdzaro- Basanez

Averin la de los Maestros y Mariano Barcenas, Guadalajara 44260, Jalisco, Mexico.
Departamento de Geografia, Universidad de Guadalajara-CUCSH. pachy88@prodigy.net. mx

J. Antonio Vazquez- Garcia

Km 15.5 carr. Guadalajara a Nogales, Las Agujas, Nextipac, Zapopan, 45110, Jalisco,
Mexico. Instituto de Botanica, Departamento de Botanica y Zoologia,
Universidad de Guadalajara-GUCBA. jvazquez@cucba.udg.mx

Yalma Luisa Vargas- Rodriguez

107 Life Sciences Building, Department of Biological Sciences, Louisiana State University,
Baton Rouge, Louisiana 70803, U.S.A. yvargal@lsu.edu

ABSTRACT. Agave valenciana (Agavaceae), a new
species from Jalisco, western Mexico, is described
and illustrated. This species is endemic to a small
section of the canyons of the Mascota and Talpa
rivers. It belongs to the Agave subgenus Agave L.
and to the Marmoratae species group. It appears to
be closely related to A. marmorata Roezl from
Puebla and Oaxaca. It differs from the latter in hav¬
ing a much larger rosette diameter, it has fewer but
longer and wider leaves, much larger flowers, and
it grows in less arid environments. This discovery
has implications for “raieilla" production, an al¬
coholic tequila-like beverage traditionally manu¬
factured in western Jalisco, Mexico.

Rksumkn. Se describe y se ilustra Agave valen¬
ciana (Agavaceae), una especie nueva de Jalisco,
en el occidente de Mexico. Esta especie es ende-
mica tie una porcion de las cuencas de los nos
Mascota y Talpa. Pertenece al subgenero Agave L.
y al grupo Marmoratae. Su pariente mas cercano
parece ser A. marmorata Roezl de Puebla y Oa¬
xaca. Difiere de esta ultima en tener un mayor dia-
metro de roseta, menos hojas pero mas largas y
anchas, flores mas grandes y creee en ambientes
menos aridos. Este descubrimiento tiene implica-
ciones en la produccion de “raieilla”, una bebida
similar al tequila que se produce tradicionalmente
en el oeste de Jalisco, Mexico.

Key words: Agave, century plant, maguey, Mar¬
moratae, mescal, Mexico, mezcal, raieilla.

An alcoholic tequila-like beverage, a type of
mescal, called “raieilla” or “ximat” is traditionally

Novon 15: l

prepared from A. maximiliana Baker (“lechugui-
11a”) in rural highlands of western Mexico (Vaz-
quez-Garcfa et al., 1995, 2004; Valenzuela Zapata,
2003). However, the “raieilla” is occasionally man¬
ufactured from, or mixed with, a different century
plant called “maguey relisero” (Rafael Castillon &
Oscar Valencia, pers. comm.). After examining w ild
populations of “maguey relisero” at Mascota, Jalis¬
co, we could not place it in any currently recog¬
nized species. Thus, it is herein described as new.

Agave valenciana Chazaro & A. Vazquez, sp. nov.
TYPE: Mexico. Jalisco: Mun. Mascota. Coamil
del Naranjo, 7 km NW of Mascota, 20°35,N,
104°54'W, 1250 m, transition betw. tropical
dry forest & oak forest, 25 Apr. 2004 (fl), V.
L. Vargas- Rodriguez, M. Chazaro-B. & ./. A.
Vazquez- Garcia 436 (holotype, IEB; isotypes,
ENCB, GUADA, IBUG, MEXU, MO, WIS,
XAL). Figures 1 and 2.

Herba simplex nonsurculosa, A. marmoratae affinis sed
statura validiore et foliis el fforibus grandioribus; rosulis
usque 1. 7-2.2 m altis, 2.7-3.3(44) m diametro; foliis 7
ad 15 rosulatis, 150-230 cm longis, 37-46(53) cm latis
ad medium, atro-viridibus, supra pallide zonatis, viridibus
vel glaucis notatis; inflorescentia cum scapo 5-6(7) m alta,
Horibus 50—60(70) mm longis intense luteis; capsulis 23—
29 mm longis, 10-13 mm latis; seminibus 3-4.5 mm lon¬
gis, 2—3 mm latis.

Perennial plant, solitary, not suckering, 5—6(7) m
tall: rosettes 1.7-2. 2 m high, 2.7-3.3(44) m diam.,
short-stemmed, with 7 to 15 leaves. Mature leaves
150-230 cm long, 37-46(53) cm wide in the mid¬
dle, up to 30 cm wide at the base, up to 15 cm
thick at the base, broadly lanceolate, in various at-

25-530. Published on 12 December 2005.

526

Novon

4 cm

Figure 1. Agave valenciana Chdzaro & A. Vazquez, lint' drawing based on M. Chdzaro el al. Mill) (IBUG). — A.
Habit. — B-l). Leaf teeth at the base, middle, and top, from Vargas- Rodriguez el al. 436 (IBUG). — K. Terminal spine,
from Vargas- Rodriguez el at. 436 (IBUG). — F. Inflorescence. — G. Flowering branch. — H. Flower in cross section. —
I. Flower bud. J. Mature flower. — K— I.. fruit and seeds, from V dzquez-GureCa el al. 7V54 (IBUG).

titudes, thick, firm concave, dark green, slightly
glaucous, smooth at lower and upper surface, cross-
banded in built sides; margins frequently undulate,
distinctly crenate at the middle to closely dentate

at flic ends, with mammae 4— 6 X 5-1') nun; mam¬
milliform teeth mostly 2— 5(— 7) mm. 5-17(22) mm
apart, flattened, the cups from very broad bases 5—
()(12) mm, mostly straight, some curved toward tin*

Volume 15, Number 4
2005

Chazaro-Basanez et al.

Agave valenciana from Jalisco

527

base, dark brown to red-brown, interstitial teeth few
or none; spine 10—15(20) mm long, usually short
and eonie, robust, rarely short-decurrent, dark
brown to red-brown. Panicles 5—6(7) m high, shaft
12—35 cm (Ham. at die base, with 25 to 35 diffuse
and flat-topped umbels in the upper hall of shaft;
bracts triangular, acute at the apex, variable in size
along the shaft, spirally arranged, searious. ca. 15
cm apart. Flowers 50-60(70) mm long, brilliant yel¬
low, protandrous; ovary body 18—27 X 4—5 mm,
light green, the neck 3—5 mm long and unconstrict¬
ed; tube 5—7 mm long, 10 mm wide, somewhat lun-
nelform; tepal 10-13 X 4-5 mm, linear, fleshy,
erect, the apex galeate, yellow1; filaments 34—38 mm
long, inserted at 2—4 mm near base of tube, slender,
yellow; anthers 10-17 mm long, centric, yellow.
Capsules oblongoid 23—29 X 10—13 mm, slender
stipitate, short-beaked, thin walled; carpels acute,
23—29 X 7—8 mm, curving outward for dehiscence;
seeds triangular, but curved in one side, 3—4.5 X
2—3 mm, flat, membranous, light, black (Fig. I).

Habitat. Agave valenciana thrives on steep
slopes of basaltic rock in canyons that are part of
the Mascota and Talpa watersheds, at the north¬
western end of Sierra Madre del Sur. It is, so far,
endemic to this region although the flora of much
of this cordillera is not well known. Agave valen¬
ciana grows in the ecotone of tropical dry forest
with oak forest, together with Euphorbia tanqua-
huete Sesse & Mocino, Bursera Jacquin ex L., Ficus

petiolaris lll>k. Jacaratia mexicana DC., Plumeria
rubra I... Agonandra racemosa (DC.) Stand ley. Eu¬
phorbia pulcherrima Willdenow, Quercus L, Pseu-
dobombax ellipticum HBK. Cephalocereus alensis
(Weber) Britton & Rose, Pedilanthus calcaratus
Schleehtendal, Nopalea Salm-Dyck, Opuntia Mill¬
er, Mammillaria Torrey & Gray, Hechtia Klotzsch,
and Pittocaulon phylliaris McVaugh.

Phenology. Flowering from March to May;
fruits dehiscing from September to November (Apo-
linar Gomez-Nunez, pers. comm., 2004).

Common names. “Maguey relisero,” “maguey
marzeno,” or “maguey manso” (Oscar Valencia,
pers. comm.).

Distribution. So far. Agave valenciana is en¬
demic to the steep and rocky canyons of the Mas-
cota and Talpa rivers, both in western Jalisco, at
900-1250 m asl (Fig. 2). This species was recently
observed, but not collected, 15 km west of the type
locality, near El Mosco canyon, in the vicinity of
the Talpa river. The authors observed less than 300
adult individuals near the type locality; however,
further exploration in the entire extent of those can¬
yons is urgently needed, especially at FI Mosco
area.

Conservation status. Agave valenciana deserves
a protected status. This species has an adult mean
density of 15 individuals per ha (Apolinar Gomez-

528

Novon

Nunez, pers. comm.), it is not a suckering plant,
and it is sometimes used for “raicilla” production.
Thus, we propose its inclusion to the Mexican en¬
dangered species act (NOM-059-ECOL, 2001) un¬
der the Special Protection category. This category
includes possible threatened species and promotes
population recovery and habitat restoration.

Different conservation and reproduction efforts
for Agave valenciana have arisen. Pew plants have
been grown successfully from seeds (Oscar Valen¬
cia, pers. comm.) and specific mature individuals
have been spotted for seed collection during the
early autumn of 2005. Apolinar Gomez-Nunez, an
agronomist and teacher at the CEBETA high school
(Direccidn General de Educacidn Tecnologica
Agropeeuaria-SEP, Mascota, Jalisco), is currently
collaborating with the authors implementing de¬
mographic and pollination studies, seed collections,
and germination experiments for A. valenciana con¬
servation and cultivation purposes.

Studied populations were located on private
properties; three of the landowners, Ramon Brise¬
no. Miguel Esparza, and Margarita Nelson, are
aware of the potential of this species and the need
to protect it. For now, extraction of plants from hab¬
itat should be avoided to prevent ecosystem deg¬
radation. However, planned collection of seeds
should be allowed to promote its reproduction in
nurseries and its eventual introduction into culti¬
vation.

Potential use and economic value. Agave valen¬
ciana is a remarkable addition to the succulent flo¬
ra of western Mexico and deserves to be cultivated
at least as an ornamental species because of its
impressive size and its showy brilliant yellow flow¬
ers. Its potential for “raicilla” production is also
promising but needs to be explored and evaluated.
For instance, mature “heads" of A. valenciana used
for “raicilla" production weight far over 350 kg
(Manuel Salcedo Gutierrez, pers. comm.. 2004), at
least three times heavier than those of A. tequilana
Weber. Agave valenciana is known for having ac¬
ceptable sugar content (Apolinar Gomez-Nunez.
pers. comm.. 2004), and it is believed that the
plants reach maturity in 15 years. To the best of
our knowledge, the density of A. valenciana in the
studied area is nearly 50 individuals per ha. with
an adult density of 30%. Considering the extent of
the two canyons (20 km to their convergence) and
their relatively homogeneous habitat, we speculate
that several thousands of adult individuals could
exist in tin* area.

The social organization and infrastructure for
promoting Agave valenciana as a new source of

“raicilla" in western Jalisco already exist, since re¬
cently a Mexican board (Consejo Mexicano de Pro-
rnocion de la Raicilla) for promotion of “raicilla”
from A. maximiliana was established. This board is
now headed by Manuel Salcedo-Gutierrez and in¬
cludes 37 board members, 22 distilleries, with over
50,000 plants and an annual “raicilla” production
of 5,000 liters.

Eponymy. We are pleased to name this impres¬
sive century plant after Oscar M. Valencia- Pelayo,
a talented amateur botanist who in the last five
years has been engaged in the study and reproduc¬
tion of trees and agaves of western Mexico.

Discussion. Agave valenciana belongs to the
Agave subg. Agave L. and to the Marmoratae spe¬
cies group as assigned by H. S. Gentry (1982). Ex¬
cept for its much larger habit and leaves, this spe¬
cies shares most of the distinctive characteristics
of the Marmoratae species group (Gentry, 1982),
such as crenate margins, small terminal spines,
small bright yellow flowers with small tubes 1/4 to
1/2 as long as the tepals and cross-banded leaves
in both sides.

Agave valenciana is perhaps most related to A.
marmorata, from Puebla and Oaxaca in Mexico, for
having a robust habit, leaves much wider at the
middle than at the ends, cross-banded in both
sides, and marginal teeth spaced over 15 mm apart.
The new species differs from the latter in having a
much larger rosette diameter (cf. Table 1 for con¬
trasting states in A. marmorata ), fewer but longer
and wider leaves, smaller spines, teeth, and cap¬
sules, and growing in less arid environments of
western Jalisco in the transition between tropical
dry forest and oak forest (Table 1). We also con¬
trasted A. gypsophila H. S. Gentry and A. nayari-
tensis II. S. Gentry against A. valenciana (Table 1),
since they occur in western Mexico anti are mem¬
bers of the Marmoratae species group, sensu Gentry
(1982). However the two are far smaller in habit,
they are not distinctly wider at the middle of the
leaves than at the ends, neither are the leaves
cross-banded. Furthermore, they differ in geogra¬
phy and habitat (Gentry, 1982; McVaugh, 1989; Fi¬
ler & Kristen, 2002).

fhe Marmoratae species group sensu Gentry
(1982), w ith the addition of Agave valenciana. now
includes six species and displays notable patterns
of allopatric speciation, habitat specialization, and
narrow endemism (Fig. 2). In Mexico, Agave zebra
11. S. Gentry is only known from arid mountains in
the Sonoran Desert (Gentry, 1982); A. nayaritensis
H. S. Gentry is only known from moist tropical
cliffs at Navarit and Sinaloa (Etter & Kristen,

Comparison of Agave valenciana Chazaro & A. Vazquez with morphologically anil geographically related species of I he Marmoratae species group (Gentry, 1982; litter

Volume

2005

15, Number 4

Chazaro-Basanez et al. 529

Agave valenciana from Jalisco

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2002); A. valenciana Chdzaro & VAzquez is only
known from transitional tropical dry forest with oak
forest on basaltic slopes at western Jalisco; A. g\p-
sophila H. S. Gentry is known from limestone at the
seaward facing slopes of Sierra Madre del Sin in
Colima, Jalisco (but not from Manantlan as cited
by Vazquez-Garcfa el al., 1995), Michoacan, and
Guerrero (Gentry, 1982; McVaugh, 1989); A. mar -
morata Roezl is found in semiarid xerophytic* scrub
in Puebla and Oaxaca (Gentry, 1982); and ,4. gri-
jalvemis B. Ullrich is endemic to the tropical dry
forest at Canon del Sumidero, Depresion Central de
Chiapas (Ulrich, 1990).

Paratypes. MEXICO. Jalisco: Mun. Mascota, Coamil
del Naranjo, 7 km NW of Mascota, 12 Apr. 2001, \1.
Chdzaro-R., 0. Valeneia-P. & I. Contreras-V. 8//0(IBUG);
25 Apr. 2004, E /. Vargas- Rodriguez, M. Chdzaro-R. &
./. A. Vdz<i uez- Garc(a 438 (I 111 1(7); 25 Apr. 2004, ./. A.

I (iztjuez-dan ia, ). /,. Vargas-Rodriguez & M. Chdzaro-H.
782*81 (1BUG); 18 Die. 2004, ,/. A. Vdzquez-Carcta, 1. /,.
Vargas-Rodriguez & Apolinar Cdmez 795- / (1BUG).

Acknowledgments. Vie thank Rafael Cast i I l<5n-
I/>pez and Hamdn Segura, both from Mascota, Ja¬
lisco, for providing information that led to the dis¬
covery of A. valenciana ; Hamdn Briseno, Apolinar
Gomez, Ignacio Contreras, and Jesus Cortes for
held assistance; Theodore S. Cochrane lor prepar¬
ing the Latin diagnosis; Refugio Vazquez for the
line drawing; Eduardo Sahagun and Patricia Her¬

nandez de Chazaro for their valuable suggestions

on the manuscript; Hirineo Martinez Barragan and

Mario Alberto Ruiz I/ipez for their academic sup¬
port.

I .iterature Cited

Liter, J. & M. Kristen. 2002. Agave nayariteusis. Cad.
Succ. .1. 74(5): 241-244.

Garcia-Mendoza, A. 2002. Distribution of Agave (Agava-
ccae) in Mexico. Cad. Succ. J. 74(4): 177-187.

Gentry, H. S. 1982. Agaves of Continental North America.
Univ. Arizona Press, Tucson, Arizona.

McVaugh, It. 1989. flora Novo-Galiciana 15: Bromelia-
ceae to Dioscoreaceae. Univ. Michigan Herbarium.

N0M-059-ECOL. 2001. Norma ofieial mexicana. Protec¬
tion ambiental-especies nativas de Mexico de flora y
fauna silvestres-categorfas de riesgo y especificaciones
para su inclusion, exclusion o eambio-lista de especies
en riesgo. Diario Ofieial de la Federacidn. 06 de marzo
de 2002, Mexico, D.K

Ulrich, B. 1990. Agave grijah crisis Ullrich. Line neue Art
aus Chiapas | 1 gave grijalvensis Ullrich. A new species
from Chiapas |. Kakteen And. Sukk. 41(6): 102-108.

Valenzuela Zapata, A. <i. 2003. El agave tequilero: cultivo
e imlustria de Mexico. Mundi-Prensa.

V azi]uez-( tarda, J. A.. It. Cuevas-G., T. S. Cochrane, II.
H. litis, E J. Santana-M. L. Guzm4n-ll. 1995. f lora
de Manantlan. Sida Bot. Misc. 13. Botanical Research
Institute of Texas.

- , M. Chazaro It., <7 Nieves II.. Y. L. Vargas-Ro¬
driguez, M. Vazquez (3 & A. Flores M. 2004. Flora del
norte de Jalisco y clnohotanica huiehola. Serie Fronte-
ras de Biodiversidad I. Universidad de Guadalajara
CUCBA-CUSH.

Resurrection of Seutera Reichenbach
(Apocynaceae, Asclepiadoideae)

Mark Fish be in

Department of Biological Sciences, Mississippi State University, P.O. Box GY,
Mississippi State, Mississippi 39762, U.S.A. Current address: Portland State University,
Department of Biology, P.O. Box 751, Portland, Oregon 97207, U.S.A.

mfish@pdx.edn

W. I). Stevens

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

stevens@mobot.org

ABSTRACT. The genus Seutera Reichenbach is
resurrected to accommodate the taxa most frequent¬
ly known as Cynanchum august ifoli urn Persoon. C.
palmeri (S. Watson) S. F. Blake, and C. peninsula re
S. F. Blake, for which the combinations Seutera an-
gustifolia (Persoon) Fishbein & W. I). Stevens. S.
palmeri (S. Watson) Fishbein & W. I). Stevens, and
S. palmeri var. peninsularis (S. F. Blake) Fishbein
& W. I). Stevens are provided. The nomenclatural
and taxonomic validity of Seutera are discussed.
Ceropegia palustris Pursh and Amphistelma sali-
narum C. Wright are placed in the synonymy of
Seutera angustifolia. Cynanchum mulegense Wig¬
gins is placed in the synonymy of Seutera palmeri.

Key words: Apocynaceae, Asclepiadoideae, Cy¬
nanchum, Funastrum, Seutera.

Woodson (1941) proposed broad generic con¬
cepts for North American Aselepiadaceae, follow¬
ing which Cynanchum L. had 23 synonyms and was
divided into six subgenera. A few ol the subim >rgcd
genera, particularly the entirely South American
Ditassa R. Brown and the largely South American
Tassadia Deeaisne, were rarely accepted as syno¬
nyms by subsequent students, while others, includ¬
ing Metalepis Grisebach, Metastelma R. Brown, and
Orthosia Deeaisne have been gradually resurrected
(e.g., Stevens et ah. 2001). Liede (1997) recircum¬
scribed the New World Cynanchum and divided it
into seven sections. More recent molecular phylo¬
genetic results (Liede N Tauber, 2002; Liede &
Meve, 2002; Rapini et ah. 2003; Liede-Schumann
et ah. 2005) suggest that the relationships of New
World Cynanchum and allied genera are more am¬
biguous than previously recognized. One of the
clear messages resulting from this recent work is
that only a small part of Woodson’s concept of New

Novon 15:

World Cynanchum is likely to be retained within
the evolving concept of the genus, probably only
Cynanchum subgen. Mellichampia Woodson, which
corresponds to Sundell’s (1981) Cynanchum sect.
Roulinia (Deeaisne) Sundell and section Melli¬
champia (A. Gray) Sundell, and to Liede's (1997)
Cynanchum sect. Mellichampia, section Roulinia,
and section Telminostelma (E. Fournier) Liede. As
Cynanchum. sensu Woodson, is gradually disman¬
tled, the taxonomic and nomenclatural issues of the
residual elements are being resolved. Ditassa, Me¬
tastelma, Orthosia, Metalepis, and Tassadia arc now
generally recognized, although sometimes ambigu¬
ously defined (Stevens, 1988, 2002: Fishbein &
Levin, 1997; Rapini, 2002; Rapini et ah. 2003).

Liede (1997) placed seven North American and
Caribbean species in Cynanchum seel. Machridea
(Rafinesque) Liede, with the type C. angustifolium
Persoon. While Cynanchum angustifolium was not
specifically mentioned in Woodson’s ( 1941) circum¬
scription, it would key to his Cynanchum subgen.
Metastelma on the basis of the umbelliform inflo¬
rescence. Liede and Meve (1997) ext luded this
species from Metastelma, noting most importantly
the basally connate corona lobes. In tbe analysis of
Liede and Tauber (2002), tbe three sampled spe¬
cies included in Cynanchum sect. Machridea. (,.
angustifolium, C. peninsulare S. F. Blake, and C.
utahense (Engelmann) Woodson are nested within
Funastrum E. Fournier; Liede and Meve (2002) for¬
mally transferred those three species to hinastrum,
reduced two other names originally me luded in Cy-
nanchum sect. Machridea to the synonymy of h.
peninsulare, and left the placement of the two re¬
maining species pending the availability of “ana-
lyzable material” (Liede & Meve. 2002: 579). This
placement was reconfirmed in a broader analysis

531-533. Published on 12 December 2005.

532

Novon

with more limited sampling (one species) of section
Macbridea (I Jede-Schumann et ah, 2005). Howev¬
er. support for the placement of the three species
within Funastrum, rather than as a sister group,
was not strong and was based solely on sequence
data from a single chloroplast locus.

Funastrum as here circumscribed is a New'
World genus of about 17 species of woody vines
with a corona comprised of a fleshy, unlobed ring
at the junction of the corolla and the gynostegium
and five vesicular lobes adnate to the backs of the
anthers. I he distinctive corona of Funastrum is a
potential synapomorphy setting the genus apart
from relatives in the Americas.

In our analysis of the morphology of the three
species transferred to Funastrum , Cynanehum
utahense is a plausible Funastrum ; although the
fleshy ring is poorly developed and the vesicles are
entirely absent, the position of tin1 ring, general
shape of the anthers, and the biogeography is com¬
patible with Funastrum and we accept that transfer.

I he other two species, however, are cither thin, her¬
baceous. and rhizomatous (Seutera angustifolia
(Persoon) Eishbein & W. I). Stevens) or suffrutes-
eent and prominently corky at the base (S. palrneri
(S. Watson) hishbein & W. I). Stevens), somewhat
succulent, glabrous, I inear- leaved, twining vines
growing on or near beaches or in near-shore
swamps in North America and the Caribbean. The
corollas are small, pale green or tinted with brown
or yellow, entirely glabrous, have a short tube and
long, narrow lobes that are erect at the base and
then spreading, and have strongly contorted aesti¬
vation. I lie coronas entirely lack tin* fleshy ring and
vesicles, but rather have five basallv connate, more
or less laminar, three-toothed lobes borne at the
junction of the corolla and the gynostegium. Most
of these characters would be discordant in Funas¬
trum. It is especially difficult to conceive how such
fundamentally different coronas could have arisen
within a genus of seemingly recently diverged (pers.
obs.) species. While the molecular sampling re¬
mains limited and the eladistic support not con¬
vincing, it seems imprudent to adopt a classification
that is morphologically untenable. Rather than pre¬
tend that traditional morphology, especially of the
corona, has no value whatsoever, we reject the in¬
clusion of Cynanehum angustifolium and C. pen-
insulare into Funastrum. Instead, we propose to
recognize these two morphologically distinct spe¬
cies in the ir own genus. Although one of the spe¬
cies. Snitera angustifolia. is distributed around the
Gulf of WU jxico, and the other, S. palrneri, is found
in Baja California, both are primarily beach and

salt marsh plants and the gap in the distribution is
reasonable.

The oldest generic name for these species is
Lyonia Elliott (Sketch Rot. S. Carolina 1: 316.
181 7, nom. rej.), not Lyonia Rafinesque (published
in 1808). nor Lyonia Nuttall (published in 1818,
nom. cons.). Lyonia Elliott was renamed Macbridea
Rafinesque (Amer. Monthly Mag. & Crit. Rev. 3:
99. June 1818) and Seutera Reichenbach (pub¬
lished in 1828). but the former has ambiguous pri¬
ority relative to Macbridea Elliott ex Nuttall (pub¬
lished in late May-early June 1818. dates fide
Index Nominum Generieorum). Since Macbridea El¬
liott ex Nuttall (Lamiaceae) is a currently recog¬
nized genus, it seems best to use the unambiguous
Seutera for this genus.

Seutera Reichenbach. Consp. Regn. Veg. 131.
1828. TYPE: Seutera maritima Decaisne, in
A. de Candolle, Prodr. 8: 590. 1844, nom. il-
leg.

Pattalias S. Watson, Proc. Amer. Acad. Arts 24: 60. 1889.
TV PE: Pattalias palrneri S. Watson.

Seutera angustifolia (Persoon) Eishbein & W. I).
Stevens, comb. nov. Basionym: Cynanehum
angustifolium Persoon, Syn. PI. 1: 274. 1805,
not Cynanehum angustifolium Wight & Arnott
(1834) or Cynanehum angustifolium (I)eeais-
ne) Morillo (1986). Funastrum angustifolium
(Persoon) Liede & Meve, Nordic J. Rot. 22:
587. 2002 [2003], TYPE: U.S.A. “Hab. ad lit-
tora maris in Carolina."* Miehaux s.n. (holo-
tvpe, P not seen).

Ceropegia palustris Pursli, El. Amer. Sept. I: 184. 1813
1 1814]. Syn. nov. Vincetoxicum palustre (Pursh) A.
Cray, Syn. Id. N. Amer. 2: 102. 1878. Cynanehum
palustre (Pursh) A. Heller, Cal. N. Amer. PI. 0. 1898.
Metastelma palustre (Pursh) Schlechter, in Urban,
Symb. Antill. I: 258. 1899. Lyonia palustris (Pursh)
Small, El. Miami 149. 1913. TYPE: U.S.A. “Caro¬
lina,” Lyon s.n. (holotype, 8M not seen).

Amphistelma salinarum C. Wright, in Grisehach, Cal. PI.
Cub. 175. 1866. Syn. nov. Metastelma salinarum (C.
Wright) C. Wright, in Sauvalle, Anales Acad. Ci.
Med. Habana 7: 105. 1870. Cynanehum salinarum
(C. Wright) Alain. Mem. Soc. Cub. Mist. Nat. “Eelipe
Poey” 22: 120. 1955. TYPE: Cuba. “Cuba ooc., in
pratis salinas pr. Guanimar,” C. Wright 295ft (holo¬
type, Gil not seen; isotype, MO).

I bis species has often been named Cynanehum
palustre (e.g., Standley & Williams, 1969) and it is
clear from both habitat and morphology that the
concepts are identical. Cynanehum salinarum is
likewise identical.

Volume 15, Number 4
2005

Fishbein & Stevens
Resurrection of Seutera

533

Seutera palmeri (S. Watson) Fishbein & \\. I).
Stevens, comb. nov. Basionym: Pattalias pal¬
meri S. Watson, Proc. Amer. Acad. Arts 24:
60. 1889. Cynanchum palmeri (S. Watson) S.
F. Blake, Contr. Gray Herb. 52: 83. 1917.
TYPE: Mexico. “Near a sandy beach at Mu-
leje,” E. Palmer 424 (holotype, GH not seen).

Cynanchum mulegen.se Wiggins, Proc. Calif. Acad. Sci.,
ser. 4, 30: 247. 1965. Syn. nov. TYPE: Mexico. Baja
California Sur: mouth of Estero Mu lege. /. Wiggins
& V. Wiggins 18085. 21 Oct. 1962 (holotype, l)S not
seen).

Seutera palmeri var. pen insular is (S. F. Blake)
Fishbein & W. D. Stevens, comb. nov. Basion¬
ym: Cynanchum peninsulare S. F. Blake, Contr.
Gray Herb. 52: 83. 1917. Cynanchum palmeri
var. peninsulare (S. F. Blake) I. M. Johnston,
Proc. Calif. Acad. Sci., ser. 4, 12: 1 129. 1924.
Funastrum peninsulare (S. F. Blake) Fiede &
Meve, Nordic J. Bot. 22: 589. 2002 [2003 1.
TYPE: Mexico. “Tower California, west coast
of Cape Region,” Nov. 1902, T. S. Brandegee
s.n. (holotype GH, not seen).

Although Cynanchum peninsulare and C. palmeri
were recognized as distinct species by Fiede
(1997), they were combined under Funastrum pen¬
insulare by Fiede and Meve (2002). We have ob¬
served several populations in the field in Baja Cal¬
ifornia Sur, Mexico, and conclude that the northern,
Gulf of California ( Seutera palmeri var. peninsular-
is), and southern. Pacific Ocean (S. palmeri var.
palmeri ), populations differ only slightly, primarily
in flower size, and we follow Johnston (1924) in
treating these allopatric populations as two varieties
of a single species. Cynanchum mulegense was de¬
scribed from the type locality of Seutera palmeri
and is identical.

Acknowledgments. We thank Bruce Hansen
and Victoria Hollowed for providing review com¬
ments.

Fiterature Cited

Fishbein, M. & R. Fevin. 1997. Metastelma mexicanum
(Asclepiadaceae): A new combination and re-evaluation
of the status of Basistelma Bartlett. Madrono 44: 268—
276.

Johnston, I. M. 1924. Expedition of the California Acad¬
emy of Sciences to the Gulf of California in 1921. Proc.
Calif. Acad. Sci., ser. 4, 12: 951-1218.

Fiede, S. 1997. American Cynanchum (Asclepiadaceae) —
A preliminary infrageneric classification. Novon 7:
172-181.

- & U. Meve. 1997. Some clarifications, new spe¬
cies, and new combinations in American Cynanchinae
(Asclepiadaceae). Novon 7: 38—45.

- & - . 2002 [2003]. Dissolution of Cynan¬
chum sect. Macbridea (Apocynaceae— Asclepiadoideae).
Nordic J. Bot. 22: 579-591.

- N A. Tauber. 2002. Circumscription of the genus

Cynanchum (Apocynaceae— Asclepiadoideae). Syst. Bot.
27: 789—800.

Liede-Schumann, S., A. Rapini, I). J. Goyder & M. W.
Chase. 2005. Phylogenetics of the New World subtribes
of Aselepiadeae (Apocynaceae— Asclepiadoideae): Me-
tastelmatinae, Oxypetalinae and Gonolobinae. Syst. Bot.
30: 184-195.

Rapini, A. 2002. Six new species of Ditassa R. Br. from
the Espinhago Range, Brazil, with notes on generic de¬
limitation in Metastelmatinae (Apocynaceae— Asclepia¬
doideae). Kew' Bull. 57: 565—583.

- . M. W. Chase, I). J. Goyder & J. Griffiths. 2003.

Aselepiadeae classification: Evaluating die phylogenet¬
ic relationships of New World Asclepiadoideae (Apo¬
cynaceae). Taxon 52: 33—50.

Standley, P. C. & F. () Williams. 1969. Flora of Guate¬
mala. Asclepiadaceae. Fieldiana, Bot. 24 (8, part 4):
407-472.

Stevens, W. D. 1988. New names and combinations in
Apocynaceae, Asclepiadoideae. Phytologia 64: 333—
335.

- . 2002. A new Tassadia (Apocynaceae, Asclepia¬
daceae) from Bolivia and Brazil. Novon 12: 408-410.

- , C. Ulloa C., A. Pool & (). M. Montiel. 2001.

Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot.
Gard. 85: i— xlii. 1—2666.

Sundell. E. 1981. The New World species of Cynanchum
subgenus Mellichampia (Asclepiadaceae). Evol. Mon¬
ogr. 5: 1—63.

Woodson, R. E., Jr. 1941. North American Asclepiadaceae
I. Perspective of the genera. Ann. Missouri Bot. Gard.
28: 193-244.

New Infragenera in Baccharis (Asteraceae, Astereae)

Daniel A. Giulia no

Division Plantas Vasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata,
Argentina. giuliano@museo.fenym.unlp.edu.ar

Abstract. Five new sections of Baccharis (As-
teraeeae, Astereae) are described: B. sect. Agglo-
meratae , including 2 species: B. condensata Busby
and B. platypoda DC.; B. sect. Canescentes, 8 spe¬
cies: B. gibertii Baker, B. gnaphalioides Sprengel,
B. helichrysoides DC., B. leucocephala Dusen, B.
leucopappa DC., B. patens Baker, B. phylicifolia
DC., and B. psammophila Malme; B. seel. Curity-
benses , I species: B. curitybensis I leering ex Mal-
me; B. sect. Paleatae, 1 species: B. retamoides Phi¬
lippi; and B. sect. Tridentatae, 6 species: B.
deltoidea Baker. B. dlinita DC., B. illinitoides Mal-
agarriga, B. isabclae Soria Zardini, B. subopposita
DC., and B. tridentata Vahl. Four new series of
Baccharis are described: B. [sect. Angiistifoliae ] ser.
Uniflorae, 7 species: B. frigida Kunth. B. hutchi-
sonii Cuatrecasas. B. laxi flora Busby, B. liberta-
densis (S. B. Jones) II. Robinson, B. sternbergiana
Steudel, B. trineura Soria & Zardini, and B. uniflo¬
ra (Buiz & Pav6n) Persoon; B. (sect. Cylindricae]
ser. Axillares, 20 species: B. angusticeps Dusen ex
Malme, B. aracatubensis Malagarriga & Hatsch-
baeb. B. axillaris DC., B. concinna G. M. Barroso,
B. cultrata Baker, B. elliptica Gardner, B. hyperi-
cifolia Baker, B. incisa Hooker <!4 Amott. B. late-
ralis Baker, B. lymanii G. M. Barroso, B. minuti-
flora Martins ex Baker, B. parvidentata
Malagarriga, B. pauciflosculosa DC.. B. perdusenii
Giuliano, B. polyphylla Gardner. B. pseudovacci-
nioides Malagarriga, B. selloi Baker, B. serrula
Schultz Bipontinus ex Baker, B. truncate Gardner,
and B. xiphophylla Baker; B. |sect. Nitidae | ser.
Hirsutae, I species: B. muelleri Baker; and B. [sect.
Nitidae | ser. Lanuginosae, 4 species: B. calvescens
DC., B. elaeagnoules Steudel ex Baker, B. myrici-
Jolia DC., and B. serniserrata DC. Tin ee new com¬
binations at the series level are proposed: Baccharis
ser. Angiistifoliae (Baker) Giuliano; B. ser. Cylin-
dricae (I leering) Giuliano; and B. ser. Nitidae (Cua¬
trecasas) Giuliano. A new name, Baccharis perdu¬
senii, is proposed to replace the illegitimate name
B. dusenii G. M. Barroso. A respective lectotype is
designated for B. platypoda. B. retamoides. and B.
tridentata-, a neotype is designated for B. uniflora.

B 1st MKN. Se describen cinco nuevas secciones

de Baccharis (Asteraceae, Astereae): B. sect. Ag-
glomeratae, incluyendo 2 especies: B. condensala
Busby y B. platypoda DC.; B. sect. Canescentes, 8
especies: B. gibertii Baker, B. gnaphalioides Spren-
geh B. helichrysoides DC., B. leucocephala Dusen.
B. leucopappa DC., B. patens Baker, B. phylicifolia
DC. y B. psammophila Malme; B. sect. Curityben-
ses, I espeeie: B. curitybensis fleering ex Malme;
B. sect. Paleatae. I espeeie: B. retamoides Philippi;
y B. sect. Tridentatae, 6 especies: B. deltoidea Ba¬
ker, B. dlinita DC., B. illinitoides Malagarriga, B.
isabclae Soria tv Zardini, B. subopposita DC. y B.
tridentata Vahl. Se describen cuatro nuevas series
de Baccharis: B. |sect. Angiistifoliae | ser. I nijlorae,
7 especies: B. frigida Kunth, B. hutchisonii Cua¬
trecasas. B. laxiflora Busby. B. libertadensis (S. B.
Jones) H. Robinson. B. sternbergiana Steudel. B.
trineura Soria & Zardini y B. uniflora (Ruiz Pa-
vbn) Persoon; B. [sect. Cylindricae | ser. Axillares,
20 especies: B. angusticeps Dusen ex Malme, B.
aracatubensis Malagarriga <N, Hatschbach. B. axilla¬
ris DC., B. concinna C. M. Barroso, B. cultrata
Baker, B. elliptica Gardner, B. hypericifolia Baker.
B. incisa Hooker & Arnott, B. lateralis Baker, B.
lymanii C. M. Barroso, B. minutiflora Martins ex
Baker, B. parvidentata Malagarriga, B. pauciflos¬
culosa DC.. B. perdusenii Giuliano, B. polyphylla
Gardner, B. pseudovaccinioides Malagarriga, B. sel¬
loi Baker. B. serrula Schultz Bipontinus ex Baker,
B. truncal a Gardner y B. xipliopliylla Baker; B.
|sect. Nitidae ] ser. Ilirsutae. I espeeie: B. muelleri
Baker; y B. \ sect. Nitidae \ ser. Lanuginosae. 1 es¬
pecies: B. calvescens DC., B. elaeagnoules Steudel
ex Baker, B. myricifolia DC. y B. serniserrata DC.
Se proponen Ires nuevas eombinaciones a nivel de
series: Baccharis ser. Angiistifoliae (Baker) Giulia-
no, B. ser. Cylindricae (I leering) Giuliano y B. ser.
Nitidae (Cuatrecasas) Giuliano. Se propone tin
nombre nuevo, Baccharis perdusenii. para reempla-
zar el nombre ilegftimo B. dusenii C. M. Barroso.
Se designa un respective leetotipo para B. platy¬
poda, B. retamoides y B. tridentata: se designa un
neotipo para B. uniflora.

Key words: Asteraceae, Astereae, Baccharis.
sections, series. South America.

Novon 15:534—541. Pubi.ishkd on 12 Dkcembeh 2005.

Volume 15, Number 4
2005

Giuliano

Infragenera in Baccharis

535

Baccharis I... with ca. 375 species (Giuliano, in
prep.), is the largest genus within Asteraeeae tribe
Astereae. Its infrageneric taxonomy is under revi¬
sion; tlie sections created by classical authors (Per-
soon, 1807; Candolle, 1836; Baker, 1882) were
highly heterogeneous groups, since they were based
upon vegetative features (mainly leaf shape). Cua-
treeasas (1967) redefined and lectotypified those
infragenera, thus elucidating in part this problem.
However, nearly 90 species have never been as¬
signed to sections, whereas many others are mis¬
placed in the sections into which they were origi¬
nally classified. Here nomenclatural novelties are
presented, in advance of the overall revision of the
infrageneric taxonomy of Baccharis.

Nkw Suctions ok Baccharis

Baccharis sect. Agglomeratae Giuliano, sect,
nov. TYPE: Baccharis platypoda DC.. Prodr. 5:
409. 1836.

A ceteris sectionibus generis Baccharidis diflert loliis
penninervis numerosis nervis lateralibus et capitulescen-
tiis valde dimorphis (capitula staminata in glomerulis lon-
giter pcdunculatis ad axilam foliorum superiorum inserta;
capitula pistillata in capitulescentiis globosis densis ter-
minalihus disposita).

Erect shrubs, clothed by minute tufted tri-
chomes. Leaves petiolate, blade pinnately veined,
margins entire or with few pairs of teeth. Capitula
sessile, the staminate ones arranged in long pedi¬
cellate glomerules inserted at the axils of the distal
leaves, the carpellate ones arranged in terminal
sessile globose capitulescences; involucres eam-
panulate; receptacles flat, alveolate. Ratio of car-
pellate:staminate florets [corresponding to carpel-
late vs. staminate specimens respectively] ea. 1:1.
Carpellate florets with corollas dentate at apex.
Achenes terete, 10-ribbed, glabrous; pappus bris¬
tles in 2 series, non-aeerescent, caducous. Stami-
nate florets with corollas showing little differenti¬
ated limb; style branches attached, brief; pappus
bristles plumose at apex.

Includes two species, one from the Andes of Bo¬
livia (B. condensala ) and the other from eastern
Brazil (B. platypoda).

1. Baccharis condensata Busby, Descr. 5. Amer.

PI.: 148. 1920.

2. Baccharis platypoda DC., Prodr. 5: 409.

1836. TY PE: “N° 283, Herbier de Bresil (Mr.
Vauthier 1833), Marianne (Prov. Minas Ge-
raes), Boite N° 795 Herb, genevense” (lecto-
type, tlesignated here, G-DC). [This specimen
was selected from the three sheets at G-DC
containing type material because it is repre¬
sentative for the species, and the label clearly
provides the type locality and collection data.]

Baccharis platypoda was placed by Candolle
(1836) and Baker (1882) in the formerly heteroge¬
neous broad section Cuneifoliae DC. in Baccharis.
The other species included in section Agglomeratae
has never been classified into any section.

Section Agglomeratae belongs to subgenus Bac¬
charis because it has attached brief staminate style
branches and 10-ribbed glabrous achenes with bi-
seriate caducous pappus bristles; however, its affin¬
ities with other sections are unknown. This section
is unique within the genus because of the capitu-
lescence dimorphism and the pinnately veined
blades with numerous lateral veins.

Baccharis sect. Canescentes Giuliano, sect. nov.

TYPE: Baccharis helichrysoides DC., Prodr. 5:
415. 1836.

Suffrutiees alho-lanuginosi; folia sessilia iliscoloria uni-
nervia vel penninervia; capitula in cymis eorymbiformibus
compositis axillaribus et terminalibus vel in cymis corytti-
biformibus congestis ad apicem ramulorum disposita; in-
volucra hemisphaerica; achaenia 5-costata pi 1 is geminis
et papillis vcstita; flnrum pistillatorum setae pappi ae-
crescentes; florum staminatorum rami styli lanceolati se-
parati.

Erect or decumbent subshrubs, plants lanugi-
nous, densely clothed by uniseriate straight or
slightly curved trie homes, usually accompanied by
biseriate glandular triehomes on upper leaf surfaces
and by tangled uniseriate triehomes on lower leaf
surfaces. Leaves sessile, discolorous, 1 -veined or
with 1 to 3 pairs of lateral veins, margins entire
and revolute. Capitula pedicellate, arranged in ax¬
illary and terminal compound corymbiform cymes,
or in congested corymbiform cymes at the apex of
scapose stems; involucres hemispheric; receptacles
convex or conic, alveolate, clothed by triehomes.
Ratio of carpellate:staminate florets ea. 1:1. Car¬
pellate florets with corollas dentate at apex.
Achenes terete, 5-ribbed, with short and asymmet¬
ric or very long twin triehomes; pappus bristles in
2 series, accrescent, persistent. Staminate florets
with corollas showing well-differentiated limb; style
branches lanceolate, free; pappus bristles smooth.

Eight species, which grow1 in eastern and south¬
ern Brazil, southeastern Paraguay, northeastern Ar¬
gentina, and Uruguay.

1. Baccharis gibertii Baker, in Martins. FI. Bras.

6(3): 52. 1882.

2. Baccharis gnaphalioides Sprengel, Syst. Veg.

3: 461. 1826.

3. Baccharis helichrysoides DC., Prodr. 5: 415.

1836. TYPE: Brazil. Rio Grande do Sul: Sello

<1201. 1 (ex Herbarium of Museum Imperialis

Brasiliensis n° 811) (leetotype, designated by

Barroso (1976: 60), P).

536

Novon

4. Baeeharis leueoeephala Dusen, Ark. Bot.
9(15): 24. 1910.

5. Baeeharis leiieopappa DC.. Prodr. 5: 415.

1836.

6. Baeeharis patens Baker, in Marlins, FI. Bras.

6(3): 52. 1882.

7. Baeeharis phylieif'olia DC., Prodr. 5: 415.

1836.

8. Baeeharis psanmiopliila Malme, Kongl.

Svenska Vetenskapsakad. Handl. ser. 3, 12(2):
70. 1933.

Baeeharis leucocephala and B. psammophila
have not been assigned to any section before; the
other species have been placed by classical authors
(Candolle, 1836; Hooker & Arnott, 1841; Baker,
1882) in die formerly heterogeneous section Dis-
colores DC.; however, by virtue of the redefinition
of diis section by Cuatreeasas (1967), it lias to in¬
clude B. phylicoides Kunth and related species (B.
caprariifolia DC., B. erioclada DC., B. nummuloi-
de.s fleering ex Malme, etc.). Also, several of the
species here affiliated with section Canescentes
were placed by 1 leering ( 1904) and Giuliano (2001)
in Baeeharis sect. Paniculatae Heering, but the lat¬
ter possesses heads solitary at the apex of short
branehlets (in turn grouped to form leafy paniculate
capitulescences), usually Hat naked receptacles,
and achenes with uniseriate non-glandular tri¬
chomes and/or biseriate glandular trichomes, while
section Canescentes has heads inserted at the axils
of highly reduced bracts, conic or convex recepta¬
cles covered by long setae, and achenes with twin
trichomes.

Baeeharis sect. Curityhenses Giuliano, sect. nov.

TY PE: Baeeharis curitybensis fleering ex Mal¬
me, Kongl. Svenska Vetenskapsakad. Handl.
ser. 3, 12(2): 69. 1933.

Sectionibus subgeneris Baccharidis similis achaeniis
10-costatis et pappi setis 2-seriatis caducis, sed florum
staminatorum styli ramis lanceolatis separatis et achaeniis
pilis geminis vestilis differ!.

Erect shrubs, lanuginous, densely clothed by
uniseriate filiform trichomes. Leaves petiolate,
blades discolorous, 3-veined, margins entire or with
few pairs of teeth. Capitula pedicellate, arranged in
corymbiform cymes inserted at the axil of reduced
distal leaves, in all forming a leafy compound cor¬
ymb; involucres hemispheric; receptacles flat, al¬
veolate. Ratio of carpellate:staminate florets ca. 1:
I. Carpel late florets with corollas dentate at apex.
Achenes terete, 10-ribbed, with twin trichomes;
pappus bristles in 2 series, non-accrescent, cadu¬
cous. Staminate florets with corollas with well-dif¬

ferentiated limb; style branches lanceolate, free;
pappus bristles plumose at apex.

Includes only one species, endemic to south¬
eastern Brazil (Sao Paulo, Parand, and Santa Ca¬
tarina).

1. Baeeharis curitybensis Heering ex Malme.
Kongl. Svenska Vetenskapsakad. Handl. ser.
3, 12(2): 69. 1933. TYPE: Brazil. Parana: Cu¬
ritiba. 20 Oct. 1908, Dusdn 6906 (lectotype,
designated by Barroso (1976: 97), S).

This species was not previously affiliated with
any section. It shows contradictory characters: the
number and structure of the achene ribs and the
number of series and the caducous condition of the
pappus bristles are similar to those of subgenus
Baeeharis ; on the other hand, the style branches
are lanceolate and free as in all the rest of the
subgenera, and the achenes are clothed by twin
trichomes. f urther research is needed in order to
elucidate its affinities.

Baeeharis sect. Paleatae Giuliano, sect. nov.
TYPE: Baeeharis retamoides Philippi, Anales
Univ. Chile 34: 180. 1870.

Sectioni Angustifoliis primo adspectu similis, sed plan-
tis minutis nidis pilosis vestitis, capitulorum pistillatorum
receptaculis paleaeeis, floribus pisti 1 latis:stami nat is ratio-
lie 4:1 vel 5:1, et achaeniis pilis geminis dense vestitis
differ!.

Erect subshrubs, clothed by minute tufted tri¬
chomes. Leaves sessile, blades 1 -veined, margins
entire, the distal ones reduced to small bracts. Ca¬
pitula pedicellate, solitary at the apex of the
branches or grouped to form a corymbiform or ra-
cemiform cyme; involucres hemispherical; recep¬
tacles of carpellate capitula paleaceous. Ratio of
carpellate:staminate florets 4:1 or 5:1. Carpellate
florets with corollas truncate at apex. Achenes te¬
rete, 5-ribbed, large-sized (2.7— 4.5 mm long),
densely clothed by long twin trichomes; pappus
bristles in 2 to 3 series, accrescent, persistent. Sta¬
minate florets with corollas showing well-differen¬
tiated limb; style branches lanceolate, free; pappus
bristles denticulate.

Includes a single species, which grows in Argen¬
tina, from the provinces of Salta to Mendoza.

1. Baeeharis retamoides Philippi, Anales Univ.
Chile 34: 180. 1870. TYPE: Argentina. Men¬
doza: 1868—1869, R. Philippi s.n. (lectotype,
designated here, SGO-060829). [Although the
two original type material sheets (the other is
SGO-044062) are morphologically similar and
present identical collection data, this speci¬
men is selected because it is more profusely
branched. 1

Volume 15, Number 4
2005

Giuliano

Infragenera in Baccharis

537

This species was previously placed within sec¬
tion Angustifoliae Baker emend. Cuatrecasas (Ciu-
liano, 2001). based on leaf morphology and capi-
tuleseence architecture; nevertheless, it presents
remarkable differences, such as the vegetative ves-
liture of minute tufted hairs (vs. non-tufted unise-
riate hairs with acuminate apical cell), carpellate
capitula with paleaceous receptacles (vs. non-pa-
leaceous), ratio of earpellate:staminate florets ca. 5:
1 (vs. ca. 1:1), and achenes clothed by long twin
trichomes (vs. sparsely clothed by uniseriate tri¬
chomes). Within Baccharis, the paleaceous recep¬
tacles are found in section Oblongifoliae DC.
emend. Cuatrecasas, Pseudobaccharis (Cabrera)
Cuatrecasas, Psila (Philippi) Cuatrecasas, and Tri-
nervatae DC. emend. Cuatrecasas, whereas hemi¬
spherical involucres plus ratio of florets 5:1 are pre¬
sent in section Stephananthus (Lehmann) Heering;
nevertheless, B. retamoides shows notable differ¬
ences (especially in habit, and achene and pappus
morphology), which makes highly improbable any
relationship among any of them.

Baccharis sect. Tridentatae Giuliano, sect. nov.

TYPE: Baccharis tridentata Vahl., Symb. Bot.
3: 98. 1794.

Sectioni Cylindricis affinis planlis xylopodiferis et folds
trinervatis saepe oppositis vel suboppositis, sed capitulis
in cymis corymbiformibus compositis dispositis, capitu-
lorum pistillatomm involucris campanulatis pedicellatis,
floribus pistillatis: staminatis ratione 2:1, el florum pisti-
llatorum setae pappi non accrescentes differt.

Erect subshrubs with xylopodium, clothed by mi¬
nute tufted trichomes. Leaves pseudopetiolate, 3-
veined, margins with few pairs of teeth. Capitula
pedicellate, arranged in compound corymbiform
cymes at the apex of the branches; involucres cam-
panulate; receptacles of the carpellate capitula flat,
of the staminate ones convex, both alveolate. Ratio
of carpellate:staminate florets ca. 2:1. Carpellate
florets with corollas dentate at apex. Achenes te¬
rete, 10-ribbed, glabrous; pappus bristles in 2 se¬
ries, non-accrescent, caducous. Staminate florets
with corollas showing little differentiated limb; style
branches attached, brief; pappus bristles plumose
at apex.

Six species, which grow in two areas: Andes of
Bolivia and northwestern Argentina, and from east¬
ern Brazil to northeastern and central Argentina.

1. Baccharis delloidca Baker, in Martins, FI.

Bras. 6(3): 90. 1882.

2. Baccharis illinita DC., Prodr. 5: 412. 1836.

3. Baccharis illiniloides Malagarriga, Mian. Soc.

Ci. Nat. La Salle 37: 138. 1976.

4. Baccharis isabelae Soria & Zardini, Candollea

46(2): 537. 1991.

5. Baccharis suhopposita DC.. Prodr. 5: 413.

1836.

6. Baccharis tridentata Vahl, Symb. Bot. 3: 98.

1 794. TV PE: “ Baccharis tridentata ; Conyza
cuneifolia: Montevideo,” Commerson s.n. (lec-
totype, designated here, C). | This is the single
original specimen of this taxon deposited at C;
Vahl in the protologue solely stated "habitat in
Brasilia.”]

Baccharis illiniloides and B. isabelae have not
been assigned at the sectional level before; the oth¬
er four species were formerly placed in the large
heterogeneous section Cuneifoliae DC. by classical
authors (Candolle, 1836; Hooker & Arnott, 1841:
Baker. 1882). Recently, B. suhopposita and B. tri¬
dentata were placed by Giuliano (2001) in section
Cylindricae Heering because they present xylopo¬
dium as well as 3-veined opposite or subopposite
leaves, but they do have significant differences
such as: capitula in compound corymbs (vs. foliose
spikes or glomerules), carpellate capitula with in¬
volucres campanulate (vs. cylindrical), the ratio of
carpellate:staminate florets 2:1 (vs. 1:1). and a pap¬
pus slightly accrescent (vs. usually notably accres¬
cent): these differences justify the recognition of
this taxon as an independent section.

New Series oe Baccharis

Baccharis sect. Angustifoliae Baker emend. Cua¬
trecasas

Baccharis ser. Uniflorae Giuliano. ser. nov.
TYPE: Molina uniflora Ruiz & Pavon, Syst.
Veg. 208. 1798 |= Baccharis uniflora (Ruiz &
Pavon) Persoon].

A serie typica differt plantis minutis nidis pilosis ves-
titis et folds trinervis.

Plants clothed by minute tufted trichomes.
Leaves 3-veined. Receptacles of carpellate capitula
without paleae. Ratio of carpellate:staminate florets
ca. 1:1.

Seven species, mainly distributed in the Andean
region of Ecuador, Peru, and Bolivia.

1. Baccharis frigida Kunth, in Humboldt. Bon-

pland & Kunth, Nov. Gen. Sp. 4 | folio]: 38.
1818.

2. Baccharis hutchisonii Cuatrecasas, Caldasia

10: 25. 1967.

3. Baccharis laxiflora Rushy, Bull. New York

Bot. Card. 8(28): 129. 1912.

4. Baccharis libertadensis (S. B. Jones) H. Rob¬

inson. Phytologia 65(1): 37. 1988.

538

Novon

5. Baccharis sternhergiana Steudel, Nomencl.

Hot. (ed. 2) I: 179. 1840. Replaced name:
Baccharis semiserrata DC., Prodr. 5: 419, n°
149. 1836, non Baccharis semiserrata DC..
Prodr. 5: 404, n° 38. 1836.

6. Baccharis triueiira Soria & Zardini. Candollea

56: I 16. 2001.

7. Baccharis uiiiflora (Ruiz & Pavon) Persoon,

Syn. PI. 2: 425. 1807. Molina uniflora Ruiz &
Pavon. Syst. Veg.: 208. 1798. TYPE: Peru. Ju-
nfn: Parma, 13,000 ft., June 1922. Machride
& Featherstone 1051 (neotype, designated
here, US). | The original type material of Ruiz
and Pavon could not be located at MA or any
other herbaria where duplicates of their spec¬
imens are deposited; therefore, a neotype must
be designated. Pile specimen Machride <£*
Featherstone 1051 was selected because it is
representative for the species and was collect¬
ed in one of the original type localities.]

For the following series that includes the lec-
totypified species for Baccharis sect. August i/oliae ,
Baccharis ulicina Hooker & Arnott, the following
combination is proposed:

Baccharis ser. Angustifoliae (Baker) Giuliano,
stat. nov. Basionym: Baccharis sect. Angusti-
foliae Baker, in Martins, FI. Bras. 6(3): 55.
1882; emend. Cuatrecasas, Revista Acad. Co-
lomh. Ci. Exact. 13(49): 88. 1967. I Y PF: Bac¬
charis ulicina Hooker & Arnott. J. Bot. (Hook¬
er) 3: 38. 1841 (lectotype, designated by
Cuatrecasas (1967: 88)).

Plants loosely clothed by small uniseriate tri-
chomes with apieulate or flagellate terminal cells.
Leaves uninervate. Receptacles of carpellate capit-
ula without |)aleae. Ratio of carpellate:staminate
florets ca. 1:1.

Includes 19 North and South American species.

1. Baccharis darwinii Hooker & Arnott, J. Bot.

(Hooker) 3: 39. 1841.

2. Baccharis crosoricola Rzedowski, Bol. Soc.

Bot. Mexico 45: 111. 1983 1 1984].

3. Baccharis gracilis DC., Prodr. 5: 423. 1836.

1. Baccharis hrlrrothalmnidcs Britton, Bull.

Torrey Bot. Club 19: 4. 1892.

5. Baccharis marcctiifolia Bentham. PI. Hartw.:

202. 1845.

6. Baccharis inelanopotainica Spegazzini. Ana-

les Soc. Ci. Argent. 48: 189. 1899.

7. Baccharis niiiltisulcata Baker, in Martins, I I.

Bras. 6(3): 45. 1882.

8. Baccharis nivalis (Weddell) Schultz Bipontinus

ex Philippi, Anales Univ. Chile 87: 705. 1894.

9. Baccharis nrridcnlalis S. F. Blake. Contr.

U.S. Natl. Herb. 22(8): 595. 1924.

10. Baccharis orhignyana Klatt. Abb. Naturf.
ties. Halle 15: 327. 1882.

11. Baccharis pahueri Greenman, Proc. Amer.
Acad. Arts 41(9): 259. 1905.

12. Baccharis pctrophila R. F. Fries, Nov. Acta
Regiae. Soc. Sci. Upsal. ser. 4, 1(1): 81. 1905.

13. Baccharis squarrnsa kunth. in Humboldt.
Bonpland & Kunth, Nov. Gen. Sp. 4 [folio]: 52.
1818.

14. Baccharis laltalensis Johnston, Contr. Cray
Herb. 85: 125. 1929.

15. Baccharis tenella Hooker Arnott, J. Bot.
(Hooker) 3: 42. 1841.

16. Baccharis texana (Torrey & A. Cray) A.
Cray, Mem. Amer. Acad. Arts, n.s., 4: 75.
1849.

I 7. Baccharis idicina Hooker & Arnott, J. Bot.
(Hooker) 3: 38. 1841. TYPE: Argentina.
Woods of Cordoba. Tweedie 1125 (holotype, k
not seen).

18. Baccharis vancssac R. M. Beauchamp.
Phytologia 46(4): 216. 1980.

19. Baccharis wrightii A. Gray, Smithsonian
Contr. know!. 3(5): 101. 1852.

Baccharis sect. Cylindricae Heering
Baccharis ser. Axillares Giuliano, ser. nov.
TYPE: Baccharis axillaris DC., Prodr. 5: 407.
1836.

A seric typica differt foliis saepe sessilihus trinervis vel
penninervis, capitulis in spicis vel racemis foliosis dis-
positis, florum pistillatoruin setis pappi non aecrescenti-
l>us.

Leaves usually sessile, 3-veined or pinnately
veined. Capitula arranged in leafy spikes or ra¬
cemes. Pappus of carpellate florets non-acereseent.

Twenty species, distributed in eastern and south¬
ern Brazil.

1. Baccharis angusticeps Dusen ex Malme,
Kongl. Svenska Vetenskapsakad. Handl. ser.
3. 12(2): 77. 1933.

2. Baccharis aracatuhensis Malagarriga &
Hatschbach, Rodriguesia 28(40): 168. 1976.

3. Baccharis axillaris DC., Prodr. 5: 407. 1836.

TYPE: Brazil. “Sancti-Pauli,” 1833, Sello s.n.
(ex Herbarium of Museum Imperialis Brasi-
liensis n° 481) (holotype, P).

4. Baccharis concinna C. M. Barroso, Rodrigue¬

sia 28(40): 162. 1976.

5. Baccharis cultrata Baker, in Marlins. FI. Bras.

6(3): 93. 1882.

6. Baccharis clliptica (Gardner, London J. Bot. 7:

83. 1848.

7. Baccharis hypcririfnlia Baker, in Martins. FI.

Bras. 6(3): 69. 1882.

8. Baccharis incisa Hooker & Arnott. J. Bot.

(Hooker) 3: 29. 1841.

Volume 15, Number 4
2005

Giuliano

Infragenera in Baccharis

539

9. Baccharis lateralis Baker, in Martins, H. Bras.

6(3): 100. 1882.

10. Baccharis lyinanii G. M. Barroso, Rodrigue-
sia 28(40): 165. 1976.

1 1. Baccharis minutiflora Martins ex Baker, in
Martins, FI. Bras. 6(3): 69. 1882.

12. Baccharis parvidentata Malagarriga. Mem.
Soc. Ci. [Nat. La Salle 37(107): 138. 1976.

13. Baccharis pauciflosculosa DC.. Prodr. 5:
413. 1836.

14. Baccharis perdusenii Giuliano, nom. nov.
Replaced name: Baccharis dusenii G. M. Ba¬
rroso. Rodriguesia 28(40): 166. 1976. nom.
illeg., non B. dusenii 0. Hoffmann, in Dusen.
Wiss. Erg. Schwed. Exp. Magellansl. 3(5):
244. 1900. TYPE: Brazil. Rio de Janeiro: Ita-
tiaia. 1902. Dusen 66 0 (holotype. R not seen).

15. Baccharis polyphylla Gardner, London J.
Bot. 7: 88. 1848.

16. Baccharis pseudovaccinioides Malagarriga,
Contr. Inst. Geobiol. La Salle Canoas 2: 47.
1952.

17. Baccharis selloi Baker, in Martius, FI. Bras.
6(3): 68. 1882.

18. Baccharis serrula Schultz Bipontinus ex
Baker, in Martius, El. Bras. 6(3): 70. 1882.

19. B accharis truncata (Gardner, London .1. Bot.
7: 82. 1848.

20. Baccharis xiphophylla Baker, in Martius. El.
Bras. 6(3): 68. 1882.

For the following series that includes the lec-
totypified species for Baccharis sect. Cylindrical’ ,
Baccharis santiagensis Heering [= B. rufescens
Sprengel], designated by Ariza Espinar (197.3:
180), the following combination is proposed:

Baccharis ser. Cylindricae (Heering) Giuliano.
stat. nov. Basionym: Baccharis sect. Cylindri¬
cae Heering, in Reiehe, Anales Univ. Chile
111: 176. 1902 [1903], TYPE: Baccharis san¬
tiagensis Heering, in Reiehe, Anales Univ.
Chile 111: 176. 1902 [1903] (lectotype, des¬
ignated by Ariza Espinar (1973: 180)).

Leaves pseudopetiolate (rarely petiolate), .3-
veined. Capitula arranged in leafy glomerules,
which usually are arranged in a raceme-like eapi-
tulescence. Pappus of carpellate florets usually ac¬
crescent.

Includes 31 species mainly from Brazil, Para¬
guay, Uruguay, and Argentina; a few species have
Andean distribution.

1. Baccharis brevifolia DC.. Prodr. 5: 409. 1836.

2. Baccharis camporum DC., Prodr. 5: 399.

1836.

3. Baccharis cliilco Kunth, in Humboldt. Bon-

pland Kunth, Nov. Gen. Sp. 4 [folio|: 44.
1818.

4. Baccharis cognata DC., Prodr. 5: 413. 1836.

5. Baccharis nuheliutu Hooker & Arnotl. J. Bot.

(Hooker) 3: 28. 1841.

6. Baccharis glaucescens (Chodal tN Hassler)

Soria & Zardini, Candollea 46: 539. 1991.

7. Baccharis humilis Schultz Bipontinus ex Bak¬

er, in Martius, FI. Bras. 6(3): 92. 1882.

8. Baccharis intermixta Gardner, London .1. Bot.

7: 84. 1848.

9. Baccharis itatiaiae Wawra, Itin. Prine. S. Co-

burgi 2: 28. 1888.

10. Baccharis leptocephala DC.. Prodr. 5: 41.3.
1 836.

11. Baccharis leptophylla DC.. Prodr. 5: 42.3.
1 836.

12. Baccharis longii Govaerts, World Checklist
Seed PI. 2(1): 9. 1996.

13. Baccharis niicrodonla DC.. Prodr. 5: lit).
1 836.

14. Baccharis oreophila Malme, Kongl. Svenska
Vetenskapsakad. Handl. ser. 3, 12(2): 75.
1 933.

15. Baccharis pederscnii Cabrera, Darwiniana
16: 410. 1970.

16. Baccharis pentodonla Malme, Kongl.

Svenska Vetenskapsakad. Handl. 32(5): 52.
1 899.

17. Baccharis pcntziif nlia Schultz Bipontinus ex
Baker, in Martius, FI. Bras. 6(3): 96. 1882.

18. Baccharis petraea Heering, Jahrb. Hamburg.
Wiss. Anst. 21 (Beili. .3): 30. 1904.

19. Baccharis pflanzii Perkins, Bot. Jahrb. Syst.
49: 224. 1913.

20. Baccharis pseudohrevifolia D. J. N. Hind.
Kew Bull. 48: 26.3. 199.3.

2 1 . Baccharis pseudotenuifolia Malagarriga,
Contr. Inst. Geobiol. La Salle Canoas 2: 46.
1 952.

22. Baccharis ramosissinia Gardner. London J.
Bot. 7: 84. 1848.

23. Baccharis retieularia DC.. Prodr. 5: 409.
1836.

24. Baccharis relusa DC.. Prodr. 5: 412. 1836.

25. Baccharis rufescens Sprengel. Syst. Veg. 3:
464. 1826.

Baccharis santiagensis Heering, in Reiehe, Anales Univ.
Chile III: 176. 1902 publ. 1903. TYPE: Chile. Cor¬
dilleras de Santiago, Dt'uz 21 (holotype, SCO).

540

Novon

26. Baeeharis snlzinnimii DC., Prodr. 5: 400.
1 836.

27. Baeeharis schultzii Baker, in Martins, FI.
Bras. 6(3): 7 8. 1882.

28. Baeeharis sessiliflora Vahl. Symb. Bot. 3: 97.
1 794.

29. Baeeharis siilalentata DC., Prodr. 5: 408.
1836.

30. Baeeharis various Gardner, London J. Bot.
7: 84. 1848.

31. Baeeharis vernicosa Hooker & Arnott, J.
Bot. (Hooker) 3: 28. 1841.

Baeeharis sect. Nitidae Cuatrecasas
Baeeharis ser. Ilirsiitae Ginliano, ser. nov. TYPE:
Baeeharis muelleri Baker, in Martins. FI. Bras.
6(3): 61. 1882.

A serie typiea differ! pi 1 is hirsutis et capitulis in cymis
eorymbiformibus term inali bus dispositis.

Plants hirsute, clothed by long erect uniseriate
trichomes and minute glandular and noil-glandular
lulled trichomes; the vestiture becomes much
sparser with age. Capitula in terminal corymbiform
cymes.

Includes a single species, endemic to the state
of Santa Catarina, Brazil.

1. Baeeharis muelleri Baker, in Martins. FI.
Bras. 6(3): 61. 1882. TYPE: Brazil. Santa Ca¬
tarina: Barra do Itajaf, Mueller .755 (leetotype,
designated by Barroso (1976: 99), k).

Baeeharis ser. Laiiuginosae Ginliano, ser. nov.
TYPE: Baeeharis calvescens DC., Prodr. 5:
413. 1836.

A serie typiea diflert plantis albo-lanuginosis pi I is uni-
seriatis metis et crispis dense vestitis.

Plants lanuginous, clothed by long uniseriate tri¬
chomes accompanied by minute glandular and non-
glandular tufted trichomes on upper leaf surfaces,
and by tufts of short crisped trichomes on lower
leaf surfaces; the pubescence becomes much spars¬
er with age. Leaves eoncolorous or discolorous. Ca¬
pitula arranged in axillary corymbiform cymes.

Four species, distributed in eastern and southern
Brazil, eastern Paraguay, and the province of Mi-
siones, Argentina.

1. Baeeharis calvescens DC., Prodr. 5: 413.

1836. TYPE: Brazil. Minas Gerais: Marianna,
Vauthier 332 (leetotype, designated by Barroso
(1976: 119), G-DC).

2. Baeeharis claeagnoides Steudel ex Baker, in

Martius, FI. Bras. 6(3): 53. 1882.

3. Baeeharis myrieifolia DC.. Prodr. 5: 405.

1836.

4. Baeeharis seniiserrata DC., Prodr. 5: 404.

1836.

Baeeharis semiserrata presents vestiture sparser
than that of B. calvescens and B. elaeagmhdes,
whereas in B. myrieifolia the pubescence is only
restricted to young organs.

For the following series that includes the lec-
totypified species for Baeeharis sect. Nitidae, Mo¬
lina nitida liuiz & Pavdn, the following combina¬
tion is proposed:

Baeeharis ser. INilidnc (Cuatrecasas) Ginliano.
stat. nov. Basionym: Baeeharis sect. Nitidae
Cuatrecasas. Revista Acad. Colomb. Ci. Exact.
13(49): 70. 1967. TYPE: Molina nitida Ruiz
& Pavon, Syst. Veg.: 204. 1798 [= Baeeharis
nitida (Ruiz & Pavdn) Persoon).

Plants clothed by minute tufted trichomes.
Leaves eoncolorous. Capitula usually arranged in
axillary corymbs.

Includes six species from the Andes, the Gua-
yana Shield. Brazil, and Paraguay.

1. Baeeharis anianihayensis Zardini & Soria,

Candollea 46: 535. 1991.

2. Baeeharis dentata (Vellozo) G. M. Barroso,

Bodriguesia 28(40): 123. 1976.

3. Baeeharis nitida (Ruiz & Pavon) Persoon, Syn.

PL 2: 425. 1807. Molina nitida Ruiz & Pavdn,
Syst. Veg.: 204. 1798. TYPE: Peru. “Lima.
Clasis 19. Baeeharis axillaris. F. P. e. 1.65. c.d.
ex Chinehao" (leetotype, designated by Cabre¬
ra (1960: 199), MA).

4. Baeeharis rivularis Gardner, London J. Bot. 7:

83. 1848.

5. Baeeharis sehomhurgkii Baker, FI. Bras. 6(3):

89. 1882.

6. Baeeharis siugidaris (Vellozo) G. M. Barroso.

Bodriguesia 28(40): 96. 1976.

Acknowledgments. I thank Susana Freire for
helpful comments, and John Pruski and John Tem¬
ple for valuable reviews of the manuscript. This
study was supported by a fellowship (“Beca do For-
macidn de Postgrado") awarded by the Consejo Na¬
tional tie Investigaciones Cientfficas y Tdcnieas tie
la Republiea Argentina (CONICET).

Literature Cited

Ariza Espinar, L. 1973. Las especies de Baeeharis (Gom-
positae) de Argentina Central. Bol. Acad. Nac. Ci. 50:

1 75-305.

Baker. J. G. 1882. Compositae: Baeeharis. In: C. F. Mar¬
tius (editor), FI. Bras. 6(3): 37—100. Mtlnchen, Wien,
Leipzig.

Barroso. C. M. 1976. Compositae, Subtribo Baceharidinae
Hoffman. Estudo das espdeies ocorrentes no Brasil. Bo-
drigudsia 28(40): 3-273.

Volume 15, Number 4
2005

Giuliano

Infragenera in Baccharis

541

Cabrera, A. L. 1960. Notas sobre tipos de Compuestas
sudamericanas en Herl)arios europeos. Ill, bos tipos de
Ruiz y Pavon. Bob Sue. Argent. Bot. 8: 195—215.

Candolle, A. P. de. 1836. Cornpositae: Baccharis. In:
Prodr. 5: 398—429. Treuttel et Wiirtz, Paris.

Cuatrecasas, J. 1967. Revision de las especies colombia-
nas del g^nero Baccharis. Revista Acad. Colomb. Ci.
Exact. 13(49): 5-102.

Giuliano, I). A. 2001. Clasificacidn inlragenerica de las
especies argentinas de Baccharis (Asteraceae, Aste-
reae). Darwiniana 39: 131—154.

Heering, W. C. A. 1904. Die Baccharis- Arten des Ham¬
burger Herbars. Jahrb. Hamburg. Wiss. Anst. 21 (Beih.
3): 1-16.

Hooker, Vi. J. & G. A. W. Arnott. 1841. Cornpositae: Bac¬
charis. In: Contributions towards a Flora of South Amer¬
ica and ihe islands of the Pacific. J. Bot. (Hooker) 3:
21-43.

Persoon, C. H. 1807. Syngenesia Superflua: Baccharis. In:
Synopsis Plantarum 2: 423—425. Parisiis lutetiorum el
Tubingae.

Taxonomic Novelties in Astragalus (Leguminosae)
for South America

Edith Gomez-Sosa

Instituto <le Bolanica Darwinion, C.C. 22, B1642HYI) San Isidro, Argentina.
egomez-sosa@darwin.edu. ar

A US IK ACT. A new species in Astragalus , endemic
lor Peru, is described for Leguminosae, Galegeae.
Astragalus sagasteguii may be compared with the
related A. dillinghami Maebride in Peru and A.
diminutivus (Philippi) Gomez-Sosa in Bolivia and
Argentina. The new combination Astragalus dimi¬
nutivus is made, transferring the species from the
genus Phaca. In addition. Astragalus deminutivus
I. M. Johnston newly synonymizes to A. diminuti¬
vus. Further, Astragalus dielsii Maebride is recog¬
nized as a variety to A. diminutivus. The geograph¬
ical distribution of Astragalus diminutivus var.
dielsii extends from Peru into Bolivia and Argen¬
tina. Morphological description and comparison, il¬
lustration. as well as habitat and geographical dis¬
tribution are provided.

Rksumkn. Una nueva especie de Astragalus, en-
demica del Peru, es deseripta para Leguminosae,
Galegeae. Astragalus sagastegidi, puede ser eom-
parada eon A. dillinghami Maebride, en Peru, y A.
diminutivus (Philippi) Gomez-Sosa en Bolivia y Ar¬
gentina. Adcnuis A. dielsii Maebride es transferida
como una variedad de A. diminutivus. La distribu-
cibn geogralica de Astragalus diminutivus var. diel¬
sii se extiende desde Peru a Bolivia y Argentina.
Se ineluyen deseripeiones, tabla de relaciones,
ilustraeiones, datos sobre habitat y distribueion
geogralica.

Key words: Argentina, Astragalus, Bolivia,
Chile, Galegeae. Leguminosae, Peru.

The genus Astragalus L. includes ea. 2500 spe¬
cies (Podlech, 1986) with a worldwide distribution,
except for Australia and New Zealand. There are
about I 10 species endemic to South America; they
are found in Ecuador, Peru, Bolivia, Chile, Argen¬
tina, and Uruguay (G6mez-Sosa, 1979; Davina &
Gomez-Sosa. 1993). There is only one species of
the genus, A. distinens Maeloskie. known from Uru¬
guay (Gomez-Sosa, 2003).

There is a group of South American Astragalus
species of tiny, matted or tufted herbs. In many
instances, these laxa have unknown fruit or have

Novon IS: 542-547. Pi bushed on 12 Put

been incompletely collected; consequently, it is di-
fieult to delimit species. Inside this group are the
taxa currently presented in this manuscript.

Among Astragalus specimens received as a gift
for determination from the New York Botanical Gar¬
den, the following new species from Peru was
found.

1. Astragalus sagasteguii Gomez-Sosa. sp. nov.

TYPE: Peru. Cajabamba, Cajamarca, jalca,
3800 m, 17 Nov. 1983, A. Sagdstegui Alba et
al. 1 1181 (holotype, SI; isotypes, F, MO, NY).
Figure I.

Habitus prostratus-pulvinatus, pubescens; stipulis am-
plexieaulibus, pubescentibus; foliola 15 ad 17. elliptico-
cuneata, apiee retuso; 1 (ail 2) (lores parvi, calyx 4 mm;
vexillum ultra unguem obovatum 7—8 mm longum: legu-
inen 0.3 X 0.2 cm, subglobosum, subbiloculare, cum mar-
gine dorsale inHexo I mm lain et cum margine membra-
naceo inconspicuo 0.5 mm lato solum hasi; semina 1 ad

2. reniformia.

Depressed-pulvinate herb, 4—16 cm diam., acau-
lescent, the stoutly branched eaudex 4—8 cm, stip¬
ules amplexieaul, membranous, densely sericeous-
villous. Leaves to 1.4 cm long; leaflets 15 to 17,
opposite, decrescent distally, folded, in outline el-
liptie-euneate, retuse, subsessile, 2.5—3 X (I— )1.5
mm, cinereous-villous trichomes 0.5—1 mm long.
Bracts subulate 2.5 X I mm; bracteoles 2 X 0.5
mm, pubescent, ciliate; flowers purple 1 (to 2), sub-
sessile, 8—9 mm long; calyx 4 mm, villous, tube 2
mm long, teeth subulate, 2 mm; banner 8 X 6 mm,
obovate; wings 5 X 1.5 mm, the claws included;
keel 4 mm; ovary substipitate, scattered pilose,
style glal irons, incurved at apex, the stigma capi¬
tate; ovules 2. Legume evanescent, subglobose, 0.3
X 0.2 cm, truncate at base, apex rounded, valves
membranaceous, indexed below middle of pod, with
an incipient septum to 0.5 mm only at base, ob¬
solete in the apical half of the dorsal suture, com¬
monly concealed by petals, bract, and mature stip¬
ule; seed I or 2. reniform. brown, generally mottled.

Etymology. This species is dedicated to the
type collector, Abundio Sagastegui Alba, professor

EM HE 8 2005.

Volume 15, Number 4
2005

Gomez-Sosa

Astragalus for South America

543

Figure 1. Astragalus sagasteguii Gomez-Sosa. — A. Plan! habit. — B. Apex of branch with flower. — C. Leaflets, below
and above. — I). Calyx and internal pubescence of the teeth. — E. Banner. — F. Wing. — G. Keel. — 11. Androecium.
— I. Gynoecium. — J. Ovary, longitudinal section. — K. Fruit, lateral view. — L. Emit, longitudinal section, with an
incipient septum only at the base. — M. Seed. Based on Sagdstegui et al. 1 1 Ifil (holotype, SI).

544

Novon

Table 1 . Comparison

of A. sagasteguii, A. diminutivu.

s var. diminutivus , and A. dillingh

'ami

A. sagasteguii

A. diminutivus var.
diminutivus

A. dillinghami

Leaves

Length (cm)

to 1.4

1-2

1.5-2

Leaflets

Number

15 to 17

13 to 17(to 19)

9 to 13

Size (mm)

2.5-3 x (]-)1.5

1.5-2 X 0.6-1

4X2

Shape

elliptio-ouneate

elliptic to subcircular

ovate-lanceolate

Apex

retuse

obtuse to slightly retuse

acute

Pubescence

cinereous-villous on both
surfaces

glabrous above/villous
below

silvery-villous on both
surfaces

Trichome length (mm)

0.5-1

0.5-1

1 .5-2.5

Flowers

Number

1 (to 2)

1 (to 2)

4 to 8

Length (mm)

8-9

7-9

8-10

Calyx length (mm)

4

3-4

7

Legume

subbiloeular septum to

0.5 mm only at base

unilocular

unknown

and director of the Natural History Museum (HAO)
at Antenor Onego University, Trujillo, Peru. He is
an enthusiastic collector and student of the Peru¬
vian flora.

Habitat and distribution. Astragalus sagaste¬
guii occurs in "jalca” in the Andean mountains of
Peru, which refers to die ecological zone along the
Andean divide between the drier Paramo to the
north and the wetter Puna to the south. Found
above 3300 m. the jalca vegetation is predomi¬
nantly grassland, and the predominant grass is
"iclui" \Stipa ichu (Ruiz & Pavdn) Kunth], Native
fauna include frogs, lizards, birds, small mammals,
and deer.

Climate. The climate of the this new species is
typical of Andean regions near the equator: cool
and humid, with a distinct rainy season. Tempera¬
tures are relatively constant throughout the year,
and seldom dip below freezing even at the highest
mountain elevations. Conditions are often windy,
especially at higher elevations. The climate is char¬
acterized by a distinct rainy season (October to
April) followed bv a dry period (May to September).

The genus Astragalus is represented by 23 spe¬
cies (Zarueehi, 1993) in the flora of Peru. The new
species A. sagasteguii is in close relationship with
A. dillingliami J. F. Macbride “depressed, com¬
pact,. . . (lowers crowded or few in the axils, about
I cm long. . ."(Macbride. 1943: 396). Moreover, A.
dillinghami is distinguished by its 4 to 8 flowers,
leaflets ovate-lanceolate with acute apex, and a sil¬

very-villous pubescence with trichomes 1.5— 2.5
mm long.

In addition. Astragalus sagasteguii appears to be
related to A. diminutivus (Philippi) Gdmez-Sosa
from Peru, Bolivia, and Argentina, which it resem¬
bles by having a single flower that is 7-9 mm long.
In contrast A diminutivus has smaller leaflets that
are sericeous-pubescent, especially on the uniloc¬
ular legume. (See Table 1 for a comparision of these
three species.)

A critical examination of the type collections of
Phaca diminutiva Philippi from northern Chile, l.s-
tragalus deminutivus I. M. Johnston from southern
Bolivia, and A. dielsii .1. F. Macbride from Lima, in
northern Peru, reveals that the following new com¬
bination should be recognized.

2. Astragalus diinimitivus (Philippi) Gdmez-
Sosa, comb. nov. Basionym: Phaca diminutiva
Philippi, Anales Mus. Nac. Santiago de Chile,
2° Seccidn. Cat. PI. Itin. Tarapaca 14. 1891.
TYPE: Chile, “inter Aguas Calientes et Socai-
re, R. A. Philippi s. n, (holotype, SCO; isotype,
SI). Figure 2.

Astragalus deminutivus I. M. Johnston, J. Arnold Arbor.
28: 406. 1947. Syn. nov. TYPE: Bolivia. Tarija: Puna
de Patanea, 3700 m, 7 Jan. 1901. K. A. Fiebrig 2608
(holotype, (41; isotypes, K, P, SI).

Because of the very short original diagnosis of
this taxon (Philippi, 1891) a more complete de¬
scription of the type variety is given here.

Volume 15, Number 4
2005

Gomez-Sosa

Astragalus for South America

545

L

Figure 2. Astragalus diminutivus (Philippi) Gdmez-Sosa var. diminutivus.- — A. Plant habit. — B. Apex of branch with
flowers. — C. Leaflets, belowr and above. — I). Calyx and internal pubescence of the teeth. — E. Banner. — F. Wing.
— C. Keel. — H. Androecium. — I. Gynoecium. — J. Fruit, longitudinal section. — K. Seed. Astragalus diminutivus var.
dielsii (J. F. Macbride) Gdmez-Sosa. — L. Leaflets, below and above. A— D based on ./. //. Hunziker 10503: E— K. Macbride
& Featherstone 2479 (isotype. SI); L. V. Cabezas 60.

546

Novon

Astragalus diminutivals (Philippi) Gomez-Sosa
var. diminulivus

Herb eespitose, 2—6 cm diam., green-cinereous,
caudex 1 — 2( — 3) cm, with membranous stipules im¬
bricate, white-pubescent, 2.5— 3.5 mm long, heaves
1—2 cm long, petiole 0.2— 0.5 mm long; leaflets 13
to 1 7(to 19), elliptic to subcircular, 1.5—2 X 0.6—1
mm, obtuse to slightly refuse at apex, thickened,
folded, glabrous above, villous-pubescent below,
covered with trichomes 0.5—1 mm long. Bracts tri¬
angular-ovate, 1.5— 3(— 4.5) mm long; bracteoles the
same length, filiform, both pubescent anil eiliate;
flowers I (to 2) erect, sessile, blue to light blue; ca¬
lyx pubescent, tube campanulate, 2-2.5 mm long,
teeth subulate, 1 — 1.5 mm long; banner 6—8 X (2.5—
)3— 4.5 mm. obovate to widely obovate, emarginate
at the apex, basally narrowed to claw; wings 5— 6(—
8) X 1—1.5 mm, rectangular, obtuse; keel 4.5—5
mm long; ovary shortly stipitate, pubescent, style
glabrous, laterally compressed and indexed at apex,
stigma minutely-capitate; ovules 2(to 4). Legume
membranaceous, evanescent, subglobose, 0.2—0.25
X 0.2—0.25 cm, unilocular, covered bv remains of
bracts and perianth; seed I, subglobose, sometimes
mottled, hilum apical, 2-2.5 X 2-2.5 mm.

Distribution and habitat. A regional endemic
in western South America, cited for Peru and re¬
ported now with a southern range extension to Bo¬
livia, Chile, and northwest Argentina, inhabiting
higher elevations 3600—4300 m. Collector’s notes
(K. Ehrich 75) indicate it occurs on sandy or clayey
limestone.

Climate. This region is in the biogeographic
province of Puna (Cabrera & Will ink, 1973) be¬
tween 15° and 27° S, with the Andean mountains
to the west and the “Cordillera Real’’ mountain
range to the cast at elevations of 3200 to 4000 m.
Moreover, the Puna is in the Neotropical subregion
(Crisci et al.. 2000). The climate of this region is
dry and cold, with a large contrast in temperature
and rainfall during the year. The principal rainfall
occurs only in summer, varying from approximately
700 to 50 mm, decreasing along gradients from
north to south and from east to west.

Additional specimens examined. BOLIVIA. Tarija:

I .ago Taxara, R. Ehrich 75 (LPB, SI). Potosi: Porco, Uyu-
ui. E. \splund 3137 & 3138 (UPS). La Paz: Pacajes,
Oral. Campero, E. Asplund 2780 (UPS); Sur Chichas, Am¬
elia. E. Asplund 3025 (UPS). ARGENTINA. Jujuy: Hu-
rnaliuaea. Lag. Tres Cruces, /*. Claren 11657 (SI, STOCK);
Coehinoca, E Cahezas 60 (SI); Abra Pampa, N Mina
Aguilar, E. Ancibor & R. Mujiea 18 (BAA); Yavi, Cerro
Negro. T. Meyer 31162 (LIE. SI); arriha <le Rincon de
Cajas. II. Sleumer 3618 (LIE, SI). Salta: Caldera, suhida

al Nevado del Castillo, Ires Lagunas, II. Sleumer el id.
2981 (LIE, SI).

2a. Astragalus diminutivus (Philippi) Gomez-
Sosa var. dielsii (J. F. Macbride) Gbmez-Sosa,
stat. nov. Basionym: Astragalus dielsii .1. F.
Macbride., Publ. Field Mus. Nat. Hist., But.
Ser. II: 24. 1931. TYPE: Peru. Aneash, Ya-
nashallas, Lstacidn 35 mi. W of Huallanea, 2
Oct. 1922. "loose stony slopes, light blue f Is.,
about 16,000 It., the parts spreading,” ./. F.
Macbride & W. Featherstone 2479 (holotype,
FM; isotype, SI). Figure 21..

Astragalus diminutivus var. dielsii. with leaflets
pubescent above as well as below is thereby distin¬
guished from A. diminutivus var. diminutivus.

Distribution and habitat. Found in western
Peru, Bolivia, and Argentina, in high mountains al
elevations ol 3600—4400 m.

Additional specimens examined. BOLIN I A. Laguna de
Polos f, D'Orbigny 1432 (P). ARGENTINA. Jujuy: Cochi-
nora, Abra de Rachaite, M. Mulgura et al. 1311 (SI);
Casubindo, IV. Deginani et al. 520 (SI); Humahuaea, Mina
Aguilar, arriba del Molino, //. Sleumer 3428 (LIE, SI). J.
II. Ilunziker et at. 10503 (SI): Tres Cruces. A. Soriano 653
(SI); Susques, al pie del cerro Tuzle, A. Cabrera 8644 (SI),
1. Cabrera 8748 (LP, SI); Abra Chorrillos, A. Cabrera et
al. 31774 (SI); Rinconada, Ruta provincial 746, 1.4 km de
Mina Pirquitas cam i no a Coranzulf, 0. Morrone et at. 2589
(SI).

The name Phaca diminutiva Philippi was cited
by I. M. Johnston (1947: 407) as an "unplaced spe¬
cies." because he had not seen the type specimen
from Chile. In the same paper Johnston published
Astragalus deminutivus, a new' species for Bolivia
and Argentina, with similar vegetative and floral
characteristics to the ones that encompass F. di¬
minutiva. fhe legume was described by Johnston
(1947: 406) as “subglobosis ea. 2 mm longis et 1.5
mm crassis subtus introflexis” or as a subbiloeular
legume. A critical examination of the Astragalus
deminutivus type collections at FM, GH, K. and P
in addition to cited specimens of E. Asplund (UPS)
reveals the legume is without inflexion ol the su¬
ture, and thus the fruit is unilocular; these collec¬
tions otherwise share the same characteristics as A.
diminutivus. The Philippi type locality o {Phaca di¬
minutiva is noted between Aguas Calientes to So-
caire, in northern Chile, east of the Salar of Ata¬
cama, at 5030—5070 m near the political boundary
with Argentina, in Provinces of Jujuy and Salta,
and in the vicinity of the Tropic of Capricorn, w ith¬
in the same distributional area as A. deminutivus.

Acknowledgments. 1 thank the directors and cu¬
rators of FM. GH. K. LP. LIE. M. NY. P. SGO. and

Volume 15, Number 4
2005

Gomez-Sosa

Astragalus for South America

547

UPS for their loans of the cited material and also
for the cordial attention during my visit to each of
the herbaria. I thank the artist, Francisco Rojas, lor
the drawing. I am also grateful to Jan Saunders for
helping me with the revision of the English version.
My special gratitude to Victoria C. Hollowed and
an anonymous reviewer for offering careful sugges¬
tions toward its revision. Herbarium visits were
supported by a grant from CONICET (National
Council for Scientific and Technological Research)
of Argentina and aid from American and European
institutions.

I .iterature Cited

Cabrera. A. L. & A. Villink. 1973. Biogeografia de Ame¬
rica Latina. Serie de Biologia, Monografla 13. Organi¬
zation de los Estados Americanos (O.E.A.), Washing¬
ton. b.c.

Crisei, J. V.. E. Katinas & P. Posadas. 2000. Introduct ion

a la Teona y Praetica de la Hiogeograh'a Historica. S<>-
eiedad Argentina de Botanica, Buenos Aires.

Davina, J. R. & E. Gdmez-Sosa. 1993. Cariotipo de siete
especies del genero Astragalus (Leguminosae) de la Ar¬
gentina. Bob Soc. Argent. Bot. 29: 197-201.

Gdmez-Sosa, E. 1979. Las especies sudamericanas del ge-
nero Astragalus. I. Las especies patagonicas argenlinas.
Darwiniana 22: 313—376.

- . 2003. El genero Astragalus (Leguminosae) en la

Repilblica Oriental del Uruguay. Hiekenia 3: 161-162.

Johnston, I. M. 1947. Astragalus in Argentina, Bolivia,
and Ghi le. J. Arnold Arbor. 23: 336—409.

Macbride, J. F. 1943. Flora of Peru. Publ. Field Mus. Mat.
Hist.. Bot. Ser. 13(111): 292-293.

Philippi, R. A. 1891. Catalogo Plantas Itinerant) Tarapaca
14. Anales Mus. Nac. Santiago tie Chile.

Podlech, I). 1986. Taxonomic anti phytogeograph ical
problems in Astragalus ol Old World and Southwest
Asia. Proc. Roy. Soc. Edinburgh 89- B: 37-43.

Zarucchi, J. L. 1993. Astragalus. Pp. 430-451 in: L. Bra¬
ke & J. L. Zarucchi (editors). Catalogue of the Flower¬
ing Plants & Gymnosperms til Peru. Monogr. Syst. Bot.
Missouri Bot. Gard. 45.

Two New Species of Mikania (Asteraceae, Tribe Eupatorieae)

from Tropical America

Walter C. Holmes and Darrell S. Vodopieh
Department ol Biology, Baylor University, Waco, Texas 76798-7388, U.S.A.
walter_holmes@baylor.edu

ABSTRACT. Mikania pooleana, known only from
Oaxaca, Mexico, is similar to M. toruluzii of Mexico
and Central America, but distinguished by its lan¬
ceolate leaves and hemispherical glomerules. The
latter species has ovate to broadly ovate leaves and
spherical glomerules. Mikania saxicola ol Minas
Gerais, Brazil, is characterized by its dense spike¬
like capitulescence, small heads about 4.5 mm
long, and cypselae with forked trichomes. It has
affinity to M. microlepis, also of Brazil, which is
distinguished by its dense fulvous-tomentose na¬
ture, larger heads, and lack of forked trichomes.

Key words: Asteraceae, Eupatorieae, Mikania ,
tropical America.

IMikania pooleana W. C. Holmes & Vodopieh. sp.
nov. TYPE: Mexico. Oaxaca: ca. 21 mi. S of
Valle National. Hwy. 175 (Tuxtepec— Oaxaca),
infrequent vine, with tree ferns, 13 Mar. 1680.
./. M. Poole 2247 & ./. A. McDonald (holotype,
TEX). Figure I.

Haec species M. toruluzii sirnilis sed in foliis lanceolatis
differt.

Twining vines. Stems terete, striate, glabrate,
solid. Leaves lanceolate, 7.5—12 X 1.9— 3.6 cm;
apex caudate to caudate acuminate, margins entire
in the basal half, remotely serrate-denticulate in the
upper half, the teeth 0.7-2 cm distant, base acute
to acute-acuminate, surfaces glabrate, venation In¬
nervate with a conspicuous pair of lateral nerves
separating near 1 cm from the base of the blade,
these accompanied by another pair of obscure
nerves separating within 5—8 mm of the base of the
blade, tertiary nerves obscure, disposed in a slight¬
ly antrorse-orientated transverse pattern: petioles
2-2.5 cm long, glabrous. Capitulescence an elon¬
gate pyramidal panicle, 19 X 7.5 cm; branehlets
glabrate at base, puberulent above; bracts lanceo¬
late to linear, much reduced in size above; capilula
ca. t nun long, disposed in somewhat dense hemi¬
spherical glomerules 2-3 cm diam., ultimately the
beads in sessile cluster ol threes. Subinvolucral
bracts linear, 1-2 mm long, puberulent. Phyllaries

4, elliptic-oblong, ca. 3.5 mm long, the base fused
for about 1 mm. surfaces glabrous, apex rounded
to obtuse, puberulent. Florets 4. Corollas ca. 6 mm
long, white, tube 2-3 mm long, slightly bulbous at
base, gradually expanded into the funnelform
throat, 2—2.5 nun long, teeth debate, ca. 0.8 mm
long, sparingly glandular. Cypselae tawny, ca. 2 nun
long, glabrous, the faces reticulated. Pappus bris¬
tles white, 21 to 33, ca. 4.5 mm long, margins scab-
rid. apex slightly thickened.

In Holmes (1990). this new species keys to the
vicinity ol Mikania tonduzii B. 1.. Robinson and M.
globosa Coulter, which appear to be its nearest rel¬
atives. These two species have ovate to broadly
ovate leaves and heads disposed in very dense
spherical glomerules. In contrast, Mikania poo¬
leana has lanceolate leaves and hemispherical
glomerules. Additionally, M. globosa has phyllaries
that completely envelop the adjoining floret it sub¬
tends. The ultimate disposition of the capitula in
sessile clusters ol threes is unusual for Mikania
with pyramidal eapitulescences. This arrangement
is typical ol the Mikania parviflora (Aublet) Karsten
complex (Robinson, 1922a). Additional character¬
istics of this complex are corymbose capita lesc-
ences and densely hirsute style appendages and ba¬
ses. Mikania pooleana has none of these traits,
which suggests that this head arrangement arose
independently in other Mikania and does not in¬
dicate relationship.

Using the classification proposed by Holmes
(1996), Mikania pooleana is in Mikania sect. Mi¬
kania. This is determined from its subinvolucral
bracts that are borne at the very base of the invo¬
lucres and from the maturation pattern of the ca¬
pitula ol the hemispherical glomerules from the
center to the periphery.

Mikania pooleana is known only from the type
material.

We are privileged to name this species to com¬
memorate the collector, Jackie M. Boole. Boole, an
employee of Texas Parks and Wildlife Department,
has been a pre-eminent botanist concerned with
preserving the native flora of Texas.

Novon 15: 548-551. Published on 12 December 2005.

Volume 15, Number 4
2005

Holmes & Vodopich
Mikania from Tropical America

549

UNIVERSITY OF

HERBARIUM

liiif

Oaxaca: Ca. 21 mi 5 of Valle Nacional,

Hwy 175 (Tuxtepec-Oaxaca) ; edge of cleared
forest. Moist, huniusy soil. With tree
ferns. Infrequent vine. Discs white.

13 March 1980

Jackie M. Poole 2247 with J. A. McDonald

Figure 1. Photograph of the holotype of Mikania pooleana W. C. Holmes & Vodopich.

Mikania saxicola Schultz-Bip. ex W. C. Holmes
& Vodopich, sp. nov. TYPE: Brazil. Minas
Gerais, Serra do Chapada, June 1827, L. Rie¬
del s.n. (holotype, NY). Figure 2.

Haec species M. microlepis similis sed in planta glabra

differt.

Semiwoody vine. Stems terete, striate-costate al¬
ter drying, glabrous. Leaves deltate, 3.5—6 X 2.4—

550

Novon

h *Co/

JaJ W C. . Zt>0 I

Vw H <Vw-a ~ (Ly *v. C

H , Ar*\ P ti

A. PMAe^b: 3Kc;ieanunH J1aHrcaop4>a b Epa3nnHc,
1824—1829 rr.

m h a.

Trite H. Krascheninnikov.

Herbarium Horti Petropolitani.

'.(kisL.

a /O > ,

. . :.. ...

L Riedel: Her Brasiliomo 1824—29.

Figure 2. Photograph of the holotype of Mikania saxicola Schultz-Hip. ex W. C. Holmes & Vodopich.

5.5 cm, apex acute to a rounded point, margins
entire, occasionally with two bluntish-rounded bas¬
al lobes to ea. 3 X 5 mm, base euneate, palmately
8- to 5-nerved from insertion of the petioles; sur¬
faces glabrous, reticulate, the lower surface densely

glandular; petioles 0.8— 1.7 cm long, glabrate. Cap-
itulescence paniculate, dense, 7 X 4.5 cm; branch-
lets irregularly angled to subterete, puberulent,
glandular; bracts at base of eapitulescence similar
to leaves but reduced in size, those at summit lin-

Volume 15. Number 4
2005

Holmes & Vodopich
Mikania from Tropical America

551

ear. Capitula ca. 4.5 mm long, ultimately disposed
in spikelike raeemes ca. 5 mm long, usually num¬
bering 4 to 6 capitula. Subinvolucral bracts lance-
elliptic, ca. 1.5 mm long, puberulent, glandular,
apex acute. Phyllaries obovate, 1.8—2 mm long, gla-
brate to puberulent, sparingly glandular, margins in
the distal hall finely serrate-ciliate. Corollas ca. 2.2
mm long; tube ca. 0.8 mm long, throat funnelfonn
to semicampanulate, ca. 1 mm long, lightly glan¬
dular; teeth deltate-ovate, ca. 0.2 mm long, glan¬
dular. Cypselae ca. 1.5 mm long, puberulent,
densely so at the summit, the trichomes sometimes
forked or branched at the apex. Pappus bristles
white. 1-1.4 mm long. 15 to 20. margins scabrid.
apex occasionally slightly thickened.

Mikania saxicola is characterized by its con¬
gested capituleseenee with the capitula borne in
short spikelike racemes, heads ca. 4.5 mm long,
which is small for the genus, and presence of
branched or forked trichomes on the cypselae. The
latter trait is very unusual in the genus and in the
Eupatorieae, being reported by Holmes and Pruski
(2000) as dendroid pubescence in M. wurdackii
Pruski & W. C. Holmes. Similar trichomes are also
known from M. citriodora W. C. Holmes (Holmes,
1991) and from M. grazielae R. M. King & H. Rob¬
inson (King & Robinson, 1980), both of Brazil. In
these two species, the forked trichomes appear to
represent an additional and modified pappus, since
they are borne primarily near the summit of the
cypselae. However, the forked trichomes of M. sax¬
icola occur on all parts of the cypselae.

The disposition of capitula into short spike-like
raeemes indicates that the species belongs to Mi¬
kania sect. Slim mikania W. C. Holmes (Holmes,
1996). In Barrosos (1959) treatment of the Mikania
of Brazil. M. saxicola could be included in section
Globosae B. L Robinson (Robinson, 1922b) be¬
cause of its densely congested capituleseenee, or,
with equal facility, in section Spicato-racemosae
Baker (Baker. 1876), because of disposition of the
heads into short spike- 1 ike raeemes. I his ambigu¬
ity, which is responsible for Barrosos classification

not gaining general acceptance, is discussed in
I lolmes ( 1 996).

Mikania saxicola appears similar to M. microle-
pis, also of Brazil. Mikania microlepis is character¬
ized by its dense fulvous-tomentose nature, larger
heads, and lack of forked trichomes.

Mikania saxicola is known only from the type
specimen. The holotype was a gift from L to !N\
presented in about 1990 (John Pruski, pers.
comm.). It is not known if duplicates exist at L. I he
type specimen was annotated as '"Mikania saxicola
Sch.-Bip. in h. Riedel,” by II. Krascheninnikov, a
name never validated through publication.

Acknowledgments. We thank the curators of the
herbaria for their loans of specimens that made this
study possible. We are also indebted to Billie L.
Turner for bringing to our attention that Poole 2247
(holotype of Mikania pooleana) was an undescribed
species. Also. John Pruski provided information
about the type specimen of Mikania saxicola.

Literature Cited

Baker, J. G. 1876. Compositae II. Eupatorieae. In (7 Mar-
tius. LI. Bras. 6(2): 181-374, pi. 51-102.

Barroso, G. VI. 1958 [1959]. Mikaniae do Brasil. Anp
Jard. But. Rio de Janeiro 16: 239—333, pi. 1—31. foto
1-57.

Holmes, W. C. 1990. The genus Mikania (Compositae—
Eupatorieae) in Mexico. Sida, Bot. Misc. 5: iv + 45.
Botanical Research Institute of Texas, fort Worth.
- . 1991. Studies in Mikania (Compositae: Eupato¬
rieae)— XVII: Two new species from Minas Gerais, Bra¬
zil. Phytologia 70: 47-51.

- . 1996. A proposed sectional classification for Mi¬
kania (Eupatorieae). Pp. 621—626 in I). J. N. Hind (ed¬
itor), Compositae Systematics. Proceedings of the Inter¬
national Compositae Conference, Kew, 1994. Vol. I.
Royal Botanic Gardens, Kew.

- & J. F. Pruski. 2000. Mew species of Mikania

(Compositae: Eupatorieae) from Ecuador and Peru. Syst.
Bot. 25: 571—576.

King, R. M. & H. Robinson. 1980. Studies in the Eupa¬
torieae (Asteraeeae). CLXXXVIII. New species ol Mi¬
kania from Brasil. Phytologia 45: 124—141.

Robinson, B. I.. 1922a. Records preliminary to a general
treatment of the Eupatorieae, I. Contr. Gray Herb. n. s.
64: 1-21.

- . 1922b. The mikanias of northern and western

South America. Contr. Gray Herb. n. s. 64: 21 — 116.

Una Nueva Especie de Mexico y Centroamerica, Marsdenia
hiriartiana (Apocynaceae, Asclepiadoideae, Marsdenieae)

Veronica Juarez Jaimes

Instituto de Biologfa, UN AM, Apartado Postal 70-233, Del. Coyoaean, 04510 Mexico,
D.F., Mexico, vjuarez@ibiologia.unam.mx

W. I). Stevens

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

stevens@mobot.org

RKSUMKN. Una nueva especie, Marsdenia hiriar¬
tiana es descrita e ilustrada. Esta especie es simi¬
lar a Marsdenia macrophylla (Humboldt & Bon-
pland ex Schultes) E. Fournier y M. altissima
(Jacquin) Dugand pero se distingue de la primera
por presentar hojas suborbiculares a orbiculares y
foliculos (mire 22 y 30 cm de largo, y de la segunda
por ser glabra y tener una inflorescencia laxa y el
tubo y los ldbulos de la corola mas o menos del
mismo tamano.

Abstract. A new species, Marsdenia hiriartiana,
is described and illustrated. This species is similar
lo Marsdenia macrophylla (Humboldt & Bonpland
ex Schultes) E. Fournier, and M. altissima (Jacquin)
Dugand, but it is distinguished from the first by its
suborbicular to orbicular leaves and follicles 22 to
30 cm long, and from the second because it is gla¬
brous and has a lax inflorescence and a corolla tube
approximately equaling the corolla lobes.

Key words: Apocynaceae, Asclepiadoideae,
Central America, Marsdenieae. Marsdenia , Mexico.

Marsd« •ilia hiriartiana Juarez Jaimes & W. I). Ste¬
vens. sp. nov. T1PO: Guatemala. Peten: Tikal
National Park, in ramonal covering the ruins,
8 June 1959 (fl). C. Landed 16062 (holotipo,
LL; isotipo, MEXU). Eigura I.

Folia suborbiculata. Infforeseentia umbelliformis laxa,
pedicellis 9.0—13.0 mm longis; calycis lobi 2-3 glandibus
in quoque sinu; corollae lobi 6.0 mm longi. Foil iculus plus
quam 20 cm longus.

Plantas eon latex bianco. Tallos glabros, ramas
7.0 mm de aneho, glabras, lenticeladas. Hojas
I 1.5-22.0 X 9—20 cm, suborbiculares a orbiculares,
eon 7— fl pares de nervios secundarios, base cor-
dada, apice cortamente acuminado, membranaceas,
haz glabra o eon tricomas adpresos muy esparcidos,
eon 6 a 8 coleteres t*n la base del limbo sobre el

nervio medio, enves con zonas eon trieomas adpre¬
sos amarillos y zonas glabras; pecfolo 3.5— 6.0 cm
de largo, glabro. Inflorescencia umbel iforme, con
12 a 20 flores; pediinculo 1.0— 1.4 cm, puberulento;
braeteas lanceoladas, glabrescentes; pedieelos 0.9-

1 .3 cm. glabrescentes; lobulos del ealiz ea. 6.0 X

3.3 mm. anchamente elipticos, la superficie adaxial
glabra con 2 o 3 glandulas en cada seno, superficie
abaxial glabrescente; corola campanulada 0.8— 1.0
cm de diametro, lobulos ea. 6.5 X 3 mm, oblongos;
tubo ca. 6.0 mm de largo; ginostegio ca. 4 mm de
largo; corona estaminal con lobulos ca. 3.5 mm de
largo, oblongo-truncados, dpice agudo; membranas
de las anteras ca. 2.0 mm de largo; anteras ca. 1.3
mm de largo, oblongas; corpusculos ca. 0.6 mm de
largo; polinios ea. 1.2 mm de largo, elipsoidales a
oblongos; apice del estilo 1.5— 2.0 mm, anchamente
conico con pliegues espiralados, obtuso; carpelos
glabros. Foliculos 22—30 X 4—8 cm, muy angosta-
mente elipsoidales; semillas ca. 1.3 X 8 cm, de
forma irregular, hordes trasparentes, eon una linea
media muy marcada, coma ea. 6 cm, blanca.

Marsdenia hiriatiana esta relacionada moifolo-
gicamente con Marsdenia altissima, puesto que am-
bas tienen hojas suborbiculares a orbiculares, entre
9 y 22 cm de largo, pedunculos de alrededor de I
cm de largo, y tubo de la corola de mas de 4.5 mm
de largo, sin embargo, M. hiriartiana se distingue
por sus hojas glabrescentes, inflorescencia laxa, asf
como por la presencia de 2 a 3 coleteres en cada
seno de los lobulos del ealiz, lobulos de la corola
easi del mismo tamano que el largo del tubo de la
corola, apice del estilo 1.5—2 mm de largo, foliculos
glabros. lisos y lenosos. Porotro lado M. hiriartiana
se conoce solo del norte de Mesoam(Tica, mientras
que M. altisima se distribuye desde el norte de
Colombia, Venezuela hasta Argentina.

Marsdenia hiriartiana fue por mucho tiempo
eonsiderada como Marsdenia macrophylla o miem-

Novon 15: 552-554. Publishkd on 12 December 2005.

Volume 15, Number 4
2005

Juarez Jaimes & Stevens
Marsdenia hiriartiana

553

F igura 1. Marsdenia hiriartiana Judrez -Jaimes & W. I). Stevens. — A. Hojas. — It. Inflorescencia. l/)l)iilo <lel

caliz con coleteres. — D. Corola, ginostegio y corona estaminal. — h. Polinario. — F. F into. — G. Semilla. Dilmjo del
tipo, Lundell 16062 (F.L).

554

Novon

labia 1. Rosumen do earacteres

morfoldgicos outre Marsde

ma hiriartiana y especios morfoldgicamente eercanas.

Caraeteres

M. hiriartiana

M. altissima

M. macroph ylla

1 loja

laminas (cm)

1 1.5-22 X 9-20

9-22 X 9-16

1 1.5-20 X 5-13

peefolos (cm)

3.5-6

4-7

2. 0-3.0

forma

suborbicular a orbicular

anehamente ovada a
suborbicular

elfptica a anehamente
elfptica

pubescencia

liaz glabra, pelos adpre-
sos muy esparcidos

liaz densamente tomen-
tosa, envds tomentoso-
velloso

haz y envds glabros en
hojas adultas

Inlloreseeneia

no. de flores

12 a 20

10 a 30

18 a 25

pediinculos (mm)

10-14

8-10

5-13

pedicelos (mm)

9-13

2-6

5-8

no. de coldteres

2—3 en c/seno

sin glandulas

1 en c/seno

largo del tubo de la

6.0

4.5-6

3.4— 3.8

oorola (mm)

largo de los Idbulos de la

6.5

2.4-3

3.6— 5.3

oorola (mm)

largo del ginostegio (mm)

4

2.3-3

4.5

largo do los Idbulos de la

3.5

1.3- 1.6

2. 5-3.9

corona (mm)

largo de las anteras (mm)

1.3

0.5

1.2

largo de los corpusculos (mm)

0.6

0.15-0.18

0.5

largo de los polinios (mm)

1.2

0.7

l.l

largo del apice del estilo (mm)

1. 5-2.0

0.6

1.0- 1.2

forma del estilo

anehamente ednico

rmiico

conico

dpioe del estilo

obtuso

obtuso

obtuso

Fruto

largo (cm)

22-30

16-22

15-17

ancho (cm)

4-8

3.5-6

4-5

forma

angostamente elipsoidal

ovoide-cih'ndrieo

angostamente elipsoidal

textura y pubescencia

glabro lenoso y liso

densamente piloso y
aterciopelado, pericar-
pio grueso y duro

glabro lenoso y liso

bro del cotnplejo de especios reconocido cornu M.
macrophylla (Standley & Williams. 1968). pom M.
hiriartuina so dislinguo pur sus hojas suborbicu-
laros a orl)icularus, pediinculos de mas do I cm do
largo, prosonoia do 2 a 8 coleteres on oada sono do
I os Idbulos dol odliz, tubo y Idbulos do la oorola do
6 mm do largo, y folfculos do mas do 22 cm do
largo. Marsdenia macrophylla so distingue por sns
hojas mas pequefias, elfpticas a anohamonte olip-
lieas, tnbo y Idbulos do la oorola hasta do 4.5 mm
do largo y folfculos hasta do 17 cm do largo. Mars¬
denia macrophylla so eneuontra dosdo Costa Rica
hasta Sudamerica. Kn la labia I so rosumen las
caracterfstieas morfoldgicas do ostas 3 especios.

F.s importanto sefialar (|uo C. L. I. undoll ya habfa
reconocido como una nuova ospocio al material por
dl rocoloctado, pom nunea public^ ol nuevo taxon.

Dislrdmcidn. Ilasta el momonto Marsdenia hi-
riartiana so conoee dosdo Chiapas on Mexico hasta
I londuras.

Marsdenia hiriartiana Judroz-Jaimes & W. I).
Stevens so dedica a la memoria do Patricia Hiriart,
botanica moxicana ostudiosa do la vogotaoidn del
nordosto do Mexico, maestra y amiga.

Baratipos. MEXICO. Chiapas: 5 km NW of Turn ho,
W. Stevens & K. Martinez 25886 (MO). GUATKMAbA.
Csciiintla: below Las l.ajas. B. Standley 64768 (F). IV-
tdn: Tikal, Tikal National Park, C. Lundell 15258 (LL),
(.. Lundell 17089 (LL); Uaxactum, //. Bartlett 12555 (F,
MO). C. Lundell 16628 (LL). HONDURAS. Atlantiiia:
Uancolilla Valley, near Tela, B. Standley 54867 (F).

Agradecimientos. A Patricia Davila. Miguel Or¬
tiz y Fernando Chiang por la revision crftica <lel
manuscrito. A esto ultimo tambidn por su ayutla on
la description on latin.

Literatura Citada

Standley, PC. & L. 0. Williams. 1968. Asclepiadaceae.
Kn: Flora ol Guatemala. Fieldiana, Rot. 24(8, part 4):
407—472.

A New Species of the Genus Actinodaphne with Four New
Synonyms of the Genus Litsea (Lauraceae)

Jie Li

Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming,
Yunnan 650223, People's Republic of China, jieli@xtbg.ac.cn

Xi-wen Li

Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan 650204,

People’s Republic of China.

Abstract. Work in progress for the Lauraceae
volume of the* Flora oj China has yielded one un-
described species and several new synonyms. In
this contribution, a new species, Actinodaphne
menghaiensis J. Li, is described from Yunnan.
Southwest China, and its affinities are discussed.
Four new synonyms are proposed: Litsea garrettii
Gamble, L. euosma W. W. Smith, L. atrata S. Lee.
L. magnoliijolia Yang & P. H. Huang for L. mar-
tabanica (Kurz) Hooker f., L. nwlhs Hemsley, L.
salic if olia (INees) Hooker f., L. semecarpifolia (Nees)
Hooker (.. respectively. Lectotypes are designated
for Litsea garrettii Gamble, L. mollis Hemsley, and
L. euosma W. W. Smith.

Key words: Actinodaphne, China, Lauraceae,
Litsea.

Lauraceae are a large tropical family of trees and
shrubs with small and inconspicuous flowers, which
has rendered the family inaccessible for the non¬
specialist. Although the Chinese species of Laura¬
ceae have been reported in the Flora Reipublicae
Fopularis Sinicae (t BPS) (Li et al., 1984), the tax¬
onomic treatments are incomplete and poorly un¬
derstood because of the insufficient knowledge
about infraspecific variation and distribution, as
well as specific delimitations, and the entire family
in China needs much work. During the preparation
of the Lauraceae treatment for the Flora of China ,
a large number of Lauraceae specimens in KLIN
and the collections received on loan from various
herbaria (E, K. I BSC, YUKU) have been examined.
As a result, one previously undescribed species is
presented here, and four synonyms are made. They
are arranged in alphabetical sequence.

Actiuoduplhne menghaiensis J. Li. sp. nov.
TYPE: China. Yunnan: Jin-bo-a-ka village.
Meng-a district, Meng-hai, in dense humid
valley forest, 1500 m, 10 Sep. 1961. Y. //. Li
3603 (holotype, KUN). Figure 1.

Novon 15:

Actinodaphne obovatae affmis, sed gemmis, ratnulis fo-
liisque glabris; fructibus minoribus, ad 2.5 X 1.5 cm; pe-
dunculis fructiferis gracilioribus, ad 1 cm longis, ca. 2
mm in diam. differt.

Tree, 8 m tall; twigs terete, ca. 0.8— 1.2 cm diam.,
glabrous; terminal buds sub-globose, brownish gla¬
brous. Five to six leaves verticillate at the top of
branchlcts; lamina thin coriaceous, obovate or el¬
liptic. 15-30(-40) X 6 — 1 2 (— 1 S ) cm, the base cu-
neate, the apex acuminate, both surfaces glabrous,
lucid on the upper surface, pale green on the lower
surface, triplinerved; midrib prominently raised on
both surfaces; lateral veins 7 or 8 on each side, 2
basal lateral veins beginning at ea. 0.2— 1.2(— 1 .5)
cm of the leaf base, the parallelodromous tertiary
veins slightly raised on the lower surface. Petioles
2-4(-6) cm long, glabrous. Flower unknown. In-
fructescences racemiform, up to 4 cm long, with
yellow-brown pubescence. Fruit ellipsoid, up to 2.5
cm long, 1.5 cm diam., becoming purple-red when
mature; cupule flat, up to 1 cm diam.; fruit pedicel
slender, up to 1 cm long, ca. 2 mm diam. Fruiting
in September.

In the past, Actinodaphne obovata (Nees) Blume
was erroneously supposed to be distributed in
Northeast India, Southeast Tibet. South and South¬
east Yunnan (Hooker, 1886; Li et al., 1984). After
a careful study of the collection from South Yunnan,
we have found that it is not tin* same as those from
Northeast India, Southeast Tibet, and Southeast
Yunnan that have rusty pubescence on the branch-
lets and leaves. In fact, A. obovata is distributed at
both ends of the Tanaka-Line (Tanaka, 1954; Li X
Li, 1997); the same distribution pattern can be seen
in Litsea chingpingensis Yang X P. H. Huang (Li X
Pai, 1983), Dipterocarpus re/iisus Blume (Li et al.,
2002), and so on. Such a distribution pattern may
be a result from the northward movement of the
Burma-Malaya plate (Audley-Charles, 1987). The

555-558. Published on 12 December 2005,

556

Novon

Figure 1. Actinodaphne menghaiensis J. Li, sp. nov. —A. Vegetative branch, with leaves. — B. A portion of the branch
with three infructescences. — C. Fruit. Drawn from the type specimen, K //. Li 360.3 by Xi-lin Wu.

species Actinodaphne menghaiensis from South
Yunnan, as defined here, has a totally different Ho¬
listic origin from A. obovata, which is from North¬
east India, Southeast libet, and Southeast Yunnan.

At present, Actinodaphne menghaiensis is known
only from the type collection in Menghai, Yunnan,
China. It resembles closely A. obovata and differs
from the latter in having glabrous buds, branehlets.

Volume 15, Number 4
2005

Li & Li

Actinodaphne menghaiensis

557

and leaves, smaller fruits (up to 2.5 X 1.5 cm), as
well as slender fruit pedicels (up to 1 cm long and
2 mm diam.).

Litsea martabanica (Kurz) Hooker f., FI. Brit. In¬
dia 5: 164. 1886. Tetmnthera martabanica
Kurz, For. FI. 2: 301. 1877. TYPE: Burma.
Tenasserim & Martaban, in the drier hill for¬
est, 4000-6000 ft.. Kurz (holotype, K).

Litsea garrettii Gamble, in Kew Bull. 204. 1913. Syn. nov.
TYPE: [Thailand.] Siam, H. H. G. Garrett 63 (lecto-
type, designated here, K).

In 1972, Kostermans determined Edinburgh’s
specimens of Litsea garrettii collected from Yun¬
nan, China, Myanmar, and Thailand as L. marta¬
banica, considering that the former was a synonym
of the latter. A study of the available collections of
L. garrettii in K and KUN. including the syntypes,
shows that this species has all the characters of L
martabanica, especially for the foliage indumentum
and racemitorm pseudo-umbels. We agree with
Kostermans’s treatment and regret that he did not
publish Ins opinion during his life, and therefore
publish the new synonymy here.

Litsea martabanica is distributed in Thailand,
Myanmar, and South to Southwest Yunnan. China,
550—2500 m. It is more or less sympatric, overlap¬
ping in its range with L. semecarpifolia listed below.
Such a distribution pattern shows that there is a
close Holistic relationship between Myanmar and
South Yunnan, China, which may be the result of
the northward movement of the Burma-Malaya
plate (Audley-Charles, 1987).

II. B. G. Garrett 63 (K) is designated as a lec-
totype of Litsea garrettii Gamble for it is the only
collection with staminate (lowers among syntypes
and the epithet is named after the collector.

Additional specimens examined (all specimens at
KUN). CHINA. Yunnan: Menghai, C. W. Wang 73484,
II. T. Tsai 5910403, K. M. Feng 14240, Sino-Russ. Exped.
003, F. Y. Chiu 56449, Y. H. Li 3628, P. I. Mao 7141:
Jinghong, C. W. Wang 75500. K. M. heng 14478: Mcngla,
II. T. Tsai 59-11094, G. J. Pei 59-9481. I. H. Li 1098,
W. T. Wang 10390: Jingdong, V/. K. Li 561, C. Y. Wu 209,

S. G. Xu 3407, S. Y. Bao 366: Si mao. Chinese Forest Inst.
14)1488. P. I. Mao 5790: Menglian, Menglian Exped.
9580: Linoang, T. P. Zhu 009, J. S. Xing 366: Gengma,

T. P. Zhu 312, Y. II. Li 2108, S. T. Li 805: Gangyuan, Y.
II. Li 11843: Shuangjinag, J. S. Xing 941: Longling. H.
T. Tsai 55790: Luxi, C. Chen 452. R. L Xiong 580155:
Ruili, S. Clww 528: Ixmgehuan, G. I). Tao 13565: Fugong,
H. T. Tsai 58903.

Eitsea mollis Hemsley, in J. Linn. Soc. Bot. 26:
383. 1891, non L. mollis (Blume) Boerlage, in
Handl. FI. Ned. bid. 3: 141. 1900. TYPE: Chi¬
na. Hupeh, A. Henry 3177 (lectotype, desig¬
nated here, K).

L. euosma W. W. Smith, in Notes Bot. Card. Edinburgh
13: 166. 1921. Syn. nov. TYPE: China. Yunnan, G.
Forrest 9333 (lectotype, designated here, E).

On the basis of the descriptions, the differences
between Litsea euosma and L. mollis are as follows:
the former is characterized by ovate to elliptic
leaves, acuminate apex, round base, sparse indu¬
mentum on lower surface, and 4 fasciculate pseu¬
do-umbels; the latter by oblong leaves, acute apex,
cuneate base, dense indumentum on lower surface,
and 2 or 3 fasciculate pseudo-umbels. After check¬
ing the syntypes in K and E as well as a large
number of collections in KUN. we consider that the
differences are too overlapping to distinguish them.
Traditionally, the specimens at KUN from tropical
areas are determined as L. euosma. those from sub¬
tropical as L. mollis. After a careful study of these
two species, we consider they are the same; tin*
diagnostic characters for L. euosma fall in the range
of L. mollis, and hence the new synonymy is pro¬
posed. Litsea mollis is distributed in South to
Southwest China and North Vietnam, 200-2700 m.

A. Henry 3177 (K) is designated as the lectotype
of Litsea mollis Hemsley, for it is the more complete
collection among the three syntypes seen by the
authors. G. Forrest 9333 (E) is designated as the
lectotype of L. euosma W. W. Smith, for it is men¬
tioned first by the describing author.

Additional specimens examined (all at KUN). CHINA.
Yunnan: Jinping, K C. Viang 3. II. Q. Sha 101, ./. A.
Zhang 177, Y. C. Hsu 282. Sino-Russ. Exped. 1873: Ping-
bian. P L Mao 3478, Sino-Russ. Exped. 4495. K. M. Feng
4848: Xichou, S. Z. Wang 414, C. W. Wang 85969. C. A.
Wu 7273: Hekou, K. H. Cai 998, K. M. Feng 2 1722: Men-
gla, Y. H. Li 771, C. ./. Pei 59-9481: Jingdong, C. A. Wu
9106, P. Y. Chiu 53020. M. K. Li 3022, X. F. Deng 044:
Shuangjiang, J. S. Xing 1046: Cangyuan, ). II. Li 12232:
Longling, //. T. Tsai 56672, C. Chen 619: Tengchong, C.
Chen 301, W. C. Yin 60-1228. R. L. Xiong 580950: Luxi,

R. L. Xiong 580241: Lianghe, R. L Xiong 580861: Fu¬
gong, H. T. Tsai 54981: Gongshan, Q. Lin 790853. K. M.
Feng 24240; Yangbi, Sino-Amer. Exped. 520. Guangxi:
Rongshui, S. H. Chen 16688, T. C. Chen 797: Longzhou,

S. 11. Chen, 12845. Guangdong: Ruyuan, L. Teng 44158;
Lianxian, P. C. Tam 60118; Yangshan, P. C. Tam 60290.
Jiangxi: Anfu, J. S. Yue 3057; Xunwu, C. M. Hu 1249;
Suichuan, S. K. Lai et al. 5523. Hunan: Ningyuan, P. C.
Tsong 908; Xinning, L. H. Liu 15044. Zhejiang: Taishun,
S. Y. Zhang 5543. VIETNAM. Vinhpliu: Sino-Viet. Ex¬
ped. 2041.

Eitsea salicifolia (Nees) Hooker (.. H. Brit. India
5: 167. 1886. Tetranthera salicifolia Nees, PI.
Asiat. Bar. 2: 66. 1831. TYPE: Himalayas,
Wallich 2536 (lectotype, designated by J. 1).
Hooker, 1886: 167, K).

Litsea atrata S. Lee, in Acta Phytotax. Sin. 8(3): 195.
1963. Syn. nov. TYPE: China. Hainan: Po-ting His-
en, F. C. How 72739 (holotype, I BSC).

558

Novon

Litsea salicifolia, as accepted here, lias wide dis¬
tribution and large morphological variation, just as
Hooker (1886: 168) stated "leaves very variable,
pale brown or darker above when diy, smooth and
usually obscurely reticulated above, nerves beneath
strong with faint nervules.” Characters for Litsea
at nit a (Lee, 1963) are just iu the range of L. sali¬
cifolia , especially for L. salicifolia var. laurifolia
Hooker f.; the differences in leaf shape and color
are rather subtle. We decided to accept this species
in its broad sense and conclude that the two species
are the same; therefore the new synonymy is pro¬
posed. Litsea salicifolia is distributed in India, Ne¬
pal. Sikkim, Bangladesh, Myanmar, South China,
and North Vietnam. 200-1200 in.

Additional specimens examined f all at KUN). CHINA.
Yunnan: Pingbian. II. T. Tsai 61373, I’. I. Mao 3672.
Sino-Russ. L.xped. 3678: Jinping, Sino-Russ. L.xped. 2334.
Luchun L.xped. 1023: Luchun, Luchun L.xped. 238: t un¬
ing. C. W. Wang 80683: Malipo, .S'. /. Wang 818: Jingh¬
ong, Sino-Russ. L.xped. 0803: Mengla, ). II. Li 18641:
Cangyuan, t. II. Li 11673. Guizhou: l.undian. .S Guizhou
L.xped. 774: Anlong, Anshun L.xped. 88: Xingvi. Z. S.
Zhang 6144. Guangxi: I .ongzliou. ,S. II. Chen 11620: Fu-
sui. .S'. II. Chen 12137: Bose,/. T. 14 601024. VIETNAM.
Ninhhinh: Sino-Viet. L.xped. 2363.

Litsea seiueearpifolia (Nees) Hooker f., FI. Brit.
India 5: 165. 1886. Tetranthera semecarpijolia
Nees. PI. Asiat. Bar. 3: 31. 1832. TYPE: Bur¬
ma. Wallich 6345 B (lectotype designated by J.
I). Hooker, 1886: 165, K).

L. magnoliifolia Yang & 1*. H. Huang, in Acta Phytotax.
Sin. 16(4): 52, pi. 3, f. 2. 1978. Syn. nov. TYPE:
China. Gengma, Yunnan Univ. L.xped. 302 (holotype.
YUkU).

After comparison of the specimens from Myan¬
mar with those from China, there is no doubt that
the two species are the same, and hence the new
synonym is made. Litsea semecarpifolia. as accept¬
ed here, is distinguished by its elliptic leaves (1 1-
20 X 5—10 cm), with obtuse apex and cuneate
base; its racemiform pseudo-umbels on axillary
shortened branchlets; perianth segments 8; 12 to
I I fertile stamens; and cup-shaped eupules. Six
perianth segments are typically seen in Litsea. The

presence of eight perianth segments in Litsea se-
mecarpifolia is unusual within the genus. Four peri¬
anth segments have been reported in L. khasyana
by Meissner (in DC. Prodr. XV. I: 185. 1864); with
no perianth segments seen in L. glutinosa by .1. I).
Hooker (in FI. Brit. bid. 5: 157. 1886). Litsea se¬
mecarpifolia is distributed in South to Southwest
Yunnan, China, Myanmar, and East Bangladesh,
740-1400 m.

Additional specimens examined. CHINA. Yunnan:
Gengma, Y. II. Li 2133 (KUN); Jinghong, C. W. Wang
70136 (KUN), Chinese Lores! Inst. Lxped. 14)1614 (KUN).
MYANMAR. Maymyo, Bhamo, Subonchaung to Kun-
baung, J. II. Lice 3283 (E); C. Gilbert Rogers 667 (E).

Acknowledgments. We thank the curators of E.

I BSC, k. KUN. and YLIKU for their kind help and
Xi-lin Wu for Ins skillful drawing. Thanks are also
due to Jens Rohwer and Henk van der Werff for
their helpful comments. Jie Li acknowledges grant
support from YNSF (2001C0008K), CASTA LENTS
(20010713093959), and CEF (498) in China

Literature Cited

Audley-Cliarles, M. G. 1987. Dispersal of Gondwanaland:
Rele vanee to evolution of the Angiosperms. In: T. C.
Whitemore (editor), Biogeographical Evolution of the
Malay Archipelago. Clarendon Press. Oxford.

Hooker,.!. D. 15586. Laurineae. In: J. D. Hooker (editor).
The Flora of British India, Vol. 5. L. Reeve, London.
Lee, S. K. 1963. Notes on Hainan lauraeeous plants. Acta
Phytotax. Sin. 55: 181 — 196.

1.1. X. W. & J. Li. P >97. The Tanaka-Kaiyong Line — An
important Holistic line for the study of the flora of East
Asia. Ann. Missouri Rot. Card. 554: 888—892.

1.1, X. W. & P. Y. Pai. 1983. Lauraceae. In: C. Y. Wu &
X. W. l.i (editors). Flora Yunnanica, Vol. 3. Science
Press. Beijing.

- . P. Y. Pai. S. K. Lee, F. N. Wei, Y. T. Wei, Y. C.

Yang. P. H. Huang, H. I’. Tsui, X. I). Shia X J. I,, l.i.
19551. Lauraceae. In: X. W. l.i (editor). Flora Reipub-
ficae Popularis Sinicae, Vol. .31. Science Press. Beijing.

I.i, X. W., J. l.i X P. Ashton. 2(H)2. Notes on Dipterocar-
paceae of China. Acta Bot. Yunnan. 24: 409-420.
Meissner, C. 15564. Lauraceae. In: A. P. de Candolle (ed¬
itor). Prodromus Systematis Naturalis Regni Vegetablis,
Vol. XV. Masson et Fils, Paris.

Tanaka, 1. 1954. Species problems in Citrus. Japanese
Society for the Promotion of Science, Tokyo.

A New Species of Triaenophora (Scrophulariaceae) from China

Xiao- Dong Li and Jian-Qiang Li

Herbarium (HIB), Wuhan Botanical Garden of Chinese Academy of Sciences,
Wuhan 430074, People’s Republic of China, whlxd9msn@sina.com;
jianqiangl@hotmail.com (for correspondence)

Van yon Zan

Shennongjia National Natural Reserve, Hubei 442421, People’s Republic of China

ABSTRACT. A new species of Triaenophora Soler-
eder, T. shennongjiaensis X. I). Li, Y. Y. Zan & J.
Q. Li (Scrophulariaceae), is described and illus¬
trated from Shennongjia National Natural Reserve,
Hubei Province, China. The new species is related
to T. rupestris (Hemsley) Solereder, from which it
differs in having densely glandular leaves, dentate
bract margins, and pale yellow petals that are re¬
fuse or rarely obtuse at their apices.

Key words: China, Scrophulariaceae, Triaeno¬
phora.

Triaenophora Solereder (Scrophulariaceae) was
separated from the genus Rehmannia Liboschitz ex
Fischer & C. A. Meyer, on the basis of its three-
lobed calyx (Solereder, 1909). Triaenophora is dis¬
tributed in western and northwestern Hubei and
Chongqin in China, with only two species, T. ru¬
pestris (Hemsley) Solereder in Hubei and T. integra
(Li) Ivanina in Chongqin (Forbes & Hemsley, 1890;
Li. 1948; Ivanina, 1955; Chin, 1979). These two
species are similar in general appearance, but the
latter species has smaller leaves, which are round¬
ed at the tip and entire instead of acute and serrate.

fhe new species was collected from the Panlong
cavern, Shennongjia National Natural Reserve, Hu¬
bei, China. It appears most closely related to T.
rupestris (Hemsley) Solereder, hut differs from it in
having densely glandular leaves, dentate bract mar¬
gins, and pale yellow petals that are retuse or rarely
obtuse apically.

Triaenophora shennongjiaensis X. I). Li, Y. Y.
Zan & J. O. Li. sp. nov. TYPE; China. Hubei:
Shennongjia Nat. Nat. Res., Panlong cavern,
I lOe27'58"E, 31224'41"N, 8 Aug. 2003, Yan-
yan Zan 238 (holotype. HIB). Figure 1.

Species T. rupestri Solereder affinis, sed loins utrinque
dense glandulosis vel laxe pilosis, braeteis margine den-
tatis, corollas luteolis, apice retusis vel raro rotundatis dil-
fert.

Novon 15:

Perennial herbs, 10-60 cm tall, leaves and stems
with sparse non-glandular and dense glandular
hairs; roots pale yellow, fleshy, brittle; stem very
short; leaves in a basal rosette, oblong to elliptic,
8-20 X 5—10 cm, densely glandular or pdose on
both surfaces, base subcordate to cuneate, margins
dentate, apex obtuse; petioles 3—6 cm long. Inflo¬
rescence racemose, main raceme suberect, lateral
racemes prostrate, bracts in lower parts resembling
basal leaves, 20—50 cm long, compact, 12- to 54-
flowered; bracts similar to leaves but smaller, ob¬
long, margins dentate; pedicels 5—25 mm long;
bracteoles 2 at middle of pedicels, linear, opposite,
3-20 mm long. Flower with calyx cylindrical, tube
5—8 mm long, erect, 5-lobed with each lobe 3-
toothed, the teeth linear, irregularly acuminate. 6-
12 mm long; corolla pale yellow, 4—6.5 cm long,
pilose abaxially, glabrous adaxially, corolla tube
elongate, erect or slightly curved. 3-4.5 cm long.
1-1.5 cm diam., often with purplish red spots and
glands within the throat, corolla 2-lipped, lobes
with apices obtuse or rarely retuse, superior lobe
separated by deeper sinuses, narrower, 2-toothed,
8-15 X 6—10 mm, inferior lobes 3-toothed. 8—15
X 4—8 mm; stamens 4. included, anthers 2—3 mm
long; ovary ovoid, glabrous, ca. 3.5 cm long, styles
included, stigma bifid. Capsules oblong, 8—15 X 8
mm; seeds numerous, minute, ca. 1 X 0.5 mm,
oblong, laevigate.

Distribution. China, Hubei Province. Shenno¬
ngjia National Natural Reserve and elsewhere, in¬
cluding Badong and Xingshan counties (see para-
types); 700 to 1200 m. The distribution ol this new
species indicates that it is rare. Only 0 populations
containing a total of about 600 to 700 individuals
were found in Shennongjia and neighboring moun¬
tains. The type population consists of only about
100 individuals in nearly inaccessible sites on the
limestone cliff near Panlong cavern. Triaenophora
shennongjiaensis is an endangered species that is

559-561. Published on 12 December 2005.

560

Novon

endemic to the limestone habitats. Intensive
searches to locate additional native populations as
well as strict conservation measures to safeguard
the known populations are urgently needed. Triaen-
<>l>lior<i shennongjiaensis and T rupestris have allo-
patrie distributions: the former in Shennongjia Na¬
tional Natural Reserve, Badong, Xingshan, and
Baokang; and the latter in Fiehuan, Zhuxi, and

Fang counties of Hubei and Wuxi, Wulong, Wush-
an, and Chengkou counties of Chongqin.

Phenology. Flowering late June to mid October.

Both species have been cultivated in Wuhan Bo¬
tanical Garden lor one year, from July 2003 to July
2004. Triaenophora shennongjiaensis flowers in ear¬
ly May, whereas T. rupestris flowers in mid June.

Volume 15, Number 4
2005

Li et al.

Triaenophora (Scrophulariaceae)

561

The diagnostic characters of both species, namely,
the morphology of die bracts, the floral morphology,
the color of the petals, and the pubescence, are
basically stable and show no differences between
cultivated and native specimens.

Paratypes. CHINA. Ilubei : Badong County, Yutougou
village, 20 Aug. 200.'$, Xiaodong Li 0122 (HIB); Baokang
County, 38.5 km along road from Maqiao to Xiema, 31
July 2003, Xiaodong Li 0138 (HIB); 12 Aug. 2004, Xiao¬
dong Li 0158 (HIB); Xingshan County, Xiangping village,
16 July 2003, Xiaodong Li 0131 (HIB).

Key to the Species op Triaenophora

la. Leaves bigger, tip acute, and margin serrate.

2a. Bract margins entire; corolla purple-red, api¬
ces obtuse . I . T. rupestris

2b. Bract margins dentate; corolla pale yellow.

apices retuse or rarely obtuse .

. 2. 7. shennongjiaensis

lb. Leaves relatively smaller, tip rounded, and mar¬
gin entire . 3. T. integra

Acknowledgments. The project was supported
by the Chinese Academy of Sciences and Wuhan
Botanical Garden, CAS (KSCX2-SW-104,
05035123, 01035123).

Literature Cited

Chin, T. L. 1979. Triaenophora. In: T. P. Tsoong & 11. P.
Nang (editors). Flora Reipublicae Popularis Sinicae. To-
tnus 67(2): 220-222. Science Press. Beijing.

Forbes, F. B. & W. B. Hemsley. 1890. An enumeration of
all the plants known from China proper, Formosa, Hai-
nap, Corea, the Luchu Archipelago, and the Island of
Hongkong, together with their distribution and synon¬
ymy. J. Linn. Soc. But. 26: 195.

Ivanina, I.. 1955. Novyi rod son. Scrophlariaceae. Genus
novum Serophulariacearum. Not. Syst. Herb. Inst. Bot.
Korn. Acad. Sci. URSS 17: 393. f. 1-1.

Li, H. L. 1948. A revision of the genus Rehniannia. Tai-
wania 1: 71-82.

Solereder, H. 1909. Uber die Gattung Rehmannia. Bericht
Deutsch. Bot. Ges. 27: 390—101. f. 1—7.

Th ree New Species of Vitaceae from Mesoamerica

Julio Antonio Lombardi

Departamento de Botanica, Instituto de Biooiencias, Universidade Fstadual Paulista “Julio
de Mesquila Filho,” Av. 24-A 1515, 13506-900, Rio Claro, Sao Paulo, Brazil.

cissus@rc.unesp.br.

Abstract. Three new species, Cissus nicara¬
guensis, C. patellicalyx, and Ampelocissus mesoam-
ericana, are described from Nicaragua, Costa Rica,
and El Salvador, respectively. Cissus nicaraguensis
can be distinguished from all other Neotropical
species by the combination of urceolate calyx and
simple leaves. Cissus patellicalyx is recognized by
its subspherical floral buds, immature fruits mi¬
nutely tuberculate when dried, and the distinctly
raised nerves of the abaxial leaf surface. Ampelo-
cissus mesoaniericana is characterized by lenticels,
indument, and the ohpyriform (lower buds.

Rest MKN. Se describen tres nuevas especit*s. Cis¬
sus nicaraguensis, C. patellicalyx y Ampelocissus
mesoaniericana, de Nicaragua, Costa Ric a y El Sal¬
vador, respectivamente. Cissus nicaraguensis puede
distinguirse de todas las otras especies neotropi-
cales por la comhinacidn de un caliz urceolado y
las hojas simples. Cissus patellicalyx se reeonoee
por los botones florales subesfericos, los frutos in-
maduros diminutamente tuberculados cuando secos
y los nervios distintamente prominentes en el lado
abaxial de las hojas. Ampelocissus mesoaniericana
se caracteriza por las lenticelas y el indumento, y
los botones florales obpiriformes.

Key words: Ampelocissus, Cissus, Mesoamerica,
Vitaceae.

Cissus is t h«- largest genus of Vitaceae with per¬
haps 350 species in the entire world (Mabberley,
1987), while Ampelocissus is much smaller, with ca.
95 species in Africa, Asia, and America (Mexico,
south through Mesoamerica and the Caribbean)
(Lombardi, 2000). I revised the family, excluding
Vitis, in the Neotropics to include 80 species: Cis¬
sus (75), Ampelocissus (4), and Ampelopsis (1) (Lom¬
bardi, 2000). Since then, two new species have
been described (Lombardi, 2002. 2004) and anoth¬
er from South America is under description (Ham¬
ilton Beltram, pers. comm.).

While working on the Vitaceae for the Flora Me¬
soaniericana I found the following three new spe¬
cie's: Cissus nicaraguensis, C. patellicalyx, and Am¬
pelocissus mesoaniericana.

Cissus nicaraguensis is from Honduras and Nic¬
aragua. similar and perhaps close to the simple-
leaved and red-flowered species-group including
Cissus biformi folia Standley and C. gossypiifolia
Slandley. Cissus patellicalyx is noted for its sub-
spherical flower buds, unique among the Mesoam-
erican species, the minutely tuberculate dried fruit,
and the raised nerves on the abaxial side of leaves.
Ampelocissus mesoaniericana, which is very similar
to Ampelocissus erdvendbergiana J. E. Planchon,
was overlooked in my monograph (Lombardi,
2000), but is distinguished by its lenticels and the
ohpyriform flower buds.

Cissus nicaraguensis Lombardi, sp. nov. TYPE:
Ni< •aragua. Matagalpa: Rancherfa. I I km NE
de Muy Muy, 12°40'N, 85°31'W, ca. 280 m.
20-22 Aug. 1984 (fl), F. F Moreno 24466 (ho-
lotype, MO; isotype, HNMN not seen). Figure 1.

Liana, inter species neotropicales comhinatione foliis
simplicihus et calvee urceolalo singularis. Haec species
C. biformifoliae et C. gossypiifoliae similis sed calyee ur-
eeolato laterafiter tuberculato, cirrhis non ramosis squamis
destitutis, alabastris ampulliformibus et infloreseentiis mi-
noribus differt.

Liana; the trichomes malpighiaceous, silvery;
branches terete, striated, sericeous and glabreseent
or glabrous; tendrils unbranched, flattened, devoid
of scales. Stipules 2—4 mm, oblong, the base sag¬
ittate or cordate, persistent; leaves simple, drying
brown; petioles (1.5— )3-4.2(— 6.3) cm long, cylin¬
drical. blades (7.9 — ) 1 2— 14( — 1 8) X (2.5— )4— 6.6(-
9.3) cm, triangular on vegetative branches, elliptic
or rarely subtriangular on reproductive branches,
base reniform to truncate on vegetative branches,
cuneate or rarely reniform on reproductive branch¬
es, apex acuminate, margin denticulate, papery,
sparse sericeous in both sides chiefly along nerves
and glabreseent, or glabrous, spiculate on the ab¬
axial side. Inflorescences compound cymes, 1—1.9
X 1—2.1 cm, flat-topped; peduncle 0.4— 0.8 cm
long, sericeous and glabreseent or glabrous; bracts
0.5—1 nun long, triangulate, glabrous, the base of
proximal ones sagittate; pedicels (0.8— ) 1.5— 2 mm
long, sericeous and glabreseent or glabrous. Flower

Novon 15: 562-567. Publish ki> on 12 December 2005.

Volume 15: Number 4
2005

Lombardi

Mesoamerican Vitaceae

563

Figure I. (ass us nicuraguensis. — A. Reproductive branch with leaves and inflorescences. B. Lateral view ol flowei
bud. — C. Leaf of vegetative branch. — I). Lateral view of disc, stamens and style, the petals and two stamens removed.
— L. Lateral (left) and abaxial (right) view of stamens. (A. (. from the holotype Moreno 24466, MO; B, I). E from
Chorley 178, MO). Drawn by the author.

564

Novon

I »u< Is flask-shaped, red; calyx 1.1-1. 4 X 2-2.5 mm.
cotyliform, fleshy, glabrous or sparse sericeous on
the base and glabrescent, lobes deltate and undu¬
late, urceolate, laterally tubereulate, base truncate;
corolla in bud 1.9-2. 1 X 1.5— 1.7 mm, slightly 4-
angled in section, petals coherent, caducous after
anthesis, glabrous; stamens 4, 1—1.5 mm long, fil¬
ament 0.5— 0.8 mm long, the connective lanceolate,
granular, drying yellow, the anther 0.5—1 X 0.5
mm, ovate, latrorsely dehiscent; disc I mm diam.,
minutely tubular, covering the apex of ovary; style
ca. I mm long after anthesis, terete; stigma punc¬
tate, obscurely papillose. Fruit and seed unknown.

Distribution and ecology. Cissus nicaraguen.sis
is known only from Nicaragua and Honduras, at
altitudes of 280—600 m. Flowering in August and
September.

Cissus nicaraguensis resembles C. biformifolia
and C. gossypiifolia, both from Mexico. Mesoam-
erica. and northwestern South America, by the
blade sh ape (in C. gossypiifolia the leaves on the
reproductive shoots only), the red flowers, and the
minutely tubular disc (only in C. biformifolia). The
new species is distinguished easily by the urceolate
calyx laterally tubereulate (vs. not urceolate and not
tubereulate calyx of the other two species), the un¬
branched tendrils devoid of scales (vs. branched
with subtending triangular scales), the flask-shaped
flower buds (vs. conical in C. biformifolia and el¬
lipsoid in C. gossypiifolia), the corolla in bud slight¬
ly 4-angled in section (vs. circular), and the smaller
inflorescences (1—1.9 X 1-2.1 cm vs. (1. 8)3.5—
4. 6(6.2) X ( 1 ,8)2.3-3.4(5.5) cm).

The urceolate calyx combined with the simple
leaves of this species is unique among tin* known
Neotropical species, all other species with urceo¬
late calyces featuring (3-foliolate) leaves.

Paratypes. IIONDl HAS. < Hamlin: Catacamas, out¬
skirts of town, roadside vine on tall planted nettle hedge,

I I Sep. 1991, M. Charley !7H (BM not seen, MO). NIC¬
ARAGUA. Matagalpu: Rancherfa, 1 I km NK dr Muy
Muy, 20-22 Aug. 1984, l>. I>. Moreno 24424 (HNMN not
seen, MO). Nueva Segovia: Kl Jfcaro, 7 km NK del Jf-
caro. valle Casas Viejas, 2 Aug. 1980, P. P. Moreno 1680
(HNMN not seen. MKXU, MO).

Cissus |uilelliealyx Lombardi, sp. nov. TYPE:
(.osta Rica. Cartago: Parque Nac. Tapanti.
along road from main gate to hydroelectric
plant. 3—4 km from main gate, 09°45'20"N,
83°47'00"W, 1250-1350 m. 7 Sep. 1996 (fl,
young IV), T. H. Croat 70030 (holotype, MO | I
of 2; 2 of 2|; isotypes, BHCB. BM not seen,
EAP not seen. INB not seen. MKXU not seen.
PM A not seen). Figure 2.

Liana, inter species Mesoamericanas alabastris sub-
sphaericis et calyce patellifomii singulis. Hare species C.
verticillatae rt C. tiliaceae affinis sed trichoinatibus mal-
pighiaceis, nrrvis distinete prominentibus foliorum abax-
iali superficie, calyce patelliformi, alabastris subsphaeri-
cis et fmetibus lenticrllatis minute tuberrulatis (in sireo)
facile distincta.

Liana ; the trichomes malpighiaceous, silvery;
branches terete, lentieellate, sericeous and glabres-
cent; tendrils not seen. Stipules 2-4 mm, deltate,
the margin scarious, persistent; leaves simple, dry¬
ing green with cream-colored nerves on the abaxial
side; petioles (0.9— )3(— 5.6) cm long, canaliculate,
blades (6.9-) 1 2(— 15.5) X (4.3-)7.7(-l 1.2) cm, ob¬
long, subtriangular or rhombic, slightly asymmetric,
base truncate to reniform, apex acuminate, margin
denticulate, papery, sparsely sericeous on both
sides, chiefly along veins on the abaxial side and
on the blade base, glabrescent, veins very promi¬
nently raised on the abaxial side, secondary veins
forming domatia at the junction with primary vein.
Inflorescences compound cymes, 1.5— 4.5 X 1.5— 2.5
cm, flat-topped; peduncle 0.8— 2.8 cm long, sparsely
sericeous, chiefly in the extremities, glabrescent;
bracts 0.3-1 mm long, triangulate, glabrous; pedi¬
cels 2—3 nun long, glabrous. Flower buds subspher-
ical, green; calyx 1.4— 1.6 mm diam., patelliform,
papery, glabrous, truncate, base rounded; corolla in
bud 1.4— 1,6 X 1.6-1. 7 mm, slightly enlarged on
middle section, petals coherent, caducous after an¬
thesis, glabrous; stamens 4, 1.5 mm long, filament

l. 1 mm long, the connective deltate. granular, (Irv¬
ing yellow-brown, tin* anther 0.5 X 0.5 mm. deltate,
latrorsely dehiscent; disc 1.5 mm diam., not cov¬
ering the apex of ovary, margin minutely raised,
irregular; style ca. 0.6 mm long before anthesis,
terete; stigma punctate, obscurely convex. Imma¬
ture berry 6 X 6 mm, subspherical, green, lenti-
cellate, drying minutely tubereulate; seed 1.6X4
mm, subturbinate, transversely ribbed.

Distribution and ecology. Cissus patellicalyx is
known only by two collections from Costa Rica,
where it occurs in forest, at altitudes of 1250-1400

m. Flowering and fruiting in August and September.

Cissus patellicalyx superficially resembles C. til-

iacea Kunth and C. verticillata (L.) I). 11. Nicolson
iX C. E. Jarvis by its leaves and green flowers, hut
is easily distinguished by the patelliform calyx (vs.
cotyliform), subspherical flower buds (vs. ellipsoid),
malpighiaceous trichomes (vs. unbranched), lenti¬
eellate immature fruits drying tubereulate (vs.
smooth, not lentieellate), and by the prominently
raised nerves on the abaxial side of blades (vs. not
particularly prominent nerves).

I have encountered subspherical buds among the

Volume 15, Number 4
2005

Lombardi

Mesoamerican Vitaceae

565

Figure 2. Cissus patellicalyx. — A. Lateral view of flower bud. — B. Reproductive branch with leaves and inflores¬
cences. — C. Adaxial (left) and abaxial (right) view of stamens. — I). Lateral view of disc, stamens and style, the petals
and two stamens removed. (A— I) from the isotvpe Croat 79030, BHCB). Drawn by the author.

Neotropical species only in Cissus inundata (J. G.
Baker) J. E. Planchon, an endemic of rocky grass¬
lands (“earnpos rupestres”) from northern Minas
Gerais in Brazil. The patelliform calyx is also
unique among the other Cissus species from the

Neotropics; only the calyx of the species of Ampe-
locissus, Ampelopsis, and Vitis can he described as
patelliform.

The collections perhaps suggest this is a rare
species with a restricted distribution; fortunately

566

No von

Figure 3. Ampelocissus mcsoamericana. — A. Lateral view of disc, stamens and style, lliree petals and one stamen
removed. — B. Reproductive branch with vegetative branch leaf. — C. Lateral view of' flower bud. —I). abaxial (left)
and adaxial (right) view of seeds. (A-C from the holotype Tucker 905, US; I) from Ton 2444. F). Drawn by the author.

tin* specimens were collected inside a protected
area, the I’. N. Chirripd, in Costa Rica.

Turatypt \ COSTA RICA. San Jose; Canton de Perez
Xeleddn. I’. INI. Chirripo, Cuenca Terraba-Sierpe. Sendero
FI Llano. <> Aug. 1W7, K. Alfaro 1220 (INB not seen,
MO).

Ampclocissus niesoumcricanu Lombardi, sp.
nov. TYPE: El Salvador. San Miguel: ca. 200
m W of II acienda Potrero Santo. S side of I ,ake
Olomega, 13Q17'N. 88Q05'W, ca. 60 m. 5 Feb.
1942 (11). ./. M. Tucker 005 (holotype, l)S; iso¬
type, UC not seen). Figure 3.

Volume 15, Number 4
2005

Lombardi

Mesoamerican Vitaceae

567

liana bisexualis, A. erdvenbergianae proxima sed ramis
veteribus lenlicellis prominentibus praeditis, fobis baud
hirsutis margines versus et alabastris obpiriformibus dif-
fert.

Liana, hermaphrodite; young branches cobweb¬
by, terete, older sparsely cobwebby, glabrescent,
with prominent cleft lenticels, occasionally with
scars from shoot abscission; trichomes cobwebby,
white or rusty colored, or unbranched short tri¬
chomes. Stipules 2—3 mm, triangular; leaves simple,
caducous; petioles 2.9— 4.5 cm: blade 6.9— 9.3 X
7. 1-9.7 cm, widely-ovate, when young cobwebby
on both sides, mature blades sparsely cobwebby
and puberulent along vein and nerves in adaxial
side, cobwebby in abaxial side, base sagittate, lobes
sometimes overlapping, margin toothed, sometimes
slightly 3-lobed, apex acute or rounded, papery. In¬
florescences 20—25.4 X 1 1.2—37 cm, panicles with
long lateral branches; peduncles 3.9—12.1 cm. cob¬
webby, green; pedicels 2—3 mm, glabrous, red.
Flower buds obpyriform, red; calyx 0.7 nun diam.
gamosepalous, cotyliform, truncate, glabrous; co¬
rolla in bud 1.3-1. 5 X 1-1.3 nun, petals 5, 2.6 X
0.8 nun. elliptic, coherent by margins, caducous,
glabrous; stamens 5, 1 .6—2.3 mm. filaments red, an¬
thers 0.5 mm long, disc adherent to ovary, disc and
pistil 0.9 X 0.6 mm, red, 5-sulcate, ovary apex red,
glabrous, style obsolete, stigma punctate. Berry 8—
10 mm, spherical, purple; seeds (1)3 or 4, 4.7— 5.8
X 3.8—4 mm, heart- shaped. smooth, brown.

Distribution and ecology. Ampelocissus me-
soamericana occurs in low deciduous forests, high
subevergreen forests, and dry savannahs on slopes
at altitudes of 60—1350 m in Mexico and El Sal¬
vador. Flowering in January and February, fruiting
in February, April, and May— August.

I had overlooked this species while studying Am¬
pelocissus specimens for their treatment in Flora
Neotropica. Ampelocissus mesoamericana is very
similar to A. erdvendbergiana (illustration in Lom¬
bardi. 2000: 21) in the inflorescence with long lat¬
eral branches (vs. short branches in sympatric A.
acapulcensis (Kunth) ,|. F. Planehon) and mature

leaves absent at flowering time, but A. mesoameri¬
cana differs from A. erdvendbergiana in the prom¬
inent lenticels that mimic little emergences (vs.
smooth branches), the leaf' margin not hirsute (vs.
hirsute), and the obpyriform flower buds (vs. spher¬
ical).

Paratypes. MEXICO. Chiapas: Chiapa de Corzo, El
Chorreadero 9 kin E of Chiapa de Corzo along llwy. 190,
D. F. Breedlove 8462 (F); Tuxtla Gutierrez, along road to
El Sumidero de Tuxtla. 20 km N of Tuxtla Gutierrez, D.
F. Breedlove 9050 (E, US); La Trinitaria, along llwy. 190,
21 km S of La Trinitaria, D. E. Breedlove & I’. II. Raven
8489 (F); Tzimol, ca. 7 km SE of Gomitan. ea. 3 km SE
of Tzimol, R. ./. Hampshire, A. Reyes-Garcia & L. Hernan¬
dez /. 1148 (BM); Chiapa de Corzo, El Chorreadero 9 km
E of Chiapa de Corzo along llwy. 190, R. M . Ixiughlin
198 (F, MEXU); W of La Concordia, along the Rfo de la
Concordia, A. S. Ton 2444 (F); Avispero, Comitan-Moto-
z. i nl la. F. //. Xolocolzi X-1398 (MEXU). Veracruz,: Te-
petzintla, Tierra blanra, camino para la Sierra de San
Juan Otontepec, ./. Galzada 5528 (F). GUATEMALA. Za -
eapa: Serra de Las Minas, Loma El Picaeho, above Santa
Rosalia, ./. A. Steyermark 42782 (F). HONDl HAS. La
I’az: Ouebrada Salada. I km al N de La Paz, valle de
Comayagua. B. L. Holst 1622 (TEFII). EL SALVADOR.
Aliiineliapuu: near Salto de Atelnieefa, P. C. Standley
F. Padilla V. 2868 (E). Santa Ana: vicinity of Melapan,
P. C. Standley X F. Padilla V. 8085 (E).

Acknowledgments. 1 thank Gerrit Davidse for
the invitation to collaborate on the Flora Mesoam¬
ericana project, T. S. Filgueiras for the Latin di¬
agnosis, Conselho Naeional de Desenvolvimento
Cientifico e Tecnologico (CNPq) for a research fel¬
lowship grant (300220/2003-0), and Fundayao de
Amparo a Pesquisa do Estado de Sao Paulo (L\-
PFSP) for financial support (2005/02654-1).

Literature Cited

Lombardi, J. A. 2000. Vitaceae — Generos Ampelocissus.

Ampelopsis e Cissus. FI. Neotrop. Monogr. 80: 1-251.
- . 2002. Cissus pinnatifolia (Vitaceae), a new spe¬
cies from the Atlantic coast of brazil, brittonia 54: 175—
177.

- . 2004. Cissus xerophila (Vitaceae), a new species

from the xerophytic vegetation of northeastern Minas
Gerais, Brazil, brittonia 56: 288— 290.

Mabberley, I). .1. 1987. The Plant-book. Cambridge Univ.
Press, Cambridge.

Especie Nueva <le Croton (Euphorbiaceae) de Brasil

Martha Martinez Gordillo

Herbario de la Hacultad de Ciencias, UN AM. Apartado Postal 70-399, Delegacidn Coyoacdn
04310. Mexico, D.F. mjmg@hp.fcieneias.unam.mx

Efrai'n de Luna

Departamento de Sistematica Vegetal, Instituto de Ecologfa A.C., Apartado Postal 63,
Xalapa 91070, Veracruz, deluna@eeologia.edu. rnx

Kksi mkn. Se describe Croton longihracteatus , es¬
pecie nueva de Brasil. Esta especie perteneee a
Croton sec. Barhamia (Klotzsch) Baillon y se con-
sidera cereana a C. mnscicapa Miiller Argoviensis,
con la eual se compara y se distingue por las es-
tfpulas 1 »i«*n desarrolladas, las hracteolas mas gran-
des, los sepalos pistilados mas grandes y aerescen-
tes y los estambres exsertos. Se presenta una clave
para distinguirlas y una ilustracidn de la especie.

ABSTRACT. Croton longihracteatus (Euphorbia¬
ceae), a new species from Brazil, is described and
illustrated. It belongs to Croton sect. Barhamia
(Klotzsch) Baillon and appears similar to C. mus-
cicapa Miiller Argoviensis, but differs from the lat¬
ter by presenting well-developed stipules, larger
bracteoles, large and accrescent pistillate sepals,
and exserted stamens. A dichotomous key for dis¬
tinguishing both taxa and an illustration of the new
species are included.

Key words: Barhamia . Brazil, Croton. Euphor¬
biaceae.

El genero Croton E. esta ampliamente distribuido
en las regiones tropicales de la tierra. En America
se encuentran 33 secciones de las 40 en las euales
se dividib al genero (Webster. 1993), 29 de ellas
son endemicas de esle continente (Martinez. 1996).
Una de las secciones reconocidas de Croton es Bar¬
hamia. que presenta aproximadamente 18 especies
V se encucntra relaeionada eon dos secciones ame-
ricanas: Micranthis Baillon y Medea (Klotzsch) Bai¬
llon. Ea sect ion Barhamia se distingue por ser ar-
bustos con hojas generalmente dentadas, sin
glandulas conspicuas en cl apice del peefolo; es-
tipulas con glandulas, lobadas o ligeramente disec-
tailas; estambres 5 a 12; sepalos de las flores pis-
tiladas con glandulas marginales y/o abaxiales v los
estilos multilidos (Webster, 1993).

En cl Herbario de la Facultad de Ciencias
(FEME) se eomenzb el estudio de Croton sec. Bar-

Novon 13: 568-571. Published on 12 Dt-.ct

hamia en anos recientes y se hicieron analisis fi-
logeneticos basados en caracteres morfologicos en
donde se encontro un clado que incluye a especies
con los pedicelos femeninos largos, las hojas bri-
llantes y los tricomas, cuando presentes, debajo de
una capa de exudado, dando la impresibn de que
la lamina es glabra y vfscida en ambas caras.

En esta revision se solicitaron ejemplares de di-
ferentes herbarios de Brasil y de los Estados Llni-
dos de America y dentro de los mismos se obtuvie-
ron espeefmenes de Croton mnscicapa Miiller
Argoviensis. un taxon perteneciente al clado men-
cionado, que sin embargo presenta ejemplares con
caracteristicas diferentes y que al examinarlos con
detenimiento son suficientes para considerarla
como una especie distinta, la cual se propone en
este trabajo.

Croton longihracteatus Martinez Gordillo & de
Luna, sp. nov. TIP0: Brasil. Bahia: about 7—
10 km along Seabra-ltaberuba road W the
Eengois turning, by Bio Mucugizinho.
41°26 W, 12°28'S. 800 m, 27 May 1980. B.
M. Harley 22675. G. L. Bromley , A. M. De Car¬
valho. ./. M. Soares. J. /.. Huge & E. B. Dos
Santos (hoh)tipo, NY). Figura 1.

Frutex 1—2 alta. Folia ovato-deltata, apice acuminata,
margine denticulata; stipulae lanceolatae. Inflorescentia
8-14 cm longa; bractea 13—33 mm longa. lineari-lanceo-
lata, caduca; bracteolae lanceolatae, glandulis stipitatis
margine. Stamina 10, exserta; sepala pistilata lanceolata,
acrescentia, glandulis stipitatis dense ohsila. C. muscica-
pae at finis sed stipulis lanceolatis, inflorescentiis majus-
culis et staminibus 10 exsertis differt.

Arbustos 1—2 m, monoicos. Hojas 4.6— 6.5 X
2. 5-4.3 cm, ovado-deltadas; apice acuminado, mar-
gen denticulado, dientes con glandulas sesiles en
el apice. base cordata; 5 venas a la base; haz y
enves glabrescentes, con algunos pelos estrellados
sobre las venas; peefolo 1.5— 5.4 cm, con algunos
pelos simples; estfpulas 2.5— f) mm, lanceoladas,

iMHKK 2005.

Volume 15, Number 4
2005

Martinez Gordillo & de Luna
Croton longibracteatus de Brasil

569

Figura 1. Croton longibracteatus Martinez Gordillo & de Luna. — A. Kama con flores. — B. Flor estaminada. — C.
Flor pistilada. — D. Semilla. — E. Br&ctea de la base de la infiorescencia. — F. Margen de la hoja. — G. Eslfpula. A,
G, F y G, del holotipo (Harley et al. 22675, NY); B, Harley 18566 (NY); I), Mori <£■ Boom 14258 (NY); E, Mori el at.
13113 (NY).

570

No von

caducas, margen con glandulas botuliformes, sesi-
les, cara adaxial glandular, concentradas en la
base. Inflorescencia 8—14 cm; bractea 13-33 X 2—
5 mm. linear-lanceolada, margen con glandulas es-
tipitadas, base con una glandula circular promi-
nente, generalmente caduca; bracteolas 7-15 min.
lanceoladas. margen con glandulas estipitadas. Flo¬
res estaminadas pediceladas, pedicelo 2—4.5 mm,
sepalos 3.5— 5.5 X 2—2.8 mm. oblongo-ovados, uni-
dos solo a la base, cara adaxial glabra, cara abaxial
glabrescente, con algunos pelos estrellados en la
base. glandulas impresas presentes. con un meclidn
de pelos en el dpiee; petalos 4—6 X 2—3 mm, eltp-
ticos. cara adaxial v abaxial glabras. margen peloso,
glandulas impresas presentes; estambres 10, exser-
tos, filamentos glabros, conectivo glandular; flares
pistiladas pediceladas, pedicelo 2.5—5 mm, sepalos
unidos en la base, lanceolados, acrescentes, mar¬
gen glandular, cara adaxial glabrescente, cara ab¬
axial glandular-estipitada; ovario esferico, hirsuto,
estilos divididos mas de una vez, glabrescentes, li-
bres casi a la base, 9 a 12 esligmas. Fruto 6—9 X
5—9 mm, esferico, lobulado en el apice; semillas
3.5— 5.5 X 3—4 mm, elipsoides, cafes, lisas.

Fenologta. Se ha colectado de* febrero a di-
ciembre, en todos los casos presenta flores en di-
lerentes estados de desarrollo.

Distribncidn. Ilasla el momenta se loealiza en
el estado de Bahia donde crece en sitios con suelos
pedregosos y vegelacidn baja, de 100 a 1200 m de
altitud.

Discusuin. Es una especie perteneciente a Cro¬
ton sec. Rarhamia, que se encuentra relacionada
con cuatro especies (jut1 muestran hojas glabras y
vfscidas por el exudado que las cubre y que se
ballan en Sudanferiea. Todas, excepto C. perviscosiis
Croizat de Paraguay, se localizan solamente en el
estado de Bahia. Brasil, de donde son endemieas y
se desarrollan en lugares abiertos y pedregosos. En
los herbarios del Jardim Botanico do Rio tie Janeiro
(RB), del New York Botanical Garden (NY) y de la
Universidade de Sao Paulo (SPF), estos ejemplares
se ban identificado como Croton mnseicapa, especie
que se eonsidera mas eercana, tliliere porque pre¬
senta estfpulas bien desarrolladas, las bracteolas y
los sepalos pistilados mas grandes y los estambres
exsertos. A continuacidn se presenta una clave que
separa a las especies de esle grupo. Las caracle-
rfsticas de C. perviscosiis se obtuvieron de la des-
cripcion original porque no se tiene ningun ejem-
plar.

Ci. Avi: paha Difkhknciar a Croton i.ongibrm:te\tis.w.
BAHfA, Brasil, dk las Espkciks Ri i acionadas

la. Hojas elfptieas u obovadas, pedicelos de la Hor
pistilada de 2-3 mm de largo, estfpulas serradas

. C. perviscosiis Croizat.

II). Hojas ovadas o deltadas, pedicelos de la Hor pis¬
tilada 2.5—14 mm de largo, estfpulas no serradas.

2a. Hojas con el dpiee agudo, bracteolas ente-
ras, con glandulas s<5lo en la base ....

. C. glutinosus Midler Argoviensis.

2b. Hojas con el apice redondeado o aeuminado,
bracteolas con el margen glandular.

3a. Hojas con el apice redondeado. 14 a

IB esligmas .

. C. betulaster Muller Argoviensis.

3b. Hojas con el dpiee aeuminado, 9 a 12
esligmas.

4a. Apice de la lioja largo aeuminado,
margen denticulado, estfpulas lan¬
ceoladas, inflorescencia 8—1 1 cm.
gldndulas de las bracteolas estipi-
ladas, estambres 10. exsertos, se¬
palos de la (lor pistilada con gldn¬
dulas estipitadas .

. C. longibractealus sp. nov.

lb. Apice de la hoja corto aeuminado,
margen crenado. estfpulas repre-
sentadas por gldndulas esfericas,
inflorescencia 2.2— 5.2 cm. gldndu¬
las de las bracteolas sesiles, estam¬
bres 12. no exsertos, sepalos de hi
(lor pistilada con gldndulas no es¬
tipitadas .

.... C. muscicapa Muller Argoviensis.

Paratipos. BRASIL. Ilaliia: Mun. Mucuge. Mucu¬
ge. //. /' Bautista 1392 (RB); Refugio Ecoldgico (Cam-
po Bupestre) II. P. Bautista 1392 (BB); km 24 Mucuge—
Andorai. A. Allan 1795 & G. Vieira (NY); Serra do
Sincard, 6.5 km al SO de Mueugd, sobre la earr. al
Cascavel, B. M. Ilarley 2 1 0 1 7 (NY); 5 km al S de An-
dorai sobre la earr. a Mucuge por el puerile sobre el no
Baraguazu. R. M. Harley 185 66 (NY); por el rfo Aim-
liuca, ca. 3 km al S de Mucugd, sobre la earretera a
Oascavel. R. M. Ilarley 15919 (NY); .3 km al S de Mu-
cug6, en la entrada (|ue va para Jussiape, S. A. Mari,
R. M. Ring. T. .S’. l)os Santas & R. Pinto 12585 (NY);
mun. Andorai, Sierra de Andorai-Capa Bode, entrada
por Mucuge, campos rupestres, G. Martinelli 5417
(RB); Andorai, Rfo San Antonio, C. Toledo Rizzini &
Malt us 1066 (RB, MO); 5 km al sur de Andorai. sobre
la earr. a Mucuge, por la orilla, sobre el rfo Piraguaeii;

R. i M. Ilarley 18566 (NY ); I km al S de Andorai. eampo
rupestre, /. P. de Queiroz 1809 (NY); Nova Rodovia
Amlorai-Mucugd, a 3 km al S de Andorai, eampo ru¬
pestre, .S’. A. Mori & F. P. Benton 131 13 (NY); mun.

I .envois, a lo largo BR 242. a 15 km al NO de I. envois,
al km 225, eampo rupestre, .S. A. Mori & B. M. Boom
14258 (NY ); estrada entre I , envois e Seabra, 20 km NO
de I .envois. R. M. Harley, G. M. Sakaragui, P. T. Sarto,

S. Atkins & V. G. Souza 14037 (SPF, FCMF,).

Agradecimientos. A los cura< lores de los lier-
lmrios del Jardim Botanico do Rio de Janeiro (RB),
del New York Botanical Garden (NY) y de la Ini-

Volume 15, Number 4
2005

Martinez Gordillo & de Luna
Croton longibracteatus de Brasil

571

versidade de Sao Paulo (SPF), por los prestanios de
ejemplares de Croton sec. Barhumui. A Ramiro
Cruz Duran por la ilustracion de la espeeie. A Jose
Antonio Hernandez, del laboratorio de Miorocine.
de la Faeultail de Cieneias, UNAM. por la eaptura
en imagen digital de la ilustracion.

Literatura Cilada

Martinez C., M. 1996. FI genero Croton (Euphorbiaceae)
en Mesoamerica. Tesis. Facultad de Cieneias. UNAM.
Mexico.

Webster, G. I.. 1993. A provisional synopsis of the sections
of the genus Croton (Euphorbiaceae). Taxon 42: 793-823.

New Species and Combinations in Heterotaxis Lindley
(Orchidaceae: Maxillariinae)

Isidro Ojeda,'-2 German Carnevali Fernandez- Concha, and Gustavo A. Romero- Gonzdle zA
'Herbarium UICY, Centro de Investigation Cientffica tie Yucatan, A.C., Calle 43, Mo. 130,
Col. Chuhurnu tie Hidalgo, Merida 97200, Yucatan, Mexico, ojalis@yahoo.com
2Current address: University of British Columbia, Center for Plant Beset irch (CPR)

# 302-Macmillan Building 2357 Main Mall, Vancouver, B.C., Canada V6T 1/4.

isidro@inteniiange.ubc.ca
'Corresponding author, carneval@cicy.mx

'Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue,
Cambridge, Massachusetts 02138, U.S.A. romero@oeb.harvard.edu

Abstract. Two undescribed species were detect¬
ed while conducting a phylogenetic study of Hel-
erotaxis (Orchidaceae: Maxillariinae): H.fritzii Oje¬
da & Carnevali from western Ecuador and //.
schultesii Ojeda tK G. A. Romero from the Amazon
basin of Colombia and Brazil. These two species
are described and illustrated, and their affinities
are discussed. The first novelty is related to Het¬
erotaxis sessilis (Swartz) F. Barms but the leaves are
triquetrous, fleshier, and much narrower, while II.
schultesii is similar to II. villosa (Barbosa Ro¬
drigues) F. Barros, but with narrower, more coria¬
ceous leaves and orange-red flowers. Six species
are transferred to Heterotaxis: II. discolor (Loddiges
ex Findley) Ojeda & Carnevali, //. equitans (Schle-
chter) Ojeda & Carnevali, //. maleolens (Sehlechter)
Ojeda & Carnevali, //. microiridifolia (1). E. Ben¬
nett & E. A. Christenson) Ojeda & Carnevali, II.
santanae (Carnevali & F Ramirez) Ojeda & Car¬
nevali. and II. valenzuelana (A. Richard) Ojeda &
Carnevali. In addition, a lectotype and an epitype
are designated for Camaridium equitans Sehlechter,
while lectotypes are chosen for Camaridium van-
diforme Sehlechter and Dicrypta irisphyta Barbosa
Rodrigues. Lastly, Maxillaria valenzuelana subsp.
angustifolia J. T. Atwood is treated for the first time
as a synonym of Heterotaxis valenzuelana.

Rksumkn. Se detectaron dos especies nuevas eu
un (‘studio filogenetico de Heterotaxis : H. fritzii, de
Ecuador occidental y Heterotaxis schultesii, de la
cuenea amazdniea de Brasil y Colombia. Estas dos
nuevas especies son descritas, ilustradas y sus afi-
nidades diseutidas. Fa primera novedad esta rela-
cionuda con Heterotaxis sessilis (Swartz) F. Barros
pero las hojas son trfquetras, mas carnosas y an-
gostas, mientras que //. schultesii es similar a //.
villosa (Barbosa Rodrigues) F. Barros, pero con ho¬

jas unis angostas y eoriaceas y flores naranja rojizo.
Otrt is seis especies son transferidas a Heterotaxis :
II. discolor (Foddiges ex Findley) Ojeda cK Carne¬
vali, //. equitans (Sehlechter) Ojeda & Carnevali,
II. maleolens (Sehlechter) Ojeda & Carnevali, //.
micro-iridifolia (I). E. Bennett & Christenson) Oje¬
da & Carnevali, H. santanae (Carnevali & I. Ra¬
mirez) Ojeda & Carnevali y H. valenzuelana (A.
Richard) Ojeda & Carnevali. Adicionalmente, se
se lec ciona un lectotipo y un epitipo para Camari¬
dium equitans Sehlechter y lectotipos para Cama¬
ridium vandiforme Sehlechter y Dicrypta irisphyta
Barbosa Rodrigues. Por ultimo, Maxillaria valen¬
zuelana subsp. angustifolia J. T. Atwood es tratada
P<>r primera vez como sindnimo de lleterotaxis va¬
lenzuelana.

Key words: lleterotaxis, Maxillariinae, Orchi¬
daceae.

Maxillariinae is one of the most conspicuous and
diverse subtribes of the Neotropical Orchidaceae.
It is composed of 400 to 450 species (e.g., Atwood
& Mora de Retana, 1999; Carnevali & Ramfrez,
2003), approaching 650 species in other recent cal¬
culations (e.g., Dodson, 2002). They are common
and widespread elements in humid Neotropical for¬
ests and range from southern Florida (U.S.A.) to
northern Argentina, with centers of diversity in
southern Central America, along the Andes, and in
southeastern Brazil (Dressier, 1981, 1993; C arne-
vali, 1991; Senghas, 1993; Whitten et al.. 20(>0).

Members ol Maxillariinae are extremely variable
in vegetative morphology and size, varying in height
from I cm to 2 m. Plants bear heteroblastic pseu-
dobulbs (i.e.. of 1 internode) or lack pseudobulbs
altogether and feature I to 2(to 4) leaves per stem
or pseudobulb. Rhizomes are short, branching or

Novon 15: 572-582. Published on 12 December 2005.

Volume 15, Number 4
2005

Ojeda et al.

Heterotaxis (Orchidaceae)

573

not. forming erect, ascendent, or pendulous plants.
Plants are epiphytic or secondarily terrestrial in
habit. Growth may be either monopodial or sym-
podial. Despite all this vegetative plasticity, the
flowers are remarkably homogeneous in overall
structure throughout the subtribe (Carnevali & Ra¬
mirez, 1989; Carnevali, 1991; Holtzmeier et al..
1998; Dodson. 2002). Thus, the phylogenetic re¬
lationships of the several morphologically recogniz¬
able groups within the subtribe and their circum¬
scriptions have been difficult to ascertain, partially
because the great vegetative plasticity of the group
is not obviously associated with an equivalent, cor¬
related floral variability.

Until recently, orchid systematists had recog¬
nized the following genera in the Maxillariinae:
Cryptocentrum Bentham, Mormolyca Fenzl,
Chrysocycnis Bindley & Reichenbach f.. Cyrtidior-
chis Rauschert, Anthosiphon Schlechter, Pitiphy-
llurn Schlechter, Trigonidium Bindley, and Maxilla-
ri<i Ruiz & Pavon (Dressier. 1993; Atwood cA Mora
de Retana, 1999; Dodson. 2002; Carnevali & Ra¬
mirez. 2003). However, recent phylogenetic analy¬
ses using DNA sequences (Whitten et al., 2000;
Whitten, unpublished) have resulted in a broader
understanding of the relationships within tin* sub-
tribe and among some of the groups included within
Maxillaria. The most striking result of these new
analyses is, perhaps, evidence indicating that the
large genus Maxillaria, as currently circumscribed,
is grossly polvphyletic. Alternatively, the circum¬
scription of the genus would have to be expanded
in such a way as to be morphologically undiagnos-
able and ecologically and biogeographically mean¬
ingless, since it would include all members of sub¬
tribe Maxillariinae with conduplicate leaves.
Furthermore, the circumscription of subtribe Ma¬
xillariinae has also recently been expanded to in¬
clude subtribes Bycastinae and Bifrenariinae
(Whitten et al., 2000), in disagreement with the
currently accepted circumscription that in a narrow
sense encompasses only species with conduplicate
leaves, therefore excluding Bycastinae and Bifre¬
nariinae (the plants of which bear plicate leaves).

Thus, relationships within subtribe Maxillariinae
s. str. beg for a generic realignment to recognize
generic aggregations that, besides being monophy-
letic, would be more manageable in size, morpho¬
logically diagnosable, and that would reflect eco¬
logical preferences and biogeographieal patterns.
The circumscriptions of some genera surely will re¬
quire broadening, while other smaller monophyletie
groups, most of them formerly included within Ma¬
xillaria, will deserve generic recognition (Ojeda,

2003; Whitten, unpublished; Ojeda & Carnevali. in
prep.).

Among the several monophyletie groups recog¬
nized by the preliminary phylogenetic analyses
there is a basal clade that can be identified by a
combination of morphological features. One of the
most conspicuous of them is the presence of fleshy
flowers as opposed to the typically fibrous flowers
of most members of the subtribe. Within this clade
there are three monophyletie groups. One of them.
Heterotaxis Bindley, has been recently reinstated
(Barros. 2002), and this circumscription is strongly
supported by our phylogenetic work that places
Heterotaxis in a basal position in the phylogeny of
Maxillaria, or of the conduplieate-leaved Maxilla¬
riinae, more closely related to other taxa (such as
Ornithidium Salisbury, Mormolyca. and Cryptocen¬
trum) than with the "core” Maxillaria (Ojeda et al..
in prep.; Whitten et al., 2000; Whitten, unpub¬
lished) typified by Maxillaria platypetala Ruiz &
Pavon (e.g.. Dodson, 2002).

Heterotaxis comprises about eleven primarily
epiphytic species ranging from the southeastern
United States (Florida) and the Greater Antilles to
Brazil, with most of the species occurring in Central
anil South America (Carnevali, 1991). As in Ma¬
xillaria s. str., the inflorescence in Heterotaxis is 1-
flowered and emerges from the leaf axils. The flow¬
ers are yellow to orange or orange-red, with the
labellum in some species being totally purple, or
more rarely pinkish I i ps with calli varying in size
and texture; in some other species the lamina of
the labellum is basically yellow or orange with pur¬
ple or reddish spots or blotches. The distinctly
fleshy flowers are shared with the Ornithidium com¬
plex of Maxillaria (a related group of taxa that may
also merit generic recognition). Heterotaxis is char¬
acterized by short rhizomes and laterally com¬
pressed, oblong pseudobulbs, which are unifoliate,
and subtended by various leaf-bearing sheaths. Ex¬
ceptions are Maxillaria equitans Schlechter, and
Maxillaria valenzuelana (A. Richard) Nash, which
exhibits a pseudomonopodial growth without pseu¬
dobulbs. Based on this growth habit difference,
Hoehne (1947) proposed Marsupiaria based on Ma¬
xillaria valenzuelana. where he also included Ma¬
xillaria equitans.

It has been hypothesized (e.g.. Carnevali, 1991)
that Maxillaria witsenioides Schlechter (syn: Mar¬
supiaria witsenioides (Schleeter) Pabst) also belongs
in this group because of its monopodial growth hab¬
it with laterally compressed leaves. However, recent
evidence from the internal transcribed spacers se¬
quence (ITS) along with a detailed morphological
re-evaluation of M. witsenioides (Ojeda & Carne-

574

Novon

vali. unpublished) clearly shows this taxon to lie
unrelated to the members ol Marsupiaria and plac¬
es the species as a member of a clade that includes
Maxillaria acianlha Reiehenbach I. and Maxillaria
notylioglossa Reiehenbach f. (N. II. Williams, un¬
published).

Heterotaxis was based on a species described in
1 826 by John Lindley as Heterotaxis crassifolia.
I'his species was treated until recently as Maxilla¬
ria crassifolia (Lindley) Reiehenbach f. ( = Hetero¬
taxis sessilis Lindley). The same concept was later
redescribed by Lindley as Hi crypt a crassifolia
Lindley (Lindley, 18.10). Although Lindley argued
against his earlier generic concept, the Internation¬
al Code oj Botanical Nomenclature (Greuter et ah,
2000) should be followed in applying the Principle
of Priority. Thus, the name to be used to refer to
these taxa is the older Heterotaxis Lindley, sensu
Lindley, 1826.

Most authors have treated members of this com¬
plex ol taxa within a broadly circumscribed Maxi¬
llaria (e.g., loldats, 1070: Carnevali, 1991; Seng-
has. 1993; McLeish cl ah, 1995; Christenson,
1999), although several had pointed out the possi¬
bility of generic recognition (e.g., Rrieger & lllg.
1072; lllg, 1977; Carnevali. 1991; Carnevali iK Ra-
mfrez, 2003) or had already segregated totally or
partially the group into new genera or through the
resurrection of older genera (e.g.. I Iodine, 1947).

I he group had already been recognized as distinct
in several features, including characteristics of pol-
liuia and cytology (Carnevali, 1001) as well as by
the fleshy flowers. Recently, Christenson (1000)
based on morphological characters, recognized this
aggregate of species as Maxillaria sect. Iridifolia.
John T. Atwood (unpublished) recognized the Het¬
erotaxis alliance that, together with sub-alliances
Crassifolia and Valenzuelana, comprises the Het¬
erotaxis complex. More recently, a paper published
by Rarros (2(H)2) recognized Heterotaxis and trans¬
ferred four Brazilian species of this complex. Fi¬
nally, in the course of a systematic and phyloge¬
netic study of the Maxillariinae in general and of
the Heterotaxis complex in particular that was
based on morphology, gross foliar anatomy, and
DINA sequences (nuclear ITS I and ITS2 regions),
additional evidence was obtained that supports the
idea that Heterotaxis deserves generic recognition
(Ojeda, 2003).

I la- objective of this contribution is to describe
two new species that were detected in the course
of the systematic analysis of tin* genus Heterotaxis.
We then proceed to formalize the nomenclatural
novelties required to update the circumscription

and nomenclature of the genus, making six names
available for general systematic and florist if usage.

Heterotaxis fritzii Ojeda & Carnevali, sp. nov.
I VPK: Ecuador. Imbabura: Lita-Alto Tambo
area, probably along the road from Lita to San
Lorenzo, flowering in cultivation in Gaines¬
ville. Florida, U.S.A., 24 Feb. 2004, M. W.
Whitten 2672 (holotype, f LAS). Figures 1. 2.

Species haec II. sessili (Swartz) F. Harms similis sed
habito parviore, foliis triquetris proportione angustioriluis
brevioribus, callo parte anteriore farinoso differt; II. san-
tanae (Carnevali & I. Ramirez) Ojeda & Carnevali in men-
teni revocans sed foliis triquetris proportione angustiori-
bus, labello acuto recedit.

Plant eaespitnse, sub-terrestrial, presumably also
epiphytic, erect, ca. 25 cm tall, rhizome unknown
but presumably abbreviated; roots grayish, ca. 2
mm diam.; pseudobulbs small. 1-2 X 0.4— 0.6 cm.
narrowly ellipsoid, unifoliate, partially or totally
hidden by sheaths of which tin- innermost 2 to 3(to
4) on each side bear foliar blades. Leaves and the
foliar blades of the sheaths distichous, 6—13 X
0.45—0.55 cm. thickly fleshy, linear-elliptic, sub-
triquetrous with the upper margins rounded (thus
somewhat heart-shaped in cross section), unequally
2-lobed at apex, lobes obtuse, basally not constrict¬
ed into a pseudopetiole; base and margins of leaves
and sheaths dull green-yellow. Inflorescence one-
flowered, ca. 4.5 cm long, borne on the leaf-sheath
axils; floral bract 1.5 cm long, much shorter than
ovary, the last bract of the peduncle surpassing tin1
articulation ol the pedicel to the ovary and pedicel:
ovary and pedicel ca. 10 mm long, 3 mm thick.
Flowers medium sized or small for the genus, ca.
17 mm long, fleshy, clear yellow-green; perianth
segments subparallel to the column; sepals 16.5—

I 7.5 X 4.5— 5.5 mm, sub-similar with the laterals
somewhat oblique, oblong-elliptic, acute; petals
14.5—15.5 X 2.4— 2.6 mm, oblaneeolate to spatu-
late. margin entire; labellum 14.5-15.5 X 4. 5-5. 5
nun, slightly 3-lobed, rhombic-ellipsoid in general
outline, apex acute, basally cuneate, yellow with 2
series of large dark red or maroon spots or blotches
that run parallel to the calli (4 to 7 on each side),
articulated to the column foot: disc dull yellow, with
two linear calli that are parallel to the main axis of
the labellum, the basal callus is placed iu the basal
hall ol the disc, fleshy, the second callus is placed
in the apical half of the disc and is farinose (mealy);
column ca. 8 mm long, 2 mm thick, green-yellow,
strongly arcuate, hemi-cylindrical. wider at base,
with a poorly defined foot ca. I mm long; anther
unilocular with papillose margin, ca. 3 mm long,
dull ye I low; pollinia 4, unequal. Fruits not seen.

Volume 15, Number 4
2005

Ojeda et al.

Heterotaxis (Orchidaceae)

575

Figure I. Heterotaxis fritzii Ojeda & Carnevali. — A. Plant habit. — B. Perianth. — C. Flower with peduncle and last
two sheaths. — 1). Pollinia. — E. Sepals from top: ventral and dorsal view. — F. Petals from top: ventral and dorsal
view. — O. Leaf in cross section. — H. Apex of leaf. Drawn by Isidro Ojeda (based on paratype, Carnevali 3443, and
supplemented with photographs).

576

Novon

h igure 2. Heterotaxis Jritzii Ojeda & Carnevali. — A. Plant habit. — B. Base of plan! showing partially hidden pseu-
dobulb. — C. Close-up of llower, showing the two cal I i of the labellum. Photographs by W. M. Whitten (based on living
material later preserved as holotype).

Volume 15, Number 4
2005

Ojeda et al.

Heterotaxis (Orchidaceae)

577

Figure 3. Heterotaxis schultesii Ojeda & G. A. Romero. — A. Flower with peduncle showing two last sheaths. — II.
Column with lahellum articulated. — C. Perianth. — D. Anther. Drawn by Isidro Ojeda (based on puratype R. E. Schultes
& I. Cabrera 12788).

Eponymy. Named in honor of William Fritz of
Union, Missouri, who cultivated a plant of this spe¬
cies for several years when lie lived in St. Louis,
Missouri.

Until recently, this species was only known from
a 1994 collection made by one of us (Carnevali)
and C. H. Dodson along the orchid-laden road from
Lita to San Lorenzo. Imbabura, in western Ecuador.
Subsequent trips to this locality by C. H. Dodson
failed to reveal more plants. However, several
plants of the species have recently been collected
from the same general area and brought into cul¬
tivation. The holotype ol the species was prepared
from one of these plants. Heterotaxis fritzii at first
looks morphologically intermediate between H. va-
lenzuelarta and II. sessilis , neither of which grows
nearby. Both II. valenzuelana and H. sessilis have
broader leaves than those of II. Jritzii. Heterotaxis
sessilis is only known in Ecuador from Los Rios and
Guayas at lower elevations and drier areas (e.g.,
H aught 3128 (AMES, MO), but these Ecuadorian
collections might be referable to the entity de¬
scribed as Maxillaria gat une ns is Sehleehter, pos¬
sibly distinct from //. sessilis). Heterotaxis Jritzii su¬
perficially resembles H. sessilis, but is a smaller
plant overall with fleshier leaves that are laterally
(used and triangular in cross section. The lip is
similar in shape to that of Heterotaxis sessilis but
with two calli (versus one callus in H. sessilis ). Phe-
netically. Helerot axis jritzii is. however, more simi¬
lar to H. santanae (Carnevali & I. Ramirez) Ojeda
& Carnevali, with which it shares the small vege¬
tative size, narrow leaves, overall habit, and similar
flowers. The labellum in H. santanae has a rounded
or obtuse apex and it lacks the dark purple trans¬

versal blotches found in the labellar disk ol H. fri¬
tzii, among other floral differences. Heterotaxis san¬
tanae, furthermore, grows on the eastern side ol the
Andes, while the novelty here proposed is appar¬
ently restricted to the Pacific coast of northern Ec¬
uador.

Distribution and habitat notes. Heterotaxis fri¬
tzii is poorly known, since it has only been col¬
lected a few times and only seen alive once by the
authors. At the paratype locality it grew as a sub¬
terrestrial on a moist and shady road bank, along
with many other orchid species ( Scaphyglottis sp.
aff. sumersii C. Schweinfurth, S. prolifera Cogniaux,
Maxillaria cymbidioides Dodson, J. T. Atwood N
Carnevali, Epidendrum ramosum Jacquin). The sin¬
gle plant found then grew slowly in cultivation in
St. Louis, Missouri, for several years, but never
thrived and eventually died without ever flowering
again (see paratype).

Conservation status. Apparently a rare, narrow
endemic in coastal Ecuador. Since it has no horti¬
cultural appeal, it is probably not under any par¬
ticular collecting pressure.

Paratype. ECUADOR. Inihubura: 5—14 km SF ol
Lita along road to Ibarra, 600—850 m, plants collected
along road embankments, 27 Mar. 1994, G. Carnevali &
C. II. Dodson 3443 (CICY).

Heterotaxis schultesii Ojeda & G. A. Romero, sp.
nov. TAPE: Brazil. Amazonas: Rio Urubii. be¬
tween Cachoeira Iraeema & Manaos— Caracarai
road, 6 June 1951, G. 7 Prance, I). Philcox,
11 A. Rodrigues, ./. E Ramos & L. G. Earias
5023 (holotype, NY; isotypes. A, F, HR, INPA,
K. MG, NY. R, S, U. US). Figures 3, 4.

578

Novon

I'igure 4. Heterotaxis schullesii Ojeda & (». A. Romero, flowering habit (based on R. /.. Schultes & I. Cabrera 17488
(AMKS)).

Volume 15, Number 4
2005

Ojeda et al.

Heterotaxis (Orchidaceae)

579

Species liaec II. villosoe (Barbosa Rodrigues) K Barros
similis sed habitu parviore, foliis angustioribus erassiori-
bus, apice lohis magni-inaequilateris, floribus fuseis-cin-
nabarinis, labello angustiore, callo biseriato camoso al>-
horret.

Plant a eaespitose epiphyte, erect, 20—40 cm
high, rhizome abbreviated; roots grayish, about 0.2
cm diam.; pseudobulbs aggregate, 3.5-6 X 1-1.5
cm, oblong-ellipsoid and strongly compressed lat¬
erally. unevenly marked with dense longitudinal
scars upon drying, unifoliate, clothed with 4 to 6
sheaths, the innermost 2 to 4 bearing loliar blades,
these distichously arranged. Leaves coriaceous,
erect on the pseudobulb apex, 17—28 X 1 — 1.7 cm.
oblong, basally attenuate and forming a 1.5—6 cm
long pseudopetiole, apex acute, unequally 2-lobed.
the longer apical lobe 0.5—1 cm longer than the
shorter, both lobes acute, inwardly oblique; leal
blades of the sheaths enveloping the pseudobulb,

1.5— 5.5 X 0.8— 1.5 cm, partially fused at the base
(with a “y" cross section near the articulation),
forming a relatively long pseudopetiole. Inflores¬
cence 1 -flowered, borne on the leaf-sheath axils;
peduncle fleshy, 33—35 mm long, concealed by 5
distichous bracts, the last one surpassing the artic¬
ulation of tin' peduncle to the ovary and pedicel;
floral bract similar to peduncle bracts, membra¬
nous, ea. 5 mm long, triangular, acute, shorter than
ovary; ovary and pedicel 10—1 1 mm long, ea. 2 mm
thick. Flowers medium-sized for the genus, ea. 1.7
cm long., apparently erect on the pedicel, perianth
segments subparallel to the column, yellow brown
to orange or orange-red, fleshy; dorsal sepal 16—18
X 2.8— 3.1 nun. lanceolate to oblong lanceolate, ob¬
tuse, lateral sepals similar to dorsal but slightly
oblique; petals 14-16 X 1. 8-2.1 mm, oblanceolate
to spatulate. acute, the margins papillose from apex
to mid-length; labellum articulate to column foot,

13.5- 14.5 X 3.9-4. 1 mm. slightly 3-lobed, apex
obtuse to obtusely acute; disc with two ealli, these
fleshy, linear, the basal one originating at the base
of the disc and extending to about mid-length, the
apical callus borne near tbe attenuate point of the
labellum blade and from there extending to near
the apex; anther ca. 3.5 mm long, unilocular, the
apical surface papillose; pollinia four, unequal; col¬
umn 7—9 mm long, hemi-cylindric, arcuate, with a
poorly defined foot 1-2 mm long. Fruits not seen.

E pony my. (Named in honor of Richard Evans
Schultes, among the first collectors of this species.

Heterotaxis schultesii resembles II. villosa, but it
is recognized at once by its smaller overall vege¬
tative size (20-40 cm tall). The leaves are thicker
in texture, with fewer cell layers (Ojeda, 2003) and
narrower, compared to the wide leaves of II. villosa.

Most distinctly, the apex of the leaves is conspic¬
uously bilobed. The apical lobes of the leaf are so
strongly unequal that the larger lobe is 0.5—1 cm
longer than the shorter, a dimension that equals or
exceeds the width of the leaf at its apical portion.
This condition is not found in any other species of
Heterotaxis. The labellum of II. schultesii is similar
in shape to that of Heterotaxis sessil is. but the two
fleshy ealli are different. The brownish yellow to
red-orange color of the flowers is unique in Hetero-
taxis.

Heterotaxis schultesii is probably only distantly
related to II. sessilis. According to our results, H.
schultesii lies, within the genus, in a basal position
(Ojeda. 2003; Ojeda et al., 2003).

Distribution and habitat notes. Known from a
few collections made in Amazonian Brazil (Ama¬
zonas) and Colombia (Vaupes). It grows as an epi-
pliyte, and it has been reported as occurring on
savannah trees (e.g., Garay 108). on flooded forests
(e.g.. Garay 91) or along rivers (e.g.. Prance et al
5028). The known collections were made at alti¬
tudes ranging from 180 to 250 m.

Phenology. Flowering collections have been
made in the months of June, August, and Septem¬
ber; no fruiting specimens have been examined.

Paratopes. COLOMBIA. Vauj >es: Rid Piraparand
(trib., Rio Apaporis), Cano Teemeena, 10 Sep. 1952, H.
E. Schultes & I. Cabrera 17488 (AMES. CICY); Cerro dc
Yapoboda. Rid Kuduyarf, 15 Aug. 1960, L. A. Cara y 108
(AMES, (3CY); along Rid Kuduyarf cerea dr Mitu, 12
Aug. I960. L. A. Garay 91 (AMES); Rid Kuduyarf. Ya¬
poboda, savana con arenisca, 25 June 1958, II. G. Barri-
ga. R. E Schultes <8 II. Rlohm 15892 (AMES); Rio Apa-
poris entre Rfo Paeoa y el Rfo Kananarf, 21 June 1951.
R. E. Schultes A' /. Cabrera 12788 (AMES); Rfo Apaporis
entre Rio Paeoa y Rio Cananarf, 21 June 1951, R. E.
Schultes & I. Cabrera 12728 (AMES, K); Rfo Apaporis,
15 June 1952, R. E. Schultes <Y /. Cabrera 10717 (AMES).

New Combinations

Phylogenetic work using ITS 1 and 2 and a suite
of morphological and anatomical characters iden¬
tified 1 I taxa belonging in the Heterotaxis clade,
itself part of the larger Heterotaxis— Ornithidium
complex (Ojeda, 2003; Ojeda et al., 2003; N. II.
Williams et al., in prep.). All share the generalized
habit and vegetative architectures of Heterotaxis de¬
scribed above and the fleshy flowers characteristic
of the Heterotaxis— Ornithidium complex. These
species were: Max illaria discolor Loddiges ex Lou¬
don. M. villosa. II. schultesii. M. maleolens Sehlech-
ter, M. violaceopunctata Keiehenbach f., M. cras-
sifolia, M. superflua, M. santanae Carnevali & I.
Ramirez, II. fritzii, M. equitans, and M. valenzue-

580

Novon

lanu. As indicated above, Harms (2002) already
transferred several of these taxa. all of them occur¬
ring in Brazil, to Heterotaxis.

Two additional species, also represented in Bra¬
zil. Maxillaria equitans and M. valenzuelana, were
treated by Barros as members of the genus Mar-
supiaria. Our analysis strongly suggests that Mar-
supiaria is embedded within Heterotaxis , leaving a
balance of six other species that are yet to be trans¬
ferred to Heterotaxis. Of these, five species were
included in the phylogenetic analysis performed by
Ojeda et al. (2(X)3), while a sixth one. the recently
described Maxillaria microiridifolia I). E. Bennett
& K. A. Christenson, although unavailable to us in
the course of t lie present study, is clearly referable
to Heterotaxis and closely related to (if not conspe-
cific with) Maxillaria santanae Carnevali & I. Ra¬
mirez (transferred to Heterotaxis below). Thus, the
following six new combinations are warranted:

Heterotuxis discolor (Coddiges ex Cindley) Ojeda
& Carnevali, comb. nov. Basionym: Dicrypta
discolor l.oddiges ex Lindley, Edwards’s Bot.
Beg. 25: 91. 1839. 1858. Maxillaria discolor
(Coddiges ex Cindley) Reiehenbach (., in
Walp. Ann. Bot. Syst. 6: 529. 1863. TYPE:
Guyana ("Demerara”). Without precise locali¬
ty. ex Hurt. Coddiges (holotype, K-Cindl.).

As mentioned above, Dicrypta is without ambi¬
guity a nomenclatural synonym of Heterotaxis.

Dicrypta bicolor Paxton ex J. E. Planchon often
had been referred to the synonymy of Maxillaria
discolor (e.g., loldats, 1970: 449; Carnevali & Ra¬
mirez, 2003). We recently had the opportunity to
research this name, and it is evident that its no¬
menclatural and systematic status is unclear. First
of all. the authorship of the name cited by Planchon
(1858) should be “Hort. ex ,|. E. Planchon,” and
the name appears on page 159 (not ‘‘Paxt. ex J. E.
Planchon" or page 73, as cited by Eoldats, 1970:
449 and Carnevali & Ramirez, 2003). “ Dicrypta bi¬
color Bate.’ was also cited in the second supple¬
ment of Condon's Hortus Hritannicus (Coudon,
1839: 630). However, neither Planchon (1858: 159)
nor Coudon (1839: 630) validate the corresponding
horticultural name (k. Gandhi, pers. comm., 2005).
Furthermore, the World Checklist of Monocots
(2004) cites "Dicrypta bicolor (Ruiz & Pav.) Bale-
man ex Coudon. Hort. Brit., ed. 3. SuppC: 630
(1839)” as a synonym of Maxillaria bicolor Ruiz
Pavon, which we strongly oppose, as there is no
evidence linking Bateman’s plant from “Demerara”
(fide Coudon, 1839: 630) to the Andean plant de¬
scribed bv Ruiz and Pavon. At this point we de¬

ferred any nomenclatural action until the status of

Dicrypta bicolor is clarified.

Ileh ‘rotaxis equitans (Sehleehter) Ojeda & Car¬
nevali, comb. nov. Basionym: Camaridium
equitans Sehleehter, Repert. Spec. Nov. Regni
Veg. Beih. < : 1(6. 1920. Marsupiaria equitans
(Sehleehter) Hoehne, Arq. Bot. Estado de Sao
Paulo 2: 71. 1947. Maxillaria equitans
(Sehleehter) Garay, Bot. Mus. Ceafl. 18: 208.
1958. TYPE: Repert. Spec. Nov. Regni Veg.
Beih. 57: t. 63, fig. 242, 1929 (lectotype, des¬
ignated here). EPITYPE: Colombia. Valle:
Cauea Valley, Xarzal. 970-1050 m, 21 July
1922. F. W. Fennel. E. H. Killip & T. E. Hazen
HUM (epitype designated here, AMES, NY).

I The holotype, M. Madera s.n.. for Cam arid i uni
equitans was destroyed at B.]

Camaridium vandiforme Sehleehter, Beih. Bot. Centralbl.
42: 174. 1925. Maxillaria vandiformis (Sehleehter)
C. Schweinfurth, Bot. Mus. I.eafl. 11: 291. 1945.
TYPE: P. ;ru. Loreto: vicinity of Iquitos, 100 m,
dense forest, on living tree. Jan.— Feb. 1937, G. King
10076 (lectotype. designated here, AMES).
[Syntypes by G. Hiibner (120. 124. 140) were de¬
stroyed at B.|

Maxillaria matogrossensis Brade, Arq. Serv. Elorest. I: 46,
t. 3, Fig. 1—8. 1939. Marsupiaria matogrossensis
(Brade) Hoehne. Arq. Bot. Estado de Sao Paulo N.S.,
Formato Maior 2: 71. 1947. TYPE: Brazil. "Mato
Grosso”: ex Hort. Jardim Botanico Rio de Janeiro.

1 I June 1939, Inspectoria Agricola Federal da VIII
Regiao Mato Grosso V.14.399 (holotype, BB not
seen).

Heterotaxis muleoleiis (Sehleehter) Ojeda & Car¬
nevali, comb. nov. Basionym: Maxillaria ma-
leolens Sehleehter, Repert. Spec. Nov. Regni
Veg. Beih. 19: 233. 1933. TYPE: Costa Rica.
Alajuela: Rfo Jesus de San Ramon. 800 m,
I0S02’N, 84Q3 1 ' W, June 1922. A. M. I ire ties
2HI (lectotype designated by Barringer, Eiel-
diana, Bot. n.s., 17: 10. 1986. CR-25985).

| I he holotype of Maxillaria maleolens at B was
destroyed, and Barringer designated the only
known isotype as the lectotype.)

Iletcrotaxis microiridifolia (1). E. Bennett &
Christenson) Ojeda cK Carnevali, comb. nov.
Basionym: Maxillaria micro- iridifolia I). E.
Bennett & Christenson. leones Orchidacearum
Peruvianum. Plate 699. 2001. TYPE: Peru.
Puno: Sandia, Candamo river valley, 550 m,
15 Sep. 1998, M. Cavern H. e.v I). II. Bennett
7823 (holotype. Herb. Bennettianum, Lima,
Peru; isotype, NY).

Volume 15, Number 4
2005

Ojeda et al.

Heterotaxis (Orchidaceae)

581

Heterotaxis santanae (Carnevali & I. Ramirez)
Ojeda & Carnevali, comb. nov. Basionym: Ma¬
xillaria santanae Carnevali & I. Ramirez, Ann.
Missouri Hot. Card. 76: 377. 1689. TYPE:
Venezuela. Amazonas: Atabapo, Cerro Mara-
huaca, 1 1 Nov. 1986, G. Santana I (holotype,
VEN; isotype, MO).

Heterotaxis valenzuelana (A. Richard) Ojeda &
Carnevali, comb. nov. Basionym: Pleurothallis
valenzuelana A. Richard, Sagra, Hist. Fis.
Cuba 11: 234. 1850. Maxillaria valenzuelana
(A Richard) Nash. Bull. Torrcy Bot. Club
34(3): 121-122. 1907. Marsupiaria valenzue¬
lana (A. Richard) Garay, Arq. Jard. Bot. Rio
dc Janeiro 12: 183. 1952. TYPE: Cuba. Va¬
lenzuela, C. Wright 3314 (holotype, P not
seen).

Maxillaria iridifolia Bateman ex Reichenbach f.. Bon-
plandia 2: 16. 1854. Marsupiaria iridifolia (Bateman
ex Reichenbach f.) Hoehne, Arq. Bot. Est. Sao Paulo
2: 71. 1947. TYPE: Cuba, without precise locality,
“Wachst bei Cohubas," E. F. Poeppig s.n. (holotype,
H.O. Reichenbach I. Hb.. \\ not seen).

Dicrypta irisphyta Barbosa Rodrigues, Gen. et Sp. Orch.
Nov. I: I 29. 1877. TYPE: watercolor in Iconographie
des Orchidees du Bresil. Vol. 6: t. 292 ( fide Barbosa
Rodrigues 1996: 420; “Tab. 487 ’ fide Barbosa Ro¬
drigues 1877: 126) RB | Library] (lectotype, desig¬
nated here; reproduced in Barbosa Rodrigues, 1996,
1: 420). I'l'he holotype (,/. Barbosa -Rodrigues s.n.) of
Dicrypta irisphyta was probably destroyed at Barbosa
Rodrigues’ home in Rio de Janeiro, according to P.
Cribb & A. Toscano de Brito in Barbosa Rodrigues,
1996: 30-31.]

Maxillaria valenzuelana subsp. angustata J. T. Atwood,
Icon. PI. Trap. 1371. 1989. Syn. nov. Maxillaria va¬
lenzuelana var. angustata (J. T. Atwood) Senghas,
Schlechter, Orchideen ed. 3, l/B (29): 1779. 1994.
TYPE: Nicaragua. Zalaya, Siuna. F. Ortiz 612 (ho-
lotype, SEE).

As stated by Atwood in the protologue, Maxilla¬
ria valenzuelana subsp. angustata differs from the
type only in the “. . .generally smaller [size of], . .all
its parts, and the scapes and flowers [that] are more
exposed.” We have seen a great deal of material of
this species. The variation we detected along its
geographical range and within populations is con¬
siderable: we hypothesize that M. valenzuelana
subsp. angustata is well encompassed within this
range of size and morphological variation.

Acknowledgments. This publication is the par¬
tial result of the research conducted by the first
author in the course of his M.Sc. degree in System¬
atic Botany. These studies were conducted within
the “Mention en Ecologfa del Posgrado en Ciencias
y Biotecnologfa de Plantas del Centro de Investi-
gacion Cientlfica de Yucatan.” The first author ac¬

knowledges a scholarship by CONACyT that sup¬
ported his graduate work. The authors thank the
curators of the herbaria that loaned us material for
this study or that allowed us to study their herbar¬
ium specimens and living accessions. These are
primarily: AMO, F, FEAS, MEXU, NY, SEE, and
US, although materials from other institutions were
consulted during the course of this research. Norris
11. Williams and Mark Whitten (Herbarium TEAS,
Florida Museum of Natural History, University of
Florida) contributed in several ways, particularly
with the access to their laboratory facilities and
their assistance with technical and theoretical as¬
pects related to this research, besides contributing
their knowledge on the Orchidaceae in general and
of the Maxillariinae in particular. Mark Whitten
also contributed first-hand information on the mor¬
phology of Heterotaxis fritzii and allowed us to use
his photographs. Victoria Sosa (Herbarium XAE.
Institute de Ecologfa, Xalapa) contributed her
knowledge of phylogenetic matters and of the Or¬
chidaceae in general. Robert E. Dressier provided
discussion and miscellaneous information for which
we are enormously grateful. Ivon M. Ramirez
(CICY) carefully read and commented on an earlier
draft of this paper. We are grateful to the staff of
CICY for handling and databasing of the specimens
studied in the course of this research, particularly
Eilia Can, Silvia Hernandez, Filogonio May Pat,
and Jose Euis Tapia. Kanchi Gandhi (HUH) pro¬
vided invaluable help with the intricacies of orchid
nomenclature.

Literature Cited

Atwood, J. T. X I). Mora de Retana. 1999. Tribe Maxilla-
rieae: Subtribes Maxillariinae and Oneidiinae (Orchi¬
daceae). In: W. Burger (editor), Flora Costarieensis.
Fieldiana, Bot. n.s. 40: 1-182.

Barbosa Rodrigues, J. 1996. Iconographie des Orchitises
du Brasil. Edited by S. Sprunger et al. Friedrich Rein¬
hardt Verlag, Basle.

Barros, E. 2002. Notas taxonomicas para ospeeics brasi-
leiras dos generos Epidendrum e Heterotaxis (Orchida¬
ceae). Hoehnea 29: 109—113.

Brieger, E. G. & R. I). Illg. 1972. 0 grupo Heterotaxis do
genero Maxillaria Ruiz & Pavon (Orchidaceae). I’p.
93—99 in Anais da Sociedade Botanica do Brasil —
XXIII Congreso Naeional de Botanica. Recife.
Carnevali, G. 1991. Cytology and the pollinaria in the
Maxillarinae (Orchidaceae). Master of Science Thesis in
Biology, University of Missouri— St. Louis.

- & I. Ramirez. 1989. New or noteworthy orchids

for the Venezuelan flora. VII. Additions in Maxillaria
from the Venezuelan Guayana. Ann. Missouri Bot. Card.
76: 374-380.

- & - . 2003. Orchidaceae, Maxillaria. Pp.

426-454 in J. A. Steyermark, P. E. Berry, k. Yats-
kievych & B. K. Holst (editors). Flora of the Venezuelan

582

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Guayana, Vol. 7. Missouri Botanical Garden Press, St.
Louis, Missouri.

Christenson, K. A. 1999. Maxillaria, an overview. Pro¬
ceedings of tlie I (>t h World Orchid Conference: 279-
292.

Dodson, C. H. 2002. Maxillaria. Native Kcuadorian Or¬
chids 3: 544—574. Dodson Trust, Sarasota, Florida.

Dressier, R. L. 1981. The orchids: Natural History and
Classification. Harvard Univ. Press, Cambridge, Mas¬
sachusetts.

- . 1993. Phylogeny and classification of the orchid

family. Dioscorides Press, Portland, Oregon.

Foldats, K. 1970. Maxillaria (Orchidaceae). Pp. 384—557
in T. Lasser (editor), Flora de Venezuela, Vol. 15, part
4. 1 nst i t n to Rotunicn, Caracas.

Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras. I). 11. Nicolson, P. C. Silva,
J. K. Skog, P. Trehane, N. J. Turland I). L. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Begnum Veg. 138.

Hoelme, F. C. 1947. Reajustamento de algunas especies
de Maxillarias do Brasil, com criayao de dois novos
generos para elas. Arqiv. Bot. Estado Sao Paulo f.m. 2:
65-74.

Iloltzmeier, M. A., W. L. Stern & W. S. Judd. 1 998. Com¬
parative anatomy and systematic of Senghas’s cushion
species of Maxillaria (Orchidaceae). Bot. J. Linn. Soc.
127: 43-82.

Illg. B. I). 1977. Maxillaria brasiliensis Brieg. & I llg. uma
especie nova de orqufdeas da seyao Heterotaxis. Pp.
239—245 in Irabalhos do XXI Congreso Nacional de
Botanica. Bio de Janeiro, Brasil.

Lindley, J. 18.30—1840. Genera and Species of Orchida¬
ceous Plants. Ridgways, London.

Loudon. J. C. 1839. Hortus Britannicus. Longman, Rees,
Orme, Brown, and Green & Longman, London.

McLeish, I., N. IL Pearce, B. B. Adams & J. S. Briggs.
1995. Native orchids of Belize. AA, Balkema, Rotter¬
dam.

Ojeda, 1. 2003. Filogenia del complejo Heterolaxis Until.
(Orchidaceae): Evolueion de la arquilectura vegetaliva
y de los sfndromes de polinizacidn. Tesis de Maestrfa,
Posgrado en Ciencias y Biotecnologfa de Plantas del
CICY. Centro de Investigacion Cientffica de Yucatan.
Mt ■rida, YucatAn, Mexico.

- . G. Carnevali, W. M. Whitten & N. II. Williams.

2003. Phylogeny of Heterotaxis Lindley complex (Ma-
xillariinae): Evolution of vegetative architecture and
pollination syndromes. Lankesteriana No. 7: 15—47.

Planchon, J.-E. 1858. Hortus Donatensis. \\. Remquet et
Cie.. Paris.

Senghas. K. 1993. Maxillarinae. Pp. 1727-1776 in Brie-
ger et al. (editors), Die Orehideen Ed. 3. Band l/B Lie-
ferung 28. Bogen 108-111. Paul Parey, Berlin and
Hamburg, Germany.

Whitten, W. M.. N. II. Williams & M. W. Chase. 2000.
Subtribal and generic relationships of Maxillarieae (Or¬
chidaceae) with emphasis on Stanhopeinae: Combined
molecular evidence. Amer. J. Bot. 87: 1842—1856.

World Checklist of Monocots (2004). The Board of Trust¬
ees of the Boyal Botanic Gardens, Kew. Published on
the Internet; http://www.kew.org/monocotChecklist/ lac-
cessed 08 Mav 2(X)5],

A New Species of Croton sect. Luntia (Euphorbiaceae)
from Western Amazonian Brazil

Ricardo de S. Secco,' Paul E. Berry ,- and Carlos S. Rosario 1
1 Departamento de Botaniea, Museu Paraense Emilio Goeldi, Caixa I ostal .199,
Belem — PA, Brazil, rsecco@museu-goehli.br
^’Department of Botany, University ol Wisconsin, 132 Birge Hall, 430 Lincoln Drive,
Madison, Wisconsin 53706, U.S.A. peberry@wisc.edu

Abstract. A new species of Croton, C. subasper-
rimus, is described and illustrated from a previous¬
ly unexplored area of Amazonas Slate, Brazil. It
belongs to Croton seel. Luntia subsect. Cuneati be¬
cause of its pinnate leaf venation, slightly redupli¬
cate pistillate calyx, partially bisexual basal cy-
mules, presence of lepidote trichomes, low stamen
number, and multifid stigmas. It differs from related
species in its stellate-lepidote trichomes that some¬
times have tall porreet central rays and in its small¬
er, concolorous leaves with an obtuse base and slip-
itate glands along the margins.

Key words: Amazonia, Brazil, Croton, Euphor¬
biaceae, section Luntia.

Croton h. is the second largest genus of Eu¬
phorbiaceae, with over 1200 species distributed
across the tropics, and it is most densely distrib¬
uted in the West Indies and in Central and South
America (Webster, 1994; Govaerts et al., 2000).
The genus includes trees, shrubs, herbs, and lianas,
and plants are generally monoecious or occasion¬
ally dioecious, with various kinds of stellate and
lepidote trichomes. The inflorescences usually have
the pistillate flowers positioned basally and the sta-
minate ones apically, with the anthers indexed in
bml and the female petals generally reduced or ab¬
sent. The Brazilian species have not been revised
taxonomically since tin* treatment by Mueller
(1873) in Flora Brasiliensis, although many indi¬
vidual taxa have been described since then. Recent
treatments on South American Croton such as those
of Cordeiro (1992), Secco (1992). Murillo (1999).
and Webster et al. (1999) are still insufficient to
determine all the species of Neotropical Croton. In
the three large herbaria of Amazonian Brazil (IAN,
INPA, MG), there are still many unidentified or
misident died specimens of Croton, and many ol
these cannot be matched to any currently described
species.

The species described and illustrated below is a
distinctive one that was recently collected on a he¬

licopter trip to a remote and very poorly collected
area of western Amazonian Brazil.

Croton subasperriinus Secco. P. E. Berry & Ro¬
sario. sp. nov. TYPE: Brazil. Amazonas: inun.
Canutama, rio Mucuim, margem direita do rio
Purus, fazenda Morada do Sol, 22 Nov. 2000
(fl. fr). C. S. Rosario & A. S. L. Silva P03—7—
B559 (holotype. MG; isotypes, INPA, MO, RB,
W IS). Figure 1.

Frutex monoieus, ramis dense trichomatibus lepidotis
obtectis. Folia venatione pinnata. marginilnis crenatis,
basi glandularumpares instructa, supra sparse pubeseen-
tia. infra dense pubescentia. Inflorescentiae in thyrsis ra-
cemoideis terminalibus, cymulis inferioribus saepe bi-
sexualibus. Flores staminati pedicellati, pedicellis
tricliomatibus stellato dendriticis obsitis. Flores pistillali
breviter pedicellati, petalis 5 reductis vel obsoletis. Frue-
tus dense lepidoto indumento obtecti. Ab C. subcoriaceo
foliis minoribus concoloribus apicis acurninati et glandulis
stipitatis marginalibus differt.

Shrub 1.5-2 m tall: stems densely covered by
lepidote or stellate-lepidote, asperous trichomes.
Leaves chartaceous, alternate, 4—10.5 X 1.5—4 cm,
pinnately veined, elliptic to elliptic-lanceolate, with
lepidote to stellate-lepidote trichomes (sometimes
the trichomes with a central porreet ray 0. 5-0.6
mm tall), the adaxial side sparsely pubescent, with
trichomes concentrated along the midvein, the ab-
axial side more densely pubescent (especially when
young), apex acuminate, base obtuse, margin slight¬
ly crenate and with small stipitate cylindrical
glands ca. 0.3 mm diam. and 0.2-0. 3 nun long,
spaced 3—8 nun apart along the margin; petiole 4—
10 mm long, densely covered by ferrugineous in-
dument, with a conspicuous pair of saucer-shaped
glands 1—1.3 mm diam. at the apex on the adaxial
to medial side, occasionally with a smaller pair of
glands distal to the larger pair; stipules deciduous.
Inflorescence a terminal, raceme-like thyrse 9-17.5
cm long, the rachis densely covered by stellate-
lepidote trichomes, tin* male flowers distally
grouped in fascicles of 3 to 5. the female flowers

Novon 15: 583-585. Published on 12 December 2005.

584

Novon

ilgUrP K y°Um s"l>"*l"’rnmus Secco, P. K. Berry & Bosdrio. —A. Branch with inflorescences showing distal staminate
flowers and proximal pistillate ones. — B. Pistillate flower, with one calyx lobe removed to show the ovary. — C. Pistillate
flower, lateral view. —I). Fruit. K. Staminate flower. — F. Stamen. — G. Staminate petal, internal side shown. —II.
Staminate sepals, internal and external surfaces shown. Drawn from the holotype, K»s<iri<> <£ Silva P03-7-3559.

basal and solitary or sometimes in cymules with
subtending male flowers. Staminate flowers with a
slender pedicel 1-2 mm long, covered by stellate-
dendritic trieliomes; sepals 5, valvate to partly free
in bud, elliptic-lanceolate, 2— .1 mm long, externally
with a dense indument of stellate-lepidote tri¬

eliomes, internally with a sparse indument of villose
trieliomes; petals 5, elliptic-lanceolate, 1.5— 2.5 mm
long, covered on the inside by villose trieliomes.
especially along tin* margins and internally near the
base; stamens 10 to II, the filaments filiform and
4—4 mm long, densely covered by villose trieliomes.

Volume 15, Number 4
2005

Secco et al.

Croton subasperrimus from Brazil

585

ihe anthers hasifixed. elongate, deflexed from die
apex of the filament. 1 .2-1.6 mm long X 0.4 mm
wide, dorsally pubescent. Pistillate flowers with a
thick pedicel 2-3 mm long densely covered by stel-
late-lepidote trichomes; calyx 5-lobed. slightly re¬
duplicate in bud, the lobes 3-3.5 mm long, elliptic
to lance-elliptic, densely stellate-lepidote on both
surfaces; petals 5, nearly obsolete, 1-1.5 mm long,
formed by tufts of stellate trichomes; ovary globose,
ca. 2.5 mm diam., densely stellate-lepidote; styles
3, each branched up to 5 times, the branches 3—4
mm long, rugose, with lepidote trichomes at the
base, glabrous from the middle to the apex, fruit a
capsule ca. 8 mm diam., densely lepidote. meri-
carps 3(4); seeds not seen.

Habitat and distribution. Known only from the
type locality, in a remote and poorly collected re¬
gion of western Amazonian Brazil, along a tributary
of the rio Purus in Amazonas State. Il is apparently
a riverine species, probably subject to seasonal
flooding.

Etymology. The specific epithet refers to the
somewhat asperous leel of the leaves due to the stdl
trichomes on the lower surface.

By virtue of the mostly lepidote trichomes, pin¬
nate, subentire leaves with hasal glands, stamens
10 or I 1 . and pistillate flowers with pedicels shorter
than the calyx, and multifid stigmas, Croton subas-
perrimus most closely fits the definition of Croton
sect. Lamia (Rafinesque) G. L. Webster subsect.
Cuneati G. L. Webster, as defined in the sectional
synopsis of Webster (1993). Ibis section contains
about ten species that occur mainly in the Amazon
region and in the Guiana Shield. Both C. subcori-
aceus Jablonski and C. cuneatus Klotzsch belong in
the same subsection and can have similarly crenate
leaves. Another as yet undescribed species from
southern Amazonia apparently belongs to this sub¬
section as well and has the same kind of stipitate
glands along the leaf margins as our new species

does. Croton subasperrimus most closely resembles
C. subcoriaceus, but it differs from that species by
the smaller, coneolorous leaves with an acuminate
apex and obtuse base, the margin crenate and with
stipitate glands, and fewer stamens. The wide¬
spread C. cuneatus in the same subsection differs
in its leaves with a denser covering of smaller lep¬
idote trichomes on the abaxial side, a cuneate base,
and more numerous stamens.

Acknowledgments. The senior author wishes to
thank the Brazilian GNPq for their financial support
of this work (Project N° 301.252/86—6). \\ ill i am A.
Rodrigues, from the Universidade Federal do Pa¬
rana. Brazil, assisted us with the Latin diagnosis,
and Carlos A. Alvarez made the illustration of the
new taxon.

Literature Cited

Conleim. I. 1992. Flora da Serra do Cipd. Minas Gerais:

Euphorbiaceae. Bol. Bot. Univ. Sao Paulo Id: 169—21 1.
Govaerts, IT. 1). G. Frodin. & A. Badcliffe-Smith. 2000.
World Checklisl of Euphorbiaceae (and Pandaceae).
Vol. 2. Euphorbiaceae: Croton to Excoecariopsis. Royal
Botanic Gardens, Kew.

Mueller, J. 1873. Euphorbiaceae: Croton. In: C. f. P. Mar-
tius & A. G. Fielder (editors). Flora Brasiliensis I 1(2):

82-273.

Murillo, J. 1999. Composicion y distribucidn del genero
Croton in Colombia, con cuatro especies nuevas. Cal-
dasia 21: 141-166.

Secco. R. S. 1992. Notas sobre as lianas do genero Croton
E. (Euphorbiaceae). Bol. Mus. Paraaense "Emilio Goel-
di," ser. Bot. 8: 265—281.

Webster, G. I,. 1993. A provisional synopsis of the sec¬
tions of the genus Croton (Euphorbiaceae). Taxon 42:
793-823.

- . 1994. Synopsis of the genera and suprageneric

taxa of Euphorbiaceae. Ann. Missouri Bot. Gard. 81:
33-144.

- . P. E. Berry. W. S. Ambruster, H-J. Esscr. E. J.

Gillespie, W. J. Hayden, G. A. Levin, IT A. Secco &
A. V. Heald. 1999. Euphorbiaceae. In: P. E. Berry, k.
Yatskievych & B. k. Holst (editors). Flora of the Ve¬
nezuelan Guayana 5: 72—228. Missouri Botanical Gar¬
den Press, St. Louis.

Plinia nana (Myrlaceae), a New Species from Minas Gerais, Brazil

Marcos Sobral

Departamento de Botanica UFMG, Caixa Postal 486, 31270-910 — Brio Horizonte,

MG. Brazil, sohral@ufnig.hr

Abstract. Plinia nana (Myrtaceae), a new spe¬
cies I mm the campus rerrados of the Brazilian state
of Minas Gerais, is described, illustrated, and com¬
pared with the related Plinia cauliflora (Martins)
Kausel, from which il is set apart by the shrubby
habit and sessile cordate leaves.

Kkshmo. Plinia nana (Myrtaceae), imia nova es-
pecit' dos eampos cerrados de Minas Gerais, Brasil,
e descrita, ilustrada e eomparada com Plinia cau¬
liflora (Marinis) Kausel, da qual se distingue prin-

cipalmente pelo habito arbustivo e pelas folhas ses-
seis e cordadas.

Key words: Brazil, Myrtaceae, Plinia.

The American genus Plinia extends from Antil¬
les to Uruguay and comprises about 30 species
(Landrum & Kawasaki, 1997), although this esti¬
mate does not include some eastern Brazilian tree
species commonly known as “jaboticabas,” widely
cultivated for their edible fruits, as for example Pli-

l igurc I. Plinia main. Scanned image of isotype at MO (Hutschbach cl al. 6 957H).

Novon 15: 586-589. Published on 12 December 2005.

Volume 15, Number 4
2005

Sobral

Plinia nana (Myrtaceae)

587

figure 2. Plinia nana. Branch (Alves <£ Sobral 168).

ilia cauliflora (Martins) Kausel, I*, jabot icaba (Vel-
lo/o) Kausel, I1. Irunciflora ((). Berg) Kausel (Kau-
sel, 1956), P. phitrantha (Kiaerskou) Sobral, and P.
grand if ilia (Mattos) Sobral (Sobral. 1994). These
jaboticaba species are generally assigned to Myr¬
ciaria 0. Berg (Berg, 1855-1856. 1857-1859). but
Kausel ( 1956) proposed their inclusion in the genus
Plinia due mainly to their seeds with separate pla¬
no-convex cotyledons rather than the fused ones of
Myrciaria. This last genus is characterized mainly
by fused bracteoles. a calyx-tube deciduous at an-
thesis, and seeds with generally fused cotyledons,
while Plinia presents distinct bracteoles, a persis¬
tent calyx-tube, and seeds with separate, plano¬
convex cotyledons. Such limits unfortunately are
not clear; some Myrciaria present plano-convex
cotyledons and some Plinia may have a tardily de¬
ciduous calyx-tube. Considering such characters,
assigning some species to one genus or the other

may be a matter of taxonomic opinion. Since jabo-
licabas encompass character states also found in Pli¬
nia, I currently prefer to include them in Plinia.

During a floristic inventory of the carnpos cer-
rados and carnpos rupestres of central Minas Ger¬
ais. a shrubby species of jaboticaba was collected
and is herein described as new.

Plinia nana Sobral. sp. nov. I T 1'b,: Brazil. Minas
Gerais: Diamantina, rod. BR-367, km 71 1, 23
Oct. 1999, G. Hatschbach, R. Spiehiger, A. C.
Cervi & E. Barbosa 69578 (holotype. MBM;
isotype, MO). Figures 1-3.

Species haee Plinae eauli/lorae proxima. a i|iia habitu
suffruticoso et fo Ins sessihs eordatisque reeedit.

Shrub 0.5-1 .5 m: plants glabrous except for scat¬
tered trichomes on the flowers; cortex smooth, ex¬
foliating; young twigs smooth, gray or sometimes

588

Novon

hf?ure 3. Plirua nana. Flower hud, old (lower and immature fruit (Alves & Sobral 167). Seale har 3 mm.

markedly reddened in living material. Leaves ses¬
sile or subsessile; petioles absent or to I X 1 mm;
blades cordate, 24—50 X 15 — 40 mm. decussate,
slightly discolored or concolorous; apex acuminate
or mueronate; base cordate; midvein sulcate adax-
ially and markedly salient abaxiallv; secondary
veins 8 to 15 pairs, visible on both faces and salient
at least abaxially; marginal vein formed by the
arches of tin* secondary veins, somewhat discontin¬
uous, 2-2.5 mm from the margin, occasionally a
second vein 0.5—1 mm from the margin visible, the
margin itself slightly revolute; petioles absent or to
I 1— - aim. Inflorescences ramiflorous, uniflorous
(occasionally two or three arising from tin* same
place and apparently 2- to 3-florous), the fertile
axis 1-2 X 0.5 mm. with up to 5 series of bracts
distally increasing in si/e. the proximal ones ellip¬
tic, 0.11 — I X 1—1.2 mm and the distal ones lance¬
olate. 3—3.5 X 2—2.5 nun; pedicels absent in flow¬

ers or to I X 0.7 mm, elongated in fruits to 1 — 1 X
0. (-0.8 mm; bracteoles elliptic-lanceolate, to 2 X
1 mm; buds elliptic to pyriform, white. 4-5 X 3-4
mm. the ovaries with scattered gray trichomes 0.1
mm; calyx-lobes 4, well defined in bud but the sta-
minal ring and calyx-tube tearing open outside at
anthesis, elliptic-lanceolate to triangular-laneeo-
late. 2—3 X 2. 2-2. 5 mm; petals 4, orbicular, white.
5 X 4-5 mm; stamens 100 to 120, 3-4 mm, the
anthers elliptic, 0.4 X 0.2 mm, without apical
glands; calyx-tube about I mm; style to 5 mm, stig¬
ma punctiform; ovary bilocular with 2 centrally at¬
tached ovules per locule. Fruits globose, purple to
black when ripe, to 20 mm diam., glabrous or
sometimes softly puberulent, crowned by the per¬
sistent and somew hat enlarged calyx-lobes (to 4 X
3 mm); seeds reniform, 5-6 X 4 mm, with mem¬
branous testa and discrete, plano-convex cotyle¬
dons, without evident hypocotyl.

Volume 15, Number 4
2005

Sobral

Plinia nana (Myrtaceae)

589

Affinities. This species is related to Plinia cau-
liflora (Martius) Kausel, from which it can he dis¬
tinguished by the following characters:

la. Tree let or tree to 10 m high, inhabiting forests;
leaves evidently petiolate, the petioles 3 X 0.5
mm; blades membranous, with 2 well-defined
marginal veins and an acute to obtuse base . . .

. Plinia cauliflora

lh. Shrub to 1.5 m high from campos cerrados;
leaves sessile or with petioles, when present, to
1 X 1 mm; blade chartaceous, with 1 (rarely 2)
marginal vein, this occasionally discontinuous,
and cordate base . Plinia nana

Habitat and distribution. Plinia nana was col¬
lected in campos cerrados and campos rupestres at
the municipalities of Diamantina. Felicio dos San¬
tos, Itacambira, and Sao Gonyalo do Rio Preto in
central Minas Gerais.

Phenology. Flowers were collected in August
and October and fruits in October and November;
according to collectors, ripe fruits have a very sweet
taste.

Etymology. The epithet nana, from the Latin
word for dwarf, alludes to the small size of the spe¬
cies. It is noteworthy that no shrubby species of
jabotieaba was known until now.

Paratypes. BRAZIL. Minas Gerais: Felicio dos San¬

tos. 8 Aug. 2004, P. Viana & N. Mata 1856 (BHCB).
Itacambira: 30 Nov. 1984, R. Harley et al. (SPE 86241),
13 Nov. 2001, A. Tozzi & M. Alencar 2001-480 (UEC);

1 7° I ()'()2"S, 43°07'00"W, 9 Nov. 2002. F Mazine et al.
662 (BHCB, ESA). Sao Gonyalo do Rio Preto: Parque Est.
Rio Preto, Ribeirao das Eguas, HI Nov. 1999. ./. htmbardi
2548 (BHCB); Poyo dos Veados, 18 Oct. 2000. ./. Ixtm-
bardi 4084 (BHCB); 3 Oct. 2003, P. Viana et id. 1248
(BHCB); 15 Oct. 2003, T. Alves 17. Sobral 167 (BHCB).

Acknowledgments. Thanks to Gert Hatschbaeh
(MBM) and Fiorella Mazine (ESA) for sending me
duplicates of their collections; to Julio Lombardi
(HRCB) and Pedro Viana (BHCB) for their kind
help and discussions; to the stalf of the Rio Preto
State Park for their gracious cooperation; and to
Victoria Hollowed and an anonymous reviewer for
valuable suggestions on this paper.

Literature Cited

Berg, O. C. 1855-1856. Revisio Myrtacearum Americae.
Linnaea 27: 1—472.

- . 1857—1859. Myrtaceae. In: k. E P. von Martius

(org.) FI. Brasiliensis 14(1): 1-656.
kausel, E. 1956. Beitrage zur Systematik tier Myrtaceen.
Ark. Bot. 3: 491-516.

Landrum. L. R. N M. I .. kawasaki. I99t. Ihe genera ol
Myrtaceae in Brazil: An illustrated synoptic treatment
and identification keys. Brittonia 49: 508-536.

Sobral, M. 1994. Duas novas especies e seis novas eom-
binayoes em Plinia (Myrtaceae) do Brasil. Hoehnea 21:
199-205.

Las Especies del Genero Lonchocarpus seel. Lonchocarpus
(Leguminosae, Papilionoideae: Millettieae) para Bolivia

Mario Sousa S.

Depart amento <le Botdnica, Instiluto tie Biologfa, Universidatl Naeional Autdnoma de
Mexico, Apdo. 1 ostal 70-367, 04310 Mexico, D.F. sousa@servidor.unani.rnx

Kksumkn. Sc Ikicc una relacibn de las especies
de hmcliocarpus sect, lonchocarpus para Bolivia.
De las cinco especies conocidas, euatro resultaron
nuevas para la ciencia. Se deseriben. llustran y ma-
pean Lonchocarpus lomentaceus, /,. neei, L. riparius
y L. scricophyl I us, todas el las endemieas a Bolivia.

I -as eineo especies tie la seel. Lonchocarpus se con-
trastan en una clave dieotdmiea.

Ahstkact. Aii account of section lonchocarpus of
the genus lonchocarpus for Bolivia is given. Of the
live species found, four were new to science, all of
them endemic to Bolivia, hmcliocarpus lomenta¬
ceus, L. neei, L. riparius , and L. sericophyllus are
described, illustrated, and mapped. A key to dis¬
tinguish the species of section lonchocarpus is pro¬
vided.

Key words: Bolivia, Leguminosae, lonchocar¬
pus, sect, hmcliocarpus.

Kl genero hmcliocarpus kunth (Fabaceae: Millet¬
tieae) comprende alrededor de 140 especies neotro-
pieales y una espeeie, hmcliocarpus sericeus (Poiret)
kunth ex DC., extiende su distribut ion en la costa
occidental tit* Africa ecuatorial. La sect, hmchocar-
pus consta de alrededor de 2t especies, es la mejor
representada en Sudameriea eon 15 especies, v de
ellas 12 endemieas; en Mexico y Centro America se
euentan eon 13 spp., de ellas 9 son endemieas; y de
las euatro de Las Antillas dos son endemieas; una
espeeie esta representada en Africa.

Lonciiocarpi s skct. Lonchocarpus-. Lonchocarpus

SR RICH IS (PotKKT) kt NTH KX DC.

(’I V ITS CONSKKV ANIH s)

La sect, hmcliocarpus se caraeteriza por: corteza
interior generalmente exuda resina rojiza al eorte;
folfolos generalmente eon el apice aemninado; (lo¬
res con bracteolas grant les, retlontleadas, aplieatlas
v freeuentemente eubriendo la base tlel ealiz; ealiz
trunco o easi trunco; pdalos en general densamente
eaneseente serfeeos atlaxialmente; el estandarte
abaxialmenle eaneseente pilosulo en el centre tie
la base hasta generalmente el tereio inferior de la

lamina, asf como la una (partieularmente en las es¬
pecies tie Smlameriea); mecanismo floral fuerte-
mente explosivti; legumbre indehiseenle, la sutura
vexilar generalmente ensanchada; plantulas eon los
ctitiledones epfgeos, eofilos opuesltis, simples.

Ln este artfculo se deseriben como nuevas a cua-
tro de las eineo especies tie esta seccion para Bo¬
livia.

Ln Bolivia, ademas tie las especies tit' la sect.
hmcliocarpus existen hmcliocarpus hedyosmus Mi-
t|iiel. miembrt) tlt'I grupo eon esc mismo nombre, y
L. angulatus Ihicke, L. densiflorus Benlham y L.
negrensis Bentham, tres especies tpie Azevetlo-Toz-
zi (1989) propone deberfan st'r transferitlas a l)e-
guelia Aublet. Si bien reeientemente Muellera flu¬
vial is (Linthnan) Burkart fue transferida a
hmcliocarpus (Fortunato y Palese, 1997) esta es
una tlecision taxondmiea tpie no comparto. Fs mas.
L. nudiflorum Burkart, titra espeeie t|ue tambien se
encuentra en Bolivia, deberia pasar a Muellera L.f.,
pert) t'n el presente artfculo no tengo la intent ion
formal tie hacerlo. Sin embargo, existen earaeteres
t|ue estan en las yemas vegetativas. en las inflores-
eeneias y en las flores 1 1 nt* sustentan a Muellera
como genero aparte tie hmcliocarpus.

Lonchocarpus lomentaceus M. Sousa, sp. nov.

I IPO: Bolivia. Dept. Beni: Prov. Vaea I )iez. ca.

5 km SW of Kiberalta, heavily disturbed flood
plain forest, annually inundated, I 1°02'S,
66°07'W. alt. 220 m, 3 June 1982. ./. C. Sol¬
omon 7939 (holotipo, M EXU; ist)tipos, MO.
MEXU). Figuras I. 2.

f rutex ca. .5 m alius, glaber; folia 5(7)-foliolata, foliolis
cllipticis, epunctalis. Inflorescentiae 7-11.5 cm longae.
axillares. Legumen lomentaceum, curvatum vel arcuatum,
3. 5-9.5 cm longum, 1.2-1. 4 cm latum, sublignosum,
oblongum vel lineare, 1 ad 6-seminale, inter semina supra
suturam carinalem et valvas constrictum, ubi sejitis in
articulis uniseminalibus legumen frangentibus, sutura ve-
xill ari ad 3 mm lata, sulcata, duohus niarginibus crass is
limitata.

Arbustos ca. 3 m alto; corteza interior con algo de
(Initio resinoso al eorte; ramas tt'rt'tes. aeostillatlas,
sdlitlas, glabreseentes, eon lenticelas t>rbieulares a

Movon 15: 590-598. Published on 12 Deckmbeb 2005.

Volume 15, Number 4
2005

Sousa

Lonchocarpus sect. Lonchocarpus

591

I'igura 1. Lonchocarpus lomenlaceus M. Sousa. — A. Kama con hojas. infrutescencias y legumlucs. K. Legumlire
moslrando cl margen vexilar. (A. B, ./. (■. Solomon 7939.)

elfpticas; estipulas eaducas, no vistas; pecfolo 2.8—
3.5 cm largo; raquis foliar 3—5 cm largo, marcada-
mente canaliculado, glabro; hojas 5(7)-folioladas; fo-
liolos (4— )5. 5-9.5 cm largo, (2.5-)3-5 cm ancho,
elfpticos, cartaceos a subcoriaceos, epunteados. pero

con las nervaduras do 3° y 4° orden translucidas, la
base cuneada, el apice acuminado, el haz y el enves
glabrescentes a glabros; nervacion camptodroma,
nervaduras primaria v secundarias ligeramente real-
zadas en el enves, nervaduras laterales 7 a 9. Infru-

592

Novon

I'igura 2. Mupa (It* Bolivia con la distribucirin esparial de Ijonchocarpus lomentaceus VI. Sousa (■). L neei M. Sousa
l A I. /.. riparius M. Sousa ( + ), L. sericophyllas VI. Sousa (#|.

teseencias 7—1 1.5 cm largo, axi lares, cortamente pe-
dunculadas; floraeidn desconoeida; pedunculos
fructfferos 1.3—2 mm largo: pedicelos fruetfferos 2—
3 mm largo; bracteolas persistentes, opuestas, apli-
eadas al caliz, orbieulares, 0.9-1 mm largo, 0.9—1
mm audio. Flores deseonocidas; caliz (presente en
t‘l (into) eiatiforme, 2.9— 3. 1 mm largo, ea. 3 mm
ancho, eaneseente serfceo, epunteado, eorola des-
eonoeida. Legumbre lomentaeea. marcadamente eur-
vada a arqueada, algo aplanada, 5.5— 9.5 cm largo.
1.2— 1.4 cm audio, sublenosa, oblonga a linear, con
I a () semillas, glabra, eonstricta entre las semillas
sobre la sutura carinal y valvas, presentando septos
que fragmentan a la legumbre en artfcnlos mouos-
pemios, la base atenuada a largamente atenuada. el
apice rostrado. la sutura vexilar liasta 5 nun ancho,
surcada y bordeada »l<* 2 margenes gmesos de 1—2
mm ancho, la sutura carinal aquillada. Semillas in-
maduras.

Distribut ion y hdbitat. Conocida solamente de
la coleec ion tipo del Dept. Beni. Provincia Vaca

Diez. En planicies selvdticas alteradas. inundables
anualmente.

Fenologia. Con frutos easi maduros a principio
de junio.

El epfteto se refiere a la muy particmlar legumbre
lomentaeea de esta especie.

Esta especie es descrita eomo una novedad por
no existir representantes con legumbre lomentaeea
entre las especies bifloras y arbdreas del genero
Lonchocarpus. Cabe destacar que en el grupo de
especies de bejucos lenosos, propuestas como De-
guelia por Azevedo-Tozzi (op. cit.), Lonehocarpus
densijiorus Bentham presenta una legumbre similar.

Lonehocarpus neei M. Sousa, sp. nov. TIPO: Bo¬
livia. Dept. Santa Cruz: Prov. Florida. 3 km SW
ol Angostura, gorge ol Rib Piraf. along small
side stream on N side of canyon and highway;
disturbed areas of subtropical semideciduous
forest, 18°10'S, 63°33'W, alt. 7(K) m, 27 Die.
1992, M. Nee 43376 & I. Vargas C. (holotipo,
MEXU; isotipos. NY, US/). Figuras 2. 3.

Volume 15, Number 4
2005

Sousa

Lonchocarpus sect. Lonchocarpus

593

Figura 3. Lonchocarpus neei M. Sousa. — A. Rama con hojas e inflorescencias. — B. C^liz con pedicelo y braeteolas.
— C. Estandarte. — D. Ala. — E. Petalos de la quilla. — F. Tubo estaminal. — G. Gineceo. — H. Infrutescencia, rnos-
trando a las legumbres en diferentes caras. (A-G, M. Nee 43376 e /. Vargas ; H, /. G. Vargas C. 5083.)

594

Novon

Arbor 7-20 m alta. Rami brunneo-flavidi, velutini vel
glabrescentes. Folia 5 ad 7-foliolata, foliolis translucido-
punctatis. Inflorescentiae (4— )6— 9 cm longae, longipedun-
culatae, floribus I 1 — 12 mm longis; petalis roseo-violaceis.
Legumen 6-7 (— 13) cm longum, coriaccum vcl subligno-
sum, dense velutino-ferrugineum vcl brunneo-flavidum,
base atlcnuata; sutura vexillari ad I mm lata, sulcata, per
duo margines angustos marginata.

Arboles 7—20 m de alto; corteza interior con flui-
do resinoso al corte; ramas teretes, ligeramente
acostilladas, solidas. de moderadamente pardo-
amarillento velutinas a glabrescentes, con lentice-
las elfpticas a largamente elfpticas; estt'pulas pronto
eaducas, 0.9— 1.1 mm largo, 1 — 1.1 mm ancho, or-
biculares transversalmente elfpticas; pecfolo 2.8-
3.6 cm largo; raquis foliar 3—6.7 cm largo, marca-
damente canaliculado, velutino como las ramas;
liojas 5 a 7-folioladas; foliolos 4.5-8 cm largo, ( 1 .7—
)2-4 cm ancho, elfpticos en ocasiones lanceolados.
carlaceos a subcoriaceos. translucido pimteados. la
base cuneada a redondeada, el apice acuminado a
caudado, el liaz y el enves de moderada a espar-
cidamente velutinos, glabrescentes; nervacidn
camplddroma, nervadura primaria y secundarias li-
geramente realzadas en el enves. nervaduras late-
rales 6 a 8(a 10). Inflorescencias (4— )6— 9 cm largo,
axi lares, largamente pedunculadas; floracion tardfa;
pedunculos florales 2—2.5 mm largo, delgados, es-
paeiados; pedicelos 2.5—3 mm largo; bracteolas
opuestas. aplicadas al ealiz. orbieulares a lignlares.
0.9— 1.1 mm largo, 1-1.1 mm ancho. Flores 11-12
mm largo; caliz eiatiforme, easi trunco, 3—4 mm
largo, 4—5 mm ancho, pardo-amarillento serfceo,
Iranslticido punteado; corola rosado-violaceo, trans-
ltieido punteada, todos los pelalos canescente se-
rfceos; estandarte rcllexo, su lamina orbicular a
transversalmente elfptica. 10-12 mm ancho; ovario
densamente serfceo. Legumbre, recta, aplanada, 6-
7 ( — 1 3) cm largo, 2-3.1 cm ancho, coriacea a su-
blenosa, elfptica a oblonga, con 1 a 3 semillas, den¬
samente ferrugfnea a pardo-amarillento velntina,
constricta entre la semillas sobre las suturas, la
base atenuada a largamente atenuada, el apice api-
culado a rostrado, la sutura vexilar hasta I mm
ancho, surcada y bordeada de 2 margenes angostos
de 1.6— 1.8 mm ancho, la sutura carinal aquillada.
Semillas reniformes, pardo-amarillentas, ea. 19 mm
largo, ea. 13 mm ancho, ca. 3.8 mm grueso.

Distribucidn y habitat. Solo se conoce de las
estribaciones continentales de Los Andes, en la
Prov. Florida, en el Dept, de Santa Cruz. Crece en
habitats riparios, con Vitex cymosa Bertero ex
Sprengel, Licaria Aublet. Sapindus saponaria L. en
vegetaeidn secundaria de selvas subcaducifolias

subtropicales, y en selvas caducifolias en pendien-
tes. a elevaciones de 700 a 1150 m.

Fenologia. Se ha colectado en botones florales
a principios de noviembre, en plena floracion a fi¬
nales de diciembre, y en lructificacion a finales de
mayo y principios de agosto.

Esta espeeie se dedica a Michael Nee (1947—)
amigo y gran botanico conocedor de la flora de
Mexico y Bolivia, y reconocido especialista en la
familia de las Solanaceae, quien actualmente se en-
cuentra realizando la Flora de la Region del Parque
Nacional A mho to. en Bolivia.

l)e las especies de la sect. Lonchocarpus en Bo¬
livia. esta es la que tiene las (lores nuts gramles y
los pelalos de color rosado-violeta a diferencia de
las otras, eon flores chicas y blancas o de color
crema; sus legumbres son caracterfsticamente fe-
rrugineo a pardo-amarillento velutinas.

Paratipos. BOLIVIA. Dept. Santa Cruz. I'niv. Florida.
88 km SW of Santa Cruz on road to Samaipata, I8°08’S,
63°41'W, all. 1000 in. 4. hnvrence & I. Vargas II (LISZ);
Vic. “Puente Las Cruces” along highway from Santa Cruz
to Samaipata, 6 km W (by air) of Bermejo, along Rfo Vi-
eoqufn, I8°08'S. 63°41'W, alt. KMX) m, M. Nee 42197
(Mf, XII, NY. USZ); 17 km by dirt road S of turn off from
Bermejo-Samaipata highway, slopes above Quebrada La
Coca. 18WS, 63°4P30"W, alt. I 150 m. M. Nee 50509
(MKXU, NY); Volcancs, cerca de 6 km NE de Bermejo,

1 8°06'S, 63°39'W, alt. 990 m. M. Saldias 4740 (MKXU,
NY. LJSZ); Volcanos, 3 km al N de Bermejo. subicndo pur
margenes del Bid Colorado. 18°06'S, 63°39'W, alt. ca.
1000 m. /. G. Vargas C. 5083 (MKXU, NY, USZ).

Eonehocarpus pluvialis Busby, Mem. New York
But. Card. 7(3): 268. 1927. TIPO: Bolivia.
Road from Rurrenabaque to Reyes, (). E.
White tV' II. II. Rushy 1302 (holotipo, NY; iso-
tipos, GH!. US!). Voiuicapoua pluvialis Rusby,
nom. nud. Imp. Woods 6: 12. 1926.

l/mchoearpus semens (Poirct) I).C. subsp. paraguariensis
Hassler. Keddcs Report. Spec. Nov. Regni Veg. 8:
207. 1910. riPO; Paraguay. Rfo Apa: h. h'iebrig
4389 (holotipo, G!; isolipo, (ill!).

Derris steinbachii Harms, Notizbl. But. Cart. Berlin-l)ah-
lem, 10: 344). 1928. TIPO: liolix ia. Santa Cruz. Cer-
cado. Rfo I’iraf. ./. Steinbach 71 19 (holotipo, B, des-
truido; isotipos, K!. G!, K!).

Material representative. BOLIVIA. Dept. Santa
Cruz: Prov. Cordillera, over Rfo Seco on N side of settle¬
ment of Rfo Seco, along highway from Santa Cruz to Aba-
po, M. Nee 49082 (MKXU. NY); Prov. Nuflo de Chdvez.
K of San Javier on road to Concepcion.,/, li. Abbott 10330
(MKXU); Prov. Guarayos, 5 km de las Juntas Rfos San
Pablo y Negro de Caimanes, /. G. Vargas et al. 2034
(UCZ). Prov. Ichilo, Chore. A*. Quevedo <0: ./. Magne 254
(UCZ); Prov. Andres Ibanez. Santa Cruz de la Sierra. A.
Jimenez A. 7! (UCZ); Prov. Warnes, 9.5 km of Warnes, 2.5
km NYY of Candelaria. M. Nee 50532 (MKXU, NY). BRA-

Volume 15, Number 4
2005

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Lonchocarpus sect. Lonchocarpus

595

SIL Mato Grosso <lo Sul: Mun. Bonito, Campo Dos In-
tlios. G. Hulschbach el at. 73927 (MKXU). Parana: Terra
Rica, Vargea do Rio Paranapanerna, proximo ao porto Eu-
clides da Cunha, /,. A. Dambros 267 (UKC). Sao Paulo:
Mun. Teodoro Sampaio, Reserva I* lorestal do Mono do
Diabo, II. F. Leitdo F el a!. 12486 (UKC). PARAGUAY.
Rio Apa. Aquidaban, K. Fiebrig 4389 (G, (ill).

Lonchocarpus riparius M. Sousa, sp. uov. I IPO:
Bolivia. Depts. La Paz y Beni; Basin of Bio
Beni, upstream from Rurrenabaque, Rio Tui-
chi and Rio Beni, on gentle slope near river,
14°35'S, 67°35'W, alt. 320 m. 13 May 1990.
I). C. Daly 6423 (holotipo, MLXU; isotipo,
NY). Figuras 2, 4.

Arbor 12—15 m alta, ramis angulatis, solidis, velutinis;
folia (5 ad)7-foliolata, rhachidi tereli. foliolis late elliptieis
vel elliptieis, translueido-punctatis, infra velutinis. Inflo-
rescentiae 8—14.5 cm longae, paniculatae. Flores 7.5—10
mm longi, petalis cremeis vel albis. Legumen (immatu-
rum) glabrescens, sutura vexillari lata facta, duabus mar-
ginibus aliformibus limitata

Arboles 12—15 m alto; corteza interior eon abun-
dante fluido resinoso al corte; ramas anguladas,
aeostilladas, solidas, de amarillento velutinas a gla-
brescentes, con lenticelas eireulares a eliplieas; es-
tipulas cadueas, ea. 3 mm largo, ea. 2 mm anelio.
ligulares; peciolo (1.4— )2— 4 cm largo; raquis foliar
(4.5— )7. 5— 9.5 cm largo, terete, acostillado, pardo-
amarillento velutino; hojas (5 a) ( -folioladas; foliolos
(4.5 — )7 — 1 1 (— 1 4) cm largo, (3— )3.5— 5(— 6) cm anelio.
anchamente elipticos a eliptieos, cartaeeos a sub-
coriaceos, translueido punteados y con las nerva-
duras de 3° v 4° orden translucidas, la base gene-
ralmente redondeada, en ocasiones cuneada, el
apice obtuso a acuminado, el liaz esparcido y el
enves moderadamente pardo-amarillento velutino;
nervacion camptodroma, nervadura primaria y se-
cundarias ligeramente realzadas en el enves, ner-
vaduras laterales 7 a 1 1. Infloreseeneias 8—14.5 cm
largo, paniculadas, cortamente pedunculadas; llo-
racion coetanea; peduneulos Morales 1—2 mm largo,
gruesos. moderadamente espaciados; pedieelos
1.7— 2.7 mm largo; bracteolas opuestas, aplicadas
al caliz, ovadas a orbiculares, 0.9— 1.2 mm largo.
0.9—1 mm ancho. Flores 7.5—10 mm largo; caliz
ciatiforme, easi trunco, 1.5—3 mm largo, 3—4 mm
ancho, pardo-amarillento sericeo, translueido pun-
teado; corola Blanca a color crema, translueido
punteada, los petalos caneseente sericeos; estan-
darte reflexo, su lamina easi orbicular, 6—7 mm an¬
cho; ovario pardo-amarillento sericeo. Legumbre
inmadura, recta a ligeramente curvada, aplanada,
hasta 6.5 cm largo, hasta 1.3 cm ancho, eliptiea a
oblonga, con I a 4 semillas. esparcidamente sericea
a glabrescente, la base cortamente atenuada, el api-

ce obtuso a apiculado, la sutura vexilar ensancha-
da. de hasta 4 mm de anelio. bordeada por 2 mar-
genes aliformes, la sutura carinal aquillada.
Semillas inmaduras.

Dist ribucidn > habitat. Se cornice en los lfmites
de los departamentos de La Paz y Beni y en Santa
Cruz. Fn habitats riparios, a elevaeiones 275—
320 m.

Fenologia. Con botones, flores de febrero a
mayo y frutos inmaduros a mediados de mayo.

FI epiteto liace referencia al habitat ripario de
esta espeeie.

Fsta espeeie se caraeteriza por las ramas angu¬
ladas que solo se conocen dentro de la misma sec-
cion en Lonchocarpus spiciflorus Martins ex Ben-
tham. |>ero esta ultima espeeie tambien cuenta con
tallos fistulosos (pie albergan hormigas, mientras
epic L. riparius M. Sousa se difereneia por presentar
tallos solidos. Otro caracter a notar es su inflores-
cencia panieulada, porque las hojas no se desarro-
I Ian posteriormente. tambien presente en la misma
seccidn en L. sericophyllus M. Sousa y en L. atro-
purpureus Bentham del grupo de L. minimijlorus.

Paratipos. BOLIVIA. Depts. La Paz \ Beni. Rio Beni,
upstream from Rurrenabaque, above Arroyo Pacheco and
I, os Fdsiles, growing by shore near Arroyo l.os Fosiles. I).
C. Daly et al. 6510 (MKXU, NY); Dept . Santa Cruz. Prov.
Iehilo, the turnoff 5.6 km E of Villa San German, 17°23’S,
64°<>.TW. M. Nee 48416 (MKXU, NY).

Lonchocarpus sericophyllus M. Sousa, sp. nov.
TIPO: Bolivia. Dept. Beni; Prov. Ballivian, ca-
rretera Carinavi— San Borja, eumbre de la Se-
rrania del Pildn Lajas. Bosque primario suban-
(lino, suelos aremseas, I5°I7,S, 67°04 ' ( ). all.
850-900 m, I Nov. 1989, I). N. Smith A- V.
Garcia 13828 (holotipo, MFXU; isotipos, MO,
US/). Figuras 2, 5.

Arbor ea. 35 m alta, ramis teretibus, dense brunneo-
llavo-serieeis vel glabrescentibus; folia 5-loliolata, foliolis
elliptieis vel lanceolatis, epunctatis, dense vel moderate
brunneo-davo-sericeis. 1 nfloreseentiae (2.5 — )4.5 — 7.5( — 10)
cm longae, axillares. Flores 8—9 mm longi, petalis ere-
meis. Legumen (inunaturum) brunneo-flavo-serieeum, su-
tura vexillari lata facta, duobus marginibus aliformibus
marginata.

Arboles ea. 35 m de alto; corteza exterior lisa,
gris-marron, la interior eon algo de fluido resinoso
al corte; ramas teretes, ligeramente aeostilladas, so¬
lidas, de densamente pardo-amarillento serfeeas a
glabrescentes, con lenticelas eireulares a elipticas;
esti'|iulas cadueas, 0.8— 0.9 mm largo. 0.9-1 mm an¬
cho, transversalmente triangulares; peciolo 2.3—4
cm largo; raquis foliar 2.2^4 cm largo, terete, se¬
riceo eomo las ramas, pronto glabro; hojas 5-folio-

596

Novon

h igura 4. Dmchocarpus riparius M. Sousa. — A. Rama con hoja, infrutesccncias y lcgumbres. — B. Legumbres en
diferentes posiciones, mostrando el margen vexilar ensanchado y aliforme. — C. Inflorescencia paniculada. — I). Unidad
biflora. mostrando pedilnculo, pedicelos, bracteolas, cdlices y flores. — E. Cdliz, mostrando bracteolas. pedicelo y cdliz
ciatiforme. K Estandarte. — G. Ala. — II. IV'talos de la quilla, mostrando las puntuaeiones translucidas. — I. Gineceo
sericeo. — J. Tubo estaminal. (A, B. I). C. Daly et at. 6 510 y C-J. I). C. Duly 6 423.)

Volume 15, Number 4
2005

Sousa

Lonchocarpus sect. Lonchocarpus

597

2 mm

Figura 5. Lonchocarpus sericophyllus M. Sousa. — A. Rama con hoja e infrutescencia. — IF Infrutescencia, mostrando
legumbres en diferentes caras con el margen vexilar ensanchado y aliforme. — C. Unidad biflora mostrando, pedunculo,
pedicelos, brdcteas, bracteolas v botones florales. — D. Caliz, pedicelo y bracteolas. — E. Inflorescencia. — I". Estan-
darte. — G. Ala. — H. Petalos de la quilla. — I. Gineceo. — .1. Columna de estambres. (A. IF I). F— J, I). /V. Smith &
F Garcia 13828 y C, F. G. Hartshorn & E. Meneces 2085.)

598

Novon

ladas; foliolos (6— )7.5— 10 cm largo, 3.2— 4.2 cm au¬
dio, elfpticos a lanceolados, cartdceos a
subcoridceos, cpunteados, pero las nervaduras <lc
3° y 4g orden translucidas, la base cuneada, en
ocasiones redondeada, el apice acuminado, cl liaz
de densamente pardo-amarillento serfceo a gla-
brescentc, cl enves dc densa a moderadamente par-
do- amarillento serfceo; nervaeion eamptodroma,
nervadura primaria y sceundarias ligcramcntc real-
zadas cn cl enves, nervaduras latcralcs 7 a 9. In-
florcsccncias (2.5— )4.5— 7.5(— 10) cm largo, axilares
o paniculadas, cortamcnlc pedunculadas; floracion
eoetcinea; pcdunculos Morales I ..5— 2.5 mm largo,
delgados, rcgularmcntc espaciados; pcdicelos 2—3
mm largo; bracteolas opucstas, aplicadas al ealiz,
ovadas, ca. 1.1 mm largo, ca. 0.9 mm audio. Flores
8—9 mm largo; ealiz ciatiforme, easi trunco, 2.8—3
mm largo, 4—4.5 mm audio, pardo-amarillento se-
rfeeo, epunteado; eorola de color crema, epunteada,
los pelalos canescente serfeeos, estandarte rellexo.
su lamina transversalmente elfptica, ca. 6 mm au¬
dio; ovario pardo-amarillento serfceo. Legumbre
inmadnra. recta, aplanada, hasta 4.3 cm largo, has-
ta 0.9 cm audio, oblonga. con 2 a 3 semillas. par¬
do-amarillento serfeea, la base atenuada, el apice
rostrado, la sutura vexilar ensanchada, de hasta 3
mm audio, bordeada por 2 margenes aliformes, la
sutura carinal aipiillada. Semillas inmaduras.

Distribut ion y habitat. Conoeida de los depar-
tamentos de Beni y Pando, en selvas primarias sub-
andinas, sobre suelos de areniscas a elevaciones de
850 a 900 m.

Fenologia. Con botones Morales a finales de oc-
tubre y el termino de la floracion e inicio de la
fructificacidn a principios de noviembre.

Este epfteto se refiere a la pelosidad sedosa de
sus hojas y en general de los renuevos.

Especie caracteristica por su pelosidad densa¬
mente pardo-amarillento serfeea, tanto en brganos
vegetativos y Morales, asf como en las legumbres.
Tambien destaca entre las otras especies de Bolivia
por sus folfolos y Mores epunteados.

Paralipo. BOLIVIA. Dept. Pando, Cobija, camino a
Puerto Heath, G. Ihirlsliorn A- E. Meneces 2085 (US/).

Las especies de la sect. Lonchocarpus presentes
en Bolivia se destacan del resto de las de Suda-
merica, por conglomerar a ties especies eon Mores
blancas o cremas (L. pluvialis Busby, L. riparius M.
Sousa y L sericophyllus M. Sousa), aclarando <|ue

se desconocen las Mores de L. lomentaceus M. Sou¬
sa. Tambien es notoria la preseneia de tres especies
con folfolos y petalos translucido punteados (L. neei
M. Sousa, /.. pluvialis Busby y L. riparius IM. Sou¬
sa).

Los endemicos nuevos para Bolivia, en gran ine-
dida pudieran tratarse de tin endemismo real, pero
|>robablemente scan resultado de la eseasa recolee-
ta en la region. Asf, no se descarta en tin futuro la
preseneia de algunas de estas especies en Brasil y
Paraguay.

Cl.AVK PAR A I AS ESPKCIKS l)K I .A SKCT.

Lo\chocarpi s t)K Bolivia

la. Raquis foliar marcadamente eanalieulado.

2a. Hojas y legumbres glabras, arbustos ca. 3 m
alto, legumbre lomentdcea se fragmenta en

artfculos monospermos .

. L. lomentaceus M. Sousa

21). Hojas y legumbres velutinas, drliolcs 7—20
m alto, legumbre sin fragmentarse en artf-

culos monospermos . L. neei M. Sousa

I b. Raquis foliar terete.

3a. Folfolos epunteados, densamente pardo-

amarillento serfeeos en el envds .

. L sericophyllus M. Sousa

3b. Folfolos translucido punteados, velutinos en
cl enves.

4a. llamas teretes; inllorcsceneias axilares;
frutos pardo-amarillento velutinos . .

. b. pluvialis Rusby

4b. llamas anguladas; inllorcsceneias pani¬
culadas; frutos easi glabros .

. /,. riparius \1. Sousa

Agradecimientos. A Fernando Chiang, por tra-
dueir al latfn las diagnosis, a Gloria Andrade por
su soporte a la elaboraeidn del texto, a Ramiro Cruz
y Elvia Esparza por sus magnfficas ilustraciones
botanicas y a Gabriela Sanchez por hacer el mapa
de distribuciones. Finalmente a los euradores de
los siguientes berbarios F. G. GH. K. MEXU, MO,
NY, UEC, US, US/ que hicieron posible el estudio
del material.

Literatura Citada

A/.t ne do-I ozzi. A. M. (J. de. 1989. Kstudos taxonomicos
dos generos Ixmchocarpus Kunth c Deguelia Aubl. no
lb asil. Tese dc Doutorado. Universidade Fstadual de
Campinas, Campinas, Brasil.

Fortunato, R. 11. y R. Palese. 1997. Una nueva combi-
nacidn en el gdnero Ijonchocarpus Kunth (Legumino-
sae— Millettieae): hmchocarpus jluvialis (Liendm.) For¬
tunato & Palese. En L. Ramella y P. Perret (editores),
Notulae ad Floram paraguaiensem 62. Candollea 52:
509-51 I.

New Combinations for Chinese Bamboos (Poaceae, Bambuseae)

Chris M. A. Stapleton

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom.

c.stapleton@kew.org.uk

Li De-Zhu

Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming 650204,
Yunnan. Peoples Republic of China, dzl@puhlic.km.yn.cn

Xia Xian He

South China Institute of Botany, Chinese Academy of Sciences, Guangzhou 510650,
People’s Republic of China, tihxia@scib.ac.cn

ABSTRACT. On the basis of morphological char¬
acters inconsistent with their current taxonomic
placements, new combinations are made for the fol¬
lowing species of woody bamboos within the flora
of China: Ampelocalamus breviligulatus, Ampelo¬
calamus hirsutissimus, Ampelocalamus melicoides,
Drepanostach yum semiorbiculatum, Drepanostach¬
yum ampullarium, Drepanostachyum membrana-
ceum, and Bonin parvifloscula. To accommodate
broader generic concepts, new combinations are
made for Arundinaria qingchengshanensis and
Arundinaria spanostachya, and new status is given
to Bashania as Arundinaria subgen. Bashania, and
to Sarocalamus as Arundinaria subgen. Sarocala-
m u. s.

Key words : Ampelocalamus, Arundinaria, Bam¬
buseae, Bashania, Bonia, China, Drepanostach¬
yum, Poaceae, Sarocalamus.

During the preparation of a collaborative account
of Chinese bamboos for the English-language Flora
of China, it became apparent that there was a need
for several new combinations for species names in
Ampelocalamus S. L. Chen. T. H. Wen & G. Y.
Sheng, Drepanostachyum Keng 1.. Arundinaria Mi-
chaux, and Bonia Balansa, and for the provision of
names at subgeneric rank within Arundinaria s.l.
for Bashania Keng f. & T. P. Yi and Sarocalamus
Stapleton.

A MPELOCALAMl S

Ampelocalamus Itreviligulatus (T. P. Yi) Staple-
ton & 1). Z. hi, comb. nov. Basionyrn: Drepa¬
nostachyum breviligulatum T. P. Yi. J. Bamboo
Res. 12(4): 42. 1993. TY PE: China. Sichuan:
Jiange Xian. 18 Dec. 1983, Yi Tong- Pei 83208
(holotype, SIFS not seen).

Ampelocalamus hirsutissimus (W. I), hi & Y. C.
Zhong) Stapleton & I). Z. hi. comb. nov. Bas-
ionym: Drepanostachyum hirsutissimum W. I).
hi <X Y. C. Zhong, in J. Bamboo Res. 16(1):
52. 1997. TYPE: China. Guizhou: Guiyang.
Bamboo Garden of GF, 5 Mar. 1992. /hong
Yuan-Chun et al. 9203 (holotype. GF not
seen).

Ampelocalamus melicoides (Keng I.) I). Z. hi &
Stapleton, comb. nov. Basionyrn: Drepano¬
stachyum melicoideum Keng h, J. Bamboo
Res. 5(2): 35. 1986. TYPE: China. Sichuan:
Nanchuan Xian, 4 May 1957, C. F. Li 00330
(holotype, NJNU; isotype, SZ).

Ampelocalamus and Drepanostachyum are sub¬
tropical clump-forming bamboo genera with type
species from Hainan Island and the Himalayas of
northwest India, respectively. They differ in several
characters. The culm nodes and branching in Am-
pelocalamus are adapted for a semi-scandent
growth habit, with expanded nodal sheath scars, ge¬
niculate lateral branches, a large central branch
that often remains dormant, and a capacity for ae¬
rial root growth on the swollen bases of larger
branches. Culms of Drepanostachyum arc self-sup¬
porting and lac k these adaptations. The inflores¬
cence in Ampelocalamus also differs in having larg¬
er, broader spikelels (to 35 X 5 mm vs. 20 X 3
mm). Ampelocalamus is distributed from Taiwan
and Hainan Island to the wettest parts of the Him¬
alayas, especially on calcareous soils, while Dre¬
panostachyum is found along the Himalayas in dri¬
er subtropical forest types associated with trees in
Quercus L., Schima Reinwardt ex Blume. and Cas-
tanopsis (D. Don) Spaeh.

Phe two genera were confused by transferral of

Novon 15: 599-601. Published on 12 December 2005.

600

Novon

most species of Ampelocalamus into Drepanostach¬
yum (Kong. 1986; Yi. 1998) and eventually by the
synonymizing of Ampelocalamus within Drepano-
stachyum (Li, 1997). However, recent molecular
data (Nf Chonghaile, 2002) has suggested that they
are not so closely related, with Drepanostachyum in
a clade (Bootstrap Support 73%) that did not in¬
clude Ampelocalamus.

The generic characteristics of Ampelocalamus
hreviligulatus have not been investigated in depth,
but the large discrepancy in branch sizes with a
dominant central branch and finer lateral branches,
along with the deeply ridged culm intemodes sug¬
gest that it is more likely to belong in Ampeloca¬
lamus than in Drepanostachyum. This is reinforced
by its distribution in Sichuan and Gansu provinces
of China, rather than the Himalayas.

Described as related to Ampelocalamus scandens
Hsueh & V\. I). Li, with apically arching culms and
patelliform nodes, and illustrated as having a
branch complement with strongly geniculate lateral
branches, Ampelocalamus hirsutissimus would also
appear more appropriately placed in Ampelocala¬
mus rather than in Drepanostachyum. The ciliolate
culm sheath margins and prominent leaf sheath oral
setae arc similar to those of A. patellaris and A.
scandens, respectively.

The dominant central branch and geniculate lat¬
eral branches, along with broad spikelcts suggest
that Ampelocalamus is also more appropriate than
Drepanostachyum for 1. melicoides.

Drepanostachyi t /

DrepnnostncliYiim seiiiiorbieiilatiiin (T. 1*. i i)

Stapleton, comb. nov. Basionym: Fargesia se-
miorbiculata T. I*. Yi. J. Bamboo lies. 2(2): 40.
1983. TV PE: China. Tibet: Cona Xian, 7 Jan.
1978, Jiang Changgui I (holotype, SIFS not
seen).

Drepanostachyum aiiipiillarium (T. P. Yi) Sta¬
pleton. comb. nov. Basionym: Fargesia ampul-
laris T. P. Yi. J. Bamboo lies. 2(2): 18. 1983.
TYPE: China. Tibet: Zhangmu, 4 Nov. 1979.
M. L. Zhou 4 (holotype, SIFS not seen).
Drepanostachyum iiiembraiiaeeum (T. P. Yi) I).
Z. Li, comb. nov. Basionym: Fargesia mem-
hranacea T. P. \i. Acta Bot. Yunnan. 14(2):
135. 1992. TYPE: China. Sichuan: Mianning
Xian, Yi Tong Dei 90172 (holotype, SIFS not
seen).

flic type species of Drepanostachyum, D. falca-
tum (Nccs) Keng f., was not known at all in China,
while several well-known species of Ampelocalamus
had been transferred into Drepanostachyum. There¬

fore Drepanostachyum was misunderstood in China,
where it was considered very similar to Ampeloca¬
lamus, which was still recognized (Keng & Wang,
1996), albeit for only 2 less well-known species.
Meanwhile, when species of Drepanostachyum were
encountered in the Himalayas of southeast Tibet,
they were placed in an altogether different genus,
Fargesia Franchet. Section Sphaerigemma T. P. Yi
and series Ampullares T. P. Yi were described large¬
ly to accommodate these mainly Himalayan species
within Fargesia, because of their semi-orbicular
culm buds and swollen culm nodes, not seen in
other species of the genus.

Fargesia semiorbiculata and F ampullaris also
differed substantially from other species placed in
Fargesia in their larger number of more subequal
branches, leaf blades with indistinct transverse
veins, the narrow apex and distally scabrous inte¬
rior surface of their culm sheaths, and their sub¬
tropical rather than temperate habitats. These char¬
acteristics are shared with the type species of
Drepanostachyum, I), falcatum , into which genus
they are now transferred.

Fargesia membranacea differs from most other
species in Fargesia in its semi-orbicular branch
buds producing 13 to 33 subequal branches, its leaf
blades with indistinct transverse veins, and the nar¬
row apex to its culm sheaths, characteristics more
suggestive of Drepanostachyum than Fargesia.
Known only from southern Sichuan province, this
species is well separated from all other species of
Drepanostachyum.

A RUN DIN ARIA

Arundinaria subgen. Saroealamus (Stapleton) 1).
Z. Li, stat. nov. Basionym: Saroealamus Sta¬
pleton, Novon 14: 346. 2004. TYPE: Saroea¬
lamus racemosus (Munro) Stapleton.
Arundinaria subgen. Bashania (Keng f. & T. P.
Yi) I). Z. Li, stat. nov. Basionym: Bashania
Keng f. X T. P. Yi. J. Nanjing Univ., Nat. Sci.
Ed. 1982(3): 722. 1982. TYPE: Bashania far-
gesii (E. G. Camus) Keng f. & T. P. Yi.
Arundinaria qiiigrlieugshaiicnsis (Keng f. N T.
P. Yi) I). Z. Li, comb. nov. Basionym: Bashania
q i ngc hengshane ns is Keng f. & T. P. Yi. .1. Nan¬
jing Univ., Nat. Sci. Ed. 1982(3): 725. 1982.
TYPE: China. Sichuan: Guan Xian, 25 June
1981, Yi Tong-Pei 80037 (holotype, SIFS).
Arundinaria spanostaehya (T. P. Yi) I). Z. Li,
comb. nov. Basionym: Bashania spanostaehya
T. P. Yi, Acta Bot. Yunnan. 11: 35. 1989.
TYPE: China. Sichuan: Huili Xian, Beimu
Shan, 18 May 1987, Yi Tong-Pei 87249 (ho¬
lotype, SIFS).

Volume 15, Number 4
2005

Stapleton et al.
Chinese Bamboos

601

Although it is strongly suspected that Bashania
keng f. & T. P. Yi and Sarocalamus Stapleton are
probably not closely related to the North American
genus Arundinaria (Nf Chonghaile, 2002; Stapleton
el ah, 2004), the morphological distinctions are less
pronounced than those between Arundinaria and
all other Chinese genera. As the Flora of China
follows a primarily morphological classification sys¬
tem, these three genera w ill be merged into a rather
broadly interpreted Arundinaria. Hopefully further
molecular data will provide clarification of the true
relationships between these groups. NewT combi¬
nations in Arundinaria are required for two species
of Bashania. while B. fargesii and the remaining
two species of Sarocalamus already have combi¬
nations in that genus, and the affinities of three
further, more recently described species of Bashan¬
ia remain unclear.

BONIA

Bonin parvifloscula (W. T. kin) N. H. Xia, comb,
nov. Basionym: Monocladus parviflosculus W.
T. Lin. J. Bamboo Res. 12(3): 3. 1993. TYPE:
China. Guangdong: Zhaoqing, Qixingyan, Z.
K. Li 84126 (holotype, CANT).

The genus Monocladus Chia, H. L. Fung & Y. L.
Yang et al. was established for tropical bamboos
from south China with solitary branches and spike-
lets with a well-separated basal floret. It was not
realized that species of the earlier genus Bonia Bal-

ansa also had these characteristics (Xia. 1996). The
characteristics of Monocladus parviflosculus are not
well known as the only collections made so far are
poor, but it would appear that it also represents a
species of Bonia rather than any other genus.

Acknowledgments. The corresponding author
thanks Missouri Botanical Garden for funding this
research and the Keeper of the Herbarium of the
Royal Botanical Gardens, Kew, for providing work¬
ing facilities.

Literature Cited

Keng, P. C. 1986. A new discovery of bamboo genus Dre-
panostachyum Keng f. in China. J. Bamboo Res. 5(2):
28-40.

- & Z. P. Wang (editors). 1996. Gramineae (Po-

aceae), Bambusoideae. Flora Reipublicae Popularis
Sinicae 9(1). Science Press, Beijing. [In Chinese.]

Li, D. Z. 1997. The Flora of China Bambusoideae Project:
Problems and current understanding of bamboo taxon¬
omy in China. Pp. 61—81 in G. P. Chapman (editor).
The Bamboos. Academic Press, London.

Nf Chonghaile, G. 2002. Molecular Systematics of the
Woody Bamboos (Tribe Bambuseae). Trinity College,
Dublin [unpublished Ph.D. thesis].

Stapleton, C. M. A., G. Nf Chonghaile & T. R. Hodkinson.
2004. Sarocalamus, a new Sino-llimalayan bamboo ge¬
nus (Poaceae— Bambusoideae). Novon 14: 345—349.

Xia, N. H. 1996. A study of the genus lionia (Gramineae:

Bambusoideae). Kew Bull. 51: 565—569.

Yi, T. P. 1993. New taxa of Drepanoslachyum and other
new combinations of bamboos in China. J. Bamboo Res.
12(4): 42-47.

New and Interesting Milkweeds (Apocynaceae, Asclepiadoideae)

It! D. Stevens

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

stevens@mobot.org

Abstract. Matelea corrugata, from Costa Rica,
47. costaricensis, from Costa Rica, 47. emmartinezii,
from Mexico, 47. ferruginea, from Guatemala, M.
filipes, from El Salvador, M. haberi, from Costa
Rica, M. insolita, from Nicaragua, and 47. veluti-
noides, from Mexico and Guatemala, are described
as new. The new combinations Ditassa nigrescens
(= Ditassa tatei ), Gonolobus glaberrimus ( — Ma¬
telea tikalana), Marsdenia fruticosa, and Polystem-
ma guatemalense (= Matelea quirosii ), are pro¬
posed, and the latter two are leetotypified. Asclepias
angustifolia, A. auriculata, A. si mil is. Gonolobus
barbatus , G. erianthus (= G. macranthus), G. gua-
temalensis, Marsdenia gymnemoides, Matelea gen-
tlei , and Pherotrichis schajfneri are leetotypified or
neotypified. The typifieation, synonymy, and appli¬
cation of Pherotrichis villosa are discussed.

Key words: Apocynaceae, Asclepiadoideae,
Mesoamerica, Guianas, Asclepias , Ditassa, Gonolo¬
bus, Marsdenia, Matelea, Pherotrichis, Polystemma.

In the course ol ongoing studies of milkweeds,
several new species, new combinations, and no-
menclatural trivia have come to light and are pre¬
sented here.

Asclepias angustifolia Schweigger, Hnum. 1*1.
Hurt. Regiom. 13. 1812. TYPE: Hurt. Sal-
tzwedel, Frankfurt, Oct. 1825, G. Engelmann
s.n. (neotype, here designated, MO-2761538).

Woodson (1954) discussed the application of this
name, which he was substituting for the then better
known Asclepias rubricaulis I IRK. Asclepias angus¬
tifolia Schweigger was based on a plant cultivated
at tilt' Kiinigsberg botanical garden, acquired from
the Berlin botanical garden. If a type specimen ex¬
isted, it probably would have been at KRG and is
no longer extant. Woodson (1954) suggested that
herbarium specimens at MO. prepared from plants
cultivated in other German botanical gardens dur¬
ing that era, and bearing Schweigger's name, prob¬
ably represented the original introduc tion and serve
to fix the usage of the name. Woodson was clearly
neotypifying Asclepias angustifolia. but since there
are four specimens at MO lilting his general de¬

scription it seems prudent to select one specimen
to formalize this decision.

Woodson (1954: 151) cites " Gomphocarpus au¬
gust if oh us Link” as a synonym of Asclepias fruticosa
L. However, “ Gomphocarpus angustifolius Link” is
actually a combination based on Asclepias angus¬
tifolia Schweigger, Gomphocarpus angustifolius
(Schweigger) Link and surely was included in the
synonymy of A. fruticosa in error. Goyder and Nich¬
olas (2001) list Gomphocarpus angustifolius and its
basionym as possible synonyms of G. fruticosus (L.)
W. T. Aiton, based on Woodson’s synonymy, but
note that the type is no longer extant and the iden¬
tity uncertain. Therefore, this neotypification cla¬
rifies the synonymy ol Gomphocarpus fruticosus by
removing Asclepias angustifolia Schweigger and
Gomphocarpus angustifolius (Schweigger) Link as
candidates.

Asclepias auriculata I IRK, Nov. Gen. Sp. (quarto
ed.) 3: 191, t. 228. 1819. TYPE: Nov. Gen.
Sp. (quarto ed.) 3: t. 228. 1819 (leetotype, des¬
ignated here).

The original description cites “Creseit locis alsis
prope Ario Mexicanorum, alt. 1000 hex. . . .Floret
Septembri.” Woodson (1954) gives the type as
Bonpland s.n., but did not see a specimen. Original
material still has not been located, and in its ab¬
sence I propose the original plate as the leetotype.

Asclepias similis Hemsley, Biol. Cent.-Amer., Rot.
2(10): 326. 1881. TYPE: “South Mexico, Chia¬
pas etc.,” .4. Ghiesbreght 666 (leetotype, des¬
ignated here, K; duplicates of leetotype, G,
MO).

Hemsley cited two collections in his description
of the species. Ghiesbreght 665 and 666, both at K.
Woodson (1954) gives Ghiesbreght 665 at MO as a
type. While this could be considered a leetotypifi-
cation, Woodson's “Index to Exsiccatae” in the
same publication lists Ghiesbreght 665 as Asclepias
pellucida E. Fournier and Ghiesbreght 666 as A.
similis. The actual specimens of both collections at
MO were annotated by Woodson as Asclepias simi¬
lis. Asclepias similis and A. pellucida are sympatric

Novon 15: 602-619. Published on 12 December 2005.

Volume 15, Number 4
2005

Stevens
New Milkweeds

603

in southern Mexico and, as noted by Standley and
Williams (1969), are difficult to distinguish without
follicles. Although I agree with Woodson’s deter¬
minations of both collections as Asclepias similis,
and neither collection has follicles, it still seems
important that Asclepias similis be lectotypified, and
Ghiesbreght 6 66 is the less ambiguous choice.

Ditassa nigrescens (E. Fournier) W. D. Stevens,
comb. nov. Basionym: Astephanus nigrescens E.
Fournier, Ann. Sci. Nat., But., ser. 6, 14: 367.
1882. TYPE: Venezuela. “Guyana Anglica, pr.
Boraima,” M. Schomburgk 915 (bolotype, B
not seen).

Ditassa nigrescens is a rather common species in
tin* adjacent areas of Venezuela, Guyana, and Bra¬
zil. The species was treated as Ditassa tatei Glea¬
son & Moldenke (in Gleason, Bull. Torrey Bot. Club
58: 456. 1931. Holotype: Venezuela. Esmeralda, R.
Tate 199 , NY, = Cynanchum tatei R. W. Holm) in
the Flora of the Venezuelan Guayana (Morillo,
1997). However, the original diagnosis of Ditassa
tatei cited two collections in addition to the type:
R. Spruce 5226 (k) and Schomburgk 915 (B). Since
Schomburgk 915 at B was already the holotype of
Astephanus nigrescens, Ditassa tatei is a superfluous
name under Article 52 of the Saint Louis Code
(Greuter et ah, 2000), and clearly the two concepts
are synonymous.

Gonolobus barbatus HBk, Nov. Gen. Sp. (quarto
ed.) 3: 209. t. 239. 1819. TYPE: Nov. Gen.
Sp. (quarto ed.) 3: t. 239. 1819 (lectotype, des¬
ignated here).

The original description cites “Crescit in lilore
Mexicano, prope Campeche.” Original material has
not been located and in its absence I propose the
original plate as the lectotype.

Gonolobus eriantbus Deeaisne, in A. de Can¬
dolle, Prodr. 8: 592. 1844. V incetoxicum erian-
thum (Deeaisne) Arthur. Torreya 21: 11. 1921.
TYPE: Mexico. Veracruz: Bois pres Jalapa,
4000 ped., H. Galeotti 1519 (lectotype, des¬
ignated here, P; duplicates of lectotype, BR,
G, k. Pi.

Deeaisne published Gonolobus erianlhus either
as a replacement for a putative later homonym (“G.
grandiflorus Benth.,” not G. grandiflorus (Cavanil-
les) R. Brown ex Schultes) that was never published
or perhaps in recognition that Bentham’s applica¬
tion of the name was in error, but in either ease
must be considered to be newly described by De¬

eaisne. The selected lectotype is the best of the four
specimens at P cited by Deeaisne; the unselected
syntypes are: //. Galeotti 1552 (BR, P), 1559 (BR,
P), and K. Hartweg 21 1 (GH, k, NY, P). The name
Gonolobus macranthus Kunze (Linnaea 20: 27.
1847) is, from the diagnosis, clearly synonymous
with G. erianthus, and was used for the species in
Flora de Nicaragua (Stevens el ah, 2001); it was
based on cultivated material from Mexico and any
original specimens or illustrations, if such existed,
were at LZ and are no longer extant (Peter Otto,
pers. comm.).

Gonolobus glaberrimus (Woodson) W. 1). Ste¬
vens, comb. nov. Basionym: Matelea glaberri-
rna Woodson, Ann. Missouri Bot. Card. 28:
281. 1941. TYPE: Guatemala. Peten: Uaxac-
tun, 24 Mar. 1931, //. Bartlett 12500 (holo¬
type, MO; isotypes, MICH, MO, US).

Matelea tikalana hundell, Phytologia 16:446. 1968. Syn.
nov. Marsdenia tikalana (hundell) hundell, Wrightia
4: 49. 1968. TYPE: Guatemala. Peten: Tikal Nat.
Park, 20 Mar. 1959, C. hundell 15765 (holotype,
l.h).

Gonolobus glaberrimus is most closely related to
G. hamrnelii W. D. Stevens and G. hadrostemma W.
D. Stevens, from which it differs by having glabrous
leaves among many other obvious characters; the
three species are anomalous in Gonolobus in having
a massive corona adnate to the faucal annulus and
in lacking free appendages on the anther backs, but
seem better placed in Gonolobus than in Matelea.
Matelea tikalana is placed into the synonymy of
Gonolobus glaberrimus because the two type col¬
lections are virtually identical.

Gonolobus guatenialensis k. Schumann, in En-
gler & Prantl, Nat. Pflanzenfam. 4(2): 302.
1895. TYPE: Guatemala. Alta Verapaz: Pan-
samala, 4000 ft., Apr. 1887. //. ron Tuerckheim
1124 (lectotype, designated here, US).

Gonolobus velutinus Schlechtendal var. calycinus ,1. I).
Smith, Bot. Gaz. 13: 189. 1888. TYPE: Guatemala.
Alta Verapaz: Pansamala, 4000 It., Apr. 1887, II. ron
Tuerckheim 1124 (holotype, US).

Schumann intended Gonolobus guatenialensis to
be a new name for “G. velutinus Donn. -Smith, nicht
Schlecht.,” but technically the name must be con¬
sidered newly published by Schumann. Schumann’s
key adequately diagnoses the species and this lee-
totypification simply formalizes his obvious inten¬
tion of renaming Smith's taxon by selecting the
same type.

604

Novon

Vlursdcuia fruticosa (J. I). Smith) W. I). Stevens,
comb. nov. Basionym: Nephradenia fruticosa .).
i). Smith, Hot. Gaz. 16: 196. 1891. TYPE:
Guatemala. Alta Verapaz: rocky islands in Rfo
Rubelcruz, Apr. 1889, Smith 1742 (lecto-
type, designated here, US).

While Smith’s diagnosis intentionally distin¬
guished Nephradenia fruticosa from N. neriifolia
(Dec aisne) Bentham J. I). Hooker (= Marsdenia
neriifolia (Decaisne) Woodson), it has always been
considered a synonym of the same, e.g., by Wood-
son ( 1941), Standley and W illiams (1969). and Om-
lor (1998), probably because the two species have
an erect, Asclepias-Wke habit, unique among North
American Marsdenia, and are practically indistin¬
guishable from each other vegetatively. However,
the (lowers and fruits of the two spec ies are quite
different. Marsdenia fruticosa has cream-colored
flowers versus purple in M. neriifolia; a campanu-
late corolla with the tube longer than the lobes ver¬
sus rotate with the lobes longer than the tube; co¬
rona lobes that are widest at the base and
connected to the anther hack by a thin w ing versus
widest in the middle and directly adnate to the an¬
ther back; follicles that are short and broad versus
long and thin; and seeds that are without a coma
and have a thick, spongy wing versus normal co-
inose seeds without corky wings. Several genera of
milkweeds have a species or two that have lost the
seed coma and develop spongy wings associated
with becoming rhyophytes, and this species is ap¬
parently a rhyophyte. Marsdenia fruticosa, M. ner¬
iifolia, and M. laxiflora J. I). Smith (a twining or
trailing species) are clearly closely related, and
seem to lx* relatively isolated among North Amer¬
ican Marsdenia. The unselected syntype of Nephra¬
denia fruticosa is //. von Tuerckheim 1251, US.

\larsdenia gynmemoides W. Rothe, Rot. Jalnb.
Syst. 52: 409, t. 3. 1915. TYPE: Guatemala.
Huehuetenango: Nenldn. Uaxac kanal, July
1887, C. Seler <£■ E. Seler 2804 (lectotype, des¬
ignated here, F).

The two syntype collections (C. Seler & E. Seler
2804. 3093) were both at B and are no long* ‘r ex¬
tant. A fragment of one of the syntypes is at F and
is chosen as the lectotype. Lectotypification seems
especially important in this case because Rothes
(1915) drawing of Marsdenia gymnemoides is am¬
biguous. combining some characters of M. gymne¬
moides, as represented by the lectotype, and M.
luhularis L. 0. Williams, a similar and nearly svm-
patric species.

Matelea corrugata W. I). Stevens, sp. nov. TYPE:
Costa Rica. Puntarenas: Reserva Forestal Gol-
fo Dulce, Aguabuena, sector oeste, 50—150 m,
8°42'20"N, 83 °3 1 '30WW, 17 Dec. 1991, It
Aguilar 764 (holotype, MO). Figure I.

A speciebus Matelearum indumento siniplici foliis cor-
iaceis crassinervibus corona siniplici polliniis triqueteris
profunde excavatis indumento corollae singular! distin-
guenda.

Coarse woody vine, older stems and base un¬
known. latex white, young stems densely pilose, ob¬
scuring surface, triehomes 0.5—1 mm long, straight
or twisted, yellow' fading to dull white, erect to de-
flexed, internodes 3—11 cm. Leaves opposite,
blades elliptic or slightly ovate or obovate, coria¬
ceous, 6—13.2 X 2. 2-6.3 cm, apex abruptly acu¬
minate, base obtuse, rounded, truncate or shallowly
cordate, sinus to 2 mm deep, adaxially glossy,
sparsely appressed-scabrous, triehomes 0.2— 0.7
mm long, ahaxiallv densely appressed-pilose. ob¬
scuring surface, triehomes 0.7—1 mm long, lateral
veins 6 to 8 pairs, prominent above and below, mid¬
dle veins 45°-55° to midrib, colleters 4 to 10, long
and curved, irregularly scattered along the basal 5—
15 mm of midrib; petiole 1.2— 3.7 cm, densely pi¬
lose. Inflorescence congested-racemiform, densely
pilose, peduncle 0-2 mm, pedicel 3—8 mm. bracts
0.7-1. 5 X 0.3— 0.5 mm, debate, pubescent; calyx
tube 1—1.2 mm long, with 1 colleter per sinus with¬
in, lobes elliptic, obovate or spathulate, cucullate,
4-6.5 X 3.5— 3.6 mm, apex rounded, green, densely
appressed-pilose outside, triehomes 0.2— 0.5 mm
long, glabrous inside; corolla briefly eampanulate
and then rotate, brown, sometimes with pale purple
in center, abaxially glabrous except sparsely ap¬
pressed-pilose in the center of the lobes, triehomes
0.2— 0.3 mm long, adaxially hispidulous in a ring
in the tube and in 5 radial lines opposite the an¬
thers connecting the ring with the corona, thus
leaving 5 trapezoidal glabrous patches in the base
of the tube alternate with the anthers, triehomes
ferrugineous, ca. 0.05 mm long, corolla tube 2.5—4
mm long, lobes broadly elliptic, asymmetrical at the
base, 3.8— 5.5 X 6—7 mm, apices rounded and
emarginate, patent, one side of each lobe verrucose
within; gynostegium sessile, corona a fleshy disk
more or less hidden below gynostegium, apparently
purple or reddish purple, outer margin shallowly 5-
lobed and finely and inconspicuously dentate, ea.
0.7 mm wid<* at the lobes and 0.5 mm wide between
the lobes, with 5 raised cushions alternate with the
anthers; anthers trapezoidal in outline, dorsally
with a pair of horny ridges, 0.7— 0.9 X 1.3— 1.8 mm.
guide rails straight, flaring apart ca. 45° and tilted

Volume 15, Number 4
2005

Stevens
New Milkweeds

605

Figure 1. Matelea corrugata W. I). Stevens. Scan of holotype specimen {Aguilar 764, MO).

ca. 45° under style apex, 0.2— 0.3 mm long, terminal
appendages narrow strips appressed to margin of
style apex. 0.2— 0.4 X 0.9— 1.3 mm, white; ccrpus-
culum 0.24-0.31 X 0.10—0.13 mm, ellipsoid to
subsagittate, brown, translators 0.31—0.38 X 0.14—

0.19 mm, slightly curved, flat, pollinia 0.69-0.76
X 0.46—0.51 mm, asymmetrically ovoid, strongly 3-
dimensional (triquetrous), sterile and deeply exca¬
vated in proximal half, forming a tube into center
of the pollinium; style apex 2.3— 2. i mm wide, pen-

606

Novon

tagonal, slightly concave, apparently brown. Folli¬
cles fusiform, 12-16 X 3-3.5 cm, apex acute to
attenuate, base acute, green or green-black, dense¬
ly ferrugineous-pilosulose, with scattered broad-
based. conical prickles (ca. 3 per cm2), prickles 2—
4 mm long, 1—3 mm wide at middle; seeds obovate,
10—1 1 X 5.5-0.5 mm, brown, margin 0.6— 1.2 mm
wide, entire or inconspicuously undulate, surface
smooth, coma ca. 2.5 cm long, tawny.

Matelea corrugata is distinctive in appearance
and not clearly related to any other Mesoamerican
species, although clearly falling within the current
circumscription of Matelea. The pattern of corolla
indumentum and the pollinium shape, as described
above, are unique. Two of the collections, including
(lit* type, are from lower elevation (50-600 m) wet
forest in southeastern Costa Rica and two are from
higher elevation (900—1200 m) cloud forests in
northern Costa Rica, but the four collections are
practically identical. Matelea corrugata has been
collected in flower in September, November, and
December.

Earatypes. COSTA ItICA. Alujuclu: San Ramrin, Za-
potal. Mo Rarranquilla, E. Hello C. & E. Cruz L 5313
(MO); Monteverde Cloud Forest Reserve, Rfo Penas Blan¬
cas, W. Haber <& h. Hello C. 6399 (MO). Puiitarenas:
Osa Sierpe, entre Cerro Chocuaco y Cerro Kl Faro, 6'.
Herrera C., H. Soto & H. Garcia M. 4610 (MO).

Matelea costaricensis W. 1). Stevens, sp. nov.
TYPE: ( iosta Rica. San Jose; Canton de Acos¬
ta, Hda. Tiquires, camino a Aguas Buenas,
I600-IR00 m, 9°43'10"N, 84°10'50"W, 3 June
1995, ./. Morales 4283 (holotype, MO). Fig¬
ure 2.

Mateleae pinguifoliae et M. pseudobarbatae affinis, sed
differt corolla intra hispidulosa corona fere integra.

Vine, herbaceous or somewhat woody, base un¬
known, latex white, young stems with mixed indu¬
mentum. long trichomes moderately dense, 1—3
mm, straight, pale yellow, spreading, short tri¬
chomes dense, 0.1— 0.2 mm, straight, white, glan¬
dular trichomes dense, ca. 0.1 nun, brown, inter-
nodes 7—1 i cm. Leaves opposite, blades ovate, 6—
12.2 X 3.4— 7.2 cm, apex acuminate, base lobate,
lobes convergent to overlapping, sinus 0.8— 2.5 cm
deep, adaxially with moderately dense long and
short trichomes, long trichomes 0.5— 1.5 mm, some¬
times with sparse glandular trichomes on veins and
at base, abaxially with dense short and glandular
trichomes, sometimes with a few long trichomes on
veins, lateral veins 5 to 7 pairs, middle veins 40°-
45' to midrib, eolleters 7 to 12; petiole 2—7.5 cm.
with mixed indumentum, long trichomes sometimes

sparse. Inflorescence racemiform or congested-ra-
eemiform, with mixed indumentum, peduncle 9—34
mm, axis to 7 mm, pedicel I 1-26 mm, bracts 2.8—
6 X 0.3— 0.8 mm, linear or narrowly lanceolate; ca¬
lyx tube 0. 5-0.8 mm long, with I to 3 eolleters per
sinus, lobes lanceolate, 3.3— 5.6 X 1 .2-1.8 mm,
apex acute, green, with dense mixed indumentum
outside, long trichomes 0.4-1. 5 mm, glabrous in¬
side; corolla briefly campanulate then rotate, olive
green, yellow-green, green-brown, or orange-brown,
abaxially with mixed indumentum or sometimes
lacking glandular trichomes, adaxially hispidulose,
trichomes translucent white, 0. 1-0.2 mm, tube 1.7—
2.3 mm long, lobes ovate, 5.9— 8.4 X 3.4— 4.2 mm.
apex rounded and shallowly notched; gynostegium
sessile, outer corona stipitate below, cupuliform
above, stipe 0.6— 0.8 mm long, obconic, with color
and texture of corolla, upper part fleshy, glabrous,
pentagonal in outline, 041-1 mm long, 0. 3-0.6 mm
thick, distal margin entire or weakly 2-tiered op¬
posite anthers, deep purple-black, somewhat ex¬
ceeding base of gynostegium, inner corona of 5 ad-
nate ligules opposite outer corona lobes and
anthers, linear with free truncate tips incumbent on
backs of anthers, free tips 0.2— 0.3 X 0.2 mm; an¬
thers trapezoidal in outline, 0.5— 0.6 X 0.8— 0.9 turn,
tilted under style apex, guide rails straight, parallel,
tilted under style apex, ca. 0.1 mm long, terminal
appendages appressed to style apex, 0.4 X 0.8— 0.9
mm, white, corpusculum 0.21—0.26 X 0.09—0.13
mm, sagittate, red-brown, translators 0.15—0.19 X
0.1-0.14 mm, pollinia 0.39-0.44 X 0.24-0.28 mm.
angularly and asymmetrically obovate, sterile at at¬
tachment; style apex 1. 5-1.9 mm wide, slightly
pentagonal, nearly round, flat or slightly concave,
pink. Immature follicles divergent when double, fu¬
siform. to 7 X 1.5 cm, green, with dense mixed
indumentum, long trichomes 1 — 1.5 mm, apex long-
attenuate, base stipitate* with a flange at pedicel
attachment, with scattered prickles (ca. 70 total),
prickles ca. 12 mm long, 1 mm wide at middle;
seeds unknown.

Matelea costaricensis is most closely related to
M. pinguifolia (Standley) Woodson and M. pseiulo-
harbata (Pittier) Woodson, but differs from those in
lacking glandular trichomes on the upper leaf sur¬
face, in having larger flowers, the inside of the co¬
rolla hispidulous, and the outer corona more strong¬
ly pentagonal and nearly entire, lacking the distinct
coronal layers of the related species.

Matelea costaricensis is known only from cloud
forests in the mountains of central Costa Rica at
1500 to 2000 m elevation; flowers have been col-

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Stevens
New Milkweeds

607

Figure 2. Matelea costaricensis W. I). Stevens. Scan of holotype specimen (Morales 42811 , MO).

lected from April to November and immature fruits
in November and December.

Paratypes. COSTA RICA. Alajuela: Zarcern, 4.
Smith 1252 (F). P1970 (MO), 2786 (F). A. Weston 3078
(MO). San Jose: Z.P. Cerros de Eseazu, SE del Alto de
Hierbabuena, ./. Morales 1323 (MO).

Matelea emmartinezii W. 1). Stevens, sp. nov.
TYPE: (cultivated at Missouri Botanical Car¬
den from W. Stevens & E. Martinez S. 25819,
Mexico, Chiapas: 2 km N of Naja, 860 m,
16°59'N, 91°36'W, 4 Sep. 1988], W. Stevens
25956 (holotype, MO). Figure 3.

Differt a Matelea pusilliflora et M. ocellata lobis cor-
ollarum cucullatis.

Vine, older stems woody and covered with thick,
ridged, spongy, tan-colored cork, roots fibrous, latex
white, young stems with a mixed indumentum, long
trichomes in 2 lines, appressed, 0.3— 0.7 mm. white
with red septa, glandular trichomes 0.03—0.05 turn,
pale red-brown, internodes 2—12 cm. Eeaves op¬
posite, blades ovate, 4—5.5 X 1.5—2 cm, apex at¬
tenuate, apiculate or not, base truncate or very
shallowly cordate, pusticulate, adaxially glabrous
except with sparse glandular trichomes on base of
midrib, abaxially glabrous except with mixed in-

608

Novon

Figure 3. Mate lea emmartinezii W. I). Stevens. — A. Flowering branch. — IF Flower. — C. Pollinarium. Drawn from
the holotype, Stevens 25956 (MO).

dumentum on midrib and glandular triehomes on
secondary veins, lateral veins 4 to 5 pairs, strongly
curved, middle veins 40°-45° to midrib, colleters 2
to 4; petiole 0.5—1 cm, with mixed indumentum or
only glandular triehomes. Inflorescence congested-
raeemifomi, peduncle 0.5—1 mm, glabrous, pedicel
2-2.5 mm, with sparse glandular triehomes, bracts
1-2.5 X 0.2— 0.4 mm. deltate to ligulate, glabrous,
purple; calyx tube ca. 0.5 mm long, with 1 eolleter
per sinus, lobes elliptic, ca. 2.2 X 0.9 mm, apex
acute, purple, glabrous or with a few glandular tri-
ehomes on base outside; corolla rotate, green with
dense brownish purple reticulations, with a bright
white spol at tip of each corolla lobe within, adax-

ially glabrous, abaxially with short triehomes and
glandular triehomes on the distal half of each lobe,
tube 1—1.3 mm long, lobes elliptic, the center of
each lobe eueullate, ca. 4 X 2.7 mm, apex rounded
and notched when flattened, patent; gynostegium
with a cylindrical stipe ca. 0.7 mm long, outer co¬
rona a fleshy disk, ea. 1 mm wide, green and glis¬
tening above, base pentagonal in outline and over¬
lapping corolla tube, margin dull, radially striate,
purple, inner corona of 5 small lobes at apex of
stipe, ca. 0.2 X 0.3 mm, lips bilobulate and ap-
pressed to the bases of the anthers; anthers dorso-
ventrally flattened, ca. 0.4 X 1.5 mm. guide rails
straight, parallel, vertical, ca. 0.1 mm long, termi-

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609

nal appendages appressed to margin of style apex,
ea. 0.2 mm wide, white, corpusculum ea. 0.21 X
0.8 mm, sagittate, brown, translators ca. 0.19 mm
long, pollinia ca. 0.81 X 0.24 mm. obovoid, sterile
and excavated on proximal third near center; style
apex ca. 2.3 mm wide, star-shaped, shallowly con¬
vex with a central depression. Follicles and seeds
unknown.

Phis new species is most closely related to Ma-
telea pusilliflora L. (). Williams and M. ocellata W.
I). Stevens, which until now were the only mainland
representatives of a small group of Caribbean spe¬
cies characterized by asymmetrical, narrowly 5-
winged follicles, a corona in the form of a fleshy
disk at the base of the gynostegium stipe, and a
white, reflective eye at the tip ol each corolla lobe.
Matelea emmartinezii differs from both by having
cucullate corolla lobes, by having an outer corona
smooth and unadorned except striate on the outer
margin, and by having a small but obvious inner
corona. Additionally, Matelea ocellata has no pus-
ticulations on the leaves and longer inflorescences
(pedunc les 1—9 mm and pedicels 4—14 mm) and
M. pusilliflora has smaller flowers (corolla lobes
2.3— 2.7 mm long).

Matelea emmartinezii was collected in sterile
condition in an opening in dry forest in Chiapas
and later flowered in cultivation; it is known from
no other collections.

Matelea ferrugiuea W. 1). Stevens, sp. nov. T\ PE:
Guatemala. Chimaltenango: region of Los Pos-
itos, above Las Calderas, 2250—2400 m, 16
Dec. 1940, P. Standley 80219 (holotype, F;
isotype, MO). Figure 4.

Mateleae rnagnifoliae affinis, seel differt gynostegio stip-
itato corona praelonga.

Large vine, older stems and lease1 unknown, latex
unknown, young stems with mixed indumentum,
long trie homes moderately dense, 2—3 mm, straight,
pale yellow-brown, spreading, short triehomes
dense, 0.2— 0.4 mm, straight, pale yellow-brown,
glandular triehomes sparse to moderately dense,
0.2— 0.4 mm. stalk pale yellow-brown, tips brown,
hardly or not at all inflated, internodes 17—22 cm
long. Leaves opposite, blades ovate to elliptic, 12—
19 X 8.7—14 cm, apex acuminate, base lobate,
lobes descending to converging, sinus 1.7— 2.5 cm
deep, pilose, triehomes 0.5—1 mm, pale yellow-
brown. lateral veins 6 to 8 pairs, middle veins 45°—
50° to midrib, colleters 18 to 20; petiole 6.5—12.5
cm, with mixed indumentum. Inflorescence con-
gested-racemiform, with mixed indumentum, pe¬

duncle 55—135 mm. pedicel 18-20 mm, bracts to
1 1.5 X 2.7 mm, elliptic, acuminate; calyx tube ca.
I mm long, with I colleter per sinus, lobes elliptic
to spathulate, unequal, flat or somewhat cucullate,
7.3— 8.6 X 5.4—6 mm, apex rounded, green, with
dense, appressed. yellow-brown long triehomes out¬
side, triehomes 0.5— 1.5 mm, glabrous inside; co¬
rolla very briefly campanulate then rotate, yellowish
green, reticulate when dried, with dense, ap¬
pressed, yellow-brown long triehomes outside, tri-
chomes 0.5—1 mm, glabrous inside, tube ca. 4.8
mm long, lobes broadly elliptic, somewhat asym¬
metric, 10.5—11.5 X 8.6— 9.5 mm, apex rounded
and shallowly notched; gynostegium with stipe ca.
1.3 mm long, corona adnate to stipe, 5-lobed,
fleshy, ca. 1.1 mm tall, 0.9 mm wide at the base,
0.6 mm wide at the top, star-shaped in outline at
base and entirely under the gynostegium, dark-col¬
ored. stipe covered by the corona below, partially
exposed above between the lobes, lobes truncate at
tips and touching the anther bases: anthers rect¬
angular in outline, ca. 0.5 X 1.5 mm. guide rails
rounded, parallel, vertical, ca. 0.2 mm long, ter¬
minal appendages appressed to margin of style
apex, ca. 0.1 X 1.2 mm, white, corpusculum ca.
0.19 X 0.10 mm, sagittate, red-brown, translators
ca. 0.23 X 0.15 mm, straight, pollinia ca. 0.70 X
0.29 mm, obovate, sterile and excavated on proxi¬
mal third; style apex ca. 2.2 mm wide, pentagonal,
flat with a slight umbo in center. Follicles and seeds
unknown.

Matelea ferrugitiea is superficially similar to Ma¬
telea velutina (Schleehtendal) Woodson, from north¬
eastern Mexico, and the original collection had been
so determined. This new species, however, is prob¬
ably more closely related to the also rather similar,
and generally sympatrie, Matelea magnifolia (Pittier)
Woodson. Matelea velutina shares with the new spe¬
cies a vertically oriented corona surrounding the gy¬
nostegium stipe, but the lobes of the outer corona
are broad, tangentially oriented and covering the en¬
tire stipe, concave, yellow and glistening, rather than
narrow, radially compressed and broadest at the
base, attenuate above and not completely covering
the stipe, dark -colored and dull. The corona of Ma¬
telea velutina is similar to that of the smaller-flow¬
ered M. reticulata (Engelmann ex A. Gray) Woodson,
from southwestern U.S.A. and adjacent Mexico, and
both species have the terminal anther appendages
bright, reflective white and covering the entire style
apex, forming a distinctive eye in the center of the
flower. In contrast. Matelea magnifolia and M. fer-
ruginea have terminal anther appendages covering
only the margin of the style apex and have no re-

610

Novon

Figure 4. Malelea ferruginea W. I). Stevens. Sean of holotype specimen (Standley 80219, F).

flective eye; they also share, along with M. veluli-
noide.s, described below, relatively long glandular tri-
chomes with the terminal cell darker colored but
hardly inflated. Matelea magnifolia is easily distin¬
guished by having a faucal annulus, an essentially

sessile gynostegium and a horizontally oriented,
disk-like corona.

Although the Johnston collection is technically a
paratype, it was collected at the same place and on
the same date as the type collection and may be

Volume 15, Number 4
2005

Stevens
New Milkweeds

611

Figure 5. Matelea filipes W. I). Stevens & Monterrosa. Sean of holotype specimen (Sandoval 1 60S, MO).

part of the original gathering; this species has not
been collected in 65 years.

Paralype. GUATEMALA. Cliinialtenango: Calderas,
./. Johnston 1792 (F).

Matelea filipes W. I). Stevens <.A Monterrosa, sp.
nov. TYPE: El Salvador. Achuachapan: San
Francisco Menendez, cabecera de la vereda La
Pinera, 650 m, 13°49’N, 89°56’W, 6 June
1997, E. Sandoval 1608 (holotype, MO; iso¬
type, LAGU). Figure 5.

Mateleae pinguifoliae et M. pseudobarbatne at finis, sed
differt ramis infforescentiae delicatioribus florihus niino-
ribus.

Vine, older stems and base unknown, latex un¬
known, young stems with mixed indumentum, long
trichomes sparse to dense, 1.5— 3.5 mm, straight,
pale yellow-brown, spreading, short trichomes
dense. 0.1—0.15 mm, straight, translucent, glandu¬
lar trichomes dense, 0.07—0. 1 mm, dark brown, in¬
ternodes 15—20 cm. Leaves opposite, blades ovate.

612

Novon

10.2—14.2 X 6.2—10.8 cm, apex abruptly acumi¬
nate, base lobate, lobes convergent to overlapping,
sinus 1-2 cm deep, atlaxially with mixed indumen¬
tum, glandular trichomes sparse, abaxially with
dense short and glandular trichomes, lateral veins
6 to 7 pairs, middle veins 40°— 45° to midrib, col-
leters 7 to 10; petiole 5.8—7 cm, with mixed indu¬
mentum. Inflorescence eongested-racemiform, thin,
with mixed indumentum, peduncle 45—77 mm, axis
to 5 mm, pedicel 17—24 mm, bracts 1 — 1.5 X 0.1 —
0.2 mm, linear; calyx tube ca. 0.5 mm long, with I
colleter per sinus, lobes lanceolate, ca. 2.2 X 1.2
nun. apex acute, green, with mixed indumentum
outside, long trichomes 0.5—1 mm, glabrous inside;
corolla briefly campanulate, then rotate, pale
brown, with mixed indumentum outside, long tri¬
chomes 0.5—1 mm, inside with flexuous, pale yel¬
low trichomes 2-3 mm long, tube ca. I mm long,
lobes broadly ovate, ca. 2.7 X 2.3 mm, apex round¬
ed and notched, patent; gynostegium sessile, outer
corona obconie, circular in outline, glabrous, un¬
dulate and darker-colored on distal margin in 2
rings opposite anthers and 1 ring alternate with an¬
thers. margin equaling base of gynostegium, ca. 0.6
mm tall, inner corona ol 5 adnate ligules opposite
anthers, trapezoidal in outline, lateral margins
raised, ca. 0.6 X 0.5 mm, distal margin similar to
margin of outer corona, producing 3 layers opposite
anthers; anthers trapezoidal in outline, ca. 0.4 X
0.9 mm. tilted under style apex, guide rails straight,
parallel, tilted under style apex, ca. 0.1 mm long,
terminal appendages appressed to stvle apex, ca.
0.3 X 0.7 mm. white to translucent, corpusculum
ca. 0.19 X 0.11 mm, sagittate, red-brown, trans¬
lators ca. 0. 1 I X 0.11 mm. straight, pollinia ca.
0.41 X 0.29 mm, angularly obovate, sterile and ex¬
cavated at attachment; style apex ca. 1.3 mm wide,
slightly pentagonal, nearly round, flat. Follicles and
seeds unknown.

Matelea Jilipes is most closely related to M. pin-
guifolia , from Panama and northern South America,
and M. pseudobarbata, a rare species endemic to
central Costa Rica, with which it shares relatively
thin inflorescence branches, relatively small flow¬
ers, and a peculiar long, wavy indumentum on the
inside of the corolla, although the inflorescence of
M. Jilipes is longer (peduncles 45—77 mm vs. 5—25
mm) and even more delicate (pedicels 0.3 mm
diam. vs. 0.5— 0.7 mm diam.) and the flowers are
smaller (calyx lobes 2.2 mm long vs. 2.5— 4.1 mm
long) than the other two species. Additionally, Ma¬
telea pinguifolia and M. pseudobarbata have a stip-
itate gynostegium and a more massive, stipitate,
and more strongly colored corona, contrasting

sharply with the reflexed corolla of this new spe¬
cies. Matelea haberi is also similar, and has a ses¬
sile gynostegium, but has stouter inflorescences,
larger flowers (corolla lobes 3.9— 4.8 mm long), and
a more dissected corona margin.

Matelea Jilipes is know n only from the type col¬
lection from Fd Salvador. The isotype was studied
by the coauthor of the species. Jorge Alberto Mon-
terrosa Salomon.

Matelea gentlei (Lundell & Stand ley) Woodson.
Ann. Missouri Hot. Card. 28: 234. 1941. Bas-
ionym: Vincetoxicum, gentlei Lundell & Stan-
dley, in Standley, Field Mus. Nat. Mist., Hot.
Ser. 17: 269. 1937. TYPK: Belize. Belize: Gra¬
de Rock, Sibun River, 31 Jan. 1936. /I Gentle
1779 (lectotype. designated here, MICH; du¬
plicates of lectotype, A. F, MO. P. US).

The original diagnosis of Vincetoxicum gentlei
cited “ Lundell ” 1779 and I7IU) as the types. Lun¬
dell (1938) corrected the collector of the types to
Gentle. The lectotype is a flowering specimen and
the unselected syntype, /I Gentle 1700 (F, MICH,
W IS). I. as young fruits.

Matelea liuheri W. I). Stevens, sp. nov. T\ PE:
Costa Rica. Puntarenas: Monteverde, RfoGua-
cimal valley, 800 m, 6 Oct. 1984, IT. Haber
650 (holotype, MO). Figure 6.

Mateleae pinguifoliae el M. pseudobarbatae affinis, sed
statim differt corolla intra glabra.

Vine, older stems and base unknown, latex
white, young stems with mixed indumentum, long
trichomes sparse to dense, 1.5— 3.2 mm. straight,
pale yellow, spreading, short trichomes dense.
0.07-0.13 mm, straight, translucent, glandular tri¬
chomes sparse to moderately dense, 0.1—0.15 mm,
brown to nearly black, internodes I 1—30 cm.
Leaves opposite, blades ovate to elliptic, 6.2—13 X
3.4— 8.6 cm. apex abruptly acuminate, base lobate,
lolx •s descending to overlapping, sinus 0.7— 2.4 cm
deep, adaxially with dense short trichomes, some¬
times with sparse glandular and long trichomes on
veins and at base, abaxially with dense short and
glandular trichomes, lateral veins 5 to 7 pairs, mid¬
dle veins 35°— 40° to midrib, colleters 6 to 10; pet¬
iole 2.4— 5.7 cm. with dense short and glandular
trichomes, often with sparse to moderately dense
long trichomes. Inflorescence raeemiform or eon¬
gested-racemiform, with dense short and glandular
trichomes and sparse to dense long trichomes, pe¬
duncle 8—25 mm, axis to 10 mm, pedicel 6—14 mm.
first bract 5.5— 6.8 X 0.3—1 mm. linear, upper
bracts I. <—3.5 X 0.2— 0.7 mm; calyx tube 0.3-0. 5

Volume 15, Number 4
2005

Stevens
New Milkweeds

613

Figure 6. Matelea haberi W. D. Stevens. — A. Flowering branch. — B. Inflorescence. — C. Corona and gynostegiurn.

_ I). Pollinarium. — K. Nearly mature follicle. A-C drawn from Espinoza 607 (MO). I) drawn from Haber 199 (MO),

F drawn from Rivera 947 (MO).

mm long, with 1 to 3 colleters per sinus, lobes lan¬
ceolate. 2. 1-3. 1 X 1 . 1-1 .8 mm, apex acute, green,
with dense mixed indumentum outside, long tri-
chomes 0.5— 1.5 mm, glabrous inside; corolla ro¬
tate- reflexed, orange to dark reddish brown, tube
1 — 1.5 mm long, with mixed indumentum outside,
glabrous inside, lobes broadly ovate, 3.9— 4.8 X
2.8— 3.8 mm, apex rounded and shallowly notched.

with dense short and glandular triehomes and
sparse to dense long triehomes 0.5— 0.7 mm long
outside, glabrous inside; gynostegiurn with stipe
0. 3-0.4 mm long, outer corona short-stipitate be¬
low. eupuliform above, stipe 0.1— 0.2 nun long, cy¬
lindrical, with color and texture of corolla, upper
part fleshy, glabrous, irregularly divided on outer
surface in 2 rings, deep purple, slightly 5-lobed at

614

Novon

base, somewhat shorter than style apex, tube 0.3—
0.6 mm long, lobes 0.1— 0.2 X 0.3— 0.5 mm, inner
corona of 5 adnate ligules opposite outer corona
lobes and anthers, linear with free truncate or
slightly bifid tips incumbent on backs of anthers,
free tips 0.3— 0.5 X 0.3— 0.4 mm: anthers trapezoi¬
dal in outline, 0. 4-0.5 X 0. 8-0.0 mm, tilted under
style apex, guide rails straight, parallel, tilled under
style apex, ca. 0.1 mm long, terminal appendages
appressed to style apex, 0.3— 0.4 X 0.7— 0.8 mm.
white to translucent, corpusculum 0.15—0.2 X
0.07-0.09 mm. sagittate, pale brown, translators
0.1—0.15 X 0.07—0.1 mm, sigmoid, pollinia 0.27—
0.35 X 0.22—0.25 mm, asymmetrically obovate,
sterile and excavated at attachment; style apex 0.9—
1.2 mm wide, slightly pentagonal, nearly round, flat
or slightly concave, apparently purple. Follicles di¬
vergent when double, fusiform, 8—9 X 1.7—2 cm.
green, with dense mixed indumentum, long tri-
chomes to 2 mm long, sometimes sparse, apex long-
attenuate. base stipitate. with scattered prickles (I
to 3 per cm2, 40 to 50 total), prickles 11-17 mm
long. 0.8— 1.3 mm wide at middle, tips often clavate,
bifid or geniculate: seeds obovate, 7-7.7 X 4.8— 1.9
mm, yellow-brown, margin 0.6— 0.7 mm wide, irreg¬
ularly eremite on distal third to two-thirds, surface
slightly reticulate-rugose, coma 2.5—3 cm long,
white.

Matelea haberi is most closely related to M. pin-
guifolia, M. pseudobarluita , and M. Jilipes, but dif¬
fers most conspicuously in having the inside of the
corolla glabrous; it also lias denser inflorescences,
a more prominently lobed corona, and smaller pol¬
linia (0.27—0.35 X 0.22—0.25 mm vs. 0.40—0.53 X
0.28—0.30 mm) and style apices (0.9— 1.2 mm vs.

I . < — 2.5 nun). I he follicles of Matelea haberi have
longer prickles (11-17 mm vs. 7-11 mm) and
shorter triehomes (to 2.0 mm vs. to 3.5 mm) than
those of V/. pinguifolia and 47. pseudobarbata.

Matelea haberi is known only from cloud forests
in the volcanic mountains of northwestern Costa
Rica at <60 to 1550 m elevation, and it flowers and
fruits from July to January.

Baratypes. COS1A I1ICA. Ciianueuste: I’M. Hmn'm
tie la Vieja, Kst. Fas Pailas. R. Espinoza 6 07 (CIC IMF
MO), I). (,arn'a 36 (MO); Monteverde, A. Gentry, IF! Ilaber.

L Woodruff & B. Boyle 71600 (MO); I’.N. Itineon de la
Vieja, I Ida. Sta. Marfa, / Gdmez L 10312 (V); I’.N. Moron
de la Vieja, sector Los Naranjos, camino a San Jorge, G.
Rivera 017 (MO). I’linlarenas: Monteverde, IF Haber 199
(MO), 581 (MO).

.Matelea insolita W. 1). Stevens, sp. nov. TYPE:
Nicaragua. Estelf: Reserva Natural Miraflor.
Comarca Sontule, 1000-1100 m, 13°12’N,
86°20'W, . July 1999, R. Rueda, I. Coronado ,
H. Velasquez & M. Arroliga 1 1427 (holotvpe,
MO; isotype, HULE). Figure 7.

Ex affinitate Mateleae pinguifoliae el specierum at fi¬
ll i u m corona externa tenui alba quam corona interna bre-
viori confestim distinguenda.

Vine, older stems and base unknown, latex un¬
known, young stems with mixed indumentum, long
triehomes sparse. 0.4-2 nun. straight, yellow-
brown, spreading, short triehomes dense. 0. 1-0.2
mm, straight, white, glandular triehomes dense, ca.
0.1 mm, dark brown, internodes 10-16 cm. Leaves
opposite, blades ovate, 6.8-12.1 X 3.6-7 cm, apex
abruptly acuminate to attenuate, base lobate, lobes
descending to convergent, sinus 0.6-1. 6 cm deep,
adaxially with dense short triehomes and a few
glandular triehomes at base of midrib, abaxially
with dense short and glandular triehomes. lateral
veins 5 to 7 pairs, middle veins 35°^150 to midrib,
colleters 10 to 20: petiole 3.6— 6.9 cm, with dense
short and glandular triehomes. Inflorescence eon-
gested-racemiform, with mixed indumentum, long
triehomes sparse, peduncle 10-25 mm, axis to 5
mm, pedicel 15-35 mm, bracts 1 .5-1.8 X 0.3-0. 4
mm, linear or narrowly lanceolate; calyx tube 0.5—
0.8 mm long, with I colleter per sinus, lobes lan¬
ceolate, 3. 1-3.5 X 1.2— 1.3 mm, apex acute, green,
with mixed indumentum outside, long triehomes
sparse. 0.3— 0.5 mm. glabrous inside or hispidulose
at tip; corolla briefly eampanulate, then rotate, yel¬
lowish. with mixed indumentum outside, long tri¬
ehomes sparse, 0.3— 0.8 mm, inside with flexuous.
white triehomes 0.8— 1.2 mm long in tube and bast*
of lobes, tube ca. 2.5 mm long, lobes lanceolate,
5. 7-6.5 X 3.5— 3.6 mm, apex rounded and notched,
patent; gynostegium sessile, outer corona laminar,
flat, horizontal, dull white, 5-lobed, entirely hidden
under gynostegium, tube ca. 0.2 mm long, lobes ca.
0.4 X 0.9 mm, irregularly toothed, inner corona of
5 adnate ligules opposite anthers, rectangular in
outline, fleshy, purple-black, ca. 0.8 X 0.6 mm, tips
bifid and appressed to anther backs; anthers ellip¬
tic in outline, ca. 0.5 X 1.1 mm. guide rails
straight, parallel, slightly tilted under style apex,
ea. 0.1 mm long, terminal appendages appressed to
margin of style apex, ovate, ca. 0.3 X 0.8 mm,
translucent, corpusculum ca. 0.18 X 0.08 mm. sag¬
ittate, red-brown, translators ca. 0.1 I X 0.09 mm.
straight, pollinia ca. 0.35 X 0.16 mm, asymmetri¬
cally obovate, sterile and excavated on proximal
two-thirds; style apex ca. 1.4 mm wide, slightly

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New Milkweeds

615

1 cm

Figure 7. Matelea insolita W. I). Stevens. Scan of holotype specimen ( Rueda el al. 11427, MO).

pentagonal, nearly round, shallowly concave. Im¬
mature follicles fusiform, to 13.5 X 2.5 cm, green,
with dense short and glandular trichomes, apex
long-attenuate, base stipitate, with more or less
dense prickles (ea. 4 per cm2, ca. 150 total), prick¬

les 20—30 mm long, 0.6— 1.2 mm wide at middle,
tips sometimes slightly elavate; seeds unknown.

Matelea insolita is clearly of the M. pinguifolia
alliance, which shares nearly symmetrical fusiform

616

Novon

follicles that have relatively sparse long, thin,
curved prickles and a dense mixed indumentum,
and which ranges from Mesoameriea to northern
South America. Matelea insolita is vegetalively
(|uite similar to M. pinguifolia, hut is the only spe¬
cies of t he alliance, and perhaps in the whole ge¬
nus, where the outer corona is white and practically
vestigial and the inner corona is prominent. Mate-
led insolita lias only been found in northern Nica¬
ragua, between 900 and I 100 m elevation. The
flowers were collected in July and the nearly ma¬
ture fruits in January.

Paratype. NICARAGUA. Jinotega: R.iN. kilambe,
Santa Teresa do Kilambe, H. Rueda, I ). Paguaga, II. Men¬
doza , A. Rivera, l\. Toval & M. Garmendia 1 559 1 (HULE,
MO).

Matelea velutinoides W. I). Stevens, sp. nov.
T\ PE: Mexico. Chiapas: Mpio. Cintalapa, SE
of Cerro Haul on border with Oaxaca. 16 km
NW of Rizo de Oro along logging road to Co-
lonia Figaroa, 1600 m, 6 Sep. 1972, I). Breed¬
love 27(>12 (holotype, MO). Figure 8.

Maleleae velutinae affinis. sed lobis coronae radiantibus
lohis eorollae intra ad margineni hispidulosis distinguen-
da.

Vine, older stems and base unknown, latex
white, young stems with mixed indumentum, long
triehomes sparse to dense, 1.5— 2.5 mm, straight,
yellow to brown, spreading, short triehomes dense,
0.2— 0.5 mm, straight, yellow to brown, glandular
triehomes dense, 0.2— 0.5 mm, stalks pale brown,
tips black-brown, hardly or not at all inflated, in¬
ternodes 12—20 cm. Leaves opposite, blades ovate
to elliptic. 9.8—13.8 X 4.8— 7.2 cm, apex acumi¬
nate. base Inhate, lobes descending to overlapping,
sinus 0.7— 1.3 cm deep, adaxially hispid, triehomes
0.5— 1.5 mm, pale brown, short and glandular tri¬
ehomes present on midrib, abaxially hispid, tri¬
ehomes 0.5— 1.5 mm, short and glandular triehomes
present on veins and sometimes on surface, lateral
veins 6 to 8 pairs, middle veins 35°— 45° to midrib,
eolleters 10 to 24, thin, with light colored tips; pet¬
iole 3.7— 5.5 cm, with mixed indumentum. Inflores¬
cence racemiform or congested-racemiform, with
mixed indumentum, peduncle 35—95 mm, axis to
1 1 mm, pedicel 9—24 mm, bracts to 9 X 2 mm,
lanceolate; calyx tube 0.5—1 mm long, with 1 eol-
leter per sinus, lobes elliptic, unequal, flat or some¬
what eueullate, 4.3—6 X 2.3-4 mm, apex rounded
to acuminate, green, with mixed indumentum out¬
side, long triehomes 1-2.5 mm, sometimes restrict¬
ed to distal half, glabrous inside or with scattered
glandular triehomes; corolla very briefly eampanu-

late then rotate, brown to yellowish orange, retic¬
ulate when dried, with mixed indumentum outside,
long triehomes 0.5—1 mm, glabrous inside except
inconspicuously hispidulose in a narrow band along
I margin of each lobe, sometimes extending into
tube, triehomes 0.05—0.1 mm, translucent, tube
3.2— 3.6 mm long, lobes broadly elliptic, somewhat
asymmetric, 6.2-7. 1 X 5.7— 7.4 mm. apex rounded
and shallowly notched; gynostegium with stipe 0.9—

1.1 mm long, outer corona entire or shallowly 5-
lobed at base, base horizontal and overlapping the
corolla, erect above and entirely enclosing gynos¬
tegium stipe, fleshy, with radial crests, 0.8— 1.2 mm
tall, 0.7— 1.1 mm wide at the base, dark-colored,
inner corona borne on outer corona, erect, forming
radial wings on upper part of outer corona, lighter-
colored, 0.5— 0.6 X 0.3— 0.4 mm, tips touching the
anther bases; anthers rectangular in outline, 0.5—
0.7 X 1.4— 1.5 mm, guide rails rounded, parallel,
somewhat tilted under style apex, 0.15—0.2 mm
long, terminal appendages appressed to style apex
and covering about half of surface, 0.5— 0.8 X 1-
1.4 mm, white, eorpusculum 0.23-0.25 X 0.09
mm, sagittate, red-brown, translators 0.19—0.25 X
0.14—0.15 mm. straight, pollinia 0.65—0.69 X
0.25—0.29 mm, obovate, sterile and excavated in
proximal third; style apex 2. 1-2.6 mm wide, pen¬
tagonal. flat or slightly concave when dry, with a
slight umbo in center. Follicles and seeds un¬
known.

Matelea velutinoides is superficially similar to
Matelea vela tin a, from northeastern Mexico, and
most of the collections were distributed under this
name. Matelea velutinoides has the corona oriented
horizontally below and vertically above, but the
vertical part is dark-colored and radially oriented,
rather than yellow and tangentially oriented, as in
M. velutina. Matelea velutina also has larger (to

15.1 X I I cm vs. to 13.8 X 7.2 cm) and propor¬
tionately broader leaves, larger flowers (corolla
lobes 7.1—10.0 mm long vs. 6.2-7. 1 mm long), co¬
rollas that are entirely glabrous within, and termi¬
nal anther appendages that entirely cover the style
apex. Matelea velutinoides differs from M. ferrugi-
n ea in having a spreading, mixed indumentum on
the outside of the calyx and corolla, rather than a
simple appressed indumentum, and in the corona
covering the entire gynostegium stipe.

Matelea velutinoides is known from cloud forest
in Chiapas and Guatemala between 1600 and 2200
m elevation and is known to flower between April
and September.

Paratypes. MEXICO. Chiapas: Mpio. Motozintla, 15
km al () de Motozintla, camino a Siltepec, K. Martinez .S'.

Volume 15, Number 4
2005

Stevens
New Milkweeds

617

Figure 8. Matelea velutinoides \\. I). Stevens. Scan of holotype specimen (Breedlove 27642, MO).

et al. 20741 (MO); Carelas, E. Matuda 15517 (F); Mpio.
Fseuintla, Cacaluta, E. Matuda 17001 (1. MFXU); Mpio.
Siltepee, Cascada, E. Matuda 17642 (F); Cerro del Bo-
queron, C. Purpus 7277-B (F); Mpio. Motozintla, FI Ro¬
sario. E. Ventura & E. I^dpez 3935 (F). GUATEMALA. Kl
Quiche: falls of Rio de las Violetas, 2.5 mi. N of Nejal),
G. Proctor 25445 (CAS. F. TEX).

Pherotrichis schaffneri A. Gray, Syn. FI. N. Amer.
(ed. 2) 2(1): 462. 1886. TYPE: Mexico. “North¬
ern Mexico,” ,/. Schaffner 65 (leetotype, desig¬
nated here, GH; duplicate of leetotype, K).

(day (1886a) validly, but minimally, published

618

No von

Pherotrichis balbisii (see below) and P. schaffneri,
where lit' t iles the earlier written, but later pub¬
lished, fuller treatment ((bay, 1886b). Gray inten¬
tionally and speeifieally plaeed one syntype of Gon-
olobus pogonanthus (see below) (Ghiesbreglit 670)
in his Pherotrichis balbisii and the other (Parry &
Palmer 586) In Ins P. schaffneri , thereby effectively
lectotypifying the Hemsley name with the Ghies-
breght specimen and placing it in synonymy of his
superfluous P. balbisii (= P. villosa (Schultes) Meis-
ner). The unselected syntypes of Pherotrichis
schaffneri are: C. Parry & E. Palmer 58 6 (syntype,
GH; isosyntypes, k. MO) and ,/. Lemmon 2016
(syntype, GH). Pherotrichis schaffneri is almost cer¬
tainly a species distinct from P. villosa (Schultes)
Meisner and the leelotvpification stabilizes that us¬
age.

IMicrolrieliis villosa (Schultes) Meisner, PI. Vase.
Gen. 2: 170. 1810. Cynanchum villosum
Schultes, in Koemer & Schultes, Svst. Veg. 6:
103. 1820. based on Asclepias villosa Balbis,
M< •rn. Acad. Sci. Turin, Sci. Phys. 7: 331, t. 4.
1803-4, not A. villosa P. Miller, 1768. TYPE:
unknown provenance cultivated at the Turin
botanical garden, G. Balbis s.n. (holotype, TO,
not seen; isotypes: G, MO).

(hmolobus pogonanthus Hemsley, Biol. Cent.-Amer., But.
2(11): 333. 1882. TYPE: Mexico. Chiapas, 1864—
1870. A. Ghiesbreglit 670 (leetotype, designated hy
Gray, Proc. Amer. Acad. Arts 21: 400. 1886. k; du¬
plicate of leetotype, GH).

Pherotrichis Decaisne was validly published (De-
caisne, 1838), but the combination based on the
only included species, Asclepias villosa Balbis, now
known to be a later homonym of A. villosa P. Miller,
was not made. Later, Decaisne (1844) included
Pherotrichis in his concept of Lachnostoma and
published the name /,. balbisii Decaisne (in A. de
Candolle, Prodr. 8: 602. 1844), a superfluous name
based on Asclepias villosa Balbis. The names Pher¬
otrichis villosa (Schultes) k. Schumann (in Engler
& Prantl, Nat. Pflanzenfam. 4(2): 303. 1895), P.
balbisii A. Gray (Syn. FI. N. Amer. (ed. 2) 2(1): 462.
1886). and Matelea balbisii Woodson (Ann. Missou¬
ri Bot. Gard. 28: 231. 1941) are likewise based on
Asclepias villosa Balbis and superfluous. The spe¬
cies has generally been treated as Matelea balbisii
(Standley & Williams, 1969; Breedlove, 1986) and
Pherotrichis balbisii (Stevens, 2001), but the over¬
looked Pherotrichis villosa (Schultes) Meisner must
be used for the species when placed in the genus
Pherotrichis. Asclepias villosa Balbis was described
from material of unknown provenance cultivated at
the Turin botanical garden and specimens appar¬

ently distributed by Balbis (e.g.. a specimen at G:
"misit Balbis 1805, non indigena pedem., in ho. T
culta") are taken to be tvpe material.

Polysteinimi giiateinalense (Schlechter) W. I).
Stevens, comb. nov. Basionym: Labidostelma
guatemalense Schlechter, in Loesner, Bull.
Herb. Boissier, ser. 2, 6: 843. 1906. TYPE:
Guatemala. Huehuetenango: Nentdn, Sep.
1887. G. Seler A E. Seler 0279 (leetotype, des¬
ignated here, F).

Vincetoxicum quirosii Standley, Publ. Field Mus. Nat.
Hist., Bot. Ser. 18: 959. 1938. Syn. nov. Matelea
(juirosii (Standley) Woodson, Ann. Missouri Bot.
Gard. 28: 224. 1941. TV PE: Costa Kira. Puntarenas:
Manzanillo, 22 June 1937, M. (hunts C. 7.12 (holo¬
type, F).

Polystemma Decaisne has long been considered
a synonym of Matelea Aublet but was resurrected
by Stevens et al. (2001) for the single species P.
rindifloriim Decaisne. The only other names attri¬
buted to the genus. Polystemma rupestre Brandegee
and P. scopulorum Brandegee are considered by
this author to be synonyms of P. viridiflorum. This
is thus the second species of Polystemma: however,
about 18 additional Mexican species, currently
placed in Matelea, will be added in a fuller taxo¬
nomic treatment. Species of Polystemma can be'
distinguished from those of the current concept of
Matelea by having glandular trichomes that are ini¬
tially translucent but turn white and crystalline
with age, while those of Matelea, when present, are
red, yellow, brown, or black and do not become
crystalline. The glandular trichomes of the related
genus Dictyanthus are also translucent but remain
translucent. The follicles of Polystemma are long,
thin, symmetrical, smooth, and mottled, states in¬
dividually rare or non-existent in Matelea and nev¬
er found in this combination. The distal-dorsal mar¬
gin of the outer corona of Polystemma is usually
provided with long, filiform appendages, although
these can be short in some species or, in the case
ol Polystemma guatemalense, stout.

The holotype of hibidostelma guatemalense was
at B and is no longer extant. A fragment of the
holotype at F, although small, serves as an adequate
leetotype.

Acknowledgments. Alba Arbelaez prepared the
line drawings and Fred keusenkothen and Brigham
Fisher prepared the images. Amy Pool improved
the manuscript, especially the nomenclatural and
bibliographic details. Gordon McPherson and Vic¬
toria Hollow ell provided helpful comments.

Volume 15, Number 4
2005

Stevens
New Milkweeds

619

Literature Cited

Breedlove. I). K. 1986. Flora de Chiapas. Listados Florist.
Mexico 4: i— v, 1—246.

Deeaisne, J. 1838. fitudes sur t[uelques genres et espfeces
de la famille des Aselepiadecs. Ann. Sci. Nat.. Bot.,
s£r. 2. 9: 257-278, 321-348. pi. 9-12.

- . 1844. Asclepiadeae. In A. L. P. P. de Candolle.

Prodromus Systematis Naturalis Regni Vegetabilis 8:
490-665.

Goyder, I). J. & A. Nicholas. 2001. A revision of Gom-
phocarpus R. Br. (Apocynaceae: Asclepiadeae). Lew
Bull. 56: 769-836.

Gray, A. 1886a. Asclepiadaceae. Syn. hi. N. Amer. (ed.
2) 2(1): 85-106, 401-404, 462.

- . 1886b. Contributions to American botany. XXIV.

2. Sertum Chihuahense. Proc. Amer. Acad. Arts 21:
378-1 1 3.

Greuter, W.. J. McNeill. K. R. Barrie, H. M. Bordet. V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
,1. F. Skog. I*. Trehane. N. J. Turland & 1). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Lundell, C. L. 1938. Studies of Mexican and Central
American Plants. VI. Amer. Midi. Naturalist 20: 236-
242.

Murillo, G. N. 1997. Asclepiadaceae. In J. A. Steyermark,
P. E. Berry X B. K. Holst, Flora of the Venezuelan
Guayana 3: 129-177.

Omlor, R. 1998. Generische Revision der Marsdenieae
(Asclepiadaceae). Dissertation. Universitat Kaiserlau-
tern. Shaker Verlag, Aachen.

Rothe, \\. 1915. Cher die Gattung Marsdenia R. Br. uml
die Stammpllanze der Condurangorinde. Bot. Jahrb.
Syst. 52: 354 — 434.

Standley, P. C. & L. () Williams. 1969. Flora of Guate¬
mala. Asclepiadaceae. Fieldiana, Bot. 24 (8. part 4):
407-472.

Stevens, W. D. 2001. Asclepiadaceae. hi G. C. de Rze-
dowski & J. Rzedowski (editors). Flora Fanerogdmica
del Valle fie Mexico, ed. 2. Institute de Fcologfa. Patz-
cuaro, Mexico.

- . C. Ulloa L., A. Pool & 0. M. Montiel. editors.

2001. Flora de Nicaragua. Monogr. Syst. Bot. Missouri
Bot. Gard. 85: i— xlii. 1—2666.

Woodson, R. E., Jr. 1941. North American Asclepiadaceae
I. Perspective of the genera. Ann. Missouri Bot. Gard.
28: 193-244.

- . 1954. Lhe North American Species of Asclepias

L. Ann. Missouri Bot. Gard. 41: 1—21 I.

Novelties in Cynanchum L., sensn Woodson, in Mesoamerica

It' I). Stevens

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

stevens@niobot.org

Abstract. Eleven species and one genus from
Mesoamerica are described as new and four new
combinations are provided for taxa previously in¬
cluded in Cynanchum (Apocynaceae, Asclepiadoi-
deae) from Mesoamerica. Liedea is newly described
to accommodate L. jilisepala. Cynanchum surruhri-
florum. Metastelma brachymischum, M. thalamosi-
phon, M. thysanotum. M. triodontum, M. yucata-
nense, Orthosia cynaruhoides, 0. extra-axillaris, O.
ramosa, (). smaragdina, and 0. stipitata arc de¬
scribed. Combinations are provided for Orthosia
glaherrima, (). miscra , and 0. ruhens.

Key words: Apocynaceae, Asclepiadoideae, Cy¬
nanchum, Liedea. Metastelma. Orthosia, Mesoam¬
erica.

Fislibein and Stevens (2005 this issue) discuss
the rationale and consequences of the disarticula¬
tion of Woodson’s (1941) concept of New World Cy¬
nanchum E. Although the process is now well under
way, there are still many uncertainties in the ge¬
neric circumscriptions. In providing names for an
upcoming volume of Flora Mesoamericana, I mirror
the approach of Rapini (2002) for South American
taxa. in gradually realigning the species into genera
that better reflect our current understanding of re¬
lationships but without breaking the bonds of tra¬
ditional morphology.

Cynanchuin L.. Sp. PI. I: 212. 1753. TYPE: Cy¬
nanchum aeutum E.

It now seems clear that, in the New World, Cy¬
nanchum is represented only by Cynanchum sub¬
gen. Mellichampia Woodson (e.g., Rapini et al.,
2003), a group of about 15 species ranging through
North and South America. This new species (its
easily into that group.

I. Cyiiancliiiin siirruhriflorum W. I). Stevens,
sp. nov. TY PE: Mexico. Tabasco: Maeuspana,
Ealda del Cerro de Maeuspana, al E de la
planta de Cal, Km 2.2 de la earretera, 75 m,
17 June 1979, C. Cowan 2326 (holotype, MO:
isotypes MO, TEX). Figure I.

Cynancho jaliscano affinis sial antheris angustiorihus
alis hrevioribus caudiculis brevioribus differt.

Twining vines, lower stems and base unknown,
latex white; young stems tomentulose at the nodes
and in I line on the internodes, trichomes 0.1— 0.2
mm long, recurved, ferrugineous, internodes 8-9
cm. Leaves opposite, apparently without pseudo¬
stipules, blades ovate, 8.1—12.5 X 4.6— 5.8 cm.
apex acuminate, base shallowly lobate. sinus to 0.6
mm deep, glabrous or with a few scattered tri-
cliomes on the veins, lateral nerves 4 or 5 pairs,
middle veins 40°-50° to midrib, colleters 3 to 5;
petioles 4. 5-6.1 cm, tomentulose on adaxial sur¬
face. Inflorescence extra-axillary. 1 per node, ra-
eemiform, many-flowered, tomentulose on 1 side,
peduncle 1 .2-5.4 cm, axis to 5 cm, pedicel 6.8-8
mm, bracts 0.4— 1.5 X 0.2— 0.5 mm, debate to lan¬
ceolate; calyx with I colleter below each sinus
within, lobes elliptic with obtuse lips. 1.7— 2.3 X
0.6—1 mm, glabrous or inconspicuously tomentu¬
lose at base abaxially, ciliolate, trichomes ca. 0.2
mm long, glabrous inside, green with a purple tint;
flower buds conical, somewhat apiculale; corolla ro¬
tate or slightly campanulate, glabrous, white with a
pink tint, tube 0. 2-0.6 mm long, lobes lanceolate,
with 2 submarginal fleshy ridges, tips rounded,
nearly truncate, erect then spreading from above
the middle, 3.1—5 X 1.3— 1.7 mm; gynostegium with
slip*' 0.3— 0.4 mm long (from base to inflated anther
back), ribbed, corona borne at base of stipe, tube
0.2— 0.4 mm long, the 5 free lobes laminar, trilob-
ulate. 4—5.2 mm long, body of each lobe elliptic,

1 .6— 2 X 0.8— 1.3 mm. lateral lobules ca. 0.4 mm
long, central lobule narrowly debate, nearly linear,
twisted, terminal anther appendages ovate-retuse,
0.5— 0.7 X 0.8— 0.9 mm, inclined toward style apex,
white, anthers T-shaped in outline, 0.8—1 X I mm,
guide rails rounded, vertical, ca. 0.2 mm long; pol-
linaria pendent, corpusculum 0.1—0.19 X 0.11 —
0.13 mm, ellipsoid or subsagittate, dark red-brown,
translators 0.31—0.33 X 0.06 mm, horizontal,
slightly flattened, slightly geniculate, red-brown,
pollinia 0.73—0.74 X 0.15—0.21 mm, narrowly ob-
ovoid: style apex 1.5— 1.8 mm w ide, slightly convex,
densely papillate. Follicles and seeds unknown.

INovon 15: 620-641. Published on 12 December 2005.

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

621

Figure 1. Cynanchum surrubrifioriim W. I). Stevens. Photograph of branch and opened flower of Cowan 2326 (holotype,
MO).

This new species is most closely related to Cy¬
nanchum jaliscanum (Vail) Woodson, with which it
shares a stipitate gynostegium, short guide rails,
and long, horizontal, pigmented translators. V hen
Sundell (1981) treated Cynanchum jaliscanum,
there were available for study only the three orig¬

inal syntype collections from Mexico, two from Jal¬
isco and one from Veracruz, and one additional col¬
lection from Nayarit. There are now 16 collections
available, not counting the Veracruz collection, and
all range from Nayarit to Miehoacan at 770—1400
m elevation, making the Veracruz collection. Batten

622

No von

s.n., relatively more anomalous than it might have
seemed to Sundell. On closer examination, the Ve¬
racruz collection, with the ambiguous locality "Ori¬
zaba. generally matches a collection from lowland
Tabasco, the type designated here for Cynanchum
surrubrifiorum. In addition to the difference in el¬
evation, as lar as known, and obvious geographical
distinction, and it is rare for non-weedy milkweeds
to occur on both sides of Mexico, Cynanchum sur¬
rubrifiorum differs from C. jaliscanum in its shorter
corolla tube. 0.2— 0.6 nun versus 0.6— 2 mm, shorter
gynostegium stipe. 0.3-0. 4 mm versus 0.4— 0.8 mm.
narrower corona lobes, 0.8— 1.3 mm versus 1.3— 1.9
mm, narrower anthers, 1.0 mm versus 1.3- 1.9 nun,
shorter guide rails, 0.2 mm versus 0.3— 0.5 mm, and
shorter translators, 0.31—0.33 mm versus 0.53—0.64
mm. V\ bile most of the differences are not individ¬
ually dramatic, the totality suggests recognition of
tin* east coast lowland species as distinct from the
west coast upland species, Cynanchum jaliscanum,
and the two are relatively easily distinguished from
tin* other species of Cynanchum.

Paratype, MEXICO. Veracruz: Orizaba, 1850-1874,
M. Botleri s.n. (Oil. NY).

Tie dea W. I). Stevens, gen. nov. TYPK: Liedea jil-
isepala (Standley) W. I). Stevens.

A Cynaneho sect. Microphyllo I.iede inflorescentiis pe-
dunculatis panindiformihus eaulibus aequabiliter pubes-
centibus recedit.

(wining perennial vines with sympodial branch¬
ing, apparently woody but not corky below, latex
white: stems uniformly pubescent with multicellular
trie homes of uniform length. Leaves opposite, with¬
out pseudostipules, small, ovate, petiolate. Inflores¬
cence extra-axillary, I per node, pedunculate, cy-
tnose, fundamentally paniculiform but with some
axes reduced to produce apparently raeemiform in¬
florescences with 4 flowers and 1 axis per inflores¬
cence node or all axes reduced and nearly umbel-
liform. Calyx connate at base, with colleters below
sinuses within; corolla campanulate, glabrous with¬
in, aestivation imbricate and dextrorse; gynoste¬
gium short-stipitate, corona borne on gynostegium
stipe, laminar, connate below, briefly 5- or 10-lobed
distally; anther connectives dorsally thickened and
convex, terminal appendages appressed to style
apex; pollinia pendent, uniformly fertile, translators
delicate, round in section, without appendages, cor-
puseula triangular in outline; style apex shallowly
convex, smooth. Follicles apparently single, fusi¬
form. symmetrical, thin-walled, smooth: seeds elon¬
gate, cucullate, with an entire wing, surface smooth,
comose.

I. Liedea filisepala (Standley) W. I). Stevens,
comb. nov. Basionym: Metastelma filisepalum
Standley, Publ. Field Mus. Nat. Hist.. Bot. See.
18: 956. 1938. Cynanchum filisepalum (Stan¬
dley) L. (). Williams, Fieldiana, Bot. 32: 36.
1968. TYPE: Costa Rica. Alajuela: entre San
Ramon v La Palma de San Ramon. 30 June
1928, A. Brenes 6179 (holotype, F).

I.iede (1997) placed Cynanchum filisepalum,
from Costa Rica and Ecuador, in Cynanchum sect.
Microphyllum I.iede, along with nine high-altitude
northern Andean species. This section was defined
primarily by the “dichasial" branching pattern, un¬
bearded corollas, and single follicles. The molec¬
ular study of I.iede and Tauber (2002) suggests that
Woodsons (1941) concept of New World Cynan¬
chum is untenable and that Cynanchum sect. Mi¬
crophyllum can no longer be included in tin1 cur¬
rent concept of the genus Cynanchum. At the
generic level, the name available for Cynanchum
sect. Microphyllum is Scyphostelma Baillon (I.iede
& Meve, 2004). However, it seems clear to this
author that Cynanchum filisepalum does not belong
with the other nine species in Scyphostelma and
Liedea is here described as new with but one spe¬
cies, although the generic description is prepared
to accommodate several apparently undescribed
species from northern South America.

I he sympodial branching pattern of Liedea, and
the remaining species of Cynanchum sect. Micro¬
phyllum. is shared with many species of the related
genera Ditassa R. Brown, Orthosia Decaisne, Sten-
omeria Turczaninow, and Tassadia Decaisne. This
branching pattern is characterized by twining, in¬
determinate, more or less sterile shoots with long
internodes and more or less non-twining, determi¬
nate, fertile, lateral branches with short internodes
and distichous leaves; in the extreme, especially in
Stenomeria and Tassadia, the lateral branches be¬
come leafless and appear to be inflorescences. Lie¬
dea differs from the remainder of Cynanchum sect.
Microphyllum in inflorescence structure, stem in¬
dumentum, and habit. The other nine species have
sessile, fasciculate inflorescences, while those of
Liedea are pedunculate and paniculiform. The typ¬
ical milkweed raeemiform inflorescence, as for ex¬
ample in Cynanchum sect. Mellichampia Sundell,
has two flowers and one continuing axis at each
node, while a paniculiform inflorescence has one
flower and two axes, or even three axes and no
flowers, at each node. Liedea has three or four flow¬
ers and one axis at each node, hut on close ex¬
amination one or two of the flowers are borne on
reduced axes; thus it is a paniculiform inflores-

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

623

cence with the general appearance of a racemiform
one. Liedeu has stems that are uniformly pubescent,
while the species of Cynanchum sect. MicrophyUum
that I have been able to examine have the indu¬
mentum confined to one or two lines. Liedea Jili-
sepala seems to he a high-climbing liana in cloud
forest at 1000—1500 m in Costa Rica (e.g., A.
Skutch A 1 65, MO). Panama (e.g.. /V. Hensold 1018,
MO), and Ecuador (e.g.. A. Gentry & G. Shupp
26877, MO), while the other species of Cynanchum
sect. MicrophyUum seem to be low herbs or sub¬
shrubs in open vegetation at 2300—4200 m in the
northern Andes from Venezuela to Peru.

Melastclmu R. Brown. Asclepiadeae 41. 1810.
TYPE: Metastelma parviflorum (Swartz) R.
Rrown ex Schultes.

Metastelma, as represented hy the type, is rela¬
tively easily circumscribed in North America. Cen¬
tral America, the Caribbean, and northern South
America, and has been accepted in most modern
Holistic treatments. The inclusion of a disjunct
group of species from southern South America,
however, is more problematical and has led some
authors, e.g., Rapini (2002), Rapini et al. (2003).
and Liede-Schumann et al. (2005), to question the
genus as a whole. Eive new species of Metastelma
in the sense of the type species are described be¬
low.

I. Metastelma brachymischum \\. I). Stevens,
sp. nov. TYPE: Mexico. Chiapas: El Sumidero,
22 km N ofTuxtla Gutierrez, 1350 m, 19 Aug.
1972. 1). Breedlove 27 188 (holotype, MO), fig¬
ure 2.

A speciehus Metastelmatis stipitibus hrevibus et coron-
is laminaribus terminalibus haec lohis corouae integristri-
chomatibus corollae longiorilms iiiflorescenliae breviori-
hus distinguenda.

Twining vines, woody and corky below, cork
brown, latex unknown, underground parts un¬
known; young stems puberulenl on I side with re¬
curved white trichomes 0.1— 0.2 mm long, inter¬
nodes 1—3 cm. Leaves opposite, without
pseudostipules, blades elliptic to ovate, 2.3— 4. 1 X
0.7— 1.2 cm, apex acute to acuminate, apiculate,
base obtuse to rounded, adaxially glabrous except
inconspicuously puberulenl on midrib and margin,
abaxially glabrous, lateral nerves 10 to 12 pairs,
middle veins 40—45° to midrib, eolleters 0 to 2;
petioles 2— 4 mm, puberulent on adaxial margin. In¬
florescence extra-axillary, 1 per node, sometimes
aggregated on short leafless shoots, eongested-ra-
cemiform. 10- to 20-flowered. peduncle 0.5—1 mm.

puberulent on 1 side, axis to 3 mm, thickened by
a tight spiral of pedicel bases, pedicel 0.5—1 mm,
puberulent on 1 side, sulcate when dry, bracts 0.2—
0.4 X 0.1— 0.3 nun, ovate; calyx with 1 colleter be¬
low each sinus within, lobes ovate with rounded
tips, 0. 7-0.8 X 0.6— 0.7 mm, puberulent at base
abaxially, ciliolate, glabrous inside, green to purple
with scarious margins; corolla campanulate, white,
tube 0.8—1 mm long, glabrous, lobes lanceolate
with acute tips, erect. 1.7— 1.8 X 0.9—1 mm, abax¬
ially glabrous, adaxially with marginal trichomes
erect, white, unicellular, 0.3— 0.5 mm long distally,
becoming 0.15—0.2 mm long proximally, not reach¬
ing base of lobes, and with center trichomes dense,
retrorse, translucent, unicellular. 0.25—0.35 nun
long; gynostegium with stipe 0.4— 0.5 X 0.5— 0.7
mm, broadly winged, corona lobes borne at or near
top of stipe, lanceolate with attenuate tips, laminar,
loosely appressed to anthers and meeting over style
apex, free part 0.8—1 X 0.2— 0.4 mm, terminal an¬
ther appendages ovate, 0.3—0.35 X 0.3— 0.4 mm,
erect or inclined toward style apex, white, anthers
quadrate in outline. 0.3-0. 4 X 0.5-0.55 mm, guide
rails straight, at ea. 45° to gynostegium axis, salient
at base. ca. 0.25 mm long; pollinaria pendent, cor-
pusculuin 0.17—0.19 X 0.06—0.07 nun. obovoid.
pale brown, translators 0.06—0.09 mm, flattened,
bent at middle, pollinia 0.17—0.19 X 0.07—0.09
mm, obovoid with a narrow neck; style apex 0.6—
0.8 mm wide, strongly convex, ca. twice the height
of eorpuscula. Follicles and seeds unknown.

Metastelma brachymischum is not obviously re¬
lated to any other species in the genus. It shares
the general shape of the pollinaria and corona with
the rather polymorphic and widespread Metastelma
schlechtendalii Decaisne, but that species has a
proportionally shorter corolla tube (about a quarter
of the total corolla length vs. about half), corolla
lobes with a pair of submarginal fleshy ridges,
shorter corolla trichomes (center trichomes 0.05—
0.12 mm vs. 0.25—0.35 mm), a nearly flat style
apex, and longer inflorescences (1—15 mm vs. 1—3
mm), among other characters. The guide rails of
this new species are similar to those of Metastelma
chiapense A. Gray, M. liesnerianum (L. 0. \\ illiams)
Liede, M. longicoronatum (L. O. Williams) l.iede.
and M. thalamosiphon W. I). Stevens, described be¬
low', but each of those species is different in a va¬
riety of characters. Liede and Meve (2004) cite a
duplicate of the holotype of Metastelma brachym¬
ischum, Breedlove 27188 (TEX), under Metastelma
longicoronatum, which is also endemic to Chiapas
at similar elevations, but that species has 3-lobu-
lale rather than entire corona lobes, a rare char-

624

No von

figure 2. Metastelma brachymischum W. I). Stevens. Photograph of braneh and opened flower o (Breedlove 14010 (F).

aeter shared only with Metastelma triodontum (de¬
scribed below) among North and Central American
species ol Metastelma , in addition to other differ¬
ences; perhaps the llreedlove collection is mixed.
The same authors cite a duplicate of the paratype.

Breedlove 14010 (TEX), as Metastelma barbigerum
var. liesnerianum (L. 0. Williams) Liede & Meve,
but that concept is described below as M. thala-
mosiphon.

Metastelma brachymischum has been collected

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

625

only in the vicinity of El Sumidero, in seasonal ev¬
ergreen forest on limestone, at 1200—1350 m ele¬
vation, and flowering in August and October.

Parnlype. MEXICO. Chiapas: 17 km N of Tuxtla Gu¬
tierrez along road to El Sumidero, I). Breedlove 14010 (K).

2. Metastelma thalamosiplion W. I). Stevens, sp.
nov. TV PE: Belize. Western Hwy., 12.5 mi. W
of Belize, 10 Aug. 1970. ./. Wiley 193 (holo-
tvpe. MO). Figure 3.

A Metastehnate liesneriano trichomatibus marginalibus
albis loborum corollae longitudine trichomatibus medianis
translucidis aequantes et lobis coronae basi adnatis cor¬
ollae recedit.

Twining vines, woody and corky below, cork pale
brown, latex white, roots fibrous; young stems pu-
berulent in a broad band, sometimes with a thin
line of triehomes on opposite side, trichomes white,
recurved, 0.2— 0.4 mm long, internodes 0.5—5 cm.
Leaves opposite, without pseudostipules, blades
mostly elliptic, sometimes somewhat ovate or ob-
ovate. 1.9— 3.7 X 0.8— 1.3 cm. apex acute to round¬
ed or emarginate, apiculate, base obtuse to rounded
or rarely subeordate, adaxially sparsely puberulent
on midrib and margin, abaxially glabrous, lateral
nerves 6 to 10 pairs, middle veins 30°— 65° to mid¬
rib, oolleters 0 to 4; petioles 2.2-4 mm, puberulent
on adaxial margin. Inflorescence extra-axillary, 1
per node or sometimes with a subsidiary inflores¬
cence on a short axillary shoot, congested-racemi-
form or more commonly congested-paniculiform, 5-
to 14-flowered, peduncle 0—1.6 mm, puberulent in
I line, axis to 1 mm. pedicel 1—2.2 mm. glabrous
to sparsely puberulent, in 1 line or generally, bracts
0.25—0.5 X 0.2— 0.3 mm, ovate; calyx with 0 to 2
oolleters below each sinus within, lobes ovate with
rounded lips. 0.9-1. 2 X 0.6-0.9 mm. sparsely pu-
berulen! abaxially. especially at base, glabrous
adaxially, green to purple with scarious margins;
corolla campanulate, white or tinted pink outside,
tube 0.8—1 mm long, glabrous, lobes lanceolate
with acute tips, erect or tips somewhat reflexed. 2-
2.3 X 0.8-1 mm. abaxially glabrous, adaxially with
marginal trichomes erect, white, unicellular, 0.3—
0.4 mm long, covering the distal third to three-
quarters of the margin, and with center trichomes
erect or retrorse, translucent, unicellular, 0.2— 0.4
mm long, covering the distal half to the entire cen¬
ter of the lobes; gynostegium with stipe 0.2— 0.4 X
0.6— 0.7 nun, ovoid, corona lobe bases covering the
length of the stipe, corona lobes acicular with
broad, keeled bases adnate to the corolla tube up
to nearly the same height as the attachment to the
stipe, tips acute, erect or somewhat recurved, free

part 0.8—1 X 0.15—0.2 mm. slightly exceeding style
apex, terminal anther appendages ovate, 0.3— 0.4 X
0.2—0.35 mm, erect or somewhat inclined over style
apex, white, anthers quadrate to trapezoidal, 0.3—
0.5 X 0.4—0.65 mm, guide rails straight, salient at
base, 0.25—0.3 mm long; pollinaria pendent, cor-
pusculutn 0.15-0.19 X 0.08-0.1 mm, subsagittate,
pale brown, translators 0. Of)— 0.09 mm. thin, round
in cross section, somewhat geniculate, pollinia
0.19—0.23 X 0.07—0.1 mm. narrowly obovoid, prox-
imallv thin and sterile; style apex 0.5— 0.6 mm
wide, shallowly conical. Follicle 1, fusiform-atten¬
uate. 4.2-6 X 0.2— 0.3 cm, glabrous, seeds ca. 18;
seeds obovate, eucullate, ea. 3.8 X 1.8 mm. red¬
dish brown, margin 0. 1—0.2 mm. irregularly dentate
on distal third, concave side tuberculate, convex
side sparsely tuberculate near margin, coma 1.5—2
cm, tawny.

Liede and Meve (2004) described and provided
an excellent illustration of this new species as Me¬
tastelma barbigerum var. liesnerianum (L. 0. Wil¬
liams) Liede & Meve. Metastelma liesnerianum (L.
0. Williams) Liede. as 1 recognize the taxon, was
originally based upon a single collection from
Guanacaste, Costa Rica, and apparently Liede and
Meve saw only a photograph of the type and no
other Costa Rican material and based their descrip¬
tion and illustration on material from Belize and
southern Mexico. I have now studied seven addi¬
tional collections of Metastelma liesnerianum, all
from Guanacaste, and find that it is amply distinct
from the plants from Belize and southern Mexico.
Notwithstanding the highly unlikely disjunction,
spanning the well-collected countries of Nicaragua.
El Salvador, Honduras, and Guatemala, the char¬
acters distinguishing the two species, or for that
matter almost any species of Metastelma. are not
particularly visible on photographs. The most read¬
ily obvious difference between the two species is
in the corolla lobe pubescence; Metastelma liesner¬
ianum has the white, marginal trichomes of the co¬
rolla 0.05—0. 1 mm long and the central, translucent
trichomes 0.4— 0.5 mm long, distinctly contrasting
in length, while in M. thalamosiphon the marginal
trichomes are 0.3— 0.4 mm long and the medial tri¬
chomes are 0.2— 0.4 mm long, the central trichomes
always equal to or shorter than the marginal tri¬
chomes and more or less concealed by them. Some¬
what less obvious, but even more compelling, the
corona lobes of Metastelma thalamosiphon are
broadly keeled at the base, with the keels adnate
to the corolla tube; the broad bases of the corona
lobes, adnate to about the same level on the stipe
and the corolla tube partition the corolla tube into

626

Novon

I iginc •?. Metaslehna thalamosiphon \\. I). Slovens. Photograph of branches and opened flowers of (Aibrera & ('iibrcra
1 1X60 (MO).

5 chambers (see Liede cK Meve, 2004, p. 44. fig.
4-e ). In Metastelma liesnerianum flic corona lobes
are entirely free from the corolla and the corolla
tube is not chambered.

Metastelma thalamosiphon grows in seasonally
inundated savannas at 0 to 220 m elevation. It has
been most commonly collected in Belize and ad¬
jacent Mexico. Pwelve additional collections cited

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

627

by Liede and Meve (2004) under Metnstelma bar-
bigerum var. liesnerianum, not examined in ibis
study, may be found to pertain to ibis species and
provide the logical addition of Peten. Guatemala,
to the range. The species seems to flower through¬
out the year.

Paratypes. MEXICO. Campeche: 6 km al E de Si I -
virtue sobre la carr. Francisco Escarcega-Chetumal. E.
Cabrera C. & //. de Cabrera 10964 (MO); 8 km al SE de
Conhuas sobre cl camino al Centro Ceremonial de Calak-
mul, E. Cabrera C. & II. de Cabrera 1 1061) (MO); 10 km
al SE de Sabancuy sobre el camino Escdreega— Ciudad del
Carmen, E. Cabrera C. N II. de Cabrera 118 69 (MO); Km
20. al S de la entrada a Calakmul, E. Madrid A'.. E. I.ira.
E. Marline: .S'.. I). Alvarez M. & S. Ramirez A. 531 (MO);
9 km al SE de Dos Naeiones, camino a El Civalito, E.
Marline: S.. I). Alvarez M. & S. Ramirez A. 31430 (MO);
Km 31 de Calakmul. E. Marline: ,S„ IK Alvarez M. & S.
Ramirez A. 31631 (MO). Chiapas: \\ o( Cata/aja on road
to Yillahennosa, I). Breedlove & G. David.se 5530 7 (MO);
Aguacate, E. Matilda 3765 (F). Tabasco: I i kin al NE
de Ciudad del Carmen sobre la carr. a Champoton. Isla
del Carmen, E. Cabvera C. &■ H. de Cabrera 15072 (E.
MO). Quintana Roo: 20 km al NO de fclipe Carillo
Puerto sobre el camino a Vigfa Chico. E. Cabrera C. & ti¬
de Cabrera 1010 (MO). Yucatan: • 1 km al O de Tab! sobre
el camino Yaxcaba-Sotuta, E. Cabrera C. & II. de Cabrera
10737 ( MO). HE l .l/E. Salt Creek Estate. Mi. 18 Northern
llwy., L. I Heckman 153a (MO); Hattieville, ./. Dwyer
10402 (MO), ,/. Diner & R. Pippen 10093 (MO); 12.5 mi.
N\\ of Belize, Northern llwy.. ,/. Dwyer 10706 (F, MO);
Mi. 11, Northern llwy.../. Dwyer 10937 (MO), A. Gentry
7640 (MO); Mi. 19, Northern llwy.,./. Dwyer & R. Lies tier
1201 1 (F, MO); Mi. 5 3/4 Northern «Twy., ./. Dwyer 12771
(MO, NY); Baumgartner's Ridge Lagoon Plantation, Mi. I I
1/2 Northern Ilw\.. ,/. Dwyer 13074 (MO); Hector Creek.
P. Gentle 1503 (MO. NY): 12 mi. \\ of Belize, .S’. McDaniel
13032 (MO); I mi. V ol jet. w / road to Ferguson Bank. 8
mi. E of Hattieville, along Hector Creek Road. P. Sorensen
7031 (F, MO); 1-2 mi. S of Dales Point. G. Proctor 36515
(MO); along New Northern llwy. at jet. w V road to Crooked
free, R. Worthington 24077 (MO).

3. Metastelma thysaiioluiii W. I). Stevens, sp.
nov. TA PE: Ed Salvador. Ahuachapan: San
Benito, en el bautizadero, 850 m, 26 July
1995, M. Sandoval Ac E. Sandoval 96 (holo-
type, MO; isotypes, B, EAP, LAGU). Figure 4.

Species haec Metastelmati sepicola indumento corollae
si m i 1 is. a M. pednnculare forma loborum coronae similis.
all utroque lobis coronae longioribus distincta.

Twining vines, woody and corky below, cork pale
brown, latex white, underground parts unknown:
young stems glabrous or internodes with a thin line
of recurved white trichomes ea. 0.1 mm long, in¬
ternodes 2-1 I cm. Leaves opposite, without pseu¬
dostipules, blades lanceolate to elliptic, 2.1— 6.2 X
0. 6-1.6 cm. apex acute to attenuate, apiculate.
base acute to rounded, adaxially glabrous or
sparsely puberulent on midrib and margin, abaxi-

ally glabrous, lateral nerves 1 I to 15 pairs, middle
veins 40°— 55° to midrib, eolleters 0 to 2; petioles
3.5-10 mm, puberulent on adaxial margin. Inflo¬
rescence extra-axillary, I per node or sometimes
with a subsidiary inflorescence on a short axillary
shoot, congested-raeemiform, 4- to 20-flowered, pe¬
duncle 5-16 mm, glabrous or puberulent in 1 line,
axis to 3 mm, thickened, pedicel 1.5-3. 5 mm, gla¬
brous or puberulent in 1 line, bracts 0.35—0.6 X
ca. 0.3 mm, deltate; calyx with I eolleter below
each sinus within, lobes ovate with rounded tips,
0.6— 0.9 X 0.3— 0.6 mm, sparsely puberulent abax-
iallv. glabrous adaxially, green to purple with sear-
ious margins; corolla campanulate, white to pale
yellowish green, tube 0.7—1 mm long, somewhat
saccate below lobes, glabrous, lobes ovate with
acute lips, erect. 0.8-1. 3 X 0. 4-0.8 mm, with 2
submarginal fleshy ridges, abaxially glabrous, adax¬
ially with a thin, submarginal line of erect, white,
unicellular trichomes 0.05-0.1 mm long, center
and immediate margins of lobes glabrous; gyno-
stegium with stipe 0.5-0. 6 X 0.3-0. 5 mm. narrowly
winged, corona lobes borne at top of stipe, ligulate
with rounded tips, keeled at the base, appressed to
anthers and arching over top of style, tips overlap¬
ping above style apex, free part 0.5— 0.7 X 0. 1—0.2
mm, terminal anther appendages nearly circular,
0.15—0.25 mm diam., appressed to margin of style
apex, white, anthers rectangular, 0.18—0.25 X
0.35—0.55 mm, guide rails rounded, widest at cen¬
ter. 0.13—0.25 mm long; pollinaria pendent, eor-
puscnlum 0.07—0.13 X 0.04—0.05 mm, ellipsoid,
pale brown, translators 0.05-0.08 mm, thin, round
in cross section, strongly sigmoid, pollinia 0.12—
0.15 X 0.05—0.08 mm, narrowly obovoid, proxi-
mally thin and sterile; style apex 0.5— 0.7 mm wide,
slighllv umbonate in center. Follicle I. lusiform-
attenuate, ca. 4.7 X 0.5 cm, glabrous; seeds un¬
known.

I his new species was treated as " Metastelma sp.
B” in Flora de Nicaragua (Stevens et al.. 2001).
Liede and Meve (2004) placed Metastelma sepicola
Pittier, endemic to Costa Rica, into the synonymy
of M. pednnculare Decaisne, endemic to Guatema¬
la, stating that the two “show no constant differing
characters.” They also studied one of the paratypes
of Metastelma thysanotum. Molina 23080 , and
while noting some differences, decided that more
material was needed to make a decision. As rec¬
ognized here, Metastelma sepicola consistently dif¬
fers from M. pednnculare in having the inner (ace
of the corolla lobes with a thin line of submarginal,
erect, white trichomes (“verrucose trichomes of
Liede & Meve. 2004, p. 34) while the center of the

628

Novon

Figure I.
(MO).

Melastelma thysanotum W. I). Stevens. Photograph of branch anti opened flower of Sandoval & Rnmdn 1365

lobes are glabrous and M. pedunculare has no sub-
marginal line of triehomes, but the center of the
lobes are covered with retrorse. nearly translucent
triehomes (“smooth, hispid, downwardly directed
triehomes” of l.iede & Meve. 2004, p. 34). Fur¬
thermore, the corona lobes ol Metastelnia sepicola

are markedly spathulate, much broader at the tip.
while those ol M. pedunculare are ligulate or even
slightly narrowed tit the tip. MeUistelma thysanotum
ha s exactly the corolla indumentum of M. sepicola,
but the corona lobes are ligulate, as in M. pedun-
eulare. However, both Metastelma sepicola and M.

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

629

pedunculare have corona lobes that are shorter than
to nearly equaling the level of the style apex, while
in M. thysanotum the corona lobes are longer than,
loosely appressed to. and more or less reaching the
opposite side of the top of the style apex, crossing
at the center. Somewhat less obviously, there are
also consistent size differences: Metastelma thysan¬
otum has corolla lobes 0.8-1. 3 mm long and gv-
nostegium stipes 0.5— 0.6 mm long, M. pedunculare
lias corolla lobes 1.7-2. 1 mm long and stipes 0.9-
1.1 mm long, and M. sepicola has corolla lobes 1.5—
1.8 mm long and stipes 0.7— 1.1 mm long. All three
of these species, as well as Metastelma stenomeres
(Standlev & Steyermark) W. I). Stevens, have inflo¬
rescences that tend to flower longer than most spe¬
cies of Metastelma, and can produce thickened
axes covered with pedicel scars; Molina 23080. the
one specimen of Metastelma thysanotum examined
by Liede and Meve (2004), is not unusual in this
regard, but does have numerous inflorescences ag¬
gregated on short, leafless shoots, and perhaps is
diseased.

Metastelma thysanotum is found on volcanic
soils from southwestern Guatemala throng h El Sal¬
vador to north-central Nicaragua, and will probably
be discovered in southern Honduras. It has been
found between 350 and 1600 m elevation and prob¬
ably flowers year-round, although it has been most
frequently collected with flowers between July and
September.

Paratypes. GUATEMALA. Chimaltenango: Alame¬
da. Johnston 1022 (F). Saoatepequez: 4 km NW of
San Miguel Duenas on road to Acatenango, M. Nee, D.
Atha & ./. ,/. Castillo M. 472 II (MO, NY). Solola: Santo
Tomas, J. Cuj 7077 (MO, UVAL). EL SALVADOR. Ahu-
acliapau: along road between San Francisco Men^ndez
and Tacuba, T. Croat 42148 (MO), 42170 (MO); San Fran¬
cisco Menendez, FI Corozo, ,/. Rosales 1195 (B, BM,
FAGU, MO): San Benito, E. Sandoval & F. Chinchilla 481
(B, LAGU, MO). E. Sandoval & A. Roman 1365 (B,
LAGll. MO), E. Sandoval 1689 (B. LAP, FAGU, MO):
PN. El Imposible, A. Sermeho 1167 (FAGU, MO), R. To¬
ledo 23 (B. FAGU, MO), R. Villacorta, R. Toledo & R.
Ldpez 878 (15. FAGU, MO). Santa Ana: FI Pi rial ito. E.
Montalvo. J. Gonzalez & R. Villacorta 6328 (FAGU, MO).
NICARAGUA. Esteli: Estanzuela Creek, 8 km W of Fs-
lelf, A. Molina R. 23080 (F, MO. NY). Matagalpa: 3 km
af SE de Fsquipulas. P. \ 1 arena 25352 (MO).

4. Metastelma triodontiim W. I). Stevens, sp.
nov. TYPE: Guatemala. Huehuetenango: above
La Libertad, Cerro Pueblo Viejo, 1900 m, 20
Aug. 1942, ./. Steyermark 50999 (holotype,
MO; isotype, F). Figure 5.

Speciebus corona trilobata cum Metastelmate longico-
ronato congruens, sed differt floribus minoribus indumen-
to eorollae alis antherae curvatis.

Twining vines, woody anti corky below, cork
brown, latex unknown, underground parts un¬
known; young stems puberulent on I side with re¬
curved white Irichomes 0.15—0.2 mm long, inter¬
nodes 1-3 cm. Leaves opposite, without
pseudostipules, blades narrowly elliptic. 2—2.5 X
0.4— 0.8 cm, apex acute, apiculate, base acute,
adaxially glabrous except puberulent on midrib and
margin, abaxially glabrous or with a few Irichomes
along midrib, lateral nerves 1 to 6 pairs, middle
veins 50°— 60° to midrib, colleters 0 to 2; petioles
1 .5— 2.5 mm. puberulent on adaxial margin. Inflo¬
rescence extra-axillary, I per node, congested-ra-
cemiform, 4- to 6-flowered, peduncle 1 .5-3.5 mm,
puberulent on 1 side, pedicel 1—2 mm, puberulent
on I side, bracts 0.4-0. 7 X 0. 3-0.5 nun. ovate;
calyx with I colleter below each sinus within, lobes
ovate with rounder! tips, 0.6— 0.7 X 0.35—0.4 mm.
puberulent at base and margin abaxially. adaxially
glabrous, purple with scarious margins; corolla
campanulate, white, tube 0.5— 0.6 mm long, abaxi¬
ally glabrous, adaxially with trichomes of lobes con¬
tinuing into top of tube, lobes lanceolate with acute
tips, patent, 1-1.1 X 0. 5-0.6 nun. abaxially gla¬
brous. adaxially with broad margins and distal half
covered with erect, white, unicellular trichomes,
0.1-0.15 mm long, center trichomes restricted to a
narrow strip in basal half, dense, retrorse, translu¬
cent, unicellular, 0.1-0.15 mm long; gynostegium
with stipe 0.15-0.2 X 0.4-0. 5 mm. broadly winged,
corona lobes borne at top of stipe. 3-lobed, laminar,
free part ca. 0.65 X 0.4 mm. central lobule nar¬
rowly debate, bent sharply over style apex, lateral
lobules small, borne near middle, debate, terminal
anther appendages ovate, ca. 0.2 X 0.2 mm, ap¬
pressed to style apex, white, anthers rectangular in
outline, ca. 0.3 X 0.4 nun. guide rails nearly ver¬
tical at top, curving outward toward base. ca. 0.3
mm long; pollinaria pendent, corpusculum ca. 0.14
X 0.05 mm, subsagittate. brown, translators ca.
0.04 mm. round, bent at middle, pollinia ca. 0.16
X 0.05 mm, asymmetrically ellipsoid; style apex
ca. 0.5 mm wide, nearly flat with small umbo, fol¬
licles and seeds unknown.

Among the Mesoamerican species of Metastelma,
only this new species and Metastelma longicoron-
atum (L. 0. Williams) Liede have 3-lobulate corona
lobes. The flowers of Metastelma longicoronatum
are larger overall (2.4— 2.9 mm vs. 1.5— 1.7 mm
long) and most notably the corona lobes are longer
(1.4— 1.9 mm vs. 0.65 mm), have their lateral lob¬
ules at the base rather than the middle and curving
upward rather than straight, and have their central
lobules erect and only slightly arching over the

630

Novon

2

figure 5. Metastelma triodontum W. I). Stevens. Photograph of branch and opened flower of Steyermark 50999 (iso-
lype, F).

style apex rather than bent sharply over the style
apex. Metastelma longicoronatum also has longer
inflorescences, leaves with more secondary veins,
and guide rails that are straight and at about 45°
lo flic gynostegiuni axis rather than nearly vertical

at ihc top and gradually flaring outward. A few oili¬
er species have (lowers in the si/e range of Metas¬
telma triodontum , but all have entire corona lobes
and other differences. The indumentum of the co¬
rolla lobes, with the white, marginal trichomes cov-

Volume 15, Number 4
2005

Stevens

Novelties in Cynanchum

631

ering most of the lobes and the translucent, central
trichomes restricted to a narrow line in the basal
half, seems especially distinct.

Metastelma triodontum is known only from the
type collection from Guatemala.

5. Metastelma yucatanense W. I). Stevens, sp.
nov. T\ PE: Mexico. Quintana Roo: directly \\
of Puerto Morelos, 5 m, 4 May 1982, G. Dav-
idse. M. Sousa S., A. (dialer & E. Cabrera C.
20050 (holotype, MO; isotype, MEXLJ). fig¬
ure 6.

Species haec a Metastelmate palmeri differt stipitc brev-
iore ct a corona occulta al is antherae rectis ct hasi non
curvatis.

Twining vines, herbaceous, without cork, latex
white, underground parts unknown: young stems
glabrous or internodes with a thin line of recurved
white trichomes 0. 1—0.3 mm long, internodes 2-5
cm. Leaves opposite, without pseudostipules,
blades elliptic to ovate, 1.1— 2.8 X 0.4— 0.9 cm.
apex acuminate to rounded, apiculate, base obtuse
to rounded, adaxially glabrous except inconspicu¬
ously puberulent on midrib and margin, abaxially
glabrous, lateral nerves 5 to 8 pairs, more or less
inconspicuous, middle veins 30°— 55° to midrib,
eolleters 0 to 2: petioles 1.5—5 mm. puberulent on
adaxial margin. Inflorescence extra-axil I ary, I per
node or sometimes with a subsidiary inflorescence
on a short axillary shoot, congested-racemiform. 7-
to 16-flowered, peduncle 1—4.5 mm, glabrous or
puberulent in 1 line, axis to 2.5 mm, pedicel 2.5—
4.5 mm, bracts 0.2— 0.7 X 0.2— 0.4 mm, ovate; calyx
with 0 to 3 eolleters below each sinus within, lobes
ovate with rounded tips. 0.6— 1.1 X 0.1— 0.7 mm.
sparsely puberulent at base abaxially anil some¬
times on margin, adaxially glabrous, green to pur¬
ple with scarious margins; corolla shallowly cam-
panulate, white to cream, tube 0.4— 0.6 mm long,
glabrous or with adaxial marginal trichomes of
lobes extending into top of tube, lobes lanceolate
with rounded tips, spreading, 1.4— 2.3 X 0.6— 0.9
mm, with 2 submarginal fleshy ridges, abaxially
glabrous, adaxially with margins covered with erect,
white, unicellular trichomes, 0. 1-0.3 mm long dis-
tally, becoming 0.05—0.1 mm long proximally, cen¬
ter trichomes absent or, when present, retrorse, yel¬
lowish translucent, unicellular, 0.1 -0.2 mm long;
gynostegium with bulbous stipe 0.2— 0.4 X 0.5— 0.7
nun. broadly winged with bases of corona lobes that
cover the entire stipe, corona lobes lanceolate with
attenuate tips, keeled at the base, erect or some¬
what sigmoid, free part 0.5— 1.1 X 0.2— 0.3 mm, ter¬
minal anther appendages nearly circular, 0.25—0.35

mm diam., appressed to margin of style apex, white,
anthers trapezoidal. 0.4— 0.5 X 0.45—0.55 mm,
guide rails straight, ± tangential to anther body,
radially salient at top and somewhat inclined to¬
ward the axis below, 0.35—0.55 mm long; pollinaria
pendent, corpusculum 0.09—0.17 X 0.06-0.1 1 mm.
ovoid to ellipsoid, brown, translators 0.05—0.09
mm, thin but dilated distally, round in cross sec¬
tion, sigmoid to geniculate, pollinia 0.18—0.23 X
0.06—0.1 mm, ellipsoid or somewhat ovoid or ob-
ovoid; style apex 0.55—0.7 mm wide, convex. Fol¬
licle 1. fusiform-attenuate, 4.4— 4.8 X 0.15—0.2 cm.
glabrous, seeds ea. 14; seeds obovate, nearly ellip¬
tical. flat or cucullate, 3.8-4 X 1.4— 1.6 nun. red¬
dish brown, margin 0.1— 0.2 mm, inconspicuous,
finely and inconspicuously undulate, concave side
tuberculate, convex side verrucate near margin,
coma 2—3 cm, tawny.

Metastelma yucatanense shares the general shape
of the corona with M. palmeri S. Watson, but that
species, which in my concept ranges from Texas to
Puebla and perhaps Oaxaca, at 400—2300 m ele¬
vation. differs in a number of characters. The co¬
rolla lobes of Metastelma palmeri are generally cov¬
ered with uniformly short white trichomes (0.02—
0.1 mm) and the translucent trichomes are
restricted to a basal triangle, while the white tri¬
chomes of M. yucatanense form a discrete marginal
band and are longer at the tips of the lobes (0.1 —
0.3 mm) and become gradually shorter toward the
base of the lobes; M. palmeri has a distinct stipe
with the corona lobes attached at the top, while this
new' species has a somewhat shorter, and bulbous,
stipe that is completely covered by the broadly
keeled bases of the corona lobes; M. palmeri has
the guide rails more vertically and more radially
oriented with a prominent radial projection at the
base, while the new species has guide rails that are
somewhat inclined under the anther body, more
tangentially oriented, and are straight, with no bas¬
al projection. Metastelma turneri Liede ix Move
shares the pattern of corolla indumentum with this
newT species, but is restricted to northeastern Mex¬
ico, is more robust and pubescent, and has shorter
guide rails (0.2—0.25 mm) that tilt away from the
anther body, among other differences. The follicles
of this new species are distinctively thin and the
seeds are small for the genus. The single Guate¬
malan collection ( Contreras 9336) is the only one
cited by Liede and Meve (2004), and they tenta¬
tively assign it to Metastelma palmeri. While it
clearly is not Metastelma palmeri , this specimen
represents the low extremes of most measurements

632

Novon

figure (>. Metastelma yucataneiise W. D. Stevens. Photograph of branches and opened flowers of Cabrera & Cabrera
(>M7 (MO).

of M. yucataneiise and is somewhat disjunct from
the other collections.

Metastelma yucataneiise is primarily a coastal
species hut has been found somewhat inland, up to

about 65 m elevation, and apparently flowers year-
round.

Paratypes. MEXICO. (Juintumi Roo: 10 km al N de
Puerto Morelos, camino a Punta Caraeol. E. Cabrera C. &

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Novelties in Cynanchum

633

II. de Cabrera 3139 (MO); 15 km ill N de I’unla Sam. F.
Cabrera C. & H. de Cabrera 6 487 (MO); camino a Nizuc,
l\ Moreno 10 1 (MKXU); Punta Sam. 5 km al i\ <le Puerto
Juarez. II. Quero. II. Flores & R. (Wether 2331 (KNCB.
MO). 0. Tellez V. & F. Cabrera C. 1545 (MO); I s I a llolhox,
V. Rico-Gray & I. Fspejel 131 (F); Puerto Morelos. F. Mo¬
reno 312 (MFXU), M. Sousa S.. <). Tellez V. X F. Cabrera
C. 10793 (MO). Yucatan: 10 km al N de Telchak Pueblo,
earr. Motul-Telebak Puerto, E. Cabrera C. & II. de Ca¬
brera 9674 (MO); above Calcelitok, S. Darwin 231 / (MO);
8.5 kin al N de Dzemul camino a Xtampu, R. Ihirdn &
M. Mendez 2369 (MO). GUATEMALA. Izalml: Cadenas
(Puerto Mendez). Sarstun River, F. Contreras 9336 (MO).

Orthosia Decaisne in A. DC.. Prodr. 8: 526. 1844.
TYPE: Orthosia congesta (Vellozo) Decaisne
(lectotype, designated by A. A. Bullock, Kew
Bull. 13: 98. 1958).

Eiede (1997) first pointed out that some North
American species of Cynanchum, sensu Woodson,
belonged to the genus Orthosia, previously only
known from southern South America. In her anal¬
ysis of Orthosia and the related genera, the inflo¬
rescences were described as axillary or extra-axil¬
lary, simple and sciadioidal. While the
inflorescence description is basically correct in a
descriptive sense, it is instructive to understand
their development. Typical inflorescences of Ascle-
piadoideae are either terminal or extra-axillary, ap¬
parently developmental!) terminal, but pushed
aside by the emerging shoot, and one per node.
Many species of Orthosia have these apparently ax¬
illary inflorescences, two per node, that are eon-
gested-racemiform or congested-panieuliform. Oth¬
er species, clearly related in other regards, have
both axillary and extra-axillary inflorescences or
even exclusively extra-axillary inflorescences. Or¬
thosia glaberrima, validated below, is especially in¬
formative in understanding the origin of this inflo¬
rescence type, having axillary, extra-axillary, and
even terminal inflorescences, sometimes more than
one kind at a node, and most unusual, the appar¬
ently axillary inflorescences often revert to being
vegetative shoots, leaving a cluster of pedicel scars
at the base of lateral shoots. Clearly, the axillary
inflorescences of Orthosia, as is the case with the
related Tassadia (Stevens, 2002), are actually con¬
densed axillary branches with one, then racemi-
form, or more, then paniculiform, terminal or extra-
axillary true inflorescences. Duly Mesoamerican
species of Orthosia are considered here; five spe¬
cies are described as new' and three new combi¬
nations are provided.

I. Orthosia cynanchoides W. 1). Stevens, sp.
nov. T\ PE: Mexico. Chiapas: Rodeo, near Si 1 -
tepee, 2800 m, I Aug. 1941, E. Matilda 4562
(holotype. MO; isotypes, F, MEXU). Figure 7.

Inter species Orthosiae infloreseentiis solitariis et axil-
lari I »us et extra-axillaribus tubo corollae brevi tubo coro-
nae brevi lobis coronae brevibus areuatis gynostegio cy-
lindrico brevistipitato poliniis ellipsoideis dimulio minor
quam corpusculis dilfert.

Twining vines, woody and corky below, latex and
underground parts unknown; young stems puberu-
lent in a broad band, trichomes erect, 0.2—0.25 mm
long, white, internodes 2—13 cm. Leaves opposite,
persistent, without pseudostipules, blades lanceo¬
late. 2. 1 -3.2 X 0.4— 0.6 cm, apex acute to attenu¬
ate, apiculate or not, base obtuse to rounded, adax-
ially puberulent on midrib and margin, sparsely
puberulent on basal half of surface, abaxially gla¬
brous or with a few trichomes along midrib, lateral
nerves 3 to 4 pairs, inconspicuous, middle veins
55°— 60° to midrib, colleters 2: petiole 2—3 mm,
densely puberulent along adaxial margin. Inflores¬
cence axillary or extra-axillary, 1 per node, con-
gested-racemiform, nearly umbelliform, 2 to 4 (low ¬
ers open at a time, peduncle 0—2.5 mm. puberulent
on 1 side, pedicel 3—3.5 mm, puberulent except 1
line glabrous, bracts lanceolate, 0.5— 0.7 X 0.1— 0.2
mm. purple; calyx without colleters, lobes elliptic
with rounded tips, 0.7— 0.8 X 0.5— 0.6 mm. ap-
pressed -puberulent outside, glabrous inside, erect
to ascending, purple; corolla shallowly campanu-
late, nearly rotate, glabrous, white, tube 0.5— 0.6
mm long, lobes elliptic with rounded tips. 4—4.2 X
2—2.2 mm, spreading; gynostegium c\ lindrieal. vv ith
stipe ea. 0.3 mm, corona of 5 lobes connate at base,
borne at base of the stipe, tube ca. 0.1 mm long,
lobes lanceolate, somewhat shouldered Iowan I the
rounded tip, ca. 0.8 X 0.45 mm. center of lobe
adnate to stipe, then arching outward toward corolla
tube and then back toward gynostegium, with the
tips touching the center of the anther backs, ter¬
minal anther appendages ovate, ca. 0.3 X 0.3 mm.
arching over and nearly covering style apex; an¬
thers rectangular, ca. 0.7 X 0.5 mm. guide rails
vertical, slightly bowed outward at center, ca. 0.7
mm long; pollinaria pendent, corpusculum ellip¬
soid, ca. 0.21 X 0.09 mm, brown, translators ca.
0.06 mm, thin, round in cross section, strongly sig¬
moid. pollinia ellipsoid, ca. 0.13 X 0.08 mm; style
apex flat, ca. 0.6 mm wide. Immature follicles 2.
divergent 180°, fusiform-attenuate, to 41 X 2 mm,
smooth, sparsely puberulent, mature follicles and
seeds unknown.

fhe single collection of this species, from near
Siltepec and probably from cloud forest, was dis¬
tributed as Metastelma angustifolium Turczaninow.
a species of Orthosia from north and central Mexico
that also passes under the name Cynanchum kun-

634

Novon

J

llm (Decaisne) Standley. Orthosia cynaruhoides is
superficially similar to this species, but the latter
differs iti many details, including having a conical
gynostegium and corona lobes larger than, and

completely covering, the anthers. Among the spe¬
cies of Ortliosia with both apparently axillary and
extra-axillary inflorescences, 0. cynotulwidcs
stands out in having both the corolla and the corona

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Stevens

Novelties in Cynanchum

635

divided nearly to the base and the corona lobes
relatively short and strongly curved.

2. Orthosia extra-axillaris W. I). Stevens, sp.
nov. TYPE: ( iosta Rica. San Jose: Cordillera
de Talamanca, W ridge of Cerros Cuericf,
3160 m, 9°34'N, 83°40'W, 15 Sep. 1983, G.
Davidse 24730 (holotype, MO: isotype, CR).
Figure 8.

Inter species Orlhosiae inflorescentiis solitariis extra-
axillaribus tuho ooronae longo campanulato lobis coronae
latis gynostegio cylindrico brevistipitato poliniis asym-
metricis corpusculum fere aequantibus differt.

Twining herbaceous vines, latex white, under¬
ground parts unknown; young stems puberulent in
a narrow to broad band, triehomes erect to some-
v\ 1 i a t reflexed, 0. 2-0.4 mm long, white, internodes
1—13 cm. Leaves opposite, persistent, without pseu¬
dostipules. blades lanceolate, 2.5-4 X 0.5-1 cm,
apex attenuate, apiculate or not, base acute, adax-
ially puberulent on midrib and margin, abaxially
glabrous or puberulent on midrib, lateral nerves 4
to 6 pairs, middle veins 40°— 50° to midrib, colleters
2; petiole 2-6 mm, puberulent along adaxial mar¬
gin. Inflorescence extra-axillary, I per node, con-
gested-racemiform or congested-paniculiform, 2 to
9 flowers open at a time, peduncle 0—4 mm, pu¬
berulent on 1 side, pedicel 2.5— 5.5 mm, puberulent
in 1 line, bracts lanceolate, 0.7-1. 2 X 0.4— 0.6 mm,
green or purple; calyx with 0 to 2 colleters below
each sinus within, lobes ovate with acute to round¬
ed tips, 1.1— 1.8 X 0.7— 0.8 mm, with a few tri¬
ehomes at base and margin outside, glabrous in¬
side. ascending, green or purple; corolla shallowly
campanulate, nearly rotate, glabrous, white or out¬
side tinted with yellow, green, pink, or purple, tube
0.8—1 mm long, lobes ovate with rounded tips, 1.9—
2.5 X 1.2-1. 7 mm, ascending; gynostegium cylin¬
drical, with stipe 0.1— 0.3 mm, corona of 5 lobes
connate at base, borne at the base of the stipe, tube
0.3— 0.5 mm long, campanulate, lobes lanceolate,
somewhat to strongly shouldered toward the round¬
ed tip. 1-1.8 X 0.8-1. 2 mm, erect, somewhat short¬
er to somewhat longer than the gynostegium, ter¬
minal anther appendages ovate, 0.35—0.5 X 0.25—
0.5 mm, arching over and nearly covering style
apex; anthers rectangular, 0.65—0.8 X 0.5— 0.7 mm,
guide rails vertical, straight. 0.65—0.8 mm long;
poll inaria pendent, corpusculum ellipsoid to sub-
sagittate, 0.19—0.25 X 0.08—0.13 mm, red-brown,
translators 0.03—0.06 mm, thin, dilated at tip or
not, round in cross section, straight to geniculate,
pollinia ellipsoid, reniform or angularly obovoid,
0.14-0.18 X 0.09-0.11 mm; style apex convex.

0.6—1 mm wide. Immature follicles 2, divergent
180°, fusiform-attenuate, to 38 X 3 mm, smooth,
sparsely puberulent at base, mature follicles and
seeds unknown.

This species shares the general appearance of
Orthosia cynanchoides, especially in the narrow,
relatively coriaceous, persistent leaves and rela¬
tively large, nearly rotate corollas. However, Ortho¬
sia extra-axillaris, compared to O. cynanchoides,
has corolla lobes about twice as long as the tube,
rather than about eight times longer, and a corona
with a longer, campanulate tube free from the stipe
and larger lobes. The shapes of the corona lobes
and of the pollinia are unusually variable hut many
other characters hold this small group of specimens
together. The inflorescences are always of the typ¬
ical extra-axillary sort and one per node, apparently
the only Orthosia from North America without at
least some axillary inflorescences. This Costa Rican
endemic has been found in cloud forests between
2300 and 3160 m elevation and has been collected
with flowers between June and October.

Paratypes. COSTA RICA. Curtugo: 14 km SI*, of El
Empalme. F. Almeda, It. Wilbur & T. Daniel 3101 (MO);
Heredia: I’.N. Braulio Carillo, Sendero Cerro Cuararf, G.
Rivera 107 (MO); I’.N. Braulio Carillo. Estaeidn liana. G.
Varela 103 (MO). San Jose: Estac ion Cuericf, sendero a
La Piedra, 5 km E de Villa Mills, R. Gamboa It. 753 (MO).

3. Orthosia glaberrima (Woodson) \\. I). Stevens,
comb. nov. Basionym: Met-astelma glaberri-
mum Woodson, in Woodson & Seibert, Ann.
Missouri Bot. Card. 24: 200. 1937. TYPE:
Panama. Chiriqui: valley of Rio Chiriqui Viejo,
near Monte Eirio, 5500 ft.. II July 1935. R.
Seibert 300 (holotype, MO).

This species was treated as Cynanchum sepium
(Decaisne) Standley in the Flora oj Guatemala
(Standley & Williams, 1969) and Cynanchum gla-
berrimum (Woodson) E. (). Williams in the Flora of
Panama (Spellman. 1975). The species of Orthosia
sometimes known as Cynanchum sepium is actually
rare and endemic to a small area in the mountains
east of Oaxaca, Mexico, in the district of Ixtlan.
while O. glaberrima is common in the mountains
from Chiapas to the Panama-Costa Rica border.
Metastelma pallidum Rushy from Colombia is ob¬
viously related, and would provide an older epithet,
but has a dense, yellowish indumentum not seen in
the Mesoamerican collections, and is not consid¬
ered synonymous.

636

Novon

Figure 8. Orthosia extra-axillaris W . I). Stevens. Photograph of branches and opened flowers of Almeda el al. 3101
(MO).

4. Orthosia lnisera (I.. 0. Williams) W. I). Ste- samald, 3800 ft., July 1887,//. Von Tuerckheim

vens, comb. nov. Basionym: Cynanchum mis- 1290 (holotype, US; isotypes, K (ill).

erum U. (). Williams, Fieldiana, Bot. 32: 38.

1968. TYPE; Guatemala. Alla Verapaz: Pan-

This species was treated as Cynanchum miserum

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Novelties in Cynanchum

637

in the Flora of Guatemala (Standley & Williams,
1969). It is most similar to Orthosia ramosa , de¬
scribed and compared below. Orthosia misera is
among the species of Orthosia completely leafless
at maturity. It is found in cloud forests from Chia¬
pas to Honduras.

5. Orthosia ramosa W. I). Stevens, sp. nov.
TYPE: Honduras. Ocotepeque: vie. of El Por¬
tillo, Cordillera de Merendon, 2000 m, 31
Aug. 1975, A. Molina R. & A. Molina 30910
(holotype, MO: isotypes. FA P. F). Figure 9.

Inter species Orthosiae inflorescentiis solitariis vel gem-
inatis axillarihus tuho corollae longo tubo coronae brevi
lobis coronae brevibus arcuatis gynostegio brevistipitato
dorsis antherarum vesiculatis poliniis anguste obovoideis
longis differ!.

Twining or scandent vines, woody and corky be¬
low, latex white, underground parts unknown:
young stems puberulent in an inconspicuous line,
trichomes creed to reflexed, ca. 0.1 mm long, white,
internodes 2—1 1 cm. Leaves opposite, caducous,
without pseudostipules, when present on young
shoots, blades linear to narrowly lanceolate, 1.1—3
X 0.1— 0.3 cm, apex acute to attenuate, base obtuse
to rounded, adaxially inconspicuously puberulent
on midrib and margin, abaxially glabrous, lateral
nerves ca. 15 pairs, inconspicuous, middle veins
ca. 60° to midrib, colleters 2; petiole 1—3 mm, pu¬
berulent along adaxial margin. Inflorescence axil¬
lary, 1 or 2 per node, congested-racemiform, 2 to 4
flowers open at a time, peduncle 0—2 mm, puber¬
ulent on 1 side, pedicel 2—3.5 mm, glabrous or with
a few scattered trichomes, bracts ovate, ca. 0.2 X
0.2 mm, green; calyx with 0 to I colleters below
each sinus within, lobes ovate with rounded tips,
0.6— 0.7 X 0.5—0.55 mm, with a few trichomes at
base outside, glabrous inside, erect to ascending,
green; corolla campanulate, nearly tubular, gla¬
brous, white, tube 1.2— 1.6 mm long, lobes ovate
with rounded lips, 0.5— 0.6 X 0.5— 0.6 mm. erect;
gynostegium with anther backs inflated, broadest in
the center and tapering toward tip and base, with
stipe 0.45—0.5 mm, narrowed in middle and 5-
ribbed, corona of 5 lobes connate at base, borne at
the base of the stipe, tube ca. 0.05 mm long, lobes
debate with rounded tips. 0.25—0.3 X 0.15—0.3
mm, arching outward toward corolla tube and then
back toward gynostegium. with the tips touching
the center of the stipe, terminal anther appendages
ovate, 0.1— 0.2 X 0.1— 0.2 mm, arching over style
apex; anthers rectangular in outline, 0.25 X 0.4-
0.45 mm. guide rails somewhat tilted in at the top,
straight, ca. 0.25 mm long; pollinaria pendent, cor-
pusculum subsagittate, 0.10—0.11 X 0.05—0.06

mm, pale brown, translators 0.06—0.09 mm, thin,
round in cross section, geniculate, pollinia narrowly
obovoid. 0.13—0.14 X 0.05—0.06 mm; style flat or
slightly convex, 0.4— 0.5 mm wide. Follicles and
seeds unknown.

This new species corresponds in superficial ap¬
pearance to Orthosia misera, and both share polli¬
naria with long, geniculate translators and narrowly
obovoid pollinia that are longer than the corpus-
cula. However, Orthosia misera has a more open
campanulate corolla, a shorter stipe (ca. 0.3 mm),
and straight, erect corona lobes that are as large
as, and completely cover, the anthers. Orthosia ra¬
mosa is found in pine-oak and cloud lorests in Hon¬
duras. between 1600 and 2200 m elevation, and is
known to flower in May and August.

Paratypes. HONDURAS. Cortes: trail from Ccrro
Candles to Cerro Jilinco, T. Hawkins 971 (EAP. MO).
Lempira: Montana de Celaque, SE pari of massif, trail
from Cracias to top of mtn, G. Davidse & R. Zuniga 34697
(EAP. MO).

6. Orthosia rubens (L. 0. Williams) \\. I). Ste¬

vens. comb. nov. Basionym: Cynanchum rub¬
ens L. 0. Williams, Fieldiana, Hot. 32: 39.
1968. TYPE: Guatemala. Chimaltenango: San¬
ta Elena, 2400—2700 m, 27 July 1933, A.
Skutch 503 (holotype, US; isotype [fragment of
holotype], E).

Phis species was treated as Cynanchum rubens
in the Flora oj Guatemala (Standley & Williams,
1969) and is endemic to Guatemalan cloud forests.
It is the only Mesoamerican Orthosia with more
than a hint of red on the corolla.

7. Orthosia smaragdina W. D. Stevens, sp. nov.

TYPE: Guatemala. San Marcos: Einca Arme¬
nia, San Rafael Pie de la Cuesta to Carrizal,
past Finca Africa, 1300—1600 m, 9 to 12 Aug.
1980, J. Dwyer 15327 (holotype, MO). Figure
10.

Inter species Orthosiae inflorescentiis solitariis vel gem-
inatis axillarihus tubo corollae longo tuho coronae longo
partim obtengenti gynostegium lobis coronae longis gy¬
nostegio eylindrico poliniis ellipsoideis minor quam cor-
pusculis differ!.

Twining vines, woody and corky below, latex
and underground parts unknown: young stems
glabrous or with a few trichomes at the nodes,
trichomes erect, ca. 0.1 mm long, white, inter¬
nodes 2—5 cm. Leaves opposite, persistent, with¬
out pseudostipules, blades lanceolate, 4—6 X
1.2-1. 7 cm, apex acute, apiculate, base obtuse
to rounded, adaxially puberulent on midrib and

638

Novon

h igure 9. Orthosia ramosa Vi. I). Stevens. Photograph of branches and opened flowers of Molina & Molina .10910
(isotype, F).

margin, ahaxially puberulent on base of midrib,
lateral nerves 7 to 9 pairs, middle veins ca. 55°
to midrib, colleters 2; petiole 5—6 mm, sparsely
puberulent along adaxial margin. Inflorescence

axillary, I or 2 per node, eongested-racemiform,
sessile. 3 flowers open at a time, pedicel 3.5— 4.2
mm, sparsely puberulent, bracts ovate, 0.3— 0.5
X 0.2-0. 3 mm, green; calyx without colleters.

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Novelties in Cynanchum

639

Figure 10. Orthosia smaragdina W. I). Stevens. Photograph of branches and opened flower of Dwyer 15327 (holotype,
MO).

lobes ovate with rounded tips. ca. 0.7 X 0.7 mm.
appressed-puberulent outside, glabrous inside,
erect, green; corolla campanulate, 5-fluted, gla¬
brous, green, tube ca. 1.4 mm long, lobes elliptic
with rounded tips, ca. 2 X 0.8 mm. erect; gynos-
tegium cylindrical, sessile, corona of 5 lobes con¬

nate at base, borne at the base of the gynoste-
gium, tube ca. 0.5 mm long, clasping around the
lower third of the gynostegium, lobes narrowly
debate with acute or slightly bifid tips, ca. 1.5 X
0.7 mm. erect, exceeding the gynostegium. ter¬
minal anther appendages semicircular, ca. 0.3 X

640

Novon

1 cm

hgurr 11 Orthosia stipilatu W. I). Stevens. Photograph of branches and opened flower of Hampshire el al 466
(holotype, MO).

0.3 mm, arching over style apex; anthers rect¬
angular or slightly trapezoidal in outline, ca. 1.2
X 0.o5 mm, guide rails vertical, bowed inward,
narrowest a little above the middle and slightly
flared at base, ca. 1.1 mm long; pollinaria pen¬

dent, corpusculum ellipsoid, ca. 0.16 X 0. 1 I
mm. pale brown, translators ca. 0.05 mm, thin,
round in cross section, straight, pollinia ellip¬
soid, ca. 0. 11 X 0.09 mm; style flat. ca. 0.6 mm
wide. Follicles and seeds unknown.

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Novelties in Cynanchum

641

This distinctive species, known only from the
type, has relatively large leaves and (lowers, de¬
scribed by the collector as emerald-green. Ibis is
the only Mesoamerican species of Orthosia with a
tubular corona tightly clasping around the lower
part of the cylindrical gynostegium; pollinators
must probe the narrow gap between tbe gynoste¬
gium and the corona tube to access the opening at
the base of the guide rails. 1 he translators are short
and straight and the pollinia are symmetrical and
smaller than the corpuscula.

8. Orthosia stipitata \\. IT Stevens, sp. nov.

TYPE: Mexico. Chiapas: El Triunfo Reserve,
2100 m. 15°39'N, 92°50'W, 19 Feb. 1990. R.

,/. Hampshire , R. ./. Stafford. A. Reyes M.

Heath & A. Long 466 (holotype. MO; isolypes.
RM. MEXU). Figure 11.

Inter species Orthosiae inflorescentiis solitariis vel gem-
inatis axillarihus tubo corollae longo tubo coronae brevi
lobis coronae brevibus arcuatis gvnostegio conico longis-
tipitato dorsis antherarum vesiculatis poliniis anguste el-
lipsoideis longis differt.

Twining or scandent vines, latex and basal parts
unknown; voung stems puberulent in a fine line,
trichomes erect, ca. 0.1 mm long, white, internodes
3—5 cm. I .eaves opposite, caducous, without pseu¬
dostipules. Inflorescence axillary, 1 or 2 per node,
congested-racemifomi, sessile. 1 to 2 flowers open
at a time, pedicel 3.5—4 mm, puberulent. mostly in
I line, bracts debate, 0. 3-0.5 X 0.3 mm. green;
calyx with 1 to 3 eolleters below each sinus within,
lobes ovate with rounded tips. 0.8— 0.9 X 0. 7-0.8
mm. appressed-pu I >e ru len t outside, especially on
midrib, glabrous inside, ascending, green: corolla
campanulate. glabrous, white, tube ca. 1.5 mm
long, lobes ovate with rounded tips. 1.6— 1.7 X I-
1.1 mm, erect; gynostegium conical, anther backs
inflated, stipe ca. 1.4 nun. narrowed in middle, co¬
rona of 5 lobes connate at base, borne at the base
of the stipe, tube ca. 0.1 mm long, lobes debate
with acute tips, ca. 0.6 X 0.4 mm, arching outward
toward corolla tube and then back toward gynos¬
tegium. with the tips touching the stipe about a
quarter of the way up. terminal anther appendages
ovate, ca. 0.25 X 0.25 mm. arching over style apex;
anthers trapezoidal in outline, ca. 0.35 X 0.5 mm.
guide rails ca. 45° from vertical, straight, ca. 0.25
mm long; pollinaria pendent, corpusculum ellip¬
soid, ca. 0.17 X 0.08 mm, red-brown, translators
ca. 0.05 mm. thin, round in cross section, straight,
pollinia narrowly ellipsoid, ca. 0.23 X 0.05 nun:

style flat, 0.6 mm wide. Follicles and seeds un¬
known.

I bis new species, known only from the type, col¬
lected in cloud forest in Chiapas. Mexico, shares
the caducous leaves, inflated anther backs, coronal
structure and size, and general pollinaria charac¬
teristics w ith Orthosia ramosa. but has corolla lobes
about as long as the tube, instead of half as long,
a stipe about three times longer than that of O.
ramosa and a more strongly conical gynostegium.

Acknowledgments. Fil'd Keusenkothen and
Brigham Fisher prepared the images. Amy Fool im¬
proved the manuscript in many ways, especially in
reining in rhetorical excesses. I lie assiduous edi¬
tors of Novon took care of the rest.

Literature Cited

Fishbein, M. & W. I). Stevens. 2005. Resurrection of Seu-
tera Keichenbach (Apocynaceae, Asclepiadoideae). No¬
von 15: 531—533.

Liede, S. 1997. American Cynanchum (Asclepiadaeeae) —
A preliminary infrageneric classification. Novon 7:
172-181.

- & U. Meve. 2004. Revision of Metastelma (Apo¬
cynaceae— Asclepiadoideae) in southwestern North
America and Central America. Ann. Missouri But.
Card. 91: 31-86.

- & A. Tauber. 2002. Circumscription of the genus

Cynanchum (Apocynaceae— Asclepiadoideae). Syst. But.

27: 789-800.

Liede-Schumann, S., A. Rapini, I). J. Goyder & M. W.
Chase. 2005. Phylogenetics of the Now World subtribes
of Asclepiadeae (Apocynaceae— Asclepiadoideae): Me-
tastelmatinae, Oxypetalinae and Gonolobinae. Syst. Bot.
30: 184-195.

Rapini. A. 2002. Six new species of Ditassa R. Br. from
the Kspinhayo Range, Brazil, with notes on generic de¬
limitation in Metastelmatinae (Apocynaceae- Asclepia¬
doideae). kew Bull. 57: 565-583.

- . M. W. Chase, l). J. Goyder & J. Griffiths. 2003.

Asclepiadeae classification: Evaluating the phylogenet¬
ic relationships of New World Asclepiadoideae (Apo¬
cynaceae). Taxon 52: 33-50.

Spellman, I). I.. 1975. Asclepiadaeeae. ha R. I'.. Woodson.
Jr. et ah. Flora of Panama. Ann. Missouri Bot. Gard.
62: 103-156.

Standley. P. C. & I .. O W illiams. 1969. Flora of Guate¬
mala. Asclepiadaeeae. Fieldiana. Bot. 24(8, part 4):
107-172.

Stevens. W. D. 2002. A new Tassadia (Apocynaceae, As-
clepiadaceae) from Bolivia and Brazil. Novon 12: 408-
410.

- . C. l lloa If., A. Pool & <). VI. Montiel. 2001.

Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot.
( .a rd. 85: i— xlii. 1—2666.

Sundell, E. 1981. The New World species of Cynanchum
subgenus Mellichampia (Asclepiadaeeae). Evol. Mon¬
ogr. 5: 1-63.

Woodson, R. E.. Jr. 1941. North American Asclepiadaeeae
1. Perspective of the genera. Ann. Missouri Bot. Gard.
28: 193-244.

Johnstonia, a New Genus of Gouanieae (Rhamnaceae) from Peru

Roberto D. Tortosa

Laboratories de Bot^nica “L. R. Parodi,” Facultad de Agronomfa, Universidad de Buenos
Aires, Consejo Nacional de Investigaciones Cientfficas y Tecnicas, Av. San Marini 4453,
1417 Buenos Aires, Argentina. tortosa@agro.uba.ar.

Absi’KACT. A new genus, Johnstonia (Rhamna¬
ceae, Gouanieae), is proposed for Gouania axilli¬
flora M. (.. Johnston, a species from Peru affined
to Alvimuintha Grey-W ilson and Reissekia Endlich-
er. from which it differs by the sessile paired flow¬
ers in the axil of leaves; it is also distinct from
Alvimuintha by 1 1 it- entire menearps. I lie new com¬
bination Johnstonia axilliflora (M. C. Johnston) Tor¬
tosa is made.

RKSUMKN. Se propone un nuevo genero. Johns¬
tonia (Rhamnaceae. Gouanieae), para Gouania axi¬
lliflora M. G. Johnston, una especie de Peru affn a
Alvimuintha Grey-W ilson and Reissekia Kndlicher,
de I os cuales se diferencia por las flores geminadas
sesih-s en la axila de las hojas; tarnbien se distin¬
gue de Alvimuintha por los mericarpios enteros. Se
establece la nueva combinacion Johnstonia axilli¬
flora (M. C. Johnston) Tortosa.

Key words: Gouanieae, Johnstonia, Peru,
Rhamnaceae.

The tribe Gouanieae Reissek of the family
Rhamnaceae groups genera with inferior ovary,
generally with winged fruits (Suessenguth, 1953).
One of its genera. Gouania Jacquin. has a wide
distribution area in America (from Florida (U.S.A.)
and northern Mexico to Bolivia. Paraguay, and
northeastern Argentina and Uruguay), Africa, Asia.
Australia, and Oceania. Its species are climbers
with short shoots ending in a tendril, spike-like in-
florescences. flowers with a disk with conspicuous
bee lobules opposite to the sepals, and winged
fruits that divide into three mericarps, each sus¬
tained by two carpophore bundles.

Gouania axilliflora M. C. Johnston, proposed by
Johnston (1988) for a Peruvian plant, differs from
(’Ouania in the absence of tendril-shoots, absence
of floral disk lobules, and in the inflorescences
formed by sessile (lowers paired in the axil of
leaves. In the first two characters it matches two
other South American genera of Gouanieae, Alvi-
miantha Grey-W ilson and Reissekia Kndlicher. both
monotypic genera from the east of Brazil, with pe¬
dunculated umbel late-cymose inflorescences.

I he unique characters of the Peruvian species,
lead us to erect a new genus.

Johnstonia Tortosa, gen. nov. TYPE: Johnstonia
axilliflora (M. C. Johnston) Tortosa.

Rhamnacearurn, a genere Gouaniam lohulis disci ea-
rentihus, florihus geminis ad axilfani foliorum, absentia
ramulorum in capreolos terminantes differt: Alt imianthae
el Reissekiae affinis, sed florihus geminatis sessilihus al>
eis reeedens, al> Alvimuintha differt mericarpiis integris.

Climbers with tendrils in the flowering branches.
Leaves sparse, elliptic, entire, 3-nerved, petiolate.
Mowers in axillary, sessile, 2-flowered inflorescenc¬
es. Floral tube hemispherical; sepals 5, triangular,
externally pubescent; petals 5, concave; stamens 5,
with tin- anthers hooded by the petals; disk flat,
unlobed; ovary inferior, triloeular, with solitary
ovules in each loculus, style 3-branehed. stigmas
club-shaped, fruit schizocarp, giving rise to three
winged mericarps.

I he genus is named in honor of Marshal C. John¬
ston. who has clarified the taxonomy of many gen¬
era of Rhamnaceae from America.

1. Johnstonia axilliflora (M. C. Johnston) Tortosa,
comb. nov. Basionym: Gouania axilliflora M.
C. Johnston. Sysl. Bot. 13: 493. 1988. TYPE:
Peru. Department of Cajamarca, Province of
Cajabamba, Condebamba valley, Cajabamba—
Cajamarca road, 15 Feb. 1983, D. I\l. Smith &
R. Vdsquez M. 3396 (holotype, TEX; isotypes
(not seen) AMAZ, CPUN. HUT, MO. USM).

When Johnston (1988) proposed Gouania axilli¬
flora M. C. Johnston fora Peruvian plant, he stated
that in habit, foliage, and structure of the flowers
the new species is closely related to Alvimiantha
tricarnerata Grey-W ilson. charac terized by its flow¬
ers arranged in a pedunculate 2- to * -flowered con¬
gested cyme and by its winged seeds, lb- consid¬
ered that the characters used for recognition of the
genera in the tribe Gouanieae were so weak that he
rejected the alternative of erecting a genus for the
new species. Therefore, In- discussed the possibility

Novon 15:642-645. Purlisiikd on 12 December 2005.

Volume 15, Number 4
2005

Tortosa

New Genus from Peru

643

of including Alvimiantha within Gouania near G.
axilliflora, but considered that much field and lab¬
oratory study was necessary before such a decision.

Medan (1989) described the structure of the
fruits of the genera Alvimiantha and Gouania. both
with a 3-seeded schizocarp, with winged diaspores.
He showed that in the genus Gouania , when fruits
ripen, the three carpels separate along the planes
of the septa, and each meriearp hangs down from
the detached marginal bundles. Further, the wings
of the diaspores (mericarps) represent peric arp out¬
growths of the commissural areas of adjacent car¬
pels. In Alvimiantha. the fruits also undergo car-
pellary separation along the planes of the septa with
die detachment of marginal bundles, but a tangen¬
tial splitting of the pericarp wall occurs, gi\ing rise
to an outer coriaceous valve and a winged diaspore
(with a wingless seed included), confounded by
Grey-Wilson with a winged seed (Grey-Wilson.
1978: 290. figs. H, J. k). The fruits of Johnstonia
axilliflora conform with those of Gouania (fig. ID).

Johnstonia and Gouania differ in their habit.
Gouania bears serial buds in the leaf axi Is of or-
thotropic shoots (erect axis, with radial symmetry,
negatively geotropic) (Greiner. 1974; Inurn et ah.
1991); the distal bud of the axillary complex gives
rise to a tendril shoot (Fig. I A, B. ts). I liese tendril
shoots have determinate growth and generally hear
2 prophylls (Fig. IB, p) and a foliage leaf, in the
axil of which develop a tendril (fig. IB, I) and a
spike-like inflorescence (Fig. IB. ij (lourn el ah,
1991). Tendrils may also occur in the axis of the
inflorescences (Fig. IB, t,; see also illustration ol
G. latijolia Beissek, in Beissek. 1861, tab. 36). In
Johnstonia axilliflora, tendrils (fig. IG, t) occur
only at the nodes of plagiotropic shoots (axis ±
horizontal, with dorsiventral symmetry) (fig. 1G. ps;
see also Johnston, 1988: 494, fig. 1, a); serial buds
and tendril-shoots are lacking. Alrimiantlui and
Reissekia correspond in this aspect of habit to
Johnstonia (Fig. IE; see also Grey-Wilson, 1978:
290, fig. 1, and Beissek. 1861. tab. 40. respective¬
ly)-

Inflorescences in Gouania are spike-like ra¬
cemes (Fig. IB. r, r,), in Johnstonia 2-flowered
glomerules (Fig. IG, g). and in Alvimiantha and
Reissekia pedunculated umbellate-cymes (fig. IE,
u). In Gouania the disc of the flower is conspicuous
with 5 elevated acuminate, obtuse, truncate or
emarginate lobules, opposite to the sepals, while in
Alvimiantha. Johnstonia, and Reissekia the floral
disk is thin. flat, and unlobed.

All vegetative and reproductive characters place
Johnstonia axilliflora near Alvimiantha, but the dis¬
tinct trait of the fruit in the latter supports the de¬

cision to treat this species as belonging to a new
genus.

K I. > TO GENERA of Goi ANIEAE IN AMERICA

la. Climbers with tendrils.

2a. Floral disc with conspicuous lobes opposite
the sepals; tendril-shoots present; inflores¬
cence a spike-like raceme . Gouania

2b. Floral disc without lobes; tendril-shoots ab¬
sent.

3a. Leaves entire, 3-nerved.

4a. Mericarps undivided; inflorescence

a 2-flowered glomerule . . . Johnstonia
4b. Mericarps splitting in an outer co¬
riaceous valve and a winged dia¬
spore; inflorescence a pedunculated

umbellate-cyme . Alvimiantha

3b. Leaves finely serrate, pinnately nerved
(obscurely 3-nerved); inflorescence a
pedunculated umbellate-cyme . . Reissekia

lb. Perennial, erect or annual, decumbent herbs . .

. Crumenaria Martins

Representative specimens examined. Alvimiantha. A.
trieamerata Grey-Wilson: BKAZIL. 28 km NW of Lagoin-
ha (which is 5.5 km SW of Delfino) on side road to Minas
do Mimoso, Gaatinga, R. M. Harley 16890 (photos of iso¬
types MO, NY). Gouania. G. udenophora Pilger: BOLIV¬
IA. Pando: Madre de Dios, I’rov. Barranco de Puerto
Candelaria, 140 m s.m., 19— V III— 1985, Monies 228 (LPB.
NY). G. aptera DC.: PERU. Amazonas: l.uya, Camporre-
dondo, 1 8— V I- 1 999. Campos 6 027 (BAA). G. latijolia
Beissek: BOLIVIA. Santa Cruz: Prov. Florida, 5 km (by¬
road), 3 km (by air) NE of Bermejo, along petroleum pipe¬
line and old horse road to Santa Cruz, 17— VII— 1987, Nee
A- Coimbra S. 85252 (LPB. MO, NY). G. longipetala
llemslev: CAMEROON. Bipinde, X-1913. G. Gaubay 402
(SI). G. longispicata Engler: RHODESIA. Nyassa, Hoeh-
land. Station kymbila. 1911, A. Stolz 901 (SI). G. lupu-
loides (L) Urban.: U.S.A. Florida: Indian River Region,
27-X-1902. C. Brevard (SI); ARGENTINA. Misiones:
Iguazu, Cataratas del Iguazii. Bah on Sallo Brasilenos, 30-
\ 11-1979. Tortosa et al . 124 (BAA). G. microcarpa DC.:
PHILIPPINES. Luzon: Prov. Rizal. XII-1909. \1. Ramos
96 (SI). G. polygama (Jaequin) Urban: BOLIVIA. Prov.
Nor Yungas, 16.5 km NE, of Chuspipata, 13.6 km SW of
Yolosa, 3— IV— 1984, Solomon & Gelding 121 18 (MO). G.
trichodonta Beissek: BOLIVIA. Pando: Prov. I. Roman.
Rio Negro por el Rio Negro. 120 m s.m., 17— VI— 198/.
Solomon 17089 (MO). G. ulmifolia Hooker: PAR At, l AY.
Alto Parana: Escuela Tecniea Forestal, I ’to. Pte. Stroess-
ner. km 12. 01-1982, Fernandez Casas 5656 (G. MO):
BRAZIL. Parana: Ponta Grossa. 13-11-191 I. /’ Dressen
(SI).

Acknowledgments. Fhe author gratefully ac¬
knowledges the assistance of B. L. Turner for mak¬
ing accessible material ol the holotype of 6. axilli¬
flora M. C. Johnston; Marshal C. Johnston and an
anonymous reviewer for critical comments; Victoria
G. Hollowell for editorial advice; and Universidad
de Buenos Aires for financial support.

644

Novon

Figure I. A. B. Gouania. A. Axillary complex. — B. Vegetative and reproductive structures at a node. C. I), John-
stonia ax ill {flora. — C. Vegetative and reproductive structures at a node. — I). Cross section of a mericarp (from Smith
& Vdsquez 3396, holotype, TEX). — E. Alvimianlha tricamerata. Vegetative and reproductive structures at a node.
References: g, 2-flowered glomerule; os, orthotropic shoot; p, prophyll; ps, plagiotropic shoot; r and r,, spike-like
raceme; t and t„ tendril; ts, tendril shoot; u, umbellate-pedunculate inflorescence; y3, dormant proximal bud.

Volume 15, Number 4
2005

Tortosa

New Genus from Peru

645

Literature Cited

Greniers, G. 1974. Architecture de quelques lianes d’Af-
rique tropicale. Candollea 29(1): 57— 110.

Grey- Wilson. C. 1978. Alvimiantha, a new genus of
Rhamnaceae from Bahia, Brazil. Bradea 2(4.1): 28 7—
290.

Johnston, M. C. 1988. Gouania axilliflora (Rhamnaceae),
a new species from Peru. Syst. Bot. 13(4): 493-495.

Medan, I). 1989. Diaspore diversity in the anemochorous
Gouanieae (Rhamnaceae). PI. Syst. Evol. 168: 149—158.

Reissek, S. 1861. Rhamneae. In: Martins, H. Bras.: 80—
1 1 5, tab 24 — 4 1.

Suessenguth, K. 1953. Rhamnaceae. In: A. Engler & K.
Prantl, Die naturlichen Pflanzenfamilien 20d: 7-173.
Duncker & Humblot, Berlin.

Tourn, G. M., A. Bartoli & R. 1). Tortosa. 1991. The mor¬
phology and growth of Gouania ulmifolia Triana &
Planch. (Rhamnaceae): An architectural analysis. Pp.
666—667 in C. Edelin (editor). L'Arbre. Biologic el IV-
veloppement, Memoires 2 hme. Colloque International
L’Arbre, Montpellier, France.

Iwo l\ew Species of Senecio (Asteraceae, Senecioneae)

from Argentina

Roberto I). Tortosa

Laboratorios de Botanica “L. R. Parodi, Facultad de Agronotnia, Universidad de Buenos
Aires, Av. San Martin 4453, 1417 Buenos Aires, Argentina, tortosa@agro.uba.ar; and
Consejo Naeional de Investigaciones Cientificas y Teenicas (CONICET), Argentina.

Adriana Bartoli

Laboratories de Botanica "I.. B. Parodi,” Facultad de Agronomfa, Universidad de Buenos
Aires, Av. San Martin 4453, 141 i Buenos Aires, Argentina. cbartoli@agro.uba.ar

Abstract. Two new species of Senecio 1,. from
Argentina, S. fragrant issimus and .S', pumilus, are
described and illustrated. Both species belong to
Senecio ser. Xeroseneno (Cabrera) Cabrera A S. K.
1 re ire and are allied to S. filaginoides DC., from
which S. fragrant issimus differs by its ascending
branches with needle-like leaves crowded along the
stems, and S. pumilus diverges by its dwarf cush¬
ion-like habit and elliptic, thicker leaves. A key is
included to those species affined to S. filaginoides.

Rk.SUMKN. Se describen e ilustran dos nuevas es-
peeies de Senecio I., de Argentina, S. fragrantissi-
tn us y S. pumilus. Ambas perteneeen a Senecio se-
rie Xerosenecio (Cabrera) Cabrera & S. E. Freire y
son alines a S. filaginoides DC., de la cual S. fra-
grantissimus difiere por sus ranias aseendentes eon
hojas aciculares aglomeradas a lo largo de las ra-
mas, y .S', pumilus diverge por formar cojines eon
hojas ehpticas mas crasas. Se incluye una clave
para las especies afines a S. filaginoides.

Key words: Argentina, Asteraceae, Compositae,
Senecio.

The series Xerosenecio (Cabrera) Cabrera & S. F.
Freire of the genus Senecio I,, sect. Senecio (sensu
Cabrera el al„ I 999) is characterized by a ligneous
habit and discoid eapitula disposed in terminal cor¬
ymbs. Cabrera (1949) and Cabrera el al. (1999) in¬
cluded 39 species in this series, distributed in Ar¬
gentina and Chile, with only one species (,S. vira-vira
Hieronymus) also in Uruguay and in southeastern
Brazil in the state ol Bio Grande do Sul.

During a taxonomic revision of Senecio ser. Xe¬
rosenecio, we found two new species from Argenti¬
na. both affined in Senecio filaginoides DC. by their
woolly stems, leaves, and phyllaries, and by their
narrowly eylindrie or obeonical involucres, less
than 8 mm high, composed of 10 lo 14 phyllaries.

Senecio iVugranlissimus Tortosa cK A. Bartoli, sp.
nov. I 'l PE: Argentina. San Luis: Pringles, Sa-
ladillo, “Flores color erema, perfumadas,” 9
Nov. 1940, G. Cotas 13X7 (holotype, I.P). Fig-
ure 1.

Affinis Senecioni filaginoidi DC. sed ramis hornotinis
ascendentibus loins acieularis eongestis notabilis differt.

Shrub 50-80 cm, mostly ramified from the base,
lanuginose or sublanuginose on stems, leaves, and
phyllaries; stems ascending with very short inter-
nodes from midstem upward. Leaves alternate, ses¬
sile, acicular, ± spreading, 0.9-3. 5 X 0.04-0.06
cm. margin revolute, lanuginose or sublanuginose
on both surfaces. Capitula pedunculate, arranged 5
to 15 in terminal corymbs; peduncles 0.5-1. 3 cm
long, involucre ealyculate, eylindrie, 0-8 X 2.5-3
mm; phyllaries 10 to 12, linear ovate, 6-8 X ca. I
mm. dorsally lanuginose or sparsely lanuginose,
apex acuminate with a tuft ol short triehomes; ca-
lycular bracts ca. 4, linear, ca. 1.5 mm long, dor-
sally lanuginose. Disc florets 22 to 25, hermaph¬
roditic, corolla cream, tubular, 5-lobed, 7-7.5 mm
long, glabrous, basally broadened, limb slightly
broader than the tube and not markedly distinct,
lobes narrowly triangular, ea. 0.6 mm long, al base
0.3 mm wide; anthers minutely tailed, ea. 2-2.5
mm long; filament collars spatulate ca. 0.5 mm
long; apical anther appendage obovate, ea. 0.4 mm
long; basal tails ea. 0.15 mm long. Style branches
ea. 1.5 mm long. Achenes eylindrie. pubescent, ea.

3 mm. eostate; pappus bristles many, slender, bar-
bellate, t— 8 mm long, whitish.

Distribution and habitat. Senecio fragrantissi-
mus is known only from two neighboring localities
in the political province of San Luis of western Ar¬
gentina. in xerophytic habitats with sandy soil at
<800 m.

Novon 1 5: 646-649. Published on 12 December 2005.

Volume 15, Number 4
2005

Tortosa & Bartoli
Senecio from Argentina

647

figure 1. Senecio fragrtiritissimus Tortosa & A. Bartoli. — A. Habit. — B. Capituluni. Drawn from the type, (>. Covns
1337 (I.P).

Etymology. The specific epithet makes refer¬
ence to the fragance to which all collectors of this
species alluded.

Senecio fragantissimus is allied to S. filaginoides,

a species with a wide distribution in Argentina (Ca¬
brera el al., 1999). The new species differs from
the latter by its needle-like leaves, crowded togeth¬
er due to congestion of the upper stem internodes.

648

Novon

Figure 2. Senecio pumilus Tortosa & A. Bartoli. — A. Habit. — B. Capitulum. Drawn from the type, E. Ancibor y //.
Schwabe s.n. (BAA 5789).

giving the branches and their foliage a cylindrical
appearance. A similar aspect may be seen in some
specimens of .S', bracteolatus Hooker tS. Arnott (().
Boelcke 42(19 (BAA), A. Havrylencco 6 and 7
(BAA), Vallerini 349 (BAA)), but this species is

glabrous and its heads have a carnpanulate invo¬
lucre.

Puratypes. ARGENTINA. Sail Luis: Pringles, Sala-
dillo, 9 Nov. 1940, A. Burkart (LP); 9 Nov. 1940, Ennis
1001 (I P): General Pedernera, al pit4 del Cerro El Morro,

Volume 15, Number 4
2005

Tortosa & Bartoli
Senecio from Argentina

649

cerca de la Ea. Ea Morena. 18 Oct. 1951, .4. /.’ Hunziker
9287 (LP).

Senecio puiiiilus Tortosa & A. Bartoli, sp. nov.
TYPE: Argentina. Mendoza: Malargiie, Porte-
zuelo del Choique, 2400 m s.m., 12 Mar. 1967.
E. Ancibor & H. Schwabe s.n. (holotype, BAA
5789). Figure 2.

Inter species seriei Xerosenecionis indumento lanato,
capitulis parvis in corymbis dispositis, involucre angusto
ad S. jilaginoidem DC. accedens sed statura minorc. foliis
ellipticis erassioribus involucro obconico ab ea recedens.

Dwarf shrubs 8—30 cm, much branched at the
base, stems, leaves, and phyllaries densely whitish
lanuginose; stems prostrate subligneous giving rise
at their nodes to herbaceous ascending shoots.
Leaves alternate, sessile, 0.8—2 X 0.15—0.30 cm,
elliptic, obtuse or rounded at the apex, margin rev¬
olute, densely lanuginose on both surfaces. Capit-
ula pedunculate, peduncles 0.5-1. 5 cm long, ar¬
ranged 3 to 6 in terminal corymbs; involucre
calyculate, obconical, 6-7 X 3-3.5 mm; phyllaries
8 to 12, 6—7 X 1 — 1.2 mm, dorsally lanuginose,
apex acuminate with short triehomes; calvcular
bracts ea. 5. elliptic, ea. 1—1.5 mm long, dorsally
lanuginose. Disc florets ca. 20 to 24, hermaphro¬
ditic, corolla yellow, tubular, 6-6.5 mm long, gla¬
brous, basal ly broadened, narrowly campanulate,
lobes triangular, ca. 0.5 mm long, at base 0.3 mm
wide; anthers minutely tailed. 2—2.5 mm long: fil-
ament collars spatulate, ca. 0.25 mm long; apical
anther appendages deltoid, ca. 0.3 mm long; the
hasal tails ca. 0.1 mm long. Style branches ca. 1.2
nun long. Achenes cylindric, glabrous, ca. 3.5 mm,
costate. Pappus bristles many, slender, barbellate.
ea. 6 mm long, whitish.

The heads of Senecio pitmilus are similar to those
of S. filaginoides, but its dwarf cushion-shaped hab¬
it. not taller than 30 cm (vs. erect. 0.5—1 m tall in
S. filaginoides ), its elliptic, entire, thicker leaves
(vs. linear to linear-elliptic in S. filaginoides), and
its obconical involucre (vs. cylindric in S. filagi¬
noides) distinguish the new species.

Distribution and habitat. Senecio pumilus has

been found in the eastern Mendoza and Neuquen
in Argentina, at elevations of 1900—2500 m ol the
Andes Mountains, an arid region with cool weather
exposed to strong winds.

Etymology. The specific epithet makes refer¬
ence to the habit of the species.

Paratypes. ARCENTINA. Mendoza: Malargiie. I'or-
tezuelo del Choique, E. Ancibor A II. Schwabe s.n. (BAA
5784); Ca. Portezuelo del Choique, ,/. II. Hunziker 6881
(BAB); Malargiie, A. Burkart, Y .S. Troncoso A E. G. \i-
cora s.n. (SI 14338); Cerros al N de Calmuco, .4. Burkart
s.n. (SI 14330): Laguna Coipo-Leuquen, al S de Calmuco,
4. Burkart. N. S. Troncoso A E. 6'. Nicora s.n. (SI I 4340);
6 km N tie Calmuco. G. Corns 412 (LP); Entre Calmuco
v Pto. Los Colgados, 4. Prina, G. Alfonso A 1L Muino
1870 (SBFA); in montanis Valle del Yeso. Ceding s.n.
(LP). Neuquen: Minas. Cordillera del Viento. arroyo Dos
Puentes. E Pasture 3 (SI).

Ki'.'t to Species Ai.i.ikd to Si:\i:<:io fuagisoidks

la. Leaves acicular, less than 0.6 mm wide .

. Senecio fragrantissimus Tortosa & A. Bartoli

lb. Leaves linear or elliptic, 1.5—3 mm wide.

2a. Phyllaries 7 to 8; florets 10 to 12 .

. Senecio pycnanthus Philippi

2b. Phyllaries 10 to 14; florets It! to 26.

3a. Shrubs 0.08—0.30 m high: leaves ellip¬
tic, entire; involucre obconical ....

.... Senecio pumilus Tortosa & A. Bartoli
3b. Shrubs 0.5—1 m high; leaves linear or
linear-elliptic, entire, toothed or lobed;

involucre cylindrical .

. Senecio filaginoides DC.

Acknowledgments. We thank the anonymous re¬
viewers and Victoria C. Hollowed for reviewing the
manuscript and for useful comments. We also ac¬
knowledge the assistance of the curators of BAB,
TP, SI. and SB FA that made this study possible.
Thanks to the artist. Francisco Rojas, for the draw¬
ings. This study was supported by the Universidad
de Buenos Aires.

Literature Cited

Cabrera, A. L. 1949. El genero Senecio (Compositae) en
Chile. Lilloa 1 5: 27—501 .

- , S. E. Ereire & L. Ariza Espinar. 1999. Tribu VIII.

Senecioneae. Pp. 3-164, 171-179 in A. T. Hunziker
(editor). Flora Fanerogamica Argentina 62, 280. Aste-
raceae, parte 13.

Clematis erectisepala (Ranunculaceae), a New Species from

Eastern Tibet, China

Lei Xie*, Jing-hua Shi, and Liang-qian Li **

Laboratory of Systematic & Kvolut ionary Botany and Herbarium, Instilute of Botany,
Hie Ch inese Academy of Sciences, 20 Nanxincun. Xiangsfum, Beijing 100093,

Peoples Republic of China.

*Graduate School, The Chinese Academy of Sciences, Beijing 100039, China.

xielei@ibcas.ac.cn

^^Corresponding author: lqli@ibcas.ac.cn

Abstract. Clematis erectisepala, a new species of
Kanmiculaceae collected from Yanjing Village of
Markam County in eastern Tibet, is described and
illustrated. It is distinguished from other Clematis
by its nodding yellow (lowers with erect sepals and
glabrous stamens. This new species is similar to C.
brevicaudata DC. vegetatively, but differs from the
latter in floral structure.

Key words: China, Clematis, eastern Tibet,
Ranunculaceae.

Clematis L., containing about 300 species, is one
of the largest genera in Ranunculaceae (Wang,
1998). It is distinguished from other genera of the
family by woody climbing stems (for most Clematis
species) and opposite leaves. The genus Clematis ,
widely distributed in all continents except Antarc¬
tica, exhibits significant diversification in the
Hengduan Mountains (western Sichuan, eastern l i¬
bel. and western Yunnan provinces in China). Lin¬
naeus (1753) first described nine species of Cle¬
matis in his great work Species Plant-arum. The
genus as delimited by Candolle (1818) has been
accepted as a natural group. Piantls (1888) clas¬
sification of Clematis, with minor changes and ad¬
ditions, has been generally accepted until today.
Recently, increasing attention has been paid to Cle¬
matis due to its great horticultural value (Tamura,
1995; Johnson. 1997; Grey-Wilson, 2()()0). In the
summer and autumn of 2003, an extensive field
investigation was conducted in the Hengduan
Mountains. Based on thorough examination and
identification of the specimens of Clematis collect¬
ed during the exploration and their comparison to
the specimens deposited in PK and KUN, a new
species was recognized as Clematis erectisepala L.
Xie. J. H. Shi & L. Q. Li.

Clematis erectisepala L. Xie. J. H. Shi & L. Q.
Li. sp. nov. TYPE: China. Tibet: Markam
County, Yanjing Village, climbing shrubs along
the roadside, ca. 2650 m, 29°02,33WN,
95°45'87"E, 31 Aug. 2003, L Xie YJ01 (ho-
lotype, PE; isotypes, MO. PE |2|). Eigures 1, 2.

Ilaec species C. brevicaudatae DC. affinis, sed al> ea
floribus niitantilius atque calyce campanulato ex sepalis
luteis ereetis eonstante differ!.

Woody vines; branches shallowly 6-grooved,
sparsely puberulous. Leaves mostly ternate at up¬
per nodes, with some bi-ternate and pinnate leaves
at lower nodes; petioles 30-50 mm; leaflet blades
papery, ovate or narrowly ovate, 21-30 X 12-22
mm, 3-lobed or undivided, both surfaces glabrous,
margin coarsely dentate or entire or denticulate,
base truncate or broadly cuneate, rarely rounded;
apex acuminate or caudate; basal veins abaxially
slightly prominent. Cymes terminal and axillary, 3-
lo many-flowered, often panicle-like; peduncle 6—
10 cm. sparsely appressed puberulous; bracts op¬
posite. subsessile, narrowly ovate to lanceolate, 4—
13 mm long, 2-4 mm wide. Pedicel 5-10 mm. pu¬
berulous; calyx eampanulate, 10 mm diam.; sepals
4, ye flow, erect, oblong, 10 mm long, 3—4 mm wide,
margin abaxially velutinous, apex acute and re¬
curred: stamens numerous, 8-10 mm long, fila¬
ments linear, glabrous, anthers narrowly oblong, 2—
3 mm, glabrous, apex obtuse; ovary puberulous;
style 5 mm, densely villous. Fruit not seen.

Pollen grain prolate spheroidal (20 X 15 gm),
trieolpate; eolpi long, thin, running from pole to
pole; exilic bears a perforate tectum with mieroe-
ehinate ornamentation (Figure 2A— C).

I he shapes of leaf epidermis cells are irregular
on both the adaxial and abaxial blade surfaces, with
stomata only appearing abaxially. The stomata! ap¬
paratuses are all anomocytic. The outlines of the
guard cell pairs are suborbieulate, and the rims are

Novon 15: 650-653. Published on 12 December 2005.

Volume 15, Number 4
2005

Xie et al.

Clematis erectisepala (Ranunculaceae)

651

Figure 1. Clematis erectisepala L Xie. J. H. Slii & L (,). Li. — A. Habit. — 15. Flower. — C. Sepal (adaxial side). — I).
Sepal (abaxial side). — E. Stamen. Drawn from L. Xie YJOI (PE).

almost at the same level with the epidermis. The
wax ornamentation of the leaf cuticle is nearly
smooth on the adaxial surface and heavily striated
on the abaxial surface (Figure 21)— F).

Distribution. Endemic to the Flengduan moun¬
tains in China, it has been recorded so far from
only four counties: Nanping County, Barkam Coun¬
ty (Sichuan Province); Yanjing County (Tibet Au¬
tonomous Region); and Deqen County (Yunnan

Province). It has been found at 1500—2700 m ele¬
vation.

Flowering and fruiting. Flowering time August
to September; fruit not seen.

Ecology. The new species is found in the edge
of Yanjing Village, which is located in Me Kong
River valley at an altitude of 2650 m. It scrambles
up shrubs of Rosa L. and is associated with Cle-

652

No von

Volume 15. Number 4
2005

Xie et al.

Clematis erectisepala (Ranunculaceae)

653

mntis rehderiann Craib and C. akebioides (Maxi-
mowiez) Veitch, which are two common Clematis in
the Hengduan Mountains.

Etymology. The species epithet refers lo the
erect sepals of this species; most Clematis with gla¬
brous stamens have spreading sepals.

Clematis erectisepala L. Xie, J. H. Shi & L. Q.
Li is allied to Clematis brevicaudata DC. by its hab¬
it, inflorescence type, leal shape, and other vege¬
tative characters. However, the new species differs
by floral structure: the sepals are yellow, erect and
the flowers are bell-shaped and always nodding,
fhe new species is often misidentified as C. peterea
Handel-Mazzetti and C. brevicaudata DC. by its
leaf shape and glabrous stamens.

Paratypes. CHINA. Sichuan: Nanping County, river
valley, all. 1500 rn. 7 Aug. 1983, /.. A. Tang 1648 (PE);
Barkam County, river valley, alt. 2650 m, 3 Aug. 1977,
Anonym 23328 (PE). Tibet: Yanjing County, river valley,
alt. 2700, Comprehensive exploration team 76—767 (PE).
Yunnan: Deqen County, river valley, alt. 2100 m. 25
Aug. 1981, 1. F. Han. K. M. Deng & Y. R. Chen HI-
1765 (PE).

Acknowledgments. We express our thanks to
Prof. W. T. Wang for his invaluable advice on the
manuscript. We are indebted to Y. H. Xiao and X.
.1. Yang for their assistance on the SEM experiments
and film development. We are also grateful to Y. B.
Sun for the drawing. This work was supported by
the National Natural Science Foundation of China
(Grant No. 30470126).

Literature Cited

Candolle, A. P. tie. 1818. Clematis. In: Regni Vegetabilis
Systema Naturale I: 131-167. Paris.

Grey-Wilson, C. 2000. Clematis the Genus. Timber Press,
Portland.

Linnaeus, C. V. 1753. Species Plantarum. Stockholm.
Johnson, M. 1997. Slaktet Klematis. Sodertiilje: Magnus
Johnson’s Plantskola AB.

Prantl, k. H188. Ranunculaceae. In: A. Engler X k.
Prantl (editors). Die natiirliehen Pflanzenfamilien. 3(2):
43—66, 274. Leipzig.

Tamura, M. 1995. Clematis. In: P. Hiepko (editor). Die
natiirliehen Pflanzenfamilien. Ed. 2. 17 a l\: .368-387.
Duneker & Humblot, Berlin.

Wang, W. I. 1998. Notulae de Ranunculaceis Sinensibus
(XXII). Acta Phytotax. Sin. 36: 150—172. 1 1 n Chinese
with English abstract.]

Eigure 2. Clematis erectisepala L. Xie, J. H. Shi & L. Q. Li. Scanning electron photomicrographs of pollen grains (A-
C). — A. Polar view. — B. Equatorial view. — C. Pollen exine details. Scale bar = 7.5 /ran. Scanning electron photo¬
micrographs of leaf epidermis (D-F). — 1). Adaxial epidermis. — E. Abaxial epidermis. — F. Stomatal apparatus. Scale
bar = 50 /iun.

Volume 15, Number 1. pj ». 1—252 of NOYON was published on 29 April 2005.
Volume 15, Number 2, pp. 253—384 of NOVON was published on 13 July 2005.
Volume 15, Number 3. pp. 385—502 of NOVON was published on 21 September 2005.
Volume 15. Number 4, pp. 503—654 of NOVON was published on 12 December 2005.