Volume 12 NO VON

Number 1

2002

A Synopsis of the Central Asian Rhammatophyllum (Brassicaceae)

Ihsan A. Al-Shehbaz

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

Oliver Appel

Bredkamp 36E, 22589 Hamburg, Germany

Abstract. The genera Koeiea and Prionotrichon
are reduced to the synonymy of Rhammatophyllum.
The new combinations R. afghanicum, R. erysimo-
ides, R.flexuosum, R. gaudanense, R. ghoranum, R.
kamelinii, and R. pseudoparrya are proposed. A key
to all nine species of the central Asian Rhamma¬
tophyllum is presented.

Key Words: Asia, Brassicaceae, Koeiea, Mito-
phyllum, Prionotrichon, Rhammatophyllum.

During work on the Brassicaceae for the forth¬
coming volume of Kubitzki’s Families and Genera
of Vascular Plants, it became evident that the limits
of several genera need adjustment and that some
nomenelatural changes are necessary. A case in
point is the delimitation of the genera Rhammato¬
phyllum O. E. Schulz, Prionotrichon Botschantsev
& Vvedensky, anti Koeiea K. H. Rechinger.

Schulz (1933a) proposed the genus Mitophyllum
to accommodate a species previously placed in Ar-
abis L, A. pachyrhizum Karelin & Kirilov, which is
endemic to Kazakhstan. He (Schulz, 1933b) re¬
named the genus Rhammatophyllum because he
discovered that his Mitophyllum is a later homonym
to that of Greene (1904). Mitophyllum Greene is
now recognized as a synonym of Streptanthus Nut-
tall (Al-Shehbaz, 1985; Rollins. 1993). Rhamma¬
tophyllum remained monotypic until Botschantsev
(1952) and Vassiljeva (1969) each added another
species.

Botschantsev and Vvedensky (1948) proposed
the genus Prionotrichon and separated it from the
closely related Rhammatophyllum by flower color,

lack of the septum venation, and the type ol in¬
dumentum. The (lower color is said to he yellow in
Prionotrichon and white in Rhammatophyllum, hut
the type species of the latter genus has creamy
white to yellow (lowers. The trichomes in both R.
pachyrhizum and R. frutex are medifixed, sessile,
and their two rays are rather soft and often crisped.
Typically malpighiaceous trichomes, which are
found in Erysimum I,., Lohularia Desvaux, Farsetia
Turra, and many other genera of Brassicaceae, are
also medifixed and 2-rayed, but their rays are al¬
ways rigid and straight. In the two original species
that Botschantsev and Vvedensky (1948) assigned
to Prionotrichon, P. pseudoparrya and P. erysimo-
ides, the trichomes strongly resemble those of
Rhammatophyllum in having soft, crisped rays, hut
their two rays have a few minute lateral branches.
Finally, Botschantsev and Vvedensky (1948) indi¬
cated that the septum in Rhammatophyllum is 3-
veined, while that of Prionotrichon is not veined.
However, the septum in species of Prionotrichon
has a broad central band sometimes marked by the
presence of either a distinct midvein or three faint
veins. Veined septa are rare in the Brassicaceae,
and their occurrence in both Rhammatophyllum
and Prionotrichon further supports the union of the
two genera. Therefore, we believe that the differ¬
ences in flower color, indumentum, and septum,
which were claimed by Botschantsev and Vveden¬
sky (1948) to support the recognition of two inde¬
pendent genera, are artificial and only one genus is
represented.

Botschantsev (1966, 1987) expanded the limits

Novon 12: 1-4. 2002.

2

No von

of Prionotrichon to include most of the species that
Rechinger (1954, 1964, 1968) placed in Koeiea. It
is interesting to note that Rechinger (1954) as¬
signed Koeiea to the tribe Alysseae but later lie
(Rechinger, 1968) assigned it to the tribe Matthi-
oleae. By contrast, Rotschantsev (1966, 1987)
placed the combined genus in the Arabideae,
whereas Schulz (1936) placed Rhammatophyllum
next to Erysimum in the tribe Hesperideae. The
placement by these three authors of what we con¬
sider one genus in four tribes clearly shows the
difficulties in the evaluation of generic and tribal
relationships in the Rrassicaceae based on few mor¬
phological characters. Without molecular studies
on this complex, it is uncertain what the nearest
relatives of Rhammatophyllum (including Koeiea
and Prionotrichon) are. On the basis of habit, leaf
and fruit morphology, and indumentum, it appears
that Rhammatophyllum is perhaps more related to
Erysimum than to any other genus.

Rechinger's (1968) account of Koeiea for Elora
franica included a heterogeneous assemblage of
species now assigned to different genera. For ex¬
ample, K. altimurana K. II. Rechinger was shown
by Botsehantsev (1987) to he a synonym of A rah is
fruticulosa C. A. Meyer, a species also treated in
that same flora (Hedge, 1968) in a different tribe,
Arabideae. It is highly unlikely, however, that A.
fruticulosa would eventually be retained in Arabis
because the genus, as presently delimited, is clear¬
ly polyphyletic and needs to be divided into several
segregates (Koch cl al.. 1999).

Polatsehek and Rechinger (1968) retained Erys¬
imum gaudanense Litvinov in Erysimum despite
their observation that the trichomes are not typical
of that genus because they have soft, crisped rays
with minute lateral branches. On the basis of tri-
clmme morphology and other aspects of the plant,
Botsehantsev (1987) was correct in placing the spe¬
cies in Prionotrichon and in considering it to be
closely related to the species we recognize herein
as Rhammatophyllum erysimoides, R. ghoranum, R.
kamelinii , and R. pseudoparrya.

Rechinger (1968) transferred Parrya karatavica
Lipschitz to Koeiea , but because of its typically
stellate, sessile, rigidly rayed trichomes and den¬
tate. canescent, closely overlapping leaves, the spe¬
cies does not resemble any member of Parrya or
Rhammatophyllum (including Koeiea and Priono¬
trichon). Nabiev (1972) placed the species in his
monotypie genus Botschantzevia and indicated that
it differs from Arabis , Prionotrichon, and Rham¬
matophyllum in having different trichomes. How¬
ever, the placement of this species in a monotypie
genus is inappropriate, and we believe that it shows

more affinities to Erysimum than to any of the gen¬
era above.

As delimited herein, Rhammatophyllum includes
nine species distributed from eastern Turkmenistan
eastward through Afghanistan, northern Tajikistan,
southern Kyrgyzstan, eastern Kazakhstan, and the
Altay region of westernmost Mongolia. The genus
has not yet been collected from western Xinjiang
(China), but it is likely to be found at least in the
western parts of Dzungaria. Rhammatophyllum
consists of shrubs or subshrubs with linear to fili¬
form. entire leaves that are attenuate into a persis¬
tent base, often slightly swollen nodes, minutely
stalked, softly submalpighiaceous, or rarely sub-
dendritic trichomes, yellow, creamy white, or rarely
purple petals, latiseptate, torulose fruits, distinctly
veined fruit valves, and flattened, winged, mar¬
gined, or rarely wingless seeds. We consider the
presence of incumbent cotyledons in the species
originally placed in Rhammatophyllum and aecum-
bent ones in those assigned to Prionotrichon to he
a minor character insufficient to support the main¬
tenance of two genera.

|{|iaiiiiiialopliylliiin (). E. Schulz, Report. Sp.
Nov. Regni Veg. 33: 190. 1933. Based on Mi-
tophyllum 0. E. Schulz, Notizbl. Bot. Cart.
Berlin-Dahlem II: 872. 1933, non Greene,
Leafl. Bot. Observ. Grit. 1: 88. 1904. TYPE:
Rhammatophyllum pachyrhizum (Karelin &
Kirilov) ( ). E. Schulz.

Prionotrichon Botsehantsev & Vvedensky, Bot. Mater.
Gerh. Inst. Bot. Zool. Akad. Nauk Uzbekst. S.S.B.
12: 8. 1948. Syn. nov. TYPE: Prionotrichon pseu¬
doparrya Botsehantsev & Vvedensky.

Koeiea k. II. Rechinger, Anz. Osterr. Akad. Wiss.. Math.-
Naturwiss. kl. 91: 60. 1954. Syn. nov. 14 PE: Koeiea
afghanica k. H. Rechinger.

Plants subshrubs or shrubs. Trichomes unicel¬
lular. eglandular, softly malpighiaceous, suhmal-
pighiaeeous with a few lateral minute branches,
rarely subdendritic. Stems erect to ascending,
branched, slightly swollen at nodes. Basal leaves
absent. Catiline leaves sessile or rarely petiolate,
filiform to linear or lanceolate, not auriculate, en¬
tire. Racemes several flowered, ebracteate, corym¬
bose, elongated in fruit. Fruiting pedicels slender,
ascending to divaricate. Sepals oblong, deciduous,
ascending, subequal, base of lateral pair not sac¬
cate. margin membranous. Petals yellow, creamy
white, or rarely purple, longer than sepals, distinct¬
ly clawed; blade obovate to spatulate, apex obtuse.
Stamens 6. tetradynamous; filaments not dilated or
slightly flattened at base; anthers oblong, apiculate
at apex. Nectar glands confluent and subtending

Volume 12, Number 1
2002

Al-Shehbaz & Appel

Central Asian Rhammatophyllum

3

bases of all stamens; median glands present or ab¬
sent; lateral glands often annular. Ovules 16—40 per
ovary. Fruit dehiscent siliques, linear, latiseptate,
sessile or shortly stipitate; valves leathery, with a
distinct midvein, hairy as leaves, torulose; replum
rounded in cross section, visible; septum complete,
membranous, veinless or with 1—3 longitudinal
veins; style obsolete or distinct and to 2 mm long;
stigma capitate, entire or 2-lobed. Seeds uniseriate.
winged, margined, or wingless, oblong, strongly
flattened; seed coat not mucilaginous when wetted:
cotyledons accumbent or rarely incumbent.

1 . Rhammatophyllum pachyrhizum (Karelin &

Kirilov) (). E. Schulz, Repert. Sp. Nov. Regni
Veg. 33: 190. 1933. Arabis pachyrhiza Karelin
& Kirilov, Bull. Soc. Imp. Naturalistes Moscou
15: 144. 1842. Mitophyllum pachyrhizum (Ka¬
relin & Kirilov) 0. E. Schulz, Not izbl . Bot.
Cart. Berlin-Dahlem 11: 872. 1933. TYPE:
[Kazakhstan. Dzungaria], Bischtas Mis., be¬
tween Ayaguz and Donsyk, 1841. G. S. Karelin
& I. P. Kirilov 1193 (holotype, EE; isotypes, P,
TK).

Distribution. Kazakhstan, Kyrgyzstan.

We have not seen the type collection of Rham¬
matophyllum krascheninnikovii A. N. Vassiljeva
(Vassiljeva, 1969), and based on its similarity in
flower color and other characters to R. pachyrhizum ,
we follow Czerepanov (1995) in reducing the former
to synonymy of the latter.

2. Khnmmntophy Hum frutex Botschantsev &

Vvedensky, Bot. Mater. Gerb. Inst. Bot. Akad.
Nauk Uzbeksk. S.S.R. 13: 9. 1952. TYPE:
[Kazakhstan]. W Betpak Data, vicinity of Ken-
derlik, red clay slopes, 24 May 1936, B. Mi¬
ronov & V. Pazi 353 (holotype, TAK).
Distribution. Endemic to Kazakhstan.

3. Rhummutophy lhim erysimoides (Karelin &

Kirilov) Al-Shehbaz & 0. Appel, comb. nov.
Basionym: Arabis erysimoides Karelin & Kiri¬
lov, Bull. Soc. Imp. Naturalistes Moscou 15:
145. 1842. Prionotrichon erysimoides (Karelin
N Kirilov) Botschantsev & Vvedensky, Bot.
Mater. Gerb. Inst. Bot. Zool. Akad. Nauk Uz-
bekst. S.S.R. 12: 8. 1948. TYPE: [Kazakhstan.
Dzungaria], between Chulak and Ai rivers,
1841, G. S. Karelin & I. P. Kirilov 1192 (ho¬
lotype, EE; isotypes, B, P).

Distribution. Endemic to Kazakhstan.

4. Rhammatophyllum gaudauense (Litvinov)

Al-Shehbaz & O. Appel, comb. nov. Basionym:
Erysimum gaudanense Litvinov, Trudy Bot.
Muz. Imp. Akad. Nauk 1: 33. 1902. Priono¬
trichon gaudanense (Litvinov) Botschantsev,
Novosti Sist. Vyssh. Rast. 3: 125. (1966).
TYPE: [Turkmenistan], “in deelivibus herbosis
montium prope Gaudan,” 28 Apr. 1898, I). I.
Litvinov 518 (holotype, LE; isotype, WU).
Distribution. Turkmenistan and Afghanistan.

5. Rhammatophyllum ghoraimm (K. H. Rechin-

ger) Al-Shehbaz & 0. Appel, comb. nov. Bas¬
ionym: Koeiea ghorana K. H. Rechinger, Anz.
Osterr. Akad. Wiss., Math.-Naturwiss. Kl. 101:
425. 1964. Prionotrichon ghoranum (K. H. Re¬
chinger) Botschantsev, Novosti Sist. Vyssh.
Rast. 24: 97 (1987). TYPE: Afghanistan. Prov.
Ghorat: Shutar Khan Kotal ridge, between Qala
Ahangaren and Qala Sharak, ca. 34°20'N,
64°55'E, ca. 2725 m, 27 July 1962, K. II. Re¬
chinger 18866 (holotype, W; isotype, MO).
Distribution. Endemic to Afghanistan.

6. Rhammatophyllum kamelinii (Botschantsev)

Al-Shehl >az & O. Appel, comb. nov. Basionym:
Prionotrichon kamelinii Botschantsev, Novosti
Sist. Vyssh. Rast. 24: 98. 1987. TYPE: Mon¬
golia. Altay Mts., Uljastyjn-Gol, Bulgan-Gol,
10 July 1984, R. V. Kamelin & S. Darijmaa
331 (holotype, LE; isotype, MO).

Distribution. Endemic to the Altay region of
Mongolia.

7. Rhammalophyllum pseudoparrya (Bot¬

schantsev & Vvedensky) Al-Shehbaz & O. Ap¬
pel, comb. nov. Basionym: Prionotrichon pseu¬
doparrya Botschantsev & Vvedensky, Bot.
Mater. Gerb. Inst. Bot. Zool. Akad. Nauk Uz¬
beks!. S.S.R. 12: 8. 1948. TYPE: [Tajikistan].
Alaisky Range, Karategen, near river Kizil-Su,
above Sarigul, 10 Sep. 1927. /. A. Raikova
1563 (holotype, TAK).

Distribution. Northern Tajikistan and southern
Kyrgyzstan.

8. Rhammatophyllum flexuosum (K. II. Re¬

chinger) Al-Shehbaz & O. Appel, comb. nov.
Basionym: Koeiea flexuosa K. 11. Rechinger,
Anz. Osterr. Akad. Wiss., Math.-Naturwiss. Kl.
101: 426. 1964. Prionotrichon flexuosum (K.
H. Rechinger) Botschantsev, Novosti Sist.
Vyssh. Rast., 24: 99. 1987. TYPE: Afghani¬
stan. Prov. Kabul: Sanglakht, above Jalrez, ca.
34°30'N, 68°32'E, ca. 2400 m, 12 July 1962,
K. H. Rechinger 18027 (holotype, W).

4

Novon

Distribution. Endemic to Afghanistan.

9. Hliaiiimulopliyllum afghanicum (K. H. Re-

chinger) Al-Shehbaz & 0. Appel, comb. nov.
Basionym: Koeiea afghanica K. H. Rechinger,
Anz. Osterr. Akad. Wiss., Math.-Naturwiss. Kl.
91: 60. 1954. Prionotrichon afghanicum (K.
H. Rechinger) Botschantsev, Novosti Sist.
Vyssh. Rast. 3: 125. 1966. TYPE: Afghani¬
stan. Bend-i Amir, 2000 m, 19 Aug. 1948. M.
Koie 2845 (holotype, C; isotype, W).
Distribution. Endemic to Afghanistan.

Kky to the Species oe Rhammatophyllvm

la. Trichomes sulxlendritic, rarely mixed with (ewer

suhmalpighiaeeous ones; fruits (2.5— )3— 4.5 mm
wide; seeds broadly winged all around; Afghan¬
istan . 9. A', afghanicum

lb. Trichomes malpighiaceous or submalpighia-
ceous; fruits 1 — 2 ( — 2 .5) mrn wide; seeds wingless,
margined, or narrowly winged all around, rarely
broadly winged distally.

2a. Trichomes exclusively malpighiaceous with
unbranched 2 rays; cotyledons incumbent;
seeds wingless, rarely obscurely margined.

3a. Petals purple; shrubs ca. 50 cm tall;
leaves 0.5— 2.5 cm X 0.9— 1.2 mm; Ka¬
zakhstan . 2. R. frutex

3b. Petals creamy white; plants subshrubs
10—30 cm tall; leaves (2.5 — )3 — 7 cm X
ca. 0.5 mm; Kazakhstan and Kyrgyzstan

. 1 . /\’. pachyrhizum

2b. Trichomes suhmalpighiaeeous, with minute
lateral branches on the 2 rays; cotyledons
accumbent; seeds winged, rarely margined.

4a. Leaves with a distinct petiole 0.5—2 cm
long; fruits 5-8 cm long; replum straighl
between the seeds in mature fruit.

5a. Petals 10—13 X 5—7 mm; leaves 2—

7 mm wide; petiole 1—2 cm long;

Altay of Mongolia ... 0. A’. kamelinii
5b. Petals 7— H X 3— f mm; leaves to I
mm wide; petiole rarely to I cm
long; Kazakhstan . . 3. R. erysimoides
4b. Leaves sessile; fruits 1.5— 4(— 5) cm long;
replum often constricted between the
seeds in mature fruit.

6a. Gynophore 1—3 mm long; racemes
(lexuous in fruit; sterile shoots often
well developed, with leaves to 2—3
cm long; Afghanistan 8. R. flexuosum
6b. Gynophore obsolete, rarely to 0.7 mm
long; racemes not (lexuous in fruit;
sterile shoots absent, i( present then
with leaves rarely to 1.5 cm long.

7a. Leaves flat; seeds distinctly
winged all around; Afghanistan

. 5. R. ghoranum

7h. Leaves longitudinally plicate;
seeds narrowly margined at
least distally.

8a. Petals pilose outside; gy¬
nophore obsolete; style to

1 mm; Afghanistan and

Turkmenistan .

. 4. R. gaudanense

8b. Petals glabrous outside;
gynophore distinct, to 0.7
mm; style 1-2 mm; Taji¬
kistan and Kyrgyzstan . .
. 7. R. pseudoparrya

Acknowledgments. We are grateful to Tatyana
Shulkina for her help in the translation of Russian
text. We thank Michael (1. Gilbert and Anthony R.
Bracli for their help with the literature.

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Al-Shehbaz, I. A. 1985. The genera of Thelypodieae (Cru-
eiterae; Brassicaceae) in the southeastern United States.

J. Arnold Arbor. 66: 95—1 1 I.

Botschantsev, V. P. 1952. Plantae novae ex Asia Media.
Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk.
S.S.B. 13: 3-20.

- . 1966. De Cruciferis notae eriticae, 5. Novosti

Sisl. Vyssh. Bast. 3: 122-139.

- . 1987. He genere Prionotrichon Botsch. et Vved.

(Cruciferae— Arabideae). Novosti Sist. Vyssh. Bast. 24:
96-1 00.

- & A. I. Vvedensky. 1948. Cruciferae novae ex

Asia Media. Bot. Mater. Gerb. Inst. Bot. Zool. Akad.
Nauk Uzhekst. S.S.B. 12: 3-12.

Czerepanov, S. K. 1995. Vascular Plants of Bussia and
Adjacent States (the former USSB). Cambridge Univ.
Press, Cambridge.

Greene, E. L. 1904. Certain West American Cruciferae.

Lead. Bot. Observ. Grit. I: 81—90.

Hedge, I. C. 1968. Arabideae. In: K. H. Rechinger (edi¬
tor), Flora Iraniea 57; 193—218. Akademische Druck-
u. Verlagsanstalt, Graz.

Koch, M.. J. Bishop & T. Mitchell-Olds. 1999. Molecular
systematies and evolution of Arahidopsis and Arabis. PI.
Biol. I: 529-537.

Nabiev, M. 1972. Botschantzevia Nabiev — Genus novum
Crueiferarum. Novosti Sisl. Vyssh. Bast. 9: 186—187.
Polatschek, A. & K. II. Rechinger. 1968. Erysimum. In:

K. H. Rechinger (editor). Flora Iraniea 57: 285—05.
Akademische Druck- u. Verlagsanstalt. Graz.

Rechinger, K. II. 1954. Cruciferae novae Afghanicae. Anz.
Osterr. Akad. Wiss.. Math.-Naturwiss. Kl. 91: 58—64.

- . 1964. Notizen zur Orient-Flora, 63—69: Neue

Crueiferen aus Afghanistan. Anz. Osterr. Akad. Wiss.,
Math.-Naturwiss. Kl. 101: 122—127.

- . 1968. Matthioleae. In: K. H. Rechinger (editor).

Flora Iraniea 57: 219-250. Akademische Druck- u.
Verlagsanstalt. Graz.

Rollins. B. C. 1993. The Cruciferae of Continental North
America. Stanford Univ. Press, Stanford.

Schulz, (). E. 1933a. Eine neue Cruciferen-Gattung in Mit-
telasien. Notizbl. Bot. Cart. Berlin-Dahlem I I: 872—873.

- . 1933b. fiber verschiedene Crueiferen. Bepert.

Sp. Nov. Begni Veg. 33: 183-191.

- . 1936. Cruciferae. In: A. Engler & K. Prantl (ed¬
itors), Hie naturlichen Pflanzenfamilien, ed. 2, 17B:
227-658. Verlag von Wilhelm Englemann, Leipzig.
Vassiljeva. A. N. 1969. Species novae generum Parrya B.
Br„ Ermania Cham., Rhammatophyllum 0. E. Schulz
ex Asia Media. Bot. Mater. Gerb. Inst. Bot. Akad. Nauk
Kazakhsk. S.S.B. 6: 17-26.

Cardaria, Coronopus, and Stroganowia are United with
Lepidium (Brassicaceae)

Ihsan A. Al-Shehbaz

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

Klaus Mummenhoff

Department of Botany, Faculty of Biology/Chemistry, University of Osnabriick, Barbarastr.

11, D-49069 Osnabriick, Germany

Oliver Appel

Bredkamp 36E, 22589 Hamburg, Germany

ABSTRACT. The genera Cardaria (including / / v-
menophysa), Coronopus, and Stroganowia are hard¬
ly distinct morphologically from the larger h'pi-
dium, and t he distinctions of all four genera are
based solely on a few fruit characters of doubtful
value. Recent molecular data strongly suggest that
the genera Coronopus and Stroganowia are poly-
phyletic and, together with Cardaria, are nested
within Lepidium. Therefore, they are herein for¬
mally united with Lepidium, the earliest published
generic name in this group. The new names L. ap-
pelianum, L. botschantsevianum , L. buschianum, L.
karelinianum, and L. mummenhoffianum are pro¬
posed because the transfer of their replaced names
to Lepidium would create later homonyms. The fol¬
lowing 20 new combinations are proposed: L.
brachyotum, L. cardiophyllum, L. englerianum, L.
lepidioides, L. litwinowii, L. longifolium, L. minor,
L. navasii, L. paniculatum , L. rhytidocarpum, L. ro-
bustum, L. sagittatum, L. saravschanicum, L. ser¬
ration, L. tianschanicurn, L. tiehmii, L. tolmaczovii,
L. trautvetteri, L. violaceum, and L. zambiensis.

Key words: Brassicaceae, Cardaria, Coronopus,
II ymenophysa, Lepidium, Stroganowia.

During work by two of us (Al-Shehbaz and Ap¬
pel) on the forthcoming account of the Brassicaceae
(Cruciferae) for K. Kubitzki's Families and Genera
of Vascular Plants, and on the basis of molecular
studies on the genus Lepidium L. and relatives,
such as Cardaria Desvaux, Coronopus Zinn, and
Stroganowia Karelin & Kirilov (Bowman et ah,
1999; Bruggemann, 2000; Mummenhoff, 1995;
Mummenhoff et ah. in press; Mummenhoff, unpub¬
lished), it has become amply evident that only one
genus is involved, and that nomenclatural adjust¬

ments are needed to make the names available lor
several floristic works in progress.

Iwpidium is a cosmopolitan genus of about 175
species distributed on all continents except Ant¬
arctica (Al-Shehbaz, 1986). It is one of the most
natural and readily distinguished of all genera in
the family. As presently delimited, all species have
angustiseptate fruits with two subapical ovules (one
in each locule), and the trichomes (when present)
are always simple. The flowers in more than half of
the species have just two stamens, and the petals
in most of those are rudimentary or lacking. In the
remaining species of Lepidium, the flowers are al¬
ways petaliferous and have either six or four sta¬
mens (Thellung, 1906; Al-Shehbaz, 1986). For the
reduction of stamen number in Lepidium, the in¬
terested reader should consult Bowman and Smyth
(1998) and Bowman et al. (1999).

The genera Cardaria, Coronopus, and Stroga¬
nowia resemble Lepidium in having similar fruit
structure, two ovules per ovary, and simple tri¬
chomes. Most of them have six stamens, but a few
species of Coronopus have two stamens, just like
the majority of Lepidium. As shown below, the prin¬
cipal characters used to distinguish these genera
from Lepidium are based solely on minor differenc¬
es in fruit morphology. However, recent molecular
studies (Mummenhoff et al., in press) have clearly
shown that the genera Cardaria, Coronopus, and
Stroganowia are nested within Lepidium, that Co¬
ronopus and Stroganowia are definitely polyphylet-
ic, and that Lepidium is paraphyletic.

Cardaria

Cardaria is said to differ from Lepidium by hav¬
ing indehiscent instead of dehiscent fruits, and both
genera were recognized in most of the major floris-

Novon 12; 5-11. 2002.

6

Novon

tic or monographic works, including those by
Schulz (1936), Hedge (1968), Hewson (1982), Al-
Shehbaz (1986), Schultze-Motel (1986). Hollins
(1993), and Tulin et al. (1993). However, as indi¬
cated by Thellung (1906), there are some species
of Lepidium with tardily dehiscent or even indehis-
cent fruits. In fact, some authors (e.g., Thellung,
1906; Busch, 1939a; Rich, 1991) reduced Cardaria
to synonymy of Lepidium, though Busch (1939a,
1939b) also recognized Hymenophysa C. A. Meyer
as a related genus with two species. By contrast,
Schulz (1936) recognized Cardaria as a monotypie
genus that he placed with Lepidium in the tribe
Lepidieae and treated Hymenophysa as a monotypie
genus that he placed in the tribe Euelidieae. The
single character that Schulz (1936) used to separate
Cardaria from Hymenophysa (and the two tribes in
which he placed them) is the presence of angusti-
septate (flattened at a right angle to the septum) vs.
inflated fruits. The example of Cardaria-Hymeno-
physa shows that heavy reliance on the type of com¬
pression of fruit led to the artificial delimitation of
genera and tribes. For another example, the vast
majority ol the approximately 170-190 species of
Alyssum I.. have latiseptate fruits (flattened parallel
to tin1 septum), and only a few species (e.g., A. tur-
gidum T. R. Dudley and A. glohosum Grossheim)
have distinctly inflated fruits (Dudley, 1964; Ave-
lisian, 1980). Alyssum glohosum was placed in the
monotypie genus Takhtajaniella V. E. Avetisian
(Avetisian, 1980), but we believe that it is perfectly
at home in Alyssum.

Molecular data (Mummenhoff, 1995; Briigge-
mann, 2000; Mummenhoff et al., in press) clearly
demonstrated that Cardaria is nested within Lepi-
dium, and that it is most closely related to L. cam-
pestre E. and its relatives, which Thellung (1906)
assigned to Lepidium sect. Lepia (Desvaux) DC. The
cpDNA data (Mummenhoff et al.. in press) indicate
monophyly of C. dralxi (L.) Desvaux and C. pubes-
cens (C. A. Meyer) Jarmolenko, whereas the ITS
data (Briiggemann, 2000) suggest polyphyly of Car¬
daria because C. pubescens appears to be more
closely related to members of Lepidium sect. Lepia
than does C. draba. However, both the cpDNA and
ITS data show that Cardaria is nested well within
Lepidium next to section Lepia. It is interesting to note
that the type species of Cardaria was initially de¬
scribed by Linnaeus (1753) as Lepidium draba L.

There is disagreement as to the number of spe¬
cies recognized in Cardaria. Mulligan and Frank-
ton (1962) and Czerepanov (1995) recognized as
many as five and six species, respectively. Bv con¬
trast. other authors (e.g.. Hedge, 1968; Al-Shehbaz,
1986) recognized two, while Rollins (1993) recog¬

nized three. We believe that only two species merit
recognition, and both should be placed in Lepi¬
dium. These are L. draba, which consists of sub¬
species draba and chalapense (L.) Thellung, and L.
appelianum Al-Shehbaz, a new' name proposed
herein because the transfer of Hymenophysa pubes¬
cens C. A. Meyer to Lepidium would create a later
homonym of the South American L. pubescens Des¬
vaux.

CoROMm .s

Coronopus consists of 10 species native to South
America, Africa, Southwest Asia, and adjacent Eu¬
rope (Al-Shehbaz, 1986). It differs from Lepidium
primarily by having didvmous fruits with thickened,
reticulate, rugose, to tuberculate (rarely smooth),
indehiscent fruit valves. In three species (C. navasii
Ban. C. squamatus (Forsskal) Ascherson, and C.
violaceus (Munby) Kuntze) the fruits are not didy-
mous while in the remaining seven ( C. didymus (L.)
Smith, C. integrifolius (DC.) Sprengel, C. niloticus
(Delile) Sprengel, C. lepidioides (Cosson & Durieu)
Kuntze, C. serratus (Poiret) Desvaux, C. rhytidocar-
pus (Hooker) Macloskie, and C. zambiensis Jonsell)
they are distinctly didymous. Coronopus wrightii H.
Hara (Japan, Taiwan) was reduced by Cheo et al.
(2001) to synonymy of C. integrifolius (herein as L.
englerianum). The valve orifice that faces the re¬
plum is usually smaller than the seed and, there¬
fore. the seeds are not readily released, and these
valves act as the dispersal unit. In Lepidium the
fruit is not didymous, anil the valves are generally
thin and with orifices often wider than the seeds.
Therefore, the seeds are easily released from the
valves. However, in the southern Argentinian and
Chilean L. pseudodidymum Thellung, the fruits
somewhat approach C. didymus in their reticulation
and thickness. In fact, L. pseudodidymum was treat¬
ed by Muschler (1908) as C. didymus subsp. aus¬
tralis (J. D. Hooker) Muschler and was later re¬
described by Boelcke (1975) as Coronopus
leptocarpus Boelcke. This shows that Lepidium and
Coronopus can hardly be delimited on morpholog¬
ical grounds. In our opinion, features of the fruit
valves in Coronopus are adaptations for dispersal
and are insignificant in the delimitation of the ge¬
nus. fhe development of thick, reticulate, rugose,
or tuberculate fruit walls apparently evolved inde¬
pendently within Lepidium (including Coronopus).
Molecular data by one of us (Mummenhoff, unpub¬
lished) clearly show that Coronopus is poly phy let ic
and nested within Lepidium.

Although molecular studies have not yet been
done on the monotypie Delpinophytum Spegazzini,

Volume 12, Number 1
2002

Al-Shehbaz et al.

Genera Transferred to Lepidium

7

it is very likely that D. patagonicum (Spegazzini)
Spegazzini is only a dwarf, pulvinate species o{ Lep¬
idium. The species was subsequently treated by
Muschler (1908) as Coronopus patagonicus (Spe¬
gazzini) Muschler. Both C. didymus and C. squa-
matus were originally described as /.. didymum L.
and L. squamatum Forsskal, respectively. Corono¬
pus niloticus is known in Lepidium as L. niloticum
(Delile) Sieber ex Steudel. For the remaining seven
species of Coronopus , new combinations are pro¬
posed herein to accommodate their transfer to Lep¬
idium.

Stroganowia

Both Pavlov (1933, 1939) and Botschantsev
(1984) provided brief synopses on Stroganowia,
and the latter author recognized 21 species in the
genus. The characters used to distinguish Stroga¬
nowia from Lepidium are the perennial habit and
slightly angustiseptate or nearly quadrangular
fruits. However, these features are tenuous at best,
and many species of Lepidium (e.g., L. latifolium
L. and relatives) resemble Stroganowia in habit.
I'he degree of the fruit flattening is not a reliable
feature either, and there are no other differences
between the two genera. In fact, some species rec¬
ognized by Botschantsev (1984) as Stroganowia
(e.g., S. affghana, S. subalpina) were treated by
Thellung in Lepidium. who (1906: 157) suggested
that Stroganowia is best treated as a section of Lep¬
idium. Molecular data available so far for seven
species (Mummenhoff, unpublished) show that the
Old World Stroganowia forms separate groups nest¬
ed within Lepidium. What was explicitly mentioned
by Botschantsev (1984) about the dubious merit of
Stroganowia has now been substantiated by Mum-
menhofF s molecular studies. As for ,S. tiehmii Rol¬
lins. Price (pers. comm.) indicated that the species
falls within the American species of Lepidium
based on molecular studies. Therefore, Stroganowia
is polyphvletie, and the alleged relationship of S.
tiehmii with its nearest disjunct relatives in Central
Asia (Rollins, 1982) has not been supported by mo¬
lecular data.

All of Botschantsev s (1984) 21 species of Stro¬
ganowia merit recognition as species of Lepidium.
Three species, S. affghana (Boissier) Pavlov, S. bu-
pleuroides (k. H. Rechinger) Botschantsev, anil S.
subalpina (Komarov) Thellung ex Pavlov, were orig¬
inally described in Lepidium , and their retention in
that genus poses no nomenclatural problems. The
transfer to Lepidium of .S', angustifolia Botschantsev
& Vvedensky, S. intermedia Karelin & Kirilov, S.
persica N. Busch, and S. rubtzovii Botschantsev

would create later homonyms, and the following
new names are proposed herein to accommodate
them in Lepidium: L. botschantsevianum Al-Sheh-
baz, L. harelinianum Al-Shehbaz, L. buschianum
Al-Shehbaz, and L. mummenhoffianum Al-Sheh¬
baz, respectively. Upon its transfer to Lepidium, S.
puberula Kitamura would become an illegitimate
later homonym of L. puberulum Bunge. Therefore,
S. puberula should be recognized in Lepidium as L.
altissimum K. H. Rechinger, which is the earliest
legitimate name for the species in Lepidium. New
combinations in Lepidium are proposed herein to
accommodate the remaining 13 species of Stroga¬
nowia: S. brachyota Karelin & Kirilov, S. cardi-
ophylla Pavlov, S. litwinowii Lipsky, S. longifolia
(Boissier) Botschantsev & Vvedensky (originally
described as Heldreichia longifolia Boissier), S. mi¬
nor Botschantsev & Vvedensky, S. paniculata Re¬
gel & Schmalhausen, S. robusta Pavlov, S. sagittata
Karelin & Kirilov, S. saravschanica Bulgakova, S.
tianschanica Botschantsev & Vvedensky, .S’, tiehmii
Rollins, S. tolmaczovii Junussov, and S. trautvetteri
Botschantsev.

Lepidium L., Sp. PI. 2: 643. 1753. TYPE: Lepi¬
dium latifolium L. (lectotype, designated by
Britton & Brown, 1913).

Coronopus Zinn, Cat. PI. Hort. Gott. 325. 1757, nom. cons.

TYPE: Coronopus squamatus (Forsskiil) Asoherson.
Cardaria Desvaux, J. Bot. Agric. 3: 163. 1815. TYPE:

Cardaria draba (L.) Desvaux.

Stroganowia Karelin & Kirilov, Bull. Soc. Imp. Natural-
istes Moscou 14: 386. 1841. TYPE: Stroganowia
sagittata Karelin & Kirilov (lectotype, designated by
Botschantsev, 1984).

Although Greuter et al. (1993, 2000) listed the
type species of Coronopus as C. ruellii Allioni.
which is based on Cochlearia coronopus I ... both of
these specific epithets are synonyms of Lepidium
squamatum Forrskal. Cochlearia coronopus and Co¬
ronopus ruellii are correctly listed as synonyms of
L. squamatum by Hedge (1968) and Greuter et al.
(1986), respectively.

Lepidium appelianum Al-Shehbaz, nom. nov. Re¬
placed name: Hymenophysa pubescens C. A.
Meyer, in Ledebour, Icon. PI. 2: 20. 1830. non
Lepidium pubescens Desvaux, J. Bot. Agric. 3:
180. 1815, nec L. pubescens Tineo. Cat. PI.
Hort. Panorm. 150. 1827. TYPE: [Kazakhs¬
tan.] “Locis humidis subsalsis deserti Soon-
goro-kirghisici orientalis versus montes Ar-
kaul.” 14 May 1826, C. A. Meyer s.n.
(lectotype, designated here, I ,E).

The choice of C. A. Meyer s.n. as the lectotype

8

Novon

was based on the fact that L. appelianum is a new
name replacing the later homonym L. pubescens,
and this collection was among those cited in the
original publication.

The species is named in honor of Oliver Appel,
one of the authors of the present paper and an ex¬
pert on the Brassieaceae.

Lepidium botschuntsevianum Al-Shehbaz. nom.
nov. Replaced name: Stroganowia angustifolia
Botschantsev & Vvedensky, Rot. Mater. Gerh.
Inst. Rot. Zool. Akad. Nauk Uzhekst. S.S.R.
12: II. 1948, non Lepidium angustifolium
Rushy, Deser. S. Amer. PI. 23. 1920. TYPE:
| Uzbekistan. | Samarkand: Zarafshon, Katta
Gorgansky, Ml. Actao, near village Shamani.
24 May 1925, M. G. Popov 349 (holotype,
TASH).

The new name commemorates Victor Petrovich
Botschantsev (1910—1990), an outstanding expert
on the Brassieaceae of the former Soviet Union.

Lepidium hraeliyotuin (Karelin & Kirilov) Al-
Shehhaz, comb. nov. Basionym: Stroganowia
brachyota Karelin & Kirilov, Bull. Soc. Imp.
Naturalistes Moscou 14: 387. 1841. TYPE: “In
montosis deserti Soongoro-Kirghisiei prope
Ajagus," June 1840, G. S. Karelin & I. P. Kir¬
ilov s.n. (holotype, LE).

Lepidium hiiscliiuniim Al-Shehhaz, nom. nov.
Replaced name: Stroganowia persica IN.
Busch, Zliurn. Russk. Rot. Obshch. Akad.
Nauk S.S.S.R. II: 225. 1926, non Lepidium
persicum Boissier, Ann. Sci. Nat. ser. 2, 17:
196. 1842. TY PE: N Iran. Tabris (Atropatan-
ia): Mt. Scher-Dara, between Sofian and Se-
jvan, 1430-1500 m. 18 June 1924, A. Gross-
heirn s.n. (holotype, LE).

The new name commemorates Nicolai Adolfo-
witscli Busch (1869—1941), an outstanding expert
of the Brassieaceae of Asia and the author of most
of the accounts of Brassieaceae for the Flora of the
former Soviet Union (see Busch, 1939a, 1939b).

Lepidiuni cardiophyllum (Pavlov) Al-Shehbaz,
comb. nov. Basionym: Stroganowia cardiophyl-
la Pavlov, Rot. Zhurn. S.S.S.R. 18: 364. 1933.
TYPE: [Turkmenistan.] “Turkestania Ross.,
prov. Syr-Darja, montes Maschat-tau (prae-
montorium Alatau Talassici), in decliviis glar-
eosis pratensibus ail trajeetum Dau-baba,
1200 m,” 31 Aug. 1931, N. V. Pavlov 1226
(holotype, MW: isotype, LE).

Lepidium eiiglerinnuiii (Muschler) Al-Shehbaz.
comb. nov. Basionym: Coronopus englerianus
Muschler. Rot. Jahrb. Syst. 41: 139. 1908.
TYPE: Mozambique. Mouth of Zambezi River,
Peters s.n. (holotype, B).

The earliest name for the species is Senebiera
integrifolia I)C. (Mem. Soc. Hist. Nat. Paris I: 144.
1799), but the transfer of its specific epithet to Isp-
idium would create a later homonym of L. integri-
folium Nuttall ex Torrey A. Gray. The second
earliest name for the species is Senebiera linoides
DC., but the transfer of its specific epithet to Lep-
idium would create a later homonym of L. linoides
Thunberg. According to Marais (1970), the last spe¬
cies is a synonym of Lepidiuni divaricatum Aiton.
Jonsell (1974) was the first to reduce Coronopus
englerianus to synonymy of S. integrifolia , and the
epithet “ englerianus " is the earliest available name
that can be used for the species in Lepidium.

Lepidium karelinianuin Al-Shehbaz, nom. nov.
Replaced name: Stroganowia intermedia Ka¬
relin & Kirilov, Bull. Soc. Imp. Naturalistes
Moscou 15: 162. 1842, non Lepidium inter¬
medium A. Richard, Tent. FI. Abyss. 1: 21.
1847. TYPE: “In montosis apricis Alatau inter
fluvios Baskan et Sarchan,” July 1841. G. S.
Karelin A /. P. Kirilov s.n. (lectotype, desig¬
nated by Pavlov (1933), LE; isolectotype,
MW).

The new name honors Grigorij Siliovitsch Kare¬
lin (1801—1872), one of the famous botanical ex¬
plorers of central Asia.

Lepidium lepidioides (Gosson & Durieu) Al-
Shehbaz. comb. nov. Basionym: Senebiera lep¬
idioides Cosson & Durieu, Bull. Soc. Bot.
France 2: 245. 1855. TYPE: Algeria. “In di-
tione Mzab prope Gucrrara." 3 Nov. 1854, V.
G. Rehoud s.n. (holotype, P).

Lepidium litwinowii (Lipsky) Al-Shehbaz, comb,
nov. Basionym: Stroganowia litwinowii Lipsky,
Trudy Imp. S.-IVterburgsk. Bot. Sada 26: 121.
1910. TYPE: Turkmenistan. Mountains near
Gaudan, 30 May 1898. D. Litwinow 625 (ho¬
lotype. LE).

Lepidium longifolium (Boissier) Al-Shehbaz.
comb. nov. Basionym: Heldreichia longifolia
Boissier, Ann. Sci. Nat. ser. 2. 17: 187. 1842.
TYPE: Iran. Mount Zerdakou [Zard Kuh],
Aucher 320 (holotype, G).

Hedge (1968) had correctly questioned the in-

Volume 12, Number 1
2002

Al-Shehbaz et al.

Genera Transferred to Lepidium

9

elusion of the species in Heldreichia Boissier be¬
cause of its habit, non-fleshy leaves, and toothless
filaments, lie indicated that the species should ei¬
ther be placed in a monotypic genus or in Stroga-
nowia. As indicated by Botschantsev and Vveden¬
sky (1948), the species is morphologically very
close to S. angustifolia (herein as Lepidium bot-
schantsevianum).

Lepidium minor (Botschantsev & Vvedensky) Al-
Shehbaz, comb. nov. Basionym: Stroganowia
minor Botschantsev & Vvedensky, Bot. Mater.
Gerb. Inst. Bot. Zool. Akad. Nauk Uzbekst.
S.S.R. 3: 19. 1941. TYPE: [Tajikistan.] Pamir:

| up B iver Yakabag Darya, around village Tash-
korgan], 18 June 1936, Botschantsev & Butkov
8 (holotype, FASH # 161584).

It is quite possible that the collection was actu¬
ally made from near Taxkorgan, a town in the south¬
western Xinjiang Autonomous region at the Taji-
kistan-China border, as such town no longer exists
in Tajikistan.

Lepidium mummenhoffianum Al-Shehbaz, nom.
nov. Replaced name: Stroganowia rubtzovii
Botschantsev, Novosti Sist. Vyssh. Rast. 21:
79. 1984, non Lepidium rubtzovii Vassilczenko,
Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Ka-
zakhsk. S.S.R. 4: 40. 1966. TYPE: “Juguni
Alatau Dzhungaricus, distr. inter pagos Sar-
kand et Kopal, locus Bel-Bulak.” 20 July
1934, N. /. Rubtzov s.n. (holotype, EE).

The species is named in honor of Klaus Mum-
menhoff, one of the authors of this paper, in rec¬
ognition for his extensive molecular work on Lepi¬
dium and related genera.

Lepidium navasii (Pau) Al-Shehbaz, comb. nov.
Basionym: Coronopus navasii Pau, Butl. Inst.
Catalana Hist. Nat. 22: 31. 1922. TYPE:
Spain. Sierra de Gador, 2000 m, B. Navas s.n.
(holotype, MA not seen).

Lepidium paniculatum (Regel & Schmalhausen)
Al-Shehbaz, comb. nov. Basionym: Stroganow¬
ia paniculata Regel & Schmalhausen, Trudy
Imp. S.-Peterburgsk. Bot. Sada 5: 242. 1877.
TYPE: Raskrask, “in valle fluvii” Tschirtschik,
8000-9000 ft., Aug. 1876, A. Regel s.n. (lec-
totype, designated by Pavlov (1933), EE).

Lepidium rhytidocarpum (Hooker) Al-Shehbaz,
comb. nov. Basionym: Senebiera rhytidocarpa
Hooker, London ,1. Bot. 2: 506. 1843. TYPE:
Patagonia, Tweede s.n. (holotype, K).

Lepidium robustum (Pavlov) Al-Shehbaz, comb,
nov. Basionym: Stroganowia robusta Pavlov,
Bot. Zhurn. S.S.S.R. 18: 367. 1933. TYPE:
Turkmenistan. Mt. Boroldaj-tau, summit of Bu-
kuj-tau, 1600 m, 27 June 1931, N. Pavlov 397
(holotype, MW; isotype, EE).

Lepidium sagitlatum (Karelin & Kirilov) Al-
Shehbaz, comb. nov. Basionym: Stroganowia
sagittata Karelin & Kirilov, Bull. Soc. Imp.
Naturalistes Moscou 14: 387. 1841. TYPE: “In
arenosis ad radicem montium Tarbagati prae-
sertim ad torrentes Dschany-bek et Terekty,”
May 1840, G. S. Karelin & /. P. Kirilov s.n.
(holotype, EE).

Lepidium saravschanicum (Bulgakova) Al-Sheh-
baz. comb. nov. Basionym: Stroganowia sar-
avschanica Bulgakova, Bot. Mater. Gerb. Inst.
Bot. Akad. Nauk Uzbeks. S.S.R. 20: 8. 1982.
TYPE: Uzbekistan. Jugum Seravschanicum,
above Sarykol, vicinity of Tillja-Tikan, ca.
1000 m, 13 June 1980, L. L. Bulgakova s.n.
(holotype, TASH; isotype, MO).

Lepidium serratum (Poiret) Al-Shehbaz, comb,
nov. Basionym: Senebiera serrata Poiret, in Ea-
marck, Encycl. 7: 76. 1806. TYPE: [Uruguay].
Montevideo, Commerson s.n. (holotype, P-JU).

Lepidium serratum is unrelated to and should not

be confused wi th the H awaiian L. serra H. Mann.

Lepidium tianschanicum (Botschantsev & Vved¬
ensky) Al-Shehbaz, comb. nov. Basionym:
Stroganowia tianschanica Botschantsev &
Vvedensky, Bot. Mater. Gerb. Inst. Bot. Zool.
Akad. Nauk Uzbekst. S.S.R. 12: 12. 1948.
TYPE: [Kyrgyzstan.] W Tian Shan, Maidantal-
sky, Chotan-gutsai, 24 July 1940, V. Makar-
chuk s.n. (holotype, TASH).

Lepidium tiehmii (Rollins) Al-Shehbaz, comb,
nov. Basionym: Stroganowia tiehmii Rollins,
Syst. Bot. 7: 215. 1982. TYPE: U.S.A. Nevada:
Lyon County, Virginia Range, SE of Talapoosa
Peak, T19N, R24E, sect. 34, 5900 ft., 1 June
1980, A. Tiehtn 5783, F. Almeda & M. Wil¬
liams (holotype, Gil; isotype, MO).

Lepidium tolmaezovii (Junussov) Al-Shehbaz,
comb. nov. Basionym: Stroganowia tolmaezovii
Junussov, Dokl. Akad. Nauk Tadzh. SSR 18(2):
62. 1975. TYPE: W Tajikistan. Karatau ridge,
4 km N of Mt. Chodzha-Maston, 17 May 1960,
A. Meczislavskij & V. N. Zavedeev 675 (holo¬
type, LE).

10

Novon

Ecpiilium trautveiteri (Botschantsev) Al-Sheli-
baz, comb. nov. Basionym: Stroganowia traut-
vetteri Botschantsev, in Komarov, FI. URSS 8:
652. 1939. TYPE: [Kazakhstan.] Ad lacnm
Balchasch, Betpak-dala, 13 June 1843. A.
Schrenk 399 (holotype, EE).

Gcpidiiiiu violuccum (Munby) Al-Shchbaz. comb,
nov. Basionym: Senebiera violacea Munby,
Bull. Soc. Bot. France 2: 282. 1855. TYPE:
Algeria. Dhaya-Baalis, Ain-Turck, near Oran,
G. Munby s.n. (holotype, K?).

Lcpidiiiin zanibiensis (Jonscll) Al-Shchbaz,
comb. nov. Basionym: Coronopus zanibiensis
Jonscll, Bot. Not. 127: 1 16. 1 974. TYPE: Zam¬
bia. Kalabo District: Liuwa Plain, Paramount
Chiefs Game Reserve, ca. 45 km N of Kalabo,
14 Sep. 1959, Drummond & Cookson 6/53
(holotype, BM; isotypes, BR, COI. I .ISC.
SRGH; see Jonscll. 1974).

Acknowledgments. We arc grateful to Tatyana
Shulkina and Eidiya Toropova for their help in the
translation of Russian literature and herbarium la¬
bels. We thank Michael G. Gilbert and Anthony
R. Brach for providing photocopies of some of the
original literature. We arc indebted to Neil llar-
riman. an anonymous reviewer, and Victoria C.
Hollowed for tbeir critical reviews of the manu¬
script.

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moulin. T. S. Filgueiras, I). H. Nieolson, P. C. Silva,
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species from tropical Africa. Bot. Not. 127: I 13—1 18.

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Slavii, Holmiae [Stockholm].

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1411-1425.

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Al-Shehbaz et al.

Genera Transferred to Lepidium

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Iwo New Species of Nanuza (Velloziaceae) from Brazil

Ruy Jose Vdlka Alves

Departamento de Botanica, Herbario, Museu Nacional, Quinta da Boa Vista s.n.,
Sao Cristovao, Rio de Janeiro, RJ, Brazil

Austkact. Nanuza almeidae Alves an<l N. luetz-
elburgii Alves are described as new species of the
previously monotypie Nanuza L. 15. Smith & Ay-
ensu (Velloziaceae). Nanuza almeidae is distin¬
guished (rom Nanuza plicata and N. luetzelburgii
mainly by leaf blades basally narrowed into a pseu¬
dopetiole, linear-lanceolate tepals, triangular fila¬
ments with a flattened base, and presence of lab-
dane-type diterpenes. Nanuza luetzelburgii is
distinguished from both other species by rounded
edges on the trigonous caudex, lamina with furrows
with abundant, sturdy hairs, vestigial adaxial fur¬
rows, and serrulate sheath margins, lacking the fil¬
aments typical of the closely related N. plicata and
N. almeidae, and by presence of heptadeeenie acid.
Xerophytolic acid was found in Nanuza plicata but
not in both new species.

Rk.SUMO. Nanuza almeidae Alves e N. luetzelbur¬
gii Alves sao descritas como especies novas do ge-
nero ate entao considerado monotfpico, com apenas
Nanuza plicata (Martins) L 15. Smith & Ayensu
(Velloziaceae). Nanuza almeidae se distingue de
Nanuza plicata e N. luetzelburgii pelas tepalas lin¬
ear— lantieoladas, limbo foliar na base estreitado em
pseudopecfolo, mais largo e eurto, adaxialmente
glabrescente, filetes triangulares com base achatada
e pela presenya de diterpenos do tipo labdano. Na¬
nuza luetzelburgii se distingue de N. plicata e N.
almeidae pelas margens arredondadas do caudex
trfgono, preenchimento dos sulcos no limbo por
abundantes pelos rigidos, sulcos vesligiais na face
adaxial, Iminhas foliares com margens serreadas,
sem as cerdas caracteristicas de N. plicata e N.
almeidae, bem como pela presenya de acido hep-
tadec enico. Nanuza plicata se distingue das espe¬
cies novas pela presenya de acido xerofitolico.

Key words: Brazil, heptadeeenie acid, leaf anat¬
omy, Nanuza, tetracosanoic acid, Velloziaceae.

I lie Velloziaceae are a predominantly Neotropi¬
cal family with a major center of diversity in Bra¬
zil s savannic vegetation (campo rupestre, cerrado.
and caatinga). Many species are known only from
their holotypes, duplicates are rare in herbaria, and
many populations are narrowly endemic. A few spe-

Novon 12: 12-17. 2002.

t ies, such as Nanuza plicata, are apparently more
widespread, occurring in several populations dis¬
persed as a mosaic throughout their ranges. Though
the populations of Nanuza are reported by some
collectors as dense and common, the genus is poor¬
ly represented in herbaria, with the exception of
SPF (over 50 specimens) and BB (ca. 20). The ma¬
terial of Nanuza sent by VIES (Herbario. Museu de
Biologia Mello Leitao, Santa Teresa, State of Hspi'r-
ito Santo) got lost in the mail during a flood, and
no material (even under earlier synonyms) was lo¬
cated in (ill A and HB.

The plicate leaves and trigonous caudex led Mar¬
lins to describe Vellozia plicata (Martins, 1823)
based on a plant lie had collected in 1810 in Bahia.
The species was subsequently transferred to Xero-
phyta by Sprengel (1827). A spec imen collected in
1820 by Bold in Minas Gerais was described as
Vellozia triquetra Bold (1827). Baker (1875) trans¬
ferred Pohl s species to Xerophyta. file genus Na¬
nuza was segregated from Vellozia Vandelli by
Smith and Ayensu (1976), based on Vellozia plicata
Martins (Martins. 1823). Smith and Ayensu based
the new genus on plicate leaves with more deeply
furrowed adaxial surfaces, trigonous caudex, and
pollen dispersed in single grains, as opposed to Vel¬
lozia, which is characterized by terete leaves with
furrows restricted to the abaxial surface and pollen
in tetrads. The originally proposed differential
characters still hold true for Nanuza as circum¬
scribed here, with the exception of the relative
depth of the furrows on the lamina, which are deep¬
er on the abaxial side in N. luetzelburgii and N.
almeidae.

A further distinctive character of Nanuza is the
neatly tristiehous leaf set (it is spiral in Vellozia).
Though Vellozia Candida Mikan has a slightly tri¬
gonous stem (Alves, 1994) hidden by a layer of
sheaths forming a cylindrical caudex, and tristi¬
ehous leaf rosettes, the clearly trigonous caudex
distinctive of Nanuza conceals a cylindrical stem.
A distinct pseudopetiole is present in Nanuza al¬
meidae, and I have not found a reference to one in
other Velloziaceae. Characters such as leaf indu-
ment, anatomy, and chemical constituents of epi-

Volume 12, Number 1
2002

Alves

Nanuza from Brazil

13

cuticular wax are believed to vary more in wide¬
spread speeies of Velloziaceae (Mello-Silva. 1990a,
1995b, 2000), although it seems early to consider
this a rule. Chemical characters were found useful
in delimitation of taxa (genera) by Salatino et al.

(1989).

A recent phylogenetic study using chloroplast
l)NA (Salatino, 1999) supports the distinct position
of Nanuza. still considering it a monotypic genus
of Nanuza plicata (Martius) L. B. Smith & Ayensu.
Though collections are relatively infrequent, the
studied material indicates that the variability in
Nanuza can be partly accounted for by segregation
of two species herein described as new. in which
morphological, anatomical, and chemical differenc¬
es are consistent with known geographical distri¬
bution. It is probable that, with more collections,
additional species will be described from among
the specimens left for now within Nanuza plicata.

The leaves of all species known so far are con-
duplicate-plicate and not simply plicate as stated
in the description of the genus by Smith and Ay¬
ensu (1976). Furthermore, the furrows resulting
from the plication in the lamina of all examined
Nanuza plicata are deeper on the abaxial side, and
not the adaxial as postulated by Smith and Ayensn
( 1976). This is also true for Nanuza almeidae, while
the inverse occurs in N. luetzelburgii. The plication
is most evident in dried leaves of N. plicata (Fig.
1M. N), and least perceptible in N. luetzelburgii
(Fig. IK, L). The leaf blades vary in dimensions,
indument, and presence of a pseudopetiole. Leaf
sheaths can have filamentous (Nanuza plicata and
N. almeidae) or serrulate (N. luetzelburgii) margins.

Flowers are known from a single specimen of
both Nanuza almeidae and N. luetzelburgii and
from less than 20% of the N. plicata vouchers in
the consulted herbaria. All species typically have
flowers with white tepals, though purple stripes
were reported in buds ol one Nanuza plicata pop¬
ulation ( G . E. Valente 148 & A. Azevedo, SPF). An
apical excavation was found in the hypanthium of
all examined herbarium specimens of both new
species, though Mello-Silva (1995a) observed a hy¬
panthium entirely filled by the ovary on material
from Pico das Almas in the State of Bahia.

fhe distinguishing characters of both new spe¬
cies herein are based essentially on the same mor¬
phological character set used to circumscribe other
species of Velloziaceae in recent descriptions (Mel¬
lo-Silva. 1990b, 1992). Anatomical differences in
the transversal sections of leaf lamina and chemical
constituents of selected specimens provided valu¬
able additional characters for delimitation of the

two new species and their distinction from Nanuza
plicata.

Nanuza almeidae Alves, sp. nov. TYPE: Brazil.
Rio de Janeiro: Municfpio de Mage, Distrito
de Piabeta, Fragoso/Tocaia, encosta rochosade
pedra proxima a base do Morro do Urubu, so-
bre a roeha em local ensolarado, 500 m s. m.,
tepalas brancas, 12 Nov. 2000 (fl. fr). Paulo
Roberto Almeida de Jesus s.n. (holotype, R).
Figure 1 A — J .

llaec species N. plicata (Martius) I.. It. Smith A Ay¬
ensu, tepalis lineari-lanceolatis, filamentis triarigularibus
hasibus applanatis, foliis latioribus, adaxialiter nervis
ecristatis ad basim in pseudopetiolum angustatis atque
seminum testa profunde foveolata statim diagnoscitur.

Heliophyllous shrubs up to 150 cm tall; caudex
basally cylindric, thickened by adventive root pen¬
etration, branching along the entire length, upper
half and branches trigonous with rounded wedges,
consisting externally of 3 series of leaf sheaths and
ending in tristichous, apical leaf rosettes. Apical
leaves in each rosette up to twice the size of those
at the base; leaves deciduous, light green, glabres-
cent, parchment-like; lamina linear-lanceolate, 8—
9 X 90—95 mm, conduplicate-plicate, abaxial fur¬
rows slightly deeper than adaxial, margins serrulate
with the teeth curved toward the apex, base nar¬
rowed to 1—1.5 mm wide, conduplicate, with a
curved pseudopetiole furrowed only on the abaxial
side; leaf sheaths triangular, conduplicate, 20 mm
long, base (when flattened) 5 mm wide, margins of
apical half emitting long, vinaceous fibers that in¬
terlace with those of adjacent sheaths; inflorescence
terminal, subtended by 2 linear-triangular, basally
sheathing bracts equaling the tepals and later, by
elongation, the apex of the capsule; flower single in
each rosette; pedicel glabrous, trigonous, 1.5—2 X
80 mm; tepals 6, white, membranous, glabrous, lin¬
ear-lanceolate, in 2 whorls, subequal, with acro-
dromous venation; outer tepals 38 X 7 mm wide,
with conspicuous midribs; inner tepals more deli¬
cate, 37 X 10 mm wide; hypanthium 34 mm long,
glabrous, conspicuously trigonous, basally 2 mm
broad and hardly delimited from the pedicel, api¬
cal ly 5 mm diam.. with a 2 mm depression; ovary,
except for the apical excavation, filling 90% of the
hypanthium; style trigonous, in the center of the
depression, 1 X 35 mm; stigma capitate, 3 X 4
mm, lobes fleshv; stamens 6, 20 mm long; filaments
triangular, 3 mm long, basally 1 mm wide, inserted
atop the hypanthium margins; anthers 17 mm long,
basally auriculate, thecae subequal; pollen dis¬
persed in monosulcate monads, 35.5 X 25.7 pan,
with the aperture at the distal pole, with reticulate

14

Novon

200[jm

5mm

5mm

M

200(jm

5mm

figure I. A-J. Namiza almeidae Alves drawn from the holotype Paulo Roberto Almeida de Jesus s.n. (R). — A. Habit.
— B. Real apex. — C. flower subtended by 2 bracts. — I). Tepal from outer whorl. — E. Stamen. — F. Style. — G.
Seed. H. Pollen monad. — I. Transverse section of vascular bundle topped by sturdy hair. — J. Median transverse

Volume 12, Number 1
2002

Alves

Nanuza from Brazil

15

exine; capsule 45 mm long, 8 mm diam., longitu¬
dinally dehiscent from apex to Vz length, externally
sulcate, similar to, though smaller, than that on ex¬
amined specimens of N. plicata ; seeds pale brown,
pyramidal, testa retic-ulate-foveolate, slightly trans¬
lucent.

Transversal section of lamina. Vascular bundles
1 1; midvein and the 2 vicinal vascular bundles cir¬
cular, with sclerenchyma girders only abaxial, the
4 subsequent pairs of bundles pear-shaped, with
conspicuous abaxial U-shaped and adaxial sagit¬
tate girders; marginal trace with abaxial girder large
and asymmetrical, adaxial girder lacking or vesti¬
gial: mesophyll adaxially with palisade grading into
spongy tissue. Bundle sheath distinct, in contact
with abaxial and adaxial epidermis.

The type specimen was collected on a granitic
inselberg known as Mono do Urubu emerging from
degraded Atlantic rainforest in the state of Rio de
Janeiro, from the municipality of the type citation.

Nanuza almeidae is most easily distinguished in
herbarium material by leaf blades with a distinct
pseudopetiole, lamina wider than I cm lacking ser¬
rulate crests on the adaxial surface, and by linear-
lanceolate tepals. In contrast, broadly lanceolate te-
pals are shared by Nanuza almeidae and N. plicata .
and the latter has serrulate crests. The triangular
filaments with a flattened base and seeds with
deeper pits in the reticulate seed coat distinguish
it further from N. plicata. The specific epithet refers
to the collector of the holotype, Paulo Roberto Al¬
meida de Jesus, an enthusiastic potential field nat¬
uralist who frequently surprises us in the herbarium
bringing large bags filled with curious plants, ar¬
thropods, mollusks, and minerals, which we dis¬
tribute to specialists and collections. His first col¬
lection, now cultivated at our institution, inspired
this study and part became the holotype of Nanuza
almeidae.

Paratypes. BRAZIL. Bahia: Pedra Azul, 4 km da ci-
dade na estrada MG-105 para Jequitinhonha, Inselberg &
beira da estrada, R. M. Silva 440 et al. (SIT ). Espfrito
Santo: Vale do Rio Pancas, cultivated in Pedra de Guar-
atiba. Rio de Janeiro by Burle Marx, N. L Menezes s.n.
(SPF); Colatina, BR-259 a 17 km de Colatina para Joao
Neiva, Grande afloramento rochoso, 13 Jan. 1985, J. R.
Pirani A I). Zappi 1117 (SPF); Vitoria, Guajura, Praia do
Canto. A. G. Silva 463 (R); Vitoria, Morro do Guajura,
Praia do Canto, 27 Aug. 1984. A. G. Silva 449 (RB). Rio

de Janeiro: Mun. MagG Distrito de Piabeta, Fragoso/To-
caia, base do Morro do Urubu, 500 m s. m., 20 Nov. 2000
(41), Paulo Roberto Almeida de Jesus s.n. (R); s.l., s.d.,
Carlos Viana Freire s.n. (R-50224).

Nanuza luetzellmrgii Alves, sp. nov. TYPE: Bra¬
zil. Pernambuco: Serra Talhada, Serra da Car-
naubeira, Serrote Redondo. Eagedo no Topo da
Serra, arbusto 50-80 cm, 22 May 1071 (fr), E.
P. Heringer R26, I). Andrade Tima, ,/. de I3. L.
Sobrinho & A. C. Sarmento (holotype, R; iso¬
type, RB). Figure IK. L.

Haec species caudice obscure triquetro aristis rotun-
datis et foliorum vagina serrulata esetosa atque laminae
sulcis utrinque triehomatibus brevibus rigidis numerosis
instructis abaxialibus, quam adaxialibus 4-plo profundior-
ibus statim distinguitur.

Shrubs to 80 cm tall: caudex branches trigonous
with rounded edges, 12-15 mm diam., densely
packed with 3 series of leaf sheaths with serrulate
margins; leaves rigid; lamina linear-lanceolate, 3—
5 mm wide, 14—25 cm long, without pseudopetiole,
conduplicate, conspicuously plicate (in dry material
plication barely detectable and lamina condupli¬
cate along midrib, then bent back outward along
4th rib), in smaller scale vest igial ly plicate along
all ribs: abaxial and adaxial furrows internally with
abundant short and sturdy hairs; flowers solitary;
tepals lanceolate-elliptic, 7 mm wide. 27 mm long,
white \\L. Emperaire s.n., SPF]; pedicel 0.2 cm
wide, 1 1 cm long, grading into a 70 mm long, 3
mm wide trigonous ovary; stamens 6, erect, yellow;
style with stigma exceeds stamens by 5 mm. Cap¬
sule (A. Fernandes s.n., SPF) dry, dehiscent, cla-
vate, basal half 3-7 cm long, to 2-3 mm diam.. the
apical half thickens abruptly to 8—9 mm diam.; sur¬
face sulcate-striate as in both other species.

Transversal section of lamina. Spongy tissue
filling almost the entire mesophyll; midvein and
marginal veins with only abaxial sclerenchyma
girder, the remaining 7 pairs with both; the adaxial
girder with clavate apex and Y-shaped base. Ad¬
axial furrows vestigial, abaxial four times deeper;
midvein distinct with only abaxial sclerenchyma
bundle; mesophyll with spongy tissue; bundle
sheath distinct, not in contact with either epider¬
mis; cuticle layer discontinuous.

Lamina of material from Pernambuco (E. P. Her-

section of lamina, k. L: N. luetzelburgii Alves drawn from E. P. Heringer R26. I). Andrade Tima, J. de P. T. Sobrinho
& A. C. Sarmento (holotype, R; isotype. RB). — K. Transverse section of vascular bundle; note hairs in abaxial furrows
and on adaxial surface. — L. Median transverse section of lamina. M, N. N. plicata. — M. Transverse section of vascular
bundle. — N. Median transverse section of lamina with bundle topped by sturdy hair. In 1, k. and M. sclerenchyma
and cuticle are in black.

16

Novon

inger 826, I). Andrade Lima, ./. de R. L. Sobrinho
& A. C. Sarmento, R, RB) with 17 vascular bundles
and adaxial cuticle 2—3 cells thick; material from
Piauf (L. Emperaire 715, I PA) has 13 bundles and
a thicker cuticle (4—5 cells). This seems related to
tliffi irences in annual rainfall (ea. 500 mm in Per¬
nambuco and < 300 mm in Piauf).

The specific epithet refers to the first collector,
Philip von Eiitzelburg, a naturalist and important
plant collector in northeastern Brazil.

Paratypes. BRAZIL. Pernambuco: Serra Talhada,
Serra da Camaubeira, Serrote Redondo, 22 May 1971. E,
P Heringer, I). Andrade Lima, J. de P. L. Sobrinho & A.
C. Sarmento s.n. (R, RB); Petrolina, July 1974, I). Andra¬
de Lima s.n. (SPF); Flores, Serra do Cruzeiro na estrada
Triunfo-Sftio dos Nunes, 28 Mar. 1970, I). Andrade -Lima
70—5848 (I PA). Piauf: Sao Raimundo Nonato, Caldeirao
dos Rodrigues, 7 Jan. 1985, hiure Emperaire s.n. | (I |. Fx
Herbdrio da Missao Franco-Brasileira no Piauf (SPF); Ser¬
ra da Capivara, 1979, Laure Emperaire 715 [fr] (IPX, RB);
Serra dos Ca_?, 6 July 1912, iMtzelburg 1210 (RB); Serra
da Capivara, 1979, Laure Emperaire 494 (IPA); Guaribas,
Parque Nacional da Serra das Confusoes, Gruta do Boi,
ca. de 20 km de Caracol. 1 — 15 Oct. 2000 [st], Percequillo,
A. R. & Rodrigues, M. T. U. s.n. (SPF); Caracol, Serra das
Confusoes, Jorge, A. s.n. [st] (SPF), A. Fernandes s.n. | fr]
(SPF); Municfpio Sao Raimundo Nonato, Margern direita
da Fstrada Canto do Buriti, S. Raimundo Nonato, 13 Oct.
1999, Franca, A. R. et al. s.n. |st|, Fx Herbdrio TEGB
(SPF); Piauf?: Nordeste, Fxeursao com Bigarela, Aug.
1972, I). Andrade- Uma 72—6958 (IPA).

Gas chromatographic analysis coupled with mass
spectrometry of the crude extract obtained in di-
chloromethane was conducted on samples of all
three species. Nanuza almeidae revealed diterpenie
skeletons of the labdane type not found in the other
species and tetraeosanoic acid {C2>H )002, molecular
mass = 382}; Almeida s.n. (R). Nanuza luelzellmr-
gu revealed 8-heptadecenic aeid {Cl8H;i402, molec¬
ular mass = 282}; Heringer 826 et al., R Laure
Emperaire 715 (IPA). No long-chain acids were
found in N. plicata, which revealed only a mixture
of carbohydrates with chain lengths of 12, 22. 27,
and 35 carbons in a recent sample ( Riehl s.n.. R),
from the same population in Nova Friburgo, Rio de
Janeiro, from which Riehl et al. (2000) had de¬
scribed xerophytolic acid.

As all species of Nanuza are insufficiently rep¬
resented in collections, their distribution remains
insufficiently known. At present, it seems that Na¬
nuza almeidae occurs on gneiss-granitic inselbergs
composing mountain chains of the first belt parallel
to the coast, in the states of Bahia, Espfrito Santo,
and Rio de Janeiro. Nanuza luetzelburgii is known
only from basaltic outcrops in the caatinga (dry sa¬
vanna) of Pernambuco and Piauf in northeastern
Brazil. Nanuza plicata has a broader range, growing

on rock outcrops of various types throughout the
Espinhaco and Chapada Diamantina mountain
chains.

Color images complementing this article will be
available at the internet address (http://acd.ufrj.br/
~mndh/nanuza.html) immediately after printing.

Key to thk Species of Nanuza

la. Caudex distinctly trigonous to triquetrous, with
sharp edges; abaxial furrows of lamina well de¬
veloped, without hairs; leaf sheath margins emit¬
ting long libers that interlace on the caudex.

2a. Lamina linear to lanceolate, without pseu¬
dopetiole, tepals elliptic . N. plicata

2b. Lamina with distinct pseudopetiole, adaxi-
ally glabrescent, tepals linear-lanceolate . .

. N. almeidae

lh. Caudex obscurely trigonous with rounded edges,
abaxial furrows ol lamina with abundant sca¬
brous hairs; adaxial furrows half the depth of ab¬
axial; leaf sheath margins serrulate, without fi¬
bers . N. luetzelburgii

Selected additional examined material of N. plicata.
BRAZIL. Italiia: Mun. Agua (Juente, Pico das Almas,
Vertente oeste, entre Paramirim das Crioulas e a face
NNVt do Pico, 13°31'S, 42°00,W, 1250 m s. m., R. M.
Harley 27535 (SPF); Mun. Bom Jesus da Fapa. Caatinga
sobre afloramentos (lagedos) rochosos, 14 Feb. 1991, G.
M. Halsehbach 55144 & (). S. Ribas (SPF); Monte Santo,
3 Oct. 1972, I). Andrade Lima s.n. (SPF); Monte Santo
[localidade tfpica], 4 Feb. 1972, I). Andrade Lima s.n.
(RB); Mun. Gentio do Ouro, 27 Nov. 1992. M. M. Arbo,
R. M. Silva & Jodo Vicente 5337 (RB); Mun. Barra da
Estiva, 16 km NE de Barra da Estiva, 790 m s. m„ 24
Nov. 1992, AL AL Arbo, R. Al. Silva & Jodo Vicente 802
(RB); Mun. Santo Inacio, Serra do A^urua, Arredores da
cidade, 26 Oct. 1970, D. Andrade-Lima 70—6150 (IPA);
Mun. Acobina, Campo Formoso, 21 July 1978, I). Andra¬
de-Lima 78-8577 (IPA). Espfrito Santo: Mun. Colatina,
Angelo freschiani, ca. 3 km apds Ponte do Pancas, ca.
200 m s. m., veg. campestre em afloramento gneissico, H.

Q. Boudet Fernandes 2684 el al. (SPF); Mun. Colatina, 15
km N de Colatina, 4 Dec. 1994, ,/. R. Pirani, AL A. G.
Magenta & A. A. Goncefdio s.n. (RB); Colatina, Rodovia
FS-080, ca. 15 km a Norte de Colatina, 4 Dec. 1994, ,/.

R. Pirani et al. s.n. (SPF). Minas Gerais: Jufz de Fora,
Linhares, Pedreira Santo Cristo, 15 May 1996,4. A. Caia-
fa & G. L. G. Soares s.n. (SPF); 10 km a Sul de Medina,
rodovia BR-I 16, 20 Nov. 1985, 6’. Hatschbach 50013 &
./. M. Silva (SPF); Mun. Itinga, 5 km de Itinga, R. M. Silva
427 et al. (RB, SPF): Mun. Itinga, Santana do Ara^uari,
28 Aug. 1982, L. G. Mautone, E. E Guimaraes C. T.
Ri zzini s.n. (RB); Parque Estadual da Serra do Brigadeiro,
Serra da Araponga, Fazenda Neblina. Proximo ao labora-
tdrio de Campo, trilha subindo mono, 30 Sep. 1995, ,/. A.
hmdmrdi 983 (SPF); Mun. Mantena, Rio Preto, Fazenda
Boa Vista do Sr. Antonio Ferreira Valente, 14 Oct. 1995,
G. E. Valente 148 & A. Azevedo (SPF), 14 Oc t. 1995, G.
E. Valente 147 & A. Azevedo (SPF); Pedra Azul, 15 km da
cidade, V. C. Souza 5193 et al. (SPF); Mun. Carlos Chagas,
Rodovia BR418, Rio Quegeme, paredoes rochosos, G.
Hatschbach 47086 (SPF); Aguas Vermelhas, proximo ao
Curral de Dentro, margern da Rodovia 251, 15 Aug. 1974,
I). Andrade Lima s.n. (SPF); Mun. Varzelandia, Serra da

Volume 12, Number 1
2002

Alves

Nanuza from Brazil

17

Caatinga, sobre a Lapa do Varal, 1 Feb. 1985. J. G. da
Silva 1069 & M. Menezes (R); Mun. Varzelandia, Barrei-
rinho, 7 Feb. 1985, ./. G. da Silva 1143 & M. Menezes
(R); Serra do Torreao, July 1897, F. Brandao s.n. (RB).
Rio de Janeiro: Mun. Petropolis, Araras, Malta, 8 Nov.
1968, P. I. S. Braga 1354 (RB); Mun. Petropolis, Pedro
do Rio, Secretario, Nascente do Rio Capirn Roxo, 7 Dec.
1986, G. Martinelli 11985 & C. Alvim (RB).

Acknowledgments. I thank colleague Jorge Fon-
tella Pereira for the Latin diagnoses, Vania Gon¬
salves Esteves and Claudia Barbieri Ferreira Men-
donya for aid with the pollen description and
image, Jenilce R. Martins, Claudia M. Rezende,
and Angelo C. Pinto for aid with the chemical anal¬
yses and the curators of GUA, HB, I PA, R, RB,
SPF, and VIES for support.

Literature Cited

Alves, R. J. V. 1994. Morphological age determination &
longevity in some Vellozia populations in Brazil. Folia
Geobot. Phytotax. 29: 55-59.

Baker, J. G. 1875. Synopsis of the African species of Xe-
rophyta. J. But. 13: 236.

Martius, C. F. P. von. 1823. Velloziaceae. In: Nov. Gen.
Sp. PI. 1: 16, t.9.

Mello-Silva. R. de. 1990a. Morphological and anatomical
differentiation of Vellozia hirsuta populations (Vellozia¬
ceae). PI. Syst. Evol. 173: 197-208.

- . 1990b. A new species of Vellozia from the Es-

pinhayo Range, Brazil, with some considerations on the
section Xerophytoides. Kew Bull. 46: 321—326.

- . 1992. Three new species of Vellozia from Pico

das Almas, Bahia, Brazil, with an account of their leal
anatomy. Kew Bull. 48: 1-8.

- . 1995a. Velloziaceae. In: B. L. Stannard (editor),

Flora of the Pico das Almas, Chapada Diamantina, Ba¬
hia, Brazil. Royal Botanic Gardens, Kew\

- . 1995b. Aspectos taxonomicos, biogeogrdficos,

morfologicos e biologicos das Velloziaceae de Grao-Mo-
gol, Minas Gerais, Brasil. Bol. Bot. Univ. Sao Paulo 14:
49-79.

- . 2000. Partial cladistic analysis of Vellozia and

characters for the phylogeny of Velloziaceae. In: K. A.
Wilson & D. A. Morrison (editors), Monocots: System-
atics and Evolution. CSIRO, Melbourne. | Abstract.]

Pohl, J. E. 1827. Velloziaceae. In: PI. Bras. 1: 129.

Riehl, C. A. S., A. C. Pinto & C. R. Kaiser. 2000. Isolation
and structural determination of xerophytolic acid, a nov¬
el 3-geranyl-4-hydroxybenzoate derivative from Xero-
phyta plicata. Spectroscopy Lett. 33(5): 643—651.

Salatino, A. 1999. Main results from trnV-V sequencing
of Velloziaceae and allied taxa. Anais Acad. Brasil. Ci.
71: 203-206.

Salatino, M. L. F., A. Salatino, N. L. de Menezes X R. de
Mello Silva. 1989. Alkanes of foliar epicuticular waxes
of Velloziaceae. Phytochemistry 28: 1105—1114.

Smith, L. B. & E. S. Ayensu. 1976. A revision of Amer¬
ican Velloziaceae. Smithsonian Contr. Bot. 30: 1—172.

Sprengel, K. P. J. 1827. Velloziaceae. In: Syst. Veg. 4(2):
137.

Two New Species of Phacelia (Hydrophyllaceae) from the
Southwestern United States

N. Duane Atwood

I afe Science Museum, Brigham Young University, Provo, Utah 84602, IJ.S.A.

Frank ./. Smith

Western Ecological Services, Inc., Box 422, Millville, Utah 84326, U.S.A.

Teri A. knight

The Nature Conservancy, 1771 E. Flamingo Road, Suite 111 B, Uas Vegas. Nevada 89119,
U.S.A. Present address: 1924 Quimper Court, Henderson, Nevada 89014, U.S.A.

ABSTRACT. Phacelia filiae N. I). Atwood, F. J.
Smith & T. A. Knight is an undescribed species
from Clark, Nye, and Lincoln Counties, Nevada. It
is closely related to Phacelia parishii of California,
Arizona, and Nevada and Phacelia beatleyae of
southern Nevada. Phacelia petrosa N. D. Atwood.
F. J. Smith & T. A. Knight from Arizona, Nevada,
and Utah is also described as new. It is most closely
related to Phacelia crenulata, which is known from
Arizona, California, Nevada, and Utah. The most
prominent differences among these species are the
shaf )e of the seeds and appearance of the seed
coats.

Key words: Arizona, Hydrophyllaceae, Nevada,
Phacelia , U.S.A., Utah.

The genus Phacelia comprises about 250 taxa
mostly native to the New World and is best devel¬
oped in the western United States and northern
Mexico. This genus is known for its large number
of endemic taxa. The two species described below
are narrow endemics, with Phacelia filiae restricted
to the Mojave Desert and the transition zone of the
Mojave Desert and the Great Basin, and Phacelia
petrosa to the lower drainages of the Colorado River
in southern Nevada, southern Utah, and northwest¬
ern Arizona. One of the key characters used to dis¬
tinguish species of Phacelia is seed morphology.
Other morphological features that are useful are:
size, shape, and color of flowers; duration; leaf size,
shape, and pubescence; as well as stamen and style
length in contrast to the corolla length. In addition,
geographic distributions are useful in distinguish¬
ing taxa.

Novon 12: 18-26. 2002.

Phacelia filiae N. D. Atwood, F. J. Smith & T. A.
Knight, sp. nov. TYPE: U.S.A. Nevada: Clark
Co., Nellis Range, 36°38'02"N, 1 15°30'57"W,
Three Lakes Valley, 45 air km NW of I .as Vegas,
3080 ft. elev., 2 May 1998, F ./. Smith & P.
Dwyer 4217 (holotype. UNLV; isotypes, ARIZ,
ASC, BRY, CAS, GH, MO. NY. N FS. OSU. RM.
RSA. UNLV, US, UTC). Figures 1-3.

Similis Phacelia parishii sed mi floribus et lobis eorol-
larum majoribus foliis caulinum plus numerosis et semi-
nibus grandibus paucioribus differt et a /I beatleyae in
foliis plerumque basalibus nee vulgo caulinis et planis
nee revolutis admodum integris seminibus parvioribus et
plus numerosis absiniilis.

Plants annual, 2.3— 6.0 cm high, branched at or
near the base; stems 2.5—7 cm long, curved up¬
ward; leaves mostly basal, petiolate, the blade 0.7—
3.0 X 0.5— 1.5 cm. ovate, elliptic, or oblong, entire
to few toothed, the petiole 1.5—15 mm long; inflo¬
rescence of terminal secund cymes, 0.6— 3.5 cm
long; calyx united basally, the lobes heteromorphic,
of unequal widths with 3 distinct veins, from nar¬
rowly spathulate to broadly obovate, 4—5 X 1.2— 2.7
mm in flower, fruiting calyx 4.5-8 X 1.2— 3.5 mm;
corolla tubular-campanulate, 6—7 X 2. 5-3. 5 mm,
the lobes 1 .5—2.4 mm long, finely puberulent ex¬
ternally, the tube pale yellow, the limb lavender;
stamens and style included, filaments 3—4 nun long,
glabrous, attached at the base of the corolla tube;
the style 1.1 — 1.2 mm long, bifid, glabrous; capsules
4.2-5. 5 X 3.2— 3.5 mm, finely glandular puberu¬
lent; seeds 18—26, 1.3-1. 8 X 0. 5-0.6 nun, angular,
and with rounded or narrowed tips, black with ev¬
ident pits.

Distribution. Phacelia filiae occurs in southern
Nevada on Nellis Range (Clark County), Nevada

Volume 12, Number 1
2002

Atwood et al.

Phacelia from Southwestern U.S.A.

19

Figure I. Phacelia filiae N. I). Atwood, F. J. Smith & T. A. Knight. — A. Habit. — B. Flower. — C. Fruit and
calyx. — 1). Seed. — E. Seed detail. Drawn by Linda A. Vorobik (plant based on the holotype Smith & Dwyer 4217,
seed based on Smith & Ackerman 3880).

20

Novon

lest Site (Nye County), Desert National Wildlife
Refuge (Clark and Lincoln Counties) and in urban
Las Vegas (Fig. 2).

Habitat. Plants of Phacelia filiae are found on
relatively flat areas or on low knolls of the valley
floor, mostly above the playas and in the foothills
of desert mountains. This species grows in light-
colored soils of calcareous sandstone, siltstone,
tuffaceous claystone, and limestone substrates. L I -
evation ranges from 2000 to 4000 feet, where it
occurs in shadseale. blackbrush, and creosote bush
scrub.

Paratype. s. U.S.A. Nevada: Clark Co., near Las Ve¬
gas. ft May 1.944, A. Eastwood A' J. Howell 8958 (CAS.
(ill. POM. RM, UTC); foothills of Spotted Range. 13 Mav
1941, //. I). Ripley A R. C. Barneby 3425 (CAS. RSA);
along Cheyenne Ave. near the drive-in theaters. Las Ve¬
gas. 25 Apr. 1979, P. Leary 2941 (UNLV); Indian Springs
Valley, TI3S, R57E, S29, 15 May 1993, E J. Smith A E.
Watkins .1670 (UNLV); Pintwater Range, T15S R56E SI.
3660 ft. elev., 23 Apr. 1995. E ./. Smith A R. Schofield
4847 (UNLV); Indian Springs Valley, T15S, R56K, S3. 23
Apr. 1995, E J. Smith A R. Schofield 3848 (UNLV); In¬
dian Springs Valley, T13S, R57K S29, 6 May 1995, E ./.
Smith A' T. Cox 3861 (UNLV); W of Pintwater Range,
TI 3S, R56E S28, 7 May 1995, E ./. Smith A T. Cox 3862
(UNLV); W of Pintwater Range. T13S R56E S25, 7 May

1995, E ./■ Smith & T. Cox 3863 (UNLV); Spotted Range,
T15S, R55E, S5. 23 May 1995, E ./■ Smith. ./. Heers A'
E. Watkins 3893 (UNLV); Three Lakes Valley, T16S R57E
S3, 30 Apr. 1995. F. J. Smith & E. Watkins 3858 (UNLV);
Lincoln Co., E side of Desert Lake, 28 Mav 1983, T. Ack¬
erman 83-562 (UNLV); SE end of Desert Lake. T11S,
R60E, 10 May 1987, 4. Tiehrn 10994 (RM. RSA, UNLV.
UTC); Desert Lake, I I IS, R60E SI, 22 Apr. 1995, F. J.
Smith. R. Schofield A- B. Benjamin 3845 (UNLV); E side
of Desert Lake, T1 IS. R60E, 16 May 1995, E J. Smith
A 7.' Ackerman 3880 (UNLV); Desert Lake. I I IS. R60E,
S36. 28 Apr. 1995. E J. Smith A S. Sheldon 3854 (BRY,
MO, NY, UTC); Nye Co.. NW side of Rd. 28-03, 0.8 mi.
NE of its junction with Jackass Flats Rd., E Rock Valley,
26 May 1976, M. Williams 145 (UC); Rd. 28-03. 0.8 mi.
NE of junction with Jackass Flat Road, E Rock Valley. 20
May 1976, M. Williams 120 ( NTS); I mi. E of Pink Holes
Hill W of f renchman Elat. 8 June 1976 , M. Williams 173
(NTS); \\ side of low hills 1 .0 mi. S of Little Skull Moun¬
tain. Vi of Jackass Flat Road, Rock Valley, 26 Apr. 1978,
S. Cochrane & M. Williams 1010 (UNLV); NW Rock Val¬
ley off the W side of Jackass f lats Road S of the pass
into Jackass Flats, 26 Apr. 1978. M. Williams 450
(UNLV); 3.7 mi. W of 28—03 Road, W Rock Valley, 9 May
1978. .S’. Cochrane A Timbrook 1092 (UNLV, RENO); E
side of Burma Road, 1.4 mi. NW from Mercury Highway
at Mercury Pass, NW' slopes of Mercury Ridge, Red
Mountain, 28 Apr. 1978. M. Williams 461 (UNLV); Rock
Valley, UTM 567482E, 4064091 N, 4 May 1995, G. Lyon
300 (UNLV); Rock Valley, along N and S sides of Jackass

Volume 12, Number 1
2002

Atwood et al.

Phacelia from Southwestern U.S.A.

21

Figure 3. Scanning electron micrographs of seed and seed coat. A, B. Phacelia filiae from Smith & Ackerman 3880
(UNLV). — A. Seed. — B. Seed coat. C, D. Phacelia parishii from Anderson 95—9 (ASU). — C. Seed. — D. Seed coat.
E, F. Phacelia beatleyae from Bentley 5585 (UTC). — E. Seed. — F. Seed coat. Scale bar in A, C, E = 0.5 mm. Scale
bar in B, I), F = 0.075 mm.

Flats Road, S of Skull Mtn., UTM 570164E, 4062190N,
8 May 1995, T. Lindemann 47 (UNEV); Rock Valley, UTM
576247E, 4061394N, 8 May 1995, T. Lindemann 48
(UNLV); Rock Valley, UTM 577080E 4060820N, 8 May
1995, T. Lindemann 50 (UNLV); Rock Valley, UTM
580561 E, 4060888N, 8 May 1995, T. Lindemann 52
(UNLV); Rock Valley, UTM 581013E 4060575N, 9 May
1995, G. Lyon 301 (UNLV); Rock Valley, UTM 579200E
4061100N, 1130 111 elev., 9 May 1995, G. Lyon 302
(UNLV); Rock Valley, UTM 578490E 4061200N, 9 May
1995, G. Lyon 303 (UNLV); Rock Valley, UTM 577820E
4060960N, 9 May 1995, G. Lyon 304 (UNLV); Frenchman
Flat, UTM 590270E 4063720N, 10 May 1995, G. Lyon
305 (UNLV); Frenchman Flat, UTM 587780E 4065920N,
10 May 1995, G. Lyon 306 (UNLV).

In the spring of 1995, a new annual species of
Phacelia was discovered on the Nellis Ranges by
the second author. During that same year, the sec¬
ond author was conducting an inventory of Phacelia
parishii for the Nevada Natural Heritage Program.
In May, Smith visited known sites (Fig. 2) of P
parishii and P. beatleyae on the Nevada Test Site.
Plants from there did not match the description or
fit the key for P. parishii or P beatleyae. They ap¬
peared to be the same undescribed Phacelia found
on the Nellis Ranges. Surveys for this new Phacelia
continued on Nellis lands through 1999, with pop-

22

Novon

illations appearing in high precipitation years. Ap¬
proximately 30 populations comprise the currently
known range.

The most significant differences among these
species are the seeds: their number, shape, and the
appearance of the seed coats. The seeds of these
three species are distinct. The seeds of P. filiae are
1.3— 1.8 mm long, 18 to 26 per capsule, long-an¬
gular in shape and rounded or narrowed to the
ends, with more and deeper pits. The seeds of P.
parishii are smaller, more numerous (30 to 40 per
capsule), mostly rounded in shape with round tips
and shallow pits; and the seeds of P. beatleyae are
40 to 50 per capsule, angular in shape, round at
the tips, and deeply pitted. These distinguishing
characters are observed readily in the electron mi¬
crographs (Fig. 3). In addition, the stems ol P. filiae
and P beatleyae are always leafy, but some stems
of P. parishii can be leafless. Also, the stems of P.
beatleyae are erect while those of P. filiae and P.
parishii are decumbent. Phacelia filiae differs from
P. parishii in having larger tubular-campanulate
flowers (6— 7 mm long vs. 4-5 mm long), larger co¬
rolla lobes (1.5— 2.4 mm long vs. 0.7—1 .3 mm long),
and more cauline leaves. It further differs from P.
beatleyae, which lacks a basal rosette and has
strongly revolute and essentially entire leaves.

Phacelia Jiliae should be placed w ith P. parishii
and P. beatleyae in the subgenus Eutoca. According
to Howell (1943), Phacelia subg. Eutoca is char¬
acterized by plants that are chiefly annual (some
perennial) with leaves entire to shallowly lobed, co¬
rollas tubular or tubular-campanulate, stamens
shorter than the corolla, style bifid or at most parted
Vi its length, and by capsules that are generally
elliptic or oblong and obtuse or truncate below the
terminal apieulation. The species name filiae is
used to honor the third author’s daughter, with
Clarke’s phacelia the suggested common name.

The following key separates P filiae from closely
related species of Phacelia that occur in Nevada.

la. heaves essentially entire, revolute, all cauline;
seeds 40 to 50 per capsule, 0.6-1. 0 mm long,

short angular or rounded .

. P. beatleyae Constance & Reveal

II). heaves entire-orenate, not revolute, basal and
cauline; seeds 40 or fewer per capsule, 1.1 — 1.8
mm long, roundish oblong or long angular.

2a. Corolla 6—7 mm long, the lobes 1.5— 2.4 mm
long; seeds 18—26 per capsule, 1.3— 1.8 mm
long, deeply pitted (crater-like), long angular
with narrowed or rounded ends and deeper

pits .

P. filiae N. I). Atwood, F. J. Smith & T. A.

. Knight

2b. Corolla 4—5 mm long, the lobes 0.7— 1.3 mm
long; seeds 30 to 40 per capsule, 0.9— 1.1
mm long, roundish oblong with round ends
and shallow pits . P. parishii A. Gray

Phacelia petrosa N. I). Atwood, F. J. Smith N T.
A. Knight, sp. nov. TYPE: U.S.A. Arizona: Mo¬
have Co., T32N, R9W, S24, Mohave Strip, 1
mi. N of the Colorado River, E of Whitmore
Canyon-Colorado River overlook on limestone
talus slopes, 22 Apr. 1999. I). Atwood & H.
Furniss 24210 (holotype, BRY; isotypes,
ARIZ, ASC, RM, CAS, GH. K, MO. NTS, NY.
OSU. POM. RM. RSA. UNLV, US, UTC). Fig¬
ures 4—6.

A Phacelia crenulata in pilis hispidis et glandularibus
floribus minoribus et pedicellis longioribus infra, stami-
nibus exsertis minoribus seminibus foveis nullis differt, a
P. ambigua in seminibus pagina ventrali corrugata intus
el cristam latum unum pagina dorsali bruneo pallide e
margine stramineo pal lido cicatricibus dorsalis rotundatis
el elevatis et e radiis stellatis et Horibus parvioribus et
pubescentibus glandularibus infra inflorescentiis praeci-
pue differt.

Plants annual. 1.0— 3.2 dm tall: basal leaves 3.3—
7.5 cm long, rounded to oblong, petiole up to 3 cm
long, the blade 2.5— 5.0 cm long, toothed, pinnatifid;
cauline leaves irregularly dentate, reduced upward,
0.8—3 cm long, 0.8— 2.0 cm broad, petiole 1.1— 2.4
cm long; pubescence densely hispid with spread¬
ing, shiny hairs, and glandular villous; inflores¬
cence of paired terminal cymes w ith capitate mul¬
ticellular hairs, cymes elongating to 6 cm long in
fruit; pedicels 0.8— 1.5 mm long in flower, lower
pedicel to 5.4 mm long; calyx 3.0— 3. 4(4.2) mm
long, the lobes oblanceolate, separate to near the
base, 0.9— 1.5 mm wide, not enlarging in fruit; co¬
rolla campanulate, 5.8— 6.0 mm long and broad,
blue, lighter at base, the lobes glabrous to pubes¬
cent externally; stamens and style exserted, sta¬
mens subequal to the style, 3. 7^1.3 mm long, fil¬
aments purple and glabrous, style 2.8-4. 5 mm long,
bifid, lower portion pubescent with glandular hairs;
capsule globose, mature capsules 3.0— 4. 0(4. 6) mm
broad anti long, equaling to exceeding the calyx
lobes, finely puberulent with some glandular hairs
especially on the upper half; seeds 4. oblong, 3.3—
3.8 mm long, 1.5— 3.1 mm wide, the ventral surface
corrugated along the inside margin and one side of
the ridge, dorsal surface with a lighter stramineous
margin and light brown surface, dorsal cells round
in shape and raised with star-like rays.

Distribution. Phacelia petrosa occurs in south¬
western Utah (Washington County), southeastern
Utah along the San Juan River (San Juan County),
northwestern Arizona along the Colorado River in
Mohave and Coconino Counties, and southern Ne¬
vada in Lincoln anti Clark Counties (Fig. 5).

Habitat. The species is generally found on dry7
limestone and volcanic talus slopes of foothills,

Volume 12, Number 1
2002

Atwood et al.

Phacelia from Southwestern U.S.A.

23

Figure 4. Phacelia petrosa N. I). Atwood, F. J. Smith & T. A. Knight. — A. Dorsal surface of seed. — B. Ventral
surface of seed. — C. Capsule, sepals, and style. — D. Flower with exserted stamens and style. Drawn by Shannon
Workman (based on the type specimen Atwood & Furniss 24295 (BRY)).

washes, and gravelly canyon bottoms. It grows
mostly on calcareous parent material but has been
collected on dry volcanic talus slopes. The eleva¬
tion range is 2500 to 5800 feet, where it occurs in
mixed desert shrub, creosote bush, and blackbrush
communities.

Paratypes. U.S.A. Arizona: Coconino Co., Bright An¬
gel trail, 16 Apr. 1938, Harrow s.n. (UC); Bright Angel

trail near first creek crossing above river, 9 May 1940,
Bailey & Bailey 1097 (UC); beyond Miner's cabins. Hav¬
asu Canyon. 3 May 1947, Dearer 20B5 (ASC); Hualapai
Canyon, 28 May 1950, J. Howell 265BB (CAS); above
House Rock Rapids and Hot Na Na Wash, 8 May 1973,
A. Phillips s.n. (ARIZ); about 1 mi. downstream from
House Rock Bapids and Rider Canyon, 9 May 1973, A.
Phillips s.n. (AIHZ); first canyon downstream from Unkar
Canyon on N, 13 May 1973, A. Phillips s.n. (AKIZ); 18.5
mi. campground, Colorado River, 18 May 1976, Hevly s.n.

24

Novon

(ASC); Cottonwood Canyon, 22 Apr. 1979, Van Devender,
Cole, Markgraf & Marlin s.n. (AR1Z); Mohave Co., Torow-
eap Valley, ea. 4250 ft. rlev., 50 Apr. 1952, K. McClintock
S2—2/7a (CAS); S ol Toroweap Point, halfway between
Devil’s Bathtub and Burro Canyon, 1 May 1952, E.
McClintock 52—348 (CAS); inner gorge ol Grand Canyon,
along lava slide on trail to river, 5 May 1952, E. Mc¬
Clintock 52 — i06 (CAS); below Emery Falls, opposite
Rampart Cave, 8 Apr. 1971. Martin s.n. (AR1Z); side can¬
yon at Grand Canyon mile 254.6R, 8 Apr. 1992. 4. Phil¬
lips & II Phillips 9381 (ASC); T35N, R15W, S31 NW Va,
black lava ridge SE of Pakoon Ranch. 15 May 1985, I).
Atwood 10945 (BRY); T32N, RIOW. SO, Andrus Canyon
3 mi. W ol Andrus Point, 20 Apr. 1999, I). Atwood & B.
Eurniss 24295 (BRY, MO, NY, US); T32N, R9W, Sll
NWSE, 2 mi. E of Whitmore Point, 22 Apr. 1999. I).
Atwood & B. Eurniss 24222 (ARIZ, ASLJ, BRY, MO, NY,
UNEV). Nevada; Clark Co., SE end of Las Vegas Range,
14 May 1976, T. Ackerman 70-56 (UTC); NNW foothills
ol Charleston Mis., T17S, R50E, S4 SW Va, 5 air mi. S of
Indian Springs, 2 May 1985, I). Atwood & E. Thorne
10854 (BRY, RENO); l.ns Vegas Range, Second Canyon

5 ol Peek-a-boo Canyon, 29 May 1976, T. Ackerman 4990
(UC, UNI.V); Desert Range, T13S, R59E S20. 22 May
1993, E ./. Smith. T. .4. Knight & J. Pedrick 3680 (UNEV);
Valley of Eire. T18S, ROOF. SI, 17 May 1995, E ,/. Smith

6 T. Ackerman 3882 (UNEV); Lincoln County, Eas Vegas
Range, Elbow Canyon, T14S, R02E, SI NE!4, 15 May
1995, E ./. Smith & T. Ackerman 3877 (UNEV, UTC);
Desert Range, T12S. R59E, SO, 1 May 1993. E ./. Smith

& B. Schofield 3034 (UNEV). Utah: San Juan Co., Mile
40.0 at the lower end of the Goosenecks of the San Juan
River, 6 May 1997, 1\. I). Atwood et al. 2191 1 (BRY, MO);
mi. 59.2, 0.7 mi. below Johns Canyon along the San Juan
River, 7 May 1997, N. I). Atwood et al. 21934 (BRY);
Washington Co., T42S, R19W, SI 8, Beaver Dam Wash 3
mi. S of Lytle Ranch, 17 Apr. 1986, I). Atwood & K.
Thorne 12000a (BRY); SW ol Bulldog Canyon, 17 May
1980. /.. C. Iliggins 10552 (BRY); Diamond Valley vol¬
cano, 20 May 1986, L. C. Higgins 10005 (BRY), 2 June
1998, L. C. Higgins 19041 (BRY, GH, MO, NY, OSU,
RSA, US); TIES, RIOW', S3 SWSE, NE corner of Snow
Canyon State Park at Winter Quarters, 20 May 1992, M.
E Franklin 7484 (BRY); T43S, R18W, S33, S of Bull Dog
Canyon, 3 May 1980. K. Thorne et al. 4505 (BRY); T43S,
R18W, S28 SE Va, Bull Dog Knolls, 1 May 1986, O'. Baird
& S. Welsh 2395 (BRY); T43S, T18W, S28, S slope of Bull
Dog Knolls, 30 Apr. 1986, S. Welsh & O’. Baird 23098,
O'. Baird 2323 do (BRY).

In 1970, llie senior author examined collections
ol this species from the Grand Canyon. An attempt
was made to find the species in the field in 1971
without success. The species became a forgotten
entity until Smith and Knight discovered it while
doing fieldwork on Nellis Ranges in 1993. They
collected a Phacelia in the Desert Range (Lincoln
County, Nevada) with pubescence much different

Volume 12, Number 1
2002

Atwood et al.

Phacelia from Southwestern U.S.A.

25

Figure 6. Scanning electron micrographs of seeds and seed coat. A. 15. Phacelia petrosa from Atwood 10945 (BRY).
— A. Seed. — B. Seed coat. C. D. Phacelia crenulata from Holmgren & Reveal 1031 (UTC). — C. Seed. — I). Seed
coat. E. K Phacelia amhigua from Higgins 15475 (BUY). — E. Seed. — F. Seed coat. Scale har in A, C. E = 1.5 mm.
Scale bar in B, I), F = 0.3 mm.

than typical Phacelia amhigua and P. crenulata.
Based on close examination, the seed morphology
of P. petrosa differs from that of Phacelia amhigua
and Phacelia crenulata. Phacelia petrosa has been
identified on herbarium sheets in the past as P.
crenulata, P. pedicellata, P. amhigua, P. minutiflo-
ra. and P. amhigua var. minutiflora.

Phacelia petrosa belongs to the Crenulatae group
of species in the subgenus Phacelia in section Pha¬
celia. The Crenulatae group is distinguished by the
four-seeded capsule and excavated ventral surface
of the seeds (Atwood, 1975). Phacelia petrosa is
most closely related to P. crenulata and P. amhigua.

The most significant differences among these spe¬
cies are the appearance of the seed coat, flower
size, and pubescence. As can be seen on electron
micrographs, the seed coat morphology for each of
the species is distinctive (Fig. 6A— F). Phacelia pe¬
trosa differs from P. crenulata in being more hispid,
with shorter and less glandular pubescence, smaller
flowers, longer pedicels on the lower flowers, sta¬
mens less exserted, and the seeds not foveolate.
The new7 species differs from P. amhigua in having
the ventral surface of the seeds corrugated along
the inside margin and one side of the ridge, dorsal
surface with a lighter stramineous margin and light

26

Novon

brown surface, dorsal cells round in shape and
raised with star-like rays, smaller flowers, and glan-
dular pubescence, especially below the inflores¬
cence. I he species is named for the rocky places
where it grows on limestone and volcanic sub¬
strates. The suggested common name is rock pha-
celia.

The following key distinguishes Phacelia petrosa
and its closely related species.

la. Stems mostly glandular with some short non-

glandular hairs (not hispid) .

. Phacelia crenulata Torrey ex S. Watson

lb. Stems with spreading, hispid hairs as well as vis¬
cid -puberulent or finely stipitate-glandular pu¬
bescence.

2a. Seeds corrugated only along one side of the
ventral ridge, no stramineous margin on the
dorsal surface Phacelia ambigua M. E. Jones
2b. Seeds corrugated on the margins as well as
along one side of the ventral surface, stra¬
mineous margin present on the dorsal sur¬
face .

Phacelia petrosa IN. I). Atwood. F. J. Smith &

T. A. Knight

Conservation status. Phacelia filiae is limited to
a small area in southern Nevada from the urban
Las Vegas area to lands managed by the Depart¬
ments ol Defense, Energy, and Interior. Populations
in or near the urban environment of Tas Vegas are
threatened by direct loss from development and
habitat fragmentation, or are already extirpated.
Populations on federal lands (Defense and Energy)
are largely protected by their remoteness and re¬
stricted access. A few populations occur in valley
bottoms and may be vulnerable to defense-related
activities. Conservation of these areas is being ad¬
dressed by Nellis Air Force Base (Keystone Dia¬
logue. 1998). The conservation status of Phacelia

petrosa is unknown. Since most locations were not
visited, an evaluation needs to be made.

Acknowledgments. The authors thank Linda
Ann Vorobik and Shannon Workman for ink illus¬
trations of P. filiae and P. petrosa , respectively, Wil¬
liam McManus and Heather Leary for SEM photog¬
raphy, Stanley Welsh and students of the 1999
winter Botanical Latin class for the Latin descrip¬
tions, and Marti Aikens and Gerald Hughes of Geo-
Graphics for production of the distribution maps.
Review by Noel Holmgren strengthened the man¬
uscript and is greatly appreciated. Kevin Blomquist
donated time, assistance, and access to the Nevada
Test Site. Curators at ARIZ. ASC. BRV CAS. GIL
NTS. NY. POM. RENO, RM. RSA. UC. UNLV, and
UTC generously loaned specimens for review. De¬
partment of Defense funded inventory work by The
Nature Conservancy was conducted through a grant
from the Legacy Resource Management Program.
We are grateful to Colonels Thomas Lillie and
Douglas Ripley for their support, advice, and en¬
thusiasm. Nellis Air Force stall, and Eric Watkins
in particular, facilitated trips onto the restricted
Ranges for over half a decade. Overlapping surveys
for the Nevada Natural Heritage Program provided
additional support for Phacelia inventory in Nevada.

Literature Cited

Atwood, N. I). 1975. A revision of the Phacelia crenulatae
group (Hydrophyllaceae) for North America. Great lia-
sin Naturalist 35: 127-190.

Howell, J. T. 1943. Studies in Phacelia: A revision of
species related to P. pulchella and P. rotundifolia. Amer.
Midi. Naturalist 29: 1-26.

Keystone Dialogue. 1998. Keystone dialogue on Nellis Air
Force Range stewardship. Unpublished manuscript,
Nellis Air Force, Nevada.

Ipomoea electrina (Convolvulaceae): A New Name for
Exogonium luteum House

Daniel F. Austin

Arizona- Sonora Desert Museum, 2021 N. Kinney Road, Tucson, Arizona 85743, U.S.A.

,/. Andrew McDonald

Department of Botany, University of Texas, Austin, Texas 78713, U.S.A.

ABSTRACT. The history of a southern Mexican en¬
demic Ipomoea is summarized, beginning with its
publication in 1908 as Exogonium luteum House.
Because of a previously existing Ipomoea lutea
Hemsley, the new name Ipomoea electrina is pro¬
posed to refer to t he unique amber-colored flowers
of these plants. Since two distantly related conge¬
ners, Ipomoea lutea and /. urbinei, have nomencla-
tural and misidentifieation histories interlaced with
/. electrina, they are also discussed and compared
will) this taxon.

Kksumen. Se presenta un resumen de la historia
de una Ipomoea endemiea del sur de Mexico, em-
pezando con su publicacion en 1908 como Exo¬
gonium luteum House. Debido a la previa existen-
cia de una Ipomoea lutea Hemsley, se propone el
nuevo nombre Ipomoea electrina en referencia al
caracterfstico color arnbar que poseen las llores de
estas plantas. Dos eongeneres lejanamente rela-
cionados, Ipomoea lutea e Ipomoea urbinei, con his-
torias de nomenclatura y erroneas identificaciones
entrelazadas con Ipomoea electrina, tambien se dis-
cuten y comparan con este taxon.

Key words: biodiversity, convergence, Convol¬
vulaceae, Exogonium, Ipomoea, Mesoamerica,
morning glories. New World.

Homer I). House described numerous New
World Convolvulaceae (House, 1905, 1906a— d,
1907a— e, 1908a— d, 1909), among them Exogonium
luteum , a name and species that has proven con¬
troversial in recent years (Austin. 1978, 1983; Mc¬
Pherson. 1979). House (1908d) originally put the
species in Exogonium, a genus that he considered
to be distinct and closely related to Ipomoea, but
distinguishable on the basis of its exserted stamens
and style (i.e., ex o-gonia). New World morning glo¬
ries of this type, usually presenting yellow1, orange,
or red. salverform corollas, are generally associated
with bird pollination, or the occasional visitations
of butterflies (Austin, 1978, 1997; McDonald,

1987, 1991). Subsequent authors have recognized
that this ornithophilous syndrome has arisen on nu¬
merous occasions within New World Convolvula¬
ceae, especially within the genus Ipomoea s.h, in¬
cluding most of the original elements that House
assigned to Exogonium. Consequently, modern stu¬
dents of the family have considered the exogonioid
taxa largely as derivative species that should be
accommodated in various sections within Ipomoea
(Austin, 1997; McDonald 1987, 1991; McPherson,
1979). Some species of Exogonium sensu House
were retained in Ipomoea sect. Exogonium (Choisy)
Grisebach (McDonald, 1987), while others were
transferred to Ipomoea sect. Mina (Cervantes) Gri¬
sebach (Austin, 1978, 1980) and Ipomoea sect. Er-
iospermum Hallier f. (Austin, 1980), and yet others
to Turbina (Austin & Staples, 1991), and even
Ruellia (Austin & Wasshausen, 1973) of the Aean-
thaceae. Exogonium luteum requires the same con¬
sideration, though its legitimate transfer to Ipomoea
has yet to take place.

Taxonomic Problem

Austin (1978) maintained Exogonium luteum in
Ipomoea, but noted that the specific epithet of the
basionym could not be applied to the genus due to
the priority of Ipomoea lutea Hemsley. Austin
therefore proposed a new name for the species, /.
shinnersii, under which was also recognized Ipo¬
moea shinnersii var. woronovii (Standley) I). F. Aus¬
tin (1978). Unfortunately, the epithet woronovii in
fact has priority over the epithet shinnersii due to
its earlier publication date, and therefore renders
I. shinnersii as an incorrect name under Article
52.3 of the Code (Greater et al, 2000). Cater. Austin
(1983) corrected that mistake by creating /. woron¬
ovii (Standley) 1). F. Austin, while still interpreting
E. luteum as a synonym of I. woronovii.

McPherson (1979) also realized the misinterpre¬
tation, and proposed Ipomoea crocea as a new name
for Exogonium luteum in his dissertation. Unhap-

Novon 12: 27-30. 2002.

28

No von

pily, he did not formally publish this name, breed-
love (1986), apparently finding the McPherson
name on an annotation label, published it as a no¬
men nudum. Thus, the third attempt to provide a
name for these Mexican plants created yet another
unusable name.

McDonald (1987) later recognized Ipomoea wo¬
ronovii as a synonym of /. urbinei, but did not dis¬
cuss Exogonium luteum, having recognized that it
referred to a distinct, and indeed, distantly related
species. McDonald (1987) did not propose a new
name for House's species, as his treatment did not
discuss the group to which E. luteum belongs.
Thus, E. luteum has been recognized as an Ipomoea
by all modern students of the genus, even though
it still lacks a legitimate name.

Taxonomic Resolution

Because three distinctive species are involved
with this confusion, we present the nomenclature
and a brief description of each, to help clarify the
matter for our upcoming treatment of the Convol-
vulaeeae in Elora Mesoamericana.

I. Ipomoea lutea Hemsley, Diagn. PI. Nov. Mex-
ic. 2: 34, tab. 60. 1879, non Exogonium lu¬
teum House, Bull. Toney Bot. Club 35: 103.
1908. Quamoclit lutea (Hemsley) Hallier f.,
Bot. Jahrb. Syst. 16: 537. 1893. TYPE: Gua¬
temala. 0. Salvin & C. Godman (holotype, K
not seen, photo).

Ipomoea lutea f. rubra O'Donell. Lilloa 29: 67. 1959.
TV PE: Mexico. Chiapas: from Chicharras, alt. 3000—
6000 ft., 6 Feb. 1896. E. It Nelson .1768 (holotype,
(ill; isotype, US).

Twining herbs, probably perennial, stems ( limb¬
ing or prostrate, probably to 5 m, glabrous or some¬
times pilose on the nodes. Leaves 3—15 X 3—15
cm, ovate, entire, with undulate borders, irregularly
dentate, 3-lobed or 3-parted, glabrous or pilose on
the base, the base cordate, the apex acute to acu¬
minate, at times obtuse, mucronate. Inflorescences
in cymes, ± corymbiform. Flowers 4 to 14. rarely
solitary; sepals 2-3 mm, the outer ovate to elliptic,
glabrous, with subterminal arista 1—8 mm, glabrous
or finely pilose, the inner 3—4 nun, ovate, narrowly
elliptic to ± orbicular, obtuse to truncate or emar-
ginate, glabrous, borders hyaline, arista 3—7 mm,
glabrous or pilose; corolla 4—6 cm, ± tubular, yel¬
low, red, or orange, somewhat curved, the limb 5-
lobed, the lobes 4—6 mm long, narrow, obtuse, gla¬
brous; stamens 5.5—7 cm, exserted from corolla
tube 1 cm or more. Fruits 8—10 mm, globose, cap¬

sular; seeds I to 4, 4—4.5 mm, globose, brown, to-
mentose, with patches of longer clear trichomes.

Distribution. Forests; 1200—1500 m. Endemic
to southern Mexico and nearby Guatemala. Flow¬
ering December— February.

Following the taxonomic precedent set by
O’Donell (1959), we retain Ipomoea lutea in Ipo¬
moea sect. Mina on the basis of its highly diagnos¬
tic sepals, which present long and fleshy, subter¬
minal aristae. This taxon is the sister species of /.
hastigera Kunth, which is distinguished primarily
by its smaller corolla and pseudo-umbellate cymes.

Specimens examined. GUATEMALA. Kellerman 557
((41, US). MEXICO. Chiapas: Matuda 16190 (F, (.11.
MEXU, US).

2. Ipomoea urbinei House, Muhlenbergia 3: 41,
pi. 2, fig. 2. 1907. TYPE: Mexico. Colima:
slopes of Volcan de Colima, 1881. M. Bdrcena
214 (plate selected as lectotype by McDonald,
1987: 51).

Quamoclit tubulosa M. Martens & Galeotti, bull. Acad.
Roy. Sue. Bruxelles 12: 270. 1845. Ipomoea tubulosa
(M. Martens X Galeotti) Hemsley, Biol. Centr. Amer.
Bot. 2: 395. 1882, non Ipomoea tubulosa Roemer &
Schultes, Syst. Veg. 4: 789. 1819. Exogonium
uhdeanum Fenzl ex Hallier f., nom. illegit., Bot.
Jahrb. Syst. 16: 559. 1893. Ipomoea uhdeana (Fenzl
ex Hallier f.) I). F. Austin, nom. illegit., Ann. Mis¬
souri Bot. Card. 64: 332. 1978. TYPE: Mexico. Mi-
choacdn: “dans les champs d’Uruapan, a 4000 pieds,
FI. rouge,” II. Galeotti 1393 (holotype, BR: isotypes,
BR, P, W).

Ipomoea woronovii (Standley) I). F. Austin, Taxon 32: 626.
1983. Exogonium woronovii Standley, Field Mus.
Natl. Mist., Bot. Ser. II: 171. 1932. Ipomoea sbin-
nersii I). F. Austin var. woronovii (Standley) I). F.
Austin, Ann. Missouri Bot. Card. 64: 332. 1978.
TYPE: Mexico. Michoacan: G. ./. N. Worn note 2906
(holotype, l |.

Herbs, perennials; the stems woody at least near
the base, to 3 m, often purple-pigmented, mostly
glabrous. Leaves 4—7 X 3—5.5 cm, ovate or ovate-
elongate, glabrous or densely puberulent with mi¬
nute yellow trichomes on lower surface, the base
cordate and occasionally enveloping inflorescences
on fertile branches, the apex attenuate, mucronu-
late. Inflorescence in monochasial cymes. Flowers
2 to 5, with peduncles 1 .4—3.3 cm long, with pu¬
bescence like the leaves; sepals unequal, the outer
shorter, 3—4 mm, coriaceous, the inner 7—9 mm.
membranous, the apex acute to obtuse on outer,
obtuse to emarginate on inner, glabrous; corolla 3—
4 cm long, ± saberform, red, the tube almost con¬
stituting the whole perianth, the limb 5 distinct

Volume 12, Number 1
2002

Austin & McDonald
Ipomoea electrina

29

short triangulate lobes, 5—7 mm long; stamens ex-
serted, exceeding the limb by 3—5 mm. Fruits cap¬
sular, conical, I 1 — 12 mm long, 2-locular, 4-valvate;
seeds 4, 6-10 mm long, 4—5 mm wide, dark brown,
puberulent.

Distribution. Mountain forests; 1300-1640 in.
Endemic to Colima and Michoacan, Mexico. Flow¬
ering September— January.

Illustrations. House (1907d: 41, pi. 2, fig. 2);
McDonald (1987: 80, fig. 9b).

Hallier created an illegitimate superfluous name
because he included only indirectly (he did not cite
the basionym directly, only the combination based
on it) the type of the earlier name Quamoclit tub-
ulosa M. Martens & Galeotti. In other words, Hal¬
lier should have used the available epithet tubu-
losum instead of adopting the herbarium name
written by Fenzl on the sheet as a determination.
Therefore, Ipomoea uhdeana (Fenzl ex Hallier f.)
D. F. Austin was based on the illegitimate basionym
Exogonium uhdeanum Fenzl ex Hallier 1. Since us¬
age of an illegitimate basionym is prohibited by the
Code (Greuter et al., 2000), we use the next avail¬
able name. Ipomoea tubulosa (M. Martens & Gal¬
eotti) Hemsley cannot be used because Ipomoea
tubulosa Roemer & Schultes has priority over this
binomial. In the absence of specimen at MFXU,
McDonald (4987) designated the plate as the lec-
totype.

As indicated by McDonald (1987). Ipomoea ur-
binei is a member of I. sect. Exogonium. The spe¬
cies shares many traits, including unequal, acute,
membranaceous sepals, salverform corolla, and ex-
serted stamens, with I. dumosa (Bentham) F. 0.
Williams and that is surely the sister taxon.

Additional specimen. MEXICO. Michoacan: G. Hin¬
ton et al. 12254 (NY. TEX, US).

3. Ipomoea electrina D. F. Austin & J. A.
McDonald, nom. nov. Replaced name: Exo¬
gonium luteum House, Bull. Torrey Bot. Club
35: 103. 1908, non Ipomoea lutea Hemsley,
Diagn. PI. Nov. Mexic. 2: 34, t. 60. 1879. Ip¬
omoea shinnersii D. F. Austin, nom illegit. sed
non superfl. [see Taxon 32: 626. 1983], Ann.
Missouri Bot. Card. 64: 337. 1977 [1978]. Ip¬
omoea woronovii (Standley) D. F. Austin var.
lutea (House) I). F. Austin, Taxon 32: 626.
1983. TYPE: Mexico. Oaxaca: Cuesta de Chi-
quihuetlan, 3300 ft. alt.. 2 Sep. 1895, C. Con-
zatti & Gonzalez 668 (holotype, GH; isotype,

NY).

Herbs, perennial; the stems woody at the base.

herbaceous toward the apex, reaching 2—3 m or
more, densely or sparsely pubescent. Leaves 4—9
X 2.5—7 cm, ovate, pubescent above, the base cor¬
date, the apex acuminate. Inflorescence in dicha-
sial cymes. Flowers 3—18, with peduncles 1.5—
4(10) cm. with pubescence like the leaves; sepals
unequal, the outer 5—6 mm, the inner 6.6— 8.5 mm,
coriaceous, often verrucose at least on the base, the
apex acute on the outer, obtuse on the inner; corolla
5—6.5 cm, hippocrateriform, yellow or orange-gold,
the tube 3.5-4 X 4—6 mm, the limb with 5 distinct
linear lobes 1.5— 2.3 cm long; stamens exserted >

5 mm. Fruits seen only in immature stage, conic,
2-locular, 4-valvate; seeds dark brown, pubescent
with long, brown trichomes.

Distribution. Mesophyllous forests, pine-oak
forests, low deciduous forests, oak forests; 700—
2100 m. Endemic to Mexico. Flowering July— Sep¬
tember. January.

Illustrations. House (1908b: pi. 2. fig. c).

Common name. Erijol de la virgen (Oaxaca);
(lowers are said to be edible.

Etymology. The epithet electrina refers to the
“amber” color (yellow-orange) of the corollas. The
base electrinus, -a, -urn derived by combining the
noun electrum (Greek elektron, usually meaning
amber), with the Greek adjectival suffix -inns, -a, -
um indicating the color of amber. Information on
orthography was extracted partly from Nicolson and
Brooks (1974).

Ipomoea electrina bears only a superficial resem¬
blance to I. urbinei. House described and illustrat¬
ed the species as having linear limb segments,
much as in agreement with the type specimen. This
feature contrasts dramatically with the much re¬
duced, scarcely triangulate limb segments of I. ur¬
binei (see McDonald, 1987: 80, fig. 9b). Moreover,
the corollas of /. urbinei are red, with stamens ex¬
serted but a few millimeters beyond the plane of
the limb. In contrast, corollas of I. electrina are
bright yellow-orange, the stamens and style ex¬
ceeding the corolla by more than 5 mm. In all like¬
lihood I. electrina is a relative of /. conzattii Green-
man and allies, belonging to Ipomoea sect.
Eriospermum. With them, it shares the long woolly
trichomes on the margins of the seeds.

Specimens examined. MEXICO. Oaxaca: Distr. Te¬
huantepec, Cerro Marimba, Martinez 1035 (MEXU); S del
Cerro Guiengola. Torres & Cabrera 6 310 (MEXU). Torres

6 Torres 194 (MEXU). Torres A' Torres 202 (MEXU); Santa
Lucia, Martinez 133 (MEXU); SO de Buenos Aires, Torres
& Martinez 7349 (MEXU); San Carlos Yautepee, SE de el
Cameron. Torres & Martinez 12582 (MEXU); Santiago Ea¬
ch iguiri, Cerro de Las Flores, Campos 3803 (MEXU).
Chiapas: Purpus 9189 (US).

30

Novon

Acknowledgments. We thank Fernando Cliiang
(MKXU), Gerrit Davidse (MO), Guillermina Mur-
guia S. (MKXU), Dan Nicolson (US), Mario Sousa
(MKXU), and Rafael Torres C. (MKXU) for nomen-
elatural and editing comments, and information re¬
garding these plants. Ingrid Curril (Florida Atlantic
University) kindly provided the Spanish version of
the abstract.

Literature Cited

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Kxogonium (Convolvulaceae). Ann. Missouri Rot . Card.
64: 330-339.

- . 1980. Additional comments on infrageneric taxa

in Ipomoea (Convolvulaceae). Taxon 29: 501—502.

- . 1983. New combinations in Mexican Ipomoea

(Convolvulaceae). Taxon 32: 625, 626.

- . 1997. Dissolution oi Ipomoea ser. Anisomerae

(Convolvulaceae). Bull. Torrey Bot. Club 124: 140—159.
- & G. W. Staples. 1991. A revision of the neotrop¬
ical species of Turbina Raf. (Convolvulaceae). Bull. Tor¬
rey Bot. Club. I 18: 265-280.

- & 1). C. Wasshausen 1973. A Ruellia (Acantha-

ceae) species in Kxogonium (Convolvulaceae). Phyto-
logia 25: 433— 137.

Breedlove, 1). E. 1986. Convolvulaceae. Pp. 74—77 in Lis-
tados Florfsticos de Mexico IV. Flora de Chiapas. In¬
stitute de Biologfa, Universidad Nat ional Autonoma de
Mexico, Mexico ( lily.

Grenier, W.. J. McNeill. F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland X I). L. Hawks-
worth (editors). 2000. International Code ol Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
House, H. D. 1905. Two new species of Convolvulus from
the western United States. Bull. Torrey Bot. Club 32:
139-140.

- . 1906a. A new southern Convolvulus. Torrey a 6:

149, 150.

- . 1906b. A new species of Dichondra. Muhlenber-

gia 1: 130-131.

- . 1906c. Studies in the North American Convol¬
vulaceae — I. Bull. Torrey Bot. Club 33: 313—318.

- . I906d. Studies in the North American Convol¬
vulaceae — II. The genus Operculina. Bull. Torrey Bot.
Club 33: 495-503.

- . 1907a. A new species of Evolvulus from Colom¬
bia. Torreya 7: 61—62.

- . 1907b. New species ol Ipomoea from Mexico and

Central America. Muhlenbergia 3: 37—16. pi. 31—33.

- . 1907c. Studies in the North American Convol¬
vulaceae — III. Bull. Torrey Bot. Club 34: 143—149.

- . 1907d. New species of Ipomoea from Mexico and

Central America. Muhlenbergia 3: 37—46.

- . 1907e. New or noteworthy North American Con¬
volvulaceae. Bot. Caz. 43: 408—414.

- . 1908a. Two Bahamian species of Evolvulus. Bull.

Torrey Bot. Club 35: 89-90.

- . 1908b. Studies in the North American Convol¬
vulaceae — IV. Bull. Torrey Bot. Club 35: 97—107, pi.
101-102.

- . 1908c. The North American species of the genus

Ipomoea. Ann. New York Acad. Sci. 18: 181—263.

- . 1908d. Studies in the North American Convol¬
vulaceae — IV. The genus Kxogonium. Bull. Torrey Bot.
Club 35: 97-107, pi. 1-2.

- . 1909. Studies in the North American Convolvu¬
laceae — V. Bull. Torrey Bot. Club 36: 595—603.

McDonald, J. A. 1987. Revision of Ipomoea section Kxo¬
gonium (Choisy) Griseb. (Convolvulaceae). Brenesia 28:
41-87.

- . 1991. Origin and diversity of Mexican Convol¬
vulaceae. Anales Inst. Biol. Univ. Auton. Mexico, Ser.
Bot. 62: 65—82.

McPherson, G. 1). 1979. Studies in the Genus Ipomoea
(Convolvulaceae). University of Michigan University
Microfilms, Ann Arbor.

Nicolson. D. H. & R. A. Brooks. 1974. Orthography of
names and epithets: Stems and compound words. Faxon
23: 163-177.

O’Donell, C. A. 1959. Las especies americanas de Ipo¬
moea L. sect. Quamoclit (Moench) Griseb. Lilloa 29:
1 9-86.

Una Nueva Especie Sudamericana del Genero Rytidosperma
(Poaceae: Arundinoideae: Danthonieae)

Carlos Baeza

Departamento de Botanica, Universidad de Concepcion, Casilla 160-C, Concepcion, Chile.

cbaeza@udec.cl

RKSUMEN. Se describe e ilustra una nueva espe¬
cie de Rytidosperma (R. quirihuense; Poaceae:
Arundinoideae: Danthonieae) para sudamerica.
Esta especie crece en el centro-sur de Chile
(36° Id'S, 72°45'W). Morfoldgicamente es muy di-
ferente al resto del grupo sudamericano. Solo pre-
senta afinidades feneticas con Rytidosperma pas-
chale, de la cual se diferencia por la panicula
distica, por el tamano de la espiguilla terminal, por
la densidad y longitud de pelos de los fascfculos
de la lemma, entre los caracteres mas importantes.

Abstract. A new species of Rytidosperma ( R.
quirihuense; Poaceae: Arundinoideae: Danthonieae)
from south-central Chile (36°13'S, 72°45'W) is tle-
scribed and illustrated. It differs morphologically
from other South American representatives of the
genus. Its closest phenetic affinities are with Ryti¬
dosperma paschale, from which it differs in posse¬
sion of a distichous panicule, as well as differences
in size of the terminal spikelet, and in the density
and length of hairs on the fascicules of the lemma.

Key words: Arundinoideae, Chile. Danthonieae,
Poaceae, Rytidosperma.

El genero Rytidosperma Steudel comprende unas
45 especies, la mayorfa de ellas distribuidas en
Australia. Nueva Zelandia y Tasmania (Nicora &
Rugolo, 1987). En el continente americano crecen
8 taxa, los cuales se encuentran en Chile y Argen¬
tina. En Chile la distribucidn abarca desde la pro-
vincia de Liman en la IV Region hasta la provincia
de Ultima Esperanza en la XII Region, incluvendo
a Rytidosperma paschale (Pilger) C. Baeza [publi-
cado como Rytidosperma paschalis (Pilger) Baeza).
especie endemica de Isla de Pascua (Baeza. 1990).
Numerosos son los autores que ban contribuido al
conocimiento de este genero, ya sea a uivel mor-
lologico, anatomico, citologico, biogeografico, fe-
netico y filogenetico (Nicora, 1973; Connor X Ed¬
gar. 1979; Baeza, 1996a, 1996b, 1997. 1998;
Linder & Verboom, 1996; Hsiao et al., 1998; Bark¬
er et al.. 2000; Linder & Barker. 2000). Durante el
desarrollo del proyecto “Analisis de la flora vas¬
cular de la zona de transition climatica mediter-

raneo-templada de Chile (36°31'S, 38°30'S): dis-
tribucion de la riqueza taxonomica y determinat ion
de areas prioritarias” se recolecto material de esta
nueva especie.

Para la description de la nueva especie todas las
metlidas referidas al antecio se hicieron consider-
ando el ultimo de la espiguilla terminal, debido a
que hay una gran variation dentro de la espiguilla,
en cuanto al tamano tie este. Para la description
anatomica tie la lamina se considero la zona media
tie la penultima innovacion esteril siguientlt) los
criterios propuestos por Baeza (1997).

Rytidosperma quiriliuense C. Baeza, sp. nov.
TIPO: Chile. Nuble: Camino Las Achiras hacia
Quirihue (36°13'S, 72°45'W), 355 m, 20 Nov.
2000, C. Baeza, P. Ldpez & M. Parra 2112
(holotipo, CONC; isotipo, SCO). Figura 1.

Planta perennis, culmi 2—1 nodi. 25^45 cm alti. Inno-
vationes intravaginales. Ligulae dense ciliatae, ciliis 0.2—
0.4 mm longis. Paniculae distichae, 4.5-8 cm longae;
spiculae 5— 1 9-flores; glumae lanceolatae, subaequales vel
inferior superiorem paullo excedens, plerumque 10.5—14
mm longae, 5— 6-nervatae; lemma in dorso pilorum serie-
bus duobus distinctis, serie inferiore prope supra callum,
1.5—2 mm longa, serie superiore 0.7—2 mm longa; lem-
matis corpus 11—13 mm longum 1 — 1.2 mm callum int lu-
tlens: lobi laterales 6.5—8 mm longi, setae 3— ^ 1 mm longae;
arista tirma, 12-16 mm longa; palea firma, lanceolata, 5-
6.5 X 1.4— 1.8 mm longa, bidenticulata, in carinis supra
minute ciliolata; antherae 0.8— 1.8 mm longae, luteae.
Carvopsis 2 X 0.8—1 mm. fulva; hilum punctiforme, ova-
tum, 0.6 mm longum.

Planta perenne, eanas florfferas 2 a 4 nodes, tie
25^45 cm tie altura. Innovaciones intravaginales.
Hojas basales de hasta 10 cm de largo, pilosas.
Vainas pilosas, con mechones de pelos de hasta 3
mm a ambos costados tie la h'gula. Ligula pestan-
osa, con pelos tie 0.2— 0.4 mm. Laminas pilosas ra-
las, tie hasta 9 cm de largo por 1.5—2 mm de aneho.
Panicula distica, tie 4.5—8 cm tie largo, estrecha,
con 4 a 10 espiguillas. Pedicelos pubescentes, tie
1—10 mm de longitud. Espiguillas 5- a 9-floras, sin
considerar el ultimo antecio esteril, la espiguilla
terminal tie 2-2.4 cm de largo. Glumas verdosas,
lanceoladas, 5- 6 6-nervadas, la inferior tie 10.5—

Novon 12: 31-34. 2002.

32

Novon

l igura I. Rytidosperma quirihuense C. Baeza. — A. Planta. — B. Lemma. — C. Palea. — I). I.odfenlas. — E. Espiguilla.
— F. Epidermis abaxial lie la lamina. — G. Corte transversal de la lamina. Basado en el holotipo C. Baeza. B. Ldpez &
M. Barra 2112 (CONG).

Volume 12, Number 1
2002

Baeza

Rytidosperma quirihuense de Chile

33

14 mm tie longitud, la superior de 1 1-14 mm, ma-
yores que el conjunto de los antecios. Antecio in¬
ferior desde el eallo hasta las aristas de los lobulos
de 1 1 — 13 mm y desde el eallo liasta el apice de la
arista dorsal de 17—21 mm, eallo piloso, de 1—1.2
mm, pelos de 0.3-1. 2 mm, arista dorsal con la base
geniculada, de 12—16 mm de longitud. Lemma pi-
losa en el dorso, con pelos fasciculados dispuestos
en dos hileras paralelas Lien definidas, la superior
con 6 fasciculus, cuatro laterales y dos dorsales, los
pelos de 0.7—2 mm de largo, los fascfculos dorsales
con una baja densidad de pelos, a veces caedizos,
los laterales siempre bien visibles y persistentes.
Hilera inferior eompuesta de 10 fascfculos de pelos
persistentes, muy densos, de 1 .3—2 mm de largo,
sin pelos ralos entre ambas hileras de fascfculos
transversales. Lema desde su base (sin incluir el
eallo) liasta la insercion de la arista dorsal de 3—4
mm. Lblnilos laterales de 6.5—8 mm. las aristas de
3—4 mm de largo. Palea lanceolada, de 5—6.5 X
1.4— 1.8 mm, carinas finamente ciliadas, base del
dorso con pelos ralos, cortos, a veces caedizos en
la madurez, apice bidenticulado. Lodfculas 2, pi-
losas, cuneadas, de 0.4— 0.6 mm, con pelos de 0.2—
0.6 mm. Estambres 3, anteras de 0.8-1. 8 mm. Ova-
rio de 1-1.5 mm. Estigmas de 1.5—2 mm. Cariopsis
elfptica, castana, de 2 X 0.8—1 mm. Embrion de
0.8—1 mm. Hilo punctiforme, ovalado, de 0.6 mm.

Epidermis abaxial de la lamina. Celulas largas
intercostales con paredes onduladas, de 40—146 X
10—18 p.m; estomas redondeados a algo romboi-
dales, numerosos; celulas cortas intercostales pre-
sentes; pelos bicelulares presentes, Irecuentes, con
la celula apical de menor tamano que la celula bas¬
al; macropelos presentes, frecuentes; celulas inter-
estomaticas de 40—75 X 12—18 pm. Celulas cos¬
tales con paredes onduladas iguales a las
intercostales, de 30—210 X 6—14 pm, con celulas
sflico-suberosas alternadas, las silfeeas redondea-
das a algo halteriformes, de 4—23 X 6—15 pm, las
suberosas rectangulares, de 2—12 X 8—17 pm.
Aguijones presentes, numerosos, ubicados en los
nuirgenes de la lamina. Figura I F.

Carte transversal de la lamina. Contomo de la
lamina en forma de U abierta. Haces vasculares de
primer orden 7, trabados, con bandas de tejido es-
clerenquimatico conectado a ambas epidermis, los
de segundo orden 2, con un casquete de tejido es-
clerenquimatieo en la epidermis adaxial, no conec¬
tado a la vaina parenquimatiea, los de tercer orden
4, libres, sin esclerenquima. Tejido esclerenqui-
matico discontinue en ambas epidermis. Celulas
buliformes en forma de abanico, 2 6 3 veces ma-
yores que las celulas epidermicas, disminuyendo
en tamano hacia los lados. Figura 1C.

Distribution y habitat. Esta especie habita en
la Cordillera de la Costa de Chile centro-sur, en la
localidad de Las Achiras, camino hacia (^uirihue,
Provincia de INuble, VIII Regidn. El suelo es de
naturaleza arcilloso. Esta nueva especie erece en
sectores donde aun permanecen fragmentos de
bosque nativo y flora herbacea y arbustiva asocia-
da. La poblacion observada no sobrepasaba los 20
individuos. En la actualidad, esta localidad esta al-
tamente intervenida por plantaciones artificiales de
Dinas radiata D. Don y Eucalyptus globulus Labil-
lardiere. Solo ha sido colectada en la localidad tipo.
Sin embargo, es muy probable que otras pobla-
ciones de la misma especie esten creciendo en sec-
tores cercanos donde aun hay fragmentos de veg-
etacion nativa. Esta zona de la Cordillera de la
costa ha sido historicamente poco colectada, lo que
se demuestra por el bajo numero de colectas de-
positadas en los herbarios de CO INC y SGO.

Morfologieamente, esta especie es muy diferente
al resto de los taxa que erecen en Sudamerica con¬
tinental. La panfcula dfstica, la morfologfa de la
lemma, la distribucion de los fascfculos de pelos,
su numero y densidad son caracteres propios de
esta especie, que la separan completamente del
resto. Solo Rytidosperma paschale, especie insular,
presenta algunas similitudes, sobre todo respecto a
la densidad de pelos en el dorso de la hilera su¬
perior de la lemma y en la poea pilosidad de la
palea, sin embargo es lacilmente diferenciable de
R. quirihuense porque las espiguillas terminales no
sobrepasan los 1.5 cm de longitud, no posee una
panfcula dfstica, por la baja densidad de pelos en
la hilera inferior de la lemma, por la prcsencia de
pelos bicelulares con la celula apical de mayor la-
mano que la basal, por el contorno de la lamina en
forma dc V, convoluto, por presentar solo 3 haces
vasculares de primer orden, por la ausencia de ha¬
ces vasculares tie segundo orden, etc. En R. qui¬
rihuense, la seccion transversal de la lamina pre¬
senta una gran cantidad de haces vasculares de
primer orden trabados como tambien presencia de
haces vasculares de tercer orden, lo que se repite
en el resto de las especies americanas. Las celulas
buliformes mayores en tamano que las celulas ep¬
idermicas es un earacter que tambien esta presente
en Rytidosperma paschale, R. sorianoi Nicora y R.
virescens (E. Desvaux) Nicora var. parvispiculum Ni¬
cora. La epidermis abaxial presenta las mismas car-
acterfsticas de las otras especies sudamericanas,
esto es celulas largas intercostales de paredes on¬
duladas, macropelos y pelos bicelulares presentes,
celulas suberosas rectangulares, etc. La forma re-
dondeada de las celulas silfeeas es un earacter que
se repite en todas las especies del genero, aunque

34

Novon

en Rytidosperma quirihuense tienden a ser algo hal-
teriformes, situat ion tambien observada en R. pas-
dude.

Clave para Rytidosperma quirihuense y las Especiks
SUDAMERICANAS (MoDIKICADA l)E B AEZA, 1996a)

la. Hilera superior de fascfculos pilosos de la lemma
con 2—6 haces de pelos.

2a. I ’an feu la dfstica; espiguilla terminal de 2-
2.4 cm de longitud; planta del continente

. R. quirihuen.se

2b. Panfcula no dfstica; espiguilla terminal de
1.1— 1.5 cm de longitud; planta de Isla de

Pascua . R. paschale

1 1). Hilera superior de fascfculos pilosos de la lemma
con 8—10 haces de pelos.

3a. Glumas linear-lanceoladas, muy agudas y

violac-eas . . . R. violaceum (E. Desvaux) Nicora
3b. Glumas lanceoladas.

4a. Hilera inferior de fascfculos pilosos de
la lemma poco definidos, con baja den-
sidad de pelos, a veces sblo hay pelos
ralos.

5a. Apiee de la palea bimucronado,

plantas muy pilosas .

... R. pictum (Nees & Meyen) Nicora
var. bimucronalum Nicora
5b. Apiee de la palea obtuso, 2-4-den-
ticulado, t rune ado o algo agudo, las
carinas nunca prolongadas en mu-
crones, plantas glabras a pilosas ra-

las . R. pictum var. pictum

4b. Hilera inferior de fascfculos pilosos de
la lemma bien definidos, con alta den-
sidad de pelos.

6a. Palea de 2.3—1 mm de largo, aristas
de los ldbulos de la lemma de 0.2-
2 mm, largo de la lemma desde el
eallo hasta las aristas de los ldbulos
de 3.6-7(-8) mm.

7a. Apiee de la palea agudo, vain-

as muy pilosas . R. sorianoi

7b. Apiee de la palea obtuso o lev-
emente bident iculado, vainas
glabras, muy raro con pelos en
los mdrgenes R. lechleri Steudel
6b. Pdlea de (4— )4.7— 6.5 mm de largo,
aristas de los lobulos de la lemma
de 1.8— 5(— 5.8) mm, largo de la
lemma desde el eallo hasta las aris¬
tas de los ldbulos de 7.5— 14( — 15)
mm.

8a. Apiee de la palea obtuso a
subtridenticulado, vainas pilo¬

sas a pilosas ralas .

. . . R. virescens var. parvispiculum
8b. Apiee de la pdlea 2— 3-denti-
culado, vainas glabras ....
. R. virescens var. virescens

Agradecimientos. Se agradece el apoyo presta-
do por el p rover to Fondecyt N° 1000364 y por la
Direction de Investigation de la Universidad de
Concepcion, DIUC N° 99.111.020-1-3 y DIUC
97.111.017—1.0. Un agradecimiento especial a Pe¬
dro Arias por la confeccion de los dibujos.

Literatura Citada

Baeza, C. 1990. Rytidosperma paschalis (Pilger) Baeza,
una nueva combinacidn para la flora agrostoldgica de
Chile. Gayana, But. 47(3-4): 83—84.

- . 1996a. Los generos Danthonia DC. y Rytidosper¬
ma Steud. (Poaceae) en America. Una revisidn. Sendt-
nera 3: 11-93.

- . 1996b. Numero de cromosomas en algunas es-

pecies chilenas de Danthonia DC. y Rytidosperma
Steud. (Poaceae). Gayana, Bot. 53(2): 329-333.

- . 1997. Anatomfa foliar y epidermis ahaxial de las

especies americanas de Danthonia DC. y Rytidosperma
Steud. (Poaceae). Gayana, Bot. 54(1): 61—88.

- . 1998. Analisis fenetico y de componentes prin-

cipales en las especies sudamericanas de Rytidosperma
Steud. (Poaceae). Gayana, Bot. 55(1): 1 1-15.

Barker, N., C. Morton & H. binder. 2(KK). The Danthon-
ieae: Generic composition and relationships. Pp. 22 1 —
2.30 en S. W. L. Jacobs & J. Everett (editors). Grasses:
Systematies and Evolution. CS1RO, Melbourne.

Connor, H. E. & E. Edgar. 1979. Rytidosperma Steudel
( Notodanthonia Zotov) in New Zealand. New Zealand J.
Bot. 17: 31 1-337.

Hsiao, C., S. W. Jacobs, N. P. Barker & N. J. Chatterton.
1998. A molecular phylogeny of the subfamily Arun-
dinoideae (Poaceae) based on sequences of rl)NA. Aus¬
tral. Syst. Bot. II: 41-52.

binder, II. <X N. P. Barker. 2000. Biogeography of the
Danthonieae. Pp. 231— 238 en S. A. h. Jacobs & J. Fiv-
erett (editors). Grasses: Systematies and Evolution. CSI-
RO. Melbourne.

- & A. Verboom. 1996. Generic limits in the Ryti¬
dosperma (Danthonieae, Poaceae) complex. Telopea 6:
597-627.

Nicora, E. G. 1973. Novedades agrostologicas patagoni-
cas. Darwiniana 18: 80—106.

- & Z. Rugolo. 1987. Los generos de gramfneas de

America Austral. Argentina, Chile, Uruguay y dreas
limftrofes de Bolivia, Paraguay y Brasil. Buenos Aires.
Ed. Hemisferio Sur, Buenos Aires.

Senecio albaniae (Asteraceae: Senecioneae), a New Species from

Central Peru

Hamilton Beltran

Museo <le Historia Natural, Universidad National San Marcos, Av. Arenales 1256,

Apartado 14-0434, Lima, Peru

Abstract. A new species, Senecio albaniae (As¬
teraceae: Senecioneae), is described from central
Peru. The species is characterized by its shrubby
habit, decurrent leaf bases forming winged stems,
and solitary capitula on elongated peduncles. The
species is described and illustrated, and its possi¬
ble relationships are discussed.

Rksumkn. Una especie nueva, Senecio albaniae
(Asteraceae: Senecioneae), se describe del Peru
central. La especie es caracterizada por su habito
arbustivo, hojas decurrentes y tallos alados, capi-
tulo solitario con pedunculos 5-15 cm de longitud.
Se describe e ilustra la especie, y discuto las re-
laciones potenciales de esta especie.

Key words: Andes, Asteraceae, Peru, Senecio,
Senecioneae.

Senecio L. is an important genus of the South
American Andes, which is a center of diversity for
the genus. Senecio is found from coastal deserts to
alpine habitats in the Andes. The genus is repre¬
sented by ca. 180 species in Peru (Dillon & Hen-
sold. 1993), several of which are endemic and of
limited geographic distribution (Vision & Dillon,
1996). Recently two new species have been added
(Beltran & Galan De Mera. 1997. 1998). A species
of Senecio with decurrent leaf margins and winged
stems was discovered among unidentified Seneci¬
oneae housed at the Herbarium of the Universidad
de San Marcos in Lima (USM). Subsequent field
studies in the approximate region of the first col¬
lection confirmed the species as new.

Senecio albaniae H. Beltran, sp. nov. TV PE:
Peru. Lima: Huarochiri, San Pedro de Casta.
Mashca, Camino del pueblo San Pedro de Cas¬
ta hacia Mareahuasi, 1 1°46'S, 76°35'W, 3680
m, 21 mayo 2000, //. Beltran 3474 (holotype,
USM; isotypes, F, HAO, HUT, MO, NY, S.
TEX, US). Figure 1.

Haec species S. calcencis Cabrera & Zardini etiam .S’.
mollendoensis Cabrera primo viso similis, sed a hoc lam¬
ina foliari basi exauriculata marginibus integra, ab illo

capitulis majoribus pedicellis longioribus insidentibus, ab
ambobus foliis decurrentibus distinguitur.

Shrubs to 0.6 m high; stems much branched,
sparsely tomentose and winged from the decurrent
leaf bases, the wings 2—3 mm wide. Leaves alter¬
nate, simple, sessile, lamina membranaceous, lan¬
ceolate to oblanceolate, 6-9 X 0.5-1 cm, apex acu¬
minate, basally decurrent to 1-1.5 cm; adaxially
dark green and arachnoid tomentose, abaxially pale
green and densely tomentose, pinnately veined,
margins denticulate. Capitulescence lax, with 1 or
2 branches terminating in capitula from the upper¬
most nodes; peduncles 5—15 cm long, finely striate,
arachnoid tomentose, with bracteoles ca. 7 per pe¬
duncle, lanceolate, 15-35 X 0.5-1. 5 mm. apex
long attenuate distally. Capitula radiate: calycular
bracts ca. 9, linear, 9-11 X 0.5-0. 8 mm, apex at¬
tenuate; involucre campanulate to hemispherical,
9—10 X 10—13 mm diam.; phyllaries 12 to 14, un-
iseriate, subequal, chartaceous, lanceolate, 10—12
X 1-3 mm, free, dorsally arachnoid tomentose, in¬
side glabrous, apex acute, briefly short-pubescent,
margins scarious; ray florets 9 to 12, pistillate, well
exserted from the involucre; corolla yellow, the tube
linear, 5—6 mm long, glabrous, the limb oblong, 15—
20 X 2-3 mm, flat, glabrous, 4- or 5-nerved, apex
bi- or tridentieulate; disk florets 55 to 70. bisexual,
corolla yellow, tubular, 12—14 mm long overall,
tube 5-6 mm long, the limb 3-4 mm long, narrowly
campanulate, apically 5-dentate, teeth triangular,
1 — 1.2 X 0.4 mm, apex acute and slightly thick¬
ened: anthers ca. 4 mm long, base obtuse, with api¬
cal appendage 0.4-0. 5 mm long; style branches re¬
curved. 2.5 mm long, with papillate stigmatic lines
marginally on inner surfaces, apex truncate with a
small crown of penicillate trichomes. Achene (im¬
mature) cylindrical, 3—4 X 0.7— 0.9 mm, densely
sericeo-papillose; pappus biseriate, persistent,
white, 6-7 mm long with bristles equivalent in
length, strigillose.

Distribution and ecology. Senecio albaniae is
known only from the type locality, San Pedro de
Casta in the province of Huarochiri, Lima. Peru, on

Novon 12: 35-37. 2002.

36

Novon

figure 1. Senecio albaniae II. Beltran. — A. !• lowering branch. — B. Enlarged leal with decurrent base. — C. Bay
floret. — I). Disc floret. (Drawn from ./. Alban & N. Malca 11339, USM.)

Volume 12, Number 1
2002

Beltran

Senecio albaniae from Peru

37

the western escarpment of the Andean Cordillera
and shrubby communities that include Mutisia ac¬
uminata Ruiz & Pavon, Ophryosporus peruvianas
(Gmelin) King & Robinson, Senecio cerratei Cua-
trecasas, Senecio richii A. Gray, Senecio subcandi-
dus A. Gray (all Asteraceae), and Opuntia subulata
(Muehlenpfordt) Engehnann (Cactaceae).

Etymology. This new species is named lor its
first collector, Joaquina Alban, a Peruvian ethno-
botanist at the Universidad de San Marcos.

Discussion. This is the only known South
American species of Senecio with winged sterns.
This character, however, is also known from the Pe¬
ruvian Caxamarca sanchezii M. 0. Dillon & Sagas-
tegui in a newly described monotypic genus iu the
Senecioneae (Dillon & Sagastegui, 1999).

Since there is no modern account of the entire
genus, relationships among species of Senecio re¬
main speculative; therefore, I am not able to fit the
new species within any of the infrageneric catego¬
ries established by Cabrera (1985) and Jeffrey
(1992) for Andean taxa of Senecio. However, it
could be placed tentatively into the series Otopteri
(Cabrera, 1985) through the link with Senecio ban-
gii Rusby, a superficially similar Bolivian plant.
Curiously and according to my observations, S.
bangii Rusby is an anomalous member of the se¬
ries.

Clearly, S. albaniae would appear closest to a
Peruvian natural complex made up of S. icaensis H.
Beltran & Galan de Mera, S. abadianus DC., 5.
arnaldii Cabrera, S. okopanus Cabrera, S. lomin-
cola Cabrera, S. yauyensis Cabrera, S. subcandidus
A. Cray, 5. tovarii Cabrera, S. truxillensis Cabrera.
S. calcensis Cabrera & Zardini, and S. mollendoen-
sis Cabrera. Probably, 5. bangii Rusby could find a
better position within this latter complex.

In Peru, Senecio albaniae superficially resembles
S. calcensis Cabrera & Zardini (1975), a species
with shorter peduncles and smaller capitula, and S.
mollendoensis Cabrera (1962), a species with upper

leaves with dilated and auriculate bases and
coarsely dentate margins.

Paratype. PERU. Lima: Huarochiri, San Pedro de
Casta, Path Chaniulpo-Quinual 1 l°45.734'S,
76°35.581'W, 3200-3500 m, 21 May 1999, ./. Alban &
N. Malca 11339 (F, MO, USM, NY).

Key to Senecio albaniae and Related Peruvian Spe¬
cies

la. heaves deeurrenl on stems . S. albaniae

lb. Leaves not decurrent on stems.

2a. Basal leaves sessile, auriculate, the margins

coarsely dentate . S. mollendoensis

2b. Basal leaves with short petioles, not auric¬
ulate, the margins finely dentate . . S. calcensis

Acknowledgments. I thank Guillermo Pino for
the Latin diagnosis, Maximilian Weigend and Ar¬
turo Granda for help, and Michael Dillon for criti¬
cally reviewing an early draft of the manuscript.

Literature Cited

Beltran, H. & G. De Mera. 1997. Senecio icaensis sp. nov.
(ASTERACEAE), un nuevo endemismo de las lomas
costaneras del Peru. Anales Jard. But. Madrid 55: 168—
170.

- & - . 1998. Senecio larahuinesis sp. nov.

(ASTERACEAE,), una nueva espeeie de los andes per-
uanos Peru. Anales Jard. Bot. Madrid 56: 168—169.
Cabrera, A. L. 1962. Compuestas andinas nuevas. Bob
Soc. Argent. Bot. 10: 21^45.

- . 1985. El genero Senecio (Compositae) en Bolivia.

Darwiniana 26(1-4): 79—217.

- & E. M. Zardini. 1975. Especies nuevas o criticas

del gdnero Senecio (Compositae). Bob Soc. Argent. Bot.
16: 377-389.

Dillon, M. & N. Hensold. 1993. Family Asteraceae. In L.
Brako & J. L. Zarucchi (editors). Catalogue of the Flow¬
ering Plants and Gymnosperms of Peru. Monogr. Syst.
Bot. Missouri Bot. Gard. 45: 103-189.

- & A. Sagastegui. 1999. Caxamarca, a new mono¬
typic genus of Senecioneae (Asteraceae) from northern
Peru. Novon 9: 156—161.

Jeffrey, C. 1992. The tribe Senecioneae (Compositae) in
the Mascarene Islands witli an annotated world check¬
list of the genera of the tribe; Notes on Compositae: VI.
Kew Bull. 47: 49-109.

V'ision, T. J. & M. Dillon. 1996. Sinrtpsis de Senecio L.
(Senecioneae, Asteraceae) para el Peru. Arnaldoa 4:
23-46.

Bernardia fonsecae (Euphorbiaceae), a New Species from

Guerrero, Mexico

Angelica Cervantes M.

Departamento <le Botdnica, Instiluto de Biologfa, UNAM, Apartado Postal 70-233,

Del. Coyoacan, 04510, Mexico, D.F.

Jaime Jimenez Ramirez

Facultad de Ciencias, UNAM, Apartado Postal 70-399, Del. Coyoacan, 04510, Mexico, D.F.

ABSTRACT. Bernardia fonsecae, endemic to Guer¬
rero, Mexico, is described and illustrated, and hab¬
itat data are provided. Bernardia fonsecae is mor¬
phologically similar to B. mollis Lundell. They have
in common long petioles, similar shape and size of
the staminate bract, and a lobulate intrastaminal
disc. Bernardia fonsecae differs from B. mollis in
its leaf base and leal shape, vestiture of the ailaxial
surface of the leal, vestiture of the abaxial veins,
diameter of the cicatricoso-crateriform glands, po¬
sition of the staminate inflorescence, and number
of stamens, among other characteristics.

Key words : Bernardia , Euphorbiaceae, Mexico.

Pax and Hoffmann (1914) wrote the most recent
revision for Bernardia , a wholly Neotropical genus
o( Euphorbiaceae, recognizing only four species in
Mexii •o. As the number of collections for the genus
in Mexico has increased, new species have been
described, and the contributions of McVaugh
(1961, 1995) and Lundell (1940, 1945, 1976.
1985) are of special interest. With ea. 25 species
of 50 recorded for the genus (Webster, 1994), Mex¬
ico has been considered, together with Brazil, as a
center of diversity for the genus.

Bernardia is divided into seven sections accord¬
ing to Pax and Hoffmann (1914), three of which
occur in Mexico: sect. Tyria (Klotzsch) Muller Ar-
goviensis, sect. Alevia (Baillon) Miiller Argoviensis,
and sect. Tragantluis (Klotzsch) Muller Argoviensis.
Sections Tyria and Alevia consist of trees and
shrubs with fasciculate or divided hairs, while the
monotypic section Tragantluis has simple hairs and
includes the only herbaceous species of the genus.
Section Tyria has lacinulate-lacerate style branch¬
es, and section Alevia has non-laciniate style

branches. Tyria, with ea. 21 species, is mainly
Mexican with two representatives in the West In¬
dies. Here we describe a new species of section
Tyria from Mexico.

Rernurdiu fonsecae A. Cervantes & J. Jimenez
Ramirez, sp. nov. TYPE: Mexico. Guerrero:
Mpio. Leonardo Bravo, Puerto Soleares, 7 km
despues de Carrizal hacia Atoyac, 17°35'15"N,
99°50'W, 2500 m, montane rain forest, 5 Nov.
1998, R. M. Fonseca 2538 (holotype, M E XL:
isotypes, ECME. MO). Figure 1.

Arbor 6.0—18.0 m alta, dioica. Folia chartacea leviter
scabra, longe petiolata; costa abaxialiter trichomatibus
stellatis radiis longissimis praeditis vestita. Infloreseentia
mascula ex racemo axillari tenninelive constans, bracteis
4- ad 7-floris. Infloreseentia feminea ex spica terminali
constans, bracteis unifloris. Flos masculus staminibus (22
ad) 26 ad 44. Flos femineus stylo lacerato piloso rufo vel
rubiginoso. Capsula ca. 24 mm diametro, pericarpo mi¬
nute aeuleato, confertim stellato-hirto; seminibus globosis
leviter carinatis.

Frees 6.0—18.0 m tall; dioecious. Branchlets with
conspicuous elongate lenticels (1)2—4 mm long.
Leaf bla<le 6.0— 16.0(— 22.0) cm long, 3.3-9.5(—
12.0) cm wide, broadly elliptic to elliptic, elliptic-
obovate or rarely lanceolate; margin serrate with 3
or 4 teeth per cm, teeth glandular; base attenuate,
rounded or sometimes cordate or subtruncate; apex
abruptly acuminate or acute; 3-nerved at the base,
ailaxial midrib especially near the base with fas¬
ciculate and stellate triehomes with 2 to 8 radii,
0.2— 0.5 mm long, abaxial midrib especially near
the base with conspicuous and soft stellate tri¬
ehomes with 8 to 15 radii, 0.75 mm long; upper
foliar surface punctate under magnification; drying
ehartaeeous, darker above, slightly rough; ailaxial

— >

Figure I. Bernardia fonsecae A. Cervantes X J. Jimenez Bamfrcz. — A. Branch with staminate inflorescence (Fonseca
2538, type). — B. Branch with pistillate inflorescence ( Fonseca 2539). — C. Ovary; note the style and annular disc

Novon 12: 38-41. 2002.

Volume 12, Number 1
2002

Cervantes & Jimenez
Bernardia fonsecae from Mexico

39

( Fonseca 2539). — D. Seed: ventral, dorsal, and lateral views ( Fonseca 2540). — E— G. Fruit, equatorial and polar views
(bottom) (Fonseca 2540). — H. Stellate trichomes from the abaxial midrib (Fonseca 2533. type). — 1. Staminate flowers
and a subtending bract (Lozano 710). — J. Cieatricoso-crateriform gland on abaxial surface of the leaf (Fonseca 2539).

40

Novon

Table 1 . Morphological characteristic

s distinguishing Bernardia fonsecae

and B. mollis.

Characteristics

B. fonsecae

B. mollis

1 ,eaf shape

broadly elliptic to elliptic', elliptic-
obovate or rarely lanceolate

ovate, elliptic-ovate

I -eat base

attenuate or rounded

broadly rounded

Foliar surface

smooth, punctate under magnifica¬
tion

rugose

Abaxial vein vestiture

stellate triehomes with 8 to 15 ra¬
dii. 0.75 mm long, only near
the leaf base

fasciculate and stellate triehomes
with 2 to 6 radii, 0.5 mm long,
along the entire leaf

Adaxial vestiture

stellate-puberulent

appressed fasciculate

Cicatricoso-crateriform gland diameter

0.5-0.8 mm

0.8- 1.0 mm

Staminate inflorescence position

axillary and terminal

axillary

Staminate tepal size

3.5— 4. 5 X 1.5— 4.0 mm

2.0— 3.5 X 1 .2 mm

Staminate tepal vestiture

glabrescent on inner surface

pilose on inner surface

Staminate (lowers per bract

4—7 (lowers

3—5 flowers

Stamen number

(22)26—44

22-24

Filament length

2.5 mm

2.0 mm

Anther length

0.3 mm

0.5 mm

surface glabrescent or sparsely puberulent with
simple, stellate or multiradiate triehomes, radii 6
to 10, ea. 0.1 mm long, abaxial surface glabrescent
or sparsely puberulent with stellate and multiradi¬
ate triehomes, radii 8 to 16, ca. 0.1 mm long; I to
8 cicatricoso-crateriform glands at margin and on
each side of the midrib in abaxial surface, 0.5— 0.8
mm diam.; petioles 1.0-4. 2(— 8.0) cm long, stellate-
puberulent and tomentose; stipules deciduous, 2.0—

2.5 mm long, narrowly triangular, entire, rigid, ap-
pressed, strigulose, pale yellow. Staminate inflores¬
cence racemose, terminal and axillary, at anthesis
1.5— 7.5 cm long, the (5)10 to 15 bracts separated
or congested on the axis; bract 4- to 7-flowered,
broadly deltoid-ovate, cupuliform, 2. 5-8.0 X 3.5—

4.5 mm, apiculate, coriaceous, pilose on outer sur¬
face, glabrescent on inner surface; bracteoles in¬
conspicuous. Staminate flowers pedicellate, pedicel
2.0-3. 5 mm long at anthesis; tepals 3 or 4, valvate,
ellipt ie or oblong, 3.5— 4.5 X 1 .5— 4.0 mm, equal,
densely pilose on outer surface, glabrescent on in¬
ner surface; stamens (22)26 to 44; filaments stout.

2.5 mm long, pinkish; anthers globose, 0.3 mm
long, connective inconspicuous; iutrastaminal disc
lobulate. Pistillate inflorescence spicate, terminal,
peduncles 10 mm long, 4 to 8 bracts; bract 1 -flow¬
ered, ovate, cupuliform, 3.0— 3.5 X 3. 0-4.0 mm,
acute, coriaceous; bracteoles conspicuous. Pistil¬
late flower sessile; tepals 6, imbricate, broadly
ovate, unequal in size, densely pilose on outer sur¬
face, glabrescent on inner surface; disc annular,
flat, margin erose, glabrous; ovary densely stellate-
hirsutulous, globose, deeply 3-lobed, 3—4 X 3-5
mm, styles 3, lacerate, stigma reddish or rufous.

pilose. Fruits capsular (1)2- or 3-lobed, 24.0 mm
diam., densely stellate-hirsutulous, slightly keeled
on midrib, pericarp minutely aculeate, dehiscent.
Seeds spheroid, 14.0 mm long, 12.0 mm wide,
slightly carinate on superior and dorsal face, car¬
uncle absent, testa crustaceous, chestnut-colored,
minutely lineolate-verrucose and brown-marbled.

This species is known only from montane rain¬
forests at 2200—2500 m in Guerrero, Mexico, where
it is associated with Fuchsia sp., Clethra sp.. Sa¬
lami m sp., Abies religiosa Lindley, Chiranthoden-
dron pentadactylon Larreat. and Cornus disciflora
Sesse & Moyino ex DC.

Bernardia fonsecae is unique within the section
in its large leaf size, the vestiture of the abaxial
midrib (soft stellate triehomes with 8 or 15 radii,
0.75 mm long), and in having the largest fruit
known of Bernardia. It belongs to section Tyria and
is morphologically similar to B. mollis Lundell,
which occurs in montane rainforests of Chiapas.
These species have in common long petioles, sim¬
ilar shape and size of the staminate bract, and a
lobulate iutrastaminal disc. Bernardia fonsecae dif¬
fers from B. mollis in its leaf base and leaf shape,
vestiture of the adaxial surface of the leaf, vestiture
of the abaxial veins, diameter of the cicatricoso-
crateriform glands, position of the staminate inflo¬
rescence, and number of stamens, among other
characteristics (Table 1).

The name honors Rosa Marfa Fonseca, teacher
and botanist from the Facultad de Cieneias.
UN AM. w ho has made important contributions to
the knowledge of the flora of Guerrero state.

Volume 12, Number 1
2002

Cervantes & Jimenez
Bernardia fonsecae from Mexico

41

Paratypes. MEXICO. Guerrero: Mpio. Gral. Helio-
doro Castillo, aprox. 3 km de Puerto del Gallo en direc-
cidn NE, ladera 0 del cerro Teotepec, 18 Oct. 1999, E.
Dominguez 1250 (FCME, MEXU); Mpio. Leonardo Bravo,
Puerto Soleares, 7 km despues de Carrizal hacia Atoyac.
5 Nov. 1998. Fonseca 2539 (FCME, MEXU, MO), 2540
(FCME. MEXU, MO); Mpio. Leonardo Bravo, 7 km delan-
te de Carrizal rumbo a Atoyac y 2 km al E rumbo a Cacho
de Oro, II Sep. 1999, Fonseca 2937 (FCMF, MEXU);
Mpio. Leonardo Bravo, aprox. 500 m del Puente Las Pas-
t i I las, camino Filo de Caballos— Chichihualco, 6 Deo.
1999, Fonseca 2951 (FCME, MFXU); Mpio. Chilpanein-
go, al 0 de Omitelmi, Canada de la Laguna de Agua Frfa,
19 July 1985, Lozano 710 (FCME, MEXU, MO).

Acknowledgments. Hilda Flores Olvera, Hclga
Ochoterena, and Lawrence Kelly all greatly en¬
hanced the (juality of this article by providing nu¬
merous insightful comments. We thank Ramiro
Cruz Duran for the excellent drawing. We also ap¬
preciate the editor and anonymous reviewers,
whose comments helped improve the manuscript.

The field trips to Guerrero were supported by CON-
ABIO.

Literature Cited

Lundell, C. L. 1940. Studies of tropical American plants-
I. Contr. Univ. Michigan Herb. 4: 12—14.

- . 1945. New phanerogams from Texas, Mexico and

Central America. Wrightia 1: 53—61.

- . 1976. Euphorbiaceae. Studies of American

plants XII. Wrightia 5: 243-249.

- . 1985. Euphorbiaceae from Mesoamerica. Phyto-

logia 57: 367-368.

McVaugh, R. 1961. Euphorbiaceae Novae Novc-Galici-
anae. Brittonia 13: 145—205.

- . 1995. Euphorbiacearum Sertum Novc-Galiciana-

rum Revisarum. Contr. Univ. Michigan Herb. 20: 173—

215.

Pax, F. & K. Hoffmann. 1914. Bernardia. In Das Pflan-
zenreich IV. 147. VII (Heft 63): 21^15.

Webster, G. L. 1994. Synopsis of the genera and supra-
generic taxa of Euphorbiaceae. Ann. Missouri But.
Card. 81: 33-144.

Novelties in Croton (Euphorbiaceae) from Southeast Asia

// ans-Joach i m Esser

Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A.

ABSTRACT. A new species, Croton kongkandanus
Esser, is described from Thailand, based mainly on
its large fruits. The name of Croton lissophyllus
from India and Bangladesh is validated, and C. lan-
iflorus Geiseler is recognized as the valid name for
the poorly known and illegitimate Croton lanatus
Loureiro from Vietnam and described here. Seven
taxa of Southeast Asian Croton are lectotypified,
two new synonyms are proposed, and two potential
ones are discussed, some of them in preparation for
inclusion in the Flora of Thailand.

Key words: Asia, Bangladesh, Croton , Euphor¬
biaceae, India, Thailand, Vietnam.

The revision of Euphorbiaceae for the Flora of
Thailand , recently conducted by staff of the Forest
Herbarium, Bangkok, and collaborators, had a re¬
liable basis in the study of Thai Euphorbiaceae by
Airy Shaw (1972). Nevertheless, several novelties
and problems came to light, among them some in
the genus Croton L. In connection with this project,
taxonomic problems in Croton from Indochina and
India were studied. The complete revision of Croton
for the Flora of Thailand will be published sepa¬
rately, hut some notes on Croton in Thailand and
adjacent regions are recorded here. Thai localities
are cited in accordance with the guidelines for the
Flora of Thailand.

Croton chevalieri Gagnepain, Bull. Soc. Bot.
France 68: 550. 1921 (1922]. TYPE: Vietnam.
Bienhoa, reserve of Trang Torn, 30 Mar. 1914
(fl). A. J. H. Chevalier 32048 (holotype, P; iso¬
type. A).

In contrast to the other species of Croton he had
described in 1922, Gagnepain did not accept C.
chevalieri later in 1925 (Gagnepain, 1925), but list¬
ed it under his dubious species at the end, not
including it in his key. Croton chevalieri is based
on a single, rather poor collection. It is most prob¬
ably not a separate species. It resembles C. poila-
nei , except that the hairs are stellate to stellate-
lepidote, whereas lepidote hairs are diagnostically
important for C. poilanei. It might prove to he an
aberrant collection of the latter on further study.

Croton drlpyi Gagnepain, Bull. Soc. Bot. France

Novon 12: 42-46. 2002.

68: 552. 1921 1 1922]. TYPE: Vietnam. “Ad
Deon-ba montem prope Cay minh,” Apr. 1866
(fl, fr), L. Pierre 6231 (lectotype, designated
here, P; isolectotypes, BM, K, P).

Among the four syntvpe collections, Pierre 6231
is the one with the largest number of duplicates
found and the only one with both flowers and fruits.
The sheet with analytical drawings attached to it,
made by E. Oelpy, is chosen as the lectotype.

Croton griffithii Hooker f., Fl. Brit. Ind. 5: 392.
1887. TYPE: Malaysia. “Malacca,” s.d. (fl, fr),
W. Griffith 4781 (lectotype, designated here, K;
isolectotype, k).

Hooker noted in his protologue that his new spe¬
cies might be heterogeneous. Among the nine syn-
type collections at Kew, six ( Griffith 4778 , 4781,
King's Collector 1115, 4484, 6157, Scortechini s.n.)
agree with the current concept of C. griffithii
(leaves drying yellowish brown, cordate-rounded at
base, with abaxial glands), whereas two ( Wallich
7754, 7967) belong to C. laevifolius Blume (leaves
drying darker, acute-obtuse at base, with lateral-
adaxial basal glands). Maingay 1406 was not stud¬
ied. Airy Shaw (1972) was certainly correct to dis¬
tinguish these two species that had been united by
several authors in the past, such as Corner (1939).
The above lectotypification fixes the current use of
the name, with Griffith 4781 being the only syntype
bearing both flowers and fruits.

Among the two sheets of Griffith 4781 at Kew,
the one with analytical sketches of flowers and at¬
tached dissected (lowers is chosen here as the lec¬
totype.

Croton kongensis Gagnepain, Bull. Soc. Bot.
France 68: 555. 1921 1 1922], TYPE: Laos.
Pak-lay, leg. 1866—1868 (fl, fr), C. Thorel s.n.
(lectotype, designated here, P; isolectotype, P).

The sheet with fruits and analytical drawings,
which had already been separated in a type folder,
is selected here.

Volume 12, Number 1
2002

Esser

Novelties in Croton

43

Croton kongkandanus Esser. sp. nov. T\ EH:
Thailand. South-Eastern: Chon Buri, Khao
kieo Wildlife Sanctuary, 300 m, 25 July 1976
(fr), ./. F. Maxwell 76-443 (holotype, BE
56565; isotype, AAU).

Species Crotonis foliis ovato-ellipticis non triplinerviis
pilis stellato-lepidotis argenteo-brunneis dense obtectis ad
C. argyratum maxime accedit, sed ab eo fruotibus multo
majoribus 22—27 mm (non 12-16 mm) longis brevipedi-
cellatis (3—4 mm, non 10-13 mm) differl.

Tree to 15 m tall. DBH to 25 cm; bark gray,
smooth to very finely roughened and with promi¬
nent, scattered lenticels; young branchlets densely
pubescent, glabreseent. Indumentum consisting of
stellate to stellate-lepidote hairs (radii more fused
on leaves) with a small brownish center and hyaline
radii (appearing silvery) on the leaves, brownish on
leaf veins and floral parts, flat, sometimes with a
porrect central radius, ca. 0.2— 0.3 mm diam., with
ca. 20-30 radii. Stipules linear-elliptic, 7-20 mm
long, densely pubescent. Leaves often apically
crowded but basically alternate; petioles 25-55 mm
long, densely pubescent; blades ehartaceous. ellip¬
tic to slightly ovate, 1 1-19 X 5-8.5 cm, index 1.8-
2.3. base obtuse with the very base cordate, margin
entire, apex shortly acuminate, soon glabrous
above, beneath densely silvery-pubescent and not
glabreseent, hairs quite uniform, brown-tinged on
venation; basal glands lateral on the abaxial midrib
base, flat, sessile, 0.5-1 mm diam., marginal glands
absent; side veins distinct on both surfaces, in 8 to
10 pairs, not triplinerved, tertiary veins visible on
both surfaces. Inflorescences terminal, solitary,
densely and persistently silvery-brown pubescent
throughout, 8— 10 cm long, consisting of a basal part
of 1—2 cm with 2 to 6 pistillate flowers, without
lateral staminate flowers in the same bract, and a
larger upper staminate part, with 1 to 3 flowers per
bract; floral bracts ca. 1 mm long, eglandular, quite
persistent. Staminate flowers densely pubescent;
pedicel 1-1.5 mm long; sepals ca. 2 X 1.5 mm;
petals ca. 2-2.5 X 1 mm; stamens ca. 11, pubes¬
cent at base. Pistillate flowers densely pubescent;
pedicel ca. I mm long; sepals ca. 4—5 X 2-2.5 nun,
longer than the ovary; petals 2 X 0.5 mm; ovary
2.5 mm long; slylar column short (ca. 0.5 mm), stig¬
mas 4—5 mm long, bifid on the apical ca. 3 mm.
Fruits: pedicel 3—4 mm long; schizocarp 22—27 X
20—21 mm, elliptic to slightly obovate, not sulcate,
densely pubescent outside. Seeds flattened-elliptic,
15-16 X 8—1 1 X 6 mm, smooth, brown, sometimes
slightly variegated, caruncle not seen.

The species in named after Kongkanda Chaya-
marit of the Forest Herbarium, Royal Forest De¬

partment. Bangkok, who spent so much of her time
and energy in arranging and organizing the Eu-
phorbiaceae project for the Flora of Thailand.

This species probably belongs in Croton sect. Ar-
gyrocroton (Miiller Argoviensis) G. L. Webster ac¬
cording to Webster (1993). Croton kongkandanus
differs from all silvery-leaved species of Croton in
Southeast Asia and Malesia by the much larger
fruits, a diagnostically important difference. The
other Malesian species with silvery leaves have
fruits 4-16 mm long. In other characters, the new
species is very similar to C. argyratus Blume. The
latter has mature fruits 12—16 mm long on distinct¬
ly longer pedicels exceeding 10 mm, and is re¬
stricted to West Malesia, while in Thailand it is
only known from the southernmost Peninsular prov¬
inces. The Indo-Chinese species C. budopensis
Gagnepain and C. maieuticus Gagnepain. consid¬
ered as synonyms of C. argyratus by Airy Shaw
(1972), are all separate endemic species: C. budo¬
pensis has much smaller, subsessile fruits only 4—
5 nun long, and C. maieuticus has a fruit size sim¬
ilar to C. argyratus but stipitate glands on the leaf
bases.

Type and paratype collections of C. kongkandan¬
us were distributed under the name of Croton ar¬
gyratus.

The only Asian species comparable with C.
kongkandanus in fruit size is C. malabaricus Bed-
dome. This species is only known from southwest¬
ern India (Western Ghats) and was thoroughly de¬
scribed and illustrated by Chakrabarty and
Balakrishnan in 1997. Croton malabaricus seems
to differ in having shortly stipitate fruits and longer
pedicels in staminate and pistillate flowers and
fruits.

Croton kongkandanus is so far only known from
Thailand. It grows in mixed deciduous and ever¬
green, seasonal hardwood forest, disturbed ever¬
green forest, found over shale bedrock.

Paratypes. THAILAND. Northern: Chiang Mai.
Chiang Dao Distr., Doi Chiang Dao Animal Sanctuary,
Uuay Nah Lao Station, 600 m, 14 Aug. 1995 (fr). Maxwell
95-496 (BKF, CMU); Tak, Pha Wo, 650 m. 13 July 1972
(fr), Smitinand & Seidenfaden 11630 (BKF); Sukhotai, Sri
Satchanalai National Park, 200 m, 7 Feb. 1988 (ster).
Smitinand s.n. (BKF 87372). South-Eastern: Chon Buri.
Khao Khieo Wildlife Sanctuary, 400 m, 27 Mar. 1976 (fl).
Maxwell 76-162 (AAU. BK. L); Chanthaburi, Khao Sabap
National Park, 6 July 1927 (fr). Put 912 (BK, BM, K, L,
P).

Croton krnhus Gagnepain, Bull. Soc. Bot. France
68: 555. 1921 [1922]. TYPE: Cambodia. Me¬
kong river, Kratee, 9 Mar. 1914 (fl), A. J. H.

44

Novon

Chevalier 31882 (lectotype, designated here,

P).

Although several of the remaining seven syntype
collections (now paralectotypes) are distributed
with duplicates, the chosen lectotype is the only
sheet with attached analytical drawings, and with
the vernacular name ‘krabas prey' on the label, the
source of the epithet. It is also one of the richest
collections among the syntypes.

Croton lachnocarpus Bentham, Hooker’s J. Mot.
Kew Card. Misc. 6: 5. 1854. TYPE: Hong
Kong, ,/. G. Champion (holotype, K not seen).

Croton calococcus Km/. J. Asiat. Soc. Mengal 42: 242.
1873. Syn. nov. TYPE: Burma. Rangoon, s.d., .S'.
Kurz 1607 A (lectotype, designated by Chakrabarty &
Balakrishnan (“1992," published in 1997), CAL not
seen; isoleetotype, k not seen).

Croton bonianus Gagnepain, Bull. Soc. Rot. France 68:
549. 1921 1 1922]. SYNTYPES: Vietnam, ban Mak.
10 INov. 1883, R. I\ Ron 2260 (A, P), R. P. Ron 215
(P not seen), near Hanoi, But-son, 7 Dec. 1883, R.
R. Ron 2545 (A, P); Thinh-chau, R. R. Ron 2568 bis
(P not seen).

Croton rnurex Croizat, J. Arnold Arbor. 23: 41. 1942.
TYPE: Vietnam. 12 km N ol Dankia-Langbiang, 27
Oct. 1930 (fl), K. Poilane 18657 (holotype, A; iso-
types, K, 1’).

Croton trachycaulis Airy Shaw, Kew Bull. 23: 74. 1969.
TYPE: Thailand. South-Western: Prachuap kliiii
Khan, llui Yang, 3 Oct. 1930 ((I, fr). Put / = Put
Phraisurind / 5186 (holotype, K; isotypes. A, RK,
BM. L).

This synonymy was largely (except for C. calo¬
coccus) included by Govaerts et al. (2000) after dis¬
cussion with the author. The whole complex is best
treated as a single polymorphic species, and was
formerly divided into species mostly on the basis
of two characters, indumentum and fruit surface.
Fruit surfaces vary from smooth to muriculate
seemingly as natural variation, not correlated with
other characters or geography. The indumentum
does show a geographical pattern: Plants from Chi¬
na are usually glabreseent on the upper leaf sur¬
face, with leaf hairs that are slightlv smaller (ea.
0.8 mm diam.) with a central porrect radius not
longer than the lateral radii. Plants from Thailand
always have leaves pubescent above, and the hairs
are often larger ( (0.2— )0.8 — 1 .5 mm diam. | and with
an exceptionally long porrect radius, with the lat¬
eral radii short to nearly rudimentary on the upper
leaf surface (not on other parts). This alone is prob¬
ably not sufficient for separation at the species lev¬
el. Further studies with a better representation from
Indochina are necessary to evaluate if clear-cut dif¬
ferences exist somewhere along the dine. If the in¬

dumentum turns out to be significant, the correct
name for the Thai plants is probably C. calococcus ,
and among the Vietnamese names at least C. bon¬
ianus certainly remains a synonym ol C. lachno¬
carpus.

I he two syntypes of C. bonianus, Ron 2260 and
Bon 2343, are mounted together on one sheet in
both A and P, with one fruiting twig and one (low¬
ering but one joint label; they cannot be separated.

Croton laniflorus Geiseler, Croton. Monogr.: 44.
[Mar.) 1807. Croton lanatus Loureiro, Fl.
Cochinch, ed. 1, 2: 581. 1790, nomen illeg.,
non Lamarck, Encycl. 2: 211. 1788. Croton
lasianthus Persoon, Syn. PI. 2: 586. [Sep.|
1807, redundant name. Croton erioanthemus
Smith, in Rees, Cycl. 10: Croton no. 21. 1808.
redundant name. Triplandra lanata (Loureiro)
Rafinesque, Sylva Tellur.: 63. 1838. TYPE: Vi¬
etnam. “In sylvis montanis Cochinchinae,” s.d.
(fl, fr), ./. Loureiro s.n. (holotype, BM).

Croton limitincola Croizat, J. Arnold Arbor. 23: 45. 1942.
Syn. nov. TYPE: Vietnam. Taai Wong Mo Shan, Sep.
1939 (fl. fr), W. T. Tsang 29584 (holotype, A).

Geiseler’s name is the earliest of three names
replacing the illegitimate name of Loureiro. Croton
laniflorus was misspelled as “ laxiflorus ” by the In¬
dex Kewensis, a mistake not picked up by Govaerts
et al. (2000); neither works indicate that it was a
replacement for Loureiro’s name. Croton lasianthus
was published a few months later than C. laniflorus,
but has been the accepted name in the literature,
following the authority of Muller (1866) and Merrill
(1935).

Because the identity and many important char¬
acters of C. laniflorus have never been clarified
(even Merrill, 1935, was in doubt), it is useful to
contribute a short description of the type:

Woody plant. Indumentum consisting of stellate
hairs with a brown center and pale-hyaline radii
(appearing as brownish dots), Hat. with ca. 15 radii.
Stipules linear-elliptic, ca. 6 mm long. Leaves al¬
ternate, only immature ones on the type sheet; pet¬
ioles with few hairs; immature blades membranous,
elliptic, ca. 8 X 3.5 cm, index ca. 2.3, base obtuse,
margin shallowly serrate, apex acute to subacumi¬
nate, glabrous above, beneath with scattered but
distinct hairs; basal glands abaxial-lateral on the
junction of petiole and midrib, stipitate and quite
distinct, marginal glands not seen; side veins dis¬
tinct on both surfaces, in 10—1 I pairs, not tripli-
nerved, arching in quite distinct loops (brochidod-
romous). Inflorescences only seen in fragments.
Pistillate flowers: pedicel 4 mm long, with few hairs

Volume 12, Number 1
2002

Esser

Novelties in Croton

45

only; sepals 4—5 X 1.5— 2.5 mm, much longer than
the ovary, nearly glabrous; petals not seen; ovary
ea. 2 mm long, distinctly brown hispid-pubescent
with long porrect radii; stylar column short (ca. 0.5
mm) but present, stigmas ea. 3 mm long, bifid on
the apical ca. 2 mm, slightly Hat and broadened.
Fruits only seen in fragments; schizocarp 9 mm
long, distinctly pubescent, smooth. Seeds not seen.

Although the holotype of C. laniflorus is a some¬
what fragmentary collection, sufficient diagnostic
characters are present. The type of C. limitincola
is identical in all important characters studied here,
and < 1 i ffers in more distinctly pubescent pistillate
flowers and in much larger leaves up to 22 X 9
tan, but the leaves are mature and the smallest leaf
on the holotype is nearly identical in size with the
type of C. laniflorus.

Croton vietnamensis Rat leliffe-Smith & ( iovaerts
(the name replacing the illegitimate Croton longipes
Gagnepain non M. E. Jones), described from Nha-
trang, Vietnam, is probably also synonymous, al¬
though this needs further study.

Croton lissoplivllus Radcliffe-Smith & Covaerts,
Kew Bull. 52: 187. 1997 (nomen) ex Esser, sp.
nov. Croton laevifolius auct. non Blume, in
Hooker f., FI. Brit. India 5: 391. 1887. Croton
khasianus Hooker f., El. Brit. India 5: 392.
1887, nomen inval. Croton hookeri Croizat, J.
Arnold Arbor. 21: 498. 1940, nomen inval. et
illeg., non Croton hookeri Andre, 111. Hort. 19:
56. 1872. TV BE: India. Meghalaya: Khasi
Mountains, 2000-4000 ft. (fl. fr). ./. I). Hooker
& T. Thomson s.n., s.d. (“• Croton 17") (holo¬
type, GH).

Species Crotonis ad C. laevifolium accedit sed pilis stel-
latis, foliis serratis glabris vel parce pubescentibus, glan-
dulis basilaminaribus stipitatis, fructibus bilocularibus
distinguenda est.

Chakrabarty and Balakrishnan (“1992,” pub¬
lished in 1997) contributed an excellent descrip¬
tion and illustration of this species (as “C. hook-
eri"), anti they seem to be completely correct in
their circumscription. The bilocular ovaries and
fruits seem indeed to separate the species from sev¬
eral similar Asian Crotons. The record for Thailand
by Airy Shaw (1972) could not be confirmed.

This species has a remarkable nomenclatural
history. Hooker (1887) had proposed the name C.
khasianus only provisionally, allowing for the pos¬
sibility that the plants he had described under C.
laevifolius Blume on the previous page of his work
turned out to be different from Blume’s species.
This did not constitute a valid publication (ICBN

Art. 34.1; Greater et al., 2000). Croizat (1940) was
aware of that; he accepted the species and replaced
the name with C. hookeri. He did not realize that
he had created a later homonym of C. hookeri An¬
dre. Radcliffe-Smith and Govaerts (1997) therefore
replaced the name C. hookeri Croizat with C. lis-
sophyllus. Their literature reference for C. hookeri
Andre was unfortunately erroneous here and in Go¬
vaerts et al. (2000: 404), who also attributed the
name to Veitch. Croizat’s name, however, was not
only illegitimate but also invalid, because he did
not contribute a Latin description or diagnosis, and
did not refer to a previously published Latin diag¬
nosis, which is mandatory after I January 1935
(ICBN Art. 36.1), for which reason C. lissophyllus
also remained invalid.

Govaerts et al. (2000) cited a synonym under C.
lissophyllus, C. cnidopyllus Radcliffe-Smith & Go¬
vaerts, the name replacing the illegitimate C. urti-
cifolius Y. T. Chang & f). II. Chen non Lamarck.
It was described from China (Guizhou). Although I
was unable to examine original material ol that spe¬
cies, it is improbable that it is conspecific with C.
lissophyllus. The latter seems to be a local endemic,
and its leaves are not triplinerved. Croton cnido-
phyllus might be closely related to (or even be syn¬
onymous with) C. lachnocarpus.

The type specimens of C. lissophyllus were dis¬
tributed under the name of Croton laevifolius.

Paratypes. BANGLADESH. Silhet, Pandowah Hill,
May 1822 (fern, fl), l\. Wallich 7719 (CAL not seen, K,
K-WALL). INDIA. Meghalaya: Khasi Mountains, 13
June 1850 (fl), J. I). Hooker & Thomson s.n. (A, K); 20
June 1850 (fl), ,/. I). Hooker & T. Thomson s.n. (K); s.d.,
J. D. Hooker & T. Thomson s.n. (“Croton 17”) (A, G-DC,
NY, TCI), U); s.d. (male fl), W. Griffith 47H4 (GH, K).

Croton inekongensis Gagnepain, Bull. Soc. Bot.
France 68: 558. 1921 [1922], TYPE: Cambo¬
dia. Campong-luang, leg. 1866—1868 (fl, fr), C.
Thorel s.n. (lectotype, designated here, P; is-
olectotype, P).

The Thorel collection from Campong-luang is the
only one among the four syntype collections (two
made by Harmand and Thorel each) with fruits.
Among the two sheets at P, the one with attached
drawings is chosen here.

Croton poilanei Gagnepain, Bull. Soc. Bot.
France 68: 559. 1921 [ 1922 1. TYPE: Vietnam.
Baria: Nui-dinh, 25 Oct. 1919 (fl), E. Poilane
(leg. A. J. H. Chevalier) 636 (lectotype, des¬
ignated here, P; isolectotypes. A, K).

Several excellent syntypes with duplicates are

46

Novon

available, all with analytical drawings by Delpy and
used by Gagnepain attached to the Paris sheet. The
chosen leetotype had already been separated in a
type folder in Paris, and it is the only one among
the syntypes with a label by Poilane, to whom the
species was dedicated.

Groton tliorclii Gagnepain, Bull. Soc. Hot. France
68: 560. 1921 [1922], TYPE: Vietnam. “Ad
montem Dinh prope Baria,” Apr. 1867 (fl,
imm. fr), L. Pierre 6226 (leetotype, designated
here, P; isolectotypes, BM, GH, NY).

Croton thorelii was based on nine syntype col¬
lections. All of them belong to this species and not
to the similar C. decalvatus Esser, with which C.
thorelii was formerly confused by some authors,
such as Airy Shaw (1972) (Esser & Chayamarit,
2001). Among the syntypes in P with drawings at¬
tached, the one with the largest number of dupli¬
cates is selected here.

Gagnepain (1925: 264, fig. 28.8—10) stressed the
diagnostic importance of pubescent stamens. Sta¬
mens pubescent to a various and often variable de¬
gree were found in several Asian species of Croton,
and do not appear to be significant. The quadrifid
stigmas however, also discussed by Gagnepain,
seem to be diagnostic for C. thorelii.

Acknowledgments. This study was financed by
the Thai Biodiversity Research and Training Pro¬
gramme, a joint program supported by The Thai
Research Fund and the National Center for Genetic
Engineering and Biotechnology. It was also sup¬
ported by a Mercer Fellowship of the Arnold Ar¬
boretum, Harvard University. I bis work could not
have been done without the support of the staff of
the Forest Herbarium, Bangkok, and the Harvard
University Herbaria; in particular I have to mention
Kongkanda Chayamarit, who organized the Eu-
phorbiaceae project for the Flora of Thailand. I
enjoyed useful discussions with Walter Kittredge
and Kancheepuram Gandhi from the Harvard her¬
baria. in particular on the problems of C. lisso-
phyllns. I also received helpful comments from Paul

Berry (University of Wisconsin) and Alan Rad-
cliffe-Smith (Kew), as well as from two anonymous
reviewers. 1 am grateful to the directors and cura¬
tors of the herbaria who made it possible to study
their holdings: A. AAU, BCU, BK, BKF, BM. CMU,
FR, G. G-DC, GH, HBG, K, K-WALL, E. MICH.
MO. MT, NY. P. PH. QBG. SING. TCI), TI. U, and
Khon Kaen University.

Literature Cited

Airy Shaw. II. K. 1972. The Euphorbiaeeae of Siam, kew
Bull. 26: 191-363.

Chakrabarty, T. & IN. P. Balakrishnan. 1992 1 1 997]. A
revision of Croton L. (Euphorbiaeeae) for Indian Sub¬
continent. Bull. Bot. Surv. India 34: I— 85.

Corner, El. J. II. 1939. Notes on the systematy and distri¬
bution of Malayan Phanerogams III. Card. Bull. Sin¬
gapore It): 239-329.

Croizat, 1.. 1940. New and critical Euphorbiaeeae from
Eastern Tropical Asia. J. Arnold Arbor. 41: 490—510.
Esser, II. -J. & k. Chayamarit. 2001. Two new species and
a new name in Thai Croton (Euphorbiaeeae). Thai For.
Bull. (Bot.) 29: 51-57.

Gagnepain, F. 1921 [1922]. Euphorbiaeees nouvelles
d’lndo-Chine (Croton). Bull. Soe. Bot. France 68: 548—
562.

- . 1925. Euphorbiacees. Pp. 229—372 in M. H. Le-

eomte, Flore Generale de l’Indo-Chine 5. Masson et
Cie., Paris.

Govaerts, R., I). G. Frodin & A. Radeliffe-Smith. 2000.
World Checklist anti Bibliography of Euphorbiaeeae.
Royal Botanic Gardens, Kew.

Greuter, W„ J. McNeill, F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). H. Nicolson, P C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & I). I.. Hawks-
worth. 2000. International Code of Botanical Nomen¬
clature (Saint Louis Code). Regnum Veg. 138.

Hooker, .1. I). 1887. Euphorbiaeeae. Pp. 269—477 in FI.

Brit. India 5. Reeve, London.

Merrill, E. I). 1935. A commentary on Loureiro’s “Flora
Cochinchinensis.” Trans. Amer. Philos. Soc. Ser. 2,
24(2): 1-445.

Muller. J. 1866. Euphorbiaeeae excl. Euphorbieae. Pp.
189—1273 in A. I)e Candolle, Prodromes svstematis na-
turalis regni vegetabilis 15, 2. Fleischer, Munchen &
Masson, Paris.

Radeliffe-Smith, A. & R. Govaerts. 1997. New names and
combinations in the Crotonoideae. kew Bull. 52: 183—

1 89.

Webster, G. L. 1993. A provisional synopsis of the sec¬
tions of the genus Croton (Euphorbiaeeae). Taxon 42:
793-823.

Astragalus tuyehensis (Fabaceae), a New Species from Iran

Ahmad Ghahreman

Department of Biology, Faculty of Sciences, University of Tehran, Tehran, Iran

A li-Asgh a r Maassoum i

Research Institute of Forests and Rangelands, P.0. Box 13185-116, Tehran, Iran

Farrokh Gh ah rem a n i-neja d

Department of Biology, Faculty of Science, University of Tarbiat-Moaallem, 49 Dr. Mofatteh
Avenue, 15614 Tehran, Iran. ghahremaninejad@saba.tmu.ac.ir

ABSTRACT. A new species from Iran, Astragalus
tuyehensis, belonging to the bifurcate-haired sub¬
genus Cercidothrix, is described and illustrated. Its
relationships to A. durandianus and its distinguish¬
ing characters, such as inflated calyx and non-swol-
len fruit, are discussed.

Key words: Astragalus, Fabaceae, Iran.

Astragalus L. (Fabaceae) is probably the largest ge¬
nus of flowering plants, containing up to 3000 species
(l/K-k & Simpson, 1991). Iran is one of the biggest
centers of diversity for the genus (Ghahremani-nejad,
2(MX)), with approximately 700 species with 56% en¬
demism (Maassoumi, 1998). The new species belongs
to A. subg. Cercidothrix Bunge, which is characterized
by perennial growth and the presence of bifurcate
hail’s (Bunge, 1868). This subgenus has nearly 800
species in the Old World, 157 (20%) of which occur
in Iran. In Iran, subgenus Cercidothrix has 83 (53%)
endemic species (Ghahremani-nejad, 2000). Major ar¬
eas of species endemism for this subgenus occur in
Turkey, Iran, Kazakhstan, Afghanistan, and Gliina,
with }58. 81, 47, 38, and 32 endemic species reported,
respectively (Ghahremani-nejad, 2000).

Astragalus tuyehensis Ghahreman, Maassoumi &
Ghahremani-nejad, sp. nov. TYPE: Iran. Sem-
nan: 35 km N of Damghan, above Tuyeh (SW
slope), 2(MX) m, 16 May 1978. P. Wendellxnv &
M. Assadi 2948a (holotype, TARI). Figure 1.

Haec species ad Astragalum subg. Cercidotrichem per-
tinens A. durandiano Aitchison & Baker maxime affinis,
sed ab eo calyce inftato oblongo-ovoideo (vs. cylindrico)
atque fructu non tumido facile distinguitur.

Perennial, ca. 8-13 cm tall, vested with bifurcate
hairs; caudex branched, short, woody. Stems 1—2 cm
long, whitened by a dense coat of short appressed
white hairs. Stipules partly adnate at base to petiole,
free (non-connate), lanceolate-ovate, 2—3 mm long.

densely canescent with appressed white or mixed
white and black hairs. Leaves imparipinnate, 1.5— 3.5
cm long, both petiole and rachis densely canescent;
petiole 1 .5—2 cm long, 2. 5-3.5 times the length of the
rachis, partially persistent but not spinescent; leaflets
in 2 or 3 pairs, elliptic, elliptic-oblong to rarely ellip-
tic-obovate, acute, 5—9 X 1.5— 2.5 mm, canescent on
both sides. Peduncles 3—5.5 cm long, somewhat short¬
er than to 2 times leaf length, canescent; inflorescence
ovate-spherical, 3- to 10-flowered; bracts lanceolate-
ovate to lanceolate, (1)2—3 X 1—2 mm, canescent with
ascending black and mixed, or exclusively white
hairs; bracteoles wanting; pedicel 2 mm long. Flowers:
ealvx at first cylindric, becoming oblong-ovoid inflated
(to 6 mm diam.) in fruit, 12-14 mm long, finally rup¬
tured or not ruptured by fruit, canescent with ascend¬
ing white and black hairs, the calyx tube 4—5 times
the length of the lanceolate teeth; corolla yellow to
yellow-brown (known only from dry state). Standard
20 mm long, 10 mm broad, retuse, the limb elliptic-
ovate, not auriculate, 4 to 5 times as long as the claw;
wing 20 mm long, the limb oblong-elliptic, 7—8 X 3—
3.5 mm, obtuse; keel 19 mm long, the limb 6 X 3.5
mm; ovary 9—10 mm long, subsessile, densely white-
sericeous, imperfectly bilocular, 18-ovulate; ovules re-
niform, 0.25 X 0.2 mm; style 9-10 mm long, gla¬
brous, basally pilose. Pod 9-10 mm long, terminating
in a beak 1 — 1.5 mm long, somewhat bilocular, dense¬
ly white-sericeous.

Taxonomic and distributional remarks. Astragalus
tuyehensis has not been seen in the field by the au¬
thors, but is known to us only from four specimens,
one at TARI, and three at FUMI I. The species occurs
in the provinces of Semnan and Khorassan (Fig. 2).
The closest relative of the new species is A. dur¬
andianus Aitchison & Baker (J. E. T. Aitchison 688,
type, LE!), also from subgenus Cercidothrix, and
shares the characters of short overall height, bifur¬
cate hairs, short stem, bracteole wanting, with stip-

Novon 12: 47-49. 2002.

48

Novon

Figure I. Astragalus luyehensis Ghahreman, Maassoumi & Ghahremani-nejad. Habit with details of flowers. — 1.
Habit. — 2. Magnified view of raehis and leaflets. — 3. Calyx. — 4. Floral standard. — 5. Wing. — 6. Keel. — 7. Stamens.
— 8. Pistil. — 9. Fruit. (From the holotype: Wendelbow & Assadi 2948a.) (1, scale bar = 2 cm; 2, 9. scale bar = 6
mm; 3—8, scale bar = 1 cm.)

ules adnate at the base on one side of the petiole
and free (noil-connate) on the other side. Astragalus
durandianus, which lias a small distributional
range in western Afghanistan and eastern Iran
(Lock & Simpson, 1991; Hechinger, 1957: 133), is
a perennial plant with bladdery-swollen fruit. The
new species is readily separated from A. durandi¬
anus by its oblong-ovoid inflated calyx (not cylin-
tlric) and non-swollen fruit. Moth of these species
are excellent candidates for section Erioceras, if
only one character lor each species is ignored (the

swollen fruit ol A. durandianus and the swollen ca¬
lyx of A. tuyehensis). Future molecular systematic
work will be necessary to help us better understand
the sectional placement ot selected Old World As-
tragalus.

Paratypes. IK AN. Miorassan: between Mashhad and
Neyshabur, Piveh-jan, 29 Apr. 1985, //. Safari & M. Jo-
hartchi 12841 (FUMH); Golmakan, S of Esjil, I May 1985,
M. Johartchi 12528 (FUMH); S of ( Auchan. Yadak, 1 700
m. 15 May 1985, M. Joharchi & Zanguee 12905 (FUMH).

Acknowledgments. We thank Sepideh Parsa-pa-

Volume 12, Number 1
2002

Ghahreman et al.

Astragalus tuyehensis from Iran

49

44 46 41 50 52 54 56 51 M H »4

jouh for preparing the illustration. We also thank

John F. Gaskin (MO) for his very able assistance.

Literature Cited

Bunge, A. 1868. Generis Astragali Species Gerontogeae
1. Claves diagnosticae. Mem. Acad. Imp. Sci. Saint Pe-
tersbourg, ser. 7, 11(16): 1—254.

Ghahremani-nejad, F. 2000. Systematic and Biosystematic
Studies on the Sections of Perennial Bifurcate Astrag¬
alus L. (Fabaceae). Ph.D. Dissertation, Department ol
Biology, University of Tehran, Tehran. [Unpublished.]

Lock, J. & K. Simpson. 1991. Legumes of West Asia, a
Check-list. Royal Botanic Gardens, Kew.

Maassoumi, A. A. 1998. Astragalus in the Old World, a
Check-list. Research Institute of Forests and Range-
lands, Tehran.

Rechinger, K. H. 1957. Leguminosae. In: M. koie & K.
H. Rechinger (editors), Symbolae Afghanicae 3. Biol.
Skr. 9, 3: 1-208.

Phlomis tathamiorum (Lamiaceae), a New Species from Lebanon

Ricardns M. Haber

Department of Sciences, American University of Technology (AUT), PO Box 967,

Jounieh, Lebanon

Myrna T. Seaman

Center for Environmental Development, Awareness & Research (C.E.D.A.R.), and
Department of Sciences, Notre Dame University, PO Box 967, Jounieh, Lebanon

Abstract. A new species of the genus Phlomis
L. is described. Plilomis tathamiorum R. M. Haber
& Semaan is distinguished by the following char¬
acters: whole plant eglandular; basal leaves cor¬
date, oblong-lanceolate; indumentum almost solely
pannose, of layered unstalked, short-stalked, and
long-stalked stellate triehomes; inflorescence pa¬
niculate with lew 1 — 6( — 7) mm long subulate to lan¬
ceolate bracteoles; verticillasters and flowers nu¬
merous; calyx tubular, enlarged at middle; lower lip
of corolla longer; nutlet t ip stellate with stipitate
glands.

Key words: Lamiaceae, Lebanon, Phlomis.

For two decades the authors have joined their
efforts in reviewing the described flowering plants
of Lebanon to update their geographical distribu¬
tions and amass botanical photographic records to
be used in awareness campaigns and education kits
concerning the importance of biodiversity and eco¬
logical stability. This extensive research resulted in
the discovery of a new species of the genus Phlomis
L. (Lamiaceae).

file genus Phlomis has until now been repre¬
sented in Lebanon by eight species (Huber- Morath,
1 982; Mouterde, 1983). W bile six species (P.
amanica Vierhapper, P kurdica Rechinger f., P.
longifolia Boissier & Blanche, P. rigida Labillar-
diere, P. syriaca Rechinger f., and P. viscosa Poiret)
are also distributed in Turkey and the Levant. P.
brevilabris Ehrenberg and P. brachyodon subsp. da¬
rn ascena (Bornmiiller) Samuelsson are endemic to
Lebanon and Syria.

Phlomis tatliamiorimi R. M. Haber & Semaan,
sp. nov. T\ PE. Lebanon: North Lebanon Prov¬
ince: Ehden town, 1400 m, 34°17'32.3"N.
35°56'51.9"E, 15 Aug. 2000, K. Haber & M.
Semaan s.n. (holotype, MO; isotypes, PH,
RBGK). Figure 1.

Herba perennis eglandulata; radice lignosa; indumento

panniformi fere omnino stellato ex trichomatibus sessili-
bus stipitatisque constantei; caide erecto usque ad lit)
cm alto, quadrangulato, rigido autem fragili, infra brun-
neolo supra purpurascente, trichomatibus slellatis albis
apprime circa nodos densis floceosisque pannoso. Folia
liasalia 7-13 X 2. 3-6.0 cm, cordata, oblongo-lanceolata,
acutata, minute crenata. Inflorescentia ex verticillastris
multifloris in paniculum dispositis constans; bracteolis
paucis, 1— 6(— 7) mm longis, lineari-lanceolatis, subulatis.
Flos cafyce tubulari, 14—16 mm longo, ad medium dila-
tato, intus villoso, dentibus 4—6 mm longis, inaequalibus,
acumiuatis; corollae labio iuferiore quam superiore lon-
gioris. Nuculae ad apices trichomatibus steflatis persis-
tentibus ornatae.

Perennial eglandular herb with ligneous base; in¬
dumentum pannose, of dense white and yellow lay¬
ered unstalked, short-stalked, and long-stalked
erect stellate triehomes, with mixed adpressed stel¬
late and dendroid triehomes only at base of calyces
and on upper side of petioles; stem erect, to 110
cm high, quadrangular, rigid but brittle, brownish
below, purplish above, pannose with white stellate
triehomes, these especially dense and floccose
around nodes. Basal leaves with petiole 3—6 mm
wide, up to 15 cm long; blade 7—13 X 2.5— 6.0 cm,
oblong-lanceolate, broadly acute, usually deeply
cordate, rarely cuneate, minutely crenate, slightly
undulate, conduplicate or rarely flat, rugose espe¬
cially below, coriaceous, pannose with triehomes
white or yellow especially beneath along midvein
and main veins; cauline leaves with petioles 3—5
cm below, becoming shorter and more slender up¬
ward; blade 6—10 X 2.5— 4.0 cm, acute, slightly
conduplicate, pannose, the lower cordate-lanceo¬
late and similar to basal leaves, becoming oblong-
lanceolate upward, base cuneate to attenuate, often
unequal; floral leaves shortly petiolate, rarely ses¬
sile; blade pannose, 1. 5-5.0 X 0.4— 1.5 cm, oblong-
lanceolate, conduplicate, acute, attenuate, longer
than lower verticillasters, shorter upward. Inflores¬
cence a panicle of verticillasters; terminal axis with
5 to 10 lax verticillasters, each 8- to 14(usually 10)-
flowered; lateral branches up to 8, sometimes lon-

Novon 12: 50-52. 2002.

Volume 12, Number 1
2002

Haber & Semaan

Phlomis tathamiorum from Lebanon

51

ger than terminal, major branches with I to 5 (or
6) verticillasters each 5- to 10-flowered; petioles 0-
2 mm long; bracteoles 1 — 6( — 7) mm, few (ca. 20).
herbaceous, linear to lanceolate, subulate, rarely
clavate, sometimes irregular or dimidiate; pannose
outside with evident dendroid-stellate trichomes,
often yellow at tip, glabrescent inside. Calyx 11—
16 mm long, tubular, broadest (4-7 mm) in middle,
pannose outside, especially around base, villous in¬
side, 10-veined, the teeth unequal. 4—6 mm long,
triangular-lanceolate, acuminate; corolla 25—40 mm
long, yellow, lower lip orange-tipped, exceeding up¬
per lip; upper lip helmet-shaped, pannose outside,
villous inside at tip and lateral margins, glabrous
at center; lower lip pannose outside with sessile
stellate trichomes especially toward margin, sparse¬
ly villous inside, densely and minutely verrueulate
in middle; corolla tube annulate within; stamens
included in corolla, inserted just below annulus;
periphery of nectar disk raised into two large and
two small fringed, glandular-pubescent, succulent,
reniform-lenticular, basally joined septa, the larger
shallowly sulcate from middle to base, the smaller
brachiate between two pairs of nutlets. Nutlets
tipped with persistent stellate trichomes and many

st i pitate glandules, only one maturing per ovary;
mature seed 5-9 X 2.5— 3.5 mm. brown, obovate-
oblanceolate, rugose, slightly dorsoventrally flat¬
tened, with prominent funiculus.

Distribution and habitat. The species is rare,
localized in the rocky mountain pastures around the
town of Ehden, North Lebanon, which lies on the
western slopes of the Mount Lebanon Range at mid
altitudes (1400-1700 m). Disturbance and loss of
habitat are inferred to influence the abundance of
the species. Phlomis tathamiorum occurs today in
wild land lots patched among considerable areas
that have been claimed for development, agricul¬
ture, and diverse human activities. The remnant
habitat areas are subject to various disturbance fac¬
tors. Threats by real estate development and rec¬
reational activities seem imminent. In 1997, the au¬
thors transplanted stem bases from different
populations to privately owned land within the hab¬
itat range of P. tathamiorum. The transplants have
matured to seeding individuals. The flowering pe¬
riod of P. tathamiorum extends from July to August.

The differentiation of P. tathamiorum from the
other Phlomis species in Lebanon is emphasized in
the following key.

52

Novon

la. Corolla pink . P rigida

II). Corolla yellow, sometimes with brownish upper
lip.

2a. Shrubs; bracteoles 1 ()— 2( ) mm long, about as
long as calyx tube, numerous, dense.

3a. Calyx teeth up to I mm long . . P. amanica
3b. Calyx teeth 1—12 mm long.

4a. Calyx and bracteoles viscid . . .

. P. viscosa

4b. Calyx and bracteoles not viscid . .

. P. longifolia

2b. Herbs; bracteoles 1 .5 — 9( — 12) mm long, usu¬
ally few or absent, weak.

5a. Calyx tubular or narrowly campanulate;
tube enlarged, broadest above.

6a. Indumentum floccose, not persis¬
tent; stem and leaves become gla-
brescent; corolla lips equal . . .

.... P. brachyodon subsp. damascena
6b. Indumentum persistent; corolla lips
unequal.

7a. Indumentum yellowish gray;
calyx sub-adpressedly tomen-
to.se; bracteoles 2—6 mm long

. P. syriaca

7b. Indumentum gray; calyx erect¬
ly tomentose; bracteoles longer

. P. brevilabris

5b. Calyx tubular; tube broadest at middle.

8a. Nutlets eglandular, glabrous; brac¬
teoles 1.5-2 mm long ... P. kurdica
8b. Nutlets with stipitate glands and
stellate trichomes at apex; bracte¬
oles 1-6 mm long . . . P. tathamiorum

In addition, Phlomis tathamiorum differs from P.
kurdica by its taller stems, its more branched inflo¬
rescence with more verticillasters, and attenuate-
based floral leaves. It differs from P. syriaca by
longer-petiolate basal and cauline leaves, more
branched inflorescence, more flowers per verticil-
laster, oblong-lanceolate floral leaves, and pubes¬
cent nutlets. It is differentiated from P. brevilabris
and P. brachyodon subsp. damascena by its taller
stems, its more branched inflorescence, more flow¬
ers per verticillaster, and shorter bracteoles. It also

differs from P brevilabris by pubescent nutlets, and
from P. brachyodon subsp. damascena by the un¬
equal lips of the corolla, the inferior being longer.

The geographical distribution of P. tathamiorum
is sympatric with that of P syriaca and P. brevilabris
(Mouterde, 1983). They co-exist in the mid alti¬
tudes of the western slopes of the Mount Lebanon
Range in a Mediterranean climate associated with
a snowy season (Service Meterologique, 1966,
1967). However, the expansion range of P tath¬
amiorum is distinctly allopatric from that of P. kur-
dica and P. brachyodon subsp. damascena (Mou¬
terde, 1983). The occurrence of the latter taxa is
restricted to the inland, where climatic conditions
are semi-arid (Service Meterologique, 1966, 1967).

The species is named in gratitude and respect
after H.E. Mr. David Tatham, former United King¬
dom Ambassador to Lebanon, and to Mrs. Valerie
Tatham, for their invaluable support of conservation
efforts for Lebanon’s wildlife and natural resources,
and most especially for their love of Lebanon’s wil¬
derness.

Paratype. LEBANON. North Lebanon Province:
Ehden town, 1400 m, 34°17'38"N, 35°56'45"E, 12 Sep.
2000. R. Haber & M. Semaan s.n. (MO. I’ll, RBGK).

Literature Cited

Huber-Morath, A. 1982. Pldomis. Pp. 102-126 in P. II.
Davis, Flora of Turkey, Vol. 7. Edinburgh llniv. Press,
Edinburgh.

Mouterde, P. 1983. Pldomis. Pp. 130—136 in Nouvelle
Flore du Liban et de la Syrie, Vol. 3. Dar El-Machreq,
Beirut.

Service Meterologique. 1966. Atlas Climatique du Liban:
Pluie, Temperature, Pression, Nebulosite, Tome I. Di¬
rection de P Aviation Civil, Service Meterologique, Bey¬
routh.

- . 1967. Atlas Climatique du Liban: Humidite At-

mospherique, Statist ique Diverses de Frequence, Tome
II. Direction de V Aviation Civil, Service Meterologique,
Beyrouth.

A New Species in Potent ilia Section Leptostylae (Rosaceae) from

Yunnan, China

/tiros hi Ike da

Faculty of Informatics, Okayama University of Science, Ridai-cho 1-1, Okayama-shi,
Okayama Pref., 700-0005 Japan, ikeda@big.ous.ac.jp

Hideaki Ohba

Department of Botany, University Museum, University of Tokyo, Hongo 7-3-1, Tokyo,
113-0033 Japan, ohba@um.ii-lokyo.ae.jp

Sugong Wu

Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, Yunnan,

China, sugong@public.km.yn.cn

ABSTRACT. A new species, Potentilla polyphyllo-
ides H. Ikeda & II. Ohba, is described and illus¬
trated. It is assigned to section Leptostylae by hav¬
ing an herbaceous habit, slender or fusiform styles,
auricles of stipules of basal leaves situated at ad-
axial side of petioles, and flowering stems from ax¬
ils of basal leaves. It resembles P. polyphylla Wal-
lieli ex Lehmann, but dilfers by its glandular hairs
on the lower surface of leaflets, pedicels, and the
outside of episepals, as well as the occasional root¬
ing at the axils of cauline leaves on the lower por¬
tion of flowering stems. Potentilla polyphylloides re¬
sembles P fallens Cardot sharing the glandular
hairs, but P. fallens differs in having nearly entire
episepals that are shorter than the sepals, and lon¬
ger styles nearly twice as long as the ovaries. Po¬
tentilla polyphylloides has a chromosome number of
2n = 42, while that of P. polyphylla and P. fallens
is 2 n = 28.

key words: China, chromosome number, Poten¬
tilla, Kosaceae, section leptostylae.

Ikeda and Ohba (1993, 1999) revised the tax¬
onomy of Potentilla sect. Leptostylae in the Hima¬
layas and adjacent regions. Species of section Lep¬
tostylae are characterized by sharing an herbaceous
habit, slender or fusiform styles, auricles of stipules
of basal leaves situated at the adaxial side of pet¬
ioles, and flowering stems from the axils of basal
leaves (Ikeda & Ohba, 1999). In 1994 and 1996,
Wu et al. collected specimens of one Potentilla re¬
sembling P. polyphylhi Wallich ex Lehmann in a
meadow on Ml. Cangshan, Yunnan Province, south¬
west China. It was assigned to section Leptostylae,
but Ikeda and Ohba (1999) excluded this from their

revision due to insufficient material. After com¬
pleting the manuscript of this revision, Wu and Ike¬
da found the same Potentilla again in a meadow on
Mt. Cangshan in 1997. After the study of its mor¬
phological variation, chromosome number, and re¬
productive biology, it is concluded that the material
represents a new species.

Somatic chromosomes were counted from root
tips. In the field root tips were pretreated in 2 mM
8-hydroxyquinoline solution for 2—3 hours and
fixed in Newcomer’s fluid. They were subsequently
macerated in IN HCI at 60°C for 10.5 min., stained
with 2% lacto-propionic orcein, and squashed for
cytological observation.

Potentilla polyphylloides H. Ikeda & II. Ohba,
sp. nov. TYPE: China. Yunnan: Yangbi, Cang¬
shan— a Fruit Garden— Yangbi, 3230 m, 19
Aug. 1997, S. G. Wu, //. Ikeda, S. Akiyama, F.
Miyamoto & W. Chen 91 (holotype, K1JN; iso¬
types, A, E, HYO, MO, II). Figures I, 2.

Alt affini P. polyphylla Wallich ex Lehmann perspicue
differt hypanthio, pedicellis et foliolis subtus glandulosis.
Planta propter indumentum glandulosum P.fallenti Cardot
appropinquat, sed episepalis 3— 7-dentatis sepal is longior-
ibus et stylis ovariis brevioribus satis distineta.

Perennial acaulescent herb with thick simple or
sometimes branched rootstocks. Basal leaves ob-
lanceolate, imparipinnate, 8—20 X 2—4 cm, petio-
late, forming a rosette; lateral leaflets 10 to 15
pairs, serrate, gradually decreasing in size toward
base, with alternating smaller leaflets; petioles 0.8—
3.5 cm long, with ascending or patent unicellular
hairs; base of uppermost leaflet pair cuneate. Leaf¬
lets with appressed or ascending unicellular hairs

Novon 12: 53-57. 2002.

54

Novon

Figure 1. Potent ilia polyphylloides H. Ikcda & H. Ohba, habit. Seale bar = 2 cm. Drawn from the Wn el al. 91
isotype at TI.

on upper surface, glandular hairs and ascending
unicellular hairs on lower surface; terminal leallet
subsessile, oblong to obovate, 1.2-2. 4 X 0.7— 1.4
cm, serrate with 17 to 33 teeth per leaflet. Auricles
of stipules free. Peduncles 17—60 cm long, ascend¬
ing or prostrate, with appressed or ascending uni¬
cellular hairs, occasionally rooting from the nodes.
Catiline leaves trifoliolate or with 2 to 8 pairs of
leaflets, sometimes with alternating smaller leaflets.
Auricles of stipules entire or with a few teeth in
lower ones, 10 to 20 teeth in upper ones. Pedicels
1.2— 3.8 cm long, with glandular hairs and ap¬
pressed or ascending unicellular hairs. Flowers in
a diehasium, 1.1 -1.5 cm across. Hypanthia 4—8
mm across. Episepals oblong to widely obovate,

2. 5- 5. 5 X 2-4.5 mm, usually serrate with 3 to 7
teeth, rarely entire, apex acute, with glandular hairs
and ascending unicellular hairs on both sides, usu¬
ally longer than sepals. Sepals elliptic to ovate,

2.5- 4 X 1.8-4 mm, entire, apex acute to obtuse,
ascending unicellular hairs on outer side and mar¬

gin. lanate apieally, glabrous basally on inner side.
Petals obovate to widely obovate with round or re-
tuse apex, 5-6.5 X 3.5-7 mm. Stamens 20, in 3
whorls; antisepalous ones 5, from the inner whorl
longer than others, 1.6-2. 2 mm long; antipetalous
stamens 5, from the middle whorl shorter than oth¬
ers. 1.2— 1.3 mm long; those located between petals
and sepals 10, from the outer whorl, 1.4— 1.6 mm
long; anthers spheroidal, subbasal, with 4 locules,
0. i -0.8 mm across. Ovaries ellipsoid, smooth, 0.5—
0.6 X ca. 0.4 mm; style subbasal, 0.8-0.9 mm long;
stigma slightly inflated, papillate; placenta located
at ventro-lateral side near style base. Achenes
many, on dome-shaped receptacle, obliquely ellip¬
soidal, smooth, 1.2— 1.4 X 0.7— 0.9 mm.

Potentilla polyphylloides resembles P. polyphylla
in sharing 3- to 7-dentate episepals longer than se¬
pals and relatively short styles (ca. 1.5X as long as
ovaries). However, P. polyphylloides differs from P.
polyphylla in having glandular hairs on the lower

— »

Figure 2. Floral dissections comparing Potentilla polyphylloides (A-F), P. polyphylla (G — I), and P. fallens ( J— M ). —
A, O. J. Episepals, inner surface (upper) and outer surface (lower). — B, H, K. Sepals, inner surface (upper) and outer
surface (lower). — C. Petal. — D. Three types of stamens: antisepalous ones (upper left two), between petals and sepals

Volume 12, Number 1
2002

Ikeda et al.

Potentilla polyphylloides from China

55

(upper right two), and antipetalous (lower two). For each stamen pair, inner surface (left) and outer surface (right). —
E, I, L. Pistils. — F, M. Multicellular hairs with glandular lips. Scale bars: A, B, G, H, J, k to same scale Bar = I
mm; C, I) scale bars = 1 mm; E, I, L to same scale bar = 0.5 mm; F, M to same scale bar = 0.5 mm.

56

Novon

Figure 3. Somatic chromosomes at metaphase of Potentilla polyphylloides. Upper: microphotograph of chromosomes.
Lower: drawing based on photograph. Scale bar = 5 gm.

surface of leaves, pedicels, and the outside of ep-
isepals, as well as occasional rooting at axils of
cauline leaves along the lower portion of flowering
stems. Potentilla polyphylloides resembles P. fallens
Cardot in sharing glandular hairs, hut P. fallens dif-
lers from P. polyphylloides by its nearly entire ep-
isepals shorter than the sepals and the styles nearly
twice as long as the ovaries (Fig. 2).

Potentilla polyphylloides makes gregarious com¬
munities on grazed grassland slopes between 3200
and 3500 m altitude, and sometimes it becomes a
dominant species. It bears flowers and fruit well in
the field and may reproduce sexually by seeds as
well as vegetatively by detaching from rooted nodes
of flowering stems; P. polyphylla and P. fallens re¬
produce only by seeds.

Potentilla polyphylloides is known only from Mt.
Cangshan. Potentilla polyphylla is distributed in
the Himalaya, southwestern China (Yunnan Prov¬
ince), and southeastern Asia (Java); P. fallens oc¬
curs in southwestern China in Yunnan and Sichuan
Provinces. These two species grow on grassland,
similar to P. polyphylloides , but they have not been
recorded from Ml. Cangshan (Ikeda & Oliba, 1993).

figure 3 shows the somatic chromosomes of P.
polyphylloides. The chromosome number of P. po¬
lyphylloides is 2/i = 42. different from the 2 n =

28 of P. polyphylla (Ikeda Ohba, 1993) and P.
fallens (Ikeda & Ohba, unpublished). Since the ba¬
sic chromosome number of Potentilla is x = 7 (Shi-
motomai, 1930), P. polyphylloides is a hexaploid
species while P. polyphylla and P. fallens are tet-
raploid taxa.

W ith the addition of Potentilla polyphylloides ,
the key to the species ol Potentilla sect. Leptostylae
series Lineatae presented in Ikeda and Ohba (1993,
1999) should be changed as follows.

Key to the Species of Potentieia section Leptostyiae
series Lineatae

la. Tw o auricles of stipules of basal leaves connate

from base to middle . P. festiva Sojdk

lb. Two auricles of stipules of basal leaves free from
each other.

2a. Base of uppermost leaflet pair decurrent . .

. P. josephiana H. Ikeda & H. Ohba

2b. Base of uppermost leaflet pair euneate.

3a. Peduncles and hypanthia without glan¬
dular hairs .

. P polyphylla Wallich ex Lehmann

3b. Peduncles and hypanthia with glandular
hairs.

4a. Stigma not inflated; leaves densely
sericeous on lower surface . . .

. P. lineata Treviranus

4b. Stigma inflated; leaves sparsely
strigose on lower surface.

Volume 12, Number 1
2002

Ikeda et al.

Potentilla polyphylloides from China

57

5a. Episepals nearly entire, shorter
than sepals; styles nearly twice

as long as ovaries .

. P. fallens Cardot

5b. Episepals with 3 to 7 teeth,
longer than sepals; styles 1 .5
times as long as ovaries ....

. . P. polyphylloides H. Ikeda & H.

( )hba

Paratypes. CHINA. Yunnan: Dali, Cangshan, Zhon-
ghe Peak, 3390 m, 3 Oct. 1994, S. G. Wu et al. 1305
(KUN, TI); Dab, Cangshan, 3440 m, 1 Sep. 1996, S. G.
Wu et al. 968 (KUN, Tl); Yangbi, Cangshan— a Fruit Gar¬
den— Yangbi, 3500 m, 19 Aug. 1997, S. G. Wu et al. 92
(KUN, TI).

Acknowledgments. We thank Futoshi Miyamoto,
Tokyo University of Agriculture, for the drawing of
P. polyphylloides. This study was supported by a

Grant-in-Aid for Scientific Research (A), No.
1 1691178 in 1999. 2000, and 2001 (to Ohba) from
the Japan Society for the Promotion of Science, and
also by a fund from the Hyogo Science and Tech¬
nology Association in 1996 (to Ikeda).

Literature Cited

Ikeda, H. & H. Ohba. 1993. A systematic revision of Po¬
tentilla lineata and allied species (Rosaceae) in the
Himalaya and adjacent regions. Rot. J. Linn. Soc. 112:
159-186.

- & - . 1999. A systematic revision of Poten¬
tilla L. section Leptostylae (Rosaceae) in the Himalaya
and adjacent regions. Pp. 31—117 in H. Ohba (editor),
The Himalayan Plants, Vol. 3. Univ. Tokyo Press, To¬
kyo.

Shimotomai, N. 1930. Chromosomenzahlen und Phylogen-
ie bei der Gattung Potentilla. J. Sci. Hiroshima Univ.,
Ser. B, l)iv. 2, 1: 1—11.

Taxonomic Status of Acleisanthes , Selinocarpus , and
Ammocodon (Nyctagi naceae)

Rachel A. Levin

Department of Ecology & Evolutionary Biology, University of Arizona, Tucson, Arizona
85721, U.S.A. Current address: Laboratory of Molecular Systematics, National Museum of
Natural History, Smithsonian Museum Support Center, 4210 Silver Hill ltd.. Suitland,
Maryland 20746, U.S.A. rlevin@lms.si.edu

ABSTRACT. Recent molecular and morphological
studies suggest that neither Acleisanthes nor Seli¬
nocarpus (Nyctaginaceae) are monophyletic as cur¬
rently circumscribed. Further, these data provide
no basis lor continued recognition of the monotypic
genus Ammocodon as separate from these other two
genera. Therefore, all 8 Selinocarpus species and
the monospecific Ammocodon are transferred to
Acleisanthes ; 3 additional Selinocarpus species are
reduced to synonymy. The genus Acleisanthes is
now composed of 16 species, distributed primarily
in the Chihuahuan Desert of North America. Di¬
agnostic features of Acleisanthes include an herba¬
ceous or woody perennial habit in arid environ¬
ments, and the presence ol both eleistogamous and
chasmogamous flowers often simultaneously on an
individual plant. Further, flowers are usually axil¬
lary and subtended by 1 to 4 subulate bracts, with
chasmogamous flowers frequently white or yellow
and funnelform to salverform.

Key words: Acleisanthes, Ammocodon, Chihu¬
ahuan Desert, Four o’clock, Nyctaginaceae, Seli¬
nocarpus.

'Die genus Acleisanthes A. Gray (Nyctaginaceae)
consists of 7 species distributed from southern Cal¬
ifornia east to the Texas coast and south to San Luis
Potosf, Mexico, with its center of distribution in the
Chihuahuan Desert (Smith, 1976). Selinocarpus A.
(nay is another small genus of I I species found in
the southwestern United States and northern Mex¬
ico, with 1 species disjunct and endemic to Somalia
(Fowler & Turner, 1977; Thulin, 1993; Turner,
1993). Like Acleisanthes, Selinocarpus is most spe¬
cies-rich in the Chihuahuan Desert of New Mexico,
Texas, and Mexico. Within this region, many Seli¬
nocarpus species appear to prefer gypseous sub¬
strates (Fowler & Turner, 1977), whereas Acleis¬
anthes inhabit a variety of soils, including those
that are gypseous (Smith, 1976; Levin, pers. obs.).
Although the genera are morphologically similar,
Selinocarpus species have been characterized by

the presence of winged fruits similar to those found
in its namesake genus Selinum in Apiaeeae (Gray,
1853). Further, the majority ol Selinocarpus species
are perennial shrubs, whereas Acleisanthes species
are all herbaceous perennials. However, the genera
are similar in other aspects, including the produc¬
tion ol eleistogamous flowers in addition to the often
funnelform, hawkmolh-pol limited, chasmogamous
flowers with nocturnal anthesis. Since the original
descriptions of these genera by Gray (1853), Stan-
dley (1916) segregated Selinocarpus chenopodioides
A. Gray into the monotypic genus Ammocodon
Standley as A. chenopodioides (A. Gray) Standley
based on characters now understood to be autapo-
morphic. including a reduction in stamen number
and its many-flowered inflorescences versus the
usually solitary flowers found in other species of
Selinocarpus.

Recent phylogenetic analyses (Levin, 200(f) of
these genera based on molecular sequence data in¬
cluding the nuclear ribosomal ITS region and a re¬
gion of the chloroplast genome between the rhe L
and rtceD genes and confirmed by morphology sug¬
gest that their current taxonomic circumscriptions
need to be reevaluated. For example, there is strong
support for a sister taxon relationship between A.
wrightii (A. Gray) Bentham & Hooker and S. dif-
fusus A. Gray; in a combined parsimony analysis
of nuclear and chloroplast data, this relationship
was retrieved in 87% of bootstrap replicates (BS)
(Levin, 2000: fig. 2). Morphologically this result is
not surprising, as these species are very similar
vegetatively. Leaf shape and color are nearly con¬
gruent, and 5. diffusus shares an herbaceous habit
with Acleisanthes species, in contrast to the woody,
shrub habit commonly observed in the genus Seli¬
nocarpus. Further, phylogenetic results clearly split
Acleisanthes into two groups. One group is com¬
posed of three Acleisanthes species: A. anisophylla
A. Gray, A. longiflora A. Gray, and A. obtusa (Choi-
sy) Standley (BS = 87, Levin, 2000: fig. 2). In ad¬
dition to strong support for this elade from molec-

Novon 12: 58-63. 2002.

Volume 12, Number 1
2002

Levin

Genera in Nyctaginaceae

59

ular data, a synapomorphy for this group of
Acleisanthes is the presence of 5 stamens in cleis-
togamous flowers, in contrast to the 2 stamens
found in the cleistogamous flowers of the four other
Acleisanthes and many Selinocarpus species. Inter¬
estingly, plants of this lineage of three Acleisanthes
species also have fruits lacking resinous glands. By
contrast, all but the basal member of the other
Acleisanthes group possess resinous glands on their
fruits. The second group of Acleisanthes species in¬
cludes the basal A. crassifolia A. Gray, A. nana I.
M. Johnston, A. acutifolia Standley, A. wrightii, and
S. diffusus. These taxa are more closely related to
all other Selinocarpus species and Ammocodon
chenopodioides than to the other three Acleisanthes
species (BS = 100, Levin, 2000: fig. 2), which are
sister to the second group of Acleisanthes plus Se¬
linocarpus species and Ammocodon chenopodioides.
Thus, there is robust evidence that Selinocarpus
species and Ammocodon chenopodioides are nested
within a paraphyletic Acleisanthes. Consequently, I
suggest that a single taxon composed of Acleisan¬
thes and Selinocarpus species and Ammocodon
chenopodioides be recognized.

Asa Gray (1853) first described both Acleisanthes
and Selinocarpus at the same time; thus, neither
name has clear priority. Therefore, l propose that eight
Selinocarpus species, including S. angustifol ms I < urey,
S. diffusus , S. lanceolatus Wooton, S. palmeri Hemsley,
5. parvifolius Torrey, 5. purpusianus Heimerl, 5. so-
malensis Chiovenda, and S. undidatus B. A. Fowler
& B. L. Turner, and Ammocodon chenopodioides be
transferred into Acleisanthes. Based on morphological
evidence (see below), an additional three Selino¬
carpus species, S. nevadensis (Standley) B. A. Fowl¬
er & B. L. Turner, S. maloneanus B. L. Turner, and
S. megaphyllus B. L. Turner, are synonymized. With
the proposed combinations and synonymy, Acleis¬
anthes would now be composed of 16 species, all
characterized by a perennial habit and the presence
of both cleistogamous and chasmogamous flowers
that are often present simultaneously. All species
produce funnelform to salverform chasmogamous
flowers that are frequently white or yellow, less of¬
ten pink or purple, and each flower is subtended
by 1 to 4 subulate bracts. Flowers are generally
solitary and axillary, most are hawkmoth-pollinated
and fragrant, and their single-seeded Iruils are ei¬
ther ribbed or winged. Many of these species can
be locally common, but are rare at the landscape
level and tend to be limited to certain substrates.
The following key to Acleisanthes is based upon my
study of both fresh and herbarium material and rel¬
evant literature (Smith, 1976; Fowler & Turner,
1977; Timlin. 1993). The fruit characters included

in the key to Acleisanthes are not difficult to ob¬
serve, because plants are rarely found without fruits
from either chasmogamous or cleistogamous flowers
in various stages of maturity.

Key to the Species ok Acleisanthes (encompassing the
former taxa Selinocarpus and Ammocodon)

1 a. Plants herbaceous perennials.

2a. Fruits winged, not ribbed.

3a. Flowers in cymose inflorescences of
few to numerous (10 or more) flow¬
ers; chasmogamous perianth magen¬
ta-purple, 4—5 mm long; stamens in
chasmogamous flowers usually 2

(rarely 3) .

... A. chenopodioides (A. Gray) R. A.

Levin

3b. Flowers solitary to paired; chasmo¬
gamous perianth white, 20—50 mm
long; stamens in chasmogamous
flowers usually 5 (rarely 4).

4a. Chasmogamous perianth 30—

50 mm long; plants found on
various substrates in the
southwestern United States . .

.... A. diffusus (A. Gray) R. A.

Levin

4b. Chasmogamous perianth 20—

35 mm long; plants found in

Somalia .

A. somalensis (Chiovenda) It. A.

I .eviri

2b. Fruits ribbed, not winged.

5a. Fruits with resinous glands at or
near proximal end.

6a. Plants with stems 2—8 cm
long; chasmogamous perianth
purple, 8—10 mm long ....

. A. nana 1. M. Johnston

61). Plants with stems 10—40 cm
long; chasmogamous perianth
white, 25—45 mm long.

7a. Plants upright; resinous
glands occurring below
proximal end of fruit;
plants found from 750 to
2500 m in elevation . .

. A. acutifolia Standley

7b. Plants more prostrate;
resinous glands occur¬
ring at proximal end of
fruit; plants found from
250 to 1000 m in ele¬
vation .

A. wrightii (A. Gray) Ben-
tham & Hooker

5I>. Fruits without resinous glands.

8a. Veins of upper leaf surfaces
with white, conspicuous tri-
chomes; cleistogamous flowers

with 2 stamens .

. A. crassifolia A. Gray

8b. Veins of upper leaf suriaces
glabrous; cleistogamous flow¬
ers with 5 stamens.

60

Novon

9a. Plants prostrate, stems
to 15 cm long; leal pairs
at a node strongly un¬
equal; veins deeply in¬
cising upper leaf sur¬
face; fruits with 1(1 ribs
... A. anisophylla A. Gray
9b. Plants prostrate to clam¬
bering, stems to 100 cm
long; leaf pairs equal to
subequal; veins of upper
leaf surface inconspicu¬
ous; fruits with 5 ribs.

10a. Flowers solitary;
e h a s m oga m o u s
perianth to 17 cm

long .

A. longiflora A. Gray
10b. Flowers solitary or
in cymes of 2-5
flowers; chasmoga-
mous perianth
2.5—5 cm long . .

... A. obtrna (Choisy)
Stam I ley

lb. Plants woody perennial shrubs.

11a. Plants with divaricating branching, hav¬
ing profusely branched stems; chasmoga-
mous perianth usually yellow, 30—50 mm
long.

12a. Shrubs small, to 3 dm tall; leaves to
40 mm long; chasmogamous peri¬
anth yellow to white .

. . A. lanceolatus (Wooton) K. A. Levin
12b. Shrubs larger, to 6 dm tall; leaves
to 20 mm long; chasmogamous peri¬
anth usually yellow, rarely white or
pink.

13a. Stems grayish purple; leaves
linear to lanceolate; chasmo-
gamous perianth yellow, white,
or pink; plants occurring on
gypsum.

14a. Chasmogamous perianth
pink; mature fruits 5—7

mm long .

.... A. palmeri (Hemsley)
R. A. Levin

14b. Chasmogamous perianth
yellow or white; mature
fruits 7.5—10 mm long
. . A. purpusianus (Heimerl)
IL A. Levin

13b. Stems not gray; leaves ovate;
chasmogamous perianth yel¬
low; plants found on sand¬
stone and limestone in addi¬
tion to gypseous soil .

... A. parvifolias (Torrey) R. A.

Levin

I lb. Plants virgate, stems having few branch¬
es; chasmogamous perianth green-yellow
to red-orange, 6—12 mm long.

15a. Leaf margins sinuous; chasmoga¬
mous perianth ochreous .

. . A. undulatus (R. A. Fowler & R. L.

Turner) R. A. Levin

15b. Leaf margins straight; chasmoga¬
mous perianth green-yellow . . .

. . A. angustifolius (Torrey) R. A. Levin

Aeleisaiithes angiistif olius (Torrey) R. A. Levin,
comb. nov. Basionym: Selinocarpus angustifol¬
ius Torrey, Rep. U. S. Mex. Bound. Surv.. Hot.,
170. 1859. TYPE: Mexico. Chihuahua: Presi¬
dio del Norte (Ojinaga), July 1852, C. C. Parry
et al. s.n. (leetotype, designated by Fowler &
Turner (1977), NY not seen; isoleetotype, US).

This species, together with A. undulatus, is dis¬
tinctive among Acleisanthes due to the presence of
short, relatively inconspicuous chasmogamous flow¬
ers. It is distinguished from its sister taxon, A. un¬
dulatus, in having flowers that are green-yellow
rather than orange, straight leal margins, greater
edaphic tolerance, and occurring over a w ider geo¬
graphical range. I have observed this species grow¬
ing in sympatry with A. undulatus , and in such ar¬
eas both species are readily discerned.

Specimens examined. U.S.A. Texas: Brewster Co.,
Tornillo Creek in Rig Bend National Park, Levin 97-1
(ARIZ); ca. 75 mi. S of Alpine, Powell 2IH0 (NMC); Pre¬
sidio Co., 2 mi. N of Presidio, Spellenberg A- Moore 2623
(NMC). MEXICO. Chihuahua: base of IN end of Sierra
del Cuchillo Parado where crossed by Ojinaga— Cd. Chi¬
huahua Hwy., 60 km W of Ojinaga, Chiang, Wendt A
Johnston 9772 (NMC). Coahuila: ca. 6.9 mi. 5W of Cua-
tro Cienegas on Uwy. to San Pedro, then 0.7 mi. F of Hwy.
in first canyon on N side of limestone Sierra San Marcos
del Pino, Henrickson 20402 (ARIZ). Durango: ca. 0.5
mi. SW of Lerdo on first limestone road cut, Spellenberg
A' Syvertsen 3763 (NMC).

Aeleisaiithes chenopodioides (A. Gray) R. A.
Levin, comb. nov. Basionym: Selinocarpus
chenopodioides A. Gray, Amer. J. Sci. Arts, ser.
2, 15: 262. 1853. Ammocodon chenopodioides
(A. Gray) Standley, J. Wash. Acad. Sci. 6: 629.
1916. TYPE: ll.S.A. Texas or New Mexico,
1851 or 1852, C. Wright 1707, left side of
sheet (leetotype, designated by Fowler & Turn¬
er (1977), “left hand specimen on sheet,” GH).

This taxon was one of the original species that
defined the genus Selinocarpus by Gray (1853),
having the characteristic winged fruits. Later this
species was segregated into its own genus. Ammo-
codon, based upon flowers that were unique among
Selinocarpus ; flowers occur in many-flowered, ey-
mose inflorescences, whereas solitary, axillary flow¬
ers are usual within the genus (Standley, 1916).
Further, chasmogamous flowers are small and have
a reduced number of stamens. However, other char¬
acters unite this species with Acleisanthes as here
circumscribed. Plants are herbaceous, a habit

Volume 12, Number 1
2002

Levin

Genera in Nyctaginaceae

61

shared with Acleisanthes s. str. as well as A. dijfusus
and A. somalensis. Molecular data suggest that this
taxon clearly belongs nested within a broadly de-
fined Acleisanthes, and that it may be sister to A.
somalensis (Levin, 2000).

Specimens examined. U.S.A. Arizona: Cochise Co.,
Portal— Rodeo Rd., 4 mi. NW of Rodeo, near cattle guard,
McCormick A Assoc. 215 (ARIZ). New Mexico: Dona
Ana Co., The Jornada LTER, G-BASN NPP site, NE cor¬
ner, Lower Summerford Mtn. Bajada slope, Levin 99—1
(ARIZ). Texas: Hudspeth Co., N end of the Quitman
Mtns. at a roadside park along 1-10, Worthington 6694
(ARIZ).

Acleisanthes difTusus (A. Gray) R. A. Levin,
comb. nov. Basionym: Selinocarpus dijfusus A.
Gray, Atner. J. Sci. Arts, ser. 2, 15: 262. 1853.
TYPE: U.S.A. Texas: Pecos Co., 1851 or 1852,
C. Wright 1708 (lectotype, designated by
Fowler & Turner (1977), GH).

Selinocarpus nevadensis (Standley) B. A. Fowler & IS. E.
Turner, Phytologia 37: 201. 1977. Syn. nov. Selino¬
carpus dijfusus A. Gray subsp. nevadensis Standley,
Contr. U.S. Natl. Herb. 12: 388. 1909. TYPE: U.S.A.
Nevada: Lincoln Co., Overton, May 1891, V. Bailey
1932 (holotype, US).

Fowler and Turner (1977) defined .S’, nevadensis
as distinct from S. dijfusus on the basis of having
wider, rounder leaves and a more narrow, delicate
chasmogamous perianth. However, these differenc¬
es are likely due to environmental effects on plastic
morphological traits. Therefore, I do not believe
there is sufficient evidence that these species are
separate, and 1 am reducing S. nevadensis to syn¬
onymy with A. dijfusus.

Specimens examined. U.S.A. Arizona: Mohave Co.,
along US Hwy. 93 ca. 30 mi. NW of Chloride, Barr 68—
283 (ARIZ). Nevada: Clark Co., ca. 2 mi. N of Jean, off
frontage road by 1-15, Levin 98—1 (ARIZ); Ml. Springs
Summit, 2.1 mi. below Blue Diamond cutoff, Niles A Blum
972 (ARIZ); Mojave Desert, ca. 30 mi. N of Las Vegas,
19 mi. S of Indian Springs, E side of IJS Hwy. 95, Spel-
lenberg 9482 (NMC). New Mexico: l)e Baca Co., 18 air
km S of Fort Sumner, E side Pecos River, summit of Espia
Peak, Spellenberg A /acker 12404 (NMC). Oklahoma:
Harmon Co., on gypsum 6 mi. S of Hollis, Waterfall 9001
(ARIZ).

Acleisanthes lanceolatus (Wooton) R. A. Levin,
comb. nov. Basionym: Selinocarpus lanceolatus
Wooton, Bull. Torrey Bot. Club 25: 304. 1898.
TYPE: U.S.A. New Mexico: Dona Ana Co.,
just S of the White Sands, 26 Aug. 1897, E.
O. Wooton 389 (holotype, LIS; isotypes, NMC,
US).

Selinocarpus maloneanus B. L. Turner, Phytologia 75: 239.
1993 [1994]. Syn. nov. TYPE: U.S.A. Texas: Hud¬

speth Co., N end of Malone Mtns., near Finley, 29
July 1958, I). S. Correll A /. M. Johnston 20358
(holotype, EE not seen).

Selinocarpus megaphyllus (B. A. Fowler A B. E. Turner)
B. E. Turner, Phytologia 75: 242. 1993 [ 1 994]. Syn.
nov. Selinocarpus lanceolatus Wooton var. megaphyl¬
lus B. A. Fowler & B. E. Turner, Phytologia 37: Dll.
1977. TYPE: Mexico. Chihuahua: ca. 15 mi. SW of
Estacion Moreon on Rio Com hos Eake Road, Sierra
de las Monillas. 25 May 1971, A. M. Bowed 2105
(holotype, LL not seen).

The main distinctions between S. maloneanus, S.
megaphyllus, and A. lanceolatus are in leaf shape
and width, and S. megaphyllus is reportedly distin¬
guished from the other two species by a larger,
white chasmogamous perianth (Fowler A Turner,
1977; Turner, 1993). However, Fowler and Turner
(1977) noted that there is morphological overlap
between .S', megaphyllus and A. lanceolatus, and the
vegetative characters that have been used to sep¬
arate these three species are known to be quite
plastic. An excellent example of the wide variation
in leaf shape within a population is the NMC col¬
lection of Spellenberg & Syvertsen 3758, listed be¬
low, that includes specimens from multiple individ¬
uals at the same locality. These individuals were
clearly nonspecific, with gradations in leaf width as
the only distinction among the individuals. 1 have
examined a specimen ( Spellenberg & Syvertsen
3753) from the same locality as one ( Powell 211 6;
Turner, 1993: 241) considered nonspecific with S.
maloneanus and a specimen ( Henrickson 6799) col¬
lected from the type locality (Turner, 1993: 242) of
S. megaphyllus. In addition to these, I have ex¬
amined other specimens of A. lanceolatus, and I do
not believe that the distinctions outlined by Turner
(1993) warrant specific recognition.

Specimens examined. U.S.A. New Mexico: Otero Co.,
about 14 mi. W of Alamogordo on US Hwy., at Point of
Sands just W of White Sands National Monument, Spel¬
lenberg 3910 (NMC). Texas: Culberson Co., 39 mi. N of
Van Horn on Hwy. 54, Levin 97—5 (ARIZ); State Hwy. 54
N of Van Horn, 15 mi. S of junction with US Hwy. 62—
180 in NW corner of county, Spellenberg A Ward 9711
(NMC). MEXICO. Chihuahua: 12 mi. NE of Cuehillo
Parado along old road in a gypsum outcropping N of Rio
Conehos, Henrickson 6799 (NMC); 13.6 kin W of Camargo
on road to I .ago Boquilla, on gypsum river terraces, Spel¬
lenberg A Syvertsen 3758 (NMC); 17.4 mi. SW of Coyame,
8.8 mi. NE of railroad crossing, Chihuahua Hwy. 16, Spel¬
lenberg A Syvertsen 3753 (NMC).

Acleisanthes palmeri (Hemsley) R. A. Levin,
comb. nov. Basionym: Selinocarpus palmeri
Hemsley, Biol. Cent.-Amer., Bot., 3: 6. 1882.
TYPE: Mexico. Coahuila: San Lorenzo de La¬
guna, May 1880, E. Palmer 1118 (holotype, k
not seen; isotype, US).

62

No von

The characters that appear to distinguish this
species from A. purpiisiamis are a smaller fruit and
the presence of a pink chasmogamous perianth,
rather than the yellow or white perianth found in
A. purpusianus. In addition to these morphological
differences, A. palmeri is known only from two
proximate localities in Coahuila, Mexico. Therefore,
it may be more appropriate to synonymize A. pal¬
meri with .4. purpusianus, a similar species also lim¬
ited to a few gypseous localities in Coahuila, al¬
though nomenelatural priority would he given to 4.
palmeri. However, as I have not observed 4. palmeri
in the field and have only viewed two herbarium
specimens ol this species, it does not seem appro¬
priate at the present time to reduce these two spe¬
cies to synonymy.

Specimens examined. MEXICO. Coahuila: ca. 15
road ini. E of Torreon and about 2.6 mi. by winding road
E of El Coyote, at the NW end of a small mtn. range, the
Sierra de Solis, mostly on the tops ol knolls. Spellenberg
X' Syvertsen .3768 (NMC).

Acleisanthes parvifolius (Torrey) R. A. Levin,
comb. nov. Rasionym: Selinocarpus dijfusus A.
Gray var. parvifolius Torrey, Rep. U. S. Mex.
Bound. Surv., Bot., 168. 1859. Selinocarpus
parvifolius (Torrey) Standley, Contr. U.S. Natl.
Herb. 12: 388. 1909. TYPE: U.S.A. Texas:
Canons of the Rio Grande or Presidio del Nor¬
te, 1852, C. C. Parry el al. s.n. (lectotype, des¬
ignated by Fowler & Turner (1977), GH; iso-
lectotvpe, IIS).

This species is similar morphologically to 4. lan-
ceolatus, A. purpusianus, and 4. palmeri. Most no¬
tably, these four taxa share a distinctive divaricat¬
ing branching pattern within a woody shrub, and
they also have very similar flowers. However,
w hereas these other species appear confined to gyp¬
seous soils, 4. parvifolius is not so restricted.

Specimens examined. U.S.A. Texas: Brewster Co.,
Tornillo Creek, dig Bend National Park, levin 97-2
(Alt I/); Hudspeth Co., 33 mi. SE from freeway on Texas
Rd. 192, on county portion of road, between Quitman Mts.
& tlie Rio Grande, Spellenberg <£• Ward 9688 (NMC).
MEXICO. Chihuahua: 10.7 road mi. S ol Ojinaga along
Hwy. IB (Ojinaga— C. Camargo), open sandstone hill along
I hw., Hendrickson 7701 (NMC).

Acleisanthes purpusianus (Heitnerl) R. A. Levin,
comb. nov. Basionym: Selinocarpus purpusi¬
anus Heitnerl, Oesterr. Bot. Z. 63: 353. 1913.
TYPE: Mexico. Coahuila: Sierra (ltd Rey, June
1910. C. A. Purpus 4505 (holotype, US).

As discussed in the entry for 4. palmeri, these
two taxa are very similar except for a difference in

perianth color. This may likely be due to popula¬
tion-level differences, especially as there is known
variation among populations of 4. purpusianus,
some ol which have a yellow chasmogamous peri¬
anth, others a cream to white perianth (Fowler &
Turner, 1977).

Specimens examined, MEXICO. Coahuila: Km 134
nn Hwy. 30 towards Torredn, levin 98—6 (A 111/); Hwy. 30,
68 mi. N of San Pedro, on gypseous flat, about 7 mi. S of
Puente Arroyo Seco, Spellenberg. Willson A' Feather 4050
(NMC); ca. 12 (air) mi. SW ol Cuatro Cienegas (13.5 road
mi.) along Hwy. to San Pedro and 4.4 mi. W on trail,
Henrickson 12552 (NMC); Hwy. 57. 1.2 mi. S of Herman¬
ns, Spellenberg A Syvertsen 8779 (NMC).

Acleisanthes somalensis (Chiovenda) R. A. Lev¬
in, comb. nov. Basionym: Selinocarpus soma¬
lensis Chiovenda, f lora Somala 284. 1929.
TYPE: Somalia. Costa dei Migiurtini, dintorni
di Biaddo, June 1924, I\. Puccioni & G. Ste-
fanini 814 (holotype, FT not seen).

Despite its disjunct occurrence in Somalia, both
morphological and molecular evidence support its
inclusion within the genus Selinocarpus (Levin,
2000) and, thus, within a newly expanded Acleis¬
anthes. With the other species formerly placed in
Selinocarpus, this taxon shares a presence of
winged fruits and a preference for gypseous soils.
Surprisingly, A. somalensis looks remarkably simi¬
lar to 4. dijfusus, although molecular sequence data
suggest a close relationship with 4. chenopodioides.

Specimens examined. SOMALIA. Buran (Sorl): 10—
I3°N. 48-47°E, Collenette 125 (K). Calgiiduud: 4 km S
of Ceelbuur(El Bur) on road to Gal Hareeri, gypsum plain,
Timlin & Dahir 6517 (K).

Acleisanthes undulatiis (B. A. Fowler <X B. L.
Turner) R. A. Levin, comb. nov. Basionym: Se¬
linocarpus undulatus B. A. Fowler & B. L.
Turner, Phytologia 37: 194. 1977. TYPE: Mex¬
ico. Coahuila: 4 mi. W of Cuatro Cienegas,
mouth of canyon, 24—26 Aug. 1938, /. M.
Johnston 7159 (holotype, GH).

Although this species occurs in geographically
proximate locations to 4. palmeri and A. purpusi¬
anus, both morphological and molecular evidence
closely ally 4. undulatus with 4. angustifolius (Lev¬
in, 2000). These sister taxa share a short chasmo¬
gamous perianth and virgate growth form, lacking
the divaricating branching pattern of A. lanceolatus,
A. parvifolius, A. purpusianus, and A. palmeri.
Acleisanthes undulatus can be distinguished from
A. angustifolius by its sinuate leaf margins and
ochreous perianth.

Volume 12, Number 1
2002

Levin

Genera in Nyctaginaceae

63

Specimens examined. MEXICO. Coahuila: outside of
Cuatro Cienegas at Km 94 on Hwy. 30 towards Torreon,
Levin 9 tt-7 (AKIZ); 8 km W of Cuatro Cienegas on S
slopes and foothills of Sierra de la Madera, Johnston ,
Chiang & Morofka 12076 (NMC); Mpio. Parras, S side of
Sierra Parras along road toward Menchaea, ea. 14 air km
S of Parras, just N of Sierra Prieta, Nesom 7651 (NMC).

Acknowledgments. I thank Big Bend National
Park and the Jornada LTER lor permission to col¬
lect plants. R. Spellenberg for help in locating plant
populations, and the staff at AR1Z for their assis¬
tance. I'he following herbaria allowed me to borrow
specimens: GH, K, NIVIC, US. I thank I,. A. Me-
Dade for critically reading an earlier version of this
manuscript, financial support was provided by the
National Science Foundation DEB9806840 to L. A.
McDade and R. A. Raguso and by small grants
from the American Society of Plant Taxonomists,
Sigma Xi. the Department of Ecology N Evolution¬
ary Biology at the University of Arizona, and the
University of Arizona Research Training Group in
the Analysis of Biological Diversification (NSE
DIR-91 13362, BIR-9602246). This paper is part of
a dissertation submitted to the University of Ari¬
zona in partial fulfillment of the requirements for
the degree of doctor of philosophy.

Literature Cited

Chiovenda, E. 1929. Flora Somala. Sindicato Italiano Arti

Grafiehe, Rome.

F owler, B. A. & B. L. Turner. 1977. Taxonomy of Selino-
carpus and Ammocodon (Nyctaginaceae). Phytologia 37:
177-208.

Gray, A. 1853. Brief characters of some new genera and
species of Nyctaginaceae, principally collected in Texas
and New Mexico. Amer. J. Sri. Arts, ser. 2, 15: 259—
263.

Heimerl, A. von. 1913. Eine neue Art der Gattung Seli-
nocarpus. Oesterr. Bot. Z. 63: 353—356.

Hemsley, W. B. 1882. Biologia Centrali-Americana; or,
contributions to the knowledge of the fauna and flora of
Mexico and Central America, Botany, Vol. 3. R. H. Por¬
ter & Dulau, London.

Levin, R. A. 2000. Phylogenetic relationships within Nyc¬
taginaceae tribe Nyctagineae: Evidence from nuclear
and chloroplast genomes. Syst. Bot. 25: 738—750.

Smith, J. M. 1976. A taxonomic study of Acleisanthes
(Nyctaginaceae). Wrightia 5: 261—276.

Standley, P. C. 1909. The Allioniaceae of the United
States with notes on the Mexican species. Contr. U.S.
Natl. Herb. 12: 303-389.

- . 1916. Ammocodon, a new genus ol Allioniaceae,

from the southwestern United States. J. Wash. Acad.
Sci. 6: 629—631.

Thulin, M. 1993. Nyctaginaceae. Pp. 168-175 in M. Tim¬
lin (editor). Flora of Somalia, Vol. 1. Royal Botanic Gar¬
dens, Kew.

Torrey, J. 1859. Report on the United States and Mexican
Boundary Survey, Botany, Vol. 2. A. O. P. Nicholson,
Washington, D.C.

Turner. B. L. 1993. New species and combinations in Se-
linocarpus (Nyctaginaceae). Phytologia 75: 239—242.

Wooton. E. <). 1898. New plants from New Mexico. — II.
Bull. Torrey Bot. Club 25: 304-310.

Three New Taxa of Guadua (Poaceae: Bambusoideae) from

South America

Ximena Londono

Institute) Vallecaucano de Investigaciones Cientfficas, Apartado Aereo 5660, Cali, Colombia

Lynn G. Clark

Department of Botany, Iowa State University, Ames, Iowa 5001 1-1020, U.S.A

Abstract. Two new species of Guadua from
South America are described, illustrated, anti com¬
pared with putatively related species. The two new
species are Guadua macrospiculata from the west¬
ern Amazon basin in southeastern Colombia, north¬
western Brazil, and northern Peru, and G. uncinata
from southern Colombia and central-eastern Ecua¬
dor. A description of the Brazilian species G. ta-
goara is included, establishing one new subspe¬
cies, G. tagoara subsp. glaziovii, from the Atlantic
(orests ol the slate ol Bio de Janeiro. The empty
bracts at the base of the spikelet proper ol the pseu-
dospikelet in Guadua are reinterpreted as sterile
lemmas and not glumes.

Kksumkn. Dos nuevas especies de Guadua y una
subespeeie tie G. tagoara son descritas para Sur
America, incluyendo ilustraciones y datos compar-
ativos con especies afines. Las nuevas especies son
Guadua macrospiculata, que ocurre en la cuenca
occidental Amazonica. especfficamente en el sur-
oriente de Colombia, noroccidente de Brasil y norte
del Peru, y G. uncinata que habita el sur de Co¬
lombia y la region centro-oriental del Ecuador. Se
hace una revision de la especie brasilena G. ta¬
goara estableciendo una nueva subespeeie, G. ta¬
goara subsp. glaziovii, que ocurre en la Mata At-
lantica del estado de Rio de Janeiro en Brasil. Se
reinterpretan las bracteas vaefas en la base de la
llamada espiguilla de la pseudoespiguilla en Guad¬
ua conio lemas esteriles y no glumas.

Key words: Andean foothill swamps, Atlantic
forest, Bambusoideae, Guadua, igapo, Poaceae,
pseudospikelet morphology, South America, ta¬
li uarripa.

Guadua Kunth is distinguished among native
New World bamboos by its large size, triangular
culm leaves, light-colored supra- and infranodal
bands of hair, thorny branches, pseudospikelets,
and paleas with winged keels (Judziewicz et al.,
1999). Its species are broadly distributed from

Mexico to northern Argentina and Uruguay, with
the greatest diversity below 1500 m in elevation in
a variety of habitats, including lowland tropic al and
lower-montane forests, savannas, cerrado, and gal¬
lery forests. We here describe two new species, G.
macrospiculata Londono & L. G. Clark from the
western Amazon basin of southeastern Colombia,
northwestern Brazil, and northern Peru, and G. un¬
cinata Londono & L. G. Clark from southern Co¬
lombia and central-eastern Ecuador. We also pro¬
vide a detailed description of G. tagoara (Nees)
Kunth and establish a new subspecies. Including
the 2 new species, Guadua comprises 26 described
species, although a few names remain to be clari¬
fied and some species remain undescribed.

Like other woody bamboos, species of Guadua
can be distinguished readily by a suite of vegetative
and reproductive characters once complete material
is available. It is often the case in Guadua, how¬
ever, that pairs of species may be very similar veg-
etatively but distinct in their reproductive mor¬
phology (e.g., G. macrospiculata and G. glomerate
Munro) or very similar in their reproductive mor¬
phology but distinct in their vegetative morphology
(e.g.. G. paniculata Munro and G. weberbaueri Pi 1-
ger or G. macrospiculata and G. ciliata Londono &
Davidse). Thus, lack ol flowering material or col¬
lection of only flowering branches without addition¬
al structures (usually due to the large size and
thorniness of the plants) are especially problematic
for elucidating the systematics of this genus. An
additional compounding factor, which holds for all
bamboos with pseudospikelets, is that reproductive
morphology changes significantly during develop¬
ment, more dramatically than in bamboos with true
spikelets. A corollary of this is that it is difficult to
compare developmental stages among herbarium
specimens. While the developmental problem is not
easily resolved, the value and utility of Guadua
specimens can be increased greatly by the inclu¬
sion of label data on culm size, thorn morphology,
size, and distribution of supra- and infranodal

Novon 12: 64-76. 2002.

Volume 12, Number 1
2002

Londono & Clark
New Taxa of Guadua

65

bands, even if these structures are not actually col¬
lected.

Interpretation of the bamboo pseudospikelet is
still debated. As discussed in Judziewicz et al.
(1999: 39), McClure (1934) originally proposed the
term to describe the “units in iterauctant synflo-
rescences of woody bamboos that rebranch to pro¬
duce successive orders of spikelets.” Young and
Judd (1992: 757) defined the pseudospikelet as a
“shortened vegetative axis or branch that termi¬
nates in a single spike of florets.” While definitions
differ on what the pseudospikelet actually repre¬
sents, most of the interpretations agree that the
pseudospikelet includes at least a prophyll, a series
of gemmiparous bracts, and a spikelet proper, this
last considered homologous to the spikelets of other
bamboos and grasses. Because of this presumed ho¬
mology, any empty bracts al the base of the spikelet
proper usually have been regarded as glumes (Jud¬
ziewicz et al., 1999).

According to our present interpretation, a pseu¬
dospikelet of Guadua consists of a prophyll, one to
several gemmiparous bracts, none to several sterile
lemmas, one to several fertile florets, and a rudi¬
mentary floret; each coflorescence and pseudospi¬
kelet is subtended by a subtending bract. In con¬
trast to previous interpretations (e.g., Soderstrom,
1981; Young & Judd, 1992; Judziewicz et al.,
1999), we argue that the “empty” bracts distal to
the gemmiparous bracts and proximal to the fertile
florets are best regarded as sterile lemmas rather
than glumes. Usually these “empty” bracts are
comparable in size to the fertile lemmas, but most
important, they often enclose a rudimentary palea
in Guadua , evidence that they are comparable to
or derived from fertile lemmas. Subtending bracts,
gemmiparous bracts, glumes, and both sterile and
fertile lemmas are all homologous at some level, but
what are normally called glumes in bamboos and
other grasses may have arisen in different ways in
different lineages (Stapleton, 1997; Grass Phylog-
eny Working Group, 2001).

Guadua macrospiculala Londono & L. G. Clark,
sp. nov. TYPK: Colombia. Amazonas: Mpio.
Puerto Narino, lago de Tarapoto, crece en var-
zea, a orilla del lago, 16 Nov. 1990, X. Lon¬
dono & M. Kobayashi 577 (holotype, COL; iso¬
types. ISC, US), f igure I.

Bambusa lignosa, spinosa. Rhizoma sympodiale, pachy-
morphum. Culmi 8-15 m alti, 1.8-4 cm diam.; internodia
(1 1— )20— 32 cm longa, solida vel fere solida. Folia cul-
morum leviter eoriacea, decidua; vagina 1.5— 2.8-plo lon-
gior quam laminam; lamina (2— )5— 10 cm longa, persis¬
ted, erecta, triangularis. Ramificatio intravaginalis. Folia

cujusquisque complementi (5 — )8 1 5; vagina pubescens
vel glabrescens, biauriculata vel fimbriata, auriculis fal-
cato-lanceolatis, fimbrialis; pseudopetiolus 1-1.5 mm lon-
gus, plerumque reflexus; figula interior 0.25 mm longa;
lamina (2— )5— 10(— 12) cm longa, (0.8 — ) 1 — 2( — 2.8) cm lata,
ovato- vel lineari-lanceolata. Synflorescentiae ramos ter-
minantes cum 1—9 coflorescentiis. Pseudospiculae (3-)6-
1 2( — 1 7) cm longae, lineari-lanceolatae, prophyllo singuli,
bracteis gemmiferis l-3(-6), lemmata sterilia 1-2, llos-
culis 6—1 1 ( — 1 7); lemma 15—24 mm longum, ovato-lanceo-
latum, 18-25-nervatum, abaxialiter glabrum; palea sulco
pubescenti, carinis alatis 2, alis 0.25-0.5 mm latis, pub-
erulis. Lodiculae 3. Stamina 6. Ovarium fusiforme, infra
glabrum, supra hispidulum, stigmatibus tribus plumosis.

Woody, thorny bamboo. Rhizomes sympodial, pa-
chymorph, short-necked, the neck 3—5 cm long.
Culms 8—15 m tall, 1.8—4 cm diam., clambering or
seandent. the distal portion cascading down from
trees, forming clumps 2 X 2 m with ca. 90—100
culms/clump, dark purple when young turning
green later and yellow at maturity; internodes (11 —
)20— 32 cm long, cylindrical, solid or with a small
central lumen, slightly strigose, covered by white-
tinged pubescence; nodes solitary, the nodal line
horizontal, with appressed white hairs in a band 7—
12 mm wide above and 3-8 mm wide below the
nodal line; supranodal ridge manifest ca. 1 cm
above the nodal line; bud single, triangular, the
shoulders of the prophyll ciliate. Culm leaves (8—
) 1 1 — 1 8(— 26.5) cm long, (4.5—) 1 2— 1 6 cm wide at
base, slightly coriaceous, deciduous, sometimes the
basal ones rolling on the culm, triangular, the youn¬
gest green-purple then stramineous; sheaths 6—14.5
cm long, 1.5— 2.8 limes as long as the blade, adax-
ially glabrous, shiny, with a band of hairs 1 cm wide
close to the margin, abaxially pubescent, covered
with mixed coarse, rigid, hyaline or red-tinged, eas¬
ily removed hairs 1 mm long, and soft, wavy, white-
tinged, shorter hairs, ciliolate on one margin, the
summit incurved at the middle, continuous with the
blade and confluent with the inner ligule, bearing
small fimbriate auricles or not; auricles 1—3 mm
long, brown, fimbriate, the fimbriae 3 mm long, ba-
sally scabrid and erect, apically smooth and curled;
inner ligule 1-1.5 mm long, truncate, stramineous,
abaxially pubescent, incurved at the middle, the
margin ciliolate; blades (2— )5— 10 X 2.5—9 cm, per¬
sistent, triangular, erect, abaxially pubescent, cov¬
ered by soil, wavy, while appressed hairs less than
0.2 mm long, adaxially conspicuously nerved and
pubescent between nerves, marginally ciliate es¬
pecially toward the base, the apex mucronate, the
mucro 2—2.5 mm long. Branching intravaginal, typ¬
ically with one strong dominant branch, armed;
thorns (1 to) 3 per node, the central one dominant
and bigger, the other two usually slightly reflexed.
Foliage leaves (5 to) 8 to 15 per complement, ±

66

Novon

Figure I. Guadtut macrospiculata Londono & L. G. Clark. — A. Mid-culm node with branch complement of leafy
flowering axis. — 15. Ligular area of the foliage leaf showing blade base and sheath summit with long fimbriae. — C.
Culm leaf, adaxial view. — I). Detail of culm leaf inner ligule, adaxial surface. — E. Detail of culm leaf blade apex,
adaxial view. — F. Actively growing new culm showing inflated blades. — G. Thorny culm node. — H. Pseudospikelet.
— 1. Lemma, adaxial view. — J. Detail of lemma apex, adaxial view. — K. Lemma, abaxial view. — L. Detail of lemma
apex, abaxial view. — M . Palea showing the ciliolate winged keels, abaxial view. — N. Palea, abaxial/lateral view.

Volume 12, Number 1
2002

Londono & Clark
New Taxa of Guadua

67

pendent or drooping; sheath densely pubescent to
glabrescent, green when young becoming strami¬
neous, marginally ciliate, bearing fimbriae or fim¬
briate auricles at the summit; auricles 0.5-1.25 mm
long, usually present, falcate-lanceolate, dark
brown to stramineous, fimbriate; fimbriae 3—10 mm
long, ivory or brown, basally scabrid and erect, api-
callv smooth and erect or curled; inner ligule 0.25
mm long, membranous, stramineous to brown,
abaxially densely strigillose, the margin minutely
eiliolate; outer ligule 0.5—0.75 mm long, strami¬
neous, glabrous, shiny, the margin eiliolate to
smooth; pseudopetioles 1—1.5 mm long, usually re¬
flexed, adaxially hispidulous, abaxially hirsute to
glabrous and pulvinate, the pulvinus yellow, shiny,
hirsute to glabrous; blade (2— )5— 10(— 12) cm long,
(0.8-)l-2(-2.8) cm wide, 12- to 14-nerved, ovate-
or linear-lanceolate, adaxially scabrid to glabrous,
with scattered, sparsely strigose, hyaline hairs 0.3—
0.5 mm long, with 3 or 4 rows of prickle-hairs along
one edge, abaxially pilose to glabrescent, covered
with wavy hairs 0.2— 0.3 mm long, the midnerve
yellow and prominent in the middle lower portion,
the base rounded-attenuate, the margins scabrous,
the apex acuminate, with a muero 1—2 mm long.
Synflorescences terminating leafy or leafless branch¬
es, polytelic, consisting of 1 to 9 coflorescences
each with I to 3 multiflowered pseudospikelets, oc¬
casionally forming ± dense aggregations of up to
10 pseudospikelets on leafless axes, coflorescences
and pseudospikelets subtended by bracts; main
axes pubescent; subtending bracts gemmiparous or
not. varying throughout the main axes, fully devel¬
oped and similar to a foliage leaf with a small blade
or the blade a reduced apicule, the blades decid¬
uous, the sheath abaxially pubescent to glabres¬
cent, the margin eiliolate, the summit fimbriate.
Pseudospikelets (3 — )6 — 1 2( — 1 7) cm long, 3 — If — 5 )
mm wide, linear-lanceolate, usually straight, some¬
times slightly curved at the base, green with purple
spots when young, later stramineous, consisting of

1 prophyll, I to 3 (to 6) gemmiparous bracts, I or

2 sterile lemmas, 6 to 11 (to 17) fertile florets, and
a terminal rudimentary floret; prophyll 0.75—2.5 X
1.3—3 mm, abaxially puberulent between the
winged eiliolate keels; gemmiparous bracts I to 3
(to 6), 4— 5(-8) mm long, 3— 4(— 6) mm wide, 11- to
17-nerved, ovate to ovate-lanceolate, sometimes
empty, abaxially pubescent to glabrescent, adaxi¬
ally densely strigillose toward the apex with incon¬

spicuous transverse veins, the margins smooth, the
apex apiculate with a muero ca. 0.5 mm long; sterile
lemmas I or 2, 6—14 X 5—7.5 mm, 17- to 20-
nerved, rounded on the back, abaxially glabrous
and shiny, adaxially densely strigillose toward the
apex, always enclosing a rudimentary palea, the
margins smooth and purple-tinged to stramineous,
the apex acuminate with a muero 0.25—0.5 mm
long; rachilla segments (4— )7— 9 mm long, 2 mm
wide, segments between the lemmas glabrous, those
between the prophyll and the gemmiparous bracts
pubescent, usually longer between fertile florets,
not disarticulating easily at the junction with the
floret. Fertile florets 6 to 11 (to 17) with lemmas
15-24 X 7-9(— 10) mm, 18- to 25-nerved, evate-
lanceolate, rounded on the back, abaxially glabrous
with inconspicuous transverse veins, adaxially pu¬
bescent on the upper VS, the margins smooth and
purple-tinged, completely embracing the palea. the
apex acuminate with a muero 0.25—0.5 mm long;
paleas 10-16 X 2—3 mm, I I - to 13-nerved, shorter
than the lemma, 2-keeled, the sulcus 6- to 8-nerved
with inconspicuous transverse veins, pubescent, the
enfolding margins glabrous, 3-nerved, the keels
winged, the wings 0.25—0.5 mm wide, 1- or 2-
nerved, puberulent on both surfaces toward the up¬
per V2—V3, marginally eiliolate above the middle, the
cilia hyaline, the wings ending 1-1.5 mm before
the apex. Lodicules 3, 4—8 X 1—2 mm, many-
nerved, membranous, puberulent on the upper half,
eiliolate on the upper part of the margin, the apex
acute, the anterior pair a little longer and wider
than the posterior one. Stamens 6, the anthers 6—7
mm long, ochre, basally sagittate, apically apicu¬
late. Ovary 2—7 mm long, fusiform, basally thick¬
ened and glabrous, apically densely hispidulous to
glabrescent at maturity; style 1. 4—5 mm long,
densely antrorse hispidulous; stigmas 3, plumose,
ca. 4—5 mm long, hyaline. Caryopsis 7 X 1.25 mm,
fusiform, with a persistent hispidulous style base.

Etymology. The specific epithet refers to the
very long pseudospikelets, which reach up to 17
cm in length.

Distribution and habitat. Western Amazon ba¬
sin in southeastern Colombia (Amazonas), north¬
western Brazil (Amazonas), and northern Peru (Lor¬
eto). This species occurs in the lowland forests of
the Amazon region, mainly on riverbanks, in sta-

e-

— O. Lodicules. — P. An anther. — (j. Gynoecium with three plumose stigmas. (A— E and H— Q based on Londono &
Kobayashi 577 ; F, G based on Londono & Kobayashi 565.)

68

Novon

Table 1 . Morphological comparison of Guadua macrospiculata, G. glomerata, and G.

ciliata.

Character

G. macrospiculata

G. glomerata

G. ciliata

Culms

solid or hollow wifli a

solid

hollow

small lumen

Thorns per node

1-3

( 1 — )5 — 7

absent

Internodes

slightly strigose

strigose

usually glabrous

Culm leaf sheath, abaxial indument

pubescent

glabrous

glabrescent

Culm leal sheath girdle

absent

absent

present

Foliage leal sheath, abaxial indu-

densely pubescent to

glabrous

hirtel Ions to glabrous

ment

glabrescent

Foliage leal blade position

usually reflexed

erect

erect

Pseudopetiole, abaxial indument

hirsute to glabrous

subglabrous

glabrous

Foliage leaf blade size (cm), abaxi-

2-12 X 0.8-2.8, pi-

7-22 X 1 .3-4.5, gla-

9-27.5 X 1 .2-4.3, glabrous

al indument

lose to glabrescent

brous

Pseudospikelet size (cm)

3-17 X 0.3-0.5

4-7 X 0.35—0.6

3-1 1 x 0.3-0.5

Lemma margins

glabrous

glabrous

ciliate

Palea keel wing width (mm), indu-

0.25—0.5, pubenilent

ea. 1 , glabrous to gla-

0.7— 0.9, pubenilent

merit

brescent

lionary flooded forests called igapo in Brazil or ta-
huampa in IVrn.

Common rmnu\ N um-chi in Ticuna dialect
(Brazil, Colombia), and maronilla (Peru).

Uses. The Ticuna community in Colombia uses
the culms of G. macrospiculata to make frames for
drying and painting bark fabric, as planting sticks
for yuca ( Manihot esculenta Crantz), and to make
arrows for bow hunting; the thorns are used to lance
boils.

Phenology. The flowering cycle of G. macros¬
piculata is a very short interval for a woody bam¬
boo. Observations of the flowering cycle in a cul¬
tivated stand in Quindio, Colombia, from 1990 to
2000 were made annually by the first author, who
noted flowering approximately every two years. All
the culms in a single clump flowered without in¬
terruption during the flowering cycle and remained
green, and after flowering the culms did not die.
The flowering cycle apparently does not respond to
external weather conditions such as the alternation
of wet and dry seasons. Flowering dates as deter¬
mined from herbarium specimens from a single lo¬
cality are consistent with a short flowering cycle but
arc not definitive.

Guadua macrospiculata is most similar to G.
glomerata and G. ciliata. The three species share
a climbing habit, relatively slender culms, and tri¬
angular culm leaves. They also all occur in t he Am¬
azon basin, along riverbanks that are flooded sea¬
sonally, with the bases of the plants submerged in
water. The three species are compared and con¬
trasted in Table I .

Guadua macrospiculata differs from G. glomer¬
ata by its denser and bigger clumps (90 to 100

culms/clump, 2 X 2 m vs. 4 to 13 culms/clump, 1
X 0.5 m); the delicate and mostly uniform size of
the foliage leaf blade (vs. conspicuous size varia¬
tion); the glabrous enfolding margins of the palea
(vs. pubescent); and the narrower palea keel wings,
ending 1—1.5 mm before the apex with the margins
ciliolate from the middle toward the apex (vs. end¬
ing at the apex with a tuft of hairs at the tip, the
margins smooth in the lower and middle portions,
but ciliolate at the apex). These two species have
in common the inner ligule of the culm leaf curled
in the middle, lowermost basal culm leaf sheaths
that rot on the culm, green pseudospikelets with
purple spots when young, a glabrous, many-nerved
lemma with smooth margins, and a pubescent palea
sulcus.

Guadua macrospiculata is distinguished from G.
ciliata by having dark purple culms when young (vs.
green when young), foliage leaf blades usually ovate-
lanceolate and (2-)5-l0 X 2.5-9 cm |vs. usually lin¬
ear-lanceolate and (9-) 12-27.5 X (1.2— )2- 3.5( — 4.3)
cm], the main axes of synflorescence pubescent (vs.
glabrescent), pseudospikelets green with purple spots
when young (vs. olive green when young), and lemmas
with smooth margins (vs. marginally ciliate with pur¬
ple-tinged and hyaline hairs). These two species ex¬
hibit an inflated culm leaf blade (Fig. IF, as seen
in side view) when the new shoot is growing ac¬
tively, but this condition disappears later as the
culm leaf dries out. Both species also have a pul-
vinate loliage leaf pseudopetiole, the palea smaller
than the lemma, and usually I to 3 pseudospikelets
per coflorescence.

The new culms of G. macrospiculata are hosts to
butterflies of the family Nymphalidae, subfamily

Volume 12, Number 1
2002

Londono & Clark
New Taxa of Guadua

69

Satyrinae, tribe Pronophilini (D. Harvey, pers.
comm.). The adult females deposit their eggs at the
apex of the new culms. Several other bamboos of
the genera Otatea (McClure & E. W. Smith) C. Cal¬
deron & Soderstrom, Chusquea Kunth, Guadua ,
and Merostachys Sprengel are also hosts to a wide
variety of insects (Judziewicz et al., 1999).

Paratypes. BRAZIL. Amazonas: Rio Javari above
mouth of Rio Taquari, 29 Oct. 1976 (fl). Prance et al.
24202 (US); Igarape Umarizal, Esperantja, 8 Dec. 1945
(fl), Murga & Black 842 (US). COLOMBIA. Amazonas:
Mun. Leticia, Parque Nacional Natural de Amacayacu-
INDERENA, a orilla de la Q. Bacaba, 100 m, 8 Nov. 1990
(fl), londono & Kobayashi 536 (COL, ISC, TULV, US);
Parque Nacional Natural Amacayacu-INDERENA, a or¬
illa de Q. Matamata, 100 m, 8 Nov. 1990 (fl), londono &
Kobayashi 537 (COL, ISC. TULV, US); a 30 m de distaneia
del INDERENA, sobre trocha Amacayacu-Matamata, eer-
ca de Q. Matamata, 1 1 Nov. 1990 (fl), londono A Kobay¬
ashi 563 (COL, ISC. TULV, US); San Martin de Amacay-
acu, 100 m. 12 Nov. 1990 (fl), londono & Kobayashi 569
(COL. ISC, TULV, US). PERU. Loreto: Maynas, Iquitos,
rio Nanay, Cocha de la Marina, 90 m. 2 Jan. 1983 (II).
McDaniel & Rimachi 26601 (AMAZ, NY, US); Pto. Al-
mendras, rio Nanay, 20 May 1981 (fl), Ruiz 168 (AMAZ);
Bio Nanay, Q. Moropon above Bellavista, 7 Jan. 1976 (fl),
McDaniel & Rimachi 20381 (AMAZ, NY); rio Nanay,
slightly below Bellavista on left margin, 90 m, 7 Oct.
1980, Rimachi 5344 (USM); Punchana, Rio Nanay, trocha
de la Cocha a Manga Posa, frente al puerto de Picuruyacu,
90 m, 20 July 1994 (fl), Rimachi 1 1035 (US); Rio Nanay,
Albergue Isabel Loro Park, 3 Aug. 1991 (fl), Londono 675
(TULV, US); Tahuampa near rio Amazonas between Pun¬
chana y Sta. Clara de Nanay, outskirts of Iquitos, 120 m,
3 Feb. 1977 (fl). Gentry et al. 21614 (MO, TULV, US);
Alto rio Nanay, between Santa Maria de Nanay y Diamante
Azul, 140 m, 25 Mar. 1979 (fl). Gentry et al. 26213 (MO,
TULV, US); Rio Itaya, below San Juan de Muniches, 120
m, 19 Mar. 1977 (fl). Gentry et al. 18396 (MO, TULV, US);
Rio Itaya from Iquitos to San Juan de Muniches, 100 m,
9 Mar. 1973 (11), McDaniel & Rimachi 16927 (AMAZ,
MO); Rio Itaya, Moena cano, 4 km de Iquitos, 1 Apr. 1976
(fl). Revilla 470 (MO. TULV, US); Yanomono explorarna
tourist camp, rio Amazonas, above mouth of rio Napo, 120
m, 28 Dec. 1982 (fl). Gentry & Emmons 38746 (USM);
Petropolis, rio Yavaris, 12 Aug. 1976. Revilla 1090 (MO).
Requena: Cano Iricahua, abajo de Jenaro Herrera, mar-
gen izquierda rio Ucayali, 26 Nov. 1982 (11), Encarnacidn
25086 (AMAZ, NY, US); Rio Tapiche, tributario del Uca¬
yali, ca. I hr. above Requena, 8 Dec. 1977 (fl). Gentry et
al. 21236 (MO, TULV, US, USM).

Guadua uncinata Londono & L. G. Clark, sp. nov.
TYPE: Ecuador. Tungurahua: found wild be¬
tween San Francisco and Machay, along the
road in the valley of the Pastaza River, 3500
ft., 13 July 1945 (fl), F A. McClure 21364 (ho-
lotype, US; isotypes, COL, ISC, MO, NY). Fig¬
ure 2.

Bambusa lignosa, spinosa. Rhizoma sympodiale, pachy-
morphum. Culmi 3—8 m alti, 2— 4(— 6) cm diam.; internodia
20-40 cm longa, ad basim et apicem culmorum solida

aliter cava. Folia culmorum leviter coriacea, decidua; va¬
gina 2.8-7.5-plo longior quam laminam; lamina ( 1 .7 — )3—
5(-8.5) cm longa, persistens, ereeta, triangularis. Ramifi-
catio intravaginalis; ramus dominans unus ramis
minoribus 1-3, rami conspicue spinosi; spinae uncinalae.
Folia cujusquisque complementi 7 — 9( — 1 6); vagina glabra
vel pubescens, biauriculata, fimbriata, auriculis (1— )2-4
mm longis; lamina (6.5—) 1 0— 1 5(— 1 8) cm longa, (1.4— )2-
3.5(— 4.2) cm lata, ovato- vel lineari-lanceolata; pseudo-
petiolus 3-6 mm longus; ligula interior 0.2-0.5(-0.8) mm
longa. Synflorescentiae ramos terminantes cum I— 5(— 7)
coflorescentiis. Pseudospiculae 3— 6(— 8) cm longae, li-
neari-lanceolatae, prophyllo singuli, bracteis gemmiferis
1— 4(— 6), lemmata sterilia 1— 2(— 3), flosculis (3— )4— 5; lem¬
ma 10—15 mm longum, ovato-lanceolatum, 12-25-nerva-
tum, supra glabrum vel puberulum; palea sulco pubes-
centi, carinis conspicue alatis 2, alis 0.5—1 mm latis,
glabris. Lodiculae 3. Stamina 6. Ovarium fusiforme, infra
glabrum, supra hispidulum, stigmatibus tribus plumosis.

Woody, thorny bamboo. Rhizomes pachymorph.
Culms 3-8 in tall. 2-4(-6) cm diam., erect at the
base, immediately arching to broadly arching api-
cally and climbing on trees, white-green to green
when young becoming stramineous; internodes 20—
40 cm long, cylindrical, the basal and upper ones
solid, the middle ones hollow with a small lumen
ca. 5 mm diam., densely pubescent when young,
covered by very oppressed strigose, antrorse, hya¬
line or brown hairs, glabrous at maturity; nodes sol¬
itary, the nodal line horizontal, with oppressed,
white, soft, short, hairs in a band 5-10 nun wide
above and 8-12 mm wide below die nodal line;
supranodal ridge manifest; bud single, triangular,
the shoulders of the prophyll ciliale. Culm leaves
(14.5-)19-26(-35.5) cm long, 11-14 cm wide at
the base, slightly coriaceous, deciduous, triangular,
green-stramineous with yellow spots when young
becoming stramineous; sheaths 13—28 cm long,
2.8— 7.5 times as long as the blade, abaxially pu¬
bescent, covered by (1) strigose, antrorse, transpar¬
ent or brown hairs 0.8— 1.5 mm long that rub off
easily, (2) shorter, strigose, mostly retrorse, hyaline
or brown hairs, and (3) appressed, white-tinged,
wavy, tiny hairs, the margins papery, ciliate or
smooth, the cilia transparent to brown, up to 3 mm
long, rubbing off easily, bearing fimbriate auricles
and fimbriae at the summit; auricles 4—6 X 1—1.2
mm, falcate, puberulous, fimbriate; fimbriae 3—7
mm long, ivory, straight to curled, basally scabrid,
apically smooth; inner ligule 0.5—1 mm long, trun¬
cate, straight, slightly curled, or inclined, dark
brown to stramineous, abaxially pubescent, some¬
times not extending completely from margin to mar¬
gin, the margin ciliolate; blades (1 .7— )3— 5(— 8.5) cm
long, 1-6 cm wide, triangular, erect, abaxially pu¬
bescent. with white, wavy, appressed, tiny hairs,
adaxially densely pubescent between the nerves,
the margins basally fimbriate, otherwise ciliate to

70

Novon

Mf;iire 2. Guadua uncinata Londofio & L. G. Clark. — A. Flowering branch. — 15. Culm leaf, adaxial view. — C. Mid-
culm node with branch complement showing single dominant branch and recurved thorns. — I). Ligular area of foliage
leal, showing pseudopetiole, sheath summit, and auricles. — E. Pseudospikelet. — F. Prophyll, abaxial view. — G.

Gemmiparous bract, adaxial view. — II. Lemma, adaxial (left) and abaxial (right) views. — I. Palea, abaxial view. J.

Apex ol palea showing winged keels, abaxial view. — K. l.odicules. — L. Young androeeium and gynoecium surrounded

Volume 12, Number 1
2002

Londono & Clark
New Taxa of Guadua

71

smooth, tlie cilia easily removed, the apex mucro-
nate, the mucro 1-1.5 mm long. Branching intra-
vaginal. the branches manifestly thorny, consisting
of one main branch and 1 to 3 secondary branches;
thorns (1 to) 3 to 5 per node, recurved. Foliage
leaves 7 to 9 (to 16) per complement. ± horizontal-
drooping; sheatli abaxially usually glabrous or
sparsely covered by appressed, strigose, antrorse,
transparent or brown hairs up to I mm long on the
lower portion, bearing fimbriate auricles and fim¬
briae at the summit, the overlapping margin con¬
spicuously ciliate, the underlapping margin cilio-
late to smooth; auricles (1— )2— 4 X 0.25—0.5 mm,
ligulate, puberulous, dark purple to stramineous,
fimbriate; fimbriae (3— )6 — 1 1 mm long, ivory, basally
seabrid, straight, apieally smooth, wavy to curled;
inner ligule 0.2-0.5(-0.8) nun long, truncate, mem¬
branous, abaxially puberulous, the margin eiliolate
to smooth; outer ligule 0. 2-0.7 mm long, glabrous,
shiny, stramineous, the margin eiliolate to smooth;
pseudopetioles 3—6 mm long, adaxially glabrous ex¬
cel)! puberulous on the lower portion, abaxially gla¬
brous. pulvinate, the pulvinus yellow, shiny, puber¬
ulous or glabrous; blades (6.5— ) 10—1 5(— 18) X (1.4—
)2— 3.5( — 4.2) cm, 15- to 17- (to 20)-nerved, juvenile
leaves wider than adult leaves, ovate-lanceolate or
linear-lanceolate, adaxially glabrous except with a
few sparsely strigose. transparent hairs up to 1 mm
long, with 3 or 4 raised seabrid submarginal nerves
on one side, abaxially sparsely hairy, covered by
strigose. straight, antrorse, hyaline or white-tinged
hairs up to 1 mm long, tessellate. lighter green than
adaxially, the midnerve yellow and conspicuous in
the middle lower portion, the margins scabrous, the
apex acuminate with a mucro 1 .5—3 mm long. Syn-
fiorescences terminating lealy and leafless branches
of all orders, polytelic, consisting ol 1 to 5 (to 7)
coflorescences each with 1 to 3 (to 15) multiflow-
ered pseudospikelets, coflorescences and pseudos-
pikelets subtended by bracts, the main axes his-
pidulous or glabrous; subtending bracts varying in
size and shape throughout the main axes. fully de¬
veloped and similar to a small foliage leaf with a
small blade to triangular and apiculate, the blade
when present deciduous, the sheath abaxially gla¬
brous, eiliolate to glabrous along one margin and
bearing fimbriae or not at the summit. Pseudospi¬
kelets 3 — 6(— 8) X 0.3-0.5(-0.6) cm, linear-lanceo¬
late, straight to slightly curved, green with purple
spots when young becoming brown to stramineous,

consisting of 1 prophvll. I to 4 (to 6) gemmiparous
bracts, 1 to 2 (to 3) sterile lemmas, (3 to) 4 to 5
fertile florets, and a terminal rudimentary floret;
prophyll 2-4 X 1.5—4 mm. abaxially puberulous
between the winged keels, the wings eiliolate, pu¬
bescent on both sides, occasionally gemmiparous;
gemmiparous bracts 1 to 4 (to 6), 3-8 X 3—4 mm,
10- to 15-nerved. ovate to ovate-lanceolate, some¬
times with a small blade, deciduous or not, abaxi¬
ally puberulous or glabrous, adaxially tessellate,
densely pubescent toward the apex, the margins
minutely eiliolate or smooth, the apex shortly mu-
cronate, the mucro 0.2— ().5(— 0.8) mm long; sterile
lemmas 1 or 2 (to 3), 7— 9(— I 1) X 5-6 mm, 15- to
16-nerved, ovate-lanceolate, lighter in color than
the lemmas, abaxially puberulous or glabrous,
adaxially pubescent, densely so toward the apex,
the margins smooth, enclosing a rudimentary palea,
mucronate, the mucro 0.5-0.8(-1) mm long; rach-
illa segments 4—6(-9) mm long, glabrous and shiny,
apieally with a rim of puberulous hairs, straight to
slightly zigzag along the pseudospikelet. disarticu¬
lating below the attachment of each lemma and fall¬
ing attached to the floret. Fertile florets (3 to) 4 to
5 with winged palea keels exceeding the lemma
margins, basally thickened; lemma 10-15 X 6-9
mm, 12- to 25-nerved, ovate-lanceolate, abaxially
puberulous or glabrous, adaxially densely pubes¬
cent at the tip otherwise puberulous or glabrous,
the margins smooth and papery, the apex shortly
mucronate, the mucro (0.2-)0.5-0.7 mm long; pa¬
lea 9-14 X 2—2.5 mm, 9- to 13-nerved, 2-keeled,
the sulcus 2 mm wide, 4- to 7-nerved, densely pu¬
bescent, tessellate. the enfolding margins 3- to 4-
nerved, glabrous, tessellate, the keels winged, the
wings 0.5— 0.8(— 1) mm wide, 2-nerved, shiny, gla¬
brous except puberulous at the junction with the
keels, the margins minutely eiliolate for the upper
y3. Lodicules 3, 4-6 X 1-3.5 mm, many-nerved,
concave, thicker below, membranous for the upper
bearing dark brown prickle hairs on the upper
membranous portion, the margins apieally eiliolate,
acuminate, the posterior one symmetrical, narrower
and slightly longer than the asymmetrical anterior
pair. Stamens 6. the anthers 4 — 6(— 7) mm long,
when young white-yellow becoming creamy yellow
with purple spots and oehraceous to stramineous,
basally sagittate, apieally apiculate. Ovary ( 1 .5— )2-
4 mm long, fusiform, basally yellow or purple and
glabrous, apieally densely hispidulous; style 1.5—

by lodicules. — M. Gynoecium with three plumose stigmas. (A. D— M based on McClure 21364: B, C based on Londono
& Quintero 109.)

72

Novon

3.5 nun long, densely antrorse hispidulous; stigmas
3, plumose, ca. 3—6 nun long, with the main nerve
yellow to purple. Caryopsis not seen.

Etymology. The specific epithet refers to the
strongly recurved thorns of the branches, like cat’s
claws, which help tin* plant to climb on trees.

Distribution and habitat. This species occurs
in southern Colombia and central-eastern Ecuador,
on the eastern side of the Andes (Eastern Cordillera
in Colombia), in the foothills of the mountains, at
elevations between 280 and 1200(-1500) m. It
grows in swamps (locally called chucuas ) associated
with Mauritia flexuosa E. f., Heliconia L., and Ficus
L., where it forms very dense and hard to penetrate
clumps.

Common name. Cachupenda.

Uses. In Colombia this species is not used
probably because of its poor wood ( ]iial ity. In Ec¬
uador. however, according to McClure 21364, the
species has many local uses.

Phenology. The flowering cycle of C. uncinata
apparently is 10 years long. In late 1987, a plantlet
from a flowering clump was taken from its natural
habitat in Putumayo, Colombia, and planted at the
Bamboo Germplasm bank of the Juan Maria Ces-
pedes Botanical Garden, in Tulua, Valle del Cauca,
Colombia (Londono & Quintero 203). During the
first four years (1988-1992), the plantlet did not
increase very much in size, reaching 0.6 cm in di¬
ameter and 2.5 m in height, but it continued to
flower. Between 1992 and 1996 the plantlet in¬
creased in size, reaching 2.5 cm in diameter and 8
m in height, but it ceased flowering. In 1997, how¬
ever, it started to flower again. Based on the 13
years of observations made by the first author, it
can be inferred that C. uncinata has a flowering
cycle of 10 years, it remains green and does not
die after flowering, and it can continue to flower for
approximately 5 years.

Guadua uncinata is most similar to G. angusti-
folia. The two have foliage leaf blades of similar
size and shape, with sparse hairs on both surfaces;
ciliolate margins on the foliage leaf sheath with the
outer ligule abaxially pubescent; straight or slightly
curved pseudospikelets; mature florets with the
winged keels of the palea exceeding the margins of
the lemma; lemmas with smooth, papery margins;
paleas with a hairy sulcus and glabrous enfolded
margins; and a basally glabrous, apically hispidu¬
lous, fusiform ovary.

Guadua uncinata is distinguished from G. an-
gustifolia by having culms erect at the base then
broadly nodding above and climbing apically (vs.
erect); much shorter and thinner culms [3—8 m X

2— 4(— 6) cm vs. 10—30 m X 6—22 cmj; solid basal
and apical internodes (vs. hollow); smaller culm
leaves (14.5—35.5 X I 1—14 cm vs. 30—60 X 20—
50 cm) that are much less pubescent abaxially,
lighter in color, and bear fimbriate auricles or fim¬
briae at the summit of the culm leaf sheath (vs.
absence of auricles); fimbriate auricles at the sum¬
mit of the foliage leaf sheath (vs. auricles absent);
more congested coflorescences at maturity | 1 to 3
(to 15) pseudospikelets/cofloreseence vs. I to 4];
and the winged keels of the palea surpassing the
apex of the palea (vs. ending at the apex). Addi¬
tionally, although the strongly recurved thorns of G.
uncinata are found in other climbing species, this
feature contrasts with the longer, straighter thorns
of G. angustifolia.

The size difference between G. uncinata and G.
angustifolia persists even when both species are
grown under the same conditions. As described un¬
der Phenology, plantlets of G. uncinata grown in
cultivation reached to 2.5 cm in diameter and to 8
m in height after 13 years. Plantlets of G. angus¬
tifolia (londono & Quintero 23.5) planted at the
same time (Mar. 1987), in the same soil and weath¬
er conditions, and coming from the same region
(Putumayo), developed culms 12 cm in diameter
and 18 m tall over the same period of time.

Paratypes. COLOMBIA. Putumayo: Puerto Caicedo,

1 km de Pto. Caicedo en la via a Garzon, sohre la Q. El
Achote, 340 m. 5 Mar. 198i (fl). londono A’ Quintero 203
(COL. TULV, US). Caqueta: vfa Florencia— Bel6n de los
Andaqufes, desvi&ndose por la via Morelia-San Jorge, a

2 km de Morelia, tinea San Francisco, 280 m, 8 Feb.
1987, londono & Quintero 109 (COL, TULV, US). EC¬
UADOR. Tiingiiraliu a: on tile trail up to Hacienda La
Gloria from “Rio Negro,” 5000 ft.. 12 July 1945, McClure
21361 (US); found wild along the trail leading from the
"Bio Negro” bridge across the Pastaza River, 4000 ft.. 12
July 1945 (fl), McClure 21362 (COL, ISC. MO, l)S); valley
of the Pastaza River, along the road from Rio Negro to the
Hacienda La Gloria. 4000 ft., 12 July 1945 (11). McClure
21363 (NY, MO. US).

Giiuduu tagoara (Nccs) kiuitli. Enum. PI. I: 434.
1833. Bamhusa tagoara Nccs, Fl. Bras. Enum.
PI. 2: 532. 1829. Arundarbor tagoara (Nees)
kimtze, Revis. Gen. PI. 2: 761. 1891. TYPE.
Brazil. "Habitat in sylvis altitudine: 1800 ped
in adscensu montis, Serra do mar diet i. eun-
tibus versus Guarantingueta, provinciae S.
Pauli,” Marlins s.n. (holotype, BR not seen).

Bamhusa barbata Trinius, Mem. Acad. Imp. Sci. Saint-
Petersbourg, Ser. 0, Sci. Math., Seconde Pt. Sci. Nat.
3. I ; 62 i . 1835. Nastus barbatus (Trinius) Ruprecht,
Bambuseae 42, I. 17. 1939 | preprint |. TYPE. Brazil.
“Hah. in sylvis provinc. Minarum Bras.,” /,. Riedel
520 (holotype, LE not seen; isotypes, G, k. P not
seen, US-fragment).

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Londono & Clark
New Taxa of Guadua

73

Bambusa distorta Nees, Linnaea 9: 470. 1834. Arundarbor
distorta (Nees) kuntze, Bevis. Gen. PI. 2: 761. 1891.
Guadua distorta (Nees) Ruprecht, Bambuseae 41, 1.
16, f. 59. 1939 [preprint]. TYPE. Brazil. S.d., V. Bel¬
low s.n. (holotype, B not seen; isotype, US [flowering
piece is Guadua, sterile pieces are Poaceae, not
Guadua]).

Bambusa spinosissima Hackel, Oesterr. Bot. Z. 53: 197.
1903. Guadua spinosissima (Hackel) E.-G. Camus,
Ees Bambusees 1: 112. 1913. TYPE. Brazil. “In
provincia Santa Catliarina, prope Blumenau,” July
1888, E. Ule 878 (holotype, W; isotvpe, US-frag-
ment).

Rhizomes sympodial, pachymorph, 30—40 cm
long. Culms 10— 15(-20) m tall, 5—10 cm diam.,
erect at the base, the apical portion arching after
developing foliage leaves, with the secondary
branches very elongate, leaning on or pendent from
trees, glaucous when young, becoming green mot¬
tled with light yellow-green dashes at maturity; in¬
ternodes (16— )20— 60(— 80) cm long, cylindrical, hol¬
low, some filled with water, walls 5—10 mm thick,
the surface roughened, striate and slightly tuber-
culate, covered by short, brown, antrorse hairs be¬
low the nodes when young then becoming glabrous;
nodes solitary, the nodal line horizontal, with a
supranodal band (0.8 — ) 1.1 — 1 ,5( — 1 .9) cm wide of
dense, white, antrorse, appressed hairs, and an in-
franodal band 0.7— 1(— 1.5) cm wide of brown-red to
white retrorse, appressed hairs; supranodal ridge
pronounced; hud single, positioned ca. 2 mm above
the nodal line, triangular, plano-convex, the shoul¬
ders of the prophyll glabrous. Culm leaves slightly
coriaceous, deciduous, pushed away by the devel¬
oping axillary branches, but those at the lowermost
nodes strongly attached to the culm, purple-green
and maculate with yellow when young becoming
brown to stramineous, usually lacking auricles and
fimbriae; sheaths 17—38 X 10—44 cm, 5—12 times
as long as the blade, abaxially pubescent with a
mix of (1) coarse, rigid, irritating, brown-red hairs
to 2.5 mm long, (2) prickly, retrorse, brown-red
hairs less than 1 mm long, and (3) soft, wavy hy¬
aline hairs, the margins entire; inner ligule 1—2 mm
long, truncate, densely ciliolate, not extending to
the culm leaf margins; blades (2-)3-5.5(-7) cm
long, 9— 10(— 13) cm wide, very broadly triangular,
erect, persistent, abaxially less pubescent than the
sheath but densely covered with soft, appressed,
hyaline hairs and some scattered coarse, shorter,
rigid, antrorse, brown-red hairs, adaxially conspic¬
uously nerved anti densely strigose between the
nerves, the margins entire, the apex strongly mu-
cronate usually split in two parts, the mucro up to
2 mm long. Branching intravaginal, branches soli¬
tary and armed, usually not developing at the lower

nodes, the middle and upper nodes with a dominant
central branch held at a 45°— 50° angle, with age
developing 1 to 4 secondary branches from the bas¬
al nodes of the central branch, the secondary
branches extending outward, clambering over trees
to 2-6 m long; thorns usually on the lateral branch¬
es, 1 to 5 per node, absent from the lowermost
nodes of the lateral branches on the main culm.
Foliage leaves 5 to 8 (to 10) per complement;
sheaths abaxially densely pubescent to glabrous,
usually with a patch ol hyaline hairs on one side
of the midnerve near the apex, green when young,
becoming stramineous, tessellate venation through
the margins, the margins eiliate, the summit fim¬
briate, auricles usually absent; auricles when pre¬
sent 1—5 mm long, dark brown, marginally fimbri¬
ate; fimbriae 7—15 mm long, ivory, curled distally;
inner ligule 0.5—2 mm long, pubescent, the margin
finely ciliolate; outer ligule 0.5-1 mm long, usually
glabrous and shiny, stramineous, the margin mi¬
nutely ciliolate when young, becoming smooth;
pseudopetioles 4— 10(— 15) mm long, pulvinate, adax¬
ially hispidulous to glabrous, abaxially glabrous;
blades (9 — ) 1 5 — 25( — 4 1 ) X (1.1— )2— 5(-6) cm, L:W =
4-9:1, 15- to 29-nerved, linear-lanceolate to broad¬
ly ovate-lanceolate, adaxially glabrous, abaxially
glabrous and strongly tessellate, the midnerve and
primary nerves prominent and yellow, one margin
scabrous, the other seabrid, the apex acuminate,
ending in a mucro. Synflorescences terminating leal-
less or leafy branches of all orders, polytelic, con¬
sisting of 7 to 11 (to 13) coflorescences, each with
1 to 6 (to 9) multiflowered pseudospikelets, coflo¬
rescences and pseudospikelets subtended by
bracts, the main axes glabrous; subtending bracts
7—13 X 4—6 mm, triangular, deciduous, mucronate,
generally gemmiparous, abaxially pubescent to gla¬
brous, adaxially pubescent toward the apex. Pseu¬
dospikelets (0.5— )1— 3(— 4) X 2— 4(— 6) mm, linear-
lanceolate, glabrescent. yellow-green mottled with
purple to stramineous, consisting of 1 prophyll, 2
to 3 (to 4) usually gemmiparous bracts, 1 (or/to 2)
sterile lemma, 1 to 4 (to 6) fertile florets, and a
terminal rudimentary floret; prophyll 1.5—2 X 3^4
mm, abaxially pubescent, usually gemmiparous, the
keels eiliate; gemmiparous bracts 2 or 3 (to 4), 2—
4 X 1.5—4 mm, 4- to 8- (to 10)-nerved, broadly
ovate, abaxially puberulent to glabrous with the
midnerve strongly marked, adaxially shortly pubes¬
cent throughout the upper % with antrorse, ap¬
pressed hyaline hairs, the apex shortly mucronate;
sterile lemmas 1 (or 2), 4.5— 6(— 7) X 4—6 mm, 10-
to 12-nerved, broadly ovate to ovate, abaxially pu¬
berulent to glabrous, adaxially pubescent for the
upper %, usually enclosing a rudimentary palea.

74

Novon

marginally ciliate especially on the upper Vs. the
cilia wavy and deciduous, the apex mucronate;
rachilla segments less than 1 mm long between pro-
phyll and glumes then elongated after the first floret
up to 5 mm, hispid, disarticulating just below the
juncture with the lemma. Fertile florets 1 to 4 (to
6) with the winged pa lea keels exceeding the lem¬
ma margins; lemmas (6 — )7— 1 l ( — 14) X (4— )5— 7(— 8)
mm, (9- to) I 1- to 13- (to l8)-nerved, ovate, round¬
ed on the back, when mature not embracing the
palea completely, ahaxially glabrescent, usually
glabrous with the nerves strongly manifest, adaxi-
ally pubescent for the upper % with antrorse, ap-
pressed, hyaline hairs, the margins ciliate, bearing
a tuft of hyaline hairs distally helow the mucro, the
cilia wavy, soft, and deciduous, the mucro 0.5—1
mm long, the apex mucronate; / miens 7-11 (—13) X
4 — i5 mm, 9- to 15-nerved, 2-keeled, ahaxially and
adaxially puberulent, the apex acute with a tuft of
hyaline hairs, the sulcus 5- to 9-nerved, puberu¬
lent. the enfolding margins 2- to 4-nerved, glabrous
and shiny, the keels winged, the wings 0.8— 1.2 mm
wide, wider apically than basally, puberulent on
both surfaces, with very short, appressed, hyaline
hairs, 1- to 2-nerved, eiliolate on the upper Vs. Lod-
icules 3, 2.5—4 X 1-3 mm, 12- to 15-nerved ba¬
sally, hyaline to yellow-brown, acute, the upper half
covered with brown-red to hyaline prickles, cilio-
late on the upper margin, thickened and darker he¬
low, thinner and membranous above, the anterior
pair slightly asymmetrical, the posterior one sym¬
metrical. Stamens 6, the anthers 5—6.5 mm long,
yellow-brown, basally sagittate, apically emargin-
ate. Ovary 3—6 mm long, fusiform, brown-red, an¬
trorse hispidulous; style 1, hispidulous; stigmas 3,
plumose. Caryopsis not seen.

Distribution. Atlantic and montane forests of
the eastern coast of Brazil, from Bahia to Santa Ca¬
tarina; 50—1200 m.

Uses. Split culms are used in the construction
of mud huts and shelters, water pipes, sheathing of
cart sides, and as a fire starter.

Common names. Taquaraqu; taquara grossa
(Tu pi -guarani').

Phenology. The flowering cycle of this species
has not been studied or reported. Based on her¬
barium specimens, however, it is probable that
flowering occurs every two years. This species does
not die after flowering, and only a few' culms of the
clump flower at the same time.

Guadua tagoara is characterized by apically
climbing culms, glaucous new culms (Judziewicz et
ah, 1999: fig. 153), very broadly triangular culm
leaf blades often with a split apex (Judziewicz et

ah. 1999: fig. 7E, F), and usually a patch of hyaline
hairs toward the apex of the foliage leaf sheath. It
most closely resembles G. weberbaueri and G. sar-
cocarpa Londono & P. Peterson, which are restrict¬
ed to the Amazon basin and eastern slope of the
Andes. All three species share basally erect culms
that are apically arching and branching, reaching
over trees for support; a prominent supranodal
ridge; adaxially appressed pubescence on lemmas
and bracts; the pubescent sulcus of the palea;
prickles on the upper portion of the lodieules; and
antrorse-hispidulous ovary and styles.

Guadua tagoara is distinguished from G. weber¬
baueri and G. sareoearpa by its culm leaves with
the margins continuous at the junction of the sheath
and blade (vs. culm leaves with the margins slightly
discontinuous at the junction of the sheath and
blade); and the very broadly triangular culm leaf
blade with the apex usually split in two (or more)
parts (vs. a triangular culm leaf blade with a strong
mucro at the apex). In addition, G. tagoara differs
from G. weberbaueri by the ahaxially glabrous and
conspicuously tessellate foliage leaf blades (vs.
ahaxially pilose and inconspicuously tessellate
blades).

Considerable morphological variation exists
among and within the populations of G. tagoara,
perhaps related to the apparently short flowering
cycle of this species. A study to evaluate the ge¬
netic variability of G. tagoara would he worthwhile,
especially if these data could he related to flowering
specimens and morphological variation. All vari¬
ants show the distinctive culm leaf and other fea¬
tures mentioned above as characteristic of G. ta¬
goara, and thus we have chosen to recognize a
single species. Within the range of variation of G.
tagoara, however, some specimens representing
perhaps three or four populations from Rio de Ja¬
neiro show a distinctive correlation among extremes
of variation in coflorescence congestion, pseudos-
pikelet size, and the occurrence of fimbriate auri¬
cles on the foliage leaf sheaths. We recognize this
set of populations as G. tagoara subsp. glaziovii.
This subspecies is known from a limited number of
populations from a restricted geographic range
nested within the distribution of the autonymic G.
tagoara subsp. tagoara; it is unknown whether
populations of the two subspecies are strictly sym-
patric.

Key to the Si hsci- cies ok Guadua tagoara

la. Pseudospikelets 9 to 15 (to 40) per cofloresc¬
ence; pseudospikelets lanceolate, 1— 3(— 4) cm
long, 3—6 mm wide; foliage leaf sheaths fimbriate

Volume 12, Number 1
2002

Londono & Clark
New Taxa of Guadua

75

bill not auriculate al the summit .

. G. tagoara subsp. tagoara

lb. Pseudospikelets usually 1 to 6 (to 9) per coflo¬
rescence; pseudospikelets narrowly lanceolate,
2.5—4 cm long, 2-3 mm wide; foliage leaf
sheaths auriculate-fimbriate at the summit . . .
. G. tagoara subsp. glaziovii

Guadiia tagoara (Nees) Kunth subsp. tagoara

Foliage leaf sheaths bearing only fimbriae at the
summit, the fimbriae 8—15 mm long. Pseudospi¬
kelets 9 to 15 (to 40) per coflorescence, 1— 3 (—4) cm
long, 3-6 mm wide, lanceolate, with I to 4 (ertile
florets; sterile lemmas 5— 6(— 7) mm long; lemmas
7—1 1 ( — 14) mm long, 5— 7(— 8) mm wide, the macro
0.5—1 mm long; paleas 7 — 1 1 ( — 13) mm long. 4-5
mm wide. Lodicules 2.5-4 mm long. 1-3 mm wide.
Ovary 3—6 mm long.

Distribution. Atlantic and montane forests of
the eastern coast of Brazil from Bahia to Santa Ca¬
tarina; 50—1200 m.

One collection from Serra do Mar in Sao Paulo
(Medina 5), with shorter pseudospikelets and very
congested coflorescences with up to 40 pseudos¬
pikelets, and another from the Serra dos Organs in
Rio de Janeiro (Reitz 6926). with robust pseudos¬
pikelets up to 3.6 cm long and 6 mm wide, (filler
from the remaining collections. However, popula¬
tions of G. tagoara in the two areas were visited by
both authors; only minor variations in vegetative
features were observed and the specimens lit well
within this subspecies in every other respect.

Specimens examined. BRAZIL. Bahia: Mun. Gandu,
2 kin N of Gandu City, Faz. Santo Antonio, 90 m, 19 Apr.
1972, Calderon A Pinheiro 2228 (CEPEC, k. MO. US);
Mun. Itajuipe, 2 km S of Pimenteira on road to Uniao
Oueimada. Fazenda Sao Jorge, 500 m, 5 May 1976, Sod-
erstrom el al. 2186 (CEPEC, F, K. MO, SP. US); Mun.
Jequie, road Jequie-piau, ca. 40 km SE ot Jequie, I 75 m.
16 May 1976, Calderon et al. 2439 (INPA, k. MO, US);
Mun. Mascote, 20 km E of Camara, 9 km W of Santa
l.uzia do Salobro on road to Canavieiras, edge of rio Be¬
lem, 7 Apr. 1976. Soderstrom et al. 2128 (CEPEC. E. k.
SP, US). Espirito Santo: Vitoria. Viana. 12 June 19B7
(fl). Gomez 84 (SP); Mun. Domingo Martins, km 42 before
Marechal Floriano, 530 m, 21 Feb. 1990, Clark A Morel
69 1 (ISC. MBME. MO, NY. RB, SJRP, SP. US). Minas
Gerais: Mun. Marlieria, Parque Estadual do Rio Doce,
400 m. 18 Sep. 1975. Heringer A Eiten 15024 (MO, US).
Parana: Capao Bonito, in silva primaria, 27 Mar. 1915
(fl). Dusen 16878 (F, k, MO. NY, US); Mun. Campina Gde.
do Sul, Eigueira. rio Capivari, 19 Dec. 1972 (II). Hatsch-
bach 31014 (ISC. MBM, SP, US); Mun. Antonina, rio Pe-
queno, 50 m, 10 Jan. 1974 (fl). Hatschbach 33640 (ISC.
MBM, US); Mun. Lima Duarte, Serra do Ibitipoea, about
1 10 m before the entrance to the Parque Estadual (Elo-
restal) do Ibitipoea, 1220 m, 3 Feb. 1991, (dark A Morel
784 (BHCB. ISC, MO, SJRP, SP, US). Rio de Janeiro:
Mun. Resende, 27 Apr. 1926, Gehrt A Hoehne 17589

(US); Parque Nac. de Itatiaia, 12 Nov. 1943 (fl), McClure
21285 (ISC. k. US), McClure 21291 (k, US); Parque Nac.
de Itatiaia, road to Maromba, near Hotel Simon, 1000 m,
12 Feb. 1990, Clark et al. 670 (AAU, ISC. k. MO. NY,
RB. SJRP, SP, US); between Nova Friburgo and Caehoeiro
de Macaco, 980 m, 30 Apr. 1972 (fl), Soderstrom A Sucre
1978 (CEPEC, INPA, k. RB. US); Mun. Nova Friburgo,
Macae de Cima, nascente do Rio Macae, 1000 m, 28 Nov.
1986, Martinelli et al. 12008 (RB); Reserva Macae de
Cima, sit io Taquara-oca, trail to Caseada Preta, 710 m, 8
Mar. 1992, Londono A Sarahyba 720 (IBGE. RB. FUEV,
US); Parque Nac. de Itatiaia. 1050-1250 m, 30 Apr. 1985
(fl). Zuloaga et al. 2344 (MO. US, VV IS). 1 1 10 m. 6 Mar.
1992, Irmdono A Sarahyba 707 (IBGE. RB. TULV, US);
Nova Friburgo, 9 Dec. 1918, Curran 622 (US); Mun. Ter-
esopolis, Serra dos Orgaos, Rio Paquequer, 1000 m, 15
Aug. 1940 (fl), Brade 16566 (IAN, RB), 12 June 1940 (fl),
Grade 16280 (IAN, R); Parque Nacional da Serra dos Or¬
gans. 16 Feb. 1990, Clark et al. 685 (ISC. MO, NY. RB,
SJRP, SP. US), 1 170 m, 6 Mar. 1992, Londono & Sarahyba
713 (IBGE. RB. TULV, US); 16 km S of Itaipara, Serra
dos Organs. 950 m, 26 Mar. 1976 (II), Davidse et al. 1 1420
(k, MBM. MO, NY, US); Serra dos Organs, s.d. (11), Reitz
6925 (SP. US); Soberbo, 13 July 1968 (11). Braga 28 (RB);
Mun. Petrdpolis, vale de Bom Sucesso, 720 m, 6 May
1972, Soderstrom A Sucre 1985 (INPA, RB. US); Vas-
souras, 400 m, Apr. 1976 (11). Sucre 11192 (k. RB, US);
Bom Jesus de Itabapoana, Carabugu, 7 July 1982 (fl). Car-
auto el al. 4293 (R); Rio de Janeiro, s.d. (fl). Glaziou
20153 (k, SP, US); Rio de Janeiro (11), s.d.. Glaziou 13324
(E. k); Rio, s.d. (fl), Sello 17 (US). Santa Catarina: Ter¬
esa polis, S. Amaro, 400 m, 24 Aug. 1958 (fl), Reitz di-
Klein 7058 (US); Rio Tavares, Ilha de S. Catarina, Flori-
anopolis, 200 m, 28 July 1965 (fl), Klein A Bresolin 61 19
(LIS): Vargem Grande, Lauro Miiller, 350 m, 11 July 1958
(fl). Reitz A Klein 6732 (US); Mata da Cia. Hering, Blu-
menau, 23 Nov. 1954 (fl), Reitz A Klein 2283 (US); Novo
Horizonte, Lauro Muller, 350 m, 22 Aug. 1958 (fl). Reitz
A Klein 7036 (MBM, SP, UB, US); Parque Botanieo do
Mono Bau, Ilhota, 300 m, 1 Nov. 1968 (fl), Reitz 6890
(SP. US); ca. 10 km from Aiure, 25 Feb. 1992, lx>ndoho
A Clark 705 (IBGE, TULV, US); about 12-13 km along
turn off from BR-101 to Brusque near Cunhas, 20 m, 9
Mar. 1991, Clark A Oliveira 886 (HRB. ISC, SJRP, SP.
US); road Sao Bento do Sul— Corupa, 17 km NW of Co-
rupa. 590 in. 4 Mar. 1991. Clark A Windisch 857 (HRB.
ISC. MO. RB, SJRP, SP. US); Serra Geral. road Timbe do
Sul to S. Jos6 dos Ausentes, 9.8 km from Timbe do Sul,
Serra da Roeinha, 425 m. 6 Mar. 1991, Clark A Windisch
873 (HRB. ISC, SJRP, SP, US). Sao Paulo: Taubale et
Lorena, Od. 1833, Riedel 1636 (k, NY, US-fragment);
road between Piedade and Tapirai at km 107 marker, 3
Mar. 1960, Medina 10 (SP); Serra da Cantareira. 18 May
1901 (fl), Hammar 6116 (SP, US), 14 June 1948 (fl). Kuhl-
mann s.n. (SP); between Taubat6 & Ubatuba, Serra do
Mar, 20 Nov. 1962 (fl), Medina 5 (SP, US); Serra da Can¬
tareira, 22 Oct. 1948 (11), Pickel 5891 (SP. US); Carlos
Botelho State Park, Ribeirao Branco, km 55, ca. 23 km S
of Park headquarters, 2 Sep. 1987, Gentry et al. 58817
(MO): Reserva Forestal Carlos Botelho, Serra de Parana-
piacaba, past Rio Bonito, 28 Jan. 1990, Clark A Windisch
648 (ISC. k. MO, NY. RB, SJRP, SP. US), 185 m, 16 Feb.
1992, Clark A Oliveira 1025 (ISC, SJRP. SP, US); Sta.
Izabel. Igarata, 26 June 1952 (fl). Kuhlmann A Gonzalves
3147 (SP. UB), 26 Sep. 1950 (11), Kuhlmann 2549 (SP);
Mun. de Salesopolis, BoracOa, 28 Apr. 1949 (fl), Kuhl¬
mann 3146 (SP); base of Serra da Bocaina, road Areias-

76

Novon

Sao Jos£ de Barreiro, km 251.5, 15 Jan. 1990, Clark A
Morel 624 (AAU, ISC, MO, NY, KB. SJRP, SR US); Mini.
Biritiba Mirim, Estagao Biol. Boraceia, Casa Grande, 800
m, 24 Eel). 1991, Clark A Morel HIS (ISC. MBM, MO,
NY, RB, SJRR SR US); SI'- 183, road Piquete— Cruzeiro,
about 10 km E of Piquete, near the gate of a fazenda, 570
m, 16 Jan. 1991, Clark & Morel 749 (ISC. k. MO. RB,
SJRP, SP, US); 5.5 km from the Paraiui border, route
BR 1 16, 630 m, 17 Feb. 1992, LondoOo A- Clark 6 H9
(IBGE, SR US).

Giiuduu tagoarn subsp. glaziovii (Hackel) Lon-
dono & L. G. Clark, comb, et stat. nov. Bas-
ionym: Bambusa glaziovii Hackel, Oesterr.
Rot. Zeit. 53(5): 194. 1903. Guadua glaziovii
(Hackel) E.-G. Camus (as glaziowii ), Res Barn-
bushes I: 108, pi. 66, fig. A. 1913. TYPE.
Rra/.il. Rio de Janeiro, s.d., A. F. M. Glaziou
17450 (holotype, W; isotypes, k, MR. P. US-
fragment).

Guadua longifimbriata E.-G. Camus, Ees Bambusees I:
113, pi. 64, fig. C. 1913. Bambusa longifimbriata
(E.-G. Camus) McClure, Smithsonian Contr. But. 9;
66. 1973. TYPE: Brazil. Rio de Janeiro: Petropolis.
Sao Cristovao, 29 Feb. 1872, A. F. M. Glaziou 5717
(holotype, P not found; isotypes, (7 IAN, k. US-frag-
ment).

Foliage leaf sheaths bearing fimbriate auricles at
the summit, the auricles 2-5 mm long, dark brown,
the fimbriae 6— 15(— 20) mm long. Pseudospikelets
1 to 6 (to 9) per coflorescence, 2.5—4 cm long, 2—
3 mm wide, narrowly lanceolate, with 2-6 fertile
llorets; sterile lemmas 4.5—6 mm long; lemmas 7.5—
9 mm long, 5—6 mm wide, the mucro 0.5 mm long;
paleas 7—8.5 mm long, 3.5— 4.5 mm wide. Lodicules
2. 5-3.2 mm long, 1—2.5 mm wide. Ovary 3—4 mm
long.

Etymology. Phis subspecies is named for Au¬
guste F. M. Glaziou, a prolific French collector in
Brazil during the 19th century.

Distribution. This subspecies occurs in the At¬
lantic forests only of the state of Rio de Janeiro.

Paratypes. BRAZIL. Rio de Janeiro: Mun. Nova
friburgo, Distr. de Maeah de Cima, a 3 km do Hotel Fa¬
zenda Sao Joan, 1000 m, 6 June 1989 (II), Lima et at.
ASH l (MO, NY, B. RB, US). 960 m, 8 Mar. 1992. Londono
A Sarahyba 71 6 (IBGE, RB, TULV, US); Barque Nacional
da Tijuca, Bom Betiro, 575 m, 25 Feb. 1972, Soderstrom
et al. 1H54 (CEPEC, INPA. k. RB, US): Rio de Janeiro,
24 May 1895 (fl). Hie 4115 (IAN, US).

Acknowledgments. We are grateful to Jairo Lar-
rahondo lor his excellent illustrations. XL thanks
lh(‘ International Timber Organization (ITTO) of Ja¬
pan for support (Grant No. 047/99 A). XL also
thanks Paul Peterson (Smithsonian Institution) and
Lynn Clark for the use of laboratory facilities. Spe¬
cial thanks are given to Pedro Acevedo and Lynn
Clark for their support of XL throughout this study.
Fieldwork by XL in Brazil was made possible by a
grant to Clark and Londono from the National Geo¬
graphic Society; Sergio Sarahyba, Haroldo C. de
Lima, and Paulo Windisch provided logistical sup¬
port. XL thanks Mikio Kobayashi, Lars P. Kvist.
and INC1VA for support for fieldwork in Amazonian
Colombia, northern Amazonian Peru, and in the
Cordillera Oriental of Colombia. Fieldwork in Bra¬
zil by LC was supported by NSF grant BSR-
8906340, with subsequent support for manuscript
preparation from NSF grant DEB-9806877.

I .iterature Cited

Grass Phylogeny Working Group. 2001. Phylogeny and
subfamilial classification of the grasses (Boaceae). Ann.
Missouri Bot. Gard. 88: 373-457.

Judziewicz, E. .1., L. G. Clark. X. Londono A M. J. Stem.
1999. American Bamboos. Smithsonian Institution
Bress, Washington, D.C.

McClure, F. A. 1934. The inflorescence in Schizostachyum
Nees. J. Wash. Acad. Sci. 24: 541-548.

Soderstrom, T. B. 1981. Some evolutionary trends in the
Bambusoideae. Ann. Missouri Bot. Gard. 68: 15—47.
Stapleton. C. M. A. 1997. The morphology of woody bam¬
boos. Bp. 251-267 in G. B. Chapman (editor). The Bam¬
boos. Academic Bress, London.

Young, S. M. A W. S. Judd. 1992. Systematics of the
Guadua angustifolia complex (Boaceae: Bambuso¬
ideae). Ann. Missouri Bot. Gard. 79: 737—769.

Una Nueva Especie de Echeandia (Anthericaceae)
de Guerrero, Mexico

Ana Rosa Lopez- Ferrari, Adolfo Fspejo Serna, y Jacqueline Ceja Romero
Herbario Metropolitano, Departamento de Biologfa, C. B. S., Universidad Autonoma
Metropolitana, Unidad Tztapalapa, Apartado postal 55-535, 09340 Mexico, D.F.

aes@xan u m.uam . mx

RksUMKN. Se describe e ilustra Echeandia mag-
nifica s|>. nov. (Anthericaceae) con material prov-
eniente del estado de Guerrero, Mexico. Ea nueva
especie, perteneciente al subgenero Echeandia,
esta relaeionada con E. michoacensis (Poellnitz)
Cruden y con E. drepanoides (Greenman) Cruden,
pero es claramente distinguible de la primera por
sus hojas con el margen largamente ciliado, por su
infloreseencia ramificada y por sus flores grandes
con los tepalos internos ampliamente eliptieos. I)e
E. drepanoides se distingue por el mayor tamafio de
sus anteras y porque estas abren por una hendidura
apical.

ABSTRACT. A new species, Echeandia magnifica
(Anthericaceae), from Guerrero, Mexico, is de¬
scribed and illustrated. The new species, included
in the subgenus Echeandia, is related to E. mi-
choacensis (Poellnitz) Cruden and E. drepanoides
(Greenman) Cruden but is clearly distinguished
from the former by its leaves long ciliate in the
margins, its branched inflorescence, and its large
flowers with the internal tepals widely elliptic in
shape. From E. drepanoides it is distinguished by
its larger anthers that dehisce through apical slits.

Key words: Anthericaceae, Echeandia, Guer¬
rero, Mexico.

K1 genero Echeandia con 78 especies (Cruden,
1999), es el segundo mas grande dentro de las An¬
thericaceae (Dahlgren et al., 1985) y esta restrin-
gido a America. En los ultimos 20 anos se han
deserito cerca de 50 especies (Cruden, 1981, 1986,
1987, 1993, 1999). lo que equivale aproximada-
mente a la mitad de los laxa conocidos y al 75%
de los presentes en Mexico (Espejo & Lopez-fer-
rari, 1993).

En recientes viajes de campo al estado de Guer¬
rero, recolectamos material del genero que no pudo
ser asignado a ninguna de las especies descritas y
la revision cuidadosa del mismo nos llevo a con-
cluir que se trata de una especie nueva que a con-
t i n uacion ( leseril >i mos.

Echeandia magiiilica Lopez-Eerrari, Espejo & Ceja,
sp. nov. TIPO: Mexico. Guerrero: mun. Chilpan-
cingo de los Bravo, ca. 2 km despues de X<x:-
omanatlan, rurnbo a Omiltemi, vegetaeion ar-
vense en bosque de pino-eneino, 17°33'49"N,
99°38' 1 9"W, 1950 m, 4 Sep. 1999, J. Ceja, A.
Espejo, A. R. Lopez-Eerrari y A. Mendoza 907
(holdtipo, IJAMIZ; isotipos, BRIT, ENCB,
FCME, IEB, MO, UC). Eigura 1.

Herba ex radicibus 12—15 cm longis, folia basalia, li-
nearia vel lanceolata, 50—60 cm longa, 4.5—5 cm lata,
marginibus ciliatis. Inflorescentia scapo lereti, glabro,
usque ad 1(K) cm lougo; insidens tepalis flos flavis, I f > —

1 8.5 mm longis, 6-12 mm latis; filarnentis muricatis; an-
theris liberis. Capsula oblongoidea, 12-15 mm longa, 7-
10 mm diametro.

Hierba perenne, glabra, de hasta 1.8 (2.1) m de
alto incluida la infloreseencia; raices cih'ndricas, de
12—15 cm de largo, engrosadas en la poreidn distal;
cormo cubierto por las vainas persistentes de las
hojas, que se vuelven Hbrosas con la edad; hojas
basales lineares a lanceoladas, de 50—60 X 4.5—5
cm, verdes en el haz, glaucas en el enves, sinuadas
y largamente ciliadas en el margen con pelos de
hasta 1 mm de largo; infloreseencia una pamcula
con numerosas flores, de ca. 90 cm de largo; escape
cilmdrieo, glabro, de 90-160 cm de largo; bracteas
del escapo triangulares, de 10-14 cm de largo;
bracteas de la infloreseencia triangulares, de 5-7
cm de largo; bracteas florales angostamente trian¬
gulares, de ca. 5 mm de largo; pedicelos de ca. 1
cm de largo, articulados por debajo de la mitad,
acrescentes en el fruto; flores amardlo-anaranjadas,
divaricadas; tepalos extemos eliptieos, de 16-17 X
6-6.6 mm, acuminados, enteros, trinervados; te¬
palos intemos ampliamente eliptieos, de 18—18.5
X 12 mm, redondeados, enteros, trinervados; fila-
mentos muricados, mas conspicuamente en su mi¬
tad apical, de ca. 7.4 mm de largo; anteras libres,
oblongas, de 4.5—5 mm de largo, con dehiscencia
por una hendidura apical; ovario oblongo, de 4 mm
de largo; estilo de ca. 1 cm de largo. Capsula ob-
longa, triquetra, de 12—15 mm de largo, de 7—10

Novon 12: 77-79. 2002.

78

No von

^-^o'^0eP

Figiira I. Echeandia magnified Ldpez-Ferrari, Espejo & Ceja. —A. Hdbito. — B. Detalle de la inflorescencia. — C y
D. Hor. E. Hor disecada. — F. Detalle <lel androeeo y el gineceo. — G. Fruto. (Ilustraeidn basada pii el ejemplar j.
Ceja el al. 907, holdtipo.)

Volume 12, Number 1
2002

Lopez-Ferrari et al.
Echeandia magnifica

79

mm de diametro; semi lias numerosas, de ca. 2 mm
de diametro, con la testa negra, caliculosa. Numero
cromosomico n = 8 ( Crude n I 724).

La especie acjui descrita pertenece al subgenero
Echeandia ya que presenta flores amarillas y te-
palos elfpticos. Echeandia magnifica muestra si¬
militudes, tanto en caracteres vegetativos eomo flor-
ales, con E. michoacensis (Cruden, 1993; von
Poellnitz, 1943); sin embargo esta ultima presenta
el escapo generalmente simple y el margen de las
liojas entero a mas o menos muricado o denticu-
lado, a diferencia de la primera que lo tiene lar-
gamente ciliado.

Echeandia magnifica presenta tambien semejan-
zas con E. drepanoides, a la eual se parece mueho
vegetativamente debido a sus liojas con el margen
largamente ciliado, pero de la eual difiere en el
tamano mayor de las anteras y en que estas abren
por una hendidura apical. Ademas, E. drepanoides
se conoce solo de algunas localidades dispersas en
el estado de Oaxaca.

Otro taxon con el que la nueva especie podri'a
eonfundirse, dada su distribucion geografica, es
Echeandia echeandioides (Schlechtendal) Cruden;
sin embargo, las anteras de esta especie presentan
dehiscencia lateral y no apical y las hojas son mu-
cho mas angostas y tienen el margen diminuta-
mente denticulado a cortamente ciliado.

Echeandia magnifica se conoce hasta ahora un-
icamente de los alrededores de Chilpancingo, en el
estado de Guerrero, en donde crece abundante-
mente en sitios encharcados dentro de las areas
perturbadas del bosque de pino-encino. HI epfteto
especffico hace referenda al tamano de las plantas
y de las ilores que caracterizan a la especie.

Pardtipos. MEXICO. Guerrero: man. Chilpancingo
de los Bravo, ca. 2 km despues de Xocomanatlan, rumbo
a Omiltemi, 4 Nov. 1999, J. Ceja, A. Espejo, A. R. Ldpez-
Ferrari y A. Mendoza 1023 (ENCB, MEXIJ, UAMIZ);
mun. Chilpancingo de los Bravo, ca. 23 km S of Chilpan¬
cingo. ca. 1090 m. 2 Aug. 1970. R. W. Cruden 1/24
(ENCB. GH, k, UC).

Agradecimientos. Agradecemos a Victoria Sosa
la revision crftica del manuscrito. Robert W. Cru¬
den examino el trabajo y puso amablemente a nues-
tra disposition sus notas asf eomo material adi-
cional de la nueva especie aquf descrita. Roy
Gereau corrigio y complemento la diagnosis latina
y Rolando Jimenez Machorro elabord la ilustracion
que acompana al trabajo.

Eiteratura Citada

Cruden, R. 1981 . New Echeandia (Liliaceae) from Mexico.
Sida 9: 139-146.

- . 1986. New species of Echeandia (Liliaceae) from

Central America. Phytologia 59: 373-380.

- . 1987. New species of Echeandia (Liliaceae) from

Nueva Galicia. Contr. Univ. Mich. Herb. 16: 129-133.

- . 1993. New species of Echeandia (Liliaceae) from

Oaxaca, Mexico. Phytologia 74: 128—137.

- . 1999. A new subgenus and fifteen new species

of Echeandia (Anthericaceae) from Mexico and the
United States. Novon 9: 325—338.

Dahlgren, B. M. T.. H. T. Clifford & P. F. Yeo. 1985. The
Families of the Monocotyledons. Springer- Verlag. Ber¬
lin.

Espejo, A. & A. R. Lopez-Ferrari. 1993. Las Monocoti-
ledoneas Mexicanas una Sinopsis Flonstiea 1. Lista de
Referenda PARTE II. Anthericaceae, Araceae, Areca-
ceae, Asphodelaceae y Asteliaceae. Consejo Nacional
de la Flora de Mexico, A. C., Universidad Autonoma
Metropolitana-lztapalapa, Comision Nacional para el
Conocimiento y Uso de la Biodiversidad. Mexico, D.F.
Poellnitz, k. von. 1943. Neue Formen der Gattung An-
thericum L. au.s America. Revista Sudamer. Bot. 7:
157-162.

Macrocarpaea august [folia (Gentianaceae), a New Species

from Peru

James S. Pringle

Koyal Botanical Gardens, P.0. Box 399, Hamilton, Ontario, Canada LBN 3H8

ABSTRACT. Macrocarpaea angustifolia, known
only from the Departamento de Pasco, Peru, differs
from die sympatric M. revoluta in having glabrous,
narrower, acuminate leaves with obscure secondary
venation. It further differs in its verrucose (Rus-
byanthus- type) pollen, which suggests a closer re¬
lationship to such species as M. corymbosa and M.
viscosa.

Key words: Gentianaceae, Macrocarpaea, Peru.

Macrocarpaea angustifolia J. S. Pringle, sp. nov.
TYPE: Peru. Pasco: Prov. Oxapampa, head¬
waters of Rio Tunqui, trail to Chuchurras-Pal-
cazu, 75°28'W, I (FIT'S, shrubby vegetation on
ridge, 1900 in, 2 Jan. 1984, R. Foster, M.
Chanco, J. Alban * D. I\. Smith 7747 (holo-
type, MO; isotype, USM photocopy seen). Fig¬
ure 1 .

Frutex glabra. Folia subcoriacea, laminis elliptico-
lanceolatis, 10—14 X 1.5—3 cm, marginibus incrassatis
revolutisque, apicem versus peracuminatis, et petiole an-
gusto circa I cm longo. Calyx 6—10 mm, lobis semiorbi-
cularibus tube duplo brevioribus, margine scarioso. Co¬
rolla 35—45 mm, lobis ovato-oblongis, 7—10 mm longis.
Exinium poll inis verruoosum.

Shrub to ca. 3 m, glabrous throughout. Stems
nearly terete, striate. Leaves firm, blades dark
green above, paler beneath, elliptic-lanceolate, 10—
14 X 1 .5—3 cm, only midrib conspicuous, second¬
ary and lesser veins obscure; margins thickened,
revolute; base tapering to a slender petiole up to
ca. 1 cm long; apex strongly acuminate. Inflores¬
cence a terminal, open, paniculate cyme of ca. 15—
25 (lowers. Flowers ± nodding, on pedicels 15—25
mm long. Calyx campanulate, 6-10 mm long, lobed
ca. Vi of its length; lobes equal or nearly so, semi¬
circular, slightly overlapping, scarious-margined.
Corolla pale yellowish green, 35—45 mm long; tube
expanding rather abruptly 2-4 mm above summit
of calyx; lobes oblong-ovate, 7-10 mm long. An¬
thers seen after dehiscence, estimated ca. 5.0 mm
long when straight. Exine of pollen verrucose. Ma¬
ture fruit not seen.

Known only from the type collection.

Superficially, M. angustifolia somewhat resem-

Novon 12: 80—8 1 . 2002.

hies the Venezuelan species formerly known as M.
salici/olia Ewan. The latter, however, has pollen in
tetrads, and is now placed in Chelonanthus or, by
those more narrowly defining the genera in this com¬
plex, in Rogersonantbus, whereas M. angustifolia has
separate pollen grains, like all other species of Ma¬
crocarpaea as the genus is now circumscribed. Also,
in M. angustifolia as in Macrocarpaea species gen¬
erally, the bracts in the inflorescence are gradually
reduced, the proximal bracts being distinctly leaflike,
whereas in those species transferred to Chelonanthus
or Rogersonanthus all of the inflorescence bracts are
greatly reduced and scalelike. Among species re¬
tained in Macrocarpaea, M. angustifolia exhibits
some similarity to M. revoluta (Ruiz & Pavdn)
Gilg. which is native to the same part of Peru. The
leaves of M. angustifolia differ in being glabrous,
much narrower, and strongly acuminate, with ob¬
scure secondary venation and with more slender
petioles, and, at least as seen through comparison
of the respective type specimens, its inflorescence
is fewer-flowered and more open. It also resembles
M. subcaudata Ewan, of Costa Rica and Panama,
especially in its narrow, acuminate leaves but dif¬
fers in its terrestrial rather than epiphytic habit,
its much longer pedicels, and its glabrous calyx.

Another somewhat similar species is M. ban-
giana Gilg of Bolivia, from which M. angustifolia
differs in its narrower, glabrous, strongly acumi¬
nate leaves of thicker texture, without the con¬
spicuous secondary venation of the Bolivian spe¬
cies, and in its smaller flowers. The pollen of M.
angustifolia is conspicuously verrucose, thus be¬
ing of the M. corymbosa -type or Rusbyanthus- type
of Nilsson (1968, 1972), whereas that of M. revo¬
luta, M. subcaudata, and M. bangiana is of Nils¬
son’s M. glabra- type (Nilsson, 1968, 1972; Weav¬
er, 1972). Palynology suggests that the closest
relationships of M. angustifolia are with M. cin-
chonifolia (Gilg) Weaver (Ecuador to Bolivia). M.
corymbosa (Ruiz & Pavon) Ewan (S Peru), M.
pachystyla Gilg (S Peru), and M. viscosa (Ruiz &
Pavdn) Gilg (Peru), which also have the M. cor¬
ymbosa pollen type.

Acknowledgments. I am very grateful to Jason

Volume 12, Number 1
2002

Pringle

Macrocarpaea angustifolia from Peru

81

Dept. Pasco. Prov. Oxapampa.
Headwaters of Rio Tunqui. trail to
Chuchurras-Palcazu.

7 5* 2 8 1 W » 1 0 * 1 4 ' S .

Shrubby vegetation on ridge.

Alt. 1 900 m.

Shrub. 3 r; flowerr pole yelloviri-

Pobin Foster. M. Chanco.2 Jan.

J. Alban 4 D.N. Smith 7747
MISSOURI BOTANICAL GARDEN HERBARIA

figure I. Macrocarpaea angustifolia J. S. Pringle, Foster et al. 7747 (holotype. MO).

R. Grant of l’Universite de Neuchatel, Switzerland,
for discussions about Macrocarpaea, comments on
the manuscript, and photocopies of the isotype. The
type specimen of the name M. angustifolia was col¬
lected under the auspices of the Flora of Peru pro¬
ject, a collaborative effort of the Missouri Botanical
Garden, the Universidad Nacional Mayor de San
Marcos, and the Universidad Nacional de Amazon¬
ia Peruana. Fieldwork by Robin Foster et al. was
supported by US/MI). This is Contribution no. 88

from the Royal Botanical Gardens, Hamilton, On¬
tario.

Literature Cited

Nilsson, S. 1968. Pollen morphology in the genus Macro¬
carpaea (Gentianaceae) and its taxonomical signifi¬
cance. Svensk But. Tidskr. 62: 338—364 + 4 pi.

- . 1972. Pollen morphological contributions to the

taxonomy of Usianthus |sic) s. lat. (Gentianaceae).
Svensk Bot. Tidskr. 64: 1 — 13 + 8 pi.

Weaver, R. E., Jr. 1972. The genus Macrocarpaea in Costa
Rica. J. Arnold Arbor. 53: 553-557.

Pseudostellaria sierrae (Caryophyllaceae), a New Species

from California

Richard K. Rabeler

University of Michigan Herbarium, 1205 N University Ave., Ann Arbor, Michigan

48109-1057, U.S.A.

Ronald L. Hartman

Rocky Mountain Herbarium, Department of Botany, University of Wyoming, Laramie,

Wyoming 82071-3165, U.S.A.

ABSTRACT. Pseudostellaria sierrae is described as
new from northern California. Glabrous stems and
leaves, a V-shaped apical notch in each ligulate
petal, live yellow anthers, and seeds with minute
projections on each tubercle are features useful in
distinguishing P. sierrae from P. jamesiana (Torrey)
W. A. Weber & R. L. Hartman, the congener widely
distributed in the western United States. Pseudos¬
tellaria sierrae is found in mixed oak or conifer
forests.

Key words: California, Caryophyllaceae, North
America, Pseudostellaria, Sierra Nevada, Stellaria ,
United States.

Pseudostellaria is a genus of 16 (Bittrich, 1993)
to 20 species (Rabeler, pers. compilation), all but
3 of which are restricted to Asia (Afghanistan and
Siberia east to Japan and Korea). Two additional
species from Korea were recently described and ef¬
fectively, but not validly, published (bee, 1998).
Pseudostellaria europaea Schaeftlein occurs in
southeastern Europe, while two species, P. jamesi¬
ana (Torrey) W. A. Weber & R. L. Hartman and
this novelty, are known in western North America,
fhe chief feature distinguishing Pseudostellaria is
the presence of thickened root sections, often as
small tubers near the base of the stem (P. jamesi¬
ana, Weber & Hartman, 1979) or as tuberous, ci¬
gar-like, thickened roots arising periodically along
the rhizome (P. sierrae. Fig. I). While most species
of the genus have both chasmogamous and cleis-
togamous flowers, the North American species lack
the latter. Pseudostellaria jamesiana and P. sierrae
share an unusual pattern of fruit dehiscence with
the six valves usually rolling back 2 or 3 revolu¬
tions, forming a shallow dish (Fig. 1C; Weber &
Hartman, 1979). This mechanism may insure the
release of the 1 or 2 seeds that are quite large.

Novon 12: 82-86. 2002.

Pseudostellaria sierrae Rabeler & R. F. Hart¬
man, sp. nov. TV PF: U.S.A. California: Plumas
Co., in litter under Pinus lambertiana, mixed
oak-pine-fir forest, 150 ft. S of FS road
26N49Y, 0.9 mi. F of Plumas Co. 204 (Dixie
Canyon Rd.), 1.95 mi. N of California 89. W
of Crescent Mills, 40°06'24"N, 120°55'45"W,
T26N, R9F. sec. 14, SF V4, ca. 1456 m, 14
June 1997, R. K. R<d)eler 1290 | wit h B. Ertter,
T. Morosco, J. Shevock] (holotype, MICH; iso¬
types. GH, MO, RM. RSA, UC). Figure 1.

Herbae perennis omnino glabra; radices vertieales car-
nosi rhizomatibus. Folia lanceolata ad elliptica marginibus
integris term iter scariosis. Flores perfeeti actinomorphi 1 —
3 per caulem vel ramum; sepala lanceolata ad anguste
ovata marginibus scariosis; petala 5, ligulata apiee inciso
V-formo; stamina 5, antherae flava; sty li 3 distincti, stig-
matibus quasi terminalibus. Capsule ovoidea, valvis 6
pauxillum recurvis ad arete 2- vel 3-plo revolutis; semina
1 vel fortasse 2 brunneola circularia ad oblonga lateraliter
complanata tuberculata aliquot tuberculis 6-12 verrucis
rubris minutis; verrucae saepe 5—8 projecturis conicis
minutis; radicula prominens.

Perennial, glabrous herb 9-27 cm tall, with
stems arising singly, at vary ing intervals along rhi¬
zome. Roots vertical or nearly so, 5—15 cm long,
ca. 0.3 mm thick near point of attachment to rhi¬
zome, enlarging to 2-3 mm thick below, fleshy; rhi¬
zomes white to tan, 0.5—2 mm diam., shiny, usually
with 1 to several scaly buds at the nodes. Stems
simple or branched, rounded, obtusely angled or
grooved when pressed, internodes 1—4 cm long or
more below, often 0.3—1 mm long in upper V3—V2,
axillary branches 1 to 3(7), 1— 5(9.5) cm long.
Leaves opposite, exstipulate, the lowest 2 to 6 and
those of the rhizome rudimentary, connate for 0.5—
1.5 mm, 3—8 mm long, the primary leaves in 5 to
10 pairs on main stem with bases weakly fused,
those of the upper stem and especially branches
generally much more closely spaced, narrowly lan¬
ceolate or elliptic, 0.7-3. 5 cm long, 1-5(8) mm

Volume 12, Number 1
2002

Rabeler & Hartman
Pseudostellaria sierrae from California

83

84

Novon

figure 2. Scanning electron micrograph of seed of Pseudostellaria sierrae ( Hartman 58546. KM) showing the minute
projections on each tubercle. Scale line represents 10 gin.

wide, lower surface of blade finely reticulate, pus¬
tulate (30 X magnification), margin entire, thinly
scarious. Flowers perfeet, chasmogamous, regular,
solitary, terminal and axillary on main stem and
often 1 or 2 (to 6) axillary branches, sometimes
abortive; sepals lanceolate to narrowly ovate, 4—6.5
mm long, to 7 mm in fruit, shiny, glabrous, margins
scarious, with midvein and 4 to 6 weaker veins
arising from base, apex acute or acuminate; petals
5, while, ligulate, 5-8(9) X 1-1.5 mm, apex with
a V-shaped notch 0.2— 0.7 mm deep; stamens 5,
each with a circular, thickened, basal gland 2-2.5
times the filament width, anthers yellow. 0.6— 0.7
mm long; styles 3, distinct, 2—3.5 mm long, stigma
terminal, 0. 1-0.2 mm long; ovary 3-carpellate;
ovules 3 to 6, most abortive. Capsule ovoid, 4—4.5
mm long, valves 6. dehiscing by recurving some¬
what to rolling back tightly 2 or 3 revolutions or for
2A of length with the lower portions forming a shal¬
low dish, exposing the basal placenta, a broad, ir¬
regular column 0.5-1 mm tall. Seeds I, possibly 2,
light brown, 3—3.4 mm long, circular to somewhat
oblong, flattened laterally, radicle prominent, ma¬
ture seeds with most tubercles broadly conical to
elongate, rounded, interdigitating, each with 6 to
12 minute red humps that develop into 5 to 8 con¬
ical projections ea. 0.02 mm long (50X magnifi¬
cation).

Pseudostellaria sierrae exhibits two features that
make placement within the most recent infragener-
ic classification (Mizushima, 1965) problematic,
first, 5 rather than (8) 10 stamens appears to he

unique in the genus. Mizushima (1965) divided
Pseudostellaria into two sections on the basis of
seed tubercle ornamentation. In section Pseudos¬
tellaria , including only P. rupestris (Turczaninow)
Pax, the seed tubercles each end in an apically
barbed glochid, while in section Mamillatae (Ohw i)
M. Mizushima the seed tubercles end in a single
deciduous spinule. Second, the minute conical pro¬
jections from the end of each tubercle in P. sierrae
may be an intermediate condition (Fig. 2).

Flic paratypes cited below include all other col¬
lections we have found. Habitat information sug¬
gests P. sierrae to he a plant of the woodland floor,
either in mixed oak-pine or fir forests, at 1250—
1970 m elevation. At both sites we visited, the
ground cover was sparse and mostly consisted of P.
sierrae. All collections occur in the Northern High
Sierra Nevada subregion of the California Floristie
Province (Hickman, 1993), the eight localities
spanning a latitudinal range of 285 km (Fig. 3).

Pseudostellaria sierrae may be more common
than suggested by the few collections cited. In de¬
scribing his efforts to locate P. sierrae, Powell Ahart
(pers. comm.), an experienced amateur familiar
with the Bowman Lake area, stated: “Both sides of
the road under Red Firs [Abies magnifica A. Murray
his] are massive stands of Pseudostellaria .” Given
the similarity of the range of these taxa (see Hick¬
man, 1993, and Map 8W in Little. 1971), we sus¬
pect additional fieldwork may show P. sierrae oc¬
cupying a somewhat larger area.

Botanists have collected P. sierrae occasionally

Volume 12, Number 1
2002

Rabeler & Hartman
Pseudostellaria sierrae from California

85

Figure 3. Distribution of Pseudostellaria sierrae. Area shown is the northern two-thirds of California.

since 1878. The most widely distributed collection
prior to our own collections, M. E. Jones 3287, in¬
troduced each of the authors independently to this
species. Rabeler first noticed P. sierrae when he
received seven specimens of Jones 328 7 among the
loans requested for his dissertation research on the
Stellaria calycantha (Ledebour) Bongard complex
(Rabeler, 1986). Although they were clearly not
Stellaria crispa Chamisso & Schlechtendal. Rabeler
was not able to identify them beyond “cf. Pseudos¬
tellaria ?” in 1986. Hartman was introduced to
Pseudostellaria sierrae via the UC specimen of
Jones 3287. When the UC loan to Rabeler was re¬
turned in late 1986, Hartman was working on the
Caryophyllaceae treatment for the Jepson Manual
(Hartman, 1993). The UC specimen of Jones 3287
that Rabeler had annotated as “cf. Pseudostellaria"
caught Hartman’s eye as well as the following cu¬
ratorial note: “ Pseudostellaria is from Asia. File in
Caryophyllaceae indet.” Hartman also was unable
to place it in an existing taxon.

With the exception of Jones 3287, all of the col¬
lections of P. sierrae identified to species we have
seen have been either misidentified as Stellaria ja-
mesiana Torrey (= P. jamesiana) or Arenaria ma-
crophylla Hooker (= Moehringia macrophylla
(Hooker) Fenzl) or identified only to one of these
genera (see Hartman. 1993, for generic delimita¬
tions). The only evidence that we have seen of pre¬

vious recognition of this species as new' was from
the USFS (at RM) sheet of Bolt’s 1934 collection
from Tadpole Spring. Identifying it as “ Moehrin -
giaf", M. L. Bombard had penciled a series of
notes on the sheet in 1939 ending with “possibly a
new species.” While all three taxa can he found in
Sierran coniferous forests, they can he easily sep¬
arated bv the characters given in die following key.

Key to Pseudosteiiaria sierrae ani> Species with
Which It Has Been Confused

la. Plants glabrous throughout; flowers solitary,
mostly terminal; stamens 5 Pseudostellaria sierrae

lb. Plants either with dense glandular hairs (espe¬
cially above) or peg-like hairs; flowers in cymes;
stamens 10.

2a. Plants with dense glandular hairs (some¬
times glabrous below); inflorescence bracts
large, leafy; petals shallowly notched ....

. Pseudostellaria jamesiana

2b. Plants with short, peg-like hairs; inflores¬
cence bracts tiny, scarious; petals entire . .
. Moehringia macrophylla

Paratypes. U.S.A. California: Nevada Co.. Truckee,
May 1878, J. P Moore s.n. (CAS); dry soil, shaded slopes
below sphagnum bog, Fall Creek, just E of Fake Bowman
Hoad, ca. 5 mi. N of Yuba Gap, 14 July 1965, 6. II. True
2257 [with J. T. Howell] (CAS, MICH, NY); in yellow pine
forest, dry rocky slope on mountain near dirt road anil dry
flat S of cement dam across Fall Creek, ca. lA mi. F, of
Bowman Lake Road, T1 7N, K12F, NW V* of sec. 6, 5600
ft., 10 Aug. 2000, L. Ahart 8650 (BRIT, CDA, CHSC,

86

Novon

DAV, II). JEPS, MICH, MSC, RM, IIC, V); Placer Co.,
Emigrant Cap, 28 June 1882, M. E. Jones 3287 (CAS,
MO, MSC, NY, POM, UC, US); timberland-chaparral, 1
mi. N of Tadpole Spring, T15N, RITE, sec. 5. 6500 ft.,
19 July 1934, H. Holt s.n. (USES at RM) but H. Holt 146
(UC); under sparse canopy of Abies magnified, especially
common in shaded areas or along large logs in the drip
zone, often forming a monoculture, near campsite, ea. 200
yards from foresthill Road, 2—2.5 road mi. W from Sailor
Elat trail turn-off, T15N, R13E, sec. 5, ca. 6500 ft.. 1 I
July 1997, R. L flartman 58546 (BRY, CAS, COLO,
MICH, NY, RM. US); Plumas Co., rocky hillside, Grizzly
Ridge, 6500 ft., 29 June 1919, W. W. Wagner s.n. (DS);
forest of mixed conifer, oak, maple, E side of mountain
above Crescent Mills, E of Round Valley Reservoir, T26N,
R9E, sec. 23, ca. 4500 ft., 14 May 1990, H. Enter 8984
(with G. Schoolcraft) (OSC, RM); shaded slope s in conif¬
erous forest surrounding Snake Lake, 39°58'N, 120°59'W,
1250 m, 30 May 1993, G. E Hrusa 1 1 139 & E. Sandoval
(CDA, JEPS); Tuolumne Co., Tioga Road, 6 June 1951,
//. & M. Hearing 6923 (SBBG); Gin Elat near Yosemite
Valley, June 1964. M. Ackley s.n. (CAS).

Acknowledgments. We thank David J. Keil for
assistance with the Latin description; Barbara Hit¬
ter (UC) for hospitality and logistics support in ar¬
ranging the expedition for Rabeler to collect the
type; James R. Shevock and Kathy Van Zuuk (IJ.S.
forest Service) lor arranging appropriate permitting
for Rabeler and Hartman, respectively; Larry Ja¬
neway (CHSC) for introducing the plant to Lowell
Ahart and for forwarding his collection and corre¬
spondence to MICH; Greg Brown for assistance
with the SHM photo; and Carolyn Crawford for the
illustration. We also thank the curators of CAS,

MO. RSA/POM, SBBG, and UC/JEPS for hospital¬
ity during visits by Rabeler and/or Hartman and
CAS, CHSC, DAY, F. GH. HSC. MO. MSC. NSMC,
NY, RENO, RSA/POM, SBBG, UC/JEPS, and US

for loans to MICH (all) or RM (UC) lor this study.

Literature Cited

Bittrich, V. 1993. Caryophyllaceae. Pp. 206-236 in K.
Kubitzki, J. G. Rohwer & V. Bittrich (editors). The Fam¬
ilies and Genera of Vascular Plants. II. Flowering
Plants, Dicotyledons, Magnoliid, Hamamelid and Car-
yophyllid Families. Springer- Verlag, Berlin.

Hartman, R. L. 1993. Caryophyllaceae (except Silene). Pp.
475—489, 493—497 in J. C. Hickman (editor). The Jep-
son Manual: Higher Plants of California. Univ. Califor¬
nia Press, Berkeley.

Hickman. J. C. 1993. Geographic subdivisions of Califor¬
nia. Pp. 37—46 in J. C. Hickman (editor). The Jepson
Manual: Higher Plants of California. Univ. California
Press, Berkeley.

Lee, Y. N. 1998. f lora of Korea, 3rd rev. ed. Kyo-Hak
Publishing, Seoul.

Little, E. L., Jr. 1971. Atlas of United States Trees, Vol.
1. Conifers and Important Hardwoods. U.S.D.A. Forest
Serv. Misc. Publ. I 146.

Mizushima, M. 1965. Critical studies on Japanese plants,

1 1: The genus Pseudostellaria Pax in Japan. Bull. Bot.
Surv. India 7: 62—72.

Rabeler, R. K. 1986. Revision of the Stellaria calycanlha
(Caryophyllaceae) Complex and Taxonomic Notes on the
Genus. Ph.D. Dissertation, Michigan State University,
East Lansing.

Weber, W. A. A R. Hartman. 1979. Pseudostellaria ja-
mesiana, comb, nov., a North American representative
of a Eurasian genus. Phytologia 44: 313-314.

Eriogonum diatomaceum (Polygonaceae: Eriogonoideae), a New
Species from Western Nevada, U.S.A.

James L. Reveal

Professor Emeritus, University of Maryland, College Park, Maryland 20742-5815, U.S.A.
Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, 11. S. A.

jrl9@umail.umd.edu

Joan Reynolds

Botanical Consultant, 4900 Grande Road, Reno, Nevada 89511, U.S.A.
JReynoldsBotany@aol.com

Jacquelyn Ricci a n i

3530 Deodar Street, Silver Springs, Nevada 89429, U.S.A.

ABSTRACT. A new species of’ Eriogonum subg. Eu-
cycla (Polygonaceae, Eriogonoideae), E. diatoma-
ceum , is described from llie Pine Nut Range. I, yon
County, Nevada, U.S.A. It is an edaphic endemic
species restricted to chalky, diatomaeeous outcrops
of the Coal Valley Formation in Churchill Narrows
south of Fort Churchill State Park. The new species
is outwardly similar to the Great Plains species E.
pauciflorum (sect. Capital a ), but is more closely re¬
lated to E. ochrocephalum and its relatives. Eriogon¬
um diatomaceum may be distinguished by its
sheathing, elliptic, densely gray-tomentose leaves,
erect scapose and white-tinged tomentose flowering
stems, capitate inflorescences, glabrous and creamy
white flowers, and glabrous achenes.

Key words: Eriogonoideae, Eriogonum , Nevada,
Polygonaceae, U.S.A.

Pieciani and Reynolds first found Eriogonum
diatomaceum (Polygonaceae Jussieu; Eriogono¬
ideae Arnott) in June 1997 near Churchill Narrows
in the Pine Nut Range of Lyon County, Nevada. The
material was identified subsequently as an undes¬
cribed species by Reveal during a workshop ses¬
sion on wild buckwheats sponsored by the Jepson
Herbarium in July 1997.

Eriogoiuini diatomaceum Reveal, J. Reynolds &
Pieciani, sp. nov. TYPE: U.S.A. Nevada: Lyon
Co.. 2.6 mi. (1.5 air mi.) SSW of Churchill (aban¬
doned historic site along the railroad), 0.6 air mi.
SW of Mill Canyon Creek crossing along the dirt
road into Churchill Narrows, on barren slopes
associated with Atriplex, T16N, R24E, sec. 21
SE.V4 of the NE‘/4 (N39°14'30", W119°17'15"),
4425 ft. elev., 29 June 2001, J. L Reveal 8306
(holotype, US; isotypes, BRY, GH. K. MARY,
MO. NY, RENO, UC. US, UTC). Figure 1.

Ah E. ochrocepkalo floribus alhis et ah h. anemophilo
foliis caulihus vaginantibus differt.

Low, matted, herbaceous perennials with a
branched, woody caudex forming a loosely
branched mat 0.5— 2.5 dm across, arising from a
stout woody taproot; leaves sheathing up the stems
for (1.5)2— 4 cm with the lower stems densely white-
tinged tomentose among the petiole bases and per¬
sistent leaves, the leaf blade elliptic1, (0.5)0. 8—
2(2.3) X (0. 3)0.5- 1 .2(1.8) cm. equally densely
grayish tomentose on both surfaces, the margins en¬
tire and plane, the apex broadly acute, the base
cuneate, the petiole slender, (0. 3)0.5— 1.5(1. 8) cm
long, tomentose, the petiole base elongate-triangu¬
lar to triangular, 3-5 X 2-4 mm, densely white-
tomentose abaxially, glabrous adaxially except for
a band of long hairs across the base: flowering stems
scapose, erect, (0.4)0.5-l .5(2) dm long, white-
tinged tomentose; inflorescences capitate, the* heads
1—1.5 cm across: bracts scale-like, 3 to 8, elongate-
triangular to triangular, 1—3 X 0.7— 1.2 mm. while
tomentose abaxially, green and essentially glabrous
adaxially except for the thinly tomentose margins
and sparsely tomentose apices, slightly acute ba-
sally; peduncles lacking; involucres congested, 5 to
10 per head, turbinate, rigid. (2.5)3— 4.5 X 2—3
mm, tomentose abaxially, glabrous adaxially, the 5
to 7 acute teeth 0.6—1 mm long, the bractlets linear.
3—4 mm long, minutely fringed with minute gland-
tipped cells, the pedicel 3-5 mm long, glabrous;
flowers numerous, creamy white with green-tan (in
anthesis) to red-tinged (in fruit) bases and midribs,
becoming rose-red in late fruit, (1.5)2. 5— 3 mm long,
glabrous except for minute glands along the midribs
adaxially, the tepals essentially similar, ovate-ob¬
long, those of the inner whorl slightly narrower and

Novon 12: 87-89. 2002.

88

Novon

Figure 1. Habit of Eriogonum diatomaceum Reveal, J. Reynolds & Picciani. Photograph by J. L. Reveal.

more distinctly oblong than those of the outer
whorl, united V3-V2 their length with the fused basal
portion distinctly pustulose in fruit; stamens ex-
serted, 3—3.5 mm long, the filaments sparsely pi¬
lose basally, the anthers cream-white to pale yellow,
0. 6-0.7 mm long, broadly oblong to ovate; achenes
light brown, 2-2.5 mm long, glabrous, the subglo-
bose base tapering to a long, 3-angled beak.

Etymology. From diatoma INI, (New Latin), di¬
atom, and by extension diatomacea NL for diato-
maceous earth, as to the substrate on which the
plant is restricted. Roth words are derived from the
Greek diatomos, cut in half.

Eriogonum diatomaceum is known from a single
population on low bills south and southwest of
Churchill, Lyon Co., Nevada. This single popula¬
tion encompasses 15 more or less distinct occur¬
rences restricted to the northeastern portion of the
Pine Nut Range. The population is scattered over
approximately three square miles of Bureau of Land
Management lands. Less than 50.000 individual
plants are known (Reynolds, 2001).

The new species, informally called Churchill
Narrows buckwheat, is restricted to diatomaceous
soils of the Coal Valley Formation on relatively un¬
disturbed dry barren exposed knolls and drainages
on all aspects between 4300 and 4560 feet in el¬
evation. The habitat supports a sparse vegetation

cover dominated by Churchill Narrows buckwheat
in association with shadscale (Atriplex confertifolia
(Torrey & Fremont) S. Watson), princes plume
(Stanleya pinnata Nuttall var. pinnata ), Bailey’s
greasewood (Sarcobates baileyi Coville), and bud-
sage (Picrothamnus desertorum Nuttall) anti other
species such as Kochia americana S. Watson, Te-
tradymia glabrata Torrey & A. Gray, Psathyrotes
annua (Nuttall) A. Gray, Eriogonum defiexum Tor¬
rey var. nevadense Reveal and E. lemmonii S. Wat¬
son.

The Coal Valley Formation is a fluviolacustrine
deposit of soft lake beds, sandstone, conglomerate,
and sedimentary breccia. Andesite and andesitic
sediments are commonly imbedded. The formation
contains a unique fossil flora of aquatic plants and
mammalian remains (Berry. 1927). The deposit is
part of the series of Tertiary diatomites that filled
the subsidiary basins within the Basin anil Range
extensional tetonic system of Nevada. The known
aerial extent of outcrop where the Churchill Nar¬
rows buckwheat is found includes approximately
150 acres of variable thickness (8 to 30 feet) (Car¬
penter, 1997). The volcanic sedimentary rock as¬
sociated with this formation is generally described
as late Miocene to early Pliocene in age. and con¬
sists of stream and lake deposits of silty shale, di¬
atomaceous shale, pebbly conglomerate, siltstone,

Volume 12, Number 1
2002

Reveal et al.

Eriogonum diatomaceum from Nevada

89

and sandstone. Pile major components are fossil di¬
atoms (amorphous silica), calcium montmorillonite,
feldspar, anti gypsum.

The new species was found during an impact
survey for the Bureau of Land Management prior
to granting a mining lease. II mining were permit¬
ted, Eriogonum diatomaceum instantly would he
entlangered. Even if the area is not mined, current
off-road vehicle activity in the region is sufficient
to threaten the long-term survival of the species.
Current consultation between the Bureau of Land
Management and the State of Nevada (which al¬
ready considers the taxon to be a “sensitive plant")
may mean that a formal proposal for federal pro¬
tection may not be necessary. Nonetheless, moni¬
toring will be necessary to assure the species’ long¬
term survival.

Eriogonum diatomaceum is a member of the sub¬
genus Eucycla (Nu ttal 1) Kuntze. The new species is
outwardly similar to the Great Plains species, E.
pauciflorum Pursh, and especially the densely to-
mentose, more compact phase, variety gnaphaloides
(Bentham) Reveal of southeastern Wyoming and ad¬
jacent western Nebraska. The similarity of the
sheathing leaves is particularly striking. These
leaves, plus the long scapose, tomentose flowering
stems, capitate inflorescences, and creamy white
flowers add to the impression. Nonetheless, E. dia¬
tomaceum is more closely allietl to E. ochrocephal-
um S. Watson and its numerous relatives common
to the Great Basin of Nevada and adjacent states.
Also outwardly similar to the new' species in gen¬
eral appearance are E. loganum A. Nelson, a nar¬
row endemic of northern Utah, and E. brandegei
Rydberg of Colorado. Both belong to the E. brevi-
caule Nuttall complex and as such are unrelated.

The newr species is allied to those species of the
Eriogonum ochrocephalum complex that may he
characterized as matted perennials with scapose
stems and capitate inflorescences, and specifically
with those that have a rigid, usually turbinate in-
volucral tube. The majority of these eight species
have bright yellow flowers and generally glabrous
and/or glandular flowering stems. Of the various en¬
tities found in western Nevada, E. ochrocephalum
var. alexandrae Reveal has tomentose stems, and
both E. anemophilum Greene and E. tiehmii Reveal
have creamy white flowers. None of these species
have sheathing leaves. The long scapose flowering
stem of E. diatomaceum rivals the lengths observed

in E. ochrocephalum var. sceptrum Reveal of south¬
western Idaho, and the two share the long, narrow,
turbinate involucre. From all of these species, the
new' species also may be readily distinguished by
the dense, essentially lanate tomentum of the
leaves, which is evenly distributed on both surfac¬
es. The remaining 12 species of the E. ochroce¬
phalum complex have membranaceous rather than
rigid involucres.

In Reveal’s (1985) key to Nevada wild buck¬
wheats, the new species may he keyed as follows:

20(16). Involucres 2—2.5 mm long, floccose without:

leaf blades broadly elliptic to obovate or
suborbicular; flowers glabrous; Eureka,
Lander, Pershing, and southwestern Hum¬
boldt Cos . 14. E. anemophilum

Involucres (2.5)3— 4.5 mm long, densely to¬
mentose or sparsely floccose without; leaf
blades elliptic to oblong; flowers glabrous or

sparsely glandular . 20a

20a(20). Flowers glabrous; leaves sheathing up the
stems (1.5)2— 4 cm; involucres turbinate, 2—

3 mm wide, the teeth 0.6—1 mm long; Lyon

Co . 14a. E. diatomaceum

Flowers sparsely glandular; leaves strictly
basal; involucres turbinate-campanulate, 3—

4 mm wide, I he teeth 1.5—2 mm long; Es¬
meralda Co . 15. E. tiehmii

Paratypes. U.S.A. Nevada: Lyon County, S end of
Churchill Narrows, 1.5 air mi. S of Churchill, TI6N,
R24E, sec. 22, 4275 It. elev., Picciani I (K. MARY, NY,
RENO, UC, US); 2 air mi. W of Churchill Narrows, 2 air
mi. SSW of Churchill, T16N, R24E, see. 21, 4480 ft.
elev., Picciani 2 (BRY, GH, K, MARY, MO, NY, RENO,
UC, US).

Acknowledgments. We thank James Morefield
of the Nevada Natural Heritage Program, Depart¬
ment of Conservation and Natural Resources, in
Carson City, for his support and encouragement.

Literature Cited

Berry, E. W. 1927. The flora of Esmeralda Formation in
western Nevada. Proc. U.S. Natl. Mus. 72(23): 1-15.
Carpenter, A. 1997. The Ecology of Churchill Narrows
Buckwheat, Physiography & Geology. Unpublished Re¬
port .

Reveal, J. E. 1985. An annotated key to Eriogonum (Po-
lygonaceae) of Nevada. Great Basin Naturalist 45: 493—
519.

Reynolds, J. 2001. Current knowledge and conservation
status of Eriogonum diatomaceum Reveal. J. Reynolds
& Picciani (Polygonaceae: Eriogonoideae), Churchill
Narrows buckwheat. Status report prepared for U.S. I).
I. Bureau of Land Management, Nevada State Office
and Nevada Natural Heritage Program, Department of
Conservation and Natural Resources, Carson City.

A New Species of Oxalis (Oxalidaceae) from El Salvador

Karen Sidwell and Sandra Knapp

Department of Botany, The Natural History Museum, Cromwell Hoad, London, SW7 5BD,

United Kingdom

ABSTRACT. Oxalis salvadorensis, a new species
from the mixed lowland forests of western El Sal¬
vador. is described. An illustration is provided, and
the differences between this and other morpholog¬
ically similar Mesoamerican species ol Oxalis are
discussed.

Key words: El Salvador, Mesoamerica, Oxali¬
daceae, Oxalis.

The genus Oxalis L. consists of around 500—800
species, mainly herbs, often with underground
bulbs or tubers, distributed worldwide. Current es¬
timates of the number of species of Oxalis cover a
large range and highlight the need for research on
species delimitation and relationships in this group.
In his comprehensive treatment of the family Ox¬
alidaceae, Knuth (1930) recognized 7 genera in the
family, of which Oxalis was by far the largest with
791 species. In his subgeneric classification, Knuth
divided Oxalis into 37 sections, based principally
on habit and leaf characters; his treatment is the
only worldwide subgeneric classification of the ge¬
nus. Since publication ol Knuth’s work, research on
Oxalis has largely consisted ol traditional taxonom¬
ic revisions ol Knuth’s sections (for example, Eiten,
1963; Denton, 1973; Lourteig, 1975, 1979). More
recently, Lourteig (1994, 1995, 2000) has revised
the American species of the genus and has provid¬
ed a comprehensive new subgeneric and sectional
classification.

While preparing a treatment of Oxalidaceae for
Flora Mesoamericana, a new species, herein de¬
scribed as Oxalis salvadorensis, was found.

Oxalis salvadorensis Sidwell. sp. nov. TV PE: El
Salvador. Ahuachapan: P.N. El Imposible, San
Benito al N del pie del bajadero de los esco-
bos, 13°49'N, 89°56'W, 19 June 1992. F. San-
doval & F. Chinchilla 453 (holotype, LAGU;
isotypes, B, E, MO). Figure 1.

Herba aeaulis seaposa bulbo squamoso; foliola termin-
alia symmetrica obcordata sinu non profundo; foliola la-
teralia asymmetrica basiscopo reducto; llores albi; semina
lateraliter longitudinaliterque pocata.

Stemless herb, 25—50 cm; bulbs ca. 20 X 20

mm; bulb scales 2-15 X 2.4—22 mm. linear to nar¬
rowly triangular, with 1—3 veins converging just
short ol apex, drying reddish brown, membranous
al edges, the margins ciliate. Leaves basal. 1-5 per
plant, palmate; petioles 8—32 cm, sparsely pubes¬
cent with simple, slender, transparent trichomes 2—
5 mm long; leaflets 3, unequal, glabrous, or sparse¬
ly pubescent with slender trichomes like those of
the petiole, the lamina minutely pitted, especially
below; terminal leaflet 30— 53(— 80) X 32— 48(— 64)
mm, obeordate, symmetrical, the base acute, the
apex with rounded lobes 21— 35(— 66) mm apart, and
a shallow sinus 2—5 mm; lateral leaflets 23— 35(—
62) X 30— 40(— 100) mm, obeordate, asymmetrical,
more elongate aeroseopieally, the sinus 1—3 mm.
Inflorescences umbelliform, with 4—11 flowers; pe¬
duncles 11—27 cm; bracts 1.5— 2.5 X 0.4— 0.6 mm,
triangular to narrowly triangular, the apex acute,
with scattered oxalate deposits. Pedicels 8—1 2(— 14)
mm, glabrous; sepals to 2.5—3 X 0.7—1 mm, linear,
the apex obtuse to slightly rounded, with oxalate
deposits appearing as scattered orange dots, the
margins sparsely ciliate with eiliae ca. 0.5 mm
long; petals 8—11 X 3.5—4 mm, white; filaments in
two whorls, one whorl ca. 2.5 mm, the other 4—5.5
mm; anthers globose, ca. 0.75 mm long; styles 5,
0.5— 0.8 mm or 2.5—3 mm, pubescent, the plants
distylous or semi-homostylous(?). Capsules 8—10 X
2—3 mm, ovoid, glabrous, pubescent at apex and
on styles; seeds ca. 6 per locule, the surfaces ap¬
pearing pitted due to the presence ol both longi¬
tudinal and lateral ridges.

Distribution. Known only from mixed lowland
forest in the El Imposible National Park in western
El Salvador at ca. 700 m.

The species is easily distinguished from other
stemless Oxalis species in Mesoamerica by die
large, clearly asymmetric lateral leaflets with
rounded lobes. Oxalis salvadorensis is morphologi¬
cally similar to several other Mesoamerican spe¬
cies: 0. gregaria (Rose) Knuth (W Mexico), O. dirn-
idiata Donnell Smith (Guatemala), O. debilis Kunth
(Guatemala to Peru), and the widespread and var¬
iable O. latifolia Kunth s.l. (see Table 1 ).

Heterostyly is important taxonomically in Oxalis,

Novon 12: 90-93. 2002.

Volume 12, Number 1
2002

Sidwell & Knapp
Mesoamerican Oxalis

91

ISOTYPE

Oxalis salvadorensis Sidwell
Karen Sidwell BM July 2000

Oxalidaceac

Oxalis debit is Kunth

Dupl del W Burger (F) 1995

EL IMPOSIBLE (EL SALVADOR)

Oxalidaccac

Oxalis cf. debilix var. corymbosa (Dc.) Ixuirt.

Depto Ahu»chap4n San Benito, al N del pie del bajadcro de k* cscoboc.

25cm dc largo, rosados, hojuela terminal hast a 6cm dc anebo y 6
largo. Pcduneulo hasfa 28cm dc largo. Flores biancas. Plants no
cootun. [ES/aaJ No. Herb. LAGU: ES-00453
Del. W Bcrendsohn 10/92
Herbaria: MO B LAGU ['

Lc&: KSandirvul 453 con Chinchilla 19.6.1992

M B

F igure 1. Oxalis salvadorensis K. Sidwell. I ’holograph of the isotype at B.

Sectional membership according to Section Oxalis subsect. Dimidiatae Section lonoxalis Section lonoxalis Section lonoxalis

Lourteig (2000) (knuth) Lourteig (Small) Knuth (Small) Knuth (Small) Knuth

Elevation 700 m 1200-1450 m 20-1900 m, mossy 0-1400 m 800-3050 m

92

Novon

C — bfc
cd C

b£ rrt

— G

3 £ * a.

cd co bD

cd

a;

u.

a?

but it can be difficult to distinguish flower forms on
herbarium sheets. Flowers are generally tristylous,
distylous, or homostylous; all of these forms, except
homostylous, have whorls of stamens of different
lengths (see Denton, 1973). The flowers of 0. sal-
vadorensis appear to be distylous or semi-homos-
tylous (sensu Denton, 1973, where one but not both
of the stamen whorls equals the stigma length), but
further observations with more specimens and more
complete material are needed.

Based on the scapose habit and scaly bulbs, Ox¬
alis salvadorensis belongs in Oxalis sect, lonoxalis
(Small) Knuth. In the recent monograph of Oxalis
subg. Oxalis (Lourteig, 2000), 0. salvadorensis
would key out as belonging to section lonoxalis,
and within that section would key near to 0. cua-
trescasasii Lourteig from western Colombia (couplet
“Fob anchamente obcordados o subtriangulares,
enteros”; Lourteig, 2000: 534). In the recently com¬
pleted treatment of Oxalidaceae for Flora de Nic¬
aragua (Sullivan, 2001), where 0. salvadorensis
might be expected to occur, it would key out in
couplet 1 with 0. latifolia and 0. debilis, also mem¬
bers of section lonoxalis (see Table I). Denton
(1973: 488) proposed a putative phylogenetic
scheme of relationships for North American taxa in
section lonoxalis, in which 0. salvadorensis might
be placed close to her O. gregaria-O. alpina (Rose)
Knuth group. A great deal more taxonomic and
phylogenetic investigation is clearly required in
these interesting and relatively poorly collected
herbs.

Paratypes. EL SALVADOR. Ahuachapan: 1 \ . El
Imposible, San Benito, 700 m, 13°49'N, 89°56'W, 9 July
1993, E. Sandoval & F. Perez 1344 (B, E, LAGU, MO),
28 May 1994, F. Chinchilla & ./. M. Chinchilla s.n. (B),
23 June 1994, F Chinchilla s.n. (MO).

Acknowledgments. We thank the herbaria cited
in the text for permission to borrow or examine their
specimens; Norman Robson for providing the Latin
description; and Pat Hart of the Photographic Unit
in the NHIVI for the preparation of Figure 1.

Literature Cited

Burger, W. 1991. Oxalidaceae. In Flora Costaricensis.

Fieldiana, Bot. n.s. 28: 2—16.

Denton, M. 1973. A monograph of Oxalis section lonoxalis
(Oxalidaceae) in North America. Michigan State Univ.
Publ. Biol. Ser. 4: 456-615.

Eiten, G. 1963. Taxonomy and regional variation of Ox¬
alis sect. Corniculatae 1 . Introduction, keys and syn¬
opsis of the species. Amer. Midi. Naturalist 69: 257-
309.

Knuth, R. 1930. Oxalidaceae. In Das Pflanzenreich 4:

130. Wilhelm Engelmann, Leipzig.

Lourteig, A. 1975. Oxalidaceae Extra- Austroainericanae I.

Volume 12, Number 1
2002

Sidwell & Knapp
Mesoamerican Oxalis

93

Oxalis L. sectio Thamnoxys Planchon. Phytologia 29:
449-471 .

- . 1979. Oxalidaceae Extra-Austroamericanae II.

Oxalis L. sectio Corniculatae DC. Phytologia 42: 57—
198.

- . 1994. Oxalis L. subgenero Thamnoxys (Emil.)

Reiche emend. Lourt. Bradea 7: 1-199.

- . 1995. Oxalis L. subgenus Trifidus Lourt. subgen.

nov. Bradea 6: 389—392.

- . 2000. Oxalis L. subgenero Manaxalis (Small)

Lourt., Oxalis y Trifidus Lourt. Bradea 7: 201—629.
Sullivan, G. 2001. Oxalidaceae. In W. 1). Stevens. C. Ulloa
U., A. Pool & 0. M. Montiel (editors). Flora de Nica¬
ragua 2: 1861-1864.

Conservation and Ecology of Lilium pyrophilum , a New Species of
Liliaceae from the Sandhills Region of the Carolinas
and Virginia, U.S.A.

Mark W. Skinner

USDA-NRCS, National Plant Data Center. P.0. Box 74490, Baton Rouge, I Louisiana 70874,

U.S.A. mskinner@po.nrcs.usda.gov

Bruce A. Sorrie

3076 Niagara-Carthage Road, Whispering Pines, North Carolina 28327, U.S.A.

1 )sorri e@eart hlink.net

Abstract. Lilium pyrophilum M. W. Skinner &
Sorrie is a new taxon from the Sandhills region of
the Atlantic Coastal Plain, where it is known from
16 counties in the Carolinas and adjacent Virginia.
It occurs within the longleaf pine ecosystem in
sandy, wet to dry ecotonal situations such as
streamheads, seeps, swampy streams, and wet,
maintained powerlines. Lilium pyrophilum is most
closely allied to L. superhum but is allopatrically
distributed and confined to a restricted habitat.
Compared to L. superhum it blooms later, is smaller,
has fewer and smaller flowers with slightly longer
tubes, and it has shorter and relatively broader
leaves that are ascending and concentrated in fewer
whorls toward the bottom of the stem. Frequent fires
are essential lor habitat maintenance in natural set¬
tings, and thus the Latin epithet pyrophilum (= fire
loving) is used for the new' taxon. We suggest the
common name Sandhills lily for this rare lily, and
urge its fullest protection.

Key words: fire dependence, Liliaceae, Lilium .
long-leaf pine ecosystem. North Carolina, South
Carolina, U.S.A., Virginia.

For some time botanists in the Sandhills region
have known of the existence of an unusual and per¬
haps undescribed true lily. In their flora of the Car¬
olinas, Radford et al. (1968) mentioned robust Lil¬
ium michauxii Poiret plants in bogs that resembled
Lilium superhum L. It is these plants, previously
masquerading in herbaria under both those names,
that we describe here as L. pyrophilum.

Since its recognition as distinctive in the mid
1970s (Russo, 1997). the taxonomic identity of this
new lily has been debated. As Radford et al. (1968)
suggested, it is similar to the other pendent-flow-
ere d lilies in the region, L. michauxii and L super-
hum. However, regional botanists recognized that

the turks-cap lily (L. superhum ) per se does not
occur in the lower part of the Piedmont, which is
the hilly region between the Appalachian Moun¬
tains and the inner Atlantic Coastal Plain of which
the Sandhills are a part. The Carolina lily (L. mi¬
chauxii) is quite recognizable by its compact bulb,
striking obovate leaves, fragrant flowers, and drier
habitat. Speculation therefore centered on a link
between the unknown lily and L. iridollae M. G.
Henry, a rare endemic lily from the Panhandle re¬
gion of Florida and adjacent Alabama (Henry,
1946). Indeed, natural heritage programs recorded
Carolina populations of L. pyrophilum as L. iridol¬
lae (e.g.. North Carolina Natural Heritage Program.
1999), and early status reports commissioned be¬
cause of L. pyrophilum s rarity assumed the Caro¬
lina Sandhills lily populations represented a sig¬
nificant range extension for L. iridollae. This
suggestion was well founded because both overall
morphology and the baygall ( Magnolia , Ilex). Sar-
racenia bog, and streamhead habitats of these two
species are closely similar. Others (Moretz & Smith,
1995) suggested that the unknown lily had been
previously described from Mississippi as /,. gaza-
ruhrum M. k. Roane & J. N. Henry (1980). How¬
ever, we have examined lilies from very near the
type locality of L. gazaruhrum in Choctaw County,
Mississippi, and have also seen the type, and like
Adams (1981), we conclude that L. gazaruhrum can
be accommodated within L. superhum. Extended
field and herbarium study and unpublished mor¬
phometric analysis ultimately led us to determine
that the lily described here is distinctive and new.

Liliiim pyrophilum M. W. Skinner & Sorrie, sp.

nov. TV PE: U.S.A. North Carolina: Moore Co..

Highway 73 near Pinehurst, 149 m. 1 Aug.

1995 (fl), M. W. Skinner 272 (holotype, GH;

isotype, MO). Figures I, 2.

Novon 12: 94-105. 2002.

Volume 12, Number 1
2002

Skinner & Sorrie

Conservation of Lilium pyrophilum

95

Figure 1. Lilium pyrophilum M. W. Skinner & Sorrie. — A. Habit with bulb and flowers, based on the type collection
(M. W. Skinner 272) from Moore County, North Carolina. — B. Capsule before opening, from a B. A. Sorrie photograph
taken at the type locality.

96

Novon

Figure 2. Lilium pyrophilum, photographs A— I) by M. W. Skinner, E by It. A. Sorrie. — A. Hal)it of the type specimen
(M. W. Skinner 272). — It. Habit. — C. Rhizome, the next year's bulb at left (M. If! Skinner 267). — I). Flower. — E.
Fruiting stalk from the type locality. A, E from Moore County, North Carolina, R— I) from Ft. Jackson, Richland County,
South Carolina.

Volume 12, Number 1
2002

Skinner & Sorrie

Conservation of Lilium pyrophilum

97

Haec species Lilio superbo L. arete affinis, sed ab eo
anthesi serotina, liabitu multo minore, foliis brevioribus
atque tubo ttorali longiore distinguitur; locos perculationis
ortusque rivulomm saepe incensos habitat.

Bulb a slowly growing scaly rhizome, horizontal
and elongate, 2. 4-2. 8 X 5. 2-8. 6 cm, 0.3-0. 5 times
taller than long, lacking scales between the 2—3
bulb units that represent annual growth, the young
end sometimes dichotomously branched at 120°
from main axis; scales (modified leaves) numerous,
fleshy and starchy, unsegmented, longest 1. 1-1.9
cm, white; roots on each bulb mostly contractile
and thus thick (to 3 mm) and often concentrically
wrinkled, a few thinner and fibrous. Stems to 1.6
m, erect. ± glabrous, green, adventitious roots
(stem roots) above the bulb sometimes present.
Leaves numerous, simple, exstipulate and sessile,
the lower leaves scarcely ascending or ascending,
drooping at tip or not, upper ascending and ap-
pressed. (scattered or) in 0-12 ± proximal whorls
of 3 or more leaves, 3—1 1 (—15) per whorl; blade
narrowly elliptic, 2.3— 10.3(— 12.2) X 0.8— 2.4 cm,
1.6— 7.6(— 10.3) times longer than broad, apex acute
(scarcely acuminate on upper stem), margin entire,
usually glabrous, not undulate, green and somewhat
lighter below, the major veins 3, these glabrous be¬
low. Inflorescences terminal, racemose (umbellate in
small plants) and open, maturing acropetally, 1-7-
flowered; bracts generally 1-2 per flower, often one
broadly lanceolate and basal to the pedicel, the oth¬
er narrowly lanceolate, adaxial, and attached near
the middle of the pedicel; pedicels 6.8-16.5 cm.
Flowers bisexual, pendent, shaped as a “turk’s-
cap.” radial (or slightly vertically bilateral as flower
matures), not fragrant; tepals free, re flexed V5—V4
along length from hase, red-orange or dusky red
(magenta or pinkish, pale orange, red) apically to
(pale) yellow (yellow-orange) centers to visibly
green basallv, magenta spots concentrated in prox¬
imal %— %, sepals 3, ± lanceolate and narrowed at
the hase, 6.7-8. 9 X 1.1-1. 7 cm, glabrous, each
with an adaxial nectar-bearing surface basally and
with two sharp parallel longitudinal median ridges
abaxially, apex usually acute; petals 3 and similar
to the sepals, 6.3-8. 7 X 1 .5-2.2 cm, apex acute,
the hasal nectaries less extensive than those on the
sepals, with two rounded longitudinal median ridg¬
es adaxially; stamens 6, free, opposite the perianth
parts, quite exserted; filaments parallel at first then
widely spreading at 12—28°, 4.5— 5.9 cm; usually
pale green; anthers versatile, oblong, 1.1-1. 8 cm
when fresh, magenta (purple), becoming darker;
pollen rust-colored, becoming lighter; pistil 1, com¬
pound, 3-lobed with 3 chambers, oblong, 3.4— 6.4
cm; ovary superior, 1.5— 2.8 cm; axile placentae 6;

ovules as many as seeds, a few developing without
embryos; style 1, round in cross section, initially
parallel to flower axis but generally elongating and
curving toward periphery, pale green and often
spotted purple; stigma 3-lobed. Capsules loculici-
dal, the valves 3, erect, ± oblong-obovate, with
constricted hase, not strongly winged, 241-4.7 X
(1.3— )1. 5—1.9 cm, 1.7-2. 8 times longer than broad,
green maturing brown. Seeds many, in 6 ranks,
shape a flat 60° wedge, surface verrucose, light
brown with darker ovate embryo in center.

Etymology. The epithet is from the Latin for
“fire loving,” and is employed here because of the
very frequent fires that are essential to maintain
proper habitats for L. pyrophilum. Because the spe¬
cies is restricted to the Sandhills of the Carolinas
and adjacent Virginia we propose the vernacular
name of Sandhills lily.

Phenology. Flowers in summer (late July-mid
August); capsules mature in late October.

Distribution. The known range of Lilium pyro¬
philum extends from southeastern Virginia to south-
central South Carolina, wholly within the Atlantic
Coastal Plain. The great majority of populations oc¬
cur within the Sandhills region, an area of rolling
topography dissected by abundant blackwater
streams. Although this physiographic region ex¬
tends from the Carolinas to central Alabama, in
terms of floristics and phytogeography the core area
extends from Johnston County in east-central North
Carolina to Richmond County in extreme east-cen¬
tral Georgia (Sorrie & Weakley, 2001). We have
documented the lily from ten counties in North Car¬
olina, four in South Carolina, and two in Virginia
(Fig. 3), at elevations ranging from 25 to 150 m.

The Sandhills region is a significant area of spe-
ciation within the North American coastal plain
(Sorrie & Weakley, 2001), and many endemics have
been recognized there. The following are distrib¬
uted exclusively within the core Sandhills region or
have the majority of their populations there: As-
tragalus michauxii (Kuntze) F. J. Hermann, Liatris
cokeri Pyne & Stueky, Lycopus cokeri Allies ex Sor¬
rie, Physalis lanceolata Michaux, Pityopsis pinifolia
(Elliott) Nuttall, Pyxidanthera barbulata Michaux
var. brevifolia (Wells) Allies, Stylisma pickeringii
(Torrey ex M. A. Curtis) A. Gray var. pickeringii,
Vaccinium crassifolium Andreanszky subsp. sem-
pervirens (Anderson & Rayner) Kirkman & Balling-
ton. Lycopus cokeri is a very frequent associate of
Lilium pyrophilum. whereas Liatris cokeri, Physalis
lanceolata, and Pityopsis pinifolia occur on adja¬
cent dry to xeric slopes.

Habitat: the physical setting. The Sandhills re-

98

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Figure 3. Lilium fryrophilum distribution in Virginia, North Carolina, and South Carolina. Solid circles represent
counties with extant populations documented since 1990, open circles counties where the species is historical (pre-
1990) and presumed extirpated.

gion (also known as the Fall-line Sandhills) consti¬
tutes the innermost portion of the coastal plain and
abuts the Piedmont physiographic province. The
fall-line is a distinct geological boundary separating
the younger Cretaceous and Tertiary age sediments
to the east and the older Paleozoic formations to
the west (Fenneman, 1938). Parent materials of the
Sandhills are unconsolidated to partly consolidated
sands, gravels, and clays.

Compared with the rest of the coastal plain, to¬
pography in the Sandhills is highly varied, with el¬
evations that range from 30 to 180 m above sea
level. The term “rolling hills” is apt; although the
region is highly dissected, there are few sharp geo-
morphic features. Rock outcrops are scarce. Drain¬
age creeks are abundant; most lie 10-30 m below
the surrounding hilltops and small plateaus, where¬
as larger streams and rivers lie significantly lower.
A number of large rivers traverse the Sandhills from
their origins in the Piedmont or mountains. All
drainages that originate in the Sandhills are dark
tea in color due to high tannin content and are
termed blackwater streams.

Upland soils in the Sandhills are acidic well-
drained sands and loamy sands; subsoils may be
sandy throughout, or have a clayey or loamy layer.

Soils of streamheads and creeks are finer loams and
loamy sands; those along rivers contain consider¬
able organic matter.

Habitat: plant communities. Lilium pyrophilum
occurs almost exclusively in ecotonal situations
within the longleaf pine (Finns palustris Miller)
ecosystem. It inhabits narrow transition zones —
usually 2 to 10 m — between dry longleaf pine-oak-
wiregrass uplands and wet, wooded creeks and
streamheads. Upland communities (usually longleaf
pine-oak-w iregrass with scattered low shrubs)
quickly give way to a shrub-cane-fern-herb eco-
tone, in turn replaced by a tree-shrub-sphagnum
streamhead with flowing water. Due to the hilly to¬
pography of the Sandhills region and myriad drain¬
age creeks, ecotonal habitats once were abundant
across the landscape, but widespread fire suppres¬
sion and conversion of streamheads to fishing
ponds, farm impoundments, swimming ponds, and
water reservoirs have severely reduced these plant
communities.

Ecologists usually ally ecotonal communities of
the Sandhills region with the adjacent wetland com¬
munities based on the substantial percentage of
shared species versus the much smaller percentage
of species in common w ith adjacent uplands (Schaf-

Volume 12, Number 1
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Skinner & Sorrie

Conservation of Lilium pyrophilum

99

ale & Weakley, 1990). Indeed, there is a mix of
upland and wetland plants, but Sandhills ecotones
support a significant number of species that do not
normally occur in either “parent" community. In
fact, some of these species such as pine barren
reedgrass ( Calamovilfa brevipilis (Torrey) Scribner)
and savanna cow bane (Oxypolis ternata (Nuttall)
Heller) occur in the Sandhills only in ecotones and
related seepage habitats. Phytogeographically,
these plants may be considered a subset of those
that are characteristic of wet savannas and flat-
woods of the outer coastal plain of the Carolinas,
particularly the colorful pitcherplants ( Sarracenia ).
meadow-beauties ( Rhexia ), seedboxes (. Ludwigia ),
milkworts (Polygala), orchids, sundews (. Drosera ),
and yellow-eyed-grasses ( Xyris ).

Lilium pyrophilum is most closely associated
with three distinctive wetland communities, as fol¬
lows:

1. Streamhead Pocosin. This community oc¬
curs along headwaters of creeks and stream
branches where seepage water from adjacent eco¬
tones forms definite rivulets. It extends downstream
as long as such seepage is important to plants rel¬
ative to stream flooding. The term “pocosin" is used
because the usually dense evergreen shrubs and
sparse canopy are reminiscent of true pocosins that
develop on peat that accumulates on flatter and
younger portions of the coastal plain (Schafale &
Weakley, 1990; Sharitz & Gibbons, 1982). Typical
shrubs and vines are titi ( Cyrilla racemiflora L.),
fetterbush (Lyonia lucida (Lamarck) K. Koch), ink-
berry (Ilex glabra (L.) A. Gray and /. coriacea
(Pursh) Chapman), highbush blueberry ( Vaccinium
formosum Andreanszky and V. fuscatum Aiton).
sweet pepperbush (Clethra alnifolia L.), evergreen
bayberry (Myrica heterophylla Rafinesque), redhay
(Persea palustris (Rafinesque) Sargent), poison su¬
mac (Toxicodendron vernix (L.) Kuntze), and blas¬
pheme vine (, Smilax laurifolia L.). Trees may be
sparse or fairlv dense, with tuliptree (Liriodendron
tulipifera L.), pond pine (Pinus serotina Michaux),
swamp black gum (Nyssa biflora Walter), red maple
(Acer rubrum L.), sweetbay (Magnolia virginiana
L.), and occasionally Atlantic white cedar (Cha-
maecyparis thyoides (L.) Britton, Stems & Poggen-
burg). Sphagnum moss is usually abundant and
cane (Arundinaria tecta (Walter) Muhlenberg) fre¬
quent to common.

Streamhead Pocosin ecotones are normally dom¬
inated by low shrubs and cane down-slope and
herbs and graminoids up-slope, but complex pat¬
terns may be produced depending on soil moisture
and disturbance. Woody plants are kept short by
recurring fires (or mowing in powerlines). Promi¬

nent shrubs are dangleberry (Gaylussacia frondosa
(L.) Torrey & A. Gray ex Torrey), Clethra alnifolia ,
Ilex glabra, and I. coriacea, maleberry (Lyonia li-
gustrina (L.) DC.), swamp azalea (Rhododendron
viscosum (L.) Torrey), blueberries (Vaccinium cras-
sifolium and V. tenellum Aiton), dwarf witchalder
(Fothergilla gardenii L.), and honeycups (. Zenobia
pulverulenta (Bartram ex Willdenow) Pollard). Im¬
portant grasses include Ctenium aromaticum (Wal¬
ter) Wood, Muhlenbergia expansa (Poirel) Trinius,
Aristida virgata Trinius, A. stricta Michaux, l)i-
chanthelium spp., Panicum virgatum L. var. cub-
ense Grisebach, Andropogon glomeratus (Walter)
Britton, Sterns & Poggenburg, Calamagrostis coarc-
tata (Torrey) Eaton, and Calamovilfa brevipilis.
Conspicuous sedges include Rhynchospora spp. (up
to a dozen taxa), Carex glaucescens Elliott, C. tur-
gescens Torrey, Eleocharis tuberculosa (Michaux)
Roemer & Schultes, E. tortilis (Link) Schultes, and
occasionally Eriophorum virginicum L. Other fre¬
quent monocots include Juncus trigonocarpus Steu-
del, Eriocaulon decangulare L., and Lachnocaulon
anceps (Walter) Morong. The majority of Lilium pyr¬
ophilum populations occur in the mid to lower por¬
tions of this ecotonal habitat type, and only rarely
in the Streamhead Pocosin proper.

La. Canebrake Variant. Where burned very
frequently, Streamhead Pocosins support fewer
trees and shrubs. These are replaced by dense
stands of Arundinaria tecta called canebrakes. Spe¬
cies diversity is low relative to normal streamhead
communities, although ectones remain diverse.
Only small populations (1 to 5 plants) of Lilium
pyrophilum occur in this habitat type.

2. Sandhill Seep. This community occurs on
slopes where the clay layer abruptly forces water
to the surface, usually where there is a sudden in¬
crease in the angle of the slope. Sandhill Seeps
occur within the upland pine-oak -wiregrass com¬
munity as a patch of shrubs, ferns, and cane that
form a “rim” across the slope. They may be con¬
nected to Streamhead Pocosins and Small Stream
Swamps, and may have perennially active seepage
or be merely moist. Once numerous in the Sandhills
region, high quality examples are now rare due to
fire-suppression. Lilium pyrophilum appears to be
rare in this habitat.

3. Coastal Plain Small Stream Swamp. This
community occurs downstream from Streamhead
Pocosins where water volume is great enough to
cause frequent flooding following rainstorms. It is
dominant along the major blackwater streams
through the Sandhills region. Canopy dominants are
Nyssa biflora, loblolly pine (Pinus taeda L.), Acer
rubrum, willow oak (Quercus phellos L.), water oak

100

No von

Table 1. Summary of protec

lion of known L. pyrophilum populations.

Ownership

Type of protection

Populations

Individuals

Private

none

3

16

National wildlife refuge

passive

1

2 4-

Military reservation

Camp Maekall

passive

2

2

Fort Bragg

passive

21

63

Fort Jackson

passive

1

~

State forest

passive

1

1 +

State game land

passive

5

15

Private with utility easement

management agreement

6

K0

Slate park

active

1

7

Private conservation

active

1

42

3

205+

(Q. nigra L.), sweetgum (Liquidambar styraciflua
L.), and scattered pond cypress (Taxodium ascen-
dercs Brongniart). Subcanopy trees include Ameri¬
can holly ( Hex apnea Aiton), Cyrilla racemifiora,
Persea palustris, and Magnolia virginiana. Shrubs
and vines vary from sparse to dense and include
doghobble ( Ixnicothoe axillaris (Lamarck) I). Don),
blackberry ( Riibus argutus Link), Smilax laurifolia,
greenbrier (Smilax rotundifolia L.), sawbrier (Smi¬
lax glauca Walter), poison ivy (Toxicodendron rad-
icans (L.) Kuntze), sweetspire ( I tea virginica L.),
and crossvine (Bignonia capreolata L.). Herbs are
poorly represented, except in light gaps and right
along the streambanks. Due to the persistently sat¬
urated soils, fire is only a minor component of the
ecology of this community. Ecotones to Small
Stream Swamps are generally less diverse than
those of slreamheads due to diminished seepage
from adjacent uplands. Only small populations (I
to 5 plants) of I ilium pyrophilum occur in these
ecotones.

Rarity, management, and conservation. Lilium
pyrophilum is a very rare plant with clearly defined
threats. It is highly vulnerable due to rarity within
its limited range and patchy habitat, widespread
land conversion to other uses, and modern sup¬
pression of fires. Today it survives on properties on
which prescribed fire is a management tool or
where periodic cutting or mowing takes place, such
as in powerline and gasline rights-of-way.

The total number of individual plants docu¬
mented at the 42 known extant ( 1990— present) pop¬
ulations is approximately 205; there are 1 I popu¬
lations represented only by historical (pre-1990)
collections. The 6 largest extant populations num¬
ber 42, 25, 15, 12, II. and 10 individual plants;

all others hold fewer than 10 plants each, with most
limited to 1 to 3 individuals.

Fortunately, nearly all populations receive pro¬
tection (Table 1) that varies from active conserva¬
tion to management agreements to passive conser¬
vation. Thirty-nine of 42 extant populations and
189 of 205+ total plants are protected at some lev¬
el. Although this degree of protection is encourag¬
ing despite the small overall population, future
prospects remain uncertain. Even if landowners are
aware of the lily on their property and wish to foster
if, most do not have the resources available to mon¬
itor populations nor to conduct specific manage¬
ment protocols. Lilies survive on several properties
not as a result of management activities targeted at
the plants, but because of the general use of pre¬
scribed fire to improve habitat for federally endan¬
gered red-eoekaded woodpeckers, for improved
military troop maneuverability, and for forestry
management. On Fort Jackson in South Carolina,
exploding ordnance starts the frequent fires that
contribute most to habitat maintenance. In utility
rights-of-way, lilies survive because periodic cut¬
ting to promote human access reduces competition
from encroaching woody plants. More sobering is
the realization that all four unprotected populations
occur on private land in fire-suppressed situations.
The general lack of burning within the private sec¬
tor foretells slim prospects for finding significantly
more lilies.

The Sandhills lily currently occupies an ex¬
tremely narrow range of plant communities, and
there is no evidence to suggest that it formerly had
wider ecological amplitude. Like its close relative
L. iridollae, it appears to be a habitat specialist that
requires the unusual combination of saturated soils

Volume 12, Number 1
2002

Skinner & Sorrie

Conservation of Lilium pyrophilum

101

and periodic fire. Within the Sandhills region other
rare Carolina endemics such as Kalmia amenta
Michaux and the federally endangered Lysimachia
asperulifolia Poiret also share these requirements.
Each inhabits the inner portions of the ecotone
where shrubs and/or cane are dominant, rather than
laterally where graminoids prevail. Disturbance in
the form of periodic fire (or cutting of utility rights-
of-way) provides a release from competing shrubs
and tree saplings, followed by a brief period of a
few years when flowering and fruiting take place.

Although we acknowledge the ecological impor¬
tance of fire to this new lily, we also realize that
frequent fire is not a panacea for all members of
the longleaf pine ecosystem. Each has its own lim¬
its of fire tolerance with regard to frequency, sea¬
sonality, and intensity. Research is needed on the
specific effects of fire on Lilium pyrophilum. That
nearly as many lilies exist in powerlines and gas¬
lines as in areas that burn frequently suggests that
mechanical disturbance may be important as well.
But what form, if any, did this disturbance take in
pre-settlement times? Is it an adequate long-term
substitute for fire? To our knowledge no public util¬
ity uses fire as a management tool; it should be
tried where practicable and compared with tradi¬
tional cutting. The use of herbicides in rights-of-
way is a concern — does it affect the lily? If so,
widespread use of herbicides may significantly re¬
duce chances of finding additional populations of
the Sandhills lily.

Populations of rare lilies in (North America have
been decimated in the recent past by collectors and
fanciers who hope to appreciate these plants in
their home gardens (Skinner & Pavlik, 1994).
These misguided efforts usually end in failure due
to the highly specific habitat requirements of most
geographically restricted lilies. We encourage bot¬
anists and plant lovers to appreciate these plants
in the field, and caution that removal of plants from
most populations would be in violation of existing
state or federal laws.

Biologists and land managers continue to find
new populations, but overall numbers remain ex¬
tremely low. Therefore, we recommend that the U.S.
Fish and Wildlife Service consider Lilium pyro¬
philum for listing under the federal Endangered
Species Act. and urge state protection as well.

Similar species. Lilium pyrophilum is one of a
monophyletic group of lilies (Skinner, in press) that
also includes L. superhum (turk’s-cap lily), L. mi-
chauxii (Carolina lily), and L. iridollae (Panhandle
lily). This species group is characterized by a gen¬
erally southern distribution in the United States,
green styles, large flowers (sepals 5.7—10.5 cm)

with large anthers (1.0— 2.0 cm), buds that are tri¬
angular in cross section, generally smooth-mar¬
gined and smooth- veined leaves, sepals with two
abaxial longitudinal ridges, and whitish bulbs. The
remaining Lilium of eastern North America with
pendent flowers also represent a well-formed clade
marked by a primarily northern distribution, red
styles, smaller flowers (sepals 3.2— 9.3 cm) with
smaller anthers (0.4— 1.3 cm), round buds with
smooth backs, scarious-margined and scarious-
veined leaves that are therefore noticeably rough,
and bulbs that are yellow or become so with age.
This group includes Gray’s lily (L. grayi Watson),
Canada lily ( L . canadense L.), and Michigan lily (L.
michiganense Farwell). A key to L. pyrophilum and
allies follows:

Key to the Lilium Species or the Eastern United
States with Pendent Flowers, Green Styles, Tri¬
angular Buds, and Twin Abaxial Sepal Ridges

1. Leaves strongly oblanceolate, noticeably pale be¬
neath, somewhat fleshy; flowers strongly fragrant

. Lilium michauxii

1'. Leaves elliptic or weakly oblanceolate, scarcely
pale beneath, thin; flowers not fragrant.

2. Rhizomes with 3—4 annual bulbs, 9.6—18.4
cm, scaleless sections between annual bulbs
2. 7-5.4 cm; bulb scale leaves or their ab¬
scission scars present; leaves subtly oblan¬
ceolate; flowers yellow-orange, 1—3; coastal
Alabama and western Florida . . Lilium iridollae
2'. Rhizomes with 2(— 3) annual bulbs, 5.2—10.2
cm, scaleless sections between annual bulbs
0.3-3.8(-4.6) cm; bulb scale leaves or their
abscission scars absent; leaves ± narrowly
elliptic; flowers orange or reddish, 1-20 or
more; southern and eastern United States.

3. Plants 1.2— 2.8 m; leaves 7.1—26.1 cm,
3.9-18.4 times longer than broad, usually
horizontal or drooping, in 6—24 ± evenly
distributed whorls of 3 or more; flowers
red-orange (red, orange) to yellow (yel¬
low-orange) to green basally, 1—20 or
more; Louisiana to Missouri, east to Flor¬
ida and New Hampshire Lilium superbum
3'. Plants 0.63—1.6 m; leaves 2.3— 10. 3(—

12.2) cm, 1.6— 7.6(— 10.3) times longer
than broad, usually ascending, in 0—12
± proximal whorls of 3 or more; red-or¬
ange or dusky red (magenta or pinkish,
pale orange, red) apically to (pale) yellow'
(yellow-orange) centers to green basally,

1—7; Sandhills of Virginia and North and
South Carolina . Lilium pyrophilum

Of the four taxa, L. michauxii is the most deriv¬
ative and is easilv distinguished by its strongly ob-
ovate or oblanceolate leaves that are few' in number,
darker green above and lighter below, fleshy, and
undulate along the margins. Its compact bulb, pref¬
erence for well-drained sites, and delicately scent¬
ed flowers with exceedingly wide petals (1.8— 2.9

102

Novon

cm) are also distinctive. It is the only pendent lily
sympatric with L. pyrophilum. Lilium iridollae is
narrowly endemic to Escambia, Santa Rosa, Oka¬
loosa, and Walton Counties in the western Florida
panhandle and Baldwin, Escambia, and Covington
Counties in adjacent Alabama, and it is allopatric
to L. pyrophilum. Although its habitat within the
longleaf pine ecosystem is similar, it is morpholog¬
ically divergent by virtue of an elongate rhizome
that retains 3—4 years’ growth, the presence of basal
(winter) leaves (otherwise known in North Ameri¬
can lilies only in the distantly related pine lily, L.
catesbaei Walter), subtly oblanceolate stem leaves,
and particularly long pedicels (to 23 cm vs. a max¬
imum of 19 in the other three species). Its vibrant
and uniformly orange-yellow flowers are usually di¬
agnostic as well.

Lilium superbum is more similar to the Sandhills
lily than other congeners. It occurs directly to the
north and east of the limited range of L. pyrophilum
but is absent to the west through the Piedmont; it
then reappears along the Appalachian spine. Over¬
lap in blooming times of the two laxa is moderate
(Table 2); thus reproductive isolation is achieved
geographically and to some degree phenologically.
As its name suggests, L. superbum is taller and
more robust than the other lilies within its general
range, including L. pyrophilum (Table 2). It bears
more and larger flowers, has longer and relatively
narrower leaves, and has more whorls of 5 or more
leaves (Table 2), and these are rather evenly spaced
along the stem. Lilium pyrophilum typically has the
leaf whorls concentrated toward the bottom of the
stem, and the leaves are strongly ascending, which
is often characteristic of lilies that occur in very
high light environments. Of subtle importance is
the shape of the floral tube, which is slightly longer
and more tubular in L. pyrophilum than in L. su¬
perbum (Table 2). This may be related to pollinator
effectiveness (Skinner, 1988), as the Sandhills lily
is pollinated at least partially by hummingbirds
whereas L. superbum is almost entirely swallowtail
butterfly-pollinated, notwithstanding infrequent vis¬
its from hummingbirds. This longer, thinner tube
sometimes obscures the noticeable “green star” that
is formed by the six areas of nectary tissue in the
widely flaring L. superbum ; instead L. pyrophilum
often displays a green triangle formed only by the
sepal nectaries.

The rhizomes of L. pyrophilum anil L. superbum
are closely similar, though the latter tends to grow
from a larger bulb as befits its greater stature. Lil¬
ium superbum also has a tendency toward longer
scales that are two-segmented rather than entire,
and a more marked tendency toward clonal growth

that is a result of repeated dichotomous branching
within the rhizomes.

Phytogeography and theories of origin. We pro¬
pose three alternatives for the origin of the new
taxon. The authors do not necessarily agree on the
likelihood of each, but we feel that carefully chosen
molecular evidence might be conclusive.

1. As a peripherally isolated derivative of L. su¬
perbum or its ancestor. This theory acknowledges
the phenotypic similarity between these two species
and their current lack of sympatry.

2. Hybrid origin. Throughout its limited range L.
pyrophilum co-occurs with L. michauxii, but it over¬
laps scarcely if at all with L. superbum in the coast¬
al plain of southeastern Virginia. It is conceivable
that L. pyrophilum arose through past contact be¬
tween these other two species and has evolved its
own suite of morphological, biological, and ecolog¬
ical characteristics. Today, L. superbum and L. mi¬
chauxii are sympatric over a large area of moun¬
tains and piedmont from Virginia to Alabama.
Though they rarely produce hybrids (Adams, 1982),
the possibility ol hybrid origin cannot be excluded
as physical conditions and plant genotypes are flu¬
id. Hybrids between L. michauxii and L. pyrophil¬
um are known from disturbed habitats associated
with utility rights-of-way, and are best recognized
by the fine tepal spots and broad petals of the for¬
mer species. Whether they occur in more natural
settings is uncertain.

3. Common ancestry with L. iridollae. Eleven
vascular plant species occur in the Sandhills region
of the Carolinas as disjuncts from the Gulf region
w here L. iridollae grows (Sorrie et al., 1997; Sorrie
& Weakley, 2001), and a number of others share
essentially the same pattern of disjunction. Some
of these species occur in the same streamhead hab¬
itats as L. pyrophilum, including Carex turgescens
Torrey, Eriocaulon texense Kornicke, Rhynchospora
leptocarpa (Chapman ex Britton) Small. R. macro
(C. B. Clarke) Small, R. oligantha A. Gray. Xyris
chapmanii Bridges & Orzell, and X. scabrifolia
Harper. It may be significant that these same spe¬
cies also co-occur with Lilium iridollae in the Gulf
Coastal Plain, and it suggests that the historical
events that introduced these grasslike species to
the Sandhills may also have brought a common an¬
cestor of L. iridollae and L. pyrophilum into that
region.

The specimens cited below (herbarium acronyms
are listed in full in the Acknowledgments) repre¬
sent seven extant (1990— present) populations, each
in a distinct county (the Lee Co., North Carolina,
historical collection from 1961 has recently been
verified to represent an extant population), and

Volume 12, Number 1
2002

Skinner & Sorrie

Conservation of Lilium pyrophilum

103

Table 2. Major differences between L. pyrophilum and L. superbum. Measurements are means, (ranges), and [sample
sizes]; p values indicate the probability that the samples are from the same statistical population, and are from inde¬
pendent sample T-tests performed using Statistica (1999). Fire frequencies are from Frost (1998).

L. pyrophilum

/,. superhum

Distribution

Sandhills of SC, NC, and VA

Coast anil mountains from NH to

NC, MO, LA, and FL

Habitat

streamhead pocosins, sandhill seeps, swampy
streams, wet utility lines; fires extremely fre¬
quent (everyr 1—3 years)

rich, moist woods, roadsides, stream-
sides; fires vary (every' 1-100
years)

Blooming period

peak in early Aug. (late July to mid Aug.)

peak in mid— late July (July to early
Aug.)

Pollinators

ruby-throated hummingbirds ( Archilochus colu-
hri.s Linnaeus) and palamedes swallowtails
( Pnpilio palamedes Drury), probably other
swallowtails including spicebush (R. Iroilus
Linnaeus); relative importance of bird and
butterfly pollinators unknown

large swallowtails, esp. spicebush (R.
Iroilus ), also tiger (R. glaucus Lin¬
naeus) and pipevine ( Batlus phile-
nor Linnaeus)

Plant height (cm)
p <C 0.001

103.4 (63-160) [27 1

174.4 (1 18-280) [31]

Rhizome height

p = 0.02

26.0 (23.9-28.1) [4|

33.4 (23.5-43.9) 1 17]

Rhizome longest scale
(mm)

p = 0.09 (not signifi¬
cant)

15.5 (11-18.9) [4]

21.8 (1 1.8-39.3) [17]

Leaf arrangement

usually whorled, whorls often clustered toward
bottom of plant, lower leaves scarcely ascend¬
ing or ascending, drooping at tip or not, up¬
per ascending and appressed In stem

always whorled, whorls evenly

spaced on stem, leaves horizontal
and drooping at tips, upper leaves
often ascending in sun

Leaf shape

narrowly (to broadly) elliptic

(very) narrowly elliptic (to scarcely
oblanceolate)

Leaves per plant
p C 0.001

51.0 (29-102) 1 27 1

96.5 (40-186) [31 1

Whorls per plant of 5 or
more leaves
p C 0.001

3.9 (0-10) 1 27 1

8.4 (3-14) [31 1

Leaf length (mm)
p <C 0.001

62.2 [23- 103(- 122)| [206]

138.8 (71-261) [205]

Leaf length/width ratio
p <C 0.001

4.1 [1.6— 7.6(— 10.3)] [206]

9.9 (3.9-18.4) [205]

Flower color

muted orange-red or yellow-orange with red
dusting on tips (variable and sometimes red¬
dish pink, red-orange, magenta, dusky red, or
salmon), grading to (pale) yellow (yellow-or¬
ange) centers to green at base; maroon spots
often large

(pale) red-orange (pale red. red, or¬
ange, yellow scarcely suffused
with red) grading to yellow (yel¬
low-orange) to green at base; ma¬
genta (maroon) spots only in yel¬
low and often large

Flowers per plant
p <c 0.001

1.7 (1-7) [57]

3.7 (0-22) |299|

Sepal length (mm)
p = 0.014

78.1 (67-89) [25]

83.5 (68-105) [37]

Floral tube length (mm)

p « 0.001

17.9 (13.1-24.3) [25]

13.0 (8.9-16.5) [37]

Capsule length (mm)

p = 0.001

38.9 (28.2-47.0) [21]

45.3 (28.8-61.5) [23]

104

Novon

eleven historical populations (collected prior to
1990). Most ol these historical populations have
been surveyed but habitat is generally degraded,
and we assume the plant is extirpated at each lo¬
cation. Four counties — Nash and Northampton in
North Carolina and Kershaw and Orangeburg in
South Carolina — are represented only by historical
specimens collected before 1990. There are current
sight records (North Carolina Natural Heritage Pro¬
gram and South Carolina Heritage Trust databases,
1999) from 35 additional populations that add 4
additional Carolina counties to the total with extant
populations, as well as a sight record from 1 ad¬
ditional county of Virginia, Sussex County (B. Van
Eerden, pers. comm.). Thus L. pyrophilum is cur¬
rently known from 42 populations in 12 counties,
and historically at 1 1 populations in these and 4
additional counties.

Paratypes. U.S.A. North Carolina: Cumberland Co.,

6.8 mi. S of Fayetteville on INC 87, 7 Aug. 1657. II. E.
Allies 33525 (CA, NCU); Harnett Co., wet soil by route
53, 1 mi. S of Pineview, I Aug. 1927, II. R. Totten s.n.
(NCU); Fort Bragg Military Reservation, Northern Training
Area, seep and pocosin complex, 6 Aug. 1993, R. A. Sorrie
7512 with R. Von Eerden & T. Hippensteel (NCU), 3 Aug.
1995, M. IF Skinner 270 ((41); Hoke Co., pocosin border

3.9 mi. W of Montrose, 9 Aug. 1957, II. E. Aides 33802
(NCU); Johnston Co., streamhead under powerline, 20 July
1999, I1. McMillan 3900-a with E. Hojnos (CLEMS); Lee
Co., wet seepage along railroad, 2 mi. S of Lemon Springs,
29 July 1961, A. E. Radford 44148 (NCU); Moore Co.,
open shrub bog. Hog Island, SR 2026, 31 July 1974. ,/.
II. Carter III 1003 (wwh); seepage areas in powerline near
Pinehurst, 9 Aug. 1992, R. A. Sorrie 6746 (bas), 7 Aug.
1993, R. A. Sorrie 7513 (NCU); Nash Co., recently burned
dry pocosin on US 64, 2 Vi mi. SW of Nashville, 22 July
1949, W. R. Cox & R. k. Godfrey 2734 (NCSC); North¬
ampton Co., low undrained roadside 3 mi. N of Jackson,
2 Aug. 1958, ./. IL Hardin 910 (NCSC); Richmond Co.,
peat sedge bog near US 1, 5 mi. N of Rockingham, 24
July 1956, A. E. Radford 14323 (NCU); seepage bog in
powerline near Sandhills Game Land, 27 July 1997, R. A.
Sorrie 9367.5 (bas), 4 Aug. 2000, B. A. Sorrie 10584
(NCU); Richmond/Scotland Co., near Pine Lake, 30 July
1933, E Smith s.n. (DUKE). South Carolina: Chester¬
field Co., peat sedge bog 8 mi. E of Patrick, 1 1 Aug. 1956,
A. E. Radford 15809 (NCU); Kershaw Co., powerline
crossing of a cleared pond pine pocosin, dirt road off US
I, 3 Sep. 1982, I). A. Rayner 1496 (USCH); Orangeburg
Co., boggy woodland border 3 mi. NE of Orangeburg on
SC 33. 19 July 1957, //. E. Aides 31685 (NCU); Richland
Co., Fort Jackson Army Installation, South Impact Area,
15 Apr. 1992,./. R. Nelson 12269 (USCH), I Aug. 1995,
M. W. Skinner 267 (GH). Virginia: Greenville Co., W of
Jarratt in powerline, diverse seepage wetland, 8 Aug.
1991, T. J. Rawinski 11471 (VPI).

Acknowledgments. We thank Cary' Norquist of
the II. S. Fish and Wildlife Service for inspiration
and financial support, A1 Sehotz for illuminating
discussions about relationships in Lilium, and Al,

Jame Amoroso, Scott Gunn, and Bert Pittman for
their expertise in the field. Victoria Hollowell’s crit¬
ical reading was salutary, and (Jerald Smith and two
anonymous reviewers significantly improved the
manuscript. We appreciate Yevonn Wilson-Ramsey
for her lovely illustration and Roy Gereau for his
clever Latin translation, and we gratefully acknowl¬
edge the curators of the herbaria we visited [bas
(pers. herb. B. A. Sorrie, Whispering Pines, NC),
CLEMS, DUKE, FLAS, FSU, CA, HUH. IBE, LSU.
MISSA, MO. NGSC, NCU. NO. PH, US, USCH.
USNM, VDB (BRIT), and wwh (Weymouth Woods
Sandhills Preserve Herbarium, Southern Pines,
NC)] and their institutions.

Literature Cited

Adams, R. M. II. 1981. Comments on the three new L il¬
ium species described by Roane and Henry. N. Amer.
Lily Soc. Yearb. 34: 21-24.

- . 1982. Nodding Lilium species of eastern North

America [Liliaceae]. Baileya 21: 165—188.

Eenneman, N. M. 1938. Physiography of the Eastern Unit¬
ed States. McGraw-Hill, New York.

Frost, C. 1998. Presettlement fire frequency regimes of the
United States: A first approximation. Pp. 70-81 in T. L.
Pruden & L. A. Brennan (editors). Fire in Ecosystem
Management: Shifting the Paradigm from Suppression
to Prescription. Tall Timbers Fire Ecology Conference
Proceedings, No. 20. Tall Timbers Research Station,
Tallahassee, Elorida.

Henry, M. G. 1946. A new lily from southern Alabama
and northern Florida. Bartonia 24: 1—5.

Moretz, C. C. & G. L. Smith. 1995. Lilium gazarubrum
Roane and J. N. Henry discovered in North Carolina
(abstract). ASB Bull. 42: 143.

North Carolina Natural Heritage Program. 1999. Database
records for Lilium “ iridollae Division of Parks and
Recreation, Raleigh, North Carolina.

Radford, A. E„ H. E. Ahles & C. R. Bell. 1968. Manual
of the Vascular Flora of the Carolinas. Univ. North Car¬
olina Press, Chapel Hill.

Roane, M. K. & J. N. Henry. 1980. Notes on four lilies
from southeastern United States. Quart. Bull. N. Amer.
Lily Soc. 34: 25—32.

Russo. M. J. 1997. Status Survey of Lilium sp. nov. [for¬
merly considered Lilium iridollae Henry], Unpublished
report prepared for North Carolina Plant Conservation
Program and U.S. Fish and Wildlife Service.

Schafale, M. P. & A. S. Weakley. 1990. Classification of
the Natural Communities of North Carolina. Third Ap¬
proximation. North Carolina Natural Heritage Program,
Division of Parks and Recreation. Raleigh.

Sharitz, R. R. A J. W. Gibbons. 1982. The Ecology of
Southeastern Shrub Bogs (Pocosins) and Carolina Bays:
A Community Profile. U.S. Fish & Wildlife Service,
Washington. i).C. FWS/OBS-82/04.

Skinner, M. W. 1988. Comparative Pollination Ecology
and Floral Evolution in Pacific Coast Lilium. Ph.l). Dis¬
sertation, Harvard University, Cambridge.

- . In press. Lilium. In Flora of North America Ed¬
itorial Committee (editors). Flora of North America
North of Mexico, Vol. 26. Oxford Univ. Press, New York.
- & B. M. Pavlik. 1994. Inventory of the Rare and

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2002

Skinner & Sorrie

Conservation of Lilium pyrophilum

105

Endangered Vascular Plants of California. California
Native Plant Society, Sacramento.

Some, If. A. & A. S. Weakley. 2001. Coastal plain vas¬
cular plant endemics: Phytogeographic patterns. Cas-
tanea 66: 50—82.

- , B. van Eert len & M. J. Russo. 1997. Noteworthy

plants from Fort Bragg and Camp MacKall, North Car¬
olina. Castanea 62: 239-259.

South Carolina Heritage Trust. 1999. Database records for
Lilium “ iridollae .” Department of Natural Resources,
Columbia, South Carolina.

Statistica. 1999. StatSoft, Inc. Tulsa, Oklahoma.

New Combinations within North American Schoenoplectus smithii
and S. purshianus (sect. Actaeogeton, Cyperaceae) and Comparison

with Eastern Asian Allies

S. Galen Smith

Department of Biological Sciences, University of Wisconsin-Whitewater, Whitewater,
Wisconsin 53190, U.S.A. sgsmith2@facstaff.wisc.edu

Eisuke Hayasaka

Botanical Garden, Graduate School of Science, Tohoku University, Sendai, 980-0862 Japan.

hayasaka@biology.tohoku.ac.jp

ABSTRACT. The new combinations Schoenoplectus
purshianus (Fernald) M. T. Strong var. williamsii
(Fernald) S. G. Smith, .S'. smithii (A. Cray) Sojak
var. setosus (Fernald) S. G. Smith, and S. smithii
var. levisetus (Fassett) S. G. Smith are made. Di¬
agnostic features, mainly from spikelet dimensions,
perianth bristles, and aehenes, are provided that
distinguish the eastern North American .S’, smithii
and S. purshianus, and the eastern Asian and Pa¬
cific island S. hotarui (Ohwi) Holuh, .S', juncoides
(Roxburgh) Falla, and .S’, komarovii (Roshevitz) So¬
jak, all five of which have been confused with each
other. A description of Schoenoplectus sect. Actaeo¬
geton (Reichenbach) J. Raynal is provided based
mostly on North American and eastern Asian spe¬
cies.

Key words: Asia. Cyperaceae, North America,
Schoenoplectus, Schoenoplectus sect. Actaeogeton.

This paper deriv es from the senior authors work
on his treatment of Schoenoplectus (Reichenbach)
Falla for the upcoming volume 23, Cyperaceae, of
the Flora of North America (FNA), during which he
studied numerous herbarium specimens from about
52 herbaria (Smith, 1995), as well as from the ju¬
nior authors ongoing work toward a monograph of
Schoenoplectus, during which he studied numerous
herbarium specimens from the herbaria listed in
the acknowledgments.

Scirpus L. 8.1. has been segregated worldwide in
recent years based on embryo as well as vegetative
and reproductive characters (Bruhl, 1995; Goetgh-
ebeur & Simpson, 1991; Hayasaka & Ohashi,
2000; Smith & Hayasaka, 2001; Smith & Yatskiev-
ych, 1996; Wilson, 1981). Nine segregate genera
will be recognized for the upcoming Flora of North
America: Amphiscirpus, Blysmus, Bolboschoenus, Is-
olepis, Oxycaryum, Schoenoplectus, Scirpus, Tricho-

Novon 12: 106-1 1 1. 2002.

pliorum, and Wehsteria. The largest of these in
North America is Schoenoplectus with about 70 spe¬
cies worldwide and 17 in North America.

Schoenoplectus is diverse morphologically and
has been divided into several infrageneric taxa
(Oteng-Yeboah, 1974; Raynal, 1976; Smith & Hay¬
asaka, 2001). Smith and Hayasaka (2001) recog¬
nized and provided a key to four sections within
Schoenoplectus: sect. Schoenoplectus, seel. Actaeo¬
geton (Reichenbach) J. Raynal, sect. Malacogeton
(Ohwi) S. G. Smith & H ayasaka, and sect. Supini
(Chermezon) J. Raynal.

Upon their transfer from Scirpus to Schoenoplec¬
tus, the infraspecific taxa of the two native North
American species of Schoenoplectus sect. Actaeo¬
geton, S. purshianus (Fernald) M. T. Strong and S.
smithii (A. Gray) Sojak. require new combinations.

I liese species also have been confused with each
other as well as with three species of eastern Asia
and adjacent western Pacific islands (Koyama,
1958, 1962). In addition, the section Actaeogeton
has never been clearly delineated. This paper ad-
dr* *sses these nomenclatorial and taxonomic prob¬
lems.

b Schoenoplectus purshianus (Fernald) M. T.
Strong, Novon 3: 202. 1993. Scirpus purshi¬
anus Fernald, Rhodora 44: 479. 1942. Scirpus
debilis Pursh, 11. Amer. Sept. 1: 55. 1814. non
Scirpus debilis Lamarck, lllustr. 1: 141. 1791.
Scirpus erectus Poiret var. debilis Camus, FI.
Gen. Indo-Chine 7: 136. 1912. TYPE: U.S.A.
Pennsylvania: Muhlenberg Herbarium, “No.
65 and “in. 33 ’ (leetotype, designated by
Strong (1993: 203), PH, Herb. Muhl. sheet
345—74, photograph seen; isoleetotype, PH,
Herb. Muhl. sheet 345—75, photograph seen).

Muhlenberg first published the name Scirpus de-

Volume 12, Number 1
2002

Smith & Hayasaka

North American Schoenoplectus

107

bilis without a description in his 1813 catalog, and
Pursh apparently appropriated Muhlenberg’s earlier
name. The lectotype and duplicate sheet may have
been seen bv Pursh (Merrill & Hu, 1949), or the
specimens may even have been collected by Pursh
(C. E. Smith, 1962: 458). A. E. Schuyler and J.
Macklin at PH have kindly provided us with pho¬
tographs of die types and closely examined the
specimens. On the lectotype sheet, “Scirpus debilis
lupulinus Sm.” is written on the label; “345-74” is
at the bottom of the sheet. The annotation directly
on the sheet reads “Scirpus purshianus Fern. Type
of Scirpus debilis Pursh, AE Schuyler 1976”; this
sheet bears 15 inflorescence-bearing culms, 3 of
which are attached together at their bases, and 12
are without bases. A second sheet bears labels and
annotations similar to those on t he lectotype with
“345—75” written at the bottom of the sheet, and
the annotation on the sheet reads “Scirpus purshi¬
anus Fern. TYPE MATERIAL of Scirpus debilis
Pursh, FI. Am. Sept. 1: 55. 1813|1814|. AES
1976.” It bears 14 culms with inflorescences, and
is evidently part of the same collection as the lec¬
totype. All culms on both sheets are very similar,
and several bear achenes with attached bristles,
which are typical of Schoenoplectus purshianus (A.
E. Schuyler & J. Macklin, pers. comm. 2001).

I a. Schoenoplectus purshianus var. purshi¬
anus

Perianth bristles 6. stout, tapered distally, about
equaling achene, densely spinulose. Reported hab¬
itats: shores of inland ponds and ditches. Distrib¬
uted throughout the range of the species in north¬
eastern North America from Quebec to Georgia
westward to Minnesota, eastern Missouri, and
northern Louisiana.

1 h. Schoenoplectus purshianus var. williumsii

(Fernald) S. G. Smith, comb. nov. Basionym:
Scirpus debilis Pursh var. williamsii fernald,
Rhodora 3: 252. 1901. Scirpus smithii A. Gray
var. williamsii (Fernald) Beetle, Amer. J. Rot.
29: 655. Oct. 1942. Scirpus purshianus Fer¬
nald f. williamsii (Fernald) Fernald, Rhodora
44: 479. Dec. 1942. Scirpus juncoides Rox¬
burgh var. williamsii (Fernald) T. Koyama,
Canad. J. Bot. 40: 914. 1962. Schoenoplectus
smithii (A. Gray) Sojak subsp. williamsii (Fer¬
nald) Sojak. Cas. Nar. Mus. Odd. Prir. 148:
194. 1980. TYPE: U.S.A. Massachusetts:
sandy shore of Massapoag Lake, Sharon, 7
Sep. 1901, Plantae Exsiccatae Grayanae 26,
E. F. Williams & M. L. Fernald s.n. (holotype,
GH; isotypes, BRIT, GAS, (41, ILL, MO,
NEBC, SAPS, UC, WTU).

The names Scirpus smithii A. Gray var. williamsii
(Fernald) Beetle and Scirpus juncoides Roxburgh
var. williamsii (Fernald) T. Koyama were misap¬
plied by their authors to the entire species S. pur¬
shianus. Koyama (1963: 457) stated that his new
combination Scirpus juncoides Roxburgh var. digyn-
us (Bockler) T. Koyama has priority over Scirpus
debilis var. williamsii. As Koyama referred to his
1962 paper, he apparently intended his new vari¬
etal combination to apply to the whole species ,S.
purshianus. Bockler’s type of the basionym Scirpus
supinus L. [var.] digynus Bockler needs to he ex¬
amined to determine the specific and varietal iden¬
tity of Scirpus juncoides var. digynus (Bockler) T.
Koyama.

Perianth bristles absent. Reported habitats:
shores of lakes, reportedly with substantial water-
level fluctuations. This variety is known only from
the type locality in eastern Massachusetts, several
counties in southwestern Michigan, one county in
adjacent Indiana, and two counties in Wisconsin.

2. Schoenoplectus smithii (A. Gray) Sojak, Gas.
Nar. Mus. Odd. Prir. Ill: 62. 1972. not
Schoenoplectus smithii (A. Gray) ,1. Raynal,
Adansonia, n.s., 16(4): 530. 1977. Scirpus smi¬
thii A. Gray, Manual ed. 5: 563. 1867. T\ PE:
U.S.A. New Jersey: in tidal mud on the N.J.
shore of the Delaware below Red Bank oppo¬
site the River Schuylkill, July 1865. C. F.
Smith s.n. (holotype, GH; isotype, PH).

2a. Schoenoplectus smithii var. smithii

Perianth bristles absent or rudimentary. Report¬
ed habitats: fresh-water tidal shores of rivers and
inland shores with large fluctuations in water level.

I bis variety is known from Quebec (St. Laurence
River near Quebec City) to New Jersey westward
to Minnesota.

2b. Schoenoplectus smithii var. setosus (Fer¬
nald) S. G. Smith, comb. nov. Basionym: Scir¬
pus smithii A. Gray var. setosus Fernald, Rho¬
dora 3: 252. 1901. Scirpus smithii A. Gray f.
setosus (Fernald) Fernald, Rhodora 44: 479.
1942. TYPE: U.S.A. Illinois: Augusta, 1845.
S. B. Mead s.n. (holotype, (41: isotypes, GH.
NY. PH).

Perianth bristles 4 to 6, equaling or slightly ex¬
ceeding achene, densely spinulose. Chromosome
number n = 19 (Schuyler, 1972). Reported habi¬
tats: wet places with little fluctuation in water level
such as mires and shores of small lakes or ponds.
This variety is known from Quebec (near Quebec

108

Novon

City) to Delaware and western North Carolina west¬
ward to Wisconsin and Minnesota. In Michigan,
Wisconsin, and Minnesota it is much more common
than S. smithii var. smithii.

2c. Schoenoplectus smithii var. levisetus (Fas-
sett) S. G. Smith, comb. nov. Basionym: Scirpus
smithii A. Gray var. levisetus Fassett. Rhodora
19: 42. 1921. Scirpus smithii A. Gray f. levi¬
setus (Fassett) Fernald, Rhodora 44: 479.
1942. TYPE: 1J.S.A. Maine: tidal Hats of the
Cathance Biver at Bowdoinham and at its
mouth in Merrymeeting Bay, 25 Ang.-2 Sep.
1920, N. C. Fassett s.n. (holotype, GH: isotype,
NY).

Perianth bristles 1 to 3 (4). mostly much shorter
than achene, mostly without spinules. Chromosome
number n — 19 (Schuyler, 1972). Habitat: restrict¬
ed to fresh-water (or slightly brackish?) tidal muddy
or rocky shores of rivers. This variety is very locally
distributed but sometimes locally common (Schuy¬
ler, 1972). Smith has seen specimens from New
Brunswick (Northumberland Co.), Quebec (Port-
neuf Co., near Quebec City), Connecticut (New Ha¬
ven), Massachusetts (Middleton Co.), Maine (Pen¬
obscot! and Sagadahoc Cos.), and Virginia (Fairfax
Co.). It is known to be extirpated from Virginia
(Strong, 1994).

Koyama (1962: 919) treated the eastern Asian
Schoenoplectus komarovii (Roshevitz) Sojak as Scir¬
pus smithii var. leiocarpus (Komarov) T. Koyama, =
Schoenoplectus smithii subsp. leiocarpus (Komarov)
Sojak, basionym Scirpus supinus L. var. leiocarpus
Komarov (other synonyms and type given by Koy¬
ama, 1958, 1962). However, our herbarium obser¬
vations and published descriptions (Koyama, 1958.
1962; Ohwi, 1965) indicate that, pending a revision
of the S. juncoides complex, S. smithii var. leiocar¬
pus should be treated as the distinct species S. ko-
rnarovii following Ohwi (1965). Although S. koma¬
rovii is similar to S. smithii var. setosus in most
characters, including those of perianth bristles and
achene shape and surface, it differs from S. smithii
in its smaller achenes and its frequently compound
inflorescences (Table 1).

While Smith lias not detected morphological dif¬
ferences between the varieties other than those of
the perianth bristles as given above, in his opinion
these live infraspecific taxa should be treated as
varieties rather than forms because the perianth
bristle differences appear to be correlated with hab¬
itat differences, and thus these taxa can be inter¬
preted as ecotypic varieties and may be valuable
ecological indicators. Also, form status implies that

the morphological differences are completely trivi¬
al.

Schoenoplectus smithii var. setosus, S. smithii var.
levisetus, and S. purshianus var. williamsii were first
described with no mention of ecological differences
except for S. smithii var. levisetus, for which Fassett
{1921) found the fresh-water tidal habitat remark¬
able. \\ hen Fernald (1942: 483) reduced the vari¬
eties to forms he merely stated: “. . . their differ¬
ences are in the presence or absence of
perianth-bristles, or in the case of S. smithii, forma
levisetus, the lack of retrorse barbs on the bristles.”
However, Schuyler (1972: 398—399) stated: “In
both 5. purshianus and S. smithii the presence or
absence of barbed bristles appears to be correlated
with the stability of the habitat. For example in
eastern New York, New Jersey, eastern Pennsylva¬
nia and Delaware. S. smithii f. smithii is almost
completely restricted to the specialized conditions
of fresh inter-tidal zones of the Hudson, Raritan
and Delaware Rivers while in the same area f. se-
tosus occurs in and around lakes, ponds and quag¬
mires. . . . Thus it appears that plants of S. smithii
which lack barbed bristles are better adapted to the
periodic fluctuations in water levels which are char¬
acteristic of the estuarine environment than are
those which have barbed bristles. . . . S. smithii f.
levisetus is common along the estuaries of the Ken¬
nebec River in Maine and the St. Lawrence River
in Quebec.” Ferren and Schuyler (1980) and Strong
(1994) essentially repeated these statements for S.
smithii var. smithii and S. smithii var. setosus and
provided more detailed ecological information for
S. smithii var. smithii.

fhe few1 data Smith have seen on herbarium la¬
bels support these reported habitat differences. For
S. smithii var. smithii , in addition to coastal estu¬
aries, reported habitats include protected beaches
of the very large Lakes Huron, Erie, Cayuga (New
York), Oneida (Newr York) and Champlain. In con¬
trast. reported habitats for S. smithii var. setosus are
“bog” pools and peaty quagmires, marsh edges,
sandy or muddy lake and river shores, marshy
shore of drying shallow seepage lake, recently
scraped road through wet meadow near lake shore,
floating muck in lake in center of bog, mill pond,
and muck of small drying-up alkaline lake.

Schoenoplectus purshianus and S. smithii have
been widely confused. Many specimens are mis-
identified, and some authors have treated .S’, pur¬
shianus and S. smithii as nonspecific (Beetle, 1942;
Gleason, 1963; Gleason & Cronquist, 1963; Voss,
1972). Most published distributions are therefore
erroneous, and the distributions given here are
based almost entirely on specimens that the senior

Volume 12, Number 1
2002

Smith & Hayasaka 109

North American Schoenoplectus

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Novon

author has seen. The taxonomic confusion is due to
the subtle nature of the diagnostic characteristics
of achenes and perianth bristles as illustrated by
Strong (1994), Blondeau et al. (1996), and in the
upcoming volume 23 of the Flora of North America,
and is summarized in Table 1 herein. Confusion is
also caused by the presence in both species of
plants with or without perianth bristles as well as
plants witli erect proximal involucral bracts. Plants
without bristles may be identified to species using
the differences in achene shape and surface sculp¬
turing as given in Table 1.

Our herbarium studies and the literature indicate
that Schoenoplectus purshianus and S. smithii
should be treated as distinct species restricted to
eastern North America and distinct from their rel¬
atives, which are found in eastern Asia and the
Pacific islands. Some authors, however, have treat¬
ed S. purshianus and S . smithii as conspecific with
eastern Asian species of the “S. juncoides complex”
(Koyama, 1958. 1962. under Scirpus ), which is
comprised ol perhaps 10 taxonomically difficult
species of eastern Asia and tin* Pacific islands. The
distinguishing characteristics of S. purshianus and
S. smithii and the eastern Asian relatives with
which they have been confused are summarized in
Table 1.

Dei. imitation of Schoenoplectus sect.
Actaeogeton

The species of Schoenoplectus sect. Actaeogeton
anti section Supini were all included in Scirpus
sect. Actaeogeton by beetle (1942). Fernald (1942).
and Koyama (1958, 1962). Raynal (1976) segre¬
gated Schoenoplectus sect. Supini from other sec¬
tions based on the perianth bristles reduced or ab¬
sent, and the ability to produce amphiearpie flowers
in the basal leaf sheaths. Although species of
Schoenoplectus sects. Actaeogeton and Supini are
mostly very similar in both vegetative and sexual
structures, our studies support their treatment as
separate sections based mainly on the ability for
amphiearpy and the usual presence of a node above
the basal leaves in the latter (Smith & llayasaka,
2001). All plants of the North American species of
these two sections can be distinguished by these
characters (Smith & Yatskievych, 1996), but some
Old World species of Schoenoplectus sect. Supini,
e.g., S. supinus (L.) Palla, lack a cauline leaf.

The following description of Schoenoplectus sect.
Actaeogeton is based on about 300 specimens of
the North American species and about 500 speci¬
mens of eastern Asian species as well as the de¬
scriptions and illustrations in Koyama (1958. 1962)

and Ohwi (1965). The achene SEM micrographs
provided by Oh and Ham (1998) show that the epi¬
dermal surface sculpturing of S. hotarui (Ohwi)
Holub, S. lineolatus (Franchet & Savatier) T. Koy¬
ama. S. triangulatus (Roxburgh) Sojak. and S. wal-
lichii (Nees) T. Koyama. all of which we include in
section Actaeogeton (Smith & Hayasaka. 2001), are
very similar to each other, sharing longitudinally
oriented, linear epidermal cells.

Schoenoplectus (Reichenbach) Palla sect. Ac-
tueogeton (Reichenbach) J. Raynal. Adanso-
nia 16: 130. 1976. Scirpus I,, sect. Actaeogeton
Reichenbach, FI. Germ. Excurs.: 78. 1830.
Schoenoplectus subg. Actaeogeton (Reichen¬
bach) Oteng-Yeboah, Notes Roy. Bot. Card.
Edinburgh 33: 315. 1974. TYPE: Schoeno¬
plectus mucronatus (E.) Palla.

Annuals or perennials, culms densely tufted, ca.
5—200 cm; rhizomes present or apparently absent,
very short, hidden among culm bases and roots, or
rhizomes long and horizontally creeping and culms
single (in .S’, lineolatus). Culms cylindrie or trigon¬
ous, ca. 1—200 cm long X 1—8 mm thick. Leaves
all basal, Vi of to rarely ca. equaling culm, ca. 2
leaves well developed; ventral band hyaline or
membranous; ligules minute; blades from a mere
mucro to ca. 8 cm long X 0.5-1 mm wide, cross
section C-shaped or trigonous. Inflorescence of a
single spikelet or usually capitate; spikelets 2 to
20: involucral bracts 1 to 2. the proximal erect to
divergent, ehanneled-subcylindrie or trigonous,
usually greatly exceeding the inflorescence. Floral
scales elliptic to ovate; membranous with thicker
midrib, often faintly manv-nerved: margins cilio-
late; apices entire, mueronate. Perianth present or
absent, of 6 (or fewer) bristles that are retrorsely to
spreading spinulose or rarely smooth. Stvles bifid
or trifid. Achenes dark brown to blackish when
ripe, obovoid, lenticular to trigonous, abruptly
short-beaked, prominently to obscurely rugose or
nearly smooth at 10 to 20X. Reported chromosome
numbers n = 19. 20. 21. 22. 30. 36, 37.

Habitats and distribution. Fresh-water wet¬
lands, often emergent; worldwide, with the main
center of diversity in eastern Asia.

This description is tentative pending further
study of species, especially from Asia and Africa,
where a taxonomic revision is needed. Fourteen
species that we include in Schoenoplectus sect, Ac-
taeogeton from North America and eastern Asia are
listed in Smith and Hayasaka (2001). but it is pos¬
sible that more species would be included after fur¬
ther study. Although S. lineolatus has long, liori-

Volume 12, Number 1
2002

Smith & Hayasaka

North American Schoenoplectus

111

zontally creeping rhizomes in sharp contrast to the
short, hidden rhizomes of all of the other species
we include in Schoenoplectus sect. Actaeogeton, S.
lineolatus closely resembles the other species in its
leaves, floral scales, achene surfaces, and pericarp
internal structure (Hayasaka, unpublished data). In
addition, hybrids between S. lineolatus and both .S'.
hotarui and S. triangulatus have been reported
from Japan (Koyama, 1958; Hayasaka & Ohashi,
2000).

Acknowledgments. The senior author especially
thanks A. E. Schuyler for generously sharing his
insights and unpublished research materials on
Schoenoplectus, and A. Reznicek and M. Oldham
for providing information and voucher specimens,
especially from Canada. He also thanks the cura¬
tors of the many herbaria that loaned specimens
and accommodated bis visits as listed in Smith
(1995). We thank the directors and staff of the her¬
baria at the University of Wisconsin (W IS) and lo-
hoku University (TUS), and the Department of Bi¬
ological Sciences of the University of
Wisconsin-Whitewater for their essential logistical
support and kind help in many ways. Two reviewers
provided valuable critical comments, and Victoria
Hollowed, scientific editor at Novon, provided valu¬
able information, critical comments, and skillful
editing. The junior author thanks Mitsuo Suzuki
and Hiroyoshi Ohashi for their continuous help in
many ways and warm encouragement since he be¬
gan his work on Schoenoplectus in 1997, and the
curators of A, B1SH, BM. BRI, FLAS, GH. IBSC,
KUN, L, NOU, P, SAPS. SING, SKK. SNU. TUS,
TUSG, UPS. and URO for the loan of specimens or
accommodating his visits.

I .iterature Cited

Beetle, A. A. 1942. Studies in the genus Scirpus L. V.
Notes on the section Actaeogeton Reich. Arner. J. Rot.
29: 563—565.

Blondeau, M., C. Roy & M. Garneau. 1996. Schoenoplec¬
tus purshianus. Une Cyperaceae nouvelle pour le Que¬
bec. Nat. Canad. 120: 1 1-16.

Bruhl, J. J. 1995. Sedge genera of the world: Relationships
and a new classification of the Cyperaceae. Austral.
Syst. Bot. 8: 125-305.

Fassett, N. C. 1921. An estuarian variety of Scirpus smi-
thii. Rhodora 23: 4 1 —13.

Fernald, M. L. 1942. Transfers in Scirpus [sect.| Actaeo¬
geton. Rhodora 44: 479—484.

Ferren, W. R„ Jr. & A. F. Schuyler. 1980. Intertidal vas¬

cular plants of river systems near Philadelphia. Proc.
Acad. Nat. Sei. Philadelphia 123: 86—120.

Gleason, H. A. 1963. The New Britton and Brown Illus¬
trated Flora of the Northeastern United States and Ad¬
jacent Canada, Vol. 1. Third printing, revised. Hafner,
New York.

- & A. Cronquist. 1963. Manual of Vascular Plants

of Northeastern United States and Adjacent Canada. I).
Van Nostrand. Princeton, New Jersey.

Goetghebeur, P. & 1). A. Simpson. 1991. Critical notes on
Actinoscirpus, Bolboschoenus, Isolepis , Bhylloscirpus and
Amphiscirpus (Cyperaceae). Kew Bull. 46: 168-178.

Hayasaka, E. & H. Ohashi. 2000. New combinations in
Japanese Schoenoplectus (Cyperaceae). J. Jap. Bot. 75:
223-225.

Koyama, T. 1958. Taxonomic study of the genus Scirpus
LinnG J. Fac. Sci. Univ. Tokyo, sect. 3, Bot. 7: 271—
366.

- . 1962. The genus Scirpus Linn. Some North

American aphylloid species. Canad. J. Bot. 40: 913—
937.

- . 1963. Nomenclatural remarks on the genus Scir¬
pus. Phytologia 8: 457-458.

Merrill. E. I). & S.-Y. Hu. 1949. Work and publications
of Henry Muhlenberg, with special attention to unre¬
corded or incorrectly recorded binomials. Bartonia 25:

1 —66.

Oh. Y. C. & E. J. Ham. 1998. A taxonomic study on
Scirpus Linn£ (Cyperaceae) of Korea. Korean J. PI. Tax¬
on. 28: 217—247.

Ohwi. J. 1965. Flora of Japan [in English). Edited by F
G. Meyer & E. 11. Walker. Smithsonian Institution,
Washington, D.C.

Oteng-Yeboah, A. A. 1974. Taxonomic studies in Cyper-
aceae— Cyperoideae. Notes Roy. Bot. Gard. Edinburgh
33: 31 1-316.

Raynal, J. 1976. Notes Cyperologiques: 26. Le genre
Schoenoplectus II. L’amphicarpie et la sect. Supini. Ad-
ansonia, ser. 2, 16: 119—155.

Schuyler, A. E. 1972. Chromosome numbers of Scirpus
purshianus and S. smithii. Rhodora 74: 398-402.

Smith, C. E.. Jr. 1962. Henry Muhlenberg — Botanical pi¬
oneer. Proc. Amer. Philos. Soc. 106: 443—460.

Smith, S. G. 1995. New combinations in North American
Schoenoplectus. Bolboschoenus. Isolepis. and Trichopho-
rum (Cyperaceae). Novon 5: 97—102.

- & E. Hayasaka. 2001. Delineation of Schoeno¬
plectus sect. Malacogeton (Cyperaceae), new combina¬
tion, and distinctions of species. J. Jap. Bot. 76: 339-
343.

- & G. Yatskievych. 1996. Notes on the genus Scir¬
pus s. lat. in Missouri. Rhodora 98: 168-179.

Strong, M. T. 1993. NewT combinations in Schoenoplectus
(Cyperaceae). Novon 3: 202—203.

- . 1994. Taxonomy of Scirpus, Trichophorum, and

Schoenoplectus (Cyperaceae) in Virginia. Bartonia 58:
29-68.

Voss, E. G. 1972. Michigan Flora, Vol. I. Cranbrook In¬
stitute of Science, Bloomfield Hills, Michigan.

Wilson, K. I,. 1981. A synopsis of the genus Scirpus sens,
lat. (Cyperaceae) in Australia. Telopea 2: 153—1 i2.

Neomitranthes obtusa (Myrtaceae), a New Species from Espirito

Santo, Brazil

Marcos Sobral

Faculdade de Farntacia UFRGS, Av. Ipiranga 2752, 90610-010, Porto Alegre, RS, Brazil.

sobral@farmacia.ufrgs.br

Oberdan Za inborn

Departamento de Botaniea UFES, Av. Mareclial Campos 1468, 29040-090, Vitoria,

ES, Brazil

ABSTRACT. Neomitranthes obtusa (Myrtaceae), a
treelet from restinga forests in Espirito Santo,
southeastern Brazil, is described, illustrated, and
compared with the apparently related Neomitran¬
thes glomerata, from which it is distinguished hv
its obtuse leaves, inconspicuous venation, fewer
secondary veins, and smaller petioles.

RESUMED Neomitranthes obtusa (Myrtaceae), un
arbolito de las restingas de Espirito Santo, en el
sudeste de Brasil, es descrita, ilustrada y compar-
ada con la especie aparentemente afin Neomitran¬
thes glomerata , de la eual se distingue por las hojas
obtusas, la venacion inconspicua, el niimero menor
de nervios secundarios y los peciolos mas cortos.

Key words: Brazil, Myrtaceae, Neomitranthes.

Neomitranthes was erected bv Eegrand (Legrand
& Klein. 1977) in order to accommodate three
southern Brazilian species formerly described un¬
der Mitranthes. I he main distinction between these
two genera lies in the embryo morphology: Mitran¬
thes presents well-developed cotyledons and hypo-
cotyl, which are characteristic of subtribe Myrci-
inae in Myrtaceae; in Neomitranthes the cotyledons
are well developed but the hypocotyl is absent or
inconspicuous, characteristic of subtribe Eugeni-
inae. Since then, although the independence of
Neomitranthes is clear, its generic boundaries are
far from confidently established. The genus is iden¬
tified chiefly by its axillary flowers borne in ra¬
cemes or dichasia that open through a calyptra.
Ovaries of Neomitranthes have 2 loeules with 2 to
6 ovules per locule, and its embryos reveal 2 dis¬
tinct plano-convex cotyledons occasionally with a
small hypocotyl; the wall of tin* loeules in some
species is internally pilose, a puzzling character oc¬
casionally found in other Eugeniinae such as Eu¬
genia and Hexachlamys. All these characters ex¬
cept the hairy loeules overlap those present in other

Novon 12: 112-114. 2002.

more widespread Eugeniinae such as Plinia and
Siphoneugena (Eandrum & Kawasaki, 1997; Proen-
ya, 1990). Neomitranthes is ultimately distin¬
guished from Plinia by its calyptrate calyx and the
ovary with generally more than 2 ovules per locule,
and from Siphoneugena by the persistence of a ca¬
lyx tube with the staminal ring in the inner surface
after anthesis.

The taxonomic value of the opening of the calyx
has been severely challenged (Landrum, 1984) and
can be considered at best a very weak generic char¬
acter; the persistence or not of a calyx tube as a
useful character has not yet heen properly assessed.

1 his situation points to an impasse in establishing
definite generic boundaries; it is plausible that fu¬
ture studies may relegate Neomitranthes to synon¬
ymy either under Plinia or Siphoneugena — if not
merge these three genera into one inclusive genus,
in this case Plinia.

I he number of species in Neomitranthes is un¬
known. Landrum and Kawasaki (1997) estimated
fewer than 5 species, a number surely too conser¬
vative, and Mattos, in a series of papers (Mattos,
1981, 1989, 1990. 1997), attained a list of 19 spe¬
cies, an excessive number since some of these spe¬
cies are surely synonyms or are better placed in
other genera (e.g.. A', maria-aemiliae (I). Legrand)
Mattos is better treated as Myrceugenia ovalifolia
((). Berg) Landrum and N. hatschbachii (I). Le¬
grand) Mattos as Myrceugenia gertii Landrum; see
Landrum, 1984). Ihe species of Neomitranthes oc¬
cur mainly in coastal forest formations from north¬
eastern to southern Brazil, from the states of Bahia
to Rio Grande do Sul.

Neomitranthes obtusa Sobral & Zambom, sp.
nov. IA PE: Brazil. Espirito Santo: Guarapari,
Parque Estadual Paulo Cesar Vinha, 28 July
1998. 0. Zambom & A. Assis 338 (holotype,
VIES; isotype, MBM). Figure I.

Volume 12, Number 1
2002

Sobral & Zambom
Neomitranthes obtusa from Brazil

113

114

No von

Species N. glomeratae proxima, a qua foliis crassis, ob-
tusis, margine forte revoluta, nervis lateralis usque 16,
inconspieuis, petiolis brevioribusque recedit.

Treelet 3—4 ni high. Plants glabrous. Bark
smooth, exfoliating. Leaves elliptic to elliptic-ob¬
long, coneolorous to slightly discolorous, charta-
ceous to coriaceous, (45)65-90 X 30—50 mm, the
length/width ratio 1.5— 2.5:1, sometimes slightly un¬
equal in the same pair; apex obtuse or rounded;
base obtuse or rounded, rarely the very base some¬
what acute; central vein strongly convex at the ad-
axial surface but scarcely convex, plane, or even
slightly sulcate abaxially; secondary nerves 8 to 16
pairs, scarcely evident and occasionally convex
adaxially, hardly or not at all visible abaxiallv; mar¬
ginal vein 1.2—2 mm from the margin, the margin
itself strongly revolute; petioles 1.3—2 X 0.5— 0.7
mm. Inflorescences axillary or ramiflorous racemes,
1.5—2 X 1 mm, 4- to 6-florous, the flowers whitish
at anthcsis; bracts rounded-ovate, eiliate, 0.6— 0.8
X 0. 6-0.7 mm; pedicels 1.3—2 X 0.6— 0.7 mm;
bracteoles ovate, eiliate, occasionally carinate, de¬
ciduous at anthesis, equaling the bracts; buds glo¬
bose to obovate, 3-5 X 2. 5-3. 5 mm, apiculate, the
apiculum up to 5 mm long and eventually with a
tuft of hairs up to 0.1 mm long at the tip; calyx
completely fused, tearing irregularly at anthesis
and occasionally forming one larger, ± rounded
lobe up to 2 mm diam. (“calyptra”); calyx tube 2—3
mm deep; petals absent or 1, generally adnate to
the larger calyx lobe, rounded or elliptic, 1.5—3 X
1—2 mm, slightly eiliate; stamens white or creamy,
about 80, up to 4 mm long; anthers globose, 0.3-
0.4 X 0.3 mm; style up to 7 mm long; ovary 2(3)-
locular with 1 or 2 centrally attached ovules per
locule. Fruits globose, 1- or 2-seeded, 10-12 mm
diam., crowned by the calyx tube, vinaceous when
ripe. Seeds with testa easily detachable; embryos
reniform, up to 8 X 5 mm, with 2 distinct and
sometimes very unequal plano-convex cotyledons
and occasionally a hypocotyl 2—2.5 X 1 mm.

Phenology. Flowers collected in June and Julv;
fruits collected in September.

Distribution. The new species has been col¬
lected only in the municipality of Guarapari, in the
state of Espfrito Santo.

Ecology. Neomitranthes obtusa is a treelet of
restinga formations (coastal forests growing on
sandy soils).

Etymology. The specific epithet refers to the

plant's obtuse leaves. Elsewhere in the genus, acu¬
minate or rostrate leaves are more commonly found.

Neomitranthes obtusa is apparently related to N.
glomerata (1). Legrand) 1). Legrand, a species from
southern Brazilian coastal forests ranging from Sao
Paulo to Santa Catarina, from which it can be dis¬
tinguished by the following characters:

1. Leaves with apex obtuse, margins strongly revo¬
lute; secondary nerves 8 to 16 pairs, scarcely vis¬
ible on both faces, generally less evident abaxi¬

ally; texture chartaceous or occasionally
coriaceous; petioles up to 2 mm long; ovary with
inner walls of locules glabrous; restingas of Es-
pfrito Santo . Neomitranthes obtusa

I . Leaves with apex acuminate or attenuate, mar¬
gins scarcely or not at all revolute; secondary
nerves 20 to 25 pairs, visible on both sides, gen¬
erally visible abaxially; texture at most charta¬
ceous; petioles 5-10 mm long; ovary with inner
walls of locules occasionally pilose; tall coastal
forests (mata atlantica) of Sao Paulo to Santa Ca¬

tarina . Neomitranthes glomerata

Paratypes. BRAZIL. Espfrito Santo: Guarapari,

Parque Estadual Paulo Cesar Vinha, 30 July 1996, Gomes
2192 (1CN, VIES), 9 June 1998, Assis 500 (1CN, VIES),
28 July 1998, Zambom & ,4.s\sis 339 (MO, VIES), 9 Sep.
1999, Assis 723 (1CN, VIES).

Acknowledgments. We thank Andre Assis for
his kind cooperation in collecting additional ma¬
terial of this species, and Victoria Hollowell and
two anonymous reviewers for valuable and patient
observations that improved our paper.

I .iterature Cited

Landrum, L. K. 1984. Taxonomic implications of the dis¬
covery of calyptrate species of Myrceugenia (Myrta-
ceae). Brittonia 36: 161-166.

- & M. L. Kawasaki. 1997. The genera of Myrta-

eeae in Brazil: An illustrated synoptic treatment and
identification keys. Brittonia 49: 508—536.

Legrand, C. I). & R. \1. Klein. 1977. Mirtdceas: Campo-
manesia, Feijoa, Hritoa , Myrrhinium, Hexachlamys, Si-
phoneugena, Myrcianthes, Neomitranthes , Psidium. Pp.
572—730 in R. Reitz (organizer). Flora Ilustrada Catar-
i Dense. I tajaf.

Mattos, J. R. 1981. Novidades taxonomicas em plantas do
Brasil. Loefgrenia 76: 1—3.

- . 1989. Novidades taxonomicas em Myrtaceae —

VI: SecySes do genero Neomitranthes Legr. Loefgrenia
95: 1-2.

- . 1990. Novidades taxonomicas em Myrtaceae —

VI (sic). Loefgrenia 99: 1—6.

- . 1997. Novidades taxonomicas em Myrtaceae —

XIV. Loefgrenia 1 10: 1—4.

Proenya, C. 1990. A revision of Siphoneugena Berg. Fid-
inburgh J. Bot. 47: 239-271.

Swartzia mexicana (Fabaceae, Swartzieae), a New Species from

Oaxaca, Mexico

Mario Sousa S.

Departamento <le Botanica, Instituto de Biologfa, Universidad Nacional Autononia de
Mexico, Apdo. Postal 70-367, 04510 Mexico, D.F. Mexico. sousa@servidor.unam.mx

Rosaura Grether

Departamento de Biologia. Division de Ciencias Biologicas y de la Salud, Universidad
Autonoma Metropolitana-Iztapalapa, Apdo. Postal 55-535, 09340 Mexico, D.F. Mexico.

rogg@xanu m . uam . nix

Abstract. Swartzia mexicana. a new legume
species from the State of Oaxaca, Mexico, is de¬
scribed and illustrated. It is presently known only
from the districts of Ixtlan and Tuxtepec. The new
species belongs to section Swartzia, subsection
Swartzia, and is assigned to series Tounateae. With¬
in that series, it is closely related to S. sumorum
by the leaflets not prominently reticulate, the style
uncinate, and the stigma truncate, as well as by the
legume elliptic in shape. A key to distinguish the
new species from S. sumorum and S. wurdackii is
provided.

Resum EN. Se describe e ilustra Swartzia mexi¬
cana, una especie de leguminosas nueva para la
ciencia, del estado de Oaxaca. Mexico. Hasta ahora
se conoce solo de los distritos de Ixtlan y luxtepec.
La especie nueva corresponde a la sect. Swartzia,
subsect. Swartzia y se incluye en la serie Touna¬
teae. Dentro de esa serie, se relaciona estrecha-
mente con .S', sumorum por los foliolos sin nervacion
reticulada prominente, el estilo uncinado y el es-
tigma truneado, asf como por la legumbre de forma
elfptica. Se presenta una clave para distinguir a la
especie nueva de S. sumorum y de S. wurdackii.

Key wor<ls: Fabaceae, Mesoamerica, Mexico,
Oaxaca, series Tounateae, Swartzia.

Swartzia is a neotropical genus of about 155 spe¬
cies, most of them found in South America. Only
two species are known to occur elsewhere, in trop¬
ical Africa, according to Cowan (1981)- Species of
the genus vary from small trees or shrubs (1.5— )3—
18 m high, such as S. invenusta Barneby and S.
alato-sericea Barneby, to large trees up to 35-40 m
high, such as S. gigantea R. S. Cowan and .S'. po-
lyphylla DC. Some slender trees, such as .S. pal-
ustris Barneby, have occasionally sarmentose
branches, but the only species with a clambering

habit is S. froesii R. S. Cowan. The genus is a com¬
mon component of lowland forests, especially along
rivers anil in swampy forests, but in the Andean
area it can be found from sea level up to 2100 m
elevation, as S. amplifolia Harms var. colombiana
R. S. Cowan (Cowan, 1968; Barneby, 1991. 1996).

In Mexico, Swartzia is found from Nayarit to
Chiapas along the Pacific slope, and it extends
eastward to Veracruz, Tabasco, and the \ucatan
Peninsula. Swartzia habitats include tropical rain¬
forests, semideciduous and deciduous forests, as
well as cloud forests. Besides Swartzia mexicana,
three other species of the genus are known to occur
in Mexico: S. cubensis (Britton & Wilson) P. C.
Standley (Chiapas, Tabasco, Campeche, Yucatan,
and Quintana Roo), S. myrtifolia J. E. Smith (Ve¬
racruz, Oaxaca, Chiapas, Tabasco), and .S’, simplex
(Swartz) Sprengel (Nayarit, Jalisco, Miehoacan,
Guerrero, Veracruz, Oaxaca, and Chiapas). In the
State of Oaxaca, the latter two are allopatric to the
new species; they are found along the Pacific slope
in the districts of Jamiltepec, Pochutla, and Juclii-
tan, growing mainly in lowland forests, 200 to 780
m.s.m., although S. simplex has been collected up
to 1660 m elevation in the District of Pochutla.

Swartzia cubensis also belongs to series Touna-
tae: it is distinguished from S’, mexicana by its mar-
ginate to winged leaf rachis, shorter inflorescences,
and yellow to orange, long stipitate, subglobose or
ovate, rostrate fruits. Swartzia myrtifolia and .S', sim¬
plex are members of series Possira. They are dis¬
tinguished from the new species by their petalif-
erous flowers, by the narrowly margined or winged
petiole and leaf rachis, and by the subglobose or
oblong fruits, very similar to those of S. cubensis.

Mesoamerica is a more diverse region: about 9
to 12 species of Swartzia have been found from
northern Oaxaca and the Isthmus of Tehuantepec,

Novon 12: 115-119. 2002.

116

Novon

figure 1. Swartzia mexicana M. Sousa & H. Grether. — A. Branch with leaves and inflorescences. — B. Close-up of
stipules. — C. A group of inflorescences in bud. — I). Floral bud and bract. — E. Flower. — F. Stamens. — G. Gynoecium

Volume 12, Number 1
2002

Sousa & Grether

Swartzia mexicana from Mexico

117

and in Tabasco, Chiapas, and die Yucatan Penin¬
sula in Mexico, to Panama. The new species is de¬
scribed from the northwestern phytogeographic lim¬
it of the Mesoamerican region, as part of the
ongoing Flora Mesoamericana project coordinated
by the Missouri Botanical Garden, the Instituto de
Biologfa, Universidad Nacional Autonoma de Mex¬
ico, and The Natural History Museum (London)
(Davidse et al., 1994).

Swartzia mexicana M. Sousa & B. Grether, sp.
nov. TYPE: Mexico. Oaxaca: Distrito de Ixtlan.
Municipio de Ixtlan de Juarez, 12 km NE de
Vista Hermosa, carretera de Ixtlan a Valle Na¬
cional. 17 Apr. 1982. R. Cedillo-Trigos 1234
& R. Torres-CoUn (holotype, MEXU; isotypes,
BM, MEXU, MO, UAM1Z). Figures 1-3.

Arbor 20-40 m alta; stipulae 1-2.5 mm longae, 0.4-1
mm latae, axis foliaris (petiolus cum rachis) 8— 18(— 20) cm
longus; foliola (5— )7, lanceolato-oblonga, raro elliptica,
utrinque glabra, apice acuminata; racemi cauliflori (7— )9-
16(— 20) cm longi, bracteae 0.6— 0.9 mm longae; pediceli
3-7 mm longi, alabastra ad 6X5 mm; legumen oblique
ellipticum, base atenuata, verrucosum, brunneum, semine
uno. elliptico.

Trees 20—40 m high, the bark striate to rugose,
grayish brown, with prominent lentieels; branchlets
strigulose, rufous, glabrescent. Leaf buds strigose,
rufous; young leaves purple, strigulose; mature
leaves green, glabrous; stipules 1-2.5 X 0.4—1 mm,
subulate to deltoid, strigose to strigulose, decidu¬
ous; petioles ( 1 — )2.5 — 5 cm long, terete to flattened
or slightly sulcate on the upper surface, glabrous;
rachis (3.5— )5.5 — 13(— 1 5) cm long, flattened to sul¬
cate on the upper surface, very sparsely strigulose
to glabrous, not stipellate; leaflets (5 to)7. the pet-
iolules 3—5 mm, sulcate, glabrous, the blades of the
lowermost pair (3.5— )4. 5— 10 X (2— )2.5— 4.5(— 5.5)
cm, the other pairs (6— )7— 14.5(— 16) X (2.5— )3— 5(—
5.5) cm. lanceolate-oblong, rarely elliptic, glabrous
on both surfaces, venation tenuous, the main vein
prominent on the lower surface, very sparsely stri¬
gulose at the base, glabrescent, the base cuneate,
the margins entire, thickened, sometimes very
slightly revolute, the apex acuminate. Inflorescenc¬
es racemose, (7— )9— 16(— 20) cm long, strigose, ru¬
fous, cauliflorous, the young racemes growing on
very compressed brachyblasts, and disposed in
groups of (3 to)6 to 9(to 12), rarely axillary, and
solitary; peduncle (0.3— )0.5— 1.5(— 2) cm long, the
peduncle and the floriferous axis strigulose, rufous;

bracts 0.6— 0.9 X 0.4— 0.6 mm, deltoid, strigose, ru¬
fous, caducous; bracteoles lacking; pedicels 3—7
mm long, strigose, rufous; floral buds 3.5—6 X 3—
5 mm, almost globose, strigulose, rufous, the apex
shortly apiculate. Flowers apetalous, calyx 5—5.5
mm long, strigulose, with 4(or 5) irregular lobes,
free or shortly fused at the base, green, persistent;
corolla lacking; stamens numerous (29 or 30), the
longer filaments 6.5-8 mm, free to base, the smaller
ones 4-5 mm, white, exceptionally fused in groups
of 2 or 3 at the base (Fig. IF), glabrous, anthers
1.5—2 X 0.8— 1.2 mm, oblong to elliptic, nearly bas-
ifixed; gynoecium glabrous, ovary stipitate, 4-6 mm
long, including the gynophore, the latter 2 mm long,
sparsely strigulose at the base, with 4 or 5 cam-
pylotropous ovules, obliquely disposed, style 1 .5—2
mm, arcuate, glabrous, stigma truncate to obliquely
truncate. Legume 4.5—5 X 2—2.3 cm, obliquely el¬
liptic, the base attenuate, the valves 2-5 mm thick,
ligneous, verrucose, dark brown, 1 -seeded, the
apex obliquely obtuse. Seed 3.2 X 1.7 X 1.4 cm,
elliptic, the testa chartaceous, rugose, brown, bril¬
liant, the aril probably red. thin.

Common names. “Jarrito silvestre," “palo de
jarrito de monte.”

Distribution and habitat. As far as we know,
the new species is endemic to northern Oaxaca. It
occurs in cloud forest with Lauraceae and Rheedia,
premontane wet forest, and montane tropical rain¬
forest. at elevations of 725—1300 m.

Phenology. It has been collected in flower dur¬
ing March and April, and the fruits have been
found in May.

The new species belongs to section Swartzia,
subsection Swartzia, and is assigned to series Toun-
ateae, which is characterized by the floral buds
usually 5 mm or less in diameter, as well as by the
lack of a petal and bracteoles.

Swartzia mexicana is closely related to S. su-
morurn A. R. Molina. Both species share the leaf¬
lets not prominently reticulate, the style uncinate,
and the stigma truncate, as well as the legume el¬
liptic in shape. It is distinguished from S. sumorum
by the narrower leaflets, the larger stipules and flo¬
ral buds, the larger number of leaflets, as well as
the longer racemes and pedicels (see key). The new
species is also related to S. wurdackii R. S. Cowan
by the leaf rachis lacking wings or stipels, and the
gynoecium glabrous; the latter can be distinguished
by the longer stipules, the leaflets with prominent

with campylotropous ovules. — H. Legume.
Hernandez 142).

I. Seed. (A-G, R. Cedillo-Trigos 1234 & R. Torres -Colin; H, I, P. Osorio-

118

Novon

Figure 2. Swartzia mexicana M. Sousa & l{. Grether. Close-up of racemes, floral buds, and flowers (photographed
from R. Cedillo-Trigos 1234 <V R. Torres-CoUn).

reticulate venation, and the shorter racemes (see
key).

Series Tounateae comprises 22 species, includ¬
ing Swartzia mexicana. Other species of this group
willi leal raohis lacking wings or stipels but having
a densely pubescent gynoecium are Swartzia ama-
zonica Moore occurring in Brazil, S. bannia Sand-
with known from Suriname and Guyana, and .S', car-
ibaea Grisebaeh from the West Indies.

The new species can be distinguished by adding
it to Cowan’s key to species of series Tounateae

(1968: 23—24) in his treatment for Flora Neotropi-
ca, at couplet I I (p. 23), in the following way:

I I. Gynoecium glabrous or sparsely strigose basally.

12. Stipules 5 mm long; leaflets with prominent
reticulate venation; racemes 5—6.5 cm long;

Venezuela . 3. wurdackii

12. Stipules 0.1— 2.5 mm long; leaflets with one
prominent vein and tenuous, pinnate vena¬
tion on ihi' lower surface; racemes (7— )8—
16(~20) cm long.

13. Stipules 0.4— 0.5 mm long; leaflets (3
to)5; racemes 8-10 cm long; floral buds

Volume 12, Number 1
2002

Sousa & Grether

Swartzia mexicana from Mexico

119

Figure 3. Swartzia mexicana M. Sousa & R. Grether. Close-up of open fruit and seed (photographed from R. Osorio-
Herndndez 142).

3 X 3 mm; pedicels 1-1.5 mm long; 2,
rarely 1 , globose seeds, 1—1.5 cm diam.;

[Nicaragua . S. sumorum

13. Stipules 1-2.5 mm long; leaflets 7(less
often, to 5); racemes (7— )9— 16(— 20) cm
long; floral buds 3.5-6 X 3-5 mm; ped¬
icels 3-7 mm long; 1 elliptic seed, 3.2
X 1.7 X 1.4 cm; Mexico ... .S', mexicana

Raratypes. MEXICO. Oaxaca: Distrito de Ixtlan,
Mun. Comaltepec along trail that descends from Hwy. 175
at 1280 m elevation (Puerto Antonio) to settlement of Cua-
je. 2-6 Dec. 1994, H. Hoyle & S. Acosta 3960 (MEXU,
MO, OAX), H. Hoyle & S. Acosta 4011 (MEXU, MO);
Puerto Antonio, 25 Mar. 1989, R. Lopez-Luna 0438
(MEXU); Distrito de Tuxtepec, Mun. San Felipe Usila.
Monte Pan, 3.1 km en Ifnea recta al SE (135°) de Santa
Cruz Tepetotutla, I I Apr. 1994, P. Osorio-Herndndez 37
(MEXU), 7 May 1994, R. Osorio-Herndndez 142 (MEXU),
R. Osorio-Herndndez 152 (MEXU); 5.1 km en Ifnea recta
al SE (154°) de Santa Cruz Tepetotutla, 29 Mar. 1995, A.
Rincon-Gut ierrez 6 07. C. Gallardo & E. Perez-Garcia
(MEXU).

Acknowledgments. We express our gratitude to
Fernando Cliiang. Institute de Biologfa, UNAM, for
the Latin translation of the diagnosis, to Lawrence
M. Kelly, Institute de Biologfa, UNAM, for His re¬

view of the English version of the manuscript, to
Gloria Andrade, Institute de Biologfa, UNAM, and
to Elvia Esparza, Institute de Biologfa, UNAM, for
tin1 excellent illustration. Special thanks are given
to Victoria C. Hollowed, Missouri Botanical Gar¬
den. and David A. Neill. Missouri Botanical Gar¬
den. Herbario Nacional de Ecuador, for critical
comment improving the manuscript.

Literature Cited

Barneby, R. C. 1991. Notes on Swartzia (Leguminosae:
Swartzieae) preliminary to the Flora of the Venezuelan
Guayana. Ann. Missouri Hot. Gard. 78: 1 17— 183.

- . 1996. Neotropical Fabales at NY: Asides and

oversights. Brittonia 48: 174—187.

Cowan, R. S. 1968. Swartzia (Leguminosae, Gaesalpinioi-
deae, Swartzieae). Flora Neotropiea, Monogr. 1: 1—228.

- . 1981. Swartzieae. Pp. 209—212 in R. M. Polhill

& P. H. Raven (editors). Advances in Legume System-
atics, Part 1 . Royal Botanic Gardens, Kew.

Davidse, G„ M. Sousa & A. 0. Chater (editors). 1994.
Flora Mesoamericana 6: XII 1 — XIV; Mapa del area de la
Flora Mesoamericana. Universidad Nacional Autonoma
de Mdxico. Mexico, D.F.; Missouri Botanical Garden,
St. Louis; The Natural History Museum, London.

Rubiacearum Americanarum Magna Hama Pars VI: New Species
of and Morphological Notes on Psychotria subg. Psychotria
(Psychotrieae) from Mesoamerica and Western South America

Charlotte XL Taylor

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

( !!harlotte.Taylor@mobot.org

Abstract. Eight new Neotropical species of Psy¬
chotria subg. Psychotria are described and illus¬
trated here: P. cerrocoloradensis of western Panama
and P. matagalpensis of northern Nicaragua have
both previously been confused with P. sylvivaga
Standley and P. ch iru/uina Standley; P. cornejoi of
western Ecuador has been previously confused with
P trivial is Rushy; P. jefensis of Costa Rica and Pan¬
ama and P. orosioides of Costa Rica have both been
previously confused with P. orosiana Standley; P.
lorenciana of southern Mexico and Guatemala lias
been variously confused with P. grandis Swartz, P.
flava Oersted ex Standley, and P costivenia Grise-
baeh; and P. romolerouxiana of Amazonian Ecuador
and Peru and P. sacciformis of lowland Colombia,
Ecuador, and Peru have both previously been con¬
fused with P. horjcnsis Kunth. The interpretations
of stipule morphology and the arrangement of the
flowers on the higher-order inflorescence axes as
taxonomically informative characters have varied
among previous authors and are re-evaluated here.

hey words: Neotropies, Psychotria, Rubiaceae.

Psychotria L. (Psychotrieae) in its broad sense
comprises approximately 1000 species of shrubs
and small trees (or much less often herbs, climbers,
or epiphytes) found throughout the tropics. This ge¬
nus is characterized bv its usually woody habit; its
relatively small, entomophilous, usually white or
cream-colored flowers that are typically distylous;
its valvate corolla lobe aestivation; its inferior ovary
with usually two loeules, each with one basal ovule;
and its drupaceous fruits with usually two plano¬
convex pyrenes. Recently, several studies employ¬
ing morphological and molecular characters have
indicated that Neotropical Psychotria as it is cur¬
rently circumscribed is systematically more com¬
plex than has usually been appreciated (Taylor,
1996; Nepokroeff et ah. 1999; Andersson & Rova,
1999; Piesschaert, 2001). These studies have dis¬
tinguished within traditional Psychotria several lin¬
eages that are not all closely related. The species
discussed and described below all belong to Psy-

Novon 12: 120-132. 2002.

chotria subg. Psychotria ; all of these studies have
considered this a monophyletic group (Steyermark,
1972; Hamilton, 1989; Taylor, 1996; Nepokroeff el
ah, 1999; Andersson & Rova, 1999; Piesschaert,
2001).

Neotropical Psychotria subg. Psychotria is char¬
acterized by the distinctive color of its dried spec¬
imens. which usually are strongly tinged with
brown, red-brown, red-gray, gray, or gray-green; its
stipules of widely varied form but that are usually
quickly caducous exposing a ring of well-developed
infrastipular colleters (i.e., glandular trichomes),
which are up to 3 mm long and typically persistent
on the stem as a chestnut-brown fringe; its white
flowers that are relatively short, with tubes 1-8 mm
long, and pubescent in the throat and/or at the
point of stamen insertion; and its orange to red
fruits that bear two planoconvex, usually longitu¬
dinally ridged pyrenes. The pyrenes are character¬
ized by an alcohol-soluble reddish pigment and a
lack of preformed germination slits, and both of
these characters are shared with paleotropical spe¬
cies of Psychotria subg. Psychotria (Petit, 1964;
Nepokroeff et al., 1999; Piesschaert, 2001). In con¬
trast, the other two large Neotropical groups within
Psychotria s.h, section Notopleura [ = Notopleura
(Bentham & Hooker f.) Bremekamp: Taylor, 2001b]
and subgenus Heteropsychotria Steyermark, have
dried specimens with a usually green or brown-
green color, stipules of various forms but that are
usually persistent and without colleters or with
these caducous or drying clear, fruits that may be
orange or rod al maturity but are more often white,
blue, or purple, and pyrenes two to five per fruit
and widely varied in form but lacking alcohol-sol¬
uble red pigments and bearing preformed germi¬
nation slits.

Below I re-evaluate some morphological char¬
acters that have and also have not been emphasized
by previous authors to distinguish among species
of Psychotria subg. Psychotria, and describe sev¬
eral new' species.

Volume 12, Number 1
2002

Taylor

Psychotria subg. Psychotria

121

Reconsideration of some Morphol.ogical
Characters

Hamilton (1989) monographed 61 species of
Psychotria subg. Psychotria found from Mexico
through Panama, and contributed significantly to
the knowledge of the Neotropical species of this
group in several aspects. He presented the first
careful consideration of morphological characters
with taxonomic significance in this group, and for
the most part his work is extremely useful. He re¬
lied strongly on details of inflorescence arrange¬
ment, leaf venation patterns, anti specimen drying
colors to separate species and delimit species
groups, with much success. However, the use of
these characters is problematic at times. For ex¬
ample, in some species there is wide variation in
the pattern of leaf venation, as Hamilton noted in
his descriptions (e.g., P panamensis Standley).
Also, the specimen drying color may depend on
preparation of the material, for example, treatment
of it with alcohol, as much as on the biology of the
plant, and specimens of subgenus Heteropsychotria
occasionally show the same drying colors for either
of these reasons. And, Hamilton’s characterization
of inflorescence arrangement is based on fully de¬
veloped inflorescences, so plants with poorly de¬
veloped or broken inflorescences may be difficult
to identify.

In the years since the publication of Hamilton’s
treatment, numerous additional good collections of
these plants have been made, contributing much
new morphological and floristic information. Study
of Psychotria subg. Psychotria for the upcoming
Flora Mesoamericana treatment incorporated these
specimens and found that some of the characters
Hamilton used, in particular flower arrangement,
are more variable and therefore less informative
than has been assumed while others, in particular
stipule morphology, are more informative than has
been recognized.

Much work remains to be done to understand
inflorescence structure in Psychotria, and a de¬
tailed analysis will not be presented here, but some
aspects of Hamilton’s work require comment in the
light of these new collections. In particular, Ham¬
ilton (1989) distinguished two general floral ar¬
rangements: “flowers sessile in glomerules” in con¬
trast with “flowers pedicellate” in cymules. which
he used to separate species and species groups.
However, this distinction can be problematic to ap¬
ply in many species of Psychotria. The flowers of
these species are arranged in generally dichasial
cymules that bear one sessile or subsessile terminal
flower subtended by one or two developed higher-

order axes, which in turn produce another terminal
flower or group of flowers and frequently also a still
higher-order set of axes that in turn produce anoth¬
er flower or cymule. The individual flowers may be
borne separately along the axes, or in some species
are consistently arranged in glomerulate groups of
two to several flowers. In this latter case the glo¬
merulate arrangement recognized by Hamilton is
usually clear. In the former case, the terminal flow¬
er as well as the flowers produced on the higher-
order axes are sometimes clearly pedicellate, so
their arrangement is clear. However, in many spe¬
cies the terminal flowers are subsessile and the ar¬
rangement is ambiguous for the flowers at the ends
of the higher-order axes, because they are separat¬
ed along these axes and the structures that support
them can be, and variously have been, considered
pedicels or higher-order axes. Hamilton (1989) dis¬
tinguished several species groups based on whether
the flowers are pedicellate or sessile, but he did not
clarify his definition of these higher-order axes and
seems himself to have called them pedicels in some
species groups but axes in others. The position of
the flowers produced on these higher-order axes has
also been variously described because sometimes
the higher-order axes elongate as the fruits develop,
so the flowers at anthesis are subsessile in glom¬
erules while the fruits are clearly separated on the
tardily developing axes, or pedicels.

Below and in the upcoming Flora Mesoameri¬
cana treatment, the term “sessile” is applied to
flowers that are attached directly (i.e., without a
pedicel) to developed higher-order axes at anthesis
and have the floral bracts borne directly at the base
of the hypanthium (i.e.. ovary portion; Fig. 2b).
Pedicels can be distinguished from higher-order
axes by the location of floral bracts at their bases
or in their lower portions but not next to the flower
(Figs. 2H. 3D).

Another complicating factor in the interpretation
of “sessile” vs. “pedicellate” flowers in this group
is the presence in some species of a stipe on the
fruit (e.g., Psychotria panamensis) even though the
flowers are sessile at anthesis. These structures
have been called pedicels by most previous authors
but are interpreted here instead as a stipitate pro¬
longation of the fruit base, because the elongation
begins with and parallels fruit development; be¬
cause this structure elongates distal ly leaving the
floral bracts at its base; and because the zone of
fruit dehiscence is located at the base of this struc¬
ture rather than at its connection to the fruit base.

Hamilton described and very nicely illustrated
(1989: fig. 2) the variety of stipule forms found in
this group, and for most species he described the

122

No von

stipules accurately according to li is classification.
He commented (1989: 68) that “Stipules can pro¬
vide valuable information as to interspecific rela¬
tionships but prove almost worthless as a source of
key characters, since in this subgenus they are usu¬
ally caducous, or deciduous,” and did not empha¬
size stipule form in the keys or descriptions. How¬
ever, stipule morphology actually appears to be
highly consistent within a species, highly informa¬
tive taxonomically, and usually evident on herbar¬
ium specimens (and always evident on living
plants). Below and in the upcoming Florn Mesoam-
ericana treatment, stipule morphology is strongly
emphasized in the characterization of species.

Hamilton (1989) classified stipules as “sheath¬
ing” when there was any degree of union across the
intrapetiolar portions of the stipules, from very
short fusion at the bases to complete fusion into a
single continuous structure, and whether or not the
united portion breaks apart easily when the leaves
expand (e.g., F. carthagenensis Jacquin) or remains
permanently fused (e.g.. P. panamensis). Below and
in lh(“ upcoming Flora Mesoamericana treatment, a
distinction is made among his various forms of
“sheathing” stipules to recognize the following
types:

• stipules that are interpetiolar and also shortly
and/or weakly united in the intrapetiolar portion
and quickly separate along this intrapetiolar line
(e.g., P. carthagenensis ; Hamilton 1989: fig. 2a,
e), here called “interpetiolar and shortly intra¬
petiolar”;

• stipules that are fused around the stem into a
tube that is open at the top, and fall off as a tube
that is complete or sometimes split open on one
side (e.g., P. chagrensis Standley; Hamilton
1989: fig. 2g). here called “tubular”;

• and stipules that are completely fused into a con¬
ical cap that falls off as a single unit (e.g.. P
trichotoma M. Martens & Galeotti; Hamilton
1989: fig. 2d), here called “calyptrate.”

Hamilton (1989) used the term “hiaristate” rath¬
er generally for any stipules with two aristas or nar¬
row lobes in some part, combining under this term
both the condition with the lobes solitary on each
interpetiolar side for a total of two at each node and
the condition with two lobes on each interpetiolar
side* for a total of four at each node (e.g., Hamilton,
1989: figs. 2e and 2g, both dr ‘scribed in the* re¬
spective species descriptions as “biaristate”). Be¬
low and in the upcoming Flora Mesoamericana
treatment, the varied stipule forms in Psychotria
subg. Psychotria are all considered to be derived
from two interpetiolar. bilobed stipules, so (lit* num¬
ber of aristas or lobes is counted with respect to

each interpetiolar portion. Thus, stipules with two
aristas at each node (Hamilton, 1989: fig. 2g) are
here considered different from those with four aris¬
tas at each node (Hamilton, 1989: fig. 2e). No sin¬
gle Neotropical species of Psychotria subg. Psycho¬
tria has yet been seen with both of these stipule
forms.

New Species

Psychotria eerrocoloradensis Dwyer ex C. M.
Taylor, sp. nov. TYPE: Panama. Chiriqm: Cerro
Colorado, on road. 31.6 km from Bio San Felix
bridge, 1690 m. 15 July 1976, G. A. Sullivan
332 (holotype, MO-2616700). Figure 1A, B.

Haec species a Psychotria amplifronde pedunculo long-
iore, a // rujiramea inflorescentia ampliore atque a P. syl-
vivaga bracteis floralibus pedicellis et limbo ealycino lon-
gioribus distinguitur.

Shrubs to 1.5 m tall; stems densely lanuginose-
hirtellous to glabrescent. Leaves elliptic, 10-22 X
3—10 cm, chartaceous, at apex acuminate, at base
euneate to obtuse or truncate, above glabrous, be¬
low densely strigillose to sericeous on veins and
glabrous to sparsely puberulous on lamina; second¬
ary veins 10 to 16 pairs, united in a somewhat loop¬
ing submarginal vein, abaxially without domatia in
axils, adaxially venation plane or sometimes a little
impressed, abaxially costa prominent and second¬
ary and tertiary venation prominulous; petioles 1.4—
3.5 cm long; stipules caducous, interpetiolar and
shortly intrapetiolar, lanceolate, 13—17 mm long, at
apex acute and shortly bidentate with the teeth to
3 mm long and ligulate to triangular, abaxially
densely lanuginose to strigillose except glabrescent
toward margins. Inflorescences terminal, strigillose
to lanuginose, peduncle 3.5— 7.5 cm long, branched
portion 6—14 X 7—20 cm. pyramidal; secondary
axes 2 per node, branched; bracts puberulous, 1.5—
8 mm long, narrowly triangular, floral bracts 1.5—3
mm long, ovate to lanceolate; pedicels 0—0.5 mm
long; flowers sessile to subsessile in dichasial to
umbelliform cymules of 3 to 7; hypanthium 0.8-1
mm long, turbinate, glabrous; calyx limb 1 .8— 2 mm
long, glabrous, lobed for V4—V2. lobes triangular, cil-
iolate; corolla tubular- funnelform, white, externally
glabrous, internally glabrous except densely pilo-
sulous at stamen insertion, tube ca. 3.5 mm long,
lobes 5. ca. 1.5 mm long, triangular; anthers ca. 1
mm long, included; stigmas ca. 0.8 mm long, ex-
serted. Infructescences similar in shape and size to
inflorescences; immature fruits ellipsoid to obovoid.

Habitat , distribution , and phenology. In wet
montane forest. 1630—1700 m, western Panama;
collected in flower in July and August.

Volume 12, Number 1
2002

Taylor

Psychotria subg. Psychotria

123

Figure 1. A, B. Psychotria cerrocoloradensis C. M. Taylor. — A. I* lowering stern. — B. Mower at anthesis. C E.
Psychotria lorenciana C. M. Taylor. — C. Flowering stem. — D. Flower at anthesis. — F. Stipule. A, based on Croat
69148 ; B. based on Sullivan 382; C, E, based on Contreras 10014; D, based on Lundell & Contreras 19332. A, C to
5-cm scale; B, D, E to 5-mm scale.

124

No von

John Dwyer originally recognized this species as
new and proposed this epithet, which refers to t he
collection locality of the Panamanian specimens.
Psychotria cerrocoloradensis is similar to P. amplif-
rons Standley, which differs by its shorter pedun¬
cles 0.3—2 cm long; to P. rufiramea Standley, which
differs hy its smaller inflorescences with the
branched portion 1.5-4 X 2-^ cm; and to Psycho¬
tria sylvivaga, which differs hy its shorter floral
bracts 0.5— 1.5 mm long, longer pedicels 0.5—2 mm
long, and shorter calyx limbs 1.2— 1.5 mm long. The
flowers seen are all similar to the long-styled form
of distylous Psychotria species in the arrangement
of their anthers and stigmas.

Paralypes. PANAMA, lincus del Toro: Cerro Colo¬
rado, 6.5 mi. W of Chame, steep forested slopes N of road,
8°35'N, 81°50'W, Croat 69148 (MO), Boras del Toro/
Chiriquf Border: Cerro Colorado, 1 1 .2 km along ridge
road from main road to Escopeta, Folsom 4888 (MO). Chl-
riqui: Cerro Colorado, along mining road 31.6 km beyond
bridge over Rfo San F6lix (10.6 km beyond turnoff to Es¬
copeta), Croat 87158 (MO).

Psychotria cornejoi C. M. Taylor, sp. nov. TYPE:
Ecuador. Guayas: 3 km E of Ol6n (5 km IN of
Manglarolto, on the beach), 75-190 m, 20
Dec. 1961. C. //. Dodson & L. R. Thien 1669
(holotype, MO-3849660). Figure 24-1).

Haec species a congeneris stipulis interpetiolaribus un-
iaristatis ac plerumque dense hirtellis atque foliorum sat
parvorum supra medium latiorum nervis secundariis
eamptodromis distinguitur.

Shrubs and small trees to 6 m tall; stems gla¬
brous. Leaves elliptic to usually obovate or oblan-
ceolate, 2.5—11.5 X 1.5—5 cm, papyraceous, at
apex acute to a little acuminate, at base cuneate to
acute and attenuate, adaxially glabrous, abaxially
glabrous except sometimes hirtellous on costa and
occasionally also secondary veins; secondary veins
4 to 6 pairs, not looping to interconnect, abaxially
without or often with pit domatia in axils of 2 or 3
apical pairs of secondary veins, domatia sacciform,
0.5— 1.5 mm long, a little elongated, generally ap-
pressed to blade, adaxially costa prominulous and
remaining venation plane, abaxially costa and sec¬
ondary veins prominulous and remaining venation
plane; petioles 1.5—20 mm long; stipules caducous,
interpetiolar, lanceolate to triangular, 3—4 mm long
(excluding aristas), glabrous to densely hirtellous
and often becoming glabrescent, acute to obtuse
and 1 -aristate. aristas 0.8—3 mm long, puberulous
to hirtellous. Inflorescences terminal, glabrous, pe¬
duncles 0.2—6 cm long, branched portion 2.5—9 X
3-10 cm, pyramidal, secondary axes 2 per node,
branched; bracts 0.3— 1.5 mm long, elliptic to tri¬

angular or linear, ciliolate; pedicels 1-2 mm long;
flowers pedicellate in umbelliform cymules of 2 to
5; hypanthium turbinate to cylindrical, ca. 0.8 mm
long, glabrous; calyx limb 0.5-0. 8 mm long, gla¬
brous, entire to denticulate; corolla funnelform,
white, externally glabrous, barbate in throat, lube
1.5—2 mm long, lobes 5, 1.5—2 mm long, narrowly
triangular; anthers ca. 1.2 mm long, exserted; stig¬
mas ca. 1 mm long, partially exserted. Fruits ellip¬
soid, ca. 5X4 mm, apparently red; pyrenes 2,
planoconvex, dorsally with ca. 5 rounded longitu¬
dinal ridges.

Habitat , distribution, and phenology. In pre-
montane dry forest and seasonal cloud forest at 50—

I 155 m. central western Ecuador; collected in flow¬
er in January, March, September, November, and
December, in fruit July through September and in
November.

I his species is distinguished by its combination
of interpetiolar stipules that are aristate and usually
densely hirtellous, its relatively small leaves that
are broadest above the middle, its secondary leaf
veins not looping to interconnect, and its elongated
sacciform domatia. In form these domatia are un¬
usual in this genus but somewhat similar to those
of Psychotria sacciformis of Amazonian Ecuador
and Peru; they are discussed further under that
species, below. The flowers seen are generally sim¬
ilar to the short-styled form of distylous Psychotria
species in the arrangement of their anthers and
stigmas, except the stigmas appear to be partially
exserted. The specific epithet honors Xavier Cor¬
nejo, an Ecuadorian botanist who is exploring ex¬
tensively in the western part ol the country and
documenting its flora, including this new species.

Paratypes. ECUADOR. Without location, “Coast
plain,’ Rimbach 99 (F). El Oro: Hacienda Daucay,
3°29'S, 79°45' W, Cornejo & Bonifaz 2587 (MO); Hacienda
Buenaventura, 12 kin W of Pinas on road to Machala,
3°43'S, 79°46'W, Kessler 2588 (MO); trail between Por-
tovelo and Zaruma, Steyermark 58984 (E). Guayas: Cor¬
dillera Chongon-Colonche, Bosque Protector Loma Alta,

I °48'S. 8( ) 47 V\ . Cornejo & Bonifaz 5242 (MO); Cerro de
Cana, Cordillera de Colonche, Valverde 1014 (MO). Eos
Bios: near Quevedo, Canton Vinces, Mdxia 6602A (E).
Manahi: San Sebastian, Parque Nacional Maclialilla,
l°35'S, 80°42'W, Cornejo & Bonifaz 1285 (MO), 1°36'S,
80°42 W, Gentry el al. 72502 (MO); Cerro Montecristi,
Dodson & Thien 1720 (MO). Gilmartin 826 (MO); canton
Montecristi. Cerro Montecristi. carretera Manta— Jipijapa,
entrada por Montecristi o El Chorrillo, 1°()2'S. 80°41'W,
Nunez et al. 891 (MO); canton Pedernales, Cerro Pata de
Pajaro a 10 km al E de Pedernales, a 5 km del rancho
de la familia Arroyo. (KIO UN, 79°57'W, Vargas 1289
(MO).

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Psychotria subg. Psychotria

125

abaxial leaf surface, showing two domatia situated in the abaxial axils ol two secondary veins with the costa. L
Detail of cymule with (lower at anthesis. A, based on Gentry el al. 72502', B, I), based on Dodson & Thien 1 669,
E, based on Alvarez et al. 36; F, H. G, based on Romoleronx et al. 2575; I, based on Croat 19475: J-L, based on
Romoleroux & Foster 1684. A, E, I, to 5-cm scale; all others to 5-mm scale.

126

Novon

Psychotria jefensis Dwyer ex C. M. Taylor, sp.
nov. TYPE: Costa Rica. Puntarenas: canton de
Osa. hila Costena, Fila Cruces, cabeceras del
Rio Piedras Blancas, Cerro Anguciana, faldas
al O, 8°49'N, 83°1 l'W, 1000-1400 in, 8 Dee.
1988, B. Hummel, R. Aguilar & M. Grayum
19214 (holotype, INB: isotype, MO-5169726).
Figure 3F, G.

Ilaec species a Psychotria orosiana el P. laselvensi lob-
u I is stipularibus hirtellis, foliis domatiis foveolatis caren-
tibus atque limbo calyeino longiore distinguitur.

Small trees to 4 m tall; stems glabrous. Leaves
narrowly elliptic-oblong, 6.5-19.5 X 2-4.5 cm,
papyraceous to chartaceous, at apex acute to some¬
what acuminate, at base acute to cuneate, adaxially
and abaxially glabrous; secondary veins 10 to 21
pairs, looping al least loosely to interconnect, abax¬
ially without domatia, adaxially costa plane to
thickened and remaining venation plane, abaxially
costa prominulous, secondary veins thickened, and
remaining venation plane; petioles 5-8 mm long,
glabrous; stipules caducous, interpetiolar, lanceo¬
late, ca. 6 mm long, glabrous, acute and 1-aristate,
aristas ca. 3 mm long, hirtellous. Inflorescences not
seen. Infructescences terminal, glabrous, peduncle
1. 5-6.5 cm long, branched portion 4—9 X 6— 10 cm.
pyramidal; secondary axes 2 per node, branched;
bracts 0.5—2 mm long, deltoid, glabrous, deciduous;
pedicels 1-3 mm long; fruits pedicellate in umbel-
liform cymules of 3 to 8. subglobose, 4 X 4—4.5
mm, glabrous, with persistent calyx limb 1-1.2 mm
long, shallowly lobed, glabrous; pyrenes 2. plano¬
convex, dorsally with 4 to 5 rounded longitudinal
ribs.

Habitat, distribution, and phenology. In wet
forest at 650—900 m, Osa Peninsula of Costa Rica
and central to eastern Panama; collected in fruit in
April and December.

This new species is similar to Psychotria orosi¬
ana and P. laselvensis C. W. Hamilton, both of
which can be distinguished from this new species
by their glabrous to puberulous stipule lobes,
leaves usually with foveolate domatia in the abaxial
vein axils, and shorter calyx limbs 0.2— 0.8 mm
long. Although flowers have not been seen, the
fruiting collections show adequate morphological
features to distinguish this species. John Dwyer
originally recognized this species as new and pro¬
posed this epithet, which refers to the collection
locality of one of the Panamanian specimens.

Paratypes. PANAMA. Panama: vicinity of Cerro
Jefe, Altos de Pacora, Antonio 3228 (MO). San Bias: Cer-
ro Brewster, headwaters of Rfo Cangandf, 9°I8,N,
79°16'W, de Severs et al. 5489 (MO).

Psychotria lorenciana C. M. Taylor, sp. nov.
TYPE: Guatemala. Izabal: El Estor, bordering
Rio Sareo, El Milagro Ranch, 22 May 1975,
C. L. Lundell & E. Contreras 19332 (holotype,
MO-3763295; isotype, F). Figure 1C— E.

Haec species a Psychotria grandi, P costivenia et I’,
flava fructu subgloboso in diam. 4—5 mm atque inflores-
centiae rotundato-corymbifonnis axibus secundariis imis
insigniter bene evolutis distinguitur.

Shrubs or small trees to 15 m tall; stems gla¬
brous. Leaves elliptic to oblanceolate or obovate,
15—25 X 5.5—1 I cm, papyraceous, glabrous above
and below or sometimes below puberulous on costa
and less often also on secondary veins, at apex ob¬
tuse to deltoid, at base cuneate to acute; secondary
veins 14 to 19 pairs, sometimes looping to inter¬
connect at least in distal portion of blade, abaxially
without domatia, adaxially venation plane or costa
sometimes prominulous, abaxially costa prominent,
secondary veins prominulous and lesser venation
plane; petioles 1.5—5 cm long; stipules interpetiolar,
caducous, 5-12 mm long, ovate to elliptic, glabrous
except margins eiliolate, rounded to usually etnar-
ginate or sometimes 1-aristate with aristas to 2 mm
long and inserted at or below apex. Inflorescences
terminal, glabrous, paniculate, peduncles 4—14 cm
long, branched portion 4—17 X 5— 26 cm. rounded-
corymbiform, secondary axes 2 per node or often 4
and subequal, extensively branched with first in¬
ternode usually well developed; bracts 0.5—5 mm
long, triangular to rounded; pedicels to I mm long;
flowers distylous. mixed sessile and pedicellate in
dichasial cymules of 2 to II; hypanthium turbinate,
ca. 0.8 mm long, glabrous; calyx limb truncate to
denticulate, ca. 0.5 mm long, glabrous; corolla tu-
bular-infundibuliform, while, externally glabrous,
barbate in throat, tube 2-2.5 mm long, lobes 5,
deltoid to narrowly triangular, 1.2-2 mm long,
acute, smooth abaxially; anthers ca. 1 mm long,
included; stigmas ca. 0.5 mm long, exserted. Fruits
subglobose, 4—5 mm diam., color unknown; pyrenes
2, planoconvex, dorsally slightly ridged to smooth.

Habitat, distribution, and phenology. In wet
forests of southern Mexico and Guatemala at 120—
800 m, collected in flower in March and May
through July, in fruit in February, March, and May.

This new species has previously been confused
with the sympatric species Psychotria grandis, P.
costivenia, and P. flava, by both Standley and Wil¬
liams (1972) and Hamilton (1989). Psychotria lor¬
enciana is distinguished from these three species
by its subglobose fruits 4—5 mm in diameter and
its different inflorescence arrangement, with the
secondary axes two at each node or often four, in

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2002

Taylor

Psychotria subg. Psychotria

127

either ease with the secondary axes all ascending
and with their basalmost internodes all generally as
well developed as the comparable internode ol tbe
primary axis. In contrast, these other species have
subglobose to usually ellipsoid fruits 5-13 X 4-9
mm and corymbiform to usually pyramidal inflores¬
cences with the secondary axes two at each node
or usually four or six. with all of the secondary axes
less well developed than the primary axis and when
four or six. then also markedly unequal in pairs.
The flowers seen are all similar to the long-styled
form of distylous Psychotria species in the arrange¬
ment of their anthers and stigmas. It is a pleasure
to dedicate this species to David H. Lorence, a
long-time and distinguished student ol the Mexican
and Mesoamerican Rubiaceae.

Paratypes. GUATEMALA. Alta Verapaz: 9 mi.
uproad to Oxec along gravel road which turns N off High¬
way 7E between Tucuru and El Estor ca. 6 km NE of
Panzos, Croat 41672 (MO); between Einca Cubilgiiitz and
Hacienda Yaxcabanal, Steyermark 44844 (E). Steyermark
44847 (E). Huehuetenango: around Ixcan at Patcushfs,
Sierra de los Cuchumatanes, Steyermark 49197 (E). Iza-
bal: Cadenas/Puerto Mendez, bordering Rfo Dulee, Con¬
treras 9887 (E); Puerto Mendez/Cadenas, on Rfo Dulee
Road, 19 km, Contreras 10014 (MO); Puerto Mendez on
Rfo Dulee Road, Contreras 10309 (F); between Seja/Cien-
aga, 5 km from Seja, 200 m E from Rfo Dulee Road,
Contreras 10204 (F, MO); between Cienaga/Seja, on Pcten/
Guatemala Road, Contreras 10728 (E); El Estor, bordering
Lake Izabal. E. in El Zapotillo, Lundell A Contreras
19299 (F); La Jagua Creek. Murcielago, ca. 10 mi. E of
El Estor, Lake Izabal, Rowland & Snedaker B-5 (F), Sne-
daker D-127 (F). Peten: La Cumbre, on old Pusila Road,
3 km, top of hill. Lundell & Contreras 20011 (F). MEXI¬
CO. Chiapas: mpio. Las Margaritas, western side of La¬
guna Miramar E of San Quintfn, Breedlove 44160 (MO):
mpio. Oeosingo, en el vert ice del Rfo Chixoy, E. Martinez
S. 18941 (E, MO).

Psychotria matagalpensis C. M. Taylor, sp. nov.
TYPE: Nicaragua. Malagalpa: ridge between
Cerro Bravo and Cerro Picacho, mountains N
of Hotel Selva Negra, 13°01'N, 85°55'W,
1490-1550 m, 26 May 1985, G. Davidse, A.
Grijalva & M. Sousa 30484 (holotype, M0-
5059845). Figure 3A, B.

Ilaec species a Psychotria chiriquina el P. sylvivaga
limbo calycino ac corolla brevioribus alque foliis domatiis
foveolatis munitis distinguitur.

Shrubs or small trees to 8 m tall; stems glabrous.
leaves elliptic, 4-10.5 X 1-3 cm. papyraceous, at
apex acute to acuminate, at base acute, abaxially
and adaxially glabrous; secondary veins 4 to 5
pairs, not looping to interconnect, abaxially with 2
to 4 pairs of veins bearing well developed, glabrous
to occasionally pilosulous, foveolate domatia, adax¬
ially venation plane, abaxially costa and secondary

veins thickened and remaining venation plane; pet¬
ioles 1—2.5 cm long, glabrous; stipules caducous,
interpetiolar and shortly also intrapetiolar. elliptic
to obovate, 4—18 mm long, obtuse to acute and
sometimes shortly bidentate, glabrous, margins cil-
iolate and often membranaceous. Inflorescences ter¬
minal, glabrous, sessile or subsessile and appearing
fasciculate, 3-5 X 3-5 cm, broadly pyramidal to
corymbiform-rounded; secondary axes 2 per node,
branched, with basalmost pair generally equally
well developed as primary axis; bracts 0.2— 0.5 mm
long; pedicels 1-3 mm long ; flowers pedicellate in
umbelliform cymules of 3 to 7; hypanthinm 0.8—1
mm long, glabrous, turbinate; calyx limb 0.6—1 mm
long, glabrous, subtruncate to broadly and shallow¬
ly dentate; corolla salverform, white, externally gla¬
brous, internally densely pilosulous at stamen in¬
sertion, tube ca. 3 mm long, lobes 5, ca. 2 mm long,
triangular to narrowly so; anthers ca. 0.8 mm long,
exserted; stigmas ca. 0.8 mm long, exserted. In-
fructescences similar in size and shape to inflores¬
cences; fruits ellipsoid to ellipsoid-obovoid, 6—7 X
4.5—5 mm, red; pyrenes 2, planoconvex, dorsally
with 4 to 5 rounded longitudinal ribs.

Habitat, distribution , and phenology. In wet
montane, cloud, and dwarf forest with boreal ele¬
ments at 920-1700 m, central-western Nicaragua;
collected in flower May through August, in fruit
January through May anti August through Decem¬
ber.

This new species is similar to Psychotria chiri-
quina of Costa Rica and Panama, with which it was
confused by Hami Ron (1989) and Taylor (2001a);
P. chiriquina differs from this new species by its
longer calyx limbs 1-1.6 mm long, its longer corolla
tubes 4—7 mm long, anil its leaves without domatia
in the abaxial vein axils or with only a tuft of pu¬
bescence here. The specific epithet refers to the
region from which this new species is known. I he
(lowers seen are all similar to the short-styled form
of distylous Psychotria species in the arrangement
of their anthers and stigmas. This new species is
also similar to P. sylvivaga, also of Costa Rica and
Panama, which differs from P. matagalpensis by its
longer calyx limbs 1.2-1. 5 mm long, its longer co¬
rolla tubes 4-5 mm long, and its leaves without
domatia in the abaxial vein axils or with only a tuft
of pubescence here.

Paratypes. NICARAGUA. Jinotega: la entrada a
“Aranjuez”, 13°01'-02'N, 85°55'-56'W, Araquistain
3645 (MO); carretera Matagalpa-Jinotega, entre el Km
133-134, a 10 km al SE de la ciudad de Jinotega,
13°01'30"N, 85°55'W, Grijalva & Araquistain 175 (MO);
Kilambe, “Paricuttn”, 4 km al SE del Cerro Kilambe,
13°35'N, 85°40'W, Moreno 7459 (MO); Cerro Kilambe,

128

Novon

Figure 3. A, li. Psycholria matagalpensis C. M. Taylor. —A. Flowering branch. — IT Mature flower bud, partially
opened. C-h. Psychotria orosioides V.. M. Taylor. — C. Flowering branch. — I). Detail of cymule with flower at anthesis.

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Psychotria subg. Psychotria

129

fa I das K del Pico Piedra Pelona, 13°34’N, 85o40< W, Mo¬
reno 7752 (MO). Moreno 7759 (MO); Montana Cuspire,

1 3°1 7'N. 86°09'W, Moreno 8036 (MO); along Hwy. 3 ca.
1.9 km NW of Aranjuez road entrance, 13°02,N, 85°56,W.
Stevens 5625 (BM, MO), Stevens 9 178 (BM, MO); Maci/.os
de Penas Blancas, along trail between finca of Socorro
Meji'a and finca of Luis Manzanares, 13°F6'— 17,N,
85°40'— 11 'W, Stevens 11317 (MO); along Hwy. 3 ca. I km
NW of La Fundadora entrance, unnamed peak ca. 500 m
W of hwy., 13°01'N, 85°56'W, Stevens & Henrich 20445
(MO), Stevens & Miller 22461 (BM. MO). Madriz: Ccrro
Pataste, Neill 129 (MO); Cerro Volcan Somoto (Volcan Te-
pesomoto), 13°26'N, 86°35'W, Moreno 2870 (MO),
I3°25'N. 86°34'W, Moreno 20073 (MO). Matagalpa: Fin¬
ca Los Alpes, 9 km NE de Matagalpa, 12°57 N. 85°21W.
I). Castro 2536B (MO); El Porvenir, Cordillera Central of
Nicaragua, A. Molina R. 20516 (F); Cordillera Central of
Nicaragua, vicinity El Porvenir 4 km E of Santa Maria de
Ostuma. A. Molina R. 22920 (F, MO); El Picacho, E of
Santa Marfa de Ostuma, Cordillera Dariense, .4. Molina
R. & A. R. Molina 30521 (F); Cerro El Picacho. N del
Hotel Selva Negra, I3°00'N, 85°55'W, Moreno 21678
(MO), Nee 27621 (MO), Nee 27633 (MO); Cordillera Dar¬
iense, Hacienda Santa Marfa de Ostuma, 10 km N of Ma-
tagalpa, Neill 2304 (BM. MO). Neill 3167 (BM. MO), Tom¬
lin 24 (MO). Tomlin 60 (BM. MO). Tomlin 126 (MO),
Tomlin 145 (MO), Tomlin 157 (MO); Macizos de Penas
Blancas, SE side, drainage of Oiieluada El Quebradhn,
slopes N and W of Hacienda San Martin, 13°14'— 15'N,
85°38'-39'W, Stevens el at. 21145B (MO); NW slope of
Cerro El Picacho, 13°00'N, 85°55'W, Stevens 22157 (MO);
“Disparate de Potter,” Santa Marfa de Ostuma, Cordillera
Central of Nicaragua, L. O. Williams & T. P. Williams
25051 (E), Williams et al. 27594 (BM. F). Williams et al.
29176 (F); Santa Maria de Ostuma, Cordillera Central de
Nicaragua between Matagalpa and Jinotega. Williams et
al. 23342 (F); Finca Santa Maria de Ostuma, Cordillera
Central of Nicaragua, L. O. Williams & 4. Molina R.
42652 (BM. F).

Psychotria orosioi«les C. M. Taylor, sp. nov. TYPE:
Costa Rica. Alajuela: Reserva Forestal de Ar-
enal, Quebrada San Gerardo, Rio Cano Negro,
Finca de los Mejias, 10°22'N. 131° 18' \\. 12(H) m,
17 Feb. 1990, E. Bello 1894 (holotype, CR; iso-
type, MO-4362126). Figure 3C-E.

Haec species a Psychotria orosiana limbo calycino
longiore distinguitur.

Shrubs to 2 m; stems hirtellous to glabrescent or
glabrous, somewhat flattened. Reaves oblanceolate,
8-17 X 3—6 cm, papyraceous, adaxially glabrous,
abaxially glabrous except hirtellous on costa and
sometimes also secondary veins, at apex acuminate,
at base narrowed then abruptly truncate to rounded;
secondary veins 10 to 13 pairs, united in a contin¬
uous, slightly looping submarginal vein, abaxially

with domatia, adaxially and abaxially costa promi-
nulous, secondary veins thickened, and remaining
venation plane; petioles 0.5—2 mm long, hirtellous
to glabrous; stipules caducous, interpetiolar and
shortly intrapetiolar, hirtellous, ovate to lanceolate,
5—8 mm long, interpetiolarly 2-aristate, aristas 0.5—
2 mm long, hirtellous. Inflorescences terminal, pu-
berulous to pilosulous, peduncles 4-10 cm long,
branched portion 2-5.5 X 3-10 cm, pyramidal to
rounded-corymbiform, secondary axes 2 or 4 per
node, branched; bracts 0.5-3 mm long, lanceolate
to narrowly triangular; pedicels 0.5—1 mm long;
flowers pedicellate in umbelliform to dichasial cy-
mules of 3 to 7; hypanthium ca. 0.5 mm long, pu-
berulous to hirtellous, cupuliform to turbinate; ca¬
lyx limb 1.2-2 mm long, puberulous, lobed lor 1/2-
2/3, lobes sometimes rather unequal in length on a
single flower, narrowly triangular, acute to acumi¬
nate, entire to ciliolate; corolla tubular to lunnel-
form, apparently white, externally glabrous, inter¬
nally densely pilosulous at stamen insertion, tube
2.5—3 mm long, lobes 5, ca. 1.2 mm long, trian¬
gular. thickened to shortly hooked at adaxial apex;
anthers in short-styled form ca. 1.2 mm long, par¬
tially exserted, in long-styled form not seen; stig¬
mas in short-styled form ca. 0.5 mm long and in¬
cluded, in long-styled form ca. 0.8 mm long and
exserted. Fruits ellipsoid to subglobose, 4—5 X 4—
5 mm, red; pyrenes 2, planoconvex, dorsally with
4 to 5 rounded longitudinal ridges.

Habitat . distribution , and phenology. In under-
story of wet lower montane forest at 900—1400 m
in northwestern Costa Rica; collected in flower in
February, June, and July, in fruit in April and May.

This new species is similar in overall aspeet to
Psychotria orosiana , which has been collected in
the same region and can be distinguished by its
shorter calyx limb 0.5-0. 8 mm long. The specific
epithet refers to the similarity between these spe¬
cies.

Paratypes. COSTA RICA. Alajuela: K< iserva Biolb-
gica Monteverde, Rfo Penas Blancas, Altos de Duel i tad a
Portal, 10°17'N, 84°43'W, Bello 902 (MO); Reserva Biol-
dgica Monteverde, Rfo Penas Blancas, parcela de Aleman,
10°18'N, 84°45'W. Bello 962 (MO); Bosque Eterno de los
Ninos, valley of Rfo Agua (Jala, Atlantic slope. 13 km NF,
of Monteverde, 10°23,N, 84°42'W, Haber & Zuchowski
10550 (MO). Guanaeaste: Rfo Chiquito de lilaran, Rfo
Negro valley, Atlantic slope. 10°22,N. 84°52’W, Haber ex
Bello 5879 (MO).

— E. Stipule and adjacent leaf bast'. F. G. Psychotria jefensis C. M. Taylor. — F. Fruiting branch. — G. Stipule. A, based
on Neill 2304 ; B. based on Davidse et al. 30484: C, based on Bello 962: I). E. based on Bello 1894: F. G, based on
Hummel et al. 19214. A, C, F, to 5-cm scale; all others to 5-mm scale.

130

Novon

Psychotria romolerouxiana C. M. Taylor, sp.
nov. I\ 1*K: F.cuador, Napo: Estacidn Cientffica
Yasunf, parcela de 50 ha, Plata #50094,
00°38'S, 76°30'W, 200-300 m, 10 June [with¬
out year; 1995], K. Romoleroux R. Foster
1684 (holotype, QCA; isotype. MO-4933514).
Figure 2G. I, J, K.

Haee species a Psychotria borjensi dornatiis foveolatis
bene evolutis at<|ue stipulis hilohis longioribus distingui-
tur.

Shrubs to 2.5 in; stems puberulous to glabres-
cent. halves elliptic to narrowly elliptic or oblan-
ceolate, 4.5—20 X 0.9—7 cm, at apex slenderly acu¬
minate, at base acute to sometimes abruptly and
shortly obtuse, adaxially glabrous, abaxially gla¬
brous on lamina and densely spreading-puberulous
on costa and secondary veins, margins undulate at
least when dry; secondary veins 6 to 10 pairs, loop¬
ing broadly to interconnect with I to 2 secondary
loops, abaxially 2 to 5 secondary veins near apex
usually with domatia in axils, domatia foveolate, 2-
5 mm long, with swollen portion fused to costa,
adaxially venation plane or costa prominulous,
abaxially costa prominulous, secondary veins thick¬
ened to prominulous, and higher-order venation
plane; petioles 2—9 mm long, sometimes slightly
winged; stipules persistent on 2 to 5 most distal
nodes, interpctiolar and shortly intrapetiolar, ob-
ovate to elliptic, 6.5—14 mm long, glabrous except
sometimes hirtellous to puberulous in a central line
and/or with margins ciliolate to ciliate, at apex bi-
dentate with teeth narrowly triangular, 1—4 mm
long, acute to acuminate. Inflorescences terminal,
puberulous, peduncles 2—4 cm long, branched por¬
tion 1—4 X 1 .2-8 cm, ovoid to broadly pyramidal
or subglobose, secondary axes 4 or 6 per node,
shorter and reflexed in pairs, branched one time or
terminating in a glomerule; bracts reduced or to 0.8
mm long, triangular; flowers sessile in glomerules
of 2 to 3, these glomerules often arranged on short
diehasial axes; hypanthium ea. 0.5 mm long, cylin¬
drical to turbinate, puberulous; calyx limb ca. 0.5
mm long, denticulate, puberulous; corolla funnel-
form, white, externally glabrous, internally densely
hirtellous at stamen attachment, tube ca. 2.5 mm
long, lobes 5, ca. 1 .5 mm long, triangular; anthers
ea. I mm long, included; stigmas 0.8-1 mm long,
exserted. Fruits ellipsoid, 5-6 X 4. 5-5. 5 mm, red;
pyrenes 2, planoconvex, dorsally with 4 to 5 re¬
duced, rounded to acute, longitudinal ridges.

Habitat, distribution, and phenology. In wet
“tierra firme” forest and seasonally inundated "var-
zea forest at 100—300 m, Amazonian Colombia to
Peru; collected in (lower in April. June through Au¬

gust, October, and November, in fruit January
through March and May through December.

Tin s new species keys out to Psychotria borjensis
in several flora treatments (e.g., Steyermark. 1972,
1974), and has been combined with this species in
recent works (Taylor & Pool. 1993; Taylor, 1997,
1999). However, P. borjensis is apparently allopatric
in the northeastern Amazon basin and differs from
this new species by the combination of its leaves
abaxially without domatia or with only a few flat
foveolate domatia up to I mm long and its acute
triangular stipules 5-8 mm long. This new species
is also similar to P. sacciformis' see tin1 discussion
under the description of this latter species below.
The flowers seen are all similar to the long-styled
form of distylous Psychotria species in the arrange¬
ment of the anthers and stigmas. The specific epi¬
thet honors Katya Romoleroux, an Ecuadorian bot¬
anist who is studying the Ecuadorian flora.

Paratypes. COLOMBIA. Amazonas: Parque Nacion-
al Natural Amacayacu, Centro Administrative Amaeayacu
INDERENA, a la orilla del Rfo Amacayacu, 3°47'S,
70°15'W, Pipoly 15309 (MO); Parque Nacional Natural
Amacayacu, Quebrada de Agua Pudre, ca. 1.5 km N L de
desembocadura sobre el Rio Amacayacu, Muestreo Per-
manente E’strategiea del MO, 3°47'S, 70°15'W. Pipoly et
al. I600S (MO), 16519 (MO); Parque Nacional Natural
Amacayacu. Mala-Mata, en trocba hacia donde el “Abuelo
Panduro”, 3°47'S, 70°15'W, Rudas & del Aguila 1.383
(MO). ECUADOR. Napo: Yasunf Forest reserve, 1—3 km
E of Pontiffcia Universidad Catblica del Ecuador Science
Station, by Tiputini River, 00°4I'S, 76°24'W, Acevedo-
Rodriguez & (Fdeno 7344 (MO), sendero alras de la casa,
Jaramillo 8514 (F); Estacibn Cientffica Yasunf, Rfo Ti¬
putini, al NO de la confluencia con el Rfo Tivacuno. (>
km E de la carretera Maxus, Km 44, desvfo hacia el pozo,
00 38 S. 76°30'W. Romoleroux 2230 (F), Romoleroux &
hosier 2134 (F. MO). Pastaza: Lorocachi, ca. 5 km SSW
of the military camp, l°38'S, 75°58'W. Brandbyge A: A.s-
anza 31051 (MO); Curaray, SE of the airstrip, 1°22'S,
76°57'W, Holm-Nielsen et at. 22274 (MO). PERU. Lor¬
eto: Prov. Maynas, Varadero de Ma/.an from Rfo Amazon¬
as to Rfo Napo. Croat 19475 (F. MO), Croat 20762 (MO);
Santa Marfa del Ojeal on Rfo Amazonas, ca. 15 km
downriver from Iquitos, Davidson 5328 (MO); Rfo Gueppi
[sic; Guepf], tributary of Rfo Putumayo, northernmost t ip
of Peru on border with Ecuador, trail from Puerto Peru (8
km from mouth of river) toward Rfo Napo, (Fairy et al.
21946 (t. MO); Quebrada Sucusari, Ulachapa camp of Ex-
plorama, N side of Rfo Napo below Mazan, Centry et al.
27576 (MO); Yanomono, Expforama Tourist Camp, 25 km
NE of Iquitos along Rfo Amazonas between Indiana and
mouth of Rfo Napo, 3°28'-30'S, 72°48'-50'W, (Fairy et
al. 31466 (F, MO). (Fairy et al. 37213 (MO), (Fairy et <d.
37987 (MO). (Fairy et al. 39718 (MO), (Fairy et al.
42253 (MO), Gentry et al. 42438 (MO). Pipoly et al. 12427
(M0), Pipoly et al. 12485 (MO), Vdsquez et al. 4699 (MO),
Vdsquez et at. 5166 (MO), Vdsquez et al. 14102 (MO),
Vdsquez et al. 14142 (MO), Rushmaster Trail, Webster
23334 (MO); Rfo Ampiyaeu. Pc has and vicinity, near
town, 3°10'S, 71°49'W, Plowman et at. 6538 (F); Cahuide
(Rfo Itaya), Vdsquez & Jaramillo 5635 (F. MO); Iquitos,

Volume 12, Number 1
2002

Taylor

Psychotria subg. Psychotria

131

Allpahuayo, Estaeion Experimental del Instituto de Inves-
tigaciones de la Amazonia Peruana (IIAP), Vdsquez & Jar-
amillo 14477 (MO); Nanay, Monte real [sic |. Woytkowski
5154 (MO): Prov. Requena: Sinchicuy, Rio Amazonas,
3°35'S, 73°15'W, Vdsquez et al. 7801 (F).

Psychotria saeciformis C. M. Taylor, sp. nov.
TYPE: Ecuador. Napo: Estaeion Cientffica \a-
sunf. Rfo Tiputini, al NO de la conflucncia con
el Rio Tivacuno, 6 km E de la carretera Max-
us. Km 44, desvio hacia el pozo, 00°38'S,
76°30'W. 200-300 m, 8 Oct. 1996, K. Romo-
leroux, M. Bass, R. Foster & G. Villa 25/5 (ho-
lotvpe, QCA; isotypes, F, MO-5094912). Fig¬
ure 2E, F. G, H.

Haec species a congeneris foliis domatiis foveolatis in
saccum clausum Q.5-2.0 mm longum prolongatis munitis
distinguitur.

Shrubs or small trees to 4 m; stems glabrous or
sometimes hirtellous in a longitudinal interpetiolar
line. Leaves oblanceolate to elliptic-oblanceolate or
elliptic, 6—14 X 1.8-6 cm, papyraceous, adaxially
and abaxiallv glabrous or adaxially hirtellous along
costa, at apex acuminate, at base cuueate to obtuse
or acute; secondary veins 3 to 9 pairs, united in a
looping, weak to well developed submarginal vein,
abaxiallv with domatia in axils, domatia foveolate,
0.5—2 mm long, tubular to sacciform and prolonged,
emergent, adaxially venation plane or costa some¬
times prominulous, abaxiallv costa prominent, sec¬
ondary veins prominulous, and remaining venation
plane; petioles 3—6 mm long, glabrous to sparsely
hirtellous; stipules caducous, interpetiolar and
shortly intrapet tolar, narrowly lanceolate to narrow¬
ly triangular, 3—12 mm long, glabrous or usually
hirtellous at least in a central interpetiolar line,
slenderly acuminate. Inflorescences terminal, gla¬
brous to puberulous, peduncles 3.5— 7.5 cm long,
flexuous, branched portion 3.5—6 X 3. .5— 6 cm,
ovoid to rounded-corymbiform, secondary axes 4
jter node with 2 shorter and reflexed, branched 1—
3 times, with higher-order axes and pedicels
spreading to 90°-100°; bracts reduced or to 1 mm
long, triangular, glabrous; pedicels 1—2 mm long;
flowers in umbelliform cymules of 3 to 7; hypan-
thium ca. 0.5 mm long, turbinate to somewhat cu-
puliform, glabrous; calyx limb 0.5— 0.8 mm long,
glabrous, denticulate; corolla tubular, cream to
white, externally glabrous, barbate in throat, tube
ca. 2.5 mm long, lobes 5, ca. 1.5 mm long, trian¬
gular; anthers in both forms ca. 1 mm long, in
short-styled form exserted, in long-styled form in¬
cluded to partially exserted; stigmas in short-styled
form ca. 0.8 mm long and included, in long-styled
form ca. I mm long and exserted. Fruit ellipsoid to

obovoid, 5-5.5 X 4 mm, red to purple, with stipes
to 4 mm long; pyrenes 2, planoconvex, dorsally with
4 to 5 rounded longitudinal ridges.

Habitat, distribution, and phenology. In wet
forest at 200—750 m in Amazonian Ecuador and
Peru; collected in flower in January, July, October,
September, and November, in fruit March through
May and in September.

This new species is distinguished by its leaves
abaxiallv with foveolate domatia that are prolonged
into tubular, emergent sacs 0.5—2 mm long and
completely closed or sometimes often tardily at the
very top; the specific epithet refers to these struc¬
tures. These structures are highly unusual among
the South American Rubiaceae. Similar domatia
are known to me only in Psychotria cornejoi of
South America (described above), and have been
reported from one African species. Coffea dactyli-
fera Robbrecbt & Stoffelen, where they are also
considered unusual (Stoffelen et al.. 1999). Psycho¬
tria sacciform is will key out to Psychotria horjensis
in several flora treatments (Steyermark. 1972.
1974; Taylor. 1997), and has been confused with
this species in other recent works (Taylor & Pool,
1993; Taylor, 1999). However, P. horjensis is ap¬
parently allopatric in the northeastern Amazon ba¬
sin. and differs from this new species by the com¬
bination of its leaves abaxiallv without domatia or
with only a few flat foveolate domatia up to 1 mm
long and its acute stipules. This new species is also
similar to P. romolerouxiana, described above,
which differs in its domatia that are fused along
their lengths with the costa and its bidentate stip¬
ules.

Paratypes. COLOMBIA. Antioquia: mpio. San Luis,
Quebrada La Cristalina, 6°N, 74°45'W, ./. 6. Ramirez &
Cardenas L 1923 (JAUM, MO): mpio. de San Luis, cor-
regimiento El Prodigio, vereda Las Confusas, hacienda La
Esperanza, margen izquierda del Rfo El ligre, Toro 15
(MEDEL, MO). Caqueta: mpio. San Vicente del Caguan.
trazado de la carretera entre Neiva y San Vicente, vereda
Las Perlas, bajo Rfo Pato, finca Galicia, <*n la vega del
Rfo Pato, Betancur et al. 2266 (HLJA. MO). Chord: E
side of Serranfa del Darien, from A rand u 8°30 N,
77°2()'S, Juncosa 655 (MO). Meta: mpio. La Macarena, I
hora sobre el Rfo Guayabero en la desembocadura del Rfo
Losada, 2°20'N, 74°60'W, Callejas A Mantlanda 7074
(HUA, MO). Putumayo: mpio. Mocoa. El A fan. 11. R.
Ramirez P. 769 (MO, PSO). ECUADOR. Morona-Sauti-
ago: pozo petrolero “Garza” de TENNECO, 35 km
(approx.) al NE de Montalvo, 1°49'S, 76°42’W, /ok &
Espinoza 4767 (MO). Napo: Loreto, al N de Galeras,
Bloque 19, lfnea sfsmica 22. helipuerto 3, Companfa Iri-
ton, 00°47'S, 77°28'W, Ereire & Cerda 220 (MO); via Pay-
amino-Loreto, 4—6 km del rfo, 00°26'S, 77°02'W, Zaruma
733 (MO); canton Aguarico, Parque Nacional Yasnnf, La¬
gunas de Garza Cocha, 01°01'S, 75°47'W, Cerdn A Calla
4962 (MO); canton Archidona, Parque Nacional Sumaco,

132

No von

carretera Hollfn-Loreto, comunidad Avila Vieja, 00°38'S,
77°26'W, Alvarado 485 (MO); Avila Viejo, 00°38'S,
77°25'S, Kohn & Alvarado 1581 (MO); canton La Joya de
los Sachas, comunidad de Pompeya, lado S del Rio Napo,
campamento de Maxus, Rfo Jivino, carretera de Maxus,
Km 1—5, 00°25'S, 76°37'W, Grijalva el al. 217 (MO); can-
tdn Orellana, Parque Nacional Yasunf, carretera y oleo-
ducto de Maxus, Km 40, parcela permanente #10.
(K)°39'S, 76°26'S, Aulestia 2715 (MO, QCNE); Parque Na-
cional Yasunf, carretera y oleoducto de Maxus en con-
struccidn, Km 53—54, Dik 416 (MO); Parque Nacional Ya¬
sunf, Tiputini Km 41, parcela 1, Maria el al. 3074 (MO);
Kstaeion Cientffica Yasunf, Rfo Tiputini, al NO de la con-
fluencia con el Rfo Tivacuno, E de la carretera Maxus,
Km 44, desvfo hacia el pozo Ivacuno, Sendero Dicaron,
entre la carretera y la parcela de 50 ha, 00°38'S, 76°30'W,
Romoleroux & Foster 1632 (K MO, OLA); Kstaeion Cien¬
tffica Yasunf, parcela de 50 ha, Sendero Saino, 00°38'S,
76°30'W, Romoleroux & Foster 1909 (F, MO. QCA); Es-
tacidn Cientffica Yasunf, Rfo Tiputini, al NO de la con-
fluencia con el Rfo rivacuno, 6 km E de la carretera Max¬
us, Km 44, desvfo hacia el pozo Tivacuno, parcela de 50
ha, 00°38'S, 76°30'W. Romoleroux et al. 2191 (MO,
QCA), 2231 (F, QCA), 2271 (F, MO, QCA); Kstaeion Cien-
tffica Yasunf, Rfo liputini, al NO de la confluencia con el
Rfo Tivacuno, 6 km E de la carretera Maxus, Km 44.
desvio hacia el pozo, 00°38'S, 76°30,W, Romoleroux et al.
2645 (F, MO, QCA); canton Tena, Reserva Bioldgica Jatun
Sacha, Rfo Napo, 8 km al E tie Misahuallf, 1 °04'S,
77°36'W, Alvarez 25 (MO), Alvarez et al. 36 (MO). Cerdn
946 (MO), 1222 (MO), Cerdn & Iguago 5548 (MO), Pa¬
lacios 4216 (MO); Parque Nacional Yasunf, pozo petrolero
Daimi I, CONOCO, 1°02'S, 76°10'W, Hurtado et al. 42
(MO). Pastaza: Lorocachi, zona O del campamento rnili-
tar, a 3 km del Rfo Curarav. 1°38’S, 75°58fW, Jaramillo
et al. 30796 (MO).

Acknowledgments. I thank the curators of BM,
CR, DUKE, F, HUA, INB, JAUM, MEDEL, NY.
I*M A, and PSO for access to specimens and infor¬
mation, and in particular 1M. Correa; B. E. Gereau
for preparation of (he Latin diagnoses; I). Eorence,
W. I). Stevens, C. Ulloa U., R. Foster, and T. Wach-
ter for helpful discussions; S. Knapp, G. Davidse,
0. M. Montiel, and If. Magill for significant en¬
couragement of this work; and the Andew Mellon
Foundation through a grant to the Field Museum of
Natural History for support for travel to consult that
collection.

Literature Cited

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ganization of the genus Psychotria and tribe Psycho-
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. 1997. Rubiaceae. In: R. Vdsquez M., Kldrula de
las Reservas Bioldgicas de Iquitos, Peru. Monogr. Syst.
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- . 2001b. Overview of the Neotropical genus No-

topleura (Rubiaceae: Psychotrieae), with the description
of some new species. Ann. Missouri Rot. Card. 88: 478-
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- & A. Pool. 1993. Rubiaceae. In: L. Brako & J.

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Gard. 45: 1010-1053.

Emended Circumscription of Begonia silletensis (Begoniaceae) and
Description of a New Subspecies from Yunnan, China

Mark C. Tebbitt

Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York,

1 1225-1099, U.S.A.

Guan Kaiyun

Kunming Institute of Botany, The Academy of Sciences of China, Kunming, Yunnan, China

Abstract. The circumscription of Casparya sil¬
letensis A. DC. ( = Begonia silhetensis (A. DC.) C.
B. Clarke) is emended so as to accurately describe
the type material of this species and exclude ma¬
terial of B. aborensis S. T. Dunn. Begonia silletensis
subsp. mengyangensis, a taxon endemic to t lie
Xishuangbanna region of Yunnan Province, China,
is newly described and illustrated. Begonia sille¬
tensis subsp. mengyangensis differs from subspe¬
cies silletensis by its larger leaves (14—27 X 20-27
cm vs. 10-17 X 10—15 cm) and unequally ridged
fruit. Begonia silletensis subsp. silletensis is report¬
ed new to Burma and Thailand. A key is provided
to distinguish the two subspecies.

Key words: Begonia, Begoniaceae, China.

Approximately 70 species of Begonia L. are
found in Yunnan Province, representing about 00%
of the total number found in China and 0 of the 18
Asiatic sections (Doorenbos et ah, 1998). Begonia
silletensis, a species previously considered restrict¬
ed to northeastern India, was discovered in the
Xishuangbanna region of Yunnan Province in 1977
during a Kunming Botanic Garden expedition and
introduced into cultivation. Examination of the Be¬
goniaceae collections of the Kunming Botanic Gar¬
den Herbarium has revealed additional material
collected from the same region by an earlier Sino-
Soviet expedition in 1957. As these disjunct Chi¬
nese populations of B. silletensis differ from the
northeast Indian populations by having larger
leaves and unequally ridged fruit, the Chinese pop¬
ulations are recognized here as a new subspecies.
Before we can compare the new subspecies with
the typical B. silletensis, the description of B. sil¬
letensis requires emending.

I)e Candolle (1864) originally described Begonia
silletensis as a species of the genus Casparya. The
original circumscription of Casparya silletensis A.
DC. does not match its type ( Wallich 9107). De
Candolle (1864) described C. silletensis as having

velutinous petioles, peduncles, outer surfaces of the
male perianth segments, and ovaries and hispidu-
lous leaf blades. However, the type collection that
he cited along with the description is glabrous
throughout. Apart from this discrepancy, De Can¬
dolle’s description of C. silletensis matches the type
collection. We therefore suggest that De Candolle
based his species on the Wallich 9107 collection
and additional material of a different taxon. As no
additional material is cited in his description of 6.
silletensis, the identity of this additional material is
not certain. However, it is likely to have been B.
aborensis S. T. Dunn, as this species is covered with
velvety hairs and is closely related to B. silletensis.
Begonia aborensis was not recognized as a distinct
species until 1920. In 1879. Clarke synonymized
the genus Casparya with the genus Begonia and
arbitrarily changed the spelling of the epithet ''sil¬
letensis" to "silhetensis.” Clarke (1879) certainly
did include material of both B. silletensis and B.
aborensis in his description of B. silhetensis. lbs
circumscription of Begonia silhetensis ( = B. sille¬
tensis) is based on Wallich 9107 and two other col¬
lections, Griffith 2569 and Wallich 3677B. Our re¬
search shows that Griffith 2569 and Wallich 3677B
are nonspecific with Burkill’s collections from the
Abor Hills of northeastern India upon which Dunn
(1920) based his Begonia aborensis. Clarke, in fact,
suspected that his description was based on mate¬
rial of more than one species as he stated: “Grif¬
fith’s No. 2569 (with which Wallich’s No. 3677B
agrees), differs considerably in hairiness and in size
of the flower from Wallich’s No. 9107 on which
A. DC. founded his Casparya V silhetensis : two spe¬
cies are possibly here mixed, but the material is
not sufficient to justify a new species” (Clarke,
1879: 636). The circumscription of Begonia sille¬
tensis is emended here so as to accurately describe
the type ( Wallich 9107) and exclude material that
Dunn much later described as B. aborensis. Follow-

Novon 12: 133-136. 2002.

134

Novon

ing Clarke (1879), the species is classified in the
genus Begonia.

Begonia silletensis (A. DC.) C. B. Clarke emend.

M. C. Tebhitt. Casparya silletensis A. DC.,

Prod. 15(1): 277. 1864. Begonia silhetensis (A.

DC.) C. B. Clarke, in J. I). Hooker FI. Brit.

Ind. 2: 636. 1879. TYPE: Sillet Mts., Wallich

9107 (holotype, G-DC; isotypes, BM, K-
WALL).

a. Begonia silletensis subsp. silletensis

Dioecious perennial herbs. Stems prostrate, ca. 1
cm diam., internodes 1-2 cm, glabrous. Stipules
persistent, lanceolate, 1.4-2. 1 X ca. 0.4 cm, apex
acute, margin entire, both surfaces glabrous. Leaves
lew, drooping: petioles 22—43 cm, glabrous; blade
green, glabrous on both surfaces, ovate, 10—17 X
10—15 cm, apex shortly acuminate, base asymmet¬
ric, sinus 3—4.5 cm deep, margin undulate, with
short serrate teeth, primary veins 7-9, palmate. In¬
florescence axillary, an erect dichasium in which the
primary peduncle branches vary from 0.1 to 5 cm
so that the inflorescence is either umbellate or cy-
mose in appearance, flowers fragrant, 1-10; pedun¬
cles 8—1 1 cm in both male and female inflores¬
cences; bracts deciduous, ovate to elliptic, ca. 1.1
X 0.2 cm, margin entire, both surfaces glabrous.
Pedicels: those of male flowers 1-2 cm, those of
female flowers 4-5 cm. Male flowers: bracteoles ab¬
sent; perianth segments white, greenish white or
| >in k , 4, outer 2 ovate, oblong or obovate, concave,
thick, 4.5-15 X ca. 10 mm, apex blunt, inner 2
ovate-obovate, 3.4—17.5 X 7.5—8 mm, apex round¬
ed; stamens 100+, arranged in a symmetrical dome¬
shaped mass, filaments 1.75—2.5 mm, free to base,
attached to a raised receptacle, anthers linear,
1.75—2.75 mm, dehiscing via vertical slits along
side of anther, connective projecting 0.5 mm, apex
rounded. Female flowers: bracteoles absent , perianth
segments white or pink, 4, oblong to obovate, apex
rounded, outer 2 slightly longer; ovary ovoid to el¬
lipsoidal, ca. I X 1 cm, circular in cross section,
4-locular, placentation axillary, placentae bifid,
rhomboidal in cross section, ca. 4X4 mm, bearing
ovules on both surfaces ol branches; styles decid¬
uous, 4, very broad, base 0.5 cm across, 0.8— 1.2
cm long, fused just below half way, and then bifid,
band ol stigmatic papillae twice spirally twisted.
Infructescences I -2-1 in i ted; fruiting pedicels ca. 3.5
cm; fruit indehiscent, usually cork-like in texture
or rarely leathery (in sicco), ovoid to ellipsoidal, 2.3
X 1.5 cm. walls ca. 2.5 mm thick, lacking ridges
or wings.

Northeastern India, Burma, and northern Thai¬
land. We report it as new to Burma and Thailand.
Wallich proposed the invalidly published name
"Begonia gigantea” for this species based on the
collection “ Wallich 3677B ” (Wallich, 1831).

Representative specimens. INDIA. Assam: Chanduar
forest, Mann 7/87 (K); Luckimpore, Makum 300 ft.,
12.iv.1885, Clarke 37805 [specimen annotated by C. B.
Clarke as "H. sphaerocarpa” nomen nudum] (K); Nam-
chung, 155 ft., Luchimpore, I8.iv.1885. Clarke 37937A
(specimen annotated by C. B. Clarke as “B. sphaerocarpa”
nomen nudum] (K); Cachar, moist shade of Shapose Bom-
ara, Nov. 1873, Keenan s.n. (K); Katakhal Forest, Mann
s.n. (k). BURMA. Nammeen to Namma, Myitkina District,
1000 ft., 7. i i i . 1910, Lace 5170 (E). THAILAND. Northern
Chiengmai, Trang, ca. 800 m, 11. vi. 1960, herb scattered
in evergreen forest, by stream, fruits purplish red, angular,
T. Smitinand & II. St. John 6 832 (k).

h. Begonia silletensis subsp. meiigyangensis M.

C. Tebbitt & K.-Y. Guan, subsp. nov. TYPE:
China. Yunnan: Xishuangbanna, on way from
Puurem to Mengyang, bottom of valley in wet
area in slope facing N. dense forest, 21 Apr.
1957, Sino-Soviet Union expedition 9633 (ho¬
lotype, kUN; isotype, KUN). Figure 1.

A B. silletensis subsp. silletensis fob is majoribus (ad 27
X 27 cm); fructus globosus interdum carinatus.

Dioecious, perennial herbs, 20—50 cm tall. Stems
prostrate, 1.5— 4.0 cm diam., cortex red as seen in
cross section, internodes 2—3 cm, with sparse short
hairs. Stipules persistent, reddish, not keeled,
asymmetrically triangular-lanceolate, 2—2.5 cm
long, dorsal surfaces with short hairs. Petioles green
with white lenticels, red at base, 30—55 cm long,
with short white hairs throughout; blades shiny
green, leathery, glabrous above, paler green, hairy
beneath, broadly ovate, 14-27 X 20—27 cm, apex
acute, base cordate, slightly asymmetric, sinus 4.5—

1 I cm deep, leaf bases overlapping, margin undu¬
late, with short serrate teeth, primary veins 8, pal¬
mate. Inflorescence axillary, an erect dichasium in
which the primary peduncle branches are reduced
to 1—3 mm long so that the inflorescence appears
umbellate, flowers fragrant, ca. 15; peduncles: those
of male inflorescence 10—15 cm long, with short
white hairs, those of female inflorescence ca. 17 cm
long, with short white hairs; bracts deciduous, nar¬
rowly to broadly ovate, 0.8—2 X 0.2-1. 2 cm. both
surfaces glabrous, margins shortly eiliate. Pedicels
of male flowers 12—35 mm long. Male flowers: brac¬
teoles absent; perianth segments white, sometimes
flushed pink, 4, outer 2 broadly ovate to broadly
elliptic, ca. 15 X 10—15 mm, apex obtuse, outer
surfaces pubescent, inner two broadly elliptic to
broadly obovate, ca. 10 X 12 mm, apex obtuse,

Volume 12, Number 1
2002

Tebbitt & Guan

Begonia silletensis from Yunnan

135

136

Novon

outer surfaces glabrous. Stamens ca. 60, arranged
in a symmetrical dome-shaped mass, filaments free
to base, ca. 2 mm long, attached to a raised recep¬
tacle, ca. 1 mm high, anthers elliptic, ca. 2.2 mm
long, dehiscing via vertical slits along side of an¬
ther, connective projecting 0.5 mm, apex rounded.
Pedicels of female flowers 1—2 cm long. Female
flowers-, bracteoles absent, perianth segments white,
sometimes flushed pink, 4—6, outer 2—3 ovate, ca.
15 mm long, apex obtuse, inner 2-3 elliptic to ob-
ovate, ca. 12 mm long, apex obtuse; ovary ellipsoi¬
dal, ca. I X 1 cm, 4-locular, placentation axillary,
placentae bifid, bearing ovules on both surfaces of
branches; styles long persisting but eventually de¬
ciduous, 4, fused at base for 1.5—2 nun, bifid, band
of stigmatic papillae once spirally twisted. Infruc-
tescence 4— 6-fruited. Fruiting pedicels green, erect,
to 3 cm long; fruit indehiscent. fleshy, ellipsoidal,
(0.5— )1— 2 cm diam., with short hairs, lacking ridges
or wings or with up to 4 blunt unequal, triangular
ridges (on same plant), ridges 2-3 X 3-5 mm high.
Seed ca. 300 pan long.

Begonia silletensis subsp. mengyangensis grows
among tall herbaceous vegetation in damp, shady
areas in primary tropical forest at altitudes between
570 and 1200 m; it has been collected with flowers
and fruit in May and April. This taxon has proved
amenable to cultivation, both at Kunming Botanic
Garden, China, and Glasgow Botanic Gardens,
Scotland.

Paratypes. CfflNA. Yunnan: Xishuangbanna, Cheli
liuslia lie, along riverside wet areas in shady forest. 600-
800 in, 30 Apr. 1957, Sino-Soviet Union expedition 9834
(KUN); Xishuangbanna, Mengla, Mengyang, in valley un¬
der dense forest in wet areas, 1200 m, 5 Apr. 1957, Sino-

Soviet Union expedition 5869 (KUN-3 sheets); Mengla,
Mengxin he, in valley, alt. 570 m, 19 Mar. 1977, Xha-
ngjianhou 13666 (KUN). CULTIVATED. Glasgow Botanic
Garden National Begonia Collection, 18 Jan. 2000. L. L.
Forrest 99 (E).

Key to the Subspecies of Begonia silletensis

la. Leaves 10—17 X 10—15 cm; fruit lacking ridges

. subsp. silletensis

lb. Leaves 14—27 X 20—27 cm; fruit unequally

ridged . subsp. mengyangensis

Acknowledgments. We thank Adele Rossetti
Morosini who provided the illustration and Kerry
Barringer, Laura Forrest, Katherine Gould, James
Yeadon, and two anonymous reviewers who provid¬
ed comments on the manuscript. We also thank Da¬
vid Chamberlain for helping to arrange llie field¬
work and Ewan Donaldson and John Stevenson for
providing cultivated material. This research was
supported by the Biotechnology & Biological Sci¬
ences Research Council (grant no. 9370131 1) and
represents a portion of a Ph.D. dissertation sub¬
mitted by the first author to the University of Glas¬
gow. Scotland.

Literature Cited

Candolle, A. de. 1864. Begoniaceae. Prodr. 15: 266—408.
Clarke, C. B. 1879. Begonia. P. 636 in C. B. Clarke (ed¬
itor), Flora ol British India, Vol. 2. I,. Reeve, London.
Doorenboos. .1.. M. S. M. Sosef & J. .1. F. E. de Wilde.
1998. The sections of Begonia including descriptions,
keys and species lists (Studies in Begoniaceae VI).
Agric. Univ. Wageningen Pap. 98(2): 1-266.

Dunn, S. T. 1920. Plantarum Novarum in Herbario Horti
Regii Conservatarum. Bull. Misc. Inform. 109—1 10.
Wallich, N. 1831. A Numerical List of Dried Specimens
of Plants in the East India Company’s Museum, Col¬
lected under the Superintendence of Dr. Wallich of the
Company’s Botanic Garden at Calcutta. P. 129. London.

Magnolia pugana (Magnoliaceae): Una Nueva Combinacion en el

Complejo M. pacific a

./. Antonio Vazquez G., Servando Carvajal y Leticia Hernandez L.

Km 15.5 earn Guadalajara a Nogales, Las Agujas, Nextipac, Zapopan 451 10, Jalisco,
Mexico. Herbario IBIJG. Institute do Botanica, Departamento de Botanica y Zoologfa,

Universidad de Guadalajara

Rush MEN. Investigacidn morfologica y ecologies
reciente, tanto en la costa norte de Jalisco como en
la Zona Centro del mismo Estado, apoyan el recon-
ocimiento de Magnolia pugana (H. H. litis & A.
Vazquez) A. Vazquez & Carvajal a nivel de especie.

Abstract. Recent morphological and ecological
investigation in both the northern Jaliscan coast
and central Jalisco support the recognition of Mag¬
nolia pugana (H. H. litis & A. Vazquez) A. Vaz¬
quez & Carvajal at the species level.

Key words: Magnolia, Magnoliaceae, Mexico.

I a seccion neotropical Theorhodon (Spach. 1839;
MO-W'-Tropieos, (http://mobot.mohot.org/W3T/
Search/vast. html), 09 julio 2(X)1) (caracterizada por
tener hojas siempre verdes, estfpulas libres del pe-
ciolo, gineceo sesil, tepalos subsimilares y estambres
sin conectivo extendido) del genero Magnolia presen-
ta un patron marcado de especiaeion alopatrica (How¬
ard, 1948; Vazquez G., 1990, 1994). con considerable
variaeion entre poblaciones, tal vez relacionado con
la distancia geografica y/o ambiental entre las mismas.
De tal suerte que el intrineado relieve y diversidad
climatica parecen favorecer dicho fenomeno de es-
peciacion. En la actualidad esta seccion incluye 11
especies publicadas, la mayoria de Mexico (6) y varias
mas estan sin describirse, con escaso material fertil
de llores y frutos. Vazquez (1994) y Eelger (1971)
ilustran respect ivamente importantes aspectos de var-
iacion tanto morfologica en hojas v frutos como ecol-
ogiea de lo que podnamos denominar el complejo
Magnolia pacified A. Vazquez. Este complejo consiste
de tres subespecies alopatricas (Fig. 1): M. pacifica
subsp. pacifica, M. pacifica subsp. tarahumara y M.
pacifica subsp. pugana. las cuales por falta de sufi-
cientes espec fineries fertiles procedentes de la Sierra
Madre Occidental fueron tratadas en forma conser-
vadora como subespecie por Vazquez G. (1994). La
disponibilidad de mayor cantidad de frutos maduros
procedentes de poblaciones representativas de la cos¬
ta norte de Jalisco y del centro de Jalisco, aunado a
exploraciones adicionales en habitats poteneiales,
permitio reconocer y confirmar una discontinuidad

morfologica, geografica y ecologica entre Magnolia
pacifica subsp. pacifica y Magnolia pacifica subsp.
pugana (Eig. I), por lo que esta ultima se propone
como una especie independiente. fan solo los car-
acteres del numero de carpelos por fruto y mirnero de
estambres por flor permiten separarla con claridad.

La ( lave publicada para taxa del complejo M.
pacifica (Vazquez G., 1994: 13) <]uedarfa, traducida
y corregida, de la siguiente forma:

Pedunculos generalmente glabros; polifolfculos maduros
oblongoides, con 25—35 folfculos; hojas de angostamente
alargadas o lanceoladas a angostamente elfpticas, 2.5—8
cm de ancho (Centro y N de Jalisco a S de Zacatecas,

“region de barrancas”) . M. pugana

Pedunculos pubescentes, al menos en los nodos; polifolf¬
culos maduros de angostamente oblongoides a ovoides o
subglobosos, con 18—26 folfculos; hojas anchamente elfp¬
ticas a lanceolado elfpticas, 3—10 cm de ancho.

- Polifolfculos maduros angostamente oblongoides, con
18-25 folfculos, estos, cuando maduros y abiertos, con
loculos angostos, mas largos que anchos; hojas usual-
mente anchamente elfptico obovadas, 4—10 cm de ancho
(Nayarit, SW Jalisco) ... M. pacifica subsp. pacifica

— Polifolfculos maduros ovoides a subglobosos, con 21—26
folfculos, estos, cuando maduros y abiertos, con loculos
ovados a redondeados (excepto los de la base del jxilifol-
fculo); hojas elfpticas a lanceolado elfpticas, 3—7 cm de
ancho (SE de Sonora, SW de Chihuahua, NW de Sinaloa,
NW de Durango) ... M. pacifica subsp. tarahumara

Magnolia pugana (H. II. litis & A. Vazquez) A.
Vazquez & Carvajal, comb. nov. Bason i mo:
Magnolia pacifica A. Vazquez subsp. pugana
H. II. litis & A. Vazquez, Brittonia 46: 14.
1994. TIPO: Mexico. Jalisco: margen de bosque
en pequeno tributario del arroyo San Girenzo,
35 km al NW de Guadalajara, 8 km NW de
Tesistan, Zapopan, 20°50'N, 103°34'W, 1450
m, 18 Mar. 1987. litis, Chdzaro />'. y R. Ixfpez
V. 29722 (holotipo, WIS; isotipos, I BUG,
MEXU, MICH, MO, US). Figura 2.

Arboles de 10-15(20) m de alto, 50-80(150) cm
dap, con las primeras ramas de 2—15 m de el piso;
ramillas eseneialmente glabras, corteza rugosa, de
1—2 cm de grosor, gris oscura a castano. Hojas eor-

Novon 12; 137-141. 2002.

138

Novon

iaceas, angostamente oblongas o lanceoladas a an-
gostamente elfptieas, 12-22 X 2.5-8 cm; agudas a
obtusas en el dpice, cuneadas eu la base; estfpulas
lineares de 4.5—7 X 3—6(8) cm de audio, palido
serfceas en el exterior, con nervaduras reticuladas;
pecfolos canal iculados, de 1— 1.5(3) cm de largo,
glabros o rara vez pubescentes. Flores con botones
ovoides a oblongoides, protegidos por 2 a 3 brac-
teas, la superior espatacea de 4—6.5 X 1.5—2 cm;
la bractea inferior (perula) dehiscente, 1.5—2 X 6—
9 mm, densamente palido-velutina. Flores abiertas

9—15 cm en diametro, blancas; pediinculos glabros,
3.5 X 3.5—6 mm de diametro, consiste de dos en-
trenudos, 0.5—1 .5 mm de largo, el menor de 33 mm
de largo; sepalos, 3, obovado-oblongos, concavos,
3—7 X 2—3.5 cm, verdosos en el exterior, blancos
o color crema en el interior, glabros en ambas car-
as; petalos 6 a 7, anchamente obovados, abrupta-
mente atenuados hacia su delgado pecfolo, los ex-
teriores 5—7 X 2. 5-5. 5 cm, los interiores 3—5.5 X
1—4 cm de audio, blancos, glabros en ambas caras;
estambres 92 a 104, 8-12 X 1-2 mm de ancho,

Volume 12, Number 1
2002

Vazquez et al.
Magnolia pugana

139

Cuadro 1. Diferencias entre

Magnolia pugana y M. pacifica

sensu stricto.

Caracterfsticas

M. pugana

M. pacifica sensu stricto

Pedunculos

esencialmente glabros

pubescentes al menos en los nudos

Pol i fo 1 f culos mad u ros

oblongoides, con 25—35 folf-
culos

estrechamente oblongoides, con 18—25 folfcu¬
los

# de estanibres

92-1 (M)

100-1 10

Ldculos de folfculos

ovales a easi redondos, poco
mas largos que anchos

angostos, mucho mas largos que anchos

Forma de hojas

angostamente oblongas a
lanceoladas, a estrecha-
mente elfpticas

angostas a ampliamente elfpticas o obovadas

Ancho de la hoja

2.5—8 cm

4-10 cm

Habitat

bosque de galena en barran¬
cas subtropicales

bosque mesofilo de montana

Es])eeies arboreas eonumes

Astronium graveolens Jae-
(]uin, Ficus, Nectandra y
Salix

Podocarpus reichei Buchholz & Gray, Matu-
daea trinervia Lundell, Pinas jaliscana

Perez de la Rosa, Dion edule var. sotwrense
(De Luca, Sabato & Vdzquez Torres) Mc-
Vaugh & J. A. Perez de La Rosa, Styrax
argenteus C. Presl, Persea hintonii G. K.

Allen e Inga laurina (Swartz) Willdenow

Geograffa

Jalisco: centra y norte; Zaca¬
tecas: sur

Jalisco: suroeste; Navarit: centra

las anteras acuminadas a cuspidadas en el apice,
color crema; gineceo oblongoide-elipsoide, verde,
glabro; estilos encorvados, 2—3 mm de largo, ver-
dosos, con papilas estigmaticas inconspicuas. f ru-
tos de tipo polifolfculos oblongoides, glabros. eon
25—35 folfculos, estos, cuando maduros y abiertos,
con loculos ovales casi redondos, un poco mas lar¬
gos que anchos, algunos tenidos de color rojizo.
Semillas prismatico triangulares, subcilfndricas o
rotundo eomprimidas de 9—12 X 7—8 mm, con sar-
cotesta de color rojo escarlata.

Variabilidad. Es muy posible que la obtencion
de suficientes especimenes con llores y frutos ma¬
duros de poblaciones del complejo M. pacifica en
las estribaciones de toda la Sierra Madre Occiden¬
tal apoye la mocion de que M. pacifica subsp. tar-
ahumara deba tambien ser reconocida a nivel de
especie. Las poblaciones de las montanas del sur
de Durango y Sinaloa son excluidas por aliora ya
que podrfan, incluso representar una especie dis-
tinta, sin embargo todavfa no son bien conocidas.

Distribucidn, habitat y fenologui. Mexico, cer-
canfas de Guadalajara en Zapopan. Jalisco, y hacia
el Norte hasta Moyahua, Zacatecas, 1300—1800 m
(Fig. 1). En canadas profundas (barrancas) con re-
manentes de bosque en arroyos permanentes, con
arboles de Astronium, Ficus, Clethra, Nectandra y
Salix ; arbustos de Malvaviscus y Randia ; y en oca-
siones especies raras como Psilotum y Equisetum.
Florece de marzo a principios de julio, con algunas
Mores durante la mayor parte del aim; los frutos por
lo comun permanecen unidos y su dehiscencia se

presenta en marzo o en abril del siguiente ano. Al¬
gunos arboles tiran sus frutos en mayo, despues de
su dehiscencia, mientras que otros los mantienen
durante la florae ion.

Eponimo. La especie en honor a Luz Maria Vil¬
larreal de Puga, eximia profesora y exploradora bo-
tanica del Instituto de Botanica de la Universidad
de Guadalajara.

Nombres y usos locales. “Almacasusco” [Rze-
dowski 9135), “magnolia” y “manolia” ( Vazquez G.
et al. 5019a-c). A los petalos de Magnolia pugana
se les atribuyen propiedades medicinales, y se pre-
paran como tisana para tratar padecimientos car-
diacos. Las Mores se colectan y lit i I ix.au en peregri-
naciones de cuaresma para adornar liogares y
recintos religiosos y aromatizar el ambiente.

Estado de conservacidn. Magnolia pugana es
endemica a la region de barrancas (Rzedowski &
Me Vaugh, 1966) del centra de Jalisco y zonas ad-
yacentes de Zacatecas. Esta especie, por el redu-
cido tamano de sus poblaciones y por su distribu¬
cidn restringida se encuentra en peligro de
extincion. Aunque en el Bosque La Primavera. de-
cretado como Area de Proteccidn Forestal y Refu¬
gio de Fauna Silvestre, se ha reportado una pob-
lacion de M. pugana, esta no contiene toda la
variabilidad genetica de la especie. Por eso, se re-
comienda proteger de manera prioritaria otras pob¬
laciones in situ antes de que sean exterminadas por
los cambios del uso del suelo, por incendios lores-
tales o bien por aprovechamientos forestales de au-
toconsumo. Tambien se recomienda su reproduc-

140

Novon

' 4

jfir

■K

Hi %

m

v* \ ./dBm

IMi

iiV

Figura 2. Magnolia pugana (H. H. litis & A. Vazquez) A. Vazquez & Carvajal. — A. Flor. — B. Fruto inmaduro. —
C. Fruto en dehiscencia. — D. Fruto despues de la dehiscencia.

Volume 12, Number 1
2002

Vazquez et al.
Magnolia pugana

141

cion y propagation para su uso en zonas urbanas
(con base en la experiencia <le plantaciones exito-
sas realizatlas en el Centro Universitario de Cien-
cias Biologicas y Agropecuarias de la Universidad
de Guadalajara).

Ejemplares representatives examinados. MEXICO. Jal¬
isco: (1) Zapopan, Arroyo San Lorenzo, 8 km al NW de
Tesistan, 1530 m, 30 June 1984, Samz-Ch. & Nieves-H.
134 (IBUG, W IS, XAL); 8 km al NW de Tesistdn,
20°50'N, 1 03°34'W, 1450 m, 18 Mar. 1987. litis et al.
29724 (WIS); 1550 m, 12 Oct. 1984, Samz-Ch. 203
1 1 BUG), 14 July 1989, Vdzquez-G. et al. 5019a-c (WIS),
17 Jan. 1999 , Sanchez- M. s.n. (IBUG); 1460 m. 6 Jan.
1974, Villarreal de Puga 0126 (IBUG, MEXU, MICH);
1450 m, W de Santa Lucia, 18 Mar. 1987, Chdzaro-B. et
al. 4545 (IBUG); 12 km al NL de Tesistan, camino a San
Cristobal de la Barranca, 1600 m, 12 Aug. 1985, Soltero-
Q. & Contreras 240 (IBUG); La Mesita de Santa l.ucfa,
1600 m, 15 July 1989, Ram(rez-D. & Camacho 1333
(IBUG); 10 km antes de San Cristobal de la Barranca,
1560 m, 14 Feb. 1982. Castro-C. 91 (IBUG); Arroyo Agua
Caliente, Banos El Encanto, Arroyo Los Pajaritos, 1350
m, 29 Dec. 1974. Villarreal de Puga 7102 (IBUG); La
Sabana, camino a El Taray, N de Tesistan, 1800 m, 28
June 1986, Rami'rez-D. 332 (IBUG); Reserva Forestal y
de la Fauna Bosque La Primavera, 4 June 1974, Becerra-
C. 1540 (IBUG). (2) Amatitan. 17 km de Santa l.ucla,
brecha de Tesistan a presa Santa Rosa, 1355 m, 1 Mar.
1995, Silva-V. et al. 1-3 (IBUG); (3) San Cristobal de la
Barranca, Arroyo Milpillas, 1400 m, 13 June 1987, ()r-
nelas-U. & Flores-M. 355 (IBUG); 9 km al NE de San
Cristobal de la Barranca, earretera a Colotlan, 1300 m, 15

Oct. 1986, Santana- M. et al. 2561 (IBUG); earretera Tes¬
istan— San Cristobal de la Barranca, 16 July 1987, 1450
tn. 12 July 1987, Chdzaro-B. et al. 4372 (IBUG); 10 km
despues de el Balneario Los Camachos, 1400 m, 20 Dec.
1980, Cutierrez-O. s.n. (IBUG); El Savial. 3 km al W de
la earretera Guadalajara— Ixtlahuacdn del Rio, 1630 m, 6
May 1984. Gallegos s.n. (IBUG). Zacatecas: Moyahua, 1 i
km al S de Moyahua, 1400 m, 25 June 1957, Rzedowski
9135 (ENCB, MEXU, MICH); Santa Rosa, arroyo Los
Aquacastizcos, 21 Oct. 1992, Ennquez-E. 327 (IBUG).

Agradecimientos. A los curadores dc los her-
barios ENCB, IBUG, JAL MEXU, MICH, MO. US,
WIS, ZEA por los espeefmenes facilitados eu cal-
idad de prestamo. A Manuel Flores Martinez por la
elaboration de la ilustracion.

Literatura Citada

Felger, R. S. 1971. The distribution of Magnolia in north¬
western Mexico. J. Arizona Acad. Sci. 6: 251—253.
Howard, R. A. 1948. The morphology and systematics of
West Indian Magnoliaceae. Bull. Torrey Rot. Club 78:
458-463.

Rzedowski, J. & R. McVaugh. 1966. Vegetacidn de Nueva
Galicia. Contr. Univ. Mich. Herb. 91: 1—123.

Spach, E. 1839. Magnolia sect. Theorhodon. Hist. Nat.
Veg. Phan. 7: 470.

Vazquez-G., J. A. 1990. Taxonomy of the Genus Magnolia
in Mexico and Central America. MS Thesis, University
of Wisconsin, Madison.

- . 1994. Magnolia (Magnoliaceae) in Mexico and

Central America: A synopsis. Brittonia 46: 1-23.

Iaxonomic and Morphological Notes on Hypoxis angustifolia
(Hypoxidaceae) from Africa, Madagascar, and Mauritius

Justyn a Wil and-Szym a riska

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A., and
Department of Geobotany, A. Miekiewicz University, Al. NiepodlegloSci 14. 61-713 Poznan,

Poland, wiland@amu.edu.pl

Zbign lew A da mski

Electron Microscopy Laboratory, A. Miekiewicz University, ul. Grunwaldzka 6, 60-780

Poznan. Poland

ABSTRACT. Pour varieties within //. angustifolia
Lamarck, the most common species of the genus
Hypoxis L. in Africa, are recognized, based in part
on differences of seed testa features between pop¬
ulations in Mauritius, Madagascar, and continental
Africa. SUM micrographs ot seeds of H. angusti¬
folia var. angustifolia , variety luzuloides (Robyns X
four nay) Wiland, and variety madagascariensis Wi-
land are included, and full descriptions of the
above varieties as well as //. angustifolia var. buch-
ananii Baker are given.

Key words: Africa, Hypoxidaceae, Hypoxis ,
Madagascar, Mauritius.

Hypoxis angustifolia is the most common and
widely distributed species of Hypoxis (Hypoxida¬
ceae) in Africa and on West Indian Ocean islands
(Baker, 1877: 369, 1878a: 265, 1878b: 111, 1896:
180—181, 1898: 378; de Cordemoy, 1895: 188; Du¬
rand X Schinz, 1895: 231, 1896: 260; Engler,
1908: 352; Durand X Durand, 1909: 554; Ncl,
1914b: 303; De Wildeman, 1921: 33; Bews, 1921:
64; Hutchinson X Dalziel, 1936: 394; Guinea Lo¬
pez, 1945: 258; Williams. 1949: 302; Robyns X
Tournay, 1955b: 388; Troupin, 1956: 207; An¬
drews, 1956: 306; Binns, 1968: 53; Hepper, 1968:
172; Morton, 1968: 31: Geerinck, 1971: 5—6; Jacot
Guillarmod, 1971: 148; Troupin, 1971: VI. 277;
Ross, 1972: 132; Wickens, 1976: 160; Marais,
1978; Nordal et al., 1985: 24; Champluvier, 1987:
81; Nordal X Iversen, 1987: 34; Agnew X Agnew,
1994: 313; Thulin, 1995: 31; Zimudzi, 1996: 15;
Nordal, 1997: 87). Throughout its range from Guin-
ea-Gonakry to Mauritius and to South Africa (for a
map with general distribution sec Wickens, 1976:
316) this species occurs from sea level to about
2000 m (Nordal et al., 1985, suggested even to
3000 m). It is common in a wide variety of open

Novon 12: 142-151. 2002.

habitats, including secondary habitats such as road
borders, grazed pastures, fields, and plantations.

Hypoxis angustifolia is regarded as a “good” spe¬
cies on the basis of cytologieal observations: its
karyotype is established at the diploid (rarely tri-
poloid) level, with no indications of apomixis (Nor¬
dal et al., 1985). It is characterized by wide mor¬
phological variation but was usually treated as a
single species without infraspecific taxa (e.g., Geer¬
inck. 1971). Some authors (Nel, 1914b; Marais,
1978) facing the morphological variation within this
species observed that it should probably be divided
into microspecies. Until the present, however, only
the variety H. angustifolia var. buchananii Baker
had been described from the southeastern African
coast.

for this paper a thorough study of morphology
of //. angustifolia was conducted on specimens
from Central Africa, Madagascar, and Mauritius
and compared with specimens from other regions
of the continent. Characters shared by almost all
populations of the species include a membranous
tunic, corymbose inflorescences w ith long pedicels,
and two-branched trichomes on the leaves. Field
observations in Tanzania show that the plant habit
of //. angustifolia is highly influenced by its habi¬
tat. Plants growing in places exposed to sun and
wind, on rocks, especially along disturbed areas
like paths, tend to have rather rigid, narrow leaves.
Plants occurring on the forest bottom tend to have
wider leaves, which are soft in texture. Specimens
in numerous herbaria (B, BR, BRLU, DAR, k. MO.
NHT, P, POZG, TAN, UPS, WAG) often differ wide¬
ly in dimensions and sometimes also shapes of te-
pals, anthers, pistils, and bracts; it seemed possible
al first that this morphological variation might he
as great within a single population as between
widely separated populations.

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Wiland-Szymariska & Adamski
Hypoxis angustifolia

143

Figure I. Hypoxis angustifolia var. luxuloides (Robyns & Toumay) Wiland. — A— I. Morphological forms of style with
stigma. — J. Anomalous style and stigma. — k. Stamen, form «. — L. Stamen, form fi. — M-l’. Bracts. A & N from Bamps
& Malaisse 8052 , BR; B from Quarre 5056, BR; C from Noiifalise 826, BR; D from Gillet s.n., BR; E from Troupin 1243,
BR; F from Lejoly 82/1013. BRLU; G from Liben 620. BR; H from Van Oosten 120, BR; I from Pauwels 6009. BR; J
from Vanderyst s.n.. BR; K & 0 from Williams 18. BR; I. & M from lisowski 86119, POZG; P from Lewalle 5486, BR.

Studies conducted on mature flowers from 51
herbarium sheets of H. angustifolia from Central
Africa provided the observation that morphologies
of style and stigma are the most variable characters

in this species. Ten forms of gynoecium were ob¬
served. differing in a ratio of stigma to style length
atid stigma shape (Fig. I A— I). they can be divided
into two groups according to the ratio between style

144

Novon

and stigma: the first group, when the style is much
longer than the stigma (Fig. 1A, C, I). F, H), anil
the second group when both are of equal size (Fig.
IB, G, I) or the stigma is longer than the style (Fig.
IE). The style can be trigonal in cross section (Fig.
1 A, B, C, I), G, I) or terete (Fig. IE, F, H), visibly
tapering toward the apex (Fig. I A, C, E) or wider
only in the basal part and then of a constant width
(Fig. II), F, H, I) or equal throughout its length
(Fig. IB, G). It can be massive in shape (Fig. IB.
E, F) or thin and delicate (Fig. II). H). The stigma
is usually composed of three lobes except in one
case where it is divided in four (F ig. 1C). The lobes
can be totally fused, and in this case the papillae
will be distributed in three stripes (F ig. I A. B. D,
F„ G, 1). All of these stripes can be of an equal
length (Fig. 1A, B. I). E) or unequal (Fig. 1G. I).
They might be of uniform width (F ig. 1A, D), ta¬
pering toward the stigma’s apex (Fig. IB. FA or ta¬
pering toward its base (Fig. 1G, I). Another variable
feature is stigma shape. It tends to be more or less
pyramidal in shape, tapering toward the apex (Fig.
IB, I), E), ovoid (Fig. I A), or rather oblong (Fig.
1G. I). Sometimes the stigma consists of three free
lobes (Fig. IF". H) covered with papillae on their
edges and adaxially. Eobes can be triangular in
shape (Fig. 1 F) or oblong (Fig. 1 IT). In several spec¬
imens tb<‘ stigma was divided in the apical [tart into
4 obtuse lobes covered with stripes of papillae on
margins (Fig. 1C). In one case an anomaly in the
gynoecial structure was observed (Fig. 1J): one
stripe of papillae on a stigma is much shorter and
there is a spherical appendage covered with papil¬
lae below it, and 2-branehed trichomes grow out of
the style. I he occurrence of trichomes on (he style
is extremely rare and may be considered a devel¬
opmental anomaly. The most common forms of gy-
noecium are depicted in F igure I A and II). F re¬
quently plants with a different gynoecial type look
alike otherwise. No correlation among gynoecial
structure, other morphological features, or geo¬
graphical range was observed. Palynological and
molecular studies might provide insights into the
origin of this natural infraspecifie diversity.

I he fact that the ratio of style to stigma length
is not constant within //. angustifolia negates the
taxonomic value of the sections proposed by Nel
(1914a, 1914b), who emphasized this character as
one of the most important in his subdivision of Hy-
poxis. It was already observed as a common situa¬
tion in this genus (Nordal et al.. 1985). In the de¬
scription of section Angustifoliae, Nel (1911a)
implied that species of this group posses a gynoe-
cium with a stigma much shorter than the style, and
only exceptionally equal. Although Nel gave //.

dinteri Nel as an example for this section, the sec¬
tional epithet is taken from //. angustifolia , a spe¬
cies with a much greater variability in these char¬
acters. Nel's observations of variability within //.
angustifolia were limited to dimensions of leaves.

I lie morphological variability of the reproductive
organs in II. angustifolia involves stamens as well.
I he stamens are biseriate, with the inner cycle usu¬
ally shorter than the outer, with a difference of
about 0.5-1. 0 mm, entirely due to the length of
filaments. Equally long stamens were observed only
in a few plants. Filaments are subulate, with one
distinct vein and almost membranous edges. Bare¬
ly, the y are linear and wider only in the basal part.
There are two anther forms named here a and /3
(Fig. IK. E). Both are emarginate at the apex, ex¬
actly as Nel (1914a. 1914b) observed, and sagittate
at the base. Anthers a are usually relatively long
in comparison to the filaments and reach 2—3 mm
in length (exceptionally I mm). Thecae are oblong
and obtuse on both ends (Fig. IK). Raphides are
often visible in the outer tissue. Anthers /3 are usu¬
ally shorter. 1.2-2. 5 mm long (exceptionally 3 mm
long). I heeae are obtuse in the basal part. The con¬
nective is thickened in the apical part and often
prolonged into tiny appendages, a phenomenon that
is especially distinct after dehiscence (Fig. IE).
Anthers ir are much more common than (3 and were
observed in 60% of studied plants.

I here is usually a difference between bracts sub¬
tending older and younger flowers in one inflores¬
cence in most species of Hypoxis. Upper younger
bracts in //. angustifolia are usually one-veined
and similar to each other. Older bracts are more
variable. All bracts are subulate, acute and more
or less keeled, covered with trichomes along the
midrib abaxially (Figs. 1M— P). The most common
bracts are 5-15 mm long, usually less than I mm
wide; lightly keeled or plane in the middle, flatly
hooded al apex, single veined, villous along the
midrib, hispid at apex, exceptionally with simple
trichomes on edges near apex; sometimes with
some raphide cells or red stripes on midrib (Fig.
IN, ()). Less common are strongly keeled bracts
with two smaller veins parallel to the midrib; 12—
17 mm long and 1.5— 2.0 mm wide; with oppressed
trichomes and occasionally with several simple tri¬
chomes on edges at apex, very often with raphide
cells in tissue (Fig. 1M). The least common are
bracts ca. 12.5 mm long and 0.5-1. 7 wide, single-
veined. pubescent with long (to 4 mm) trichomes,
almost glabrous in basal part, sparsely ciliate on
margins with short simple trichomes (Fig. IP).

During the study of Madagascar plants it was
observed that in many specimens of H. angustifolia ,

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Wiland-Szymariska & Adamski
Hypoxis angustifolia

145

especially II. angustifolia var. madagascariensis,
the apices of the inner tepals are covered with club-
shaped one- or few-celled papillae (Fig. 3B). Such
papillae sometimes also occur on the tepals of con¬
tinental plants; however, they are usually not as
visible as in Madagascar populations. This feature
of inner tepals has so far not been observed in any
other species of Hypoxis ; in other species the inner
tepals are usually apically flat and entire, although
they are sometimes acute. The outer tepals are al¬
ways somewhat hooded at the apex and possess a
small appendage covered with papillae or short tri-
chomes (see Wiland. 1997).

A sculptured seed testa has often been noted as
a shared character in //. angustifolia (Nordal et al..
1985; Nordal & Iversen. 1987). It is usually brown
in coloration, in some specimens showing a bit of
iridescence, that makes it similar to the testa of
some American species (Brackett. 1923). SFM mi¬
crographs from African populations show ovoid
seeds to 2 mm long with a short papillose append¬
age at the apex, and the testa with pyramidal point¬
ed projections covered with a wrinkled brown cu¬
ticle (Fig. 2A. B) (see also Nordal et al.. 1985: 19;
Nordal & Iversen, 1987: 45). Seeds of //. angusti¬
folia var. buchananii are black (Y. Singh, pers.
comm.), but unfortunately they were not available
for a closer examination.

An examination of materials from Madagascar
and Mauritius at MO revealed that numerous spec¬
imens of H. angustifolia possess seeds with a testa
sculpture different from that described above. No
other authors dealing with Hypoxidaceae from
these islands (Baker, 1877; de Cordemoy, 1895;
Perrier de la Bathie, 1950) except Marais (1978)
have noted the seed testa sculpture of II. angusti¬
folia. Specimens from Mauritius, including the type
specimen, have black seeds with colliculate testa
and smooth cuticle (Fig. 2C). Semi-spherical pa¬
pillae are minutely mieropapillate on their surface
(Fig. 2D). Such seeds were mentioned by Marais
(1978) for the plants of Reunion and Mauritius and
depicted on the plate in his paper. In plants from
Madagascar two kinds of testa sculpture are pre¬
sent. A large proportion of specimens have testa
with pyramidal papillae covered with a wrinkled
brown cuticle, as is common in continental plants.
Some specimens, however, possess black seeds
with slightly longitudinally furrowed tuberculate
papillae (Fig. 2E, F). The latter sculpture pebbling
is different from that observed in Mauritius as well
as on the African continent.

Because of the morphological variation of the
seed testa sculpture in //. angustifolia from Maur¬
itius, Madagascar, and continental Africa, recog¬

nition of four varieties is justified. Hypoxis angus¬
tifolia var. buchananii moreover is easily
distinguishable because of its larger habit and be¬
cause it grows in tufts versus the smaller and sol¬
itary-growing plants of other varieties. The island
taxa are probably descendants of the continental
populations. However, H. angustifolia was first de¬
scribed from Mauritius, so the type specimen pos¬
sesses seeds with the least common type ol testa
sculpture.

Hypoxis angustifolia Lamarck, Encycl. 3: 182.
1789. TYPE: Mauritius. Commerson s.n. (ho-
lotype, P, computer image seen).

Perennial herb, 10-53 cm high; rhizome globose,
0.5-1 .2 cm diam. (when dry), white inside, sur¬
mounted by inconspicuous membranous remains of
old leaves, sometimes with some thin fibers; roots
white, ± thick. Leaves grouped in a whitish pseu¬
dostem, outer leaves rarely present, if present not
numerous, ovoid and spathe-like in basal part, lin¬
ear in upper part, acute, to 10 cm long, ca. 0.7 cm
wide, pilose along midrib and margins beneath with
2-branched white trichomes; nervation composed of
5 to 13 veins of unequal size; inner leaves 3 to 10.
grass-like, linear or broadly linear, tapering toward
apex, keeled, often recurved along two prominent
lateral veins, acute at apex, 7—50 cm long, 0.2— 1.0
cm wide, ciliate on margins and midrib beneath or
very sparsely pilose on entire surface; trichomes
2(-3)-branched. 1.3-2. 5 mm long, golden or white,
soft; nervation composed of 5 to 29 veins ol un¬
equal size with 2 or 4 lateral veins prominently
larger than the others. Scapes I to 7. 4-20 cm high.
0.5-1. 1 mm wide, compressed, winged and gla¬
brous in lower half, ciliate in upper half and pilose
only beneath inflorescence; trichomes 2- or 3-
branched; flowers single or in a lax 2- to 6-fiowered
corymbose cyme; bracts subulate or sword-shaped,
acute, often two subtending a solitary flower, some¬
times those subtending first two flowers much larger
than upper ones, 3—17 mm long, 0.2— 1.< mm wide,
keeled, 1-, 3-, or 5-veined, glabrous, villous or
sparsely pilose on midrib abaxiallv, sometimes cil¬
iate on margins; pedicels 5.5—35 mm long, pubes¬
cent or pilose. Tepals 6 (exceptionally 4), yellow,
bright yellow, or outer tepals green and inner yel¬
low, sometimes with a red or brownish stripe along
midrib; outer tepals ovate, acute at apex, 3.5—10.0
mm long, 1.0-3. 3 mm wide, adaxially with a ela-
vate appendage below apex, 5- to 7-veined with
irregular veins, villous or sparsely pilose abaxially;
inner tepals ovate, obtuse, 3. 5-9.0 mm long, 1.2—
4.0 mm wide, 5- to 7-veined with irregular veins.

146

Novon

A

B

C

E

Figure 2. A, li. Hypoxis angustifolia var. luzuloides. — A. Seed. — B. Seed tesla sculpture. C, I). II. angustifolia var.
angustifolia. — C. Seed. — I). Seed testa sculpture. E, F. //. angustifolia var. madagascariensis. — E. Seed. — F. Seed

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Wiland-Szymariska & Adamski
Hypoxis angustifolia

147

glabrous or pilose along midrib abaxially; stamens
usually unequal, outer longer than inner or rarely
equal; filaments subulate; outer stamens 2.2— 4.0
mm long with filaments 1 .3-3.0 mm long; inner sta¬
mens 1.7— 3.5 mm long with filaments 1-2 mm long;
anthers linear, deeply sagittate at base, retuse at
apex, 1 .0—3.2 mm long; ovary obconical, 2—5 mm
long, 1—3 mm wide, almost glabrous or villous; style
0.5-3.0 mm long; stigma 0.6-2. 5 mm long, both
variable in shape. Capsule turbinate, 3.5-14.0 mm
long, 2. 5-4.0 mm diam., glabrous or pubescent,
surmounted by persistent perigon, often splitting in
three lobes; seeds numerous, ovoid, 0.8— 2.0 mm
long, 0.65—0.80 mm wide, with short papillose ap¬
pendage at apex, black or brown; for cuticle and
testa features, see varietal descriptions.

Kky to thi: Varieties of Hypoxis angustifolia

la. Plants growing in tufts; inner leaves 9-10 nun
wide, flat, with at least 4 prominent lateral veins

. H. angustifolia var. buchananii

lb. Plants growing solitary; inner leaves 3-f5 mm
wide, with 2 prominent lateral veins.

2a. Seeds brown; papillae pyramidal; cuticle

wrinkled . //. angustifolia var. luzuloides

2h. Seeds black; papillae semi-spherical or tu-
berculate; cuticle smooth.

3a. Papillae semi-spherical .

. //. angustifolia var. angustifolia

3b. Papillae tuberculate .

.... H. angustifolia var. madagascariensis

Hypoxis angustifolia var. angustifolia

Herb to 5-15 cm high; rhizome 0.8-1. 2 cm diam.
(when dry). Inner leaves 3 to 10, linear, 7—15 cm
long, ca. 0.2 cm wide, ciliate on margins and mid¬
rib beneath or very sparsely pilose on entire sur¬
face; trichomes golden; nervation composed of 7 to
I 1 veins with two lateral veins prominently larger
than the others. Scapes 2 to 7, 4.0-7. 5 cm high,
0.5-0. 8 mm wide, ciliate in lower half, pilose in
upper half; flowers single or in a lax 2-flowered
cyme; bracts subulate, usually two subtending a sol¬
itary flower, 3-7 mm long, ca. 0.2 mm wide, 1-
veined, villous on midrib abaxially; pedicels 7—1 1
mm long, pubescent. Tepals 6, yellow or bright yel¬
low; outer tepals 4-5 mm long, ca. 2 mm wide, 5-
veined, villous abaxially; inner tepals 3.7— 4.5 mm
long, 1.2-1. 4 mm w ide, 5-veined, pilose along mid¬
rib abaxially; stamens unequal; outer stamens 2.4—
2.8 mm long with filaments 2.0-2. 1 mm long; inner
stamens 1.7— 2.5 mm long with filaments 1. 8-2.0

mm long; anthers ca. 1 mm long; ovary ca. 2.5 mm
long, ca. 2 mm wide, villous; style subulate, 1.5—
2.0 mm long; stigma 0.6-0. 7 mm long, composed
of three fused lobes. Capsule 3.5— 4.5 mm long, ca.
3 mm diam., pubescent; seeds ca. 1 mm long, ca.
0.8 mm wide, black; testa covered with smooth cu¬
ticle; semi-spherical papillae minutely micropapil-
late on their surface (Fig. 2C, D).

Habitat. Gravelly soil, along path and roadside
in sun with grasses; in open, humid places (Marais.
1978); 660-770 m altitude.

Observations. This typical subspecies occurs on
Mauritius and probably also on Reunion.

Flowering January and September.

Specimens examined. MAURI PIUS. Montane
Cocotte nature reserve, Lorence 2248 (MO); Ma-
cambe forest, Lorence 660 (MO).

Hypoxis angustifolia var. huchananii Baker, J.
Linn. Soe., Bot. 17: 111. 1878b. TYPE: South
Africa. Natal: Nov. 1875, Buchanan s.n. (ho-
lotype, K).

Herb to 35 cm high; rhizome not seen. Inner
leaves 4 to 6, broadly linear. 16-35 cm long, 0.9-
1 .0 cm w ide, sparsely ciliate on margins and midrib
beneath; trichomes golden; nervation composed of
25-29 veins with 4 lateral veins prominently larger
than others. Scapes 4 to 5, 13.0-19.5 cm high. ca.

1 mm wide, winged and glabrous in lower half, cil¬
iate in upper half, sparsely pilose on the whole sur¬
face below inflorescence; flowers in a very lax 2- to
3-fiowered cyme; bracts subulate, 6—9 mm long, ea.

1 mm wide, 1 -veined, pilose on midrib abaxially;
pedicels 20—35 mm long, sparsely pilose. Tepals 6.
yellow'; outer tepals ovate, acute at apex, ca. 10 mm
long, ca. 3.3 mm wide, with an oblong appendage,
7-veined, sparsely villous abaxially with 2- or 3-
branehed trichomes; inner tepals ca. 9 mm long, ca.
3 mm w ide, 7-veined, pilose along midrib abaxially
to half of its length; stamens unequal; outer stamens
ca. 4 mm long w ith filaments ca. 2.5 mm long; inner
stamens ca. 3.5 mm long with filaments 2.0 mm
long; anthers ca. 3.2 mm long; ovary 4.0— 4.5 mm
long, ca. 1.2 mm wide, villous; style ca. 0.5 mm
long; stigma ca. 2 mm long, pyramidal, obtuse at
apex, wider than style, with three wide stripes of
papillae. Fruits and seeds not seen.

Observations. On the sheet with the type spec¬
imen three plants are present. Two of them corre-

testa sculpture. A and B from Lewalle 5486. MO; C and I) from Lorence 1421. MO; E and F from Lowry, Rakotozafy
& Nicoll 4208, MO.

148

Novon

spond to die description above, whereas one lias
much narrower leaves with fewer veins and belongs
to the widespread continental variety. So far. //. an-
gustifolia var. buchananii was cited from only two
other collections: from Inanda Wood 426, 771 (Bak¬
er, 1896: 180-181). Whereas //. angustifolia var.
luzuloides occurs mainly in open places with sun¬
light, Baker (1896: 180) observed that this is "a
large shade grown variety, with longer leaves of
very thin texture; pedicels longer, very slender.”
Pedicels of this taxon are indeed usually longer
than in the other varieties; however, this feature
overlaps somewhat between 11. angustifolia var.
buchananii and //. angustifolia var. luzuloides. This
variety, which occurs on the very edge of the range
of //. angustifolia, certainly deserves further study.

Specimen examined. REPUBLIC OF SOUTH AFRI¬
CA. Natal: Umzinto, Rvnie Park, Strey 5974 (BR).

Hypoxis angustifolia var. luzuloides (Robyns &
lournay) Wiland, comb, et stat. nov. Basionym:
Hypoxis luzuloides Robyns & Tournay, Bull.
Jard. Bot. Plat 25: 254. 1955. TYPE: Congo-
Kinshasa. Kivu, Tschambi, G. F. de Witte 1130
(holotype, BR).

Herb 10—53 cm high; rhizome 0.8— 1.2 cm diam.
(when dry). Inner leaves 3 to 12, grass-like, linear,
tapering toward apex, usually keeled, often re¬
curved along two prominent lateral veins, acute at
apex, 10—50 cm long, 0.3— 0.8 cm wide, filiate on
margins and midrib beneath or very sparsely pilose
on entire surface; trichomes golden or white; ner¬
vation composed of 5 to 13(23) veins with 2 thicker
than others. Scapes I to 6, 5-20 cm high. ca. 1 mm
wide, ciliate in lower half, pilose in upper half;
flowers single or in a lax 2- to 6-flowered cyme;
bracts subulate, 12-17 mm long, (0.5—) 1 .0— 1 .7 mm
wide, I- or 3-veined, villous on midrib abaxially,
sometimes ciliate on margins; pedicels 12—25 mm
long, pubescent. Tepals 6 (exceptionally 4), yellow,
bright yellow, or outer tepals green and inner yel¬
low, sometimes with a red stripe along midrib; outer
tepals 5-8 mm long, 2-3 mm wide, 5- to 7-veined,
villous abaxially; inner tepals 4—7 mm long, 3-4
mm wide, 5- to 7-veined, pilose along midrib abax¬
ially to % of its length; stamens usually unequal;
outer stamens 3-4 mm long with filaments 2. 5-3.0
mm long; inner stamens 2-3 mm long with fila¬
ments 1.5— 2.0 mm long; anthers 1.5— 3.0 mm long;
ovary 2—5 mm long, 1—3 mm wide, villous; style 1 —

3 mm long, stigma 0.7-2. 5 mm long, both variable
in shape. Capsule 7—14 mm long, ca. 3 mm diam.,
pubescent; seeds ca. 2 mm long, 0.65-0.80 mm

wide, brown; papillae pyramidal, testa covered with
wrinkled cuticle (Fig. 2A, B).

General distribution. Intertropical and southern
Africa, Madagascar, Mascarenes.

Habitat. Miombo; dry forest; bush; various
grasslands; wooded grassland in river valleys; river
banks; lakeshores, moist depressions; dunes; on
base of termite mounds; pastured grassland on
rocky soil; granite slabs on slopes; secondary grass¬
land on laterite; fallow fields in grassland, shrubby
fallow fields on sand; moist roadsides, roadside
ditches, along trails and roads in the forest, dis¬
turbed forest, rocky gorge on steep grade; cultiva¬
tion ol manioc; loamy soil, with kaolin, rocky soil.
Altitude 0-2800 m.

Observations. Flowering from January to May.
Medicinal plant; pulp from the rhizome is useful
for healing pustules and infected wounds (Gillet &
Paque, 1910). Its chemical compounds were stud¬
ied by Sibanda et al. (1990). Hypoxis luzuloides Ro¬
byns & lournay (1955a) was described from Cen¬
tral Africa as a separate species in Nel’s section
Argenteae based on the fact that in the type spec¬
imen the stigma was longer than the style. This is,
however, weak support as discussed above. Various
authors included this name in the synonymy of //.
angustifolia (Geerinck, 1971; Nordal el al., 1985;
Zimudzi, 1996; Nordal, 1997; Wiland-Szymariska,
2001). 'I'he type specimen of H. luzuloides possess¬
es all features common in //. angustifolia and seeds
with the testa sculpture and wrinkled cuticle typi¬
cal of continental populations. I have therefore
transferred this specific epithet to the varietal rank
rather than choosing a new epithet and type spec¬
imen.

Selected examined specimens. SIERRA LEONE. Ibis-
well 37 (K), Morton SL3532 (K). NIGERIA. Kaduna: lg-
abi Dist., Anara, Keay 25779 (K). Plateau: Jos Plateau.
Naraguta, Lawlor A- Hall 101 (K). Taraba: Mambila Pla¬
teau. Chapman 66 ( K). Adamawa: Chappal Hendu,
Chapman 4424 ( K ). CAMEROON. North-West: Bame-
noa, Balut-Ngemba, Richards 5303 (K). CHAD. Gluiri-
Baguirmi: Region du Lac Fitri entre Boullai et Djannaw,
Chevalier 9485 (K). CENTRAL AFRICAN REPUBLIC.
Baniiiigui-Burigoruu: 8°28'N, 20°53'E, Fay 3105 (K).
SUDAN. Darfur: Jabel Maria. Widens 1945 (K). ETHI¬
OPIA. Timlin, Hynde A Tadesse 3369 (UPS). Amhara:
Blue Nile Gorge, Gilbert 2162 (K). Oroiniya: 5 km N of
Neghelle, Vollesen 26-10 (K). Southern: 1 I km S of Dilla.
Friis, Tadesse A Jefford 4274 (K). CONGO-KINSHASA.
Bas-Zai’re: Lutete, Hens 221 (BR, K, P). Kinshasa: Ki-
muenza, Lejoly 82/1013 (BR, BRLU). Bandundu: Chutes
remho. Rreyne 2590 (BR). Haut-Zaire: Ahok, Scops 168
(BR); Nioka, Lejoly 3338 (BREL ). Lisowski 48484
(POZG). Kasai Oriental: Lomami, Dandoy 206 (BRLU).
Kivu: Kikanga, Requaert 4211 (BR). Shaba: Tshibonde,
Mullenders 2346 (BR); Kapolowe, Lisowski R- 72 76
(POZG). RWANDA. Parc National de LAkagera, Rouxin

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2002

Wiland-Szymariska & Adamski
Hypoxis angustifolia

149

& Radoux 434 (BR); Gabiro, Burgeon 17 & 34 (BR); ki-
bugabuga, Liben 620 (BR); Nemba, Bouxin & Radoux
1415 (BK); Bugesera, Augier 2898 (BR); Matinza, Bouxin
& Radoux 1008 (BR); Lulama, Bouxin A: Radoux 418
(BR); Mimuli. Troupin 4230 (BR); Lukionji, Troupin 0776
(BR); Kakitumba, Christiaensen 736 (BR). BURUNDI.
Route Bubanza— Musigati, Ia- walle 4 189 (BR); Bujumbura.
Ijewalle 5486 (BR. MO); plaine de la Ruzizi. Heckmans
2906 (BR, MO), Lambinon 75/44 (WAG); Bulamata, Ger¬
main 7191 (BR); Rumonge, Lewalle 5029 (BR). UGAN¬
DA. Buslienyi: Kasharara, Rwaburindore 1872 (MO).
Mbarara: Mbarara, Purseglove 602 (K); Kilaro, Rwabu¬
rindore 2064 (MO). Masaka: 1-2 km E of Kikoma. Lye
4440 (k, UPS). Mpigi: Entebbe, Brown 293 (k). Muko-
no: Busana, Bugerere, Eggeling 527 (k). luanga: on low¬
er NVY slopes of Rugiri hill. Wood 702 (k). kumi: Bu-
kedea, Katende K226 (K). Moroto: base ol Mt. Debasien,
near Mornita, Eggeling 5798 (k). Kapchorwa: NE Mt.
El gon, Tweedie 1110 (K). Mbale: N Bugisu, Muyembe
escarpment, Norman 256 (k). KENYA. Western: kaka-
mega, Carrall 429 (k). Nyanza: Oyugis— Kendu Road,
Hanid & Kiniaruh 651 (MO); Tinderet Forest Res.. Maas
Geesteranus 5220 (BR). Kif’t Valley: Kipkarren, Brod-
hurst-Hill 173 (k); 8 km S of Nandi Hills. Davidse 7121
(MO); Menengai, Maas Geesteranus 4573 (BR). Central:
Vlei near 14 Falls, Thika— Donyo Sabuk Road, Verdcourt
1965 (k). Eastern: Moyale, Gillet 12799 (BR, k). Coast:
Maji ya Chumvi, Kassner 453 (k); Shimba Hills, Longo
Mwagandi area, Magogo <£• Glover 311 (MO); Shimba
Hills, Pengo Hill, Bamps 6331 (BR). TANZANIA. Ka-
gera: kabogo, near Shanga, Tanner 5857 (K). Mara: Wo-
gakuria Hill, Greenway & Turner 12007 (K). Shinyanga:
Shinyanga, Bax s.n. (K). Arusha: Essimingori Mountain,
Wiland & Mboya 174 (MO); Mt. Meru. Bjprnstad 233 (k.
DAR); between Magugu & Babati, Polhill & Paulo 2373
(BR). Kilimanjaro: road to Horombo, McCusker 98 (k);
Moshi— Arusha road. Polhill 6k Paulo 984 (BR). Tanga:
12 km E of Old Korogwe Eriksson, Kalema & Leliyo 572
(NHT); Machui, Tanga— Pangani road, Faulkner 1844 (B,
BR. k); Mandundu, Archbold 1007 (DAR). Dodoma:
Mpwapwa, kiboriani Mountains, Hornby & Hornby 2096
(k). Morogoro: Uluguru Gebirge, Sehlieben 3245 (B, BR,
k, MO); Mgeta, Wigfield 2313 (DAR). I’wani: 59 mi. from
Dar es Salaam on main road to Morogoro, Welch 296 (BR,
k); about 15 mi. S of Dar es Saalam, between kambiji
and Mjimwema. Batty 1347 (MO); Mafia Island, Rufiji.
Wallace 811 (k); Dar es Salaam, Harris 244 & 1234
(DAR); kibaha Research Station, Banda 314 (NHT). Zan¬
zibar: Zanzibar Island, Faulkner 2940 (BR, k); Pemba
Island, Williams 18 (BR, k). Iriuga: near Isele village,
Wiland & Mboya 73 (MO); Chimala escarpment, Richards
7513 (k); kidatu. Mhoro 424 (DAR). Lindi: Nachingwea,
Anderson 947 (NHT). ZAMBIA, van Rensburg 1317 (P).
MOZAMBIQUE, de Honing 7406 (BR), Quintas 157 (BK),
Bates 520 (BR), Pope & Miller 588 (BR). REPUBLIC OF
SOUTH AFRICA. Sehlieben 7345 (BR), Ackock 10832
(BR). MADAGASCAR. Fianarantsoa: 10 km W of Ivato,
Croat 29634 (MO, TAN); Ranomafana Res., NE of Fian-
arantsoa, Leeuwenberg et al. 14147 (MO, TAN). Tanana¬
rive: Forest Station of Manjakatiompo, Groat 28950 (MO,
I AN). Toamasina: swamp of Torotonsifotsy, Rakotomal-
aza, Andrianasolo, Rasolomanana & Andraintsiferana
1340 (MO, TAN); 7 km E of Anjozorobe, Schatz, Stevens
& Lowry 3483 (MO, TAN). Toliara: 5 km S of Manam-
baro, 23 km W of Fort Dauphin, Miller & Randrianasolo
6218 (MO, TAN). Tulear: Chain Anosyennes from Fort
Dauphin to Ranamafona, Croat 31840 (MO).

Hypoxis angustifolia var. nuulagascariensis Wi¬
land, var. nov. TYPE: Madagascar. Toamasina:
Ambanizana, Masoala Peninsula, along Andro-
ka River S of MBG house, 15°39'30"S.
49°57'30"E, June 1993, Zjhra cQ Hutcheon
273 (holotype, MO). Figures 2E. F. 3.

flaec varietas a 11. angustifolia var. angustifolia et //.
angustifolia var. luzuloides seminis testa tubereulato-acu-
leata differt.

Herb to 10—15 cm high; rhizome 0.5— 1.2 cm
diam. (when dry), surmounted by inconspicuous
membranous remains of old leaves to 1 cm high.
Inner leaves 5 to 16, not grouped in a distinctive
pseudostem, linear, tapering toward apex, keeled,
acute at apex, 9—38 cm long, 0.2— 0.4 cm wide, al¬
most glabrous or sparsely ciliate on margins and
midrib beneath; trichomes white; nervation com¬
posed of 5 to 15 veins. Scapes 1 to 6. 2.5—13.0 cm
high, 0.5— 1.1 mm wide, elongate with age, ciliate
in lower half, sparsely pilose in upper half; flowers
single or in a lax 2- or 3-flowered cyme (Fig. 3C);
bracts sword-shaped, w'ith membranous edges,
those subtending the first two flowers much larger
than the third, often two subtending a solitary (low¬
er, the lowest 12-17 mm long, 1.0— 1.7 mm wide,
3- or 5-veined, the uppermost 5—7 mm long,
sparsely pilose abaxially or glabrous; pedicels 5.5—
12.0 mm long, glabrous or sparsely pilose. Tepals
6 (exceptionally 4), yellow; outer tepals 3.5— 4.3 mm
long, 1.0— 1.5 mm wide, 5-veined, sparsely pilose
abaxially; inner tepals 3.5— 4.2 mm long, 1.2— 1.6
mm wide, with numerous papillae around apex (Fig.
3B), 5-veined, glabrous; stamens equal, ea. 2 mm
long with filaments 1.4— 1.6 mm long, or unequal
with outer stamens ca. 2.2 mm long with filaments
1.3— 1.4 mm long, and inner stamens ca. 1.8 mm
long with filaments 1.0— 1.1 mm long; anthers 1.2—
1.4 mm long; ovary 3-4 mm long, 1.0— 1.7 mm
wide, almost glabrous; style 1.2— 1.3 mm long, tri¬
gonous; stigma 0.7— 0.8 mm long, composed of three
oblong, partially fused lobes. Capsule 4.5— 8.0 mm
long, ca. 2.5— 4.0 mm diam., glabrous; seeds 0.8—
0.9 mm long, 0.65—0.7 mm wide; testa black with
slightly longitudinally furrowed tuberculate papil¬
lae, cuticle smooth (Fig. 2E, F).

Habitat. Wet rocks; on trail in primary forest;
475—850 m altitude.

Vernacular name. Mangitrampango (Miller c fc
Lowry 3905).

Observations. Flowering in February, April.
June, and October. From //. angustifolia var. luzu¬
loides it differs in the seed testa sculpture, larger
bracts, smaller flowers, and more distinctive pap-
illation on the apex ol the inner tepals. In Mada-

150

Novon

Figure 3. Hypoxis angnstifolia var. madagascariensis VC i land . — A. Habit. — B. Papillae on apex of inner tepal. — C
Inflorescence. A & B from Rakoto & Turk fid, MO; C from Zjhra & Hutcheon 273 (MO).

gascar there is no distinct difference in geograph¬
ical distribution or habitat preference between
these two taxu. Field studies are needed to evaluate
the relationship between them.

Paralypes. MADAGASCAR. Toamasina: Maroantse-
tra. mountains NE of village Ambanizana, hurry, Rako-
tofazy & Nicoll 420fi (MO). Fiunaruntsou: National Park
of Ranomafana, Rakoto <£ Turk 33 (MO. TAN). Antsiran-
ana: Nature Reserve of Marojejy, NW of Mandena, Miller
& hurry 3906 (MO). Tainatave: ca. 50 km NW of Tan -
atave, trail from Fotsimavo to Nature Reserve of Betam-
pona. Gentry 11311 (MO. TAN).

Acknowledgments. We thank the curators and
workers of all above-mentioned herbaria for their

help and hospitality. Our special thanks are ad¬
dressed to Petra de Block and Yashica Singh for
their contributions, to Roy Gereau for his comments
and suggestions during preparation of the manu¬
script, to Porter P. Lowry II and the MBG employees
in TAN, and to the editors of the MBG Press for
their help. Fieldwork was possible thanks to Nation¬
al Geographic Society grant number 6891-00. Ad¬
ministrative divisions used in the text were mainly
based on maps available on http://www.fao.org.

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New Combinations in Chinese Pittosporum (Pittosporaceae)

/hang Zhiyun

Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

zhangzy@ns.ibcas.ac.cn

Nicholas J. Turland

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

nicholas.turland@mobot.org

Abstract. As part of a revision of Chinese Pit¬
tosporum Banks ex Gaertner lor the forthcoming ac¬
count of Pittosporaceae in the Flora of China, Vol¬
ume 9, it is necessary to publish seven new
combinations. All of the taxa were originally de¬
scribed at specific rank. Six are morphologically
similar to five other species, under which they are
reduced to varieties: Pittosporum balansae Aug.
DC. var. chatterjeeanum (Gowda) Zhi Y. Zhang &
Turland, comb, et stat. nov., Pittosporum glahratum
Lindley var. wenxianense (G. H. Wang & Y. S. Lian)
Zhi Y. Zhang X Turland, comb, et stat. nov., Pit¬
tosporum kweichowense Gowda var. buxifolium (K.
M. Feng ex C. Y. Wu) Zhi Y. Zhang X Turland,
comb, et stat. nov., Pittosporum kweichowense var.
podocarpifolium (C. \. Wu) Zhi Y. Zhang & Tur¬
land, comb, et stat. nov., Pittosporum podocarpum
Gagnepain var. hejiangense (H. Y. Su) Zhi Y. Zhang
& Turland, comb, et stat. nov., and Pittosporum reh-
derianum Gowda var. ternstroemioides (C. Y. Wu)
Zhi Y. Zhang X' Turland, comb, et stat. nov. The
remaining new combination is Pittosporum pentan-
drum (Blanco) Merrill var. formosanum (Hayata)
Zhi Y. Zhang & Turland, comb, nov., which is the
correct name at varietal rank under P. pentandrum
when P. formosanum Hayata var. formosanum and
variety hainanense Gagnepain are considered to be
synonyms.

Key words: China, Pittosporaceae, Pittosporum.

As part of a revision of Chinese Pittosporum
Banks ex Gaertner for the forthcoming account of
Pittosporaceae in the Flora of China, Volume 9
(Zhang et ah, in press), all names described in this
family based on Chinese types were carefully eval¬
uated. It was decided that seven new combinations
were necessary: six of them originally described as
species, here reduced to varietal rank, and the sev¬
enth the correct name for a variety where an au¬

tonym has priority over the varietal epithet cur¬
rently in use.

1. Pittosporum balansae Aug. DC., Bull. Herb.

Boissier, ser. 2. 4: 1071. 1904.

la. Pittosporum balansae var. chatterjeean-

um (Gowda) Zhi Y. Zhang & Turland, comb,
et stat. nov. Basionyrn: Pittosporum chatter¬
jeeanum Gowda, J. Arnold Arbor. 32: 318.
1951. TYPE: “Burma: Kang-fang” [actually
China. Yunnan: Tengehong Xian, Gangfang],
1500 m, thickets along river banks, “4-XIII-
1938" [sic!], Kingdon Ward 75 (holotype, A).

Pittosporum chatterjeeanum is sufficiently similar
morphologically to P balansae to be nonspecific. It
differs in having lanceolate leaves 1—3.5 cm wide,
vs. oblong leaves 2-5 cm wide in P. balansae var.
balansae and narrowly lanceolate leaves 1 — 1.5 cm
wide in P. balansae var. angustifolium Gagnepain.
Therefore, treating P. chatterjeeanum as a variety of
P. balansae is appropriate. Both P. balansae var.
balansae and variety angustifolium occur in west¬
ern Guangdong, southwestern Guangxi, and Hai¬
nan, as well as in Vietnam, while variety chatter¬
jeeanum is disjunct from the other varieties,
growing in western Yunnan (Tengehong Xian) and
adjacent northern Myanmar.

2. Pittosporum glabratum Lindley. . I. Unit. Soe.

London 1: 230. 1846.

2a. Piltosporum glahratum var. wenxianense

(G. H. Wang & Y. S. Lian) Zhi Y. Zhang &
Turland, comb, et stat. nov. Basionyrn: Pittos¬
porum wenxianense G. H. Wang & Y. S. Lian,
Acta Phytotax. Sin. 34: 210. 1996. I V PE: Chi¬
na. Gansu: Wen Xian, Bikou, 1 100 m, 16 Aug.
1993, K S. Lian et id. 93—452 (holotype,
NWTC not seen).

Novon 12: 152-154. 2002.

Volume 12, Number 1
2002

Zhang & Turland
Chinese Pittosporum

153

From its original description, Pittosporum wen-
xianense is sufficiently similar morphologically to
P. glabratum to he conspecific. It differs in having
leaves obovate to narrowly obovate, with a truncate
to obtuse apex, vs. leaves oblong, oblanceolate, nar¬
rowly lanceolate, or beltlike, with an acute apex, in
both P. glabratum var. glabratum and P. glabratum
var. neriifolium Rehder & E. H. Wilson. Therefore,
treating P. wenxianense as a variety of P. glabratum
is appropriate. Pittosporum glabratum var. glabra¬
tum occurs in Fujian, Guangdong, Guangxi, Gui¬
zhou, Hainan, and Hunan, as well as in Vietnam,
while variety neriifolium has a similar range, oc¬
curring also in Hubei, Jiangxi, and Sichuan, but is
not yet known in either Hainan or Vietnam. Pittos¬
porum glabratum var. wenxianense is known only
from southern Gansu (Wen Xian), which is at the
northern edge of the distribution of Pittosporum in
China.

3. Pittosporum kweichowense Gowda, J. Ar¬
nold Arbor. 32: 296. 1951.

3a. Pittosporum kweichowense var. huxifol-
■ him (K. M. Feng ex C. Y. Wu) Zhi Y. Zhang
& Turland, comb, et stat. nov. Basionym: Pit¬
tosporum buxifolium K. M. Feng ex C. Y. Wu,
FI. Yunnan. 3: 318. 1983. TYPE: China. Yun¬
nan: Xichou Xian, 26 Sep. 1947 (fr), A. M.
Feng 12083 (holotype, KUN not seen).

3b. Pittosporiun kweichowense var. podoear-
pifolium (C. Y. Wu) Zhi V. Zhang & Turland,
comb, et stat. nov. Basionym: Pittosporum po-
docarpifolium C. Y. Wu, FI. Yunnan. 3: 318.
1983. TYPE: China. Yunnan: Funing Xian,
foot of limestone mountain, I I Apr. 1940 (II),
C. W. Wang 88305 (holotype, KUN not seen).

From their original descriptions, Pittosporum
buxifolium and P. podocarpifolium are sufficiently
similar morphologically to P. kweichowense to lie
conspecific. Pittosporum buxifolium differs in hav¬
ing obovoid capsules, vs. cylindric capsules in P.
kweichowense , while P. podocarpifolium differs in
having lanceolate leaves 0.5— 0.8 cm w ide, vs. nar¬
rowly oblong leaves 1—2.2 cm wide in P kweichow¬
ense. Therefore, treating P. buxifolium and P. po¬
docarpifolium as varieties of P. kweichowense is
appropriate. Pittosporum kweichowense var. kwei¬
chowense occurs in southwestern Guizhou (Anlong
Xian), western Hunan (Zhijiang Xian), and south¬
eastern Yunnan (Wenshan Xian), while both variety
buxifolium and variety podocarpifolium are known
only from southeastern Yunnan (Guangnan Xian
and Xichou Xian, and Funing Xian, respectively).

4. Pittosporum pentandrum (Blanco) Merrill,

Bur. Gov. Lab. (Manila) 27: 19. 1905. Basion¬
ym: Aquilaria pentandra Blanco, FI. Filip.
373. 1837.

4a, Pittosporum pentandrum var. formosan-
um (Hayata) Zhi Y. Zhang & Turland, comb,
nov. Basionym: Pittosporum formosanum llay-
ata, in Matsumura & Hayata, .). Coll. Sei. Imp.
Univ. Tokyo 22: 32. 1906. TYPE: China. Tai¬
wan: “Kaisa et kiko,” 23 Nov. 1896, C. Owa-
tari s.n. (syntype, TI); “Takaw,” without date,
A. Henry 52 (syntype, TI); “Reigaryo,” 31 Feb.
1898. C. Owatari s.n. (syntype, TI).

Pittosporum formosanum Hayata var. hainanen.se Gagne-
pain. in Lecomte, FI. Indo-Chine 1: 238. 1909. Pit¬
tosporum pentandrum var. hainanense (Gagnepain)
H. I.. Li, J. Wash. Acad. Sci. 43: 45. 1953. TYPE:
Vietnam: “rapides de Bac-ninh, Qiiang-yen.” without
date, Balansa s.n. (syntype, P not seen); “vers Ninli-
binh,” without date. Bon s.n. (syntype, P not seen).

W hen Pittosporum formosanum var. formosanum
and variety hainanense are considered to be syno¬
nyms (the view adopted by the present authors), the
correct name for this taxon at varietal rank under
P. pentandrum is variety formosanum. Chang and
Yan (1979: 35) adopted the same taxonomy, but
were incorrect to use the combination P. pentan¬
drum var. hainanense because the autonym, P. for¬
mosanum var. formosanum, was established simul¬
taneously with variety hainanense and has priority
under Article 1 1.6 of the Saint Louis Code (Grenier
el al., 2000). Hatusima (in Hatusima & Amano,
1994: 74) published the combination P. pentan¬
drum var. formosanum, but invalidly because no
basionym was cited (Art. 33.3).

Pittosporum pentandrum var. formosanum occurs
in southern Guangxi (Hepu Xian), Hainan, and
southern Taiwan (including Fan Yu), as well as in
Vietnam.

5. Pittosporum podocarpum Gagnepain, Notul.

Syst. (Paris) 8: 211. 1939.

5a. Pittosporum podocarpum var. hejiaug-
ense (II. Y. Su) Zhi Y. Zhang & Turland,
comb, et stat. nov. Basionym: Pittosporum he-
jiangense H. Y. Su, Bull. Bot. Res., Harbin
4(4): 201. 1984. TYPE: China. Sichuan: lle-
jiang Xian. Fubao-qu, Qilong Commune, 75 m,
2 Aug. 1977. Hejiang Expedition 0777 (holo¬
type, YBDC not seen).

From its original description, Pittosporum hejiang-
ense is sufficiently similar morphologically to P. po¬
docarpum to be conspecific. It differs in having

154

No von

sparsely puberulent young branches, vs. glabrous
branches in both P. podocarpum var. podocarpum
and P. podocarpum var. angustatum Gowda, and
densely brown pubescent branches in P. podocar¬
pum var. molle W. D. Han. Therefore, treating P.
hejiangense as a variety of P. podocarpum is appro¬
priate. Pittosporum podocarpum var. podocarpum
occurs in Fujian, Gansu, Guangxi, Guizhou, Hubei.
Hunan, Shaanxi, Sichuan, southeastern Xizang, and
Yunnan, as well as in northeastern India, Myanmar,
and northern Vietnam, while variety angustatum
has a very similar range, occurring also in Guang¬
dong, but as yet unknown in either Xizang or Vi¬
etnam. Pittosporum podocarpum var. i nolle is known
only from southern Fujian (Hua’an Xian) and south¬
ern Guizhou (Libo Xian), while variety hejiangense
is known only from southeastern Sichuan (Hejiang
Xian).

6. Pittosporum rchderiaimm Gowda, J. Arnold
Arbor. 32: 297. 1951.

6a. Pittosporum rehderiaiiuni var. ternstroe-
mioides (C. Y. Wu) Zhi Y. Zhang & Turland.
comb, et stat. nov. Basionym: Pittosporum tern-
stroemioides G. Y. Wu, FI. Yunnan. 3: 320.
1983. TYPE: China. Yunnan: Fongling Xian,
foot of mountain, among shrubs, 10 Aug. 1941
(fr), C. IT Wang 90010 (holotype, KUN not
seen).

From its original description, Pittosporum tern-
stroemioides is sufficiently similar morphologically
to P. rehderianum to be conspecific. It differs in

having capsules each with ca. 8 seeds, vs. capsules
each with 21-24 seeds in P. rehderianum. There¬
fore, treating P. ternstroemioides as a variety of P.
rehderianum is appropriate. Pittosporum rehderian¬
um var. rehderianum occurs in southeastern Gansu,
western Hubei, southern Shaanxi, and western
Sichuan, whereas variety ternstroemioides is dis¬
junct, known only from western Yunnan (Fongling
Xian).

Acknowledgments. We thank Anthony IF Bracli
(Harvard University Herbaria) and Akiko Shimizu
(University of Tokyo) for checking type specimens
at their respective institutions. Vie also thank Peng
Hua (Kunming Institute of Botany) for advice and
William J. Hess (The Morton Arboretum) for re¬
viewing the manuscript. This project has been sup¬
ported by the National Natural Science Foundation
of China (NSFC-39970055) and the China Schol¬
arship Council (98491005).

Fiterature Cited

Chang, II. T. X S. Z. Yan. 1979. Pittosporaceae. Pp. I —
36 in: II. T. Chang (editor), FI. Reipubl. Popularis Sin.
35(2). Academia Sinica, Beijing.

Greuter, VC. .1. McNeill. F. IF Barrie, H. M. Bordet. V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
.1. K. Skog. P. Trehane, N. J. Turland X I). F. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Ilalusima, S. X T. Amano. 1994. Flora ol the Ryukyus,
South of Amatni Island, ed. 2. Biological Society of Oki¬
nawa.

Zhang, Z. Y., IF I). Zhang |H. T. Chang] X N. J. Turland.
In press. Pittosporaceae. In: Z. Y. Wu X P. IF Raven
(editors), Flora of China, 9. Science Press. Beijing X
Missouri Botanical Garden Press, St. Louis.

Volume 12, Number I, pp. 1—154 of NOVON was published on 2 April 2002.

Volume 12 NO VON
Number 2
2002

A New Species of Thamnobryum (Musci: Neckeraceae) from
Venezuela, with a Key to the Neotropical Species of Thamnobryum

Bruce Allen

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

bruce.allen@mobot.org

Steven P. Churchill

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

steve. church i I l@mobot.org

Abstract. Thamnobryum liesneri differs from all
other species of Thamnobryum in its ligulate to Un¬
gulate secondary stem and branch leaves that have
exceptionally broad (to 160 gm wide and a third
or more of the leaf base width), strongly forked and
spurred, subpercurrent costae. It differs from all
species of Porotrichum and Porothamnium in hav¬
ing leaves with very' broad costae and short upper
cells. A key to the three species of Thamnobryum
in the Neotropics is given.

Key words: Musci. Neckeraceae. Thamnob¬
ryum, Venezuela.

Thamnobryum is a handsome genus of frondose
or stipitate mosses with 42 species (Crosby et al..
2000) distributed throughout the world. The genus
is usually included in the subfamily Thamnioideae
of the Neckeraceae (Fleischer, 1905-1906; Broth-
erus, 1906, 1925; Robinson, 1975; Crum & An¬
derson. 1981; Vitt, 1984; Enroth, 1989. 1994;
Buck, 1998; Buck & Goffinet, 2000). Monkemeyer
(1927), however, placed the genus (as Thamnium,
an illegitimate name) in its own family. This family
(now the Thamnobryaceae) has been expanded by
Margadant and During (1982), Walther (1983),
Buck and Vitt (1986), Sastre-De J esus (1987), and
Churchill and Linares (1995) to include all mem¬
bers of the Neckeraceae with a frondose or stipitate
habit and striate exostomes. As discussed by Buck

(1998). however, some thamnoid genera are weakly
stipitate. and both striate and papillose exostomes
can be found in the thamnoid as well as the neck-
eroid genera.

I he taxonomy of the Thamnioideae is so unset¬
tled that it is difficult to find fault with Robinson’s
(1975: 56) characterization of them as a “. . . vile
group of genera. ... I his is because the important
character states within the group are not discrete,
anil within this mainly dioicous group that often
lacks sporophytes many of its critical characters are
sporophytic. In the Neotropics Thamnobryum is
close to Porotrichum and Porothamnium. It differs
from both genera mainly in having leaves with
stronger costae and shorter upper cells (see Rob¬
inson. 1975; Sastre-De Jesus, 1987; Buck. 1998).

I here are three species of Thamnobryum in neo¬
tropical South America, one of which is described
below as new to science.

Thamnobryum liesneri B. H. Allen & S. P. Chur¬
chill. sp. nov. TV PE: Venezuela. Amazonas:
Atures, lower forested E slope of unnamed
1760 m peak. 8 km NW of settlement of Yu-
taje, 3 km W of Rio Coro-Coro, W of Serranfa
de Yutaje, 05°41'N, 66°09'W. Ronald Liesner
& Bruce Holst 21891 (holotype, MO; isotypes,
H. NY, US, VEN). Figure 1A-L.

Species haec a congeneris foliis ligulatis-lingulatis, cos-

Novon 12: 155-158. 2002.

156

No von

Eigure I. Thamnobryum liesneri 1$. II. Allen & S. I’. Churchill. —A. Upper secondary stem leaf apex. — B. Koliose
pseudoparaphyllia. — C, 1). Upper secondary stem leaves. — E. Cross section of secondary stem. — E. Upper secondary

Volume 12, Number 2
2002

Allen & Churchill

Thamnobryum liesneri from Venezuela

157

lis latissimis, caloaratis et furc-atis, subpercurrentibusque
differt.

Plants slender, dull, light-green, older plants
black-brown, in dense, wiry tufts to 4 cm high. Pri¬
mary stems yellow-red, becoming red-black with
age, creeping, in cross section with central strand
poorly developed, cortical cells thin-walled, and
epidermal cells very thick-walled, to 6 cells thick;
rhizoids in circular clusters on the parts of the
stems that touch die substrate, red, smooth. Leaves
widely spaced, reduced, hastate, to 0.5 mm long,
costa single, strong. Secondary stems yellow-red,
becoming red-black with age, erect, stipitate below,
weakly frondose above, irregularly 1—2 pinnately
branched, in cross section central strand weakly
developed, cortical cells firm-walled, hyaline, epi¬
dermal cells small, thick-walled, golden-yellow to
red, in 6—8 rows; foliose pseudoparaphyllia present;
axillary hairs 6—7 cells long, lower 3^4 quadrate,
upper 2—3 cylindrical, all cells often light red-
brown; rhizoids densely clustered at base of sec¬
ondary stems, red, smooth; secondary stems and
branches ending in flagellate tips. Sti[ >e leaves
erect to erect-spreading when dry, erect-spreading
when wet, widely spaced, reduced, stipe leaves
near base hastate, becoming linear-lanceolate
above, 1.0— 1.8 mm long. Leaves erect to erect-in¬
curved when dry, more spreading when wet. oth¬
erwise little changed, well-spaced, ligulate to Un¬
gulate, 1.7— 2.0 mm long, 0.3— 0.4 mm wide, flat to
variously concave; apices acute; margins plane to
reflexed below, entire in lower V2 to %, irregularly
toothed to crenulate at apex; lamina unistratose
above, uni- or bistratose at base; costae single, very
broad (to 160 /xm wide at base), subpercurrent
(ending 10-20 cells below the apex), often forked
at apex, usually spurred below; leaf cells firm-
walled, smooth, not porose, apical cells irregularly
rhomboid to hexagonal 12—20 X 6—10 /xm, upper
median cells fusiform-rhomboid, usually obliquely
arranged from the costa, basal cells oblong-rect¬
angular, 20—40 X 6—8 /xm, alar cells short, weakly
differentiated. Dioicous (?). Sporophytes unknown.

Etymology. This species is named for Ronald
Lee Liesner. an outstanding vascular plant and
bryophyte collector at the Missouri Botanical Gar¬
den. Ron Liesner was the actual collector of Tham¬
nobryum liesneri.

<—

Habitat. On boulder near stream; 1050-1200
m.

Thamnobryum liesneri differs from all species of
Thamnobryum familiar to us by its ligulate to lin-
gulate secondary stem and branch leaves that have
exceptionally broad (to 160 /xm wide and a third
or more of the leaf base width), strongly forked and
spurred, subpercurrent costae. There are some spe¬
cies of Thamnobryum from the Juan Fernandez Is¬
lands with costae to 100 /xm wide, but these spe¬
cies have ovate-lanceolate or lanceolate leaves w ith
percurrent or excurrent costae. Although the pres¬
ence of flagellate tips on nearly all the secondary
stems and branches of T. liesneri seems a distinc¬
tive feature of the species, there are many other
species of Thamnobryum that occasionally have fla¬
gellate stem and branch tips. Thamnobryum liesneri
differs from all species of Porotrichum and Poro-
thamniurn in having leaves with very broad costae
and short upper cells.

Key to the Neotropical Species of Thamnobryum

I. Secondary stem and branch leaves ligulate to 1 in¬
sulate; costae to 160 /xm wide at base T. liesneri B.

H. Allen & S. P. Churchill
1 . Secondary7 stem and branch leaves ovate-ligulate
or oblong-ovate; costae 40—60 /xm wide at base

. 2

2. Secondary stem and branch leaves plicate
when dry, ovate-ligulate, 2-4 mm long; stipe

leaves 2—3 mm long .

. T. fasciculatum (Hedwig) I. Sastre

2. Secondary stem and branch leaves smooth
when dry, oblong-ovate, 2-3 mm long; stipe

leaves to 1 mm long .

. ... T. tumidicaule (K. A. Wagner) F. 1). Bowers

Acknowledgments. This work was supported by
National Science Foundation Grant DEB-0072433
(to the Missouri Botanical Garden). We thank Jo¬
hannes Enroth (H) for his evaluations on this new
species.

Literature Cited

Brotherus, V. F. 1906. Neckeraceae. In: A. Engler & K.

Prantl (editors), Nat. Pflanzenfam. 1(3): 775—816.
- . 1925. Musci. In: A. Engler (editor), Nat. Pflan¬
zenfam. ed. 2, 11.

Buck, W. R. 1998. Pleurocarpous Mosses of the West In¬
dies. Mem. New York Bot. Card. 82.

- & B. Goffinet. 2000. Morphology and classifica¬
tion of mosses. Pp. 71—123 in A. J. Shaw & B. Goffinet

stem leaf median cells and margin. — G. Cross section, upper part of secondary stem leaf. — H. Cross section, lower
part of secondary stem leaf. — I. Upper secondary stem leaf basal cells and margin. — J. Axillary hair. — K. Stipe leaf.
— L. Primary stem leaf. Scale in mm: bar = 0.05 (B. J); bar = 0.06 (A, E-I); bar = 0.34 (K. L); bar = 0.4 (C, I)).
All figures from Liesner & Holst 21891 (holotype).

158

Novon

(editors), Bryophyte Biology. Cambridge Univ. Press,
Cambridge.

- & 1). H. Vitt. 1986. Suggestions lor a new familial

classification of pleurocarpous mosses. Taxon 35: 21 —
60.

Churchill, S. P. & E. Linares C. 1995. Prodromus Bryol-
ogiae Novo-Granatensis. Bibliot. Jose Jeronimo Triana
12.

Crosby, M. R„ R. K. Magill. B. Allen & S. He. 2000. A
Checklist of the Mosses. Missouri Botanical Garden, St.
Louis. | Distributed by the authors.]

Crum, H. A. & L. E. Anderson. 1981. Mosses of Eastern
North America. Columbia Univ. Press, New York.

Enroth, J. 1989. Bryophyte flora of the Huon Peninsula.
Papua New Guinea. XXVII. Neckeraceae (Musei). Acta
Bot. Fenn. 137: 41—80.

- . 1994. On the evolution and circumscription of

the Neckeraceae (Musei). J. Hattori Bot. Lab. 76: 13-

20.

Fleischer, M. 1905—1906. Neue Familien, Gattungen und
Arten der Laubinoose. I. Teil. Hedwigia 45: 53—64
[1905], 65-87 [1906].

Margadant, \\. I). & H. During. 1982. Beknopte flora van
Nederlandse blad- en levermossen. W. J. Thieme, Zut-
phen.

Monkcmeyer, W. 1927. Die Laubmosse Europas. In: L.
Rabenhorst, Kryptogamen-Flora von Deutschlands, 0s-
terreichs und der Schweiz. Vol 4(1). Leipzig.

Robinson, H. 1975. The mosses of Juan Fernandez Is¬
lands. Smithsonian Contr. Bot. 27.

Sastre-De Jesus, I. 1987. A Revision of the Neckeraceae
Schimp. and the Thamnobryaeeae Marg. & Dur. in the
Neotropics. Ph.D. Dissertation, City University of New
York, New York.

Vitt, D. fl. 1984. Classification of the Bryopsida. Pp. 696—
759 in R. M. Schuster (editor). New Manual of Bryology.
Hattori Botanical Laboratory, Nichinan.

Walthcr. K. 1983. Syllabus der Pflanzenfamilien. Vol. 2.
Bryophytina, Laubmoose. G. Borntraeger, Berlin.

Overview of Sphaeralcea (Malvaceae) in Southern Utah and
Northern Arizona, U.S.A., and Description of a New Species

N. Duane Atwood and Stanley L. Welsh

S. L. Welsh Herbarium, Life Science Museum, Brigham Young University, Provo, Utah
84602, U.S.A. nda@email.byu.edu; slslwelsh@aol.com

ABSTRACT. Described is Sphaeralcea gierischii N.
D. Atwood & S. L. Welsh, sp. nov., which is distin¬
guished from S. rusbyi by the glabrous or glabres-
cent herbage, with few or no stellate hairs confined
to the leaf margin, larger (15-25 mm long) flowers,
and restricted range and habitat, and from S. moor-
ei in the 3- to 5-parted, narrow leaf lobes, bright
green leaves sometimes suffused with red-purple,
and different habitat. Two varieties are evaluated at
the species level within Sphaeralcea based on dif¬
ferent morphological features, geographical distri¬
bution, and habitat: S. moorei (S. L. Welsh) N. I).
Atwood & S. L. Welsh, stat. nov., and S. fumariensis
(S. L. Welsh & N. D. Atwood) N. D. Atwood & S.
L. Welsh, stat. nov. A key to the closely related taxa
is included, and each of the taxa is provided with
a description. The geographical distribution and
ecology in southern Utah and northern Arizona,
U.S.A. , are also discussed.

Key wards: Arizona, Malvaceae, North Ameri¬
ca, Sphaeralcea, Utah.

Collections taken over the past half century give
indication that southern Utah is an important center
for diversity and speciation in the genus Sphaer¬
alcea. The genus as a whole consists of taxa whose
morphological distinctions are obscured by overlap
of practically every characteristic. Floral features
tend to be very similar, and their usefulness as di¬
agnostic characters is clouded by intermediates
among the constituent taxa. Characteristics of the
mature fruiting carpels have been considered as
perhaps more important than other floral features.
However, mature carpels are seldom collected. As
soon as the main flush of flowering is finished most
collectors tend to lose interest. Thus only a small
fraction of the huge number of specimens collected
in this attractive genus bear mature carpels. Some
of the intermediacy of morphological features is the
result of hybridization among the various taxa, often
across sectional lines as defined by Kearney. Thus,
numerous specimens intermediate between the taxa
tend to accumulate in herbaria without proper iden¬
tification. The result of the problematic morphology

has been the definition of taxa by use of multiple,
overlapping characteristics. By such a procedure it
has been possible to recognize the main subunits
within what is more or less a mosaic of variation.
The following short summary is confined to those
species involved within sections Laxae and Mun-
roanae as defined by Kearney (1935). This includes
those species that are thought to have been most
important in contributing to the diversity of forms
present in southern Utah and adjacent Arizona.

The following key is presented to allow discrim¬
ination of most specimens of the related and similar
taxa known to occur in southern Utah and imme¬
diately adjacent northern Arizona.

la. Herbage only sparingly pubescent, the herbage
bright green (the stems sometimes suffused with
red-purple).

2a. Herbage glabrous or glabrescent, stellate
pubescence, if any, confined to leaf margins
(rarely a few stellate triehomes on the blade
surface); calyces glabrous externally; flowers
large, the petals 15-25 mm long; plants
known only from northern Mohave Co., Ar¬
izona, and immediately adjacent Washington
Co., Utah, on the Harrisburg Formation . .

. 3. 5. gierischii

2b. Herbage definitely, though sometimes
sparsely, stellate hairy overall (if sparsely so,
of different distribution); calyces stellate ex¬
ternally; flowers large to small, petals mostly
less than 20 mm long; plants of other dis¬
tribution or habitat.

3a. Leaves 3- to 5-parted or -divided, the
lobes with narrow, regularly pinnatifid
margins, the teeth nearly right angles to
the vein: carpels 4—6 mm high, often
with transparent lacunae, reticulate
mainly in the lower ca. 1/3; plants local
in Washington Co., Utah, and south to

southern Arizona . . 7. 5. rusbyi

3b. Leaves variously lobed, divided, or part¬
ed, the lobes with broader margins ir¬
regularly toothed or lobed, but not as
above; carpels 3-4.5 mm high, with
opaque or translucent lacunae, reticu¬
late in the lower 2/5— 3/5; plants of var¬
ious or broader distribution.

4a. Calyx pubescence almost or quite
as dense as that of the herbage,
mainly 4.5— 7.5 mm long (if longer,

Novon 12: 159-166. 2002.

160

Novon

not of Washington Co., Utah) at an-
lliesis; leaf blades with lobes fre¬
quently not cut to the base, often
wider than long, the terminal lobe
mostly less than twice as long as
broad; carpels with lacunae in the
lower 2/5- 1/2 ... 4. S. grossidariifolia
4b. Calyx pubescence more dense than
that of the herbage, mainly 6.8-9
mm long at anthesis; leaf blades
rather uniformly dissected to the
base, mostly longer than broad, the
terminal lobe mostly more than hall
as broad as long; carpels with la¬
cunae in the lower ca. 1/3 . . .

. 7. S. rusbyi

lb. Herbage moderately to densely pubescent, shad¬
ed yellow, white, or gray (the stems sometimes
suffused with red-purple).

5a. Inflorescence loosely thyrsoid (appearing pa¬
niculate), leafy; flowers not numerous at
each node; peduncles generally elongate;
calyx surpassing the fruit; carpels with re-
ticulae extending onto back of carpel; leaves
with conspicuous, elevated veins beneath,
the pubescence often turning yellow in age;
plants of Washington and Kane Cos., Utah,
and adjacent Nevada, Arizona, and Califor¬
nia; Mexico .

. I . S. ambigua

5b. Inflorescence contracted thyrsoid-glomerate,
not especially leafy; flowers often numerous
at each node; peduncles generally contract¬
ed: calyx often shorter than the fruit; carpels
with reticulae confined to lateral face ol car¬
pel: leaves conspicuously veined or not, the
pubescence seldom if ever turning yellow in
age; plants of various or other distribution.

6a. Leaves shallowly 3- to 5-lobed; carpels
with well-defined to nearly obscure re¬
ticulae on the lower 1/3 ol the carpel;
plants mainly of western Colorado, west
across Utah to Nevada and eastern Cal¬
ifornia. and south to northwestern New

Mexico and central Arizona .

. 6. .S'. jKirvifolia

6b. leaves 3- to 5-cleft, -parted, or -divid¬
ed. carpels with well-defined reticulae
on less than half of carpel face; plants
of various distribution.

7a. Plants copiously stellate-pubes¬
cent, the trichome branches not ra¬
diating in a single plane, appearing
gray, known only from Smoky
Mountain in eastern Kane Co.,

Utah . 2. .S’, fumariensis

71). Plants not copiously pubescent or
appearing especially gray, the tri¬
chome branches mostly radiating in
a single plane, of other distribution.

8a. Herbage bright green; leaves
thin-textured, often narrowly
Inbed; plants of Glen Canyon
of the Colorado River and its
tributaries, in eastern Kane
anil Garfield, and western San
Juan Cos., Utah ... 5. .S', gioorei

fib. Herbage gray-green to whitish
canescent; leaves often thick-
textured, seldom narrowly
lobed; plants widespread in the
western United States ....
. 4. S. grossulariifolia

1. Sphaeralcea ambigua A. Gray, Proe. Anier.

Acad. 22: 292. 1887.

Stems arising from a woody caudex, several to
numerous, 3—10 dm tall, shaded white to ye llow
canescent, the triehomes becoming yellow in age;
leaf blades 1-6 cm long (from sinus to apex), 0.8-
5 cm wide, thickened, usually rugose, with veins
prominent beneath, ovate, debate, or nearly orbic¬
ular. the base cordate to deeply cordate, obscurely
to definitely 3- to 5-lobed, the lobes crenate; inflo¬
rescence commonly open glomerate-paniculate, but
sometimes narrowly thyrsoid; pedicels usually
shorter than calyx; calyx 6-12(-20) mm long at an¬
thesis, usually uniformly and densely stellate-pu¬
bescent but sometimes glabrescent on the outer
surface, the lobes lanceolate to acuminate; petals
15-22(-35) mm long, orange (grenadine) to orange-
pink (fading pink); carpels 3.5—6 mm high, 2—4 mm
wide, the reticulate portion comprising ca. 1/3 of
the carpel; n = 5.

Habitat and distribution. Creosote bush, Josh¬
ua tree, ambrosia, blackbrush, mixed warm desert
shrub, and pinyon-juniper communities at 700 to
1750 m in western Kane and Washington Counties,
Utah; Nevada, Arizona, and California; Mexico.

Our materials belong to Sphueralcea ambigua
var. ambigua. The common name slippery elm has
been applied due to the mucilaginous condition
that allows the bark to peel easily from the stem.
The plants are components of forage for both live¬
stock and desert tortoise.

2. Spliaeraleea fumariensis (S. L. Welsh & N.

D. Atwood) N. I). Atwood & S. L. Welsh, stat.
nov. Basionym: Spliaeraleea grossulariifolia
(Hooker & Arnott) Rydberg var. fumariensis S.
I.. Welsh & N. I). Atwood, Rhodora: 103: 83.
2001. TYPE: U.S.A. Utah: Kane Co., W side
of Little Valley, N of Lake Powell, 6 May 1998.
S. L. Welsh & N. I). Atwood 26968 (holotype.
BRY).

Stems few to many from a woody caudex, 2.3—
5.5 dm tall or more, the stems typically dark red-
purple at least near the base, the herbage gray-
green and abundantly stellate pubescent, the
triehomes with longest ray 0.8— 1.2 mm long, the
rays radiating in more than one plane; leaf blades

Volume 12, Number 2
2002

Atwood & Welsh
Sphaeralcea in Utah and Arizona

161

1.2— 3.7 cm long, 2—7.8 cm wide, often wider than
long, cordate-ovate or cordate in outline, the base
cordate to truncate or obtuse, digitately 3- to 5-
lobed, the main lobe 1—3.5 cm long and 0.8— 2.5
cm wide, entire or cleft or parted to irregularly
toothed; inflorescence rather compactly thyrsoid,
with usually more than 1 (lower per node, or rather
compactly glomerate-paniculate with 2 to 5 flowers
on lateral peduncles; pedicels shorter than to longer
than the calyx; bracteoles linear, sometimes red-
purple, but not especially contrasting with the ca¬
lyx; calyx 7.5— 9. 5(— 12) mm long at anthesis, green,
becoming stramineous in fruit, uniformly and
densely stellate-hairy, the rays of hairs radiating in
more than a single plane, the lobes ovate to lance-
acuminate; petals 12-17 mm long, orange (grena¬
dine); carpels 10-14, 3.3— 4.1 mm high, 2. 1-2.8
mm wide, the reticulate portion forming ea. 1/2 of
the carpel, the lacunae ± opaque.

Habitat and distribution. Match weed, ephedra,
blackbrush, galleta, shadscale, and juniper com¬
munities, at 1340 to 1650 m, endemic on the
Straight Cliffs, Tropic Shale, and Dakota Forma¬
tions, and on alluvium derived from those forma¬
tions, on Smoky Mountain and vicinity, eastern
Kane County, Utah.

The copiously stellate-hirsute pubescence gives
the plants of Sphaeralcea fumariensis a gray-green
appearance when dry, and a truly gray appearance
in the field. Many of the trichomes are caducous,
with abundant residue of stellate trichomes accu¬
mulating in the pressing paper. It is possibly most
closely allied to S. moorei, which occurs nearby in
sandy soils along Glen Canyon in Kane County and
east through Garfield and San Juan Counties, but
differs from that entity not only in the harsh pu¬
bescence but in the larger calyces. These features,
along with the restricted habitat, markedly separate
it from the green, thin-leaved, and narrowly lobed
plants of S. moorei.

Representative specimens examined. U.S.A. Utah: Kane
Co., head of Smoky Hollow dugway, T41S, R4E, S8, 17
June 1998, /V. I). Atwood & S. I,. Welsh 23859 (BUY);
Tibbett Spring, T41S, R3E, S32, 20 May 1999, N. I).
Atwood et al. 24598 (BRY); oa. 7 mi. W of Smoky Moun¬
tain. head of Creeks road, T40S, R3E, S36, 20 May 1999,
N. I). Atwood et al. 24614 (BRY); ea. 7 mi. W of Kelly
Grade, Croton Canyon Junction, I43S, R3E, S14, 20 May
1999. N. I). Atwood et al. 24627 (BRY); Drip Tank Can¬
yon. E of Four Mile Bench. T40S, R3E, S24, 6 May 1998,
.S'. L. Welsh & A. I). Atwood 26938 (BRY); Reeses Canyon,
ea. 2 mi. N of confluence of Last Chance Canyon/Reeses
Canyon, T40S, R4E, S36, 21 May 1999, A. D. Atwood et
al. 24638 (BRY).

3. Sphaeralcea gierischii N. 1). Atwood & S. U.
Welsh, sp. nov. TYPE: U.S.A. Arizona: Mohave
Co., ca. 0.2 mi. N of Black Rock Gulch, T40N.
R13W. S3, 24 Apr. 2000, N. I). Atwood. H.
Furniss & L. C. Higgins 25293 (holotype,
BRY; isotypes, ARIZ, ASU, BRY, GH, MO.
NY, RM, US). Figure 1.

Similis S. moorei in habitu sed in floribus majoribus
pubescentiis paucioribus vel nullis in laniinis et calyces
externis et carpelis majoribus discrepantibus, et e S. rus-
byi in floribus (petalisque et calycibus) majoribus calycis
glabris externis divergentihus.

Stems few to many from a woody caudex, 4.3-
10.3 dm tall, the stems often dark red-purple, the
herbage bright green and subglabrous to almost or
quite glabrous; leaf blades 1.2—4 cm long, 1—5 cm
wide, usually longer than wide, ovate to cordate-
ovate in outline, the base cordate to truncate or
obtuse, 3- to 5-lobed, the main division entire or
cleft or parted to irregularly toothed; inflorescence
thyrsoid, with usually more than I flower per node,
or glomerate-paniculate with 2 to 5 flowers on ax¬
illary peduncles; pedicels shorter than to much lon¬
ger than the calyx (to 7 cm long); bracteoles linear,
often red-purple and contrasting with the calyx; ca¬
lyx 5—10 mm long, green, becoming stramineous in
fruit, uniformly glabrous externally (or rarely with
one or few stellate hai rs), the rays of hairs mainly
radiating in a single plane, the lobes ovate to lance-
acuminate; petals 15—25 mm long, orange (grena¬
dine); carpels 10—15, 4.5— 5.5 mm high, the retic¬
ulate portion forming from 2/5 to 3/5 of the lower
portion of the carpel, reticulate on the sides.

Habitat and distribution. Warm desert shrub
community, mainly on gypsiferous outcrops of the
Harrisburg Member of the Kail tab Formation, at ca.
1090 m, in northern Mohave County, Arizona (near
Black Rock), and in closely adjacent Washington
County, Utah.

Pubescence is a variable feature throughout the
genus, but seldom is it so lacking as in Sphaeralcea
gierischii, wherein only the inner portion of the ca¬
lyx and the carpels are routinely copiously stellate.
The stems are produced in tall, open clumps, and
are only sparingly leafy. Main loliage leaves in the
lower portion of the stems are large, with the central
lobe greatly elongated and having a long-cuneate
base. Pubescence on the herbage is very sparse,
and on the calyces consists ol one or a few tri¬
chomes when any hairs are present at all. The open
inflorescence is shared with S. rusbyi, and in both
cases the feature might be derived in some part or
entirely from the contiguous S. arnbigua A. Gray,
which frequently has large flowers (15—22 mm

162

Novon

B.

1CM

Kigure 1. Sphaeralcea gierischii N. I). Atwood & S. L Welsh. — A. Habit. — B. Carpel (longitudinal view). — C. Calyx
and schizocarp. Drawn from the holotype collection, N. D. Atwood. B. Furniss & /,. C. Higgins 25293 (BUY).

Volume 12, Number 2
2002

Atwood & Welsh

Sphaeralcea in Utah and Arizona

163

long), but differs vastly in its dense white to yellow
canescent, thick, usually rugose, prominent veined,
deltoid to nearly orbicular cordate-based leaves,
short pedicels, and larger prominent reticulate car¬
pels (12-16 mm high).

Paratypes. U.S.A. Arizona: Mohave Co., Black Knolls.
T41N, R12W, S7, 26 May 1978, R. K. Gierisch 4355 (BRY);
Black Knolls. T41N, R12W, S7, 23 Apr. 2000. N. I). At¬
wood & H. Fumiss 25233 (BRY); Black Knolls, T40N,
R 13W, S3, 10 May 2000, l\. I). Atwood 25543 (BRY); 1.2
mi. N of Black Rock Gulch, 24 Apr. 2(XK). N. D. Atwood
25287 (BRY); S of Black Knolls, 6 June 1980, C. E. Handy
119 (BRY). Utah: Washington Co., T4.3S, R16W, S28, 3/4
mi. S of Little Round Valley, ca. 2 mi. S of Atkinville, 29
Apr. 1985, K. Thorne & N. I). Atwood 3749 (BRY).

The species is named in memory of the late
Ralph K. Gierisch, ardent collector of plants in the
Intermountain West for more than five decades, who
spent several years in the latter part of the century
(1970s and early 1980s) investigating the flora of
the Mohave Strip, Mohave County, Arizona. He
made the first collection of the species from the
Black Knolls in 1978.

4. Sphaeralcea grossulariifolia (Hooker & Ar-
nott) Rydberg, Bull. Torrey Bot. Club 40: 58.
1913. Sida grossulariifolia Hooker & Arnott,
Bot. Beechey Voy. 326. 1840. Malvastrum gros-
sulariifolium (Hooker & Arnott) A. Gray, Mem.
Amer. Acad. II. 4: 21. 1849. Sphaeralcea pe¬
dal a Torrey, in A. ( fray, Mem. Amer. Acad. II,
4: 23. 1849. Sphaeralcea grossulariifolia subsp.
pedata (Torrey) Kearney, Univ. Calif. Publ. Bot.
19: 88. 1935; protologue locality: “California.
Fremont.” TYPE: “Fremont's Expedition to Cal¬
ifornia 41 1. 1845. Sphaeralcea new (S. Pedata
call it). Moving Fork, 1st Camp. Utah” (holo-
type, NY-Torrey). [Moving Fork is probably in
western Colorado (Welsh. 1998: 368).]

Stems few to many from a woody caudex, mostly
3.5— 7.5 dm tall or more, the herbage shaded white
or gray to yellow canescent to rather sparsely stel¬
late-hairy and green; leaf blades 1.2-5 cm long, 1 .3—
5.2 cm wide, from longer than wide to wider than
long, cuneate-ovate to ovate or cordate-ovate in out¬
line, the base cordate to truncate or obtuse, 3- to 5-
lobed, the lobes distinct to the base or confluent to
well above the base, the main lobe again few-lobed
or less commonly parted or irregularly toothed; in¬
florescence compact-thyrsoid, with usually more
than 1 flower per node; pedicels shorter than to lon¬
ger than the calyx; calyx 4.5—9 mm long, uniformly
stellate or rarely the calyx lobes glabrate, the rays
of hairs not or less commonly radiating in a single
plane, the longest rays mainly less than 0.5 mm long

except on some calyx bases, the calyx lobes ovate to
lance-acuminate; petals 8-18 mm long, orange or
rarely rose pink; caqiels 10—14, 2.9— 3.7 mm high,
reticulate portion forming from 2/5 to 3/5 of the car¬
pel, the lacunae opaque or nearly so.

Habitat and distribution. Salt, warm, and cool
desert shrub, piny on-juniper, and ponderosa pine
communities at ca. 790 to 2290 m in Beaver, Box
Elder. Carbon, Emery, Garfield, Grand, Juab, Iron,
Kane, Millard. Piute, Salt Lake, San Juan, Sanpete,
Tooele, Uintah, Utah, Washington, and Wayne
Counties, Utah; Washington, Oregon, California,
Nevada, and Arizona.

This is the most common and w idely distributed
species in Utah except for the only distantly related
Sphaeralcea coccinea (Nuttall) Rydberg, which is
widespread, occurring from Texas to the Western
U.S., north to Canada. However, Sphaeralcea gros-
sulariifolia forms intermediates with .5'. coccinea, S.
parvifolia, ami with the more northerly distributed
S. munroana (Douglas) Spach ex A. Gray, which
occurs in northern Utah northward. The report by
Kearney ( 1 935) of -S', digitata (Greene) Rydberg ap¬
parently belongs here with Sphaeralcea grossulari¬
ifolia. Leaf variation is great, but the blade dissec¬
tion is less complex than in the similar and
undoubtedly closely allied S. moorei, S. fumarien-
sis, S. rusbyi, and S. gierischii. The Colorado Can¬
yons and Kaiparowits endemics, S. moorei and S.
fumariensis, share with S. grossulariifolia the fea¬
ture of the reticulate portion of the carpel covering
2/5 to 3/5 of the surface.

There is much variation within the widely dis¬
tributed S. grossulariifolia, with herbage varying in
aspect from gray to gray-green or brownish, to yel¬
low-green, to green. Leaf blades tend not to be cut
completely to the base as is often the case in the
related S. moorei, S. fumariensis, S. rusbyi, and S.
gierischii. Most specimens range in size from 3.5 to
5.5 dm in height, while all of the other species,
except S. fumariensis, commonly tend to exceed
that height. It seems probable that most Utah spec¬
imens from Washington County, previously identi¬
fied as S. grossulariifolia, are better placed with the
taller .S’, rusbyi, which tends to have green or green¬
er herbage, more uniform and more uniformly deep¬
ly cut leaf blades, and proportionally less of the
carpel base occupied by reticulae.

5. Sphaeralcea moorei (S. L. Welsh) N. D. At¬
wood & S. L. Welsh, stat. nov. Basionym:
Sphaeralcea grossuliariifolia var. moorei S. L.
Welsh, Great Basin Naturalist 40: 35. 1980.
TYPE: U.S.A. Utah: Kane Co., E side of Last
Chance Bay, Lake Powell, Entrada sandstone,
2 May 1972. S. L. Welsh & N. D. Atwood
11597 (holotype, BRY). Figure 2.

164

Novon

/:•

Figure 2. Sphneralcea moorei (S. L. Welsh) IN. I). Atwood A S. I,. Welsh. — A. Habit. — It. Carpel, closed carpel (left)
and open carpel (right). — C. Calyx and schizoearp. Drawn from the holotype collection, S. L. Welsh & N. I). Atwood
11597 (ItllV).

Volume 12, Number 2
2002

Atwood & Welsh

Sphaeralcea in Utah and Arizona

165

Stems few to many from a woody caudex, 3-7.5
dm tall or more, the stems typically dark red-purple
at least near the base, the herbage green and rather
sparsely stellate pubescent, the trichomes 0.3-0.8
mm wide, the rays radiating in a horizontal plane;
leaf blades 0.9— 3. 5(— 7.4) cm long, 1.0— 2.6 (15, a
rare exception) cm wide, often longer than wide,
ovate to cordate-ovate or cordate in outline, the base
cordate to truncate or obtuse, digitately 3- to 5-
lobed, the main lobe 0.8— 3. 5(— 7.4) cm long and 0.8—
1.5 cm wide, entire or cleft or parted to irregularly
toothed, or rarely the blade confluent and merely
palmately lobed; inflorescence rather compactly
thyrsoid, with usually more than 1 flower per node,
or rather compactly glomerate-paniculate with 2 to
5 flowers on axillary peduncles; pedicels shorter
than to much longer than the calyx; bracteoles linear,
sometimes red-purple, but not especially contrasting
with the calyx; calyx 4.5— 7. 2(— 8.5) mm long at an-
thesis, green, becoming stramineous in fruit, uni¬
formly stellate-hairy or less commonly glabrous ex¬
ternally only near the apex, the rays of hairs mainly
radiating in a single plane, the lobes ovate to lance-
acuminate; petals 11-15 mm long, orange (grena¬
dine); carpels 10-14, 4.5-5. 1 mm high, 2. 1-2.3 mm
wide, the reticulate portion forming ca. 1/2 of the
carpel, the lacunae opaque.

Habitat and distribution. Blackbrush, ephedra,
matchweed, Vanclevea, yucca, pinyon-juniper, and
sagebrush communities, on alluvium and colluvi¬
um, and on the Navajo, Moenkopi, and Cutler, and
undoubtedly other. Formations, at 850 to 1830 m,
along the Colorado River in Glen Canyon and its
tributaries in eastern Garfield and Kane Counties,
and in southern San Juan County, Utah, along the
San Juan River; endemic.

This is mainly a low-elevation species of sandy
tracts along the Fake Powell portion of the Colorado
and San Juan Rivers on the Entrada Sandstone For¬
mation. Sphaeralcea moorei does extend upward
somewhat along the east escarpment of Fifty Mile
Mountain, west of Glen Canyon, and evidently
reaches its northern limit at Calf Creek, east of Es¬
calante. Some plants with uncut leaves are includ¬
ed here also, especially some from the Three Gar¬
den Reentry along Lake Powell. The species tends
to intergrade at least morphologically with S. gros-
sulariifolia, but not more so than for laxa as a whole
in this genus, and it is reasonable to accept it at
specific rank.

Representative specimens examined. U.S.A. Utah:
Kane (io„ mouth of Kscalante River, Lake Powell, 5 June
1972, S. L. Welsh & G. Moore 11810 (BRY); Willow Tank.
4 May 1962, I). A. White 111 (BRY); Escalante Canyon,

5 June 1972, S. L Welsh & G. Moore 11827 (BRY); Es¬
calante Canyon, 24 Aug. 1971, N. I). Atwood & R. Allen
3211 (BRY); Hole-in-the-Rock, 14 May 1953, B. F. Har¬
rison 12112 (BRY); San Juan Co., 1 mi. E of Hole-in-the-
Rock, 16 June 1972, S. L. Welsh & C. A. Toft 11869
(BRY); Three Garden, Lake Powell, ca. 1 mi. N of conflu¬
ence with San Juan River Ann, 5 May 1974, S. L. Welsh
12420 (BRY); Comb Wash, 6 June 1970, S. L Welsh &
N. II Atwood 9972 (BRY).

6. Sphaeralcea parvifolia A. Nelson, Proc. Biol.
Soc. Wash. 17: 94-95. 1904. TYPE: U.S.A.
Nevada: Lincoln Co., Caliente, 22 May 1902,
Goodding 916 (RM not seen).

Sphaeralcea marginata York ex Rydberg, Bull. Torrey Bol.
Club 33: 145-146. 1906. TYPE: U.S.A. Colorado:
Grand Junction, 1901, C. F. Baker 93 (NY not seen).
Sphaeralcea arizonica Heller ex Rydberg, Bull. Torrey But.
Club 40: 59. 1913. TYPE: U.S.A. Arizona: Flagstaff
(Coconino Co.), 16 June 1898, McDougal 120 (ho-
lotype, NY not seen).

Stems few to many from branching woody cau¬
dex; leaf blades 1.5-5. 5 cm long, 1.2-5.2 cm wide,
ovate to orbicular, reniform, or cordate-ovate, the
base cordate to truncate or obtuse, usually shallow¬
ly 3- to 5-lobed, the sinuses usually shallow, the
lobes crenate-dentate; inflorescence commonly nar¬
rowly thyrsoid, typically with more than 1 flower
per node; pedicels usually shorter than the calyx;
calyx uniformly stellate, the rays of hairs not in a
single plane, the lobes lance-ovate to deltoid; petals
7—15 mm long, orange; carpels 10—12, 3-4 mm
high, the indehiscent part forming from 1/4 to 1/3
of the carpel, faintly reticulate on the sides.

Habitat and distribution. Blackbrush, other
warm desert shrub, salt desert shrub, sagebrush,
pinyon-juniper, and mountain brush communities
at 820 to 2700 m in Daggett, Duchesne, Emery,
Garfield, Grand, Iron, Juab, Kane, Millard, Piute,
San Juan, Sevier. Tooele, Uintah, Washington, and
Wayne Counties, Utah; Western Colorado, Nevada,
Arizona, New Mexico, and California.

Sphaeralcea parvifolia has been compared by
Kearney (1935) with S. ambigua, which it resem¬
bles in habit and leaf morphology. It is, however,
the more southern and lower elevation counterpart
of S. munroana, from which S. parvifolia cannot
always be distinguished, and with which it could
be included without serious damage to the taxon¬
omy of the genus. Otherwise, the relationships of
S. parvifolia in Utah seem to lie with the largely
sympatric S. grossulariifolia, but it is apparently
transitional with both ,S. ambigua and S. rusbyi. The
confluent leaf blades of S. parvifolia are diagnostic
for most specimens from those of .5’. grossulariifolia,
and the smaller leaf blades and less apparent ve-

166

Novon

nation allow segregation of most specimens from S.
ambigua , but in neither instance should blade size
and confluence be taken as tools for positive iden¬
tification. As in most taxa within the genus the mor¬
phological features blend where the taxa are in geo¬
graphical contact, and indeed many of the
diagnostic features are vegetative ones. However,
the characteristics of the fruiting carpels are not
absolute as diagnostic tools either.

7. Spliaeraleea rusbyi A. Gray, Proc. Amer.
Acad. 22: 293. 1887. TYPE: U.S.A. Arizona:
Yavapai Co., Prescott, May 1883, II. //. Rushy
537 (holotype. NY).

Stems few to many from a woody eaudex, 2.4 — fi.2
dm tall or more, the stems typically dark red-purple
at least near the base, the herbage green and rather
sparsely stellate-pubescent to rather densely so. the
trichomes 0.4— 0.8 mm wide, the rays radiating in
more than a single plane: leaf blades 1 .6—1.3 cm long.
0.6— 3.5 cm wide, usually longer than wide, ovate to
cordate-ovate in outline, the base cordate to truncate
or obtuse, digitatelv 3- to 5-lobed, the main 1. 3-3.8
cm long and 5-2.6 cm wide, entire or cleft or parted
to irregularly toothed, or rarely the blade confluent
and merely digilately lobed; inflorescence rather com¬
pactly thyrsoid, with usually more than 1 flower per
node, or compactly glomerate-paniculate with 2 to 5
flowers on axillary peduncles; pedicels shorter than to
much longer than the calyx; bracteoles linear, some¬
times red-purple, but not especially contrasting with
the calyx; calyx 6.5— 9.5 mm long at anthesis, green,
becoming stramineous in fruit, uniformly stellate-hairy
or glabrous externally only near the apex, the rays of
hairs mainly radiating in more than a single plane,
the lobes ovate to lance-acuminate; petals 12—19 mm
long, orange (grenadine); carpels 10-14, 4. .5-5.2 mm
high, 2.4— 2.8 mm wide. the reticulate portion forming
ca. 1/3 of the carpel, the lacunae opaque.

Habitat and distribution. Mixed desert shrub
and floodplain communities, with cottonwood, oak.
juniper, creosote bush, rattany, burrobush, black-
brush. ephedra, snakeweed, rabbitbrush, and four-
wing saltbush on Quaternary alluvium and collu¬
vium. Chinle, Moenkopi, and Kaibab and other
mostly calcareous formations, at ca. 1030 to 1220
m in Iron and Washington Counties, Utah, and Mo¬
have and other counties, Arizona.

In addition to the features cited in the key, the
herbage of S. rusbyi is ordinarily green or suffused
with dark purple and with rather sparsely stellate

hairs, much less so than the calyx; the inflorescence
tends to he open glomerate-paniculate, and the flow¬
ers average larger (to 18 mm long) than those of the
similar ,S. grossidariifolia (mainly less than 15 mm
long). The species is apparently most closely allied to
S. gierischii, however. Its smaller flowers (both calyces
and petals) and rather densely stellate calyces are
apparently diagnostic. Until recently S. rusbyi has
been a very obscure and poorly understood species
in Utah (Welsh et ah, 1993), with specimens exam¬
ined not at all uniform and many of them included
with the previous concept of S. grossidariifolia. The
main foliage leaves of S. rusbyi are rather uniformly
ovate-oval in outline, and are rather uniformly peda-
telv lobed. Stems of S. gierischii tend to he sparsely
leafy, with the blades more broadly ovate in outline,
and with the lobes much attenuated at the base. In
this manner, .S', gierischii differs from all other closely
allied taxa in southern Utah and northern Arizona,
including moorei, which it somewhat resembles in
the narrowly lobed leaves and stem color. The larger
set available now allows a better understanding of the
taxon within Utah, where it is confined to Washington
County, with the exception of one population overlap¬
ping the very southeast comer of Iron County.

Representative specimens examined. U.S.A. Arizona:
Mohave Co.. 38 mi. SE of Littlefield, T33N, R15W, 25
Apr. 1997, A. IK Atwood A- S. L. Welsh 21804 (HRY).
I tali: Iron Co., Camp Creek I mi. ESE of 1-55, T38S,
R12W, S10. 23 Sep. 1987, K. H. Thorne et al. 5601
(BRY); Washington Co., ea. I mi. NE of La Verkin, 30
Apr. 1993, .S'. /,. Welsh & /.. C. Higgins (BRY); 6 mi. E of
Hurricane, 27 May 1968, A. D. Atwood 1405 (BRY); Ko-
lob Terrace. T39V, R11W, S23, 7 July 1981, .S. L. Welsh
20718 (BRY).

Acknowledgments. The authors are indebted to
Noel 11. Holmgren and John La Duke for their most
helpful reviews of the manuscript and to BRY il¬
lustrator Shannie Workman for the excellent illus¬
trations. We thank Randy Baker in the Monte L.
Bean Museum for help with the graphics. Also spe¬
cial thanks are given to Victoria Hollowell (MO) for
her invaluable help during the editing process.

I .iterature Cited

Kearney, T. H. 1935. The North American species of

Sphaeralcea subgenus Eusphaeralcea. Univ. Calif. Publ.

Bot. 19(1): 1-128, pi. 1-12.

Welsh. S. I.. 1998. John Charles Fremont, Botanical Ex¬
plorer. Monogr. Syst. Bot. Missouri Bot. Card. 66.
- . N. I). Atwood, S. Goodrich & L. C. Higgins.

1993. A Utah Flora. Monte I.. Bean Life Science Mu¬
seum, Brigham Young University, Provo, Utah.

A New Species of Abronia (Nyctagi naceae) from San Juan County,

New Mexico, U.S.A.

N. Duane Atwood, Stanley L. Welsh
Herbarium, Life Science Museum, Brigham Young University,

Pi ovo, Ltah 84602-0200, U.S.A.

Kenneth D. Heil

Herbarium, San Juan College, Farmington, New Mexico 87402, U.S.A.

ABSTRACT. A new species of Abronia is described
from northwestern New Mexico, U.S.A., as Abronia
bolackii N. D. Atwood, S. L. Welsh & K. I). Heil.
It is compared to A. nana S. Watson based on sim¬
ilarities in stature and anthocarp structure and to
A. fragrans Nuttall in the caulescent stems. A key
to closely related taxa is included.

Key words: Abronia, New Mexico, nomencla¬
ture, North America, Nyctaginaceae.

The new Abronia is part of voucher specimens
collected during the 1999 and 2000 field seasons for
the San Juan Basin Flora Project. The flora covers
all the drainage system and lands in the four corners
area of Arizona, Colorado, New Mexico, and Utah
that drain into the San Juan River. The target date
for publication is 2006. Exploration of the unique
habitats in the San Juan Basin area has yielded new
state and county records, as well as new taxa. As
this study progresses we expect additional taxa will
be discovered and named over the next decade or
so. Initial attempts at identification were made by
use of the early treatments of Abronia in New Mexico
by Standley (1909), Tidestrom and Kittell (1941),
Martin and Hutchins (1980), and the treatment by
Galloway (1975). The specimens proved contrary to
all of the existing treatments of the genus. The col¬
lections of an unidentified species of Abronia were
subsequently compared to the large set of reference
material at Brigham Young University (BRY), as well
as that at San Juan College (SJNM), and identified
as a new species. Two additional collections from
New Mexico were provided by Richard Spellenberg,
New Mexico State University (NMC).

Abronia bolackii N. D. Atwood, S. L. Welsh & K.
D. Heil, sp. nov. TYPE: U.S.A. New Mexico: San
Juan Co., tributary of Stuart Canyon S of San
Juan River on Bolack Ranch, 36°40'27.8"N,
108°8'51.4"W, 18 May 2000, N. Duane Atwood ,
Kenneth Heil, Steve O'Kane & Arnold Clifford
25673 (holotype, BRY; isotypes, ARIZ, ASU,
GH, MO, NY, RM, SJNM, US). Figure 1.

Similis A. nana in habitu generali sed rhizomatibus
gracilis repentibus, caulibus longioribus, foliis et fructi-
feris bracteis majoribus, corollae tubo breviori et floribus
viridi-albidis, et inflorescentiis axillaribus nec scaposis
absiniilis; et similis A. fraganti in caulibus elongatis sed
in rhizomatibus elongatis perianthiis brevioribus et antho-
carpis parvioribus et pubescentiis differt.

Perennial, the stems 9—18 cm high above
ground, arising from vertical branches from cord¬
like creeping rhizomes and forming extensive col¬
onies; aerial stems short-caulescent with 1—3 ap¬
parent internodes, these shorter than the leaves,
puberulent; leaf blades, 1—2.8 X 0.5-1. 6 cm wide,
elliptic, oblong to ovate, entire, the margins puber¬
ulent; petiole 0.5— 3.2 cm long; inflorescence glan-
dular-puberulent; the peduncles 1.8— 5.5 cm long;
bracts 5, green becoming white at maturity, broadly
ovate, rounded or apiculate, 6—10 X 4—10 mm,
with evident white interrupted veins; flowers 16 to
25 per head, the perianth green-white, the tube 7—
11 mm long, glandular, the lobes ca. 3 mm wide;
anthocarps obconic, (3- to) 5-winged or angled, pu¬
berulent. 5—6.7 mm long.

Habitat and distribution. Gypsiferous clay lens
of the Ojo Alamo Formation, from 1678 to 1724 m
elevation, in Juniperus, Hilaria, Amelanchier, Pur-
shia, and Ephedra community. Known only from
San Juan County, New Mexico (Fig. 2).

The species is named lor Tommy Bolack, the
owner and operator of the B-Square Ranch. For
years Mr. Bolack has practiced conservation and
watershed management and has helped protect the
native flora against habitat loss and habitat frag¬
mentation due to development. Along with San
Juan College, Mr. Bolack is helping fund the San
Juan Flora Project. The type locality of A. bolackii
is situated in the southeastern portion of his ranch.

Abronia bolackii is similar in stature and antho¬
carp structure (size and shape) to A. nana as in¬
dicated in the diagnosis. However, the new taxon
differs from A. nana, in being caulescent and lack-

Novon 12: 167-169. 2002.

1cm

168

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Volume 12, Number 2
2002

Atwood et al.

Abronia bolackii from New Mexico

169

Figure 2. Collection locations of Abronia bolackii in San Juan County, New Mexico, U.S.A.

ing persistent leaf bases, not caespitose-subacau-
lescent and with persistent leaf bases, and in pos¬
sessing a slender, creeping rhizome. The caulescent
nature of A. bolackii is similar to and probably in¬
dicative of relationship with caulescent Abronia fra-
grans, which occurs in the general vicinity. It dif¬
fers from that species in being rhizomatous and
having shorter and smaller perianths, with the tube
7—11 mm long in A. bolackii and 10—25 mm in A.
fragrans. Additionally, A. fragrans is a much larger
plant with different anthocarp morphology, larger
thicker leaves, usually larger flowers, a taproot, and
longer, multicellular, dense triehomes on the an-
thocarps. Closely allied species, both geographi¬
cally and possibly genetically based on its morpho¬
logic similarities, can be distinguished by the
following key:

Kky to Related Taxa

la. Plants definitely caulescent; caudex arising from
deep-seated rhizomes or from a taproot, seldom
with a thatch of marcescent leaf bases.

2a. Plants with slender, cordlike deep-seated
rhizomes, forming extensive colonies; flow¬
ers greenish white, the tube 7-1 1 mm long;
anthocarps 5—6.7 mm long, the apex short

hairy with simple hairs . A. bolackii

2b. Plants lacking rhizomes, arising from tap¬
roots; flowers greenish to rose or purplish,
the tube 10—25 mm long; anthocarps 5—12
mm long, the apex more densely pubescent
with longer multicellular hairs .... 4. fragrans
11). Plants subscapose, the leaves mostly all basal;

caudex often with a thatch of marcescent leaf
bases, situated atop a thickened taproot . . A. nana

Paratypes. U.S.A. New Mexico: San Juan County, B-
Square Ranch, first road SE near mouth of Stewart Can¬
yon. up road ca 1.5 mi., 36°40'23.5"N, 108°08'57.3”W,
18 May 20(H). K. Heil 14289 (SJNM); 10 mi. SSW of Wa-
terfiow on the Navajo Mine W ol the Neck, 2 June 1985,
R. Spellenberg & R. Corral 8213 (NMC); 6 mi. N of Fruit-
land, T29N, RI4W, S7, 12 May 1994, R. Sivinski * K.
Lightfoot 2665 (NMC).

Acknowledgments. The authors are indebted to
Noel H. Holmgren and Larry C. Higgins (or their
constructive reviews of the manuscript and to BRY
illustrator Shannie Workman for the excellent illus¬
tration. We thank Richard Spellenberg for the loan
of NMC Abronia collections and Randy Baker in
the Monte L. Bean Museum for help with the graph¬
ics. Also special thanks are given to Victoria Hol-
lowell (MO) for her invaluable help during the ed¬
iting process.

Literature Cited

Galloway, L. A. 1975. Systematics of the North American
desert species of Abronia and Tripterocalyx (Nyctagi-
naceae). Brittonia 27: 328—347.

Martin, W. C. & C. R. Hutchins. 1980. A Flora of New
Mexico, 2 vols. J. Cramer, Germany.

Standley, P. C. 1909. The Allioniaceae of the United
States with notes on Mexican species. Contr. IJ.S. Natl.
Herb. 12: 303-389.

Tidestrom, I. & T. Kittell. 1941. A Flora of Arizona and
New Mexico. Catholic Univ. America Press, Washing¬
ton, D.C.

Una Nueva Especie de Croizatia (Euphorbiaceae) del Ecuador

Carlos E. Ceron

Herbario “Alfredo Paredes”, Escuela de Biologfa de la Universidad Central, Apartado

17.01.2177, Quito, Ecuador

Grady L. Webster

Section of Plant Biology, University of California, Davis, California 95616, U.S.A.

RESUMEN. Se describe e ilustra una nueva espe¬
cie de ( ] roizalia (Euphorbiaceae) de la Costa Ecua-
toriana. Croizatia cimalonia es una especie arbus-
tiva o pequena arborea que se caracteriza por
formar colonias entre alturas de 300-800 m en el
Cerro Cimalon de la Reserva Ecologica Manglares-
Churute. Se caracteriza por los tallos con corteza
externa fisurada. pecfolos cortos de 5—13 nun. es-
triados, flores pentameras, verdosas, pedicelos fe-
meninos 25—40 nun de largo, estrigillosos, frutos
capsulas 3-lobada.

Abstract. A new species of Croizatia (Euphor¬
biaceae) from the costal lowlands of Ecuador is de¬
scribed anil illustrated. Croizatia cimalonia is a
shrub or small tree that forms stands between 300
and 800 m on Cerro Cimalon in the Reserva Eco-
Idgica Manglares-Churute. It is characterized by
trunks with fissured external bark; short, trans¬
versely striate petioles (5—13 mm long); greenish
pentamerous (lowers, shortly strigose pistillate ped¬
icels (25-40 mm long); and 3-lobed, capsular
fruits.

Key words: Croizatia , Ecuador, Euphorbiaceae.

Croizatia Steyermark es un genero de aproxi-
madamente 5 especies de arbustos y arboles pe-
quenos de bosques piemontanos en Sudamerica; se
conocen especies de Panama. Venezuela y Colom¬
bia (Webster et ah, 1987; Dorr. 1999). En Ecuador
una especie no descrita de la Amazonia ecuatoria-
na, 0llgaard et id. 38956 (AAU), es citada por
Webster et al. (1987) y Jorgensen y Leon-Yanez
(1999). Para la Costa Ecuatoriana, el genero es la
primera vez que se registra. Ea localidad donde se
ha registrado esta especie es un remanente ubicado
en uno de los cerros tjne forma parte de la Cordil¬
lera Costanera, protegida en la Reserva Ecologica
Manglares-Churute a unos 30 km del puerto mas
importante del Ecuador, Guayaquil.

Croizatia cimalonia C. E. Ceron & G. L. Webster,
sp. nov. TIPO: Ecuador. Provincia del Guayas:
Canton Naranjal, Reserva Ecologica Mangla¬
res-Churute, Cerro Cimalon, 300—400 m,
79°33'W, 02°37'S, Bosque siempre-verde Pie-
Montano, 26 julio 1999, C. E. Ceron , C. Car-
vajal & R. Romero 38915 (holotipo, QAP fe-
menino; isdlipos, AAU, DAV. MO. QCA.
QCNE). Eigura 1.

Frutex arborescens 1—8 m altus; a C. neotropica differt
pedicel is fructiferis hrevioribus, strigosis; a C. naiguatensi
differt pedicellis staminatis longioribus, foliis spathulatis
non elliptieis; a C. panamensi differt stipulis minoribus.

Arbusto o arbol de 1—8 m de alto X 3—5 cm de
DAP. algunos arbustos ramifieados desde la base,
corteza externa fisurada color blancuzco, con pre-
sencia de Briophytos, corteza interna rosada, olor
acre similar a Daphnopsis Martins (Thymeleaceae),
madera rosada. Lamina de las hojas generalmente
espatulada, apice ± acuminado, base atenuada,
14—38 cm de largo. 4—9.5 cm de ancho, glabra o
inconspieuamente estrigosa en el enves. nervios la-
terales 9—14 en cada lado, nervios terciarios for-
mando retfculos con los secundarios, pecfolo de 5—
13 mm de largo, curvado, estrfas finas transversa-
les; estfpulas escariosas, triangular-acuminadas,
erectas en el apice de las ramitas, color vinoso, 5—
8 mm ile largo, 3—5 mm de ancho, semideeiduas.
Flores masculinas color crema, ligero ramifloras;
pedicelo de 8—13 mm de largo, hirto; s^palos 5.
elfpticos, enteros, 3—4.5 mm de largo, 2—3 mm de
ancho, hirsutos; petalos 5. obovados, subenteros
con los bordes ciliados, 2-3 mm de largo, 1-1.5
mm de ancho; disco masivo, liso y glabro, 1 mm
de alto. 2 mm de ancho; estambres 5, filamcntos
1.5-2 mm de largo, hirsutos; anteras elfpticas de 1
mm de largo, pistilodio 3-lobado, denso hirsute, 2
mm de alto. Flores femeninas solitarias en cada
axila; pedicelo estrigilloso, de 25—40 mm de largo
en el fruto; sepal os 5, persistentes en el fruto, lan-
ceolados, apice obtuso, color verde, estrigillosos, 5—

Novon 12: 170-172. 2002.

Volume 12, Number 2
2002

Ceron & Webster

Croizatia cimalonia from Ecuador

171

Corte transversal de fruto. (Cerdn et al. 38925, QAP.)

10 mm de largo, 2—5 mm de aneho; petalos 5, al-
ternos con los sepalos, redondeado obovados o su-
borbiculares, 1—2.5 mm de largo, 2 mm de aneho,
escaso ciliados en el cuerpo y densamente en los
hordes, color verde; disco anular, glabro, entero. 5
mm de aneho; ovario denso sericeo, 3 mm de largo
y aneho, estilo expandido, 1.5— 2.5 mm de largo,
tres veces bffido, la porcion no ramificada 0.5—1
mm de largo, la porcion ramificada media de 1 mm

de largo, la porcion ramificada apical de 0.3-0.5
mm de largo. Capsula oblada, 3-lobada, 10—15 mm
de alto, 17—23 mm de aneho, glabra, color verde,
ligeramente estrigosa en las suturas, estilo persis-
tente tie 2—3 mm tie largo; eje central de 6 mm de
alto, apice dilatado en 3 alas oblonga-obovadas con
el apice redondeado de 7 mm de largo, 5 mm de
aneho. Semillas dos por loculo, trigonas, lisas, color
cafe, 8—12 mm de largo, 5—7 mm de aneho, hilo

172

Novon

medio de 2 mm de ancho, eotiledones de 10 mm
de largo.

Variation y afinidad. Croizatia cimalonia difie-
re de C. neotropica Steyermark (Distrito de Bolivar,
Venezuela), en los siguientes earaeteres: La corteza
exterior fisurada, hojas mas largas, pedicelo feme-
nino estrigilloso, menos largo, y hay mayor varia-
cion en el luibito de la planta, mayor numero de
los nervios secundarios de las hojas y en las se-
millas 8—12 mm de longitud. En C. neotropica, el
pedicelo fomenino es glabro, mas largo. Menor nu¬
mero de nervios secundarios en las hojas y en las
semillas 10.5—12 mm de longitud.

Clave de las Especies Descritas de Croizatia

la. Pedicelos de los frutos glabros, 35—45 mm de

largo, frutos 1 o (2) por axila . C. neotropica

11). Pedicelos de los frutos strigosos o hirtellos, 8—

40 mm de largo.

2a. Flores pistiladas 2 o 3 por axila; filamentos

libres . C. brevipetiolata (Secco) Dorr

2b. Flores pistiladas solitarias; filamentos libres
o connados.

3a. Filamentos connados; estipulas 10—20

mm de largo, ± persistentes .

. C. panamensis G. L. Webster

3b. Filamentos libres; estipulas 4—8 mm de
largo, ± decfduas.

4a. Pedicelos estaminados 3—4 mm de
largo; pedicelos de los frutos 8—12
mm de largo; pbtalos estaminados

1 — 1.5 mm de largo .

. C. naiguatensis Steyermark

4b. Pedicelos estaminados 8—13 mm de
largo; pedicelos de los frutos 25—

40 mm de largo; petalos estamina¬
dos 2-3 mm de largo . . . C. cimalonia

Distribucidn y ecologia. La especie nueva es
conocida solamente de dos poblaciones en la Re-
serva Ecoldgica Manglares-Churute. En el Cerro
Cimaldn entre 300—400 m, se encuentra en el so-
tobosque formando densos rodales. En muestreos
de transectos en 0.05 ha se ha encontrado hasta 46
individuos (29.6%) y es la especie mas frecuente,
igual entre 420—470 m, es la especie mas frecuente
con 82 individuos (45%). La topograffa es muy ac-

eidentada en el Cerro Cimalon, aunque la especie
a veces esta en pequenas mesetas planas <le este
cerro; el suelo es rojo y pedregoso con muy poco
humus, en la transicion de los bosques seeos a los
de neblina. La especie se encuentra tambien en
forma menos abundante en el Cerro Masvale de la
misma Reserva Manglares-Churute entre altitudes
de 780—800 m, con una presencia de 8 individuos
(6.1%) en 0.05 ha. Los dos cerros se encuentran
separados entre si por una planicie dislurbada y
cultivada de aproximadamente unos 4 km de dis-
tancia (Ceron, 1996). Se eneontrd con flores y frutos
en agosto— septiembre de 1992 y en julio de 1999.

Esta especie es nombrada por el Cerro Cimalon
de la Reserva Ecoldgica Manglares-Churute, loca-
lidad tipo.

Paratipos. ECUADOR. Guavas: Canton Naranjal,
Reserva Ecoldgica Manglares-Churute, Cerro Cimaldn,
79°33'W, 02°37'S, bosque siempre-verde Pie-Montano,
300-400 m. 26 jul. 1999, Cerdn et at. 38918, 38925
(DAV, MO, DAP, QCNE); Cumbre del Cimaldn subiendo
por el Polfgono, 350—470 m, 8 ago. 1992, Cerdn et at.
19900 (QAP), 28 sep. 1992, Cerdn et at. 20442 (QAP,
QCNE); Cerro Masvale, 780— 800 m. 15 ago. 1992, Cerdn
et at. 20240 (QAP. QCNE).

Agradecimientos. Nosotros estamos agradecidos
a los senores Ramon Romero y Maximo Reyes,
guardaparques de la Reserva Ecoldgica Manglares-
Churute por su asistencia en el campo; al herbario
QCNE por permitirnos util izar el mismo; al Simon
Laegaard del Herbario AAU, por la revision del
artfculo, comentarios y traduccibn del Rcsumen; y
al Nigel Pitman de Duke University, por sus co¬
mentarios y revision general del artfculo.

Literatura Citada

Cerdn, C. E. 1996. Diversidad, especies vegetales y usos
en la Reserva Ecoldgica Manglares-Churute, Provincia
del Guayas, Ecuador. Revista Geogr. (Quito) 36: 1—92.
Dorr, L. J. 1999. A new combination in Croizatia (Eu-
pborbiaceae). Sida 18: 831—836.

Jorgensen, P. M. & S. Ledn- Yanez (editors). 1999. Cata¬
logue of the Vascular Plants of Ecuador. Monogr. Syst.
Rot. Missouri Rot. Gard. 75.

Webster, G. I... L. Gillespie & J. Steyermark. 1987. System-
aties of Croizatia (Euphorbiaceae). Syst. Rot. 12: 1-8.

Novae Gesneriaceae Neotropicarum VIII: Alloplectus grandicalyx ,
a New Species from Colombia and Ecuador

John L. Clark12 and Laurence E. Skog2

'George Washington University, Biological Sciences, 2023 G Street, N.W., Washington, l).C.

20052, U.S.A. clark.john@nmnh.si.edu

2Botany Section-Systematic Biology, Smithsonian Institution, P.O. Box 37012, National
Museum of Natural History, MRC-166, Washington, D.C. 20013-7012, U.S.A.

skog.larry@nmnh.si.edu

Abstract. A new species, Alloplectus grandica¬
lyx J. L. Clark & L. E. Skog, is described from
northern Ecuador and southern Colombia. Alloplec¬
tus grandicalyx is differentiated from other similar
species on the basis of larger calyx lobes, longer
pedicels, and villous pubescence on the stems,
leaves, petioles, and calyx lobes. A key is provided
to help distinguish A. grandicalyx from other
shrubby and resupinate flowered Alloplectus spe¬
cies. In addition, the commonly confused and wide¬
ly spread Alloplectus tetragonoides Mansfeld is dis¬
cussed and neotypified.

Key words: Alloplectus, Colombia, Ecuador,
Gesneriaceae.

An ongoing study of Alloplectus Martius has re¬
vealed a new species from Ecuador and Colombia
that can be distinguished from congeners by the
combined characters of calyx lobes that cover the
bottom third to half of the corolla, pedicels that
usually exceed the petioles, and villous pubescence
on the stems, leaves, petioles, and calyx lobes.

Alloplectus is a genus in need of revision as the
most recent treatment of the entire genus is over
100 years old (Hanstein, 1865). Hanstein’s publi¬
cation focused on the Gesneriaceae at the botanical
garden in Berlin and provided an overview of the
family. Alloplectus was only a part of an entire
monograph of the Gesneriaceae known at that time,
but Hanstein recognized 30 Alloplectus species in
his treatment. More recent estimates of the genus
range from 75 (Burtt & Wiehler, 1995) to 65 (Wieh-
ler, 1973). After a thorough review of the 119
names listed in Index Kewensis, only 32 are con¬
sidered by us to be currently accepted names in
Alloplectus. Below are described two species of Al¬
loplectus that have been confused, one new and the
other poorly known and needing neotypification.

Alloplectus grandicalyx J. L. Clark & l„ E. Skog,
sp. nov. TYPE: Ecuador. Napo: Canton Archi-
dona, Beserva Ecologica Antisana, Sector
Guacamayos, Sendero Jumandy-La Virgen
(km 30), path that follows potential oil pipe¬
line, 00°38'S, 77°49'W, 1700-1800 m, 1 May
1998, ./. L. Clark, E. Narvaez A T. Pauchi
5449 (holotype, QCNE; isotypes, AAE, COL.
E, MO, QCA, SRP, US). Eigure 1.

Haec species A. tetragonoides similis sed pedicellis
longioribus et calycibus grandioribus differt.

Terrestrial suffrutescent shrub; stem erect, qua¬
drangular, succulent, becoming woody, 0.5— 1.5 m
tall, to ca. 1 cm diam., usually glabrescent basally,
villous pubescent distally, unbranched, internode
length 3-15 cm, nodes and leaf scars swollen.
Leaves opposite, nearly equal in a pair; blades el¬
liptic, 8—22 X 4—10 cm, apex acuminate, base
acute, margin serrate, above green, rugose, gla¬
brous, below all green, all red, or green with red
tinge, sparsely to densely pilose; petioles terete, 3—
8 cm long, green, sparsely to densely pilose. Inflo¬
rescences axillary, of 1 to 4 pendent, epedunculate
flowers per axil; bracts caducous at anthesis, ovate,

1 .5-3 X 1-2 cm, red; pedicels 4-9 cm long, red,
villous. Flowers resupinate, showy, and zygomor-
phic; calyx lobes 5, conduplicate, erect, nearly
equal, ventral lobe narrowly ovate, other lobes
ovate, 3-4 X 2-3 cm, red, base cordate, apex ob¬
tuse, margin serrate; corolla tubular, 4—5 cm long,
dark red with or without yellow lobes, outside
sparsely pilose becoming densely pilose at apex,
inside glabrous, base gibbous, ampliate toward the
limb, the mouth ventricose on dorsal surface, limb
1-1.5 cm wide, the lobes free, nearly equal. 3—5
X 3-8 mm, entire; stamens 4, included, filaments
glabrous, connate at base and adnate to corolla tube
base for 3—5 mm, anthers included, coherent by
their apices, ca. 3.5 X 1.5 mm; disc of a single
bilobed, unlobed, or multilobed gland, glabrous;

Novon 12: 173-178. 2002.

174

Novon

Figure 1. Alloplectus gmndiealyx J. I.. Clark & l.. K. Skog. — A. Habit. — B. Leaf margin. — C. Corolla. — I). Corolla
opened to show stamens. — E. Calyx. — F. Calyx opened and corolla removed to show bilobed nectary. — G. Young
fruit. — II. Seeds. A from A'. A. Schultes & M. Villarreal 7667: 11 from ./. Cuatreeasas 8620: C-f from 1/. A. Bristol
266: G from the isotype, ./. A. Clark. A'. Narvaez it' T. Pauchi 5440: II from 4. Gentry , M. Monsalve it' P. Silverstone
47657 V

Volume 12, Number 2
2002

Clark & Skog

Novae Gesneriaceae Neotropicarum VIII

175

ovary ca. 0.6 X 0.4 cm, woolly, style included, 4
cm long, glabrous. Fruit a fleshy capsule usually
hidden in persistent calyx lobes, pendent, ovoid,
ca. 1.5 X 1.0 cm, woolly; seeds 0.8-1. 1 mm long,
numerous, fusiform, dark brown, and longitudinally
striate.

Distribution and ecology. Alloplectus grandica-
lyx is found in the montane forests of Ecuador and
southern Colombia, but is most common between
1500 and 2500 m on the northeastern slopes of the
Ecuadorian Andes. The species frequently overlaps
with A. tetragonoides, which is trans-Andean and
more common at lower elevations. At mid eleva¬
tions on the eastern slopes of the Andes, these two
species are apparently sympatric. In areas where
these two species overlap A. grandicalyx can be
distinguished from A. tetragonoides by having larg¬
er calyx lobes, longer pedicels, and villous pubes¬
cence. The resupinate flowers in these two species
are typical of most Alloplectus species with tubular
ventricose corollas. However, little is known about
the adaptive functions of resupinate flowers in Al¬
loplectus or other genera in Gesneriaceae where it
is known; e.g., Nematanthus Schrader (Chautems,
1988).

Alloplectus grandicalyx is distinguished from the
similar Alloplectus tetragonoides and other species
of Alloplectus by having unusually large calyx lobes
that cover the bottom third to half of the corolla
tube. The pedicel is also longer than that normally
found in other species of this genus and usually
exceeds the length of the petiole. The calyx will
commonly appear entire on herbarium sheets be¬
cause the lobes are imbricate with involute mar¬
gins. Some characters that are difficult to see on
herbarium collections but readily available in the
field include the dark red to maroon corollas with
or without yellow lobes and the tendency for the
leaf blades to be bullate. Also, most other similar
Alloplectus species will have a lighter red or yellow
corolla and non-bullate leaves (e.g., many popula¬
tions of A. tetragonoides from the Napo and Tun-
gurahua provinces of Ecuador). Most Alloplectus
species have a single or two-lobed nectary gland,
whereas A. grandicalyx sometimes has a multilobed
nectary gland. We are including below a description
of A. tetragonoides because of the possible confu¬
sion with A. grandicalyx and A. tetragonus (Han-
stein) Hanstein. In order to stabilize the species
concept of A. tetragonoides we have selected a neo¬
type.

Paratypes. COLOMBIA. Caqueta: Rfo Hacha, ./.
Cuatrecasas 8553 (US); Guadalupe— Florencia road, km
14, W of pass, X. Londoho & L. P. Kvist 99 (COL, US).

Huila: eastern slope of Cordillera Central, Finca Meren-
berg, km 101 La Plata-Popayan road, B. A. Stein 3730
(MO, US); Cordillera Oriental, San Andres, ,/. Cuatrecasas
8620 (COL, US). Putumayo: Sihundoy, near Rio Blanco
(via Mocoa), E. Hernandez, A. Guerrero & A. Estrada 401-
A (US), 404 (US); Rfo Mocoa drainage, between Scbamate
and San Antonio, J. A. Ewan 16693 (MO. US); valley of
Sibundov. R. E. Schultes A M. Villarreal 7475 (US), R. E.
Schultes A M. Villarreal 7667 (US). ,/. Cuatrecasas 1 1460
(US), M. L. Bristol 366 (COL, GH. US). ECUADOR. Car-
elii: 12 km E of Maldonado on road to Tulcan, A. 11.
Gentry A G. Shupp 26626 (MO, SEE): Rfo Verde and ridge
flanking Cerro Golondrinas, U. .S’. Hoover 2354 (MO). Mo-
rona-Santiago: Tumbes, 17-18 km N of Gualaquiza on
road to Indanza, G. Hurling A L. Andersson 24222 (GB,
US); Limon Indanza, Cordillera de Huaracayo, E of Cor¬
dillera del Condor and Rfo Coangos, E of Shuar village of
Tinkimints, D. Neill A ./. Manzanares 13126 (MO. QCNE,
US). Napo: Baeza Tena road, S slope of Cordillera de los
Guacamayos, J. /., Luteyn & M. Lebron-Luteyn 5669 (CAS,
MO, NY, SEE, US), G. Hurling & Andersson 19501 (GB,
US), W. G. D'Arcy 14089 (MO, US), W. Palacios 6896
(MO, QCNE, US),’ W. Palacios 5931 (MO, QCNE, US). B.
A. Stein 2647 (MO, US), J. H. Kirkbride , ,/r. A 11. Chamba
R. 4191 (U, US); Comunidad de Santa Lucfa de Bermejo,
.4. Alvarez, (). Brito & S. Romero 867 (MO, QCNE, US);
Cosanga, trail E of town, ./. D. Boeke & J. B. McElory
(GH, NY, SEE. US); Yanayacu Biological Station and Cen¬
ter for Creative Studies, 5 km W of Cosanga, J. L. Clark
A H. F. Greeney 5774 (QCNE, US). Orellana: Sumaco
Napo Galeras National Park, Volcan Sumaco, J. L. Clark
2282 (QCNE, US). Sucunihios: Reserva Ecoldgiea Cay-
ambe Coca, Volcan Reventador, trail between Rfo Quijos
and refugio, J. /.. Clark 4448 (QCNE, US). Tungiirahiia:
Hacienda San Antonio, 2 km E of Banos on Rfo Ulba, B.
A. Stein 2929 (NY); Rfo Verde, forested trail from Banos
Puyo road towards Cascada de San Miguel, J. L. Clark, E.
Narvaez A E. Varros 5719 (QCNE, US). Zainora-Chin-
chipe: W of Zamora, J. Hart 1172 (A); Agua Rica, along
main road from Limon (Gral Leonidas Plaza Gutierrez) to
Gualaceo, ,/. L. Clark . L Jost A F Sanchez 5939 (AAU,
AZUAY, COL, K, MO, NY, QCA, QCNE, US).

Alloplectus tetragonoides Mansfeld, Biblioth.
But. 116: 144-145. 1937. TYPE: Ecuador.
Tungurahua: Canton Banos, Parroquia Rfo Ne¬
gro, near Rfo Topo, 01°23'S, 78°10'W, 1400
m, 17 Jan. 2001, /. L. Clark & V. Duran 6000
(neotype, designated here, QCNE; isoneotypes,
AAU, COL, F, K, MO, NY, QCA, S, US).

Terrestrial suffrutescent shrub or rarely epiphytic
herb; stem erect, quadrangular, succulent, becom¬
ing woody, 0. 5-2.0 m tall, to ca. 1 cm diam., usu¬
ally glabrescent, sometimes sparsely pubescent dis-
tally, unbranched, internode length 3-12 cm, nodes
and leaf scars swollen. L eaves opposite, nearly
equal in a pair; blades elliptic, sometimes ovate,
9—39 X 3.5-24 cm, apex acuminate, base cuneate
or acute, margin serrate, above green and sparsely
pil ose, below green, red, or green with red tinge,
sparsely pilose to densely pilose (especially pilose
on young foliage); petioles sulcate, 2—14 cm long,

176

Novon

green, glabrous to densely pilose. Inflorescences ax¬
illary. of 1 to 4 pendent, epeduneulate flowers per
axil; bracts caducous at anthesis, ovate, 2-3 X 0.5—

1.5 cm, red or green; pedicels 1—5 cm long, gla¬
brous to pilose. Flowers resupinate, showy, and zy-
gomorphic; calyx lobes 5, erect, nearly equal,
ventral lobe narrowly ovate, other lobes ovate, 1.5—

2.5 X 1-2 cm, all green, all red, red with green
margins, or green with red margins, sparsely pilose
to hirsute, midvein especially pilose, base truncate
to slightly cordate, apex acute to acuminate, margin
serrate to serrulate; corolla tubular, 3—6 cm long,
red, yellow, or red with yellow lobes, outside dense¬
ly pilose, inside glabrous, the base gibbous, am-
pliate toward the limb, the mouth ventricose, limb
1.4— 2.0 cm wide, the lobes free, nearly equal, ca.
4—5 X 5—8 mm, entire; stamens 4, included, fila¬
ments glabrous, connate at base and adnate to co¬
rolla tube base for 3—5 mm, anthers included, co¬
herent by their apices, ca. 2.0 X 2.5 mm; disc of
a single bilobed or unlobed gland, glabrous; ovary
ca. 0.7 X 0.5 cm, hirsute, style included, 5 cm
long, glabrous. Fruit a fleshy capsule usually hid¬
den in persistent calyx lobes, pendent, ovoid, ea.

1 .5 X 1 cm, green; seeds ca. 0.9 X 0.3 mm, nu¬
merous, fusiform, dark brown, anti longitudinally
striate.

Distribution and ecology. Alloplectus tetrago-
noides is common in Andean montane forests
throughout Ecuador and Colombia. It occurs on the
Western and Central Cordilleras of Colombia and
both eastern and western Andean slopes of Ecua¬
dor. flic species is frequent between 1500 and
1800 m, but occurs occasionally to 2200 m. The
low mountains along the border with Panama and
the drier forests of northern Colombia probably act
as an ecological barrier to this species’ range and
may explain why it has not been collected in mon¬
tane forests north of Colombia.

Alloplectus tetragonoides is highly variable, lead¬
ing to its confusion with other species (see key be¬
low). The undersurface of the leaves can be totally
green, green with a tinge of red, to totally red. Ca¬
lyces can be green or red, and with either com¬
pletely yellow, completely red, or yellow with red
lobed corollas. It is common to find individuals with
green calyces and yellow corollas with red lobes
growing sympatrically with individuals that have
completely red perianths. The most useful charac¬
ter in differentiating A. tetragonoides from similar
species such as .4. tetragonus and A. grandicalyx
is its smaller calyx with lobes having serrulate mar¬
gins (f ig. 2). In comparison to 4. tetragonoides. the
calyx lobe margin of 4. tetragonus is more dentate.

The calyx of 4. grandicalyx is two to three times
larger than that of 4. tetragonoides and 4. tetragon¬
us (viz. Eig. 2). All of the illustrated species in
Figure 2 differ from Alloplectus cucullatus C. V.
Morton (included in key, but not in illustration) by
having calyx lobes that are plane relative to the
eucullate calyx lobes of 4. cucullatus.

The first author visited the Rio Topo area in the
Tungurahua province of Ecuador in January of
2001. This area is where Ludwig Diels made the
original collection selected for the holotype of Al¬
loplectus tetragonoides (Mansfeld, 1937). No dupli¬
cates of L. Diels 1002 are known to exist. Although
many pre-World War II photos of Gesneriaceae type
specimens from the Berlin herbarium are extant, we
have not been able to locate photos ol the collec¬
tions that Mansfeld designated as type specimens.
Most of Mansfeld’s publications on Gesneriaceae
were in the later part of the 1930s. Therefore, the
collections that Mansfeld used for his species con¬
cepts probably only existed as types for a few years
until they were subsequently destroyed in the af¬
termath of Allied bombing of Berlin in 1943. Thus,
to stabilize the concept of the species, we have des¬
ignated a neotype collected from the type locality
where the species is extremely common.

Representative specimens. COLOMBIA. Cauea: Car-
retera Mococa Pitalito, sitio Sajonia, F. Betancur et al.
5538 (COL, US). Choco: Mpio. de San Just5 del Palmar,
SW of town, F. Forero et al. 3402 (COL, US(2)). Iluila:
Rfo Villalobos, region of the confluence of Rfos Villalobos
and Cauchos, R. F. Scludtes & M. Villarreal 5204 ((41.
US). Narino: 10 km de Guayacana por la via Tumaco
Junfn, X. Londono et al. 510 (US). Putumayo: along road
between Pasto and Mocoa, El Selencia, H. Kennedy 210
(LIS). Valle del Cauea: Cordillera Occidental, hoya del
Rfo Sanquininf, J. Cuatrecasas 15503 (US); Mpio. Dagua,
Corregimiento el Carmen, M. Amaya & J. F. Smith 581
(COL). ECUADOR. Carchi: cloud forest above Maldon¬
ado, C. Luer et al. 3374 (SEL). Imbahura: Carretera Ota-
valo Selva Alegre, //. van der Werjf & fL Palacios 10558
(MO, US). Maiialn': Carmen, G. II. II. Tale 472 (US). Mo-
rona-Santiago: Cordillera del Cbndor, Cuangos, 20 km
E of Gualaquiza, 4. Gentry 80029 (MO. US (2)); Cordillera
Cut Lieu, above Rfo Tzontza, F. Prieto CuP-12 (NY). Napo:
Cantdn Archidona, Reserva Ecologica Antisana, comuni-
dad Shamato, entrada por km 21 Shamato. J. L. Clark
5033 (AAU. COL, E. MO, QCNE, SRP. US). Orellana:
Southern slopes ol Volc4n Sumaco, Hollfn Loreto road, Km
25, I). Neill & M. Asanza 8882 (MO. QCNE, US). Pas-
taza: Banos Puyo road, 5 km W ol Mora. /.. P. Kvist 6 0334
(QCA, US). Pichincha: along old road between Santo Do¬
mingo— Quito and Chiriboga, Mendoza et al. 625 (QCA,
QCNE, US). Sucumbios: Reserva Ecologica Cavambe
Coca, Volcan Reventador, trail between Rfo Quijos and
refugio, J. L Clark 4454 (AAU, COL, E. MO, NY, QCA,
QCNE, SRP, US). Tiinguruhua: 5 km W ol Banos, C. II.
Dodson & I.. II. Thien 977 (LIS(2)); between Rfo Maputo
and Rfo Margaritas. C. W. Pen/and & R. //. Summers 186
((41. US); roadside between Ambato and Banos, W. DArcy

Volume 12, Number 2
2002

Clark & Skog

Novae Gesneriaceae Neotropicarum VIII

177

1cm

Figure 2. Lateral calyx lobe variation in Alloplectus. — A. A. tetragonus. — B. A. panamensis. — C. A. grnndicalyx.
— I). A. tetragonoides. A from E. Hernandez et al. 751 ; B from A. Sytsma & L. Andersson 4611; C from the isotype, J.
L Clark, E. Narvaez & T. Pauchi 5449; D from D. Neill M. Asanza 8882.

178

Novon

14017 (MO, US). Zamora-Chinchipe: 10 km h of Pa-
quisha, (>. Hurling & /,. Arulersson 24095 (US).

Kl-.'i to CoMMONIA CONRSKI) SHRUBBY AU.OI’l.FA'.TlS
Spi ck s with 10 si pinati: Flowers

(This key is not intended for the non-resupinate or lianoid
species of Alloplectus.)

la. Distal leaves and calyx lobes covered with vil¬
lous or puberulous-pilose pubescence.

2a. Calyx lobe margin denticulate to dentate,
lobes plane, ovate, and acute at apex; corolla
red or appearing yellow due to the dense
yellow pubescence; widespread throughout
the mountains of Panama and Costa Rica,
rarely collected in Colombia ... ,4. tetragonus
2b. Calvx lobe margin serrulate to serrate, lobes
plane to cueullate, ovate to oblong, and apex
acute to obtuse; corolla completely red, com¬
pletely yellow, or yellow with red lobes,
da. Pedicels longer than petioles; calyx at
leasl one third the length of the corolla
tube, red; corollas dark red to maroon,
with or without yellow lobes; leaves
sometimes bullate; montane forests of
northern Kcuador and southern Colom¬
bia . .4. grandicalyx

3b. Pedicels shorter than petioles; calyx
less than one third the length of the co¬
rolla tube, color variable; corolla color
variable, red. yellow, and sometimes
yellow with red lobes; leaves not bul¬
late; common in Andean montane for¬
ests throughout Ecuador and Colombia
. 4. tetragonoides

lb. Distal leaves and calyx lobes covered with to-
men lose- woolh pubescence.

4a. Calyx lobes cueullate and ovate; corollas red
with dense yellow pubescence; transitional
and montane forests of northern Nicaragua,

Honduras, and Guatemala . 4. cucullalus

4b. Calyx lobes plane and oblong; corolla red

with white hirsute pubescence; common in
wet forests from Panama to Colombia, and

occasionally reaching Ecuador .

. A. panamensis C. V. Morton

Acknowledgments. Vie are grateful to Cathy
Pasquale-Johnson for preparing the illustration of
the new species. The first author thanks Alice Tan-
gerini for providing guidance in making the illus¬
tration of calyx lobes. We are especially grateful to
John K. Boggan, Harold Robinson, Peter M.
Jprgensen, J. F. Smith, and V. Hollowed for their
helpful comments on the manuscript. Fieldwork for
the first author was made possible by the Explorers
Club Washington Group, George Washington Uni¬
versity Chapter of the Society of Sigma Xi. and the
Jose Cuatreeasas Botanical Fund of the Department
of Systematic Biology- Botany, U.S. National Muse¬
um of Natural History.

I .iterature Cited

Rmtl. If. I.. X II. Wiehler. 1995. Classification of the fam¬
ily Gesneriaceae. Gesneriana 1: 1—4.

Chautems, A. 1988. Revision taxonomique et possibilities
(Thybridalions de Alematanthus Schrader (Gesneri¬
aceae), genre endemique de la foret cohere biTsilienne.
Dissertationes Botanicae, 112: 1—226. J. Cramer: Ber¬
lin. Germany.

Ilanstein, J. 1865. Die Gesneraceen des Kiiniglichen Her¬
bariums und der Garten zu Berlin, nebst monogra-
phischer Uebersicht der Familie im Ganzen, II.
Abschnitt. Gattungen und Arten. Drittes Stuck. Die Eu-
gesnercen. Rhvtidophylleen, und Beslerieen. binnaea

34: 225-462.

Mansfeld, R. 1937. Gesneriaceae, In: Beitrage zur Ketml-
nis der Vegetation und Flora von Ecuador. Biblioth. Rot.
116: 144-147.

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

A New Genus and Species in Cupressaceae (Coniferales) from
Northern Vietnam, Xanthocyparis vietnamensis

A. Farjon

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom.

a.farjon@rbgkew.org.uk

Nguyen Tien Hiep

Institute of Ecology and Biological Resources of the National Center for Natural Sciences
and Technology of Vietnam, Nghia Do, Can Giay, Hanoi, Vietnam. ntienhiep@hn.vnn.vn

l). A. Harder

Arboretum, UC Santa Cruz, 1 156 High Street, Santa Cruz, California 95064, U.S.A.

dkharder@cats.ucsc.edu

Phan Ke Loe

Department of Botany, Vietnam National University. Hanoi, Vietnam, pkeloc@yahoo.com

L. Averyanov

Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov Street 2. St.
Petersburg, 197376, Russia, av@herb.bin.ras.spb.ru

Abstract. Botanical explorations of heretofore
poorly inventoried limestone formations in the
northern Vietnamese border province of Ha Giang
have yielded a new taxon of conifer that is to be
classified in a cupressoid clade of Cupressaceae. It
has foliage and ovuliferous cone characters that
compare closely to Chamaecyparis nootkalensis (D.
Don) Spach, as well as a feature peculiar to it and
rare in conifers: the simultaneous occurrence ol ju¬
venile and mature leaves on normal plagiotropic
branching systems of mature trees. It is proposed
in this paper to unite C. nootkatensis with the newly
discovered species in a new genus Xanthocyparis ;
of this new genus and species a full description anil
illustrations are provided. Additional contributions
to this paper include observations on its habitat,
conservation status, and a taxonomic discussion.

Key words: conifers, conservation, Cupressa¬
ceae, North America, taxonomy, Vietnam, Xantho¬
cyparis.

The discovery of a new living conifer that is suf¬
ficiently distinct so as not to be accommodated
comfortably in any known genus is an exceptional
event which has only occurred a few times in the
last 50 years. The conifers have received much at¬
tention from taxonomists and others over a long

time even though they are a small group of plants
with no more than 650 species worldwide. This dis¬
covery has become even more interesting because
detailed comparisons and subsequent analysis have
revealed it to be related to a species of which the
generic placement has been controversial. It has
thus not only enlarged our knowledge of biodiver¬
sity, but also increased our understanding of phy¬
logenetic relationships of the conifers in the wider
context of the family Cupressaceae to which it be¬
longs. Implied in these relationships is a biogeo¬
graphic history encompassing eastern Asia and
North America that has been known from numerous
previously discovered relationships among plants of
both continents. The new conifer, its characters,
and taxonomic relationships are presented below.

Xanthocyparis Farjon & Hiep, gen. nov. TV PE:

Xanthocyparis vietnamensis Farjon & Hiep.

A genere Chamaecypari stomatibus utrinque in super-
ficiebus foliorum maturorum lateralium equaliter disposi-
tis, foliis maturiis lateralibus apices liberos (non adpres-
sos) gerentibus, strobilis bracteis-squamis 4 (raro 6)
suffultis; a genere Cupresso strobilis minoribus bracteis-
squamis seminibusque minus numerosis diagnoscenda.

Small or large evergreen trees, monoecious, with
fibrous bark exfoliating in longitudinal strips; heart

Novon 12: 179-189. 2002.

180

Novon

wood yellowish, slow growing. Foliage branches
spreading in plagiotropic sprays or drooping, form¬
ing a pyramidal, conical, or irregular flat-topped
crown. P’oliage of three types: juvenile linear
leaves, transitional leaves, and mature scale leaves.
Juvenile leaves present on seedlings only or also in
mature trees, radially disposed in alternating
whorls of four: foliage branchlets with this type of
leaves always sterile. Transitional leaves present on
seedlings only or also in mature trees, decussate,
divided in facials and laterals of nearly ecpial size,
the laterals weakly disposed in a plane; foliage
branchlets with this type usually sterile. Mature
leaves present in mature trees, decussate, dimor¬
phic in shape and size with the laterals strongly
flattened and disposed in a plane; foliage branch-
lets with this type often fertile. Leaves in whorls of
four or decussate. Pollen cones terminal and soli¬
tary on lateral branchlets with small scale leaves.
2.5—5 X 2-2.5 mm; mierosporophylls 10 to 16. de¬
cussate. peltate, bearing 2 (or 3) relatively large
microsporangia. Seed cones terminal and solitary
on lateral branchlets with unmodified scale leaves,
maturing to 7-11 X 10—12 mm, opening wider.
Bract -scale complexes in 2 (sometimes 3) decussate
pairs, fused at base, the upper pair(s) connate,
spreading wide to release the seeds, valvate to sub-
peltate. with a prominent central umbo. Central col¬
umella present or absent, small. Ovules axillary to
bracts, 1 to 5 per bract; seeds usually fewer, con¬
centrated on the upper pair (if 2 pairs) or middle
pair (if 3 pairs) of scales, flattened, with two thin
lateral wings. Seedlings with 2 cotyledons, followed
by juvenile linear leaves in whorls of four.

Eponymy. From Creek xantho — yellow, the
color of the wood, and cyparis — cypress.

1. Xanthocyparis vietnaineiisis Farjon & Hiep,
sp. nov. T\ PE: Vietnam. Fla (dang: Ouan Ba.
Bat Dai Son. Bat Dai Son Provincial Protected
Area, 10 Feb. 2001, I). K. Harder , /V. T. Hiep ,
l\ K. Loc, 1 .. V. Averyanov, G. E. Schulz & S.
Bodine DKH 6091 (holotype, ILN; isotypes.
HN, k, LE. MO). Figure 1.

Arbor 10— 15-metralis, sempervirens, monoica; cortex
librosus. Kami lunges, horizontaliter dispositi; ramulorum
frondes dimorphae foliis adultis et juvenilibus praeditae.
Folia adulta decussata, inconspicue glandulifera: ea ra¬
mulorum (sub)ultimorum 1.5—3 mm longa, 1—1.3 mm lata,
dimorpha: folia dorsi-ventralia (anglice “the facials”) ad-
pressa, anguste angulato-ovata vel rhombiea, in parte dis-
tali carinata, margine minute denticulata praeter apicem
versus integra. apiee acuto; folia lateralia moderate lon-
giora, conduplicata. basin versus deeurrentia, recta vel
falcata, margine minute denticulata praeter apicem versus
integra. apice libero acuto vel pungenti; stomata foliorum

adultorum inconspicua plerumque adaxialia, pauca in par¬
te proximali superficiei abaxialiae. Folia juvenilia in ver-
licillis quadrifolii disposita, basin versus deeurrentia, pa-
lentia, 15—20 mm longa, 1.5—2 mm lata, monomorphia,
linearia, eglandulifera, margine integra, apice acuto; sto¬
mata foliorum juvenilium in vittis 2 in superfieie abaxiali
solum disposita. Amenta maseula solitaria, in ramulis lolia
adulta gerentibus terminalia, 2.5— 3.5 mm longa, 2-2.5
mm lata: microsporophylla 10-12, peltata; microsporangia
2(-3), abaxialia. Strobili feminei solitares, in ramulis folia
adulta gerentibus terminales vel subterminales, post duos
annos maturi, subglobosi, post dehiscentia 9—1 1 mm lon-
gi. 10-12 mm lati; braeteae-squamae 2( — 3) paribus, op-
positae, decussatae, pare supero majore connato vel re-
cluso valvato vel subpeltato, distaliter latiores, rugosae et
recurvo-umbonatae. Semina ca. K— 9. ovoidea vel irregu-
laria, 4.5—6 mm longa, 4—5 mm lata, tesla in alas 0.5—1
mm latas expansa.

Small to medium-sized tree to 10—15 m; trunk
monopodial, terete, up to 50 cm diant.; bark smooth
and thin on branches, purplish to red-brown, ex¬
foliating in thin flakes and strips, on the trunk of
larger trees becoming soft and fibrous, brown to
gray-brown, exfoliating in numerous thin strips.
Branches long, spreading ± horizontally; foliage
branches numerous, spreading mostly in plagiotro-
pic overlapping sprays or slightly drooping, forming
a pyramidal crown in young trees but a spreading,
irregular or flat-topped crown in old trees. Foliage
in mature trees predominantly with mature leaves,
also with juvenile leaves, often also with transition¬
al leaves. Foliage sprays with juvenile leaves bushy,
sparsely branched, ultimate branchlets 20—50 mm
long, not flattened. Foliage sprays with mature
leaves flattened, with rounded outline; leading fo¬
liage branches quadrangular to terete, with ca. 4
orders of branching, still covered with green leaves
in 3rd to 4th year of growth; subultimate branchlets
pinnate, ultimate branchlets of unequal and irreg¬
ular length and spreading at 30°— 45°, 5-20 X 1.5-
3 mm. distinctly flattened. Juvenile leaves in whorls
of four, decurrent, monomorphic, the distal part
spreading at nearly 90°, the proximal decurrent part
4-5 mm long, the distal free parts 15—20 X 1.5-2
mm. linear, margins entire, tapering to a fine point.
Stomata in two whitish bands on the abaxial side
only, in each band ca. 6—7 irregular rows, oriented
parallel to the leaf axis. Transitional leaves similar
to mature leaves but longer (5-7 mm), lanceolate,
the laterals spreading at 45°. Mature leaves decus¬
sate. short deeurrent, imbricate, dimorphic, on
(sub)ultimate branchlets 1.5—3 X 1—1.3 mm (the
laterals slightly longer than the facials); the facials
narrowly ovate-rhombic, keeled, ± appressed, mar¬
gins minutely denticulate to entire toward the acute
or acuminate and free apex; the laterals condupli-
eate, the proximal part decurrent, the distal part

Volume 12, Number 2
2002

Farjon et al.

New Vietnamese Genus

181

MISSOURI

BOTANICAL GARDEN
HERBARIUM

N? 5194647

Ha Gi ang

Ouan Ba District, Bat Dai Son
Municipality; Bat Dai Son Provincial
Protected Area (2000); on limestone
ridge of degraded forest just
northwest of Bat Dai Son border
station; collections in forest above
cu 1 t i vat l ons . (Forest possibly burned
ca. 25 years ago or so, only small
trees and few fire tolerant herbs).

1 3°09 ' 1 3 "N i04°59'36'lE 1064 m

D.K. Harder, N.T. Hiep, P.K. Loc , L.V, ft
ver yanov, G.E. 5chat2 & S. Bcdine 6091

Supported by National Science Foundation
and National Geographic Society
MISSOURI BOTANICAL GARDEN HERBARIUM <h0>

MISSOURI BOIANICA! GARDEN HERBARTIJM (Mb;

Figure 1. Photograph of the holotype oi Xanthocyparis vietnamensis Farjon & Hiep. D. k. Harder, N. T. Hiep, R K.
Loc, L. V. Averyanov, G. L. Schatz & S. Bodine Dkll 6091(HN).

182

Novon

spreading free from the leaf above at ea. 30°,
straight or falcate, margins minutely dentic ulate ex¬
cept toward the acute or pungent apex. Stomata on
mature leaves inconspicuous, mostly adaxial, a few
scattered on the proximal abaxial faces, covered
with a layer of euticular wax. Glands inconspicu¬
ous, in a depression below the keeled distal part of
some of the facials. Pollen cones 2.5— 3.5 X 2—2.5
mm. oval-terete; microsporophylls 10 to 12, ca. I
X 1 mm, with erose-denticulate margins and mu-
cron ate apex, green turning yellow-brown; each
bearing abaxially two large, subglobose yellow mi¬
crosporangia containing spherical pollen. Seed
cones sparse but sometimes grouped with 2 or 3
together at the outer margins or nearer the base of
foliage sprays with mature leaves, initially consist¬
ing of the 2 upper pairs of green leaves (bracts)
with axillary ovules. Mature cones developing in
two years, green, turning dark or dull brown, subgl¬
obose, 9—1 l X 10—12 mm when open, some ± per¬
sistent after seed dispersal. Bract-scale complexes
in 2 (sometimes 3) decussate pairs in normally de¬
veloped cones (irregular or underdeveloped cones
are found), valvate to subpeltate (the third pair if
present ± peltate and 4—5-angled), the lower pair
oblong, all widest distally, with rounded but irreg¬
ular upper margin; outer surface smooth, becoming
rugose or radiately furrowed from a prominent, 1—
2.5 mm long umbo (including the bract apex); inner
surface red-brown marked proximally with white or
gray seed scars; a small columella present or absent
at the shoot apex. Ovules I to 3 per fertile bract
(upper bracts in 6-scaled cones sterile); seeds max.
8 or 9 per cone, ovoid or irregular, flattened (1.5—
2 mm thick), 4.5—6 X 4-5 mm including two lateral
wings, body of seed ± pustulate, light brown or red-
brown, with white hilum at base and micropylar
beak often persistent at the apex; seed wings 0.5—
1 mm wide, thinly membranous, lighter colored.
Seedlings not seen.

Distribution. Vietnam: North Vietnam, Ha
Giang Province, very locally in the Bat Dai Son
mountain system near the Chinese border; altitu¬
dinal range 1060—1180 m.

Ecology. In mixed angiosperm-eonifer cloud
forest with the conifers Amentotaxus argotaenia
(1 lance) Pilger, Nageia wallichiana (C. Presl) kun-
tze, Pseudotsuga sinensis Dode var. brevifolia (W. C.
Cheng & l„ k. Fu) Farjon & Silba, Podocarpus
pilgeri Foxworthy, and Tnxus chinensis (Pilger) Reh-
der. Dominant among angiosperms are species of
Acer. Carpinus, Lithocarpus, Quercus , and Him us:
frequent are Pistacea uieinmannifolia J. Poisson ex
Franchet and Platycarya strobilacea Siebold &

Zuccarini. In a second stratum under the ca. 20 m
tall canopy species of Elaeocarpus, Eriobotrya, Sor-
bus. Schefflera, and many others frequently occur.
Shrubs and herbs abound; among the latter are nu¬
merous species of Orchidaceae, terrestrial as well
as epiphytic, sometimes determining the aspect of
the ground cover vegetation. Ferns and especially
bryophytes arc similarly abundant both as litho-
plivtes and as epiphytes. The limestone ridges on
which Xanthocyparis occurs are extremely eroded,
composed of resistant, marble-like rock outcrops
interspersed with thin soil pockets. The climate is
subtropical but damp and wet much of the year.

Conservation. Newly discovered Xanthocypar¬
is vietnamensis is restricted to a few localities in
close proximity, mostly now in inaccessible sites
on steep limestone ridges. Logging has increased
in recent years and is estimated to have caused
serious decline in numbers of larger, well-growing
trees. This practice may have had negative effects
on genetic diversity. Regeneration is poor due to
heavy competition in remaining populations. This
species is Critically Endangered under the IUCN
Red List Categories Version 3.1 (IUCN. 2001): CR
(B2a-c).

Uses. This species produces fine, yellow-brown,
very hard, fragrant timber. The superb quality of
the wood, in conjunction with the widespread de¬
sirability of cupressaceous wood in traditional uses
of many kinds in eastern Asia, combined with slow
growth, has made it a highly prized timber. Due to
lack of transport facilities and other factors, most
of the timber has been traded locally.

Paratypes. \ IETNAM. Ha Citing: Quan Ba, Bat Dai
Son. Bat Dai Son I’rov ineial Protected Area. 10 Feb. 2001 .
I ). K. Harder et td. HKH 6 090 (IIN. MO, LE), 12 Feb.
2001. I). K. Harder et al. DKH 6 224 (HN, k, MO. I.E);
Can Ty, Sing Xuoi Ho, 12 Oct. 1999, Nguyen Tien Hiep,
I, L Averyanov & P. ./. Cribb NTH .1594 (HN, MO. LE,
k).

In October 1999 a conifer was found in North
Vietnam (A. T. Hiep et al. NTH 3594) that appeared
to have morphological characters suggesting a re¬
lationship with species in the cupressoid clade of
Cupressaceae s.l. (Gadek et al.. 2000). The speci¬
men compared in particular with Chamaecyparis
and Cupressus . yet showed some traits not found in
any known species in these or related genera, im¬
mediately suggesting that a new taxon could have
been found. The only known species in that group
that are certain to be indigenous in this part of Asia
(including most of Myanmar, North Laos, South
1111111011. Guangxi, Guangdong, and Hainan |Cliina|)
are Calocedrus rnacrolepis kurz and Fokienia liodg-
msii (Dunn) A. Henry N H. Thomas. The new co-

Volume 12, Number 2
2002

Farjon et al.

New Vietnamese Genus

183

Figure 2. — A. Tree of Xanthocyparis vietnamensis al lial Dai Son (photograph by L. Averyanov). — B. Branch of X.

vietnamensis with two different foliage types (photograph hy D. K. Harder). — C. Branchlet of X. vietnamensis with
mature seed cones (photograph by D. K. Harder). — D. Branchlet of A. vietnamensis with pollen cones (photograph by
D. K. Harder).

nifer (Figs. 2, 4) appeared to occur very locally on
limestone karst ridges known to have yielded other
narrow endemics, some of which have turned out
to be new species, e.g., orchids. Such habitats are
also frequently the refuges of relict conifer taxa,
presumably because these conifers have adapted to
the poor growing conditions associated with karst
where competitor broad-leaf tree species could not
follow. The most conspicuous feature of this new
conifer taxon is the occurrence of juvenile leaves.

transitional leaves, and mature leaves in the foliage
of crowns of mature trees (Figs. 2B, 4). These ju¬
venile, monomorphic leaves 15 X 1.8 mm can al¬
ternate on a branch with mature dimorphic leaves
2.5-3 X 1.2- 1.3 mm (I). K. Harder et al. 6224).
This trait is reminiscent of the foliage in mature
trees of Callitris macleyana F. Mueller; however,
unlike that species, Xanthocyparis vietnamensis
does not produce cones on branchlets with these
juvenile leaves. In Widdringtonia nodiflora (L.) E.

184

Novon

Figure 3. Branchlet of X. nootkatensis with mature seed cones (photograph by A. Farjon).

Powrie, coppiced (fire-damaged) plants produce
similar juvenile leaves usually on basal reiterated
branches (Pauw Linder, 1997). A mixture of nee-
dle-like and scale-like leaves is common in Juni-
perus chinensis Roxburgh, where the needle leaves
represent the juvenile stage because such leaves
are the first to appear after the cotyledons. Indeed,
in all species of Cupressaceae s.l. (Farjon, 1998,
2001) this transition from juvenile (via transitional
forms) to mature leaves takes place in the seed¬
lings, often accompanied by a change in phyllotax-
is. Normally, in most cupressoid taxa. juvenile
leaves do not reoccur; when they are present these
branches are often the result of reiteration. This
seems to be the case in some specimens of X. viel-
namensis (e.g., N. T. Hiep et al. NTH 3594), but in
others (e.g.. I). K. Harder et al. DKH 6091 holotype.
D. K. Harder et al. DKH 6224 ) the branching pat¬
tern of both foliage types shows a normal alterna¬
tion not suggesting reiteration. In the known spe¬
cies of Chamaecyparis and Cupressus juvenile
leaves are restricted to the seedling stage. However,
juvenile leaf characters have been retained in some
eullivars of Chamaecyparis. It is possibly a neotenic
trait controlled by certain genes that can be
“switched on or ofF' at different stages of growth in
several taxa. By itsell, this would be a somewhat
doubtful character to indicate the existence of a

distinct species; however, its coexistence with ma¬
ture foliage throughout the life history of the tree
is rare in conifers.

Careful comparison of the morphology of the newr
discovery in Vietnam with that of other, more or
less similar, species in Cupressaceae leads to Cha¬
maecyparis nootkatensis (Fig. 3) as the most similar
species. This taxon has very similar seed cones
with 4(to 6) bract-scale complexes (“—12" is cer¬
tainly an error for C. nootkatensis in Taylor & Szik-
lai, 1976), which are (|uite distinct from both its
congeners and from those of Cupressus, with which
Chamaecyparis has been united from time to time
(see, e.g., Camus, 1914, for a monographic treat¬
ment). There has been debate concerning the
placement of C. nootkatensis in either genus, with
some authors arguing for inclusion in Cupressus
based on characters of the ovuliferous cones
(Frankis, 1993; Jagel & Stiitzel. 2001). Recent cla-
< list ie evidence based on molecular data (mat K
gene) gives only weak support for its inclusion in
Cupressus (Gadek et al., 2000); similar evidence us¬
ing a combined data set (mat K 4- noil-molecular
data) does not and places it as a sister group to
Cupressus + Juniperus with stronger bootstrap sup¬
port (Gadek et al., 2000). Inclusion of the new Viet¬
namese species in a phylogenetic analysis of Cu¬
pressaceae s.l. based on morphological data (a full

Volume 12, Number 2
2002

Farjon et al.

New Vietnamese Genus

185

account of which will be published later) resulted
in a separate clade for C. nootkatensis + X. viet-
namensis within a major clade distinct from both
Chamaecyparis and Cupressus (Fig. 5). Sequencing
of DNA of X. vietnamensis has yet to be undertaken.

Apart from the markedly different ages of plants
between C. nootkatensis and X. vietnamensis in
which juvenile leaves still occur, the leaf morphol¬
ogy of both is very similar, both of juvenile, tran¬
sitional and mature (Figs. 2D, 3, 4) leaves. In both
species the mature leaves are markedly dimorphic
(differently shaped facials and laterals) on plagio-
tropic foliage branchlets. This is common in all Cu-
pressaceae with such branching systems, causing a
bilateral flattening of the leaf-covered branchlets.
Growth of branches profoundly determines leaf
shape in Cupressaceae (Daguillon, 1899; Rouane,
1973; Offler, 1984). Unlike other taxa with this
marked leaf dimorphism (e.g., Chamaecyparis s.
str., Calocedrus, Fokienia, Platycladus, Thuja, Thu-
jopsis), in which the distribution of abaxial stomata
is asymmetrical, i.e., largely confined to the phys¬
iological underside of the lateral leaves and the ab¬
axial side of facials on that side of the branchlet,
both Chamaecyparis nootkatensis and Xanthocypar-
is vietnamensis have only a weakly developed sto-
matal asymmetry. In this respect they resemble cer¬
tain species of Asian Cupressus with (weakly)
dimorphic adult leaves on pendulous branchlets (C.
cashmeriana Royle ex Carriere, C. funebris End-
licher). Leaf dimorphism in Cupressaceae is strong¬
ly correlated (i.e., adaptive) with moist climates
(e.g., Chamaecyparis , Fokienia, Thuja ); monomor¬
phism of mature leaves resulting in quadrangular
to terete branchlets is correlated with (seasonal)
aridity (e.g., Cupressus arizonica, Juniperus sect.
Sabina). Glands on the facial mature leaves of X.
vietnamensis are less conspicuous than those of
Chamaecyparis nootkatensis; neither normally pro¬
duce droplets of resin.

Pollen cones of both species are very similar (in
C. nootkatensis slightly larger) with 2(or 3) relative¬
ly large microsporangia; in Chamaecyparis s. str.
and especially Cupressus the latter are more nu¬
merous (3 to 6) and relatively smaller (Figs. 2D, 4).
Seed cones of Xanthocyparis vietnamensis (Fig. 2C)
and Chamaecyparis nootkatensis (Fig. 3) are also
similar; the most striking feature is the predomi¬
nance of only two decussate pairs of bract-scale
complexes in the mature cones. In both species an
occasional third distal pair develops (or sometimes
only one of these); although the sampling of X. viet¬
namensis studied is still somewhat limited (ca. 30
cones seen) it seems that this is somewhat more
frequent in this species (Figs. 2C, 4). In both taxa

a small columella develops at the shoot apex of 4-
scaled cones, sometimes consisting of two parts. As
in all Cupressaceae s.l. (inch Taxodiaceae) the ovu-
liferous cone starts with ovule development borne
on, subtended by, or surrounded by leaves (bracts);
the scale forms by secondary (intercalary) growth.
In both species this growth is more pronounced on
the adaxial side of the bracts and more rapid in the
second pair of the bract-scale complexes, resulting
in semi-valvate cone scales that remain largely
fused proximally. The only other taxon in the eu-
pressoid clade (Gadek et ah, 2000) with somewhat
similar cone development is Tetraclinis; this type is
more fully developed in the callitroid clade (Gadek
et ah. 2000) with which both (ossil (e.g.. Engelhardt
& Kinkelin, 1908) and Recent species (Ei, 1953)
of Tetraclinis have erroneously been identified.
Whereas in Tetraclinis the number of bract-scale
complexes (2 pairs) seems primary in origin, in
Chamaecyparis nootkatensis and Xanthocyparis
vietnamensis it is almost certainly reduced, as evi¬
denced by the occasional third pair that is much
smaller and never associated with the ovules. Poor¬
ly developed, smaller cones of Cupressus lusitanica
Miller sometimes have only 2 pairs (instead of 3 to
4, rarely 5 pairs) but they are more peltate. The
ontogeny of the bract-scale complexes and the
placement of ovules of X. vietnamensis and Cha¬
maecyparis nootkatensis are likely to be similar but
were only studied in detail for the latter species
(Jagel & Stiitzel, 2001). Seedling development and
seedling characters of X. vietnamensis remain to be
studied.

Considering the evidence based on morphology,
it is very likely that there is a closer relationship
of Xanthocyparis vietnamensis w ith Chamaecyparis
nootkatensis than with any other Recent taxon in
Cupressaceae. We appear to have found a very in¬
teresting relict taxon on the western margin of the
Pacific that belongs to a lineage distinct from both
Chamaecyparis and Cupressus and is possibly more
ancient than either of these. A comparison with the
Late Cretaceous taxon Mesocyparis Mclver & Bas¬
inger, which has been found in northwestern North
America and northeastern Asia (Mclver & Basin¬
ger, 1987; Mclver & Aulenback, 1994) is of inter¬
est. In this fossil genus short dimorphic leaves ap¬
pear on opposite (pen)ultimate lateral branchlets,
and long dimorphic leaves follow a series of short
leaves on leading shoots. Although the opposite
branching pattern contrasts with the alternate
branching in Recent cupressoid genera, the alter¬
nation of two leaf shapes is also found in Xantho¬
cyparis vietnamensis, but with monomorphic (juve¬
nile) longer leaves. The seed cones of Mesocyparis

186

Novon

Figure 4. Xanthocyparis vietnamensis Farjon & fliep. — A. Branchlet with scale leaves and seed cones. — B. Branchlet
with scale leaves and needle leaves. — C. Branchlet with scale leaves. — 1). Branchlet with needle leaves. — h. Detail
of upper and lower side of needle leaf. — F. Branchlet with pollen cone. — (>. Microsporophyll with two microsporangia.
— H. I. Seed cones, closed and open. — J. Seeds. [A. C, F— J = Harder el al. 6091 (HN. k); B. I). F = Harder el al.
6224 (k). Illustration by Aljos Farjon.

Volume 12, Number 2
2002

Farjon et al.

New Vietnamese Genus

187

Callitris drummondii
Callitris oblonga
Callitris macleana
Widdringtonia nodiflora
Actinostrobus pyramidalis
Neocallitropsis pancheri
Fitzroya cupressoides
Diselma archeri
Austrocedrus chilensis
Libocedrus plumosa
Papuacedrus papuana
Pilgerodendron uviferum
Chamaecyparis nootkatensis
Xanthocyparis vietnamensis
Microbiota decussata
Platycladus orientalis
Tetraclinis articulata
Calocedrus decurrens
Cupressus bakeri
Juniperus californica
Juniperus drupacea
Juniperus oxycedrus
Juniperus indica
Juniperus recurva
Juniperus phoenicea
Juniperus virginiana
Cupressus funebris
Chamaecyparis lawsoniana
Fokienia hodginsii
Thuja plicata
Thujopsis dolobrata
Metasequoia glyptostroboides
Cryptomeria japonica
Glyptostrobus pensilis
Taxodium distichum

, — Sequoia semprevirens
i— Sequoiadendron giganteum
r— Cunninghamia lanceolata
"L_ Taiwania cryptomerioides

— Athrotaxis laxifolia

— Sciadopitys verticillata

— Pinus sylvestris

figure 5. Stiiel consensus of 144 equally parsimonious trees <>1 523 steps found from a heuristic search of morpho¬
logical and anatomical data (53 characters) in Cupressaceae s.l. and Sciadopitys verticillata , using Pinus sylvestris as
outgroup. Cl — 0.2428; RI = 0.3077; RC = 0.0747. |A fidl account of the characters and the data set will he given
by Farjon in his monograph of Cupressaceae.]

umbonata Mclver & Basinger are similar to those
of Chamaecyparis nootkatensis and X. vietnamensis
with two paired bract-scale complexes. However,
the position of seed cones on the foliage branches
of Mesocyparis umbonata is quite different espe¬
cially from C. nootkatensis. They are not terminal
on foliage sprays but appear lower on foliage
shoots, on opposite, very short branehlets axillary
to long leaves. A lower position does also occur in
X. vietnamensis, but these cones are not in decus¬
sate pairs. Early developmental stages observed in
fossil seed cone specimens of Mesocyparis umbon¬
ata are essentially similar to those in C. nootkaten¬
sis (Mclver & Aulenbaek, 1994) but in the fossil
the cones remain notably smaller, i.e., development
after pollination of the ovules lags behind. In both
C. nootkatensis and X. vietnamensis seed cone

growth after pollination alters the relative sizes of
bract and scale, considerably influencing the size
and shape of these bract-scale complexes at ma¬
turity. The seeds of fossil Mesocyparis and of Cha¬
maecyparis and Xanthocyparis are very similar. An¬
other bate Cretaceous fossil, Chamaecyparis
corpulenta (Bell) Mclver (Mclver, 1994) from Van¬
couver Island. Canada, has leaves more similar to
other Recent species in Chamaecyparis . but again
very small seed cones with four scales. This low
number of bract-scale complexes is apparently
primitive, as it occurs in all earliest (Cretaceous)
members of Cupressaceae thus far known (Mclver.
1994).

We conclude from the above evidence that Cha¬
maecyparis nootkatensis should be placed with
Xanthocyparis vietnamensis in a separate genus.

188

Novon

Xanthocyparis lias retained several characters or
traits that were evident in similar cupressoid mem¬
bers of Cupressaceae s.l. in the bate Cretaceous,
Init which have been lost or replaced by other fea¬
tures in more advanced members ol Chamaecyparis
and Cupressus during the Tertiary.

fhe following new combination has to be made
as a result of the new taxonomy here proposed:

2. Xanthocyparis noolkatensis (I). Don) Farjon
& Harder, comb. nov. Basionym: Cupressus
nootkatensis I). Don. in Lambert, Deser. Linus
2: 18. 1824. Chamaecyparis nootkatensis (I).
Don) Spaeh, Hist. Nat. Veg. Phan. II: 388.
1841. Cupressus nutkatensis Hooker. 11. Bor.
Amer. 2 (10): 165. 1838. Chamaecyparis nut-
kaensis Lindley & Gordon. J. Holt. Soc. Lon¬
don 5: 207. 1850. TYPE: Canada. British Co¬
lumbia: Hecate Strait. Banks Island

[“northwest coast of North America behind
Bank’s Island"], 1787. A. Menzies s.n. (holo-
type, not seen; isotypes, K. MO).

Thuja excelsa Bongard, Mem. Acad. Imp. Sei. Saint-Pe-
tersbourg, ser. (>. Sei. Math. 2: 164. 1832. Cupressus
americana Trautvetter, PI. I mag. Deser. FI. Russ. I
(1/2): 12. I. 7. 1844. TYPE: U.S.A. Alaska: Baranof
Island. Sitka. “Dr. Mertens” s.n. (no date) (holotype.
not seen; isotype, MO).

Thujopsis borealis hort. ex Carrifere, Traite Gen. Conif.:

I 13. 1855. TYPE: Unknown, based on cultivated
plant.

The species Cupressus macrocarpa Hartweg ex
Gordon is considered to be the male parent ol the
generic hybrid XCupressocyparis leylandii (A. B.
Jackson & Dallimore) Dallimore & A. B. Jackson;
the female parent, from which hybrid seeds were
collected, is thought to be Chamaecyparis noolka¬
tensis (I). Don) Spaeh (Jackson cK Dallimore, 1926).
Reversed parentage was also reported to have oc¬
curred (Jackson & Dallimore, 1926), but both the
protologue and the type specimens at k indicate
that this hybrid was lust raised from seed obtained
in 1888 from C. noolkatensis. Two further nothos-
peeies with male parentage from C. noolkatensis.
but involving as female parents Cupressus arizomea
Greene var. glabra (Sudworth) Little and Cupressus
lusitanica Miller, were describee I by Mitchell
(1970). Under tin* rules of the Code (Art. 116.2;
Greuter et al., 2000) the name of a nothogenerie
hybrid is to be combined from the first part or the
whole of one and the last part or the whole of the
other parent. The reclassification of Chamaecyparis
noolkatensis in a new genus Xanthocyparis there¬
fore requires a name change of the hybrid genus ol
which it is one ol the parents; after ample delib¬

eration it has been considered best practice to ef¬
fect as minimal a change as possible. The neces¬
sary new combinations are given below.

XCuproeyparis Farjon, nom. nothogen. nov.

Cupressocyparis Dallimore & A. B. Jackson, Forestry
1 I: 3. 1937; Boy. Bot. Gard. kew Hand-list Conif., ed. 4:
37. 1938.

3. X Cuprocyparis leylandii (A. B. Jackson &

Dallimore) Farjon, comb. nov. Basionym: Cu¬
pressus leylandii A. B. Jackson Dallimore,
kew Bull. 1926: 111. 1926. XCupressocyparis
leylandii (A. B. Jackson & Dallimore) Dalli¬
more & A. B. Jackson, Forestry 11: 3. 1937;
Roy. Bot. Gard. kew Hand-list Conif., ed. 4:
37. 1938. TY PE: England. Northumberland:
Haggerston Castle (cultivated), 26 Nov. 1925,
hi. ./. Leyland s.n. ( specimen marked I). with
cones) (leetotype, designated here, k).

Hybrid formula: 8 Cupressus macrocarpa Hartweg
ex Gordon X 9 Xanthocyparis nootkatensis (D.
Don) Farjon & Harder | Cupressus macrocarpa
Hartweg ex Gordon X Cupressus nootkatensis
I). Don; Cupressus macrocarpa Hartweg ex
Gordon X Chamaecyparis nootkatensis (I).
Don) S| )ach]

4. XCuproeyparis notahilis (A. F. Mitchell) Far¬

jon. comb. nov. Basionym: XCupressocyparis
notahilis A. F. Mitchell, J. Roy. Hort. Soc. 95
(10): 453. 1970. TYPF1: England. Hampshire:
Forest Research Station. Alice Holt Lodge, 31
July 1963, A. F. Mitchell s.n. (holotype, k).
Hybrid formula: 9 Cupressus arizomea Greene var.
glabra (Sudworth) Little X 8 Xanthocyparis
nootkatensis (I). Don) Farjon & Harder.

5. XCuproeyparis ovensii (A. F. Mitchell) Far¬

jon. comb. nov. Basionym: XCupressocyparis
ovensii A. F’. Mitchell, J. Roy. Hort. Soc.
95(10): 454. 1970. TYPE: England. Hamp¬
shire: Forest Research Station, Alice Holt
Lodge, 1970, A. F. Mitchell s.n. (holotype, k).
Hybrid formula: 9 Cupressus lusitanica Miller X
8 Xanthocyparis nootkatensis (I). Don) Farjon
& Harder.

Acknowledgments. The authors thank Marc
Goode of kew for his kind help with tin- Latin di¬
agnosis and description, and two anonymous re¬
viewers for their constructive remarks. Fieldwork
supporting this discovery was provided by the Na¬
tional Science Foundation ( DF.B-98 < 023 1 to the
Missouri Botanical Garden. D. Harder PI.) and the
National Geographic Society (grant # 6733-00,
"Botanical inventory of unexplored areas in Viet

Volume 12, Number 2
2002

Farjon et al.

New Vietnamese Genus

189

Nam: the north” to DKH). This support is gratefully

acknowledged.

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Nomenclatural Clarification in Aristea Section Racemosae
(Iridaceae) in the Cape Flora of South Africa

Peter Goldblatt

It. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299,
St. I.ouis, Missouri 63166-0299, U.S.A. peter.goldblatt@mobot.org

John C. Manning

National Botanical Institute, P. Bag XT. Claremont 7735, South Africa.
manning@nbict.nbi.ac.za

Roy E. Gere an

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

roy.gereau@mobot.org

Abstract. We demonstrate that the correct
names for three species of section Racemosae of the
Afro-Magadagascan genus Aristea are: Aristea cap-
itata (L.) ker Gawler, A. braeteata Persoon, and A.
bakeri Klatt. A neotype is selected for Gladiolus
capitatus L. (1753). t tie basionym of A. capitata (L.)
ker Gawler (1802). Aristea major Andrews (1801)
is placed in taxonomic synonymy with A. capitata.
The typification and identity of Ixia thyrsiflora I).
Delaroehe (1766), basionvm of Aristea thyrsiflora
(I). Delaroehe) N. K. Brown (1929), often regarded
as conspecific with A. major, remain in question;
however, due to priority of publication of the bas-
ionyms A. thyrsiflora presents no threat to the sta¬
bility of A. capitata. Weimarck effectively lectotvp-
ified the superfluous and illegitimate name Moraea
caerulea Thunberg (1787). basionym of Aristea ca-
erulea (Thunberg) Vahl (1805). Aristea braeteata
Persoon (1805) is accepted as a replacement name
for M. caerulea, and is the oldest legitimate name
available for this species. Aristea monticola Gold¬
blatt (1971) was also published as a replacement
name for M. caerulea. and thus falls into synonymy
with A. braeteata. Aristea bakeri klatt (1894) was
published as a replacement name for the illegiti¬
mate later homonym A. paniculata Baker (1892).
Aristea macrocarpa G. J. Lewis (1940) and A. con-
fusa Goldblatt (1970) are placed in taxonomic syn¬
onymy with A. bakeri.

Key words: Aristea, Cape f lora. Iridaceae.
South Africa.

The identity of Gladiolus capitatus L. (1753), the
basionym of Aristea capitata (L.) Ker Gawler, lias
always been uncertain because no type has ever

been found. Early botanists dealing with the south¬
ern African Iridaceae, including Linnaeus's con¬
temporary, C. P. Thunberg, were unanimous that the
plant represented a robust species of a group that
is now recognized as the Afro- Madagascan genus
Aristea Aiton (ca. 50 species). The name Gladiolus
capitatus has long been associated with the species
currently known as A. major Andrews (1801) or A.
thyrsiflora (I). Delaroehe) N. E. Brown (1929), both
of which represent the same plant (ker Gawler,
1802; Baker. 1892. 1896), or with two other spe¬
cies, A. macrocarpa G. J. Lewis or A. confusa Gold¬
blatt. The last two names were published for plants
known at the time as A. capitata, which the re¬
spective authors rejected because this name lacked
a type anil could not be reliably identified to spe¬
cies (Lewis in Weimarck, 1940; Goldblatt. 1671).
Confusion about A. capitata was such that Wei¬
marck (1940) recognized A. capitata despite in¬
cluding its intended replacement name A. macro¬
carpa in th<‘ same account. In her account of Aristea
for the Flora of the Gape Peninsula, Lewis (1950c)
also regarded ,4. capitata and A. macrocarpa as sep¬
arate species, despite her earlier treatment (Lewis
in Weimarck, 1940). To add to the confusion, Gold¬
blatt (1971) published yet another name, A. con¬
fusa, for the plants called 4. capitata by Lewis

(1950c).

These species are fairly closely related and were
all included by Weimarck (1940) in Aristea sect.
Racemosae, a taxon defined by woody capsules with
three broad, radial wings that develop from the loc¬
ales of the ovary with each locule containing be¬
tween two and twelve radially compressed, lamel¬
late seeds (seed number depending on the species)

Noyon 12: 190-195. 2002.

Volume 12, Number 2
2002

Goldblatt et al.

Aristea Section Racemosae

191

(Goldblatt & Manning, 1997). Most members of the
section are robust plants, often 1 m or more in
height, with tough fibrous leaves, and a highly
branched compound inflorescence (a synfloresc-
ence of binate rhipidia), but a few species are rel¬
atively small plants. Goldblatt and Le Thomas
(1997) maintained section Racemosae in a phylo¬
genetic analysis of Aristea that included several
pollen characters. They showed that all the species
of the section examined have unspecialized pollen
grains with a single, smooth aperture and reticulate
exine with small lumina. Goldblatt and Le Thomas
included section Racemosae in Aristea subg. Aris¬
tea, which has two more sections, Aristea and Sin¬
gulars. These two sections comprise small plants
that resemble section Racemosae in their winged
capsules and lamellate seeds (doughnut-shaped in
Aristea singularis of sect. Singulars ), but they have
a dichotomously branched flowering stem, and in
section Aristea unusually large pollen grains with
complex, trisulcate to spiral, confluent apertures
and reticulate exine with large lumina.

Aristea capitata was one of the first two species
of the genus to be described, the other being A.
africana (L.) Hoffmannsegg: basionym Ixia africana
(Linnaeus, 1753). The latter is the type species of
Aristea, formally named in 1789 in Aiton’s Hortus
Kewensis, actually as we now know by the Swedish
botanist Daniel Garl Solander, who was never for¬
mally credited with authorship of the genus (Ker
Gawler. 1802: Krok, 1925). At the time of its de¬
scription, Aristea included only A. cyanea [Solander
in] Aiton, an illegitimate synonym of Ixia africana.
The British petaloid monocot expert John Ker Gavv-
ler (1802) transferred Gladiolus capitatus to Aristea
and included Ixia thyrsiflora (Delaroche, 1766). Ar¬
istea major (Andrews, 1801), and Moraea caerulea
Thunberg (now ,4. bracteata Persoon) as synonyms,
indicating a broader interpretation of the species
than we now believe is correct. In order to establish
a stable taxonomy for Aristea, we have the choice
of formally proposing the rejection of Gladiolus cap¬
itatus, an undertaking not certain of favorable out¬
come, or of choosing a suitable neotype for the spe¬
cies. We have decided on the latter alternative.

N KOTY I’l FICATION OK GLADIOLUS CAPITATUS

As outlined above, the identity of Gladiolus cap¬
itatus, the earliest species referable to Aristea sect.
Racemosae, has always been uncertain because of
the absence of a type. The name has consistently
been used for species of Aristea sect. Racemosae
and has been applied either to the plant currently
known as A. major or to another species, variously

called A. bakeri, A. macrocarpa, or A. confusa. The
most likely species that Linnaeus had in mind
when he described G. capitatus is the species later
called Aristea major (Goldblatt & Manning, 2000).
which has also been called A. thyrsiflora (Brown,
1929: Weimarck. 1940). The protologue of G. cap¬
itatus (Linnaeus, 1753: 37) refers to a large plant
with blue flowers (j)lanta maxima, floribus caeru-
leis ), a branched stem ( caule rarnoso), a capitate
flowering stem ( capital is pedunculatis), and a tu¬
berous rootstock (radice tuberosa). Aristea major
closely matches this description, particularly re¬
garding the crowded and relatively compact synflo-
rescence of numerous, blue flowers borne at the
apex of an unbranched flowering stem (or pedun¬
cle). usually 1-1.5 m high (Goldblatt & Manning,
2000). The distinctive, dense synflorescenee may
well have prompted the specific epithet capitatus,
meaning head-like. Aristea major has a range ex¬
tending from the Cape Peninsula north to the Pi-
ketberg and east through the coastal Cape moun¬
tains to George. It is likely that a plant from the
Cape Peninsula or nearby was one that came to
Linnaeus’s attention before 1753, for exploration of
the then Dutch colony had not extended signifi¬
cantly into the interior at that time. Thus, from both
morphological and geographical points of view the
selection of a specimen currently assignable to A.
major seems the ideal choice as a neotype for A.
capitata, and A. major thus falls into synonymy.

This selection will stabilize the taxonomy of Ar¬
istea sect. Racemosae and has the added benefit of
circumventing a nomenclatural controversy about
the identity of Ixia thyrsiflora D. Delaroche (1766),
which is most likely a species of Aristea. N. E.
Brown (1929) maintained that a specimen in the
Burman Collection at Geneva was the type of I.
thyrsiflora, anti that it represented A. major, making
this an earlier name for that species. Goldblatt and
Barnard (1970) disagreed that the Geneva speci¬
men was the type, citing discrepancies in the pro-
tologue between the description, Delaroche’s man¬
uscript notes (in the Leiden Herbarium), and the
Burman specimen. We are not prepared to clarify
the typification of I. thyrsiflora in this paper; how¬
ever, due to the priority of publication the name
Aristea thyrsiflora presents no threat to the stability
of A. capitata.

Aristea capitata (L.) Ker Gawler. Curtis’s Bot.
Mag. 17: pi. 605. 1802. Gladiolus capitatus L.,
Sp. pi. 37. 1753. TV PE: South Africa. Western
Cape: foot of Du Toit’s Kloof, 15 Oct. 1949, W.
F. Barker 6075 (neotype, designated here,
NBG; isoneotypes, K, MO. PBE).

192

Novon

A rislra major Andrews, Bot. Repos. 3: pi. 160. 1801.
TYPE: South Africa. Western Cape: without precise
locality or collector, illustration in Andrews, Bot. Re¬
pos. 3: pi. 160. 1801.

Arislea thyrsi flora sensu N. K. Brown, Kew Bull. 1929: 36.
1929, non Ixia thyrsiflora I). Delaroche, Diss. pi.
nov. 20. 1 766.

In llie imperfect 19th century taxonomy ol Aris-
tea sect. Racemosae more than one species was in¬
cluded under the name Aristea capitata, most com¬
monly A. bakeri or A. bracteata. Weimarck (1940)
made significant progress in distinguishing the spe¬
cies of the section, admitting in addition to A. cap¬
itata i, A. thyrsiflora (with A. major as a synonym),
A. macrocarpa (desc ribed in this monograph by G.
.). Lewis), the nomenclaturally superfluous A. ca-
erulea (Thunberg) Vahl, and A. bakeri klatt. The
name ,4. capitata was applied to plants that we can¬
not at present distinguish in any way from Lewis’s
A. macrocarpa except for the smooth spathes and
bracts, but specimens cited by Weimarck include
plants that we would now refer to A. major. More¬
over. the reference illustration cited by Weimarck
(1940) for A. capitata, Curtis's Botanical Magazine
plate 605, also seems to differ in no significant way
from A. major. Other members of Aristea sect. Ra¬
cemosae that Weimarck recognized, but do not di¬
rectly concern the history of A. capitata, are A.jitn-
cifolia Baker, A. racemosa Baker, and A. rigidifolia
G. J. Lewis.

This paper also deals with two nomenclatural
and taxonomic questions in Aristea sect. Racemosae
related to the neotypification of Gladiolus capitatus:
the correct name for A. confusa Goldblatt, a sub¬
stitute name for the illegitimate and superfluous A.
caerulea (Thunberg) Vahl (basionym Moraea caeru-
lea Thunberg); and the circumscription and earliest
names for plants currently called A. bakeri Klatt.
A. confusa , and A. macrocarpa (Goldblatt & Man¬
ning. 2000).

Thk Correct N ame eor Aristea caeri i.ea

In 1787 Thunberg described Moraea caerulea for
what we now know was the second species of Ar¬
istea sect. Racemosae to be named, the first being
4. capitata. In fact, specimens in the Thunberg col¬
lection at Uppsala, Sweden, show that his concept
of M. caerulea included two species, the lectotype
(effectively designated by Weimarck in 1940).
which is now Aristea bracteata, and the species that
we now call A. capitata. Moraea caerulea is super¬
fluous under ICBN Art. 52.1 (Greuter et ah, 2000),
because Thunberg cited the name Gladiolus capi¬
tatus L. in the protologue which, under ICBN Art.
52.2, constitutes inclusion of its type. Since M. ca¬

erulea is a superfluous basionym, it is illegitimate
and not available for later use. ker Gawler (1802)
included M. caerulea in the synonymy of A. capi¬
tata, but other contemporaries evidently disagreed
with him, and Vahl made the combination A. ca¬
erulea in 1805. South African born Hendrik Per-
soon (1805) also recognized Aristea. admitting five
species to the genus, one of them A. bracteata, a
new species, in which he included Thunberg’s Mo¬
raea caerulea. Inclusion of that name in synonymy
would seem to make A. bracteata a superfluous
name, but inasmuch as M. caerulea is itself super¬
fluous, Persoon’s new name is valid and legitimate.
Persoon cited the figure of M. caerulea in Thun¬
berg’s Dissertatio de Moraea (1787), which makes
it easy to interpret A. bracteata, since Persoon prob¬
ably did not see the specimens in Thunberg’s col¬
lection, given the difficulties of travel and commu¬
nication at the time. Thus, A. bracteata Persoon is
a nomenclaturally acceptable name for A. caerulea
(Thunberg) Vahl. The new name A. monticola pro¬
posed for Moraea caerulea (Goldblatt, 1971) be¬
comes an unnecessary nomen novum and later syn¬
onym.

J. G. Baker followed ker Gawler (1802) in treat¬
ing Aristea caerulea as a synonym of A. capitata in
his accounts of the genus in Handbook of the Iri-
deae (1892) and Flora Capensis (1896), but Wei¬
marck (1940) definitively recognized A. caerulea,
pointing out its morphological differences including
the conspicuously hispid-papillate, firm-textured
spathes, relatively narrow leaves 3—6 mm wide,
broad spathes and bracts, and relatively short cap¬
sules.

Aristea bracteata Persoon, Syn. pi. 1: 41. 1805,
nom. nov. for Moraea caerulea Thunberg, Dis¬
sertatio de Moraea no. 12. 1787, nom. illeg.
superfl. pro Gladiolus capitatus L. Aristea ca¬
erulea (Thunberg) Vahl. Enum. pi. 2: 124.
1805. Aristea monticola Goldblatt, J. S. Afri¬
can Bot. 37: 234. 1971. TYPE: South Africa.
Without precise locality or date, Thunberg s.n.
"M. caerulea (lectotype, designated by Wei¬
marck (1940), UPS).

Aristea caerulea var. rolmsta Weimarck, Acta Univ. Lund,
n.s. 36: 74. 1940. Syn. nov. TYPE: South Africa.
Western Cape: without precise locality, Oct. 1915. R.
Marloth 7173 (holotype, PRE).

Aristea caerulea var. elongata Weimarck, Acta Univ. Lund,
n.s. 36: 73. 1940. Syn. nov. TYPE: South Africa.
Western Cape: Saron, Sep. 1919, L. Bolus s.n. (syn-
tvpes. BOL 16739, k).

As outlined above, Aristea bracteata is readily
recognized by the relatively large individual flower

Volume 12, Number 2
2002

Goldblatt et al.

Aristea Section Racemosae

193

clusters enclosed by broad, more or less dry, brown,
firm-textured spathes anti tfie relatively narrow
leaves with prominent, hyaline margins. The
spathes are also conspicuously scabrid or hispid, a
feature shared with A. rupicola Goldblatt & .). C.
Manning (1997) and A. macrocarpa (here included
in A. baked), in which the character is weakly ex¬
pressed and sometimes evidently absent. The latter
species has distinctive, elongate capsules mostly
20-30 mm long and large flowers, the tepals ca. 20
X 15 mm versus 12—17 X 12—15 mm in A. brac-
teata (Lewis, 1950a, 1950b). As in other tall spe¬
cies of section Racemosae, there is some variation
in branching pattern. Of the two additional varieties
included in A. caerulea by Weimarck, A. caerulea
var. elongata represents a plant with short lateral
branches and sessile upper flower clusters, while
variety robusta also has sessile upper flower clus¬
ters and capsules to 14 mm long, compared with
capsules ca. 10 mm in the typical variety (Wei¬
marck, 1940). We see no need to recognize these
local variants.

The lectotypification of Moraea caerulea (Wei¬
marck, 1940) is somewhat unsatisfactory, because
Weimarck clearly designated the sheet annotated
by Thunberg as “47. caerulea a," which consists
only of the leafy part of the stem. In the protologue
of 47. caerulea, Thunberg listed two localities for
his two collections (now mounted on three sheets),
one from “between the Outeniqua Mts and the
Langkloof” and the other from "hills around the
Cape,” without linking the localities to specific her¬
barium sheets. The narrow leaves of “47. caerulea
a ’ belong with the flowering stem of the sheet
marked “47. caerulea which is the plant illus¬
trated in the Dissertatio (plate 2, fig. 2) and specif¬
ically cited by Persoon. This plant corresponds to
our interpretation of A. bracteata, and Thunberg’s
cited locality, “bills around the Cape,” agrees with
this interpretation. The specimen “47. caerulea /3
is the eastern form of Aristea capitata (presumably
from “between the Outeniqua Mts and the Lang¬
kloof”). This form has darker rhipidial spathes and
bracts and a somewhat less crowded synflorescence
than the typical, western form.

Delimitation ok Aristea rarer/

Among the robust members of Aristea sect. Ra¬
cemosae are plants with a panicle-like, compound
inflorescence, broad, fibrous, strap-like leaves, and
capsules at least three times as long as wide and
mostly 18—28 mm long, that contain 3 or 4 seeds
in each locule. Originally plants with such com¬
pound inflorescences and broad leaves were in¬

cluded in A. capitata sensu Ker Gawler (1802),
probably because the capsules were not known.
Baker (1892. Aug.) was the first to distinguish the
species as A. paniculata, a later homonym of A.
paniculata Pax (1892. Apr.). Klatt (1894) provided
the replacement name A. bakeri shortly thereafter
for Baker’s homonym, A. paniculata. Aristea macro¬
carpa was a new species described by the South
African G. J. Lewis in Weimarck (1940) for plants
until then included in A. capitata. Important fea¬
tures that Lewis used to distinguish A. macrocarpa
were the elongate capsules, bristly abaxial surface
of the spathes, and the entire, rust-brown bracts.

In view of Lewis’s recognition of Aristea macro¬
carpa, the inclusion by Weimarck (1940) of A. cap¬
itata in his account is puzzling. Weimarck associ¬
ated the name with plants from the southern Cape
and elsewhere, and cited, as a reference specimen,
the illustration in Curtis's Botanical Magazine pi.
605 (Ker Gawler, 1802), which we regard as A. cap¬
itata as the species is neotypified here. Specimens
cited by Weimarck are either that species or few-
branched specimens of A. bakeri. Equally puzzling
is Lewis’s (1950c) account of Aristea in Flora of the
Cape Peninsula, where she provided descriptions
for both A. macrocarpa and A. capitata, and accom¬
modated them in her key. Given the background
Lewis gave for describing A. macrocarpa the treat¬
ment is, at best, confusing. Nevertheless, Lewis’s
field understanding of the Iridaceae and of Aristea
was widely held to be sound. Respecting that,
Goldblatt (1971) provided an alternative. A. con-
fusa, for plants from the Cape Peninsula called A.
capitata. Lewis (1950c) distinguished A. capitata
from A. macrocarpa by the smooth rather than his-
pidulous bracts, narrower tepals 16—20 X 6— 8 mm
(vs. 18 X 14 mm for the outer tepals in A. macro¬
carpa), and capsules 12—20 mm long. The habit
and details of the spathes (except for the scabridity)
were essentially identical between the two species
while the capsules of A. macrocarpa are longer, 18—
35 mm.

Weimarck (1940), however, recognized one more
species, Aristea bakeri, that closely resembled A.
macrocarpa in general habit, spathe morphology,
and in the diagnostically critical elongate capsules.
The range of A. bakeri extended from Riversdale
and knysna eastward to the Vanstaden’s River
Mountains, thus overlapping that of Weimarck’s A.
capitata (i.e., A. confusa sensu Goldblatt) in the
west, the range of which was seen as extending
from the Cape Peninsula eastward to Knysna. As
mentioned in our key to the species of section Ra¬
cemosae (Goldblatt & Manning, 1997), Aristea bak¬
eri is separated from A. capitata sensu Weimarck

194

Novon

(and A. confiisa Goldblatt) on doubtful grounds.
Weimarck’s key criteria were: bracts brown and
persisting; branches of the synflorescence erect;
seeds dark, 3— 4(— 5) min long for A. capitata : versus
bracts gray-green and caducous; branches of the
synflorescence spreading; seeds brown. 1.5—2 mm
long for A. bakeri.

I he distinction simply does not hold when the
specimens in southern African herbaria, most of
them not seen by Weimarck and collected after the
1950s, are examined. Branching in the species is
somewhat variable, and caducous bracts are mis¬
leading. The bracts (by which we assume Weimarck
meant leaves subtending the branches, not the
spat lies or floral bracts) in fact rarely fall. Lastly,
the seed size distinction does not seem correct, for
seeds of plants collected since Weimareks time do
not fall into his two size classes. We conclude that
there is but one wild species for plant specimens
variously called Aristea bakeri, A. confusa, or A.
macrocurpa. 'flic earliest available and legitimate
name is klatt’s (1894) A. bakeri.

Aristea bakeri klatt, in Durand & Schinz, Con-
spect. fl. afr. 5: 169. 1895 [as 1894). nom. nov.
for Aristea panic ulata Baker, Handb. Irid. 144.
1892, Aug., nom. illeg., non A. paniculata Pax,
But. Jahrb. Syst. 15: 151. 1892. April. TYPE:
South Africa. Western Cape: Galgebosch, near
Vanstaden's Kiver, Dec. 1872, P. MacOwan
2077 (holotypc, k; isotypes, GRA. SAM).

Aristea macrocurpa G. J. Lewis, in Weimarck, Acta Univ.
Lund., n.s. 36: 74. 1940. TYPE: South Africa. West¬
ern Cape: Cape Peninsula, Kirstenhosch, Nov. 1935,
G. ,/. Lewis s.n. (holotype, BOL 21665).

Aristea confusa Goldblatt. J. S. African Bot. 36: 308. 1970.
Svn. nov. TYPE: South Africa. Western Cape: Table
Mountain, Nov. 1930, T. P. Stokoe s.n. (holotype,
BOL 17518).

As circumscribed here, Aristea bakeri may be
distinguished by the elongate, broadly winged cap¬
sules somewhat truncate at the apex and (12— )18—
28(-35) mm long that contain 3 or 4 lamellate
seeds per locule. The rhipidial spalhes and bracts
arc broadly ovate, dry, rusty brown, and entire, al¬
though often becoming irregularly torn with age.
Specimens from the west of its range, previously
included in A. macrocarpa, have the abaxial sur¬
faces of the spathes and sometimes the bracts light¬
ly bristly or scabrid, a feature often obscure and
visible only under the microscope, and even then
sometimes only with careful examination. The open
branching of the flexuose flowering stem helps dis¬
tinguish the species from A. capitata, which has
short capsules mostly 8—19 mm long and about

twice as long as wide, and narrow, lanceolate
spathes and bracts, the margins of which are trans¬
lucent and the midlines darkly pigmented. Aristea
bracteata, which has prominently bristly spathes
and bracts, can be distinguished by the larger rhip-
idia. slightly smaller flowers, short capsules 10—
1 2( — 1 4) mm long, and narrow leaves 3—6 mm wide.

Recent research in connection with the prepa¬
ration of an account of Aristea in Cape Plants
(Goldblatt & Manning, 2000) led us to conclude
initially that A. confusa was nonspecific with plants
from the southeastern Cape called A. bakeri by
klatt (1894). At the time we still believed that A.
macrocarpa was distinct from A. bakeri I confusa and
separated them on the basis of the rhipidial spathes
being scabrid-papillate (A. macrocarpa) or smooth
(Goldblatt & Manning, 1997. 2000). That distinc¬
tion is weak because tin* papillae are often poorly
developed and seem to be absent in old inflores¬
cences, and sometimes even in plants of the same
collection. I here appears to be no other distinction
between the two species. Although we now include
A. macrocarpa in .4. bakeri, we recognize that the
distinction in the spathe and bract vestiture may
subsequently be linked with one or more characters
that vs ill prove our decision incorrect.

Acknowledgments. We thank Charles Jarvis,
British Museum of Natural History, and Dick Brum-
mill, Royal Botanic Gardens, kew, for advice on
nomenclatural questions. Support for field research
by grants 5994-97 and 6704-00 from the National
Geographic Society is gratefully recognized.

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Baker, J. G. 1892. Handbook of the Irideae. Reeve, Ash¬
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- . 1896. lridaceae. In W. T. Thiselton-Dyer (editor),

F lora Capensis 6. Reeve, Covenl Garden, London.
Brown, N. E. 1929. The lridaceae of Burman's Florae Ca¬
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Delaroche, I). 1766. Deseriptiones Plantarum Aliquot No-
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Goldblatt, I’. 1971. A new species of Gladiolus and some
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- & T. T. Barnard. 1970. The lridaceae of Daniel

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- & A. Le Thomas. 1997. Palynology, eladistie

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- & J. C. Manning. 1997. New species of Aristea

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Goldblatt et al.

Aristea Section Racemosae

195

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Greuter, W„ J. McNeill. F. R. Barrie, H. M. Burdet, V.
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J. E. Skog. P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (St. Louis Code). Regnum Veg. 138.
ker Gawler, J. 1802. Aristea capitata. Tallest aristea. Cur¬
tis’s But. Mag. 18: pi. 605.

klatt. F W. 1894 (as 1895). Iridaceae. Pp. 143-230 in T.
Durand & U. Schinz (editors). Conspectus Florae Af-
rieae. Jardin Botanique de I'Etat, Bruxelles. Berlin,
krok, T. (). B. N. 1925. Bibliotheca botaniea suecana ah
antiquissimis temporibus ad linem anni MCMXVI11/
Svensk botanisk litteratur fr&n aldsta tider t.o.in. 1918.
Uppsala & Stockholm: Almqvist & Wiksell.

Lewis, G. J. 1950a. Aristea macrocarpa. El. PL Africa 23:
pi. 1084.

- . 1950b. Aristea coerulea. FI. PI. Africa 23: pi.

1083.

- . 1950c. Iridaceae. Pp. 217-264 in R. S. Adamson

& T. M. Salter (editors). Flora of the Cape Peninsula.
Juta, Cape Town.

Linnaeus, C. 1753. Species Plantarum 37. L. Salvius,
Stockholm.

Pax, F. 1892. Iridaceae africanae. Bot. Jahrb. Syst. 15:
1 5( )- 1 57.

Persoon, C. H. 1805. Synopsis Plantarum seu Enchiri-
dium Botanicum. C. F. Cramer, Paris,
rhunberg, C. P. 1787. Dissertatio de Moraea. ,1. Edman,
Uppsala.

Vahl, M. 1805. Enumeratio Plantarum, vol. 2. N. Moller
& Sons, Copenhagen.

Weimarek, H. 1940. Monograph of the genus Aristea. Acta
Univ. Lund (Lunds Univ. Arsskr.), n.s. 36: 1-140.

A New Species of Homozeugos (Poaceae) from Angola

Gerald F. Guala

Fairchild Tropical Garden Research Center, 1 1935 Old Cutler Rd., Coral Cables (Miami),
Florida 33165, U.S.A. (addrt jss for correspondence), and Department of Biological Sciences,
Florida International University, Miami, Florida 33199, U.S.A. Stinger@fairchildgarden.org

Abstkact. As a result of recent revisionary stud¬
ies in the genus, a new species, Homozeugos con¬
ciliation, is described from the central highlands of
Angola in western Africa. It is differentiated from
tht' more common llornozeugos eylesii C. K. Hub-
hard by a much shorter ligule, inrolled leaves, and
longer microhairs on the abaxial surface of the
leaves with a lower ratio of basal cell length to
distal cell length. A full description of the new spe¬
cies and its predicted habitat is given.

Key words: Angola, Homozeugos, Poaceae.

Recent revisionary (Guala, 1997, 1998a, 1998b)
and biogeographic (Guala, 1998a, 1998b, 2000)
studies by the author on the genus Homozeugos
(Panieoideae: Andropogoneae) have shown that a
specimen collected in the central highlands of An¬
gola in 1969 during the war for independence is of
a species previously unknown to science. I he cur¬
rent civil war has precluded new fieldwork on the
species for the last 26 years, but sufficient herbar¬
ium material exists from the original collection to
determine that it is indeed a previously undescri¬
bed species. A description of the predicted habitat
is also given here. It is based on extrapolation from
a G I S database of 44 layers of spatial environmen¬
tal parameters for the point of collection. I lit' orig¬
inal data for the G1S composite were drawn from
the NOAA-KPA Global Ecosystems Database Pro¬
ject (1992). A complete discussion of the materials
and methods is given by Guala (1998a, 2000).

Homozeugos conciliation (7. F. Guala, sp. nov.
TYPE: Angola. Huambo: Chianga, “frequente
no terreno que ladeia, a esquerda, a estrada
para a Chianga,” ca. 1710 m, 14 Apr. 1969,
li. Teixeira 13216 (holotype, LISC; isotype,
I.ISJC). Figure 1.

Homozeugos eylesii affine sed ligulis minoribus, laminis
involutis el micropilis longioribus differt.

Caespitose perennial. Rhizomes generally short
and knotty. Bud scales sparsely pubescent, becom¬
ing shiny indurate and then stramineous. Culms ca.

1 m tall, 1.3— 3.6 mm wide; nodes villous, 1.8— 3.7
mm wide. Leaves green; sheath shorter than the
lamina in the midculm leaves, 14-47.2 cm long;
lamina inrolled, glabrous to sparsely hirsute, 13—
59 cm long (excluding the most distal leaf). 2.3—5
mm w ide (when flattened), linear to lanceolate with
a prominent midvein; margin minutely antrorsely
scabrous; ligule 4-5.4 mm long on the penultimate
leaf, 1.4— 5.4 mm long overall, domed to truncate.
Inflorescence of 4 to 5 racemose segments; each
segment 8.5-14.5 cm long with 20 to 35 spikelets;
rachis segments between the spikelet pairs 5.35—
5.6 mm long; pedicels 4.3— 5.2 mm long, hairs to
2.5 mm. Sessile and pedicellate spikelets weakly
appressed to the rachis; callus 1 .2-1.4 mm long,
pungent, velutinous, with the hairs becoming pro¬
gressively longer distally, to 3 mm; first glume 7.3—
8 mm long, 2.3— 2.4 mm wide, the lamina with 2
minutely scabrous keels appearing as the margins,
each ca. 0.6 mm wide distally grading into the mar¬
gin basally; the abaxial surface of the lamina gla-
brate to villous with hairs to 3.6 mm long; second
glume 7.3— 8.1 mm long, symmetrical, similar in
texture to the first glume, glabrous to villous with
hairs to 1.7 mm long; sterile lemma 5.5-5. 7 mm
long, hyaline, ovate to linear, 2-keeled, the apex
acute to minutely bifid, the margins villous; fertile
lemma 6.2—7 mm long, bifid at the apex with teeth
ca. 0.3 mm long, the awn straight to geniculate, 15—
24 mm long, hirsute to villous with hairs to 0.5 mm
long; palea 1.1 — 1.2 mm long; anthers 3.9— 4.3 mm
long, dehiscing from teardrop-shaped apical pores;
lodicules 0.4— 0.5 mm long. Diaspore composed of

— y

f igure I. Homozeugos conciliatum G. f. Guala. — A. Habit. Bar — 1 cm. — B. transverse section of a portion of the
leaf lamina. Bar - 1 mm. — C. Dissection of a spikelet: left to right; first glume, second glume, sterile lemma, fertile
lemma, pistil, lodicule, palea, lodicule, three anthers, callus. Bar = 1 mm. Drawn from Teixeira 13216 (holotype, LISC;
isotype, I.ISJC).

Novon 12: 196-199. 2002.

Volume 12, Number 2
2002

Guala

Homozeugos conciliatum from Angola

197

198

No von

the hermaphrodite spikelet with ils awn. Caryopsis
ca. 5 mm long.

Leaves in cross section involute, with generally
3 sizes of bundles: the largest and midsize bundles
with bundle caps on both the adaxial and abaxial
sides and spanning the thickness of t lie leaf, the
largest with conspicuously large vessels, the mid¬
size bundles in sets ol 1 to 3 intercalated between
each [tair of large bundles, without conspicuously
large vessels, the smallest bundles intercalated in
sets ol 1 to 3, between each pair of midsize bun¬
dles, these always with an abaxial bundle cap, but
without an adaxial one. Bulliform cells in discrete
Ian-shaped groups ol ea. 5 to 10. in a single adaxial
layer with smaller ones stacked and progressively
smaller abaxially forming narrowly triangular arms
between the vascular bundles, sometimes contain¬
ing tannin-like substances. Both surfaces of the
lamina similar in appearance with respect to the
costal areas, smooth, occasionally with a few
hooked prickles near the margins, the stomata in
lines alternating with single or rarely pairs of fu¬
siform cells; bicellular microhairs generally present
on both surfaces at the edges of the costae; abaxial
microhairs 54.24—55.14 gm long with their basal
cell 22.97—23.86 gill long and their distal cell
31.26—31.28 gun long, the ratio ol basal to distal
cell length 0.73—0.76; adaxial microhairs 45.64—
59.95 gun long with their basal cell 20.71-29.6 gun
long and their distal cell 24.92—31.29 gun long, the
ratio of basal to distal cell length 0.84—0.98;
straight triehomes to ea. 5 mm long also sometimes
present. Silica bodies occurring singly in cells that
occur singly or in lines, dumbbell- to 4-leaf-clover-
shaped.

Phenology. The type, collected in April, is at
anthesis.

Distribution and habitat. Homozeugos concilia¬
tion is known from a single locality on the road to
Chianga in Huambo Province at 1710—1715 m el¬
evation. Estimated coordinates: 12.80°S latitude
and 1 5.75°W longitude. Because there is only one
known locality for the species, a single point (the
type locality) was queried in the CIS database (see
Guala, 2000. 1998a) and the values are presented
here. Habitat parameters: the percentage of possi¬
ble sunshine hours that are actually sunny in Jan.
36%, Feb. 40%. Mar. 38%, Apr. 49%. May 69%.
June 80%, July 76%, Aug. 71%, Sep. 56%. Oct.
43%, Nov. 35%, Dec. 35%. The mean monthly pre¬
cipitation in Jan. is 219 mm, Feb. 179 mm. Mar.
238 mm, Apr. 146 mm. May 14 mm. June 0 mm,
July 0 mm. Aug. I mm, Sep. 20 mm, Oct. 121 mm,
Nov. 228 mm, Dec. 233 mm; with a mean annual

precipitation of 131 I mm. The mean monthly tem¬
perature in Jan. is 19.7°C, Feb. I9.9°C, Mar.
I9.9°C. Apr. 19.8°C. May 18.2°C, June 16.4°C.July
16.7°C. Aug. 19°C. Sep. 21°C, Oct. 20.9°C, Nov.
20°C. Dec. 19.8°C; with a mean annual temperature
of 19.2°C. and a surface albedo in Oct. ol 18%.
The generalized vegetation index (GVI) in Jan.
1990 was 134, in July 1990, 124. The percentage
of sand in the top I m of soil was 51%; percentage
of silt in the top I m of soil 17%; percentage of
clay in the top I m of soil 32%. In summary, the
habitat is characterized by a very dry and sunny
winter (May— September), with a warm wet growing
season.

Eponymy. The name is a plea for peace and
reconciliation in Angola. The home of this species
in Huambo is the site of some ol the most intense
fighting amidst the general hostilities paralyzing
Angola (INS Resource Information Center, 2000;
Slandley, 1999). As a result of the war. scientific
research on the country’s biota has been stifled for
decades.

Disci ssion

Homozeugos conciliation, known only from the
type collection, is clearly distinct from its five co-
genitors (Guala, 1998a, 1998b) and easily recog¬
nized by its short ligules and involute leaves. It was
not mentioned by Clayton (1973), and he appar¬
ently never saw the collection.

In a cladistic analysis of the entire genus (Guala,
1997. 1998a). II. conciliation shares long distal
cells ol the adaxial microhairs only with Homozeu¬
gos katakton Clayton, a generally much larger spe¬
cies that has awns 45—76 mm long, culms 1.2— 1.9
m tall, glumes > 9 mm long, and flat leaves 2.8—
6.3 mm wide. Although H. conciliation most closely
resembles H. eylesii morphologically, that species
is known only from several localities in Zambia.
Democratic Republic of Congo, Tanzania, and Ma¬
lawi. in contrast to all other species of Homozeugos,
which are restricted to Angola or (in the ease of //.
katakton) Angola and western Zambia.

Acknowledgments. I am grateful to the curators
of FISC and L1SCJ for their loans and to the staff
of FLAS for housing and maintaining those loans
during my studies at the University of Florida. I
also thank Walter Judd for support and encourage¬
ment during this work and the staff of tlx* Univer¬
sity of Florida Electron Microscopy Core Facility
for assistance with the SEM work that was integral
to this study.

Volume 12, Number 2
2002

Guala

Homozeugos conciliatum from Angola

199

Literature Cited

Clayton, VC D. 1973. A new species of Homozeugos (Gra-
mineae). Garcia de Orta 1: 11-12.

Guala, G. F. 1997. The integration of cladistic analyses
with GIS in the revision of two genera of tropical grasses
(Poaceae), Agenium Nees (from South America) and
Homozeugos Stapf. (from Africa). Amer. .). Mot. 84: 199.
| Abstract.]

- . 1998a. Revisions of Agenium and Homozeugos

(Poaceae: Andropogoneae): Integrating Cladistic Anal¬
ysis and Geographic Information Systems. Dissertation,
University of Florida, Gainesville.

- . 1998b. The relation of spatial environmental pa¬
rameters to phytogeny and species distribution in Agen¬
ium and Homozeugos in South America and Africa.
Amer. J. Hot. 85: 104. [Abstract.]

- . 2000. The relation of space and geography to

eladogenic events in Agenium and Homozeugos (Po¬

aceae: Andropogoneae) in South America and Africa.
Pp. 159-166 in S. W. L. Jacobs & J. Everett (editors).
Grasses: Systematics and Evolution. Volume 2 of Pro¬
ceedings. 2nd International conference on Comparative
Biology of Monocotyledons, Sydney, Australia 1998.
CSIRO, Victoria.

INS Resource Information Center. 2000. Angola: Current
Political and Human Rights Conditions in Angola. Re¬
sponse to information request number
AG001003.EXM. US Immigration and Naturalization
Service Resource Information Center, Washington. U.C.

NOAA-EPA Global Ecosystems Database Project. 1992.
Global Ecosystems Database Version 1.0. User's Guide,
Documentation, Reprints, and Digital Data on CD-
ROM. USDOC/NOAA National Geophysical Data Cen¬
ter, Boulder, Colorado.

Standley, J. 1999. “War and misery in Huambo." BBC
Online network 13 Aug. 1999.

A (New Combination in Morelia (Myricaceae) in Mesoamerica

Sandra Knapp

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD,

United Kingdom, sk@nhm.ac.uk

ABSTRACT. While completing the account of Myr-
icaceae for Flora Mesoamericana it was discovered
that a new combination was necessary in the genus
Morelia: M. lindeniana. 1 1 is here provided along
with a short synopsis of the distribution of the spe¬
cies and its differences from the widespread Mo¬
relia cerifera.

Key words: Mesoamerica, Morelia, Myrica ,
Myricaceae.

the small family Myricaceae contains much
morphological variation, and currently taxonomists
recognize four genera: My rica L., Morelia Loureiro,
CompUmia I'lleritier ex Aiton, and Canacomyrica
Guillaumin (Wilbur, 2001). These genera corre¬
spond to previously recognized subgeneric group¬
ings used by the last monographer of tin* family
(Chevalier, 1901), but recently it has come to light
that Chevalier (1901) incorrectly lectotypified the
genus Myrica with Myrica cerifera I,, (see Wilbur,
2001, for a full discussion). Verdcourt and Polhill's
(1997) proposal to conserve the generic name Myr¬
ica with a conserved type of Myrica cerifera was
rejected by the Committee for Spermatophyta
(Brummit, 1999). This necessitates the transfer ol
specific epithets for a large number of species from
Myrica to Morelia. Many of these transfers have
already been made (Wilbur, 1994; Killick el ah.
1998; Wilbur, 2001), or are in the process of being
made (Parra-Osorio, pers. comm.), but one such
transfer is necessary now to provide a valid name
for a species recognized in the treatment of Myri¬
caceae for Flora Mesoamericana.

Morelia liiuleniana ((7 DC.) S. Knapp, comb. nov.
Basionym: Myrica lindeniana C. DC.. Prodr.
10(2): 150. 1864. TYPE: Mexico. Chiapas: ./.
Linden 10 (holotype, G).

Morelia lindeniana is morphologically similar to
the more common and widespread Morelia cerifera
(L.) Small, but differs from it in its uniformly serrate
leaves, with the serrations not confined to the distal
margins, its longer leaves with more numerous and
prominent secondary veins (Burger, 1977). and in
its higher-elevation distribution. Morelia lindeniana
is found in the Mexican state of Chiapas, Guate¬
mala, and Honduras from 1600 to 2700 m in moist
mixed forests, while M. cerifera is primarily found
below 1600 m in a wide variety of drier habitats.

Acknowledgments. I thank C. Parra O. and R.
L. Wilbur for helpful reviews of the manuscript and
the curators of herbaria mentioned in tin1 text for
loan of specimens.

I .iterature Cited

Brummit, IT K. 1999. Proposals to conserve or reject. Re¬
port of the Committee on Spermatophyta. Taxon 48:
367.

Burger, W. 1977. Myricaceae. In Mora Costaricensis. Fiel-
diana, Hot. 40: 21-27.

Chevalier, A. 1901. Monographic des Myrieaeees. Mem.

Soc. Sci. & Nat. Math. Cherbourg 32: 85—341.

Killick, D. J. IT. R. M. Polhill & IT Verdcourt. 1998. New
combinations in African Myricaceae. Kew Bull. 53:
993-995.

Verdcourt, B. & 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 anil series. Sida 16:
93-107.

- . 2001. f ive new combinations in the genus Mo¬
relia (Myricaceae) for Neotropical species. Rhodora
103: 120-122.

Novon 12: 200. 2002.

New Combinations and a New Name in Syzygium (Myrtaeeae) from
Madagascar and the Comoro Islands

Jean- Noel hi bat

Laboratoire de Phanerogamie, Institul <le Systematique (FR 1541), Museum national
d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France, labat@mnhn.fr

George K. Schatz

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

george.schatz@mobot.org

Abstract. Acceptance of current generic con¬
cepts in Myrtaeeae results in the proposal of 14
new combinations and I new name in Syzygium
Gaertner. Those taxa from Madagascar and Mayotte
originally described in Eugenia L. that have ter¬
minal inflorescences and calyptrate corollas should
be accommodated within Syzygium. Complete syn¬
onymy and typifieation are given for all accepted
taxa.

RfiSUMfi. Les nouveaux concepts generiques chez
les Myrtaeeae nous amenent a proposer 14 com¬
bi naisons nouvelles et un nouveau tiom dans le
genre Syzygium Gaertner. Ces taxa de Madagascar
et Mayotte, originellement decrits dans Eugenia L.
et qui possedent des inflorescences terminales el
des corolles en calyptre, doivent etre places dans
le genre Syzygium. La synonymie complete et la
typifieation sont donnees pour tons les taxa accep¬
ts.

Key words: Comoro Islands, Eugenia, Mada¬
gascar. Myrtaeeae, Syzygium.

In both his treatment ol Myrtaeeae for the Flore
de Madagascar et des Comores, as well as its [(re¬
cursor in which new taxa were described. Perrier
de la Bathie (1952, 1953) adopted the broad view
of Eugenia L., including those taxa with terminal
inflorescences and calyptrate corollas, which he as¬
cribed to “Section Syzygium .” More recent studies
(Schmid. 1972) have definitively shown that nu¬
merous characters, including both gross floral mor¬
phological as well as anatomical characters, can be
utilized to separate Eugenia from Syzygium Gaert¬
ner. This concept of two distinct genera is now gen¬
erally accepted in modern treatments of the Myr-
taceae, as evidenced by numerous accounts for
regional floras, including Elora Zambesiaca (White,
1977), Flore des Mascareignes (Scott. 1990), and
Flore des Seychelles (Friedmann, 1994). Thus, in

line with current generic concepts, and to conform
with the adjacent regional floristic treatments cited
above, we propose the following new combinations
and a replacement name for Malagasy Syzygium.

Comprised of over 500 species, the genus Syzy¬
gium is strictly Old World with a center of diversity
in the Australasian region. Within Myrtaeeae, it is
allied to a group of Australasian— Malesian genera
that together form the Acmena Alliance (Briggs &
Johnson, 1979), sharing predominantly terminal in¬
florescences, the absence ol standard myrtaceous
hairs, and a mostly glabrous condition. The wood
anatomy and unitegmic ovules of Syzygium argue
against a close relationship with Eugenia s. str.
(Johnson & Briggs, 1984). Apart from the Syzygium
taxa enumerated below, the remaining native Mal¬
agasy Myrtaeeae all belong to Eugenia s. sir., a
genus of ca. 1000 mostly New World species. With
the transfer of several species originally described
in Myrtus L. (Scott, 1980), and the description of
several new species (Miller, 2000), Eugenia now
numbers ca. 39 species in Madagascar and the
Comoro Islands. Although not all Syzygium species
possess a calyptrate corolla, all Malagasy Syzygium
can be distinguished from Eugenia by their calyp-
trate corollas and terminal (never cauliflorous) in¬
florescences, versus free petals and axillary or cau¬
liflorous (never terminal) inflorescences in Eugenia.

Syzygium aurantiacum (H. Perrier) J.-N. Labat
& G. E. Schatz, comb. nov. Basionym: Eugenia
aurantiaca II. Perrier, Mem. Inst. Sci. Mada¬
gascar, Ser. B. Biol. Veg. 4(2): 190. 1952.
TYPE: Madagascar. Bords de la Lokoho
(N.E.), 1000 m, 2 Jan. 1949, G. Cours 342H
(holotype, P).

This species is only known from the type spec¬
imen collected in northeastern Madagascar, within
or near the Marojejy Strict Nature Reserve.

Novon 12: 201-205. 2002.

202

Novon

Syzygium baronii J.-N. La hat & G. K. Schatz,
noin. nov. Replaced name: Eugenia cyclophyl-
la Baker, J. But. 20: III. 1882; non Eugenia
cyclophylla Berg, in Marti us, FI. Bras. 14(1):
287. 1857; non Eugenia cyclophylla Thwaites
ex Duthie, in Hooker f., FI. Bril. hid. 2: 494.
1878. TYPE: Madagascar. Central, sine loc..
R. Baron 219 (holotype, k).

Eugenia vacciniifolia Baker, J. Finn. Soe., Bot. 20: 145.

1883. Syn. nov. Non Syzygium vaecinifolium Merrill,
Philipp. J. Sei. 70: 420. 1951. TYPE: Madagascar.
Central, sine loc., R. Baron 1919 (holotype, K; iso-
tvpe. P).

Eugenia loiseleuroides Baker. J. Linn. Soe.. Bot. 21: 341.

1884. Syn. nov. TYPE: Madagascar. Sine loc., R.
Baron 2641 (holotype, k: isotype, P).

Perrier de la Balltie (1952, 1953) used the name
E. vacciniifolia Baker for this species, perhaps Be¬
cause he was aware that the name Eugenia cyclo¬
phylla had been used previously by two other au¬
thors before Baker. A nomen novum is now
necessary because Syzygium vaecinifolium Merrill
already exists. Syzygium baronii is distributed in
humid montane forest of the Imerina region of the
Central High Plateau, and is apparently common in
the Ankaratra Massif between 1600 and 1800 m
altitude.

Syzygium bernieri (Drake) J.-N. I.abat N C. K.
Schatz. comb. nov. Basionym: Eugenia bernieri
Drake (as "Bernieri"). Bull. Mens. Soe. Linn.
Paris 2: 1221. 1896. T\ PF: Madagascar. Ste.-
Marie, 1834. A. C. ,/. Bernier 1 95 (as "155")
(lectotype, designated by Perrier de la Bathie
(1952). P; isolectotype, P).

Eugenia chapelieri Drake (as “ Chapelieri ”), Bull. Mens.
Soe. Linn. Paris 2: 1222. 1896. Syn. nov. TYPE:
Madagascar. [Environs de Tamatave], /.. A. Chapelier
s.n. (holotype, P; isotype, P).

Eugenia maroana Aug. DC., Bull. Herb. Boissier, Ser. 2,
1(6): 573. 1901. Syn. nov. I V PE: Madagascar. En¬
virons de Maroantsetra. 1897, 4. Mocquerys 348 (ho¬
lotype, G).

This species varies considerably throughout its
range (mainly in leaf size and shape and flower
size), and, like .S', emirnense, requires a thorough
systematic revision prior to the recognition of ad¬
ditional specific or infraspecific taxa. Therefore, the
three varieties (Eugenia bernieri var. oblanceolata.
Eugenia bernieri var. latericolor. Eugenia bernieri
var. tampinensis) described by Perrier de la Bathie
(1952), which moreover are invalid in lacking Latin
diagnoses, are not recognized here. Syzygium ber¬
nieri is widely distributed in tin* humid forest along
the east coast of Madagascar as well as the Sam-

birano region including Nosy Be, from sea level to
ca. 1000 m altitude, but more common at the lower
elevations.

Syzygium condcnsatum (Baker) J.-N. Lahat & G.
E. Schatz, comb. nov. Basionym: Eugenia con-
densata Baker. J. Bot. 20: I 12. 1882. TYPE:
Madagascar. Central. Betsileo-land. R. Baron
237 (holotype, K: isotype, P).

Syzygium condensatum, S. micropodum, and the
very polymorphic species S. emirnense together
constitute a difficult complex of species that re¬
quires a thorough systematic revision. We provi¬
sionally adopt Perrier de la Bathie's concept of spe¬
cific limits within the complex, but exclude E.
aggregata from synonymy under S. condensatum.
As treated below, further study of the lectotype of
E. aggregata (R. Baron 4917) clearly indicates that
it is conspecifie with S. phillyreifolium. Syzygium
condensatum is distributed in humid montane forest
on the Central High Plateau around Antananarivo,
as well as in the Andapa region including the Ma-
rojejy Strict Nature Reserve.

Syzygium duuguyaiium (H. Perrier) J.-N. Cabal
& G. E. Schatz, comb. nov. Basionym: Eugenia
danguyana II. Perrier (as “ Danguyana ”),
Mem. lust. Sei. Madagascar. Ser. B, Biol. Veg.
4(2): 195. 1952. TYPE: Madagascar. Ambod-
imanga a Antanambao [near Lac Alaotra].

1 200 m, I I Oct. 1945. G. Cours 2800 (lecto¬
type. designated here, P).

This species is known from relatively few collec¬
tions in low- to mid-elevation humid forest up to
1200 m altitude, from Farafangana to the Masoala
Peninsula as far west as Lac Alaotra. The variety
Eugenia danguyana var. rotranala, invalidly de¬
scribed by Perrier de la Bathie (1952), is not rec¬
ognized here.

Syzygium emirnense (Baker) J.-N. Labat N G. E.
Schatz, comb. nov. Basionym: Eugenia emir-
nensis Baker, J. Linn. Soe., Bot. 20: 145. 1883.
TYPE: Madagascar. Central, sine loc.. R. Bar¬
on 1076 (lectotype, designated here, k: isolec¬
totype, P).

Eugenia cuneifoliu Bojer ex Baker, J. Linn. Soe.. lint. 20:
144. 1883. Syn. nov. Eugenia em i mens is f. euneifolia
(Bojer ex Baker) H. Perrier, Mem. Inst. Sei. Mada¬
gascar. Ser. B. Biol. Veg. 4(2): 187. 1952. TY PE:
Madagascar, Central, sine loc., R. Baron 1254 (lec¬
totype, designated by Perrier de la Bathie (1952), K;
isolectotype, P).

Eugenia riehardiana Cordemoy, EL lie Reunion: 430.
1895. Syn. nov. Syzygium richardianum (Cordemoy)

Volume 12, Number 2
2002

Labat & Schatz 203

Syzygium from Madagascar and Comoro Islands

Gueho & A. .1. Scott, Lew Bull. 34: 494. 1980.
TYPE: Reunion. Versants des montagnes, ./. M. C.
Richard s.n. (holotype, MARS).

Eugenia emirnensis var. elongata Hochreutiner, Annuaire
Conserv. Jard. Rot. Geneve 1 I and 12: 76. 1908.
Syn. nov. TYPE: Madagascar. District of Vatomandry,
21 Eeh. 1904, J. Guillot 101 (holotype, G; isotype,
K).

Eugenia condensata Baker var. thouvenotii P. Danguy ex
Leeomte, Ees Bois de la foret d’Analamazaotra: 106.
1922. Syn. nov. TYPE: Madagascar. Analamazoatra,
1919, E. Thouvenot 124 (holotype, P; isotvpes, MO,

P).

After examination of the abundant material
available, it appears that the considerable variabil¬
ity in both vegetative and reproductive characters
may correspond to habitat differences and thus lo¬
cal ecological specialization. Comprehensive field
studies are needed to clarify the patterns of varia¬
tion prior to possible recognition of additional taxa
(see also notes under S. condensatum and S. micro-
podum). Eugenia richardiana, which is based on a
J. M. C. Richard specimen that is probably incor¬
rectly indicated as having been collected in Re¬
union. corresponds to S. emirnen.se (see the editor's
note in Scott. 1990: 29). In its broad conception as
here accepted, S. emirnense is widely distributed
and common throughout all of the humid to mon¬
tane forest of eastern and central Madagascar. The
variety Eugenia emirnensis var. submaritima and
forma Eugenia emirnensis f. podocarpifolia invalidly
described by Perrier de la Bathie (1952) are here
considered to lie within the overall variability of
Syzygi u rn emi rn ense.

Syzygium huiublotii (H. Perrier) J.-IY Rabat G.
E. Schatz, comb. nov. Basionym: Eugenia
humblotii H. Perrier (as “ Humblotii ”), Mem.
Inst. Sci. Madagascar, Ser. B, Biol. Veg. 4(2):
193. 1952. TYPE: Comores. Mayotte, Foret de
Combani, 19 Oct. 1884, L. Humblol 1329 [in¬
correctly cited by Perrier de la Bathie as
“Madagascar, Nossive, L. Ilumblol 329 vel
1 3293" \ (holotype. P: isotype. P).

This species, which is known only from the type
specimen, is endemic to the island of Mayotte in
the Comoros Archipelago. In the protologue of E.
humblotii, Perrier de la Bathie (1952) cited the type
specimen as "Hum blot 329 vel 1329?" indicating
the locality for the collection as Nossive in Mada¬
gascar. Examination of Humblots field books at P
reveals that Humblot 329 from Madagascar repre¬
sents “ Psychotria lantzii" a Drake manuscript
name that was later published as Cremocarpon lan¬
tzii Bremekamp (Rubiaceae). Humblots collections
from the Comoros were also numbered beginning

with 1, but then subsequently renumbered at P to
avoid confusion with his Malagasy collections (see
Dorr, 1997). Thus, his Mayotte collection from Oc¬
tober 1884 bearing field number 329 (but inexpli¬
cably numbered 2329 in the second of three Com¬
oros field books at P) was then renumbered 1329,
and is therefore the correct citation for the type of
E. humblotii.

Syzygiuin luguhre (II. Perrier) J.-N. Labat & C.
b. Schatz, comb. nov. Basionym: Eugenia lu-
gubris H. IVrrier. Mem. Inst. Sci. Madagascar,
Ser. B, Biol. Veg. 4(2): 187. 1952. TYPE:
Madagascar. Foret d’Analamazoatra, 800 rn,
Feb. 1912, //. Perrier de la Bathie 6487 (ho¬
lotype, P).

This species is distributed in humid forests of
central eastern Madagascar at mid-elevations, from
the regions of Andramasina and Tsinjoarivo to the
southeast of Antananarivo to the Lac Alaotra re¬
gion. The forma Eugenia phillyreaefolia f. obscuri-
folia invalidly described by Perrier de la Bathie
(1952) and then later presented as a variety (Perrier
de la Bathie, 1953), represented by M. Louvel 87,
is here considered to belong to Syzygium lugubre.

Syzygium micropodum (Baker) J.-N. Labat cK C.
E. Schatz, comb. nov. Basionym: Eugenia mi-
cropoda Baker, J. Linn. Soc.. Bot. 20: 143.
1883. TYPE: Madagascar. Central, sine loe.,
R. Baron 388 (holotype, K; isotype, P).

This species is widely distributed in mid-eleva-
tional to montane humid forest, mainly above 1000
m altitude, from Andohahela to Tsaratanana in¬
cluding the higher elevations of the Masoala Pen¬
insula. Plants collected in the littoral forest on sand
along the eastern Malagasy coast between Ft. Dau¬
phin and Ambila-Lemaitso correspond to the spec¬
imens cited by Perrier de la Bathie (1952) (H. Per¬
rier de la Bathie 14299 ; E. Urseh 107) under
invalidly described Eugenia micropoda var. littor-
alis and may represent a distinct taxon. Specimens
representing the forma Eugenia emirnensis f. sub-
rotundifolia, invalidly described by Perrier de la
Bathie (1952), are here placed under Syzygium mi¬
cropodum. Further detailed studies are necessary
before the naming of additional taxa within this
species complex that includes S. condensatum, S.
emirnense, and S. micropodum.

204

Novon

Syzygiuin onivense (II. IVrrier) J.-N. Lahat & G.
E. Schatz, comb. nov. Basionym: Eugenia on-
ivensis H. Perrier, Mem. Inst. Sci. Madagascar,
Ser. B, Biol. Veg. 4(2): 188. 1952. TYPE:
Madagascar. Environs du confluent de POnive
et du Mangoro, 600 m, Feb. 1926, //. Perrier
de la Bdthie 16990 (holotype, P).

This species is known from relatively few collec¬
tions in mid-elevation humid forest between 600
and 1 100 m altitude from the Tanala region to Lac
Alaotra.

Syzygiuni parkeri (Baker) J.-N. Lahat & G. E.
Schatz, comb. nov. Basionym: Eugenia parkeri
Baker (as "Parkeri”), J. Linn. Soe., Bot. 20:

144. 1883. TYPE: Madagascar. Central, sine
loc., G. 1L Parker s.n. (holotype. K).

Eugenia ibitensis Drake, Bull. Mus. Hist. Nat. (Paris) 9:
42. 1903. Syn. nov. Eugenia parkeri Baker var. ibi¬
tensis (Drake) II. Perrier, Mem. Inst. Sci. Madagas¬
car, Ser. B, Biol. V^g. 4(2): 194. 1952. TYPE: Mad¬
agascar. Imerina, Mont Ibity, 13 Jan. 1901, G.
Grandidier s.n. (holotype, P).

This species is widely distributed in humid to
montane forest from the Andringitra Massif to Mon-
tagne d’Ambre, and from sea level along the eastern
coast to the Ibity Massil in central Madagascar.
Like S. emirnense, S. parkeri is here accepted in its
broad conception of Perrier de la Bathie (1953), (he
confirmation of which requires further field inves¬
tigations and systematic study.

Syzygium pliillyreifnlium (Baker) J.-N. Lahat &
G. E. Schatz, comb. nov. Basionym: Eugenia
phillyreaefolia Baker, J. Linn. Soc., Bot. 20:

145. 1883. TYPE: Madagascar. Central, sine
loc.. R. Baron 95B (holotype, k; isotype, P).

Eugenia aggregata Baker, J. Linn. Soc., Bot. 22: 475.
1886. Syn. nov. TYPE: Madagascar. Sine loc., R.
Baron 4917 (lectotype. designated by Perrier de la
Bathie (1952), k; isolectotype, P).

This species is widely distributed throughout the
Central High Plateau from the Andringitra and Is-
alo Massifs to the Tampoketsa d’Ankazobe, and as
far west as the Bongolava region to the west of Tsi-
roanomandidy.

Syzygiiim sakalavarum (H. Perrier) J.-N. Lahat
& G. E. Schatz, comb. nov. Basionym: Eugenia
sakalavarum H. Perrier (as "Sakalavarum”),
Mem. Inst. Sci. Madagascar, Ser. B. Biol. Veg.
4(2): 189, pi. 13. 1952. TYPE: Madagascar.
Tsarasaotra, Nov. 1897 [as “Quest. Secteur
Ambongo-Boina: bords de tous les cours d’eau
du secteur”], // Perrier de la Bdthie 369 (lec¬
totype, designated here, P; isoleetotypes, P, P
(Drake)).

This species is widely distributed throughout
western Madagascar from Ft. Dauphin to Antsiran-
ana in dry to sub-arid deciduous forest mainly
along watercourses up to 1000 m altitude. Among
the syntypes originally cited, three of the sheets of
//. Perrier de la Bdthie 369 at P bear labels with a
precise locality and date, two of which also bear
type stickers probably affixed by Perrier de la Bath¬
ie himself; one of these latter two is thus here des¬
ignated as the lectotype. Two other sheets at P also
numbered as H. Perrier de la Bdthie 369 bear la¬
bels without the precise locality, but instead cor¬
respond to the more regional designation given for
the collection included in the protologue. Moreover,
on these more general labels, one sheet with (lowers
only is ascribed to October, and the other with fruits
to January. It is therefore likely that although these
two sheets are numbered as //. Perrier de la Bdthie
369, they were probably collected at different times
and places than the specimen bearing the Type
sticker, and are thus not considered here to rep¬
resent isoleetotypes. Capuron (1966) suggested that
E. sakalavarum was a synonym of 5. guineense
(Willdenow) DC. Lacking more detailed systematic
studies, we prefer to retain Perrier de la Bathie’s
concept.

Sy/.ygiiiin sainbiranense (II. Perrier) J.-N. Lahat

6 G. E. Schatz, comb. nov. Basionym: Eugenia
samhiranensis H. Perrier, Mem. Inst. Sci. Mad¬
agascar. Ser. B. Biol. Veg. 4(2): 193. 1952.
TYPE: Madagascar. Sambirano: valine du
Sambirano, 200—800 m, Dec. 1922, //. Perrier
de la Bdthie 15137 (holotype, P; isotype, P).

This species is only known from the type spec¬
imen.

Syzygiiim tapiaka (H. Perrier) J.-N. Lahat & G.
E. Schatz, comb. nov. Basionym: Eugenia tap¬
iaka H. Perrier (as "Tapiaka”), Mem. Inst. Sci.
Madagascar, S£r. B, Biol. Veg. 4(2): 190.
1952. TYPE: Madagascar. Tsaramandroso,
Ambato-Boeni, Bevaza (Reserve Naturelle n°

7 [Ankarafantsika]), 13 Oct. 1947, Reserves
Nat ure lies 1094-RN (holotype, P; isotypes, k,
MO, P).

This species is restricted to the dry deciduous
forest on sand at the Ankarafantsika Strict Nature
Reserve.

Taxa of Uncertain Systematic Position

Eugenia tanalensis Baker, J. Bot. 20: 111. 1882. TYPE:
Madagascar. Forests of Tanala, R. Baron 295 (holo¬
type, k not found).

Volume 12, Number 2
2002

Labat & Schatz 205

Syzygium from Madagascar and Comoro Islands

Perrier de la Bathie (1952) initially considered
E. tanalensis to be a synonym of E. phillyreaefolia,
stating that “Le type de E. tanalensis Baker (R.
Baron 1477) n’a que des fleurs deformees, parasi-
tees, indeterminables; mais les feuilles intaetes
sont celles de V Eugenia phillyreaefolia .” However,
the type of E. tanalensis is R. Baron 295; R. Baron
1477 must surely have been determined by Baker
as E. tanalensis at some later date. Moreover, al¬
though some branches of the Kew specimen of R.
Baron 1477 clearly show signs of insect parasite
deformation, others do not and can be identified as
Syzygium onivense, a conclusion Perrier de la Bath¬
ie also shared later (1953) in removing E. tanalen¬
sis from synonymy under E. phillyreaefolia and des¬
ignating it as an “incompletely known species.”
Unfortunately, we have been unable to locate any
material of R. Baron 295, and it is therefore im¬
possible to clarify the identity and status of E. tan¬
alensis.

Three other species described by Perrier de la
Bathie (1952) in Eugenia sect. Syzygium have sub¬
sequently been treated by other authors:

Eugenia cotinifolioides H. Perrier, Mem. Inst. Sei. Mada¬
gascar, Sdr. B, Biol. Veg. 4(2): 185. 1952. TYPE:
Madagascar. Pentes orientales du Marojejy, a l’ouest
de la riviere Manantenina, affluent de la Lokoho,
vers 1600 m, Dec. 1948, H. Humbert 22634 (holo-
type, P). = Memeeylon cotinifolioides (H. Perrier)
Jacques-Felix, Bull. Mus. Natl. Hist. Nat., ser. B,
Adansonia, 7(1): 14. 1985.

Eugenia cupulifera H. Perrier, Mem. Inst. Sci. Madagas¬
car, Ser. B, Biol. Veg. 4(2): 190. 1952. TYPE: Mad¬
agascar. Foret d’Analamazoatra, Gouv. General de
Madagascar 3 (holotype, P). = Carallia brachiata

(Loureiro) Merrill, Phillipp. J. Sci. 15(3): 249. 1919;
Capuron, Adansonia, sdr. 2(1): 128. 1962.

Eugenia goudotiana H. Perrier, Mem. Inst. Sci. Madagas¬
car, Ser. B, Biol. V6g. 4(2): 193. 1952. TYPE: Mad¬
agascar. Sine loc., J. P. Goudot 914 (holotype, P). =
Syzygium zeylanicum (L.) DC., Prodr. 3: 260.
1828; Capuron, Adansonia, ser. 2 (1): 128. 1962.

Literature Cited

Briggs, B. G. & L. A. S. Johnson. 1979. Evolution in
Myrtaceae — Evidence from inflorescence structure.
Proc. Linn. Soc. New South Wales, Ser. 2, 102: 1 57 —
256.

Capuron, R. 1966. Etude sur les Essences Forestihres de
Madagascar, Rotra, Fiche Botanique. Centre Technique
Forestier Tropical, Antananarivo.

Dorr, L. 1997. Plant Collectors in Madagascar and the
Comoro Islands. Royal Botanic Gardens, Kew.

Friedmann, F. 1994. Flore des Seychelles — Dicotyledo-
nes. ORSTOM, Paris.

Johnson, L. A. S. & B. G. Briggs. 1984. Myrtales and
Myrtaceae — A phylogenetic analysis. Ann. Missouri
Bot. Card. 71: 700-756.

Miller, J. S. 2000. New taxa and nomenclatural notes on
the flora of the Marojejy Massif, Madagascar. IV. Myr¬
taceae: New species of Eugenia L. Adansonia, s£r. 3,
22: 111-116.

Perrier de la Bathie, H. 1952. Les Myrtacees de Mada¬
gascar et des Comores. Revision, diagnoses et biologie.
Mem. Inst. Sci. Madagascar, Ser. B, Biol. Veg. 4(2):
161-202.

- . 1953. Myrtacees. FI. Madagasc. 152: 1—80.

Schmid, R. 1972. A resolution of the Eugenia— Syzygium
controversy (Myrtaceae). Amer. J. Bot. 59: 423^436.

Scott, A. J. 1980. A note on Myrtus (Myrtaceae) in Mad¬
agascar. Kew Bull. 34: 546.

- . 1990. Myrtacees. Flore des Mascareignes 92: 1-

70. M.S.R.I., ORSTOM and Kew.

White, F. 1977. Myrtaceae. In FI. Zambesiaca 4: 183—
212. Flora Zambesiaca Managing Committee, London.

Seventeen New Species of Lobelioideae (Campanulaceae) from

South America

Thomas G. Lammers

Department of Biology and Microbiology, University of Wisconsin Oshkosh, Oshkosh,
Wisconsin 54901, U.S.A. lammers@uwosh.edu

ABSTRACT. Almost I 1% of the 400 unidentified
specimens of South American Lobelioideae (Cam¬
panulaceae) examined recently cannot be equated
with any known species. They are described here
as 1 7 new species: 6 of Burmeistera, 6 of Centro-
pogon, and 5 of Siphocampylus. Several are char¬
acterized by morphological characters or combina¬
tions of characters that are unique or unusual in
their respective genera or in the subfamily as a
whole. Six of the novelties are endemic to Ecuador,
four to Peru, three each to Venezuela and Colombia,
and one to Bolivia.

Key words: Burmeistera , Campanulaceae, Cen-
tropogon, Lobelioideae, Siphocampylus , South
America.

The neotropical endemics Burmeistera Triana,
Centropogon C. Presl, and Siphocampylus Pohl are
the largest genera of Lobelioideae (Campanulaceae)
after cosmopolitan lobelia L. Together, these three
closely related genera (Pepper et al., 1997; Lam¬
mers, 1998a; Buss et al., 2001) comprise well over
500 species, found from southern Mexico to north¬
ern Argentina and in the West Indies, with their
greatest diversity in high-elevation habitats of the
Andes. All are large robust plants (herbaceous or
suffruticose perennials, shrubs, subshrubs, lianas,
and woody rosette plants), with large flowers (co¬
rollas averaging 30—60 mm long) borne singly in
the axils of the upper leaves or aggregated into ter¬
minal foliose or bracteate inflorescences (racemes,
corymbs, umbels); the corolla tube is neither fe¬
nestrate nor cleft dorsally and if the lobes are di¬
morphic, it is the dorsal pair that is larger (Lam¬
mers, 1998a, in press a, in press b).

Between 1990 and 1997, I examined approxi¬
mately 750 specimens of neotropical Lobelioideae
that had been sent to me for identification (Lam¬
mers, 1997); 28 of these (3.7%) could not be equat¬
ed with known species and were described as 19
new species (Lammers, 1993, 1998a). At that time,
1 indicated that several hundred additional speci¬
mens remained to be identified; 400 of these have
now been studied. Of these, 43 (10.8%) could not

be referred to any known species. They are here
described as 17 species new to science: 6 in Bur¬
meistera, 6 in Centropogon, and 5 in Siphoeampy-
lus. With these additions, the 3 genera collectively
total 545 species.

The incredible diversity evident from the fore¬
going discussion merits some comment. Altogether.

1 150 specimens of neotropical lobelioids were ex¬
amined over the past decade. Of these, 6.2% or 1
out of every 16 specimens sent to me for identifi¬
cation represented a species previously unknown to
science. Furthermore, many ol these novelties pos¬
sess features or combinations of features that are
unique or unusual (Lammers, 1997, 1998a). Among
those described here, for example, Burmeistera ar-
busculifera is the first species ol its genus with ar-
busculiform hairs and B. fimbriata the first with a
fimbriate leaf margin, while B. brighamioides and
B. hippobromoides have flowers far longer than any
of their congeners. Centropogon candidatus is
unique in its combination of two character states
rare within the group: white arbusculiform tri¬
e-homes and pinnately lobed leaves. Similarly, Siph¬
ocampylus ambivalens is the first species in the
subfamily to combine an umbellate inflorescence
with whorled leaves, S. prevaricator the first to com¬
bine an umbel with arbusculiform trichomes, while
S. plegmatocaulis is unique in its dimorphic twin¬
ing stems and dimorphic trichomes surrounding the
orifice of the anther tube.

My experience is by no means unique. A gen¬
eration ago, one of the foremost students of Lobe¬
lioideae. Rogers McVaugh (b. 1909), described
many new species of South American Lobelioideae
on the basis of specimens sent to him with requests
for identification. Several (e.g., Burmeistera pteri-
dioides McVaugh, Centropogon various McVaugh)
were characterized by unique features or combi¬
nations of features. He commented that “almost ev¬
ery mountain in the Andes seems to have different
species [of Lobelioideae] on it ... a host of new
and puzzling things turn up in every new collection
from the Andes, and it is apparent that there is still
a lifetime of work for someone interested in these

Novon 12: 206-233. 2002.

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

207

bizarre and beautiful plants” (McVaugh, 1965:
400).

McVaugh’s quote is as true today as it was 35
years ago. Diversity in these genera is far from ex¬
hausted. and further exploration of the Andes will
continue to yield new species of Burmeistera, Cen-
tropogon, and Siphocampylus for many years to
come. Furthermore, if these genera are at all rep¬
resentative of the neotropical flora as a whole, it is
clear that a very significant proportion ol the biotic
diversity of that region remains to be named and
described.

I. Burmeistera Triana, Nuev. Jen. Esp. 13. 1855.
TYPE: Burmeistera ibaguensis Triana.

With the addition of the 6 novelties described
here, Burmeistera comprises 102 species, distrib¬
uted from Guatemala to Peru (Lammers, 1998a;
Lammers & Maas, 1998). The genus may be dis¬
tinguished from Centropogon and Siphocampylus by
its combination of usually ebracteolate pedicels,
green or yellow corolla often suffused with maroon
or purple, large falcate or reflexed dorsal corolla
lobes, the wide open orifice of the anther tube, bac¬
cate often inflated fruit, and oblong to fusiform
seeds much longer than broad. The genus was di¬
vided into two sections by Wimmer (1932, 1943,
1968): Burmeistera (“Imberbes,” nom. invalid.),
with all five anthers sparsely soft-hairy or glabrous
at apex; and Barbatae F. E. Wimmer, with the ven¬
tral pair of anthers densely bearded at apex.

1. Burmeistera arbusculifera Lammers, sp.
nov. T\ PE: Ecuador. Carchi: Espejo, El Gual-
tal, Cerro Golondrinas Hembra, 00°5I'N,
78°08'W, bosque muy humedo montano bajo,
bosque alto dominado por Clusia, dr boles de
25 m de altura, 2800 m, 21 Aug. 1994, W.
Palacios & ./. Clark 12465 (holotype, OSH;
isotypes, MO, QCNE not seen). Figure 1.

Ab omnibus caeteris speciebus Burmeisterae indumento
trichomatum arbusculiformium differt; species sect. Bar-
batarum affinis B. asperae , sed all hae specie hypanthio
obconico 3.5—5 mm diametro basi cuneato, calycis lobis
longioribus 7—1 1 mm longis, corolla breviora 30 mm longa
cum tubo 15 mm longo sed ventralibus lobis 10—1 I mm
longis, et dorsalibus antheris brevioribus 4.2 mm longis
distinguenda.

Hemiepiphytic liana; stems moderately leafy,
1.8— 2.2 mm diam. toward apex, up to 26 mm diam.
at base, branched, sparsely to moderately pubes¬
cent with arbusculiform trichomes toward the apex,
scabrid (through abrasion of these trichomes) be¬
low; latex c ream-colore'd. Lamina ovate, 2.3— 5.5 X
1.1— 2.8 cm, subcoriaceous; adaxial surface dull

8 2cm

Figure 1. Burmeistera arbusculifera Lammers. — A. Ar-
busculiform trichomes on abaxial leaf surface. — B. Upper
portion of stem with flower. (Drawn from t lie holotype, Pa¬
lacios Clark 12465.)

dark green, glabrous; abaxial surface dull green-
white, pubescent especially on the veins with ar¬
busculiform trichomes; margin somewhat revolute,
denticulate with remote dark callosities; apex acu¬
minate; base rounded or obtuse; petiole 5—10 mm
long, 0.6— 1.3 mm diam., 1/4— 1/7 as long as the
lamina, pubescent with arbusculiform trichomes.
Flowers solitary in the upper leaf axils; pedicels
straight or somewhat curved, ascending, 17—29 mm
long, 0.6— 0.8 mm diam., ebracteolate, glabrous.
Hypanthium obconic, 6—9 mm long, 3.5—5 mm
diam., 1/5— 1/3 as long as the corolla, glabrous; base
cuneate, distinct from pedicel. Calyx lobes lance¬
olate, ascending, 7—11 X 2—3.5 mm, as long as the
hypanthium up to half again as long, 1/2— 3/4 as
long as the corolla tube, glabrous; apex acute; mar¬
gin denticulate with 3 dark callosities on each side.
Corolla bilabiate, pale green irregularly blotched
with purple, 30 mm long, glabrous; tube suberect,
15 mm long, 6 mm diam. at base, gradually taper¬
ing to 3 mm diam. at middle and mouth; dorsal
lobes lanceolate, falcate, 15 X 3.5 mm, as long as
the tube, the apex acute; lateral lobes ovate, de-
flexed, 1 1 X 5—5.5 mm, the apex acute; ventral
lobe 10 X 5.5 mm, the apex acute. Staminal col¬
umn exserted between the dorsal lobes; filament

208

Novon

tube suberect, 20 mm long, 1 .7 mm diam. at mouth
of corolla, flaring to .1 mm diam. at base of anther
tube, glabrous; anther tube 3.2 mm diam., slightly
deflexed, its surfaces glabrous; dorsal anthers 4.2
mm long, 1/7 as long as the filament tube; ventral
anthers 2.8 mm long, their apex sparsely pubescent
with soft wispy hairs 0.5 mm long. Fruit and seeds
not seen, but apex of ovary flat.

Distribution , habitat, and phenology. Appar¬
ently endemic to northern Ecuador, collected twice
in the Cerro Golondrinas of Carchi. Growing in wet
mossy upper montane forests at elevations of 2750—
2800 m, and flowering during July and August. Ac¬
cording to brad Boyle (pers. comm.), the paratype
was collected in a sheltered forested cove between
ridges. The canopy was 15—18 m tall and consisted
of Weinmannia (Gunoniaceae), Miconia (Melasto-
mataceae), Clusia (Clusiaceae), Myrsinaceae, and
Lauraceae, while the shrub layer comprised a va¬
riety of Melastomataceae and Rubiaceae.

Etymology. The specific epithet alludes to the
distinctive branched trichomes; from the Latin
noun arbusculae, little trees, and the adjectival suf¬
fix -fer, to bear.

Relationships. Arbuscul il’orm or dendritic hairs
(i.e., trichomes irregularly branched above a
stalked base) characterize Centropogon sect. Siph-
ocampyloides and section Niveopsis (see below), and
also occur in four species of Siphocampylus (one
described herein) and one species of the endemic
Hawaiian genus Cyanea Gaudichaud (hammers,
1998a, 1998b). However, these unusual trichomes
have not previously been reported from species cur¬
rently assigned to Burmeistera. Although W dinner's
(1943) monograph did include three species of Bur¬
meistera with arbusculiform hairs \B. asteriscus F.
E. Wimmer, B. macrocarpa (Zahlbruckner) F. E.
Wimmer, and B. peruviana F. E. Wimmer], Mc-
Vaugh (1949) later transferred them to Centropo¬
gon.

On tilt' basis of l lit- pubescent apex of its ventral
anthers, B. arbusculifera is assigned to Burmeistera
sect. Barbatae. In Jeppesen’s (1981) treatment of
Ecuadorean Burmeistera, B. arbusculifera keys
readily to B. aspera F. E. Wimmer, a poorly known
species of that section that occurs in nearby Im-
babura as well as in southern Colombia. The two
species are similar in habit; size, shape, and other
details of their leaves; length ol pedicels; shape and
margin of the calyx lobes; anti pigmentation of the
corolla. However, B. aspera differs from the new
species in having the upper portions ol the stem
and the abaxial leaf surface pubescent with tlistally
curved articulate hairs (vs. arbusculiform hairs in

B. arbusculifera)-, hypanthium subglobose, 5—6 mm
diam., truncate at base (vs. obconic, 3.5-5 mm
diam., cuneate at base); calyx lobes 4—5 mm (vs.
7-11 mm) long; corolla 39 mm (vs. 30 mm) long,
with tube 25 mm (vs. 15 mm) long and the lateral
and ventral lobes 5—6 mm (vs. 1 0-1 1 nun) long;
and dorsal anthers 6 mm (vs. 4.2 mm) long. Overall,
on the basis of morphology anti geographic prox¬
imity, B. aspera would appear to be the closest
known relative of B. arbusculifera. Ihtj possibility
of a developmental relationship between the artic¬
ulate hairs of the former anti the arbusculiform tri¬
chomes of the latter is intriguing anil merits further
study.

Paratypes. ECUADOR. Carchi : Cerro Golondrinas,
valley bottom ca. 1.5 km NNE of summit, transect 2750—
2, 00°51'N, 78°08'W, mossy upper montane forest, with
tall trees to 30 m high near creek margin, stunted elfin
forest on ridge crest. 2750 m. 25 July 1994, H. Boyle. A.
Boyle, ./. Bradford & A '. Skinner 3451 (MO. OSfl. QCNE
not seen).

2. Burnicistcra brigbaniioitles hammers, sp.
nov. TYPE: Ecuador. Carchi: further ascent of
Rio Verde past stream and waterfall entering
from SW and continuing beyond principal
drainage stream of large Cerro Golondrinas
into drainage streams of medium Golondrinas
mountains, 00°52'N, 78°07'W. forest area at
stream embankment above stream and forest
area, terminating at main stream division,
1200 m, I Dec. 1987. W. S. Hoover 2182 (ho-
lotype, MO). Figure 2.

Species insignis egregie distincta ah omnibus caeteris
speciebus Burmeisterae corolla longissima 78 mm longa
duplo vel triplo longiore speciebus generis cognitis cum
tuho gracili decemplo longiore quam latiore et calycis lob-
is at! marginem ciliatis cum trichomatihus isabellinis.

Suffruticose herb, 2 m tall; stems moderately
leafy, 7 mm diam., the portions seen unbranched,
glabrous; latex white. Lamina obovate, 10.5—21 X
5.2-1 1 cm. ebartaceous; atlaxial surface dull green,
glabrous; abaxial surface dull white-green, pubes¬
cent along the veins with sordid yellow trichomes;
margin minutely denticulate with remote dark cal¬
losities; apex rounded with a small acute lip; base
cuneate; petiole 12—35 mm long, 1.5— 3.5 mm
diam., 1/5-1/10 as long as the lamina, glabrous.
Flowers solitary in the upper leaf axils; pedicels
straight, ascending, 62 mm long, 1.4 mm diam.,
ebracteolate, glabrous. Hypanthium oblate, 8 mm
long. 9 mm diam., 1/10 as long as the corolla, gla¬
brous; base truncate. Calyx lobes debate, erect,
2.5—3 X 2.5 mm, ca. 1/3 as long as the hypanthium
anti 1/20 as long as the corolla tube; apex obtuse;

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

209

Figure 2. Burmeistera brighamioides Lammers. — A. Or¬
ifice of anther tube. — B. F lower. — C. Pubescence along
abaxial midrib. — D. Leaf. (Drawn from the holotype. Hoo¬
ver 2182.)

margin entire, densely ciliate with sordid yellow tri-
chomes. Corolla tubular, bilabiate, green, 78 nun
long, glabrous; tube suberect, 58 mm long, 5.5 mm
diant. at base and middle, very gradually tapering
to 8 mm diam. at mouth; dorsal lobes lanceolate,
falcate, 20 X 4.5 mm. 1/3 as long as the tube, the
apex acuminate; ventral lobes ovate, deflexed, 8 X
4 mm. the apex acuminate. Staminal column slight¬
ly exserted between the dorsal lobes; filament tube
suberect, 64 mm long, 3 mm diam., glabrous; an¬
ther tube 4.5 mm diam., slightly deflexed, densely
pubescent on the sutures with long spreading sor¬
did yellow hairs; dorsal anthers 9 mm long, 1/7 as
long as the filament tube; ventral anthers 7 trim
long, with dense apical tufts of sordid yellow hairs
1.6— 1.8 mm long. Fruit and seeds not seen, but
apex of ovary flat.

Distribution, habitat, and phenology. Appar¬
ently endemic to northern Ecuador and known only
from the type specimen.

Etymology. The specific epithet calls attention
to the uncanny resemblance of the long slender
flower and obovate leaf to those of the endemic Ha¬
waiian genus Brighamia A. Gray (cf. Lammers,
1989).

Relationships. Burmeistera brighamioides dif¬
fers from all previously known members of the ge¬
nus in the incredible length of its corolla (but see
below). Most congeners have corollas between 25
and 35 mm long; reported extremes are as little as
12 mm in B. kirkbridei Wilbur and as much as 45
mm in B. utleyi Wilbur. At 78 mm, the corolla of
B. brighamioides is two to three times longer than
the average congener. Furthermore, the tube of the
corolla is extremely slender, about 10 times longer
than broad, lacking basal or apical distensions, and
is three times longer than the dorsal lobes; corolla
tubes elsewhere in the genus are typically two to
seven times longer than broad, distended at base
and/or throat, and about as long as the dorsal lobes
or up to twice as long. The dense edging of sordid
yellow' trichomes on the margin of the calyx lobes
also appears to be unique within the genus.

On the basis of its densely bearded ventral an¬
thers, B. brighamioides would be assigned to Bur¬
meistera sect. Barbatae. Beyond that, it is not pos¬
sible to infer a close relationship to any previously
described species (but see below). In Jeppesen’s
(1981) treatment, it keys with difficulty to B. cer-
atocarpa H. Karsten of western Colombia and
northern Ecuador (including Carchi), which shares
with it an oblate, basally truncate hypanthium.
However, that species differs in having leaves wili¬
est at or below the middle, longer (13—16 mm) lin¬
ear glabrous calyx lobes, and shorter (5-6 mm) dor¬
sal anthers, as well as by its far shorter (20—25 mm)
corolla with basally distended tube 4—7 times lon¬
ger than wide and about as long as the lobes.

3. Ilurineistera fimhriula Lammers, sp. nov.
TYPE: Colombia. Huila: W slope of Cordillera
Oriental, 29 km above (SE of) Guadalupe on
road to Florencia, ca. 1.5 km W of Caqueta
Dept, limit, 01°55'N, 75°43'W, 2250-2350 m,
22 Mar. 1986, B. A. Stein 3706 (holotype,
MO). Figure 3.

Ah omnibus caeteris speciebus Burmeisterae foliorum
marginibus fimbriatis differt; species sect. Barbatarum ob
pedicellos bibracteolatos et calycis lobos basi connatos
affiriis B. venezuelensi, sed ab hac specie lamina elliptica
basi attenuata, bracteolis longioribus 8—9 mm longis, hy-
panthio obconico, calycis lobis longioribus 26-30 mm lon¬
gis distinguenda.

F leshy herb to 1 m tall; stems moderately leafy.
6—6.5 mm diam., the portions seen unbranched,
densely pubescent with long crisped septate tri¬
chomes; latex white. Lamina elliptic, 12-23 X 4.5—
8.8 cm; adaxial surface dull green, sparsely pubes¬
cent with long crisped septate trichomes; abaxial
surface dull white-green, sparsely pubescent with

210

Novon

Figure 3. Bur me is ter a Jim b ria t a Lammers. — A. Upper
portion of stem with flower. — B. Leaf margin. (Drawn from
the holotype, Stein 3706.)

long crisped septate trichomes; margin fimbriate
with 4 to 8 fimbriae per cm. flic fimbriae a mixture
of short (0.8— 1.5 mm long) and long (2.5— 3.4 mm
long) and each pubescent with long crisped septate
trichomes; apex cuspidate; base attenuate; petiole
20—40 mm long. 145—2.2 mm diam., 1/5— I /8 as
long as the lamina, pubescent with long crisped
septate trichomes. f lowers solitary in the upper leaf
axils; pedicels straight, ascending, 110—125 mm
long. 1.2— 1.3 mm diam.. bibraeteolate at base, pu¬
bescent with long crisped septate trichomes; brac-
teoles linear, 8—9 X 0.5 mm, the apex acuminate.
Hypanthium obeonic. 14 mm long, 9 mm diam., 1/3
as long as die corolla, pubescent with long crisped
septate trichomes; base cuneate. Calyx lobes linear-
triangular, erect or spreading slightly, 26—30 X 3
mm, about twice as long as the hypanthium and
equaling the corolla tube, connate at base for 1-2
mm, pubescent with long crisped septate trichomes;
apex acuminate; margin sparsely dentate with 6 to
10 long callosities on each side. Corolla bilabiate,
green, 45 mm long, glabrous; tube subereet, 30 mm
long, 9 mm diam. at base, gradually tapering to 4.5
mm diam. at middle before gradually increasing to
12 mm diam. at mouth; dorsal lobes triangular, fal¬
cate, 15 X 5 mm, hall as long as the tube, the apex
acuminate; lateral lobes triangular, deflexed. 13 X
6 mm, the apex acuminate. Staminal column slight¬

ly exserted between the dorsal lobes; filament tube
suberect. 36 mm long; anther tube 5 mm diam.,
gently deflexed, its surfaces glabrous; dorsal an¬
thers 10 mm long, ca. 1/4 as long as the filament
tube; ventral anthers 8 mm long, their apex densely
bearded with sordid hairs 1.5 mm long. Fruit and
seeds not seen, but apex ol ovary flat.

Distribution, habitat , and phenology. Appar¬
ently endemic to the western slope of the Cordillera
Oriental in southwestern Colombia, and known only
from the type specimen.

Etymology. The specific epithet alludes to the
distinctive leaf margin; from the Latin adjective
jimbriatus, fringed.

Relationships. On the basis of its densely
bearded ventral anthers, Burmeistera Jimbriata is
referable to section Rarbatae. Its fimbriate leaf mar¬
gin is unique in the genus and thus not helpful in
assessing relationships more precisely. However, its
bibraeteolate pedicels and basally connate calyx
lobes, while unusual, are not unique. These fea¬
tures are found in two other members ol section
Barbatae : B. caldasensis (Gleason) F. K. Wimmer
of west-central Colombia and B. venezuelensis Lam¬
mers of southwestern Venezuela. Like B. Jimbriata,
these two species are fleshy understory herbs up to
1 m tall with toothed calyx lobes equaling or ex¬
ceeding the hypanthium. Based on these similari¬
ties. it seems clear that B. caldasensis, B. Jimbriata,
and B. venezuelensis form a close-knit if not raono-
phyletie group of species.

Of the two previously described species, B. ve¬
nezuelensis seems more similar to the novelty, re¬
sembling it in its septate trichomes, large leaves
(lamina 8.5-1 1.5 X 4.5— 6.8 cm), and long pedicels
(80—135 mm). In contrast, the other previously de¬
scribed species, B. caldasensis. bears only unicel¬
lular trichomes and has smaller leaves (lamina 4.5—
6.5 X 2-2.5 cm) and shorter pedicels (4 cm long).
However, B. caldasensis does resemble the novelty
in its longer bracteoles (8 mm long) and obeonic
hypanthium; B. venezuelensis has bracteoles just
2.7—4 mm long and an oblate hypanthium.

t. Burmeistera liippobronioides Lammers, sp.
nov. I V PF: Colombia. Valle: Yataeue, CVC
camp at Anchicaya, 03°40'N, 76°50'W, mature
premontane wet forest, 700—900 m, 25 Feb.
1983, A. Gentry, A. Juncosa & F. Gomez 40746
(holotype, MO). Figure 4.

Species Burmeisterne sect. Barbatarum ol) corollam gra-
eilem longissimam affinis B. brighamioidi, sed ah hac spe¬
cie laminis oblanceolatis angustioribus 3.8— 4.4 cm latis,
hypantliio obeomco angustiore 5 mm diametro basi cu-

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

211

Figure 4. Burmeistera hippobromoides Lammers. (Drawn
from the holotype. Gentry el al. 40746).

rieato, calyois lobis lanceolatis vel anguste triangularis
longioribus 13,5-16 mm longis ad marginem glabris, et
corolla pubescenli longiore 62 mm longa statim distin-
guenda.

Shrub, 2 m tall; stems moderately leafy, 9 mm
diam., the portions seen unbranched, sparsely
short-pubescent. Lamina oblanceolate, 12—15 X
3.8-4. 4 cm; adaxial surface dark green, glabrous;
abaxial surface green, sparsely short-pubescent;
margin subentire, somewhat sinuate; apex acute or
cuspidate; base attenuate; petiole 10-18 mm long.
1.5— 1.9 mm diam., 1/8—1/12 as long as the lamina,
sparsely short-pubescent. Flowers solitary' in the
upper leaf axils; pedicels straight, ascending, 35-
60 mm long, 1.4— 1.5 mm diam., ebracteolate,
sparsely short-pubescent. Hypanthium obconic, 9
mm long, 5 mm diam., 1/10 as long as the corolla,
sparsely short-pubescent; base cuneate. Calyx
lobes lanceolate or narrowly triangular, erect, 13.5—
16 X 2—3.5 mm, ca. 1 I/2X as long as the hypan¬
thium and 1/4— 1/5 as long as the corolla tube; apex
acuminate; margin entire. Corolla tubular, bilabi¬
ate, green, 92 mm long, densely short-pubescent;
tube suberect, 67 mm long, 4.5 mm diam. at base
and middle, very gradually tapering to 8 mm diam.
at mouth; dorsal lobes lanceolate or narrowly tri¬
angular, falcate, 25 X 4—4.5 mm, ca. 1/3 as long
as the tube, the apex acuminate; lateral lobes nar¬
rowly triangular, falcate, 18 X 3—3.5 mm, the apex

acuminate; ventral lobe narrowly triangular, de-
flexed. 10 X 4.5 mm. the apex acuminate. Staminal
column slightly exserted between the dorsal lobes;
filament lube suberect, 77 mm long; anther tube 5
mm diam., slightly deflexed; dorsal anthers 9 mm
long. ca. 1/8 as long as the filament tube; ventral
anthers with dense apical tufts of white curly hairs
2 mm long. Berry globose, 12 mm diam.; seeds ob¬
long. 0.8 mm long. 0.3 mm diam., honey brown, the
testa minutely reticulate.

Distribution , habitat . and phenology. Apparently
endemic to the Cordillera Occidental of southwestern
Colombia and known only from the type specimen.

Etymology. It seems appropriate to christen the
only two species of Burmeistera with long slender
flowers with the names of the only two genera in
the subfamily with long slender flowers. Just as R.
brighamioides takes its name from the Hawaiian
endemic Brighamia, this novelty takes its name
from the Jamaican endemic Hippobroma G. Don.

Relationships. The discovery of one species of
Burmeistera with inordinately long flowers, B.
brighamioides, was quite remarkable. The discov¬
ery of a second species with such flowers taxes cre¬
dulity. Like Ecuadorean B. brighamioides (see
above), B. hippobromoides of Colombia differs from
all other known members of the genus in bearing a
corolla two to three times longer than the average
congener, with a slender lube that is almost three
times longer than the dorsal lobes and lacks basal
or apical distensions. The two species are also sim¬
ilar in length and posture of their pedicels, diam¬
eter of the corolla tube, size and shape of the co¬
rolla lobes, and size and apical pubescence of the
anther tube. Both are members of Burmeistera sect.
Barbatae.

However, the Colombian novelty differs from B.
brighamioides in a number of features. The leaves
of B. hippobromoides are oblanceolate (vs. obovate
in B. brighamioides ) and narrower (3.8-44 cm
wide vs. 5.2—1 1 cm). The hypanthium is obconic
(vs. broadly ovoid) and narrower (5 mm diam. vs.

9 mm), with a cuneate (vs. truncate) base. The ealvx
lobes are lanceolate or narrowly triangular (vs. del-
tate) and far longer (13.5—16 mm vs. 2.5—3 mm),
and the margin lacks the dense edging of sordid
yellow trichomes seen in B. brighamioides. The co¬
rolla is even longer than that of B. brighamioides
(92 mm vs. 78 mm), and densely short-pubescent
(vs. glabrous).

In light of the unique corolla they share, it seems
reasonable to regard B. brighamioides and B. hip¬
pobromoides as sister-species, i.e., to hypothesize
that their unique floral morphology has evolved just

212

Novon

Figure 5. Burmeislera knaphusii hammers. (Drawn from
the holotype, lloover 2276.)

once in the genus. One is further tempted to hy¬
pothesize that this extraordinary shift in corolla
morphology is correlated with a shift in pollen vec¬
tor. Unfortunately, almost nothing is known about
pollination in Burmeistera. Vogel ( 1969) document¬
ed chiropterophily in B. fuscoapicata F. E. Wimmer,
while Euteyn (1986) suggested that H. pinnatisecta
Luteyn was hummingbird pollinated. Conceivably,
the long slender straight corollas of B. brighamioi-
des and H. hippobromoides could be an adaptation
to pollination by hawkmoths (Lepidoptera: Sphin-
gidae), a syndrome hypothesized for both Brigh-
amia and Hippobroma (cf. hammers. 1989).

5. Burmeistera kiiaplmsii hammers, sp. nov.
TYPE: Ecuador. Carchi: mountain E & NE,
forest area along slope of mountain ENE of
Rafael Quindfs mountain finca and above Rfo
Verde including top of mountain which is sim¬
ilar in geologic format to Golondrinas Moun¬
tains, 00°52'N, 78°07'W, 1870-2400 m, 3
Dec. 1987, W. S. Hoover 2276 (holotype, MO).
F igure 5.

Species Burmeisterae sect. Barbatarum affinis B. h n
acamayensi, sed ah hae specie septatis trichomatibus lon-
giusculis patentihus in caulibus pedicellisque, laminis
ovatis vet anguste ovatis brevioribus 2.5-5 cm longis ap-
ice acuminatis basi rotundis ve! truncatis, pedicellis lirc-
viorit)us 10—12 mm longis, et floribus parvioribus cum
hypanthio 3.5-4 mm longo, corolla 29-32 mm tonga, cor-
oilac tubo 3—3.8 mm basi diametro, et lobis venlralibus
4.5—6 mm longis distinguenda.

Hemiepiphytic liana to 4 m long; stems moder¬
ately leafy, 1.4-2. 2 mm diam.. the portions seen
unbranched, pubescent with long spreading septate
trichomes. hamina ovate or narrowly ovate, 2.5—5
X 1.2-2. 2 cm, subcoriaceous; adaxial surface dull
green, glabrous; abaxial surface dull pale green,
pubescent primarily on the veins with long spread¬
ing septate trichomes; margin minutely crenulate
with remote callosities, a pair toward the apex es¬
pecially prominent; apex acuminate; base rounded
or truncate; petiole 2—3.5 mm long, 0.5 mm diam.,
1/12-1/20 as long as the lamina, pubescent with
long spreading septate trichomes. Flowers solitary
in the upper leaf axils; pedicels straight or some¬
what curved, ascending. 10-12 mm long. 0.5-0. 6
mm diam.. ebracteolate, pubescent with long
spreading septate trichomes, at least at base, lly-
panthium obeonic, 3.5—4 mm long, 1.8—3 mm
diam.. 1/8 as long as the corolla, glabrous or with
a few scattered long spreading septate trichomes;
base cuneate, ± distinct from pedicel. Calyx lobes
narrowly triangular or linear-triangular, reflexed, 4—
6 X 0.7— 1.3 mm, as long as the hypanthium up to
half again as long, 1/5— 1/3 as long as the corolla
tube, glabrous or with a few scattered long spread¬
ing septate trichomes; apex acute; margin dentic¬
ulate with 2—3 callosities on each side. Corolla bi¬
labiate. pale green faintly suffused with lavender or
red, 29—32 nun long, glabrous; tube suberect, 19—
21 mm long, 3—3.8 mm diam. at base, gradually
tapering to 2.2—3 mm diam. at middle and 2.8— 3.6
mm diam. at mouth; dorsal lobes lanceolate,
spreading, 10-12 X 3—3.7 mm. 1/2-3/5 as long as
the tube, the apex acuminate; lateral lobes ovate,
deflexed, 5-6 X 2. 2-3. 5 mm. the apex acuminate;
ventral lobe 4.5-5 X 1. 8-2.6 mm, the apex acu¬
minate. Staminal column slightly exserted between
the dorsal lobes; filament tube suberect, 24—28 mm
long, 0.8— 1.1 mm diam. at mouth of corolla, flaring
to 1.3— 1.5 mm diam. at base of anther tube, gla¬
brous; anther tube 2.5-3 mm diam.. deflexed, its
surfaces glabrous; dorsal anthers 3.2— 4.5 mm long,
1/5-1/10 as long as the filament tube; ventral an¬
thers 2.2-3 mm long, their apex sparsely pubescent
with soft wispy hairs 0.2-0. 4 mm long. Fruit and
seeds not seen, but apex of ovary flat.

Distribution , habitat, and phenology. Apparently
endemic to northern Ecuador, collected twice in the
Cerro Golondrinas of Carchi. Growing in montane
forests at elevations of 1870—2430 m, and flowering
during December.

Etymology. This species is respectfully dedi¬
cated to the memory of George Knaphus (1924-
2000), botanist and educator extraordinaire. Erom

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

213

1965 until li is death, “Dr. K.” was professor of bot¬
any at Iowa State University. In classroom, labora¬
tory, and field, he opened the eyes of thousands of
undergraduate students to the marvels of the plant
and fungal kingdoms. Furthermore, his personal
approach to academic advising and genial sponsor¬
ship of Botany Club (with colleague Fois II. Til(any)
provided a level of support, guidance, and encour¬
agement to young men and women that is rare in¬
deed in today’s academic world. The epithet coined
here is in accordance with Recommendation 60C.2
of the International Code of Botanical Nomencla¬
ture (ICBN; Greuter et al., 2000). Although his sur¬
name resembles a second declension Latin noun, it
is actually ol Norwegian origin; the initial k is pro¬
nounced and ph is not a digraph.

Relationships. On the basis ol its apically pu¬
bescent ventral anthers, Burmeistera knaphusii is
assigned to section Barbatae. In Jeppesen’s (1981)
treatment, it keys with difficulty to B. huacamay-
ensis Jeppesen, a member of that section endemic
to the Cordillera de Huacamayos in the Ecuadorean
province ol Napo. The two species share a number
of features, including habit; hypanthium shape;
size, shape, ami reflexion of the calyx lobes; color
and shape ol the corolla; and various aspects of the
staminal column. However, B. huacamayensis dif¬
fers from the novelty in its lack of hairs on stems
and pedicels, longer (9—12 cm vs. 2.5—5 cm in B.
knaphusii) lanceolate (vs. ovate or narrowly ovate)
leaves that are cuspidate (vs. acuminate) at apex
and cuneate or obtuse (vs. rounded or truncate) at
base; longer (6—7 cm vs. 1 — 1.2 cm) pedicels; and
generally larger flowers, with the hypanthium 5—6
mm long and corolla 37 mm long with the tube 6
mm in diameter at base and the ventral lobes 9 mm
long. Overall, B. huacamayensis would appear to
be the closest known relative of B. knaphusii.

Paratype. ECU A DOR. Carehi: forest and ridge area
above Rfo Verde and ridge flanking medium Cerro Golon-
drinas, 00°52'IN, 78°07'W, 2070-2430 m, 4 Dee. 1987,
W. S. Hoover 2322 (MO).

6. Burmeistera smaragdi Lammers, sp. nov.
TYPE: Ecuador. Esmeraldas: Quininde, Rilsa
Biological Station, Montanas de Mache, 35 km
W of Quininde, 5 km W of Santa Isabel, road
between the station and the SE Ridge Trail,
00°21'N, 79°44'W, premontane wet forest,
400—600 m, 19 Sep. 1994, M. S. Bass & N.
Pittman 22 (holotype, OSH; isotypes, MO,
QCNE not seen). Figure 6.

Species Burmeisterae sect. Burmeisterae affinis B. gla-
bratae et B. truneatae, sed ah bis speciebus petiolis bre-
vioribus 3—7 mm longis, foliorum apice caudato, hypan-

Figure 6. Burmeistera smaragdi Lammers. (Drawn from
the holotype, Bass & Pittman 22.)

thio anguste obconico basi attenuato, corolla breviore 24—
30 mm longa, dorsal i bus antheris brevioribus f> — 7 mm lon¬
gis basi crinito in suturis, et antheris omnibus ad apicem
glabris distinguenda.

Scandent shrub, 0.5—1 m tall; stems moderately
leafy, 1.3—2 mm diam., the portions seen un¬
branched, glabrous; latex white. Lamina narrowly
elliptic or narrowly oblong, 6.5—12.5 X 2—4.2 cm,
chartaceous; adaxial surface dull olive green, gla¬
brous; abaxial surface dull white-green, the midrib
pubescent with long septate hairs; margin subentire
or minutely denticulate with remote tiny dark cal¬
losities; apex caudate; base cuneate; petiole 3—7
mm long, 0.6— 0.8 mm diam., 1/12—1/20 as long as
the lamina, pubescent with long septate hairs.
Flowers solitary in the upper leaf axils; pedicels
straight, ascending, 37—66 mm long, 0.8- 1.4 mm
diam., ebracteolate, glabrous. Hypanthium narrow¬
ly obconic, 6—1 1 mm long, 4.5— 5.5 mm diam., 1/4—
2/5 as long as the corolla, glabrous; base cuneate,
tapering imperceptibly into the pedicel. Calyx lobes
narrowly triangular or linear-triangular, straight, as¬
cending or spreading, 12—19 X 1—2.5 mm, 2—3
times as long as the hypanthium, equaling to 3/4
again as long as the corolla tube, glabrous; apex
acuminate; margin minutely denticulate with 3 to 5
tiny dark callosities on each side. Corolla bilabiate,
green or cream-colored, often tinged or striate with
maroon or purple, 24—30 mm long, glabrous; tube

214

Novon

suberect, 1 1.5—13 mm long, 3.5—4 mm diam. al the
abruptly inflated base, 2.3— 2.5 mm diam. at mid¬
dle, gradually flaring to 2.5— 3.3 mm diam. at
mouth; dorsal lobes lanceolate, falcate, 12—18.5 X
2.3^4 mm, about as long as the tube to 2/3 again
as long, the apex acuminate; lateral lobes lanceo¬
late, dellexed, 8—13 X 2.2—3 mm, the apex acu¬
minate; ventral lobe 6.5—11.5 X 2.2—3 mm, the
apex acuminate. Staminal column exserted between
the dorsal lobes; filament Lube suberect, 21—21.5
mm long, 1.2— 1.3 nun diam. at mouth of corolla,
flaring to 2.7—3 mm diam. at base of anther tube,
glabrous; anther tube 3.5— 4.5 mm diam., slightly
deflexed, pubescent at the base of the dorsal su¬
tures with long crisped hairs, the orifice glabrous;
dorsal anthers 6—7 mm long, 1/5—3/10 as long as
the filament tube; ventral anthers 3.5— 4.5 mm long.
Fruit and seeds not seen, but apex of ovary flat.

Distribution , habitat , and phenology. Apparently
endemic to northern Ecuatlor, where it has been col¬
lected several times at the Hi Isa biological Station in
the Mache Mountains of Esmeraldas. It grows in wet
premontane forests at elevations of 100-600 m, and
dowel's during September.

Etymology. The specific epithet is the genitive
of the Eatin noun smaragdus, emerald, in reference
to the Ecuadorean province to which the species is
apparently endemic.

Relationships. With its glabrous anther tube or¬
ifice, Burmeistera smaragdi is referable to section
Burmeistera. In Jeppesen s (1981 ) treatment, it keys
to two species of that section: B. glabrata (Kunth)
Bentham & Hooker ex B. I). Jackson, which is
found from central Colombia to southeastern Ec¬
uador; and B. truncata Zahlbruckner, endemic to
the Ecuadorean province of Pichincha. These three
species share a number of features, including gen¬
eral size and shape of their leaves, pedicels shorter
than the subtending leaf, erect or spreading calyx
lobes not overlapping at base and longer than both
the hypanthium and corolla tube, and glabrous
green corollas often suffused with darker hues.
However, B. glabrata and B. truncata differ from
B. smaragdi in their longer petioles (1-2 cm vs.
0.3— 0.7 cm in B. smaragdi ); acute or acuminate (vs.
caudate) leaf apices; globose, urceolate, or semi-
ovoid hypanthium rounded or obtuse at base (vs.
narrowly obconic and attenuate at base); corolla
30—37 mm long (vs. 24—30 mm long); and anther
tube glabrous on the surface (vs. pubescent at base
of dorsal sutures) and villous at the apex (vs. gla¬
brous), with the dorsal three 8—10 mm long (vs. 6—
7 mm long). Overall, B. smaragdi seems most
closely related to B. glabrata and B. truncata.

Paratypes. ECUADOR. Esmeraldas: Quininde, Hilsa
Biological Station, Montanas de Mache, 20 km INVt of
Quininde and 3 km W of Santa Isabela, southern bound¬
ary, SW of cabin, 00°22'N, 79°45'W, premontane wet for¬
est, open area, 000 m, 24 Sep. 1994. ,/■ A’. Abbott 15243
(MO, QCNE not seen); Quininde, Bilsa Biological Station,
Mache Mountains, 35 km W of Quininde, 5 km W of
Santa Isabel, 00°21'N. 79°44'W, premontane wet forest,
on edge of clear cut and mature forest, 400—600 m, 13
Sep. 1994, ,/. L. Clark N B. Adnepos 27 (MO. OSH. QCNE
not seen): Quininde Canton, Mache-Chindul Ecological
Reserve, Bilsa Biological Station, Mache mountains, 35
km W of Quininde, 5 km W of Santa Isabel, 00°2I’IN,
79°44'W, premontane wet lores!, 500 m, 24 Sep. 1996,

I. . Clark 2902 (MO, OSH. QCNE not seen).

II. Centropogon C. Presl, Prodr. Monogr. Lobel.

48. 1836. TYPE (lectotype, designated by

Pfeiffer (1873: 650)): Lobelia surinamensis E.

[= Centropogon cornutus (L.) Druce].

With the addition of the six novelties described
here, Centropogon comprises 222 species, distrib¬
uted from southern Mexico to Bolivia and Brazil
and in the Lesser Antilles (Cam triers, 1998a). The
genus differs from Burmeistera and resembles Siph¬
ocampylus in its usually bibracteolate pedicels;
commonly red, orange or yellow (sometimes green
or white) corolla; the occluded orifice of the anther
tube; and isodiametric seeds. Centropogon and
Siphocampylus are distinguished on the basis of
fruit type: baccate, from a flat-topped ovary, in Cen¬
tropogon ; capsular, from an ovary conical at apex,
in Siphocampylus.

Centropogon has been divided (hammers, 1998a)
into five sections primarily on the basis of floral
morphology anti pubescence: Centropogon , Sipho-
campyloides Bentham (divided into subsects. Brev-
ilimbati E. E. Wimmer and Peruviani McVaugh),
Wimmeriopsis McVaugh (divided into subsects. Fal-
cali McVaugh and Colombiani McVaugh), Burmeis-
teroides Gleason, and Niveopsis Lammers. A key to
these taxa is provided by Lammers (1998a), and
their salient features are summarized here.

Three major types of gross floral morphology may
be recognized in Centropogon : (1) “short-lobed” —
corolla tube 3-8 times longer than broad and 2-5
times longer than the triangular or oblong lobes; (2)
“long-lobed” — corolla tube 3—8 times longer than
broad and about as long as the triangular or oblong
lobes; and (3) “stout” — corolla tube twice as long
as broad and four times longer than the deflate
lobes, fhe “short-lobed” corolla type characterizes
section Centropogon, section Wimmeriopsis, and
subsection Brevilimbati ; the “long-lobed” corolla
type characterizes section Bunneisteroides and sub¬
section Peruviani-, and the “stout” corolla type is
unique to section Niveopsis.

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

215

Figure 7. Centropogon beniteziae Lammers. (Drawn from
the holotype, Luteyn & Cotton 9709.)

Two types of pubescence are important in the
sectional classification of Centropogon: the tuft of
triehomes at the apex of the ventral pair of anthers,
and those forming the indumentum on stems,
leaves, hypanthia, calyx lobes, and corollas. In tin1
autonymic section, the apical anther triehomes are
concrescent into a distinct triangular scale; these
hairs are free and brush- or beard-like in the re¬
maining sections. In section Siphocampyloides and
section Niveop.sis, the surface indumentum is com¬
posed primarily of arbusculiform (dendritic) tri¬
ehomes; in the other sections, any hairs that occur
are simple (with one exception, described below).

1. Centropogon beniteziae Lammers, sp. nov.
TYPE: Venezuela. Trujillo: Dpto. Bocono, Par¬
amo Cuaramaeal, 20—21 km beyond jet. with
hwy. NE of Bocono, 09°13'N, 70°13'W, shaded
very wet streambed with large boulders, infre¬
quent, 2640—2700 m, 14 Mar. 1984, L. Lu¬
teyn & E. Cotton 9709 (holotype, OSH; iso¬
types, NY, PORT not seen, YEN not seen).
Figure 7.

Species Centropogonis subsect. Hrevilimbatorum affinis
C. elmano et C. ewanio, sed all his speciebus lamina el-
liptica, pedicellis longioribus 5.8—12.5 cm longis, et ffo-
ribus majoribus cum hypanthio 6-8 mm longo 8—10 mm
diametro, calycis lobis 12—29 mm longis, corolla 48— 60
mm longa tubo 35—45 mm longo ad eonstrictionem 3—3.8
mm diametro ad orem 7.7—10.5 mm diametro dorsalibus
lobis 3—5 mm latis ventralibus lobis 2.8— 3.5 mm latis, et
tubo antherarum glabro distinguenda.

Erect or scandent suffruticose herb, 0.5— 1.5 m
tall; stems moderately leafy, 2—3.7 mm diam. above,
up to 6 mm diam. below, branched, pubescent with
arbusculiform hairs toward apex, glabrate below;
latex white. Lamina elliptic or widely elliptic (rare¬
ly ovate), 5—15.5 X 2.5— 7.7 cm, chartaceous; ad-
axial surface dull dark green, sparsely pubescent
(densest on midrib) with simple or sparingly
branched hairs; abaxial surface dull green often
suffused with maroon, densely pubescent with ar¬
busculiform hairs; margin denticulate or serrulate
with dark callosities; apex acuminate; base cune-
ate; petiole 10—30 mm long, 0.9— 1.5 mm diam.,
1/4— 1/8 as long as the lamina, pubescent with ar¬
busculiform hairs. Flowers solitary in the upper leaf
axils; pedicels ascending, 58—125 mm long, 0.9-
1.4 mm diam., bibracteolate at or just above tin*
base, pubescent with arbusculiform hairs. Hypan-
thium subglobose or very widely depressed globose,
6—8 mm long, 8—10 mm diam., 1/6— 1/9 as long as
the corolla, pubescent with arbusculiform hairs.
Calyx lobes narrowly triangular or linear-triangular,
ascending, 12—29 X 1.5—3 mm, half again as long
to almost 4 times as long as the hypanthium, 1/3—
1/2 as long as the corolla tube, pubescent with ar¬
busculiform hairs; apex acuminate; margin entire
or serrulate denticulate with I to 5 tiny curved
teeth on each side. Corolla tubular, somewhat ven-
tricose, orange, red, or dark red, paler or yellow
within, 48—60 mm long, pubescent with arbuseu-
liform hairs; tube suberect, 35-45 mm long, 5—7
mm diam. at the base, narrowing abruptly to 2.5—
3.8 mm diam. just above the base, then flaring
gradually to 7.5—10.5 mm diam. just below the
mouth; dorsal lobes narrowly triangular, falcate,
13—17 X 3—5 mm, ca. 1/3— 2/5 as long as the tube,
the apex acuminate; ventral lobes lanceolate, de-
flexed, 9-12 X 2.8— 3.5 mm, the apex acuminate.
Staminal column somewhat exserted between the
dorsal lobes; filament tube suberect, 43—54 mm
long, 1-1.4 mm diam., glabrous or sparsely pubes¬
cent with simple white crisped hairs; anther tube
2—2.6 mm diam., suberect, its surfaces glabrous;
dorsal anthers 7—9 mm long, 1/5— 1/7 as long as the
filament tube; ventral anthers 6-7 mm long, pu¬
bescent at apex with tufts of stiff yellow-white hairs
1.2— 1.7 mm long. Berry very widely depressed glo¬
bose, 8—12 mm long, 12—15 mm diam., the calyx
lobes tardily deciduous; seeds ellipsoid, golden
brown, 1.4 mm long, 0.4 mm diam., the testa mi¬
nutely reticulate.

Distribution, habitat, and phenology. Apparently
endemic to western Venezuela, where in recent years
it has been collected frequently in wet rocky stream-

216

Novon

beds, on steep roadside embankments, and in mon¬
tane forest and subparamo habitats in the area around
Guaramacdl and Bocon6 in die state of Trujillo, at
elevations of 1600-3175 m. Plants flower from late
October to July, and bear fruit from Apri 1 to July.

Etymology. This species is named in honor of
Carmen E. Benitez de Rojas (b. 1937) of the Univ-
ersidad Central de Venezuela in Maracay, curator
of the Herbario Victor Manuel Badillo (MY). I)ra.
Benitez is an authority on the Venezuelan flora, es¬
pecially Solanaceae, and has been very helpful to
me on several occasions in locating material of Ve¬
nezuelan Lobelioideae. Earlier authors have re¬
ferred to C. beniteziae as “Centropogon sp. " (Ortega
et ah, 1987) and “Centropogon sp. A” (Dorr et ah,
2000).

Relationships. With its arbusculiform trichomes
and “short-lobed ' corolla, Centropogon beniteziae is
clearly a member of section Siphocampyloides, sub¬
section Brevilimbati. However, it is unusual among
the members of that subsection in having at least
some simple hairs on the adaxial leal surface. Most
Brevilimbati bear only arbusculiform hairs on veg¬
etative surfaces; just two previously known species
bear simple hairs on the adaxial leaf surface: C.
ewanii F. E. Wimmer of western Venezuela and C.
heteropilis F. E. Wimmer of Colombia. In fact, many
of the specimens of C. beniteziae cited here were
originally identified as C. ewanii (ch Dorr et ah,
2000). However, in C. ewanii and C. heteropilis, all
of the hairs on the adaxial surface are strictly sim¬
ple, while C. beniteziae is variable, showing a con¬
tinuum from simple to sparingly branched hairs,
often on a single leal. Furthermore, these two spe¬
cies collectively differ from C. beniteziae in their
shorter pedicels (4—6 cm vs. 5.8—12.5 cm in C.
beniteziae ), shorter calyx lobes (4—7 mm vs. 12—29
mm), shorter corollas (35—44 mm vs. — 60 mm),
and pubescent (vs. glabrous) anthers.

Centropogon beniteziae is also quite similar to
another western Venezuelan endemic, C. e! man us
F. E. Wimmer. Both species are suffruticose plants
with dark-callose leaf margins, denticulate linear
calyx lobes 1.5 to 4 times longer than the depressed
globose hypanthium, and constricted orange corol¬
las. However, C. elmanus differs in its oblong
leaves (vs. elliptic in C. beniteziae ) bearing only
arbusculiform trichomes (vs. some simple hairs
adaxially), shorter pedicels (5 cm vs. 5.8—12.5 cm),
and smaller (lowers. Specifically, the hypanthium of
C. elmanus is 5 mm long (vs. 6-41 mm in C. beni¬
teziae) and 7 mm diam. (vs. 8-10 mm diam.), with
calyx lobes 10 mm long (vs. 12-29 mm long) and
1 mm wide (vs. 1.5—3 mm). The corolla is 43—44
mm long (vs. 48—60 mm); its tube 30 mm long (vs.

35 — 45 mm), 2 mm diam. at the constriction (vs.
2. 5-3.8 mm), and 5-6 mm at the mouth (vs. 7.5-
10.5 mm); and its lobes 2 mm wide (vs. 3—5 mm
in the dorsals, 2.8— 3.5 mm in the ventrals). The
staminal column is long exserted from the corolla
(vs. scarcely exserted) and the anther tube long pi¬
lose on the dorsal sutures (vs. glabrous). On the
basis of morphology and geography, C. elmanus and
C. ewanii appear to be the closest known relatives
of C. beniteziae.

Earatypes. VENEZUELA. Trujillo: Distr. Bocond,
Pdramo de ( iuaramacdl, SE of television towers, 09°14'N,
70° 1 l'W, along stream cascading over rocks, 2000 m, 28
Apr. 1988, /.. ./. Dorr , /,. C. Barnett. N. Cuello & G. M.
Diggs. Jr. 4987 (PORT); Distr. Bocond, Paramo de Cuar-
amaeal, \\ ol mad summit. 09° I t’N. 70°1 1 'YY. 2800-2900
m, 28 Apr. 1988, L. J. Dorr. L. C. Barnett. N. Cuello &•
G. M. Diggs. Jr. 5015 (NY not seen, PORT); Man. Bocond,
Parque Nacional Guaramacdl, road from Bocond to Guar-
amaeal, SE of Bocond, N slope of mountain, 09°13'N,
70°12'W, montane forest, 2550 m, 7 July 1995, L. ./. Dorr
& L. C. Barnett 8048 (PORT); Mun. Bocond, Parque Na-
eional Guaramacdl, road from Bocond to Guaramacdl, SE
of Guaramacdl, N slope of mountain. 09°13'N, 70°12'W,
along stream, 2650 m, 20 July 1995, /,. J. Dorr & I,. G.
Barnett 8180 (F, NY not seen, PORT not seen, US): Mun.
Bocond, Parque Nacional Guaramacdl, road from Bocond
to Guaramacdl, SE of Bocond, S slope of mountain,
09 |.3'\. 70°I2'YY. montane forest. 2500 m. 22 Job 1995.
/.. ,/. Dorr A L. C. Barnett 8240 (E. NY not seen. PORT
not seen, US); Dpto. Bocond, Parque Nacional Guarama-
edl, trail from La Laguna de las Aguas Negras to La Qda.
Salvaje, N slope of mountain, 09°19'N, 70°l l'W, 27 Oct.
1998, /,. ./. Dorr. E. Briceno, G. Briceno & B. Cracas 8291
(PORT); Mini. Bocond, Parque Nacional Guaramacdl, road
from Bocond to Guaramacdl, SE of Bocond, ea. 15 km from
the post of the park guards, S slope of mountain, 09°13'N,
70°I2'W, montane and lower montane forest, 3 Nov. 1998,
L. J. Dorr. E. Brieeno, G. Briceno & R. Cracas 8486 (C
not seen, COL not seen, CTES not seen, E. MO not seen,
NY not seen, OSH. PORT not seen. US); Mun. Bocond,
Parque Nacional Guaramacdl, Bocond— easerfo de Guara¬
maeal road, S slope, ((Jda. Jirajara) from turnoff to anten¬
nas to just above El Campamento, 15 June 2001, E. ,/.
Dorr. S. M. Niflo & R. Caracas 9014 (OSH); 12 km ESE
ol Bocond, I km N to 4 km NNE of Guaramacdl, 09° 12'—

1 3 ' N . 70°09'W, 1600—1900 m, 15 Mar. 1982. R. Liesner,
A. Gonzalez. B. Stergios A G. Aymard 12954 (MO. PORT,
VEN not seen); Bocond— Guaramacdl road, kms 16—22
SSE of Bocond, subpdramo dominated by Espeletia par-
toneoides and Neurolepis, 2775—3175 m, 20 Jan. 1978, ./.
L. Luteyn, M. D'hrdn-Luteyn, I,. Ruiz-Teran & ./. A. Du-
garte 5206 (NY not seen. MERE not seen, OSH, Y EN not
seen); Dpto. Bocond, Paramo Guaramacdl, 12.2 km be¬
yond jet. outside Bocond, 09°I3'N, 70°12'W, steep road¬
side slopes dominated by Clusia and bamboo, ca. 2600
m. 19 Jan. 1984, ./. L. Luteyn & J. J. Pipoly 9296 (NY
not seen. OSH. YEN not seen); Dpto. Bocond, Paramo
Guaramacdl, 20—21 km beyond jet. with hwy. NE of Bo¬
cond. 09°13'N, 70' 1 3' Vi, shaded very' wet streambed with
large boulders, 2640-2700 m. 14 Mar. 1984. ./. I,. Luteyn
& E. Cotton 9710 (NY not seen, OSH. PORT not seen.
Y EN not seen). J. L. Luteyn & E. Colton 9711 (NY not
seen. OSH. PORT not seen, US, Y EN not seen); Dpto.

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

217

Figure 8. Centropogon candidatus Lammers. — A. Upper
portion of stem with flower. — B. Arbusculiform triehomes
on abaxial leaf surface. (Drawn from the holotype, Bristol
1182.)

Carache, via Paramo Lend# sitio denominado “Los Bar-
reales,” 9°32'N, 70°08'W, bosque subparameros. 2800-
2900 m, 29 Feb. 1988. R. Rivero & I\. Rondon 1576 (MO.
PORT not seen); Mun. Bocono, camino al Cerro Guara-
macdl via la laguna de “Los Cedros,” bosque nublado, 21
Mar. 1981, B. Stergios 2591 (PORT); Cerro Guaramacal,
Bocono, bajando hacia el caserfo de Guaramacal, selva
humeda montano alto, 25—26 Nov. 1982, B. Stergios, G.
Aymard & A. Smith 4726 (PORT); Mun. Bocono. Parque
Nacional Guaramacdl, camino Laguna de los Cedros, ver-
tiente norte, 09°07'N, 70°16'W, bosque humedo montano
alto, cresta del paramo. 2000— 24(H) m, Jan. 1991. B. Ster¬
gios 18643 (PORT not seen, US); Mun. Bocono, camino
al Cerro Guaramacdl via la laguna de “Los Cedros,”
bosque nublado, 21 Mar. 1981, B. Stergios 2591 (PORT).

2. Centropogon candidatus Lammers, sp. nov.
TYPE: Colombia. Narino: E Aponte, Rfo Ma-
jinsanoy, forest trail, 2700 m, 29 June 1963,
M. L. Bristol 1182 (holotype, US). Figure 8.

Species insignis egregie distincta ab omnibus caeteris
speciebus Centropogonis subsect. Brevilimbatorum tri-
chomatibus arbusculiformibus candidis, laminis pinnati-
lobatis, et calycis lobis ellipticis erectis 13-15 mm longis
5—6 mm latis marginibus 1—3-spinulosis.

Shrult, 3 m tall; stems 2-2.6 mm diam., moder¬
ately leafy, densely pubescent with white arbuscu¬
liform hairs. Lamina ovate, 6.4-8 cm long, 2.5—3

cm wide between the lobes, 3—3.9 cm wide across
the lobes, coriaceous; adaxial surface dull dark
green, very sparsely pubescent with white arbus¬
culiform hairs; abaxial surface dull green-white,
very densely pubescent with white arbusculiform
hairs, the surface almost wholly obscured; margin
pinnately lobed, the segments 3 to 5 per side, tri¬
angular to deltate. 3-8 X 3—9 mm, acute and mu-
cronate at apex; apex acuminate; base rounded;
petiole 14—20 mm long, 1-E5 mm diam., 1/4— 1/5
as long as the lamina, very densely pubescent with
white arbusculiform hairs. Flowers solitary in the
upper leaf axils; pedicels ascending, 27—35 mm
long, 0.8— 0.9 mm diam., ebracteolate (or the brac-
teoles obscured by the indumentum?), very densely
pubescent with white arbusculiform hairs. Hypan-
thium oblate, 6 mm long, 9—10 mm diam., 1/8 as
long as the corolla, very densely pubescent with
white arbusculiform hairs. Calyx lobes elliptic,
erect, 13—15 X 5-6 mm, more than twice as long
as the hypanthium, about half as long as the corolla
tube, densely pubescent with white arbusculiform
hairs toward base, becoming sparser toward apex;
apex acuminate; margin denticulate with 1 to 3 spi-
nulose teeth on each side. Corolla tubular, bilabi¬
ate, coral pink externally, yellow within, 46-48 mm
long, pubescent with white arbusculiform hairs;
tube gently curved, 31—32 mm long, 4 — 4.5 mm
diam. at the base, not constricted, flaring gradually
to 6.5—8 mm diam. at mouth; dorsal lobes narrowly
triangular, falcate, 15—16 X 3.7-4 mm, half as long
as the tube, the apex acuminate; ventral lobes nar¬
rowly triangular, deflexed. 8-10 X 2.5— 3.5 mm, the
apex acuminate. Staminal column exserted, emerg¬
ing above the dorsal lobes; filament tube yellow,
slightly curved, 45-47 mm long, 1 mm diam.,
densely pubescent with long spreading simple
white hairs; anther tube gray-blue, 2.3— 2.4 mm
diam., slightly curved, sparsely pubescent on the
connectives with long simple white hairs; dorsal an¬
thers 7 mm long, 1/6 as long as the filament tube;
ventral anthers 5 mm long, pubescent at apex with
tufts of yellow-white hairs 1.5-2 mm long. Fruit
and seeds not seen, but apex of ovary flat.

Distribution, habitat, and phenology. Apparently
endemic to the Cordillera Central of southwestern Co¬
lombia, and known only from the type specimen.

Etymology. The specific epithet alludes to the
plant’s dense white indumentum; from the Latin ad¬
jective candidatus, clothed in white.

Relationships. Based on its arbusculiform tri-
chomes and “short-lobed” corolla, Centropogon
candidatus is a member of section Siphocampylo-
ides, subsection Brevilimbati. Within that subsec¬
tion, however, it is remarkable for its white (pre-

218

No von

sumably unpigmented) indumentum. The
arbuseuliform hairs that eliaraeterize subseetion
Brevilimbali typically are darkly pigmented: yellow-
brown. red-brown, chocolate-brown, purple, gray, or
black. Only two other species of that subseetion are
characterized by white triehornes: C. albostellatus
Jeppesen and C. llanganatensis Jeppesen, both of
central Ecuador. Neither is very similar to C. can-
didatus in other features. Taken together, these two
species differ from tin1 novelty in their euneate or
attenuate leaf bases (vs. rounded in C. candidatus),
entire or minutely toothed (vs. lobed) leaf margins,
longer pedicels (40—80 mm vs. 27—35 mm), shorter
calyx lobes (2-10 mm vs. 13—15 mm), shorter co¬
rollas (20—40 mm vs. 46—48 mm long) constricted
toward base (vs. not constricted), and glabrous (vs.
pubescent) staminal column.

As an aside, it should be noted that while color
of the triehornes generally has been useful as a tax¬
onomic character among subsection Brevilimhati
(Jep| >esen, 1981), variation within a species is pos¬
sible. Centropogon erythraeus Drake of southern
Ecuador typically has a dense indumentum that is
purple or chocolate-brown. Recently, I examined a
specimen (Stein & D Alessandro 2730, MO) from a
known locality of C. erythraeus, which conformed
in every way to that species, except that the pu¬
bescence was pure white. Apparently, just as many
plants produce occasional individuals with unpig¬
mented flowers, so, too, do some members of sub¬
seetion Brevilimhati occasionally produce individ¬
uals with unpigmented triehornes. For this reason,
it would not be wise to distinguish a new species
solely on an unusual indumentum color.

Fortunately, Centropogon candidatus is also re¬
markable for its [(innately lobed leaf margin. In
most of subsection Brevilimhati, the margins are en¬
tire or minutely toothed (crermlate, denticulate, ser¬
rulate). Only one other species has leaves divided
into definite lobes: C. australis (F. E. Wimmer)
Gleason of Venezuela. This species does resemble
C. candidatus in lamina size, petiole length, hy-
panthium shape, and pubescence of the staminal
column. However, C. australis differs from C. can¬
didatus in its brown (vs. while) indumentum; calyx
lobes that are sublinear (vs. elliptic) and only 1—5
mm long (vs. 13—15 mm); and corolla that is purple
(vs. coral externally and yellow internally), only
30-36 mm long (vs. 46-48 mm), and constricted
(vs. not constricted) toward base. Overall, these two
species do not seem closely related, and it may be
hypothesized that leaf-lobing has evolved at least
twice in this subsection.

In addition to the white indumentum and lobed
leaves, C. candidatus is distinctive by virtue of its
calyx lobes. In the majority of subsection Brevilim¬

hati, calyx lobes are only 1—9 mm long (vs. 13—15
mm long in C. candidatus) and broadest at base
(vs. above the base) with entire margins (vs. 1 to 3
spinulose dentieulations per side). However, one
known species possesses calyx lobes similar to
those of C. candidatus: C. karstenii Zahlbruckner
has calyx lobes that are 12-15 mm long and lan¬
ceolate (i.e., broadest above the base), with 3 den-
ticulations on each margin. This species further re¬
sembles the novelty in size and general shape of
the lamina; length of the pedicels; size and shape
of the hypanthium; in the size, exsertion, and pu¬
bescence of the staminal column; and is likewise
endemic to Narino, growing in the vicinity ofPasto.
It differs from the novelty only in its red-brown
indumentum (vs. white in C. candidatus), shorter
petioles (8—10 mm vs. 14—20 mm), denticulate (vs.
[(innately lobed) lamina, narrower calyx lobes (2-4
mm vs. 5-6 mm) that are lanceolate (vs. elliptic),
and red (vs. coral and yellow) corollas with shorter
lob. ;s (dorsal lobes 10 mm vs. 15—16 mm, ventral
ones 5 vs. 8-10 mm). Overall, on the basis of mor¬
phology and geography, C. karstenii would appear
to be the closest known relative of C. candidatus.

3. Centropogon eilersii hammers & M. (). Dil¬
lon, sp. nov. TYPE: Peru. Cuzco: La Conven-
eion. road from Cuzco to Ouillabamba, after
pass Abra Malaga and before Ipal. I3°04'S,
72°22'W, remnants of former cloud forest
along the road, 2600— 2900 m, 24 Feb. 2000,
\1. Weigend & K. Weigend 2000/441 (holotype,
NY; isotype, Herbario Universidad National
de San Augustin not seen). Figure 9.

Species Centropogonis sect. Burmeisteroidis affinis C.
david-smithio et C. isabellino, sc.l ah his speciebus ped-
ieellis longiorihus 250 mm longis, ealycis lobis longiori-
hns 40 mm longis, coroll is luteolis majoribus cum corolla
85 mm longa tubo 50 mm longo, et tubo filamentorum
longiore 65 mm longo bene distinguenda.

Apparently an erect suffrutieose herb or shrub;
stems 8 mm diam., pubescent with brown arachnoid
hairs. Lamina elliptic, 215-29 X 9-10.8 cm. char-
taceous; adaxial surface dull dark green, pubescent
with short stiff white hairs; abaxial surface didl
light green, more densely pubescent with long stiff
spreading brown hairs; margin minutely denticulate
with yellow callosities; apex acute; base euneate;
petiole adaxially canaliculate, 25—38 mm long, 3—
3.5 mm diam., 1/7— 1/1 I as long as the lamina, pu¬
bescent with brown arachnoid hairs. Flowers soli¬
tary in the upper leaf axils; pedicels ascending,
curved, 250 mm long, 2.7 mm diam., bibracteolate
at base, moderately pubescent with brown arach¬
noid hairs; braeteoles linear, 7 X 0.5 mm. Hypan¬
thium narrowly eampanulate, 18 mm long. 12 mm

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

219

Figure 9. Centropogon eilersii Lammers & M. 0. Dillon.
(Drawn from the holotype, Weigend & Weigend 2000/441 .)

diain.. 1/5 as long as the corolla, pubescent with
brown arachnoid hairs. Calyx lobes linear-triangu¬
lar, erect. 40 X 3—5 mm, more than twice as long
as the hypanthium, 4/5 as long as the corolla tube,
pubescent with brown arachnoid hairs; apex acu¬
minate; margin serrulate with 8 to 12 tiny curved
teeth on each side. Corolla tubular, pale yellow, 85
mm long, pubescent with short still simple colorless
hairs, these often clustered so as to appear stellate;
tube slightly curved, 50 mm long, 12 mm diam. at
the base, narrowing imperceptibly to 10 mm diam.
at middle, then flaring gradually to 22 mm diam.
at mouth; dorsal lobes narrowly triangular, falcate,
35 X 9 mm, 7/10 as long as the tube, the apex
acute; ventral lobes narrowly triangular, deflexed,
23 X 9 mm, the apex acute. Staminal column ap¬
parently little exserted from the tube, the anthers
positioned between the dorsal lobes; filament tube
slightly curved, 65 mm long, 2 mm diam., pubes¬
cent with long soft spreading white hairs; anther
tube 7 mm diam., curved, densely pubescent with
long shaggy brown hairs; dorsal anthers 15 mm
long, 1/4— 1/5 as long as the filament tube; ventral
anthers 13 mm long, pubescent at apex with tufts
of soft brown hairs 3.5—4 mm long. Fruit and seeds
not seen, but apex of ovary flat.

Distribution, habitat, and phenology. Apparently

endemic to southern Peru, and known only from the
type collection.

Etymology. This species is respectfully dedi¬
cated to the memory of Lawrence J. Filers (1927—
2000), plant taxonomist, ardent conservationist,
and botanical educator. From 1968 to 1989, Larry
was professor of biology at the University of North¬
ern Iowa and curator of that school’s herbarium
(ISTC). His research program focused on the flora
of Iowa, and culminated in the publication of The
Vascular Plants of Iowa (Eilers & Roosa, 1994). As
much as Larry enjoyed floristic research, it was as
a teacher that he truly excelled. Students who were
having trouble with academics or with life in gen¬
eral seemed especially to flourish and blossom un¬
der Larry’s gentle tutelage. In addition to classroom
teaching, he directed the master’s theses of several
future botanists, including the author and Michael
(). Dillon of the Field Museum of Natural History
(F), who joins me in naming this species in our
professor’s honor.

Relationships. On the basis of its simple tri-
chomes and “long-lobed” corolla, Centropogon ei¬
lersii is a member of section Burmeisteroides. With¬
in that section, it belongs to a group of four species
characterized by denticulate leaves 2 to 6 times
longer than wide; green or yellow corollas; and stiff
persistent simple (often clustered) trichomes at
least on the corolla (McVaugh, 1949; Lammers,
1998a): C. altus F. E. Wimmer of northern Peru, C.
ayavacensis (Willdenow ex Schultes) Lammers of
the Cordillera Central of Colombia (C. willdenow-
ianus F. E. Wimmer, notn. superfl.; cf. Lammers,
1999), C. david-smithii Lammers of central Peru,
and C. isabellinus F. E. Wimmer, also found in Cuz¬
co. Among these species, the novelty seems most
similar to the last two, particularly in size, shape,
and pubescence of the hypanthium and anther
tube. Centropogon david-smithii and C. isabellinus
only differ from C. eilersii in their shorter pedicels
(90—160 mm vs. 250 mm in C. eilersii ); shorter ca¬
lyx lobes (7—35 mm vs. 40 mm); green (vs. pale
yellow) corollas that are shorter (56—64 mm vs. 85
mm) due to their shorter tube (20—33 mm vs. 50
mm); and shorter filament tube (46—50 mm vs. 65
mm) that is strongly (vs. scarcely) exserted.

4. Centropogon joergensenii Lammers, sp. nov.
TYPE: Ecuador. Zamora-Chinchipe: Ghinchi-
pe, Parque Nacional Podocarpus, La Esmer¬
alda (Cooperativa San Francisco de Numbala
Alto), 04°22'S, 79°03'W, bosque primario alto,
2250 m, Jan. 1994, W. Palacios & M. Tirado
13042 (holotype, OSH; isotypes, MO. QCNE
not seen). Figure 10.

220

Novon

Figure 10. Centropogon joergensenii Lammers. — A. Up¬
per portion of stem with flower. — B. Pubescence on stem
and petiole. (Drawn from the holotype, Palacios & Tirado

13042.)

Species Centropogonis subseet. Falcatorum ob trichom¬
ata septata longa affinis C. trichodi, sed ah hac specie
trichomatibus longioribus 2.5— 3.5 mm longis, pedicellis
longioribus 6.5— 7.2 cm longis, calycis lobis paree pubes-
centibus brevioribus 6-7 mm longis, corolla majore 45—
48 mm longa, columna staminali vix exserta, et tubo an-
therarum glabro statirn distinguenda.

Scandent shrub; sterns moderately leafy. 2—2.5
mm diam., branched, densely pubescent with stiff
yellow spreading multicellular hairs 2.5— 3.5 mm
long. Lamina elliptic, 5.8—8 X 2.6— 3.6 cm, char-
taceous; adaxial surface dull dark green, moderate¬
ly pubescent with long stiff yellow multicellular
hairs; ahaxial surface dull white-green, densely pu¬
bescent with long stiff yellow multicellular hairs;
margin denticulate with dark callosities; apex
acute; base obtuse; petiole 1.3— 1.8 mm long, 0.8—
1.2 mm diam., 1/4— 1/6 as long as the lamina,
densely pubescent with long stiff yellow multicel¬
lular hairs. Flowers solitary in the upper leaf axils;
pedicels ascending, 65—72 mm long, 1—1.3 mm
diam., bibracteolate at or just above the base, mod¬
erately pubescent with long stiff spreading yellow
multicellular hairs. Hypanthium depressed hemi¬
spheric, 3—4 mm long, 7—8 mm diam., 1/10—1/15
as long as the corolla, densely pubescent with long
stiff spreading yellow' multicellular hairs. Calyx
lobes triangular, erect, 6-7 X 2.5—3 mm, half again
as long to almost 2 I/2X as long as the hypanthium.

1/5— 1/6 as long as the corolla tube, moderately pu¬
bescent with long stiff spreading yellow multicel¬
lular hairs; apex acuminate; margin entire. Corolla
tubular, slightly ventrieose, red with yellow limb,
45—48 mm long, moderately pubescent with long
still spreading yellow multicellular hairs; tube
slightly curved, 40 mm long, 6 mm diam. at the
base, narrowing abruptly to 2.5 mm diam. just
above the base, then flaring gradually to 5.5 mm
diam. just below the mouth; dorsal lobes narrowly
triangular, falcate, 8 X 2.3 mm, 1/5 as long as the
tube, the apex acuminate; ventral lobes narrowly
triangular, deflexed, 7 X 1.8 mm, the apex acu¬
minate. Staminal column slightly exserted between
the dorsal lobes; filament tube slightly curved, 43
mm long, 1.5 mm diam., densely pubescent with
long soft white hairs; anther tube 2 mm diam.,
slightly curved, gray, its surfaces glabrous; dorsal
anthers 6 mm long, 1/7 as long as the filament tube;
ventral anthers 4.5 nun long, pubescent at apex
with tufts of stiff dirty-white hairs 0.8-1 mm long.
Fruit and seeds not seen, but apex of ovary flat.

Distribution, habitat, and phenology. Apparently
endemic to southern Ecuador and known only from
the type collection.

Etymology. It is a pleasure to name this spe¬
cies for Peter Mpller Jprgensen (b. 1958), from
1986 to 1989 director of the herbarium at Ponlilicia
Universidad Catoliea del Ecuador (QCA) and since
1992 a member of the curatorial staff at the Mis¬
souri Botanical Garden (MO). Peter is actively en¬
gaged in studies of the flora and biogeography of
the Andes, was co-editor ol the encyclopedic Cat¬
alogue of the Vascular Plants of Ecuador (Jprgensen
& Leon-Yanez, 1999), and, with his wife Carmen
Ulloa Ulloa (see below), is a contributor to the Flo¬
ra of Ecuador project. In forming the epithet, the
spelling of bis surname has been altered to comply
with Article 60.6 of the ICRIN (Greuter et al., 2000).

Relationships. On the basis of its simple tri-
ebomes and “short-lobed" corolla with deflexed fal¬
cate lobes, Centropogon joergensenii is a member of
section Wimmeriopsis, subsection Falcati. Though 1
did not comment upon it in my earlier review of
the genus (Lammers, 1998a), the species of sub¬
section Falcati fall into two discrete geographic
groups. The first includes four species endemic to
Costa Rica | C. irazuensis Wilbur. C. palmanus
(Donnell Smith) F. E. Wimmer, C. valerii Standley,
and C. wimmeri Standley] plus a fifth one \ C. gran-
didentatus (Sehlechtendal) Zahlbruekner] that is
distributed from southern Mexico to northern Co¬
lombia and western Venezuela. The second group
includes 12 Ecuadorean endemics [C. aequitorialis

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2002

Lammers

New South American Lobelioideae

221

F. E. Wimmer, C. brachysiphoniatus Zahlbruckner,
C. calycinus Bentham, C. jeppesenii Lammers, C.
phoeniceus Jeppesen, C. pilalensis Jeppesen, C.
rimbachii F. E. Wimmer, C. rubrodentatus Jeppe¬
sen, C. sodiroanus Zahlbruckner, C. subandinus
Zahlbruckner, C. trichodes F. E. Wimmer, and C.
uncinatus Zahlbruckner] plus one \C. reticulatus
Drake] t hat occurs in both Ecuador and northern
Peru.

The new species differs from all these in various
features, most notably in the dense indumentum of
very long stiff yellow multicellular trichomes; most
of the species are glabrous or pubescent with short¬
er, softer, unicellular hairs. Only C. trichodes F. E.
Wimmer, which is also endemic to Zamora-Chin-
chipe (valley of the Rio Valladolid), bears trichomes
at all similar to those of C. joergensenii ; they are,
however, only 1—1.5 mm long (vs. 2.5— 3.5 mm long
in C. joergensenii). While the two species are sim¬
ilar in size and shape of the leaves and general
floral structure, C. trichodes differs in its shorter
pedicels (4—6 cm vs. 6.5— 7.2 cm in C. joergensenii).
longer calyx lobes (8—15 mm vs. 6-7 mm) that are
subglabrous (vs. moderately pubescent), shorter co¬
rolla (30-35 mm vs. 45—48 mm), strongly exserted
(vs. scarcely exserted) staminal column, and anther
tube pilose with purple-blue trichomes in the su¬
tures (vs. glabrous). On the basis of morphology and
geography, C. trichodes appears to be the closest
known relative of C. joergensenii.

,5. Centropogon ulloae Lammers. sp. nov.
TYPE: Ecuador. Napo: road from Tulcan to
Santa Barbara and La Bonita, 21 km below
Santa Barbara. 00°35-45'S, 77°30'W, cloud
forest, 2400 m, 28 May 1985, B. A. Stein 2B75
(holotype, MO). Figure I 1.

Ah omnibus caeteris speciebus Centropogonis seel.
Wimmeriopsidis indumento sparse trichomatum arbuscu-
liformium minimorum differt; species ob calycis lobos basi
connatos hypanthio oblato longiores aftinis Centropogoni
calycino et speciebus similaribus, sed ab his speciebus
lamina laneeolata vel anguste elliptica longiore 12.5—17.5
cm longa, calycis lobis connatis per dimidium longitudinis
ad apicem obtusis vel rotundatis ad marginem integris, et
corolla aurantiaco-rubra cum lobis luteis bene distinguen-
da.

Shrub forming thickets to 2.5 m tall; stems pur¬
ple, somewhat densely leafy toward apex, 3 mm
diam., subglabrous; latex white. Lamina lanceolate
or narrowly elliptic, 12.5—17.5 X 4—4.6 cm, sub-
coriaceous; adaxial surface dull dark green, very
sparsely scabrid with minute arbusculiform tri¬
chomes; abaxial surface dull white-green, sparsely
scabrid with minute arbusculiform trichomes; mar-

Figure 11. Centropogon ulloae Lammers. — A. Upper
portion of stem with flower. — B, C. Arbusculiform tri¬
chomes on abaxial leaf surface. (Drawn from the holotype,
Stein 2H 75. )

gin minutely denticulate with dark intramarginal
callosities; apex acuminate; base obtuse or cuneate;
petiole 20—40 mm long, 1.1-1. 6 mm diam., 1/4—
1/6 as long as the lamina, subglabrous or sparsely
scabrid with minute arbusculiform trichomes. Flow¬
ers solitary in the upper leal axils; pedicels as¬
cending, 55 mm long, 1 mm diam., bibracteolate
just above the base, subglabrous or sparsely scabrid
with minute arbusculiform trichomes; bracteoles
linear, 5 X 0.3 mm. Hypanthium oblate, 8 mm
long, 13 mm diam., 1/6 as long as the corolla, gla¬
brous. Calyx lobes narrowly oblong, erect, 18-19
X 3.5— 4.5 mm, connate for half their length, more
than twice as long as the hypanthium, about half
as long as the corolla tube, moderately scabrid with
minute arbusculiform trichomes; apex rounded or
obtuse; margin entire. Corolla tubular, bilabiate, or¬
ange-red with yellow' limb, 50 mm long, moderately
scabrid with minute arbusculiform trichomes; tube
bent above the base, 38 mm long, 10 mm diam. at
mouth; dorsal lobes narrowly triangular, falcate, 12
X 4 mm, 1/3 as long as the tube, the apex acu¬
minate; lateral lobes narrowly triangular, falcate, 9
X 2.5 mm. the apex acuminate; ventral lobe nar¬
rowly triangular, deflexed. 8 X 2.2 mm. the apex
acuminate. Staminal column somewhat exserted be-

222

Novon

tween the dorsal lobes; filament tube suberect, 48
mm long, 1.9 mm diam., glabrous; anther tube 3
mm diam., suberect, gray, its surfaces glabrous;
dorsal anthers 7 mm long, 1/7 as long as the fila¬
ment tube; ventral anthers 5.3 mm long, pubescent
at apex with tufts of still white hairs 1.3— 1.5 mm
long. Fruit and seeds not seen, but apex of ovary
flat.

Distribution, habitat, and phenology. Apparently
endemic to northern Ecuador and known only from
the type specimen.

Etymology. It is a pleasure to name this spe¬
cies for Carmen Ulloa Ulloa (b. 1963), who served
at the Pontificia Universidad Catolica del Ecuador
((,)CA) as a research assistant (1986—1988) and pro-
fesor agregado (1990—1991) before joining the cu¬
ratorial stall at Missouri Botanical Garden (MO) in
1992. Carmen is actively engaged in studies of the
flora and biogeography of Andean forests and the
paramos of Ecuador, eo-edited Flora de Nicaragua
(Stevens et al., 2001) and. with her husband Peter
Mpllcr Jprgcnsen (see above), is a contributor to the
Flora of Ecuador project. It is an honor to com¬
memorate this couples many contributions to Latin
American botany with a bouquet of Centropogon.

Relationships. Centropogon ulloae presents yet
another challenge to MeVaugh's (1949) classifica¬
tion of the genus. The combination of arbusculiform
trichomes and “short-lobed” corolla would place
this species in section Siphocampyloides, subsec¬
tion Rrevilimbati. However, the novelty shows little
resemblance to any of those species. Its distinctive
connate calyx lobes are almost unknown in section
Siphocampyloides; within that group, only the re¬
cently described C. connatilobatus hammers of
Venezuela has basally connate calyx lobes. How¬
ever, that species differs (com C. ulloae in its much
smaller leaves (2.6— 6 X 0.6— 1.9 cm vs. 12.5—17.5
X 4—4.6 cm in C. ulloae) with shorter petioles (4—
17 mm vs. 20—40 mm); smaller hypanthium (3 mm
vs. 8 mm long, 5 mm vs. 13 mm diam.) that is
pubescent (vs. glabrous); shorter calyx lobes (12
mm vs. If?— 19 mm long) that are acuminate at apex
(vs. rounded or obtuse); and shorter corolla (42 mm
vs. 50 mm long) with red limb (vs. yellow). Fur¬
thermore, the branched hairs of the novelty differ
from those of section Siphocampyloides in their mi¬
nuteness. The former are only 1/10 as long as those
ol the Siphocampyloides; 0.1— 0.3 mm vs. 1-3 mm.
Overall, it does not appear that C. ulloae is related
to any member of section Siphocampyloides.

Though C. ulloae seems out of place in section
Siphocampyloides, it does show great similarity to
a group ol four species within the Ecuadorean

branch ol section Wimmeriopsis, subsection Falcati
(see above): C. aequitorialis F. E. Wimmer (Carehi
and Pichincha), C. brachysiphoniatus Zahlbruckner
(Pichincha), C. calycinus Bentham (Imbabura and
Pichincha), and C. phoeniceus Jeppesen (Napo).
Like C. ulloae, these species possess calyx lobes
that are longer than the hypanthium and connate.
They further resemble C. ulloae in their pedicels
equaling or longer than the (lower, relatively long
subbasal bracteoles, hypanthia broader than long,
“short-lobed” corolla morphology, and glabrous lil-
aments. In an earlier classification of the genus,
Wimmer (1931) applied the rankless infraseclional
name “Caligati F. E. Wimmer” to these plants; that
name (which was included in the synonymy of sub¬
section Falcati by McVaugh [ 1 949]) will be used
here to informally designate this species group.

I he sole impediment to the assignment of C. ul¬
loae to section Wimmeriopsis is its indumentum of
arbusculiform hairs. Throughout section Wimmer¬
iopsis, any hairs that are present are unbranched
(though sometimes multicellular). However, iu light
of the great overall similarity of C. ulloae to the
“Caligati,” and the pronounced size difference be¬
tween its arbusculiform trichomes and those of sec¬
tion Siphocampyloides, it seems best to assign the
novelty to Centropogon sect. Wimmeriopsis, subsect.
Falcati. Its minute arbusculiform hairs are thus hy¬
pothesized to represent yet another independent or¬
igin ol branched trichomes among the Lobelioideae.

In addition to its sparse indumentum ol minute
arbusculiform trichomes, the new species differs
from the other “Caligati” in details of its connate
calyx lobes. In C. ulloae, the lobes are connate for
hall their length (vs. 2/3 or more of their length in
C. aequitorialis and C. calycinus or 1/3 or less of
their length in C. brachysiphoniatus and C. phoen¬
iceus), rounded or obtuse (vs. acute) at apex, with
the free margins entire (vs. toothed). The new spe¬
cies further differs from the other four “Caligati” in
its longer (12.5—17.5 cm vs. 3—13 cm long) lance¬
olate or narrowly elliptic (vs. ovate, elliptic, or
widely elliptic) leaves and orange-red corolla with
yellow lobes (vs. red or purple only occasionally
with yellow or white lobes).

6. Centropogon vitifolius Lammers, sp. nov.
TYPE: l\ ■ru. Amazonas: Condoreanqui Prov.,
Distr. El Cenepa, Region Nororiental del Mar-
anon. comunidad de Kusu-kulmim, Rio Co-
maina, OITA'S, 78” 1 6’ W, bosque primario,
700 m. 17 Aug. 1994. R. Vasquez, N. Jaram-
illo, R. Apanu & M. Ugkuch I HR 1 6 (holotype.
OSH; isotypes, AMAZ not seen. MO). Figure
12.

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

223

Figure 12. Centropogon vitijolius Lammers. — A. Leaf.
— B. Flower. — C. Immature berry. (Drawn from the ho-
lotype, Vasquez el al. 18816.)

Species Centropogonis subsect. Colornbianorum affinis
C. fimbriatulo et C. tessmanniis , sed al) his speciebus ab¬
sentia (praeter antheras) trichomatum longorum, lamina
cordata interdum subtrilobata, et corollae lobis longioribus
(dorsalibus 18 mm longis, ventralibus 13 mm longis) facile
distinguenda.

Suffruticose lierl); stems moderately leafy, 1.5—3
mm diam., glabrous; latex white. Lamina ovate or
widely ovate, appearing vaguely 3-lobed, 5-7.5 X
3.5— 6.5 cm, chartaceous, glabrous; adaxial surface
dull light green, glabrous; abaxial surface dull
white-green; margin coarsely dentate with 4 to 7
teeth per cm, some distinctly larger than the others;
apex acuminate; base cordate; petiole 15—32 mm
long, 0.5- 1.2 mm diam., 1/4— 1/2 as long as the
lamina, glabrous. Flowers solitary in the upper leaf
axils; pedicels ascending, recurved after anthesis,
38—50 mm long, 0.8— 1.2 mm diam., ebracteolate,
minutely puberulent. Hypanthium obovoid or ob-
conic, 7-8 mm long, 6-7 mm diam., 1/8— 1/9 as
long as the corolla, minutely puberulent. Calyx
lobes triangular, recurved at apex, 2-2.2 X 0.7-1
mm, 1/4— 1/3 as long as the hypanthium, 1/25 as
long as the corolla tube, minutely puberulent; apex
acute; margin entire. Corolla tubular, rose-colored,
65-68 mm long, minutely puberulent; tube curved,
50 mm long, 5—6 mm diam. at the base, narrowing

abruptly to 2.2— 2.6 mm diam. above the hase, then
flaring gradually to 9-1 1 mm diam. just below the
mouth; dorsal lobes linear-triangular, spreading, 18
X 2.5 mm, 1/3 as long as the tube, the apex acute;
ventral lobes linear-triangular, deflexed. 13 X 2.5
mm. the apex acute. Staminal column slightly ex-
serted between the dorsal lobes; filament tube
curved, 58 mm long, 0.6 mm diam., glabrous; an¬
ther tube 2.2 mm diam., slightly curved, pubescent
toward apex with long white hairs; dorsal anthers 7
mm long, 1/8 as long as the filament tube; ventral
anthers 6.2 mm long, pubescent at apex with tufts
of stiff dirty-white hairs 1 mm long. Fruit and seeds
not seen, but apex of ovary flat.

Distribution, habitat, and phenology. Apparently
endemic to northern Peni and known only from the
type collection.

Etymology. The specific epithet is given in al¬
lusion to the resemblance of the leaves to those of
certain species of grapes; from the genus Vitis L.
(Vitaceae) and the Latin noun folium, leaf.

Relationships. On the basis of its simple tri -
chomes and “short-lobed” corolla with spreading
triangular lobes, Centropogon vitijolius is assigned
to section Wimmeriopsis, subsection Colombiani.
This group comprises 18 species distributed from
southern Mexico to central Peru, plus an additional
species endemic to Dominica and Guadeloupe in
the Lesser Antilles (McVaugh, 1949; Lammers,
1998a). The new species is most similar to the only
two members of the subsection found south of Co¬
lombia: C. tessmannii F. F. Wimmer (Amazonian
lowlands of Ecuador and Peru) and C. fimbriatulus
McVaugh (montane forests of southeastern Ecua¬
dor). These three species share a number of fea¬
tures, including habit, chartaceous ovate leaves
with conspicuously toothed margins, ebracteolate
pedicels, recurved calyx lobes, scarcely exserted
staminal column, and pubescent anther tube. Cen¬
tropogon tessmannii and C. fimbriatulus differ from
the novelty in the increased pubescence of their
vegetative organs (simple multicellular hairs up to
1.5 mm long vs. glabrous or minutely puberulent
in C. vitijolius); cuneate, obtuse, or truncate (vs.
cordate) leaf base; and shorter corolla lobes, both
dorsal (6—1 1 mm vs. 18 mm) and ventral (6—9 mm
vs. 13 mm). Neither species shows the tendency
toward a trilobate leaf as seen in C. vitifolius.

Between C. tessmannii and C. fimbriatulus, the
latter seems more similar to C. vitifolius, particu¬
larly in the length of its pedicels (4-7 cm), calyx
lobes (2 mm), corolla (60—65 mm), and filament
tube (45—55 mm). However, in addition to the fea¬
tures mentioned above, C. fimbriatulus further dif-

224

Novon

fers in its fimbriate leaf margin (vs. coarsely dentate
in C. vitifolius) with 10 to 16 (vs. 4 to 7) teeth per
cm, shorter (4—5 mm vs. 7-8 mm) hemispheric (vs.
obeonic or obovoid) hypanthium, pale lavender (vs.
rose) corolla, and shorter (5—6 mm vs. 7 mm) dorsal
anthers. Centropogon tessmannii does resemble C.
vitifolius in its coarsely dentate leal margin, longer
obeonic hypanthium, and longer dorsal anthers, hut
differs in its shorter (2—2.5 cm vs. 3.8—5 cm) ped¬
icels, longer (3—10 mm vs. 2—2.2 mm) calyx lobes,
shorter (46—60 mm vs. 65—68 mm) red (vs. rose-
colored) corolla, and shorter (40-45 mm vs. 58 mm)
filament tube. Overall, it seems best to regard the
three species of Centropogon subsect. Colombiani
occurring south of Colombia as a close-knit ii not
monophyletic group.

111. Siphocampylus Pohl, PI. Bras. Icon. Deser.
2: 104. 1831. TYPE (lectotype, designated by
MeVaugh (1943: 100)): Lobelia westiniana
Thunberg ( = Siphocampylus westinianus
(Thunberg) Pohl |.

With the addition of the five novelties described
here, Siphocampylus comprises 221 species, dis¬
tributed from Costa Rica to Argentina, with several
endemic to Cuba and Hispaniola (hammers,
1998a). I'lie genus resembles Centropogon overall,
but differs in fruit type: baccate, from a flat-topped
ovary, in Centropogon-, capsular, from an ovary con¬
ical at apex, in Siphocampylus.

The most recent classification of Siphocampylus
(Wimmer, 1953, 1968) is unsatisfactory for a vari¬
ety of reasons (Pepper et al., 1997; Lammers,
1998a; also see below), but in the absence of an
improved version, it must suffice here for purposes
of discussion. In this classification, Siphocampylus
was divided into into two sections: sect. Siphocam¬
pylus (“Maerosiphon,” nom. invalid.) and sect. Bra-
chysiphon F. E. Wimmer. The latter section was di¬
vided into four subsections: subsect. Secundiflori F.
E. Wimmer, subsect. Altofissi F. E. Wimmer, sub¬
sect. Megastomi F. E. Wimmer, and subsect. Me-
galandri F E. Wimmer. The autonymie section was
divided into four subsections: subsect. Hemisipho-
campylus (A. DC.) F E. Wimmer, subsect. Byrsan-
thes ((7 Presl) F. E. Wimmer, subsect. Isochilus F.
E. Wimmer, and subsect. Siphocampylus (“Eusi-
phocampylus,” nom. invalid.). This last subsection
was then subdivided into five greges: grex Ectropici
(F. E. Wimmer) F. E. Wimmer, grex Siphocampylus
(“Verticillati,” nom. invalid.), grex Umbellati F. E.
Wimmer, grex Botryoides F. E. Wimmer, and grex
Dissitiflori F. E. Wimmer. Several of the subsections

B 5mm

Figure 13. Siphocampylus adhaerens Lammers. — A.
Leaf. — 8. Flower. — C. Leaf margin. (Drawn from the
holotype, Luteyn et al. 5299.)

and greges were further divided into subordinate
taxa.

1, Sipbocanipy Ins adhaerens Lammers, sp. nov.
TYPE: Venezuela. Tachira: Hwy. 9. Ouebrada
de Cocoroco, 1 I km S of Delicias, 1800 m, 28
.Ian. 1978. ,/. L. Luteyn , M. Lebron- Luteyn, L.
Ruiz-Teran & ./. A. Dugarte 5299 (holotype,
OSH; isotype, MERF not seen, NY. VEN not
seen). Figure 13.

Species oh oorollam infundibularem et foliorum latni-
nam cordatam vel subcordatam affinis Siphocampylo me-
galantho et speciebus similaribus, sed ab bis speciebus
pedicelfis bibraeteolatis brevioribus 1.9-2. 3 cm longis,
corollae tubo ad orem angustiore 4.6— 6.7 mm diametro,
antheris dorsalibus brevioribus 6—6.5 mm longis, et cor¬
ollae lobis dorsalibus linearibus strictis longioribus 15—
19 mm longis ventralibus dimidia parte longioribus sed
dimidio latitudine adhaerentibus staminali columna statim
distinguenda.

Suffruteseent herb; stems scandent, moderately
leafy, 2.3—3 mm diam., moderately pubescent with
long weak spreading white hairs; latex white. Lam¬
ina ovate, 4.5—9 X 3.5— 5.5 cm, ehartaceous; ad-
axial surface dull olive green, sparsely pubescent
with short weak appressed white hairs; abaxial sur¬
face dull, paler, moderately pubescent (densest on
the veins) with long weak spreading white or yellow
hairs; margin conspicuously fimbriate with 7 to 10
teetb per cm, teeth 0.5— 0.7 mm long alternating

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

225

with teeth 1.3— 1.5 mm long; apex acuminate; base
cordate (often asymmetric), subcordate, or truncate;
petiole 14—27 mm long, 0.7— 0.9 mm diam., 1/3—
1/5 as long as the lamina, densely pubescent with
long weak spreading white or yellow hairs. Flowers
solitary in distal leaf axils; pedicels straight (curved
if scandent stem declinate), ascending, 19—28 mm
long, 0.6— 0.9 mm diam., bibracteolate at base,
densely pubescent with long weak spreading yellow
hairs; bracteoles subulate, 1.5-2 X 0.2— 0.3 mm.
Hypanthium turbinate, 5 mm long. 5—7 mm diam..
ca. l/l 0 as long as the corolla, densely pubescent
with long weak spreading yellow hairs. Calyx lobes
narrowly triangular, erect, 6.5-7. 7 X 2.5—3 mm,
1/3— 1/2 again as long as the hypanthium, ca. 1/5
as long as the corolla tube, densely pubescent with
long weak spreading yellow hairs; apex acuminate;
margin entire. Corolla bilabiate, infundibular, dark
pink outside, paler within, 50—55 mm long, sparse¬
ly pubescent along the five major veins with long
weak spreading white hairs; tube slightly curved,
33—37 mm long, 3.8-4 mm diam. at base, abruptly
narrowing to an isthmus 1.9— 2.4 mm diam. just
above base, then gradually expanding to 4.6— 6.7
mm diam. at mouth; dorsal lobes linear, strictly
erect, closely adherent to the staminal column, 15—
19 X 0.9-1 .2 mm, 2/5— 1/2 as long as the tube, the
apex acuminate; ventral lobes narrowly triangular,
recurved, 9—12 X 1.8— 2.5 mm, the apex acuminate.
Staminal column exserted between the adherent
dorsal lobes; filament tube slightly curved, 45—52
mm long, 0.9—1 mm diam., glabrous; anther lube
2—2.3 mm diam., slightly curved, sparsely pubes¬
cent along the connectives with long weak ap-
pressed white hairs; dorsal anthers 6-6.5 mm long,
1/7— 1/8 as long as the filament tube; ventral an¬
thers 4—4.5 mm long, their apex pubescent with
tufts of stiff white hairs 1.5—2 mm long. Capsule
rhomboid, 12 mm long, 9 mm diam., half-inferior;
seeds broadly ellipsoid, light brown, 0.6 mm long,
0.4 mm diam., the testa minutely reticulate.

Distribution, habitat, and phenology: Apparently
endemic to western Venezuela and known only from
the type collection.

Etymology. The specific epithet is the Latin
participle adhaerens, adherent, given in reference
to the unusual adherence of the dorsal corolla lobes
to the staminal column.

Relationships. In Wimmer’s (1953, 1968) treat¬
ment of Siphocampylus, S. adhaerens keys to a
group of six Colombian species in section Sipho¬
campylus, subsection Siphocampylus, grex Dissiti-
flori F. E. Wimmer, subgrex Pyriformes F. E. Wim-

mer: S. coronatus Gleason, S. hypsophilus F. E.
Wimmer, S. lecomtei E. E. Wimmer, S. megalanthus
Zahlbruekner, S. sissii F. E. Wimmer, and S. venosus
Gleason. These plants share with the novelty a
pink, red, or red-purple distinctly infundibular co¬
rolla (vs. subcylindric in other species of subgrex
Pyriformes ) 50—62 mm long, and ovate laminas cor¬
date or subcordate at base, and are likely its closest
relatives.

Siphocampylus adhaerens may be distinguished
from these six species by several characters. The
pedicels of all six lack bracteoles and are longer
(3—15 cm vs. 1.9— 2.8 cm in S. adhaerens ); their
corolla tubes are broader in diameter at the mouth
(9—12 mm vs. 4.6— 6.7 mm); and their dorsal an¬
thers are longer (7—10 mm vs. 6—6.5 mm). However,
the most conspicuous characters distinguishing S.
adhaerens from its allies involve the dorsal lobes of
the corolla.

First, the dorsal corolla lobes of .S', megalanthus
and its allies are shorter than those of S. adhaerens :
9—15 mm vs. 15—19 mm. Second, they are similar
to the ventral lobes in size, shape, and posture.
Siphocampylus adhaerens, in contrast, has distinct¬
ly dimorphic corolla lobes: the dorsal pair are
broadly linear, the ventral narrowly triangular. Fur¬
thermore, the dorsal pair are more than half again
as long as the ventral three (15—19 mm vs. 9—12
mm) but only hall as wide (0.9— 1.2 mm vs. 1.8— 2.5
mm). In almost all other species ol the genus, the
dorsal and ventral corolla lobes are of approxi¬
mately equal width. Three species have dorsal
lobes wider than the ventral: S. planchonis F. E.
Wimmer, S. polyanthus F. E. Wimmer, ,S. queta-
mensis F. E. Wimmer. Only in S. isochilus F. E.
Wimmer and S. sceptrum Decaisne [which comprise
Siphocampylus sect. Siphocampylus, subsect. Iso¬
chilus F. E. Wimmer (Wimmer, 1968)] are the ven¬
tral corolla lobes wider than the dorsal. However,
these two species differ from S. adhaerens in nu¬
merous features (including the open orifice ol their
Burmeistera- like anther tube; cf. Stein, 1987; Lam¬
mers. 1998a), and are not closely related to the new
species.

Finally, whereas all live corolla lobes have a
spreading or recurved posture in S. megalanthus
and its allies, this is true for only the ventral three
in S. adhaerens. The dorsal pair are strictly erect
and adhere closely to the exserted staminal column,
hiding it from sight. This close adherence of the
dorsal corolla lobes to the staminal column appears
to be unique in the genus, and perhaps in the sub¬
family.

226

No von

2cm

Figure 14. Siphocampylus ambivalens Lammers. (Drawn
from the holotype, Vargas 5061.)

2. Siphocampylus ambivalens Lammers, sp.
nov. TYPE: Bolivia. Florida: Bermejo, 5 km al
E subiendo haeia el sillar Tres Cruees, eamino
del gaseoducto y zona de la antena, I8°()B'S.
63°35.5'W, Bosque alio en laderas y ondona-
das con Cariniana estrellensis, Pachistroma
longifolia y Myroxylom peruiferum, oa. 500 m,
2 Aug. 1996, /. G. Vargas C. 5061 (holotype,
OSH; isotypes, NY, US/ not seen). Figure 14.

Species Siphocampyli positione ambivalente umbellam
apicalem gregis Umbellatontm el folia verticillala gregis
Siphocampyli combi nans.

Shrub, 2—3 m lull; stems seandent. moderately
leafy, branched from the base, 3 cm diam. at base,
2.7—5 mm diam. toward apex, moderately pubes¬
cent with short scurfy brown hairs; latex white.
Leaves whorled, 3 per node, gradually decreasing
in size toward apex; lamina narrowly ovate, 6.5—1 1
X 2.2— 4.8 cm, subeoriaceous; adaxial surface dull
dark green, sparsely pubescent on the midrib with
short scurfy brown hairs; abaxial surface dull light
green, moderately pubescent with longer crisped
white or brown hairs; margin doubly serrate; apex
acuminate; base rounded or subeordate; petiole 5—
7 mm long, 1 — 1.5 mm diam., 1/10—1/20 as long as
the lamina, densely pubescent with short scurfy
brown hairs. Flowers 7 to 10 in a terminal sessile
umbel subtended by a whorl of 3 foliage leaves;
bracts linear, 14—15 mm long, 1 mm diam., the
apex acuminate; pedicels straight, ascending, 18—
22 mm long, 0.5— 0.6 mm diam.. ebracteolate,
densely pubescent with short scurfy brown hairs.
Hypanthium broadly obconic or hemispheric, 3-4

mm long, 5 mm diam., 1/12—1/20 as long as the
corolla, moderately pubescent with short scurfy
brown hairs. Calyx lobes linear-triangular, erect or
slightly spreading, 10—12 X 0.7-1 mm, 3— 3X as
long as the hypanthium, 1/3— 1/4 as long as the co¬
rolla tube, densely pubescent with short scurfy
brown hairs; apex acuminate; margin entire. Corolla
tubular, bilabiate, pink-purple with green-yellow
limb, 52-56 mm long, densely pubescent with short
still pale hairs; tube suberect, 38—43 mm long, 5—
5.5 mm diam. at base, narrowing to an isthmus 3—
4 mm diam. just above base, then gradually ex¬
panding to 6.5—8 mm diam. at mouth; dorsal lobes
linear-triangular, 13—14 X 1.3— 2.3 mm, ca. 1/3 as
long as the tube, the apex acuminate; lateral lobes
linear-triangular, 1 1-13 X 2—2.5 mm, the apex
acuminate; ventral lobe linear-triangular, 15—16 X

2.5— 2.8 mm, the apex acuminate. Staminal column
somewhat exserted between the dorsal lobes; fila¬
ment tube suberect, 47—50 mm long, I mm diam.,
glabrous; anther tube 2—2.1 mm diam., gently
curved, with a few long weak white hairs dorsally;
dorsal anthers 9—10 mm long, ca. 1/5 as long as
the filament tube; ventral anthers 6—7 mm long,
their apex pubescent with tufts ol stiff white hairs

1.5- 2 mm long. Fruit and seeds not seen, but apex
of ovary conic, pubescent.

Distribution, habitat, and phenology. Apparently
endemic to central Bolivia and known only from the
type collection.

Etymology. The specific epi t bet refers to the
species" equivocal taxonomic position in Wimmer’s
(1953, 1968) classification; from the Latin partici¬
ple ambivalens, ambivalent.

Relationships. Siphocampylus ambivalens clear¬
ly is referable to section Siphocampylus. subsection
Siphocampylus in Wimmer’s (1953, 1968) classifi¬
cation. Within that subsection, however, it com¬
bines features of two of the five greges recognized
by Wimmer (1953): the terminal umbellate inflo¬
rescence of grex Umbellati and the whorled leaves
of grex Siphocampylus. Siphocampylus grex Umbel¬
lati comprises nine species endemic to Colombia
| .S’, amalfiensis F. E. Wimmer, ,S. densiflorus Plan-
chon, S. glareosus Zahlbruckner, S. lindleyi Le-
maire, S. longibracteolatus F. E. Wimmer, S. lucifer
F. E. Wimmer, S. microstoma Hooker] or Venezuela
[S. moritzianus F. E. Wimmer, S. reticulatus (Willd-
enow ex Schultes) Klotzseh & H. Karsten ex Vatke|,
plus one additional species [,S. corymbiferus Pohl|
distributed in Peru, Bolivia, anil Brazil (Wimmer,
1953). Siphocampylus grex Siphocampylus compris¬
es seven species endemic to southern Brazil [5. du-
ploserratus Pohl, S. fluminensis (Vellozo) F. E. Wim-

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

227

mer, S. psilophyllus Pohl, S. sulfureus F. E.
Wimmer, and S. westinianus (Thunberg) Poll 1 1 or
Bolivia [5. orbignianus A. DC., S. tenuisepalus F.
E. Wimmer], plus one [S’, verticillatus (Chamisso)
G. Don] distributed in Brazil, Paraguay, Uruguay,
and Argentina.

All known species of Siphocampylus grex Um -
bellati Have alternate leaves. If one ignores its ver-
ticillate leaves, S. ambivalens will key to S. reti-
culatus of western Venezuela in Wi miner's (1953)
treatment of grex Umbellati. The two species are
similar in a number of features, including size and
shape of their leaves, their linear bracts, ebracteo-
late pedicels, pubescent corollas, and glabrous sta-
minal columns. However, S. reticulatus differs in its
shorter (1 — 1.8 m vs. 2—3 m) stature, turbinate (vs.
broadly obconic or hemispheric) hypanthium,
shorter (35—40 mm vs. 52-56 mm) corolla, and
shorter anthers (dorsal pair 5 mm vs. 9—10 mm).

The only species of Siphocampylus grex Umbel¬
lati known from Bolivia is S. corymbiferus, whose
range extends into Peru and Brazil. This species
likewise shows some similarity to S. ambivalens in
size and shape of its leaves, its ebracteolate pedi¬
cels, and glabrous staminal column. However, S.
corymbiferus differs in its shorter stature (0.6— 1.5
m vs. 2-3 m in S. ambivalens ), longer (8—15 mm
vs. 5-7 mm) petioles, ovate or lanceolate (vs. lin¬
ear) bracts, longer (30—45 mm vs. 18—22 mm) ped¬
icels, short turbinate (vs. broadly obconic or hemi¬
spheric) hypanthium, shorter (4—6 mm vs. 10-12
mm) calyx lobes, and shorter (40—50 mm vs. 52—
56 mm) glabrous (vs. pubescent) corolla.

All known species of Siphocampylus grex Siph¬
ocampylus have flowers solitary in leaf axi Is. If one
ignores its umbellate inflorescence, S. ambivalens
will key to 5. westinianus of southern Brazil in
Wimmer’s (1953) treatment of grex Siphocampylus.
The two are similar in a number of features, in¬
cluding size and shape of their leaves, size and
color of their corollas, and glabrous staminal col¬
umns. However, S. westinianus differs in its shorter
stature (0.6— 0.8 m vs. 2—3 m in S. ambivalens ),
bibracteolate (vs. ebracteolate) pedicels, turbinate
(vs. broadly obconic or hemispheric) hypanthium,
and shorter (3-4 mm vs. 10-12 mm) calyx lobes.

Two other verticillate-leaved species are endemic
to Bolivia: S. orbignianus and 5. tenuisepalus. Both
share a number of characters with the new species,
e.g., leaf size and shape and corolla size and color
are similar in S. ambivalens and S. orbignianus,
hypanthium shape and pedicel length in 5. ambi¬
valens and S. tenuisepalus. However, S. orbignianus
differs from S. ambivalens in its longer (35—40 mm
vs. 18—22 mm in S. ambivalens ) divergent (vs. as¬

cending) bibracteolate (vs. ebracteolate) pedicels,
broadly turbinate (vs. broadly obconic or hemi¬
spheric) hypanthium, and spreading or reflexed (vs.
erect or slightly spreading) calyx lobes. Siphocam¬
pylus tenuisepalus differs from S. ambivalens in its
glabrous (vs. pubescent) stems, narrower (2 cm vs.
2. 2^4.8 cm) lanceolate (vs. narrowly ovate) laminas
cuneate (vs. rounded or subcordate) at base, longer
(10 mm vs. 5—7 mm) glabrous (vs. pubescent) pet¬
ioles, longer (12-17 mm vs. 10-12 mm) spreading
or reflexed (vs. erect or slightly spreading) calyx
lobes, and longer (65 mm vs. 52—56 mm) corolla.

Aftercareful comparison, it is concluded that de¬
spite its umbellate inflorescence, S. ambivalens is
most likely allied to the verticillate-leaved species
that make up Siphocampylus grex Siphocampylus.
Geographic relationships support this view: all hut
one of the species of grex Umbellati are endemic
to Colombia or Venezuela while all but one of the
species of grex Siphocampylus are endemic to Bra¬
zil or Bolivia. Further, condensation of stem apices
to form corymbiform or umbellate inflorescences
appears to have occurred independently in several
genera of Lobelioideae ( Burrneistera , Centropogon,
Lobelia) while the sole occurrence of verticillate
leaves in the subfamily is in the species of grex
Siphocampylus. It is thus more parsimonious to hy¬
pothesize that S. ambivalens is a member of Siph¬
ocampylus grex Siphocampylus that has evolved an
umbellate inflorescence from solitary axillary flow¬
ers, rather than a member of grex Umbellati that
has evolved whorled leaves from alternate.

3. Siphocampylus longior Lammers, sp. nov.
TYPE: Peru. San Martin: Prov. Rioja, Buenos
Aires, along road Pedro Ruiz— Rioja,

05°42'09"S, 77°53'06"W, wet cloud forest, ca.
2000 m, 21 Mar. 1998, II. van der Werff B.
Gray, R. Vasquez & R. Rojas 15376 (holotype,
OSH; isotype, MO). Figure 15.

Species Siphocampyli subsect. Secundiflorum al> omni¬
bus caeteris speciebus hujus subsectionis petiolis lon¬
gioribus 15—23 mm longis, pedicellis longioribus 40—65
mm longis, calycis lobis longioribus 7—10 mm longis, et
antheris dorsalibus longioribus 11-14 mm longis facile
distinguenda.

Suffrutescenl liana; stems scandent, often hori¬
zontal or pendent, branched, moderately leafy, 2—5
mm diam., minutely scabrid; latex white. Leaves
secund; lamina ovate or oblong, 3.5—6 X 2.2— 3.2
cm, chartaceous; adaxial surface dull olive green,
glabrous; abaxial surface dull, paler, minutely sca¬
brid on the veins; margin minutely serrulate; apex
acute or obtuse; base rounded, truncate, or subcor¬
date, and sometimes asymmetric; petiole 15—23

228

Novon

A 2cm

Figure 15. Siphocampylus longior hammers. — A. Upper
portion of stem with flowers. (Drawn from the holotype,
van der Werff et al. 15376.) — B. Flower. (Drawn from the
paratype, van der Werff et al. 15746.)

mm long, 0.8-1 mm diam., 1/2— 1/3 as long as the
lamina, minutely scabrid. Flowers in a secund ter¬
minal bracteate raceme or sometimes in the axils
of unreduced leaves; bracts elliptic, 1.4— 1.9 X 0.4—
0.7 cm, minutely scabrid, acute or acuminate at
apex, attenuate at base, the margin minutely ser¬
rulate, on petioles 6—7 mm long; pedicels ascend¬
ing, curved toward apex, 40—65 mm long, 1.2— 1.8
mm diam., ebracteolate, minutely scabrid. Hypan-
thium turbinate, 5—8 mm long, 7—8 mm diam., 1/5—
1/7 as long as the corolla, minutely puberulent. Ca¬
lyx lobes subulate, ascending, sometimes recurved
toward apex, 7—10 X 0.8— 1.2 mm above the base,
slightly longer than the hypanthium, minutely sca¬
brid; apex acuminate; margin entire. Corolla (im¬
perfect in the material seen) green or maroon, 35—
40 mm long, puberulent, more densely so toward
base. Filament tube suberect, 32 mm long, 1 mm
diam., glabrous; anther tube 3 mm diam., slightly
curved, its surfaces glabrous; dorsal anthers 1 1—14
mm long, 1/3 as long as the filament tube; ventral
anthers 8—10 mm long, their apex pubescent with
tufts of stiff white hairs 2.5—3 mm long. Fruit and
seeds not seen, but apex of ovary conic.

Distribution , habitat , and phenology. Apparently
endemic to northern Peru, with both known collec¬
tions obtained along the road between Pedro Ruiz and

Rioja in Departamento de San Martin. Growing in wet
cloud forest at 1900-2000 m and flowering in March.

Etymology. The specific epithet is the compar¬
ative of the Latin adjective longus, i.e., longer, in
reference to the fact that this species' petioles, ped¬
icels. calyx lobes, and anthers are longer than those
of any other member of the subsection.

Relationships. With its secund terminal brac¬
teate raceme of relatively short-pedicel led flowers,
Siphocampylus longior is clearly a member of sec¬
tion Brachysiphon, subsection Secundiflori. This
subsection comprises nine species: S. purdieanus
Planchon is endemic to Colombia, S. ayersiae ham¬
mers to Bolivia, and the remainder are distributed
in Peru and southern Ecuador (Winuner, 1953;
hammers, 1998a).

Although the specimens at hand are imperfect,
with flowers either in bud or past anthesis, it is
clear that they cannot be equated with any previ¬
ously described member of subsection Secundiflori,
as their petioles, pedicels, calyx lobes, and anthers
are all significantly longer. In the other species of
the subsection, petioles are 4—15 mm long (vs. 15—
2.3 nun in S. longior), pedicels 8^12 long (vs. 40—
65 mm), calyx lobes 1—5 mm long (vs. 7-10 mm),
and dorsal anthers 7-1 1 mm long (vs. 1 1-14 mm).

Paratypes. PERU. Sail Martin: Prov. Rioja, along
road Rioja— Pedro Ruiz, El Mirador, 1900 m, 26 Mar.
1998, //. van der Werff R. Gray, R. Vasquez & R. Rojas
15746 (MO, OSH).

4. Siphoeampyliis plegmaloenidis hammers,
sp. nov. TYPE: Peru. San Martin: Prov. Rioja,
along road Rioja-Pedro Ruiz, ceja de la mon-
tana, El Mirador, 05°40'29"S. 77°46'25"W,
1850 m, 25 Mar. 1998, //. van der Werff, B.
Gray, R. Vasquez & R. Rojas I562B (holotype,
OSH; isotype, MO). Figure 16.

Specie's Siphocampyli subgregis Pyriformium ob caules
dimorphos et trichomata dimorpha ad orificium tubi an-
therarum insignis; caules primageni horizontales 1.2— 1.8
mm diametro volubiles dextrorsum (externe visus) sparsim
foliacei vel demum nudi, sed rami erecti 0.8—1 mm dia¬
metro confertim foliacei basin versus el remote foliacei
llagelliformes apicem versus; trichomata ad apicetn an-
therarum dorsalium 0.1— 0.3 mm longa, sed haec anther-
arum ventralium 1—1.4 mm longa.

Vine with dimorphic stems; main stems horizon¬
tal, elongate, dextrorsely twining (in external view),
1 .2-1.8 mm diam., sparsely leafy or eventually leaf¬
less, minutely scabrid; branches erect, 0.8—1 mm
diam., highly condensed and densely leafy at base,
whip-like anti sparsely leafy distally. Lamina nar¬
rowly elliptic (sometimes lanceolate on the elongate
portions of branches), 1.9—4 X 0.5— 1.3 cm, char-

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2002

Lammers

New South American Lobelioideae

229

Figure 16. Siphocampylus plegmatocaulis Lammers. —
A. Portion of horizontal twining main stem with condensed
young branch. — B. Young capsule, topped by marceseent
corolla. — C. Portion of main stem with erect branches
older, elongated, whip-like. (Drawn from the holotype, van
der Werjf el al. 15628.)

taceous; adaxial surface dull olive green, glabrous;
abaxial surface dull, paler, sparsely scabrid with
minute dark antrorse trichomes; margin minutely
serrulate or crenulate with 1 to 4 exserted dark
callosities per side; apex acute (sometimes acumi¬
nate on the elongate portions of branches), mucro-
nate; base cuneate; petiole 3—7 mm long, 0.6— 0.8
mm diam., 1/4—1/10 as long as the lamina, mi¬
nutely scabrid. Flowers solitary in axils of leaves
on the branches; pedicels ascending, 15—38 mm
long, 0.5— 0.7 mm diam., ebracteolate, hispidulous
or minutely scabrid. Hypanthium broadly obeonie
or turbinate, 2.5— 3.5 mm long, 3.1— 3.6 mm diam.,
ca. 1/10 as long as the corolla, hispidulous. Calyx
lobes narrowly triangular, erect, slightly recurved
at tips, 0.8— 1.5 X 0.5— 0.7 mm, 3/10—2/5 as long
as the hypanthium, ca. 1/20 as long as the corolla
tube, hispidulous at base; apex acuminate; margin
entire. Corolla deep [jink, red, or red-purple, tu¬
bular, bilabiate, 27-32 mm long, sparsely hispi¬
dulous; tube comprising a proximal isthmus ca. 8
mm long, 2.5— 3.4 mm diam. at base and 1.5-2. 2
mm diam. at apex, which bends and flares abruptly
into an expanded distal portion 8—16 mm long and
4.5— 5.5 mm diam. at mouth; dorsal lobes linear-

triangular, 7-10 X 1.5-2 mm, spreading, 1/2 — 1/3
as long as the tube, acuminate at apex; ventral
lobes linear-triangular, 7—9 X 1.5— 1.9 mm, spread¬
ing, acuminate at apex. Staminal column slightly
exserted; filament tube suberect, 23—26 mm long,
1 mm diam., glabrous; anther tube 1.6— 1.7 mm
diam., suberect, its surfaces glabrous; dorsal an¬
thers 4.5—5 mm long, 1/5 as long as the filament
tube, their apex pubescent with tufts of stiff white
hairs 0.1— 0.3 mm long; ventral anthers 3.5-4 mm
long, their apex pubescent with tufts of stiff white
hairs 1—1.4 mm long. Capsule obovoid, 8—9 mm
long, 5.5—6 mm diam., 2/3— 3/4-inferior; mature
seeds not seen.

Distribution , habitat, and phenology. Apparently
endemic to northern Peru, collected a few times in
the vinity of Pedro Ruiz in Departamento de San Mar¬
tin. Growing in pajonal vegetation on exposed ridges
and in shrubs bordering cloud forest, 1850-2200 m.
Flowering from February to August; fruit maturing
from March to August.

Etymology. The specific epithet refers to the
dextrorsely twining stems; from the Greek nouns
TrAeypta, twist, and kcioAos, stem.

Relationships. Though many species of Sipho¬
campylus are scandent, a truly twining habit is rare
in the genus, known from only S. convolvulaceus
(Chamisso) G. Don of Brazil and S. cordatus (Willd-
enow ex Schultes) F. E. Wimmer of Venezuela and
Colombia. These two species are referable to Siph-
ocampylus sect. Siphocampylus, subsect. Siphocam¬
pylus, grex Dissitifiori, subgrex Pyriformes, as is the
newr species. These three species further share
ebracteolate pedicels, obconic or turbinate hypan-
thia, and glabrous filament tubes. However, S. con¬
volvulaceus and S. cordatus differ from S. plegma¬
tocaulis in their longer petioles (5—20 mm vs. 3—7
mm in S. plegmatocaulis); larger laminas (4—9 X
1.2—3 cm vs. 1.9-4 X 0.5— 1.3 cm) that are round¬
ed, truncate, or subcordate (vs. cuneate) at base;
longer calyx lobes (2—7 mm vs. 0.8— 1.5 mm); co¬
rollas that are longer (45—60 mm long vs. 27—32
mm long) and broader (2—3 mm at isthmus vs. 1 .5—
2 mm. 5-9 mm at mouth vs. 4.5— 5.5 mm); longer
dorsal anthers (6—7 mm vs. 4.5—5 mm) that are gla¬
brous (vs. bearded) at apex; and longer capsules
( I ..5-2 cm vs. 0.8-0. 9 cm).

Though S. convolvulaceus and S. cordatus are
twining vines, neither produces the unique dimor¬
phic stems of S. plegmatocaulis. For this reason, il
seems unlikely that the new species is closely re¬
lated to either of the other two twiners. Instead. .S'.
plegmatocaulis may represent an independent ori¬
gin of the twining habit, perhaps as an adaptation

230

No von

to life in exposed, nutrient-poor pajonal, an unusual
lml>itat for the genus.

In addition to the dimorphic twining stems, the
dimorphic pubescence surrounding the orifice of
the anther tube in S. plegmatocaulis is unique iti
Siphocampylus. Most members of the genus only
have apical tufts of trichomes on the ventral pair
of anthers. Just five species bear tufts of trichomes
at the apex of all five anthers: S. isochiliis, S. os-
citans B. A. Stein, S. sceplrum (sometimes all five
glabrous), S. rusbyanus Britton, and S. splendens (F.
K. Wimmer) Jeppesen ex B. A. Stein. However, in
these five species, the hairs of the dorsal and ven¬
tral anthers are similar iti size, while those of S.
plegmatocaulis differ significantly: 0.1— 0.3 mm on
the three dorsal anthers, 1-1.4 mm long on the ven¬
tral pair. Furthermore, these five species all have a
Burmeistera- like anther tube with a wide open or¬
ifice (cf. Stein, 1987; I ,a miners, 1998a). In S. pleg¬
matocaulis, the orifice of the anther tube is occlud¬
ed, as is typical lor Siphocampylus and most genera
of Lobelioideae. None of these five species is re¬
ferable to Siphocampylus subsect. Siphocampylus ,
and none is believed to be related to .S'. plegmato¬
caulis. In summary, the affinities of this new species
remain obscure.

Paratypes. PERU. Sun Martin: Prov. Rioja, Pedro
Ruiz— Moyobamba road, km 368, Campamento Oarcfa,
05°45'S, 77°43'W, pajoruil vegetation, a vegetation type
developing due to exposure and poor soil, here found on
ridges, 1900 m, 14 Aug. 1983, D. A. Smith 4803 (MO.
OSH); Prov. Rioja, Rfo Nieva valley, ca. 100 km W of
Rioja on road to Pedro Ruiz, I km E of Puente Rfo Nieva,
05°47'S, 77°40'W, twining through shrubs along road in
low cloud forest, 2000 m, 16 Feb. 1985, 8. Stein & C.
Todzia 2204 (MO).

3. Siphocampylus praevaricator hammers, sp.
nov. TYPE: Venezuela. Merida: Paramo de los
Colorados, road to San Isidro Alto, 4 km from
junction with Estanques— Paramo did Molino
road, 2600 m, 3 Feb. 1978, ./. L. Luteyn, M.
Lebron-Luteyn & /,. Ruiz-Teran 5408 (holotype,
OSH; isotypes, MERE not seen, NY, VEN not
seen). Figure I 7.

Species ob trichomata arbusculiformia Centropogonem
subsect. Brevilimbatos (praecipue C. acrodentatum et C.
(datum) simulans sed ovarii apice conico et floribus in
racemo terminali subumbellato, ergo in Siphocampylum
gregem Umbellalos disposita; in hoc grege .5’. reticulata et
.S’, moritziano affinis, sed ab his speciebus triehomatibus
arbuscuiiformibus in caule loliis inflorescentia corollaque,
calycis lobis triangularis erectis 3^f mm longis, tilamen-
torum tubo minute puberulo, et antherarum tubo apicem
versus hirto statirn distinguenda. Ab omnibus caeteris
speciebus Siphocampyli cum triehomatibus arbuscuiifor¬
mibus racemo terminali subumbellato differl.

Figure 17. Siphocampylus praevaricator hammers. — A.
Arbusculiform trichomes on abaxial leaf surface. — R. Up¬
per portion of fertile stem. — C. Upper portion of sterile
stem. (Drawn from the holotype, Luteyn et id. 5408.)

Suffrutescent herb; stems scandent, moderately
leafy, 2—2.5 mm diam., densely pubescent with ar¬
busculiform trichomes; latex white. Lamina ovate
or narrowly ovate, 4.5— 6.5 X 2.8— 3.5 cm. coria¬
ceous. those on the stem below the inflorescence
smaller; adaxial surface shiny olive green, glabrous
or with a few arbusculiform hairs along the midrib;
abaxial surface dull ochreous, densely pubescent
with arbusculiform trichomes; margin irregularly
dentate with smaller and larger teeth alternating;
apex acute or acuminate; base cordate; petiole 14—
18 mm long, 1.3— 1.5 mm diam., 1/3— 1/4 as long as
the lamina, sparsely pubescent with arbusculiform
trichomes. Flowers 30 to 35 in a terminal subum-
bellate raceme; rachis congested, condensed, 25
mm long; bracts linear, 4—6 X 0.2— 0.3 mm, the
apex acuminate; pedicels straight or somewhat
curved, ascending, 22—27 mm long, 0.7—1 mm
diam., ebracteolate, densely pubescent with arbus¬
culiform trichomes. Hypanthium turbinate, 5 mm
long, 5 mm diam., 1/8—1/10 as long as the corolla,
densely pubescent with arbusculiform trichomes.
Calyx lobes triangular, erect, 3-4 X 1.5—2 mm,
3/5— 4/5 as long as the hypanthium, 1/8—1/10 as
long as the corolla tube, densely pubescent with
arbusculiform trichomes; apex acuminate; margin

Volume 12, Number 2
2002

Lammers

New South American Lobelioideae

231

entire. Corolla bilabiate, deep violet outside, paler
within, 42-47 mm long, moderately pubescent with
arbusculiform trichomes; tube suberect, 30—34 mm
long, 4.5— 4.8 mm diam. at base, abruptly narrowing
to an isthmus 2. 2-2.3 mm diam. just above base,
then gradually expanding to 6. 7-7. 5 mm diam. at
mouth; dorsal lobes narrowly triangular, erect, 12-
13 X 2. 3-3. 5 mm, 2/5 as long as the tube, the apex
acuminate; ventral lobes narrowly triangular,
spreading, 7-10 X 2-2.8 mm, the apex acuminate.
Staminal column exserted between the dorsal lobes;
filament tube suberect, pale violet. 43—45 mm long,
0.7-1 mm diam., minutely puberulous; anther tube
gray, 2-2.2 mm diam., suberect, sparsely pubescent
with straight simple white hairs; dorsal anthers 6
mm long, 1/7— 1/8 as long as the filament tube; ven¬
tral anthers 4.5—5 mm long, their apex pubescent
with tufts of stiff white hairs 0.8-1 mm long. Fruit
and seeds not seen, but apex of ovary conic.

Distribution, habitat , and phenology. Apparently
endemic to western Venezuela and known only from
the type collection.

Etymology. The specific epithet is the Latin
noun praevaricator, one who speaks or behaves in
a deceitful, ambiguous, or equivocal fashion. While
the newT species belongs to Siphocampylus, its ar¬
busculiform trichomes might mislead one into as¬
suming that it is a member of Centropogon.

Relationships. As noted under Burmeistera ar-
busculifera, arbusculiform bairs are common in
Centropogon but very rare in other genera of Lob¬
elioideae. Just three species of Siphocampylus have
been reported to bear such trichomes (McVaugh,
1949; Lammers, 1998a): S. columnae (L. f.) G. Don
and 5'. furax F. E. Wimmer of northern Ecuador,
and S. fallax Lammers of northern Peru. All are
referable to Siphocampylus sect. Siphocampylus.
subsect. Siphocampylus, grex Dissitiflori, a group
that is characterized in part by flowers solitary in
the upper leaf axils (Wimmer, 1953). Siphocampy¬
lus praevaricator. on the other hand, bears its flow¬
ers in a congested terminal subumbellate raceme,
and so is referable to a different element of sub¬
section Siphocampylus, grex Umbellati (described
above under S. ambivalens).

In Wimmer’s (1953) treatment of Siphocampylus
grex Umbellati, S. praevaricator would key with dif¬
ficulty to S. moritzianus and S. reticulatus. both
these species are endemic to Venezuela and occur
in Merida; the former species is endemic to that
state, the latter more widely distributed. However,
these species differ from S. praevaricator in their
simple (vs. arbusculiform) trichomes on stems,
leaves, and inflorescences; filiform or sublinear (vs.

triangular) calyx lobes which are longer (6-10 mm
vs. 3—4 mm) in S. reticulatus and recurved at apex
(vs. erect) in S. moritzianus ; glabrous (vs. minutely
puberulent) filaments; and glabrous (vs. pubescent)
anther tubes.

Because arbusculiform hairs are so much more
common in Centropogon than in Siphocampylus,
one must consider the possibility that this anoma¬
lous Siphocampylus has been described already as
a species of Centropogon, An author who did not
have mature fruit or did not dissect a flower to ex¬
amine the ovary might well be led to assign this
plant to Centropogon. The two genera clearly are
closely related (Pepper et ah, 1997; Lammers,
1998a; Buss et ah, 2001), and there are many ex¬
amples of species of one genus being described un¬
der the other.

Siphocampylus praevaricator does indeed show
some resemblance in general size and shape of the
leaves and flowers to two species of Centropogon
subsect. Brevilimbati endemic to high-elevation
paramo habitats in Merida: C. acrodentatus F. E.
Wimmer and C. alatus Gleason. However, both
these species (as well as all other species of sub¬
sect. Brevilimbati ) bear solitary axillary flowers
rather than terminal subumbellate racemes. Fur¬
thermore, these two species of Centropogon have
longer pedicels (8 cm in C. acrodentatus, 5—6 cm
in C. alatus, vs. 2. 2-2. 7 cm in S. praevaricator),
which are bibracteolate toward the base (vs. ebrac-
teolate). Thus it does not appear that 5. praevari¬
cator has been described already under the genus
Centropogon,

One last hypothesis to consider is that S. prae¬
varicator, with the terminal subumbellate raceme
characteristic of Siphocampylus grex Umbellati and
the arbusculiform hairs characteristic of Centropo¬
gon subsect. Brevilimbati, is the result of hybrid¬
ization between S. moritzianus or S. reticulatus on
the one hand and C. acrodentatus or C. alatus on
the other. The co-occurrence of all four species in
Merida certainly makes this possibility tempting.
However, this seems unlikely. Neither of the salient
characters of S. praevaricator is intermediate be¬
tween the hypothesized parents. The inflorescence
is as tight and congested as that of any member of
Siphocampylus grex Umbellati, with highly reduced
bracts; there is no tendency whatsoever toward the
production of flowers in the axils of unreduced fo¬
liage leaves. Similarly, the hairs are fully as dense
and arbusculiform as any member of subsection
Brevilimbati, showing no tendency toward reduction
in branching or density. It thus seems best to hy¬
pothesize that in S. praevaricator ; the presence of
these unusual trichomes is not due to gene flow

232

Novon

from a species of Centropogon but rather is the re¬
sult of yet another independent evolutionary origin
of these structures.

Addendum

In addition to the new species described here,
the 4(H) specimens of neotropical Lobelioideae
identified over the past four years have included
some additional collections of species described as
new in a previous paper (hammers, 1998a), most
of which were based solely on the type gathering.

Ilurmeistera racemiflora hammers, Brittonia 50:
258. 1998.

COLOMBIA. Narino: en la vfa Tumaco— Pasto, km
105, a 7 km de el Diviso, 850 in, en bosque, 12 June
1986, H. Ledn & I*. Maas 1622 (U). The species was de¬
scribed solely from the type, collected in Carehi, Ecuador.
This second collection, made five years earlier, extends
the species range across the border into adjacent Colom¬
bia.

Centropogon dianae hammers. Brittonia 50: 257.

1998.

PERU. Cuzco: kins. 143—152, ( Hlantaytambo— A I la-
mayo Rd„ 5fi — <57 km NW of Ollantaytambo, 3110-3415
m, montane cloud forest, infrequent, 1 I Dee. 1978, L.
Lute yn & M. Lebrdn-Luteyn 6456 (1 ). I he species was
described solely on the basis of the type. This second
gathering was collected five years earlier in the same gen¬
eral region.

Centropogon jeppesenii hammers, Brittonia 50:
252. 1998.

ECUADOR. IMupo: trail to ha Bonita ca. 1—2 km from
end of Tulcdn-Sta. Barbara rd., ca. 25 km below Sta. Bar¬
bara, 2100-2150 m. 0°35'N, 77°30'W. cloud forest &
clearings above Rfo Chingual, 2(5 May 1985, II. .4. Stein
2871 (MO). Die species was described solely on the basis
of the type gathering. This second collection was made
three years later at or near the type locality.

Siphoeampylus sinilux hammers, Brittonia 50:
240. 1998.

BOLIVIA. Santa < ,ruz: Prov. Ichilo, Parque Naeional
Amborb, ca. 15 km (SE) up the Rfo Pitasama from the
Rfo Surutii, moist tropical forest on lower montane slopes,
sandstone, 7(H) m, I7°44'5, 63°40'W. 26 Aug 1985, J. C.
Solomon & S. Urcullo 14(174 (MO). This collection ex¬
tends the species’ distribution to a fourth province in Dep-
to. Santa Cruz; it was previously reported from localities
in Prov. Andres Ibanez. Cordillera, and Florida.

Acknowledgments. My sincere thanks to the
collectors and institutions who have made this pa¬
per possible by continuing to send me specimens
ol hobelioideae for identification. This research was

supported by Faculty Development Research Grant
FDR-737 from the University ol Wisconsin Osh¬
kosh (UWO). The illustrations represent the first
published work of UWO student Yvette Evrard, to
whom 1 extend my sincere appreciation. Thanks
also to my colleague Neil A. Harriman, who re¬
viewed an earlier version of the manuscript; and to
Brad Boyle, harry Dorr, Jim huteyn, Jon Ricketson,
Zachary Rogers, and Jim Solomon, who provided
additional information on collectors and collections.

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Dyckia delicata (Bromeliaceae), a New Species from Rio Grande

do Sul, Brazil

Jodo Larocca

Centro de Ciencias <la Saude, Unisinos, Caixa Postal 275, 93022-000-Sao Leopoldo, MS,

Brazil. jlaroeea@bios. imisinos.br

Marcos Sobral

Faculdade de Farmacia UFRGS, Ipiranga 2752, 906 10-01 0-Port o Alegre, MS, Brazil.

sobral@farmaeia.ufrgs.br

Abstract. Dyckia delicata, a narrow-endemic
bromeliad from basaltic outcrops in the state of Rio
Grande do Sul. Brazil, is described and illustrated.
It is similar to Dyckia hebdingii, from which it is
distinguished by its leaves with soil, non-injurious
spines, glabrous and few-branched or unbranched
inflorescences, and triangular-shaped seeds.

Resumkn. Se describe e ilustra Dyckia delicata,
una Bromelidcea endemica restringida a los aflo-
ramientos basal ticos del estado de Rio Grande do
Sul. Brasil. Fa especie se parece a Dyckia hebdin¬
gii, de la cual se difereneia por las hojas con es-
pinos suaves, no punzantes, por la inflorescencia
glabra y poco o no ramificada y por las semillas
triangulares.

key words: Brazil. Bromeliaceae, Dyckia.

The first thorough study of Bromeliaceae in Rio
Grande do Sid (Winkler, 1982), tin1 southernmost
Brazilian state, listed 15 native species of Dyckia.
8 of w Inch are endemic. Since then, 4 more were
described (Irgang & Sobral, 1987; Smith, 1988,
1989; Strehl, 1998), so that now there are 12 en¬
demic species, at least half of which are known
from oidy one collection site — that is, can be con¬
sidered as narrow endemic's. This relatively high
rate of endemism could be due to the occurrence
of pluvial forests on about half of the state, resulting
in the isolation of some mountain tops and rocky
outcrops; this, in turn, may have caused some iso¬
lated populations of plants to evolve as species of
their own.

During collecting trips to die hinterland of Rio
Grande do Sul, a geographically restricted, small,
and soft-spined species of Dyckia growing in iso¬
lated basaltic outcrops came to our attention and is
herein described as new.

Dyckia delicata Larocca & Sobral, sp. nov. TYPE:
Brazil. Rio Grande do Sul: Barros Cassal, Lin-
ha Pessegueiro, ea. 5 km da sede do munief-
pio, 29°07'S, 52°35'W, 350 m s.m., Dec. 1996,
./. Larocca , ,4. I). Nil son A: M. Sobral 96/00!
(holotype, ICN; isotypes, MBM, US, ZSS). Fig¬
ures 1 , 2.

| Live specimens at Porto Alegre Botanical Garden
and Fundayao Gaia at Pantano Grande.]

Species I). hebdingi proxima, a qua spinis foliorum
papyraceis, non pungentibus, inflorescentiis pauciramosis
glabrisque et seminibus triangularibus recedit.

Rosetted herb, 0. 2-0.6 m diam. Leaves succu¬
lent. cinereous to reddish-cinereous, eventually
curled toward the soil in some isolated individuals,
densely covered on both faces with pale scales up
to I mm long; leaf sheaths wide-triangular; blades
linear-triangular, 5-20 cm long and 0.7-2 cm wide;
apex forming a rigid spine; margin undulate with
soft, flexible spines 5-7 X 1-1.5 mm, straight, re-
trorse or anlrorse, spaced 3—5 mm from each other.
Inflort •scences 80— 1 50- flowered, erect, simple or
compound spikes, glabrous, with tin* axis softly
ridged longitudinally, to 1.2 m long, the main inflo¬
rescence axis up to 5 mm diam. at the base; sec¬
ondary branches when present 1-9, to 35 cm long.
Scape bracts linear-lanceolate, papyraceous, soft to
the touch, with 10—20 parallel nerves, glabrous,
longer than the internodes in the lower half of the
axis and shorter in the upper one, 30-55 X 6-10
mm. with spines up to 3 mm long in the upper
portion. Floral bracts elliptic-ovate, rounded or
acute at apex, glabrous, 5.5-8 X 5-6 mm. Galyx
green, the sepals ovate-oblong, rounded at apex, 6—
7 X 3.5-4 mm, the margins hyaline for about I
mm. Corolla campanulate or urceolate, yellow, the
petals lanceolate-spathulate to oblaneeolate, 10-13
mm long, 3—4 mm wide. Stamens 6, 9—10 mm. the
antisepalous and epipetalous somewhat unequally

Novon 12; 234-236. 2002.

Volume 12, Number 2
2002

Larocca & Sobral
Dyckia delicata from Brazil

235

Figure I. Flowering individuals of Dyckia delicata 1 1, a r-
occa et al. 96/01). Scale bar = 100 nun. Photograph J.
I .aroeca.

f igure 2. Dyckia delicata. — A. Schematic drawing of
flower in abaxial view ( Sobral et al. 9204). — B. Seed
( Larocca & Zdchia 95/025). Scales: A: 3 mm; B: I mm.

fused with the base of the corolla; anthers lanceo¬
late-ovate, ca. 2X1 mm. Style 7—10 X 0.2 mm;
stigmatic lobes 3, papillose. Ovary triquetrous, ob¬
long-acute, 7 mm long and 2.5—3 mm diam.. triloe-
ular with numerous ovules centrally attached in two
longitudinal rows per locule. Capsules elliptic, 9—
12 X 3—5 mm. Seeds triangular-shaped, 1.2— 1.7 X
0.5—1 mm, with a reduced wing 0.1— 0.2 mm wide
along the longitudinal axis.

Habitat, distribution, and phenology. Rupicolous,
collected on steep, 60°— 90° basaltic cliffs with north¬
eastern exposure, where it occurs with Parodia ha-
selbergii (Rumpler) F. H. Brandt (Cactaceae), Sin¬
ning to macrostachya (Lindley) Chautems
(Gesneriaceae), and Tillandsia recurvata (L.) L.
(Bromeliaceae). This species is known only from
the type locality, and appears to be a narrow en¬
demic of basaltic outcrops in Rio Grande do Sul.
fhe collection site at Linha Pessegueiro consists of
two small mountains with basaltic outcrops some
100 m apart. We have found the species growing
on both outcrops, and we think it is probable that
it also grows on other near outcrops, although we
were not able to verify this. Flowers were observed

in November, December, and January, and fruits in
January and February (pers. obs. from the field and
cultivated material).

Etymology. fhe species was named after the
Latin word “delicata,” delicate, in allusion to its
soft, non-injurious spines, the most evident anil
readily verifiable character of the species.

Affinities. Dyckia delicata is apparently related
to D. hebdingii L. B. Smith (Smith & Downs. 1974;
Winkler, 1982), another narrow endemic known
from only one locality on the southern slopes of the
southern Brazilian plateau known as Serra Geral,
250 km southeast of the collection site of D. deli¬
cata. fhe two species can be set apart by the fol¬
lowing characters:

la. Leaves with rigid spines, injurious to the touch;
inflorescences profusely branched, with ramifi¬
cations up to third degree; scape bracts reddish
and divergent from the axis; scapes lepidote;
flowers pilose; floral bracts always shorter than

the sepals; seeds linear-shaped . . Dyckia hebdingii

lb. Leaves with soft spines, not injurious to the
touch (except the apical spine); inflorescences
simple or with ramifications up to second degree,
in this case with fewer than 10 branches; scape
bracts green, not divergent from the axis; scapes

236

Novon

and flowers glabrous; floral bracts varying from
shorter than to longer than sepals; seeds trian¬
gular-shaped . Dyckia delicata

Paratype. BRAZIL. Rio Grande do Sul; Banos Cas-
sal, Linha Pessegueiro, ca. 5 km da sede do municfpio,
29°07'S, 52°35'W, 350 m s.m., Dec. 1995, Larocca &
Zdchia 95/025 (ICN).

Acknowledgments. We thank Ari Nilson and
Renato A. Zachia lor their kind help in collecting
Dyckia delicata, Daniel B. Falkenberg for useful
comments on the text, and two anonymous review¬
ers for insightful improvements to the manuscript.

Literature Cited

Irgang, B. E. & M. Sobral. 1987. Drckiu agudensis (Bro-
ineliaceae), nova espeeie do Rio Grande do Sul, Brasil.
Napaea 3: 5-7.

Smith, L. B. 1988. A giant Dyckia mystery. J. Bromeliad
Soc. 38: 248-249.

- . 1989. Another giant Dyckia mystery. J. Brome¬
liad Soc. 39: 206-207.

- & R. J. Downs. 1974. Bromeliaceae— Pitcairnioi-

deae. Flora Neotropica 14: 1-660.

Strehl. T. 1998. Novas bromelias do Rio Grande do Sul.
Bromtdia 4: 15—20.

Winkler, S. 1982. Die Bromeliaceae von Rio Grande do
Sul. Documenta Naturae 3: 1-90.

Notes on the Species of Cycas (Cycadaceae) from Sri Lanka and
Islands of the Andaman Sea

A. ,/. Lindstrom

Nong Nooch Tropical Garden, 34/1 Sukhumvit Hwy., Najomtien, Sattahip,
Chonburi, 20250, Thailand

K. D. Hill

National Herbarium of New South Wales, Royal Botanic Gardens, Mrs Macquaries Road,

Sydney 2000, Australia

ABSTRACT. Two cycad species, Cycas nathorstii
endemic to Sri Lanka and Cycas zeylanica native
to Sri Lanka and islands of the Andaman Sea, are
discussed. A new combination, Cycas zeylanica (J.
Schuster) Lindstrom & K. D. Hill, is established,
elevating this taxon from subspecies status on the
basis of cataphyll and leaflet characters. Both Cycas
species are lectotypified. Distribution of the two
species is investigated, and a key to the species is
provided.

Key words: Andamans, Cycadaceae, Cycas, Sri
Lanka.

The identity of the cycads native to Sri Lanka
has never been clearly understood from the earliest
recording of these plants in the region. The first
record for this country was in Flora Zeylanica, pub¬
lished by Linnaeus in 1747. Linnaeus (1753) later
treated this as one element of Cycas circinalis L.
Schuster (1932) separated the Sri Lankan material
as C. nathorstii J. Schuster, and de Laubenfels and
Adema (1998) included some but not all Sri Lan¬
kan material in C. sphaerica Roxburgh.

Cycas sphaerica is endemic to southeastern In¬
dia, and only C. nathorstii anti C. zeylanica are
therefore here recognized as being native to Sri
Lanka. The latter also occurs in the Andamans and
Nicobar Islands where its populations are consid¬
ered vulnerable.

About 90 species are recorded for the genus Cy¬
cas L. worldwide. Of these, 27 species are Austra¬
lian and about 35 are Indo-Chinese. Cycas also oc¬
curs in the Malesian region, Japan, India, and Sri
Lanka extending to Micronesia and Polynesia,
Madagascar, and East Africa. Plants are commonly
understory shrubs in forest, woodland, or savanna
habitats. Four sections have been recognized (Hill,
1995). but only one of these sections with two sub¬
sections occurs naturally in Sri Lanka. Three sub¬
sections are recognized, circumscription following

Hill (1995), with two occurring in Sri Lanka: sub¬
section Cycas and subsection Rumphiae. The full
range of the section is from India and southern In¬
dochina south to Australia, and from East Africa
east to Tonga.

Key to the Species of Cycas in Sri Lanka

I a. Lateral spines on megasporophylls distinct; cat-
aphylls thinly sericeous or glabrous with age, not
pungent, 50—60 mm long; midrib of median leaf¬
lets raised above and below; 6—10 ovules; no

spongy layer . C. nathorstii

lb. Lateral spines on megasporophylls indistinct;
cataphylls pilose, pungent, 100-120 mm long;
midrib of median leaflets raised above and Hal

below; 2—5 ovules; spongy layer present .

. C. zeylanica

Cycas subsect. Cycas (approx. 5 species) is de¬
fined by the absence of a spongy endocarp, the
presence of fibers in the sarcotesta, and a narrow
megasporophyll lamina. It ranges from India to Sri
Lanka. A single endemic species is found in Sri
Lanka.

Cycas nathorstii J. Schuster, Pflanzenr. 99: 76,
fig. 10E. 1932. TYPE: Sri Lanka. Central and
northern parts of the isle, 1866, G. Thwaites
36 89 in Herb. Barbey-Boisser (lectotype, des¬
ignated here, G; isolectotypes. A, K, LE, P).

Stems arborescent, 1—4.5 m tall. Leaves bright
green, semiglossy, 160—180 cm long, flat (not
keeled) in cross section (opposing pinnae inserted
at 180° on raehis), with 140 to 170 leaflets, tomen-
tum shedding as leaf expands. Petiole 45—55 cm
long (25—30% of total leaf), petiole glabrous, spi-
nescent for 90—95% of length. Basal leaflets not
gradually reducing to spines. Median leaflets sim¬
ple, weakly discolorous, 190—310 X 9—14 mm, nar¬
rowed to 3-4 mm at base (to 25—35% of maximum
width), 17—20 mm apart on raehis; median leaflets

Novon 12: 237-240. 2002.

238

Novon

flat; margins flat; apex softly acuminate, not spi-
nescent; midrib raised above, raised below. Cata-
phylls narrowly triangular, soft, sparsely sericeous
or lacking tomentum, 50—60 mm long, persistent.
Pollen cones narrowly ovoid, orange; microsporo-
phyll lamina firm, not dorsiventrally thickened, 30—
40 X 15—22 mm, apical spine prominent, gradually
raised, 10 mm long. Megasporophylls 15—30 cm
long; ovules 6 to 10, glabrous; lamina lanceolate,
40—65 X 18—25 mm, shortly dentate with 26 to 40
lateral spines 1—4 mm long; apical spine distinct
from lateral spines, 10—14 mm long. Seeds flat-
tened-ovoid; sarcotesta yellow; fibrous layer pre-
stmt; sclerotesta smooth; spongy layer absent.

Historical notes. This upland cycad from Sri
banka has been generally known as C. circinalis
(Trimen, 1898) and was in fact one element of the
protologue of the latter (Linnaeus, 1747). German
historian and sometime botanist Julius Schuster
distinguished C. nathorstii from C. circinalis in
1932 on the basis of specimens collected by George
Thwaites and distributed to European herbaria by
Swiss botanist and philanthropist William Barbey-
Boissier (1842—1914). Thwaites (1812—1882) was
superintendent of the Royal Botanic Garden at Per-
adeniya in Ceylon from 1849 to 1880. Collection
detail cited (Schuster, 1932: 76) was “Ceylon:
Thwaites 1866 n. 3689 in Herb. Barbey-Boissier."
Uie Barbey-Boissier herbarium and types are in G,
including Thwaites’s specimen. This may not have
been the specimen examined by Schuster, as it has
no annotation by him and was filed as undeter¬
mined. Schuster’s types were mainly in B and were
destroyed during World War II. This sheet at G in¬
cludes bodi leaflets and megasporophylls, and is
here designated the lectotype. I)e Laubenfels and
Adema (1998) included C. nathorstii in the syn¬
onymy of C. sphaerica, but also placed some Sri
Lankan collections of the same taxon in C. circin¬
alis.

Distinguishing features. Cyras nathorstii is dis¬
tinguished from C. circinalis and C. sphaerica of the
Indian mainland by its more robust habit, wider
leaflets, and larger male cones with longer and
more curved apical spines on mierosporophylls.
Megasporophyll apices of C. nathorstii are narrowly
triangular with numerous fine lateral spines ex¬
tending almost to the very tip, whereas the Indian
species possess broader apices with a distinct ex¬
tended apical spine that is free from lateral teeth.
Cycas nathorstii is distinguished from C. zeylanica
by the more closely spaced and more ehartaceous
leaflets, the shorter, softer and less pubescent cat-
aphylls, and lack of a spongy layer in the seeds.

Distribution and habitat. Known only from Sri
Lanka, where it occurs in inland and upland forests
in the north ol the island, usually in somewhat dry
sites.

Reproductive biology. No confirmed or potential
pollinator has been recorded for Cycas nathorstii.

Conservation status. Still locally frequent, al¬
though not in great numbers, this species is re¬
garded as vulnerable (IUCN, 1998. Red List cate¬
gory VU).

Vernacular. Madu.

Etymology. Honoring Swedish palaeobotanist
Alfred Gabriel Nathorst (1850—1921), professor at
the Natural History Museum in Stockholm.

Specimens examined. SRI LANKA. Ceylon, herb. P.
Perm. s.n. (L); hort. Kew. ex Ceylon, 1875, Thwaites 4230
(K); between Kuda Patessa and Maha Patessa, 15 July

1969, Camay iC Wirawan 1145 (A, K. I., NY); ridge S of
Na-Ulpota, 8 Aug. 1973, Jayasuriya 1278 (K); Maneran-
gata district. Bibile, 18 July 1972, Pepper & de Silva 4723
(k); Manerangata district, 3.2 kin K of Bibile, 28 Nov.

1970, Eosherg & Sachet 53148 (K. NY); Mandae, Old-
manas 53 (L); Moraragale district, Nilgala, Gal Oya Na¬
tional park, I May 1975, Jayasuriya 1951 (L); NW Prov¬
ince, Puttalam district, Wilpattu National Park, Kudu
patassa, 1 Nov. 1974, Davidse 8237 & Sumithraarachi
(BRI, K. L).

Cycas subsect. Rumphiae K. D. Hill is uniquely
defined by the presence of a layer of spongy tissue
within the seed. Its distribution is very wide, ex¬
tending from Africa to Fiji and Tonga, and from
New Guinea north to southern coastal Indochina.
One species, Cycas zeylanica, occurs in Sri Lanka
and islands in the Andaman Sea. This subsection
contains approximately 11 species, but the taxon¬
omy of the group is difficult and unclear. All spe¬
cies in the C. rumphii complex have appeared in
the literature erroneously as C. circinalis at some
stage (see Hill, 1994). A spongy layer causes seeds
to be buoyant, and this has been proposetl as a
dispersal mechanism (Dehgan & Yuen, 1983). This
dispersal mode has been interpreted as a cause of
the taxonomic complexity of this group (lor discus¬
sion, see Hill, 1994; Fosberg & Sachet, 1975).

Cycas zeylanica (J. Schuster) A. Lindstrom & K.
D. Hill, comb, et stat. nov. Basionym: Cycas
rumphii subsp. zeylanica J. Schuster, Pflan-
zenr. 99: 75, figs. 10C— D, I 1 K— M. 1932.
TV PE: Sri Lanka. 1866, Thwaites 3862 in
Herb. Barbey-Boissier (lectotype, designated
here, G; isolectotypes. A, K, LE).

Stems arborescent, to 3 m tall. Leaves bright to
deep green, semiglossy, 140—190 cm long, flat (not
keeled) in cross section (opposing pinnae inserted

Volume 12, Number 2
2002

Lindstrom & Hill
Notes on Cycas

239

at 180° on rachis), 70 to 100 leaflets, with white or
orange tomentum shedding as leaf expands; rachis
consistently terminated by paired leaflets. Petiole
50—70 cm long (30—40% of total leal), petiole gla¬
brous, spinescent for 30-100% of length. Basal leaf¬
lets not gradually reducing to spines. Median leaf¬
lets simple, strongly discolorous, 180-300 X 12-
15 mm, narrowed to 6-7 mm at base (to 40—50%
of maximum width), 22—27 mm apart on rachis; me¬
dian leaflets flat; margins slightly recurved; apex
acute, not spinescent; midrib raised above, flat be¬
low. Cataphylls linear, pungent, pilose, 100—120
mm long, persistent. Pollen cones fusiform; micro-
sporophyll lamina firm, not dorsiventrally thick¬
ened, 35—45 X 17-19 mm, apical spine prominent,
sharply upturned, 3—10 mm long. Megasporophylls
17—30 cm long; ovules 2 to 5, glabrous; lamina
lanceolate, 60—120 X 10—17 mm, obscurely den¬
tate with 6 to 12 lateral bumps or short spines to
2 mm long; apical spine distinct from lateral spines.
40—60 mm long. Seeds flattened-ovoid; sarcotesta
orange-brown; fibrous layer absent; sclerotesta
smooth; spongy layer present.

Historical notes. Collection detail cited (Schus¬
ter, 1932: 75) was “Ceylon: Thwaites I860 n. 3862
in Herbar. Barbey-Boissier.” Schuster’s herbarium
and types were in B, and were destroyed in World
War II. The Barbey-Boissier herbarium and types
are in G, including this collection. Three sheets
exist, with mierosporophylls, megasporophylls, and
leaflets, each on a separate sheet. More than one
plant is clearly involved, and it is impossible at this
stage to associate the leaf specimen with either of
the fertile specimens. These may not have been tin1
specimens examined by Schuster, as they have no
annotations. The sheet bearing the megasporophylls
is here designated the lectotype.

Distinguishing features. Cycas zeylanica is al¬
lied to both C. thouarsii Desfontaines ex Gaudi-
chaud and C. edentata de Laubenfels, sharing the
character lacking an apical wing and mierosporo¬
phylls with an apical spine. The numerous, long
and pungent cataphylls, an exceptionally long and
drawn out megasporophyll tip. and the widely
spaced leaflets distinguish C. zeylanica.

Distribution and habitat. Known from Sri Lan¬
ka, in southern coastal regions, and the Andamans
and Nicobar Island group. The habitat is in littoral
forest near the sea in sandy soil.

Reproductive biology. Surprisingly little is
known about the pollinators of Cycas subsect. Rum-
phiae. Weevils ( Curculionidae , Tychiodes) have
been found in male cones both in the Andamans
and Nicobar Islands (Tang et ah, 1999). Two male

cones collected in the South Andamans were found
full of these weevils ( Lindstrom 357, Lindstrom
353). but they are apparently of a different species
than those found on Cycas edentata on the eastern
coast of peninsular Thailand (Tang et al.. 1999).

Conservation status. SKI LANKA. Populations
have been totally decimated, and a survey done by
the first author failed to find any viable regenerat¬
ing populations. Only widely scattered, large, old
specimens remain. No plants were found within any
protected areas. The correct IliCN Red List cate¬
gory for the species on Sri Lanka should be Criti¬
cally Endangered (CR) (IUCN. 1998). INDIA. The
species is widespread and common on the Anda¬
mans and Nicobar Islands. However, although still
growing in large populations at several locations,
severe damage has been done by repeated strong
cyclones. The recent development of seaside resorts
on the islands has also had an impact on some
populations, and several previously large popula¬
tions have been eradicated to make space for tourist
accommodations. There are populations within pro¬
tected areas as well as in off-limit, military areas.
The recommended IIJCN Red List category would
be Vulnerable (VU) (IUCN, 1998).

Vernacular. Maha-madu (Sri Lanka).

Ctyinology. From Zeylona, the Latinized ren¬
dering of Ceylon (Sri Lanka), from where the type
of this species was collected.

Specimens examined. INDIA. South Andamans:
hurz s.n. (K, P); Havelock Island, E coast, beach No. 5,
2 m ASL, tall evergreen littoral forest, app. 50 m from the
beach, 2 Dec. 2000. Lindstrom 357 (NSW, SINC); cult,
near the jetty, 5 m ASL, 2 Dec. 2000. Lindstrom 359
(NSW, SING); W coast, beach No. 7, 3 m ASL, tall sec¬
ondary evergreen littoral forest, 3 Dec. 2000, Lindstrom
395 (NSW, SING); Kodiaghat, beach vegetation, sandy
soil, sea level, few plants, 25 Sep. 1973, Bulakrishnan
332 (SING); hort. Kew, raised from seed coll. Nieobars bv
Col. Nan in 1875, 1881 leg. ign. (K). SKI LANKA. Araiya
Ylawtha, Maddawattha, Mataya, wild near the seashore
along a stream, 27 m ASL, 10 June 2000. Lindstrom 287
(NSW. SING): cultivated along the road, 31 m ASL. 10
June 2000, Lindstrom 288 (NSW. SING); Hikkaduwa, De-
wadalla, Waulegnda, cult. I I June 2000. Lindstrom 289
(NSW. SING); Wtage, Alangama, cult., I I June 2000.
Lindstrom 290 (NSW. SING).

Acknowledgments. Kampon Tansaeha, owner
and Director of the Nong Nooch Tropical Garden,
Thailand, is gratefully acknowledged for financial
support, hospitality, and assistance. In India, A. K.
Singh and his family are gratefully acknowledged
for logistic and wonderful hospitality during the
Andaman field research bv the first author. The
Hermon Slade Foundation is thanked for financial
support. The keepers of the herbaria BKF, K, L,

240

No von

and I* are acknowledged for access to their collec¬
tions of Asian material.

Literature Cited

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relation to dispersal, evolution and propagation of Cy¬
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Fosberg, F. R. & M.-H. Sachet. 1975. Flora of Micronesia
pail 1, Gymnospennae. Smithsonian Contr. Bot. 20: 1-5.
Hill, K. I). 1994. Cyan rumphii in New Guinea and I lie
Western Pacific. Austral. Syst. Bot. 7: 544—567.

- . 1995. Infrageneric relationships, phylogeny and

biogeography of the genus Cycas (Cycadaceae). Pp.
139—162 in P. Vorster (editor), CYCAD 93. The 3rd
International Conference on Cycad Biology, Proceed¬
ings. Cycad Society of South Africa, Stellenbosch.

IUCN. 1998. 1997 IUCN Red List of Threatened Plants.
IUCN, Gland, Switzerland.

Laubenfels, 1). .1. de & F. Adema. 1998. A taxonomic
revision of the genera Cycas and Epicycas gen. nov. (Cy¬
cadaceae). Blumea 43: 351-400.

Linnaeus, C. 1747. Flora Zeylaniea. Stockholm.

- . 1753. Species Plantarum, Vol. I. Stockholm.

Schuster, ,1. 1932. Cycadaceae. Pp. 1-168 in A. Kngler
(editor), Das Pflanzenreich 99(4,1). W. Engelmann,
Leipzig.

Tang, W., G. R. Oberprieler & L-S. \ang. 1999. Beetles
(Coleoptera) in cones of Asian Cycas: Diversity, evolu¬
tionary patterns, and implications for Cycas taxonomy.
Pp. 280-297 in C. J. Chen (editor). Biology and Con¬
servation of Cycads: Proceedings of the Fourth Inter¬
national Conference on Cycad Biology. International
Academic Publishers, Beijing.

Trimen, II. 1898. Flora of Ceylon, vol. 4. Dulau. London.

A New Species of Holomitrium (Musci: Dicranaceae) from
South America, and a Key to Holomitrium in the Northern and
Central Andes of South America

Michelle Price

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Current
address: Conservatoire et Jardin botaniques. Case Postale 60, Cl I- 1292 Chambesy-

Geneve, Switzerland

ABSTRACT. Holomitrium azuayensis differs from
all other species of Holomitrium known from South
America, Central America, and the Caribbean by
its completely entire leaf margins. Holomitrium xol-
ocotzianum H. A. Crum from Mexico is a similar
species that has entire leaf margins below, but the
leaves are toothed at the apex. It is a much smaller
plant with shorter leaves that have a cucullate apex.
Also similar is H. pulchellum Mitten from Central
America and northern South America. It has sub¬
entire or weakly crenulate to sometimes serrate leaf
margins, but it has more slender and erect leaves
than //. azuayensis. The alar region in H. pulchel¬
lum is only weakly differentiated, and often fuga¬
cious, which also distinguishes it from H. azuay¬
ensis. This new moss species is currently known
from Ecuador.

Key words: Dicranaceae, Ecuador, Holomi¬
trium, Neotropics.

With around 30 species Holomitrium Bridel is
one ol the largest and most morphologically diverse
of the epiphytic genera in the Dicranaceae. This
moss genus is characterized by differentiated alar
regions; strong, single costae with central guide
cells; long, sheathing perichaetial leaves; cucullate
calyptrae; long-rostrate opercula; erect, cylindrical
capsules; weakly divided to entire peristome teeth
that are fenestrate and papillose; and 2 to 3 rows
ol cell development above the capsule mouth con¬
stituting a persistent annulus. The genus has been
recently treated from Mexico (Ireland, 1994), Cen¬
tral America (Allen. 1990, 1994), the Caribbean
(Hegewald, 1978). and Colombia (Churchill & bin-
ares, 1995). There are currently 15 species of Hol¬
omitrium in the Neotropics, 7 of which are known
from the northern and central Andes. In this region
of South America Holomitrium species are gener¬
ally robust plants, predominantly epiphytic, and
found at mid to high elevations. They tend to form
cushions on tree trunks and branches in montane
woodlands. During recent revisionary studies of the

genus a new species was discovered from Ecuador.
This new species. H. azuayensis, is distinct from all
the other species in this region by its completely
entire leaf margins. It also has peristome teeth that
are sometimes joined at the tips, a feature that has
not previously been recorded for Holomitrium.

Holomitrium azuayensis M. J. Price, sp. nov.
TA PE: Ecuador. Azuay: El Cajas National
Recreation Park, 30 km W of Cuenca, in Po-
lylepis woodlands on N facing slope of E-W
valley to the N of bake Toreadora, 3600-4000
m, 02°54'S, 79°17'W, 16 Sep. 1995, M. J.
Price 7 (holotype, MO; isotypes, BM, G,
QCNE). Figure 1.

A H. xolocotziano statura grandi, foliis lanceolatis et
grandioribus, marginibusque integris omnino differt.

Plants medium in size, light green-yellow above,
brown-green to brown below, glossy, growing in
tufts. Stems erect, 2.5— 5.5 cm tall, radiculose
throughout; branches irregularly bi-pinnate; (omen¬
tum dark red-orange, rhizoids hyaline at tips,
smooth; stem cross section with scleroderm, outer
cells small and thick-walled, inner cells large and
thin-walled, central strand small; axillary hairs 2 to
4 per axil, 6 to 8 cells long, cells elongate, hyaline
throughout, beaves moderately spaced, erect at
base, crisped above, secund or somewhat erect
when dry, secund to erect-spreading when wet, lan¬
ceolate, 3-6 X 0.5— 1.1 mm, tubulose throughout,
unistratose; base short -decurrent; apex gradually
acuminate; margins plane belowT, broadly incurved
above, sometimes slightly undulate, entire through¬
out; costae single, slender, strong, subpercurrent,
smooth at back, in cross section at mid-leaf with 4
central guide cells, dense dorsal and ventral stereid
bands; median leaf cells below shoulder rectangu¬
lar to oblate, incrassate, not to weakly porose, 15—
53 X 7-10 gm, becoming longer toward costae;
cells above shoulder rounded, quadrate, oblate or
sometimes short-rectangular, incrassate, not porose.

Novon 12: 241-244. 2002.

242

No von

Figure ]. Holomitrium azuayensis M. J. Price. — A. Plants wet and dry. — B. Leaf. — C. Stem cross section. — I).
Peristome teeth (external surface). — E. Cells at leal shoulder. — F. Basal leal cells. — C. Alar cells. — II. Cells al leaf
apex. — l. I.eal margin and upper leal cells. All from Price 7 (MO).

Volume 12, Number 2
2002

Price 243

Holomitrium azuayensis from South America

12-27 X 7—13 /am, becoming smaller and quadrate
toward apex; basal cells long-rectangular to rect¬
angular, incrassate, porose, 32-113 X 13-20 /cm;
alar cells distinct, reddish orange, rectangular to
subrectangular, 25—68 X 22—38 /cm. persistent, in¬
crassate, non-porose; marginal cells undifferenti¬
ated. Asexual reproduction by fragile flagellate
branches, 5—12 mm long, clustered at stem apex or
in branch axils, leaves to 1 mm long, costae sub-
percurrent. Pseudautoicous. Perigonia on dwarf
males, growing on stem tomentum, in leaf axils or
perichaetial bracts of female plants, plants 1-7 mm
long, often with two antheridial bracts, leaves lan¬
ceolate, 1.5— 1.7 mm long. Perichaetia terminal be¬
coming lateral by subperichaetial branching, outer
leaves erect below, erect to erect-spreading above,
lanceolate to linear-lanceolate; inner perichaetial
leaves long-sheathing, linear-lanceolate, 6—12 mm
long; margins entire throughout, sometimes weakly
crenulate at apex; costae subpercurrent; cells sim¬
ilar to vegetative leaves. Setae single, light yellow,
erect, 20—25 mm long, twisted counterclockwise
just below capsule, smooth. Capsules long-exserted,
erect; urn long-cylindrical, symmetric, 2.0— 3.5 mm
long, to 1 mm diam., smooth; neck short; mouth
slightly constricted; exothecial cells irregularly
rectangular, thin-walled, yellow tinged with several
rows of small red, incrassate cells at mouth. Sto¬
mata 6 to 10 per capsule, superficial, at base of
capsule. Annuli persistent, as 2 to 3 rows of cells
developed above the capsule mouth, cells homo¬
geneous. quadrate to triangular. Opercula long-ros¬
trate, 2.5 mm long. Peristome teeth 16, irregular,
inserted at mouth, weakly divided or entire, teeth
sometimes united at tips by inner peristomial layer
deposition, fenestrate, irregularly thickened on out¬
er surface, dark orange-red at base, fragile and hy¬
aline above, narrowly triangular, 137—175 /an long
without hyaline tips, when hyaline tips present
237—250 /irn long, finely papillose below, papillose
above. Calyptrae cucuilate, smooth, 3—4 mm long,
base entire. Spores spherical. 10-13 /an diam..
lightly roughened.

Etymology. The specific epithet azuayensis was
derived from the province Azuay, in Ecuador.

Distribution. Known from Ecuador.

Habitat. On trunks and branches of Polylepis
in montane woodlands, growing as cushions mixed
with Chorisodontium between 3600 and 4200 m.

Holomitrium azuayensis is a medium-sized plant,
growing to 5.5 cm in height, with lanceolate leaves
that are somewhat secund, moderately spaced, and
crisped when dry. The leaf margins in this species
are broadly incurved above, and completely entire

throughout. Its alar cells are well developed and
persistent, forming a distinct colored group at the
basal angles of the leaf. The median cells near the
leaf shoulders are rectangular to oblate, incrassate,
and not to weakly porose. Its upper leaf cells are
small, rounded, and quadrate to oblate or some¬
times short -rectangular. These cells are incrassate,
non-porose, becoming smaller and more regularly
quadrate toward the leaf apex. The peristome mor¬
phology of this species is unusual within Holomi¬
trium. A peristome tooth in //. azuayensis is split
from one-third to one-half of its length, forming two
hyaline filaments above. Some of these filaments
are joined at the tips, to an adjacent tooth, by their
ventral (inner) surfaces. So far, this feature has
been observed in only a couple of other species of
Holomitrium (H. flexuosum Mitten and //. olfersian-
um Hornschueh), and has not previously been doc¬
umented for the genus. This peristome morphology
is more typical for the genus Dicnemon Schwagrich-
en, where it is known in all species except one.
namely D. calycinum (Hooker) Schwagrichen (Al¬
len, 1987).

In overall aspect this newr species most closely
resembles the Mexican species //. xoloeotzianum,
described by Crum (1952). Both have secund, lan¬
ceolate leaves, similar upper leaf cell areolation,
subpercurrent costae, and flagellate branches.
However, H. xoloeotzianum is a much smaller plant,
growing only to 2.5 cm in height. It also has shorter,
more sheathing leaves with a more pronounced
shoulder than the new species. Leaf cells at the leaf
shoulder in H. xoloeotzianum are more uniform in
shape, and are smaller than those in H. azuayensis.
The cucuilate leaf apices and apical teeth of //.
xoloeotzianum also distinguish this species from II.
azuayensis. Capsule and peristome morphology of
these two species differ. In H. xoloeotzianum the
persistent annulus consists of 6 to 8 rows of cells,
compared with 2 to 3 rows of cells in H. azuayensis.
I’lie peristome teeth are densely papillose through¬
out and are not joined at their t i ps in //. xoloeot¬
zianum.

Holomitrium pulchellum and the closely related
species H. hawkinsii B. 11. Allen, from Central
America (Allen, 1997), are similar to //. azuayensis
in their leaf cell areolation, and in having nearly
entire leaf margins. In //. pulchellum and H. hawk¬
insii leaf margins can vary from crenulate to ser¬
rulate or almost entire and toothed only at the very
apex; they are never completely entire. Holomi¬
trium azuayensis is distinguished from these two
species by its lanceolate leaves with a completely
entire leal apex; well-developed, persistent alar re¬
gions; subpercurrent costae; and the presence of

244

Novon

flagellate branches. The Central American species
//. williamsii E. B. Hart ram shares with II. azuay-
ensis the presence of flagellate branches, but it lias
serrate, irregularly bistratose leaf margins. All other
species from the northern and central Andes are
distinct from //. aziiayensis by their serrulate to ser¬
rate or dentate leaf margins, while some others dif¬
fer by their elongate cells throughout the lamina.

When H. azuayensis is found fertile the rather
striking long, sheathing perichaetial leaves and
crisped stem leaves when dry confirm that it be¬
longs to Holomitrium. Two other genera in the l)i-
cranaceae diat are found in this region. Eucamp¬
todontopsis Brotherus and Schliephackea Carl
Miiller, also have long and sheathing perichaetial
leaves. Schliephackea is a pendent moss, with stems
to 30 cm in length, and it has wide-spreading some¬
what distant leaves that distinguish it immediately
from H. azuayensis. Eucamptodontopsis is predom¬
inantly a Central American-Caribbean-Guyana
Shield genus. It contains species with subentire,
cretin late or serrate leaf margins, but the genus is
characterized by leaves that are weakly spirally
twisted or contorted when dry, and elongate, rect¬
angular leaf cells throughout. The new species H.
azuayensis can be distinguished from members of
Eucamptodontopsis by its entire leaf margins, and
by its small, rounded, and quadrate to oblate or
sometimes short-rectangular upper leaf cells. When
the new species is lacking perichaetia, the crisped
leaves when dry. the differentiated and colored alar
region, and differentiated lower and upper leaf cells
(basal cells rectangular and upper leaf cells small
and rounded to quadrate or oblate) can be used to
identify it as belonging to Holomitrium.

Paratypes. ECUADOR. Imbabura: on old trail to In¬
lag. 1 1,000 ft., 24 Nov. 1943, IE C. Steere 9224 (MO.
NY).

Kky to ri t k Species ok Holomitrium in the Northern

and Central Andean Region of South America

la. Leaf margins entire throughout .... H. azuayensis
11>. Leaf margins subentire, crenulate, serrate, ser¬
rulate, or toothed above leaf shoulder . 2

2a. Cell walls of upper leaf porose or sinuose

. 3

2b. Cell walls of upper leaf straight-walled ... 4

3a. Cell walls strongly sinuose from just above alar

region . H. sinuosum

3b. Cell walls straight to leaf shoulder, mostly weakly

sinuose above . H. flexuosum

4a. Leaf margins and/or lamina irregularly bis¬
tratose . 5

4b. Leaf unistratose throughout . 6

5a. Leaves bistratose at margins and in patches

across lamina . H. crispulum

5b. Leaves bistratose only at margins ...//. arboreum

6a. Leaf margins subentire, crenulate or serru¬
late, serrate at apex . II. pulchellum

6b. Leaf margins serrate or dentate . 7

7a. Leaves over 9 mm long . //. moritzianum

7h. Leaves under 9 mm long . //. arboreum

Acknowledgments. This work was supported by
National Science Foundation, grant 9522034
(PEET: Partnerships for Enhancing Expertise in
Taxonomy) to the Missouri Botanical Garden, and
by an award from The Botanical Research Fund,
U.K. I thank all the individuals that have helped
in my efforts to complete this work, including Bruce
Allen, Robert Magill. Bill Buck, Victoria Hollowell,
and an anonymous reviewer for comments on ear¬
lier drafts of this manuscript. I thank Bruce Allen
and Robert Magill for their help and encourage¬
ment.

Literature Cited

Allen. B. H. 1987. A revision of the Dicnemonaceae
(Musci). J. Hattori Bot. Lab 62: 1—100.

- . 1990. A preliminary treatment of the Holomi-

/rmm-complex (Musci: Dicranaceae) in Central Ameri¬
ca. Trop. Bryol. 3: 59-71.

- . 1994. Moss Flora of Central America Part 1.

Sphagnaceae— Calymperaceae. Monogr. Syst. Bot. Mis¬
souri Bot. Card. 49.

- . 1997. Holomitrium hawkinsii (Musci: Dicrana¬
ceae), a new species from Central America. Novon 7:
329-332.

Churchill, S. P. & K. L. Linares C. 1995. Prodromous
Brvologiae Novo-Granatensis. Introduceion a la flora de
rnusgos de Colombia. Part I. Bibliot. Josd Jerdnimo Tri-
ana 12: 1-454.

Crum, H. A. 1952. Two new species of Mexican mosses
collected by A. J. Sharp. Bryologist 55: 51-58.
Hegewald, E. H. 1978. Critical notes on Holomitrium (Di¬
cranaceae) from the Antilles. Bryologist 81: 524—531.
Ireland. If. 1994. Holomitrium. Pp. 156—158 in A. J.
Sharp, H. Crum <!4 P. M. Eckel (editors). The Moss flora
of Mexico. Mem. New York Bot. Card. 69.

Lycianthes manantlanensis (Solanaceae), a New Species from

Mexico

Aaron Rodriguez and Ofelia Vargas

Departamento de Botaniea y Zoologia, Universidad de Guadalajara, Apartado Postal 1-139,

45101 Zapopan, Jalisco, Mexico

ABSTRACT. Lycianthes manantlanensis is de¬
scribed as a new species from Mexico. Lycianthes
manantlanensis is recognized by its glabrous, co¬
riaceous, and shiny leaves, long slender pedicels at
anthesis and in fruit, and minute calyx appendages.
It is morphologically similar to L. barbatula, L.
geminifiora, and L. orogenes and is here compared
with them.

Resumen. Se describe e ilustra a Lycianthes ma¬
nantlanensis como una especie nueva que crece en
Mexico. Lycianthes manantlanensis se reconoce por
sus hojas glabras, coriaeeas y brillosas; pedicelos
largos y delgados en la antesis y fructificaeidn y
caliz con pequenos apendiees. El taxon propuesto
es morfologicamente similar a L. barbatula, L. ge¬
minifiora y L. orogenes con las cuales se compara.

Key words: Jalisco, Lycianthes , Mexico, Sola¬
naceae.

An inventory of the family Solanaceae for the
flora of Jalisco, in western Mexico, is under way
(Rodriguez & Vargas, 1994; Vargas & Rodriguez,
1993, 1995). In the process, we found some mor¬
phologically unfamiliar specimens that we describe
as a new species.

Lycianthes manantlanensis A. Rodriguez & ().
Vargas, sp. nov. TYPE: Mexico. Jalisco: Mpio.
Cuautitlan de Garcia Barragan, Majada de las
Avellanas, comunidad indfgena de Guzalapa,
3—4 km al NNW de El Durazno, 800—1000 m,
6 Nov. 1995 (fl), R. Cuevas, E. Sanchez & F.
./. Santana 5009 (holotype, I RUG: isotypes,
ENCB, MEXU, MO, WIS, ZEA). Figure 1.

Frutex vel arbor parva 1—6 m alta, caulibus teretibus
glabris; sympodia bifoliata geminata anisophylla; lamina
foliorum maximorum coriacea glabra obovata vel elliptica
4.2-12 cm longa 2.4-^t.2 cm lata, apice acuto vel acu¬
minate, basi oblique cuneata vel cuneata interdum decu-
rrenti, petiolo gracili glabro 0.7-2 cm longo; folia minora
coriacea, glabra, obovata vel elliptica 2.5-6 cm longa 1.3—
2.7 cm lata, apice acuto vel acuminate, basi oblique cu¬
neata vel cuneata interdum decurrenti, petiolo grac ili gla¬
bro 2—1 mm longo; inflorescentia axillaris fasciculata, 3—
5 floribus; gemmae globosae vel ovoideae, 2-4 mm latae;

pedicellus sub anthesi gracillimus deflexus vel ereetus 2—
3.5 cm longus glaber; calyx campanulatus 2 mm longus
glaber tmneatus prope apicem 5 minutis appendicibus <
0.5 mm longis instructus; corolla late campanulata alba
1.2-2 cm diametro, lobis sub anthesi reflexis; stamina su-
baequalia, filamentis glabris 0.5-1 mm longis, antheris
2.5—3 mm longis oblongis; ovarium glabrum; stylus glaber
6-8 mm longus; stigma clavatum; bacca globosa, viridis 5
mm diametro; pedicellus fruetifer lignosus ereetus 2—3.5
cm longus.

Shrub or small tree, 1-6 m tall: stems terete,
glabrous. Sympodia bifoliate, geminate, the leaf
pair markedly anisophyllous. Branches glabrous or
with some simple uniseriate trichomes less than 0.5
mm long. Leaves coriaceous, flat, glabrous and
shiny on both sides; major leaves 4.2—12 X 2.4-
4.2 cm; lamina obovate to elliptic, the apex acute
or acuminate, the base euneate, sometimes oblique;
petioles 0.7-2 cm long; minor leaves not differing
from the major ones in shape, 2.5-6 X 1.3— 2.7 cm;
petioles 2-4 mm long. Inflorescences axillarv. fas¬
ciculate, 3— 5-flowered. Pedicels slender, glabrous,
erect or deflexed at anthesis, 2-3.5 cm long. Buds
globose to ovoid, 2-4 mm wide. Calyx campanulate,
truncate, glabrous, 2 mm long, with five minute
knob-like appendages below the erose margin, less
than 0.5 mm long. Corolla white, campanulate, 1 .2—
2 cm diam., the lobes minutely pilose at the tips
and along margins. Stamens subequal in length; fil¬
aments 0.5—1 mm long, glabrous; anthers 2.5—3
mm long. Ovary glabrous. Style glabrous, 6-8 mm
long. Stigma clavale. Fruit globose, green, 5 mm
diam.

Distribution, lycianthes manantlanensis has
been collected in the Sierra de Cacoma, Jalisco, at
the western end of the Trans-Mexican Volcanic
Belt. It is also known from the Sierras de Manantlan
and Coalcoman in the Sierra Madre del Sur, as well
as the states of Oaxaca and Chiapas in southern
Mexico.

Etymology. The name of the species refers to
the Sierra de Manantlan, where this species is com¬
mon.

Habitat. Lycianthes manantlanensis grows in
the transition zone between the tropical subdeci-

Novon 12: 245-248. 2002.

246

Novon

— I). Detail of pedicel and calyx (/». Cuevas el al. 5009).

(luous forest and the cloud forest. It grows in as¬
sociation with Ardisia revoluta Kunth. Inga eriocar-
pa Bentham, Magnolia iltisiana A. Vazquez,
Populus giizmanantlensis A. Vazquez & Cuevas,
Quercus insign is M. Martens & Caleotti, and Si-
paruna andina (Tulasne) A. DC., 800—1150 in. Il

also grows in the cloud forest where it is associated
w ith Baltnea stormae Martinez, Beilschmiedia pen-
dida (Swartz) Hemsley, Calatola laevigata Standley,
Capparis mollicella Standley, Cedrela odorala I ...
Cinnamomum pachypodum (Nees) Kostermans,
Coccoloba barbadensis Jacquin, Dendropanax at-

Volume 1
2002

, Number 2

Rodriguez & Vargas 247

Lycianthes manantlanensis from Mexico

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248

Novon

boreus (L.) Decaisne & Planchon, Guarea glabra
Vali I . Juglans major (Torrey) A. Heller, Licaria
triandra (Swartz) Kostermans. Malvaviscus arboreus
Cavanilles, Nectandra glabrescens Bentham, Pi¬
er as ma mexicana Brandegee, Pranas serotina Ehr-
liart subsp. capuli (Cavanilles) McVaugh, Robinso-
nella speciosa Fryxell, Triehillia americana (Sesse
& Moyino) Pennington, Trophis racemosa (L.) Ur¬
ban, and Urera earacasana (Jaequin) Grisebach.
1450—1650 m. Occasionally, it has been collected
in Abies forest, ea. 2000 m elevation.

Common name. Naranjillo.

Discussion. Lycianthes manantlanensis is rec¬
ognized by its glabrous, coriaceous, and shiny
leaves, long slender pedicels at anthesis and fruit,
and minute knob-like calyx appendages. The calyx
appendages are small and difficult to distinguish,
but they emerge from a little below the erose mar¬
gin. The calyx then splits irregularly and forms un¬
equal lobes with eorky-searred margins. Lycianllies
manantlanensis shares its coriaceous leaves with L.
barbatula Standley & Steyermark and L. orogenes
Standley & Steyermark. However, it differs from
them by lacking conspicuous calyx appendages and
other aspects seen in Table 1. Uikewise, L. ma¬
nantlanensis resembles L. geminiflora (M. Martens

Galeotti) Bitter in shape and size of the calyx.
I'he pedicel length, however, is always longer in L.
manantlanensis. In addition, the leaves in L. ge¬
miniflora are puberulent, not coriaceous and asym¬
metric.

Paratypes. MEXICO. Jalisco: Ayulla, near Santa Mo¬
nica, ea. 20°N, I04°30'W, 1950-2050 m, 12-15 Nov.

1952 (fl), R. McVaugh A: ./. Sooby, Jr. 14060 (MICH); Ca-
simiro Castillo, por la Canada El Tecolote, 19°36'49"N,
1 04° 1 8' 56" W, 1650 m, 27 Oct. 1998 (fl), R. Cuevas. L.
Guzman & ./. Aragon 6246 (/EA), 1550 m, 29 Nov. 1998
((1). R. Cuevas. I,. Guzman & ./. Aragon 6274 (ZEA), 1550
m. 30 Nov. 1998 (11), R. Cuevas, /.. Guzman & ./. Aragon
6 294 (/EA), 1450 m, 13 Dec. 1998 (sterile), R. Cuevas.
L Guzman & ,/. Aragon 6 .'180 (ZEA), 1450 in. 14 Dec.
1998 (fl). R. Cuevas, /.. Guzman & ./. Aragon 6290 (ZEA),
14 Dec. 1998 (sterile), R. Cuevas, L. Guzman & ./. Aragon
6400 (ZEA); en la Canada de La Naranjera, cerca del
arbol de aguacate, 19°37'25"N. I04°20'49"W. 1 150 m. 22
Eeb. 1999 (fr), R. Cuevas, L. Guzman & ./. Aragon 6554
(ZEA), 1 150 m, 24 Feb. 1999 (sterile), R. Cuevas, b. Guz-
mdn & ./. Aragon 6 589 (ZEA). Michoacan: Distrito de
Coalcoman, I KK) m. G. R. Hinton et al. 15942 (MICH).
Oaxaca: Villacorzo, Paraje Ea Botella, ejido Plan de Aya¬
la, 1420 m, 7 Jan. 1999 (fr), R. Ra m irez- Ca m acho 41
(CHAP, MEXD). Chiapas: Monte Ovando, 14—18 Nov.
1939 (II). K. Manilla 398! (MEXU. NY); Santa Marfa Chi-
malapa, cerro Salomon, ca. 2 km WNW del cerro Gua-
yabitos, al N de la cima, ca. 43 km en lfnea recta al N
de San Pedro Tapanatepee, I6°45'N, 94°ll'W, 27 Oct.
1986 (fr), S. Maya 4129 (CHAPA. MEXU).

Acknowledgments. We thank Refugio Vazquez
Velasco for the illustration and the staff at ZEA and
MEXU for specimen loans. We also thank Michael
Nee and Roberto Gonzalez Tamayo for their com¬
ments on the original manuscript.

Literature Cited

Rodriguez. A. & (). Vargas. 1994. Las especies de papa
silvestre (Solarium L. seecidn Petal a Dumortier) en Ja¬
lisco. Bob Inst. Hot. 2: 1—68.

Vargas, 0. & A. Rodriguez. 1993. La seecidn Solarium del
genero Solarium en Jalisco. Bol. Inst. Rot. 1: 423—439.

- & - . 1995. Nuevos registros de Solarium

(Solanaceae) para el estado de Jalisco. Bol. Inst. Bot.
3: 27-40.

A New Species of Weinmannia (Cunoniaceae: Cunonieae) from

Southern Ecuador

Zachary S. Rogers

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

zachary.rogers@mobot.org

ABSTRACT. Weinmannia auriformis is described
and illustrated from the Cordillera del Condor re¬
gion, Ecuador. It is distinguished by its small, au¬
riform, revolute, entire-margined leaflets and by the
dense tomentose pubescence on its branchlets,
leaves, stipules, inflorescences, and calyx lobes.
This new endemic species is only known from one
sandstone mountain of the Cordillera de Huaraca-
yo, east of the Cordillera del Condor.

Resum EN. Se describe y se ilustra Weinmannia
auriformis de la region de la Cordillera de Condor.
Ecuador. Esta especie se distingue por sus folfolos
pequenos que se asemejan a unas orejas con mar-
genes enteros y un hirsute dense, pubescente, en
sus ramas pequenas, sus hojas, sus estipulas, sus
inflorescencias y los lobulos del caliz. Esta nueva
especie se encuentra en un cerro formado por roea
arenisca en la Cordillera de Huaracayo al este de
la Cordillera del Condor.

Key words: Cunoniaceae, Cunonieae, Ecuador,
Neotropics, South America, Weinmannia.

The family Cunoniaceae is composed of 26 gen¬
era and about 300 species (Bradford & Barnes,
2001). The widespread genus Weinmannia L. is
made up of approximately 130 species of trees and
shrubs, more than half of which are found in the
tropics (Bradford & Barnes, 2001). In the Neotrop¬
ics, the genus ranges from southern Mexico to
southern Chile (Harling, 1999). Ecuadorian Wein¬
mannia species have leaves that are opposite, de¬
cussate, and simple or imparipinnately compound
(all Ecuadorian species with compound leaves have
winged rachises except for W. trianaea Weddell),
and interpetiolar stipules (Marling, 1999). All
Ecuadorian species of the genus have stereotypical
flowers that are small, white, bisexual, and 4-me-
rous, borne on unbranched spicate or elongate ra¬
cemose inflorescences (i.e., pseudoracemes). Wein¬
mannia species also have distinctive fruits that are
small, septicidal, 2-valved capsules that have 2
persistent stylar beaks, and lobed nectary disks lo¬
cated beneath the gynoeeium (Harling, 1999). liar-
ling’s (1999) treatment of Cunoniaceae for the Flora

of Ecuador included 26 species; however, an ad¬
ditional undescribed species has been collected
from the Cordillera del Condor region near the
Peru— Ecuador border. A color photograph of the
type collection can be found on the searchable W3
TROPIC OS database at: (http://mobot.mobot.org/
W3T/search/ vast . h tml) .

Weinmannia auriformis Z. Rogers, sp. nov.
TYPE: Ecuador. Morona Santiago: Canton Pi¬
nion Indanza, Cordillera del Huaracayo, E of
Cordillera del Condor and Rio Coangos, Cerro
Ijiach Naint, flat-topped sandstone mountain,
E of Shuar village of Tinkimints, 3°I5'49”S,
78°10'13"W, 2000 m, 21 Mar. 2001, I). Neill,
P. Berry, ./. Manzanares & L. Jost 13112 (ho-
lotype, QCNE; isotypes, GB. M 0-5300000,
NY, QCA). Figure 1.

Haec species W. loxensi et W. mariquilae maxime si-
milis, sed a hac ramulis foliis inflorescentia calyceque in-
dumento tomentoso denso vestitis, foliolis ] .3—6.6 X 1 .0—
5.0 mm, margine semper integris praeserlim apiee revo-
lutioribus, pseudoracemo eompaeto 1.1— 3.1 cm tantum
longo atque florum minorum lobulis calyeinis 0.8—1. 2 X
0.4-0. 6 mm, ab ilia foliolis et rhachide foliari valde con-
cavis distinguitur.

Shrubs or trees 0.5—4 m. Branchlets slender, te¬
rete, red-black, tomentose, becoming less dense
with age, denser at nodes, the trichomes simple,
unicellular to 1.0 mm long. Leaves imparipinnate,
1 .3-3.3 X 0.3-0.9 cm; leaflets 5- or 6-paired (rare¬
ly 4- or 7-paired), 1. 3-6.6 X 1. 0-5.0 mm, auriform
to cochleariform, strongly concave, coriaceous,
waxy and shiny, apex slightly apiculate, base ob¬
tuse, margin entire, strongly revolute, especially at
apex, upper surface scrobiculate and rugose,
sparsely tomentose, denser along lower half of mid¬
vein, lower surface smooth, tomentose along mid¬
vein, denser along lower half, the trichomes simple,
unicellular to 1.0 mm long, midvein slightly im¬
pressed above, slightly raised below, secondary
veins inconspicuous above and below, all leaflets
similar in size and shape; rachis winged, tomentose
above, denser below, especially pubescent at point
of petiolule attachment, trichomes similar to those

Novon 12: 249-252. 2002.

250

No von

Figure I. Weinmannia uuriformis Z. Rogers. — A. lirariehlet with pseudoracemes. — B. Abaxial surface of compound
leaf. — C. Adaxial surface ol leaflet. — I). Flower bud showing the corolla cap separation. — E. Mature flower after the
corolla has separated. — F. Mature capsule with persistent nectary disk and styles. — G. Seed. All illustrations drawn
by Zachary Rogers from the MO isotype ( Neill el al. 13112. MO-5300000).

Volume 12, Number 2
2002

Rogers

Weinmannia auriformis from Ecuador

251

of the leaflets, rachis wings 2.4— 5.0 X 0.8— 1.7 mm,
elliptical to obovate, concave, strongly revolute, co¬
riaceous, often obscured by dense pubescence; pet¬
ioles terete, 2. 5-4.5 mm long, tomentose to slightly
hirsute; stipules interpetiolar, free, caducous, 2.3—
4.7 X 2. 1^4.5 mm, broadly ovate, membranaceous
to subcoriaceous, red-tinged, apex rounded, base
truncate, margin entire, outside surface densely to¬
mentose except along margin, inside surface gla¬
brous and rugose, the triehomes similar to those of
the leaflets, veins inconspicuous. Pseudoracemes
usually in pairs, 1.1— 3.1 cm long; rachis densely
tomentose, triehomes simple, unicellular to 0.8 mm
long; fascicles densely congested, 2- or 3- to 4-
flowered; hracteoles 0.5— 1.1 X 0.3— 0.8 mm, nar¬
rowly oblong, membranaceous, apex acute, base
truncate, margin entire, completely glabrous, usu¬
ally persistent; pedicels 0.5— 1.2 mm long, densely
tomentose, triehomes similar to those ol the inflo¬
rescence. blowers 4-merous, actinomorphic, bisex¬
ual, diplostemonous; calyx lobes 4, 0.8— 1.2 X 0.4—
0.6 mm, narrowly ovate, membranaceous, red- or
green-tinged, apex acute, base truncate, margin en¬
tire, outside surface tomentose, denser along lower
half of midvein, inside surface glabrous, triehomes
simple, unicellular to 0.5 mm long; petals 4. free,
caducous, often falling as a unit before completely
open, 1.1 -1.7 X 0.7— 1.4 nun. broadly obovate to
broadly elliptic, membranaceous, white, apex
rounded, sessile, margin entire, ciliate, midvein
conspicuous; stamens 8. 2.4— 4.0 mm long; fila¬
ments free, slender, flattened, 1.5— 3.8 X 0.2— 0.4
mm, glabrous; anthers introrse, 0.2— 0.5 X 0.2— 0.4
mm. orbicular, apex short acuminate, base slightly
cordate, longitudinally dehiscent, versatile; nectary
disk annular, consisting of 8 concrescent lobes,
persistent in fruit; ovary biearpellate, synearpous,
0.8-1.4 X 0.5— 0.8 mm, red-tinged, glabrous except
for filiate hairs where the styles meet; styles 2,
divergent, 1.4— 1.5 mm long; stigmas simple, capi¬
tate, persistent in fruit; ovules 4 to 6 per locule.
Fruits septieidal capsules, 3.9— 5.5 X 1.7-2. 2 mm
(length measurement includes persistent styles),
ovoid-oblong, red- or brown-tinged, eostate; seeds
0.9— 1.1 X 0.5— 0.6 mm, widely elliptic to elliptic,
longitudinally ribbed, testae with few triehomes,
simple, unicellular, 0.3— 0.5 mm long.

Distribution and habitat. Weinmannia aurifor¬
mis is an endemic species known only from the
summit of Cerro Ijiach Naint, which is an isolated
steep-sided sandstone mountain with a summit
around 2000 m. The Cerro is part of the Cordillera
de Huaracayo mountain range to the east of the
Cordillera del Condor range. The locality is covered

by a dense windswept “dwarf” forest, characterized
by a low tree canopy layer to 4 m and a dense,
scrubby, shrub layer to 2 m (I). Neill, pers. comm.
2001).

Affinities. According to the key in the treatment
of the Cunoniaceae of Ecuador (Hailing, 1999),
Weinmannia auriformis would fit somewhere be¬
tween W. mariquitae Szyszytowicz and W. loxensis
Hailing, but the new species more closely resem¬
bles W. mariquitae. All three species have impari-
pinnate leaves shorter than 5 cm in length with 4—
8 leaflet pairs per leaf, and all three have elongated
pseudoracemes. However, W. auriformis is easily
distinguishable from W. mariquitae by its distinc¬
tive leaflets, which are ear-shaped, small (1.3— 6.6
X 1.0— 5.0 nun), slightly apiculate at the apices,
revolute, entire, lacking prominent secondary ve¬
nation, and adaxially pubescent. In contrast, W.
mariquitae has larger, more elongated leaflets (0.9-
2.0 X 0.5— 1.2 cm) that are adaxially glabrous with
crenate-dentate margins and prominent secondary
venation.

Other important distinctions are that W. aurifor¬
mis has much shorter (i.e., more densely compact¬
ed) pseudoracemes (1. 1-3.1 cm), than IE mariqui¬
tae, which has 3.5—12 cm long pseudoracemes; IE
auriformis has smaller (0.8— 1.2 X 0/4— 0.6 mm) pu¬
bescent calyx lobes, IE mariquitae has larger gla¬
brous lobes (1.5-1. 8 X 0.9— 1.0 mm); and IE au¬
riformis has pubescence on the rachis wings,
whereas the wings of IE mariquitae are glabrous.

On the other hand, IE auriformis is clearly dis¬
tinct from IE loxensis because the leaflets of IE lo¬
xensis are smaller (0.2— 0.4 X 0.15—0.2 cm), flat,
glabrous, and tridentate along the margin. Wein¬
mannia loxensis also has flat rachis wings and
smaller capsules that only reach 2.5 mm in length.

Ecology. This species, like many others that
are evidently endemic to the Cordillera del Condor
region, appears to be edaphieally restricted to the
nutrient-poor sandstone table-top mountains that
characterize the area. The new species has been
found on only one such summit so far, about I km2
in area, but further floristic exploration of the area
may reveal additional localities. The area is not
protected formally, and potential threats from min¬
ing and other activities could affect this and many
other locally endemic species. The entire region
merits conservation actions, which should include
participation from the indigenous Shuar who inhab¬
it the valleys and lower slopes of the mountains
(David Neill, pers. comm.).

Etymology. The specific epithet refers to the
ear-like shape of the small leaflets of the species.

252

Novon

Paratypes. ECUADOR. Morona Santiago: Canton
Limdn Indanza, Cordillera de Huaracayo, E of Cordillera
del Cdndor and Rfo Coangos, Cerro Ijiach Naint, Hal-
topped sandstone mountain. If of Shuar village of Tinki-
mints, ridge below W side of summit, 03a15'49"S,
78Q10'28"W, 1950 m, 20 Mar. 2001. I). Neill, P Perry, ,/.
Manzanares & L .lost 13083 (AAU, EOJA, M0-5290000.
QCNE, US).

Acknowledgments. The author thanks David
Neill (MO, QCNE) for calling attention to this new
species and for many helpful comments on the
manuscript; the author also thanks tin* following
people at (MO): Jon Kicketson, George Schatz, Ge-

rrit Davidse, Jason Bradford, and Ron Liesner. Spe¬
cial thanks go to Alina Freire-Fierro (QCNE) and
one anonymous reviewer for helpful comments and
suggestions on the manuscript. Roy Gereau (MO)
prepared the Latin diagnosis.

Literature Cited

Bradford. J. C. & R. W. Barnes. 2001. Phylogenetics and
classification of Cunoniaceae (Oxalidales) using chlo-
roplast DNA sequences and morphology. Syst. Bot. 26:
354-385.

Hailing, G. 1999. Cunoniaceae. In: G. Hailing E. An-
dersson (editors), FI. Ecuador 61: 1-74.

Nomenclatural Changes in North American Lilium (Liliaeeae)

Mark W. Skinner

USDA-NRCS. [National Plant Data Center, P.0. Box 74490, Baton Rouge,
Louisiana 70874, U.S.A.
mskinner@po.nres.usda.gov

ABSTRACT. The approaching release of the Flora
of North America Volume 26 requires the formal
publication of several nomenclatural changes that
will appear there. The proposed changes standard¬
ize infraspecific nomenclature in North American
Lilium at the subspecies level, which is appropriate
for these geographically segregated yet morpho¬
metrically overlapping and intergrading taxonomic
units. The changes include a new: rank for Lilium
washinglonianum Kellogg subsp. purpurascens
(Steam) M . W. Skinner, and several new combina¬
tions: L. pardalinum Kellogg subsp. pitkinen.se (Be¬
ane & Vollmer) VI. W. Skinner, L. pardalinum Kel¬
logg subsp. shasten.se (Eastwood) M. W. Skinner, L.
pardalinum Kellogg subsp. vollmeri (Eastwood) M.
W. Skinner, and L. pardalinum Kellogg subsp. wig-
gins ii (Beane & Vollmer) M. W. Skinner. Eectotypes
are designated for L. pardalinum and L. washing-
tonianum and for the synonym L. parvurn Kellogg
var. luteum Purdy, and a neotype is selected for L.
washinglonianum subsp. purpurascens.

Key words: Liliaeeae, Lilium. North America.

Ongoing biosystematic investigations of North
American Lilium (Skinner, 1988. 1.993) compel the
publication of several new combinations that will
appear in the Flora of North America. The taxa
recognized and named below display significant
and geographically coherent variation that is best
expressed at the subspecies level (Thorne, 1978):
morphological dimensions overlap between geo¬
graphically adjacent taxa, variation is roughly clin-
al across zones of intergradation, and reproductive
isolation between subspecies is primarily achieved
geographically. The new names achieve uniformity
at the subspecific rank for infraspecific nomencla¬
ture in North American Lilium. Types are supplied
for one synonym, and for accepted names that cur¬
rently lack them. Because prior workers often ig¬
nored or were unfamiliar with each other’s work
and/or the formalities of type designation, and fre¬
quently misapplied names with or without attribu¬
tion to prior circumscriptions, the nomenclatural
literature and history in western North American

Lilium is abstruse. This prompts the olten lengthy
discussions below.

I, ilium pardalinum Kellogg. Hesperian 3: 300.
1859. TYPE: Illustration. Hesperian 3(1) fron¬
tispiece, 1859 (lectotype, designated here.
University of California at San Diego).

Lilium pardalinum var. angustifolium Kellogg. Hesperian
3: 300. 1859. TYPE: not designated.

Lilium roezlii Regel, Gartenflora 19: 321. 1. 667. 1870. as
“6. roezli non L. roezlii Purdy, J. Int. Gard. Club
3(4): 522. 1919. TYPE: not designated.

Lilium harrisianum Beane & Vollmer. Contr. Dudley Herb.
4: 357. 1955. TYPE: not designated.

A. M. Kellogg (1859a) did not formally designate
type material for Lilium pardalinum. At the Cali¬
fornia Academy of Sciences wThere Kellogg worked,
most of the types were saved from the fire following
the earthquake of 1906 by Alice Eastwood, but
none of Kellogg's lily types (Vollmer. 1939) or other
original specimens survived. In his monograph, El-
vves (1877—1880) mentioned receiving a letter from
Sereno Watson in 1878 in which Watson cited some
Kellogg material of L. pardalinum and L. pardal¬
inum var. angustifolium Kellogg at his disposal. In
the letter Watson noted that Kellogg's specimen of
L. pardalinum was from Alameda County, Califor¬
nia (just across the San Francisco Bay from CAS);
thus Alameda County can be inferred as the prob¬
able L. pardalinum type locality. Watson would
have been at Harvard’s Gray Herbarium when he
wrote that letter (Stafleu & Cowan, 1988), but his
specimens of L. pardalinum and its variety angus¬
tifolium are not at GH or other likely herbaria in¬
cluding CAS. K. MO. UC/JEPS. or US. It is likely
they were returned by Watson to Kellogg and de¬
stroyed in the 1906 event.

The Hesperian illustrations of the plants that Kel¬
logg described were prepared at his direction (Cur¬
ran. 1885), and an illustration of Lilium pardalinum
accompanies the protologue (Kellogg. 1859a: fron¬
tispiece). Although it comes after the title page and
table of contents, this illustration has been cited as
the frontispiece (Woodcock & Stearn, 1950) and is
so cited here. It is one of two such plates for Hes-

Novon 12: 253-261. 2002.

254

Novon

perian 3(1); the other is a drawing of a Captain John
Paty (K. Gandhi and University of California ref¬
erence staff, pers. comm. 2001). The illustration of
L. pardalinum is missing from most of the few sur¬
viving copies ol Hesperian (the University of Cali¬
fornia at San Diego does have this volume intact
with the illustration), hut it is reproduced in some
copies of Curran (1885) and in Woodcock and
Stearn (1950: 300). Since it is all that remains of
the original material that was associated with the
protologue for L pardalinum , it is here chosen as
the lectotype.

Lilium pardalinum Kellogg occurs from sea level
to 6000 leet west of the deserts in California and
in the Klamath Mountains of Oregon; it is known
to he “exceedingly variable” (Munz, 1968: 1344).
The following systematic arrangement of the L. par¬
dalinum complex organizes some of this variation,
and is based on extensive fieldwork supported by
herbarium and literature study. Five subordinate
entities of L. pardalinum are recognized below as
subspecies, as befits such genetically based infra-
specific taxa with geographic, ecologic, and mor¬
phologic significance (Thorne, 1978). These sub¬
species are distributed parapatrically: each
occupies a discrete range to the exclusion of any
other, and hybridizes with adjacent subspecies in
peripheral areas of contact, producing roughly elin-
al variation in which floral and vegetative measure¬
ments of adjacent subspecies often overlap consid¬
erably. In addition to their geographic coherence,
the subspecies of L. pardalinum occupy similar pe¬
rennially moist or wet habitats and share several
morphological features including a continuously
scaly, often branching rhizomatous bulb with joint¬
ed scales, clonal habit without the glaucous stem
or undulate leaves found in certain other lilies, an
odorless or scarcely fragrant, brightly colored pen¬
dent flower in which red- or red-orange-tipped te-
pals are strongly reflexed to resemble a Turk’s cap
and anthers are exserted on divergent filaments,
and capsules similar in shape and proportions. The
new combinations acknowledge this genetic and
morphological affinity; the resulting subspecies are
quite recognizable if still somewhat variable, and
confusion is likely to be profound only where they
meet geographically.

Flower morphology and orientation typically out¬
weigh other features in Lilium alpha taxonomy, in
part because these are closely correlated with pol¬
lination and likewise with reproductive isolation. In
Lilium pardalinum subspecies these floral features
arc basically similar, which underlies the finding
that all live are pollinated by a combination of sev¬
eral species ol hummingbirds and large swallowtail

butterflies (Papilio spp.; Skinner, 1988). In con¬
trast, L. canadense L. and L. michiganense 0. A.
Farwell, two closely related taxa from the eastern
United States, are allopatrically distributed with a
common zone ol morphological intergradation, but
their flowers are of different forms, and consequent¬
ly are pollinated primarily by different agents: eam-
panulate L. canadense by hummingbirds and
Turk's -cap L. michiganense by swallowtail butter¬
flies (Skinner, pers. obs.). Maintaining these as sep¬
arate species is thus supported despite a pattern of
parapatry with some intergradation along the con¬
tact zone (Adams & Dress, 1982; Braun, 1967;
Skinner, pers. obs.) that is similar to that displayed
by the subspecies of L. pardalinum named here.

Kky to tiii: Si hspkcies ot Lilium pardauni w Kellogg

la. Sepals and petals uniformly yellow or yellow-or¬
ange; sepals 3.5— 7.1 cm*; anthers pale yellow,

0.5— 1.3 cm; pollen yellow or orange; pistil 3.1 —

4.3 cm; capsules 2. 3-4. 2 cm; northern California
and southern Oregon .

. Lilium pardalinum subsp. wigginsii

lb. Sepals and petals — 2-toned, with yellow or or¬
ange proximal ly, distal 1/5 — 3/5 darker orange to
red; sepals 3.7—10.4 cm; anthers magenta, oc¬
casionally purple or orange, 0.5— 2.2 cm; pollen
yellow to rust; pistil 3. 3-7. 5 cm; capsules 2.2—

5.7 cm; California and southern Oregon.

2a. Sepals (5.9— )6.6— 10.4 cm; anthers 1. 1-2.2
cm; capsules 2.9— 5.7 cm; leaves 3—12 times
longer than wide, blade ± elliptic; plants
usually in large clones with many stems;

California .

. Lilium pardalinum subsp. pardalinum

2b. Sepals 3.7— 8.3 cm; anthers 0.5— 1.8 cm; cap¬
sules 2. 2^4. 8 cm; leaves 3—34 times longer
than wide, blade elliptic1 to linear; plants
usually in small clones with few stems or
single; northern California and southern
( begun.

3a. Leaves 7—34 times longer than w ide, of¬
ten concentrated proximally, often as¬
cending, sometimes horizontal, blade ±
linear; sepals (4.9— )5. 3— 8.3 cm; anthers
0.6— 1.8 cm; pollen usually dark orange;
extreme northwestern California and ad¬
jacent southern Oregon .

. Lilium pardalinum subsp. i oilmen

31). Leaves 3—17 times longer than w ide, ±
evenly distributed along stem, ± as¬
cending or horizontal, blade ± elliptic;
sepals 3.7— 7.6 cm; anthers 0.5— 1.4 cm;
pollen yellow to brown-orange; northern
California anti southern Oregon.

4a. Pollen red- or brown-orange; an¬
thers magenta; bulb scales usually
2-segmented; sepals 4.9-7. 1 ;
northern Coast Ranges near Sebas¬
topol, California .

. . Lilium pardalinum subsp. pilkinense
4b. Pollen usually yellow or bright or¬
ange; anthers orange to magenta;

Volume 12, Number 2
2002

Skinner

North American Lilium

255

bulb scales (1— )2^t-segmented; se¬
pals 3. 7-7.6 cm; northern Califor¬
nia and adjacent southern Oregon
. . . Lilium pardalinum subsp. shastense
* Measurements are based on fresh
material and are lengths unless oth¬
erwise noted.

1 a. I, ilium pardalinum subsp. pardalinum

Lilium pardalinum Kellogg subsp. pardalinum is
confined to California, and is variable in flower size,
color, and leaf size and shape. Along the length of
the Sierra Nevada this subspecies is replaced at
higher elevations by a congener; in the north by L.
pardalinum Kellogg subsp. shastense (Eastwood) M.
W. Skinner, then by L. parvum Kellogg, and finally
by L. kelle yanum Lemmon in the high Sierra to the
south. In the Klamath Mountains it gives way to L.
pardalinum subsp. vollrneri in Del Norte County, L.
pardalinum subsp. wigginsii in northern Humboldt
County, and to L. pardalinum subsp. shastense in
northeastern Trinity County. The range disjunction
between the south coast ranges in Santa Barbara
County and the Peninsular ranges in San Diego
County probably exists because L. parryi S. Watson
is the sister group to L. pardalinum (Skinner, 1988)
and replaces it in the high mountains of southern
California.

Representative specimens. U.S.A. California: Colusa
Co., Box Springs, 10 July 1972, Alice (J. Howard It
(JEPS); Humboldt Co., on Butte Creek Rd. |near Bridge-
ville, 1.1 mi. from intersection with Highway 36], 2500
ft.. 24 July 1983, M. W. Skinner 128 (GH); Placer Co.,
Bear Valley, July 1951, A. M. Vollmer & L. Beane l till
(US); Sierra Co., Goodyear’s Bar, T19N R10E SW 1/4 of
S6, 26(H) ft., 16 July 1983, M. W. Skinner 115 (GH).

lb. Lilium paialaliimm subsp. pitkinense (Be¬
ane & Vollmer) M. W. Skinner, comb, et stat.
nov. Basionym: Lilium pitkinense Beane &
Vollmer, Contr. Dudley Herb. 4: 356. 1955.
TYPE: U.S.A. California: Sonoma Co., Pitkin
Marsh, 20 July 1954. L. Beane & A. M. Voll¬
mer 1270 (holotype, DS not seen).

The Pitkin Marsh lily is a local expression of
Lilium pardalinum with small flowers, short sta¬
mens, and brighter anthers than most members of
subspecies pardalinum. It is known only from the
vicinity of Sebastopol in Sonoma County, California.
Plants that match the type concept in Beane (1955)
are extremely rare and found only in Pitkin Marsh.
Plants f. om nearby marshes are somewhat more
variable, and some clones with larger flowers, lon¬
ger stamens, and darker rust-colored pollen begin
to approach subspecies pardalinum in flower mor¬
phology; hybrids or plants virtually indistinguish¬

able from subspecies pardalinum occur a few miles
from Pitkin Marsh. Lilium pardalinum subsp. pit¬
kinense can be distinguished with some difficulty
from L. pardalinum subsp. shastense by its darker
red-orange or brown-orange pollen.

Representative specimen. U.S.A. California: Contra
Costa Co., cultivated at Canyon by Bert Johnson (propa¬
gated from material collected in Pitkin Marsh, Sonoma
Co., California, 3 June 1983, by J. Bartel and A. Q. How¬
ard), 20 June 1987, A. (A Howard 20 (GH).

lc. Lilium pardalinum subsp. shastense (East-
wood) M. W. Skinner, comb, et stat. nov. Bas¬
ionym: Lilium nevadense var. shastense East-
wood, Leaf!. W. Bot. 1: 42. 1933. Lilium
shastense (Eastwood) Beane, Contr. Dudley
Herb. 4: 357. 1955. TYPE: U.S.A. California:
Shasta Co., Goose Valley, George Dillman
ranch, July 1912,4. Eastwood 799 A (holotype,
CAS not seen).

Lilium nevadense Eastwood var. nevadense, Leaf!. W. Bot.
1: 41. 1933. Syn. nov. TYPE: U.S.A. California:
Shasta Co., Goose Valley, George Dillman ranch,
July 1912, Eastwood 799 (holotype, CAS not seen).
Lilium parvum Kellogg var. luteum Purdy, Erythea 5: 105.
1897. Lilium parviflorum (Hooker) Purdy var. luteum
(Purdy) Purdy, Card, and Purest 10: 502. 1897.
TYPE: U.S.A. California: Plumas Co., Swamps-But-
terlly Valley, 12 June 1897, R. M. Austin 1025 (lec-
totype, designated here, US).

For t he synonym Lilium parvum Kellogg var. lu¬
teum Purdy I have selected as lectotype a Plumas
County, California, collection by Mrs. R. M. Austin.
Purdy evidently based his name and description at
least in part on Austin’s collection or her descrip¬
tion of it, since he noted (1897a: 105), “This showy
form of L. parvum is from Plumas County, where it
was collected by Mrs. Austin.” Austin’s collection
is the only known original material of L. parvum
var. luteum.

In California, Lilium pardalinum subsp. shas¬
tense occurs in the Sierra Nevada from Sierra and
Butte Counties north to the region surrounding Mt.
Shasta, and thence west to the Klamath Mountains
in eastern Trinity County, and to western Siskiyou
County where it intergrades with L. pardalinum
subsp. wigginsii. A collection at US (Mrs. M. M.
Hardy 1509) indicates that the southern limit of L.
pardalinum subsp. shastense may extend as far as
Placer County at moderately high elevations (ca.
1800 m) in the Sierra, although L. pardalinum
subsp. pardalinum occurs nearby at lower eleva¬
tions, for example. Bear Valley (also in Placer Co.
at ca. 1400 m). Plants from near Klamath Falls,
Oregon, are presumed on the basis of morphology

256

Novon

!o I>e this subspecies, though live material ol them
has not been studied.

Eastwood (1933: 42) considered that “ Lilium
nevadense and its varieties include the small-flow¬
ered tiger lilies of the Sierra Nevada." 'Phis initial
publication of Lilium nevadense Eastwood included
descriptions of three varieties: L. nevadense var.
shastense Eastwood, L. nevadense var. monetise
Eastwood, and L. nevadense var. fresnense East-
wood. Based on geographical range and morpholog¬
ical description, the latter two names refer to the
southern Sierran lily now known as L. kelleyanum
Lemmon (1903), so they are of little further interest
here. Eastwood (1933) designated a single Shasta
County location (George Dillman ranch. Goose Val¬
ley, Shasta Co., California) as the type locality for
both L. nevadense and L. nevadense var. shastense,
though she did designate different collections as
types ( Eastwood 799 for the species, Eastwood
799A for the variety). She assigned no unique geo¬
graphical range to the variety shastense, although
specimen citations in the protologue indicate that
it extended from a northern limit in Siskiyou Coun¬
ty south to Plumas County. Though she did not
mention the autonym by name, specimens cited un¬
der L. nevadense exclusive of the named varieties
indicate that its range coincided more or less with
that of variety shastense, though extending farther
south and encompassing Sierra County as well. Her
Latin diagnosis (Eastwood, 1933) and subsequent
English description (Eastwood, 1948c: 136) indi¬
cate that L. nevadense var. shastense is character¬
ized by “narrower, more numerous and erect
leaves” than the “typical" variety. In fact, Lilium
pardalinum subsp. shastense includes individuals
with narrow leaves and others with broad leaves,
often within the same population, so Eastwood’s
distinction of entities with differently shaped leaves
is not maintained here.

Thus Lilium nevadense var. nevadense anti variety
shastense were published simultaneously (East-
wood, 1933), and both names apply to the material
encompassed under the new combination. The au¬
tonym has priority over variety shastense (Greuter
et al., 20(H). Art. 1 1.6), but only within the named
rank (Art. 1 1 .2). As there are no applicable epithets
at the subspecies level, shastense is selected here
for the small -flowered subspecies of Lilium pardal¬
inum in northeastern California. This is appropriate
for several reasons, f irst. Beane (1955) published
Lilium shastense to represent the small-flowered /..
pardalinum growing in Siskiyou, Shasta, and Plu¬
mas Counties. Because of Beane’s action it is also
the epithet I used — though ditl not publish formal¬
ly — in The Jepson Manual (Hickman, 1993). Fi¬

nally, shastense is preferable because the plant is
common both in Shasta County and around the base
of Mt. Shasta in Siskiyou County, anti because the
epithet nevadense has been erroneously linked to
anti therefore confused with L. kelleyanum via the
varieties monetise, fresnense, and inyoense Eastwood
(Eastwood, 1941).

Other earlier epithets that might have been
adopted for ibis purpose are rejected for various
reasons. Purdy (1897a: 105) first published Lilium
parvum var. luteum for the “showy form [of L. par-
vum ] . . . from Plumas County” with “segments of
the perianth revolute at the base," thus clearly des¬
ignating the plant recognized here as L. pardalinum
subsp. shastense and distinguishing it from the tu¬
bular-floweret! L. parvum. Purdy (1897b) subse¬
quently decided that L. parvum Kellogg should be
restricted to plants with horizontal to ascending
trumpet -shaped flowers (its current application),
and transferred the plants with revolute perianth
and pendent flowers from subalpine Plumas County
to a new* small -flowered Sierran taxon that can be
cited as L. parviflorum (Hooker) Purdy var. luteum
(Purdy) Purdy. Lilium parviflorum (Hooker) Purdy
[var. parviflorum ] encompassed similar plants
(1897b: 502) from “the middle belt of the Sierra
Nevada from Tulare County, to the south, to the
base of Mt. Shasta.” But Purdy’s (1897b) use of
Lilium parviflorum to describe the plant here called
L. pardalinum subsp. shastense was inappropriate.
Based on geography and flower size. L. eanadense
L. var. parviflorum Hooker (1838: 181), “[from the]
Columbia and Walamet [sic] Bivers." clearly refers
to what is now known as L. columbianum hort. Lei-
chtlin in Ducharte (1871; see end note for disc us¬
sion of this apparently new citation for L. colum¬
bianum). With this meaning it was first elevated to
species level as L. parviflorum by W. G. Smith
(1874), who evidently was not aware of L. colum¬
bianum ; thus L. parviflorum (Hooker) Purdy is not
only a misapplication but is objectionable as a later
homonym (Greuter et al.. 2000, Art. 53.1). Despite
the illegitimate binomial, there is no inherent ob¬
jection to reusing the final epithet of Purdy’s
(1897b) name L. parviflorum var. luteum (Purdy)
Purdy within L. pardalinum for the plant named
here (Greuter et al.. 2000. Art. 55.2). Vet this, too,
is precluded (Greuter et al., 2000, Art. 53.4) by the
very' different and prior-named L. pardalinum Kel¬
logg var. luteum hort. T. S. Ware (in Marshall.
1889), with “yellow flowers spotted with brownish
purple” that occurs “in the south, as well as more
inland [in California]” (Woodcock & Steam, 1950:
301, 303). Eastwood (1902) employed the epithet
parviflorum in an entirely new context with publi-

Volume 12, Number 2
2002

Skinner

North American Lilium

257

cation of L. pardalinum Kellogg [var.] parviflorum
Eastwood (— L. kelleyanum Lemmon; see end note
re: priority of L. kelleyanum), and Johnston (1923)
later misapplied Eastwood’s name to the taxon
named here. Eastwood’s action renders the epithet
parviflorum, like luteum, unavailable for the pre¬
sent use at any infraspecific rank within L. pardal¬
inum (Greater et ah, 2000. Art. 53.4).

Represen I alive specimens. U.S.A. California: Butte
Co., Little Chico Creek. 1000 ft., 5 July 1900, John B.
Leiherg 5011 (US); Lassen Co., Susanville, Perkin’s
Ranch, 4800 ft.. 26 June 1897. Marcus E. Jones. A.M. s.n.
(US); Placer Co.. 1893, Mrs. M. M. Hardy 1509 (US); Plu¬
mas Co.. Butterfly Valley. 25 June 1912. II. M. Hall 9275
(US); Shasta Co., Goose Valley (near Burney), 29 June, I I
July 1912. Alice Eastuood 857 (US); Sierra Co., Salmon
Lake, July 1918, Mrs. E. C. Sutcliffe s.n. (US); Siskiyou
Co., McCloud River above Big Springs, T39N B2VV S\\
1/4 of NW 1/4 of SI 3, 6 Aug. 1984. M. W. Skinner 195
(GH); town of Mt. Shasta Kampgrounds of America,
swampy ground near RR tracks, T40N R4W S9 3900 ft.,
2 7 July 1988. M. W. Skinner 266 (GH); in spring, Cayenne
Ridge, Marble Mountain, 6 July 1940, Marion Ownbey
2200, Ered G. Meyer (US); Tehama Co., Brookside, Butt
Mt.. Deer Creek Canyon, 1500 m, 16 Julv 1911, IL W.
Eggleston 7249 (US); Trinity Co., Dan Rice Cr. on Scott
Mt., Shasta- Trinity National Forest. T39N R7V\ NK 1/16
of S9. 4600 ft., 26 July 1988, M. W. Skinner 264 (GH).
Oregon: Klamath Co., meadows and woods near Lake of
the Woods, Cascade Mts., 5000—6000 ft., 5 JuL 1936. ./.
William Thompson 18102 (US).

1*1. Lilium pardalinum subsp. vollmeri (East-
wood) M. W. Skinner, comb, et stat. nov. Bas-
ionym: Lilium vollmeri Eastwood, I, call. W.
Rot. 5: 121. 1948b. TYPE: U.S.A. California:
Del Norte Co., Darlingtonia bog 1 mi. W of
Eleven Mile Creek on the old Gasquet-Grants
Pass road, 3 July 1940, A. M. Vollmer & I. L.
Wiggins 3 (holotype, DS not seen: isotype,
UC).

Lilium pardalinum subsp. vollmeri (Eastwood) VI.
W. Skinner is a narrow serpentine endemic lily of
the Siskiyou Mountains in extreme northwestern
California and adjacent Oregon. A collection (E. W.
Hammond 384, US) from near Wimer in Jackson
County, Oregon, evidently represents the northern¬
most extent of this taxon. To the south it intergrades
with L. pardalinum subsp. pardalinum, but can
generally be distinguished by its somewhat smaller
flowers (sepal length typically is 5.3— 8.3 cm vs.
6.6-10.4 cm in subsp. pardalinum), which are often
redder apically, and its frequently ascending, nar¬
rowly elliptic or linear leaves. Northern populations
in Curry County, Oregon, as well as those in the
shade throughout its range, are rather similar to L.
pardalinum subsp. pardalinum, though generally
the plants are less overtly clonal and therefore less

clumped, fn the eastern part of its range, for ex¬
ample west of Grayback Mountain in Josephine
County, Oregon, and near Sanger Peak in Del Norte
County, California, it intergrades extensively with
L. pardalinum Kellogg subsp. wigginsii (Beane &
Vollmer) M. W. Skinner, producing swarms of in¬
termediate individuals that vary greatly in leaf ar¬
rangement and shape, and flower and anther col¬
oration.

Purdy (1919: 522) applied the name Lilium roe-
zlu to this plant, but this was an orthographic cor¬
rection and misapplication of Lilium roezlii Regel
(1870: 521. as “L. roezli ”), a taxon of uncertain
provenance and identity. Although the illustration
(Regel. 1870: t. 667; it is labeled "L. roezlel ’) ac¬
companying Regel’s lily shows a narrow-leaved Lil¬
ium pardalinum- type lily, the implied locality of
Utah (from “Mormon States’’) is clearly in error as
no native Lilium occur in Utah. Based on Roezl’s
travels, Eastwood (1948a) decided that Roezl col¬
lected it in the Sierra Nevada; thus Regel’s Lilium
roezlii is generally considered to be synonymous
with L. pardalinum var. angustifolium Kellogg
(1859a: 300), itself a synonym of L. pardalinum
subsp. pardalinum.

Purdy (1919) determined that there were two
forms ol his Lilium roezlii. Me first (1901: 354)
identified the form with flowers “orange dotted with
maroon,” which he subsequently (1919: 522) called
the “type.” This plant was later named L. wigginsii
Beane & Vollmer (in Beane, 1955), and is com¬
bined below as another subspecies of L. pardalinum
Kellogg. The other he described (1919: 522) as
growing farther to the west and “orange in the cen¬
ter and deep crimson on the outer two thirds.” It is
this plant that Eastwood later described as L. voll¬
meri. the type of which she said (1948b: 122) was
initially (mis)identified by Albert M. Vollmer as L.
roezlii Regel.

Eastwood’s suggestion (1948b: 123) that her new
species Lilium vollmeri is perhaps only an ecolog¬
ical variety of L. occidentale Purdy is not support¬
able; the more or less parallel stamens and un¬
branched rhizome of Lilium occidentale are
uniquely at odds with L. pardalinum.

Representative specimens. U.S.A. California: Del
Norte Co., Old Gasquet Toll Rd. at mile marker 10.75, in
meadow, T17N R3E NE 1/4 of SW 1/4, 1800 ft., 22 July
1985. M. W. Skinner 199 (GH). Oregon: Currv Co., U.S.
f orest Service Rd. 3698, 0.4 mi. before 3795, T37S
R12W NE 1/4 of SE 1/4, 3700 ft., 24 July 1985. M. If.
Skinner 201 (GH).

258

Novon

le. I, ilium punlalinum subsp. wifjftinsii (Beane
& Vollmer) M. W. Skinner, comb, et stat. nov.
Basionym: Lilium wigginsii Beane N Vollmer,
Contr. Dudley Herb. 4: 355. 1955. TYPE:
U.S.A. California: Siskiyou Co., Siskiyou
Mountains near the Oregon border W of High¬
way 99, 16 July 1954, L. Beane & A. M. Voll¬
mer 1266 (holotype, l)S not seen; isotypes,
MO, US).

Lilium pardalinum subsp. wigginsii is a Klamath
Mountain endemic lily that occurs broadly along
the boundary between Del Norte and Siskiyou
Counties, California, and east through southeast Jo¬
sephine County, Oregon, to Ml. Ashland in Jackson
County (Ballantyne, 1980). Il intergrades with L.
pardalinum subsp. shastense in the Marble Moun¬
tains of Siskiyou County, California. The solid or¬
ange or yellow-orange flowers and yellow anthers
are distinctive. Genetic instability in this subspe¬
cies is frequently expressed as malformed (lowers
with shrunken or missing reproductive structures,
and is probably due to introgression with other L.
pardalinum subspecies; similarly malformed flow¬
ers occur where L. canadense and L. grayi S. Wat¬
son intergrade in Virginia (Skinner, pers. obs.).

Purdy’s (1901 , 1919) application of the name Lil-
ium roezlii Regel to this taxon is discussed above
under L. pardalinum subsp. vollmeri. In the proto-
logue for L. wigginsii, here reduced to a subspecies,
Beane (1955: 355) noted that "this [may have been
t lie Idyl offered to the florist trade by Mr. Carl Pur¬
dy under the name Lilium roezlii (yellow) Regel.”
This is certainly consistent with Purdy’s (1919,
1935) own writings.

Representative specimens. U.S.A. California: Siskiyou
Co., Happy Camp to O’Brien Rd. at mile marker 7.5,
TI8N R6E SIO, 4100 ft., 21 July 1985, M. W. Skinner
198 ((41). Oregon: Ja< ■kson Co., Rogue River National
Eorest, 1.5 mi. S of Ashland Campground, T40S RIP SW
1/4 of SW 1/4 of S20. 23 July 1988, M. It. Skinner 255
((44); Josephine Co., ( i ray back Rd., 6.2 mi. from junction
with f orest Service Rd. 4611/070, T39S R7E w 1/4 of
S25, 3200 ft., 22 July 1988, M. W. Skinner 254 ((41).

2. Lilium washingtonianum Kellogg, Hesperian
3: 340. 1859. TYPE: Illustration, Hesperian 3:
341. 1859 (lectotype, designated here).

A. M. Kellogg (1859b) did not formally designate
type material for Lilium washingtonianum, and
specimen material associated with the protologue is
unknown. There are no Kellogg specimens of L.
washingtonianum at likely major herbaria includ¬
ing K, MO, UC/JEPS, or US. The only known Kel¬
logg specimen of L. washingtonianum [4. M. Kel¬
logg & Harford 898 (or 998, the number is

illegible), GH] lacks locality information (Walter
Kittredge, pers. comm. 2001) and is therefore un¬
suitable. The protologue contains no indication of
any specimens that Kellogg relied on other than his
own. Il appears that the illustration from the pro¬
tologue (Kellogg, 1859b: 341) is the earliest sur¬
viving original material associated with Kellogg's
description, so it is chosen here as the lectotype
following tin1 rationale for typification of Lilium par¬
dalinum discussed above. It is a black and white
drawing of a mature fruit (Kellogg, 1859b: 341).

Kl V TO 1111'. SlIBSl’Kt IKS OF LttJl M WASHIKGTONIAM M Kf.I -
i .< x a ;

la. Sepals and petals aging light pink or remaining
white: bulbs usually (36%) without notched or
segmented scales; sepals 8— 1 1.3 cm; Sierra Ne¬
vada and southern Cascades of California ....
Lilium washingtonianum subsp. washingtonianum
II). Sepals and petals aging deep pink or lavender;
bulbs usually (88%) with some notched or seg¬
mented scales; sepals (6. 1 — )6.7— 9.5 cm; moun¬
tains of northern California and Oregon . . .

.... Lilium washingtonianum subsp. purpurascens

2a. Lilium washingtonianum subsp. washing-
tonianuin

Distribution. California: the Sierra Nevada from
Fresno County north to the southern Cascade
Ranges at Ml. Shasta.

Representative specimens. U.S.A. California: Butte
Co., cut-over yellow pine forest now grown up to dense
brush about 1/2 mi. W of Chaparral, very fragrant, 5100
ft., 6 July 1940, Albert M. Vollmer & Ira L. Wiggins It)
(US); Mariposa Co., near Yosemite, Sierra Nevada, 1875.
John Muir 8095 (MO): Nevada Co.. Highway 20, 2.7 mi.
NE of White Cloud Campground, 14 July 1983, M. W.
Skinner 104 ((41): Tuolumne Co., hill S of Camp 127,
7500 ft.. 21 July 1863, W. II. Brewer 1981 (US).

2I>. Lilium washingtonianum subsp. purpur-
ascens (Steam) M. W. Skinner, stat. nov. Bas¬
ionym: Lilium washingtonianum var. purpur¬
ascens Steam, Card. Chron. 124: 13. 1948.
Lilium purpureum Purdy, J. Roy. Hort. Soc. 26:
351-362. 1901. Lilium washingtonianum var.
purpureum (Purdy) Purdy, J. lot. Gard. Club 3:
509. 1919. non Lilium washingtonianum var.
purpureum hort. W. Bull ex Baker. J. Linn. Soc.
Bot. 14: 233. 1874. TYPE: U.S.A. California:
Humboldt Co., 0.4 mi. S of Onion Lake on
U.S. Forest Service road 1 3 NO 1 . TUN R4F
SE 1/4 of SE 1/4 of S6, 4400 ft., 5 Aug. 1983,
M. W. Skinner 1 61 a <V h (neotype, designated
here, Gil).

W ith this taxon. Carl Purdy (1901, 1919) pol¬
ished his habit of appropriating previously pub-

Volume 12, Number 2
2002

Skinner

North American Lilium

259

lished names and misapplying them to other taxo¬
nomic concepts, often without attribution to the
original authors. To describe the pink-flowered
northern plants of Lilium washingtonianum Kellogg
here called Lilium washingtonianum Kellogg
subsp. purpurascens (Stearn) M. W. Skinner. Purdy
(1901) evidently recycled the final epithet of L.
washingtonianum Kellogg var. purpureum hort. W.
Hull ex Baker (1874) (= L. rubescens Watson, 1879)
and elevated it to species as L. purpureum Purdy.
[According to T. Masters (1874), L. washingtonian¬
um Kellogg var. purpureum hort. W. Bull ex Baker
had first been distributed (but not formally pub¬
lished) by Bull as L. purpureum in 1873: subse¬
quent mentions of the latter name by Masters
(1874) and T. Moore (1874) did not validly affect
publication since neither author accepted the taxon
(Greuter et al.. 2000, Art. 34.1).] Curiously, Purdy
(1901) reproduced in support of his L. purpureum
the illustration from Masters’s (1874) supplemental
description of L. washingtonianum var. purpureum
hort W. Bull ex Baker (1874), the ascending and
erect flowers of which clearly diagnose L. rubescens
S. Watson. Watson (1879) had c ited Baker's name
in synonymy and Masters’s use of it and his work
links the name L. rubescens to that particular illus¬
tration. Despite illustrating the wrong taxon — for
which he confusingly retained the original Masters
(1874) label of L. washingtonianum var. purpu¬
reum — Purdy’s (1901) description of L. purpureum
unequivocally indicates what is here called Lilium
washingtonianum subsp. purpurascens. (Purdy
notes a distribution extending through Oregon to
the Columbia River and heavy bulb with jointed
scales, a combination which could diagnose no oth¬
er taxon.) Purdy subsequently (1919) reevaluated
his new species and reduced it to L. washington¬
ianum var. purpureum (Purdy) Purdy, although he
(1901: 361; 1919: 51 1) and others considered this
same name sensu Baker (1874) to be a synonym of
L. rubescens. Stearn (1948) recognized that L. wash¬
ingtonianum var. purpureum sensu Purdy conflicted
with the earlier Baker name, and rectified this with
a new name, L. washingtonianum var. purpurascens
Stearn. The present action elevates Stearns name
to subspecies.

W. T. Stearn did not designate type material of
L. washingtonianum var. purpurascens nor was
there a diagnostic illustration accompanying the
protologue. According to Boy Vickery (BM, pers.
comm. 2001), there are no specimens annotated by
Stearn to variety purpurascens at BM. where Stearn
worked and deposited his collections, nor is there
any known material of this taxon annotated or col¬
lected by Carl Purdy. The lack of original type ma¬

terial necessitates designation of a neotype; for this
I have chosen a collection from Humboldt County,
California, an area that Purdy knew well and which
was undoubtedly close to the place that provided
the plants on which his taxon was based.

Lilium washingtonianum subsp. purpurascens
supplants the Sierran subspecies washingtonianum
near Mt. Shasta in Siskiyou County, California,
where intermediates occur, and extends west
through the Klamath Mountains and north through
the Cascade ranges to Mt. Hood in Clackamas
County, Oregon. In addition to the characters men¬
tioned in the key, subspecies purpurascens is dis¬
tinguished from subspecies washingtonianum by
more compact bulbs with longer scales. These mod¬
est differences indicate that these taxa are best rec¬
ognized as subspecies rather than species.

Representative specimens. U.S.A. California: Hum¬
boldt Co., Forest Service 15N01 (the Gasquet— Orleans
road], 3.8 mi. S of intersection with Forest Service I3N01,
T12N R5E NF 1/4 of NW 1/4 of S16. 44(H) ft., 30 July
1983, M. W. Skinner 145 (GH); 0.4 mi. S of Onion Lake
on LJ.S. Forest Service road 13N01, T1 1 IN K4K SE 1/4 of
SE 1/4 of S6, 44(H) ft., 5 Aug. 1983, M. W. Skinner 162a
A* b (GH); Siskiyou Co., among shrubs, summit of Cayenne
Ridge, Marble Mountain, flowers white with purple spots,
6 July 1940, Marion Ownbey 2196 , with Fred (1. Meyer
(US). Oregon: Klamath Co., Swan Lake Valley, 7 July
1905, Elmer I. Applegate 705 (US).

A Note on the Correct Citation of L.

COLUMBIA NUM

Ducharte’s (1871) recapitulation in July 1871 of
a letter from M. Leiehtlin apparently represents the
first confirmed and valid publication of Lilium col-
umbianum; thus the name may be cited as L. col-
umbianum hort. Leiehtlin in Ducharte, J. Soc. Cent.
Hort. France ser. 2, 5: 98 (1871). Leichtlin’s letter
and Ducharte’s commentary provide the distribu¬
tion and distinguishing features of the taxon, and
precede by two months a short, unauthored notice
(Anonymous, 1871) published in September 1871
in Gardeners' Chronicle, which also validly estab¬
lished the name. This brief notice, sometimes at¬
tributed to .). G. Baker (e.g., Watson, 1880: 167)
but more likely the work of M. Thomas Masters
(John Wiersema and Brent Elliott, pers. comm.
2001), attributes the name to earlier “New York
publications,” but that precise reference — which
may well be the first instance of the name- — has not
been found. Baker (1873) subsequently published
L. columbianum hort. Leiehtlin without an adequate
description: only the distribution was given, and
this could not have been considered by him to dis¬
tinguish the plant from others (Greuter et al., 2(H)(),
Art. 32.2). In his revision of the Tulipae, Baker

260

Novon

(1874) then validly published the name as L. col¬
umbianum Hanson in hurt., Leichtlin. It is this last
that is usually cited in floras and other works, the
authority variously given as Hanson, or Baker, or
Hanson ex Baker. Although Baker (1874) ascribed
the name to Hanson rather than to himself as he
does for other nomina nova in this work, no indi¬
cation is given that Hanson contributed to the de¬
scription; this later homonym is perhaps best cited
(Greuter et al., 2000, Art. 46.4) as L. columbianum
hort. Hanson ex Baker (1874).

A Noth; on tiik Priority ok Liuijm kelleyanum
Lkmmon

Kast wood’s (1902) Lilium pardalinurn Kellogg
[var. | parviflorum Eastwood precedes Lemmon’s
(1903) publication ol L. kelleyanum, but as noted
above, al the rank ol species her epithet parviflorum
becomes a later homonym of L. parviflorum (Hook¬
er) W. G. Smith (1874) (= /.. columbianum hort.
Leichtlin, in Ducharte, 1871). Therefore the next
legitimate specific epithet must be adopted (Greuter
et ah, 2000. Art. 1 1.3), and this is Lemmon’s (1903)
L. kelleyanum. Lemmon’s description is somewhat
incomplete but it well fits the plant now known as
L. kelleyanum, especially his flower size [segment
length 1 — 1.5 inches (2.5—4 cm)]. His obscure ref¬
erence (1903: 300) to “plants of the larger, yellow-
flowered lily | that] abound in tin1 vicinity” is prob¬
ably to L. pardalinurn Kellogg subsp. pardalinurn
growing at lower elevations. The type localities that
lit' cited in the protologue — along Bubb’s Creek and
Copper Creek in Fresno County, California — are lo¬
cated near the center of the range of L. kelleyanum,
and there are no other Lilium with small flowers
(segments < ca. 7 cm) until the southern limit of
L. parvum Kellogg is reached about 60 miles to the
north.

Acknowledgments. My assistants Manuel Ler-
dau, Shawn Lum. David Posner, and Jennifer Talbot
made my dissertation fieldwork enjoyable, and 1 am
grateful for their hard work, much of which was
applied to this effort. I especially thank Peter F.
Stevens for his help, encouragement, and friend¬
ship throughout the years and for nomenclatural as¬
sistance with this paper, and John Wiersema for
cogent interpretation of parts of the International
Code of Botanical Nomenclature. Victoria Hollow-
ell’s critical reading was very much appreciated,
and Gerald Smith and two anonymous reviewers
significantly improved the manuscript. Brent Elliott
at the Royal Horticultural Society (London), Vicki
McMichael al MO, and curators al BM (Boy Vick¬
ery), ( IAS (Mona Bourell), Gil (Emily Wood), K

(Aaron Davis), and UC/JEPS (Barbara Fitter) were
most helpful in checking holdings anil providing
information. Kaneheepuram Gandhi and Walter
Kittredge at Gil provided essential help with typ-
ification issues. The Hardman Foundation funded
part ol this work, as did the Harvard University
Department of Organismic and Evolutionary Biol¬
ogy and its Fernald Fund.

Literature Cited

Adams, If. M. & A. J. Dress. 1982. Nodding Lilium spe¬
cies of eastern North America (Liliaceae). Baileya 21:
165-188.

Anonymous. 1871. Untitled. Card. Chron. ser. 1(39):
1257.

Baker, J. G. 1873. A classified synonymic list of all the
known lilies, with their native countries, and references
to the works where they are figured. J. Roy. Hort. Soc.
4(13): 45.

- . 1874. Revision of the genera and species of Tu-

lipae. J. Linn. Soc.. Rot. 14: 21 1—310.

Ballantyne, 0. 1980. A preliminary study of Liliums bo-
landeri, occidentale, vollmeri, and wigginsii. Prepared
for Six Rivers National Forest, unpublished.

Beane. L. 1955. Some undescribed lilies from the Pacific
Coast and a preliminary revision of the southern Cali¬
fornia species formerly associated with Lilium humbold-
tii. Contr. Dudley Herb. 4(8): 355—360 + illus.

Braun, E. L. 1967. The Monocotyledoneae: Cat-Tails to
Orchids. Vascular Flora of Ohio, Vol. I. Ohio State
Univ. Press, Columbus.

Curran. M. K. 1885. I.ist of the plants described by Dr.
Albert Kellogg. Dr. H.H. Behr. and Mr. H.N. Bolander;
with an attempt at their identification. Bull. California
Acad. 1: 128-151.

Ducharte, M. P. 1871. Observations sur le genre I, is ( Lil¬
ium Tourn.), & propos du catalogue de la collection de
ces plantes qui a ete form^e par M. Max. Leichtlin, de
Carlsruhe. J. Soc. Cent. Hort. France ser. 2. 5: 87—106.
Eastwood, A. 1902. A flora of the South Fork of Kings
River from Millwood to the head waters of Bubhs Creek.
Publ. Sierra Club 27: 1—96.

- . 1933. An undescribed lily from the Sierra Ne¬
vada of California. Lead. W. Rot. 1(5): 41—1.3.

- . 1941. New Western plants. Lead. W. But. 3(1):

18.

- . 1948a. Studies of Pacific Coast lilies— I. Lead.

W. Rot. 5(6): 103-104.

- . 1948b. Studies of Pacific Coast lilies — II. Lead.

W. Rot. 5(7): 120-123.

- . 1948c. Studies of Pacific Coast lilies— II. Lead.

W. Rot. 5(8): 133-138.

Elwes, H. J. 1877—1880. A Monograph of the Genus Lil¬
ium. Illus. by W. H. Fitch. Taylor and Francis, London.
Greuter, W., J. McNeill. F. R. Barrie, H. M. Runlet, V.
Demoulin, T. S. Filgueiras. I). H. Nicolson, P. C. Silva,
J. E. Skog. P. Trehane, N. J. Turland & 1). I.. Hawk-
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Nomenclature (St. Louis Code). Regnum Veg. 138.
Hickman, J. C. 1993. The Jepson Manual: Higher Plants
of California. Univ. California Press, Berkeley.

Hooker, J. D. 1838. Flora Boreali-Americana. Bohn, Lon¬
don.

Johnston, I. M. 1923. Diagnoses and notes relating to the

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spermatophytes chiefly of North America. Contr. Gray
Herb. 68: 81 .

Kellogg, A. M. 1859a. Leopard lily , L pardalinum. — (Kel¬
logg). Hesperian 3: 300—301, illus. as “frontispiece” on
unnumbered page.

- . 1859b. Lady Washington lily, L. washingtonian-

um. (Kellogg). Hesperian 3: 340—341.

Lemmon, J. G. 1903. Kings River Outing, 1902 — Botan¬
ical notes, including an important discovery. Sierra
Club Bull. 4: 292-300.

Marshall. W. 1889. f loral Committee. J. Roy. Hort. Soc.
1 1 : 94.

Masters, T. 1874. Lilium u/ashingtonianum purpureum.
Card. Chron. ser. 2, 2: 322.

Moore, T. 1874. Garden lilies. — Chapter VII. f lorist and
Pomol., ser. 3, 7: 253—256.

Munz, P. A. 1968. A California Flora and Supplement.

Univ. California Press, Berkeley.

Purdy, C. 1897a. New west American lilies. Erythea 5:
103-105.

- . 1897b. Lilium parvum and L. parviflorum. Gard.

and Forest 10: 502.

- . 1901. The lilies of the western United States and

British Columbia. J. Roy. Hort. Soc. 26: 351-362.

- . 1919. Pacific Coast lilies and their culture. J.

Int. Gard. Club 3(4): 497-532.

- . 1935. Western American lilies. Lily Year-Book

No. 4: 43-54.

Regel, E. A. von. 1870. I. Originalabhandlungen, 1)
Abgebildete Pflanzen, a) Lilium roezli Rgl. Gartenflora
19: 321, I. 667.

Skinner, M. W. 1988. Pollination Ecology and Floral Evo¬
lution in Pacific Coast Lilium. Ph .14. Dissertation, Har¬
vard University, Cambridge.

- . 1993. Lilium. Pp. I 198-1200 in J. C. Hickman

(editor), The Jepson Manual. Univ. California Press,
Berkeley.

Smith, W. G. 1874. Lilium parviflorum— L. dalmaticum—L.

avenaceum. FI. Mag. 13, t. 136.

Stearn, W. T. 1948. The botanical names of some lilies.

Gard. Chron. ser. 3. 124: 13.

Stafleu, F. A. & R. S. Cowan. 1988. Taxonomic Literature,
Vol. 7, 2nd ed. Regnum Veg. 116.

Thorne, R. F. 1978. New subspecific combinations for
Southern California plants. Aliso 9: 189—196.

Vollmer, A. M. 1939. Lilium pardalinum and its allies.
Lily Year-Book 8: 108-118.

Watson, S. 1879. Contributions to American botany. IX. I.
Revision of the North American Liliaceae. Proc. Amer.
Acad. Sci. 14: 213-288.

- . 1880. Geological Survey of California. Botany.

Vol. 11. Cambridge, Massachusetts.

Woodcock, H. D. & W. T. Stearn. 1950. Lilies of the
World: Their Cultivation and Classification. Country
Life, London.

Novae Gesneriaeeae Neotropicarum IX: Cremospermopsis, A New

Genus from Colombia

Laurence E. Skog

U.S. National Herbarium, MRC-166, Smithsonian Institution, P.0. Box 37012,
Washington, D.C. 20013-7012, U.S. A.
skog.larry@nmnh.si.edu

l Airs l\ Kvist

Unit of Forestry, Royal Veterinary and Agricultural University,

Dk-1871 Frederic ksberg C (Copenhagen), Denmark.
lars_peter_kvist@hotmail.com

Abstract. A new genus in the Gesneriaeeae,
Cremospermopsis L. E. Skog & L. P. kvist, is de¬
scribed from the very humid forests in the Colom¬
bian departments of Antioquia and adjacent Boli¬
var. The new genus of two species, C. cestroides and
C. parvijlora , is similar and may be related to Cre-
mosperma, but (litters in having inflorescence
bracts, unequal calyx lobes, and spherical papillate
seeds.

Key words: Colombia, Cremosperma, Cremos¬
permopsis, Gesneriaeeae.

The flowering plant family Gesneriaeeae is better
represented in Colombia than in any other neotrop¬
ical country, with approximately 400 known species
according to a recent survey by Kvist et al. (1998).
While identifying Colombian Gesneriaeeae for that
survey, the authors found collections of an undes¬
cribed gesneriad that is apparently endemic. This
new genus, Cremospermopsis, is from a small area
in the northwestern Colombian department of An¬
tioquia, with two collections from adjacent Bolivar.
Specimens of the new genus had often been filed
in herbaria among unidentified material of Bubi-
aceae or (less frequently) Acanthaceae. A closer
look demonstrates that Cremospermopsis cannot be
a member of either of those families because of the
presence of hypogynous flowers and the absence of
stipules on the stems of the new genus. In addition,
plants have unilocular capsules with many liny
seeds, four anthers with coherent stamens, and a
more or less regular or divided nectary surrounding
the ovary, characteristics that place I he genus in
the Gesneriaeeae.

Other specimens of Cremospermopsis were rec¬
ognized as Gesneriaeeae, and mostly as unknown
species of Cremosperma Bentham. This was reason¬

able since the two genera are similar in having con¬
nate calyx lobes and filaments adnate to the corolla
tube base. The name of the new genus alludes to
the similarity of Cremospermopsis to Cremosperma.

The two species of Cremospermopsis have a
unique combination of characters, such as zygo-
morphic calyces, corolla limbs with glandular tri-
chomes, and spherical papillate seeds, which jus¬
tifies placing them in a new genus. They share more
characteristics (e.g., erect habit, opposite leaves,
calyx lobes connate, etc.) with Cremosperma than
with any other genus, but are clearly distinguished
by the presence of floral bracts.

We will not here assign Cremospermopsis to a
tribe. Possible tribes are Beslerieae and Napean-
llieae, as defined by Wiehler (1983). The presence
or absence of inflorescence bracts has been given
much importance in the taxonomy of neotropical
Gesneriaeeae, mostly because Wiehler (1983)
maintained an amended tribe Beslerieae, including
Hesleria L., Cremosperma Bentham, Gasteranthus
Bentham, Reldia Wiehler, and Resia H. E. Moore,
on the absence of floral bracts, a superior ovary, the
lack of tubers or rhizomes, and seeds without fu-
nicles, etc., in the included genera. At the present
time, however, the circumscription of Beslerieae
may be open to question. For example, the discov¬
ery of a new subspecies of Resia ichthyoides Eeeu-
wenberg having floral bracts led Skog and de Jesus
(1997) to transfer the genus Resia from the Besler¬
ieae to the formerly monotypie tribe Napeantheae
where it seems better placed. The fact that species
of Cremospermopsis also have bracts despite their
close resemblance and possible relationship to Cre¬
mosperma makes it even clearer that tribal limits
may well have to be reexamined. Table 1 compares
Cremospermopsis with Reldia and Cremosperma

Novon 12: 262-269. 2002.

Volume 12, Number 2
2002

Skog & Kvist

New Genus from Colombia

263

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(both included in Beslerieae) as well as with Na-
peanthus and Resin (tli«* two genera currently in the
tribe Napeantheae).

Although Crem os perm ops is appears to he most
similar to Cremosperma , further studies are neces¬
sary to determine if Beslerieae might have to he
expanded to include genera and species having flo¬
ral bracts, or alternatively if Cremosperma should
he transferred to Napeantheae together with Cre-
rnospermopsis.

Cremospermopsis I,. E. Skog & L. P. Kvist. gen.
nov. TYPE: Cremospermopsis cestroides
(Fritseh) L. E. Skog E. P Kvist.

Ad Cremosperma affinis sed in infiorescentiis congestis
bracteatis in lobis calycis inaequalibus zygomorphis, cum
trichomatibus glandularibus in capsulis in sicco fatiscen-
libus et seminibus sphaerieis papillatis differt.

Plants herbs to subshrubs, erect or rarely ap-
pressed to ground. Leaves opposite, subisophyllous
in pairs, blades mostly ovate, elliptic, obovate or
oblanceolate, adaxially darker green than abaxially,
lateral veins 8 to 12 per side; petiolate. Inflores¬
cences umbel-like cymes, usually congested but oc¬
casionally relatively open, of several flowers, pe¬
dunculate and pedicellate; with mostly 4 lanceolate
to ovate bracts from the apex of the peduncle. Ca¬
lyx lobes connate between 1/3 and 2/3 of their
length, subequal to strongly unequal, 3 or 4 lobes
usually lanceolate, the remaining 1 or 2 much wid¬
er and ovate but variable even within the same in¬
florescence and even more unequal in fruiting than
during flowering, all lobes directed forward and
with entire margins. Corolla ecalcarate, funnelform
to tubular, outside glabrous or glandular hairy, es¬
pecially on the limb, inside glabrous; limb ± bi¬
labiate. lobes rotund, 3 lower lobes larger than 2
upper lobes. Filaments adnate to corolla tube base
at least 2/3 of their length, glabrous, anthers co¬
herent, thecae confluent, staminode sometimes but
apparently not always present. Nectary annular or
higher on both sides of ovary and low or absent in
between. Ovary and style glabrous, stigma capitate.
Fruit a dry capsule, compressed and with two
grooves, dehiscing irregularly and disintegrating in
the persistent calyx, which may function as a
splash-cup; seeds very numerous, ± regularly
spherical, very small, surface papillate.

Cremospermopsis is easily recognizable by the
combination of congested, bracteate inflorescences
with small to tiny corollas lacking spurs, but having
densely glandular hairy throats and limbs. Both
species are particularly similar to species of Cre¬
mosperma, where the type species was formerly

placed. The unequal calyx lobes and the resulting
zygomorphic calyces is an unusual feature, and oth¬
er differences from Cremosperma include the pres¬
ence of spherical, papillate seeds vs. narrowly el¬
liptic non-papillate seeds in Cremosperma, and the
abundance of glandular trichomes on the limb and
in the throat of Cremospermopsis. The fruits are also
different. The capsules in Cremosperma split (more
or less regularly) but their valves remain surround¬
ing the seed mass during dispersal, while the cap¬
sules of Cremospermopsis apparently disintegrate
and virtually disappear leaving the seeds free in
the persistent calyx.

The two Cremospermopsis species have similar
geographical ranges almost completely restricted to
the Colombian department of Antioquia ( C . cestro¬
ides has also been collected twice in adjacent Bo¬
livar). The species have both been collected at sev¬
eral different localities suggesting that they may
also occur in the same habitat, viz., very humid
forests and often c lose to streams. Plants have been
collected from near sea level (in the Cauca River
valley) to 1550 m elevation, but most collections
come from between 500 to 900 m elevation, pos¬
sibly corresponding to the low-elevation cloud for¬
ests with a very high Gesneriaceae diversity like
that found in western Ecuador (Skog & Kvist.
2000).

little is known about the ecology of the Cremos¬
permopsis spec ies, which have not yet been studied
by the authors in their natural habitats. Both spe¬
cies have small flowers, not particularly conspicu¬
ous by their coloration, which varies from white to
yellow, but the corolla limbs and throats have nu¬
merous glandular tric homes suggesting the libera¬
tion of substances that might attract pollinators. In
addition, the expanded calyx lobes, which tend to
be more conspic uous than the corollas particularly
in Cremospermopsis parviflora, may also help to at¬
tract pollinators. The corolla tubes of Cremosper¬
mopsis flowers seem nearly blocked by the coherent
anthers, which tend to be located about where the
corolla tube is narrowest. However, both species
have well-developed glands around the ovary sug¬
gesting that pollinators may indeed find nectar in
the flowers.

Cremospermopsis has numerous tiny seeds like
many other understory Gesneriaceae, and the sur¬
rounding persistent calyces may function as splash-
cups, as the capsules seem to disintegrate at ma¬
turity. Water transport, and possibly small animals
to which the papillate seeds may adhere, may dis¬
perse the seeds.

The status of the two Cremospermopsis species in
the wild is unknown. It is possible, however, that

Volume 12, Number 2
2002

Skog & Kvist

New Genus from Colombia

265

both species are endangered. Low-elevational hu¬
mid forests in Antioquia have mostly disappeared,
and humid forest Gesneriaceae tend to be very vul¬
nerable to deforestation, as they cannot survive the
exposure to dryer microclimates.

Key to phi: Species of Cremospermopsis

la. Corolla nearly 2 times as long as the calyx and not
hidden by the calyx lobes; leaf blades oblanceolate or

less commonly elliptic, apex acuminate .

. C. cestroides

lb. Corolla shorter than the calyx and hidden by the calyx
lobes; leaf blades obovate or occasionally elliptic,
apex obtuse or less frequently acute . . C. parviflora

Cremospermopsis cestroides (Fritsch) I,. E.
Skog & L. P. Kvist, comb. nov. Basionym: Bes-
leria cestroides Fritsch, Notizbl. Bot. Gart. Ber¬
lin 11: 962. 1934. Cremosperma cestroides
(Fritsch) C. V. Morton, J. Wash. Acad. Sci. 2B:
348. 1938. TYPE: Colombia. Antioquia:
Guadalito, growing in forest shade, 2000 ft.. 4
Feb. 1880. W. Kalbreyer 1391 (lectotype. des¬
ignated here. K: B photo, US). Figure 1.

Plants herbs to subshrubs, stems erect (0.3-)
0.70-1. 5(— 2.0) m tall, to 7 mm diam.. glabrescent
below to distally pilose or villous and occasionally
strigose, intemodes (0.3— )1— 5(— 9) cm long. Leaves
with blades oblanceolate or less commonly elliptic,
8— 18(— 22) X (2— )4— 7(— 9) cm. apex acuminate, base
cuneate, rarely acute, margin remotely serrate to
subentire, adaxially pilose to glabrous, abaxially ±
sparsely pilose or strigose, secondary veins 8 to 12
per side, pilose or strigose below. Inflorescences of
umbel-like cymes, of 3 to 15 flowers, pilose, stri¬
gose or puberulent: peduncles 2— 5(— 8) cm long,
distally with 2 to 4 bracts, each ovate, 5—10 X 1—
4 mm; pedicels 2—8 mm long. Calyx lobes green,
occasionally red tinged, outside pilose especially
along margin, inside glabrous, connate, subequal to
unequal, lanceolate to ovate with acute to acumi¬
nate apex, all 8—10 mm long and up to 12 mm long
in fruiting stage, 3 or 4 of the lobes only 2—4 mm
wide and the other 1 or 2 lobes to 6 mm wide.
Corolla tubular, 1.2— 1.7 cm long, tube 1.0— 1.4 cm,

1— 2 mm wide at base, diam. 2.5— 3.5 mm basally,
narrowing to 2.0-2. 5 mm and then ampliate to 3—
4 mm distally, yellow or white, outside upper 2/3
of length glandular pilose to villous, inside glabrous
except apically; throat 3.5—5 mm diam.. limb yel¬
low, densely glandular hairy with trichomes that
may be short or have a more extended stalk of sev¬
eral cells, dorsal lobes often with pink or violet
markings, 1.5-2 mm long, lateral and ventral lobes

2— 3.5 mm long. Filaments adnate to corolla tube

base for 5—8 mm, above free for 1 .5—3 mm; anthers
0.7— 1.3 X 0.5— 0.8 mm; nectary ± regularly an¬
nular. 0.3— 0.7 mm high. Ovary 2 mm high. Capsule
5X3X2 mm.

Distribution and ecology. Plants are endemic to
the Colombian departments of Antioquia and Boli¬
var, growing in disturbed primary or old secondary
wet forest, along streams and in shady areas; (20—)
500-1 000(— 1550) m. and collected in flower in all
months of the year, except April and November; in
fruit in June, but probably most of the year.

Cremospermopsis cestroides is distinguished from
the apparently more abundant C. parviflora (see be¬
low) by the acuminate leaf apices and by larger
corollas of the former.

This species, described originally by Fritsch as
a Besleria species anti later transferred to Cremos¬
perma by Morton, was typified by a specimen at B
( Kalbreyer 1391). Unfortunately, the holotype is no
longer extant, although a photograph of the speci¬
men at B taken in the 1930s is lodged at US and
probably in other herbaria. The duplicate of Kal¬
breyer 1391 at K is selected here as lectotype. It is
remarkable that neither Fritsch nor Morton com¬
mented on the bracteate inflorescences on the types
anil clearly evident in the photograph of the holo¬
type and on the lectotype specimen at K.

Additional specimens examined. COLOMBIA. Antio¬
quia; Mpio. Anon, 3 km from Providencia hydroelectric
plant, valley near confluence of Tirana stream with Rfo
Anorf, 28 km SW of Zaragoza, Alverson et al. 130 (COL);
Mpio. Amalfi, 8—15 km from Amalfi to Rumazon, Salazar
and La Playa sites. Betancur et al. 776 (F, MO. US); bor¬
der with Bolivar department, near the confluence of the
lte and Tamar rivers into Cimitarra river, ca. 38 km W of
Barrancabermeja, de Bruijn 1537 (MO, US); Mpio. San
Luis, bank of Rio Samand Norte, above Medellin— Bogota
road. Callejas et al. 4104 (MO, US (2)); Mpio. Anorf, Dos
Bocas— Providencia road, near Rio Nechi, Callejas et al.
4466 (HUA, NY); Mpio. Remedios, vereda Santa Lucia.
9-18 km SW of Remedios along road to Puerto Berrio,
Callejas et al. 8106 (HUA. US); Mpio. San Carlos, correg.
Alto de Samana, vereda Miraflores, Callejas et al. 8603
(HUA): Mpio. San Luis, Villa del Sol site, vereda La Cris-
talina on Medellin-Bogotd road, Callejas et al. 11159
(HUA); Mpio. San Francisco, Carretera de Aquitania, na-
cimiento de la quebrada La Cristalina, Cardenas L. et al.
2629 (MO); Mpio. San Carlos, correg. Alto de Samana,
vereda Miraflores, Fonnegra et id. 3018 (HUA, MO, US);
Mpio. San Luis, Rio Samand-Rio Claro sector near Me¬
dellin-Bogotd road, Hernandez el al. 124 (HUA, SEL),
469(4) (HUA), 675 (HUA); Mpio. San Luis, 8 km E of
bridge over Rio Caldera on Medellin road, Juncosa & Es¬
cobar 724 (MO, US); La Cuchilla, correg. El Prodigio, ver¬
eda Los Medios, lAzaro T. 99 (MO); San Luis, near Rio
Claro, Lo aiza & Cogollo 33 (MEI)EL); Mpio. San Luis,
quebrada La Cristalina, Ramirez & Cdrdenas 393 (COL,
HUA), 543 (COL, HUA). 1123 (COL, HUA, MO); Mpio.
Anon, correg. Providencia, Rfo Anorf valley, between Dos
Bocas and Anorf, Soejarto et id. 3283 (MEDEL); Mpio.

266

Novon

Figure I. Cremospermopsis cestroides L. E. Skog & L. I*. kvist. — A. Stem apex. — B. Abaxial leal margin. — C. Corolla
exterior. — I). Opened corolla willi 4 stamens and I staminode at lower left. — E. Infructescence with bracts. — F.
Calyx. — G. Pistil with calyx and nectary. — H. Capsule surrounded by persistent nectary. — I. Seeds. A from Fonnegra
el al. 3018; B. C, I) from Roldan & Betancur 718; E, II. I from Callejas el al. 4104; F. G from Callejas el al. 4390.

Volume 12, Number 2
2002

Skog & Kvist

New Genus from Colombia

267

Anon, Buenos Aires, between Providencia and Alhibe,
Soejarto et al. 4505 (Ml. 1)1’ l.|; Cordillera Central Auto-
pista from Medellin to Bogota, ridge above Bio Cocorna,
Stein et al. 3182 (MO); Mpio. Anori, valley of Rio Anorf
along road 3—5 km upstream from Providencia Hydroelec¬
tric Plant, between Dos Bocas and Anori, Zarucchi 3322
(MO, US). Bolivar: Mpio. Aclii, correg. Ua Raya, refugio
El Paraiso, near Cienega Grande, 3—4 hours from Cau¬
casia above Cauca river, Callejas et al. 4390 (HUA, MO,
US(2)); Mpio. Achi, I.a Raya, Cuadros & Gentry 3587
(US).

Cremospermopsis parviflora L. E. Skog & E. P.
Kvist. sj>. nov. TYPE: Colombia. Antioquia:
Mpio. San Carlos, Embalse Punchina, margin
of streams leading into reservoir near site of
dam, 6°12'N, 74°52'W, 780 m, 18 May 1988,
,/. L. Zarucchi, (). Escobar & A. Ayala 6651
(holotype, HUA; isotypes, MO, US). Figure 2.

C. cestroidis affinis sed foliis plerumque ad apicern ob-
tusibus, tubo corollae a calycis lobis occultis.

Plants herbs, stems mostly erect, occasionally
oppressed to ground, 20— 40(— 60) cm tall, to 5 mm
diani.. glabreseent proximally to tomentose distally;
internodes 0.5— 1.5(— 3) cm long. Leaves with ob-
ovate or occasionally elliptic blades. 8—20 X 4—12
cm. apex obtuse, or less frequently acute, base
acute, margin subentire, occasionally rugose, adax-
ially glabrous, occasionally sparsely pilose, abaxi-
ally sparsely appressed pilose, secondary veins (8
to) 10 to 12 per side, below with appressed pilose
to tomentose indumentum; petioles (0.5 — ) 1 — 2 cm
long, mostly tomentose. Inflorescences congested
cymes but occasionally more open with an extended
axis, of 5 to 12 (to 18) flowers; peduncles 2—9 cm
long, pilose, sometimes dark purple, distally with
mostly 4 bracts with often variable shapes and sizes
in same inflorescence, each ovate to lanceolate,
0.7— 1.2 cm long, mostly tomentose and light green
to whitish; pedicels to 3 mm long. Calyx lobes light
green to yellowish, outside pilose to tomentose, in¬
side glabrous, connate and unequal, narrowly to
broadly ovate, all 4—5 mm long, 3 or 4 of the lobes
ca. 1 mm and the other I or 2 lobes to 3 mm wide.
Corolla nearly hidden by surrounding calyx lobes
and only the limb conspicuous, 5—6 mm long, tube
funnelform to tubular, 3—4 mm long, ca. 1.5 mm
wide at base, narrowing to nearly 1 mm and then
ampliate to nearly 2 mm distally, white, glabrous
both outside and inside; throat ca. 2 mm diam.,
densely glandular hairy with short triehomes, dorsal
lobes often pink, up to I mm long, lateral and ven¬
tral lobe white or yellow, up to 2 mm long. Fila¬
ments adnate to corolla tube base for ca. 2 mm,
above free for 0.5— 1.5 mm; anthers ca. 0.6 X 0.4
mm; nectary ca. 0.8 mm high at both sides of ovary

but in between much lower or absent. Ovary 1 .5
mm high. Capsule 3 X 2 X 1.5 mm.

Distribution and ecology. Plants of Cremosper¬
mopsis parviflora are endemic to Colombia (Antio¬
quia), growing on streamsides in disturbed primary
wet forest or old secondary rain forest at (400—
)500— 900(— 1250) m, and collected in (lower in Jan¬
uary to June, October and December, in fruit in
April, August, and November.

The tiny corollas of Cremospermopsis parviflora
are nearly hidden by the calyx lobes, a character
that distinguishes the species from C. cestroides,
which has corollas approximately twice as large and
not hidden by the calyx lobes. The calyces and
bracts of C. parviflora are apparently also mostly
pale green to white while these may be darker
green to yellow or red in the other species. In ad¬
dition, C. parviflora is a small herb that rarely ex¬
ceeds 50 cm, while C. cestroides can often be more
than 1 m tall, and may be somewhat subshrubby.
Cremospermopsis parviflora also tends to have ob-
ovate leaves with an obtuse apex and a nearly en¬
tire margin contrasting with the mostly oblaneeolate
leaves with an acute apex and a remotely serrate
margin in C. cestroides. The indumentum of the for¬
mer tends toward pilose while the latter has a rather
tomentose indumentum.

Paratypes. COLOMBIA. Antioquia: Lake Punchina,
47.5 km E of San Carlos, Brant el al. 1601 (HUA. MO.
US); Mpio. San Luis, vereda La Josefina, above the Medel¬
lin— Bogota road, 16 km SW of San Luis, Callejas et al. 4186
(NY, US); Mpio. San Luis, veredas La Cristalina & La Jo¬
sefina, Rio Claro region, 98 km SE of Medellin. Callejas &
Acevedo 1 1 167 (HUA); Mpio. San Luis, vereda las Confu-
sas, Cardenas et al. 2670 (MO); Mpio. San Luis, vereda La
Josefina. quebrada La Mariola, Cardenas & Ramirez 2729
(MO); Mpio. San Luis, carretera to Aquitania, 12 km from
Medellin— Bogota road, Cogollo el al. 3756 (MO); Mpio. San
Luis, vereda La Josefina, quebrada La Mariola, Cogollo et
al. 4285 (MO); Mpio. San Eraneiseo, correg. Aquitania, Rio
Venado, Fonnegra et <d. 4183 (MO. US); Mpio. San Luis,
Bio Samand-Rfo Claro sector near Medellin— Bogota road,
Hernandez et al. 191 (HUA. SEL), 211 (HUA, SEL), 280
(HUA); San Luis, Rio Claro, hniiza & Cogollo 42 (MED-
EL); Mpio. Amalfi, NE of Salazar, along road between
Amalfi and Fraguas 23—26.5 km from Amalfi, MaeDougal
et al. 4025 (US); bank of Guatep6 river between San Rafael
and Holanda, Orozco et at. 729 (COL (2)); Mpio. San Carlos,
near ISA hydroelectric dam reservoir, McPherson 13330
(HUA); Mpio. San Luis, quebrada La Cristalina, Ramirez &
Cardenas 232 (HUA, MO), 367 (COL, HUA, MO). 599
I HUA). 1127 (COL, HUA), 1350 (MO), 1517 (COL. HUA),
1922 (HUA. MO); road from Independence to Santa Bita.
Romero-Castaheda 1573 (COL); Mpio. Anorf, Rio Anori
valley near Planta Providencia, Shepherd 456 p.p. (MO),
564 p.p. (WIS); Mpio. Anori, Rio Anori valley between Dos
Bocas and Anori, close to Planta Providencia, SW of Za¬
ragoza, Shepherd 456 p.p. (COL), 561 p.p. (COL, MEDEL);
M) jio. Anori, near Providencia hydroelectric plant, Soejarto
& Villa 2692 ((ill (2)), 2693 (GH (2)); Mpio. Amalfi, mina

268

Novon

Figure 2. Cremospermopsis parviflora E. E. Skog & L. P. Kvist. — A. Stem apex. — B. Stem base. — C. Abaxial leaf
margin. — 1). Corolla exterior. — E. Opened corolla with 4 stamens and 1 staminode at lower left. — E. Infructescenee
with bracts. — G. Calyx. — H. Pistil with calyx and nectary. — I. Capsule surrounded by persistent nectary. — J. Seeds.
A from Ramirez & Cardenas .167', B, I), E, G, II from MacDougal el at, 4025', C from Zarucchi el al. 6651; F from
Soejarto A- Villa 2692; I from Shepherd 564; .1 from Romero- Castaneda 1573.

Volume 12, Number 2
2002

Skog & Kvist

New Genus from Colombia

269

La Vetilla, edge of quebrada Vet ill ita, Tuberquia & Gomez
20 (HUA); Mpio. Anorf, Anorf river valley near Providencia
electric plant ca. 35 km SW of Zaragoza, Waide 62408 (US).

Acknowledgments. We are grateful to Cathy
Pasquale-Johnson for preparing the illustrations, to
the reviewers for their valuable comments, and to
the herbaria that loaned specimens for our study.
Research travel by the second author was support¬
ed by the Carlsberg Foundation of Denmark.

Literature Cited

Kvist, L. P., L. E. Skog & M. Amaya-Marquez. 1998. Los
generos de Gesneriaceas de Colombia. Caldasia 20: 12—
28.

Skog, L. E. & F. de Jesus. 1997. A review of Resin (Ges-
neriaceae). BioLlania Edicibn Esp. 6: 515—525.

- & L. P. Kvist. 2000. Revision of Gasteranthus

(Gesneriaceae). Syst. Rot. Monogr. 59: I -I 18.

Wiehler, H. 1983. A synopsis of the neotropical Gesner¬
iaceae. Selbyana 6: 1—129.

Varietal Status lor a Hirsute Phase of Mirabilis linearis

(Nyctaginaeeae)

Richard Spellenberg

Department of Biology, New Mexico State University, Las Cruces,

New Mexico 88003, U.S.A.
rspellen@nmsu.edu

ABSTRACT. For the upcoming treatment of the ge¬
nus Mirabilis for the Flora oj North America North
of Mexico, M. linearis (Pursh) Heimerl is treated as
having three thoroughly intergrading varieties: M.
linearis var. linearis, M. linearis var. decipiens
(Standley) S. I Welsh, and M. linearis var. suhhis-
pida (Heimerl) Spellenberg, a new combination
published here, based on M. linearis f. subhispida
Heimerl. Mirabilis linearis var. subhispida is com¬
mon on the Great Plains, where it has most often
passed as narrow-leaved phases of M. hirsuta
(Pursh) Macmillan, or as M. gausapoides (Standley)
Standley. A key is provided to distinguish the three
varieties of M. linearis.

Key words: Mirabilis, North America, Nyctagi-
naceae.

While I was preparing a taxonomic treatment for
Mirabilis for volume 4 of the Flora of North Amer¬
ica, I followed the lead of S. L. Welsh (1986) in
considering narrow-leaved plants of Mirabilis with
pubescent fruits that have low tubercles as belong¬
ing within the large and variable complex of forms
encompassed by a broadly delineated M. linearis
(Pursh) Heimerl. Welsh's variety is distinguished in
the key below; another variety, formerly recognized
within Oxybaphus (Dayton, 1959), requires a new
combination in Mirabilis and is also distinguished
by the key.

IVlirahilis linearis (Pursh) Heimerl var. suhhispi-
da (Heimerl) Spellenberg, comb, et stat. nov.
Basionym: Mirabilis linearis (Pursh) Heimerl f.
subhispida Heimerl, Ann. Cons. Jard. Bot. Ge-
n&ve 5: 186. 1901. TYPE: U.S.A. New Mexico:
Lincoln Co., N of Gapitan Mts., 31 Aug. 1900,
F. S. Fade & E. S. Earle 383 (holotype, G-
DEL not seen; isotypes, MO, NMC. US not
seen).

Standley ( 1909: 342) seems to have been the first
to bring in Heimerl’s infraspecific epithet into eas¬
ily accessible North American literature for the
Nyctaginaeeae, presenting as a new combination

without an indication of rank the trinomial “ Allion -
in linearis subhispida (Heimerl) Standley.” He
wrote, “It seems to be a smaller plant, too, and is
probably a good species, but the author has seen
no very good material.” The plant has sparse, hir¬
sute pubescence on the basal parts of the stem. The
name has been rarely used, but within the taxon
herein covered by the infraspecific epithet “ subhis¬
pida ,” density of pubescence and stature vary con¬
siderably. Standley cited, without rank for the epi¬
thet, ‘"'Mirabilis linearis subhispida Heimerl” as the
basionym. although Heimerl had clearly used “for¬
ma” in his protologue. Standley, in Wooton and
Standley (1913: 120), recombined Heimerl’s trino¬
mial as Allionia subhispida (Heimerl) Standley and,
in 1918, Standley considered those plants covered
by the epithet “ subhispida ” at any rank to be part
of A. gausapoides (p. 222), a species he described
in Standley (1911: 406). In none of these publica¬
tions did Standley suggest rank for the epithet when
used iufraspeeifieally.

Dayton (1959) published the combination Oxy¬
baphus linearis var. subhispida (Heimerl) W. Day-
ton, Oxybaphus being a segregate genus of Mira¬
bilis. Reed (1969: 170) incorrectly used “var.” for
the rank of the epithet “ subhispida ” in all infraspe¬
cific combinations in Allionia and Mirabilis, which
were proposed as synonyms of Mirabilis gausapo¬
ides (Standley) Standley in his work. Varietal rank
had never been established for the epithet “ subhis¬
pida 3 at the infraspecific level within Allionia or
Mirabilis.

Mirabilis sect. Oxybaphus, to which this variety
belongs, is notorious for its taxonomic difficulty
(briefly reviewed by Spellenberg, 1998) stemming
from extensive intergradation and variation of all
taxa. Most plants with hispid pubescence on the
Great Plains are often considered to belong to M.
hirsuta (Pursh) Macmillan (in the treatment for
FNA this is a synonym of M. albida (T. Walter)
Heimerl) if leaves are comparatively broad, or M.
gausapoides if leaves are linear. The forms with
narrow leaves and fruits with tubercles not tall and

Novon 12: 270-271. 2002.

Volume 12, Number 2
2002

Spellenberg

Mirabilis linearis (Nyctaginaceae)

271

bract-like are considered all to be M. linearis, ac¬
creted into varieties as indicated in the following
key. Mirabilis gausapoides is considered a synonym
of M. linearis var. subhispida.

Key to the Varieties of Mirabilis linearis

la. Stems hirsute, at least basally .

. M. linearis var. subhispida

lb. Stems minutely puberulent, glabrate, or glabrous
basally.

2a. Leaves linear, grayish or bluish green, glau¬
cous; perianth white to deep rose-pink . . .

. M. linearis var. linearis

2b. Leaves linear-lanceolate to lanceolate,

green; perianth deep pink .

_ M. tin earis var. decipiens (Standley) S. L.

Welsh

Acknowledgments. I am grateful to Ronald
McGregor and Craig Freeman for pointing out this
nomenclatural oversight in their review of the man¬
uscript for Mirabilis for the Flora of North America.

Fred Barrie and George Filz provide helpful com¬
ments in their reviews of this article.

Literature Cited

Dayton, W. 1959. A new varietal combination in Oxyba-
phus. Rhodora 61: 85.

Reed, C. F. 1969. Nyctaginaceae. Pp. 1510-220 in C. L.
1, undell. Flora of Texas, vol. 2, pt. 1. Texas Research
Foundation, Renner.

Spellenberg, R. 1998. Mirabilis melanotricha (Nyctagina¬
ceae), a new combination for a common four o’clock
from southwestern North America. Phytologia 85- 99-
104.

Standley, P. C. 1909. The Allioniaceae of the United
States with notes on Mexican species. Conlr. U.S. Nall
Herb. 12: 303-389.

. 1911. The Allioniaceae of Mexico and Central
America. Conlr. U.S. Nall. Herb. 13: 377-430.

- . 1918. Allioniaceae. N. Artier. FI. 21: 171-254.

Welsh, S. L. 1986. New taxa and combinations in the Utah
flora. Great Basin Naturalist 46: 254—260.

Wooton, E. O. & P. C. Standley. 1913. Descriptions of new
plants preliminary to a report upon the flora of New
Mexico. Conlr. U.S. Natl. Herb. 16: 109-196.

Rubiacearum Americanarum Magna Hama Pars VII. New Species
of Palicourea (Psychotrieae) from Central America and Western

South America

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Gharlotte.Taylor@mobot.org

ABSTRACT. Six new species of Palicourea Aublet
are described and illustrated: P calophlebioid.es
from Costa Rica differs from P calophlebia Stan-
dley of the Andes in part by its purple inflores¬
cences; P ianthina from western Panama differs
from P. chiriquina Standley of the same region in
[tart by its purple inflorescences and longer calyx
limbs; P. premontana from Ecuador dillers from P.
sernirasa Standley of Venezuela and Bolivia in part
by its glabrous corollas; P. quimpiepyrena from
northern Peru is closely related to P. ulloana C. M.
Taylor of Ecuador but differs from that species in
part by its larger, subsessile leaves; P. roseofaucis
from western Panama dilfers from P sulphured
(Ruiz & Pavdn) DC. of the Andes in part by its
white to pink corollas; and P. stellata from east-
central Colombia, with yellow flowers, is unusual
in its well-developed fleshy appendages on the co¬
rolla lobes.

Key words: Neotropics, Palicourea, Psycho¬
trieae, Rubiaceae.

As circumscribed by Taylor (1997), Palicourea
(Psychotrieae) is distinguished within the Rubi-
aceae by its persistent stipules that are interpetiolar
to usually united around the stems into a continu¬
ous sheath and usually bilobed in their interpetiolar
portions; generally brightly colored inflorescen¬
ces — blue, purple, red, orange, or yellow — with the
flowers usually pedicellate; corollas that are usually
similarly brightly colored, five-lobed, and with well-
developed tubes that are somewhat swollen at the
base and glabrous internally except for a dense ring
of pubescence situated just above this basal swell¬
ing; and drupaceous fruits with usually two py¬
renes. This neotropical genus includes about 200
species of shrubs and small trees found from sea
level to montane forests. The species are typically
distylous and generally pollinated by humming¬
birds.

Palicourea calophlebioides C. M. I ay lor. sp.
nov. TYPE: Costa Rica. Litnbn: Canton de Tal-
amanca, Parque Nacional Cordillera de I’ala-
manca, Rfo Lori, 300 m aguas arriba union
Quebrada Luisa, entre Ujarras y San Jose Ca-
becar, 9°21'35"N, 83°13'45"W, 1800 m, 23
Mar. 1993, G. Herrera 5994 (holotype, CR-
5952; isotype, MO-4999475). Figure I A. B.

Haec species a Palicourea calophlebia (ex Aequatoria
et Colombia occidental^ foliorum ampliorum petiolis pler-
umque longioribus, stipulis brevioribus atque inflorescen-
tia corollisque purpureis lilacinisve distinguitur.

Shrubs flowering at 2.5 m tall, to 3 m tall; stems
quadrate and often becoming channeled, tomentel-
lous to glabrescent. Leaves opposite; blades broadly
elliptic to elliptic-obovate, 25—26 X 14—16 cm, at
apex rather abruptly acuminate with tips 5-12 mm
long, at base acute to obtuse, drying papyraceous,
adaxially glabrous, abaxially pilosulous; secondary
veins 26 to 29 pairs, usually looping to interconnect
at least in distal 1/2 of blade, without or with l(to
3) very weak intersecondary vein(s) present be¬
tween paiis of secondary veins, adaxially costa and
secondary veins prominulous and remaining vena¬
tion plane, abaxially costa prominent, secondary
veins prominulous, and reticulated higher-order ve¬
nation plane to thickened; margins thinly to dis¬
tinctly cartilaginous, entire; petioles 2. 5-3. 5 cm
long, tomentellous to velutinous; stipules pilosulous
to tomentellous, persistent at least with leaves,
united around stem into a continuous truncate
sheath 5—8 mm long, lobes 2 on each side, narrowly
triangular to deltate, 4-6 mm long, acute, entire.
Inflorescences terminal, erect; peduncles 2.5—10.5
cm long; panicles narrowly pyramidal, 10—24 X
3.5— 6.5 cm, with 20+ pairs of developed secondary
axes, with flowers pedicellate in eymules of 3 to 7;
bracts entire, those subtending secondary axes nar¬
rowly triangular to lanceolate, 6—20 X 4— i mm,
acute, those subtending pedicels lanceolate to lig-
ulate, 4—7 X 1-2 mm, acute to obtuse; pedicels l—
6 mm long; peduncle, axes, bracts, and pedicels

Novon 12: 272-280. 2002.

Volume 12, Number 2
2002

Taylor

Palicourea

273

Figure 1. A, B, Palicourea calophlebioides C. M. Taylor. —A. Flowering Branch. — B. Flower bud. C-E, Palicourea
lanthina C. M. Taylor. — C. Flowering branch. —I). Corolla. — E. Calyx, pedicel, and floral bract. F, G, Palicourea
roseofaucis C. M. Taylor. — F. Flowering branch. — G. Flower at anthesis. A. C, F to 5-cm scale; B, I). E, G to 5-min
scale. A, B. based on Herrera 5994 ; C. based on Knapp & Sytsrna 2658; 1). E, based on Croat 48509; F, based on
McPherson 9143; G, based on McPherson 9086.

lilac to purple, densely pilosulous except bracts fre¬
quently glabrous; flowers in bud with hypanthium
turbinate, ca. 1 nun long, glabrous; calyx limb gla¬
brous, 2.5—3 mm long, divided nearly to base, lobes

ligulate to lanceolate, acute, entire; corollas tubu¬
lar, purple to lilac, a little swollen at base, generally
straight there and in tube, externally glabrous ex¬
cept hirtellous on lobes, internally glabrous except

274

Novon

for a pilosulous ring ca. 1 mm wide just above basal
swelling, tube (‘a. 10 mm long, ca. 1.5 mm diam.
near middle, lobes deltoid, ca. 2 mm long, acute,
a little thickened adaxially; anthers and stigmas not
seen. Infructescences not seen.

Distribution , habitat , and phenology. In wet
forest at 1800 m, Costa Rica; collected with young
flowers in March.

This new species is similar in overall aspect to
Palicourea calophlebia Standley of western Colom¬
bia and Ecuador, and the specific epithet refers to
this similarity. Palicourea calophlebia differs from
P. calophlebioides by its leaves 14—25 X 5—13.5
cm, its petioles 1.5-2. 5 cm long, its stipules with
the sheaths 10—14 nun long and the lobes 10-15
nun long, its green inflorescences, and its white co¬
rollas that usually turn pale blue with age.

Paratype. COSTA ItlCA. Limon: Canton de Tala-
manca, barque National Cordillera de lalamanea, 300 in
antes tie union Quebrada kuisa con Rio hori, entre Ujar-
rds y San Jose Cabecar. 9°21,35"N, 83°13,45”W, G. Her¬
rera 5952 (MO).

Palicourea iaiitliina C. M. Taylor, sp. nov. I V PE:
Panama. Veraguas: trail on ridge to summit of
Cerro Tute, Cordillera de lute, 1 kin past Es-
euela Agrfcola Altos de Piedras, W of Santa
Fe, 8°36'N, 81°06'W, 1250-1410 m, 15 Dec.
1981, S. Knapp & K. Sytsma 2638 (holotype,
MO-2937075). Figure 1C-E, 3E.

Haec species a Palicourea chiriquina inflorescentia flo-
ribusque purpureis, limbo calycino 2.0— 2.2 mm longo
atque corolla extus dense hirtella distinguitur.

Shrubs flowering at 1 m tall, to 1.5 in tall; stems
quadrate, hirtellous to hirsute or glabrescent.
Leaves opposite; blades elliptic to elliptic-oblong or
ob lanceolate, 6.5-10 X 2. 5-5.2 cm, at apex rather
abruptly acuminate with tips 5—8 nun long, at base
cuneate to obtuse or truncate, drying subcoria-
ceous, adaxially glabrous except sometimes densely
hirtellous along costa, abaxially glabrescent to liir-
tellous, more densely so on secondary and tertiary
venation and sometimes hirsute on costa; secondary
veins 10 to 15 pairs, usually extending to unite with
margins, with l(to 3) sometimes weak intersecon¬
dary vein(s) usually present between pairs of sec¬
ondary veins, adaxially venation plane or costa,
secondary, and tertiary venation all thickened,
abaxially costa and secondary veins prominulous to
prominent and higher-order venation prominulous;
margins rather thickly cartilaginous, entire; petioles
0.4-1. 5 cm long, hirtellous; stipules glabrous to hir¬
tellous, persistent, united around stem into a con¬
tinuous truncate sheath 4-7 nun long, lobes 2 on

each side, narrowly triangular to deltoid, 3—5 mm
long, acute, entire. Inflorescences terminal, erect to
deflexed; peduncles 1.5—7 cm long; panicles pyra¬
midal, 3-1 I X 4-10.5 cm, with 5 to 10 pairs of
developed secondary axes, with flowers pedicellate
in eymules of 3 to 7; bracts entire to ciliolate, those
subtending secondary axes narrowly triangular or
narrowly ligulate, 5-12 mm long, acute to obtuse,
those subtending pedicels narrowly lanceolate, 2—

3 mm long, acute; pedicels 2—3 mm long; peduncle,
axes, bracts, and pedicels lavender to purple, mod¬
erately to densely pilosulous or hirtellous to rather
hirsute; flowers with hypanthium turbinate, ca. I
mm long, densely hirtellous to glabrous; calyx limb
densely hirtellous to glabrous, 2—2.2 mm long, di¬
vided nearly to base, lobes narrowly triangular to
narrowly ligulate, acute to obtuse, entire; corolla
tubular-funnelform, purple to lavender, a little
swollen at base, generally straight there and in
tube, externally densely hirtellous with multiseriate
trichomes, internally glabrous except for a pilosu¬
lous ring ca. 1 mm wide at ca. 2 mm above base,
tube ca. 7 mm long, ca. 1 mm diam. near middle,
lobes triangular, ca. 2 nun long, acute, a little thick¬
ened adaxially; anthers ca. 2 nun long, positioned
in corolla throat; stigmas not seen. Infructescences
similar to inflorescences, apparently purple; im¬
mature fruits narrowly ellipsoid to narrowly ob-
ovoid, ca. 7X4 mm, somewhat flattened laterally,
glabrous to hirtellous, color unknown; pyrenes 2,
planoconvex, dorsallv with 3 to 5 low rounded lon¬
gitudinal ridges.

Distribution, habitat, and phenology. In wet
forest at 1100-2200 in, western Panama; collected
in flower in August, September, November, and De¬
cember. in young fruit in February.

Ibis new species is distinguished by its rather
thickly textured leaves with the secondary veins
uniting with the margins, its stipules united into a
continuous truncate sheath, its purple to lavender
inflorescences and flowers, its relatively well de¬
veloped calyx limbs, and its corollas that are dense¬
ly hirtellous externally. The specific epithet refers
to the inflorescence and flower color. I lie flowers
seen are similar to short-styled flowers of distylous
Palicourea species in tin* arrangement of their an¬
thers and stigmas. Palicourea ianthina is similar to
P. adusta Standley and P. chiriquina Standley. Pal¬
icourea adusta differs from P. ianthina by its leaves
drying papyraceous to chartaeeous, its stipules with
sheaths 1-3 mm long and lobes 1-3 mm long, its
calyx limbs 0.5— 1.5 nun long, and its corollas ex¬
ternally glabrous or sometimes sparsely hirtellous
on the lobes, while P. chiriquina differs from P.

Volume 12, Number 2
2002

Taylor

Palicourea

275

ianthina by its floral bracts 1—1.5 mm long, its ca¬
lyx limbs 0.8—1 mm long, and its externally gla¬
brous yellow corollas.

Paratypes. PANAMA. Chiriquf: Fortuna Dam area, N
of reservoir, ridge along continental divide and S from
Quebrada de Arena, D’Arcy & Todzia 1S937 (MO); Cerro
Pate Macho, along the continental divide, on trail to Finca
Serrano, NF of Boquete, ca. 5 mi. above Boquete, Croat
48509 (MO). Veraguas: N of Santa Fe, summit of Cerro
Arizona, Hummel 4727 (MO, SCZ); above Santa Fe and
above Alto de Piedra, on ridge-trail to top of Cerro Tute-
Arizona, 8°30'N, 81°10'W, McPherson 12077 (MO).

Palicourea premontana C. M. Taylor, sp. nov.
TYPE: Ecuador. Pastaza: along road between
Puyo and Diez de Agosto and Arajuno, 18 km
NF, of main Puyo— Macas road (beginning ,3.7
km from center of Puyo at Hotel Europa), 8.2
km NE of Diez de Agosto, 1°27'S, 77051'W.
970 m, 4 May 1984, T. Croat 59005 (holotype,
MO-3189941). Figures 2D, E, 38.

Haec species a Palicourea semirasa lobulis stipularibus
longioribus, inflorescentia pro ratione angustiore, eorollae
extus glabrae lobulis brevioribus atque fructu rotundato
distinguitur.

Shrubs flowering at 2 m tall, to 6 m tall; stems
terete to somewhat quadrate, glabrous. Leaves op¬
posite; blades elliptic, 15-27 X 5.5-10 cm, at apex
acute to usually acuminate with tips to 15 mm long,
at base cuneate to obtuse, drying papyraceous,
adaxially glabrous, abaxially glabrous or sometimes
puberulous on costa; secondary veins 11 to 15
pairs, not or sometimes weakly looping to intercon¬
nect near margins, often with I (to 3) weak inter¬
secondary vein(s) present between pairs of second¬
ary veins, adaxially costa and secondary veins
prominulous and remaining venation plane, abaxi¬
ally costa prominulous to prominent and secondary
veins and loosely reticulated higher-order venation
prominulous; margins cartilaginous; petioles 1.5-4
cm long, glabrous; stipules glabrous, persistent,
united around stem in a continuous concave sheath
0.5—2 mm long, lobes 2 on each side, ligulate to
ovale, 3—8 mm long, obtuse to rounded, entire to
ciliolate. Inflorescences terminal, erect; peduncles
0—10.5 cm long; panicles narrowly pyramidal to cy¬
lindrical, 10—11 X 4.5—8 cm (excluding corollas),
with 9 to 16 pairs of developed secondary axes,
with flowers pedicellate in cymules of 3 to 7; bracts
entire, lanceolate to triangular, acute to acuminate,
those of secondary axes 0.5-2 mm long and dis¬
placed along these axes to near middle of first in¬
ternode, those subtending pedicels 0.1-0. 8 mm
long and displaced to above base of pedicel; ped¬
icels 1-6 mm long; peduncle, axes, branches.

bracts, and pedicels puberulous, red to pink or or¬
ange; flowers distylous; hypanthium cylindrical to
turbinate, ca. 1 mm long; calyx limb glabrous to
puberulous, ca. 0.8 mm long, sinuate to shallowly
lobed, lobes triangular to semicircular, obtuse to
rounded, entire to ciliolate; corolla tubular-funnel-
form, yellow, a little swollen at base, straight there
and in tube, externally glabrous, internally glabrous
except lor a densely pilose ring ca. 1 mm wide at

1.5- 2 mm above the base, tube 10-14 mm long,

1.5- 3 mm diam. near middle, lobes deltoid, 1-1.5
mm long, acute, smooth adaxially; anthers in short-
styled form ca. 3.5 mm long, included and posi¬
tioned in uppermost part of the corolla tube, in
long-styled form 3.5—4 mm long, included and po¬
sitioned ca. 1/2 ol length of corolla tube above its
base; stigmas 2, in short-styled form ca. I mm long,
included and positioned ca. 1/2 of length of corolla
tube above the base, in long-styled form 1-1.5 nun
long and exserted. Infructescences similar to inflo¬
rescences, becoming red-purple; fruits ovoid, 5 X
4—4.5 mm, rounded laterally, glabrous, purple-
black to black; pyrenes 2, planoconvex, dorsally
with 3 to 5 low rounded longitudinal ridges.

Distribution, habitat, and phenology. In wet
forest of Ecuador and southern Colombia at 600—
1650 m; collected in flower in January, March-July,
September, and November, in fruit in August and
October.

This new species is similar to Palicourea gui-
anensis Aublet, and was included in the circum¬
scription of that species presented by Taylor (1999).
Palicourea premontana generally occupies a higher
elevational range than P. guianensis, and is even
more similar in morphology and habitat to P. se¬
mirasa Standley of Venezuela, northeastern Colom¬
bia, and Bolivia. Palicourea semirasa differs from
P. premontana by its stipule lobes 1.5-2 mm long,
its pyramidal to broadly rounded inflorescences
with the branched portion 4—10 X 5—1 1 cm and
the bracts usually positioned at the bases of the
secondary axes, its externally glabrous corollas with
lobes 2-3 mm long, and its laterally flattened fruits.
Palicourea guianensis differs from P. premontana
by its pyramidal inflorescences 9—19 X 7—15 cm
with the bracts usually positioned at the bases of
the secondary axes and its externally pubescent co¬
rollas with lobes 2-5 mm long. In general both P.
premontana and P. semirasa appear to be closely
related to P. guianensis, and to replace this third
species at middle elevations in the Andes. The spe¬
cific epithet of this new species refers to its ele¬
vational distribution and habitat. Relatively few
well-preserved flowers of this species have been

276

Novon

seen, but these are unusual in Palicourea in the
relatively long stigmas of the long-styled flowers
and the apparently included anthers of the short-
styled flowers.

Paratypes. COLOMBIA. Caqueta: mpio. de Floren-
cia, vereda Las Brisas, carretera Florencia— Suaza, Km 28,
l°36'N, 75°37'W, ./. G. Ramirez el al. 5055 (JAUM, MO).
ECUADOR. Morona— Santiago: 27 km SE of San Juan
Bosco, Gentry et al. 30918 (MO). Napo: cantrtn Jena,

Volume 12, Number 2
2002

Taylor

Palicourea

277

Figure 3. Pyrenes and Iruits of five new species of Palicourea. — A. Cross section of pyrene wall of P stellata ; based
on Lozano 2264. — B. Cross section of pyrene wall of P. premontana; based on Gentry 309 18. — C. Cross section and
partial adaxial view of pyrene wall of P. roseofaucis-, based on McPherson 11657. — L). Apical view of dried fruit of P.
roseofaucis ; based on McPherson 1 1657. — E. Cross section of pyrene wall of P ianthina; based on McPherson 12077.
— F. Cross section of pyrene wall of P. quinquepyrena-, based on Campos 4212. — G. Apical view of dried fruit of P.
quinquepyrena ; based on Campos 4212. All to same 5-mm scale.

Estaciori Biologica Jatun Sacha, 8 km abajo do Puerto
Misahuallf, margen dereeha del Rfo Napo, 1°04'S,
1 7°36' W, A. Alvarez 77 (MO. QCNE), Bensman 79 (MO),
Ceron 1653 (MO, QCNE), Persson et al. 7 (MO), Rueda et
al. 1073 (MO); canton Archidona, comunidad de Pacto
Sumaco, lfmite del Parque Naeional Sumaco, (K)°38'56"S,
77°35'49"W, A. Alvarez et al. I HR I (MO, QCNE); canton
Orellana, Sector Huashito, 20 km al N de Coca, propiedad
de PALMORIENTE, 00°20'S, 77°05'W. Gudino 149 { MO,
QCNE); Chonta Punta at Rfo Napo, vicinity of Santa Rosa,
//. Lugo S. 2140 (US). Pastaza: Mera, along Rfo Allpay-
acu. Hurling A: Andersson 16964 (MO); 2 km N of Shell-
Mera, 1°29'S, 78o03'W. Holm-Wielsen & Jeppesen 369
(MO); Pindo, ea. 6 km E of Mera, II. Lugo S. 23HH (MO);
Parayacu, ca. 10 km E of Canelos, 11. Lugo S. 4562 (MO.
US); El Porvenir, ca. 5 km N of Puyopungu, 11. Lugo 5.
4887 (MO).

Palicourea quinquepyrena C. M. Taylor, sp. nov.
TYPE: Peru. Amazonas: Distrito Bagua, Ara-
mango, Chorro Blanco, 5°29'30"S, 78°20'04"W,
1500-1800 m. I I Feb. 1998, R. 1 if isqiiez & R.
Rojas 25275 (holotype, MO-5333613). Figures
2A-C, 3F, G.

Haec species a Palicourea ulloana foliorum ampliorum
subsessilium brevipetiolatorumve venis secundariis plu-
ribus brochidodromis distinguitur.

Shrubs flowering at 0.8 m tall, to I in tall, ap¬
parently little or not branched, apparently accu¬
mulating litter in leaf bases; stems terete, glabrous.
Leaves opposite; blades elliptic to usually oblan-
ceolate, 14.5—30.5 X 4.5—12 cm, at apex acute to
usually acuminate with tips to 20 mm long, at base
obtuse to truncate or rounded, drying papyraceous,
adaxially glabrous, abaxially puberulous at least on
veins; secondary veins 24 to 28 pairs, spreading
sometimes to more than 90°. usually looping to in¬
terconnect at least weakly, with 1 (to 2) weak inter¬

secondary vein(s) usually present between pairs of
secondary veins, adaxially costa and remaining ve¬
nation prominulous, abaxially costa prominulous to
prominent and secondary veins and reticulated
higher-order venation prominulous; margins thinly
cartilaginous; petioles 3—6 mm long, glabrous; stip¬
ules puberulous, persistent or sometimes fragment¬
ing. united around stem into a continuous truncate
sheath 3-4 mm long, with interpetiolar portion tri¬
angular, lobes 2 to 4 on each side, narrowly trian¬
gular. 3-7 mm long, acute, entire, closely grouped,
inserted below top of sheath at intersection of two
diagonal costae. Inflorescences terminal, apparently
erect; peduncle 14—25 cm long; panicle cylindrical
to narrowly pyramidal, 10—20 X 3.5—14 cm (ex¬
cluding corollas), with 14 to 16 pairs of developed
secondary axes, with flowers pedicellate in cymules
of 3 to 7; bracts entire to ciliolate, narrowly trian¬
gular. acute, those subtending secondary axes 3—6
mm long, those subtending pedicels 1.5—3 mm
long; pedicels 2-8 mm long; peduncle, axes, bracts,
and pedicels densely puberulous, color not noted;
flowers with hypanthium cylindrical to turbinate,
ca. 0.8 mm long; calyx limb glabrous, 0.8— 1.5 mm
long, divided nearly to base, lobes elliptic to nar¬
rowly ligulate, often somewhat unequal on a single
flower, acute to obtuse, entire; corolla slenderly
funnelform, red, swollen and a little bent at base,
straight in tube, externally papillose to puberulous,
internally glabrous except for a sparsely pilosulous
ring ca. 2 mm wide at ca. 1.5 mm above base, tube
ca. 12 mm long, ca. 1.5 mm diam. near middle,
lobes triangular, ca. 2 mm long, acute, smooth
adaxially; anthers ca. 3.5 mm long, positioned ca.
2/3 of length of corolla tube above its base; stigmas

278

Novon

5, ca. 1 mm long, exserted. Infructescences similar
to inflorescences except apparently deflexed to pen¬
dulous, purple; fruits ellipsoid to oblate, ca. 3.5 X

4.5 mm, glabrous, color unknown; pyrenes 5, tri¬
angular in horizontal section, the internal laces pla¬
nar. dorsally smooth, rounded.

Distribution, habitat , and phenology. In wet
forests at 1500-1800 m in northern Peru; collected
in (lower in February, in fruit in July.

This new species is distinguished by its subses-
sile to shortly petiolate leaves with relatively nu¬
merous secondary veins and obtuse to rounded ba¬
se's, its stipules with the lobes lightly grouped and
inserted near the middle of the interpetiolar portion
of the sheath, its relatively long peduncles, its well-
developed calyx lobes, and its fruits with five
smooth pyrenes. The specific epithet refers to these
fruits. This species shares its unusual stipule mor¬
phology and fruits with several pyrenes with Pali-
courea ulloana C. M. Taylor of Ecuador; I*, ulloana
differs from P. quinquepyrena by its leaves 4.5—14
X 1.4—5 cm with acute to cuneate bases and 10 to
12 pairs of secondary veins that extend to unite
with the margins, its stipules with the sheaths I —

2.5 mm long and two lobes 2—3.5 mm long on each
side, its peduncles 3—9 cm long, its inflorescences
with the branched portion 3.5—10.5 X 2-3 cm (ex¬
cluding the corollas), and its fruits with four py¬
renes. The few flowers of P. quinquepyrena seen are
similar to the long-styled form of distylous Palicou-
rea species in the arrangement of their anthers and
stigmas. The leaves of P. quinquepyrena apparently
accumulate litter in their bases, similar to some
other species of Rubiaceae (e.g.. Pentagonia wen-
dlandii Hooker f.) and other families; this has been
called at times a “trash bucket habit.

Paratypes. PERU. Cajamarca: prov. San Ignacio,
(ftto. Uuarango, Nuevo Mundo-Caserfo Gosen. ba Col-
mena, 5°18'30"S, 78°43'00"W, ./. Campos et al. 4212
(MO); Nuevo Mundo, Caserfo GosOi, arriba del margen
izquierdo de la quebrada I, as Juntas, Rodriguez R. &
P. Reyes M. I74R (MO).

Palicourea roseofaucis C. M. Taylor, sp. nov.
TYPE: Panama. Chiriquf: vicinity of Fortuna
Dam, along trail across valley of Quebrada
Hornito, 8°40'04"N, 79°50'04"W, I 150 m, 20
Feb. 1987, G. McPherson 10579 (holotype,
MO-4390379). Figures IF. G, 3C, I).

Haec species a Palicourea sulphured corolla ex alba
rosea suffusa distinguitur, etiarn eae areas geographicas
disjunctas habitant.

Shrubs and trees flowering at 5 m tall, to 12 m
tall; stems quadrate, glabrous. leaves opposite;

blades elliptic, 6-20 X 2-0.5 cm, at apex acute to
acuminate with tips to 10 mm long, at base acute
to cuneate, drying chartaceous to subeoriaceous,
adaxially glabrous and shiny, abaxially glabrous ex¬
cept hirtellous in a line along each side of costa;
secondary veins 8 to 10 pairs, usually extending to
near margins or sometimes looping weakly to in¬
terconnect near margins, without or infrequently
with 1 weak intersecondary vein usually present
between pairs of secondary veins, adaxially costa
and secondary veins prominulous and reticulated
higher-order venation plane to impressed, abaxially
costa prominent, secondary veins prominulous, and
reticulated higher-order venation plane; margins
thinly to distinctly cartilaginous, entire; petioles 1-

2.5 cm long, glabrous; stipules glabrous, persistent,
united around stem in a continuous sheath, inter¬
petiolar portion 5—7 mm long, emarginate to shal¬
lowly bilobed, lobes 1-2 mm long, rounded, entire.
Inflorescences terminal, erect, with peduncles 0.5—

2.5 cm long or sessile and seemingly “tripartite”;
branched portion narrowly pyramidal, 8—16 X 5—9
cm (excluding corollas), with 5 to 6 pairs of devel¬
oped secondary axes, with flowers sessile to shortly
pedicellate in cymules of 3 to 9: bracts entire, those
subtending secondary axes narrowly triangular to
lanceolate, 3—13 mm long, acute, those subtending
pedicels ligulate to triangular, 1-2 mm long, acute;
pedicels 0-1 mm long; peduncle, axes, bracts, and
pedicels strigillose to hirtellous, apparently green;
flowers distylous; hypanthium turbinate, ca. 1 mm
long, strigillose; calyx limb glabrous to strigillose,
ca. 1 mm long, divided nearly to base, lobes tri¬
angular to deltoid, acute to obtuse, entire; corolla
tubular-funnelform, externally white, internally
pink, markedly swollen at base, bent ca. 90° at base
and then again ea. 90° just above basal swelling,
externally puberulous to glabrous, internally gla¬
brous except for a pilose ring ea. 1 mm wide just
above basal swelling, tube ca. 6 mm long, ca. 2.5
mm diam. near middle, lobes triangular, ca. 3 mm
long, triangular, acute, a little thickened adaxially;
anthers in short-styled form 1-1.2 mm long, par¬
tially exserted. in long-styled form ca. 2 mm long,
included and positioned near middle of tube; stig¬
mas 2, in short-styled form ca. 2.5 mm long, in¬
cluded and positioned ea. 2/3 of length of tube
above base, in long-styled form ea. 0.3 mm long
and exserted. Infructescences similar to inflores¬
cences, red-purple; fruit ellipsoid to obovoid, ca. 5
X 5.5 mm, somewhat flattened laterally, glabrous,
blue; pyrenes 2 to 3, planoconvex, dorsally smooth
or with 3 to 5 longitudinal angles.

Distribution and habitat. In wet forest in west-

Volume 12, Number 2
2002

Taylor

Palicourea

279

ern Panama at 1100-1350 m; collected in flower
in February and April, in fruit in June and Septem¬
ber.

This new species is distinguished by its stipules
with the interpetiolar portion emarginate or with at
most short rounded lobes, its rather tough-textured
leaves that are usually shiny on the adaxial surface,
its corollas that are white externally, pink internal¬
ly, and strongly gibbous and bent near the base,
and its dorsally smooth or longitudinally angled py¬
renes. The specific epithet refers to the corolla col¬
or. This new species is similar to Palicourea sul-
phurea (Ruiz & Pavon) DC. of Andean Colombia to
Peru; P. sulphurea differs from P. roseofaucis by its
yellow corollas, and these two species appear to be
allopatric. Palicourea tumidonodosa Dwyer from
the Panama— Colombia border is also similar to both
of these species, but P. tumidonodosa is known only
from specimens with very young inflorescences, on
which several important characters are not evident,
in particular the colors of the inflorescences and
corollas, the mature sizes of the calyx and corolla,
and the size and morphology of the pyrenes. Pali¬
courea tumidonodosa may be nonspecific with P.
roseofaucis or P. sulphurea, or it may represent a
third species, but because none of these possibili¬
ties can be excluded, P tumidonodosa is here pro¬
visionally recognized as distinct.

Paratypes. PANAMA. Koras del Toro: Fortuna Dam
region, along continental divide W of highway pass,
8°45'N, 82°15'W, McPherson 9688 (MO). ( liiriqm: along
road between Gualaca and the Fortuna Dam site, at 8.3
mi. NW of Los Planes de Homito, Antonio 4146 (MO);
Fortuna Dam region, above northern edge of lake, 8°45'N,
82°1 5'W. McPherson 9086 (MO); vicinity of Fortuna Dam,
along trail from highway down to the reservoir, McPherson
9143 (MO); vicinity of Fortuna Dam, along trail from high¬
way across llio Hornito, S of reservoir, 8°45'N, 82° 1 .V W,
McPherson 11657 (MO).

Palicourea slellala C. M. Taylor, sp. nov. TYPE:
Colombia. Santander: mpio. de Gambita, ver-
eda K1 Taladro, carretera Duitama— Charala,
2100-1900 m, 14 Feb. 1983, S. Diaz P 4053
(COL-239056). Figures 2F— I, 3 A.

Haec species a congeneris foliorum venatione utrinque
prominula ac venis intersecundariis bene evolutis, limbo
calycino bene evoluto atque corollae llavae quoque lobulo
appendicem carnosam 3-4 nun longam gerente distingui-
tur.

Shrubs flowering at 1 .5 m tall, to 5 m tall; stems
terete to rather quadrate, hirtellous to glabrous.
leaves opposite; blades elliptic to oblanceolate,
3.5—1 1 .5 X 1—4 cm, at apex acute to usually acu¬
minate with tips to 10 nun long, at base acute to
cuneate, drying chartaeeous, adaxial ly glabrous or

sometimes sparsely hirtellous along sides of costa,
abaxially glabrous or hirtellous along sides of costa;
secondary veins 10 to 23 pairs, reticulating near
margins or usually extending to unite with margins,
with l(to 3) we 1 1 -developed intersecondary vcin(s)
usually present between pairs of secondary veins
and often difficult to distinguish front these, adax-
ially and abaxially costa prominulous to prominent
and secondary veins and strongly reticulated tertia¬
ry venation prominulous; margins thinly to distinct¬
ly cartilaginous, entire to usually ciliolate; petioles

3— 7 mm long, glabrous to hirtellous; stipules gla¬
brous to hirtellous, persistent, interpetiolar, 5—9
mm long, ovate to ligulate, bilobed for 1/3 — 1/2,
lobes narrowly triangular to deflate, rounded to ob¬
tuse or acute, entire to ciliolate. Inflorescences ter¬
minal, erect, seemingly “tripartite” with bracts sub¬
tending the basalrnost pair of axes well developed
and foliaceous; branched portion pyramidal to nar¬
rowly pyramidal, 6—10 X 3-7 cm (excluding co¬
rollas), with 4 to 5 pairs of developed secondary
axes, with flowers pedicellate in cymules of 3 to 7;
bracts entire to ciliolate, those subtending second¬
ary axes narrowly lanceolate to narrowly triangular,

4— 10 mm long, acute, those subtending pedicels
triangular, 1—2 mm long, acute, sometimes dis¬
placed by up to 1/3 ol length ol pedicel above its
base; pedicels 1 .5—7 mm long; peduncle, axes,
bracts, and pedicels glabrous to pilosulous, yellow;
flowers with hypanthium turbinate, ca. 2 mm long;
calyx limb glabrous, 1.2-2 mm long, divided for 1/
3—2/3, lobes triangular to ovate and overlapping on
lateral margins, obtuse to rounded, entire to cilio¬
late; corolla tubular to tubular-funnelform, cream to
usually yellow, a little swollen at base, generally
straight there and in tube, externally glabrous, in¬
ternally glabrous except for a pilose ring ca. 1.5
nun wide at ca. 1 mm above base, tube 7-8 mm
long, ea. 2 mm diam. near middle, lobes triangular,
ca. 2 mm long, acute, adaxial ly with a fleshy, con¬
ical to spatulale, reflexed projection 3—4 mm long,
the projections together forming a star-like shield
on lop of the corolla in bud; anthers ca. 2.5 mm
long, partially exserted; stigmas 2, ca. 2 mm long,
included and positioned ca. 1/2 length of corolla
tube above base. Infructescences similar to inflores¬
cences, becoming red to red-purple; fruit obovoid,
ca. 7X6 mm, somewhat flattened laterally, gla¬
brous, bright blue; pyrenes 2, planoconvex, dorsally
with 3 to 5 longitudinal planar angles.

Distribution, habitat, and phenology. In wet
forests at 1800—2750 m of the Eastern Andean Cor¬
dillera of Colombia; collected in flower in February,

280

Novon

May. June, and October, in fruit in May, June, and
December.

This new species is distinguished by its inter-
petiolar stipules, its leaves with the venation prom-
inulous on both surfaces and the intersecondary
veins well developed, its rather well-developed ca¬
lyx limb, its corollas with unusually well developed
appendages on the lobes, its obovate fruits, and its
pyrenes dorsal ly with planar angles. The corolla
lobe appendages are unusual in Palicourea, anti the
grouped appendages give the corollas in bud a star-
like appearance; the specific epithet refers to this
structure. Similarly well developed corolla lobe ap¬
pendages are found on a few other species of Pal¬
icourea: P. lehmannii (Rusby) Standley differs from
P. stellata by its stipules united around the stem
into a well-developed sheath and its blue to purple
corollas with tubes 24—30 mm long, and is known
ordy from a small region near Cali in the Western
Cordillera of the Andes; P. cornigera C. M. Taylor
differs from P. stellata by its pilosulous pubescence
on both vegetative and reproductive parts, its stip¬
ules that are united around the stem in well-devel¬
oped sheaths, its calyx limbs 3.5—6 mm long, and
its white to blue corollas with tubes 15—20 mm
long, and is known only from Ecuador; P. cornicu-
lata (5 M. Taylor differs from P. stellata by its stip¬
ules united around the stem into continuous
sheaths, its subglobose hypanthia, its corollas with
tubes 10-15 mm long and the lobe appendages
1.5— 2.5 mm long, and its pyrenes with rather sharp
dorsal ridges, and is also known only from Ecuador.
Palicourea stellata is similar also to P. sulphurea
(Ruiz & Pavbn) DC., which is sympatric but differs
by its corollas with unappendaged lobes, the

leaves ol P. stellata are rather unusual in that their
abaxial and adaxial surfaces are similar in color,
texture, prominence of the venation, and especially
in having similar pubescence along the sides of the
costa on both surfaces. The flowers seen of P. stel¬
lata are similar to the short-styled form of distylous
Palicourea species in the arrangement of their an¬
thers and stigmas.

Paratypes. COLOMBIA. Boyaca: mpio. El Encino,
corregimiento de Virolfn, Hacienda La Sierra, Lozano
2264 (COL); mpio. Arcabuco, vereda Piedras Blancas,
Finca La Delicias, Rangel el al. 13175 (COL); mpio. de
G&mbita, vereda El Taladro, J. //. Torres R. 2634 (COL).
Santander: mpio. de Charald, vereda El Taladro, Km 50-
55, carretera Duitama-Virolfn, S. Diaz P. 1613 (COL);
mpio. de Charald, vereda El Volcan, L. I). Moreno & A.
V. Mendez 73 (COL), L 1). Moreno & A. V. Mendez 76
(COL); mpio. de G&mbita, vereda El Taladro. J. H. Torres
R. 2542 (COL); mpio. de Charald. corregimiento de Vi¬
rolfn, sitio Canaverales, J. //. Torres R. 2647 (COL).

Acknowledgments. I thank R. E. Gereau for
preparation of the Latin diagnoses; the curators of
COL, CR, F. I’MA, SCZ. and US for loan of or
information about specimens; R. Foster, T. Wachter,
and the Andrew Mellon Foundation through a grant
to the Field Museum of Natural History for logis¬
tical help and support for travel to this latter insti¬
tution; R. F. Gereau, M. Grayum, I). Lorence, and
C. Ulloa U. for advice and help; and 0. M. Montiel
and R. Magi 1 1 for significant encouragement and
help with this work.

Literature Cited

Taylor, C. M. 1997. Conspectus of the genus Palicourea

(Rubiaceae: Psyefiotrieae) with the description of some

new species from Ecuador and Colombia. Ann. Missouri

But. Card. 84: 224-262.

- . 1999. Palicourea. El. Ecuador 62: 134—235.

Rubiacearum Americanarum Magna Hama Pars VIII. New species
of Gonzalagunia and Sabicea from Mesoamerica and Colombia

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
charlotte.taylor@mobot.org

ABSTRACT. The new species Gonzalagunia osaen-
sis C. M. Taylor, found from central Panama to east¬
ern Costa Rica, is described and illustrated; it has
previously been confused with G. rudis (Standley)
Standley. The new species Sabicea chocoana C. M.
Taylor, found from eastern Panama to western Co¬
lombia, is also described and illustrated; it has pre¬
viously been confused with S. panamensis Wern-
ham.

Key words: Colombia, Gonzalagunia, Mesoam¬
erica, Rubiaceae, Sabicea.

The new species of Gonzalagunia Ruiz & Pavon
and Sabicea Aublel described below were discov¬
ered during preparation of tbe Rubiaceae treatment
for the Flora Mesoamericana.

Gonzalagunia (Isertieae; Robbreeht. 1993) com¬
prises about 35 (Stahl, 1999) neotropical species
distributed from central Mexico and the Antilles to
Bolivia and southern Brazil. This genus is charac¬
terized by its usually shrubby habit; its triangular
interpetiolar stipules that are frequently persistent;
its terminal, spiciform, bracteate inflorescences
with the flowers borne singly or in congested cy-
mules directly from the well-developed primary
axis; its calyx lobes four; its salverform pink or
white corollas with the lobes four and imbricated
(or reportedly infrequently valvate: Stahl, 1999);
and its fleshy fruits with the numerous small angled
seeds borne loose and dry inside two or four bony
pyrenes. The long slender inflorescences and in-
fructescences are often pendulous, giving the plants
a distinctive aspect. The flowers are generally fra¬
grant and distylous. The fruits at maturity are typ¬
ically mealy or spongy rather than juicy. The num¬
ber of pyrenes in the fully developed fruits, two vs.
four, has been considered taxonomically informa¬
tive (e.g., Stahl, 1999). Species of Gonzalagunia are
commonly found in secondary vegetation and pro¬
duce flowers and fruits for relatively long periods,
and are therefore commonly collected. The genus
has not been studied comprehensively.

Sabicea comprises about 120—130 neotropical
and African species (Andersson, 1999). This genus

has often been included in the tribe Isertieae (e.g.,
Robbreeht, 1993), although it has also been treated
in the tribe Sabiceeae when this is separated from
Isertieae (Bremer & Thulin, 1998). In the Neotrop¬
ics Sabicea is distributed from central Mexico and
the Antilles to Bolivia and southern Brazil. This
genus is characterized in the Neotropics by its usu¬
ally twining habit; its interpetiolar stipules that are
usually ovate to elliptic, persistent, and reflexed;
its axillary, bracteate, capitate to cymose inflores¬
cences; its calyx lobes five; its salverform to fun-
nelform, usually white corollas with five valvate
lobes; its stigmas and ovary locules five; and its
fleshy fruits with numerous small angled seeds dis¬
tributed through the pulp. The fruits are juicy and
usually pass through pink and red stages to finally
become red-purple to violet-black at maturity. Spe¬
cies of Sabicea are commonly found in secondary
vegetation, and produce flowers and fruits often and
for relatively long periods. Some South American
species are low pyrrhophytes in savanna and cer-
rado vegetation (e.g., S. brasiliensis Wernham, S.
humilis S. Moore), and a few apparently may grow
as self-supporting shrubs (e.g., S. camporum
Sprague). Many species of Sabicea bear white or
tan arachnoid pubescence; the presence and dis¬
tribution of this pubescence on the plant has been
considered taxonomically informative (e.g., Steyer-
mark, 1967) although its variation in living popu¬
lations has apparently not been studied. Sabicea
was monographed by Wernham (1914), but this
work is now outdated.

Gonzalagunia osaensis C. M. Taylor, sp. nov.
TYPE: Costa Rica. Puntarenas: Reserva For-
estal Golfo Dulce, Aguabuena, Rincon de Osa,
8°42'N, 83°32'W, 50-150 m, 10 Aug. 1991,
R. Aguilar 259 (holotype, CR; isotype, MO-
4234234). Figure 1C, D.

Haec species a Gonzalagunia dicocca lobulis calycinis
linearis 1.5—3 mm longis atque lobulis corollinis acutis
3.5—6 mm longis distinguitur.

Shrubs or small trees to 4 m tall, with branches
often slender and arching; stems densely strigillose.

Novon 12: 281-285. 2002.

282

Novon

Figure I. A, B, Sabicea chocoarui C. M. Taylor. — A. Flowering branch; based on Ledn 51 6. — B. Detail of inflores¬
cence showing flower and young fruit; based on Gentry X: Aguirre 1523 6. C, I), Gonzalagunia osaensis C. M. Taylor.
— (i. Flowering branch; based on Cuadros 8050. — I). Flower; based on Grayum 8634. A. C to 5 -cm scale; B, I) to 5-
mm scale.

Volume 12, Number 2
2002

Taylor

Gonzalagunia & Sabicea

283

Leaves lanceolate, 3—10 X 1-2.5 cm. at apex ta¬
pered and acuminate, at base cuneate to obtuse,
drying papyraceous, adaxially sparsely strigillose to
glabrescent on lamina and densely strigillose on
costa and secondary veins, abaxially sparsely to
moderately strigillose at least on veins; secondary
veins 4 to 6 pairs, usually at least weakly looping
to interconnect, adaxially costa thinly prominulous
and remaining venation plane or secondary veins
sometimes a little thickened, abaxially costa and
secondary veins prominulous and remaining vena¬
tion plane, without domatia; petioles 1—4 mm long,
densely strigillose; stipules interpetiolar, narrowly
triangular to subulate, 3.5-7 mm long, long-acute,
persistent, adaxially densely strigillose, abaxially
densely strigillose on basal part and midrib and
glabrescent on margins of upper part. Inflorescences
terminal on rather short lateral branches (i.e., these
with 1 to 2 or occasionally up to 4 leaf-bearing
nodes), densely strigillose, arching to pendulous,
spiciform with primary axis prolonged and flowers
borne singly directly from it; peduncles 0.5—1 cm
long, slender; flowering portion 2.5—8 X I cm;
bracts narrowly triangular to linear, 1—4 mm long,
acute, borne in clusters of 2 to 3 at bases of ped¬
icels or sometimes I to 2 bracts basal to pedicel
and I to 2 bracts borne along its length; pedicels
1—1.8 mm long; flowers distylous; hypanthium el¬
lipsoid, ca. 1 mm long, densely strigillose; calyx
limb 1.8—2 mm long, densely strigillose, lobed for
2/3— 3/4, lobes 4, linear, sometimes somewhat un¬
equal (i.e., with less than 30% difference in length);
corolla salverform, white, externally densely seri¬
ceous to strigillose, internally glabrous except
puberulous to hirtellous on bases of lobes, tube 7—
9 mm long, lobes 4, narrowly triangular, 3.5—6 mm
long, acute; anthers subsessile, narrowly oblong, in¬
cluded. in short-styled form ca. 1.5 mm long and
situated ca. 2/3 ol length of corolla tube above its
base, in long-styled form ca. 1.2 mm long and sit¬
uated just below middle of corolla tube; stigmas 2.
linear, in short-styled form situated ca. 1/3 of length
of corolla tube above its base and included, in long-
styled form ca. 1.2 mm long and situated in corolla
throat, included or with tips exserted. Fruits when
dry ellipsoid, ca. 3 X 3.5 mm, laterally flattened,
somewhat didymous, strigillose, white, at maturity
quickly becoming inflated; pyrenes 2, subglobose.

Habitat , distribution , and phenology. In wet
forest at 0—650 m, southeastern Costa Rica to west¬
ern Panama; collected in flower May through Oc¬
tober, in fruit August through February.

This new species is distinguished by its rather
low slender habit, its relatively small lanceolate

leaves, its rather short inflorescences with the flow¬
ers borne directly and singly from the primary axis,
its linear calyx lobes 1.5—3 mm long, its acute co¬
rolla lobes 3.5—6 mm long, and its fruits with 2
pyrenes. The species epithet refers to its geographic
distribution. This new species is similar to Gonza¬
lagunia dicocca Chamisso & Schleehtendal, found
in northern to central South America, in general
aspect and in its fruits with 2 pyrenes; but G. di¬
cocca differs from G. osaensis in its calyx lobes 0.2—
I mm long and its corolla lobes 3—3.5 mm long.
Gonzalagunia osaensis has also been previously
confused with G. rudis (Standley) Standley (Dwyer,
1980; Burger & Taylor, 1993), but G. rudis differs
from G. osaensis in its habitat, in usually seasonal
or moist forest; its densely sericeous hypanthium;
its corollas with tubes ca. 3.5 mm long and lobes
ca. 1.5 mm long; and its fruits with 4 pyrenes. Gon¬
zalagunia osaensis appears to be locally common,
particularly in the Osa and Burica Peninsulas, and
seasonal in its flowering and fruiting.

Faratypes. COSTA MICA. Puntarenas: canton de
Osa, Reserva Forestal Golfo Dufce, Aguilar 1216 (INB,
MO); canton de Osa, Parque Nacional Corcovado, Penin¬
sula de Osa, Estacion San Pedrillo, Aguilar 2350 (MO);
canton de Golfito, Parque Nacional Corcovado, Peninsula
de Osa, Estacion Sirena, Aguilar 2450 (IN B, MO), Aguilar
el (d. 3539 (INB, MO); canton de Golfito, Peninsula de
Osa, El Tigre, cabecera del Rio Agujas, Finca Azofeifa,
Aguilar 2654 (INB, MO); canton de Osa, Rincon de Osa,
lila Casa Loma, sendero Eas Ranas, A. Cliacon 911 (CR,
MO); ridge between Quebrada Aguabuena and Quebrada
Banegas, ca. 5 km W of Rincon de Osa, Grayum 4055
(MO); ridge between (Quebrada Banegas and Rfo Riyito,
ca. 7 km W of Rincon de Osa, Grayum 4092 (MO); Re¬
serva Forestal Golfo Dulce, Osa Peninsula, Rancho Que-
mado, ca. 15 km W of Rincbn, Hammel el al. 16814 (CR,
MO). Hammel et al. 17003 (CR, MO); canton de Osa,
Reserva Forestal Golfo Dulce, Peninsula de Osa, Playa
Campanario o San Josecito, Sierpe, Harmon 296 (INI?,
MO); 12 km E of Golfito, Hannon & Fuentes 6130 (MO);
E of Sirena headquarters, Corcovado National Park, Haris-
horn 1866 (MO); canton de Osa, Rancho Quemado, Sier¬
pe, cabeeeras del Rfo Drake, G. Herrera 4226 (CR, MO);
Corcovado National Park, Osa Peninsula, near Park head¬
quarters at Sirena, Janzen 11031 (MO); Parque Nacional
Corcovado, upper Camaronal, Kernan & Phillips 595
(MO), Ollas trail. Kernan et al. 875 (CR, MO); ridge N of
airport, Rincon de Osa, Liesner 1960 (MO); Corcovado Na¬
tional Park, 0—2 km W of park headquarters at Sirena,
Liesner 2914 (MO); canton de Osa, Reserva Forestal Golfo
Dulce, Peninsula de Osa. Rincon de Osa, Lobo 79 (INB,
MO); canton de Osa, Reserva Forestal Golfo Dulce, Ran¬
cho Quemado, Fila Guerra, Rincon, Finca G. Padilla, A.
L. Alari'n A- Gutierrez 56 (INB, MO); canton de Osa, Ran¬
cho Quemado, Fila Division, ./. Marin 24 (CR, MO); can¬
ton de Osa, tierras del Grupo de Conservaci6n de Rancho
Quemado, Sierpe, ./. Marin 221 (INB, MO); canton de
Osa, Agua Buena, Rincon [de OsaJ, al N de BOSCOSA,
Morales 113 (CR, MO); canton de Osa, camino a la toma
de agua. Rancho Quemado, Rincbn [de Osa], Quesada

284

Novon

244 (INB, MO): Canton de Osa, Reserve Forestal Golfo
Dulce, Peninsula de Osa, Rancho Quemado, Quebrada El
Brole, Quesada & Segura 762 (INB, MO); along eamino
de Altura, 2-5 mi. W of Rinebn |de OsaJ, trail to airfield
from Mile 4, Haven 21666 (MO); Parque Nacional Cor-
covado, Estaeidn Sirena, Saborlo 80 (INB. MO); Ea Gam-
ha, Parque Nacional Espinas, Will 26/07/66/71 (MO): Re¬
fugio de Fauna Silvestre Golfito, Zamora 1056 (CR, MO);
Reserva Nacional de Vida Silvestre Golfito, F i 1 a Gamba,
eamino entre Golfito y Villa Briceiio, Zamora et al. 1616
(INB. MO). I’unlarenas— San Jose Border: Tarrazu, fa 1 -
das del Cerro Nara, ca. Esquipulas, lnnite Quepos y Tar¬
razu, Gomez- Laurito el al. 1 1576 (CR. MO). Sail Jose:
Zona Protectora Ea Cangreja, along Quebrada Grande and
on adjacent ridges, ea. 2 km NNE of Mastatal de Puriscal,
Grayum 8624 (CR, MO). PANAMA. Cliiriqui: Burica
Peninsula, San Bartolo lfmite, 21 km WNW of Puerto Ar-
muelles, Busey 456 (MO); Rurica Peninsula, Quebrada
Mdrida, 4 mi. S of Puerto Armuelles, Liesner 276 (MO).
II. ■rrera: above Chepo de Eas Minas, Folsom el al. 6680
(MO); distrito de Eas Minas, Chepo, Galdames et al. 1658
(MO); between Eas Minas and el Toro, near village of Che¬
la), McPherson 10202 (MO). Los Santos: vicinity ol head¬
waters of Rio Pedregal, 25 mi. SW of Tonosf, Lewis el al.
2622 (MO); valley of Rfo Guanico, McPherson 6227 (MO).
Los Santos— Veragu as Border: southern Azuero Pen¬
insula, near proposed route of road from El Cortezo to
Arenas, Azuero, ca. 10 km SW of El Cortezo, Hammel
5412 (MO). Veraguas: 18 km W of Eas Minas, N slope
of Cerro Alto Higo, Hammel 4246 (MO); Isla de Coiba,
Playa Hermosa, Cuadros et al. 8050 (MA, MO); Isla Coi¬
ba, distrito de Montijo, Playa Brava, Galdames et al. 2244
(MO).

Sabieea chocoana C. IVI. Taylor, sp. nov. TYPE:
Colombia. Antioquia— Chocd border: zona de
Urabfi, Cerro El Cuchillo, Finea Cidon a la
Cumbre, 20-100 m, 9 Oct. 1987, I). Cardenas
584 (holotype, JAUM; isotype, MO-3822736).
Figure 1A, B.

Haec species a congeneris infloreseentiae pedunculatae
ramosae pedieellis 2—10 mm longis, praeter corollam om¬
nibus partibus vegetativis reproductivisque pubescentia
pilosula breve uniformi vestita, lobulis calycinis 1.2— 2.2
mm longis atque corolla pubescentia hirsula patente ves¬
tita distinguitur.

Suffrutescent twiners, climbing to 2 m high;
stems densely uniformly pilosulous. Leaves elliptic-
oblong to broadly so, 5.5-1 1 X 4—8 cm, at apex
obtuse to usually gently acuminate, at base cuneate
to truncate, drying papyraceous, adaxially moder¬
ately strigillose on lamina and densely pilosulous
to occasionally hirtellous on costa and secondary
veins, abaxially moderately to densely strigillose to
usually pilosulous with usually denser and always
spreading pubescence on veins; secondary veins 7
to 1 1 pairs, weakly looping to interconnect, without
domatia; petioles 3—15 mm long; stipules ligulate,
3—4 mm long, obtuse to rounded, persistent, re¬
flexed, adaxially strigillose to glabrescent, abaxially
densely pilosulous. Inflorescences 2—1 cm long.

densely pilosulous, paniculate with I to 2 pairs of
secondary axes; peduncles 5—22 mm long; hracts
subtending secondary axes narrowly ligulate, 1.5—
5 mm long, obtuse to acute, bracts subtending ped¬
icels and flowers narrowly triangular. 0.3— 1.5 mm
long, acute; pedicels 2-10 mm long; flowers with
hypanthium ellipsoid, ca. 2 mm long, densely pi¬
losulous; calyx limb adaxially strigillose to glahres-
cent, abaxially densely pilosulous, with tube ca. 1
mm long, lobes 5, ligulate to narrowly so, 1.2— 2.2
mm long, obtuse to acute; corolla salverform, w hite,
externally densely hirsute, internally not seen, tube
8-9 mm long, lobes 5, narrowly triangular. 2-3 mm
long, acute; anthers not seen; style ca. 6 mm long,
stigmas ca. 1.2 mm long; disk annular, ca. 1 mm
high. Fruits subglobose to ellipsoid, 8—9 mm diam.,
deep purple to black; seeds angled, ca. 0.3 mm
long, finely striate.

Habitat, distribution, and phenology. In wet
forest at 10—600 m, eastern Panama to northwest¬
ern and central western Colombia; collected in
flower May through July and in October, in fruit in
May. July, and September through November.

Ibis new species is distinguished by the com¬
bination of its pedunculate branched inflorescences
with pedicels 2—10 mm long; vegetative organs, in¬
florescence axes, and hypanthia that are pilosulous
with uniform spreading trichomes; strongly reflexed
calyx lobes 1 .2—2.2 mm long; and externally hirsute
corollas. The specific epithet refers to its geograph¬
ic range, which is centered in the Choco biogeo¬
graphic region of western Colombia and adjacent
eastern Panama. Only one specimen ( H aught 2210)
has mature flowers in condition adequate for study;
whether this new7 species is distylous or homosty-
lous cannot be determined. This new species is
similar to the sympatric species Sabicea panamen-
sis Wernham; 5. panamensis differs from .S', cho¬
coana in its appressed, usually strigose or strigil¬
lose pubescence on all parts and its acute calyx
lobes 2—6 mm long. This new species is also similar
to S. thyrsiflora L. Andersson of northwestern Ec¬
uador; S. thyrsiflora differs from S. chocoana in its
appressed to only weakly spreading pubescence,
longer stipules (6.5-7 mm long), longer bracts (5-
7 mm long), and longer calyx lobes (2.5— 2.7 mm
long) that are ascending in fruit.

Paratypes. COLOMBIA. Antioquia: mpio. Turbo,
carretera Tapdn del Daridn, sector Rfo Ledn-Lomas Ais-
ladas. Km 37. Brand & Gonzalez 680 (JAUM. MO); mpio.
Caucasia, along road to Neehf ca. 19 km E from Cauca-
sia-Planeta Rica road, Brant & Escobar 1185 (HUA. MO).
Bolivar: Palotal, R. Romero-Castaneda 1172 (F). Choco:
mpio. Riosucio, zona de Urabd, Cerro del Cuchillo, D.
Cardenas 787 (JAUM, MO), cerros del Cuchillo, sector

Volume 12, Number 2
2002

Taylor

Gonzalagunia & Sabicea

285

NO, I). Cardenas 254 (JAUM, MO), sector Cuehillo Negro.
D. Cardenas 363 (JAUM. MO), camino entre Cuehillo Ne¬
gro y Punta de las Flores, I). Cardenas 1944 (JAUM, MO),
Cuehillo Negro (Rio Marfa) a cumbre NO, D. Cardenas
2240 (JAUM. MO); 2—1 km NW of Teres ita. Duke 1 1039
(MO), Duke 11054 (MO); trail from Rfo Tigre base camp
up Serranfa del Darien W of Unguia, Gentry & Aguirre
15236 (MO); mpio. de Riosucio, Parque Natural National
Los katyos, Salto de Tilupo, Tebn 137 (COL, MO), camino
Tilupo. Alto via Sautata, desviando por el camino a Tilupo
Salto parte baja. Le6n 516 (COL, MO). Santander: Mad-
galena valley, Campo Capote, 30 km E of Carare, Gentry
et al. 20079 (MO); vicinity of Barranca Bermeja (El Cen¬
tro), Haughl 2210 (E, MO). PANAMA. Darien: 0.5-1. 5
mi. E of Manene, Hartman 12104 (MO); trail NW of Cana,
Sullivan 715 (MO).

Acknowledgments. 1 thank the curators of CR.
F, HUA, JAUM. PMA, and US for access to spec¬
imens and information; Roy E. Gereau for prepa¬
ration of the Latin diagnosis; and Tyana Wachter,
Robin Foster, and the Andrew Mellon Foundation

through a grant to the Field Museum of Natural
History for support for travel to that institution.

Literature Cited

Andersson, L. 1999. Sabicea. El. Ecuador 62: 101-114.
Bremer, B. & M. Thulin. 1998. Collapse of Isertieae, re¬
establishment of Mussaendeae, and a new genus of Sa-
biceeae (Rubiaceae); Phylogenetic relationships based
on rbcL data. PI. Syst. Evol. 21 I: 71—92.

Burger, W. & C. M. Taylor. 1993. Flora Costaricensis,
Family #202 Rubiaceae. Fieldiana, Bot. n.s. 33: 1—333.
Dwyer, J. I). 1980. Rubiaceae. In: R. E. Woodson, Jr., R.
W. Schery & Collaborators, Flora of Panama — Part IX.
Ann. Missouri Bot. Card. 67: 1-522.

Robbrecht, E. 1993. Genera Rubiacearum. Opera Bot.
Belg. 6: 1 73-196.

St&hl. B. 1999. Gonzalagunia. FI. Ecuador 62: 70—101.
Steyermark. J. A. 1967. Sabicea. In: B. Maguire & Col¬
laborators, The Botany of the Guyana Highland — Part
VII. Mem. New York Bot. Gard. 17: 307—316.

Werham, H. F. 1914. A Monograph of the Genus Sabicea.
British Museum, London.

Typhonium jinpingense, a New Species from Yunnan, China, with
the I .owest Diploid Chromosome Number in Araceae

Wang Zhonglang, Li Heng and Biati Fuhua
Kunming Institute of Botany, Academia Sinica, Kunming 650204, Yunnan, China

ABSTRACT. Typhonium jinpingense Z. I,. Wang. II.
Li & F. H. Ilian (Araceae) from Yunnan, China, is
described as a new species in Araceae. The kar¬
yotype ol metaphase chromosomes in somatic cells
is: 2 n = 10 = 2m + 2st + 6sm. This is the lowest
diploid number so far reported in this family.

Key words: Araceae, chromosome numbers, Ty¬
phonium.

I* ieldwork was conducted in the southeast part
of Yunnan Province, China, in October 1009. The
new species was found on a slope near a small
stream with an elevation from 1000 to 1550 m in
Jinping County, Yunnan Province, China. This
plant was first collected and identified as Typhon¬
ium hlumei Nieolson Sivadasan, as the leaves
were (juite similar to this species. The plants were
subsequently grown and propagated; so far, more
than 20 individuals are cultivated in the Kunming
Botanical Garden. From April to August 2000 four
of them flowered, displaying quite different char¬
acteristics from T. hlumei. After checking the chro¬
mosome number for several individuals, we found
the lowest diploid chromosome number in Araceae
reported to date. We checked tin' morphological
characteristics carefully and compared them to all
the species in the genus Typhonium , and realized
this taxon is an undescribed species.

Act ■ording to the revision of Typhonium (Sri -
boonma el al., 1004) and our recent investigations,
16 species and 4 varieties in the genus have been
found in China thus far, belonging to 5 sections.
Recently, Hetterseheid and Boyce (2000) merged
the genus Sauromatum Schott (all 3 species found
in China) with Typhonium Schott (Wang & Li.
1009). Thus the number of Typhonium species in
China is now thought to be 10. However, based on
species richness in neighboring countries (Hetter-
scheid et al., 2001; Hetterseheid & Nguyen. 2001 ),
this is likely to be an underestimate of the actual
number of Typhonium species in China, especially
in the southern part ol Yunnan Province.

Materials and Methods

file plants were collected in the field from Jinp¬
ing, Yunnan, China, and cultivated in the Kunming
Botanical Garden for the taxonomic and cytological
study.

For the taxonomic study, we first cheeked the
characteristics carefully, using the key for Typhon¬
ium developed by Sriboonma et al. (1904) to de¬
termine which section the species belongs to. We
then compared it to all described Typhonium spe¬
cies, including the species published subsequent to
the 1994 revision.

for the cytological study, growing root-tips were
pre-treated in 0.002 mol/l. 8-hydroxyquinoline at
room temperature for 3 hours, then fixed in Clarke
(ethanol/acetic acid, 3:1) solution at 4°C for 30
minutes. They were then hydrolyzed in 1 mol/L hy¬
drochloric acid al 60°C for 3—4 minutes. After be¬
ing rinsed in distilled water 3 times, the root lips
were dyed in Carbol f uchsin solution for 1 hour,
then squashed and sealed. Photos were taken using
the method described in Li and Zhang (1001). flu*
chromosomes were classified by arm ratio accord¬
ing to Leven and Fredga (1964). The karyotype
classification followed the criteria developed by
Stebbins (1071).

Resi its

CYTOl.OiaCAI. STUDY

The chromosome number was counted in more
than 15 somatic cells, and the karyotype analysis
was carried out in 1 good cell. The karyotype of
the new species is described in Table I and illus¬
trated in figure 2. The chromosome number in so¬
matic cells is 10, and the karyotypical formula is
2/i = 2x = 10 = 2m+ 2st+ 6sm. This species is a
diploid. I he length in somatic mitotic cells ranges
from 2.0 gm to 6.1 gim, while the relative length
of tin' chromosomes ranges from 6.5 to 13.2. The
ratio of the longest chromosome (No. 1) to the short¬
est one (No. 10) is 2.04. flit' percentage with Long
Arm/Short Arm >2:1 is 80%. The karyotypical
asymmetry belongs to the 3B type.

Novon 12: 286-289. 2002.

Volume 12, Number 2
2002

Wang et al.

Typhonium jinpingense from Yunnan

287

Table 1. The parameters of somatic chromosomes in
mitotic nietaphase of Typhonium jinpingense.

Pair

TL

AR

PC

1

13.3

2.0

sm

2

12.7

2.6

sm

3

10.1

2.4

sm

4

7.7

3.3

St

5

6.6

1.4

111

Note: TL = Total relative length; AH = arm ratio; PC
— position of centromere.

TAXONOMIC DESCRIPTION

Typhonium jinpingense /. L. Wang, H. I.i & F.
H. Bian. sp. nov. TYPE: China. Yunnan Prov¬
ince: Jinping Xian, Shuangjin Bridge, road
side by stream, 1200 m, 30 Oct. 1999, F. //.
Bian & R. Li 200 (holotype, KUN). | Cultivated
in Kunming Botanical Garden, 2 n = 10.] Fig¬
ure 1 .

Haee species Typhonio blumei affinis, sed spathae tubo
ovato, 1.5 cm longo, 5 mm diam., pallide viridi inferne,
atropurpureo superne. lamina angusta, lanceoiata, 1 I cm
longa, medio 1 cm lata, basi atropurpurea, cretacea supra
basin, florum sterilium rudimentis clavatis, chromosoma-
tibus 2/i = 10 differ!.

Perennial herb, seasonally dormant, rhizome tu¬
berous, irregularly cylindrical, brown, horizontal or
erect, producing small gemmules and many filiform
roots, 2-2.5 cm long, 0.7— 1.0 cm diam. Leaves 1—
3, petiole smooth, green, 10—17 cm long. 0.1 cm
diam., sheathing below ca. 1.5—2 cm long; blade
chartaceous, green, cordate, usually cordate-hastate
or deeply cordate at base. 5—8 X 4—10 cm. Pedun¬
cle about 1 cm long. Spathe: tube ovoid, pale green,
dark purple at the top, 1.5 cm long, 0.5 cm diam.;
the blade narrowly lanceolate, chalky-white but
dark purple at base, 1 1 cm long, 1 cm wide below
the middle, spreading and slightly arching, the
apex long-acuminate but not twisted. Spadix ca. 14
cm long, pistillate portion ca. 0.5 cm long, green¬
ish, the style short, stigma dark purple, sterile por¬
tion ca. 0.4 cm long, densely covered proximal ly,
erect, yellowish, clavate rudiments, with naked in¬
terstice to 2 cm long; staminate portion to 0.5 cm
long, densely congested, yellow; terminal appendix
cylindrical, erect, purple, ea. 10 cm long, 2 mm
diam., sessile and not swollen at base, with very
strong odor. Flowering from the end of April to Au¬
gust.

Distribution. Known only from Jinping County,
southeastern Yunnan, China, in wet fields and road¬
sides by streams, altitude 1000—1550 m.

Discussion. Typhonium is commonly accepted
as comprising five sections as described by Sri-
boonma et al. (1994). This new species belongs to
section Typhonium. From the shape of the leaf
blade, this new species is easily differentiated from
the other Chinese Typhonium species, except T.
blumei. It is distinguished from T. blumei in having
the spathe tube pale green, dark purple at the top
(not entirely dark purple), the blade narrowly lan¬
ceolate (not broadly ovate), dark purple, spreading
and arching, apex long acuminate but not twisted,
clavate sterile rudiments, erect, and in its chro¬
mosome number. 2 n = 10.

Based on recent comprehensive reviews of the
cytology of the Araceae (Grayum, 1990; Mayo et
al.. 1997; Petersen, 1989. 1994), the chromosome
numbers vary greatly between and also within gen¬
era, from 2/i = 14 ( Ulearum ) to 2 n = 168 ( Arisae -
ma). The lowest diploid number so far known in
Araceae is 2 n — 14, observed in Ulearum sagit-
tatum Engler; based on our results, Typhonium jin¬
pingense ( 2n = 10) has the lowest chromosome
number known to occur in the family. The chro¬
mosomes were easy to group into five pairs (Fig. 2),
so Typhonium jinpingense is a diploid species and
the basic chromosome number is 5. Other species
of Typhonium have chromosome numbers of: 2n =
16. 18. 20, 26, 36, 52, 54, 65, >100 (Mayo at al.,
1997), making it one of the most variable genera
in this regard in Araceae.

Primary basic chromosome number is a main
area of research in cytological studies. The discov¬
ery of the lowest diploid number in this study may
spur further investigation ol this aspect. Possibly
the following questions will be proposed: Was it
caused by extreme aneuploid reductions? Or is 2 n
= 10 some kind of “ancestral number” only kept
in Typhonium? We think the former is more likely.
It is well known that very low chromosome numbers
(as well as very high ones) are more common in
advanced genera than in basal ones. Petersen
(1994) postulated that very low chromosome num¬
bers were explained as aneuploid reductions, from
an ancestral number x — 12 or 14. Other authors
proposed that there was an “original” series with x
= 5, 6, 7, 8, and 9, and that all arose from 7 as
they speculated the primary basic number as 7
(Jones. 1957; Larsen, 1969; Marchant, 1973). As
stated by Mayo et al. (1997), reduction in diploid
number occurs in various morphologically ad¬
vanced genera. We agreed with this opinion: Ty¬
phonium is a much advanced genus in Araceae;
such a low chromosome number was possibly re¬
duced from a higher one. Other evidence came from
our recent cytological studies in 9 Typhonium spe-

S’ h ini

288

Novon

Figure 1. Typhonium jinpingense /.. L. Wang, H. Li & F. H. Bian. — 1. Plant with rhizome. — 2. Rhizome. — 3.
Spathe. — 4. Spadix. — 3. Male portion. — 6. Male flower.

I o mm

Volume 12, Number 2
2002

Wang et al.

Typhonium jinpingense from Yunnan

289

i'W® hn nun*

T. i . . . .

Illi- 3

1

I2345

Figure 2. Cytological plates of Typhonium jinpingense. — 1. Mitotic metaphase chromosomes in somatic cell. — 2.
Karyotype. — 3. Idiogram.

cies. The chromosome number in somatic cells of
these species is from 10 to 52, the karyotypieal
asymmetry for T. giganteum Kngler belongs the 2A
type, and 6 species. T. calcicolum C. Y. Wu, T. kun-
mingen.se H. hi, T. hlumei , T. trilobatum (L.) Schott,
T. roxburghii Schott, and T. umeiense H. Li, belong
to the 2B type, while T. jinpingense and triploid T.
flagelliforme (Loddiges) Blume, 2n = 3x = 24, be¬
long to the 3B type. It is commonly known that the
more asymmetric the chromosome, the more ad¬
vanced the species (Stebbins, 1971). The 3B type
is more asymmetric than the 2 A type and the 2B
type. So. from the standpoint of karyotype, T. jin¬
pingense and triploid T. flagelliforme may be con¬
sidered to be the most advanced among the 9 Ty¬
phonium species studied.

Jinping County is an interesting place lor the
cytological study of Typhonium, as well as of the
family Araceae. In addition to the new species with
its low chromosome count, triploid T. flagelliforme
2 n = 3x — 24 was also found in this county, and
the latter can easily be propagated by the stem, but
is not yet found to bear fruits both in the wild and
in the garden. The reason we chose the specific
epithet is to emphasize the importance of this
place.

Acknowledgments. The study was supported by
the Knowledge Innovation Project (grant number
KZ2000-14) and 'The Hope of West" program, the
Chinese Academy of Sciences. We are grateful to
Wu Xi-Ling (Kunming Institute of Botany, the Chi¬
nese Academy of Sciences) for preparing the illus¬
tration.

Literature Cited

Grayum, M. H. 1990. Evolution and phylogeny of the Ar¬
aceae. Ann. Missouri. Bot. Card. 77: 628—697.

Hetterscheid, W. L. A. & P. C. Boyce. 2000. A reclassi¬
fication of Sauromatum Schott and new species of Ty¬
phonium Schott (Araceae). Aroideana 23: 48—55.

- & V. D. Nguyen. 2001. Three new species of Ty¬
phonium (Araceae) from Vietnam. Aroideana 24: 24—
29.

- , D. Sookchaloem & J. Murata. 2001. Typhonium

(Araceae) of Thailand: New species and a revised key.
Aroideana 24: 30-55.

Jones, G. E. 1957. Chromosome Numbers and Phyloge¬
netic Relationships in the Araceae. Unpublished Ph.D.
Thesis, University of Virginia, Charlottesville.

Larsen, K. 1969. Cytology of vascular plants: III. A study
of Thai aroids. Dansk. Bot. Ark. 27: 39—59.

Levan, A. K. & G. Fredga. 1964. Nomenclature for cen-
tromeric position on chromosomes. Hereditas 52: 197—
200.

Li, M. X. & X. F. Zhang. 1991. Plant Chromosome Re¬
search Technology. The Press of Forest College of
Northeast China, Harbin [in Chinese].

Marchant, C. ,1. 1973. Chromosome variation in Araceae:
V. Acoreae to Lasieae. Kew Bull. 28: 199-210.

Mayo, S. J., J. Bogner & P. J. Boyce. 1997. The Genera
of Araceae. Royal Botanic Gardens, Kew.

Petersen, G. 1989. Cytology and systematics of Araceae.
Nordic J. Bot. 9: 119-166.

- . 1994. Chromosome numbers of the genera of Ar¬
aceae. Aroideana 16: 37-46.

Sriboonma, D., J. Murata & K. Iwatsuki. 1994. A revision
of Typhonium (Araceae). J. Fac. Sci. Univ. Tokyo, sect.
Ill, 14(4): 255-313.

Stebbins, G. L. 1971. Pp. 87-93 in Chromosomal Evolu¬
tion in Higher Plants. Edward Arnold, London.

Wang, Z. L. & H. Li. 1999. Sauromatum gaoligongense
(Araceae), a new species from Gaoligong Mountains.
Acta Bot. Yunnan., Suppl. XI: 61-64.

Three New Sections and a New Subgenus of
Phyl l a nth us (E u ph orb i ac e ae)

Grady L. Webster

Davis Herbarium, Section of Plant Biology, University of California, Davis,
California 95616, U.S.A.

AusTRACT. Three new sections ol Phyllanthus are
described: section Antipodanthus (subg. fsocladus;
type I*, dictyospermus Muller Argoviensis), 15 spe¬
cies of South America and Australia; section Pityr-
ocladus (subg. Emblica; type P. symphoricarpoides
Kunth), 5 species of tropical America; and section
Hylaeanthns (subg. Conami ; type P. attenuatus Mi-
quel), 6 species of tropical America. I wo new spe¬
cies are described: P. cuatrecasanus (sect. Pityro-
cladns) and P. callejasii (sect. Hylaeanthus). A new
subgenus with unique pollen structure, Cyclanthera
(type P. lindenianus Bail Ion), is proposed; it in¬
cludes sections Callitrichoides and Cyclanthera.

Key words: Euphorbiaceae, Neotropies, Phyl-
lanthus.

Phyllanthus, with over 800 species, is the third
largest genus of Euphorbiaceae (Govaerts et al.,
2000). The remarkable diversity in growth form and
floral morphology (Bancilhon, 1971; Nozeran et al.,
1984), and especially in pollen (Punt, 1967. 1987),
is reflected in its complex classification, with 10
subgenera and over 30 sections (Webster, 1956—
1958, 1967). Although about 70% of the species of
Phyllanthus are confined to the Old World, the 200-
plus neotropical species have not received exten¬
sive study except for the treatments of Miiller
(1873) for Brazil. Jablonski (1967) for the Amazon
basin, and Webster (1956—1958, 1999) for the West
Indies and Venezuelan Guayana. During the course
of a revision of the neotropical species, a consid¬
erable number of undescribed taxa have been de¬
tected. Because the names of three newly recog¬
nized sections and a new subgenus are being cited
in a study of the pollen morphology of neotropical
taxa (Webster & Carpenter, 2002), they are being
published here in advance of the overall revision
of the neotropical taxa.

INiy Haul bus subg. Isocludus G. E. Webster
IMiyllanthus sect. Antipodaiillius G. I.. Webster,
sect. nov. TYPE: Phyllanthus dictyospermus
Miiller Argoviensis.

A sectione Paraphyllaruho differ! disco 9 integro, pol-

linis grana grosse reticulata; ah aliis sectionibus subge-
nero Isoclado differ! foliis spiraliter insertis.

Monoecious or dioecious shrubs or subshrubs,
branching unspecialized, branches smooth or pa¬
pillate; phvllotaxy spiral; leaves petiolate, stipules
persistent; flowers pedicellate, in axillary glomer-
ules; sepals 5 or 6, imbricate, entire; staminate disk
of 5 or 6 segments; stamens 3, filaments free or
connate; anthers vertically or horizontally dehis¬
cent; pollen grains subglobose, 3- or 4-colporate,
mostly coarsely reticulate; pistillate disk patelli-
form; ovary glabrous, 3-locular; styles ± free, bifid;
fruits capsular, seeds smooth or verrueulose.

About 15 species can be referred to Phyllanthus
sect. Antipodanthus, which belongs to subgenus Is-
ocladus by virtue of its unspecialized (non-phyllan-
thoid) branching. It appears to be the Southern
Hemisphere vieariant of section Paraphyllanthus
Miiller Argoviensis. The two sections seem mor¬
phologically similar in many respects, but in sec¬
tion Paraphyllanthus the anthers are always verti¬
cally dehiscent and the pistillate disk dissected.
Only 5 species have been described from South
America, mostly Brazil; the others are native to
Australia (species 19, 21. 25-27, 29-32, 35-37,
and 39 in Bentham, 1873). A number of the Aus¬
tralian species have recently been treated by Hunt¬
er and Bruhl (1996, 2000). The Australian and
South American species differ in some characters,
the American species having mostly connate fila¬
ments and horizontally dehiscing stamens; it is con¬
ceivable that cladistic analysis may show that some
of the Australian species should be referred to sec¬
tion Lysiandra (F. Mueller) (7 I,. Webster. The Aus¬
tralian species are therefore listed here but not
treated in detail.

Australian species included: Phyllanthus aridus
Bentham, P. australis J. 1). Hooker, P. calycinus
Labillardifere, P. carpentariae Muller Argoviensis,
P. fuernrohrii E. Mueller, P. hirtellus Muller Argo¬
viensis, P. maitlandianus Diels, P. oblanceolatus ,|.
I. Hunter & J. J. Bruhl, P. scaber Klotzsch, P. sim-
ilis Miiller Argoviensis, and P. striaticaulis J. T.

Novon 12: 290-298. 2002.

Volume 12, Number 2
2002

Webster

Phyllanthus (Euphorbiaceae)

291

Hunter & J. J. Bruhl. It is possible that P. erwinii
J. T. Hunter & J. J. Bruhl and P. lacunarius F.
Mueller are also referable to Phyllanthus sect. An-
tipodanthus.

Key to the American Species of Phyllanthus sect.
Antipodanthus

la. Filaments connate; stems ± terete; stipules 1-6
mm long; seeds 2—2.7 mm long; monoecious or
dioecious.

2a. Stipules 3—6 mm long; stems smooth; leaves
on main stems > 5 mm long; dioecious.

3a. Leaves on main stems > 5 mm broad;
seeds reticulate or irregularly verrucu-

lose . 1. FI dictyospermus

31). Leaves on main stems < 5 mm broad;

seeds foveolate-pu nctate .

. 2. P. pinifolius

2b. Stipules 1-2.5 mm long; stems smooth or
papillose; leaves on main stems mostly < 5
mm long; monoecious or dioecious.

4a. Dioecious; stems papillose; staminate
flowers distinctly pedicellate (pedicel >

1 mm long); pistillate sepals 5; leaf
blade margins strongly revolute, tip
blunt with abruptly reflexed acumen . .

. 3. P. ramillosus

4b. Monoecious; stems smooth; staminate
flowers subsessile (pedicel < 1 mm
long); pistillate sepals 5 or 6; leal blade
margins not revolute, apically tapering
to a straight acumen . 4. P. dawsonii

lb. Filaments free; stems decurrent-angled from

nodes; stipules 2 mm long or less; seeds 1.4— 1.7
mm long; dioecious . 5. P. rosmarinifolius

1 . Phyllanthus dictyospermus Muller Argovien-

sis, DC. Prodr. 15(2): 394. 1866. TYPE: Bra¬
zil. Minas Gerais: ./. F. Widgren 1003 (holo-
type, G).

Phyllanthus regnellianus Muller Argoviensis, FI. Brasil.
11(2): 58. 1873. TYPE: Brazil. Minas Gerais: Cal-
das, A. F. Regnell II 48 (lectotype, designated here,
S).

Montane scrublands, ca. 1000—1500 m, Brazil
(Minas Gerais). Phyllanthus regnellianus, which
was distinguished by Muller only on the basis of a
questionable difference in seed sculpturing, shows
no clear differences and is here reduced to syn¬
onymy.

2. Phyllanthus pinifolius Baillon, Adansonia 1.

5: 353. 1865. TYPE; Brazil. Parana: Curitiba,
A. St. Hilaire 1559o [15590? J (holotype, P;
isotype. K).

Scrublands, 1500-2800 m, Minas Gerais, Espfr-
ito Santo, and Parana.

3. Phyllanthus ramillosus Muller Argoviensis,

Linnaea 32: 36. 1863. TYPE: Brazil, Santa
Catarina: Herb. Franqueville (syntype, G);
“Brasilia meridionali,” F Sellout (syntype, B
destroyed).

? Phyllanthus cordobensis (Kuntze) K. Schumann, Bot. Jah-
resber. 26(1): 350. 1898. Diasperus cordobensis Kun¬
tze, Rev. Gen. PI. 3(2): 285. 1898. TYPE: Argentina.
Cordoba: San Roque, O. Kuntze (holotype, NY).

Since the Sellow syntype of Phyllanthus ramil¬
losus was destroyed, the Herb. Franqueville spec¬
imen logically should become the lectotype; how¬
ever, that specimen could not be located in the
Prodromus Herbarium in Geneva. If the Herb. Fr¬
anqueville specimen cannot be located, St. Hilaire
1321 (P) could be designated as a neotype. It is
also possible that a duplicate of Sellow may yet be
discovered.

Woodlands and scrub, 15—2700 m, Bolivia,
northern Argentina, and Brazil (Santa Catarina and
Rio Grande do Sul). The status of Phyllanthus cor¬
dobensis remains uncertain; Kuntze’s description is
inadequate, and the single specimen is somewhat
different from other collections of P. ramillosus ;
possibly it might be validated at subspecific status
when better known.

4. Phyllanthus dawsonii Steyermark, Los Ange¬
les Co. Mus. Contr. Sci. 21: 13. 1958. TYPE:
Brazil. Goias: Chapada dos Veadeiros, E. Y.
Dawson 14776 (holotype, R).

Cerrado and campos rupestres, 1250—1500 m,
Goias.

6. Phyllanthus rosmarinifolius Muller Argo¬
viensis, FI. Brasil. 11(2): 60. 1873. TYPE:
Brazil. Rio de Janeiro: Serra dos Orgaos, G.
Gardner 5852 (holotype, G).

Recorded only Irom rocky slopes, 2000—2250 m,
Rio de Janeiro, Brazil.

Phyllanthus subg. Emhlica (Gaertner) Kurz
Phyllanthus sect. Pityrocladus G. L. Webster,
sect. nov. TYPE: Phyllanthus symphoricarpo-
ides Kunth.

A sectione Emhlica differt sepalis plerumque 5, an-
theris muticis, stylis liberis non dilalatis; a sectione Mi¬
cro glochidionis differt foliis non glanduliferis, antheris
muticis, pollinis grana 4- vel 5-colporatis non diporatis.

Monoecious (rarely dioecious) shrubs, often
scandent; branches and branchlets ± scabridulous
or papillate to hirtellous or scurfy; deciduous
branchlets pinnatiform (not ramified), with ea. 10—
50 leaves; leaf blades chartaceous, lacking a sub-

292

Novon

apical laminar gland; stipules persistent; flowers
pedicellate, in axillary unisexual or bisexual cy-
rnules; staminate sepals 5 or 6, subequal, free, im¬
bricate; disk segments 5; stamens 2-5(-7), fila¬
ments connate (rarely free); anthers muticous,
dehiscing horizontally or obliquely; pollen grains
subprolate, 4- or 5-colporate, colpi monorate and
unbordered, exine reticulate; pistillate sepals 5, im¬
bricate; disk erateriform or dissected; ovary 3-loc-
ular; styles free, spreading, bifid to unlobed; fruits
dehiscent or indehiscent; seeds smooth.

Phyllanthiis sect. Pityrocladus, which is a neo¬
tropical representative of Phyllanthus subg. Embli-
ca (Gaertner) kurz, includes 6 described species
(and at least I or 2 undescribed) mainly of South
America, but with one species in Costa Rica. The
4— 5-colporate reticulate pollen grains (Webster &
Carpenter, 2002) suggest placement in subgenus
Emblica, even though the shape of the grains is
scarcely prolate. The section, named for t lie char¬
acteristic scabridulous or sc urfy indumentum of the
axes of some of the species, is similar to section
Microglochidion (Muller Argoviensis) Miiller Argo-
viensis in some pollen characters but differs in its
tendency to a scandent habit, more or less scurfy
indumentum, eglandular leaves, and less elongated
pollen grains with fewer colpi.

Key to the Species ok Phyllanthus sect. Pityrocladus

la. leaves glabrous on both faces (at most scabri-
tlulous or papillate on petiole or midrib); veins
distinctly prominulous on abaxial surface;
branchlets 10—30 cm long, 0.8— 1.8 mm thick;
with 20—35 leaves; staminate pedicels 3—7 mm
long; stamens 3—6; fruiting pedicels 2-4 mm
long.

2a. Monoecious; leaf blades elliptic, 1 .5—2.5 cm
long, apically rounded and apiculate; sta¬
mens (3)4-6; fruits indehiscent; styles un¬
lobed or apically bifid; branchlets not dis¬
tinctly winged . 1. P symphoricarpoides

2b. Dioecious; leaf blades ovate-lanceolate, 3—6
cm long, acuminate; stamens 3; fruits dehis¬
cent; styles bifid; branchlets narrowly
winged . 2. P. sponiifolius

lb. Leaf blades glabrous or hirsutulous adaxially;
veins not prominulous abaxially (or if so, blades
distinctly hirsutulous abaxially); branchlets 5—25
cm long, 0.4—1 mm thick, with 5—75 leaves; sta¬
minate pedicels 0.5— 7.5 mm long; stamens 2 or
3; fruiting pedicels 1-4 mm long.

3a. Leaf blades abaxially copiously hirsutulous,
with prominent lateral veins; stamens 2;
styles unlobed.

4a. Monoecious; branchlets 0.6— 0.9 mm
thick, with 5—1 1 leaves; leaf blades
1 .5— 3.5 cm long; petioles 1—2 mm long;

stipules 3—4.5 mm long .

. 3. P. popayanensis

4b. Dioecious; branchlets 0.4— 0.6 mm
thick, with 20—45 leaves; leaf blades 1 —

1.5 cm long; petioles ca. 0.5 mm long;
stipules 1.5—2 mm long . . 4. P. ruscifolius

3b. Leaf blades abaxially glabrous or minutely
scabridulous, veins not prominulous; sta¬
mens 2 or 3.

5a. Leaf blades elliptic, strongly inaequila-
teral at base, secondary veins straight,
nearly perpendicular to midrib, abaxi¬
ally separated from midrib; branchlets
0.4— 0.7 mm thick; stipules 1—1.5 mm
long; staminate disk segments discrete,

flat . 5. P. cuatrecasanus

5b. Leaf blades oblong, not inaequilateral at
base, secondary veins distant, curving,
not abaxially separated lrom midrib;
branchlets 0.7-1 mm thick; stipules 2-

3.5 mm long; staminate disk segments

massive, ± confluent . 6. P. valerii

1. Phy llantliiis symphoricarpoides kuiith. in

Humboldt, Bonplaml & Kunth, Nov. Gen. Sp.
2: 114. 1817. Glochidion symphoricarpoides
(Kunth) Pax & K. Hoffmann, in Engler &
Harms, Nat. Pflanzenfam. ed. 2, 19c: 58.
1931. T\ PE: Ecuador. Loxa [Loja]: Humboldt
& Bonpland (holotype, P; isotype, R).

Montane thickets, 1500—2750 m. Colombia to
Peru.

2. Phyllanthus sponiifolius Miiller Argoviensis,

Linnaea 32: 25. 1863. TYPE: Ecuador. L. Era¬
ser (holotype, G).

Montane thickets, 1000—2000 m, Colombia to
Ecuador.

3. Phyllanthus popayanensis Pax, Rot. Jahrb.

26: 503. 1899. TYPE: Colombia. Popaydn:
Paramo de Guanacas, E C. Lehmann 4708
(holotype, R destroyed; lectotype, designated
here, k).

Montane thickets, 2000—2800 m, Colombia
(Cauca). The specimen at kew is chosen as lecto¬
type because no other specimens of Lehmann 4708
have been found.

4. Phyllanthus ruscifolius Miiller Argoviensis,

DC. Prodr. 15(2): 358. 1866. TYPE: Colombia.
Prov. Cauca: J. J. Triana .3659 (holotype, P).

Montane thickets, 1900-2200 m, Colombia (Val¬
le del Cauca).

5. Phyllaiilhus cuatrecasanus G. L. Webster, sp.

nov. TYPE: Colombia. Caqueta: Quebrada del
Rfo Haeha, 2100—2250 m, J. Cuatrecasas
85.3.3 (holotype, US). Eigure 1.

Volume 12, Number 2
2002

Webster

Phyllanthus (Euphorbiaceae)

293

Similis Phyllanthi valerii, differt foiiis laevibus, dola-
briformis, nervis lateralibus rectis; stipulis minoribus; seg¬
ments disci tenuioribus.

Dioecious shrub; branchlets terete, copiously
hirtellous, 8-14 cm long, 0.4-0.7 mm broad, with
ca. 20—50 leaves; leaf blades glabrous, mostly 5-7
mm broad, elliptic, abruptly acute, ± inaequilateral
at base, abaxially alveolate-reticulate, veins
straight, these and reticulum of veinlets prominu-
lous; stipules oblong, blunt, glabrous, 1—1.2 X 0.3—
0.4 mm; staminate flowers 2-5 per axil on brach-
yblasts; pedicel 1.3— 1.5 mm long; sepals 5, 1.3-
1.6 X 0.8— 1.4 mm; disk segments 5, elliptic, 0.2-
0.25 mm broad; stamens 2, filaments completely
connate into a column 0.5-0.6 mm high; anthers <
0.5 mm long, dehiscing obliquely; pistillate flowers
unknown.

Known only from the type collection. The distin¬
guished collector, Jose Cuatrecasas, described the
perianth segments as “amarillentos con nervios ro-
jizos.” The specimen of Phyllanthus cuatrecasanus
superficially resembles small-leaved forms of P.
ruscifolius, but differs sharply from that species (as
well as P. valerii) in the glabrous, strongly inae¬
quilateral apiculate leaf blades with curious glau¬
cous-reticulate bands alternating with green lateral
veins.

6. Phyllanthus valerii Stand ley. Field Mus. Publ.
Bot. 18: 619. 1937. TYPE: Costa Rica. He¬
redia: Yerba Buena, NE of San Isidro de He¬
redia, 2000 m, P. C. Standley & F. L. Valerio
49814 (holotype, F).

Montane forests, Costa Rica, 1200—2050 m; very
similar plants have been collected in Bolivia.

Phyllanthus subg. Conami (Aublet) G. L. Webster

Phyllanthus sect. Hylaeanthus G. L. Webster,
sect. nov. TYPE: Phyllanthus attenuatus Mi-
quel.

A sectione Nothoclema differt ramulis pinnatiformibus
(non bipinnatiformibus), fructibus baccatis; a sectione
Apolepis differt filamentis connatis, disco 9 integro.

Monoecious or dioecious trees or shrubs; branch-
lets pinnatiform, usually subtended by reduced
leaves (but generally not by cataphylls); flowers in
axillary unisexual or bisexual cymules borne on
brachyblasts; sepals 6 (rarely 5), usually biseriate;
disk of discrete segments or cupular; stamens 3,
filaments free or connate; anthers muticous, de¬
hiscing horizontally or obliquely; pollen grains glo¬
bose, porate (lacking distinct colpi), exine pilate;

pistillate sepals persistent in fruit; ovary 3— 5-loc-
ular; styles free or connate, bifid to unlobed; fruits
baccate; seeds smooth.

This new section of Phyllanthus subg. Conami
(Aublet) G. L. Webster, named for the prevalent
distribution of its species in lowland Amazonian
forest (Hylaea), has escaped recognition partly be¬
cause of its vegetative resemblance to species of
two sections of Phyllanthus subg. Xylophylla (L.)
Persoon: section Brachycladus G. E. Webster (Web¬
ster, 2001) and section Elutanthos Croizat (Croizat,
1943); this has led to confusion and failure to rec¬
ognize the significance of the diagnostic characters.
The resemblance of the globose pollen grains with
pilate exine ornamentation to those of section Noth¬
oclema G. L. Webster clearly supports placement
of section Hylaeanthus in subgenus Conami (Web¬
ster & Carpenter, 2002).

Muller (1866, 1873) created much of the con¬
fusion with regard to the new section Hylaeanthus
when he mistakenly identified the type species,
Phyllanthus attenuatus Miquel, with P. aubletianus
Radcliffe-Smith (2001; name replacing Meborea
guianensis Aublet). The affinities of Aublet’s Me¬
borea remain uncertain, but it seems possibly a
species of section Brachycladus. At present, section
Hylaeanthus includes 6 species, although addition¬
al species remain unpublished. It is biogeographi-
cally parallel to section Pityrocladus in its mainly
South American distribution, but with one species
in Costa Rica.

Key to the Species of Phyuanthus sect. Hylaeanthus

la. Ovary 3-locular; styles bifid; monoecious or dioecious.
2a. Leaves completely glabrous; dioecious.

3a. Leaf blades lanceolate, abruptly acuminate;

styles connate . 1. P skutchii

3b. Leaf blades usually ovate or elliptic, blunt;

styles free . 2. P manausensis

2b. Leaves hirtellous at least on petiole; monoecious
or dioecious.

4a. Fruiting pedicels 2^4 mm long; styles free,
bifid or unlobed.

5a. Dioecious; styles bifid; 8 disk entire

. 3. P. attenuatus

5b. Monoecious; styles unlobed; 8 disk dis¬
sected . 4. P. bernardii

4b. Fruiting pedicels 6—7 mm long; styles con¬
nate, twice-bifid . 5. P. callejasii

lb. Ovary 4— 5-locular; styles unlobed; filaments con¬
nate; monoecious . 6. P madeirensis

1. Phyllanthus skutchii Standley, Publ. Field
Mus. Nat. Hist., Bot. 22: 346. 1940. TYPE:
Costa Rica. San Jose: El General, A. Skutch
4375 (holotype, F).

Evergreen forests, 20-900 m, southern Costa

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Phyllanthus (Euphorbiaceae)

295

Rica (Pacific slope); possibly occurring in Colom¬
bia.

2. Phylluntlius inanausensis W. A. Rodrigues.
Acta Amazonica 1: 17, fig. 1. 1971. TYPE:
Brazil. Amazonas: Manaus, W. A. Rodrigues
7520 (holotype, IN PA 16822).

Lowland Amazonian rain forest, mostly on terra
firme, 100—200 m, Brazil (Amazonas, Para, Rorai-
ma, Amapa).

8. Phyllanthus attenuatus Miquel, Linnaea 21:
479. 1848. TYPE: Suriname. F W. R. Hostman
& A. Kappler 305 (holotype, L; isotype, W).

Terra firme or occasionally inundated rain forest,
0—1000 m, Colombia, Venezuela, the Guayanas,
Brazil, and Peru.

4. Phyllanthus bernardii Jablonski, Mem. New

York Bot. Card. 17: 112. 1967. TYPE: Vene¬
zuela. Merida: Aricagua, L. Bernardi 6253 (ho¬
lotype, NY).

Andean forests, > 2000 m, western Venezuela
(Merida). The species deviates from others in Phyl¬
lanthus sect. Hylaeanthus in its character combi¬
nation of monoecious flower production, winged
pedicels, and small seeds (< 2 mm long). It has
not yet been possible to examine the pollen, and it
may prove not to belong to the section.

5. Phyllanthus callejasii G. L. Webster, sp. nov.

H PE (pistillate): Colombia. Antioquia: Mun.
Frontino, Corr. La Blanquita, region de Murri.
Alto de Cuevas, 1850 m, 14 July 1988, R.
Callejas, A. L. Arheldez , ./. Betancur & I. I).
Castaho 6364 (holotype, HU A 52926; isotype,
l)AV). Figure 2.

Af) aliis speciebus sectionis iliffert stylis connatis bis
bifidis, sepalis fructiferis majoribus, ramulis crassioribus.

Dioecious tree 6-8 m high; branchlets terete,
smooth (minutely papillate), 15—25 cm long, 1.5—
2.5 mm thick, with 6—9 leaves; leaf blades ovate to
elliptic, 7—12 cm long, 3.5—7 cm broad, glabrous
except for the incised midrib adaxially and midrib
and major lateral veins abaxially, veins 5—7 pairs,
curving; veinlet reticulum prominulous beneath;

<—

Figure 1. Phyllanthus cuatrecasanus G. I.. Webster. —
Left-hand photo. Type specimen ( Cuatrecasas 8533, US).
— Right-hand photo. Enlarged view of branch apex with
deciduous branchlets.

petiole 5-7 X 1-1.5 mm, copiously scabrid-hirtel-
lous; stipules blackish, triangular, 2—3 mm long.
Flowers in axillary glomerules, the pistil late mostly

2- 5 per axil; bracts anti bractlets blackish, persis¬
tent. Staminate flowers seen only in bud; sepals 6;
disk undivided; stamens 3, filaments connate, an¬
thers dehiscing horizontally. Pistillate flowers with
pedicel glabrous or sparsely and minutely hirtel-
lous, 5.5—7 mm long in fruit; fruiting sepals (5)6,

3— 3.8 mm long, 2—2.2 mm broad; disk patelliform,
ea. 1 .5 mm broad; ovary 3-locular; styles connate
into a column ca. 1.5 X 0.7 mm; style branches
0.5— 0.7 mm long, ± twice-bifid. Fruits baccate, 7—
9 mm diam. (dried); seeds somewhat compressed,
acute, truncate at base, 2.5—3 X 2-2.7 mm; testa
smooth, brownish.

Cloud forest, 1600-1850 m. Cordillera Occiden¬
tal, Colombia. Phyllanthus callejasii is clearly re¬
lated to Phyllanthus attenuatus Miquel by its bifid
styles and dioecious flower production, but is easily
distinguished by its larger leaf blades and longer
petioles, thicker branchlet axes, and longer fruiting
pedicels and sepals. It differs from the lowland P.
attenuatus and P. manausensis in its upper montane
cloud forest habitats. Phyllanthus bernardii, the
only other upland species of Phyllanthus sect. Hy¬
laeanthus, has hirtellous branchlet axes, winged
pedicels, and smaller seeds.

Additional collections. COLOMBIA. Antioquia: Mun.
Frontino, Corr. Nutibara, zona de Mum, 5—8 km S de Alto
de Cuevas, 1000—1850 m, 14 Feb. 1991, R. Callejas, F.
J. Roldan & M. V. Arbeldez 9940 (staminate paratype:
HU A 79857). Nariiio: Mun. Piedraancha, Correg. Chu-
eunez, Reserva La Planada, 1650-1800 m. 18 May 1991,
./. Betancur, S. Churchill & F. J. Roldan 2576 (HUA).

6. Phylluntlius madeirensis Croizat, Trop.
Woods 78: 7. 1944. TYPE: Brazil. Amazonas:
Mun. Humaita, between Rio Livramento and
Rio Ipixuna, B. A. Krukoff 7163 (holotype. A;
isotype, NY).

Amazonian rain forest, mostly on terra firme, ca.
200—300 m, Brazil (Amazonas, Para). This species
is sharply separated by a number of characters from
the other species of the section, but the distinctive
pollen sculpturing supports placement in Phyllan¬
thus sect. Hylaeanthus.

Pliyllantliiis subg. Cyclanthera G. L. Webster,
subg. nov. TYPE: Phyllanthus lindenianus
Baillon.

Herbae monoicae, ramifieatione more sectionis Phyl-
lanthi; flores solitarii, axillares; sepala 6 5 vel 6; disci
segmentis plerumque discretis; stamina 2 vel 3, connata;
antherae discretae vel connatae, horizontaliter dehiscen-

52926

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Phyllanthus (Euphorbiaceae)

297

tes; pollinis grana exinio clypeato; sepala 9 6; discus dis-
sectus; ovarium 3-loculare; styli bifidi; fructus capsularis;
semina scabridula vel verruculosa.

Monoecious annual or perennial herbs; branch-
lets pinnatiform or with one iterative lateral axis;
flowers axillary, solitary; staminate sepals 5 or 6;
disk dissected; stamens 2 or 3, filaments connate;
anthers discrete or completely connate into a syn-
andrium, dehiscing horizontally; pollen grains
spheroidal, clypeate, the exine shields banded or
circular; pistillate sepals 6; disk dissected, often
purplish; ovary 3-locular, styles bifid; fruit capsu¬
lar; seeds scabridulous or verruculose.

Phyllanthus subg. Cyclanthera includes two sec¬
tions, section Callitrichoides G. L. Webster and sec¬
tion Cyclanthera G. L. Webster, both endemic to
the Greater Antilles. When these sections were first
treated (Webster, 1956—1958), they were included
in subgenus Phyllanthus, but it was noted that they
have extraordinary pollen grains somewhat com¬
parable to the clypeate pollen grains (with exine
shields) in subgenus Xylophylla (Halbritter & Hes¬
se, 1995). Recent studies of pollen morphology
(Webster & Carpenter, 2002) graphically indicate
the divergence of the pollen grains in section Cy¬
clanthera from all other taxa of Phyllanthus. It
therefore seems necessary to remove these odd
West Indian plants from subgenus Phyllanthus into
a separate subgenus. The relationships of subgenus
Cyclanthera still remain uncertain, although the
distinctive pollen shows some resemblance to the
pollen of subgenus Emblica.

Sect. 1. Phyllanthus sect. Callitrichoides G. L.

Webster, Contr. Gray Herb. 176: 51. 1955.

TYPE: Phyllanthus carnosulus Miiller Argo-

viensis.

One species endemic to eastern Cuba; apparent¬
ly related to Phyllanthus tenuicaulis Miiller Argo-
viensis (in sect. Cyclanthera ). This curious plant is
vegetatively distinctive in its crisp-succulent leaves
and stems rooting at the nodes. The pollen grains,
in contrast to those of section Cyclanthera, have
elongated exine shields (or perhaps could be inter¬
preted as stephanocolporate). Phyllanthus carno¬
sulus appears not to have been collected since it
was found by Ekman over 70 years ago, so it is not
certain whether it still survives.

<—

Figure 2. Phyllanthus callejasii G. L. Webster. — Left-
band photo. Type specimen with pistillate flowers ( Callejas
et al. 6864, HUA). — Right-hand photo. Specimen with
fruits ( Betancur et al. 2576, HUA).

Sect. 2. Phyllanthus sect. Cyclanthera G. E.

Webster, Contr. Gray Herb. 176: 47. 1955.
TYPE: Phyllanthus lindenianus Baillon.

Phyllanthus sect. Cyclanthera includes four spe¬
cies (and some subspecies) of Hispaniola, two of
which also occur in eastern Cuba (Webster, 1956—
1958). The section is very isolated within Phyllan¬
thus by its highly modified androecium with unique
clypeate pollen grains (illustrated by Webster &
Carpenter, 2002).

Acknowledgments. I thank Kevin Carpenter for
his assistance in obtaining critical pollen evidence
with scanning electron microscopy at the Micros¬
copy and Imaging Facility at the University of Cal¬
ifornia, Davis. Photographs of herbarium specimens
were taken by Sam Woo, at the Instructional Tech¬
nology and Digital Media Center, University of Cal¬
ifornia, Davis. Thanks are also due to the curators
of several herbaria (including A, F, Gil, NE, NY,
TEX, UC, US) for the loan or donation of speci¬
mens, and particularly to Ellen Dean, director of
the Davis Herbarium, for providing the facilities (or
this research. The manuscript has been signifi¬
cantly improved by suggestions from Petra Hoff¬
mann (Royal Botanic Gardens, Kew) and Peter van
Welzen (Rijskherbarium, Leiden).

Literature Cited

Bancilhon, L. 1971. Contribution & lYtude taxonomique
du genre Phyllanthus (Euphorbiac^es). Boissiera 18: 1—
81.

Bentham, G. 1873. Phyllanthus. Flora Australiensis 5:
93-1 12. L. Beeve, London.

Croizat, L. 1943. Notes on American Euphorbiaceae with
descriptions of eleven new species. J. Wash. Acad. Sci.
33: 11-20.

Govaerts, FL, I). G. Frodin & A. Radcliffe-Smith. 2000.
World Checklist and Bibliography of Euphorbiaceae.
Royal Botanic Gardens, Kew.

Halbritter, H. & M. Hesse. 1995. The convergent evolu¬
tion of exine shields in angiosperm pollen. Grana 34:
108-1 19.

Hunter, J. T. & J. J. Bruhl. 1996. Three new species of
Phyllanthus (Euphorbiaceae: Phyllantheae) in South
Australia. J. Adelaide Bot. Gard. 17: 127—136.

- . 2000. Phyllanthus. In N. G. Walsh & T. ,1. En-

twisle. Flora of Victoria, vol. 4: 74—78. Inkata Press,
Melbourne.

Jablonski, E. 1967. Euphorbiaceae. Botany of the Gua-
yana Highland — Part VII. Mem. New York Bot. Gard.
17(1): 80- 190.

Lanjouw, J. 1931. The Euphorbiaceae of Surinam. Proef-
sehrift, Rijks-Universiteit, Utrecht.

Muller. J. 1866. Euphorbiaceae [except Euphorbieae]. In
A. De Candolle, Prodromus systematis naturalis regni
vegetabilis 15(2): 189-1286. Victor Masson, Paris.

- . 1873. Phyllanthus. In F. von Martius (editor).

Flora Brasiliensis 11(2): 23—76.

Nozeran, R., L. Rossignol-Bancilhon & G. Mangenot.

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1984. Les recherches stir le genre Phyllanthus (Eu-
phorbiaceae): Acquis et perspectives. Bot. Helv. 94:
199-233.

Punt, W. 1967. Pollen morphology of the genus Phyllan¬
thus (Euphorbiaceae). Rev. Palaeobot. Palynol. 39:
131-160.

- . 1987. A survey of pollen morphology in Euphor¬
biaceae with special reference to Phyllanthus. Bot. J.
Linn. Soc. 94: 127-142.

Radcliffe-Smith, A. 2001. Genera Euphorbiacearum. Roy¬
al Botanic Gardens, Kew.

Webster, G. E. 1956— 1958. A monographic study of the
West Indian species of Phyllanthus. J. Arnold Arbor.

37: 91-122, 2 1 7-268. 340-359; 38: 51-80. 170-198,
295-373; 39: 49-100, 111-212.

- . 1967. Tht* genera of Euphorbiaceae in the south¬
eastern United States. J. Arnold Arbor. 48: 303—430.

- . 1999. Phyllanthus. Pp. 191-205 in P. E. Berry,

K. Yatskievych & B. Holst (editors), Elora of the Ve¬
nezuelan Guayana, Vol. 5. Missouri Botanical Garden
Press, St. Louis.

- . 2001. Synopsis of Croton and Phyllanthus (Eu¬
phorbiaceae) in western tropical Mexico. Contr. Univ.
Michigan Herb. 23: 353-388.

- & K. J. Carpenter. 2002. Pollen morphology and

phylogenetic relationships in neotropical Phyllanthus
(Euphorbiaceae). Bot. J. Linn. Soc. 138: 325—338.

Resurrection of Segregates of I lie Polyphyletic Genus Zigadenus
s.l. (L i I i ales : Melanthiaceae) and Resulting New Combinations

Wendy IS. Zomlefer

University of Georgia Herbarium, Department of Plant Biology, 2502 Plant Sciences,
Athens, Georgia 30602-7271, U.S.A.
wendyz@dogwood.botany.uga.edu

Waller S. Judd

Department of Botany, University of Florida, P.O. Box 1 18526.
Gainesville, Florida 32611-8526, U.S.A.
wj udd@botany.ufl.edu

Abstract. Reinstatement of segregate genera
and the accompanying recombinations for species
in t he former polyphyletic Zigadenus complex al¬
low for a revised, practical taxonomy that defines
monophyletic groups and reflects phylogenetic re¬
lationships within tribe Melanthieae (Melanthi¬
aceae). Nine newT species combinations result from
recircumscription of Zigadenus Michaux and Sten-
anthium (A. Gray) Kunth and from recognition of
Anticlea Kunth and Toxieoscordion Rydberg. The
genus Stenanthella Rydberg and the basionyms for
Stenanthium leimanthoides (A. Gray) Zomlefer &
Judd ( Amianthium leimanthoides A. Gray) and An¬
ticlea saehalinensis (F. Schmidt) Zomlefer & Judd
(Stenanthium sachalinense F. Schmidt) are leetotyp-
ified.

Key words: Amianthium , Filiales. Melanthi¬
aceae, Stenanthium, Zigadenus.

The Melanthiaceae sensu the Angiosperm Phy-
logeny Group (APG, 1998) comprise 11 to 16 gen¬
era (ca. 154—201+ spp.; Zomlefer et al., 2001) of
predominately woodland and/or alpine perennial
herbs occurring mainly in the temperate to arctic
zones of the Northern Hemisphere (with one spe¬
cies of Sehoenocaulon extending into South Amer¬
ica). Of concern here are certain taxa of the tribe
Melanthieae (Zomlefer et ah. 2001): Amianthium (1
sp.), Stenanthium (3—5 spp.). and Zigadenus (19
spp.). The poorly defined genus Stenanthium s.l.
has sometimes been divided into two genera: Sten¬
anthella (2—4 spp.) and a monotypic Stenanthium
(S. gramineum ; see Rydberg, 1900; Gleason, 1952;
Utech, 1987a, b). Taxonomists have divided Ziga¬
denus into several segregate genera (summaries in
Rydberg, 1903; Gates. 1918; Preece, 1956; Zom¬
lefer. 1997). However, contemporary botanists have

typically accepted only one segregate — a monotyp¬
ic Amianthium (Utech, 1986) — with the remaining
species maintained in Zigadenus s.l. (as in Walsh,
1940; Preece, 1956; Schwartz, 1994. 2002).

As part of a study on the tribe Melanthieae (Zom¬
lefer et ah. 2001), the circumscription of the Zi¬
gadenus complex was evaluated using parsimony
analyses of ITS (nuclear ribosomal) and /mL-F
(plastid) DNA sequence data. Based on the clado-
grams resulting from this study, Stenanthium is bi-
phyletic (and embedded within Zigadenus), the tra¬
ditional Zigadenus s.l. is polyphyletic, and
Amianthium is a distinct entity only distantly re¬
lated to the other Zigadenus species. The species
of these genera form five strongly supported elades
that correlate with geographical distribution, chro¬
mosome number, and certain morphological c har¬
acters (Table I ). Therefore, we recognize five gen¬
era (some with novel circumscription): (1)
Amianthium (monotypic: A. muscitoxicum ), (2) An¬
ticlea (including Stenanthella; ca. 1 1 spp.), (3)
Stenanthium ( Stenanthium gramineum, Zigadenus
densus, and Z. leimanthoides), (4) Toxieoscordion
(ca. 8 spp.), and (5) Zigadenus s. str. (monotypic:
Z. glaberrimus).

The nomenclatural consequences of these results
include reinstatement of two segregate taxa ( Anti¬
clea and Toxieoscordion ), redefinition of Zigadenus
and Stenanthium, and nine recombinations. For
convenience, we include a key and concise treat¬
ment of all five genera, as well as a list of species
(Table 2). Although a complete revision of these
genera and an investigation of species-level issues
are not the focus of this paper, some problematic
species complexes are mentioned in the generic
commentaries. Table 3 cross-indexes the combina¬
tions accepted in this paper with familiar Zigaden-

Novon 12: 299-308. 2002.

300

Novon

included within) Ziga- Rydberg Small spp. (“death camas”)

denus s.l. Zigadenus dens us

and Z. leimanthoides
perhaps conspecific

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2002

Zomlefer & Judd
Reconsideration of Zigadenus

301

Table 2. Segregate genera of the Zigadenus complex reflecting the clades supported by morphological and molecular
data (Zomlefer et ah, 2001). This list includes nine new combinations indicated here with an asterisk.

1. Amianthium A. Gray

A. muscitoxicum (Walter) A. Gray

2. Anticlea Kunth

A. elegans (Pursh) Rydberg

*A. frigida (Schlechtendal & Chamisso) Zomlefer & Judd
*A. hintoniorum (B. I,. Turner) Zomlefer & Judd
*A. mogollonensis (W. J. Hess and Sivinski) Zomlefer & Judd
*A. neglecta (Espejo, A. R. Lopez-Ferrari & Ceja) Zomlefer & Judd
*A. occidental is (A. Gray) Zomlefer & Judd
*A. sachalinensis (F. Schmidt) Zomlefer & Judd
A. sibirica (L.) Kunth
A. vaginata Rydberg
A. virescens (Kunth) Rydberg
A. volcanica (Bentham) J. G. Baker

3. Stenanthium (A. Gray) Kunth

S. granxineum (Ker Gawler) Morong

*S. densum (Desrousseaux) Zomlefer & Judd
*S. leimanthoides (A. Gray) Zomlefer & Judd

4. Toxicoscordion Rydberg

T. brevibracteatus (M. E. Jones) R. R. Gates
T. exaltatum (Eastwood) A. Heller

*T. fontanum (Eastwood) Zomlefer & Judd
/.’ fremontii (Torrey) Rydberg
T. micranthum (Eastwood) A. Heller
T. nuttallii (A. Gray) Rydberg
T. paniculatum (Nuttall) Rydberg
T. venenosum (S. Watson) Rydberg

5. Zigadenus Michaux

Z. glaberrimus Michaux

us and Stenanthium species names (in alphabetical
order) used in current literature.

1. Amianthiuni A. Gray, Ann. Lyceum Nat. Hist.
New York 4: 121. 1837, nom. cons. TYPE:
Amianthium muscitoxicum (Walter) A. Gray,
typ. cons, [basionym: Melanthium muscaetox-
icum Walter],

Chrosperma Rafinesque, Neogenyton 3. 1825, nom. rej.

TYPE: Melanthium laetum W. Aiton.

Endocles Salisbury, Gen. PI. 51. 1866. TYPE: not desig¬
nated.

Salient features. Ovoid bulb, obtuse (slightly
narrowed but not clawed) tepal bases, reduced/ab¬
sent tepal glands, partly inferior ovary, large seeds
with a reddish to purple sareotesta, distinctive al¬
kaloids: base chromosome number v = 8.

Distribution. Southeastern United States (coast¬
al plain and mountains) in acidic coniferous forests
and bogs.

Beyond the lack of glands (Utech, 1986) and the
distinctive seeds, this genus is characterized by
amianthine, a unique alkaloid, and also jervine, a
veratrum alkaloid, evidently not found in other spe¬
cies of the Zigadenus complex (Neuss, 1953; Kup-
chan et ah, 1961). Amianthium has been consid¬

ered closely related to Zigadenus s.l. (see Zomlefer,
1997); however, molecular data (Zomlefer & Per¬
kins, 1999; Zomlefer et ah, 2001) strongly support
the separation of Amianthium as only distantly re¬
lated to other Zigadenus species. As circumscribed
here (and in most current literature), this genus
comprises one species, Amianthium muscitoxicum
(Walter) A. Gray. The preferred spelling ''muscitox¬
icum" rather than “ muscaetoxicum ” for the specific
epithet is supported by the ICBN (Greuter et ah,
2000), Article 60, Recommendation 60G.

2. Aniiclea Kunth, Enum. PI. 4: 191. 1843. Gei-
seleria Kunth, Abh. Konigl. Akad. Wiss. Ber¬
lin 3. 1842, nom. illeg., non Geiseleria
Klotzsch 1843 (Euphorbiaceae). TYPE: Anti¬
clea sibirica (L.) Kunth (lectotype, designated
by Pfeiffer, 1873: 219).

Stenanthella Rydberg, Bull. Torrey Bot. Club 27: 530.
1900. Syn. nov. TYPE: Stenanthella occidentalis (A.
Gray) Rydberg (lectotype, designated here).

Although Rydberg (1900) included two species
in his new genus Stenanthella, Stenanthella oc¬
cidentalis (A. Gray) Rydberg and Stenanthella
sachalinensis (F. Schmidt) Rydberg, he based his
description of Stenanthella on cited specimens of

302

Novon

Table 3. The species of the polyphyletic genera Zigadenus s.l. and Stenanthium s.l., typically employed in contem¬
porary floras (e.g., Ahles, 1968; Hitchcock et ah. 1969; Cronquist et ah. 1977; Gleason & Cronquist, 1991; McNeal,
1993; Kartesz, 1994; Schwartz, 2002), are listed in alphabetical order with the recombination advocated in this paper
(boldface), including nine new combinations. For completeness, the monotypic genus Amianthium , a segregate of the
Zigadenus complex, is also included; its segregate status is supported by Zomlefer et ah (2001). Only three species
(, Amianthium muscitoxicum, Stenanthium gramineum , and Zigadenus glaberrimus ) are retained in their traditional gen¬
era; Amianthium and Zigadenus are monospecific.

Ainianthiiun muscitoxicum (Walter) A. Gray

Stenanthium frigidum (Schlechtendal & Chamisso) Kunth = Ant idea frigida (Schlechtendal & Chamisso) Zomlefer
& Judd

Stenanthium grarnineum (Ker Gawler) Morong

Stenanthium occidentale A. Gray = Anticlea occidentalis (A. Gray) Zomlefer & Judd

Stenanthium sachalinense F. Schmidt = Anticlea sachalinensis (F. Schmidt) Zomlefer & Judd

Zigadenus brevibracteatus (M. E. Jones) H. M. Hall = Toxicoscordion brevihracteatus (M. E. Jones) R. R. Gates

Zigadenus densus (Desrousseaux) Fernald = Stenanthium densum (Desrousseaux) Zomlefer & Judd

Zigadenus elegans Pursh = Anticlea elegans (Pursh) Rydberg

Zigadenus exaltatus Eastwood = Toxicoscordion exaltatum (Eastwood) A. Heller

Zigadenus fontanus Eastwood = Toxicoscordion font an um (Eastwood) Zomlefer & Judd

Zigadenus fremontii (Torrey) Torrey ex S. Watson = Toxicoscordion fremontii (Torrey) Rydberg

Zigadenus glaberrimus Michaux

Zigadenus hintoniorum B. L. Turner = Anticlea hintoniorum (B. L. Turner) Zomlefer & Judd
Zigadenus leimanthoides (A. Gray) A. Gray = Stenanthium leimanthoides (A. Gray) Zomlefer & Judd
Zigadenus micranthus Eastwood = Toxicoscordion micranthum (Eastwood) A. Heller

Zigadenus mogollonensis W. J. Hess & Sivinski = Anticlea mogollonensis (W. J. Hess & Sivinski) Zomlefer & Judd
Zigadenus neglectus Espejo, A. R. Lopez-Ferrari & Ceja = Anticlea neglecta (Espejo, A. R. Ldpez-Ferrari & Ceja)
Zomlefer & Judd

Zigadenus nuttallii (A. Gray) S. Watson = Toxicoscordion nuttallii (A. Gray) Rydberg
Zigadenus paniculatus (Nuttall) S. Watson = Toxicoscordion paniculatum (Nuttall) Rydberg
Zigadenus sibiricus (h.) A. Gray = Anticlea sibirica (L.) Kunth
Zigadenus vaginatus (Rydberg) J. F. Macbride = Anticlea vagina t a Rydberg
Zigadenus venenosus S. Watson = Toxicoscordion venenosum (S. Watson) Rydberg
Zigadenus virescens (Kunth) J. F. Macbride = Anticlea virescens (Kunth) Rydberg
Zigadenus volcanicus Bentham = Anticlea volcanica (Bentham) Baker

S. occidentalis. Therefore, we select this species
as the lectotype.

Salient features. Narrowly ovoid bulb, cuneate
to gradually narrowed (but not clawed) tepal bases,
one bilobed gland per tepal, half-inferior ovary;
base chromosome number x — 8.

Distribution. Asia, North America sou th to
Guatemala in acidic and/or alpine coniferous for¬
ests, prairies, calcareous shores and fens.

fhe segregate genus Anticlea (Kunth, 1843; Ryd¬
berg, 1903) has been more recently considered as
Zigadenus sect. Anticlea (Kunth) Bentham by Pree-
ce (1936) and Schwartz (2002). The one bilobed
gland per tepal is an autapomorphy for this taxon.
Resurrection of Anticlea involves six new combi¬
nations with species formerly placed in Stenanthel-
la and Zigadenus s.l. (Table 2). Stenanthella, a
Stenanthium segregate (Rydberg, 1900), shares lit¬
tle morphologically with Stenanthium s. str. but has
the diagnostic features of Anticlea , including the
distinctive nectaries and half-inferior ovary. In ad¬

dition, Stenanthella occidentalis ( Stenanthium oc¬
cidentale) resolves in the Anticlea clade in the mo¬
lecular study by Zomlefer et al. (2001).

Anticlea includes the only Asian species in the
Zigadenus complex (A. sibirica and A. sachalinen¬
sis ), the Guatemalan A. volcanica, and the wide¬
spread and variable North American species A. ele¬
gans and A. virescens and their segregates (A.
vaginata and A. hintoniorum, A. mogollonensis, and
A. neglecta, respectively). The latter are discussed
in detail by Turner (1992), Hess and Sivinski
(1995), and Frame et al. (1999). Further study of
these taxa is needed to evaluate the current specific
and infraspecific delimitations and to suggest bio¬
logically meaningful taxon circumscriptions. Sten¬
anthium rhomhipetalum (never formally transferred
to Stenanthella ) has been reduced in synonymy un¬
der Anticlea occidentalis by Utech (1987a). In ad¬
dition, the North American A. occidentalis and the
Asian A. sachalinensis may be eonspecifie (Kup-
ehan et ah, 1961), a situation that merits further
study.

Volume 12, Number 2
2002

Zomlefer & Judd
Reconsideration of Zigadenus

303

Antielea frigid a (Schlechtendal & Chamisso)
Zomlefer & Judd, comb. nov. Basionym: Ve-
ratrum frigidum Schlechtendal & Chamisso,
Linnaea 6: 46. 1831. Zigadenus frigidus
(Schlechtendal & Chamisso) D. Don, Edin¬
burgh New Bliilos. J. 13(26): 233. 1832. Sten-
anthium frigidum (Schlechtendal & Chamisso)
Kunth, Enum. PI. 4: 190. 1843. TYPE: Mex¬
ico. Veracruz: Ml. Orizaba, F. Deppe & C. ./.
W. Schiede 983 (holotype, HAL; isotypes,
HAL, MO).

Ltech (1987b: 207) listed “fragment (ill!" as a
“type" specimen (category of type not specified).
However, the handwritten notation on this frag¬
ment packet, “ Veratrum frigidum Cham.! &
Schlech.” followed by “Schiede” lacks detail to
positively identify it as Deppe A' Schiede 983.

Antielea hintonioruni (B. L. Turner) Zomlefer &
Judd, comb. nov. Basionym: Zigadenus hinto-
niorum B. L. Turner. Phytologia 72: 378. 1992.
TV PE: Mexico. Nuevo Leon: Mpio. Galeana,
Cerro Potosf, G. B. Hinton 17200 (holotype,
TEX).

Antielea niogollonensis (W. J. Hess & Sivinski)
Zomlefer & Judd, comb. nov. Basionym: Zi¬
gadenus mogollonensis W. J. Hess & Sivinski,
Sida 16: 390. 1995. TYPE: U.S.A. New Mex¬
ico: Catron Co., Mogollon Mts., Gila National
Forest, W. J. Hess 2212 (holotype, MOB; iso¬
types, NY; also ARIZ, NCU, NMC. OKI. A.
SMU. US not seen).

Antielea neglecta (Espejo, A. R. Lopez-Ferrari &
Ceja) Zomlefer & Judd, comb. nov. Basionym:
Zigadenus neglectus Espejo, A. R. Lopez- Fer-
rari & Ceja, in I). M. Frame et al., Acta Rot.
Mex. 48: 47. 1999. TYPE: Mexico. Guanajua¬
to: Mpio. Xichu, Puerto del Manzanar. Sierra
de Xichu, E. Ventura & E. Lopez 9500 (holo¬
type, UAM1Z not seen; isotype, IEB.)

Antielea oecidentalis (A. Gray) Zomlefer & Judd,
comb. nov. Basionym: Stenanthium occide.nl ale
A. Gray, Proc. Amer. Acad. Arts 8: 405. 1873.
Stenanthella oecidentalis (A. Gray) Rydberg,
Bull. Torrey Bot. Club 27: 531. 1900. TYPE:
Rocky Mountains, Palliser’s British North
American Exploratory Expedition, 1858, E.
Bourgeau s.n. (lectotype, designated by Utech
(1987a), GH; isolectotypes, NY; also B, K not
seen).

Stenanthium rhombipetalum Suksdorf, Werdenda 1 : 6.

1923. TYPE: U.S.A. Washington: Skamania Co..
Cape Horn, W. Suksdorf 10,406 (lectotype, designat¬
ed by Utech (1987a), GH: isolectotypes, MO, NY:
also DS, UC, WTU not seen).

Antielea saehalinensis (F. Schmidt) Zomlefer &
Judd, comb. nov. Basionym: Stenanthium
sachalinense F. Schmidt, Reis. Amur-Land.,
Bot. 188. 1868. Stenanthella sachalinensis (F.
Schmidt) Rydberg, Bull. Torrey Bot. Club 27:
530. 1900. TYPE: Russia. “Ins. Sachalin” (Sa¬
khalin Island]: Ktausi, I 1 July 1860. F.
Schmidt s.n. (lectotype, designated here, LE;
photograph of lectotype. GA).

In his protologue, Schmidt (1868: 188) cited four
collections from different localities on Sakhalin
Island in I860: “Dui” (June, flowering), “Adng-
iwo” (July, (lowers withered/fallen), “Ktausi” (1 I
July, immature fruit), and “Mamie" (August,
flowers withered/fallen). We have selected a
specimen at LE, the principal depository of his
collections (Stafleu & Cowan, 1985). Other prob¬
able isolectotypes (at GH, LE, and NY) have only
"Ins. Sachalin, Exped Fr. Schmidt” on the labels.

3. Stenanthium (A. Gray) Kunth. Enum. PI. 4:
189. 1843, nom. cons. Veratrum subg. Sten¬
anthium A. Gray. Ann. Lyceum Nat. Hist. New
York 4: 119. 1837. Anepsa Rafinesque, FI. Tel¬
lur. 2: 31. 1836 [1837], TYPE: Stenanthium
angustifolium (Pursh) Kunth | basionym: Vera¬
trum angustifolium Pursh],

Stenanthium gramineum (Ker Gavvler) Morong is
synonymous with, and has priority over, S. an¬
gustifolium (Pursh) Kunth. Veratrum angustifol¬
ium, described by Pursh (1814) and transferred
to Stenanthium by Kunth (1843), is the same
plant previously described by Gawler (1813), He-
lonias graminea Ker Gawler. subsequently trans¬
ferred to Stenanthium by Morong (Britton et al..
1894). There is also some confusion in the lit¬
erature concerning Anepsa. Rafinesque (1836)
based Anepsa on lour species — two transferred
from Veratrum and Stenanthium (synonyms in¬
dicated by Rafinesque, 1836: 2: 31), as well as
two others with unknown affinity (see Cowley et
al.. 2001): (1) Anepsa carinata Rafinesque [ Ve¬
ratrum angustifolium Pursh = Stenanthium gra¬
mineum (Ker Gawler) Morong; see Zomlefer,
1997]; (2) Anepsa latifolia Rafinesque ( Veratrum
parviflorum Michaux); (3) Anepsa spicata Rafin¬
esque (species of unknown affinity); and (4)
Anepsa graminifolia (species of unknown affini¬
ty). Anepsa, a genus of mixed elements, has con¬
sequently been considered a nomenclatural syn-

304

Novon

onym of Stenanlhium (Greuter et al., 2(K)0) ami
of Veratrum (Cowley et al., 2001). However, when
Rafinesque (1836: 4: 27) stated in a later part of
the same work . . his [Gray’s] Stenanthium S.
G. [subgenus] of Veratrum is my G. [genus] Anep-
sa ,” he provided a man. ami stat. nov. for Vera¬
trum subg. Stenanthium, and the type of this sub-
genus (V. angustifolium) is. therefore, the type of
Anepsa (K. Gandhi, pers. comm.).

Tracyanlhus Small, FI. S.E. U.S. 250. 1903. Syn. nov.

TYPE: Tracyanthus angustifolius (Michaux) Small.
Oceanoros Small, FI. S.E. ll.S. 252. 1903. Syn. nov.

TYPE: Oceanoros leimanthoides (A. Gray) Small.

Salient features. Slender bulb, cuneate to grad¬
ually and slightly narrowed (but not clawed) tepal
bases, 1 obscure (or lac king) gland per tepal, su¬
perior to half-inferior ovary; base chromosome
number* = 10 (reported only for S. gramineum).

Distribution. Southeastern United States (coast¬
al plain and mountains) in acidic coniferous forests
and bogs.

Until the molecular investigation (Zomleferet al.,
2001), the relationship of Stenanthium s.l. to Zi-
gadenus s.l. was unknown, although in phenetic
analyses by Ambrose (1975, 1980) S. gramineum
clustered with several Zigadenus species, 'file re¬
sults of Zomlefer et al. (2001) strongly support a
markedly different circumscription of the tradition¬
al Stenanthium: the type clement, Stenanthium gra¬
mineum, forms a clade with Zigadenus densus and
Z. leimanthoides (while the species of the Stenan-
thium segregate Stenanthella resolve with Anticlea).
Autapomorphies for this expanded concept of Sten¬
anthium are a slender (cylindrical) bulb, loss/re¬
duction of the tepal glands, and possibly also a base
chromosome number of * = 10 (reported only lor
S. gramineum ; Miller, 1930; Sato, 1942).

Therefore, this new definition of Stenanthium ne¬
cessitates the transfer of two former Zigadenus spe¬
cies, Stenanthium densum and S. leimanthoides.
These taxa have been separated from the rest of the1
Zigadenus complex as segregate genera (Oceanoros
and Tracyanthus: Small. 1903) and considered
within Zigadenus as “section Oceanoros ” (Preeee,
1956; Schwartz, 2002). (As of this writing, this sec¬
tional name has not been validly published.) Tra¬
ditionally, plant size and inflorescence type have
been used to distinguish these two sympatric and
closely related species: specimens of Stenanthium
leimanthoides at F LAS, GA, GH. MO, and NO arc
strikingly much more robust plants with compound
racemose inflorescences in comparison to the more
delicate S. densum with unbranched racemes (W.
Zomlefer, pers. obs.). However, field studies of in¬

tergrading populations by McDearman (1984) sug¬
gest that these two taxa are likely conspecific (sum¬
mary in Zomlefer, 1997). Some authors have placed
S. leimanthoides in synonymy with S. densum (M.
MacRoberts, pers. comm.; Schwartz, 2002; Sorrie
& Weakley, 2001); varietal status of .S’, leimanthoi¬
des has also been suggested (R. \\ underfill, pers.
comm.). The status of S. leimanthoides merits ad¬
ditional systematic study, as well as determination
of the chromosome number.

Stenunlliium densum (Desrousseaux) Zomlefer &
Judd, comb. nov. Basionym: Melanthium den¬
sum Desrousseaux, in Lamarck, Encycl. 4: 26.
1796 [1797]. Zigadenus densus (Desrous¬
seaux) Fernald, Rhodora 42: 254. 1940.
TYPE: U.S. A. Carolina: no date, J. Fraser s.n.
(holotype, P-LA, microfiche 6207.654: II 1.3).

Helonias angustifolia Michaux, FI. Bor.-Amer. 1: 212.
1803. Amianlhium angustifolium (Michaux) A. Gray,
Ann. Lyceum Nat. Hist. New York 4: 124. 1837.
Zigadenus angustifolius (Michaux) S. Watson, Proc.
Amer. Acad. Arts 14: 280. 1879. Tracyanthus an¬
gustifolius (Michaux) Small, 11. S.F. U.S. 251. I'M).!.
TYPE: U.S. A. “Basse Carolina” [South Carolina]: “in
herbosis et fruticetis sylvarum humidis Carolinae in-
ferioris,” no date, J. Fraser s.n. (holotype, P-M1GHX,
microfiche 6211.48: 1.2).

See Fernald (1940) for details about the types
for both basionyms.

Stenanthium leimanthoides (A. Gray) Zomlefer
& Judd, comb. nov. Basionym: Amianlhium
leimanthoides A. Gray, Ann. Lyceum Nat. Hist.
New York 4: 125. 1837. Zigadenus leiman¬
thoides (A. Gray) A. Gray, Manual (ed. 2) 476.
1856. Oceanoros leimanthoides (A. Gray)
Small, FI. S.E. U.S. 252, 1328. 1903. TYPE:
U.S. A. New' Jersey: Camden Co., “the other
side of Haddington,” 20 July 1837, E. M. Du¬
rand s.n. (lectotype, designated here. GH; is-
olectotype, NY).

Gray (1837) cited three collections in his proto-
logue (see discussion of these paratypes in
Walsh, 1940, and Preeee, 1956). Preeee (1956)
chose the Durand specimen at GH as a “lecto¬
type” in his unpublished dissertation, and we
here follow his decision.

Paratypes. U.S. A. North Carolina: Burke Co., Table
Mountain, 1835, M. A. C.urtis s.n. (GH, NY). Louisiana:
Orleans Co., New Orleans, 1834, [l)r.| Ingalls s.n. (NY).

4. Toxicoscordion Rydberg, Bull, lorrey Bot.
Club 30: 272. 1903. TYPE: Toxicoscordion in¬
termedium (Rydberg) Rydberg.

Volume 12, Number 2
2002

Zomlefer & Judd
Reconsideration of Zigadenus

305

Chitonia Salisbury, Gen. PI. 51. 1866, nom. illeg., non
Chitonia I). Don 1823 (Melastomataceae).

Chitonia has often been cited as a synonym for
Zigadenus s.l. (Walsh, 1940: Preece, 1956; Cow¬
ley et ah, 2001). Baker (1880) adopted this name
as the basis lor erecting Zigadenus sect. Chitonia
(Salisbury) Baker and included species of Toxi¬
coscordion and Stenanthium. Salisbury (1866)
did not mention any species in the protologue or
in his discussion of the genus. No specimens are
labeled Chitonia or Zigadenus among Salisbury’s
few remaining collections at BM (R. Vickery,
pers. comm.), the repository of his collections
(StaHeu & Cowan, 1985). Although the tepal
glands are not mentioned, the description of the
plant (Salisbury, 1866: 51) from the “west coast
ol North America” with “petals attenuated into a
. . . claw” best agrees with the characteristics of
Toxicoscordion .

Salient features. Ovoid bulb, conspicuously
clawed tepal bases, 1 obovate gland per tepal, su¬
perior ovary; base chromosome number x = 1 1 .

Distribution. Midwestern United States and
western North America in acid and/or alpine co¬
niferous forests, prairies, desert, chaparral, or ser¬
pentine vegetation.

Molecular and morphological data resolve Toxi¬
coscordion as distinct from the rest of the Zigadenus
complex (Zomlefer et ah, 2001). In their unpub¬
lished dissertations, both Preece (1956) and
Schwartz (1994) treated this taxon as Zigadenus
sect. Chitonia (Salisbury) Baker (see also Schwartz.
2002). The resurrection of Toxicoscordion requires
one new combination (T. fontanum ), a species de¬
scribed within Zigadenus by Eastwood (1937), long
after Rydberg ( 1903) had erected the segregate Tox¬
icoscordion and had transferred the appropriate
contemporary Zigadenus species (Table 2). Auta-
pomorphies include conspicuously clawed tepals
(especially the inner three), one obovate gland per
tepal, and a base chromosome number of 1 1 (Zom¬
lefer et ah, 2001). A citation of 2 n = 32 for 7.
nuttallii (Fedorov, 1969; Moore, 1971, 1973). orig¬
inally reported by Zakharieva and Makushenko
(1969), is based on an undescribed and unvoueh-
ered plant then growing at the Munich Botanical
Garden (original source not cited); this “ Zigaden¬
us ” species was likely an Anticlea (x = 8).

The species of Toxicoscordion (Table 2), restrict¬
ed to midwestern— western North America, include
the well known poisonous “death eamas” plants of
the rangelands such as T. nuttallii , T. paniculatum,
and T. venenosum (see Marsh et ah, 1915. 1926).
Further study is needed to determine appropriate

specific and infraspecific limits within the variable
T. mieranthum-T. fremontii species complexes, as
well as the overlapping T. paniculatum- T. venenos-
um complex. For example, in an isozyme study of
these species (Schwartz, 1994), the monophvly of
T. paniculatum is suspect because the five sampled
populations were separated from each other in
UPGMA cluster analyses and distance Wagner
trees.

Toxicoscordion fontanum (Eastwood) Zomlefer
& Judd, comb. nov. Basionym: Zigadenus fon-
tanus Eastwood, Leaf!. W. Bot. 2: 41. 1937.
Zigadenus micranthus Eastwood v ar. font anus
(Eastwood) 0. S. Walsh ex McNeal, Phytologia
73: 308. 1992. TYPE: U.S.A. California: Ma¬
rin Co., Bootjack, Mt. Tamalpais, ./. T. Howell
12656 (holotype, CAS).

5. Zigadenus Michaux. FI. Bor.-Amer. 1: 214, t.
22. 1803. TYPE: Zigadenus glaberrimus Mi¬
chaux.

Salient features. Rhizome (no bulb), slightly
(and more or less gradually) narrowed tepal bases,
2 ovate glands per tepal, superior ovary, unusual
anatomical features; base chromosome number x —
26(?).

Distribution. Southeastern United States (coast¬
al plain) in acidic coniferous forests and bogs.

Zigadenus, as defined here, comprises only Zi¬
gadenus glaberrimus (the type and the only species
in the protologue). a circumscription advocated his¬
torically by several botanists (e.g.. Rydberg. 1903;
Small, 1903, 1933; Gates, 1918). Molecular data
(Zomlefer & Perkins, 1999; Zomlefer et ah. 2001)
confirm a monospecific circumscription of Zigaden¬
us and its divergent and isolated position as the
sister taxon of the remaining members of the tribe
Melanthieae. Autapomorphies of this distinctive
taxon (discussed in Zomlefer, 1997) include an un¬
usual chromosome number ( n — 26, tentatively re¬
ported by Preece, 1956), a rhizome lacking a bulb,
two ovate glands per tepal, and several anatomical
features (foliar stomata with two aperture lips, dis¬
tinct root exodermis, bracteolate pedicels, and
dense tannin-like inclusions: Ambrose, 1975).

The original spelling of the genus has priority
over the orthographic variant “ Zygadenus ” (from
Endlicher, 1836), commonly used in herbaria and
some older floras (e.g.. Small, 1903, 1933; Gleason,
1952). Cyanotris Rafinesque, nom. rej. against Ca-
massia Lindley (Agavaceae), is miscited as a syn¬
onym of Zigadenus in Preece (1956).

306

Novon

Taxa Inckktak Skims

Monadenus R. A. Salisbury, Gen. PI. 51. 1866.

This taxon is often cited as a synonym of Ziga¬
denus s. 1. (e.g., Walsh, 1940; Preece, 1956; Cowley
et al., 2001), and therelore is presumably a syno¬
nym for one of the genera in this paper. Salisbury
(1866) did not, however, mention any species or
make any species combinations. No specimens are
labeled Monadenus or Zigadenus among Salisbury’s
few remaining collections at RM (R. Vickery, pers.
comm.), the repository of his collections (StaHeu &
Cowan, 1985). The description of the plant (from
somewhere in America and cultivated at a nursery
in England) is ambiguous: the partly inferior ovary
may indicate placement in Anticlea (rather than
other segregates of Zigadenus), but each tepal lias
“one melliferous semi-circular cavity" at the base
(Salisbury, 1866: 52) and not the characteristic
conspicuous bilobed gland of Anticlea. Another
possibility is Stenanthium gramineum, character¬
ized by a large paniculate inflorescence with "fine
slender branches” (as Salisbury mentioned) and a
partly inferior ovary — but this species lacks any
glandular tissue or cavities on the tepals. I herefore,
the description of Monadenus cannot be matched
with certainty to any speeies or segregate of the
Zigadenus assemblage.

Gornphostylis Rafinesque, 11. Tellur. 2: 30. 1836
[1837].

This taxon has been associated with elements in
the Zigadenus complex (Walsh, 1940; Zimmerman.
1958; Cowley, 2001), considered here in Anticlea ,
Toxic oscordion, and Zigadenus s. str. Rafinesque
(1836: 30) based Gornphostylis on three mixed el¬
ements, two of them indicated as ambiguously
placed with a “?”: G. bracteata Rafinesque [= “He-
lonias bracteata Rrereton mpt.' |. “Gornphostylis l
paniculata" Rafinesque [= Toxicoscordion panicu-
latum (Nuttall) Rydberg], and “Gornphostylis l or Z.
fuscatus" Rafinesque |an unidentified name; see
Preece, 1956]. The “?” notation for the two latter
species indicates that Rafinesque was unsure of
their placement in Gornphostylis. and therefore, G.
bracteata (which lacks a “?”) would be the type of
the genus (K. Gandhi, pers. comm.). However, the
identity of this species is unclear. The short and
vague description for this Virginian plant is not suf¬
ficient to distinguish the species. Three species of
the Zigadenus complex occur in Virginia: Anticlea
elegans (Pursh) Rydberg, Stenanthium leimanthoi-
des (A. Gray) Zoinlefer & Judd, and Zigadenus gla-
berrimus Michaux; all lack the spicate inflorescence

described by Rafinesque (racemose panicles in An¬
ticlea elegans and Stenanthium leimanthoides, and
spreading paniculate inflorescences in Z. glaberri-
mus). Helonias bracteata Sims is Zigadenus glaber-
rimus (rhizome with no bulb, two glands per tepal;
see Sims, 1815), and Preece (1956) included "G.
bracteata Raf.” [properly G. bracteata (Sims) Raf¬
inesque, since cited in synonymy with Sims’s name|
as a synonym for Z. glaberrimus. However, “ Helon¬
ias bracteata Rrereton mpt. ' cited by Rafinesque is
a manuscript name with no nomenclatural standing.
Therefore, the name Gornphostylis cannot be ap¬
plied with certainty to any known taxon.

The following key utilizes field characters and
institutes a new taxonomy of the former Zigadenus
complex.

Kfa to Segregate Gkni.ua ok iiii Former Zigadenus
Complex

la. Plants witli rhizome (lacking bulb), 2 conspicu¬
ous glands per tepal . Zigadenus Michaux

lb. Plants witli bulb, 1 gland per tepal or nectaries
obscure or absenl.

2a. Tepal glands obscure to absent.

3a. Bulb ovoid, ovary superior, seeds with

red to purple sarcotesta .

. Amianthium A. Gray

3b. Bulb slender (cylindrical), ovary supe¬
rior to half- inferior, seeds brown and

lacking sarcotesta .

. Stenanthium (A. Gray) Kunth

2b. Tepal glands conspicuous.

4a. Tepals conspicuously clawed, tepal
glands obovate, ovary superior ....

. Toxicoscordion Rydberg

4b. Tepals cuneate to gradually narrowed at
base (not clawed), tepal glands bilobed,
ovary' half-inferior . Anticlea Kunth

Acknowledgments. We thank the following li¬
brarians for help with problematical references:
Judy Warnement and Jessica Thomas (Harvard
University botanical libraries); Connie Wolf and
Patricia Carroll (Missouri Botanical Garden Li¬
brary); and John Reed and Marie Long (LuEsther
T. Merz Library, New York Botanical Garden). Lisa
Campbell, Noel Holmgren. Patricia Holmgren, Sa¬
rah Hunkins, Jackie Kallunki, Barbara Thiers, and
Tom Zanoni (all NY), Emily Wood ((ill). Kent 1).
Perkins (FLAS), and James Solomon (MO) gra¬
ciously assisted WBZ at their respective herbaria.
We are grateful also to K. N. Gandhi for reviewing
the nomenclatural citations: Roy Vickery for check¬
ing holdings of Salisbury specimens at RM; Dmitry
V. Geltman (LE), Uwe Rraun (HAL), and Paul E.
Ret ry (W IS) for information on type specimens at
their respective institutions; Robert J. Liebermann
for translating Russian text and for photographing

Volume 12, Number 2
2002

Zomlefer & Judd
Reconsideration of Zigadenus

307

type specimens at LE; Millie L. Turner for infor¬
mation on the basionym of Anticlea hintoniorum ;
and the curators/collection managers of CAS, (ill.
HAE. I EH. MO. MOR, NY, and TEX for loans to
WBZ of type material. Diana R. Gunter, Victoria C.
Hollowed. Amy S. McPherson, James E. Reveal,
and Frederick IE Utech suggested improvements to
the text. The University of Georgia Department of
Plant Biology generously provided funds to WRZ
for travel to FEAS, GH. MO, and NY.

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ume 12, Number 2, pp. 155-308 of NOVON was published on 8 July 2002.

Volume 12
Number 3
2002

NOVON

New Species of Alyssum , Aphragmus, Arabis, and Sinosophiopsis
(Brassicaceae) from China and India

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. The new species Alyssum hlirnesii and
Aphragmus ladakiana from Ladak (northwestern
India), Arabis setosifolia from Xizang (China), and
Sinosophiopsis furcata from Sichuan (China) are de¬
scribed, and their relationships to the most closely
related species of their respective genera are dis¬
cussed.

Key words: Alyssum , Aphragmus, Arabis, Bras¬
sicaceae, China, India, Sinosophiopsis.

The following four new species are based on ma¬
terial collected from Ladak (India) by Leos klimes
(Institute of Botany, Section of Plant Ecology, Tre-
bon, Czech Republic) and on Chinese herbarium
specimens at KLN examined during my visit in Au¬
gust 2001. The paper represents part of the author’s
ongoing research on the Brassicaceae of the Hi¬
malayan flora.

Alyssum klimesii Al-Shehbaz, sp. nov. TYPE: NW
India. Ladak: Rupshu, Lapgo River Valley,
5600 m, 32°59'N, 78°28'E, 12-13 July 2000.
Leos KlimeS s.n. (holotype, MO #5317019).

Herba perennis, subpulvinata, tomentosa, eanescens,
pilis subdendriticis, longe stipitatis. Folia dense rosulata,
persistentia, camosa, sessilia, obovata vel spathulata, 3—
10 X 1—2 mm, dense tomentosa. Racemi 2 — 4-fR)ri, ebrac-
teati. Petala rosea vel alba cum unguibus roseis, spathu¬
lata, 2—3 X 0.8— I mm. Ovula 4, subapiculata. Siliculae
ovoideae, inflatae, dense tomentosae, 2—3 X 1.5-2 mm,
apice acuto-acuminatae; stylus 0.5— 0.8 mm longus. Sem¬
ina oblonga, 1.2— 1.5 X 0.4— 0.5 mm.

Herbs perennial, somewhat pulvinate, 1—3 cm
tall, silvery canescent and tomentose throughout;

trichomes long-stalked, subdendritic, with finely
branched rays. Stems much abbreviated, distinct
and leafless sometimes in infructescence, terminat¬
ing caudex branches. Leaves grouped in compact
rosettes, persistent, fleshy, sessile; leaf blade ob-
ovate to spatulate, 3-10 X 1-2 mm, uniformly
densely tomentose on both surfaces, base attenuate,
apex obtuse. Raceme 2— 4-flowered; peduncle dis¬
tinct and fruits borne to 1 cm above leaf rosette,
sometimes obsolete and fruits hardly exserted
above rosette. Fruiting pedicels ascending, 0.5—2
mm long, slender, densely tomentose all around.
Sepals oblong, 1.5—2 X 1-1.5 mm, persistent, stel¬
late. Petals pink throughout or white with pink
claws, spatulate, 2—3 X 0.8-1 mm, glabrous out¬
side, caducous, apex rounded; claw pink, not pa¬
pillate at base. Filaments 1.5—2 mm long, dilated
and glabrous at base, neither winged nor appen-
daged; anthers ovate, 0.2— 0.4 mm long. Ovules 4
per ovary, subapical, pendulous on long funicles.
Fruit ovoid, 2—3 X 1.5—2 mm; valves not veined,
inflat ed, densely tomentose outside, glabrous in¬
side, base rounded, apex acute-acuminate; style
0.5— 0.8 mm long, slender, glabrous, pink, exserted.
Seeds oblong, 1.2— 1.5 X 0.4— 0.5 mm; cotyledons
obliquely accumbent.

Alyssum klimesii is named in honor of Leos Kli-
meS, collector of the holotype who is conducting
ecological studies in the alpine areas of Ladak (In¬
dia). The species is most closely related to A. ca-
nescens DC. of China, Kashmir, Kazakstan, Mon¬
golia, and Russia (Cheo et al., 2001). They
resemble each other in being pulvinate, canescent

Novon 12: 309-313. 2002.

310

Novon

perennials and in having fleshy leaves and similar
flower, fruit, and seed size. Alyssum klimesii is eas¬
ily distinguished by having long-stalked subden-
dritic trichomes, fruiting pedicels 0.5—2 mm long,
persistent sepals, spatulate petals 0.8—1 mm wide,
glabrous bases of petal claws and staminal fila¬
ments, and inflated, ovoid fruits. By contrast, A.
canescens has subsessile or short-stalked stellate
trichomes, fruiting pedicels (3— )4— 7 mm long, de¬
ciduous sepals, obovate petals 1 .5-2.5(-3) mm
wide, papillate bases of claws and filaments, and
slightly flattened fruits.

In the Chinese paratypes listed below, the petals
have white blades and pinkish claws much like
those of A. canescens. However, in the collections
from Ladak the petals are pink throughout. Because
of the limited number of collections examined, it is
not possible to determine whether or not such a
difference merits taxonomic recognition.

Because of its pulvinate habit, evanescent, dense¬
ly tomentose leaves, and few-seeded ovoid fruits,
one might confuse Alyssum klimesii with some spe¬
cies of Draba. However, none of the Himalayan
species of Draba has ovaries with 4 subapical
ovules pendulous on long funicles, pink petals or
petal claws, and fruit valves as densely tomentose
as the rest of the plant, whereas many species of
Alyssum have the very same combination of char¬
acters.

Paratypes. CHINA. Xizung (Tibet): Zhongba Xian.
Longgeqi, Qinghai-Tibet Team 6591 (KUN, PE). NW IN¬
DIA. Ladak: Rupshu, ca. 5800 m, 33°10'N, 78°I2.5'E,
9 July 2000, Leas KlimeS s.n. (MO); Rupshu, Kamdar ha.
5700 m, 33°10.5'N, 78°14'E, 9 July 2000, Leas KlimeS
s.n. (MO); Rupshu, ca. 5900 m, 32°59'N, 78°29.5'E, 14
July 2000, Leos KlimeS s.n. (MO).

Aphragmus ladakiana Al-Shehbaz, sp. nov.
TYPE: NW India. I.adak: Rupshu, Parang Val¬
ley, 4860 m, 32°30'N. 78°06'E, 25 July 2000,
Leas KlimeS s.n. (holotype, MO # 5317023).

Herba perennis, 1—3 cm alia, puberula, pil its simplici-
bus usque 0.5 mm longis. Folia basalia rosulata, subcar-
nosa, oblanceolata, 5—12 X 1.5—2 mm, integra, retrorse
puberula. Raeemi 3— 6-flori, corymbosi, ebracteati. Petala
alba, late spathulata, 2.5—3 X 0.7— 0.9 mm. Ovula 28—32.
Siliquae lineares, teretes, 15—17 X 1—1.2 mm, laeves;
gynophorum 0.2— 0.7 mm longum; stylus 0.5— 0.8 mm lon-
gus; septum longitudinaliter secedens. Semina oblonga,
uniseriata, 0.6— 0.8 X 0.3— 0.4 mm.

Herbs perennial, 1—3 cm tall; caudex few-
branched. Stems erect, simple, puberulent with
simple trichomes to 0.5 mm long. Basal leaves ro-
sulate, subfleshy; petiole-like base not expanded;
leaf blade oblanceolate, 5—12 X 1.5—2 mm, re-
trorsely puberulent, base attenuate, margin entire.

apex obtuse. Cauline leaves absent. Racemes 3-6-
flowered, corymbose, ebracteate, slightly elongated
in fruit. Fruiting pedicels ascending, 2—3 mm long,
puberulent all around. Sepals 1.5—2 X ca. 0.6 mm,
puberulent. Petals white, broadly spatulate, 2.5—3
X 0.7— 0.9 mm, apex rounded. Filaments 1-1.5 mm
long; anthers oblong, ca. 0.5 mm long. Ovules 28
to 32 per locule. Fruit linear, 15-17 X 1-1.2 mm,
terete, smooth; valves distinctly veined along prox¬
imal half, glabrous; gynophore 0.2— 0.7 mm long;
septum split longitudinally, hyaline; style 0.5— 0.8
mm long; stigma entire. Seeds yellow-brown, ob¬
long, uniseriate, 0. 6—0. 8 X 0.3— 0.4 mm. Cotyle¬
dons incumbent.

Aphragmus ladakiana is closely related to A. ob-
scurus (Dunn) 0. E. Schulz, a species restricted to
Sonamarg, Kashmir (Al-Shehbaz, 2000a). From that
species, A. ladakiana is easily distinguished by
having a distinct caudex, leafless stems, smooth
fruits, hyaline septum split longitudinally, 28- to
32-ovuled ovaries, ebracteate racemes, fruiting
pedicels puberulent all around, and smaller seeds
0.6— 0.8 X 0.3— 0.4 mm. Aphragmus obscurus has
slender rhizomes, leafy stems, torulose fruits, no
septum, 5- to 10-ovuled ovaries, fully braeteate ra¬
cemes, adaxially puberulent and abaxially glabrous
fruiting pedicels, and seeds 1.2- 1.5 X 0.7— 0.8 mm.
With the discovery of A. ladakiana , Aphragmus
now includes six species, the previous five of which
are dealt with by Al-Shehbaz (2000a).

Paratype. NW INDIA. Ladak: Rupshu, ca. 5800 m,
32°43'N, 77°55.5'E, 31 July 2000. Leos KlimeS s.n. (MO).

Arabis setosifolia Al-Shehbaz, sp. nov. TYPE:
China. Tibet (Xizang): Chaya Xian, Jitang,
rocky roadside, 3700 m, 13 July 1976, Qing¬
hai-Tibet Team 12388 (holotype, KLIN; iso¬
type, KUN). Figure 1.

Herba perennis scaposa, 4—7 cm alia. Caules, pedicelli
et sepala puberula, pilis furcatis brevi-stipitatis, 0. 1-0.4
mm longis. Folia basalia rosulata, oblanceolata, 2-3.5 cm
X 3-6.5 mm, pilis simplicibus setosis usque 1.5 mm lon¬
gis. Folia caulina carentia vel unum. Raeemi ebracteati.
Sepala 2.5—3 mm longa. Petala rosea, oblanceolata, 5—6
X 2-2.5 mm. Ovula 11-14. Siliquae lineares, compres-
sae, 1.8— 2.3 cm X 1—1.2 mm, glabrae; stylus usque 0.3
mm longus. Semina oblonga 1.3— 1.5 X ca. 0.7 mm, un¬
iseriata.

Scapose perennial herbs 4-7 cm tall, without
leaf remains of previous years. Trichomes on stems,
pedicels, and sepals short-stalked, forked, 0.1— 0.4
mm long. Stems erect, puberulent. Basal leaves ro-
sulate, oblanceolate, 2-3.5 cm X 3—6.5 mm, setose
with simple trichomes to 1.5 mm long, base atten¬
uate, margin entire, apex obtuse. Cauline leaf 1 or

Volume 12, Number 3
2002

Al-Shehbaz

New Species from China

311

Figure 1. Arabia setosifolia Al-Shehbaz. — A. Plant. — B. Portion of leaf margin with setose trichomes. — C. Trichomes
of stems and pedicels. — D. Petal. — E. Fruit and fruiting pedicel. Scale: A = 1 cm; B, E = 5 mm; C = 0.2 mm; D
= 1 mm. Drawn by Al-Shehbaz: A-l) from the holotype, Qinghai-Tibet Team 12388 (KUN); F from the paratype, Yan
Jinsheng 91-492 (KUN).

absent, oblong, sessile, not aurieulate, 7—13 X 1—
2 mm. Raceme ebracteate. Fruiting pedicels 3.5—7
mm long, ascending, densely puberulent, straight
or curved upward. Sepals oblong, 2.5—3 X ca. 1.5
mm, purple tinged, sparsely puberulent to subgla-
brous. base not saccate, margin membranous. Pet¬
als pink, oblanceolate, 5—6 X 2—2.5 mm, obtuse,
undifferentiated into blade and claw. Filaments
2.5—3 mm long; anthers ovate-oblong, ca. 0.7 mm
long. Ovules 1 1 to 14 per locule. Fruits linear, com¬
pressed, 1.8— 2.3 cm X 1—1.2 mm; valves glabrous,
with a prominent midvein extending full length;

style obsolete or minute and to 0.3 mm long. Seeds
oblong, 1.3— 1.5 X ca. 0.7 mm, uniseriate, oblong;
cotyledons accumbent.

Of the 14 species of Arabis L. recognized by
Cheo et al. (2001) in China, none has a combina¬
tion of setose basal leaves, puberulent stems and
pedicels with minutely forked trichomes, stems
only 3.5—7 cm tall, and wingless seeds. The Chi¬
nese endemic A. alaschanica Maximowicz (Gansu,
Nei Mongol, Ningxia, Qinghai, Shanxi, and Sichuan
Provinces) also has setose basal leaves, but these

312

Novon

Figure 2. Sinosophiopsis furcata Al-Shehbaz. — A. Portion of plant. — B. Leaf triehomes. — C. Sepal. — D. Petal. —
E. Median stamen. Scale: A = 1 cm; B = 0.4 mm; C— E = 1 mm. Drawn by Al-Shehbaz from the holotype. Jinn Shu
5031 (KUN).

Volume 12, Number 3
2002

Al-Shehbaz

New Species from China

313

hairs are mixed with short-stalked forked ones. It
also differs from A. setosifolia by having caudices
covered with stramineous, persistent petioles of
previous years, slender styles 1—3 mm long, and
larger seeds (1.4— 2.5 X 0.8— 1.4 mm) that are
winged distal ly. Arabis setosifolia is probably most
closely related to A. saxieola Edgeworth (Afghani¬
stan, India, Kashmir, Pakistan), from which it dif¬
fers by having erect, subappressed fruits and entire
leaves hirsute on both surfaces with simple, sub-
setose trichomes. Arabis saxieola has divaricate to
divaricate-ascending fruits and serrulate basal
leaves pubescent only abaxially with short-stalked,
4-rayed stellate trichomes, anti glabrous adaxially
or sparsely pubescent with stellate or forked tri¬
chomes.

Paratype. CHINA. Tibet: eastern part of Gongjue
Xian, 3650 m, 30 June 1991, Yan Jinsheng 91-492
(KUN).

Sinosophiopsis furcata Al-Shehbaz, sp. nov.
T\ PE: China. Sichuan: Kanding Xian, Sha De
Qu Ma Ti, sunny dry areas near stream, 2700
m, 29 May 1961, Jian Shu 5031 (holotype,
KUN). Figure 2.

Herba annua vel biennis, ultra 30 cm alta. Caules gla-
bri. Folia basalia ignota; folia caulina pinnatisecta, lobis
lateralibus 5—7 jugis, anguste oblongis, 0.7-1. 5 cm X 1-
3 mm, basis subdecurrentibus, pilis sessilis, furcatis et
malpighiaceis praeditis. Pedicelli floriferi tenues, divari-
cato-adscendentes, 1.3—2 cm longi. Sepala 3-4 mm longa,
glabra. Petala alba, spathulata, 4—5 X ea. 2 mm. Siliquae
et semina ignota.

Annual or biennial herbs, more than 30 cm tall,
lower part unknown. Trichomes restricted to leaves
and young stems, sessile, 2- or 3-forked, mixed
with fewer malpighiaceous ones. Stems erect, gla¬
brous. Cauline leaves pinnatisect; petiole 1-2.5 cm
long, glabrous, grooved; lateral leaf lobes 5 to 7 on
eac h side of midvein, narrowly oblong, 0.7— 1.5 cm
X 1—3 mm, base slightly decurrent, proximal mar¬
gin often minutely 1- or 2-toothed, otherwise entire,
apex subacute; terminal lobe longer than lateral,
linear-lanceolate, sessile, decurrent with distal pair
of lateral lobes. Raceme with only lowermost few
flowers bracteate. Flowering pedicels slender, di¬
varicate-ascending, glabrous or sparsely pilose,
1.3-2 cm long. Sepals oblong, 3-4 X ca. 1.5 mm.

erect, glabrous, base nonsaccate. Petals white, spat-
ulate, 4—5 X ca. 2 mm. Stamens strongly tetrady-
namous; median filaments ca. 3 mm long; lateral
filaments ca. 1.5 mm long; anthers oblong, ca. 1.5
mm long. Pistil glabrous; style distinct; stigma cap¬
itate, entire. Fruits and seeds unknown.

Two species of Sinosophiopsis Al-Shehbaz were
previously recognized (Al-Shehbaz, 2000b), and the
Chinese-endemic genus is characterized by the an¬
nual or biennial habit, white flowers, nonsaccate
sepals, branched trichomes, and pinnatisect leaves.
Sinosophiopsis furcata is most closely related to S.
heishuiensis (W. T. Wang) Al-Shehbaz, from which
it differs by having sessile, forked trichomes mixed
with fewer malpighiaceous ones, glabrous stems, 5
to 7 lateral leaf lobes on each side of midvein, se¬
pals 3-4 mm long, and oblong anthers ca. 1.5 mm
long. By contrast, S. heishuiensis has a mixture of
simple and short-stalked forked trichomes, pilose
stems, 2 to 4 lateral lobes on each side of midvein,
smaller sepals 1 .5—2.3 mm long, and ovate anthers
0.4— 0.5 mm long. Both species are readily distin¬
guished from the third species of the genus, S. bar-
tholomewii Al-Shehbaz, by having ebracteate ra¬
cemes or only basally bracteate racemes, and much
longer pedicels (0.7—2 cm long) and petals (4-5
mm long). This last species has fully bracteate ra¬
cemes, and shorter pedicels (l-3(-4) mm long) and
petals (2—2.5 mm long).

Acknowledgments. I am most grateful to Henk
van der Werff for correcting the Latin, to Leos Kli-
me§ (Czech Republic) for sending his Ladak col¬
lections for determination, to Sun Hang and Yue
Jipei for their help and support during my recent
visit to KUN, and to Zhu Guanghua and Song Hong
for their help in the translation of labels from Chi¬
nese.

Literature Cited

Al-Shehbaz. 1. A. 2(K)()a. Staintoniella is reduced to syn¬
onymy of Aphragmus (Brassicaceae). Harvard Pap. Bot.
5(1): 109-112.

- . 2000b. Sinosophiopsis (Brassicaceae), a new ge¬
nus endemic to China. Novon 10: 340—343.

Cheo, T. Y., L. L. Lu, Y. Guang & I. A. Al-Shehbaz. 2001.
Brassicaceae. Pp. 1—193 in C. Y. Wu & P. H. Raven
(editors). Flora of China, Vol. 8. Science Press, Beijing,
and Missouri Botanical Garden Press, St. Louis.

Six New Species of Draba (Brassicaceae) from the Himalayas

Ihsan A. Al-Shehbaz

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

ihsan.al-shehbaz@mobot.org

ABSTRACT. Draba nylamensis, D. kongboiana,
and D. sunhangiana from Tibet (Xizang), China, D.
macbeathiana from Nepal, D. bagmatiensis from
Nepal and adjacent Sikkim (India), and D. hima-
chalensis from India are described, and their dis¬
tinguishing characters from the nearest relatives are
discussed.

Key words: Brassicaceae, China (Tibet), Draba,
India, Nepal.

During recent visits to various herbaria and con¬
tinuing work on the treatment of Brassicaceae (Cru-
ciferae) for the flora of Nepal. I discovered the fol¬
lowing six new species of Draba. They are
described herein to make their names available for
the Flora of Nepal and other works in progress.
Illustrations of these species and scanning-electron
microscopy of the trichomes will be published in a
forthcoming revision of Draba for the Himalayas
and Central Asia.

Draba nylamensis Al-Shehbaz, sp. nov. TYPE:
China. Xizang (Tibet): Nylam Xian, Nylam,
rocky slopes, 3800 m, 23 June 1975, Qinghai-
Xizang Team 5820 (holotype, KUN; isotypes,
KUN, PE).

Herba biennis, 12—30 cm alta. Caules erecti, distaliter
glabri, basaliter dense hispidi, pilis simplicibus usque 2
mm longis. Folia basalia rosulata, subsessilia, oblanceo-
lata vel oblongo-oblanceolata, 0.6—3 cm X 2—8 mm, sub-
tus pilis stipitato-stellatis praeditis, supra pilis plerisque
simplicibus hispidis minoribus stipitato-stellatis praeditis.
Folia caulina (8— )10— 13, sessilia, nonauriculata, oblonga
vel lanceolata. Pedioelli fructiferi 1-2 cm longi, pilis stip¬
itato-stellatis, sparsis. Sepala 2.5—3 mm longa. Petala fla-
va, spathulata, 6-7.5 X 2-2.5 mm. Ovarium glabrum, 60-
70-ovulatum. Fructus immaturi lineares, glabri, 2.5—3 cm
longi, compressi. Semina ignota.

Biennial herbs 12—30 cm tall. Stems solitary or
rarely few from base, erect, simple at base,
branched distally, densely hispid basally with sub-
setose simple trichomes to 2 mm long, these some¬
times mixed with fewer, smaller, stalked forked tri¬
chomes, glabrescent distally. Basal leaves forming
dense rosettes, subsessile; leaf blade oblanceolate
to oblong-oblanceolate, 0.6—3 cm X 2—8 mm, abax-
ially densely pubescent with stalked, 4-rayed stel¬

late trichomes with simple rays, sometimes these
mixed along midvein with much coarser, fewer,
simple or forked trichomes, adaxially with predom¬
inantly simple trichomes mixed with fewer, much
smaller stalked forked ones, base cuneate, margin
entire or I— 3-toothed, ciliate with simple trichomes
to 2 mm long, apex subobtuse. Cauline leaves 10
to 13, rarely as few as 8, sessile, not auriculate,
oblong-lanceolate to oblong, with indumentum sim¬
ilar to basal leaves, margin minutely 1—3-toothed,
apex obtuse to subacute. Racemes 6— 15-flowered,
ebracteate, elongated considerably in fruit; rachis
straight, f ruiting pedicels 1—2 cm long, divaricate-
ascending, slightly curved upward or straight, slen¬
der, sparsely pubescent with stellate stalked tri¬
chomes. Sepals oblong, 2.5—3 X 1—1.5 mm, erect,
abaxially pubescent with stalked stellate trichomes
and fewer simple ones, base of lateral pair not sac¬
cate, margin narrowly membranous. Petals yellow',
spatulate, 6-7.5 X 2—2.5 mm, apex obtuse to sub-
emarginate. Median filaments 3.5—4 mm long, lat¬
eral filaments 2-3 nun long; anthers ovate, ca. 0.5
mm long. Ovules 60 to 70 per ovary. Immature fruit
linear, 2.5—3 cm X ca. 1 mm, not inflated, latisep-
tate, not twisted, straight; valves glabrous, with ob¬
scure midvein, base and apex subobtuse; style ob¬
solete. Mature seeds not seen. Flowering June.

Rocky slopes, grassy slopes; 3650—3800 m.

Draba nylamensis is most closely related to D.
stenocarpa J. I). Hooker & Thomson (Afghanistan.
China, India, Kashmir, Kazakhstan, Kyrgyzstan,
Pakistan, Tajikistan, Turkmenistan, Uzbekistan),
which it resembles in habit, flower color, and in¬
dumentum. However, it is readily distinguished by
having (8-) 10— 13 -leaved stems, glabrous fruits, se¬
pals 2.5—3 mm long, petals 6—7.5 mm long, and
fruiting pedicels sparsely pubescent with stellate
trichomes. By contrast, D. stenocarpa has 2^4(— 6)-
leaved stems, antrorsely puberulent fruits, sepals
1.5—2 mm long, petals 3-4 mm long, and glabrous
fruiting pedicels rarely with few scattered simple
trichomes. Although only immature fruits of D. ny¬
lamensis have been examined, they appear to be
distinctly longer (2.5—3 cm) and with more ovules
per ovary (60 to 70) than those of D. stenocarpa.

Novon 12: 314— 3 IB. 2002.

Volume 12, Number 3
2002

Al-Shehbaz
Himalayan Draba

315

which has fruits (0.6— )0.9— 2(— 2.5) cm long and
ovules 32 to 48 per ovary.

Paratypes. CHINA. Xizang: Nylam Xian, Nylam,
3760 m, 13 June 1966, Zhang Yongtian & Lang Kaiyong
3949 (PF); kharba, 1 1 June 1922, E. J. Norton 85 (K).

Draba macbeathiana Al-Shehbaz, sp. nov. TYPE:
Nepal. Thorong Ea, Marsyandi Valley, stable
scree at the top of the pass, 17,300 ft., 25 July
1983, R. J. I). McBeath 1484 (holotype, E).

Herba perennis, ca. 6 cm alta. Caules erecti, tomentosi,
pilis furcatis et stellatis praeditis. Folia basalia rosulata,
petiolata, oblanceolata, 5-15 X 2-5 mm, pilosa, pilis stip-
itato-stellatis, 3- vel 4-radiatis praeditis. Raeemi 3— 6-flori,
ebracteati, laxi. Pedicelli fructiferi 1—2 cm longi, divari-
cati vel leviter reflexi. Sepala 1.5—2 mm longa. Petala Ha¬
va. spathulata, ca. 3 X 1 mm. Ovula 12—14. Fructus ob-
longi, pubescentes, 8—10 X 3-4 mm, compressi, basi et
apice obtusi. Semina oblonga, ca. 1 X 0.5 mm.

Perennial herbs ca. 6 cm tall, laxly branched.
Caudex covered with leaf remains of previous
years, many branched, ultimate branches terminat¬
ed in few rosettes. Stems erect, simple, tomentose
with stalked, forked and stellate trichomes. Basal
leaves rosulate, persistent; petiole to 10 mm long,
ciliate with simple and stalked forked trichomes;
leaf blade oblanceolate, 5—15 X 2—5 mm, pilose
with stalked, 3- or 4-rayed stellate trichomes with
simple, soft rays, base euneate to attenuate, margin
1- or 2-toothed on each side, apex acute. Cauline
leaves few, similar to basal leaves. Racemes 3-6-
llowered, ebracteate, elongated in fruit. Fruiting
pedicels 1—2 cm long, horizontal to slightly re¬
flexed. straight, slender, tomentose all around;
adaxially with a well-developed axillary gland. Se¬
pals oblong, 1.5-2 X ca. 1 mm, erect, abaxially
sparsely pubescent with stalked stellate trichomes,
base of lateral pair not saccate, margin not mem¬
branous. Petals yellow, spatulate, ca. 3 X I mm,
apex subemarginate. Filaments 1.5-2 mm long; an¬
thers ovate, ca. 0.4 mm long. Ovules 12 to 14 per
ovary. Fruit oblong, 8-10 X 3-4 mm, compressed,
latiseptate, twisted; valves pubescent, not veined,
base and apex obtuse; style 0.5— 0.6 mm long.
Seeds brown, oblong, ca. I X 0.5 mm.

Draba macbeathiana, which is named after the
collector of the holotype, is known only from the
type collection above. It is closely related to D.
humillima O. E. Schulz (Xizang, China, and Sik¬
kim, India), which it resembles in indumentum,
flower color and size, and fruit shape. It is readily
distinguished by being nonpulvinate and laxly
branched and by having toothed, acute leaves, hor¬
izontal to slightly reflexed, slender fruiting pedicels
10—20 mm long and adaxially with a well-devel¬

oped axillary gland, sepals with stellate trichomes,
12— 14-ovuled ovaries, and compressed, twisted,
pubescent fruits. By contrast, D. humillima is
densely pulvinate with entire, obtuse leaves, divar¬
icate-ascending, stout fruiting pedicels 2— 5(— 8) mm
long and adaxially eglandular, sepals with simple
trichomes, 16—24 -ovuled ovaries, and inflated, un¬
twisted glabrous fruits.

Draba konghoiuna Al-Shehbaz, sp. nov. TYPE:
China. Tibet (Xizang]: Kongbo, Nyima La,
14,500 ft., 29°28'N, 94°52'E. 28 June 1947,
F. Ludlow, G. Sh err iff & H. H. Elliot 15329
(holotype, BM; isotypes, E, TI).

Herba perennis, 3-11 cm alta, caespitosa, scaposa.
Caules erecti, dense pubescentes, basi pilis simplicibus
et stellatis. Folia basalia rosulata, persistentia, spathulata
vel oblanceolata, 5—15 X 1.5—4 mm, inferne pilis stellatis,
sessilibus vel brevi-stipitatis, 3- vel 4-radiatis, superne
pilosi pilis simplicibus; petiolis (5— )10 — 20 mm longis, cil-
iatis, suberosis. Folia caulina carentia. Raeemi 8-17-flori,
ebracteati. Pedicelli fructiferi 7—15 cm longi, pilis stel¬
latis. Sepala 2—3 mm longa. Petala Hava, late obovata, 5—
7 X 3—5 mm, apice emarginata. Ovula 12—16. Fructus
oblongi vel oblongo-ovati, puberuli, 4-6 X 1.5—2 mm,
compressi; stylus 0.4— 0.6 mm longus. Semina ovata, ca. 1
X 0.6 mm.

Perennial herbs 3-1 1 cm tall, densely caespi-
tose, scapose. Caudex covered with petioles of pre¬
vious years, few branched, ultimate branches ter¬
minated in rosettes. Stems erect, simple, densely
pubescent throughout with stellate trichomes mixed
at least basally with simple ones. Basal leaves ro¬
sulate, persistent; petiole (5— )10 — 20 mm long, per¬
sistent, thickened, corky, ciliate with simple tri¬
chomes; leaf blade spatulate or oblanceolate, 5—15
X 1.5-^ mm, abaxially densely pubescent with ses¬
sile or short-stalked, 3- or 4-rayed stellate tri¬
chomes with unbranched rays, adaxially pilose with
primarily simple trichomes, margin not ciliate,
without setose or subsetose trichomes, base atten¬
uate, margin entire, apex obtuse. Cauline leaves
absent. Racemes 8— 17-flowered, ebracteate, elon¬
gated considerably in fruit. Fruiting pedicels 7-15
mm long, ascending, straight, densely pubescent
with stellate trichomes. Sepals oblong-ovate, 2-3 X
1—1.5 mm, erect, abaxially sparsely pubescent with
simple and forked trichomes, base of lateral pair
not saccate, margin narrowly membranous. Petals
deep yellow, broadly obovate, 5—7 X 3—5 mm, apex
emarginate; claw 1-2 mm long. Filaments 2-3 mm
long; anthers oblong, 0.6— 0.8 mm long. Ovules 12
to 16 per ovary. Fruit oblong to oblong-ovate, 4—6
X 1.5—2 mm, not inflated, latiseptate, not twisted;
valves puberulent with simple trichomes sometimes
mixed with much fewer forked ones, not veined.

316

Novon

base and apex obtuse to subacute; style 0.4— 0.6
mm long. Seeds brown, ovate, ea. 1 X 0.6 mm.
Flowering May and June.

Grassy slopes, alpine grasslands, rocky areas at
altitudes 4400 to 4900 m.

Draba kongboiana is mostly closely related to D.
olgae Regel & Schmalhausen (China, Kyrgyzstan,
Pakistan, Tajikistan) from which it differs by having
stems densely pubescent throughout with stellate
and fewer simple trichomes, well -developed, per¬
sistent petioles (5— )10-20 mm long that become
thickened and corky, abaxial leaf surface pubescent
with 3- or 4-rayed stellate trichomes with un¬
branched rays, non-ciliate leaf margins, densely
pubescent fruiting pedicels, deep yellow petals,
and oblong anthers 0.6-0. 8 mm long. Draba olgae
has stems glabrous throughout or only sparsely pu¬
bescent basally, sessile leaves or with petioles to I
mm long that never become thickened or corky,
basal leaves abaxially pubescent with 3— 5-rayed
stellate trichomes at least one ray of which is
branched, leaf margins distinctly eiliate with setose
or subsetose simple trichomes, glabrous fruiting
pedicels, pale yellow petals soon turning whitish,
and ovate anthers 0.3— 0.4 mm long.

Because of similarities in habit (scapose peren¬
nials with leafless stems) and flower color and size,
it is easy to confuse Draba kongboiana with D. or-
eades Schrenk, a species widely distributed in the
Himalayas and western China, west into Central
Asia, and north into Russia and Mongolia. How¬
ever, the new species has abaxial leaf surfaces pu¬
bescent with stellate instead of primarily simple tri¬
chomes, fruiting pedicels densely pubescent all
around instead of glabrous adaxially, oblong instead
of ovate anthers, and puberulent instead of glabrous
fruits. Draba kongboiana also resembles D. jucun-
da W. W. Smith, a Chinese endemic known only
from southeastern Xizang and northwestern Yun¬
nan, in having yellow petals 5—7 mm long and pu¬
berulent fruits, but it differs in having thick, non-
surculose instead of surculose caudices, petiolate
instead of sessile basal leaves, short (0.4— 0.6 mm
long) instead of long (1.5— 2.5 mm) styles, and fruit¬
ing pedicels pubescent all around instead of gla¬
brous adaxially.

Paratypes. CHINA. Xizang: Lang La, Kymidong
Dzong, Lilung, F. Ludlow & G. Sherriff 1842 (BM. E);
Lang La. 2B°58'N, 93°42'E, F. Ludlow , G. Sherriff & G.
Taylor 4284 (BM).

Draba sunhangiana Al-Shehbaz, sp. nov. TYPE:
China. SW Tibet [Xizang): Burang Xian. Kan-
grinboque Feng, grassland on slope, 4700 m,
25 Aug. 1990, Y. Fei, II. Sun [Sung Hang], D.
Z. Li & B. Bai 447 (holotype, KUN 584594;
isotype, KUN).

Herba perennis, 5—15 cm alta, caespitosa. Caules er-
erti, ramosi, pilis crispis, simplicibus, furcatis, stellatis,
radiis inaequalibus. Folia basalia rosulata. lanceolata, 4—
10 X 1—3 mm, interne pilis stellatis, stipitatis, 3 — 5-railiu-
l is, crispis, superne pilosi pilis simplicibus, crispis; pe-
tiolis 1-4 mm longis, ciliatis. Folia caulina 2-6, ovata vel
anguste oblonga, 4—8 X 1.5—4 mm. Racemi 3— 25-flori,
ebracteati, fructiferi subumbellati. Pedicelli fructiferi 0.5—
1.5(— 2.4) mm longi. glabri. Sepala ca. 0.7 X 0.4 mm.
I’etala alba, oblanceolata, ca. 1.5 mm longa. Ovula 14—
20. Fructus ovati vel ovato-lanceolati, puberuli, 2.5 — 4(— 5)
X 1 .5—2 mm, glabri, basi inflati, obtuse, apice acuti; sty¬
lus 0. 1-0.3 mm longus. Semina ovata, 0.4— 0.5 X 0. 2-0.4
mm.

Perennial herbs 5-15 cm tall, caespitose. Cau-
dex slender, covered with leaf remains of previous
years, branched, ultimate branches terminated in
rosettes. Stems erect, branched above, pubescent
with a mixture of crisped, simple, forked, and short-
stalked stellate trichomes with unequal rays. Basal
leaves rosulate; petiole 1—4 mm long, eiliate; leal
blade lanceolate, 4—10 X 1—3 mm, abaxially pilose
with short-stalked, 3— 5-rayed stellate trichomes the
rays of which are unequal, simple, and crisped,
adaxially pilose with simple, crisped trichomes,
base attenuate, margin entire, not eiliate, apex sub¬
acute. Cauline leaves 2 to 6, sessile; leaf blade
ovate to narrowly oblong, 4—8 X 1 .5^1 mm, pu¬
bescent as basal leaves, base obtuse, margin entire
or obscurely denticulate, not eiliate, apex subob-
tuse. Racemes 3-25-llowered, ebracteate, subum-
bellate, not elongated in fruit; rachis straight. Fruit¬
ing pedicels 0.5-1.5(-2.4) mm long, divaricate,
straight, glabrous. Sepals oblong, ca. 0.7 X 0.4 mm,
erect, sparsely pilose. Petals white, oblanceolate,
ca. 1.5 mm long. Filaments ca. 0.7 mm long; an¬
thers ovate, ca. 0.1 mm long. Ovules 14 to 20 per
ovary. Fruit ovate to ovate-lanceolate, 2.5— 4(— 5) X
1.5—2 mm, basally inflated, latiseptate, not twisted;
valves glabrous, not veined, base obtuse, apex
acute; style 0.1-0. 3 mm long. Seeds brown, ovate,
0.4— 0.5 X 0.2-0. 4 mm.

Draba sunhangiana is named in honor of Sun
Hang, one of the collectors of the type gathering
and director of the KUN herbarium. It is readily
distinguished from all of the Chinese, Central
Asian, and Himalayan species by having crisped
trichomes and subumbellate fruiting racemes. The
type collection of D. sunhangiana consists of two
plants with fully mature fruits and only a few flow¬
ers, the color of which was not recorded. The two
species of Himalayan, Chinese, and Central Asian
Draba that resemble the new species in fruit shape
and size are the white-flowered D. glomerata Royle
(China, India, Kashmir, Nepal, Pakistan) and yel¬
low-flowered D. korshinskyi (0. Fedtschenko) Pohle

Volume 12, Number 3
2002

Al-Shehbaz
Himalayan Draba

317

(Afghanistan, China, Kashmir, Pakistan, Tajikis¬
tan). From these two species D. sunhangiana is
easily distinguished by having stellate trichomes
with unequal, unbranched, crisped rays, adaxial
leaf surface pilose with crisped simple trichomes,
14— 20-ovulate ovaries, and smaller seeds 0.4— 0.5
X 0.2-0. 4 mm. From D. glomerata, D. sunhangi¬
ana also differs by having glabrous fruiting pedi¬
cels and basally inflated fruits, and from D. kor-
shinskyi it differs by having leafy stems pubescent
throughout. In Draba glomerata both leaf surfaces
are uniformly tomentose with sessile or subsessile,
4-rayed stellate trichomes the rays of which are
straight and with 1 or 2 lateral branches on each
side, the fruiting pedicels are tomentose all around
with stellate trichomes, the ovaries are (6— )8— 12-
ovuled, the fruits are not inflated basally, and the
seeds are 0.7-1 X 0.5— 0.7 mm. In D. korshinskyi
the stems are leafless and glabrous throughout or
at least along the distal half, the leaves are tomen¬
tose with a mixture of simple trichomes and stellate
ones with straight, not crisped rays, the ovaries are
8— 12-ovuled, and the seeds are 1—1.2 X 0.6— 0.7
mm.

Draba himaelialensis Al-Shehbaz, sp. nov. TYPE:
India. Himachal Pradesh: above Chandratal
Lake, exposed mt. ridge in scree, 5100 m, 18
Aug. 1988. R. McBeath 2154 (holotype, E).

Herba perennis, 0.5—1 cm alta, pulvinata, scaposa, to-
mentosa, canescens; pilis stellatis, 4— 6-radiatis, radiis
tenuibus, ramosis. Folia basalia rosulata, obovata vel spa-
thulata, 2—4 X ]— 1.5 mm, tomentosa, integra. Folia cau-
lina carentia. Racemi 2 — 4-flori , ebracteati. racemi fructi-
feri folia paulo longiores. Pedicelli fructiferi 1-2.5 mm
longi, tomentosi vel apice glabri. Sepala ca. 1.2 X 0.7
mm. Petala alba, obovata, 1.2— 1.5 X ca. 0.9 mm. Ovula
10—14. Fructus orbiculares, glabri, 2—3 mm diametro,
compressi, basi et apice rotundati; stylus ca. 0.1 mm lon-
gus. Semina ovata, 0.5— 0.6 X 0.3— 0.4 mm.

Perennial herbs 0.5—1 cm tall, pulvinate, sca-
pose, canescent, tomentose throughout with short-
stalked, 4— 6-rayed stellate trichomes with finely
branched rays. Caudex covered with leaves of pre¬
vious years, many branched, ultimate branches ter¬
minated in rosettes. Stems erect, simple, tomentose.
Basal leaves rosulate; petiole 2—3 mm long, ciliate
with coarse, forked trichomes; leaf blade obovate to
spatulate, 2^1 X 1-1.5 mm, uniformly tomentose
on both surfaces, base attenuate, margin entire,
apex obtuse. Cauline leaves absent. Racemes 2^4-
flowered, ebracteate, somewhat elongated, in fruit
only slightly taller than rosette. Fruiting pedicels
1-2.5 mm long, divaricate, straight, tomentose all
around or glabrescent distally. Sepals oblong, ca.
1.2 X 0.7 mm, erect, abaxially sparsely tomentose.

base of lateral pair not saccate, margin narrowly
membranous. Petals white, obovate, 1.2— 1.5 X ca.
0.9 mm, apex rounded. Filaments 0.7-1 mm long;
anthers ovate, ca. 0.4 mm long. Ovules 10 to 14
per ovary. Fruit orbicular, 2-3 mm diam., not in¬
flated, latiseptate, not twisted; valves glabrous, not
veined, base and apex rounded; style ca. 0.1 mm
long. Seeds brown, ovate, 0. 5-0.6 X 0.3-0.4 mm.

Draba himachalensis is most closely related to
D. glomerata, which it resembles in habit, indu¬
mentum, flower color, and fruit size. However, it
differs by having leafless scapes, 2— 4-flowered ra¬
cemes, obovate, apically rounded petals 1.2— 1.5
mm long, orbicular fruits rounded at apex, and
smaller seeds 0.5— 0.6 X 0.3— 0.4 mm. Draba glom¬
erata has l-3(-5)-leaved scapes, (3 — )5 — 1 0-flow-
ered racemes, spatulate, apically subemarginate
petals (1 .8— )2— 2.7(— 3) mm long, ovate or rarely ob¬
long-ovate fruits acute at apex, and seeds 0.7—1 X
0.5— 0.7 mm.

file flowering raceme in Draba himachalensis
hardly emerges from the basal rosette, and only
when the fruits mature does it become slightly taller
than the rosette. By contrast, in D. glomerata the
raceme is distinctly taller than the rosette both in
flower and fruit.

Draba baginatiensis Al-Shehbaz, sp. nov. TYPE:
Nepal. Ganesh Himal, Bagmati Zone, Rasuwa
District, Paldol Base Camp, 4400 m, sandy
soil, 3 Aug. 1994, 28°13'N, 85°12'E, F. Mi¬
yamoto, K. R. Rajbhandari, S. Akiyama, M.
Amano, H. Ikeda & H. Tsukaya 9400056 (ho¬
lotype. TI; isotype, MO).

Herba perennis, 1—6 cm alta, caespitosa; pilis stellatis,
4-radiatis, radiis tenuibus, ramosis. Folia basalia rosulata,
oblanceolata vef oblonga, 2-10 X 0.5-3 mm, tomentosa,
integra vel dentata. Folia caulina 2—5, sessilia, ovata vel
oblonga, 2—7 X 1—3 mm. Racemi 5— 13-flori, floribus bas-
ibus braeteatis. Pedicelli fructiferi 1—5 mm longi, tomen¬
tosi, saepe recurvati. Sepala 0.8—1 X 0.4— 0.5 mm. Petala
alba, spathulata, 1.2—2 X 0.5—1 mm. Ovula 16—20. Fruc¬
tus anguste elliptici, puberuli, 3—5 X 1.2— 1.7 mm, com¬
pressi; stylus 0.1 -0.4 mm longus. Semina ovata, 0.5— 0.7
X 0.3-0. 5 mm.

Perennial herbs 1—6 cm tall, caespitose. Caudex
slender, covered with petiolar remains of previous
years, few to many branched, ultimate branches ter¬
minated in rosettes. Stems several to many, simple,
ascending, tomentose with subsessile stellate tri¬
chomes. Basal leaves rosulate; petiole to 3 mm
long; leaf blade oblanceolate to oblong, 2-10 X
0.5—3 mm, tomentose with sessile, 4-rayed stellate
trichomes the rays of which with 2 lateral branches,
base cuneate, margin entire or 1 -toothed on each

318

Novon

side, often eiliate at least near base, apex obtuse.
Cauline leaves 2 to 5, sessile; leaf blade ovate to
oblong, 2—7 X 1—3 nun, pubescent as basal leaves,
obtuse, margin entire, not eiliate at base, apex sub¬
acute. Racemes 5— 13-flowered, lowermost flowers
bracteate. Fruiting pedicels 1—5 mm long, horizon¬
tal, often recurved, forming a distinct angle with
fruit, tomentose all around with subsessile, stellate
trichomes. Sepals oblong, 0.8-1 X 0.4— 0.5 mm,
erect, caducous, abaxially pilose, base of lateral
pair not saccate, margin membranous. Petals white,
spatulate, 1.2-2 X 0.5—1 mm, caducous, apex sub-
emarginate or rounded; claw absent. Filaments 0.8—
1 mm long, caducous; anthers ovate, 0.1— 0.2 mm
long. Ovules 16 to 20 per ovary. Fruit narrowly el¬
liptic, 3—5 X 1.2— 1.7 mm, not appressed to rachis,
latiseptate, untwisted; valves puberulent with sim¬
ple trichomes; style 0. 1-0.4 mm long. Seeds ovate,
0.5-0. 7 X 0.3-0.5 mm.

Draba bagmatiensis is most closely related to I).
lasiophylla Boyle (Bhutan, China. India, Kashmir,
Kazakhstan, Kyrgyzstan, Nepal, Tajikistan, Uzbeki¬
stan), from which it is easily distinguished by having
horizontal, often recurved fruiting pedicels forming
a distinct angle with fruit, sepals 0.8-1 mm long,
petals 1.2-2 mm long, elliptic, untwisted fruits not
appressed to rachis, puberulent fruit valves with

simple trichomes, and seeds 0.5-0.7 mm long. By
contrast, D. lasiophylla has erect to ascending fruit¬
ing pedicels often subappressed to rachis and form¬
ing a straight line with fruit, sepals 1.2— 1.8 mm long,
petals 2-3.5 mm long, lanceolate to lanceolate-ob¬
long twisted fruits often subappressed to rachis, to¬
mentose fruit valves with branched trichomes, and
larger seeds 0.8— 1.1 mm long. The fruits in D. la¬
siophylla are twisted two or three turns and rarely
are they twisted one or half a turn. Both I). lasio¬
phylla and D. bagmatiensis are closely related to D.
oaricarpa 0. E. Schulz (Pakistan, Nepal, Tibet, Sik¬
kim) and differ in having caducous floral parts. The
last species is easily distinguished by having persis¬
tent sepals, petals, and stamens that often persist
well after fmit dehiscence.

Paratypes. INDIA. Sikkim: west district, opposite
bambi, above Thangshing, 4130 m, 27°3I'N, 88°1 l'E, 20
July 1992, fine scree by stream, I). G. Long, R. McBeath,
II. Koine & M. Watson 540 (CAS, E).

Acknowledgments. I am most grateful to llenk
van der Werff for correcting the Latin, to Neil A.
Harriman and an anonymous reviewer for their crit¬
ical comments on the manuscript, to Zhu Guanghua
and Yue Jipei for translating the collection data
from Chinese to English, and to the directors and
curators of BM. CAS, E, K, KUN, PE. and TI for
the loans of cited specimens.

Lesquerella is United with Physaria (Brassicaceae)

Ihsan A. Al-Shehbaz

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

ihsan.al-shehbaz@mobot.org

Steve L. O' Kane, Jr.

Department of Biology, University of Northern Iowa, Cedar Falls, Iowa 30614-0421, U.S.A.

steve.okane@uni.edu

ABSTRACT. Lesquerella (Brassicaceae) is united
with the earlier-published Physaria following an
unsuccessful attempt to conserve the more speciose
Lesquerella. Molecular, morphological, distribution¬
al, and ecological data strongly support the union
of the two genera. Ninety-one names in Lesquerella,
including 75 at the specific rank, are transferred to
Physaria. The new names P. nelsonii and P. reedi-
ana are proposed to avoid the creation of later hom¬
onyms in Physaria.

Key words: Brassicaceae, Lesquerella, North
America, Paysonia, Physaria.

The genus Physaria (Nuttall ex Torrey & A.
Gray) A. Gray was initially described in Torrey and
Gray (1838) as a section of the Old World genus
Vesicaria Adanson and was later raised to the ge¬
neric rank by Gray (1848). As presently circum¬
scribed (Rollins, 1993), Physaria consists of 22
species distributed from North Dakota south into
Nebraska, Colorado, and New Mexico, and west
through the Mountain and Pacific states, with the
range of one species, P didymocarpa (Hooker) A.
Gray, extending into southern Alberta, Canada.
With the exception of P. alpestris Suksdorf, P. geyeri
(Hooker) A. Gray, and P. oregona S. Watson, most
of the remaining 19 species occur in Colorado,
Utah, or Wyoming. Both P. didymocarpa and P. gey¬
eri were originally described in Vesicaria, but this
generic name is illegitimate because it included the
type of the earlier published Alyssoides Miller. As
many as 37 taxa of North American Brassicaceae
(see Rollins & Shaw, 1973; Rollins, 1993) were
originally described in Vesicaria, but the vast ma¬
jority of those were later transferred to lesquerella
S. Watson (Watson, 1888).

lesquerella was established by Watson (1888) as
a North American genus distinguished from the ex¬
clusively Mediterranean Vesicaria by having eden¬
tate versus dentate filaments and veined instead of
veinless septa. Watson (1888) and Payson (1922)

recognized 33 and 52 species in Lesquerella, re¬
spectively. Rollins (1993) recognized 83 species of
Lesquerella in North America, with the range of one,
L. arctica (Wormskjold ex Hornemann) S. Watson,
extending into Greenland, northern Canada, Alas¬
ka, and Arctic Russia (Rollins & Shaw, 1973; Tol-
machev, 1975). Five additional North American
species were subsequently described (Rollins,
1995; Rollins et ah, 1996; Anderson et ah, 1997;
O’Kane, 1999). Although Boelcke and Romanczuk
(1984) recognized a single species from Argentina
(and Bolivia), we believe that an additional four or
five new species grow in southern South America.
The taxonomy of the South American species of
Physaria will be published at a later date.

In his Cruciferae of Continental North America,
Rollins (1993) separated Physaria from Lesquerella
primarily on the basis of Physaria having strongly
didymous fruits with deep sinuses between the
valves distally (apically), and often proximally (ba-
sally) as well, as compared to Lesquerella, which
has non-didymous fruits with no, or shallow, distal
sinuses. Additionally, Physaria was said to have
siliques compressed perpendicular to the replum
while Lesquerella has fruits compressed parallel to
the replum or not at all, a dichotomy that is violated
in several taxa. Lesquerella and Physaria are indis¬
tinguishable in basically every other morphological
aspect, including leaf morphology, trichome type,
inflorescence, flower color, fruiting pedicels, and all
aspects of seed-coat sculpture and embryo type.
This remarkable similarity was expressed by Rol¬
lins (1939: 393), who stated, “The natural relation¬
ship between Physaria and Lesquerella is very
marked. These two genera have almost exactly the
same floral pattern, habit of growth, and trichome
morphology. The siliques too are very similar."
With the exception of the South American species
and the auriculate-leaved species of lesquerella of
the southeastern United States, Physaria and les¬
querella have a coherent geographical distribution.

Novon 12: 319-329. 2002.

320

Novon

The geographic range of Physaria falls perfectly
within that of Lesquerella, which is widespread in
the dry western United States and northern Mexico.
Furthermore, species of both genera primarily oc¬
cupy arid substrates supporting sparse vegetation.
In some cases, a species of Physaria or Lesquerella
may be the only, or one of a very few, species on a
site. Many of the species of these genera are rare,
and some are included on the United States list ol
Endangered and Threatened Species.

The degree of development of the apical and/or
basal sinuses among species of Lesquerella and
Physaria , and therefore, whether the fruit becomes
didymous or not, is a quantitative character that
shows continuity from one end of the spectrum to
the other. All workers who studied these two genera
were unable to draw a natural line between them.
For example, Payson (1922: 221), who critically ex¬
amined the boundaries between these genera, stat¬
ed, “So striking, indeed, is this similarity that one
is a little perplexed at times to know to which genus
a given plant should be referred,” and “The bridge
connecting the two genera is nearly complete.” Fur¬
thermore, Rollins (1950: 46) indicated that “The
evidence now assembled shows a continuous mor¬
phological gradation from the genus Physaria into
Lesquerella. This evidence has been accumulating
almost since the time Watson (1888) founded Les-
querella as a genus distinct from Vesicaria and
“Thus the assumed gap between these genera has
been completely closed insofar as the morphology
of the various entities involved is concerned." Ma¬
guire (1942) and Maguire and Holmgren (1951)
also expressed their dissatisfaction about the delim¬
itation of both genera, as did Mulligan (1968), who
critically examined borderline species and trans¬
ferred three species from Physaria to Lesquerella.

All the authors above have clearly indicated that
there are no adequate morphological grounds to
support the maintenance of Lesquerella and Phy¬
saria as separate genera. Although Rollins and Rii-
denberg (1971) hinted that chromosome size of
Physaria might be different from that of Lesquerella,
subsequent studies (Rollins & Riidenberg, 1977,
1979) did not support that assumption. Two main
reasons explain why both genera have been main¬
tained by previous authors. First, authors have fol¬
lowed tradition and ignored the lack of convincing
morphologies separating the two genera. Second,
authors simply avoided the transfer of the numerous
species of Lesquerella to the much smaller and ear¬
lier-published Physaria. Such arguments were pre¬
sented by Rollins and Shaw (1973: 5) who, para¬
doxically, stated that “Nothing is gained for the
classification of either Lesquerella or Physaria by

abandoning the traditional line of demarcation be¬
tween them” (italics, ours) and “We interpret the
evidence ... as once again emphasizing the step-
by-step continuity between these genera. The line
to be drawn between the genera is an arbitrary one,
and for this reason we think it should remain as it
has been accepted for many years.” Such reasoning
does not justify the maintenance of both genera,
especially when molecular data are taken into con¬
sideration.

One of the present authors (SLO) has been work¬
ing on the phylogeny of the Physaria-Lesquerella
complex for the past four years using molecular
tools. The molecular data clearly show that Phy¬
saria is nested within and evolved more than once
from Lesquerella. These results are primarily from
l)NA sequences of the internal transcribed spacer
(ITS; Baldwin et al., 1995) of nuclear ribosomal
l)NA. Less complete data from length variation of
the region between microsatellite loci, called inter¬
simple sequence repeat regions (ISSR; Zietkiewicz
et ah, 1994), support these conclusions. Regardless
of the number of derivations of Physaria from Les-
querella, the former genus is polyphyletic and the
latter is definitely paraphyletic. Molecular data
(O’Kane, unpublished) do not support any regroup¬
ing of the species that would allow the recognition
of both Physaria and Lesquerella. The two genera
can no longer be maintained on molecular, mor¬
phological, cytological, biogeographic, and ecolog¬
ical grounds. Lesquerella and Physaria (hereafter
combined as Physaria ), together with Dimorpho-
carpa Rollins, Dithyrea Haney, Lyrocarpa Hooker
& Harvey, Nerisyrenia Greene, and Synthlipsis A.
Gray, are characterized by having 4— 10-colpate
pollen grains (Rollins, 1979; Rollins & Shaw, 1973;
Rollins & Banerjee, 1979), a synapomorphy that
distinguishes these genera from the rest of the fam¬
ily, which have tricolpate pollen grains. Molecular
data (O’Kane, unpublished) also support the mono-
phyly of this Physaria alliance.

In order to avoid the massive transfer of species
from the later-published Lesquerella to the earlier-
published Physaria, O’Kane et al. (1999) proposed
to conserve Lesquerella with a conserved type. By
conserving Lesquerella, one avoids the transfer of
91 names from Lesquerella to Physaria. Only 20
species names in Physaria would have been trans¬
ferred to Lesquerella. It is rather unfortunate that
the Committee for Spermatophyta (Brummitt, 2000)
did not take the above overwhelming arguments
into consideration when it decided by a vote of ten
to five to reject the proposal because (p. 804) “The
feeling of a majority of the Committee is that in the
present case the name Physaria is already well

Volume 12, Number 3
2002

Al-Shehbaz & O’Kane
Lesquerella United with Physaria

321

known and the advantages of conservation are not
sufficient to justify over-ruling the principle of pri¬
ority.” We disagree with this reasoning and are
forced to take one of two actions. First, maintain
both genera, which would mean that a paraphyletic
Lesquerella and polyphyletic Physaria should be
recognized. However, this solution would mean that
taxonomy should be practiced without regard to
evolutionary or phylogenetic considerations, a
course of action unacceptable to us. Therefore, we
are left with the second option, to unite the two
genera. We certainly would have preferred to do so
by the conservation of Lesquerella. The extensive
transfer of species from Lesquerella to Physaria is
here made so that names are available for floristic
works in progress, including the Flora of North
America, the Flora of the San Juan Basin, and the
Arizona Flora.

The eight auriculate-leaved species previously
included in Lesquerella are excluded from this
treatment and will be transferred to a new genus in
a separate publication (see this issue of Novon ).
file South American Physaria (previously l^esquer-
ella) will also be treated in another publication. No-
menelatural adjustments made here, then, are only
for North American species. Therefore, as presently
circumscribed, the genus Physaria in North Amer¬
ica consists of 98 species, including 22 maintained
in that genus by Rollins (1993), 75 transferred
herein from Lesquerella to Physaria, and 2. P. cor-
diformis Rollins and P. montana (A. Gray) E. L.
Greene, originally described in or transferred to
Physaria, respectively.

Following the new combinations and new names
given below, the basionym and the synonym in Les-
querella (when applicable) are cited. The complete
synonymy is given in Rollins and Shaw (1973) and
Rollins (1993).

Physaria august ifolia (Nuttall ex Torrey & A.
Gray) O’Kane & Al-Shehbaz, comb. nov. Bas¬
ionym: Vesicaria angustifolia Nuttall ex Torrey
& A. Gray, FI. N. Amer. 1: 101. 1838. TYPE:
U.S.A. Oklahoma: Choctaw Co., Red River,
Thomas Nuttall s.n. (lectotype, designated by
Rollins (1956), BM; isolectotype, K).

Physaria arctica (Wormskjold ex Hornemann)
O'Kane & Al-Shehbaz, comb. nov. Basionym:
Alyssum arcticum Wormskjold ex Hornemann,
FI. Dan. 9, Fase. 26, 5, t. 1520. 1818. Les¬
querella arctica (Wormskjold ex Hornemann)
S. Watson, Proc. Amer. Acad. Arts. 23: 254.
1888. TYPE: Greenland. Omenak, C. L. Gie-
secke s.n. (holotype, C).

Physaria arenosa (Richardson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria ar¬
enosa Richardson, in Franklin, App. Narr.
Journey Polar Sea 743. 1823. Lesquerella ar¬
enosa (Richardson) Rydberg, Bull. Torrey Bot.
Club 29: 236. 1902. TYPE: Canada. Plains of
Saskatchewan. John Richardson 252 (holotype,
K; isotype, GH).

Physaria arenosa subsp. argillosa (Rollins &
Shaw) O’Kane & Al-Shehbaz, comb, et stat.
nov. Basionym: Lesquerella arenosa var. argil¬
losa Rollins & Shaw, Genus Lesquerella in N.
Amer. 178. 1973. TYPE: U.S.A. South Dakota:
Pennington Co., along bed of creek near Pied¬
mont. E. J. Palmer 37016 (holotype, GH).

Physaria argentea (Sehauer) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Vesicaria argentea
Sehauer, Linnaea 20: 720. 1847. Lesquerella
argentea (Sehauer) S. Watson, Proc. Amer.
Acad. Arts 23: 252. 1888, non L. argentea
(Pursh) MacMillan, Metasp. Minnesota Valley
263. 1892. TV PE: Mexico. Without locality,
Aschenborn 278 (holotype, B).

Physaria argyraea (A. Gray) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Vesicaria argyraea
A. Gray, Bost. J. Nat. Hist. 6: 146. 1850. Ies-
querella argyraea (A. Gray) S. Watson, Proc.
Amer. Acad. Arts 23: 254. 1888. TYPE:
U.S.A. Texas: Calhoun Co., banks of Green
Take, near Matagorda Bay, F. Lindheimer 329
(lectotype, designated by Rollins & Shaw
(1973), GH; isolectotype, MO).

Physaria argyraea subsp. diffusa (Rollins)
O’Kane & Al-Shehbaz, comb, et stat. nov. Bas¬
ionym: Lesquerella diffusa Rollins, J. Arnold
Arbor. 21: 395. 1940. TYPE: Mexico. Nuevo
Leon: Galeana, 5400 ft.. Chase 7750 (holo¬
type, GH; isotypes, AR1Z, F).

Rollins (1993) reduced lesquerella diffusa to a

variety of L. argyraea, a position that we support

at the subspecific rank under Physaria.

Physaria arizonica (S. Watson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: lesquerella ar¬
izonica S. Watson, Proc. Amer. Acad. Arts 23:
254. 1888. TYPE: U.S.A. Arizona: Yavapai
Co., Juniper Mesa, near Prescott. Apr. 1876.
Edward Palmer 16 (lectotype, designated by
Rollins & Shaw (1973), GH; isolectotypes, F,
M, MO, NY, US).

322

Novon

Physaria aurea (Wooton) O'Kane & Al-Shehbaz,
comb. nov. Basionym: lesquerella aurea Woo¬
ton, Bull. Torrey Bot. Club 25: 260. 1898.
TYPE: U.S.A. New Mexico: S fork of Tularosa
Creek, 3 mi. E of Mescalero Agency, White
Mts., 30 July 1897, Elmer 0. Wooton 245 (ho-
lotype, US; isotypes, B, K, MO, NDG, NMC,
NY, RM, UC).

Physaria berlandieri (S. Watson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
berlandieri S. Watson, Proc. Amer. Acad. Arts
23: 252. 1888. TYPE: Mexico. Tamaulipas:
near Matamoros, Berlandier 884 (lectotype,
designated by Payson (1922), GH).

Physaria calcicola (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella calci¬
cola Rollins, Amer. J. Bot. 26: 419. 1939.
TY PE: U.S.A. Colorado: Eas Animas Co., 3 mi.
E of San Francisco Creek, 8 mi. NW of Trinch-
era, 13 July 1937, Reed C. Rollins 1861 (ho-
lotype, GH; isotypes, MO, NY, OKL, UC, US,
UTC).

Physaria calderi (G. Mulligan & A. Porsild)
O’Kane & Al-Shehbaz, comb. nov. Basionym:
Lesquerella calderi G. Mulligan & A. Porsild,
Canad. J. Bot. 47: 215. 1969. TYPE: Canada.
Yukon Territory: Cathedral Rocks, Ogilvie
Mts., 66°2'N, 138°44'W, 3500 ft., 29 June
1960, J. A. Calder & J. M. Gillett 26016 (ho-
lotype, DAO; isotypes, CAN, GH).

Physaria carinata (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella cari¬
nata Rollins, Contr. Gray Herb. 171: 42. 1950.
TYPE: U.S.A. Idaho: Lemhi Co., Birch Creek.
Range 29 East-Township 11 North, 3 July
1941, Ray J. Davis 3801 (holotype, GH).

Lesquerella carinata var. languida Rollins, Cruciferae of
Continental N. Ainer. 611. 1993. Syn. nov. TYPE:
U.S.A. Montana: Granite Co., Rattler Gulch, ca. 4
mi. NW of Drummond, 'Ll IN, R13W, Sec. 10, SW
14, Sec. 9, SE !4, 4050 ft., 12 June 1990, Lisa Ann
Schassberger 364 & Diane Pavek (holotype, GH).

Physaria cinerea (S. Watson) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella cinerea
S. Watson, Proc. Amer. Acad. Arts 23: 255.
1888. TYPE: U.S.A. Arizona: without locality,
1869, Edward Palmer s.n. (holotype, US).

Physaria congests (Rollins) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella conges-
ta Rollins, Contr. Gray Herb. 214: 8. 1984.
TYPE: U.S.A. Colorado: Rio Blanco Co.,
Green River Formation, North Dudley Gulch,
ca. 3/4 mi. NE of junction of Dudley Gulch
and Piceance Creek, T2S, R97W, Sec. 4, 20
June 1893. Reed C. & Kathryn W. Rollins 8394
with Scott Peterson, Aileen G. Roads, Karen Wi-
ley-Eherle & Dieter Wilken (holotype, GH; iso¬
type, MO).

Physaria densiflora (A. Gray) O' Kane & Al-Sheh¬
baz, comb. nov. Basionym: Vesicaria densiflora
A. Gray, Bust. J. Nat. Hist. 6: 145. 1850. Les-
querella densiflora (A. Gray) S. Watson, Proc.
Amer. Acad. Arts 23: 251. 1888. TYPE:
U.S.A. Texas: Gillespie Co., Fredericksburg,
May 1847. F. Lindheimer 577 (lectotype, des¬
ignated by Rollins & Shaw (1973), GH; iso-
lectotype, M).

Physaria douglasii (S. Watson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
douglasii S. Watson, Proc. A liter. Acad. Arts
23: 255. 1888. TYPE: U.S.A. Columbia Valley,
1860, Lyall s.n. (lectotype, designated by Rol¬
lins & E. Shaw (1973), GH).

Physaria douglasii subsp. tuplashensis (Rollins,
Beck & Caplow) O’Kane & Al-Shehbaz, comb,
et stat. nov. Basionym: Lesquerella tuplashensis
Rollins, Beck & Caplow, Rhodora 97: 203.
1996. TYPE: U.S.A. Washington: Franklin
Co., White Bluffs, T13N, R27E, SI 1, W 1/2,
above Columbia River, 20 July 1994, Kathryn
A. Reck & Florence E. Caplow 94001 (holo¬
type, GH; isotype, WTU).

Rollins et al. (1996) recognized that both Les¬
querella tuplashensis and L. douglasii are quite sim¬
ilar morphologically, and that they differ primarily
by the former having a raised instead of an ap-
pressed radiate portion of the trichomes. In our
opinion, these differences do not justify the recog¬
nition of two distinct species.

Physaria eiigelniaiinii (A. Cray) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria en-
gelmannii A. Gray, Gen. Amer. Bor.-Orient.
111. I: 162. pi. 70. 1848. Lesquerella engel-
mannii (A. Gray) S. Watson, Proc. Amer. Acad.
Arts 23: 254. 1888. TYPE: U.S.A. Texas:
shores of Guadaloupe, New Braunfels, May
1846, F. Lindheimer 325 (holotype, GH; iso¬
types. B, GH. K. MO).

Volume 12, Number 3
2002

Al-Shehbaz & O’Kane
Lesquerella United with Physaria

323

Physaria fendleri (A. Gray) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Vesicaria fendleri A.
Gray, Mem. Amer. Acad. Arts 4: 9. 1849. les¬
querella fendleri (A. Gray) S. Watson, Proc.
Amer. Acad. Arts 23: 254. 1888. TYPE:
U.S.A. New Mexico: around Santa Fe, 2 Apr.
1847, A. Fendler 40 (holotype, MO).

Physaria filiformis (Rollins) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella filifor¬
mis Rollins, Rhodora 58: 201. 1956. TYPE:
U.S.A. Missouri: Dade Co., Turnback, 5 May
1929. Palmer 35604 (holotype, GH; isotypes,
F, MO, NY, US).

Physaria fremontii (Rollins & E. Shaw) O’Kane
& Al-Shehbaz, comb. nov. Basionym: Lesquer¬
ella fremontii Rollins & E. Shaw, Genus Les¬
querella in N. Amer. 228. 1973. TYPE: U.S.A.
Wyoming: Fremont Co., Wind River Mts., 5
mi. E of Atlantic City, 8200 ft.. Harry I). Rip¬
ley & Rupert C. Barneby B93I (holotype, CAS;
isotypes, NY, US).

Physaria garrettii (Payson) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella garrettii
Payson, Ann. Missouri Bot. Gard. 8: 213.
1922. TYPE: U.S.A. Utah: Salt Lake Co., Big
Cottonwood Canyon, 9700 ft., 28 June 1905,
Albert O. Garrett 1344 (holotype, MO; isotypes,
GH, RM).

Physaria glohosa (Desvaux) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Vesicaria globosa
Desvaux, J. Bot. Agric. 3: 184. 1815. Lesquer-
ella globosa (Desvaux) S. Watson, Proc. Amer.
Acad. Arts 23: 252. 1888. TYPE: U.S.A. “Cet¬
te plante emit dans les lieux arides de
I’Amerique septentrionale,” without collector
name (holotype, P).

Physaria gooddingii (Rollins & E. Shaw) O' Kane
& Al-Shehbaz, comb. nov. Basionym: lesquer¬
ella gooddingii Rollins & E. Shaw, Genus les¬
querella in N. Amer. 164. 1973. TYPE: U.S.A.
New Mexico: Catron Co., along Tularosa
Creek. 5 mi. SW of Apache Creek, 4 Sep.
1956, Rupert C. Barneby 12923 (holotype,
CAS).

Physaria goodrichii (Rollins) O’Kane & A\l-Sheh-
baz, comb. nov. Basionym: lesquerella good-
richii Rollins, J. Arnold Arbor. 64: 503. 1983.
TYPE: U.S.A. Utah: Millard Co., Desert Ex¬
perimental Range, Tunnel Springs Mts., 21 mi.
SE of Garrison, T24S, R17W, S9, 8000 ft., 16
June 1982, Sherel Goodrich 1 695 1 (holotype,
GH; isotypes, BRY, NY, UTC).

Physaria gordonii (A. Gray) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Vesicaria gordonii
[as gordoni] A. Gray, Bost. J. Nat. Hist. 6: 149.
1850. Lesquerella gordonii S. Watson, Proc.
Amer. Acad. Arts 23: 253. 1888. TYPE:
U.S.A. New Mexico: Canadian River, Raton
Mts., Apr. 1848, A. Gordon 4 (holotype, GH;
isotype, MO).

Physaria gordonii subsp. densifolia (Rollins)
O’Kane & Al-Shehbaz, comb, et stat. nov. Bas¬
ionym: lesquerella gordonii var. densifolia Rol¬
lins, Cruciferae Continental N. Amer. 621.
1993. TYPE: U.S.A. New Mexico: Lincoln Co.,
ca. 2 mi. N of Lincoln, off US llwy. 380, 13
Apr. 1990, Reed C. & Kathryn W. Rollins 9046
(holotype, (HI).

Physaria gracilis (Hooker) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Vesicaria gracilis Hook¬
er, Curtis’s Bot. Mag., n.s. 10: t. 3533. 1836.
lesquerella gracilis (Hooker) S. Watson, Proc.
Amer. Acad. Arts 23: 253. 1888. TYPE:
U.S.A. Texas: San Felipe, Drummond III 13
(holotype, K; isotype, GH).

Physaria gracilis subsp. nuttallii (Torrey & A.
Gray) O'Kane & Al-Shehbaz, comb. nov. Bas¬
ionym: Vesicaria nuttallii Torrey & A. Gray, FI.
N. Amer. 1: 101. 1838. Lesquerella gracilis
subsp. nuttallii (Torrey & A. Gray) Rollins &
E. Shaw, Genus Lesquerella in N. Amer. 53.
1973. TYPE: U.S.A. “Prairies of the Red Riv¬
er,” Melines C. Leavenworth s.n. (lectotype,
designated by Rollins & Shaw (1973), GH).

Physaria hemiphysaria (Maguire) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
hemiphysaria Maguire, Amer. Midi. Naturalist
27: 466. 1942. TYPE: U.S.A. Utah: Sanpete
Co., S side Middle Forks Park, Wasatch Pla¬
teau, 10,800 ft., 10 Aug. 1940. Bassett Ma¬
guire 20053 (holotype, UTC; isotype, DS).

Physaria hemiphysaria subsp. lucens (Welsh &
Reveal) O'Kane & Al-Shehbaz, comb, et stat.
nov. Lesquerella hemiphysaria var. lucens
Welsh & Reveal, Great Basin Naturalist 37:
338. 1977. TYPE: U.S.A. Utah: Carbon Co., 7
mi. NE of Sunnyside, Range Creek Canyon,
23 June 1977, Stanley L. Welsh & K. Taylor
15139 (holotype, BRY).

324

Novon

Physaria hitchcockii (Munz) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella hitch¬
cockii Munz, Bull. Torrey But. Club 56: 163.
1929. TYPE: U.S.A. Nevada: Clark Co.,
Charleston Mts„ 1 Sep. 1927. 10,500 ft., E. C.
Jaeger & C. Leo Hitchcock s.n. (holotype,
POM).

Physaria hitchcockii subsp. rubicundula (Hol¬
lins) O’Kane & Al-Shehbaz, comb, et slat. nov.
Basionym: Lesquerella rubicundula Hollins,
Contr. Dudley Herb. 3: 178. 1941. TAPE:
U.S.A. Utah: Garfield Co., Red Canyon, ca. 12
mi. E of Bryce Junction. Sevier Forest (now
Powell National Forest), 6 July 1912. Willard
W. Eggleston HI 98 (holotype, NA).

The main differences given by Rollins (1993) to
separate Lesquerella hitchcockii from L. rubicundula
(e.g., racemes exserted vs. embedded among the
basal leaves, smooth vs. tuberculate trichomes, and
leaves attenuate at base vs. petiolate, respectively)
are variable and often not correlated. While L. rub¬
icundula does have some smooth trichomes, most
trichomes are tuberculate and indistinguishable
from those in L. hitchcockii. The two species are
also indistinguishable moleeularly (O’Kane, unpub¬
lished) and cytologically (Windham & O’Kane, un¬
published). Maguire and Holmgren (1951) and Bar-
neby (1966) treated L. rubicundula as a subspecies
of L. hitchcockii , a position we agree with. Hollins
also reduced L. tumulosa to synonymy of L. rubi¬
cundula, a relationship that is also not substanti¬
ated on molecular and cytological grounds.

Physaria humilis (Rollins) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella humilis Rol¬
lins, Contr. Gray Herb. 214: 9. 1984. TYPE:
U.S.A. Montana: Ravalli Co., near summit of
St. Mary's Peak, Bitterroot Range, 17 July
1983, Reed C. & Kathryn W. Rollins 83300,
with Klaus H. Lackschewitz, Peter Lesica &
Aileen G. Roads (holotype, GH).

Physaria inflata (Hollins & E. Shaw) O’Kane &
Al-Shehbaz, comb. nov. Basionym: Lesquerella
inflata Hollins & E. Shaw, Genus Lesquerella
in N. Amer. 150. 1973. TYPE: Mexico. Nuevo
Leon: between Saltillo and Monterrey, along
Rte. 40, km 351. ca. 200 yards E of weighing
station, 14 Feb. 1969, Spetzman 1212 (holo¬
type, GH).

Physaria intermedia (S. Watson) O’Kane & Al-
Shehbaz, comb, et stat. nov. Basionym: Les¬
querella alpina (Nuttall ex Torrey & A. Gray)
S. Watson var. intermedia S. Watson, Proc.
Amer. Acad. Arts 23: 251. 1888. TYPE:
U.S.A. New Mexico: W of Sante Fe, A. Fendler
38 (lectotype, designated by Rollins & Shaw
(1973), GH).

Physaria johnstonii (Hollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella johnsto¬
nii Rollins, Contr. Dudley Herb. 3: 179. 1941.
TYPE: Mexico. Coahuila: Sierra del Pino, ca.
10 mi. N of camp at La Noria, Ivan M. John¬
ston & Cornelius H. Muller 556 (holotype,
GH).

Physaria kingii (S. Watson) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Vesicaria kingu S.
Watson, Proc. Amer. Acad. Arts 20: 353.
1885. Lesquerella kingii (S. Watson) S. Watson,
Proc. Amer. Acad. Arts 23: 251. 1888. TYPE:
U.S.A. Nevada: Pershing Co., West Humboldt
Mis., June 1868. Sereno Watson 82 (lectotype,
designated by Payson (1922), GH; isolecto-
type, NY).

I'he following subspecies of Physaria kingii were
variously recognized by Rollins (1993) and Hollins
and Shaw (1973) as varieties and/or subspecies un¬
der Lesquerella kingii.

Physaria kingii subsp. bernardina (Munz)
O’Kane & Al-Shehbaz, comb, et stat. nov. Bas¬
ionym: Lesquerella bernardina Munz, Bull. S.
Calif. Acad. Sci. 31: 62. 1932. TYPE: U.S.A.
California: San Bernardino Co., N side of Bear
Lake, E end of Bear Valley, San Bernardino
Mts., 16 May 1924, Frank W. Peirson 4600
(holotype, POM; isotypes, GH, JEPS, RM.
RSA).

Physaria kingii subsp. cobrensis (Rollins & E.
Shaw) O'Kane & Al-Shehbaz, comb, et stat.
nov. Basionym: Lesquerella kingii subsp. kingii
var. cobrensis Rollins & E. Shaw, Genus fes-
querella in N. Amer. 255. 1973. TYPE: U.S.A.
Nevada: Elko Co., 30 mi. E of Wells on Cobre
old Hwy. paralleling Southern Pacific Rail¬
road, Percy Train 3665 (holotype, GH; iso¬
types, NY, UC).

Volume 12, Number 3
2002

Al-Shehbaz & O’Kane
Lesquerella United with Physaria

325

Physaria kingii subsp. tliversifolia (E. L. Greene)
O’Kane & Al-Shehbaz, comb, et stat. nov. Bas-
ionym: Lesquerella diversifolia E. L. Greene.
Pittonia 4: 309. 1901. TYPE: U.S.A. Oregon:
Wallowa Go., Wallowa Mts., 7000 It.. 0 Aug.
1899, Cusick 2304 (holotype, NDG; isotypes,
F, GH, MO, UC, US, WS).

Physaria kingii subsp. latifolia (A. Nelson)
O’Kane & Al-Shehbaz, comb, et stat. nov. Bas-
ionym: Lesquerella latifolia A. Nelson, Bot.
Gaz. 42: 49. 1906. TYPE: U.S.A. Nevada: Lin¬
coln Co., Karshaw, Meadow Valley Wash, 26
Apr. 1902, Leslie N. Goodding 625 (holotype,
RM; isotypes, GH, MO, US).

Physaria klausii (Rollins) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella klausii Rol¬
lins, Contr. Gray Herb. 214: 10. 1984. TYPE:
U.S.A. Montana: Lewis and Clark Co., W side
above Hwy. 200 at Rogers Pass, 5800 ft., 25
July 1982. Klaus H. Lackschewitz & David
Ramsden 10112 (holotype, MONTU).

Physaria lata (Wooton & Standley) O’Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
lata Wooton & Standley, Contr. U.S. Natl.
Herb. 16: 126. 1913. TYPE: U.S.A. New Mex¬
ico: Lincoln Co., near Lincoln National Forest.
1903, Fred G. Plummer s.n. (holotype, US).

Physaria lesicii (Rollins) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella lesicii Rol¬
lins, Novon 5: 71. 1995. TYPE: U.S.A. Mon¬
tana: Carbon Co., ridge W of Layout Creek, Vz
mi. S of Mystery Cave, Pryor Mts., 7500 ft.,
T85, R28E, S21. SW (4. 20 June 1992, Peter
Lesica 5707 & Rob DeVelica (holotype, GH).

Physaria lindheimeri (A. Gray) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria lind¬
heimeri A. Gray, Bost. J. Nat. Hist. 6: 145.
1850. Lesquerella lindheimeri (A. Gray) S.
Watson, Proc. Amer. Acad. Arts 23: 253.
1888. TY PE: U.S.A. Texas: lower Guadeloupe,
E of Victoria, Feb. 1845, F. Lindheimer 327
(holotype, GH: isotvpes. B, (ill, K, MO).

Physaria ludoviciana (Nuttall) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Alyssum lu-
dovicianum Nuttall, Gen. N. Amer. PI. 2: 63.
1818; based on Myagrum argenteum Pursh,
FI. Amer. Septen. 2: 434. 1814, not A. argen¬
teum Vitmann, Summa 4: 30. 1790. Lesquer-
ella ludoviciana (Nuttall) S. Watson, Proc.
Amer. Acad. Arts 23: 252. 1888. TYPE:
U.S.A. Banks of the Missouri. Thomas Nuttall
s.n. (lectotype, designated by Rollins & Shaw
(1973). PHI; isolectotype, BM).

Physaria macrocarpa (A. Nelson) O’Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
macrocarpa A. Nelson, Bot. Gaz. 34: 366.
1902. TYPE: U.S.A. Wyoming: Sweetwater
Co., Bush Ranch, 10 June 1900, Aven Nelson
7031 (holotype, RM; isotypes, B, COLO, GH.
K, MO, NY, US).

Physaria mcvaughiana (Rollins) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
mcvaughiana Rollins, Contr. Gray Herb. 171:
44. 1950. TYPE: U.S.A. Texas: Brewster Co.,
E of Mt. Ord. Sierra del Norte, ea. 10 mi. SE
of Alpine, 7 Apr. 1947. Rogers McVaugh 7362
(holotype, (7 H ; isotype. A).

Physaria mexicana (Rollins) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella mexi¬
cana Rollins, Bol. Soc. Bot. Mexico 23: 45.
1958. TYPE: Mexico. Coahuila: 43 mi. S of
Monclova, 14 Aug. 1948, Kenoyer & Crum
2625 (holotype, GH).

Physaria mirandiana (Rollins) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
mirandiana Rollins, Bol. Soc. Bot. Mexico 23:
43. 1958. TYPE: Mexico. Nuevo Leon: Canon
de los Capulines, above San Enrique, Haci¬
enda San Jose de Rafces, Municipio de Der-
rumbadero, 6 Aug. 1935, G. H. Muller 2412
(holotype, GH: isotypes, F. MO).

Physaria multiceps (Maguire) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella multi-
ceps Maguire, Amer. Midi. Naturalist 27: 465.
1942. TYPE: U.S.A. Utah: Cache Co., vicinity
of Tony Grove Lake, Bear River Range, 8300
ft., 4 Aug. 1838, Bassett Maguire 16030 (ho¬
lotype, UTC; isotypes, DS, GH, RM, UC).

Physaria navajoensis (O'Kane) O'Kane & Al-
Shehbaz, comb. nov. Basionym: lesquerella
navajoensis O'Kane, Madrono 46: 88. 1999.
TYPE: U.S.A. New Mexico: McKinley Co., ca.
2 air mi. NE of Thoreau on bench below and
SE of Mt. Powell. 35°26'8"N, 108°12'26"W,
7600 ft. (2316 m), 23 May 1998, Steve L.
O'Kane , Jr. 4232 & Daniela Roth (holotype,
MO; isotypes, BRY, COLO, GH. ISTC, NMC,
NY, RM, UNM).

326

Novon

Physaria nelsonii O’Kane & Al-Shehbaz, nom.
nov. Replaced name: Lesquerella condensata
A. Nelson, Bull. Torrey But. Club 26: 238.
1899, not Physaria condensata Rollins, Rho-
dora 41: 407. 1939. TYPE: U.S.A. Wyoming:
Sweetwater Co., Tipton, Avert Nelson 4797
(lectotype, designated by Payson (1922). RM;
isolectotypes, GH, NY).

The species is named in honor of Aven Nelson
(24 March 1859—31 March 1952) who described
numerous novelties in the Brassicaceae of the
Rocky Mountain flora. The transfer of Lesquerella
condensata to Physaria would create a later hom¬
onym of P. condensata Rollins (Rollins, 1939).

Physaria ohdeltala (Rollins) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella obdel-
tata Rollins, Cruciferae Continental N. Amer.
640. 1993. TYPE: U.S.A. Idaho: Binghuam
Co., near Middle Butte, off road T-4, 3.5 mi.
E of U.S. Hwy. 26. 24 June 1986. Reed C. A:
Kathryn W. Rollins 8695 (holotype, GH).

Physaria occidentalis (S. Watson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria oc¬
cidentalis S. Watson, Proc. Amer. Acad. Arts
20: 353. 1885. Lesquerella occidentalis (S.
Watson) S. Watson, Proc. Amer. Acad. Arts 23:
251. 1888. TYPE: U.S.A. California: Siskiyou
Co., Humbug Hills, near Yreka, 30 June 1876,
Edward L. Greene 902 (lectotype, designated
by Hitchcock et al. (1964), GH).

Physaria occidentalis subsp. cinerascens (Ma¬
guire & A. Holmgren) O'Kane & Al-Shehbaz,
comb, et stat. nov. Basionym: Lesquerella oc¬
cidentalis subsp. diversifolia (Greene) Maguire
& A. Holmgren var. cinerascens Maguire & A.
Holmgren, Madrono 11: 178. 1951. TYPE:
U.S.A. Nevada: White Pine Co., White Pine
Mts., NE of Hamilton, Harry D. Ripley & Ru¬
pert C. Barneby 9295 (holotype, NE; isotypes,
UC, UTC).

Physaria ovalifolia (Rydberg) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella ovali¬
folia Rydberg, in Britton & A. Brown, Ill. FI.
N. U.S. 2: 137. 1897. TYPE: U.S.A. Nebraska:
Kimball Co., hills of upper Uawrence Fork, 1 1
Aug. 1891, Per A. Rydberg 22 (holotype, NY;
isotypes, NY, US).

Physaria ovalifolia subsp. alba (Goodman)
O’Kane & Al-Shehbaz, comb. nov. Basionym:
Lesquerella ovalifolia var. alba Goodman, Rho-
dora 38: 239. 1936. TYPE: U.S.A. Oklahoma:
Murray Co., Arhuekle Mts., 7 Apr. 1934,
George J. Goodman 2077 (holotype, OKL).

Physaria pallida (Torrey & A. Gray) O’Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria
grandiflora Hooker var. pallida Torrey & A.
Gray, FI. N. Amer. I: 101. 1838. Lesquerella
pallida (Torrey & A. Gray) S. Watson, Proc.
Amer. Acad. Arts 23: 253. 1888. TYPE:
U.S.A. Texas: San Augustine Co., near St. Au¬
gustine, Melines G. Leavenworth s.n. (holotype,
NY).

Physaria palmeri (S. Watson) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella palmeri
S. Watson, Proc. Amer. Acad. Arts 23: 255.
1888. TYPE: Mexico. Baja California: Topo
Canon, Lower California, 1884, C. R. Orcutt
1099 (holotype, GH).

Physaria parviflora (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella parvi¬
flora Rollins, J. Arnold Arbor. 64: 506. 1983.
TYPE: U.S.A. Colorado: Rio Blanco Co.,
USGS Black Cabin Gulch Quadrangle, T3S,
R100W, SI 3, 3/4 mi. S of spring in Tommy’s
Draw, 2595 m, 21 July 1982. W. Baker & S.
Sigstedt 82-308 (holotype, GH; isotype, CS).

Physaria parvula (Greene) O’ Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella parvula
Greene, Pittonia 4: 308. 1901. TYPE: U.S.A.
Colorado: Grand Co., summit of Mt. Bross,
Middle Park, 29 July 1876, H. N. Patterson
s.n, (holotype, NDG; isotypes, F, MO, NY, US).

Physaria paysonii (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella paysonii
Rollins, Contr. Gray Herb. 171. 44. 1950.
TYPE: U.S.A. Wyoming: Lincoln Co., moun¬
tains near Cottonwood Lake, E of Smoot, 13
Aug. 1923, Edwin B. Payson & George M.
Armstrong 3816 (holotype, GH; isotypes,
COLO, MO).

Physaria pcndula (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: lA’squerella pendula
Rollins, Cruciferae of Continental N. Amer.
647. 1993. TYPE: U.S.A. Nevada: White Pine
Co., Snake Range, 3 km E of Murphy Wash
road to Big Springs Wash, T10N, R68E, 6450
ft., 23 May 1985, Arnold Tiehm 9480 (holo¬
type. GH).

Volume 12, Number 3
2002

Al-Shehbaz & O’Kane
Lesquerella United with Physaria

327

Physaria peninsularis (Wiggins) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
peninsularis Wiggins, Contr. Dudley Herb. 1 :
166. 1933. TYPE: Mexico. Baja California: Ea
Encantada, Sierra San Pedro Marlir, 18 Sep.
1930, Ira L. Wiggins & L. Demaree 4919 (ho-
lotype, DS; isotypes, GH, UC. US).

Physaria pinetorum (Wooton & Standley) O’Kane
& Al-Shehbaz, comb. nov. Basionym: Lesquer¬
ella pinetorum Wooton & Standley. Contr. U.S.
Natl. Herb. 16: 126. 1913. TYPE: U.S. A. New
Mexico: Lincoln Co., Gilmores Ranch on Eagle
Creek. White Mts., 2200 m, 25 Aug. 1907,
Elmer O. Wooton & Paul C. Standley .'4160
(holotype, US; isotype, NMC).

Physaria prostrata (A. Nelson) O'Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
prostrata A. Nelson, Bull. Torrey Bot. Club 26:
124. 1899. TYPE: U.S. A. Wyoming: Uinta Co.,
Piedmont, 7 June 1898, Aven Nelson 4564 (ho¬
lotype, RM; isotypes, B, NY, US).

Physaria pruinosa (Greene) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella prui¬
nosa Greene, Pittonia 4: 307. 1901. TYPE:
U.S. A. Colorado: Archuleta Co., Pagosa
Springs, 21 July 1899, Charles F. Baker s.n.
(holotype, NDG).

Physaria piiehlensis (Payson) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella pue-
blensis Payson, Ann. Missouri Bot. Card. 8:
169. 1922. TYPE: Mexico. Puebla: vicinity of
San Luis Tultitlanapa, Carl. A. Purpus 3389
(holotype, MO; isotypes, E, GH, NY, UC, US).

Physaria pulchella (Rollins) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella pul¬
chella Rollins, Novon 5: 72. 1995. TYPE:
U.S. A. Montana: Beaverhead Co., S-facing
slope along Trapper Creek just S of Hecha
Mines, Pioneer Mts., T3S, RI1W, Sec. 32,
8600 ft., 25 July 1991, Peter Lesica 5525 &
Steve Cooper (holotype, GH; isotype, MONTU).

Physaria purpurea (A. Gray) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Vesicaria purpurea
A. Gray, Smithsonian Contr. Knowl. (PL
Wright.) 5: 14. 1853. Lesquerella purpurea (A.
Gray) S. Watson, Proc. Amer. Acad. Arts 23:
253. 1888. TYPE: U.S. A. Texas: El Paso Co.,
near El Paso, Wright 1320 (holotype, GH; iso¬
types, GH, MO).

Physaria rectipes (Wooton & Standley) O'Kane &
Al-Shehbaz, comb. nov. Basionym: Lesquerella
rectipes Wooton & Standley, Contr. U.S. Natl.
Herb. 16: 127. 1913. TYPE: U.S. A. New Mex¬
ico: without locality, 6 June 1883, C. C. Marsh
81 (holotype, US).

Physaria recurvata (Engelmann ex A. Gray)
O’Kane & Al-Shehbaz, comb. nov. Basionym:
Vesicaria recurvata Engelmann ex A. Gray,
Bost. J. Nat. Hist. 6: 147. 1850. Lesquerella
recurvata (Engelmann ex A. Gray) S. Watson,
Proc. Amer. Acad. Arts 23: 253. 1888. TYPE:
U.S. A. Texas: near Cibolo River, between San
Antonio and New Braunfels, Mar. 1846, F.
Lindheimer 330 (lectotype, designated by Rol¬
lins & Shaw (1973). GH; isolectotype, MO).

Physaria reediana O’Kane & Al-Shehbaz, nom.
nov. Replaced name: Vesicaria alpina Nuttall
ex Torrey & A. Gray, EL N. Amer. 1: 102.
1838. Lesquerella alpina (Nuttall ex Torrey &
A. Gray) S. Watson, Proc. Amer. Acad. Arts
23: 251. 1888. TYPE: U.S. A. Wyoming: Na¬
trona Co.?, Rocky Mt. range toward sources of
the Platte. Thomas Nuttall s.n. (holotype, BM;
isotypes. K, PH).

This is named in honor of Reed C. Rollins (7
December 1911—28 April 1998) for his lifetime of
contributions on the North American Brassieaceae,
especially in the genera Lesquerella and Physaria.
The transfer of Vesicaria alpina to Physaria would
create a later homonym of P. alpina Rollins (Rol¬
lins, 1981).

Physaria reediana subsp. spatulata (Rydberg)
O'Kane & Al-Shehbaz, comb, et stat. nov. Bas¬
ionym: Lesquerella spatulata Rydberg, Contr.
U.S. Natl. Herb. 3: 486. 1896. TYPE: U.S.A.
South Dakota: Lawrence Co.. Deadwood, 1600
m. Per A. Rydberg 534 (holotype, US; isotype,
NY).

Lesquerella spatulata was reduced by Rollins and
Shaw (1973) to synonymy of L. alpina and was
treated by Rollins (1993) as a variety of the latter.
It is maintained herein at the subspecific level in
Physaria.

Physaria rosei (Rollins) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella rosei Rol¬
lins, Contr. Gray Herb. 214: 23. 1984. Based
on Synthlipsis lepidota Rose, Contr. U.S. Natl.
Herb. 8: 294. 1905, non Physaria lepidota
Rollins, Brittonia 33: 335. 1981. TYPE: Mex¬
ico. Hidalgo: calcareous soil near I'ula, 6800
ft., Pringle 6899 (holotype, US; isotypes, B.
GH, M. MO, NY, US).

328

Novon

Physaria rosei subsp. perotensis (Rollins)
O' Kane & Al-Shehbaz, comb, et stat. nov. Bas-
ionym: Lesquerella rosei var. perotensis Rollins,
Conti. Cray Herb. 214: 23. 1984. TYPE: Mex¬
ico. Veracruz: Bosque de Pinos, alrededores de
Perote, 2700 m, 21 Apr. 1976, Sergio Aven-
dafio R. et al. 201 (holotype, GH).

Physaria sehaffneri (S. Watson) O’Kane & Al-
Shehbaz, comb. nov. Basionym: Vesicaria
sehaffneri S. Watson, Proc. Amer. Acad. Arts
17: 320. 1882. Lesquerella sehaffneri (S. Wat¬
son) S. Watson, Proc. Amer. Acad. Arts 23:
254. 1888. TYPE: Mexico. San Euis Potosi:
San Luis Potosi, ./. G. Schaffher 150 (holotype,
CH; isotype, F).

Physaria sessilis (S. Watson) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella gracilis
(Hooker) S. Watson var. sessilis S. Watson,
Proc. Amer. Acad. Arts 23: 253. 1888. Les-
querella sessilis (S. Watson) Small, FI. SE U.S.
471. 1903. TYPE: U.S. A. Texas: Uvalde Co.,
Rio Frio and Leona, May 1851, Wright 848
(lectotype, designated by Rollins & Shaw
(1973), GH; ?isolectotypes, GH, NY).

Physaria sinuosa (Rollins) O’Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella sinuosa Rol¬
lins, Contr. Gray Herb. 214: 23. 1984. TYPE:
Mexico. Puebla: N of Tehuacan, 5650 ft., 19
Nov. 1966. Harry D. Ripley & Rupert C. Bar-
neby 14734 (holotype, GH; isotype, NY).

Physaria subumbellata (Rollins) O’Kane & Al-
Shehbaz, comb. nov. Basionym: Lesquerella
subumbellata Rollins, Amer. J. Bot. 26: 420.
1939. TYPE: U.S.A. Utah: Uintah Co., foot¬
hills of Uinta Mts., 18 mi. N of Vernal, 17 June
1937, Reed C. Rollins 1758 (holotype, GH).

Physaria tenella (A. Nelson) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella tenella
A. Nelson, Bot. Gaz. 47: 426. 1909. TYPE:
U.S.A. Nevada: Clark Co., Moapa, 8 Apr.
1905, Leslie N. Goodding 2184 (holotype, RM;
isotypes, GH, MO, UC).

Physaria thaiunopliila (Rollins & E. Shaw)
O'Kane & Al-Shehbaz, comb. nov. Basionym:
Lesquerella thamnophila Rollins & E. Shaw,
Genus Lesquerella in N. Amer. 86. 1973.
TYPE: U.S.A. Texas: Zapata Co., 3 mi. SE of
Zapata, near US Hwv. 83, Reed C. Rollins &
Donovan S. Correll 5949 (holotype, GH; iso-
tvpe, LL).

Physaria tumulosa (Bameby) O’Kane & Al-Sheh¬
baz, comb, et stat. nov. Basionym: Lesquerella
hitchcockii Munz subsp. tumulosa Barneby,
Lead. West. Bot. 10. 313. 1966. TYPE: U.S.A.
Utah: Kane Co., 6.5 mi. SE of Cannonville, just
S of the Garfield County line, 5700 ft., 12 June
1966, Rupert C. Barneby 14424 (holotype, NY;
isotypes, BRY, CAS, GH, UC, US, UTC).

Reveal (1970) raised Lesquerella hitchcockii
subsp. tumulosa to specific rank, but Rollins (1993)
reduced it to synonymy of L. rubicundula. Because
of its pulvinate habit and filiform-linear leaves 0.7—
I mm wide, we agree with Reveal in recognizing
subspecies tumulosa as a distinct species. The re¬
mainder of the Physaria hitchcockii complex (ex¬
cept L. navaoensis ) are more loosely caespitose and
with leaves distinctly wider than those of P. tumu¬
losa.

Physaria utahensis (Rydberg) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella utah¬
ensis Rydberg, Bull. Torrey Bot. Club 30. 252.
1903. TYPE: U.S.A. Utah: Utah Co., American
Fork Canyon, 31 July 1880, Marcus E. Jones
1354 (holotype, NY; isotypes, AR1Z, DS, MO,
US).

Physaria valida (Greene) O’ Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella valida
Greene, Pittonia 4: 68. 1899. TYPE: U.S.A.
New Mexico: Gray, 1898, Josephine Skehan
s.n. (holotype, NDG; isotypes, NY, US).

Physaria vicina (Anderson, Reveal & Rollins)
O' Kane & Al-Shebbaz, comb. nov. Basionym:
Lesquerella vicina Anderson, Reveal & Rol¬
lins, Novon 7: 9. 1997. TYPE: U.S.A. Colo¬
rado: Montrose Co., small flat-topped mesa
overlooking Uncompahgre River just S ol
66500 Kinikin Road, ea. 4 air mi. SE of Mon¬
trose, 6050 ft., sec. 10, T48N. R9W, 29 June
1995, James L. Reveal 7492 (holotype, GH;
isotypes, BM, BRY, COLO, MARY, MO, NY,
RM, TEX, US. US).

Physaria wardii (S. Watson) O’Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella wardii
S. Watson, Proc. Amer. Acad. Arts 23: 255.
1888. TYPE: U.S.A. Utah (as Arizona): Gar¬
field Co., Aquarius Plateau, 11,000 It.. 16
Aug. 1875, L. F. Ward 589 (holotype, GH; iso¬
types, F, MO, US).

Lesquerella kaibabensis Rollins, Contr. Gray Herb. 211:
I 10. 1982. Syn. nov. TYPE: U.S.A. Arizona: Coco-

Volume 12, Number 3
2002

Al-Shehbaz & O’Kane
Lesquerella United with Physaria

329

nino Co., Grand Canyon National park, 18.6 mi. S
of Jacob Lake on road to N entrance of the park. 8
June 1979, Reed C. & Kathryn W. Rollins 79/9/
(holotype, GH; isotype. MO).

A critical examination of the type collections of
Lesquerella wardii and L. kaibabensis reveals that
they are indistinguishable in every morphological
aspect. The alleged differences in flower color and
style length (Rollins, 1993) do not hold. On the
Kaibab Plateau of northern Arizona, populations of
L. wardii vary from all white flowered, to all yellow
flowered, to mixed white and yellow.

Physaria wyndii (Rollins & E. Shaw) O’Kane &
Al-Shehbaz, comb. nov. Basionym: Lesquerella
wyndii Rollins & E. Shaw, Genus Lesquerella
in N. Amer. 154. 1973. TYPE: Mexico. Coa-
huila: Sierra del Carmen, Canon de Sentenela
on Hacienda Piedra Blanca, Wynd & Mueller
SI 7 (holotype, GH; isotypes, ARIZ, MEXIJ,
MO, NY, OKLA, US).

Exci.udki) Species

The following taxa are excluded from lesquerella
anti now placed in the new genus Paysonia (see
O’Kane & Al-Shehbaz in this issue of Novon).

Lesquerella auriculata (Engelmann & A. Gray) S. Watson,
Proc. Amer. Acad. Arts. 23: 250. 1888.

Lesquerella densipila Rollins, Rhodora 54: 186. 1952.
Lesquerella grandiflora (Hooker) S. Watson, Proc. Amer.
Acad. Arts 23: 250. 1888.

lesquerella lasiocarpa (Hooker ex A. Gray) S. Watson,
Proc. Amer. Acad. Arts 23: 251. 1888.

Lesquerella lasiocarpa var. berlandieri (A. Gray) Payson,
Ann. Missouri Bot. Gard. 8: 139. 1922.
lesquerella lasiocarpa var. heterochroma (S. Watson) Rol¬
lins, Rhodora 57: 245. 1955.
lesquerella lescurii (A. Gray) S. Watson, Proc. Amer.

Acad. Arts 23: 250. 1888.

Lesquerella lyrata Rollins, Rhodora 57: 252. 1955.
lesquerella perjorata Rollins, Rhodora 54: 190. 1952.
Lesquerella stonensis Rollins, Rhodora 57: 255. 1955.

Acknowledgments. We are most grateful to Neil
A. Harriman. C. Donovan Bailey, and Victoria C.
Hollowell for their critical reviews of the manu¬
script. Michael Windham graciously made his un¬
published cytological data available to us.

Literature Cited

Anderson, J. L„ J. L. Reveal & B. C. Rollins. 1997. les¬
querella vicina (Brassicaceae), a new species from the
Uncompahgre River Valley in western Colorado. Novon
7: 9-12.

Baldwin. B. G., M. J. Sanderson, J. M. Porter, M. E Woj-
ciechowski, C. S. Campbell & M. J. Donoghue. 1995.
The ITS region of nuclear ribosomal l)NA: A valuable

source of evidence of angiosperm phylogeny. Ann. Mis¬
souri Bot. Gard. 82: 247—277.

Barneby, R. C. 1966. New sorts of Lesquerella, Euphorbia ,
and Viguiera from Kane County, Utah. Leaflets W. Bot.
10: 313-317.

Boelcke, 0. & M. C. Romanczuk. 1984. Cruciferae. FI.
Patagon. 4A: 373—544. Buenos Aires.

Brummitt, R. K. 2000. Report of the committee on Sper-
matophyta: 50. Taxon 49: 799-808.

Gray, A. 1848. Genera florae Americae boreali-orientalis
illustrata. George Putnam, New York.

Hitchcock, C. L., A. Cronquist, M. Ownbey & J. W.
Thompson. 1964. Vascular Plants of the Pacific North¬
west. 2: 521—526. Univ. Washington Press, Seattle.

Maguire, B. 1942. Great Basin plants — VII. Cruciferae.
Amer. Midi. Naturalist 27: 463-469.

- & A. H. Holmgren. 1951. Botany of the Intermoun¬
tain Region — II. lesquerella. Madrono 11: 172—184.

Mulligan, G. A. 1968. Transfers from Physaria to Les¬
querella (Cruciferae). Canad. J. Bot. 46: 527—530.

O’Kane, S. L.. Jr. 1999. lesquerella navajoensis (Brassi¬
caceae), a new species of the L. hitchcockii complex
from New Mexico. Madrono 46: 88—91.

- . I. A. Al-Shehbaz & N. J. Turland. 1999. (1393)

Proposal to conserve the name lesquerella against Phy¬
saria (Cruciferae). Taxon 48: 163-164.

Payson, E. B. 1922. A monograph of the genus lesquer¬
ella. Ann. Missouri Bot. Gard. 8: 103-236.

Reveal, J. L. 1970. Comments on lesquerella hitchcockii.
Great Basin Naturalist 30: 94—' 98.

Rollins, R. C. 1939. The cruciferous genus Physaria.
Rhodora 41: 392-415.

- . 1956. On the identity of lesquerella angustifolia.

Rhodora 58: 199-202.

- . 1950. Studies on some North American Cruci¬
ferae. Contr. Gray Herb. 171: 42-53.

- . 1979. Dithyrea and a related genus. Bussey Inst.

Harvard Univ. 1979: 3—32.

- . 1981. Studies in the genus Physaria (Cruciferae).

Brittonia 33: 332-341.

- . 1993. The Cruciferae of Continental North Amer¬
ica. Stanford Univ. Press, Stanford.

- . 1995. Two Lesquerellas (Cruciferae) of south

central and western Montana. Novon 5: 71—75.

- & U. Banerjee. 1979. Pollen of Cruciferae. Bus¬
sey Inst. Harvard Univ. 1979: 33-64.

- & L. Riidenberg. 1971. Chromosome numbers of

Cruciferae 11. Contr. Gray Herb. 201: 117—133.

- & - . 1977. Chromosome numbers of Cru¬
ciferae III. Contr. Gray Herb. 207: 101—116.

- & - . 1979. Chromosome numbers of Cru¬
ciferae IV. Bussey Inst. Harvard Univ. 1979: 79—91.

- & E. A. Shaw. 1973. The Genus Lesquerella (Cru¬
ciferae) in North America. Harvard Univ. Press, Cam¬
bridge, Massachusetts.

- , K. A. Beck & E. E. Caplow. 1996. An unde¬
scribed species of lesquerella (Cruciferae) from the
State of Washington. Rhodora 97: 201-207.

Tolmachev, A. I. 1975. Cruciferae. Flora Arctica URSS.
7: 38—168. Academy of Sciences of USSR. Leningrad.

Torrey, J. & A. Gray. 1838. Flora of North America, Vol.
1. Wiley & Putnam, New York.

Watson, S. 1888. Contributions to American botany. Proc.
Amer. Acad. Arts 23: 249—267.

Zietkiewicz, E., A. Rafalski & I). Labuda. 1994. Genome
fingerprinting by simple sequence repeat (SSR)-an-
chored polymerase chain reaction amplification. Gen¬
omics 20: 176—183.

Socotrella , a New Genus of Stapeliad
(Apocynaceae— Asclepiadoideae) from the Island of Socotra

P. V. Bruyns

Bolus Herbarium, University of Cape Town, 7701, South Africa. bruyns@maths.uct.ac.za

A. G. Miller

Royal Botanic Gardens, 20a Inverleith Row, EH3 SLR, Edinburgh, United Kingdom.

t.miller@rbge.org.uk

Abstract. A new monotypic genus, Socotrella
(Apocynaceae— Asclepiadoideae— Ceropegieae), is
described with a single species, S. dolichocnema
Bruyns. This is known only from the western lime¬
stone plateau of the island of Socotra. Socotrella
differs from Caralluma R. Brown by the lack of a
differentiated leaf rudiment and its accompanying
small marginal hairs, as well as the very different
pollinaria. Although some features of the pollinaria
suggest a relationship to Ballyanthus Bruyns, Du-
valia Haworth, Duvaliandra M. G. Gilbert, Huernia
B. Brown, and Whitesloanea Chiovenda, the fact
that many inflorescences are produced toward the
lips of the stems in Socotrella at once distinguishes
it from all these genera.

Key words: Apocynaceae, Asclepiadoideae,
Ceropegieae, Socotra, Socotrella , stapeliad.

Socotrella Bruyns & A. G. Miller, gen. nov. TYPE:

Socotrella dolichocnema Bruyns, sp. nov.

A ceteris speciebus Caralluma absentia folii rudimen-
talis minuti et absentia dentium stipulaceorum distin-
guenda est, a Duvalia, Duvaliandra, Ballyanthus, Huer¬
nia, Whitesloaneaque caulis floriferis cum inflorescentiis
multis prope apicem discedit.

Small, somewhat rhizomatous succulent. Stems
decumbent, 50—150 mm long, 4—7 mm thick, dark
green, smooth; tubercles arranged into 4 rows along
stem, ± rectangular, with spreading tooth 1-2 mm
long near upper end, without leaf rudiment or stip-
ular denticles. Inflorescences several per stem near
apex, each bearing 1—5 flowers opening succes¬
sively, with acuminate bracts 1—2 mm long around
bases of pedicels; pedicel 25-42 mm long, ± 1 mm

thick, ascending and holding flower facing upward;
sepals ±1.5 mm long, 0.5 mm broad at base, ovate-
lanceolate, acute. Corolla 18—22 mm diam., shal¬
lowly campanulate; outside pale yellow-green, gla¬
brous and smooth; inside yellow with broad reddish
longitudinal markings in tube, glabrous, finely pa¬
pillate; tube 2 mm deep, 5—6 mm broad, cupular
with flat base, without thickening in corolla around
mouth; lobes 7—8 mm long, 3-4 mm broad at base,
parallel-sided for much of length, obtuse, slightly
convex inside. Corona 2 mm tall, 2 mm broad,
raised up on stipe arising from swollen area in base
of corolla tube, yellow; outer lobes reduced to mi¬
nute flap beneath guide rails; inner lobes 0.5 mm
long, erect and adpressed to backs of anthers, trun-
eate-emarginate, with broad apically somewhat lu-
berculate and almost hemispherical dorsal append¬
age around base. Anthers erect alongside style
head, margins shrinking slightly only on anthesis,
almost deflate. Pollinium ellipsoidal, much longer
than broad, pellucid margin exactly along outer
edge toward apex, caudicle attached with small pad
to ventral surface. Follicles unknown.

Socotrella dolichocnema Bruyns, sp. nov. TV PE:

Socotra. Western Plateau, 660 m, Feb. 2000,
A. G. Miller 19110 (E). Figure 1.

Species unica, a Duvaliandra dioscoridi corona exter-
iore multum redacta et orificio nectareo parviore, a spe¬
ciebus Caralluma poll ini is parvis et eorpuscula grandiore
el alata differt.

When it was first collected, we suspected that
this new species probably belonged to the genus
Caralluma. flic fairly large number of inflores-

— >

Figure 1. Socotrella dolichocnema bruyns. — A. Upper portion of flowering stem with buds. — B. Side view of flower.
— C. Side view of dissected flower. — D. Side view of gynostegium. — E. Side view of gynostegium with one inner
corona lobe removed. — F. Half-flower showing center of flower only. — G. Pollinarium. a = anther; c — corolla; g =
guide rail; h = style head; i - inner corona lobe; o = outer corona lobe; s = calyx. Scale bars: A— C. 3 mm (at A);
I). 1 mm (at F); E. 0.5 mm; F, 0.5 mm; G. 0.25 mm (at F). Drawn from Miller 191 II).

Novon 12: 330-333. 2002.

Volume 12, Number 3
2002

Bruyns & Miller
Socotrella (Apocynaceae)

331

332

Novon

cences produced toward the apices of the relatively
slender stems tended to reinforce this view.

However, a closer examination revealed that this
new species differs from all known species of Car-
alluma hy the complete lack of a differentiated leaf
rudiment at the tip of each tubercle. In Caralluma
(in the sense of Gilbert, 1990) this leaf rudiment is
accompanied hy small marginal hairs, which are
also present around and often on the stipular den¬
ticles. Both these hairs and the stipular denticles
are entirely lacking in this new species. Here the
tubercle comes to a rounded end (Fig. 1A) with no
ornamentation of this area at all.

In the complete lack of a differentiated leaf ru¬
diment on the tubercles, S. dolichocnema is some¬
what reminiscent of various species of Orbea (in the
new and expanded sense of Bruyns, 2001, 2002 in
press). Many of these have tiny stipular denticles
alongside the leaf rudiment, and the leal tapers into
a fine point. The stems are also usually distinctively
mottled with dark green on a paler background.

The flowers of .S’, dolichocnema are borne on rel¬
atively long pedicels (whence the specific epithet),
which place them at the level of the tops of the
stems. They are pretty and strikingly colored in¬
side, with bright yellow on the lobes and the same
color in the tube. In the tube there are also slightly
irregular reddish markings, which somewhat resem¬
ble color that has run down the tube. This, as well
as the shape of the flowers, is somewhat reminis¬
cent of Tromotriche umdausensis (Nel) Bruyns from
the arid west of southern Africa, but this resem¬
blance is undoubtedly purely coincidental.

When one examines the gynostegium of this spe¬
cies. there appears to be no outer corona. The swol¬
len platform on the base of the corolla tube on
which the gynostegium arises is clearly not a can¬
didate since it shows no subdivision into five parts
at any stage of its development. However, the ver¬
tical section of the flower (Fig. IF) reveals that
there is a small erect flap of tissue beneath the
guide rails (indicated by an “o”), which is not vis¬
ible easily from the outside hut which is all that
there is here of the outer corona. Behind this flap
there is a slight "nectarial orifice.” Such a reduced
outer corona is not known in any other northeast
African stapeliad, and one has to look to the south¬
ern African genus Piaranthus B. Brown to find sim¬
ilarly reduced structures in the outer coronal series
(Bruyns, 1999). It was staled (Gilbert, 1980: 101)
that the outer corona of Duvaliandra dioscoridis
(Lavranos) M. G. Gilbert was absent. However, this
is not the case. In this species the outer corona
consists of a spreading lobe laterally fused to the
sides of the rather more massive inner lobes as well

as an erect ridge of tissue a little behind the bases
of the guide rails and enclosing a more significant
“nectarial orifice” than is present in S. dolichoc¬
nema. This is somewhat different from the situation
in S. dolichocnema , where the spreading lobe is
absent.

In S. dolichocnema the inner corona lobes are,
as usual, adpressed to the backs of the anthers and,
since the anthers are erect here, the inner corona
lobes are erect, too. Each inner corona lobe has a
noticeably swollen, tuberculate dorsal appendage.
An examination of early stages of the flowers shows
that these dorsal appendages are derived from the
same meristem as the inner lobe itself and are not
part of the outer series, as is the case, for example,
in many species of Piaranthus (Bruyns, 1999).

The vertical section of the flower also shows the
arrangement of the style head and the anthers. Here
the style head projects well above the level of the
corpuscles and the anthers are erect against it. This
situation is unique among the stapeliads, where the
top of the style head is generally around the level
of the corpuscle and the anthers are normally hor¬
izontal on top of it (Bruyns, 2000a).

The pollinaria of this new species are also most
unusual. The corpuscle is comparatively large and
about equally long as broad, with inordinately long
lateral wings. The pollinia are ellipsoidal and much
longer than broad with a small insertion crest run¬
ning for a short distance down the outer edge. The
pollinia are unusually small relative to the size of
the corpuscle. Pollinaria with this shape are not
known in Caralluma, where the corpuscle is always
comparatively small, with very small lateral wings,
and the pollinia are large relative to the size of the
corpuscle. Similarly, pollinaria like those of S. dol¬
ichocnema are unknown in Orbea, where the pol-
linium is always much larger and D-shaped with
the insertion crest twisting from the outer edge to
the upper surface of the pollinium.

The shape of the corpuscle is similar to w hat one
finds generally in Duvalia, Huernia, as well as in
the monotypic genera Ballyanthus and Whiteslo-
anea from northern Somalia and Duvaliandra from
Socotra. The ellipsoidal pollinia are also similar in
these genera, though they are far smaller in the new
species. Socotrella differs from Ballyanthus, Du¬
valia, Duvaliandra, Huernia, and Whitesloanea by
the fact that several inflorescences are produced on
each flowering shoot, and they all arise toward the
tips of the stems, whereas in these other genera
there is only one inflorescence per stem often to¬
ward the apex ( Ballyanthus ) or one inflorescence
usually near the base of the stem only in Duvalia,
Duvaliandra, Huernia . and Whitesloanea.

Volume 12, Number 3
2002

Bruyns & Miller
Socotrella (Apocynaceae)

333

This new genus brings to two the number of
monotypic genera of stapeliads that are known from
Soeotra. the other being Duvaliandra, whose single
species, D. dioscoridis, is found in the granitic
mountains of the northeast of the island. The pre¬
sent new genus and species was discovered in Feb¬
ruary 2000 in the west of Socotra on a limestone
plateau with dwarf shrubland dominated by Croton
socotranus Balfour f., Pulicaria stephanocarpa,
Buxus hildebrandtii Baillon, and Cissus hamade-
rohensis A. R. Smith. It might seem peculiar that
yet another monotypic genus of stapeliad is being
described. The last one described in this journal
( Bruyns, 2000b) occurred on the edge of the botan-
ically little-known area of southern Angola. Many
of the succulents of Socotra are related to others in
Somalia, where botanical exploration is also not
very advanced. Consequently, it is possible that fur¬

ther exploration may show that both these new taxa
are not as isolated in their relationships among the
stapeliads as they seem to be at present.

Literature Cited

Bruyns, P. V. 1999. The relationships of Huerniopsis and
Piaranthus. Syst. Bot. 24: 379-397.

- . 2000a. Phvlogenv and biogeography of the sta¬
peliads. 1. Phylogeny. PI. Syst. Evol. 221:199-226.

- . 2000b. Baynesia, a new genus of stapeliad from

the north-western-most corner of Namibia. Novon 10:
354-358.

- . 2001. New combinations in the genus Orbea.

Aloe 37: 72-76.

- . 2002. A Monograph of Orbea and Ballyanthus

(Apocynaceae— Asclepiadoideae— Ceropegieae). Syst.
Bot. Monogr. 63 in press.

Gilbert, M. G. 1980. Duvaliandra'. A new genus of sta¬
peliad. Caet. Succ. J. Gr. Brit. 42: 99—101.

- . 1990. A review of Carallurna R. Br. and its seg¬
regates. Bradleya 8: 1—32.

Contributions toward a Monograph of Petalophyllum
(Marchantiophyta)

Barbara J. Crandall- Stotler, Raymond E. Stotler ; and C. Heather Ford 1
Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901-6509,

U.S.A. crandall@plant.siu.edu

ABSTRACT. The simple thalloid liverwort genus
Petalophyllum J. G. C. Lehmann is removed from
the Fossombroniaceae and sequestered in the new
family Petalophyllaceae along with Sewardiella S.

R. Kashyap, a monotypic genus endemic to the In¬
dian subcontinent. Unlike the Fossombroniaceae,
the thallus in the Petalophyllaceae is never dis¬
sected, the archegonia are clustered rather than
scattered, a pseudoperianth is formed rather than a
caulocalyx, and the capsule wall is 3—4-layered
with outer wall thickenings, which are lacking in
the Fossombroniaceae. The North American rep¬
resentative of Petalophyllum previously considered
conspecific with the European taxon P ralfsii (W.
M. Wilson) J. G. C. Lehmann is here described as
a distinct species, P. americanum. This is based
upon clear differences of the thallus, pseudoper¬
ianth, and the elater spirals. Furthermore, the ap¬
plication of the name P. australis Colenso to New
Zealand plants is shown to be incorrect; those
plants previously referred there represent a new
species, P. hodgsonii. This species resembles P.
ralfsii , but differs from it in the broad, strongly flat¬
tened thallus midrib and structure of the female
inflorescence.

Key words: Austrofossombronia, Fossombroni¬
aceae, liverworts, Marchantiophyta, New Zealand,
North America, Petalophyllaceae, Petalophyllum,
Sewardiella.

The Family Petalophyllaceae

The simple thalloid liverwort genus Petalophyl¬
lum C. G. D. Nees & Gottsche ex J. G. C. Lehmann
(Lehmann, 1844) has been aligned with the genus
Fossombronia Raddi since its publication. The in¬
clusion of these two taxa and the genus Sewardiella

S. R. Kashyap in the family Fossombroniaceae 11a-
zlinszky has been generally unequivocal among he-
paticologists, although Schuster (1953: 575) stated
“. • - I would regard it [ Petalophyllum 1 as a repre¬
sentative of a separate family, the Petalophylla-

ceae.” This family was never validly published by
him, but many years later he did segregate Petal¬
ophyllum and Sewardiella from Fossombronia and
his recently validated genus, Austrofossombronia R.
M. Schuster, at the subfamily rank when he named
the Petalophylloideae (Schuster, 1991) within the
Fossombroniaceae. The genera Fossombronia and
Austrofossombronia, then, comprised the Fossom-
bronioideae of this family.

During an ongoing monographic study of Petal¬
ophyllum, a number of structural and developmen¬
tal features have been elucidated that justify ele¬
vation of the Petalophylloideae to the rank of
family. In Petalophyllum the thallus is not dissect¬
ed, and it develops unistratose dorsal lamellae. Se¬
wardiella is composed of a very fragile unistratose
thallus wing. Conversely, the thallus of all other
members of the Fossombroniaceae is highly dis¬
sected, giving the plants a leafy appearance. The
apical cell geometry of the Petalophyllaceae is re¬
ported as being tetrahedral in Sewardiella and ei¬
ther tetrahedral or lenticular in Petalophyllum, de¬
pending on the species (Leitgeb, 1877). In the
Fossombroniaceae the geometry is strictly lenticu¬
lar (Renzaglia, 1982). Archegonia develop in dis¬
tinct clusters along the midrib in the Petalophyl¬
laceae, not scattered along the stem or midrib as in
the Fossombroniaceae. These archegonial clusters
are surrounded, even in the earliest stages of de¬
velopment, by a unistratose pseudoperianth in the
Petalophyllaceae. The pseudoperianth expands in
typical fashion after fertilization to enclose the spo-
rophyte, which is further encased by a thin shoot
calyptra. In the Fossombroniaceae, the solitary, un¬
fertilized archegonia are completely devoid of any
type of enclosing structures; i.e., they are naked on
the stem. After fertilization, concomitant with the
early stages of embryo division, the stem tissue
around the venter undergoes substantial cell divi¬
sion to form a fleshy enclosure around the basal
part of the sporophyte. A leaf-like scale, which aris-

1 Current address: Integrated Microscopy Facility, Donald Danforth Plant Science Center, 975 N. Warson Rd., St.
Louis, Missouri 63132, U.S.A.

Novon 12: 334-337. 2002.

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2002

Crandall-Stotler et al.
Petalophyllum (Marchantiophyta)

335

es at the apex of this fleshy mound, enlarges as the
sporophyte matures and forms a pseudoperianth¬
like enclosure. As originally concluded bv Chalaud
(1928), this structure is not, however, homologous
to the true pseudoperianths of Petalophyllum and
Sewardiella and is better termed a caulocalyx. Fi¬
nally, the capsule wall in representatives of the Pe-
talophyllaceae is 3—4-layered with cells in all lay¬
ers having pigmented secondary wall thickenings.
The capsule wall in Fossombronia is only 2-layered
while that of Austrofossombronia is (3— )4(— 5)-lav-
ered (Schuster, 1994; Crandall-Stotler et al., 1999).
Regardless of the number of capsule wall layers,
however, in the Fossombroniaceae the cells of the
outer wall layer are always devoid of secondary wall
thickenings.

Petalophyllaceae (R. M. Schuster) Stotler & Sto-
tler-Crandall, fam. et stat. nov. Basionym: Pe-
talophylloideae R. M. Schuster, J. Hattori Bot.
Lab. 70: 148. 1991. TYPE: Petalophyllum J.
G. C. Lehmann; type species, P. preissii J. G.
C. Lehmann, Novarum et Minus Cognitarum
Stirpium Pugillus 8: 30. 1844.

A New Species of Petalophyllum from North
America

When encountered in the field, Petalophyllum is
a conspicuously distinct simple thalloid liverwort
not easily confused with other hepatic taxa. Be¬
cause of its patchy occurrence, however, it is not
frequently observed. I be first report of the genus
in North America was by Evans (1919), who iden¬
tified two Texas specimens as Petalophyllum ralfsii
(W. M. Wilson) C. G. I). Nees & Gottsche ex J. G.
C. Lehmann, the “petal wort” common in the Brit¬
ish Isles and the Mediterranean region. Since then,
scattered reports can be found in the North Amer¬
ican literature under that name or as P. lamellatum
(C. G. D. Nees) S. 0. Lindberg, a taxonomic syn¬
onym of P. ralfsii (Lindberg, 1874). Careful evalu¬
ation of the type specimen of P. ralfsii (= Junger-
mannia ralfsii W. M. Wilson), near Penzance, Ralfs
s.n. (BM), and of European populations of this spe¬
cies have revealed several distinctions between this
taxon and the North American populations of the
genus. These morphological differences warrant the
recognition of the North American populations as a
new species, which may be contrasted w ith P. ralfsii
by several distinguishing characters. Vegetatively,
in North American plants both the wing and la¬
mellar margins are entire, very rarely sinuate. In P.
ralfsii, both the wing and lamellar margins are sin¬
uate. Mature pseudoperianths in the North Ameri¬

can plants are small, barely projecting above the
lamellae, and unlobed, with the mouth margin be¬
ing sinuate to dentate, but never eiliate. These fea¬
tures may be clearly seen in Schuster’s (1992: 428)
figure 849- 1 from Texas plants. In P. ralfsii, the
pseudoperianth is much larger and sometimes
deeply lobed with the mouth margin always strongly
toothed to ciliated or even laciniate. The elaters in
both species are bi-spiraled (tri-spiraled) at their
tips, but in our North American plants 2 to 4 (oc¬
casionally 5 or 6) spirals are found through most of
the elater length. There are never more than 2 or
3 spirals formed in P. ralfsii. In North America,
Petalophyllum has been confirmed from Arkansas,
Louisiana, and Texas, where it often occupies re¬
cently disturbed areas of well-drained sandy soils
that are generally exposed. In the British Isles, in
contrast, it is typically found in stabilized sand
dunes. Thus, these two species appear also to oc¬
cupy very different habitats.

Petalophyllum americanum C. H. Ford & Sto-
tler-Crandall, sp. nov. TYPE: U.S.A. Texas:
Bastrop Co.. Bueschner State Park, on red soil
in moist depression, 13 Mar. 1997, R. E. Sto¬
tler & B. J. Crandall-Stotler 4006 (holotype,
ABSH; isotype, as dupl. in FAA).

Differ! a Petalophyllo ralfsii marginibus thalli et lamel-
larum integris, ore pseudoperianthii sinuato-dentato, ela¬
ted bus spiris 2— 4(5— 6).

Populations light green, occurring as isolated in¬
dividuals or in small scattered patches on exposed,
loose or somewhat firm gravely or sandy loam.
Plants 3.9—1 1.2 mm long, 2.7—8 mm wide, simple
or once (twice) furcate. Thallus elliptical to ob-
ovate, broadly undulate, with the wings lamellifer-
ous. Wings flattened to erect, multistratose at the
base, becoming unistratose at the margin, with the
margin plane; the median thallus cells thin-walled,
quadrate to elongate, averaging 68 X 41.2 pan, w ith
the marginal cells distinct, tangentially elongate or
quadrate, 36.1-39.9 pm long, 34.2-43.7 pm wide.
Lamellae leaf-like, erect to suberect, running
obliquely along the thallus wings, unistratose, mul¬
tistratose at the base, up to 22 cells in height at
the highest point, slightly undulate, with the la¬
mellar margin plane; the marginal cells distinct,
quadrate to tangentially elongate, averaging 36.3 X
43.5 pm. Midrib fleshy, with the dorsal surface flat¬
tened to slightly concave, with the ventral surface
narrowly convex. Tubers apical, from microphyl-
lous, geotropic apices. Rhizoids numerous, scat¬
tered along the ventral surface of the stem, hyaline.
Dioicous, dimorphic. Male plants somewhat smaller

336

Novon

than female plants. Antherklia maturing acropetal-
ly, in 2 or 3 scattered rows along the dorsal surface
of the midrib and associated with the basal portion
of the wings, overarched basipetally by lobed mar¬
ginal extensions of the lamellae. Female plants with
5 to 9 archegonia in distinct clusters on the dorsal
surface of die midrib, with each cluster surrounded
by a pseudoperianth. Pseudoperianths at maturity
campanulate, 1(2) medially positioned along the
midrib, with the outer surface occasionally bearing
leaf-like outgrowths; the mouth erect or recurved,
with t lie margin broadly sinuate and dentate to sub-
dentate, cilia lacking, with 1(2) sporophyte(s) ma¬
turing per pseudoperianth. Capsules spheroidal,
1.4— 2.3 mm diam., dark olive brown to black; the
epidermal cells quadrate, light yellow with scat¬
tered dark red-brown nodular thickenings; the cells
of the inner layers gradating from quadrate to elon¬
gate, with dark red-brown thickening bands gra¬
dating from nodular to semiannular to annular, de¬
hiscence irregular. Spores 40-52 gm diam., light
brown to dark red-brown, disassociated when ma¬
ture. areolate. with (4)5 or 6 areolae across the dis¬
tal face; the areolae 10-12 pm wide; the proximal
surface areolate, with a central depression. Elaters
numerous, smooth, elongate, rarely to occasionally
branched, with the outer wall light yellow, smooth,
with 2 to 4 (occasionally 5 or 6) dark red-brown
spirals. Gemmae absent.

Habitat. Exposed, compact or loose, sandy or
gravely soil, often in disturbed areas, or near tem¬
porary ponds.

Distribution. Scattered locations in the Gulf
Coastal Plain of North America, extending into the
southern portion of the Interior Highlands; from
southern Arkansas to southern Louisiana west to
east and central Texas.

Paratypes. U.S.A. Texas: Bastrop Co., Bastrop State
Park, well-drained sandy soil along roadside, 12 Mar.
1997, Stotler & Stotler-Crandall 3980 (ABSH; dupl. in
FAA); nr. Buesehner State Park, well-drained sandy soil,
corner of CH 180 & Park Road 1C, 13 Mar. 1997, Stotler
A Stotler-Crandall 3998 (ABSH; dupl in FAA); Buesehner
State Park, wet seep area under grass clumps, 13 Mar.
1997, Stotler A Stotler-Crandall 4003 (MO; dupl. in FAA
at ABSH).

A New Species ok Petalophyllum from New
Zealand

During the late 1800s, the Reverend William
Colenso described two New Zealand species of Pe-
talophyllum, namely P. australe Colenso as “ aus¬
tralis ” (Colenso, 1885) and P. macrocalyx Colenso
(Colenso, 1886). No specimens of these two species
have been located in the intervening years by any

hepaticologist: not in WELT, which contains many
of Colenso’s types, nor in BM, which holds numer¬
ous of his bryophyte types that he had sent to
Hooker (Galloway, pers. comm.), nor in G, where
he sent specimens to Stephani. In a revision of Col¬
enso’s Hepaticae by Stephani (1893) neither of
these two species was mentioned, but Stephani did
state that there were other Colenso species pub¬
lished of which he had not received specimens.
Hodgson (1967) appears to be the first to recognize
the publication of these two Colenso species. She
realized that several New Zealand specimens of Pe-
talophyllum that she had studied were not assign¬
able to P. preissii , a species described from Austra¬
lia. She revived the oldest Colenso name, P.
australe, to accommodate these and apparently in
order to deal with the name P. macrocalyx, simply
listed it as a synonym. This, of course, is contrary
to nomenclatural rules. She stated that she had “. . .
adopted Colenso’s name for this species because
Gottsche’s description of P. preissii . . . does not
quite lit New Zealand plants” (Hodgson, 1967:
192). Our initial hope was to designate one of the
Hodgson specimens as a neotype for P. australe,
but this was not possible since none of that material
conformed to the protologue. The Hodgson plants
are clearly not what Colenso had described as P.
australe; rather, his description of plants with large,
much lacinate pseudoperianths fits P. preissii fairly
well. Nor can the Hodgson specimens be P. macro-
calyx, which Colenso described as being small
plants with leaves and red rhizoids. That descrip¬
tion, in fact, fits a member of the genus Fossotn-
bronia rather than Petalophyllum. We found that
the Hodgson material represents an undescribed
species of the genus distinguished from all other
species of Petalophyllum by a suite of diagnostic
characters. It is here named in honor of E. Amy
Hodgson, the amateur botanist who laid the foun¬
dation for modern New Zealand hepaticology.

Petulopliyllum lindgsonii C. II. Ford & Stotler-
Crandall, sp. nov. TYPE: New Zealand. North
Island: Morere Bush, Wairoa, on bank along
path to baths, Sep. 1947. E. A. Hodgson 11297
(holotype, MPN).

Magnitudine Petalophyllum ralfsii simulans, differt a P
ralfsii costa lata complanata, archegoniis pigmentiferis,
ore pseudoperianth ii sinuato vel dentato, sine ciliis.

Populations not observed in nature; herbarium
specimens light yellow green, occurring as isolated
individuals, loose mats or in scattered small patch¬
es on sand and humus. Plants large. 9.5—13.3 mm
long, 6.6—10.2 mm w ide, often (once) twice or more

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Crandall-Stotler et al.
Petalophyllum (Marchantiophyta)

337

furcate. Thallus elliptic to ovate, slightly to highly
undulate, with the wings lamelliferous. Wings flat¬
tened to suberect, multistratose at the base, becom¬
ing unistratose at the margin, with the margin
broadly sinuate; the median cells thin-walled,
quadrate to elongate, with the marginal cells slight¬
ly distinct, quadrate to slightly tangentially elon¬
gate, 28.9—53.2 ft m long by 24.7—55.1 gm wide.
Lamellae leaf-like, suberect to erect, running
obliquely along the thallus wings, tapering to or
almost to the thallus wing margins, unistratose,
multistratose at the base, up to 20 cells in height
at the highest point, slightly to highly undulate.
Midrib broad and fleshy, with the dorsal surface
slightly compressed, with the ventral surface
strongly flattened. Tubers 0(1), apical, from micro-
phyllous, geotropic apices. Khizoids numerous,
scattered along the ventral surface of the stem, hy¬
aline. ?Dioicous. Male plants not observed. Female
plants with 8 to 15 dark red or unpigmented ar-
chegonia in distinct clusters along the dorsal sur¬
face of the midrib, with each cluster surrounded by
a pseudoperianth. Pseudoperianth at maturity sub-
campanulate, 1 to 3 terminal or medially positioned
along the midrib, with the outer surface lacking
wings or leaf-like outgrowths; the mouth erect to
incurved or recurved, with the margin undulate,
dentate to sinuate, lacking cilia, with 1 sporophyte
maturing per pseudoperianth. Capsules brown-
black; the epidermal cells quadrate, light yellow
with scattered dark red-brown nodular thickenings,
with the cells of the inner layers gradating from
quadrate to elongate, with dark red-brown nodular
wall thickenings gradating from semiannular to an¬
nular, dehiscence irregular. Spores 39.9—51.3 gm
diam., brown, disassociated when mature, areolate,
with 3 or 4 areolae across the distal face, with the
areolae 9-10 gm across; the proximal surface are¬
olate, with a central depression. Elaters abundant,
elongate, rarely to occasionally branched, attenu¬
ate, with the outer wall light yellow, smooth, with
2 or 3 red-brown spirals. Gemmae absent.

Habitat. Along stream banks among ferns and
humus.

Distribution. New Zealand.

Acknowledgments. This work was supported by
NSF-PEET grant DEB-9521883, which is grate¬
fully acknowledged. We thank the following, who
provided living plant material: James Bray, Tim
Golden, Jered Kyzer, Daniel Marsh, and Daniel

Wrench. We gratefully acknowledge Tim Galloway
(WELT) and Ella Campbell (MPN) for their com¬
ments concerning the activities of William Colenso,
and we thank David Glenny (CHR) for his assis¬
tance in locating critical specimens and place
names. The authors sincerely thank L. Ellis (BM)
and the late P. Geissler (G) for providing study fa¬
cilities at their respective institutions. We thank the
following for arranging the loan of specimens: L.
Ellis (BM). P. Heenan (CHR). the late P. Geissler
(G), L. Dixon (MPN), and T. Galloway (WELT).

Literature Cited

Chalaud, G. t928. Le cycle evolutif de Fossombronia pus-
illa Dum. Librairie Generale de L’Enseignement, Paris.
Colenso, W. 1884 [1885]. A description of some newly
discovered and rare indigenous plants; being a further
contribution towards the making known the botany of
New Zealand. Trans. & Proc. New Zealand Inst. 17:
237-265.

- . 1885 [1886], A description of some newly dis¬
covered cryptogamic plants, being a further contribution
towards the making known the botany of New Zealand.
Trans. & Proc. New Zealand Inst. 18: 219—255.
Crandall-Stotler, B., R. E. Stotler & A. V. Ereire. 1999.
On the status of Austrofossombronia peruviana (Gottsche
& Hampe) R. M. Schust. ex Stotler, Stoll. -Crandall &
A. V. Ereire (Fossombroniineae, Marchantiophyta).
Haussknechtia Beiheft 9: 67—78.

Evans, A. W. 1919. Notes on North American Hepati-
cae — VIII. Bryologist 22: 54—73.

Hodgson, E. A. 1967. New Zealand hepaticae (liver¬
worts) — XVII. A miscellany of taxonomic notes. Part 2.
Trans. Roy. Soc. New Zealand, Bot. 3: 175—198.
Lehmann, J. G. C. 1844. Novarum et Minus Cognitarum
Stirpium Pugillus Octavus. II. Novitiae Florae Novae
Hollandiae. J. A. Meissneri, Hamburg.

Leitgeb, H. 1877. Untersuchungen ueber die Lebermoose.
III. Heft. Die frondosen Jungermannieen. O. Deistung,
Jena.

Lindberg, S. 0. 1874. Manipulus muscorum secundus.
Not. Sallsk. Fauna FI. Fenn. Forh. 13: 353—117 + 1
tab.

Renzaglia, K. S. 1982. A comparative developmental in¬
vestigation of the gametophyte generation in the Metz-
geriales (Hepatophyta). Bryophvt. Biblioth. 24: 1—253.
Schuster, R. M. 1953. Boreal Hepaticae. A manual of the
liverworts of Minnesota and adjacent regions. Amer.
Midi. Naturalist 49: 257-684.

- . 1991. Diagnoses of new taxa of Hepaticae. I. Jun-

germanniidae. J. Hattori Bot. Lab. 70: 143—150.

- . 1992. The Hepaticae and Anthocerotae of North

America, East of the Hundredth Meridian, Vol. V. Field
Museum of Natural History, Chicago.

- . 1994. Studies on Metzgeriales. III. The classifi¬
cation of the Fossombroniaceae and on Austrofossom¬
bronia Schust., gen. n. Hikobia 1 1: 439—449.

Stephani, F. 1893. A revision of Colenso’s Hepaticae, with
descriptions of new species collected by him. J. Linn.
Soc.. Bot. 29: 263-280.

A New Andean Species and a New Combination in
Oreobolopsis (Cyperaceae)

Sandra Dhooge and Paul Goetghebeur

Ghent University, Department of Biology, k.L. Ledeganckstraat 35, B-9000 Gent, Belgium.
Sandra.Dhooge@rug.ac.be; Paul.Goetghebeur@rug.ac.be

Abstract. Oreobolopsis inversa , a new species of
Cyperaceae from the Andes occurring in Ecuador
and Peru, is fully described and illustrated. It dif¬
fers from Oreobolopsis tepalifera Koyama & Guag-
lianone by its stiff erect leaves and tepals shorter
than the achene. The habit of the new species is
very similar to Trichophorum rigidum (Steudel)
Goetghebeur, Muasya & Simpson, but there are
some remarkable differences in floral morphology
between the two. In addition, the combination Or¬
eobolopsis dementis (M. E. Jones) Dhooge &
Goetghebeur is made.

Key words: Andes, Cyperaceae, Oreobolopsis,
Trichophorum.

The genus Oreobolopsis was described by Koya¬
ma and Guaglianone (1987) based on a single spe¬
cies, Oreobolopsis tepalifera. The well -developed,
membranous tepals are the most conspicuous as¬
pect of the genus. During our study of a number of
scirpoid specimens from the Americas, the pres¬
ence of two additional species became evident, one
new taxon from the Andes, and another from Cal¬
ifornia, U.S.A.. which was initially described as
Scirpus dementis M. E. Jones and recently trans¬
ferred to Trichophorum Persoon by Smith (1995).

Oreobolopsis inversa Dhooge & Goetghebeur, sp.
nov. TYPE: Ecuador. Azuay: Parque Nacional
de Las Cajas, along road Soldados— Angas, ca.
13 km from Soldados, 79°17'W, 02°53'S, alt.
3900—4000 m, 2 May 1992, S. hie guard
102684 (holotype, GENT; isotypes, AAU,
QCA, QCNE). Eigure 1.

Hcrba perennis rigida. Folia erecta. Spicula solitaria
terminalis. Glumae inferiores vacuae 3. Glumae floriferae
3-4. Flos bisexualis tepalis 3 + 3, squamiformibus, per-
sistentibus, achenio brevioribus. Achenium brunneum,
obovatum, trigonum.

Perennial, rhizomatous herb, forming dense tus¬
socks (Eig. 1 A). Roots up to 1 mm wide, brownish.
Culm (4—) I 1-25 cm long, 0.5— 0.9 mm wide, erect,
terete with 8 to 10 ribs, rounded-trigonous below
the inflorescence, scabrous distally. Leaves stiff and

erect, shorter than longest culms; sheaths brown;
blades (2— )5— 11 cm long, 0.5— 0.7 mm wide, green,
acute, margins scabrous, upper part asymmetrical
(surface of adaxial side is reduced to rib toward
leaf tip). Eigule membranous, brown. Inflorescence
of a single pseudolateral spikelet, 4—6 X 2—3 mm
(Eig. 1 B). Involucral bracts 2; lower bract pseudo¬
terminal, 13—19 mm long, scabrous, with an awn
10—18 mm long; the upper bract 4—6 mm long,
glumiform, scabrous, with an awn 1-3 mm long.
Glumes spirally arranged, brown, elliptical, gla¬
brous; lower glumes 3—3.5 mm long, empty, obtuse;
fruit-bearing glumes 3 or 4, 3—4 mm long, obtuse,
sometimes bifid at tip. Flowers bisexual. Tepals 3
+ 3, scale-like, shorter than achene (Fig. 1C); outer
w horl of tepals 1—1.4 X 0.5— 0.9 mm, pale, obtuse
or bifid; inner whorl of tepals 0.5—1 X 0.2— 0.5 mm,
irregularly lobed, pale, often with a central, red¬
dish, acute tip. Stamens 3; anthers about I mm
long, opening with 2 longitudinal slits, red-tinged;
filaments pale brown. Ovary 3-carpellate; stigma
branches 3, papillose. Achene 1.5— 1.7 X 1—1.3
mm, obovate, trigonous, brown, glabrous (Eig. ID).

Paratypes. ECUADOR. Azuay: paramos, in vicinity
of Toreador, between Molleturo and Quinoas, alt. 3810—
3930 m. 15 June 1943, J. A. Steyermark 53173 (NY);
paramo de Soldadas- Angas, at highest point of road,
79°17'Vt, 02°52'S, alt. 3950-4050 m, 14 Feb. 1988, S.
Laegaard 70109 (AAU, GENT, NY): paramo de Soldados.
SW of Cuenca, 79°17'W, 2°53'S, all. 3700-3800 m, 24
Oct. 1984, S. Laegaard 532 1 1 (A \l . CENT, NY); paramo
de Soldadas, 79°18'W, 02°53'S, alt. 3700-4000 m, 28
Aug. 1985, S. hie guard 55100 (AAU, GENT, NY); near
pass in Parque Nacional las Cajas, W of Cuenca,
79° 1 4' W, 02°40'S, alt. 4150 m, 22 Oct. 1984, 5. Laegaard
53190 (AAU, GENT, NY). Bolivar: 6 km on road Sali¬
nas— los Arenales, grass paramo and open rocks, 79°0'W,
1°22'S, alt. 4000 m, 2 Oct. 1 985, .S. Laegaard 55331
(AAU, GENT, NY). PERU. Ancash: Yungay, Huascaran
National Park, 77°34'W, 8°59'S. alt. 4000-4300 m, 12
Jan. 1985. D. N. Smith el at. 9124 (U): Hnaylas, Iluas-
earan National Park, 77°46'W, 8°53'S, alt. 3960-4400 m.
1). N. Smith et al. 9987 (U).

Anatomical Obskrv ations (Terminology
According to Metcai.kk, 1971)

fhe transverse section of the lamina is thickly
crescent iform, except at the apex where ellipsoidal.

Novon 12; 338-342. 2002.

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2002

Dhooge & Goetghebeur
Oreobolopsis (Cyperaceae)

339

Figure 1. Oreobolopsis inversa Dhooge & Goetghebeur. — A. Habit (Laegaard 102684, holotype, GENT). — B. Detail
of spikelet (Laegaard 102684, GENT). — C. Spikelet with glumes removed showing fruit and perigynous scales ( Lae¬
gaard 55331, A AU). — D. Fruit (, Laegaard 55331, AAU).

340

Novon

Figure 2. Anatomical section through the lamina (A) and culm (B) of Oreobolopsis inversa (Laegaard 53190). (Drawn
by S. Dhooge.)

The lamina margins are continuously scabrous.
Epidermis: Contrary to many Cyperaceae, the size
of the epidermal cells on both surfaces is more or
less alike. The outer periclinal wall of epidermal
cells is strongly thickened. Stomata only abaxial.
Mesophyll: a homogeneous tissue of parenchyma¬
tous cells. In the leaf center, lysigenous air cavities

are present. At the periphery, the mesophyll is in¬
terrupted by strands of sclerenchyma opposite the
vascular bundles. Adaxially, the sclerenchyma
comprises only a few (2-3) columns resting against
the epidermis. Abaxially, the sclerenchyma con¬
nects the epidermis with the largest vascular bun¬
dles, forming abaxial girders (Fig. 2A). Adaxial

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Dhooge & Goetghebeur
Oreobolopsis (Cyperaceae)

341

Table 1. Main t

lifferences between Oreobolopsis inversa and Trichophorum

rigidum.

Oreobolopsis inversa

Trichophorum rigidum

Perianth segments

+

-

Fruit length (mm)

I. .5- 1.7

1.8-2

Fruit width (mm)

1.2

1

Fruit color

chestnut

dark brown— black

BHS colour chart

grayed orange group 165 A

brown group 200 A

l.igule form

straight

arched

caps often are associated with the vascular bundles.
Vascular bundles (5 or 6) are surrounded by a bun¬
dle sheath consisting of two layers: an inner sheath
composed of cells with U-shaped thickenings and
an outer sheath of thin-walled cells. Bundle sheaths
surrounding minor vascular bundles are more
clearly marked than those surrounding the major
vascular bundles. The vascular bundles are collat¬
eral and of two different shapes: the minor vascular
bundles are circular; the major ones are elliptical.
Xylem consists of xylem vessels and xylem paren¬
chyma, phloem of sieve cells and phloem paren¬
chyma.

The transverse section of the culm is circular,
with or without a wavy outline, to rounded trian¬
gular under the inflorescence. Epidermis: Cells with
a strongly thickened outer periclinal wall. The pa¬
renchyma of the culm is homogeneous but the cen¬
tral tissue is broken down and looks like a three-
dimensional meshwork (“net” type. Metcalfe, 1971:
28). Vascular bundles (11 to 18) are in the periphery
of t he culm (Fig. 2B), supported by a sclerenchyma
girder (at the phloem side, toward the epidermis)
and a sclerenchyma cap at the xylem side. The
minor vascular bundles are only provided with 2
caps. In the lower half of the culm, small air cav¬
ities appear between the vascular bundles. Bundle
sheath: see description under lamina. The vascular
bundles are collateral with endarch xylem and ex¬
arch phloem.

The Affinity of Oreobolopsis inversa and

T RtCHOPHOR l JM R I Gl DU M

Koyama (in Koyama & Guaglianone, 1987: 84)
reported the presence of scale-like perianth seg¬
ments in the bisexual flowers of certain specimens
identified by him as “ Scirpus rigidus .” This species,
invariably without perianth segments, was recently
transferred to Trichophorum, as Trichophorurn rigi-
dum (Steudel) Goetghebeur, Muasya & I). A. Simp¬
son (Muasya et al., 2000).

A number of specimens, earlier included under
Scirpus rigidus, are here described as Oreobolopsis
inversa based on the presence of perianth segments.

The epithet alludes to its opposite characters (te-
pals shorter than the achene and the larger size of
the plant) compared with the first described spe¬
cies, Oreobolopsis tepalifera. Specimens of 0. in¬
versa are contrasted with specimens of Trichopho¬
rum rigidum (without perianth segments), because
in gross morphology they look very much alike. The
Iruit and the ligule, however, are conspicuously dif¬
ferent in shape (Table 1). Future ON A studies
could be used to support or refute our morpholog¬
ical interpretation.

A Gai.ifoknian Species of Oreobolopsis

Mention should also be made of a small scirpoid
species occurring in California (U.S.A.), Scirpus de¬
mentis M. E. Jones. This species recently was trans¬
ferred to Trichophorum by S. G. Smith (1995: 102).
However, this species is distinct from a typical Tri¬
chophorum due to the presence of scale-like peri¬
anth segments, which is the most striking feature
of Oreobolopsis.

Oreobolus R. Brown looks superficially like Or¬
eobolopsis because it also has scale-like perianth
segments. The structure of the 1 -flowered spikelet
and the distichous leaves, however, make Oreobolus
clearly distinct from Oreobolopsis. In addition, the
embryos are different: Oreobolus has a caricoid em¬
bryo with a clear basal root cap, while the embryo
of Oreobolopsis (inclusive of O. dementis) has a sub-
basal root cap (Van der Veken. 1964). Therefore,
we assume that Oreobolopsis is the most appropriate
generic name for Scirpus dementis.

Oreobolopsis dementis (M. E. Jones) Dhooge &
Goetghebeur, comb. nov. Basionym: Scirpus
dementis M. E. Jones, Contr. W. Bot. 14: 21.
1912. Trichophorum dementis (M. E. Jones) S.
G. Smith, Novon 5: 102. 1995. TYPE: U.S.A.
California: Rae Lake, King’s River, alpine, 20
July 1910, Mrs Clements s.n. (lectotype, des¬
ignated by Smith (1995), POM; isolectotypes,
DS, POM).

Scirpus yosemitanus F. J. Smiley, Univ. Calif. Publ. Bot. 9:

342

Novon

108. 1921. TYPE: U.S.A. California: Yosemite Na¬
tional Park, Tuolumne Meadows, Soda Springs, 8
Aug. 1898, J. W. Congdon s.n. (holotype, GH).

Key to the Andean Scirpoid Genera

la. Inflorescence a dense head of several spikelets,
hypogynous bristles well developed . . Phylloscirpus

lb. Inflorescence of 1 terminal spikelet . 2

2a. Hypogynous scales absent . 3

3a. Leaves ligulate . Trichophorum

3b. Leaves eligulate . Scirpus boliviensis

(generic allocation under study)

2b. Hypogynous scales present . 4

4a. Spikelet 1 -flowered, leaves distichous

. Oreobolus

4b. Spikelet 3- or 4-flowered, leaves tristi-

chous . Oreobolopsis

Key to the Species of Oreobolopsis

la. Slender, rigid plants, longer culms 10—25 cm tall

. 0. inverse i

lb. Plants < 9 cm tall.

2a. Leaves acute; achene > 1 .5 mm long, sur¬
face smooth . 0. tepalifera

2b. Leaves muticous; achene < 1.5 mm long,

surface reticulate . 0. dementis

Acknowledgments. We express our gratitude to
the curators of the Herbarium Jutlandicum of Aar¬
hus University (AAU), the herbarium of Utrecht
University (U), and The New York Botanical Gar¬
den (NY) for kindly providing specimens. We thank
Marcel Verhaegen (BK) for the SKM images and
Peter Chaerle (GENT) for his assistance with the
photography.

Literature Cited

Koyama, T. & E. 11. Guaglianone. 1987. Oreobolopsis, a
new genus of Cyperaceae (Scirpeae) from Bolivia, South
America. Darwiniana 28(1^1): 79-85.

Metcalfe, C. R. 1971. Anatomy of the Monocotyledons. V.

Cyperaceae. Clarendon Press, Oxford.

Muasya, A. M.. I). A. Simpson & P Goetghebeur. 2000.
New' combinations in Trichophorum. Scirpoides and Fi-
cinia. Novon 10: 132—133.

Smith, S. G. 1995. New combinations in North American
Schoenoplectus, Bolboschoenus, Isolepis, and Trichopho¬
rum (Cyperaceae). Novon 5: 97—102.

Veken, P Van der. 1964. Bijdrage tot de systematische
embryologie der Cyperaceae— Cyperoideae. Doctoraal
proefschrift, Katholieke Universiteit, Leuven.

Bombacaceae Neotropicae Novae vel Minus Cognitae III. Nuevas
Especies de Matisia y Quararibea de Colombia

Jose Luis Fernandez- Alonso

Instituto de Cieneias Naturales, Universidad Nacional de Colombia, Apartado Aereo 7495
Santafe de Bogota D.C., Colombia. jlfem@ciencias.unal.edu.co

Resumen. Se describen e ilustran cuatro nuevas
bombacaceas de Colombia, pertenecientes a los ge-
neros Matisia (M. gentryi y M. racemifera) y Quar-
aribea (Q. cryptantha y Q. rangelii). Las dos es-
pecies de Quararibea descritas, corresponden a
grupos muy distintivos no previamente conocidos
en territorio colombiano. Una de las especies nue¬
vas de Matisia presenta flores dispuestas en ramas
cortas afilas, formando racimos bracteados; este ul¬
timo caracter, linico en todo el genero, se describe
aquf por primera vez.

ABSTRACT. Four new Bombacaceae from Colom¬
bia, two species of Matisia (M. gentryi and M. ra¬
cemifera) and two of Quararibea (Q. cryptantha and
Q. rangelii), are described and illustrated. The two
new species of Quararibea belong to two distinctive
groups, which have not been previously reported in
Colombia. One of tbe new species of Matisia, M.
racemifera, has flowers arranged in short leafless
branches that form hracteate racemes; the latter
character, unique in the genus, is here described
for the first time.

Key words: Bombacaceae, Colombia, Matisia,
Quararibea.

Continuando con los trabajos previos al trata-
miento de la tribu Quararibeae para el territorio
colombiano, iniciados hace ya varios anos (Fernan-
dez-Alonso, 1992, 1996, 1999a, 1999b) y formando
parte de la nueva serie “Bombaceae neotropicae
novae . . .” (Fernandez- Alonso, 2001a, 2001b), se
describen en esta tercera entrega, cuatro bomba¬
caceas nuevas provenientes de Colombia. Las plan-
tas estudiadas corresponden a colecciones relati-
vamente recientes efectuadas en el Chocd central.
Valle medio del Magdalena y en la Sierra Nevada
de Santa Marta.

Matisia gentryi Fernandez-AIonso. sp. nov. TIPO:
Colombia. Chocb: ca. 50 km al F de Las Ani¬
mas, 4 km al E del no Pato, Carretera Pana-
mericana, 250 m, 1 1 Ene. 1979 (fl, fr), A. Gen¬
try & E. Renteria 23984 (holotipo, COL
204320; isotipos, HUA-11533, MO 1281065).
Figura 1.

Haec species Matisiae cordatae Bonpland affinis sed ab
ea lamina foliari plerumque late obovata raro orbiculata,
lobis brevioribus usque ad 15 mm longis, basi cordata
trinervi et nervis duabus submarginalibus minoribus nm-
nita, nervis secundariis 5 ad 9 utrinseeus nervum medium
atque fructu elipsoideo-fusiformi ealyce majore craterfor-
mi margine uniformiter sinuato-lobato basi ad eum ad-
presso subtento distinguitur.

Arboles de hasta 30 m; yemas terminales con
braeteas crasas, subcoridceas, triangulares o an-
chamente triangulares de 3-4(7) X 2.5-3 mm, con
indumento muy fino, equinado-tuberculado. Hojas
laxamente agrupadas en el extremo de las ramas
jovenes; estipulas triangulares de hasta 7 mm de
longitud; peciolo de (4.5)5.5—12 cm X 2—3 mm,
pulvfnulos proximal y distal notorios, de 3—4 mm
de diametro; laminas de anchamente obovadas a
orbiculares, de (11— )13— 23 X (9— )10-20 cm, cor-
dadas en la base con senos cortos, de hasta 1.5 cm
de longitud (profundi dad), el apice obtuso, redondo,
a veces emarginado; el margen levemente sinuoso-
ondulado; nervios basales 3, siendo el central mas
desarrollado y los adyacentes de escasa longitud en
la lamina, generalmente con otros dos nervios sub-
marginales, poco desarrollados; nervios secundarios
de 5 a 8 (9) a cada lado del nervio medio principal;
nerviacibn terciaria paralela, perpendicularmente
dispuesta a los nervios secundarios; nerviacibn
cuaternaria de tipo poligonal-reticulada, escasa-
mente resaltada (Fig. I B); indumento tanto en el
peciolo como en el enves diminutamente equinado-
tuberculado, disperso y por la haz subglabra. Inflo-
rescencias ramifloras dispuestas en formaciones no-
dosas; pedicelo de 15—18 mm, con 3 braeteolas
dispuestas en la mitad basal del pedicelo, pronta-
mente caducas; caliz obcbnico de 2—2.2 cm de lon¬
gitud y 1 .5—1.7 cm de diametro en el margen distal,
el margen irregularmente rasgado; tanto el pedicelo
como la cara externa del caliz (en seco), de color
marrbn, con indumento finamente equinado-tuber¬
culado, la cara interna con indumento serfceo,
blanquecino; petalos de color crema, suberectos en
la antesis, anchamente espatulados, de hasta 3.5 X
1 .2 cm, obtusos en el apice, la cara externa den-
samente recubierta con pelos estrellados cortos, la

Novon 12: 343-351. 2002.

344

No von

Figure 1. Matisia gentryi Fem&ndez- Alonso. — A. Apice de una rama con bracteas de la yema terminal. — B. Detalle
de la superficie de la ldmina por la haz. — C. Flor. — D. Fruto en vista lateral. — E. Fruto, seccibn transversal. (Dibujado
a partir de la serie tfpiea, A. Gentry & E. Renteria 23984.)

Volume 12, Number 3
2002

Fernandez-Alonso
Bombacaceae Neotropicae Novae

345

eara interna con indumento denso serfceo acostado;
columna estaminal de 2.3— 2.8 cm con indumento
estrellado de radios muy cortos, ramas estaminales
de hasta 1.2-1. 4 cm X 2 mm de ancho, con abun-
dante indumento glandular y con pelos largos si-
nuoso-aracnoides. Pedicelo fructifero de 1.8—2 cm
X 5-7 mm; caliz fructifero cupular, cenido al fruto
en el 1/3— 1/4 basal, de ca. 2 cm de longitud X 4—
4.5 cm de diametro, con margen uniformemente
(4)5-sinuado-lobado; frutos (Figs. ID. E) marrones,
de ovales a ovoide-fusiformes, de 6—7 X 4.5-5 cm,
ligeramente truncados en el apice, con aplculo cen¬
tral inconsplcuo; pericarpo recubierto por indu¬
mento fasciculado-estrellado, mesocarpo fibroso-
carnoso con 5 pirenos corneo-lenosos, reniformes
de ca. 3 X 1.5 X 1 cm, semilla farenocotilar.

Distribution y habitat. Conocida solo por la co¬
leccion tipo, proveniente de los bosques pluviales
del Choco central en la planicie del paclfico co-
lombiano. Esta coleccion fue citada en el catalogo
de las plantas del departamento del Choco (Forero
& Gentry, 1989) bajo el nombre M. cordata Bon-
pland, nombre con el que fueron distribuidos los
duplicados.

Etimologia. Especie nombrada en recuerdo de
Alwyn H. Gentry, una de las personas que mejor
conocio la eomposicion y la estructura de los
bosques del paclfico colombiano. Gentry colecto
gran cantidad de bombacaceas en los transectos
efectuados en el litoral paclfico y llamo la atencion
sobre una particularidad de los bosques del Choco,
la presencia de hasta 6 o 7 especies de Matisia en
parcelas de 0.1 heetarea (Gentry, 1986). Parte de
sus colecciones (algunas infertiles), fueron el punto
de partida para nuevas expediciones de coleccion,
que permitieron completar la informacion descrip-
tiva de algunas especies nuevas.

Discusidn. Matisia gentry i es afln a M. cordata,
especie ampliamente distribuida en el norte de Sud-
america y en rnenor medida, afln a M. longitubu-
losa (A. Robyns) Cuatrecasas de la Amazonia per¬
uana (Robyns, 1968). De M. cordata se diferencia
por presentar esta, hojas ampliamente ovadas,
siempre con mas de 5 nervios basales y bases mas
ampliamente cordadas (lobulos generalmente de
mas de 15 mm de profundidad), tubo estaminal mas
largo (de 3.5—5 cm), ramas estaminales tambien
mas largas (de 1.4-2. 5 cm), caliz fructifero breve-
rnente cupuliforme, escasamente acrescente y fruto
marcadamente apiculado (en seco). De M. longi-
tubulosa se diferencia por presentar esta, hojas
ellpticas con 7—9 nervios basales, pedicelos florales
mas largos (de ca. 25 mm) y al igual que en M.

cordata tanto el tubo, como las ramas estaminales
son de mayor longitud.

Matisia racemifera Fernandez-Alonso, sp. nov.
TIPO: Colombia. Choco: Mpio. de Quibdo, km
7 en la via Quibdo— Yuto, bosque entresacado,
300 m, 13 Ago. 1998 (fl, fr), J. L. Fernandez-
Alonso, F. Garcia & estudiantes Univ. Choco
16719 (holotipo, COL 421861 (1 de 2, fl),
421857 (2 de 2, fr); isotipos, COL 421858,
CHOCO, MA, MO, US). Figura 2.

Haec species ad sectionem Calyculatas Fern&ndez-
Alonso praeclare pertinet, sed a ceteris speciebus sectio-
nis inflorescentiis racemosis in ramos pendulos breves
aphyllos bracteatos dispositis, stipulis magnis 2—3 cm lon-
gis, calyce exalato atque fructu longe rostrato distinguitur.

Arboles de 12-15 m; tronco con corteza externa
oscura; ramificaeion verticilada y ramas pendulas;
las jovenes densamente recubiertas de indumento
tomentoso marron-ferruglneo. Hojas dlsticas; estl-
pulas cartaceas, caudadas, linear-lanceoladas, de
2—3 X ca. 0.5 cm, densamente estrellado-tomen-
tosas (Fig. 2A); peciolo de 1.3— 3.5 cm, con indu¬
mento estrellado-ferruglneo denso, con el pulvlnulo
proximal muy poco conspicuo y el distal notorio;
laminas cartaceas, de estrechamente obovadas a
oblanceoladas, de 21-35(46) X 9-14(17) cm, base
redondeado-cordada, agudas, ligeramente acumi-
nadas en el apice, acumen de 1—3 cm; nervios ba¬
sales 5 a 7, de los cuales 4, muy poco consplcuos;
nervios secundarios 4—5 a cada lado del nervio me¬
dio principal, broquidodromos; en la haz excavados
y en el enves resaltados; nerviacion de 3° y 4° or-
den paralelo-reticulada, resaltada en el enves; haz
verde claro (en vivo), con denso indumento estre¬
llado, de radios largos, rojizo; enves con el mismo
indumento, densamente dispuesto en los nervios y
menos denso en la lamina. Racimos dispuestos en
ramas florfferas pendulas (Fig. 2B), de 25^10 cm,
con las partes jovenes tomentoso-ferrugfneas, con
bracteas y flores; por cada flor se presentan 2 brae-
teas florales de estrechamente triangulares a lan-
ceoladas, de 1 .6 a 1.8 cm de longitud; pedicelos
de 4—5 mm en boton floral, con 3 bracteolas fer-
rugfneas, estrechamente laneeoladas, de 1.2— 1.5
cm X 2—3 mm, dispuestas a modo de calfculo; caliz
fusiforme antes de la apertura del boton. levemente
estriado, sin costas, de 1.2— 1.4 cm, con indumento
amarillento, estrellado, disperso; corola blanca.
Pedicelo fructifero de 1-1.2 cm, con bracteas y
bracteolas persistentes. Fruto de 2.8— 2.9 cm (in-
cluido el caliz), caliz fructifero cupular, de 1 X 1.8
cm, irregularmente lobado en el margen distal (Fig.
2C); fruto verde, ovado-apiculado, levemente 5-sul-
cado, de 2.6 X 1.6 cm, pico (rostro) cilindrico de

346

Novon

Figura 2. Matisia racemifera Fernandez- Alonso. — A. Delalle de las estfpulas e indumenta en una rama joven. — B.
Rama florffera alila. con botones florales. — C. Detalle de un fruto desarrollado. (Fotograffas tomadas del especimen
tipo, Fernandez- Alonso et al. 16719.)

Volume 12, Number 3
2002

Fernandez-Alonso
Bombacaceae Neotropicae Novae

347

1 cm lie longitud X 0.3 cm de ancho, exocarpo
lepidoto-ferrugineo.

Distribution y habitat. Matisia racernifera lue
recolectada por primera vez con flores en 1998, en
cercanfas de Quibdd (Choeo). Crece al igual que la
especie anterior, en bosques pluviales del Chocd
central, en la cuenca del rfo Atrato. Kstas forma-
ciones vegetales, asentadas sobre lomas bien dre-
nadas, son muy ricas en Bombacaceae y en espe¬
cial en elementos de la tribu Quararibeae. Las
Bombacaceae colectadas en la localidad tipica jun¬
to con M. racernifera, en un area no mayor de 5
hectareas, fueron: Matisia bullata Fernandez-Alon¬
so, M. castano H. Karsten & Triana, M. con bit a. M.
leptandra (Cuatrecasas) Cuatrecasas, M. idroboi
Cuatrecasas sensu lato y M. valdes-bermejoi Fer¬
nandez-Alonso & Castroviejo, Pachira aquatica
Aublet, Pachira cf. sessilis Bentham, Eriotheca
squamigera (Cuatrecasas) Fernandez-Alonso, Hu-
berodendron patinoi Cuatrecasas, Patinoa almirajo
Cuatrecasas y Phragmotheca siderosa Cuatrecasas
subsp. siderosa.

Etimologia. El nombre de la especie hace re-
ferencia a sus flores dispuestas en racimos brac-
teados, infreeuentes en el genero.

Disc us ion. Matisia racernifera se ubiea elara-
mente en la recientemente establecida seecion Ca-
lyculatae (Fernandez-Alonso, 2001a), por sus brac¬
teolas caliculiformes, y se separa del resto de las
especies de la seccion por presentar inflorescencias
racemosas dispuestas en ramas largas, afilas. Ade-
mas se caracteriza por presentar estipulas inusual-
mente grandes (2-3 cm de longitud) y densamente
tomentosas, calices no alados y frutos largamente
apiculado-rostrados. En estado vegetativo recuerda
notablemente a Matisia hirta Cuatrecasas, por la
forma de las hojas y el largo indumento (Cuatre¬
casas, 1946); no obstante esta puede separarse con
facilidad de M. racernifera por presentar flores ais-
ladas, opuestas a las hojas y pedicelos llorales mas
largos, con bracteolas florales no verticiladas, dis¬
puestas en la zona media del pedicelo. En Matisia
dolichosiphon (A. Robyns & S. Nilsson) W. S. Al-
verson de Panama, planta vegetativamente muy di-
ferente a M. racernifera (asignable a la sect. Cas¬
tano Fernandez-Alonso), se han deserito tambien
inflorescencias ramifloras paniculiformes, en nada
parecidas a las que aquf se describen (Robyns &
Nilsson, 1970).

Pardtipos. COLOMBIA. Choeo: Mpio. tie Quibdo,
km 7 en la via Quibd6— Yuto, bosque entresacado, 300 m,
13 Ago. 1998, J. L Fernandez-Alonso et al. 16701 (CHO-
CO, COL, MA, MO); Mpio. de Tad6, Corregimiento de
Salero, 1998-1999, F. Garcia 299a (CHOCO); via Quib-
do— Tutunend6, ca. 3 km al oeste de Tutunendb, bosque

pluvial tropical. 80 m, 5°46'N, 76°35'W. 5 Fme. 1981, A.
Gentry et al. 30139 (JAUM), ibidem, transecto 9, 7 Fine.
1981, A. Gentry 30308 (JAUM).

Quararibea cryptantha Fernandez-Alonso, sp.
nov. TIPO: Colombia. Magdalena: Sierra Ne¬
vada de Santa Marta, Corregimiento de El
Campano, 1300 m, 5 Jul. 1986 (fl), E. Carbond
2029 (holotipo, UTMC; isotipos, COL 500088.
MO, UTMC). Figura 3.

Haec species (Juararibeae floribundae (A. Saint-Hilaire
& Naudin) K. Schumann et Q. yunckeri Standley affinis,
sed ab eis habitu arborescente, foliis membranaceo-char-
taceis, ovali-lanceolatis vel oblongo-lanceolatis; subtus
domatiis sacciformibus, papyraceis, ebarbatis ad axillas
nervorum secundariorum munitis; pedicello 0.8—1 mm
longo, calyce viridi, urceolato-campanulato 6-7 mm longo,
ad apicem irregulariter 2 vel 3 fisso atque tubo staininali
usque ad 1 1 X 1 mm, distinguitur.

Arboles de ca. 8 m con ramificacion verticilada;
yemas glabrescentes; hojas dfsticas con olor a fe-
nogreco o regaliz (en seeo); estipulas diminutas
prontamente caducas, con indumento lepidoto ad-
preso; pecfolos verde amarillentos de 0.6— 0.9 cm
de largo, con pulvfnulo distal de ca. 3 X 1.5 mm
(Fig. 3); Luninas verde palido, membranoso-carta-
ceas, generalmente inequilateras, de oval-lanceo-
ladas a oblongo-lanceoladas, de (9 — )1 1 — 14 X (4—
)5— 6.5 cm, base ligeramente desigual, de cuneada
a ligeramente cordada; de subagudas a obtusas en
el apice; haz verde palido, subglabra con tricomas
lepidotos muy dispersos (en los nervios), el enves
subglabro; con 5 a 7 nervios laterales a cada lado
del nervio medio, excavados en la haz y resaltados
en el enves; con domacios sacciformes papiraeeos,
ubicados en la confluencia de los nervios secun-
darios con el nervio medio; nerviacion de tercer y
cuarto orden ligeramente resaltada en el enves. In¬
florescencias en ramitas muy cortas, axilares o rami¬
floras, con liasta 6(8) flores sesiles; pedicelo de has-
ta 0.8—1 mm de longitud; con 3 bracteolas de ca.
0.8 mm de longitud, obtusas, dispuestas a rnodo de
calfculo, densamente lepidotas. Flores actinomor-
fas; caliz campanulado-urceolado de 6-7 mm de
longitud (Fig. 3), externamente lepidoto, intema-
mente sericeo; con 3 a 5 lobulos marginales, de-
siguales, papiraeeos, de 1.5-2 mm de longitud; pe-
talos blancos, estrechamente espatulados de 12—13
X 2—3 mm, patentes, con indumento sericeo; eo-
lumna estaminal de ca. 1 1 mm de longitud X 1
mm de grosor, subglabra en la mitad inferior y es-
trellado-tomentosa en la mitad superior; las tecas
agrupadas en la zona ensanchada distal (de ca. 3
X 2 mm); lobulos de la columna estaminal de ca.
1 mm; las tecas 35 a 40, de reniformes a elipsoides,
de ca. 0.5 mm de longitud, por lo general agrupa-

348

Novon

TT

1 I 1 1 1

TT

TT

TT

IT

1

1

2

pulgadas

Figura 3. Quararibea cryptantha Femdndez- Alonso. Detalle de una rama con flores; en el centro una flor con la
columna estaminal y petalos desprendidos del lalamo floral. (Fotografiado de la serie tfpica. E. Carbond 2029.)

das de a dos; ovario bilocular, con I o 2 ovulos por
ldculo. Fruto (muy joven) fusiforme, comprimido,
de ca. 3 X 2.2 mm, con exocarpo lepidoto. Semillas
desconocidas.

Distribucidn y habitat. Quararibea cryptantha
se conoce solo de las estribaciones del sector no-
roccidental de la Sierra Nevada de Santa Marta, en
la franja subandina y por una coleccidn de la Se-
rranfa del Perija, recolectada en ambientes simi-
lares. De acuerdo con los estudios adelantados en
la region (Rangel, 1994), Q. cryptantha , se encuen-
tra en bosques dominados por: Viburnum tinoides
L. f. (Caprifoliaceae), Parathesis adenanthera (Mi-
quel) Hooker f. ex Mez (Myrsinaceae), Primus in-
tegrifolia (C. Presl) Walpers (Rosaceae) y Cithar-
exylum subflavescens S. F. Rlake (Verbenaceae).

Etimologia. El epiteto especffico hace referencia
a las flores muy pequenas y poco conspicuas de
esta especie de Quararibea, que son las de menor
tamano entre las especies colombianas y de las mas
pequenas en todo el genero.

Discusidn. Quararibea cryptantha pertenece a
un grupo de especies que presentan flores muy pe¬
quenas — grupo de Q. yunckeri Standley de Cen-
troamerica/(A floribunda (Saint-Hilaire) K. Schu¬
mann de Brasil. Quararibea cryptantha, se
diferencia de ellas por la siguiente combinacion de
caracteres: habito arborescente; hojas membranoso-

cartaceas, de oval-lanceoladas a oblongo-lanceola-
das, en el enves con domacios sacciformes en las
axilas de los nervios secundarios; papiraceos. no
barbados; pedicelo de 0.8—1 mm de longitud, caliz
verde, urceolado-campanulado, de 6—7 mm de lon¬
gitud, irregularmente 3 a 5 lobulado hacia el apice;
tubo estaminal de 11X1 mm.

Por el reducido tamano de sus flores, Q. cryp¬
tantha representa la antttesis de Q. ruiziana Fer-
nandez-Alonso ( 1 999b), especie recientemente des-
crita de Colombia, que presenta flores grandes, con
ramas estaminales muy largas y fruto con 3—4 pi-
renos.

Aunque el genero Quararibea ya se conoce de
la Serranfa del Perija por una coleccidn reciente
de Q. foenigracea Cuatrecasas sensu lato (Fernan¬
dez 13210, COL) y de la planicie atlantica por la
especie Q. asterolepis Pittier (Warner 507, FMB),
Q. cryptantha es la linica Quararibea registrada
hasta la fecha en el macizo de la Sierra Nevada de
Santa Marta. Con respecto al patron de distribucidn
restringido a Sierra Nevada-Perija se ha observado
tambien en otros grupos, ast en las Labiatae Hyptis
diffusa Epling y Salvia codazzina Eernandez- Alon¬
so y en la Scrophulariaceae Calceolaria leucanthera
Pennell (Fernandez-Alonso, 1995a, 1995b; Molau.
1988). Estas Ires especies eran conocidas origi-
nalmente solo de uno de los dos macizos, liabien-
dose colectado mas recientemente en ambos.

Volume 12, Number 3
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Fernandez-Alonso
Bombacaceae Neotropicae Novae

349

Pardtipos. COLOMBIA. Cesar: Serrania del Perija,
Mpio. de Agustfn Codazzi, Vereda Cara de Hombre, 1750
m, 25 May 1994, 0. Rangel el al. 12415 (COL. HUA,
MA, MO, UTMC).

Quararibea rangelii Fernandez-Alonso, sp. nov.
TIPO: Colombia. Boyacd: Mpio. de Puerto
Boyaca, Quebrada La Cristalina, 400 m,
bosque entresacado, 21 Sep. 1996 (fl, fr), O.
Rangel-Ch. 13475 (holdlipo, COL 500093;
isotipos, COL 500094, HUA, MA, MO. US).
Figura 4.

Haec species quoad calycem manifeste sulcato-arista-
tum Quararibeae pterocalycis Hemsley leniter affinis, sed
ab ea stipulis longioribus (2—3 cm longis), foliis mem-
branaceis, florum minorum calyce 2. 1-2.3 cm longo, cos-
tulis valde sinuatis, petalis ad 4—4.3 cm longis et 6—7 mm
latis, sub anthesi retroflexis; columna staminali 2.2— 2.4
cm longa; fructu 2.5-2. 7 longo et 3-3.5 cm lato, saepe
inaequilatero maturitate, plerumque monopyreno distin-
guitur.

Arboles de hasta 15—20 m, troncos con corteza
marron-grisacea y madera amarillo palido; ramas
con corteza marron, estriada, muy levemente aro-
matica (en seco); apice de las ramas jovenes re-
matado por un fascfculo de bracteas estipuliformes
(Fig. 4A) estrechamente linear-lanceoladas de 2.5—
4 cm de longitud y hasta 4 mm de ant ho en la
base. Hojas dfsticas; estfpulas 2, prontamente ca-
ducas, estrechamente lanceoladas, de 2—3 cm X 3—
4 mm, membranoso-papiraceas, con indumento
lepidoto en la lamina y estrellado-fasciculado en
los margenes. Hojas con peciolos de 1.5— 2(2.8) cm
de largo, con indumento estrellado-lepidoto muy
fino, pulvfnulo proximal notorio, elfptico, de 4—8 X
2.5— 3.5 mm, con aspecto de callo, pulvfnulo distal
inconspfcuo; laminas verde palido por la haz y ver-
de amarillento por el envds, membranosas, estre-
chamente lanceoladas, por lo general escasamente
asimetricas, de 30—45 X 9-15(-17) cm, cuneadas
en la base y de agudas a acuminadas en el apice;
glabras y levemente lustrosas por la haz y glabras
por el enves; con nerviacion pinnada, el nervio me¬
dio y 2 nervios basales laterales poco desarrollados
y hasta 9 nervios secundarios a cada lado del ner¬
vio medio; impresos en la haz y resaltados en el
enves; nerviacion de 3° y 4° orden reticulada, lig-
eramente resaltada. Inflorescencias de 1 a 4 flores
subsesiles agrupadas en ramas muy cortas; en bo-
ton rodeados por varias bracteas esteriles de la
yema (Fig. 4B) y con 3 bracteolas triangulares de
3—5 mm, sobre el pedicelo, cerca de la base del
caliz; pedicelo floral de 2-4 mm de largo, con in¬
dumento estrellado-fasciculado corto; flores acti-
nomorfas (Fig. 4C); caliz cilfndrico, de 2.1— 2.3 X
1—1.2 cm, con 10 costillas sinuosas, muy conspf-

cuas a lo largo del caliz, con indumento densa-
mente estrellado-lepidoto; petalos de 4—4.3 cm X
6-7 mm, reflejos, externamente blanquecinos con
indumento estrellatlo, internamente subglabros; co¬
lumna estaminal de 2.2-24 cm de longitud, es-
trellado-serfcea, las anteras dispuestas en la zona
ensanchada distal (de 3.5 mm de longitud y 3.5
mm de ancho) levemente lobada, los lobulos de ca.
1-2 mm (Fig. 41)); anteras ca. 40, estrechamente
reniformes. Fruto verde-anaranjado o verde-ama-
rillento (en vivo), notoriamente oblicuo-asimetrico
(Fig. 4E, F), de subesferico a reniforme, compri-
mido, de 2.5— 2.7 X 3—3.5 cm. por lo regular con
un solo pireno desarrollado; exocarpo con estrias
longitudinales, marron claro (en seco), estrellado-
lepidoto; pireno corneo, de ca. 3.3 X 1.8—2 cm,
fibroso en el exterior y liso-brillante en la cara
interna. Semilla con testa marron oscuro y cotile-
dones blanquecinos; semilla criptocotilar.

Nombres vernaculos y uso. En Puerto Boyaca,
la localidad tfpica (segun las etiquetas de herbario),
se la conoce por los siguientes fitonimos: “Guamo
rosado” y “Coco cristal”. La madera es blanda y se
utilizada en la region para la construccion de co-
rrales ( Galeano et al. 5394).

Distribucion y habitat. Las colecciones cono-
cidas de Quararibea rangelii provienen del inven-
tario recientemente efectuado en la Serrania de las
Quinchas, en el sector boyacense del valle del
Magdalena. Los bosques humedos donde crece Q.
rangelii, ubicados sobre substratos calizos y con
2500—3200 mm de precipitacion anual, ban sido
caracterizados botanicamente en fechas recientes
(Balcazar- Vargas et al., 2000). Segun las parcelas
levantadas en el estudio citado, Quararibea ran¬
gelii, es una especie muy frecuente en los bosques
de Hura crepitans L. (Euphorbiaceae) y Romeroa
verticillata Dugand (Bignoniaceae), dominados por
las familias Acanthaceae, Bignoniaceae, Bomba¬
caceae y Euphorbiaceae. Otras Bombacaceae con
interes corologico para la region y presentes en es-
tas formaciones sobre sustrato calizo son: Matisia
cardata, “zapote comestible”, que es especie nati-
va, M. obliquifolia Standley, “zapote de monte”,
ampliamente distribuida en Panama y en el norte
de Sudamerica; M. longiflora Gleason (sensu stric-
to), descrita del sector de Muzo en los anos 30, y
posteriormente localizada solo del otro lado del
Magdalena en las formaciones karsticas del Canon
de Rio Claro (Fernandez-Alonso & Cogollo, 1992);
Huberodendron patinoi y una especie afin a Quar¬
aribea asterolepis, perteneciente a un complejo
cuya taxonomfa no ha sido aun plenamente resuelta
para el territorio colombiano.

350

Novon

Figura 4. Quararibea rangelii Fernandez- Alonso. — A. Aspecto de una rama con hojas, estipulas y bracteas de la
yema. — B. Boton floral mostrando las bracteolas en el pedicelo. — C. Detalle de una llor en el extreme de una rama
corta. — I). Detalle de la porcibn superior de la columna estaminal con las anteras. — E. Fruto en vista lateral. — F.
Seccibn sagital del fruto con un pireno. (Dibujado a partir de: A de P. Franco el al. 5760 ; B y E de J. Betancur 6681 ;
C, I) y F de 0. Rangel-Ch. 13475.)

Volume 12, Number 3
2002

Fernandez-Alonso
Bombacaceae Neotropicae Novae

351

Hasla (londe se conoce, Q. rangelii, se encuentra
restringida a la vertiente occidental de la Cordillera
Oriental de Colombia. Todos los registros provienen
de la Serranfa de las Quinchas, Magdalena Medio
(Boyaca), en la franja de bosques de terraza. ubi-
cada a 300—700 m. Quararibea rangelii se suma a
la importante lista de taxones endemicos del Mag¬
dalena Medio colombiano, que incluye numerosas
especies y algunos generos (Fernandez-Alonso et
al., 2000).

Etimologia. Especie dedicada al botanico co¬
lombiano y colega Orlando Rangel Churio (COL),
quien centro sus esfuerzos en la descript ion y en
el conocimiento integral de las comunidades ve-
getales de Colombia, con resultados admirables.
Fruto de una de slis numerosas exploraciones por
el territorio colombiano son las muestras de la serie
tipo de la especie aquf descrita.

Discusidn. Quararibea rangelii es ligeramente
affn a Q. pterocalyx Hemsley por presentar cal ices
marcadamente sulcado-aristados, pero se diferencia
de ella entre otros caracteres por sus estfpulas mas
largas (2—3 cm long.), hojas membranaceas, mas
alargadas; flores menores, caliz de 2.1— 2.3 cm de
longitud, con costillas marcadamente sinuadas, pe-
talos menores, de 4—4.3 cm de longitud y 6—7 mm
de ancho, reflejos en la antesis, columna estaminal
mas corta de 2.2— 2.4 cm de longitud, fruto menor
y muy comprimido, de 2. 5-2. 7 X 3—3.5 cm, a me-
nudo inequilatero y en la madurez por lo general
con un solo pireno desarrollado.

Pardtipos. COLOMBIA. Boyaca: Mpio. tie Puerto
Boyaca, Inspeccion de Puerto Romero, alrededores de la
Quebrada La Fiebrecita, 400—680 m, 11 Oct. 1996 (fr),
./. Betancur 6 681 (COL. HUA); Quebrada La Fiebrecita,
300 m, 23 Nov. 1 997 (II), G. Galeano et al. 5894 (COL,
FMB, U1TC); Vereda La Cristalina, bosque secundario en
terraza, 21 Sep. 1996, E. Linares et al. 4415 (COL,
JAIJM); Puerto Romero, Vereda Las Palomas, 380 m, 15
Jun. 1997, P. Franco et al. 5760 (COL, F. JAIJM); Que¬
brada La Cristalina, 400 m, bosque en base de ladera, 21
Sep. 1996, 0. Rangel et al. 18.489 (COL, MO).

Agradecimientos. Quiero agradecer a la Asocia-
cion Columbiana de Herbarios y a Fabio Garcia
Cossio del Herbario CHOCO el haber facilitado la
visita de coleccion de Bombacaceae al Choc 6; a
Eduino Carbono (UTMC) y Alvaro Gogollo (JAUM)
el haber prestado y facilitado el estudio de las co-
lecciones de sus respectivos herbarios; a Paula Bal-
cazar, Orlando Rangel, Edgar Linares, Julio Betan¬
cur y Gloria Galeano, el acceso a las muestras

provenientes de la Serranfa de las Quinchas (Bo¬
yaca); a German Lopez que elabord las dos laminas

que se incluyen en el trabajo.

Literatura Citada

Balcazar- Vargas, M. P., J. 0. Rangel-C & F. Linares-C.
2000. Diversidad floristica de la Serranfa de las Quin¬
chas, Magdalena Metlio (Colombia). Caldasia 22: 191 —
224.

Cuatrecasas, J. 1946. Notas a la Flora de Colombia VIII.
Revista Acad. Colomb. Ci. Exact. 6: 533—551.

Fernandez-Alonso. J. L. 1992. Nueva especie de Matisia
(Bombacaceae-Quararibeae) del occidente de Colombia.
Anales Jard. Bot. Madrid 50: 171-174.

- . 1995a. Estudios en Labiatae de Colombia I. No-

vedades en los generos Salvia e Hyptis. Revista Acad.
Colomb. Ci. Exact. 19: 469-480.

- . 1995b. Estudios en Labiatae de Colombia II. No-

vedades en Salvia sect. Longipes Epl. Anales Jard. Bot.
Madrid 53: 4 1 — 46.

- . 1996. Contribuciones al conocimiento del genero

Phragmotheca Cuatr. (Bombacaceae-Quararibeae). Cal¬
dasia 18: 253-284.

- . 1999a. Nueva especie de zapote de monte

( Phragmotheca , Bombacaceae) en bosques premontanos
del centro de Colombia. Caldasia 21: 125—131.

- . 1999b. Nueva especie y notas al genero Qua¬
raribea (Bombacaceae). Revista Acad. Colomb. Ci. Ex¬
act. 23 (Supl. espec.): 49—52.

- . 2001a. Bombacaceae neotropicae novae vel mi¬
nus cognitae I. Novedades taxonomicas y corologicas en
Matisia, Quararibea y Spirotheca. Revista Acad. Co¬
lomb. Ci. Exact. 25: 183-206.

- . 2001b. Bombacaceae neotropicae novae vel mi¬
nus cognitae 11. Novedades taxonomicas y corologicas
en Matisia y Quararibea del norte de Sudameriea. Cal¬
dasia 23: 351-382.

- & A. Cogollo. 1992. Notas sobre Matisia longiflo-

ra Gleas. (Bombacaceae). Caldasia 17: 173—176.

- , J. A. Perez-Zabala & A. Idarraga-Piedrahita.

2000. Isidodendron, un nuevo genero neotropical de ar-
boles de la lamilia Trigoniaceae. Revista Acad. Colomb.
Ci. Exact. 24: 347-357.

Eorero, E. & A. Gentry. 1989. Lista anotada de las plantas
del Departamento del Choco, Colombia. Biblioteca Jose
Jeronimo Triana 10. Institute de Ciencias Naturales,
Universidad Nacional de Colombia, Bogota.

Gentry, A. 1986. Species richness and floristic composi¬
tion of Choco region plant communities. Caldasia 15:
71-91.

Molau, U. 1988. Scrophulariaceae Part. I. Caleeolarieae.
Flora Neotropica Monogr. 47.

Rangel, 0. 1994. Caracterizacion de la vegetacion del cos-
tado norte de la Serranfa del Perija. Informe final. In¬
stitute de Ciencias Naturales-CORPOCESAR.

Robyns, A. 1968. Bombacaceae neotropicae novae II. New
species of Eriotheca, Hampea and Quararibea. Ann.
Missouri Bot. Card. 55: 51—59.

- & S. Nilsson. 1970. Bombacaceae neotropicae no¬
vae III. A new species of Quararibea from Panama.
Bull. Jard. Bot. Nat. Belg. 40: 353-356.

Notes and New Species of Moraea (Iridaceae: Iridoideae) from the
Southern African Winter-Rainfall Zone

Peter Goldblatt

B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299,
St. Louis, Missouri 63166, U.S.A. peter.goldhlatt@mobot.org

John C. Manning

National Botanical Institute, P. Bag X7, Claremont 7735, South Africa.
manning@nbict.nbi.ac.za

ABSTRACT. Moraea cantharophila and M. lilacina
are new species of the African genus Moraea (ca.
195 species), of subfamily Iridoideae of the Irida¬
ceae. Both species belong to Moraea subg. Vieus-
seuxia, which comprises 35 species restricted to
southern Africa and is centered in the southern Af¬
rican winter-rainfall zone. Moraea cantharophila,
from southwestern Western Cape Province, has un¬
usual flowers for the subgenus with entire relatively
large inner tepals with a broad dark median streak
on the claw. It is distinguished from its closest rel¬
ative, the fly-pollinated M. lurida, by the shallower
floral cup and unscented nectarless flowers. Moraea
lilacina, from the northern Langeberg foothills in
the Little Karoo of southeastern Western Cape
Province, can be distinguished from the related M.
unguiculata by its relatively large flower, pink pig¬
mentation, trilobed inner tepals with a long, erect
central cusp, and often densely hairy filament col¬
umn. Nomenclatural adjustments include the re¬
duction of M. neopavonia in M. tulbaghensis and
the provision of a new name for the homonym M.
flexuosa Goldblatt. Newly discovered populations of
M. saxicola and M. rivulicola increase their known
geographical ranges and require some adjustment
to their delimitation.

Key words: beetle pollination, Iridaceae, Iridoi¬
deae, Moraea, Moraea subg. Vieusseuxia.

Moraea Miller, now comprising some 195 species
(Goldblatt, 1998; Goldblatt & Manning, 2000a), is
the largest African genus of tribe Irideae of the
Iridoideae, one of four subfamilies of the Iridaceae
currently recognized (Goldblatt, 1990, 2001). Mo¬
raea, including Barnardiella, Gcdaxia, Gynandriris,
Hexaglottis, Homeria, and Koggeveldia, occurs
widely across sub-Saharan Africa and has two spe¬
cies in the Mediterranean— Middle East; it is now
defined by bifacial, channeled to flat leaves and a
unique type of cormous rootstock consisting of a

single enlarged internode that is derived entirely
from an axillary bud (Goldblatt, 1976b, 1990). The
genus is currently divided into five subgenera
(Goldblatt, 1986). Moraea subg. Vieusseuxia, to
which the new species belong, is restricted to
southern Africa, south of the Limpopo— Cunene
axis, and is centered in the winter-rainfall zone in
the southwest of the subcontinent. Including the
new species described here and the synonymous
reduction of M. neopavonia R. C. Foster in M. tul¬
baghensis L. Bolus, Moraea subg. Vieusseuxia now
includes 35 species.

The winter- rainfall zone of southern Africa is the
center of Moraea both in species diversity and evo¬
lutionary depth. All five subgenera and over two-
thirds of the species of the genus occur in this small
portion of the African continent, extending from
southwestern Namibia along the western and south¬
ern coast of South Africa to Port Elizabeth. This
area of highly dissected landscape and a mosaic of
different soils has very steep ecological gradients
that expand the number of ecological niches avail¬
able for plant species. This situation has promoted
extraordinary radiation in a range of genera in
many plant families there, including geophytic gen¬
era of Iridaceae like Babiana (ca. 75 species), Geis-
sorhiza (over 80 species). Gladiolus (some 145 spe¬
cies), and of course Moraea (Goldblatt & Manning,
2000b). Many species of the southern African win¬
ter-rainfall zone have narrow ranges, and as might
be expected of a flora of semi-arid to arid habitats,
flowering is often dependent on the highly variable
seasonal rainfall. The flowering of geophytes such
as Moraea is also related to the fire cycle, an im¬
portant factor in the vegetation of the region. One
of the species described here. M. cantharophila,
flowers well only after wild fires, and at some sites
evidently only in the first spring season after fire.
I’he other new Moraea described here is a narrow

Novon 12: 352-359. 2002.

Volume 12, Number 3
2002

Goldblatt & Manning
Moraea from Southern Africa

353

geographic endemic of dry valleys in the central
southern Cape. Variable rainfall, narrow distribu¬
tions, and erratic flowering make the documentation
of the southern African geophyte flora difficult. It
is not so surprising then, that new species of narrow
distribution or of specialized habitat continue to be
discovered in a part of the world that has enjoyed
the attention of botanists for over 230 years since
Linnaeus’s student Carl Peter Thunberg collected
there in the 1770s. We suspect that additional spe¬
cies, especially geophytes, will continue to be rec¬
ognized as local exploration of the flora continues.

Exsiccatae are cited below' following the quarter-
degree square system in use in southern Africa as
outlined by Edwards and Leistner (1971).

1. Moraea cantharophila Goldblatt & J. C.
Manning, sp. nov. T\ PE: South Africa. West¬
ern Cape: Sandy’s Glen, loamy clay slope on
farm Vredehoek, 16 Sep. 2000, P. Goldblatt &
J. C. Manning 11542 (holotype, NBG; iso¬
types, K, MO, PRE). Figure 1.

Plantae 20-45 cm altae, cormo globoso 15-20 mm
diam. tunicis reticulatis fibrosis pallidis obtecto, folio unico
producto lineari canaliculato 2.5— 3.5 mm lato, caule er-
ecto usitate ramoso, inflorescentiae spatha externa 35—45
mm longa, interna 45—65 mm longa, florihus albis vel cre-
meis tepalis externis prope basem limbi flavo- vel auran-
tiaco-notatis, tepalorum unguibus plerumque flavis ca. 10
mm longis, limbis patentibus, externis (15— )18— 20 X 14—
19 mm, internis 8—15 X 5—11 mm, filamentis 4—5 mm
longis in columnam omnino connatis vel superne per 1
mm liberis, antheris purpureis ca. 6 mm longis, styli ramis
6-7 mm longis cristis 2.5-4 mm longis ornatis, ovario
exserto 7—12 mm longo, capsula oblongoidea, 16—18 mm
longa.

Plants 20—45 cm high. Corm globose, 15—20 mm
diam., the tunics of coarse, pale straw-colored, net¬
ted fibers. Cataphylls papeiy, uppermost reaching
shortly above the ground, becoming dry and dark
brown by flowering time. Foliage leaf solitary, lin¬
ear, channeled, arching outward, overtopping the
stem and often dry toward the tip, mostly 2.5— 3.5
mm wide, inserted on the stem at or close to ground
level. Stem erect, straight or flexed at nodes, always
arching outward below the inflorescence, usually
with two aerial internodes, bearing an entirely
sheathing bract-like leaf at the aerial node, this
4.5— 5.5 cm long, green with a ± dry, attenuate
apex, simple or with one, rarely two branches in
the leaf axil and aerial node, the branches arching
outward above the sheathing leaf. Inflorescence a
rhipidium, terminal on the branches, mostly 2- or
3-flowered; spathes unequal, green, with dry atten¬
uate apices, the outer 35—45 mm long, the inner
45—65 mm long. Flowers white or cream, the tepals

Figure 1. Moraea cantharophila Goldblatt & J. C. Manning
(Goldblatt & Manning 11580). — A. Corm. — B. Flowering
stem and leaf. — C. Detail of stamens and style branches.
Scale bar I cm (A, B), 3 mm (G). Drawn by J. C. Manning.

354

Novon

darkly veined on the outside, the outer tepals each
with a yellow or orange nectar guide outlined in
dark purple at the limb base, the claw mostly yel¬
low, the inner tepals each with a broad, dark pur¬
ple-brown central band and pinkish edge, the tepal
limbs spreading ± horizontally or the tips curving
slightly upward, the claws ascending, forming a cup
ca. 12 mm wide at the mouth, enclosing the fila¬
ment column and anther bases; tepals clawed, the
outer larger than the inner, the claws ca. 10 mm
long, limb of the outer tepals obovate, (15— ) 18— 20
X 14—19 mm, without a basal nectary, the claw
smooth or papillate, limb of the inner tepals 8—15
X 5—11 mm. Filaments 4—5 mm long, united in a
cylindric column or free in the upper 0.5—1 mm,
smooth, greenish to dark purple; anthers ca. 6 mm
long, oblong, diverging, appressed to the style
branches, purple, the pollen orange. Ovary exsert-
ed, oblong-triquetrous, 7-12 mm long; style divid¬
ing at the apex of the filament column, yellow to
salmon pink with a dark central band, the branches
6—7 X ca. 3 mm, ascending, the stigmatic Hap bi-
lobed, the crests triangular, uniformly salmon pink.
± erect, 2.5—4 mm long. Capsules ± oblong, 16—
18 mm long; seeds unknown.

Flowering late August and September, rarely in
early October; (lowers lasting 3 days, the tepals
arching upward and partially closing at night.

Etymology. From the Latin cantharophila,
“beetle-loving,” referring to the pollinators, hopb¬
ine scarab beetles, which visit the (lowers and
transfer pollen from one plant to another.

Distribution and habitat. Restricted to the
southwestern Cape, Moraea cantharophila occurs
on mountain slopes between the foot of Sir Lowry’s
Pass near Somerset West and Sandy’s Glen in the
Bredasdorp Mountains near Napier. Il grows in
loamy ( lay and not in the coarse sandy substrate
that is more common in the Cape mountains.

Relationships. A member of subgenus Vieus-
seuxia (Goldblatt, 1976b, 1986), M. cantharophila
has a single, long, channeled (oliage leaf typical of
the subgenus, a branched stem, but unlike the ma¬
jority of the species the inner tepals are not tricus-
pidate, but are broadly lanceolate with an entire,
acute apex. It is distinctive in the shape of the in¬
ner tepals, and it is one of the few species in the
subgenus with entire inner tepals with a broad,
well-developed limb. The filaments are united to
the apex, also rare in the subgenus, in which most
species have the upper 1—2 mm of the filaments
free and diverging. The dark central streak on the
inner tepals is distinctive and is directly related to
the pollination biology. Moraea cantharophila is ev¬

idently most closely related to two other south¬
western Cape species, M. luridci Ker Gawler and
M. insolens Goldblatt, which also have entire inner
tepals with a well-developed limb. Fly-pollinated
M. lurida has flowers with a fetid, rotting scent and
typically a darker-colored perianth, both the inner
and outer tepals of which produce nectar over the
entire inner surface of the claw, and the outer tepals
do not usually have contrastingly colored markings.
An important distinction between M. lurida and M.
cantharophila is that the tepal (daws of the former
are relatively long, 13—17 mm, and slightly exceed
the claw, compared to ca. 10 mm in M. cantharo¬
phila, this shorter than the limb. The tepal cup is
thus deeper in M. lurida and includes both stamens
and style branches. The anthers are partly exserted
in M. cantharophila. Moraea insolens has rather dif¬
ferent flowers with the short tepal (daws spread al¬
most horizontally, thus not forming a cup, and the
anthers are ca. 8 mm long and exceed the short
style crests less than 1 mm long (Goldblatt, 1976b).
It is pollinated by hopbine beetles, as is M. can¬
tharophila, and lacks nectaries, thus producing no
nectar (Goldblatt et al., 1998; Goldblatt, unpub¬
lished data).

All three species occur in the same area, that is,
from Sir Lowry’s Pass eastward to Gape Agulhas,
hut M. insolens has the narrowest range, occurring
along a band of loamy soil at the foot of the Caledon
Swartberg. Moraea lurida occurs only on stony,
sandstone-derived soils. These three species flower
well after a wild fire the previous summer or au¬
tumn, and M. cantharophila and M. lurida flower
poorly, if at all, until another fire. The known pop¬
ulations of M. cantharophila vary to some extent.
I he flowers of the Sir Lowry’s Pass population lack
floral odor entirely, and the inner tepals are white
with a broad, dark median streak, like those of the
Sandy’s Glen plants, which have a broad, dark
brown streak in the midline. I he tepals of the Sir
Lowry’s Pass plants are smaller and 8—9 X 5—6 mm
vs. 12—15 X 9—1 1 mm in the Sandy’s Glen popu¬
lation.

Plants collected in the Elgin area east of Sir
Lowry’s Pass ( Goldblatt 11633, 11638) appear to
he an unusual form of Moraea cantharophila and
may represent a form intermediate between this
species and related M. lurida. Although they have
white flowers with yellow inner tepals with a dark
median streak or yellow inner and outer tepals, the
inner tepals have short limbs mostly 3—4 X 5 mm,
either entire or three-lobed with a short, acute cen¬
tral cusp. The flowers have an unpleasant, fetid
odor, and the outer tepal claws are lightly papi 1 late
and produce a sweet exudate. Typical M. canthar-

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2002

Goldblatt & Manning
Moraea from Southern Africa

355

ophila lias entire inner tepal limbs 8— 15 X 5—11
mm. The fetid odor and sweet exudate on the tepal
claws are features of M. lurida (Goldblatt, 1986,
and unpublished obs.), which has short inner tepal
limbs that are occasionally trilobed. The possibility
of hybridization between more typical M. canthar-
ophila and M. lurida also occurred to us as an ex¬
planation for the strange floral morphology in the
Elgin plants. No M. lurida was, however, found on
sandstone slopes adjacent to the shale soils on
which M. cantharophila grew.

Moraea cantharophila flowers have the features
that are now associated with pollination by hopbine
or monkey beetles (Scarabaeidae: Rutelinae: Ho-
pliini): bright coloration, bold, contrasting mark¬
ings, a salver- or bowl-shape, absence of floral odor
(usually), and little or no nectar (Steiner, 1998;
Goldblatt et ah. 1998, 2000). Two species of ho¬
pbine beetles have been captured in the flowers,
Anisonyx sp. and Peritrichia pseudoplehia, the in¬
sects typically found at rest with their body nested
in the space between the tepal claws and style
branches. All beetles captured carried heavy loads
of M. cantharophila pollen on their bodies. Hopb¬
ines normally do little damage to flowers, although
they consume some pollen, instead using the flow¬
ers as sites for assembly, competitive behavior, and
copulation. At the type site, in Sandy’s Glen in the
Bredasdorp Mountains, M. cantharophila formed a
guild with two other species with similarly colored
and shaped flowers. Drosera cistifiora L. and Aristea
teretifolia Goldblatt & J. C. Manning. Both species
at this site have cream flowers with dark brown
central markings and prominent anthers bearing
bright yellow pollen. At the Sir Lowry’s Pass site,
M. cantharophila formed a guild with Aristea can¬
tharophila Goldblatt & J. C. Manning, Drosera cis-
tijlora, and I). pauciflora L. All four species were
pollinated there by two other hopbine beetles, Per¬
itrichia pseudoplehia and Anisonyx ursus (Goldblatt
et ah, 1998). Despite the fetid odor in the plants
from Elgin, suggesting muscid fly pollination, we
noted that these plants are, like other populations
of M. cantharophila, also pollinated by hopbine
beetles, in this case Anisonyx hessei. At one site,
Die Hawe, we noted small bees visiting the flowers.

Paratypes. SOUTH AFRICA. Western Cape: 3418
(Simonstown) near the foot of Sir Lowry’s Pass (BB), 28
Aug. 1995, Goldblatt & Manning / 0281 (K, MO, NBG);
3419 (Caledon) foot of Viljoen’s Pass, clay ground (AA),
5 Sep. 1974, Goldblatt 2491 (MO); slopes of the Green¬
land Mountains N of Elgin, 10 Oct. 2000, Goldblatt 11633
(K, MO, NBG, PRE); Die Hawe, Elgin, 10 Oct. 2000,
Goldblatt 11638 (MO, NBG).

2. Moraea lilacina Goldblatt & J. C. Manning,
sp. nov. TYPE: South Africa. Western Cape:
northern foothills of the Langeberg, near IVr-
debont. Farm Graskloof, 23 Sep. 1979. P.
Goldblatt & I. Nanni 11587 (holotype, NBG:
isotypes, K, MO). F igure 2.

Plantae 20—35 cm altae, cormo usitate globoso 12-16
mm diam. tunicis reticulatis fibrosis pallidis obtecto, folio
unico producto lineari anguste canaliculato ca. 1.5 mm
lato, caule erecto usitate eramoso, inflorescentiae spatha
externa 25—30 mm longa, interna 40-45 mm longa, flori-
bus perpallide roseis, tepalorum externorum 27—30 mm
longorum abaxialiter lilacino suffusorum unguibus flavis
ca. 10 mm longis ac limbis patentibus, internorum usque
ad 18 mm longorum tricuspidatorum unguibus brunneis
ca. 9 mm longis canaliculatis, filamentis 7—8 mm longis
infra in columnam connatis superne liberis divergentibus,
antheris albescentibus ca. 3.5 mm longis, styli ramis ca.
5X2 mm cristis 5—8 mm longis gracilibus ornatis, ovario
exserto 7—8 mm longo, capsula ignota.

Plants 20—35 cm high. Corm globose or flattened
by pressure, 12-16 mm diam., tunics of light
brown, netted fibers. Leaf solitary, linear, narrowly
channeled or the margins conduplicate, ca. 1.5 mm
wide, trailing and often dry above. Stem erect, nor¬
mally unbranched, with three aerial internodes, the
nodes each bearing a sheathing, bract-like leaf AD¬
AS mm long, the apex attenuate and dry. Inflores¬
cence mostly 2-flowered; spathes unequal, green,
with dry attenuate apices, the outer 25—30 mm
long, the inner 40-45 mm long. Flowers palest
pink, flushed lilac-pink on the reverse of the outer
tepals, fading darker pink, the outer tepals each
with a bright yellow nectar guide at the base of the
limb; tepals unequal, the outer larger, 27—30 mm
long, the claw ascending, ca. 1 0 mm long, yellow,
densely hairy, bearing a nectary at the base partly
covered by a short flap of tissue ca. 1 mm long, the
limb spreading at ca. 45° below horizontal, inner
tepals to 18 mm long, tricuspidate, the claw ca. 9
mm long, brown, channeled, with a long, suberect
central cusp ca. 9 mm long, and short, obtuse lat¬
eral lobes. Filaments 7—8 mm long, united in the
lower two-thirds into a cylindrical column, smooth
or papillate, free and diverging above; anthers ap-
pressed to the narrow style branches, ca. 3.5 mm
long, reaching to within 1 mm of the stigma lobes.
Ovary oblong, 7-8 mm long, exserted from the
spathes; style dividing at the apex of the filament
column, the branches diverging, ca. 5X2 mm,
crests slender, erect, 5—8 mm long. Capsules and
seeds unknown.

Flowering September.

Etymology. From the Latin lilacina, “lilac-col¬
ored,” referring to the color of the tepals, flushed
lilac below and darkening to lilac-pink with age.

356

Novon

Figure 2. Moraea lilacina Goldblatt & J. C. Manning
( Goldblatt & Nanni 11587). — A. Corm. — B. Flowering
stem and leaf. — C. Detail of stamens and style branches.
Scale bar 1 cm (A, B). 3 mm (C). Drawn by J. C. Manning.

Distribution and habitat. Restricted to the
southern mountains of Western Cape Province, Mo¬
raea lilacina is known from the type site in the
northern foothills of the Langeberg, northwest of
Robinson’s Pass. Plants grow in loamy soil mostly
wedged in broken sandstone rock.

Relationships. Moraea lilacina is closely allied
to the widespread southern African winter-rainfall
M. unguiculata Ker Gawler. which is a remarkably
variable species, both as regards perianth color and
size. Important features of M. unguiculata are the
inner tepals, which are 10—12 mm long and trilobed
with the central cusp linear and coiled inward. The
filaments are usually 3—5 mm long, with the column
lightly to densely papillate, and the outer tepals are
typically 12—20 mm long, exceptionally to 24 mm
long in northern Namaqualand populations, with
the limb (5-)8-12(-14) mm. file outer tepals may
be white to cream, brown, violet, or even dull yel¬
low and usually have a contrasting mark at the base
of the limb. The style branches are narrow, spread¬
ing as in M. lilacina, but with erect crests 2— 4(—
5.5) mm long. Extending from Steinkopf in northern
Namaqualand in the north to Humansdorp in the
southeast, with isolated populations in the Karoo
and Eastern Cape Mountains (Goldblatt, 1986;
Goldblatt & Manning, 1995), M. unguiculata typi¬
cally favors shale slopes and grows in renosterveld
or transitional fynbos-renosterveld, but populations
are known from sandstone slopes. Moraea lilacina
favors a similar habitat. It can be recognized mainly
by the large size of the flowers, which have inner
tepals 16—18 mm long with an erect central cusp,
filaments 7-8 mm long with a papillate or smooth
column, and outer tepals 27-30 mm long. Most of
the 16 plants we examined had unbranched stems,
whereas individuals of M. unguiculata almost al¬
ways have branched stems unless they are depau¬
perate due to environmental stress.

Paratypes. SOUTH AFRICA. Western Cape: 3321
(Oudtshoorn) valley E of Perdebont, ridge SW of Beacon
90, S-facing slopes (DD). 16 Sep. 1998, E. G. H. Oliver
& I. Oliver 10874 (NBG).

Taxonomic Notes

REDUCTION OF MORAEA NEOPAVONIA IN M.
TULBAGHENSIS

Moraea tulhaghensis L. Bolus, S. African Gard.

22: 276. 1932. TYPE: South Africa. Western

Cape: Saron, //. M. L. Bolus s.n. (holotype,
BOL).

Moraea neopavonia R. Foster. Contr. Gray Herb. 165: 108.

1947. Syn. nov. Iris pavonia L. f„ Suppl. PI. 98.

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2002

Goldblatt & Manning
Moraea from Southern Africa

357

1781. Moraea pavonia (L. f.) Ker Gawler, Konig &
Sims, Ann. Bot. I: 240. 1804, nom. iileg., non M.
pavonia (L. f.) Thunberg, Diss. Moraea, 14. 1787 | =
Tigridia pavonia (L. f.) DC.|. TYPE: Soutli Africa.
Western Cape: Swartland hills, Thunberg s.n. (lec-
totype, designated by Goldblatt (1976b), UPS. Herb
Thunberg 1148A).

In the most recent account of Moraea in southern
Africa (Goldblatt, 1986) M. tulbaghensis and M.
neopavonia were regarded as closely allied species
of subgenus Vieusseuxia. They shared with a small
group ol species within 1 he subgenus, including t he
widespread M. villosa (Ker Gawler) Ker Gawler, a
solitary foliage leaf hairy on the abaxial surface and
a shortly hairy stem. Of these species, only the
blue-flowered M. gigandra L. Bolus, M. neopavon¬
ia, and M. tulbaghensis have narrow style brant lies,
as wide as or hardly wider than the anthers, and
short style crests 1—2 mm long, which are over¬
topped hy the anthers. Both M. neopavonia and M.
tulbaghensis have orange (rarely to red) tepals and
are distinguished in the key by the anthers exceed¬
ing the style crests by more than 1 mm (M. neo¬
pavonia) or by up to I mm [M. tulbaghensis). This
distinction seemed useful in 1986 (Goldblatt, 1986)
and was supported by the presence of brilliant
green nectar guides on the outer tepals and anthers
7—10 mm long in M. tulbaghensis versus navy blue
or speckled nectar guides and anthers 9-12 mm
long in M. neopavonia. The type population of the
latter species in the west of its range near Piketberg
has boat-shaped, attenuate inner tepals, but other
populations have tricuspidate inner tepals with a
long, narrow central cusp, as does M. tulbaghensis.

New collections of both species have rendered
this distinction untenable. Plants from Elandberg
between Gouda an< 1 H ermon have flowers that seem
to be most like those of Moraea tulbaghensis, but
the deep orange to reddish outer tepals have navy
blue nectar guides and anthers about 9 mm long
exceeding the style crests by more than 1 mm. We
no longer find it possible to distinguish the two spe¬
cies and therefore unite them under the earliest
valid name, M. tulbaghensis. Available chromosome
counts, for one population of each species, are 2 n
= 12 (as M. neopavonia) and 2 n = 24 (as M. tul¬
baghensis) (Goldblatt, 1971a, 1976a). The basic
chromosome number in Moraea subg. Vieusseuxia
is x — 6. Thus in its revised circumscription M.
tulbaghensis is heteroploid with both diploid and
tetraploid populations. Related M. villosa also has
diploid and tetraploid races, as do a handful of oth¬
er species in the genus (Goldblatt, 1981).

NOTES ON MORAEA SAXICOIA GOLDBLATT

A species of Namaqualand, that is the coast and
near interior of northwestern South Africa, Moraea

saxicola (subg. Moraea sect. Deserticola) has a re¬
corded range from Steinkopf in the north to near
the town of Nuwerus in the south. Over this wide
range its morphology is relatively uniform except
that in the type population, from the Spektakel
Mountains west of Springbok, plants have flowers
with pale blue-mauve tepals and deep green leaves.
Elsewhere flowers are almost white with the outside
of the tepals faintly flushed with mauve or brown,
and the leaf is gray-glaucous. Plants most closely
resembling M. saxicola, collected in 2000 and
2001, some 50 to 70 km south of the known range,
below the foot of Vanrhyns Pass near Vanrhynsdorp
[Goldblatt 11616) and at the foot of the Kobee
Mountains ( Goldblatt <£ Porter 117BB), are much
smaller than all known populations of that species
in all parts. They are mostly only 7—10 cm high
and have a trailing to almost prostrate leaf with
strongly undulate and crisped margins and a some¬
what twisted blade. The small flowers are pale
mauve with deep yellow nectar guides. Plants grow
on the slopes of low hills in clay loam among
quartzite pebbles, a common habitat in the area.
The Vanrhyns Pass plants, collected in flower, have
inflorescence spathes 25—30 mm long, and flowers
with outer tepals 22—24 mm long, inner tepals ca.
18 mm long, filaments ca. 7 mm long, anthers ca.
3 mm long, and style crests 7—8 mm long.

The recorded variation for Moraea saxicola for
these features is consistently larger: inflorescence
spathes 30-60 mm long; outer tepals 30-40 mm
long; inner tepals 25—30 mm long. Other floral
parts similarly exceed those in this southern pop¬
ulation, with filaments 7-13 mm, anthers 6—9 mm,
style branches 8—15 mm. and crests mostly 10—15
mm long. Despite the size differences, the aspect
of the plants is so consistent with M. saxicola that
for the present we think this southern population is
best regarded as an arid habitat ecotype of M. sax¬
icola and should thus be included here, despite the
argument that can be made for infraspecific rec¬
ognition.

The vegetation where the southern populations
of Moraea saxicola grow consists of a sparse cover
of succulent shrubs, mostly Aizoaceae-Mesem-
bryanthemoideae, and geophytes including Moraea
serpentina Baker, Babiana salteri G. J. Lewis, and
B. sinuata G. J. Lewis (Iridaceae), and Ornithogal-
um polyphyllum Jacquin (Hyacinthaceae). It is
probably no coincidence that flowers of M. serpen¬
tina that are similar in size, coloring, and general
morphology fade and deliquesce at about the time
that those of M. saxicola open, at ca. 16:0011. blow¬
ers of M. serpentina utilize the late morning and
early afternoon for display and those of this popu-

358

Novon

lation of M. saxicola the late afternoon. Presumably
they share the same set of pollinators. In other pop¬
ulations of M. saxicola flowers have been recorded
as opening after 13:00H and as late as 15:00H
(Goldblatt, 1976b).

Additional specimens. SOUTH AFRICA. Western
Cape : 31.118 (Vanrhynsdorp) foot of Kobee Pass, E of
Vanrhynsdorp, in quartzite pebble areas (BD), Sep. 2000,
Goldblatt & Porter 11788 (MO, NBG); foot of Vanrhyns
Pass (DB), Oct. 2000, Goldblatt 11616 (K, MO, NBG,
PRE).

MORAEA FLEXICAULIS, A NEW NAME FOR THE
HOMONYM M. FLEXUOSA GOLDBLATT

Moraea flexicaulis Goldblatt, nom. nov. pro M.
flexuosa Goldblatt, Ann. Missouri Bot. Card.
69: 356. 1982, nom. i I leg., non M. flexuosa L.
1., Suppl. PI. 100. 1781 (= M. lewisiae (Gold¬
blatt) Goldblatt). TYPE: South Africa. North¬
ern Cape: Richtersveld, Eksteenfontein road,
Goldblatt 6000A (holotype, MO; isotypes, K,
NBG, PRE, S, WAG).

The name Moraea flexuosa Goldblatt is a hom¬
onym for M. flexuosa L. f., an epithet in turn ille¬
gitimate (Goldblatt, 1971b) because it is superflu¬
ous for M. longifolia Jacquin, which dates from
1776. Moraea flexuosa L. f. is currently called M.
lewisiae (Goldblatt) Goldblatt, and the epithet lew¬
isiae was proposed for that species, then included
in the genus Hexaglottis, because the combination
H. flexuosa (L. f.) Sweet was, like the basionym,
superfluous. Nevertheless, the name M. flexuosa L.
f. is valid, and prevents the subsequent use of the
epithet in Moraea. The new name M. flexicaulis is
proposed here for the later homonym.

We also report a range extension for this rare
species of the southern Richtersveld in Namaqua-
land. The species was described from the Anenous
flats on the farm Grasvlakte in 1982. During field¬
work in the Richtersveld in September 2000, we
found plants at the type locality and some 20 km
to the north in stony ground at the foot of low hills
adjacent to the road to Eksteenfontein ( Goldblatt ,
Manning & Savolainen 11494, 11 Sep. 2000,
NBG). Plants were in fruit, but the distinctive veg¬
etative morphology makes it all but certain that
they represent 17. flexicaulis.

RANGE EXTENSIONS FOR MOKAEA RIVUL1COLA AND AN
EMENDED CIRCUMSCRIPTION

Described in 1995, Moraea rivulicola Goldblatt
& J. C. Manning (Moraea subg. Vieusseuxia ) was
then known from northern Namaqualand in season¬

al watercourses at Spektakel Pass, near Springbok,
and in the Kamiesberg (Goldblatt & Manning,
1995). Plants were distinguished from the nearly
sympatric M. unguiculata by the larger flowers with
outer tepals 20—24 mm long with claws 7-10 mm
long and limbs 14 X 10 mm, and the inner tepals
13—15 mm long. The filaments are ca. 8 mm long
with the column ca. 5.5 mm long, and the anthers
are 4—5 mm long.

Plants from the Richtersveld near Kosies accord
with the description ami represent a range exten¬
sion to the north. A population from the rocky hills
west of Bitterfontein and south of Kotzesrus col¬
lected in 2001, however, differs in their substan¬
tially larger flowers with bright yellow-green tepals
and strong foetid odor, a feature not reported in
other populations. These plants have outer tepals
ca. 28 mm long with the claw up to 14 mm and the
limb 14 X ca. 10 mm, while the inner tepals are
ca. 19 mm long and not trilobed hut have an ovate-
attenuate limb. Other organs are also larger, with
the filament column ca. 10 mm long and free for
an additional 2.5 mm, while the anthers are 5.5-6
mm long and the style crests about 5 mm long. The
inner rhipidial spathes are 6—9 cm long, contrasting
with 5.5— 6.7 cm long in other populations. Plants
ol the Kotzesrus population share with more typical
M. rivulicola the smooth tepal claws and elongate
ovary, 11—14 mm long. The habitat was not, as in
other populations, in a watercourse, but along a dry
gully, where water presumably accumulates in pe¬
riods of precipitation. While the taxonomic status
of this population remains to be determined, there
is no doubt that it represents a significant variant
and thus a distinctive race, perhaps deserving of
subspecific recognition.

Additional specimens. SOUTH AFRICA. Northern
Cape: 29.17 (Springbok) Richtersveld, base of Kosies
(BA), 26 Sep. 1987, Williamson 3751 (NBG); 30.17 (Hon-
deklipbaai) W ol Bitterfontein, 19 km S of Kotzesrus on
the road to Landplaas, edge of granite escarpment (DD),
16 Sep. 2001, Goldblatt & Porter 11892 (K, MO, NBG,
PRE).

Acknowledgments. We thank Guy Brooke-Sum-
ner for drawing our attention to the presence of wild
flowers on the burn on the farm Vredehoek and
allowing us to work there, Roy Gereau for checking
the Latin descriptions, and Ingrid Nanni and Len-
don Porter for assistance and companionship in the
field. Hopliine beetles were identified by Holger
Dombrow, Worms, Germany, and beetle vouchers
are housed at the University of Kansas Museum,
Lawrence, Kansas. Collecting permits were provid¬
ed by the Nature Conservation authorities of West¬
ern Cape Province, South Africa. Support for our

Volume 12, Number 3
2002

Goldblatt & Manning
Moraea from Southern Africa

359

study by grants 6704-00 and 7103-01 from the Na¬
tional Geographic Society is gratefully recognized.

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Strelitzia 7. National Botanical Institute, Cape Town.

- . P Bernhardt & J. C. Manning. 1998. Pollination

of petaloid geophytes by monkey beetles (Scarahaeidae:
Rutelinae: Hopliini) in southern Africa. Ann. Missouri
Bot. Card. 85: 215—230.

- , J. C. Manning & P. Bernhardt. 2000. Adaptive

radiation of pollination systems in Sparaxis Ker Gawler
(Iridaceae: lxioideae). Adansonia ser. 3, 22: 57—70.

Steiner, K. E. 1998. Beetle pollination of peacock moraeas
in South Africa. PI. Syst. Evol. 209: 47-65.

Megacorax gracielanus (Onagraceae), a New Genus and Species

from Durango, Mexico

M. Socorro Gonzalez Elizondo, I. Lorena Lopez Enriquez
CIIDIR— Institute* Politecnico Nacional, Apdo. Postal 738, Durango, Dgo., 34000, Mexico.

sgouzalez@omanet.com.mx

Warren L. Wagner

United States National Herbarium, Department of Systematic Biology, Botany, MKC-166,
P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A.

wagner.warren@nmnh.si.edu

Abstract. Megacorax gracielanus, a distinctive
new genus and species from the Sierra de Coneto
in central Durango, Mexico, is described and illus¬
trated. Its distinctive characters include: leaves lin¬
ear, petals presented to one side of the flower, and
capsule wall thin and distended by the seeds. It
appears to be most closely related to Lopezia Ca-
vanilles based on possession of 4-merous flowers,
deciduous sepals, a capitate stigma, stipules, and
a tendency toward the structural zygomorphy in the
flowers found in Lopezia. In addition, the capsules
of Megacorax gracielanus are very similar in shape
and structure to the capsules of certain species of
Lopezia. Because Megacorax has distinctive mor¬
phological characters and does not have the unique
synapomorphies of Lopezia (2 fertile stamens or 1
fertile stamen and I staminode, and petals un¬
equal), it is here described as a related genus.

Rksumkn. Megacorax gracielanus, un nuevo ge-
nero y especie notoriamente diferente, sc describe
e ilustra a partir de material coleetado en la Sierra
de Coneto, en el centra de Durango, Mexico. Sus
caracteres distintivos incluyen hojas lineares, pe-
talos orientados haeia un lado de la flor, y pared
de la capsula delgada y distendida por las semillas.
El nuevo taxon parece estar mas cercanamente re-
lacionado a Lopezia, genera con el que comparte
los caracteres de (lores tetrameras, sepalos deci-
duos, estigma capitado, y estfpulas. Manifiesta una
tendencia hacia la zigomorfia estructural de las flo¬
ras que se presenta en Lopezia , y sus eapsulas son
muy similares en forma y estructura a las eapsulas
de algunas especies de ese genera. Debido a que
Megacorax tiene caracteres morfologieos distintivos
y no presenta las sinapomorfias unicas de Lopezia
(2 estambres fertiles o 1 estambre fertil y un esta-
minodio, y petalos desiguales o iguales) se describe
aquf como un genera relacionado.

Key words: Megacorax, Mexico, Onagraceae.

fhe family Onagraceae in the order Myrtales
comprises some 650 species in 16 genera. The fam¬
ily is distributed worldwide, but it is most species-
rich in the New World (Raven, 1988). Raven (1964,
1979. 1988) divided the Onagraceae into seven
tribes; only tribe Onagreae, with nine genera, con¬
tains more than two genera. A recent molecular
phylogenetic study (Levin et ah, in press) suggests
that Gongylocarpus should be moved to a unige¬
neric tribe, Gongylocarpeae, bringing the total to
eight tribes for the family. Systematic revisions are
available for virtually the entire family (see refer¬
ences cited in Munz, 1965; Raven, 1979, 1988).
Surveys are essentially complete for many aspects
ol the biology of the family, including chromosome
numbers and morphology (e.g.. Raven & Tai, 1979;
Raven & Gregory, 1972; lloch et ah, 1993), floral,
leaf, and wood anatomy (e.g.. Eyde, 1981; Carl-
quist, 1982), as well as morphology, embryology,
palynology (e.g., Skvarla et ah. 1978; Praglowski et
ah, 1987), reproductive biology including pollina¬
tion systems and extent/mechanism of self-incom¬
patibility (summarized in Raven, 1979), chemcsys-
tematics (e.g., Averett et ah, 1988), and, most
recently, molecular systematics (e.g.. Levin et ah,
in press), providing a wealth ol characters of evo¬
lutionary interest (summarized in Raven. 1988:
lloch et ah. 1993; Conti et ah, 1993).

Because of intense scrutiny of the family for
many decades, it was unexpected to discover an
unknown Onagraceae subshrub during recent Ho¬
listic research in the Sierra de Coneto in Durango,
Mexico. The morphological characters of this plant
show that it does not fit in any of the 16 currently
recognized genera or, with current understanding,
readily in any ol the eight tribes. We here describe

Novon 12: 360-365. 2002.

Volume 12, Number 3
2002

Gonzalez et al.
Megacorax from Mexico

361

it as a new genus and species, but do not assign it
to a tribe pending further studies.

\I< •gacorax S. Gonzalez & W. L. Wagner, gen. nov.

TYPE: Megacorax gracielanus S. Gonzalez &

W. L. Wagner, sp. nov. Figures 1, 2.

Genus novum, Lopeziae affine a qua habito suffruticoso,
foliis alternatis subfasciculatis, floribus tri-tetrameris uter-
que chasmogamis vel cleistogamis, tubo floris brevi nec-
tario apicali, petalis unilateral ibus, parietibus tenuibus
capsulae differt.

Weakly woody shrubs; stems profusely branched;
bark thin, gray brown, exfoliating on older stems;
younger stems red. Stipules present, in pairs, su¬
bulate, persistent. Leaves cauline, alternate, those
on short lateral branches with arrested internodes
crowded and appearing fasciculate, linear to linear-
lanceolate. Flowers numerous, in the axils of the
upper leaves, on long pedicels, diurnal, perfect, but
with both chasmogamous and cleistogamous ones
present on same branch. Floral tube minute with
enlarged nectary disk at mouth. Sepals (3)4, regular
and equal, valvate in aestivation, reflexed in anthe-
sis. Petals (3)4, contorted in aestivation, white, en¬
tire, appearing radially arranged in dry specimens,
but displaced to one side of the flower in anthesis.
Stamens (6)8 in two unequal series, all fertile, the
antisepalous ones longer; anthers versatile, pale
yellow. Style filiform, exceeding or slightly shorter
than the stamens; stigma entire, capitate. Ovary
pluriovulate, turbinate or cylindric. Fruit a loculi-
cidal, thin-walled capsule, locales 3 or 4, ellipsoid-
cylindrical to oblong-cylindrical, dehiscing 1/8 to
nearly 1/2 the length, the dehisced valves spread¬
ing, locule partitions thin, persisting to maturity.
Seeds small, in I irregular row in each locule, dis¬
tending the capsule walls.

Morphologically, Megacorax does not fit into any
of the other genera of the Onagraeeae. It is unique
in its combination of the following characters: pres¬
ence of stipules, flowers 3- or 4-merous, presence
of chasmogamous and cleistogamous flowers, petals
presented to one side of the flower, (6)8 stamens in
two whorls, and thin-walled capsules that dehisce
1/8 to 1/2 their length. It shares some characters
with several other genera but it does not fit into any
of them. Presence of stipules excludes it from the
tribes Epilobieae and Onagreae. The capsule struc¬
ture excludes it from the tribes Hauyeae, Fuch-
sieae, and Cireaeeae. The deciduous sepals and
presence of a floral tube exclude it from tribe Jus-
siaeeae. This leaves only the tribe Lopezieae (only
genus Lopezia Cavanilles), with which it shares the
presence of stipules, 4-merous flowers, a capitate

“dry” stigma, and slight floral zygomorphy that ap¬
proaches the structural zygomorphy found in Lo-
pezia flowers. However, Lopezia is clearly marked
in Onagraeeae by having in 4-merous flowers only
two stamens (one species) or one stamen and a
staminode (Plitmann et al.. 1973). unlike Megaco¬
rax and all other Onagraeeae, which have twice as
many stamens as sepals. The capsules of Megaco¬
rax are similar in shape and structure to those of
some Lopezia , such as L. riesenbachia Plitmann, P.
H. Raven & Breedlove. The stipules of most spe¬
cies of Lopezia are generally caducous, but are per¬
sistent in the shrubby species, similar to the per¬
sistent stipules in Megacorax. Megacorax has a
nectary disk (Fig. 2D) unlike those in Lopezia. but
possibly similar to nectaries in Gongylocarpus
(Gongylocarpeae). Overall, the morphology of Me¬
gacorax places it closest to Lopezia, but it clearly
lacks the synapomorphies of that genus, and de¬
serves separate generic status. In fact, a recent rbc L
and ndhF cpDNA analysis (Levin et al., in press)
confirms the placement of Megacorax in a strongly
supported clade sister to Lopezia.

Etymology. The generic name is based on the
combination of the Greek words mega, large or
great, and korax, raven, to honor Peter H. Raven,
who has studied the Onagraeeae for nearly half a
century and has published numerous papers and
monographs on many aspects ol the family. He also
has inspired many others to contribute to the sci¬
entific knowledge of Onagraeeae, and has been one
of our foremost advocates for conservation and sus¬
tainable use of the environment.

Megacorax gracielanus S. Gonzalez & W. L.
Wagner, sp. nov. TV PE: Mexico. Durango:
Nuevo Ideal. Sierra de Coneto, al N de Dr.
Castillo del Valle, por el camino a Ignacio Za¬
ragoza, 25°5'11"N, 104°57'10"W. 2210 m, 5
Nov. 2001, S. Acevedo, L. Lopez & S. Gonzalez
1352 (holotype, US 3376228; isotypes, ANSM.
CAS, CHAPA, CIIDIR, ENCB, IER, IIZD.
MEXU. MO. TEX. UAMIZ). Figures 1. 2.

Ex speciebus generum affinium, foliis linearibus ad li-
neari-oblanceolata 8—12 X 0.8— 1.3 mm, tubo fforali 0.2—
0.6 mm longo et disco annulari, parietibus capsulae se-
minibus distentis differt.

Xeromorphic, rhizomatous, weakly woody shrub
from a branched, ascending caudex up to 1.4 cm
thick. Stems profusely branched, ascending, (15-)25—
60 cm high, covered with thin gray bark and exfoli¬
ating in longitudinal strips on the lowermost stems,
becoming pale brown to yellow and shredding or flaky
on stems further up, the bark of main stem and pri-

362

Novon

Figure 1. Megacorax gracielanus S. Gonzalez & W. 1,. Wagner. — A. Habit, showing primary stem from near the base.
— B. Leaf. — C. Stem with lower portion of a leaf and associated stipule. — I). Upper portion of stem with both
chasmogamous and cleistogamous flowers on the same branch. — E. Seedling showing cotyledons and first pair of
leaves.

Volume 12, Number 3
2002

Gonzalez et al.
Megacorax from Mexico

363

Figure 2. Megacorax gracielanus. — A. Chasmogamous flower, showing slight zygomorphy of petal presentation. — B.
Petal of chasmogamous flower. — C. Cleistogamous flower, slightly open showing small petals. — D. Flower with petals,
stamens, and style removed, showing fleshy disk. — E. Stamen. — F. Style and capitate stigma. — G. Immature capsule
with pedicel, showing slightly distended capsule walls from seeds. — H. Longitudinal view of capsule, showing arrange¬
ment of seeds. — I. Mature dehised capsule. — J. Adaxial (raphial) surface of seed. — K. Abaxial surface of seed.

364

Novon

mary branches smooth and pale brown, upper branch¬
es terete or angled, yellow to purple or red tinged,
puberulent with [latent or ascending, translucent hairs
0.07-0.2(— 0.3) mm long. Stipules 0.2—1 mm long, lin¬
ear to subulate, puberulent to glabrate, pale yellow,
persistent at least briefly after the leaf falls, leaves
alternate, usually reduced and crowded on short lat¬
eral branches anti thus appearing fasciculate, young
ones thick and somewhat fleshy, 8-12 X 0.8— 1.3 mm,
linear to linear-lanceolate, green to reddish or purple,
minutely papillate, often with minute white dots on
the upper surface, the midrib broad and evident on
the lower surface except toward the apex, narrow and
impressed on the upper surface, otherwise the veins
indistinct, puberulent with translucent hairs ca. 0.1—
0.2 mm long, becoming glabrate, margin entire to
shallowly sinuate-denticulate, whole leaf revolute or
sometimes just the margins revolute, apex obtuse to
subacute, base gradually attenuate, sessile on a thick¬
ened base that persists as a protuberant scar. Flowers
in the axils of the upper leaves, the branches resem¬
bling leafy, many-flowered racemes, both chasmoga-
mous and cleistogamous flowers present on the same
branch, the floral parts variable in size, especially of
cleistogamous ones; pedicels up to 14 mm long, fili¬
form. ascending to erect, purple, puberulent with
white hairs 0.1-0.25 mm long, or sometimes glabrate.
Floral tube inconspicuous, 0.2— 0.6 mm long, enlarged
by annular disk in upper 2/3, puberulent. Buds up to
6.2 mm long, narrowly elliptic, the free tips 0.4— 0.5
mm long, linear. Sepals regular and equal, valvate in
aestivation, reflexed in anthesis, rarely two of the se¬
pals coherent, (1 .2— )3.4 — 6.2 X 0.3— 0.8 mm, oblong-
lanceolate, deep red or reddish purple, sparsely up-
pressed to spreading puberulent on adaxial surface,
especially toward the apex, puberulent on abaxial sur¬
face, abruptly narrowed at the apex. Petals white,
(1.3— )3.8-6.2 X ]. 2-2.6 mm, narrowly obovate, rarely
with inconspicuous purple veins, shorter than or
equaling the sepals, rounded to acute at apex, taper¬
ing to the base, in ehasmogamous flowers appearing
radially arranged in dry specimens, but displaced to
one side of the flower in anthesis. Stamens all fertile.
(6)8, in two unequal series, the antisepalous ones lon¬
ger; filaments pink to purple at least toward the prox¬
imal half, the longer set 1.2-4 mm long, the shorter
set 0.9— 2.4 mm long, with smaller anthers; anthers of
longer stamens 0.6-0. 7 mm long, those of shorter ones
0.4— 0.5 mm long, broadly oblong, cream-colored, ag¬
ing to yellow. Ovary 1.9-4. 1 X 0.9— 1.1 mm, turbinate
or cylindric, gradually tapering to the base, puberu¬
lent. Style filiform, 1.8-8 mm long, pink to purple;
stigma entire, capitate, ca. 0.4— 0.6 mm diam.. whitish,
held above the stamens or surrounded by stamens.
Capsule (3— )4.4— 5.8(-6.7) X 1.3— 1.8 mm, thin-

walled, ellipsoid-cylindrical, oblong-cylindrical, near¬
ly terete or walls slightly distended by seeds, tapering
at base, truncate at summit, pink to purple, puberu¬
lent to glabrescent, 3- or 4-celled anti 3- or 4-valved,
the valves dehiscing 1/8 to nearly 1/2, spreading in
maturity, the thin partitions between the locules per¬
sisting to maturity, with several seeds in 1 irregular
row in each cell. Seeds 1—1.2 X 0.4—0.55 mm, nar¬
rowly obovoid or narrowly oblong, pale brown, glossy,
the abaxial side convex, the raphal surface (adaxial)
side slightly depressed with margins slightly incurved,
with raphe visible as a longitudinal ridge, the testa
appearing nearly smooth, but in fact minutely pitted-
reticulate (> 50 X). Apparently self-compatible judg¬
ing from the presence of cleistogamous (lowers.

Etymology. The specific epithet honors Gracie-
la Calderon Diaz-Barriga, who has written revisions
of about half of the families for the Flora del Valle
de Mexico (the only complete flora published until
now in Mexico, in which she is the editor along
with her husband, J. Rzedowski). She is also writing
many of the revisions for the Flora del Bajfo, and
she has greatly contributed to the mentoring of sev¬
eral generations of students. We are very pleased
to dedicate this species to such an inspiring bota¬
nist and friend.

Distribution and habitat. Megacorax graciela-
nus is known only from two localities in the Sierra
de Coneto, a volcanic mountain located in central
Durango, Mexico, on the western boundaries of the
Chihuahuan Desert. Sierra de Coneto constitutes
the southeastern extreme of the Sierra de Promon-
torio and the eastern limit of an endorrheic (closed)
basin (Laguna de Santiaguillo). The climate in the
area is temperate semi-dry. The predominant sub¬
stratum is formed by ignimbrites (igneous extrusive
rocks of acidic pH). Megacorax gracielanus has
been collected between 24°55'49" to 25°5'11"N
and 104°47'5" to 104°57' 10"W, at 2210 and 2.360
m. It occurs in colonies and is scarce to abundant
in patches on rocky areas with acidic soils. The
dominant vegetation is open xerophytic woodland
of Finns cembroides Zuccarini (pinyon pine) with
Quercus etnoryi Torrey and sclerophyllous shrubs,
mainly Arctostaphylos pungens Kunth and Quercus
depressipes Trelease. The two known populations
are about 18 km apart (air distance). Based on a
survey in June 2002 (SG and LL), it is likely that
Megacorax gracielanus grows in much of the inter¬
vening area as well as in other parts of the Sierra
de Coneto. It has not been possible to survey these
areas because there are no roads. The new survey
showed that at the type locality Megacorax gracie¬
lanus is much more abundant than originally

Volume 12, Number 3
2002

Gonzalez et al.
Megacorax from Mexico

365

thought, and in some areas it is one of the dominant
elements in the lower stratum. It grows along an
open canyon about I km long (at least) and 150 m
wide. In some places there are 4 to 5 plants/m2. In
the other collection site (southernmost) Megacorax
gracielanus is scarce and infrequent with only a few
plants found (< 20). It is interesting that in both
areas it grows not only under shrubs and trees, but
also in open areas along recently open roads, on
fragmented rocks. It is much more abundant and
well developed under the shrubs, but it seems to
adapt well to colonizing disturbed areas, which
raises the question of why this unique plant is not
much more widely distributed. The Sierra de Co-
neto is also the only known locality for Mammilla-
ria theresae Cutak, and perhaps the perceived lo¬
calized nature of their distribution is due to the
lack of thorough collecting in this region of Mexico.

Phenology. Some plants were found flowering
and fruiting after the rainy season, in early Novem¬
ber. but by then most of the plants were finished
fruiting, suggesting that Megacorax gracielanus
flowers primarily earlier in the year. The paratype
collection made at the higher elevation was even
more mature and senescing for the winter. Plants
in the population had indications of foraging by do¬
mestic or wild animals.

Paratypes. MEXICO. Durango: Nuevo Ideal, Sierra
de Coneto, al NE de Once de Marzo por el cam i no a
Coneto, ladera con exposicion al N, 24°55'49"N,
104°47'5"W, 2360 m. matorral esclerdfilo (Arctostaphylos
pungens y Quercus depressipes) con Pinas cembroides,
Quercus emoryi y Arbutus arizonica, suelo somero, 6 Nov.
2001, S. Gonzalez et al. 6532 (CIIDIR, 1 1 . It. MEXU, US).

Acknowledgments. Thanks are given to Satur-
nino Acevedo S., who collected the type, and to
Peter Hoch, and J. Rzedowski for insightful com¬
ments. Jorge A. Tena, Martha Gonzalez Elizondo,
Abraham Torres S., Luis Alfredo Rangel, and Mar¬
cos Pinedo provided support for the Durango au¬
thors. The two excellent plates were prepared by
Alice Tangerini. and the translations of the diag¬
noses to Latin were provided by Dan Nicolson (both
US). S. Gonzalez and L. Lopez acknowledge CO-
FA A and EDI, Institute Politecnico Nacional, for

financial support. We greatly appreciate the com¬
ments by Jorge Crisci. Ching-I Peng, and especially
Peter Hoch, who helped us in many ways with the
manuscript, and the editors of Novon for expediting
the publication of this paper.

I .ilerature Cited

Averett, J. E., S. Huang & W. L. Wagner. 1988. Flavonoid
survey of Oenothera (Onagraceae): Sects. Gauropsis,
Hartmannia, Kneiffia, Paradoxus and Xylopleurum.
Amer. J. But. 75: 476-483.

Carlquist, S. 1982. Wood anatomy of Onagraceae: Further
species; root anatomy; significance of vestured pits and
allied structures in dicotyledons. Ann. Missouri Bot.
Card. 69: 755—769.

Conti, E., A. Fischbach & K. J. Sytsma. 1993. Tribal re¬
lationships in Onagraceae: Implications from rbcL se¬
quence data. Ann. Missouri Bot. Card. 80: 672—685.
Eyde, B. H. 1981. Beproductive structures and evolution
in Ludwigia (Onagraceae). III. Vasculature, nectaries,
conclusions. Ann. Missouri Bot. Card. 68: 470—503.
Hoch, P. C.. J. V. Crisci, II. 4’obe & P. E. Berry. 1993. A
cladistie analysis of the plant family Onagraceae. Syst.
Bot. 18: 31—47.

Levin, B. A., W. L. Wagner. P. Hoch, M. Nepokroeff, .1. C.
Pires, E. A. Zimmer & K. J. Sytsma. In press. Family-
level relationships of Onagraceae based on chloroplast
rbc L and ndh F data. Amer. ,1. Bot.

Munz, P. A. 1965. Onagraceae. North Amer. FI. II. 5: 1—
278.

Plitmann, U.. P. H. Baven & D. E. Breedlove. 1973. The
systematics of Lopezieae (Onagraceae). Ann. Missouri
Bot. Card. 60: 478—563.

Praglowski, J., J. Nowicke, P. Baven, J. Skvarla & W. L.
Wagner. 1987. Onagraceae Juss.: Onagreae B. Baimann
pro parte. World Pollen & Spore Flora 15: 1—55.
Baven, P. H. 1964. The generic subdivision of Onagra¬
ceae, tribe Onagreae. Brittonia 16: 276—288.

- . 1979. A survey of reproductive biology in Ona¬
graceae. New Zealand J. Bot. 17: 575—593.

- . 1988. Onagraceae as a model of plant evolution.

Pf>. 85—107 in L. I). Gottlieb & S. K. Jain (editors).
Plant Evolutionary Biology: A Symposium Honoring G.
Ledyard Stebbins. Chapman and Hall. London.

- & D. P. Gregory. 1972. Observations of meiotic

chromosomes in Gaura (Onagraceae). Brittonia 24: 71-
BO.

- & Wr. Tai. 1979. Observations of chromosomes in

Ludwigia (Onagraceae). Ann. Missouri Bot. Card. 66:
862-879.

Skvarla, J. J., P. H. Baven, W. F. Chissoe & M. Sharp.
1978. An ultrastructrual study of viscin threads in Ona¬
graceae pollen. Pollen & Spores 20: 5—143.

Compsoneura diazii (Myristicaceae), a New Species from the Rio
Cenepa Area of Northwestern Peru

John P. Janovec

Institute of Systematic Botany, The New York Botanical Garden, Bronx,
New York 10458-5126, U.S.A.

ABSTRACT. Compsoneura diazii (Myristicaceae), a
new species from the Rio Cenepa area of Bagua
Province, Amazonas, Peru, is described, illustrated,
and discussed. The new species is segregated from
the Compsoneura capitellata complex and is easily
distinguished by the chartaeeous leaf laminas that
turn brown upon drying; brochidodromous second¬
ary veins; an elongate-tubular perianth; erect, elon¬
gate perianth lobes; erect anthers with acute apices;
a reduced filament column; presence of minute
glandular hairs on the inner perianth surface; and
a ligneous, strongly rugose to suleate pericarp cov¬
ered with ferrugineous bifid trichomes.

Key words: Compsoneura, Magnoliales, Mag-
noliidae, Magnoliophyta, Myristicaceae, Neotrop¬
ics, Peru.

The Myristicaceae are a family of arborescent flow¬
ering plants that have significant ecological and eth-
nobotanical importance worldwide in wet lowland
tropical forests. Comprised of 19 genera and up to
500 species, the Myristicaceae rank among the 10
most diverse and important tree families in this eco¬
system (Gentry, 1982, 1988; Pascal & Pelissier, 1996;
Poulsen et al., 1996; Spichiger et ah. 1996).

1 have initiated studies of Neotropical Myristi¬
caceae with a monograph of Compsoneura (DC.)
Warburg, a genus that ranges in moist tropical for¬
est from southeastern Mexico, through Central
America, and into northwestern South America. I)e
Candolle (1856) provided the first taxonomic treat¬
ment of the Myristicaceae, in which he grouped
about 90 species known at the time into 13 sections
of the single genus Myristica, one of them being
section Compsoneura A. DC. By far, the most com¬
prehensive treatment of the Myristicaceae was pro¬
duced by Warburg (1897). In his Monographic der
Myristicaceen, he assembled 240 species into 1 5
genera, anti among those was Compsoneura, erected
from Myristica sect. Compsoneura of De Candolle.
Warburg recognized a total of 5 species in Comp¬
soneura represented by 8 available specimens, and
divided the genus into 2 sections (Compsoneura and

Novon 12: 366-368. 2002.

Coniostele ) based solely on trends of staminate flo¬
ral morphology.

With increased exploration and specimen avail¬
ability, five more species of Compsoneura were rec¬
ognized and described by Smith and Wodehouse
(1937), and a taxonomic synopsis was presented.
Since 1937, little attention has been paid to Comp¬
soneura aside from descriptions of new species by
Rodrigues (1989), who also produced a taxonomic
treatment of Brazilian members of Virola Aublet
(Myristicaceae) (Rodrigues, 1980).

In preparation of a monograph of Compsoneura,
several new species have been discovered. The
purpose of this paper is to describe, illustrate, and
discuss one of these new species.

Compsoneura diazii Janovec, sp. nov. TYPE: Peru.
Amazonas: Bagua Province, Imaza District, Agu-
aruna Community Putuim, annex Yamayakat,
zone of rolling hills 24 degrees SW of Putuim,
5°37'S, 78°22W, 700-820 m, 23 Sep. 1994, C.
Diaz 7224 (holotype, MO), f igure I .

Consortio speeifico Compsoneurae capillatae affinis sed
lamina tenui vel chartacea, venatione foliari secondaria
conspicue brochidodroma, perianthio tubulari elongato ar-
recto, lobis perianthii lanceolatis arrectis, antheris arrectis
atque apicibus acutis praeditis, columna filamentosa re-
ducta, pilis in superficie perianthii interiore minutis glan-
dulosis, pericarpio fruetus valde rugoso atque trichomati-
bus bifidis ferrugineis obtecto distinguenda.

Trees to 18 m tall, the inner bark exuding profuse
red sap; branchlets terete to subterete, 0.3-0.4 cm
diam., rigid, longitudinally striate, drying dark
brown, densely ferrugineous-tomentose when
young, glabrescent or ferrugineous-tomentose in
leal axils and scattered throughout with age, the
hairs short-stalked to sessile, 2-branched. Leaves
alternate, spiral on primary shoot, distichous on
branches, simple; petioles stout, subterete, strongly
canaliculate, 0.7— 1.5 X 0.1— 0.2 cm, slightly ru¬
gose, drying dark brown, glabrescent to thinly fer¬
rugineous-tomentose, the hairs short-stalked to ses¬
sile, 2-branched; blades elliptic to slightly
elliptic-ovate or slightly to strongly obovate-elliptic,
9.6—16.1 cm long, 3.3— 5.8 cm wide at 1/4 length.

Volume 12, Number 3
2002

Janovec

Compsoneura diazii from Peru

367

F

Figure I. Compsoneura diazii J. Janovec. — A. Habit showing axillary staminate inflorescence. — B. Staminate inflo¬
rescence. — C. Staminate flower. — I). Gynoeciurn. — E. Immature fruit. — F. I-eaf base showing abaxial view and
pistillate inflorescence showing reduced axillary' hahit. (Based on C. Diaz 7749A, 7692A, and 7644A.)

4.2-6.2 cm wide at 1/2 length, 3.7— 5.7 cm wide at
3/4 length, thin to chartaceous, adaxially drying
dark brown, dull to semi-glossy, glabrous, abaxially
drying light or dull brown, weakly resinous-punc-
ticulate, glabrous or glabrescent throughout, the
hairs minute, sessile or short-stalked, 2-branched,
the base acute, the margins conspicuously revolute,
the apex acute to narrowly obtuse to weakly atten¬
uate, caudate to strongly caudate, the acumen 0.7—

1.1 cm long; venation with costa raised and gla¬
brous adaxially, prominently keeled, glabrous to
glabrescent, especially toward base abaxially, the
secondary veins 5—11 per side, arcuate-ascending,
conspicuously brochidodromous, anastomosing
near margins, shallowly impressed adaxially, prom¬
inent abaxially, 0. 7-2.9 cm apart. Staminate inflo¬
rescences axillary, 2-fasciculate, once-paniculate,
4.0— 4.5 X 1.0-1. 4 cm, glabrescent to sparsely or

368

Novon

densely ferrugineous-tomentose, the hairs 2-
branched; rachis alternately branched, the branch-
lets 15—20, 0. 1-0.2 cm long, with a minute bract
at the apex, the flowers arranged in clusters of
about 2—6 per branchlet; pedicels slender, 0. 1 5—
0.30 cm long. Pistillate inflorescences axillary, once-
pinnate, 0.7— 0.8 X 0.8— 0.9 cm, densely to sparsely
ferrugineous-tomentose, the hairs 2-branched; ra¬
chis 4 or 5 alternately branched, the branchlets
0.02—0.03 cm long, the flowers arranged in clusters
of about 1—2 per inflorescence branchlet; pedicels
slender, 0.09 X 0.06 cm. Staminate flower buds
long-ovate to long elliptic-ovate or ovate-lanceolate.
Pistillate flower buds elliptic to ovate-elliptic. Sta¬
minate flowers: perianth elongate-tubular, 0.21-
0.32 X 0.12—0.2 cm, thin-coriaceous to coriaceous,
orange-yellow when fresh, drying brown to dark
brown, thinly pubescent adaxially, the hairs short
glandular, glabrescent to ferrugineous-tomentellous
to tomentose abaxially, the hairs short-stalked to
sessile, 2-branched, the tube 0.9—0.13 cm long, the
lobes 3 or 4, narrowly ovate to lanceolate-ovate to
narrowly deltoid, 0.12-0.21 X 0.1-0.15 cm; an-
droecium 0.10—0.14 cm long, the filament column
0.01-0.02 cm long, 0.02 cm wide at apex. 0.02—
0.03 cm wide at base, the anthers 4 or 5, free,
erect, 0.09—0.12 cm long, the apices strongly acute.
Pistillate flowers: perianth elongate-tubular, 0.35 X
0.13 cm, thick coriaceous, orange-yellow when
fresh, drying brown to dark brown, thinly pubescent
adaxially, the hairs short-glandular, densely to
sparsely ferrugineous-tomentellous to ferrugineous-
tomentose abaxially, the hairs 2-branched, the tube
0.13 cm long, the lobes 3, long-ovate to lanceolate-
deltoid, 0.22 X 0.09 cm; ovary subglobose to ovate,
0.17 cm long, 0.07 cm wide at 1/4 length, 0.08 cm
wide at 1/2 length. 0.03 cm wide at 3/4 length,
drying brown, densely ferrugineous-tomentose; stig¬
ma bilobate, shallowly cleft, 0.02 X 0.03 cm, brown
to dark brown when dry. Immature fruits 2 or 3 per
infruetescence, elliptic to elliptic-ovate, 2. 0-3.3 X
I .<5 — 2. 1 cm, strongly rugose to cerebriform-rugose
or irregularly sulcate, green to green-brown when
fresh, drying light brown to brown, ferrugineous-
tomentose, the hairs simple or 2-branched, the apex
minutely acuminate, the acumen 0.1— 0.2 cm long,
the base rounded. Seeds unknown.

This species has been collected on only four sep¬
arate occasions from wet forest in the vicinity of the
Aguaruna community of Yamayakat, Bagua prov¬
ince, Amazonas, Peru. As all four collections were
made by Camilo Diaz, a Peruvian botanist who has
conducted floristie studies in every corner of Peru,
the name Compsoneura diazii is given in his honor.

Compsoneura diazii is hypothetically a member
of the Compsoneura capitellata species complex
based on the follow ing suite of characteristics: (1)
brown-drying leaves; (2) the presence of ferrugin-
eous bifid trichomes on young stems, petioles,
leaves, inflorescences, and external perianth sur¬
faces; and (3) a ligneous fruit pericarp. However, it
can be unmistakably distinguished from all known
species of this complex by its combination of (1)
chartaceous leaf lamina; (2) conspicuously brochi-
dodromous secondary leaf venation; (3) elongate-
tubular perianth; (4) erect, elongate perianth lobes;
(5) erect anthers with acute apices; (6) a reduced
filament column; (7) presence of minute glandular
hairs on the inner perianth surface; and (8) a lig¬
neous, strongly rugose to irregularly sulcate fruit
pericarp covered with ferrugineous bifid trichomes.

Paratypes. PERU. Amazonas: Bagua Province, Dis¬
trito Imaza, Comunidad Aguaruna Putuim, anexo Yama¬
yakat, zona de colinas alatas 24 degrees SW de Putuim,
5°37'S, 78°22W, 700-820 m, 18 Jan. 1996, C. Diaz et al.
7644A (MO), C. Diaz et al. 7692A (MO), 22 Jan. 1996,
C. Diaz 7749A (MO).

Acknowledgments. I am grateful to Camilo
Diaz, Rodolfo Vasquez, and Henk van der Werff for
initial discussion of this interesting species, to
Amanda K. Neill for her excellent illustration, and
to her and Scott A. Mori for reviewing the manu¬
script and providing helpful comments. I also thank
the herbarium staff of MO for making their speci¬
mens available.

Literature Cited

De Candolle, A. 1856. Myristicaceae. Prod. Syst. Nat.
Veg. 14: 187-208.

Gentry, A. II. 1982. Patterns of neotropical plant species
diversity. Evol. Biol. 15: 1—84.

- . 1988. Tree species richness of upper Amazonian

forests. Proc. Natl. Acad. Sei. U.S.A. 85: 156-159.
Pascal, J. P. & R. Pelissier. 1996. Structure and floristie
composition of a tropical evergreen forest in southwest
India. J. Trap. Ecol. 12: 191-214.

Poulsen, A. I).. 1. C. Nelson, S. Tan & If. Balslev. 1996.
A quantitative inventory of trees in one hectare of mixed
dipterocarp forest in Temburong, Brunei Darussalam.
Pp. 139—150 in D. S. Edwards, W. E. Booth & S. C.
Choy, Tropical Rainforest Research — Current Issues.
Kluwer Academic Publishers, London.

Rodrigues, W. A. 1980. Revisao taxonomica das esp^cies
de Virola Auhlet (Myristicaceae) do Brazil. Acta Amaz.
10: 1-127 (supplement).

- . 1989. Two new neotropical species of Compso¬
neura (Myristicaceae). Brittonia 41: 161.

Smith, A. C. & R. P. Wodehouse. 1937. The American
species of the Myristicaceae. Brittonia 2: 393-510.
Spichiger, R., P. A. Loizeau, C. I.atour & (7 Barriera.
1996. Tree species richness in a southwestern Amazo¬
nian forest. Candollea 51: 559—577.

Warburg. (). 1897. Monographic der Myristicaceen. Nova
Acta Acad. Caes. Leop. -Carol. 68: 1—680.

Th ree New Species of Symplocos (Symplocaceae) from Panama and

Costa Rica

Lawrence M. Kelly

Departamento de Botanica, Institute de Biologia, Universidad Nacional Autonoma de
Mexico, Apartado Postal 70-233, 04510 Mexico D.F., Mexico. lkelly@ibiologia.unam.mx

Frank Almeda

Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco,
California 941 18, U.S.A. falmeda@calacademy.org

ABSTRACT. Three species of Symplocos are newly
described from Panama and Costa Rica. Symplocos
morii Almeda & L. Kelly, known from high-eleva¬
tion western Panama, is distinguished by coria¬
ceous leaves with entire and revolute margins,
short, glabrous styles, and white fruits. Symplocos
nanifiora L. Kelly & Almeda is a low-elevation
Costa Rican species with serrate leaf margins,
small flowers, densely villous styles, and small
fruits. Symplocos elliptica L. Kelly & Almeda,
known from Cerro Horn i to and Cerro Sapo, Pana¬
ma, is distinguished by broadly elliptic leaves with
crenate-denticulate margins and solitary flowers.
All three species are likely members of Symplocos
sect. Symplocastrum Brand based on their basally
connate stamens that are adnate to the petals.

Key words: Costa Rica, Panama, Symploca¬
ceae, Symplocos.

Symplocos Jacquin consists of 250—300 species
distributed through tropical and subtropical regions
of America, southern and eastern Asia, Australia,
and the East Indies. Selected species are also found
in temperate latitudes in the Northern Hemisphere.
Not surprisingly, many species of Symplocos have
been described in the century since Brand’s (1901)
useful but now outdated monograph of the genus.
In the course of studies of Symplocos for the Flora
Mesoamericana, three new species were discovered
from Costa Rica and Panama. Two of these species
were collected from windswept, dwarf cloud-forest
vegetation near Cerro Hornito (Cerro Pata de Ma¬
cho), Panama.

1, Symplocos elliptica E. Kelly & Almeda, sp.
nov. T\ PE: Panama. Chiriqui: Cerro Pata de
Macho [Cerro Hornito], windswept ridge, 2100
m, 17 Jan. 1986, G. de Nevers & G. McPherson
6842 (holotype, CAS: isotypes, BM, MEXU,
MO. PM A). Figure I.

Arbor 2— 3(— 20) m alta, ramis junioribus sparsissime
sericeis. Laminae foliorum 5.5— 8.5 cm longae, 3—4.3 cm
latae, coriaceae, ellipticae, margine denticulato-crenatae,
basi obtusae, apice acuminatae, adultae supra glabrae,
subtus glabratae vel sparsissime strigillosae; petiolus 10—
12 mm longus. Flores pedicellati in foliorum axillis soli-
tarii, bracteolis ovatis vel deltoideis, 1—2 mm longis, 1—
1.5 mm latis, pedicello 7—10 mm longo; calycis tubus gla-
ber, lobis semicircularibus, t-i I i at is. 1-1.5 mm longis, 1 —
2 mm latis; corolla rosea, campanulata, 7—9 mm longa,
glabra, lobis oblongis vel obovatis, 3-4 mm latis; stamina
multiseriata; stylus pilosus ad basim, 4—5 mm longus.
Fructus glaber, ellipsoideus, 1 cm longus, 5 mm latus, 4-
loeularis.

Small trees 2— 3(— 20) m tall; vegetative buds and
juvenile branches sparsely sericeous, the hairs
0.25—0.5 mm long, translucent brown. Petioles
sparsely strigose abaxially. 10-12 mm long and ca.
1 mm wide. Leaves coriaceous, broadly elliptic,
5.5— 8.5 X 3—4.3 cm, margins crenate-denticulate
with minute, black, deciduous teeth, apex acumi¬
nate, base obtuse, abaxially sparsely strigose to gla-
brescent on the elevated network of veins, adaxially
glabrous with the midvein impressed. Flowers ax¬
illary, solitary; pedicels terete, 7—10 mm long,
sparsely sericeous to glabrate; bracteoles 3 or 4,
closely subtending the base of the flower, sessile,
caducous, ovate to triangular-ovate, 1—2 X 1—1.5
mm, apex acute to obtuse, abaxially minutely se¬
riceous to glabrous, adaxially glabrous, margins cil-
iate. Calyx 5-lobed, the lobes subrotund, 1—1.5 nun
long, 1—2 mm wide at the base, glabrous, margins
ciliate and also commonly bearing brown, vesicular
glands. Corolla sympetalous, glabrous, campanu-
late, 7—9 mm long, 5-lobed, pink; lobes connate
basally for 2-3 mm and adnate to the filaments for
3-4 mm at the base, oblong to obovate, apical ly
rounded, 3-4 mm wide, entire to minutely ciliate
distally. Stamens 4-seriate; filaments connate ba¬
sally for 5—6 mm, the free portions of outermost
filaments linear-oblong, 2.5— 3.5 X 0.75—0.9 mm.
Ovary inferior, villous at the summit, glabrous on

Novon 12: 369-374. 2002.

370

Novon

Figure 1. Symplocos elliptica L. Kelly & Almeda. — a. Branch with flowers. — b. Leaf, abaxial surface. — c. Flower.
— (1. Style. — e. Stamens and petals. Based on isotype de Nevers & McPherson 6842 (MEXU).

Volume 12, Number 3
2002

Kelly & Almeda

New Species of Symplocos

371

the sides; style straight, pilose basally, 4-5 mm
long; stigma conspicuously but irregularly lobed.
Fruits drupaceous, glabrous, ellipsoid, ca. 1 X 0.5
cm, 4-celled in cross section with a rounded bony
endocarp.

Known from Cerro Hornito and Cerro Sapo, Pan¬
ama. The growth form of this species appears to be
highly variable in different habitats. The collections
from Cerro Hornito indicate that it reaches 2—3 m
as a component of windswept dwarf vegetation. The
collection from the summit of Cerro Sapo was re¬
portedly a large tree of 20 m.

The combination of broadly ell i ptic leaves with
crenate-denticulate margins and an acuminate
apex, solitary flowers, and basally pilose styles dis¬
tinguishes Symplocos elliptica from all other Central
American species. The species is likely a member
of Symplocos sect. Symplocastrum Brand based on
its basally connate stamens that are adnate to the
corolla tube. Based on its solitary, 3- or 4-bracte-
olate flowers this species is somewhat similar to
Symplocos tribracteolata Almeda, a recently de¬
scribed Costa Rican species; however, S. elliptica
differs by the size and shape of its leaves and the
basally pubescent style. Leaves of S. tribracteolata
are elliptic to elliptic-lanceolate and 1.1— 2.1 cm
wide, whereas those of S. elliptica are broadly el¬
liptic and 3-4.3 cm wide. Texture anti venation of
the leaves of S. elliptica suggest that this species
may be related to S. limoncillo Humboldt & Bon-
pland, which occurs from Mexico (Veracruz and
Chiapas) to Panama. Both S. limoncillo and S. el¬
liptica have coriaceous leaves with an elaborate
network of protruding higher-order veins on the un¬
dersurface. Nevertheless the solitary flowers and
small fruits of S. elliptica serve to clearly separate
this species from S. limoncillo (which has racemose
inflorescences and fruits 1.5— 1.8 cm long).

Paratypes. PANAMA. Boras del Toro: NW ridge of
Cerro Pata de Macho [Cerro Hornito] from summit to Fin-
ca Serrano, 1200-2100 m, 27 May 1081. K. Sytsma et al.
4958 (CAS, MO). Darien: NE slope of Summit Cerro
Sapo, approach from Garachind, 3300 ft.. 8 Mav 1079, H.
Hammel 7269 (CAS. MO).

2. Symplocos morii Almeda & L. Kelly, sp. nov.
TYPE: Panama. Chiriquf: Cerro Hornito (Cerro
Pata de Macho), S rim of the Edwin Fabrega
Dam and Reserve watershed, ridge trail lead¬
ing to the summit, 1800—1950 m, 17 Jan.
1989, F Almeda et al. 6267 (holotype, CAS;
isotype, PM A). Figure 2.

Frutex vel arbor parva 2—5 m alta, ramis vegetativis
glabris. Laminae foliorum 5.5— 8( — 9) cm longae, 3—5 cm
latae, coriaceae, late ellipticae, margine integrae, basi ob-

tusae vel rotundatae, apice rotundatae vel breviter obtu-
sato-acuminatae, adultae supra subtusque glabrae, subtus
glabris; petiolus 2—7 mm longus. Inflorescentia axillaris,
racemosa, 5—7 mm longa, glabra, bracteis bracteolisque
ovatis vel oblongis, 1—3 mm longis, 1 .5—3 mm latis; ca-
lycis tubus glaber, lobis rotundis, ciliatis, 1.3— 1.7 mm lon¬
gis, 1 .8—2.2 mm latis; corolla alba, campanulata, 5—6 mm
longa, glabra, lobis oblongis, 2. 8-3. 2 mm latis; stamina
fere libera ad basim; stylus glaber, 2 mm longus. Fructus
glaber, ovoideus vel ellipsoideus, 6-8 mm longus, 4—5 mm
latus, 3-locularis.

Shrubs or small trees 2-5 m tall. Vegetative
buds, juvenile branches, and inflorescences gla¬
brous. Petioles glabrous, 2-7 mm long and 2—2.5
mm wide. Leaves coriaceous, entire and revolute
along the margins, broadly elliptic, 5.5— 8(-9) X 3—
5 cm, apex rounded to very shortly blunt-acumi¬
nate, base obtuse to rounded (rarely acute), gla¬
brous. Inflorescences short, axillary, bracteate ra¬
cemes 5—7 mm long with 3 to 5 subsessile to
short-pedicellate (pedicels 0—2 mm long) flowers
exceeded by subtending leaves; bracts and brac-
teoles sessile, persistent, ovate to oblong, 1—3 X
1.5—3 mm, glabrous, margins ciliate. Calyx 5-lobed,
the lobes very broadly ovate, 1.3— 1.7 mm long, 1.8—
2.2 mm wide at the base, glabrous, margins ciliate.
Corolla sympetalous, glabrous, ± campanulate at
anthesis, 5-6 mm long, 5-lobed, white; lobes con¬
nate basally for 1.5— 1.8 mm and adnate to the fil¬
aments for 2—2.2 mm, oblong, apically rounded and
involute distally, 2. 8-3. 2 mm wide, margins entire
to minutely erose. Stamens ± triseriate; filaments
connate basally for 2—3 mm, the free portions of
outermost filaments linear-oblong, 2-2.5 X 0.5—
0.75 mm; anthers minutely papillose, bilocular. ±
globose, 0.4— 0.6 mm long and wide. Ovary inferior,
glabrous on the sides, the apex villous but glabres-
cent in fruit; style straight, ca. 2 mm long, glabrous;
stigma conspicuously but irregularly lobed. Fruits
drupaceous, white, glabrous, ovoid to ellipsoid, 6—
8 X 4—5 mm, 3-celled in cross section with an
irregularly undulate bony endocarp.

This species is known from Bocas del Toro, Chi¬
riquf, and Veraguas Provinces, Panama, where it
occurs in exposed, windswept sites at high eleva¬
tion with dwarf cloud-forest vegetation that forms a
3—5 m canopy. Common associates, noted in the
Mori and Bolten collection, include Quercus spp.,
Podocarpus, and Drimys.

Symplocos morii is readily distinguished from all
other Central American species of Symplocos by its
broadly elliptic, entire- and revolute-margined
leaves that are 3—5 cm wide, glabrous styles ca. 2
mm long, and fruits that are white when mature.
The species appears to be a member of Symplocos
sect. Symplocastrum and is somewhat similar to

372

Novon

Figure 2. Symplocos morii Almeda & L. Kelly. — a. Branch with flowers. — b. Feaf, abaxial surface. — c. Leaf, adaxial
surface. — d. Flower. — e. Fruit. — f. Fruit, cross section. Based on the holotype, Almeda et al. 6267 (CAS).

Symplocos costaricana Hemsley, another predomi¬
nantly glabrous, entire-margined species t hat oc¬
curs at low elevations throughout Central America.
Symplocos costaricana can nevertheless be distin¬
guished from S. morii based on its flat (non-revo-

lute), narrower leaves (2—3.5 cm wide), longer
styles (7—8 mm long), and blue-green fruits.

It is our pleasure to name this species in honor
of Scott Mori (b. 1941), an expert in the Neotropical
flora at The New York Botanical Garden, in rec-

Volume 12, Number 3
2002

Kelly & Almeda

New Species of Symplocos

373

Figure 3. Symplocos naniflora L. Kelly & Almeda. — a. Branch with flowers. — b. Leaf, abaxial surface. — c. Leaf
pubescence, abaxial surface. — d. f lower. — e. Bracteoles. — f. Style. — g. Stamens and petals. — h. Fruit. — i. Fruit,
cross section. Based on isotype Herrera 4677 (MFXL).

374

Novon

ognition of his outstanding contributions to botan¬
ical research.

Paratypes. PANAMA. Boras del Toro: on Chiriquf
trail. Elfin Forest at Divide, 20 Apr. 1968. J. H. Kirkbride
& J. A. Duke 992 (MO. NY). Ghiriqiu: path from Linares
Farm ca. 1400 m to top of Cerro Hornito at 1750 m, 27
Dec. 1977. J. P. Folsom el al. 7240 (CAS 2 sheets. MO);
Cerro Hornito, 40 km NW of Gualaca, 2238 m, 27 July
1975, S. Mori & A. Pollen 7502 (CAS. MEXU. MO). Ver-
aguas: summit of Cerro Tute above Escuela Agricola Alto
de Piedra, just W of Santa F£, 1350—1410 m, 8°32'N.
81°07'W, 5 June 1982, S. Knapp & R. Dressier 5396 (CAS.
MEXU, MO).

3. Syrnplocos iiuuifloru L. Kelly & Almeda, sp.
nov. TYPE: Costa Rica. Puntarenas: Canton
del Golfito Jimenez, between Quebrada Pate-
mazo and the headwaters of Rfo Madrigal, 650
in, 30 INov. 1990, G. Herrera 4677 (holotype,
CAS; isotypes, CR, F, MEXU, MO, NY). Fig¬
ure 3.

Arbor 13—20 m alta, ratnis junioribus pilosis. Laminae
foliorum 6.5 — 1 1( — 12.5) cm longae, 2.5— 3. 5(— 4.4) cm latae,
membranaceae, oblanceolatae, margine serrulatae, basi
acutae, apice acuminatae, adultae supra glabrae, subtus
pilosae; petiolus 4—8 mm longus. lnflorescentia axillaris,
racemosa, 5—10 mm longa, sericea, bracteis bracteolisque
ovatis vel subrotundis, 1—1.5 mm longis, 1—1.5 mm latis;
calycis tubus sericeus vel glabratus, lobis triangularibus
vel semicircularibus, ciliatis, 1-1.5 mm longis, 1-1.5 mm
latis; corolla alba, campanulata, 4—6 mm longa, glabra,
lobis oblongis, 2-2.5 mm latis; stamina multiseriata; sty¬
lus pilosus, 4—6 mm longus. Fruetus glaber, ellipsoideus.
1—1.2 cm longus, 4—5 mm latus, 4-locularis.

Trees 13—20 m tall; vegetative buds and juvenile
branches moderately to sparsely pilose, the hairs 1—
1.5 mm long, soft, brown, spreading to erect. Petioles
pilose to glabrate, 4—8 mm long and ca. 1 mm wide.
Leaves membranaceous, serrulate, oblanceolate
rarelv varying to elliptic, 6.5—1 1 ( — 12.5) X 2.5— 3. 5(—
4.4) cm, apex acuminate, base acute (often narrowly
so), abaxially pilose, densely so on the protruding
midvein, adaxially glabrous. Inflorescences short, ax¬
illary, bracteate racemes 5—10 mm long with 3 to 6
sessile to short-pedicellate (pedicels 0—3 mm long)
flowers; rachis sparsely to densely sericeous; bracts
and bracteoles sessile, persistent, ovate to subrotund,
1—1.5 X 1—1.5 mm, abaxially glabrate to densely
sericeous, adaxially glabrous, margins ciliate. Calyx
5-lobed, the lobes triangular-ovate to subrotund, 1—
1.5 mm long, 1—1.5 mm wide at the base, abaxially
sparsely sericeous to glabrate, adaxially glabrous,
margins ciliate. Corolla sympetalous, glabrous, cylin-
dric-campanulate at anthesis, 4—6 mm long, 5- or 6-

lobed, white; lobes connate basally for 1.5— 2.5 mm
and adnate to the filaments for 2-3 mm at the base,
oblong, apically rounded, 2—2.5 mm wide, entire to
minutely ciliate distally. Stamens 3- or 4-seriate; fil¬
aments connate basally for 3—5 mm, the free portions
of outermost filaments linear-oblong, 2—3 X 0.25—
0.5 mm. Ovary densely villous at the summit, gla¬
brous on the sides; style straight, 4—6 mm long, mid¬
dle and lower part densely villous; stigma
inconspicuously and irregularly lobed. Fruits dru¬
paceous, glabrous, ellipsoid, 1— 1.2 X 0.4— 0.5 cm,
4-celled in cross section with a rounded to very
slightly undulate bony endocarp.

This species is distinguished based on the fol¬
lowing combination of characters: membranaceous
leaves, small flowers with densely villous styles,
anil small fruits. It is likely a member of Syrnplocos
sect. Symplocastrum and may be closely related to
S. bicolor L. 0. Williams, another low-elevation,
small-fruited species of the section. Syrnplocos bi¬
color is distributed from Chiapas to Nicaragua and
is distinguished from S. naniflora based on its gla¬
brous leaves and styles, larger flowers (corolla 6—7
mm long), and 5-lobed corolla.

Paratypes. COSTA RICA. Alajuela: Canton de Los
Chiles, Cano Negro, Las Cabas, Cuenca del Rfo Frio, 35
in, 1 1 Oct. 1996. .4. Rodriguez 1653 (F); Canton de San
Carlos, llanura de San Carlos, finca aserradero San Jorge,
l(M) m. 21 Jan. 1996. I\. Zamora & A. Zeleddn 2368
(MEXU, MO). Puntarenas: Reserva Forestal Golfo Dul-
ee, entre Rancho Ouemado y Drake, 300 m, 29 Mar. 1991.
R. Aguilar el al. 105 (MEXU, MO); Canton de Osa, R.F.
Golfo Dulce, Peninsula de Osa, Los Mogos, 200 m, 5 July
1994, R. Aguilar 3462 (MO); Canton del Golfito, Valle de
Coto Colorado, camino a las torres del ICE, cabecera del
Rfo Sorpresa, 5 km al este. 300—400 in. 1 Feb. 1992, N.
Zamora et al. 1748 (F. MO); Canton del Golfito, Valle de
Coto Colorado, camino a las torres del ICE, cabecera del
Rfo Sorpresa. 5 kin al este. 300 — MX) m, 22 Jan. 1993, A.
Zamora el al. 1924 (F. MO. NY). San Jose: basin of El
General, 675—900 m, 12 Dec. 1977, .4. F. Skutch 5523 (F
2 sheets).

Acknowledgments. We thank Albino Luna
(Figs. I & 3) and Sheva Myers (Fig. 2) lor preparing
the line drawings, Fernando Chiang for editing the
Latin descriptions, Bertil Stahl for careful revision
of the manuscript, and the curators and staffs of the
herbaria cited herein for the opportunity to examine
collections under their care.

I .iterature Cited

Brand, A. 1901. Symploeaceae. Das Pflanzenreieh IV. 242

(Heft 6): 1-100.

A New Variety of Miconia ceramicarpa (Melastomataceae) from

French Guiana

Claire V. Martin

Museum National d’Histoire Naturelle, Laboratoire de Phanerogamie, 16 rue Buffon, 75005

Paris, France, cvmartin@mnhn.fr

Abstract. Miconia ceramicarpa (Melastomata¬
ceae) is commonly found in the tropical forest of
French Guiana. A study of the taxonomy, vegetative
architecture, and inflorescence structures showed
the existence ot an undescribed variety, Miconia
ceramicarpa var. crozieriae.

Ri.SUmC. Miconia ceramicarpa est abondamment
rencontree en foret tropicale de Guyane frangaise.
One etude de la taxonomie, de l’architecture ve¬
getative et des structures inflorescent idles a montre
la presence d’une nouvelle variete, Miconia ce¬
ramicarpa var. crozieriae.

Key words: French Guiana. Melastomataceae,
Miconia.

In the series Flora of the Guianas, the treatment
of Melastomataceae subfamily Melastomoideae was
prepared by Wurdack (in Wurdack et ah, 1993).
The author recognized 39 genera and 283 species,
including 96 species of Miconia for the Guianas
(Guyana, Suriname, and French Guiana) and 72 for
French Guiana alone.

According to Wurdack (in Wurdack et ah. 1993),
Miconia ceramicarpa (DC.) Cogniaux is a subshrub
0.4 to 1 m tall, occurring in Venezuela, the Guian¬
as, and the lower Amazon basin in Brazil . Wurdack
recognized two varieties: Miconia ceramicarpa var.
ceramicarpa (= var. violacea (DC.) Cogniaux), pre¬
sent throughout the range of the species, and va¬
riety candolleana Cogniaux (= var. navioensis Wur-
dack), known from French Guiana and Brazil
(Amapa, Para).

While conducting studies in various forest hab¬
itats in French Guiana, we were especially inter¬
ested in the morphological variation of this species,
which occupies an important place in the under¬
story, particularly in sunny locations such as log¬
ging roads and forest gaps.

The vegetative architecture of M. ceramicarpa,
following the concept of Halle and Oldeman (1970)
and Halle et al. (1978), was examined by Dauchez
(1977) and Cremers (1986). These studies showed
considerable morphological diversity in this species
with each variety showing a specific model.

Dauchez (1977) and Cremers (1986) described
the architecture of M. ceramicarpa var. ceramicar¬
pa. The structure is modular and three-dimension¬
al, conforming to Leeuwenberg’s model (Halle &
Oldeman, 1970): the modules are orthotropic, with
a terminal inflorescence and apical relays (Fig. la—
c).

The same authors (Dauchez, 1977; Cremers,
1986) described the architecture of M. ceramicarpa
var. candolleana as mixed, monopodial, and mod¬
ular, conforming to Fagerlind’s model (Halle & Old¬
eman, 1970): the primary axis (trunk) is orthotro¬
pic, monopodial with rhythmic growth, and
produces lateral branches that are plagiotropic and
modular by apposition. The modules have terminal
inflorescences (Fig. Id-h). It was already noted by
Dauchez (1977) and Cremers (1986) that some oth¬
er plants of Miconia ceramicarpa have a miniatur¬
ized architecture with a very contracted, short-lived
trunk developing only one pair of plagiotropic
branches (Fig. li. j). These plants do not conform
to either of the currently recognized varieties and
are therefore described here as new.

The new variety conforms with Fagerlind’s mod¬
el, as does variety candolleana, but with the pri¬
mary axis completely reduced producing a single
plateau at ground level (Fig. li, j). The pubescence
is similar to that of Miconia ceramicarpa var. ce¬
ramicarpa, as is the general aspect of the leaves,
although they are smaller.

The three varieties grow in similar ecological
conditions and were often found to co-occur at the
same sites in open areas in the forest (clearings,
trails, etc.). They have the characteristics of pioneer
plants.

Miconia ceramicarpa var. crozieriae Cremers &
C. V. Martin, var. nov. TYPE: French Guiana.
Montsinery, Piste de Saut Leodate, PK 25, 50
m, 19 Mar. 2000, C. V. Martin 283 (holotype,
Ik isotypes, B. BBS, BM. BR, BRG. CAY, G.
HAMAB, INPA. K. MG. MO. NY, C. US). Fig¬
ures 2, 3.

A Miconia ceramicarpa var. ceramicarpa et Miconia ce-

Novon 12: 375-378. 2002.

376

Novon

F

G

H

J

l /)

fe iww i

Figure 1. Vegetative architecture. Miconia ceramicarpa (DC.) Cogniaux. — A— C. Variety ceramicarpa, Leeuwenberg’s
model. — D-H. Variety candolleana Cogniaux, Fagerliud’s model. — 1, J. Variety crozierae Cremers & C. V. Martin,
Fagerlind’s model with primary axis very contracted or not developed.

ramicarpa var. candolleana primario axe, reducto habitu
differt. A typo apposita sympodialibus ramulis differt. A
var. candolleana orthotropa reiteratione carente in secon¬
dare stratis axibus, glabro abaxiali limbo praeter nervos,
sparse tenui indumento, praecipue differt.

Creeping herb, (10 — ) 1 5 — 20(-30) cm tall; primary
axis not seen, but probably short and ephemeral:
plagiotropic axis prostrate on the ground, rooting at
nodes, sympodial by apposition; stems, veins, inflo¬
rescences, and hypanthium covered with sparse,
line, appressed pubescence. Leaves opposite, de¬
cussate, with dorsiventral leal dimorphism; larger
leaves with petiole 1—2.5 cm long, lamina 7.5—10.5
X 2.5— 4.5 cm, oval to elliptic, apex acuminate,
base acute to obtuse, margin finely dentate and fil¬
iate, pubescence between and on the veins adaxi-
ally, only on the veins abaxially, main veins 3 with
branching point 0.2— 1.8 cm from base or less often
5 with branching 0.1— 0.5 cm from base; smaller
leaf similar but with petiole 0.2-1 cm long, lamina
3.5—7 X 1.5— 2.5 cm; glandular hairs 30—50 /am
visible only with the SEM on the abaxial and ad-
axial leaf surface. Inflorescence a raceme (0.8— )2—
4.5(— 5.5) cm long; bracts persistent. Flowers 5-
merous, sessile; hypanthium 2—3 mm long; calyx

oblate, lobes not projecting, 0.2— 0.3 mm; corolla
white to pink, petals elliptic to obovate, 1-2.5 mm
long, glabrous; stamens isomorphic, glabrous; an¬
thers 2 mm long, dehiscing by a pore; style slightly
pilose at base; stigma slightly enlarged at apex.
Fruits red, becoming blackish blue and fleshy at
maturity, 10 X 6 mm; seeds 0.4 mm long.

In order to facilitate identification of material of
Miconia ceramicarpa , the following key has been
prepared.

Key to the Varieties ok Miconia ceramicarpa

1. Prostrate herbs, 0.15—0.2 m tall; stems plagiotro¬
pic; pubescence sparse, short, appressed .

. var. crozieriae

1 '. Shrubs or subshrubs, 0.5—2 m tall; stems ortho¬
tropic or plagiotropic; pubescence dense or

sparse, appressed or erect . 2

2. Shrubs 1-2 m tall; all stems orthotropic,
erect; pubescence sparse, short, appressed

. var. ceramicarpa

2'. Subshrubs 0.5—1 m tall; main stem orthotro¬
pic, lateral shoots plagiotropic, sympodial by
apposition, decumbent, rooting at nodes and
reiterating; pubescence dense, appressed or
erect . var. candolleana

Volume 12, Number 3
2002

Martin 377

Miconia ceramicarpa from French Guiana

Figure 2. Miconia ceramicarpa (DC.) Cogniaux var. crozierae Cremers & C. V. Martin. — A. Habit. — B. Detail of leaf
(adaxial surface). — C. Detail of leaf (abaxial surface). — I). Flower. — E. Flower and hypanthium, partly dissected with
view of the stamens, style, and ovary. — F. Stamens, lateral view (left), ventral view (right). — G. Seed. (A-G from the
holotype, C. V. Martin 283.)

Th is variety is named in honor of Franchise Cro-
zier for her work with the collections at the new
“Herbier de Guyane” (Cayenne), since its opening
in 1995.

Paratypes. FRENCH GUIANA. Kourou, Acevedo-
Rodriguez 4910 (CAY, US); Saul, Allorge 385 (P); Belizon,
Aubreville 295 (P); Saiil, Aumeeruddy 60 (CAY); Cacao,
Cremers 5534 (CAY, P. US); Montagne de Kaw, Cremers

5681 (CAY, P, US); FI. Mana, Cremers 7497 (BR, CAY, P,
US); Crique Kapiri, Cremers 1 1566 (CAY, US); Savane-
Roche Quatorze Juillet, Cremers 12255 (CAY. P. US); Saiil,
Descoings 20526 (CAY, P); Montagne de Kaw, Granville
212, 213 (CAY. P); Riviere Ouaqui, Granville 1752 (CAY.
P); Uayon des Emeri lions, Granville 2269 (CAY, P); Nord
du Massif des Emerillons, Granville 3942 (CAY); Saiil.
Granville 4452 (CAY. P); Riviere Ouaqui, Granville 4930
(CAY); Montagne Bellevue de ldnini, Granville 8142
(CAY, P, U); Trois Sauts. Grenand 975 (CAY, P); Uayon

378

Novon

Figure 3. SEM. Miconia ceramicarpa (DC.) Cogniaux
var. crozierae Cremers & C. V. Martin. Seeds. (From the
holotype, C. V. Marlin 283.)

Regina-kaw, Hequet 620 (CAY); Montagne de Kaw, Hoff
5550 (CAY, P, US); Montagne de kaw, Jansen-Jacobs 5214
(CAY, P, U); Trois-Sauts, Lescure 530 (CAY, P); route de
Belizon, Marlin 41 (CAY); Piste Saut Ueodate, Marlin 383
(B, BBS. BM. BBC, CAY, G, HAMAB. 1NPA, k, MG, MO,
NY, P, U, US); Saul, Mori 15628, 18340 (CAY, NY);
Fleuve Oyapock, Oldeman-T 847 (CAY, P); Fleuve Ma-
roni, Petibon 114 (P); Saul, Philippe 26919 (CAY, NY);
Kourou, Hb. L. C. Richard (P); Riviere Arataye, Riera 699
(CAY’. P); Riviere Inini. Sastre 3963 (CAY. P); Trois-Sauts,

Sastre 4653 (CAY, P); Rivifere Arataye, Sastre 5716 (CAY,
P); Maripasoula, Schnell 11518. 11567, 11657, 11685,
11699 (P), Riviere Ouaqui, Schnell 11903 (P). BRAZIL
Amapa: h)iirteig 1792 (P). Para: Jobert 102 (P); Bene-
vides, Poisson s.n. (P). Rio de Janeiro: Glaziou 9825 (P).

Acknowledgments. I thank all those who helped
me in this study, especially Mr. Sanip in the field,
J. Florence with the Latin diagnosis, C. Feuilletand
P. P. Lowry for the English translation, and many
thanks to Georges Cremers for his continuous sup¬
port and irreplaceable help.

I .ilerature Cited

Cremers, G. 1986. Architecture vegetative et structures
inllorescentielles de quelques Melastomataceae guva-
naises. ORSTOM Paris coll. Travaux et Documents
n°199; 1-248.

Dauchez, B. 1977. De quelques Melastomataceae en Gu-
yane. Montpellier, DEA de Botanique Tropicale.

Halle, F. & R. A. A. Oldeman. 1970. Essai sur
l’architecture et la dynamique de croissance des arhres
tropicaux. Masson & Cie Ed., Monographic 6; 1—178.

- , - & P. P. Tomlinson. 1978. Tropical Trees

and Forests. An Architectural Analysis. Springer Ver-
lag, Berlin.

Wurdack, J. J., T. Morley & S. Renner. 1993. Melasto¬
mataceae. In: A. R. A. Gorts-van Rijn (editor). Flora of
the Guianas, ser. A. 13. 99: 1 — 42S.

Paysonia , a New Genus Segregated from
Lesquerella (Brassicaceae)

Steve L. O' Kane, Jr.

Department of Biology, University of Northern Iowa, Cedar falls, Iowa 50614-0421, U.S.A.

steve.okane@uni.edu

I h san A. Al-Shehbaz

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

ihsan.al-shehbaz@mobot.org

ABSTRACT. The annual, auriculate-leaved species
of Lesquerella (Brassicaceae) of the southeastern
United States are transferred to the new genus Pay¬
sonia. The new combinations Paysonia auriculata,
P. densipila, P. grandiflora, P. lasiocarpa, P. lasio-
carpa subsp. berlandieri, P. lasiocarpa subsp. het¬
erochroma, P. lescurii, P. lyrata, P. perjorata, and
P. stonensis are proposed. The distinguishing char¬
acters between the combined taxon Lesquerella—
Physaria are discussed. A detailed description of
the genus and a key to the species are provided.

Key words: Brassicaceae, Lesquerella, Physar¬
ia, Paysonia, Southeastern United States.

In his account of the auriculate-leaved species
of the genus Lesquerella S. Watson, Rollins (1955:
241) stated that these species “are all more closely
related to each other than to other species of the
genus and should be formed into a separate section
or subgenus.” Based on leaf and trichome mor¬
phology, chromosome number, duration, and molec¬
ular data (see below), we concur with Rollins’s as¬
sessment of the close relationship among these
species and that they warrant taxonomic recogni¬
tion. No formal taxonomic recognition of this well-
marked group of species, however, has been pro¬
posed. Rollins and Shaw (1973) included them in
their informal “Group 10” (of 10 groups of Les¬
querella species), but in his monograph of die North
American Brassicaceae, Rollins (1993) made no
such distinction.

Rollins (1955: 241) wondered where had the au¬
riculate-leaved “Lesquerellas come from and by
what migratory route did they get here.” By “here”
he was referring to their geographic range in the
southeastern United States west to Texas and
Oklahoma. Rollins’s question presupposed that the
auriculate-leaved species were Lesquerellas, and
that they dispersed to the southeastern United
States from their main distribution in the American

West. The group containing the auriculate-leaved
species (hereafter Paysonia) also contain species of
the genera Dimorphocarpa Rollins, Dithyrea Har¬
vey, Lyrocarpa Hooker & Harvey, Serisyrenia
Greene, and Synthlipsis A. Gray. These five genera,
together with Paysonia, Lesquerella, and Physaria.
form a monophyletic group easily distinguished
from the rest of Brassicaceae by having pollen
grains with 4-10 colpi. By contrast, the remaining
ca. 96% of the family have trieolpate pollen.

In a companion paper (Al-Shehbaz & O'Kane,
2002 this issue) we combine Lesquerella and Phy¬
saria (Nuttall ex Torrey & A. Gray) A. Gray into a
much-expanded Physaria. In that paper we provide
a justification for these nomenclatural innovations
and supporting literature. The larger, monophyletic
Physaria is characterized by (1) stem leaves that are
cuneate or attenuate into a petiole-like base, (2)
chromosome numbers based on n — 4. 5, 6. or 7
(with a feu polyploids derived from these numbers;
see Rollins & Shaw. 1973), (3) perennial duration
(or biennial or rarely annual, through derivation from
perennial ancestors), (4) no documented interspecific
hybridization (Rollins & Shaw, 1973), and (5) tri-
chomes that are strictly stellate to stellate-peltate (of¬
ten webbed between the rays), and nearly always
tuberculate. We here place those species excluded
from the expanded Physaria in a new genus, Pay¬
sonia, named in honor of Edwin B. Payson (1893-
1927), the first monographer of Lesquerella (Payson.
1922). Paysonia is characterized by (1) stem leaves
that are sessile and auriculate, (2) chromosome num¬
bers based on n = 8, (3) an annual duration, (4)
frequent natural (and artificial) interspecific hybrid¬
ization (Rollins, 1957; Rollins & Shaw. 1973; Rol¬
lins & Solbrig, 1973), and (5) trichomes that lack
tubercles and are not stellate but rather are simple,
one-branched, bifurcate, or dendritic (Rollins & Ba-
nerjee, 1975, 1979).

Novon 12: 379-381. 2002.

380

Novon

Molecular data (O'Kane, unpublished) based on
DNA sequences of the nuclear ribosomal internal
transcribed spacer (ITS) support the recognition of
Paysonia and show that this genus and the com¬
bined taxon Physaria-Lesquerella shares a common
ancestor.

Paysonia O’Kane & Al-Shehbaz, gen. nov. TYPE:
Paysonia lescurii (A. Gray) O' Kane & Al-
Shehbaz.

Herba annua; pili ramosi minute stipitati et pilis sim-
pilieibus praesentibus. Folia caulina sessilia, sagittata vel
auriculata. Racemi ebracteati, valde elongati. Sepala ob-
longa, nonsaccata. Petala alba vel flava, obovata. Ovula
2— 8(— 10) per locula. Siliculis globosis, subpyriformis, su-
borbiculatis, vel subovatis, teretis vel compressis et an-
gusti- vel latiseptatis; septum completion vel perforatum;
stylus 1-3 mm longus; stigma integra. Semina eompressa;
cotyledones accumbentes.

Herbs annual. Trichomes coarse, simple, mixed
with much smaller, short -stalked, forked or sub-
dendritic ones. Stems erect to ascending, some¬
times decumbent. Basal leaves petiolate, rosulate,
lyrate or pinnatelv lobed. Cauline leaves sessile,
minutely to coarsely auriculate or sagittate, often
amplexicaul, dentate. Racemes many flowered,
ebracteate, often elongated considerably in fruit; ra-
chis straight. Fruiting pedicels slender, ascending
to divaricate or recurved, straight or slightly
curved. Sepals oblong, deciduous, suberect, as¬
cending, or spreading, equal, base of inner pair not
saccate, margin membranous. Petals yellow or
white, broadly obovate, apex rounded; claw slightly
differentiated from blade. Stamens 6, erect, tetra-
dynamous; filaments dilated or not at base; anthers
oblong, not apiculate at apex. Nectar glands con¬
fluent, subtending bases of median stamens, sur¬
rounding those of lateral stamens. Ovules 2 to 8(to
10) per locule. Fruit dehiscent silicles, globose,
subpyriform, suborbicular, or subovate, terete, la-
tiscptate or angustiseptate, not inflated, subsessile;
valves papery or leathery, veinless, glabrous or pu¬
bescent with branched and/or simple trichomes;
septum complete, perforated, or reduced to a rim,
membranous; style 1-3 mm long, slender, glabrous
or pubescent; stigma capitate, entire. Seeds sub-
biseriate, margined, orbicular to subovate, flat¬
tened; cotyledons accumbent.

Key to the Species ok Paysoma

la. Fruits orbicular or elliptic, flattened; valves often
with coarse simple trichomes and much smaller
branched ones.

2a. Fruit latiseptate; fruiting pedicels divari¬
cate-ascending; cauline leaves strongly au-

riculate to sagittate, amplexicaul; filaments

dilated at base . P. lescurii

2b. Fruits angustiseptate; fruiting pedicels re¬
curved; cauline leaves minutely auriculate,
not amplexicaul; filaments not dilated at

base . P. lasiocarpa

lb. Fruits globose to subpyriform, not flattened;
valves uniformly with small, simple and/or
branched trichomes.

3a. Flowers white; septum perforated.

4a. Fruits subpyriform, valves glabrous or
sparsely pubescent outside, densely pu¬
bescent inside; styles glabrous .

. P perforata

4b. Fruits subglobose, valves densely pu¬
bescent outside, glabrous inside; style

pubescent . P, stonensis

3b. Flowers yellow; septum complete.

5a. Fruit valves and styles pubescent .

. P. densipila

5b. Fruits valves and styles glabrous.

6a. Fruits 2.4— 3.5 mm long; ovules 2—

4 per locule; septum opaque .

. P. lyrata

6b. Fruits 4-6(-8) mm long; ovules

(4— )6— 8(-IO) per ovary; septum
translucent.

7a. Lower stems hirsute with
spreading simple trichomes;
petals 4—5 mm wide; fruiting
raceme congested, slightly

elongated . P. auriculata

7b. Lower stems pubescent with
branched trichomes; petals 6—

10 mm wide; fruiting raceme
lax, considerably elongated ....
. P. grandiflora

For a complete synonymy of the following laxa,
see Rollins (1993).

Paysonia auriculata (Fngelmann & A. Gray)
O’Kane & Al-Shehbaz, comb. nov. Basionvm:
Vesicaria auriculata Fngelmann & A. Gray,
Host. J. Nat. Hist. 5: 240. 1845. Lesquerella
auriculata (Engelmann & A. Gray) S. Watson,
Proc. Amer. Acad. Arts 23: 250. 1888. TYPE:
U.S.A. Texas: Austin Co., near San Felipe. F.
Lindheimer 217 (holotype, GH: isotypes, GH,
MO).

Paysonia densipila (Rollins) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella densi¬
pila Rollins, Rhodora 54: 186. 1952. TYPE:
U.S.A. Tennessee: Marshall Co.. Duck River
bottom, N of Verona, 10 Apr. 1949. A. ./.
Sharp, C. J. Felix & W. Adams 11187 (holo¬
type, GH; isotypes, GH. MO, NY).

Volume 12, Number 3
2002

O’Kane & Al-Shehbaz

Paysonia Segregated from Lesquerella

381

Paysonia grandiflora (Hooker) O'Kane & Al-
Shehbaz. comb. nov. Basionym: Vesicaria
grandiflora Hooker, Curtis’s Bot. Mag., n.s. 10:
pi. 3464. 1836. Lesquerella grandiflora (Hook¬
er) S. Watson. Proc. Arner. Acad. Arts 23: 250.
1888. TYPE: U.S.A. Texas: Austin Co.. San
Felipe, Drummond 20 bis (lectotype, designat¬
ed by Rollins & Shaw (1973), K; isolectotypes,
GH, NY).

Paysonia lasioearpa (Hooker ex A. Gray) O’Kane
& Al-Shehbaz, comb. nov. Basionym: Vesicaria
lasioearpa Hooker ex A. Gray, Smithsonian
Contr. Knowl. (PI. Wright.) 5: 13. 1853. Les¬
querella lasioearpa (Hooker ex A. Gray) S.
Watson, Proc. Amer. Acad. Arts 23: 251.
1888. TYPE: “Country and state unknown:”
between Bexar and Trinity River. May 1828.
Berlandier s.n. (holotype, K).

Paysonia lasioearpa subsp. berlandieri (A.

Gray) O'Kane & Al-Shehbaz, comb, et stat.
nov. Basionym: Synthlipsis berlandieri A. Gray,
in Emory, Report U.S. & Mex. Bound. Survey
34. 1859. Lesquerella lasioearpa var. berlan¬
dieri (A. Gray) Payson, Ann. Missouri Bot.
Ganl. 8: 139." 1922* TYPE: Mexico. Tamauli-
pas: Matamoros. Berlandier 301 7 (lectotype,
designated by Rollins & Shaw (1972), GH; is¬
olectotypes, GH, US).

Paysonia lasioearpa subsp. heterochroma (S.

Watson) O'Kane & Al-Shehbaz, comb, et stat.
nov. Basionym: Synthlipsis heterochroma S.
Watson, Proc. Amer. Acad. Arts 17: 321.
1882. Lesquerella lasioearpa var. heterochroma
(S. Watson) Rollins, Rhodora 57: 245. 1955.
TYPE: Mexico. Nuevo Leon: Guajuco, 27 mi.
SE from Monterrey, Edward Palmer 33 (holo¬
type, GH).

Paysonia lescurii (A. Gray) O'Kane & Al-Sheh-
baz, comb. nov. Basionym: Vesicaria lescurii A.
Gray, Manual Bot., ed. 2. 38. 1856. Lesquerella
lescurii (A. Gray) S. Watson. Proc. Amer. Acad.
Arts 23: 250. 1888. TYPE: U.S.A. Tennessee:
Davidson Co., near Nashville, Apr. 1855, Leo
Lesquereux s.n. (lectotype. designated by Rol¬
lins (1952), GH).

Paysonia lyrata (Rollins) O'Kane & Al-Shehbaz,
comb. nov. Basionym: Lesquerella lyrata Rol¬
lins, Rhodora 57: 252. 1955. TYPE: U.S.A.
Alabama: Franklin Co., near Richardson’s
Crossing, 7 mi. E of Russellville, 16 Apr.
1955. Reed C. Rollins , George R. Cooley & L.
,/. Brass 5599 (holotype, GH; isotypes, GH.
MO, NY, UC).

Paysonia perforata (Rollins) O’Kane & Al-Sheh-
baz, comb. nov. Basionym: Lesquerella perfor¬
ata Rollins, Rhodora 54: 190. 1952. TYPE:
U.S.A. Tennessee: Wilson Co., 5 mi. N of Leb¬
anon, 30 Mar. 1952. Reed C. & Diane Rollins
5207 (holotype, GH; isotypes, F. GH. MO. NY.
US).

Paysonia stonensis (Rollins) O'Kane & Al-Sheh¬
baz, comb. nov. Basionym: Lesquerella stonen¬
sis Rollins. Rhodora 57: 255. 1955. TYPE:
U.S.A. Tennessee: Rutherford Co.. E Fork of
Stones River, Walterhill, 26 Apr. 1955, Reed
C. Rollins 55176 (holotype, GH; isotypes, F,
GH. MO, NY, US).

Acknowledgments. We are grateful to Neil A.

Harriman and Victoria C. Hollowell for their criti¬
cal comments on the manuscript.

Literature Cited

Al-Shehbaz, I. A. & S. L. 0 Kane, Jr. 2002. lesquerella
is united with Physaria (Brassicaceae). Novon 12: 319-
329.

Payson, E. B. 1922. A monograph of the genus Lesquer¬
ella. Ann. Missouri Bot. Card. 8: 103—236.

Rollins, R. C. 1952. Some Cruciferae of the Nashville
Basin, Tennessee. Rhodora 54: 182-192.

- . 1955. The auriculate-leaved species of Lesquer¬
ella (Cruciferae). Rhodora 57: 241-264.

- . 1957. Interspecific hybridization in lesquerella

(Cruciferae). Contr. Gray Herb. 181: 3-40.

- . 1993. The Cruciferae of Continental North Amer¬
ica. Stanford Univ. Press, Stanford.

- & U. C. Banerjee. 1975. Atlas of the trichomes

of lesquerella (Cruciferae). Publ. Bussey Inst. Harvard
Univ. 1975: 1-48.

- & - . 1979. Trichome patterns in Physaria.

Publ. Bussey Inst. Harvard Univ. 1979: 65—77.

- & E. A. Shaw. 1972. Nomenclatural changes in

Lesquerella. Rhodora 74: 76—79.

- & - . 1973. The Genus Lesquerella (Cruci¬
ferae) in North America. Harvard Univ. Press, Cam¬
bridge.

- & 0. T. Solbrig. 1973. Interspecific hybridization

in Lesquerella. Contr. Gray Herb. 203: 3—48.

Novae Gesneriaceae Neotropicarum X: A New Species of
Achimenes from Mexico

Angelica Ramirez Ron

Herbario Nacional tie Mexico, Facultad tie Ciencias, Universidad Nacional Autonoma de

Mexico, Mexico D.F., Mexico

Laurence E. Shag

U.S. National Herbarium, MRC-166, Smithsonian Institution, P.0. Box 37012,
Washington, D.C. 20013-7012, U.S. A. skog.larry@nmnh.si.edu

ABSTRACT. A new species of Achimenes Persoon,
A. hintoniana , is described from western Mexico
and distinguished from the closely related A. mex-
icana by the pilose indumentum of the stem,
leaves, and pedicels, the linear-lanceolate calyx
lobes, and the slightly saccate purple or rarely
white corolla.

Key words: Achimenes, Gesneriaceae, Mexico.

Achimenes Persoon is a genus of neotropical Ges¬
neriaceae distributed from northern Mexico (Chi¬
huahua and Tamaulipas states) south to Ecuador,
Venezuela, and Jamaica and comprising 23 species.
About two-thirds of the species are endemic in
Mexico, where numerous collections have been
made and where the center of diversification ap¬
pears to be. A recent revision of Achimenes (Ra¬
mirez Roa, 1987) and recent plant collections have
shown that the description of a new species, A. hin¬
toniana from Mexico, is warranted.

Achimenes hintoniana Ramirez Roa & L. E.
Skog, sp. nov. TYPE: Mexico. Nayarit: Nayar,
7 km al S de Ea Mesa del Nayar, encinar en
barranca, breeha a Jesus Marfa, 22CI2,N.
104°36'W, 1400 m, 23 Sep. 1989, P. Tenorio
L. & G. Glares F. 16332 (holotype, MEXU; iso¬
types, FCME, US). Figure 1.

Herba erecta pilosa non raniosa ad 56 cm alta. Folia
petiolata opposita paribus subaequalibus, laminis ovatis
vel ovato-ellipticis longiacuminatis margine serratis. Flo¬
res axil lares solitarii; calycis lobi lineari-lanceolati pilosi;
corolla infundibuliformis ad 6.9 cm longa purpurea, fauce
albida glabra maculis purpureis, tube sursum ampliato
base ieviter saccata, limbo 5-fido; stamina 4 (quinto ad
staminodium reducto) didynama inclusa, filamentis glabris
tubo insertis; discus annularis integer rnembranaceus.
Ovarium glabrum; stylus puberulus; stigma infundibuli-
forme; capsula pilosa.

Herbs; stems erect, 5.3-56 cm tall, green or red-
tinged, pilose. Rhizome 6.4 cm long, 7.5 mm wide.

scales red-tinged. Leaves almost isophyllous; ovate-
elliptic or ovate, 3.2-12 X 1.4-8 cm, apex largely
acuminate, base oblique, margin serrate, 10-27
teeth per side, and filiate, 7—12 pairs of secondary
veins, the upper surface pilose, green, the lower
surface pilose, pale green or red; petioles 1.5— 4.5
cm long, pilose. Flowers solitary; pedicels 1.1— 2.3
cm long, pilose, green or red; floral tube 2-3 X
1 .5—2 mm, pilose or almost glabrous; calyx lobes
linear-lanceolate, 4-9 X 1-2 mm, pilose or nearly
glabrous, ciliate; corolla infundibuliform, slightly
oblique in the calyx, 4.1— 6.9 cm long, purple or
rarely white, glabrous to pilose, tube slightly sac¬
cate, 2-5 cm long, slightly ventricose, 3-6 mm
wide at the base, very narrow, 1.2— 2.5 cm wide at
the throat, white with purple spots at the throat,
limb 3. 4-4. 2 cm wide. 5-lobed, lobes oblate, 0.9—
1.9 X 1-2.2 cm, crenate, purple or rarely white;
stamens 4 (with a reduced staminode), 2. 5-2. 8 cm
long, filaments briefly adnate to corolla tube base
and free above, glabrous, anthers ovate depressed,
0.5—1 X 1-1.5 mm; annular nectariferous disc en¬
tire. membranous, 0. i mm long; ovary glabrous,
style 2. 3-3.3 cm long, glandular-puberulous, stig¬
ma funnel-shaped. Capsule 8 mm long, 4 mm wide,
green, lightly pilose to glabrous. Seeds oblong, ea.
0.5 X 0.2 mm, red-brown, longitudinally striate.

Distribution and ecology. Endemic to Mexico,
Achimenes hintoniana is known mainly from Jalisco
and Nayarit states at 800-1650 m. with outliers in
Durango and Michoacan. growing in oak forests,
mixed oak-pine forests, dry deciduous forests, and
montane rain forests. Collections have been made
of flowering plants in July through September and
fruiting in September.

The specimens included here had in most cases
been identified as the closely related Achimenes
mexicana (Seemann) Bentham & J. D. Hooker ex
Fritsch. However, A. mexicana has villous indu-

Novon 12: 382-384. 2002.

Volume 12, Number 3
2002

Ramirez Roa & Skog
Achimenes hintoniana from Mexico

383

Figure 1. Achimenes hintoniana Ramfrez Roa & L. E. Skog. — A. Habit. — B. Lower leaf surface. — C. Corolla. —
D. Opened corolla with stamens and staminode. — E. Calyx. — F. Pistil with calyx and disc. — G. Dehisced capsule
with surrounding persistent calyx and floral tube split to the base, and funicles within the capsule. — H. Seeds. — 1.
Rhizome scales. (A-F, I from R. McVaugh 16443; G, H from R. McVaugh 16336.)

384

Novon

ment with hairs of various lengths primarily on the
stems, petioles, pedicels, and floral tube. I he in¬
dumentum of A. hintoniana is pilose and much less
dense. In addition, the calyx lobes in A. mexicana
are lanceolate-ovate, and the blue-violet corollas
are ventricose with a broadly sacciform base. Achi-
menes mexicana occurs in Chihuahua, Durango,
Sinaloa, and Sonora. The two species are not sym-
patric except for an unlocalized collection of A. hin¬
toniana from Durango (Rose 2331).

The variation seen in Achimenes hintoniana is
principally in size, but the specimen collected by
McVaugh (16336A) has white corollas. McVaugh
mentioned on the specimen label that these plants
were found among plants with purple corollas. This
color variation is not unusual in the genus and is
found also in A. antirrhina (DC.) C. V. Morton, A.
erecta (Lamarck) H. P. Fuchs, A. heterophylla (Mar-
tius) DC., and A. patens Bentham. The epithet hon¬
ors George B. Hinton (1882—1943), a collector of
plants in southwestern Mexico where he collected
many specimens of Achimenes species.

Paratypes. MEXICO. Durango: 16 Aug. 1897, ./.
Rose 2331 (NY, US). Jalisco: 8.2 mi. Vi ol Chapala on
road to Jocotepec, at km 62, hills above Lake Chapala’s
N shore, IT Anderson & C. Anderson 5169 (GH, MICH.
NY); Mpio. Talpa de Allende, Texcalame, sobre la brecha
de Talpa a Cuale, M. Chdzaro et al. 7373 (MEXU); Mpio.
Tonald, Barranca de Huentit&n, microcuenca de las 7 Cas-
cadas, 2 km al E de Tonala, A. Flores-M. & P. Ramos G.
2423 (MEXU); 10.1 mi. SSW Autlan off hwy. 80, R. Krai
27657 (ENCB); 9.4 mi. from Autlan (kms 218—219) on
road to Barra de Navidad, //. Moore & G. Hunting 3725
(US); km 69-70, near Jocotepec, on hills above the W end
of Lake Chapala, E. Norman 64 (MICH): along Jocotepec—
Chapala road on the N side of Lake Chapala, ca. 2 mi. (3
km) E of San Juan CozaH, .4. Sanders 74140 (MICH);
along hwy. 110 about 5 mi., NE of Bio Tuxpan bridge and
about 20 mi. NE of Huizache, IT Stevens & M. Fairhurst
1320 (MICH, MO); trail between El Chante and Manantan
about 15 mi. SSE of Autldn, R. Wilbur & C. Wilbur 2092
(MICH, US); 10 mi. S of Autlan. R. Wilbur & C. Wilbur
2113 (MICH, US). Michoacan: Sierra Naranjillo-Puen-
tes, Hist. Coaleornan, G. Hinton et al. 15973 (US). Nay-
arit: mountain 10 mi. SE of Ahuacatlan, on road to Bar-
ranea del Oro, C. Feddema & R. King 253 (MICH, US);

3.5 mi. NW of Ahuacatlan, S of the base of volcdn Ce-
boruco and about 1 km N of the highway, C. Feddema
422 (MICH); 31 km al NE de Jesus Marfa, eamino Jesus
Marfa— Huejuquilla, G. Flores-F. et al. 2094 (MEXU, US);
Mpio. Nayar 5 km al SE de la Mesa del Nayar, G. Elores-
F & It Ramirez R. 2639 (MEXU, US); Mpio. Nayar. 32.5
km al NE de Jesus Marfa, La China, G. Flores-F. & P.
Tenorio /,. 1003 (MEXU, US); Mpio. Nayar, 22.7 km al
NW de Jesus Marfa, eamino Jesus Marfa— La Mesa del
Nayar, G. Flores-F. & P. Tenorio L 1333 (MEXU, US); 5-

8 km al NW de la Mesa del Nayar, eamino a la Mesa del
Nayar-Villa de Guadalupe (Cerro Cangrejo), G. Flores-F
et al. 1130 (MEXU, US); 2 mi. NW of Ahuacatldn, R.
McVaugh 15405 (MICH); Arroyo de la Fundieion. 5 mi.
SE of Ahuacatlan on road to Barranca del Oro, R. Mc¬
Vaugh 16336 (MICH. US), R. McVaugh 16336A (MICH);

9 mi. N of Compostela, R. McVaugh 16443 (MICH, US);
ca. 12 mi. W of Tepic, 6.5 mi. E of Jaleocotan, McVaugh
et td. 13919 (MICH), McVaugh et al. 13975 (MICH); km
15 on road from Tepic to Jaleocotan, H. Moore & G. Hun¬
ting 3699 (MEXU, UC, US); along road from Hwy. 15 to
Santa Cruz (W of Tepic), about 9.9 mi. W of Hwy. 15, IT
Stevens 1436 (ENCB, MICH. MO); km 4 de la carr. a San
Bias, del entronque con carr. Tepic— Mazatlan. O. Tellez V.
et al. 10742 (MO). Cultivated: Growing at Marie Selby
Botanical Gardens, Sarasota, EL, originally from Nayarit,
W of Tepic near La Libertad, N. Hashor 1373 (F, SEE.
US).

Acknowledgments. We are grateful to Fernando
Chiang Cabrera from the Herbario Nacional tie
Mexico for preparation of the Latin description, to
Cathy Pasquale- Johnson for her excellent rendering
of the illustration, to Eric Roalson, Smithsonian
Laboratory for Molecular Systematics, for his ob¬
servations, to M. en C. Laura Marquez for her com¬
puter assistance, to Mario Sousa for allowing the
lust author to use the herbarium facilities at
MEXU, to the reviewers, and to the herbaria that
lent their specimens lor this study. The first author
also acknowledges the support of a Smithsonian
Short-Term Visitor Grant that allowed her to pursue
her studies ol Achimenes at the National Museum
of Natural History' in Washington, D.C.

Literature Cited

Ramfrez Boa, A. 1987. Revision de Achimenes (Gesneri-
aceae). Tesis de Licenciatura. Facultad de Ciencias,
UNAM. Mexico, D.F. Mexico.

Oxypetalum rusticum (Apocynaceae, Asclepiadoideae), a New
Species from the Espinhago Range, Minas Gerais, Brazil

Alessandro Rapini

Departamento de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, Caixa
Postal 11461, CEP 05422-970, Sao Paulo, SP, Brazil, arapini@usp.br

ABSTRACT. The Latin diagnosis of Oxypetalum rus¬
ticum Rapini is provided to validate the speeies, pro¬
posed hut not validly published in the survey of the
Asclepiadoideae from the Espinhago Range of Minas
Gerais, Brazil. The speeies, endemic to the Diaman-
tina Plateau, is closely related to O. insigne (l)ecaisne)
Malme and 0. glabrum (Decaisne) Malme, differing
mainly by the wooly indumentum of the branches and
the longer corpusculum of the pollinaria.

Key words: Apocynaceae, Asclepiadoideae, Bra¬
zil, Espinhayo Range, Minas Gerais, Oxypetalum.

Oxypetalum R. Brown includes more than 150
species distributed mainly in Brazil (Hoehne, 1916)
and Argentina (Meyer, 1943), with its species rich¬
ness decreasing toward the north of its distribution.
The genus is usually recognized by the gynostegium,
which is prolonged in a rostrum, and the translator
amis of the pollinaria, which have lateral teeth.

Studies on the Asclepiadoideae from the Espin-
hayo Range of Minas Gerais, Brazil (Rapini, 2000;
Rapini et al., 2001), have suggested new species and
several taxonomic changes in the subfamily (see Ra¬
pini, 2002, for new species of Ditassa R. Brown and
discussions on taxonomic concepts of Metastelmati-
nae genera). Here, the Latin diagnosis of Oxypetalum
rusticum Rapini. a new species proposed but not val¬
idly published in the survey, is provided (for full
description and paratypes, see Rapini et al., 2001).

Oxypetalum rusticum Rapini, sp. nov. TYPE:
Brazil. Minas Gerais: Mun. Datas [“Diaman-
tina”], Diamantina-Gouveia road, km 605, 14
Feb. 1998 (fl), F /V. Costa, P. T. Sana, W. Vi¬
le gas, L. Campaner & F. P. Andrade 35 (ho-
lotype, SPF; isotype, K). Figure 1.

Oxypetalo insigni similis, ramis et inflorescentiis racem-
iformibus lanatis, foliis persaepe latioribus supeme villosis
sed nervis lanatis, laciniis corollae sursum glabrescentibus
et retinaculis maioribus, ultra 2.4 mm longis, recedens.

Twining vine; branches wooly. Leaves elliptic
or ovate, cordate to reniform at the base, pilose
adaxially, villous on the veins, villous abaxially,
wooly on the veins. Cymes raeemiform, with up

to 6 flowers. Corolla hirsute to pubescent abaxi¬
ally, puberulent adaxially on the tube, barbulate
on the throat, glabrescent toward the apex of the
lobes; lobes oblong, 0.9-1. 4 cm long, twisted,
patent. Corona lobes scale-like to widely oblong,
cleft, cornieulate adaxially. Corpusculum spath-
ulate, keeled to involute, 2.5— 2.6 mm long; pol-
linia 0.65-0.85 mm long. Apex of the gynoste¬
gium prolongation coroniform, slightly bilobate.
Follicles wooly to velutinous.

Oxypetalum rusticum, 0. insigne (Decaisne)
Malme, and 0. glabrum (Decaisne) Malme, the last
of which is sometimes considered a variety of O.
insigne (Fontella-Pereira & Schwarz, 1983; Fontel-
la-Pereira et al., 1984), form a complex of twining
species with the gynostegium prolongation cup¬
shaped or coroniform to bilobate at apex. The new
species differs from these two by its wooly (vs. gla¬
brous to hispid) indumentum of the branches and
mainly by its longer corpusculum (2.5— 2.6 mm long
vs. up to 2.1 mm long in the other two species).
Due to the leaves usually broad, villous abaxially,
and especially the wooly branches, O. rusticum is
not subtle when compared with its closely related
and relatively slender species (see Rapini, 2000,
for photos ol the three species), w hich is the reason
for the specific epithet.

Phenology. Oxypetalum rusticum has been
found with flowers between November and April,
and with lruits in April.

Distribution and habitat. The species is re¬
stricted to the Diamantina Plateau, in the northern
range of its closely related taxa: 0. glabrum extends
into the southern Diamantina Plateau and O. insig¬
ne to Serra do Cipo, also in the Espinhayo Range
of Minas Gerais. Oxypetalum rusticum occurs only
in campos rupestres, an open vegetation on quartzite
or sandstone formations that appears at elevations
above 900 m, and which is known for its high level
of plant endemism (Giulietti et al., 1987, 1997; Ra¬
pini et al., in press).

Acknowledgments. I thank Renato de Mello-
Silva for the Latin diagnosis and M. Lucia Ka-

Novon 12: 385-387. 2002.

386

Novon

Figure 1. Oxypetalum ruslicum Rapini. — A. Habit, fertile branch, with indumentum iu detail at right. — 15. Flower.
— C. Lobe of corona, adaxial view. — 1). Appendage of gynostegium. — E. I’ollinarium. Drawn by the author from the
holotype, Costa et al. 35 (SPF).

wasaki for the English review of an initial version
of the manuscript, and also for their assistance
during my Ph.D. thesis, developed at the Univer-
siilade de Sao Paulo (USP) and funded by the Fun-

dagao de Amparo a Pesquisa do Estado de Sao
Paulo (FAPESP) and Coordenagao de Aperfeigoa-
mento de Pessoal de Nivel Superior (CAPES)
foundation.

Volume 12, Number 3
2002

Rapini

Oxypetalum rusticum from Brazil

387

Literature Cited

Fontella-Pereira, J. & E. A. Schwarz. 1983. Estudos em
Asclepiadaceae, 18. Novas combinagoes e novos sino-
nimos. Bradea 4: 13—20.

- . M. C. Valente & E. A. Schwarz. 1984. Contri-

buigao ao estudo das Asclepidaceae brasileiras, 21. As¬
clepiadaceae do municipio de Ouro Preto, Estado de
Minas Gerais — Uma sinopse. Bob Mus. But. Kuhlmann
7: 63-127.

Giulietti, A. M., N. L. Menezes, J. R. Pirani, M. Meguro
& M. G. L. Wanderley. 1987. Flora da Serra do Cipo,
Minas Gerais: Caracterizagao e lista de especies. Bob
Bot. Univ. Sao Paulo 9: 1-152.

- . J. R. Pirani & R. M. Harley. 1997. Espinhago

Range region. Pp. 397^404 in S. D. Davis, V. H. Hey-
wood, O. Herrera-MacBryde, J. Villa-Lobos & A. C.
Hamilton, Centres of Plant Diversity. A Guide and
Strategy for their Conservation, Vol. 3. The Americas.
WWF/IUCN, Cambridge.

Hoehne, F. C. 1916. Monographia das Asclepiadaceas
brasileiras. Oxypetalum, R. Br. e Calostigma, Dene.
Commiss. Linhas Telegr. Estrateg. Matto-Grosso Ama¬

zonas 38(1): 1-131, tabs.: 1-59; et ib. (1), supb: 1-13,
tabs. 60—62.; et ib. (2): 1-29, tabs. 1-12.

Meyer, T. 1943. Revision de las especies argentinas del
genero Oxypetalum (Asclepiadaceae). Lilloa 9: 5—72 +
16 tabs.

Rapini, A. 2000. Espinhago Range, Minas Gerais, Brazil.
Asclepiadoideae (Apocynaeeae). In R. Forster, M. R.
Metz & B. B. Lin, Rapid Color Guide n" 19. Environ¬
ment and Conservation Programs, Field Museum, Chi¬
cago.

- . 2002. Six new species of Ditassa R. Br. from the

Espinhago Range, Brazil, with notes on generic delim¬
itation in Metastelmatinae (Apocynaeeae— Asclepiadoi-
deae). Kew Bulb 57(3).

- , R. Mello-Silva & M. L. Kawasaki. 2001. Ascle¬
piadoideae (Apocynaeeae) da Cadeia do Espinhago de
Minas Gerais. Brasil. Bob Bot. Univ. Sao Paulo 19: 55-
169.

- , - & - . In press. Richness and en¬
demism in Asclepiadoideae (Apocyanaceae) from the
Espinhago Range of Minas Gerais, Brazil — A conser¬
vationist view. Biodiversity & Conservation.

Additions to Neocuatrecasia (Eupatorieae: Asteraeeae)

Harold Robinson

Botany, National Museum of Natural History, MRC-166, Smithsonian Institution, Box 37012,
Washington, D.C. 20013-7012, U.S.A. Robinson.Harold@nmnh.si.edu

Abstract. Neocuatrecasia sandiensis II. K obin-
son and N. yungasensis H. Robinson from high el¬
evations in Peru and Bolivia and N. tysonii H. Rob¬
inson from lower elevations in Bolivia are described
as new. A key is furnished to the 12 species of the
as yet inadequately collected genus.

Key words: Asteraeeae, Bolivia, Eupatorieae,
Neocuatrecasia, Peru.

The genus Neocuatrecasia was described by King
and Robinson in 1970 on the basis of four species
from tbe eastern slopes of the Andes in Peru and
Bolivia that had superficial resemblance to small
members of the widespread genus Ageratina Spach.
Neocuatrecasia can be distinguished from Agerati¬

na by the hairs on the base of the style. This char¬
acter. along with the ornamented cell walls of the
anther collar and the usually stipitate bases of the
achenes, places the genus far from Ageratina and
in the subtribe Gyptidinae (King & Robinson,
1987). A fifth species was transferred and four ad¬
ditional species were described in following years
(King & Robinson, 1972, 1974, 1986. 1988), but
no key has been provided since 1972 when there
were only six known species in the genus. Since
new' collections of the genus have often proved to
be new species, one presumes that many further
species remain to be discovered. Three more new
species are described herein, and a key is provided
for the 12 species now known in this Peruvian and
Bolivian genus.

Key to the Species of Neocuatrecasia

la. Peduncles with numerous long-stipitate glands longer than any non-glandular hairs.

2a. Leaves sessile; pappus of ca. 20 long fragile very slender setae not contiguous at bases (Peru) .

. N. sessilifolia R. M. King & H. Robinson

2b. Leaves distinctly short-petiolate or petioliform at base; pappus of 25 to 35 persistent setae that are
short or with nearly contiguous bases.

3a. Inflorescence broadly cymiform, with alternate branching; involucral bracts mostly narrowly ellip¬
tical, outer bracts bicostate in basal half or more; corollas with throats less than twice as wide as
basal tube; base of style enlarged; style branches scarcely twice as wide as shaft; pappus bristles
a third or less as long as the corolla; plants occurring at elevations of 500 m or less (Bolivia) . . .
. N. tysonii If. Robinson, sp. nov.

3b. Heads few or single, terminal, with subtending leaves opposite; involucral bracts broadly oblong,
only weakly veined, not costate; corollas with throats nearly three times as wide as basal tubes;
base of style not broadened except by hairs; style branches three or more times as broad as shaft;
pappus bristles half as long as corolla or longer; plants occurring at elevations of 2000-3500 m
(Bolivia) . N. yungasensis H. Robinson, sp. nov.

lb. Stipitate glands of peduncles, when present, scarcely longer than non-glandular hairs.

4a. Corolla throat 3 or more times as long as lobes.

5a. Leaf blades deeply lobed or dissected (Bolivia) . N. lobata (B. L. Robinson) R. M. King

& H. Robinson

5b. Leaf blades not deeply lobed or dissected.

6a. Heads with ca. 50 florets, with 15 or more involucral bracts (Peru) . N. weddellii (B. L.

Robinson) B. M. King & II. Robinson
6b. Heads with ca. 25 to 30 florets, with 10— 1 2( — 14) involucral bracts.

7a. Involucral bracts with acute tips; veins of leaves densely hirtellous abaxially (Bolivia)

. N. feuereri R. M. King & H. Robinson

7b. Involucral bracts broadly rounded at tips; veins of leaves puberulous with small hairs

abaxially (Peru) . N. sandiensis H. Robinson, sp. nov.

4b. Corolla throat 2 or less times as long as the lobes.

8a. Stems and peduncles hirsute to hispidulous with spreading pale hairs; sessile glandular riots red;
petioles of main leaves to 15 mm long; achenes with small setulae or scabrae along ribs (Peru)
. N. dispar (B. I.. Robinson) R. M. King & H. Robinson

8b. Stems and peduncles hirtellous to puberulous with often darkened hairs; glandular dots yellowish
to brownish; petioles of leaves only 1-5 mm long; achenes with distinct elongate setulae.

Novon 12: 388-392. 2002.

Volume 12, Number 3
2002

Robinson

Neocuatrecasia

389

9a. Corolla throats abruptly expanded at base to twice or more as wide as constriction of basal
tube; style with pubescence restricted to base.

l()a. Corollas with few or no hairs on lobes; leaves with few coarse serrations, with few
hairs adaxially, abaxial puberulence of main veins not coarser than pubesence on

remainder of surface (Peru) . N. cuzcoensis R. M. King & H. Robinson

10b. Corollas with numerous hairs on lobes; leaves entire or with shallow crenations, mi¬
nutely puberulous adaxially, abaxial hairs of midvein and proximal secondary veins
larger than those of remainder of surface (Rolivia) . . N. thymifolia (Britton) R. M. King

& H. Robinson

9b. Corolla throats less than twice as wide as constriction of basal tube; style with pubescence
continuing on to lower part of shaft.

1 la. Leaf blades elliptical, subsessile; peduncles with short-stipitate glands among the non-

glandular hairs (Peru) . N. mancoana (B. L. Robinson) R. M. King & 11. Robinson

lib. Leaves blades broadly ovate with subtruncate bases, petiolate; peduncles without stip-

itate glandular hairs (Peru) . N. hirtella R. M. King & H. Robinson

The following species are described as new.

Neocuatrecasia tysonii H. Robinson, sp. nov.
TYPE: Bolivia. Beni: Mamore, frequent in tree
margin of lagoon, spreading 1 mi. SW San Joa¬
quin, 19 Mar 1964, E. L. Tyson & M. Kuns
997 (holotype, MO). Figure 1A-D.

A speciebus aliis in inflorescentiis laxe cymiformibus
et in bracteis involucri anguste ellipticis externe ad me¬
dium bicostatis et in ramis stylorum angustioribus et in
setis pappi perbreviorihus et in basibus stylorum inflatis
et in distributio non andina differt.

Herbs to I m tall, moderately branched at ca.
45° angles; stems brown, covered with dense felt of
stipitate glands; internodes mostly 2—7 cm long.
Leaves opposite; petioles 2—5 mm long; blades nar¬
rowly ovate to lanceolate, 1.5— 7.5 X ca. 0.5— 1.5
cm. base broadly acute to narrowly acuminate, mar¬
gins with 5 to 15 often irregularly sized and irreg¬
ularly spaced serrations, apex narrowly acute, ad-
axial surface pilose with slender pale hairs, abaxial
surface paler green, pilosity mostly on veins, with
numerous dark glandular dots; trinervate with slen¬
der ascending lateral veins from near base. Inflo¬
rescence laxly cymiform with alternate branching;
peduncles 2—22 mm long, covered with slender
stipitate glands. Heads broadly campanulate, ca. 5
mm high and wide; involucral bracts ca. 20. most
narrowly elliptical, innermost ones linear-lanceo¬
late, bicostate in basal half or more, ca. 3 mm long,
ca. 0.5 mm wide, acute, puberulous with ncn-glan-
dular hairs abaxially, few stipitate glands at base
and along costae. Florets 25 to 30; corollas white
to pale violet, 2.0— 2.4 mm long, tube 0.3-0. 4 mm
long, throat cylindrical to narrowly funnelform,
scarcely twice as wide as basal tube, 1.2— 1.5 mm
long, lobes narrowly triangular, 0.4— 0.5 mm long,
with glandular dots and no hairs outside; anther
thecae ca. I mm long; style base inflated, with
hairs, no hairs on shaft; style branches narrow,
scarcely twice as wide as shaft. Achenes 1.8— 2.5
mm long, with short basal stipe ca. 0.3 mm long.

with scattered short setulae on ribs and surfaces;
pappus bristles usually 10 to 12, length ca. 0.3—
0.8 mm, very slender and often not contiguous at
base.

Neocuatrecasia tysonii is the most phyletically
divergent member of the genus with its broadly cy¬
miform inflorescence, its narrow more strongly bi¬
costate involucral bracts, and its occurrence at low
elevations in Bolivia. The narrow style branches
and distinctly broadened style base are also dis¬
tinctive. Still, all essential characteristics are those
of Neocuatrecasia. There is alternate branching in
the inflorescences of other species of the genus, but
the inflorescences are not as large or spreading as
in N. tysonii. The pappus is short in some speci¬
mens of a number of the other species, but usually
not less than half as long as the corolla. The
achenes of the present species have a basal stipe,
although it is very short compared to most of those
in the genus. The higher elevation N. feuereri of
Bolivia, which lacks stipitate glands, may he the
closest relative on the basis of its pointed involucral
bracts and comparatively narrow style branches.

The species occurs at elevations of 150—500 m
in flat grasslands, grazed natural savanna with scat¬
tered shrubs and small tortuous trees, in areas bor¬
dering on granitic outcrops, and in semideciduous
forests.

Paratypes. BOLIVIA. Beni: Itenez, S side of Rio
Guapore, “Campo Verde,” ca. 15 km ESE of Costa
Marques [Edo. Rondonia, Brazil |, 150 m, 31 Mar. 1987,
M. Nee 34624 (MO, NY, US). Santa Cruz: Velasco, Re-
serva Forestal Bajo Paragua, Cero Diamantina, 10 km al
S del Aserradero Cero Pelado, laja granitica grande (in-
selberg), 350 m, 13 May 1994, T. Killeen & Welle ns 6368
(MO, US, USZ); Prov. Nuflo de Chavez, Lomerio, comu-
nidad Las Trancas, 500 m, II Nov. 1994, 7. Killeen, Jar-
dim, Freire, Vriesendorp & Medina 7124 (US, USZ); area
de estudio del proyecto “BOLEOR,” Las Trancas-95, 450
m, 12 Dec. 1994, F. Mamani & A. Jardim 404 (MO, USZ).

390

Novon

Figure 1. Neocuatrecasia. A-l). N. tysonii H. Robinson, drawn from Killeen & Wellens 6 368 (US). — A. Corolla with
style branches. — R. Corolla in longitudinal section with stamens and style. — C. Involucral bract. — I). Achene.

Volume 12, Number 3
2002

Robinson

Neocuatrecasia

391

Neocuatrecasia yuiigasensis II. Robinson, sp.
nov. TYPE: Bolivia. Dpto. Ea Paz: Prov. Sud
Yungas, Nor-Oeste de Chojlla valle del Rio
Pongo Pampa 2230 m, exposition SO. ladera
rocosa vertical, hierba —8 cm, flores blancas,
7 Sep. 1987, E. Vargas & R. Seidel 505 (ho¬
lotype, US; isotype, LPB). Eigure 1E-H.

Ad N. feuererii similis sed in caulibus et foliis et pe-
dunculis et bracteis involucri longe stipitate glanduliferis
et in faucibus corollarum late infundibularibus distincta.

Erect herbs over 8 cm tall (only a piece of stem
seen, branching not seen); stems, petioles, leal
blades, peduncles, and involucral bracts with slen¬
der stipitate glands. Leaves opposite; blades mem¬
branaceous, ovate, 0.7- 1.5 X 0.6— 1.0 cm, base
subtruncate, margins with usually 3 blunt teeth,
apex short-acute with extreme tip blunt, with scat¬
tered stipitate glands adaxially, stipitate glands on
margins and abaxially on main veins, glandular
dots on lamina abaxially; weak trinervation from
near base. Inflorescence (as seen) a single terminal
pedunculate head; peduncle ca. 14 mm long,
densely covered with stipitate glands. Heads ca. 6
mm high, ca. 4.5 mm wide; involucral bracts ca.
12, subequal, oblong, weakly veined, non-costate,
3.0— 3.5 X 1.2— 1.5 mm, apices rounded, outer
bracts with stipitate glands abaxially. Florets 20 to
25; corollas white, ca. 3.5 mm long, basal tube ca.
1.3 mm long, slender, throat broadly funnelform,
ca. 1 .3 mm long, lobes oblong-ovate, ca. 0.8 mm
long, with few short hairs outside; anther thecae ca.
0.8 mm long; style with hairs restricted to base;
style branches 5—6 times as broad as shaft when
fully expanded. Achenes 2.5— 3.0 mm long, with
basal stipe ca. 0.5 mm long, with distinct setulae;
pappus of 25 to 30 bristles 1.5— 1.8 mm long, bases
contiguous or nearly so.

The small piece from which Neocuatrecasia yun-
gasensis is described is thoroughly distinctive in its
stipitate glands on all stemlike or foliiform parts,
in the distinctly peliolate leaves, and in the broad
corolla throats. The corollas are particularly dis¬
tinct from those of the possible near relative from
the same area, N. feuereri, which has long cylin¬
drical corolla throats much longer than the lobes or
the small anthers.

Neocuatrecasia sandiensis H. Robinson, sp. nov.
TYPE: Peru. Dept. Puno: Prov. Sandia, Ura-
ayllu, trail on W-facing slope above quarry,
I4°08'S, 69°32'W. 3510 m, high elevation
grassland, plant to 40 cm, suffrutescent, flower
disc only, white, 25 Jan. 1986, B. Bennet 1994
(holotype, NY). Figure 1I-L.

A N. hirtella aspectu simili sed in faucibus corollarum
longioribus et in pilis stylorum omnino basilaribus differt.

Small subshrubs to 40 cm tall, with many lateral
branches ascending at ca. 25-30° angles; stems
reddish, densely puberulous to hispidulous with of¬
ten reddish hairs, with interspersed dark to some¬
times reddish glandular dots; internodes 0.4— 2.0
(-6.0) cm long. Leaves opposite; petioles 0.2-1. 0
cm long; blades ovate, 1—2 X 0.4— 1.0 cm, bases
abruptly obtuse, margins with 3 to 5 shallow blunt
teeth, apex short-acute, densely gland-dotted on
both surfaces, densely minutely puberulous adaxi¬
ally, densely puberulous on veins abaxially; triner-
vate with strongly ascending proximal secondary
veins from near base of blade. Inflorescence ter¬
minal on stems and branches, with 1 to 3 pedun¬
culate heads; peduncles to 2.5 cm long, slender,
puberulous with dark hairs and with glandular dots.
Heads broadly campanulate, 8.0— 9.5 mm high; in¬
volucral bracts 12 to 14, broadly oblong, non-cos¬
tate, 5. 5-8.0 X 2.2— 3.0 mm, outer bracts with
rounded tips, densely puberulous with dark hairs
and with pale glandular dots abaxially, inner bracts
paler, mostly glabrous, more scarious and gland-
dotted toward erose ti ps. Elorets 20 to 25 in a head;
corollas white, ca. 6.5 mm long, basal tube ca. 0.8
mm long, throat ca. 5 mm long, cylindrical, scarcely
twice as broad as tube, lobes ca. 0.8 mm long, ob¬
long-ovate; anther thecae ca. 1 .3 mm long; style
with hairs only at base, style branches when fully
expanded 5-6 times as broad as shah. Achenes
3.0— 3.5 mm long, basal stipe ca. 1 mm long, se¬
tulae short and widely spreading; pappus of 29 to
32 bristles, ca. 3.5 mm long, contiguous or nearly
so at base.

Phe corollas of Neocuatrecasia sandiensis are
very similar to those of N. feuereri of Bolivia, with
the long cylindrical throats and the much shorter
anther thecae. The present species differs most ob-

t—

E— II. N. yungasensis II. Robinson, Vargas & Seidel 505 (holotype, US). — E. Corolla with style branches. — 1. Corolla
in longitudinal section with stamens and style. — G. Involucral bract with stipitate glands. — H. Aehene. I— L. N.
sandiensis H. Robinson, Bennet 1994 (holotype, NY). — I. Corolla with tips of style branches. — J. Corolla in longitudinal
section with stamens, style, and nectary. — K. Involucral bract. — L. Aehene. Scale bars all = 1 mm.

392

Novon

viously by the smaller non-velvety leaf blades and
the broader non-pointed involucral bracts.

Paratype. PERU. Dept. Pnno: Sandia, trail on W-
facing slope beginning at slate quarry in Ura-ayllu,
14°08'S, 69°32'W, 3470 m, 22 Feb. 1986, Bennet 2261
(NY).

Acknowledgments. Alice Tangerini is thanked
for the illustrations, and Marjorie Knowles is
thanked lor many details regarding the preparation
of the paper.

Literature Cited

King, R. M. & H. Robinson. 1970. Studies in the Eupa-
torieae (Compositae). XXXII. A new genus Neocuatre-
casia. Phytologia 20: 332-333.

- & - . 1972. Studies in the Eupatorieae (As-

teraeeae). LXXXVI. Additions to the genus Neocuatre-
casia. Phytologia 24: 105—107.

- & - . 1974. Studies in the Eupatorieae (As-

teraceae). CXIX. Additions to the genera Cronquistian-
thus, Helogyne and Neocuatrecasia from Peru. Phytolo¬
gia 27: 395-401.

- & - . 1986. Studies in the Eupatorieae (As-

teraceae). CCXXII. New combinations and new species
from Tropical America. Phytologia 60: 80-86.

- & - . 1987. The Genera of the Eupatorieae

(Astereaceae). Monogr. Syst. Rot. Missouri Rot. Card.

22.

- & - . 1988. Studies in the Eupatorieae (As-

teraceae). CCXXIII. New combinations and new species
of Central and South America. Phytologia 65: 62—70.

Th ree New Species of Piptocarpha (Asteraceae: Vernonieae) from

Ecuador and Peru

Harold Robinson

Botany, National Museum of Natural History, MBC-166, Smithsonian Institution, Box 37012,
Washington, D.C. 20013-7012, U.S.A. Robinson.Harold@nmnh.si.edu

Abstract. Piptocarpha geraldsmithii H. Robin¬
son, P. klugii G. Lom. Smith ex H. Robinson, and
P. vasquezii H. Robinson are described as new from
northern Peru and southern Ecuador. The first re¬
lates to an otherwise strictly Guyana Highland
group of species, and the third has terminal pani¬
cles with uniquely formed urn-shaped heads.

Key words: Asteraceae, Ecuador, Peru. Pipto¬
carpha, Vernonieae.

A series of collections of Asteraceae from north¬
ern Peru, recently sent for identification by the Mis¬
souri Botanical Garden, contained a number of un¬
described species. Three of these species proved to
be in one genus, Piptocarpha R. Brown, with one
species, P. klugii, extending into southern Ecuador.
The specimens could not be determined as any of
the seven species of the genus recognized by Ger¬

ald L. Smith in his treatment of the genus for Peru
(Jones, 1980). The substantial addition to the num¬
ber of species in Peru seems to justify the presen¬
tation of a new key to the ten species of Piptocarpha
now known from the country.

The present study brings to almost 50 the num¬
ber of species in the Neotropical Vernonian genus
Piptocarpha. The genus is distributed from Central
America and the West Indies to southeastern South
America. Most species are scandent or scrambling
with some shrubby species in southern Brazil. The
habitats are montane temperate forests to savannas.

The species treated by Smith (Jones, 1 980) as P.
sprucei Baker is a synonym of P. leprosa (Lessing)
Baker. One of the species validated here has proven
to be the same as one included by Gerald Smith in
his unpublished thesis work. It is validated here as
a Smith species in order to complete the coverage
of the Peruvian species.

Key to the Species of Piptocarpha in Peru and Ecuador

la. Inflorescence a terminal branching panicle.

2a. Heads ca. 8 mm high; inner involucral bracts not broadened distal ly, obtuse to shortly acute; abaxial

surfaces of leaves discolorous, glands obscured by dense scales or hairs . P. gutierrezii Cuatrecasas

2b. Heads ca. 10 mm high; inner involucral bracts broadened distally, apical margins broadly rounded; abaxial

surfaces of leaves concolorous, without obvious hairs or scales, with obvious glandular dots ... 8 vasquezii

H. Robinson, sp. nov.

lb. Inflorescence of heads clustered in glomerules or clumps in axils of full-sized leaves.

3a. Stems strongly angular . P. leprosa (Lessing) Baker

3b. Stems terete or nearly terete.

4a. Heads with 9 to 35 florets.

5a. Branches and undersurfaces of leaves with stalked stellate hairs . . P. asterotrichia (Poeppig) Baker
5b. Branches and undersurfaces of leaves densely covered with small fringed sessile scales.

6a. Axillary groups of heads subumbellate with heads on stout short peduncles 4-8 mm long

. P. lechleri (Schultz Bipontinus) Baker

6b. Axillary groups of heads rather glomerulate with heads borne on short ramified branches

and very short peduncles 1-2 mm long . P opaca (Bcntham) Baker

4b. Heads with up to 7 florets.

7a. Heads with 3 florets; basal appendages of anthers with blunt tips . . P. geraldsmithii H. Robinson,

sp. nov.

7b. Heads with 6 or 7 florets; basal appendages of anthers with sharp points.

8a. Pappus with slender bristles darkened toward tips; heads sessile in dense glomerules .

. P. klugii G. Lom. Smith ex H. Robinson, sp. nov.

8b. Pappus white; heads with distinct peduncles or branched stalks.

9a. Lower leaf surfaces with appressed hairs or scales; stems and branches with mostly

appressed minute tomentum . P poeppigiana (DC.) Baker

9b. Lower leaf surfaces with stalked stellate hairs; stems and branches with short but coarse

spreading fascicles of hairs . P. canescens Gleason

Novon 12: 393-398. 2002.

394

Novon

The four species presently known for Ecuador
are all included in the above key, Piptocarpha gu-
tierrezii, P. lechleri, P. klugii, and P. poeppigiana.
The three new species of Piptocarpha are as fol¬
lows. All diree possess the extremely deciduous in-
volueral bracts, nodes at the bases of the styles,
and the indurated or sclerified tails of the anthers
that are characteristic of the genus. All have the
spinulose tricolporate type A pollen that is char¬
acteristic of the subtribe Piptocarphinae. All have
corolla lobes coiling or curling backward at anthe-
sis, a characteristic found in the Vernonieae most
commonly in the subtribe Piptocarphinae.

Piptocarpha gcraldsmilliii 11. Robinson, sp. nov.
TYPE: Peru. Amazonas: Bagua Prov., Distrito
Imaza, comunidad Aguaruna de Wanas, km 92
carretera Bagua— Imacita, Cherros Chinim,
hordes quebrada rocosa, ambas marge ties,
700-800 m, 29 Aug. 1996, C. Diaz , Pena, Tsa-
majain & Roca 8065 (holotype, US; isotype,
MO). Figure 1.

In habitu et in capitulis trifloribus et in caudis anther-
arum apice Iruncatis et in superficiebus adaxialibus fo-
liorum reticulate prominule venulosis P. auyantepuiensi
similis sed in caulibus rectioribus et in acheniis glabrio-
ribus distincta.

Vines or scandent shrubs to 18 m high, with
long, nearly straight, terete, brown stems and
branches spreading at 90° angles, branches very
slightly deflected at nodes, surfaces covered with
many small fimbriate-margined scales, sparse or
abraded on larger stems. Leaves alternate, petioles
slender, 0. 7-1.0 cm long; blades stiffly chartaceous,
elliptical to broadly elliptical, 4.0— 7.5 X 1.6— 2.5
cm, base acute, margins entire, apex acute to
slightly acuminate, extreme tip blunt, adaxial sur¬
face glabrous, alveolate with close reticulum of
prominulous veinlets, abaxial surface concolorous,
sparsely lepidote with minute scales and with dark
glandular punctations; venation pinnate, with 4 to
7 secondary veins on each side, spreading at ca.
60° angles. Inflorescences glomerulate, in axils of
leaves, each cluster with 8 to 10 sessile heads.
Heads ca. 1 cm high; involucral bracts ca. 15, im¬
bricate in 4—5 series, ovate to narrowly elliptical,
1.5— 5.0 X 1.0— 1.5 mm, mostly glabrous abaxially,
tomentellous at tip. Florets 3 in a head; corollas
white, ca. 7 mm long, mostly glabrous, with few
glandular dots on distal part of tube, tube ca. 4 mm
long, narrow in basal 2 mm, broadened and fun-
nelform in distal 2 mm, throat essentially lacking,
with filaments inserted just below sinuses, lobes
linear, 3.0— 3.5 X 0.7— 0.9 mm, anther thecae ca.
2.5 mm long, including basal tails ca. 0.7 mm long.

blunt at tip; apical appendage small, ca. 0.15 mm
long and wide, glabrous. Achenes 3.5— 4.0 mm long,
mostly glabrous, with few hairs distally above
sharpest 2 angles of achene; pappus white, 5.5— 6.0
mm long, of ca. 60 bristles, slightly broadened dis¬
tally, with outer series of shorter bristles 1—2 mm
long. Pollen grains ca. 37 gm diam.

Piptocarpha geraldsmithii was provisionally
identified as P. opaca (Bentham) Baker on the basis
of the 1994 collection. True relationship is closer
to two Venezuelan species, P. auyantepuiensis Ar-
isteguieta (Steyermark, 1967) and P. jauaensis Ar-
isteguieta & Steyermark (Steyermark & Brewer-
Carias, 1976), endemic to the Guayana Highland
region. The new species seems rather individually
distinct from the two relatives by the comparatively
straight stems. The related species are more de¬
flected at the nodes, especially in the branches.
The new species is more like P. jauaensis in the
lack of numerous hairs at the top of the achene just
below the callus and pappus. Hairs are present,
however, restricted to the areas over the lateral
achene margins, a situation that proves to be true
of the type of P. jauaensis also, contrary to the orig¬
inal 1976 description. The new species is most like
P. auyantepuiensis in the adaxial surfaces of the
leaves with a close reticulum of prominulous vein-
lets. The adaxial surface of the leaves of P. jauaen¬
sis has a more hardened and smooth appearance
without prominulous veinlets. Both P. auyante¬
puiensis and P. jauaensis are described with young
stems densely covered with tomentose arachnoid
hairs or arachnoid scales, but the type specimens
show the same lepidote condition with margins of
the scales shortly fimbriate as seen in the present
new species. A note provided by Gerald Smith
states that the species and its Venezuelan relatives
are in Piptocarpha subg. Hypericoides.

The new species is named after Gerald L. Smith,
student of the genus Piptocarpha who contributed
the treatment cited here from Peru (Jones, 1980).
Smith has published notes and new taxa in the ge¬
nus in two additional papers (Smith, 1981, 1982).
Unfortunately, the Smith dissertation on Piptocar¬
pha remains unpublished.

Paratype. PERU. Amazonas: Distrito Imaza, Region
Nororiental del Maranon, comunidad de Kampaenza, ri-
bera de la quebrada Shimutaz, Rfo Maranon, 04°55'S,
78°19'W, 320 m, 24 Sep. 1994, N. Jaramillo, Apanu &
Katip 498 (MO, US).

Piptocarpha klugii G. Lorn. Smith ex II. Robin¬
son, sp. nov. TYPE: Peru. Loreto: Pumayacu,
between Balsapuerto and Moyobamba, 600—
1200 m. Aug.— Sep. 1933, Klug 8167 (holo¬
type, GA; isotype, US).

centimeters

Volume 12, Number 3
2002

Robinson

Piptocarpha from Ecuador and Peru

395

UNITED STATES NATIONAL HERBARIUM

00642060

Camilo Dia2. Antonio Pena,

Luis Tsamajain y Melanio Roca 8065

Proyecto auspiciado por la John D. and
Catherine T. MacArthur Foundation
MISSOURI BOTANICAL GARDEN HERBARIUM (MO)

Figure 1. Piptocarpha geraldsmithii H. Robinson, sp. nov. Holotype, Peru: Amazonas, Diaz et al. 8065 (US).

396

Novon

A speciebus pluribus in floribus ca. 7 in capitulo et in
setis pappi fulvescentibus et in eaudis antherarum argute
aeutis distincta.

Scandent shrubs or vines, witli sparse branching
at ca. 80° angles; stems brown, terete, with slight
deflection at nodes on stems and branches, surfaces
lepidote with small scales, sparse on larger stems,
dense on branches. Leaves alternate, petioles 0.7—
0.9 cm long; blades stiffly chartaceous, oblong-el¬
liptical, mostly 6—13 X 2.0— 4.5 cm, base acute,
margins entire, apex short-acuminate with extreme
t i p narrowly obtuse, adaxial surface rather lustrous,
with sparse minute scales, with elongate areolation
formed by prominulous veinlets, abaxial surface ei¬
ther concolorous with sparse minute scales or pale
with dense cover of scales; venation pinnate, with
ca. 6 secondary veins on each side, spreading at
ca. 75° angles, arching upward nearer margins. In¬
florescences glomerulate in axils of leaves; clusters
of ca. 20 heads on short branching stalks 2-3 mm
long. Heads ca. 7 mm high; involucral bracts ca.
15, imbricated in 4—5 series, ovate to broadly ob¬
long-elliptical, 1—4 X 1.5— 2.0 mm, lips obtuse to
rounded, glabrous abaxially, darkened toward tips.
Florets ca. 7 in a head; corollas white or greenish
white, 5.5— 6.0 mm long, basal tube slender in basal
1.5— 2.0 mm, broadened and funnelform in distal
1.5 mm, glabrous below, with scattered short-
stalked capitate glands on distal part, throat essen¬
tially lacking, with filaments inserted just below si¬
nuses, lobes linear, ca. 2.5 X ca. 0.7 mm; anther
thecae 2.3— 2.5 mm long, including pale sharply
pointed basal tails ca. 0.5 mm long; apical ap¬
pendage pale, ca. 0.7 mm long, broadly oblong-
ovate, glabrous. Achenes ca. 3 mm long, glabrous,
with 2 sharper lateral angles, with 3 weak costae
outside, 2 inside; pappus ca. 5.5 mm long, of ca.
50 bristles, very slender and pale below, broadened
and brownish distally, without an evident shorter
outer series. Pollen grains ca. 37 gm diam.

Piptocarpha klugii is one of the Andean species
with axillary clusters of heads, pointed anther tails,
and the lepidote rather than stellate pubescence,
but it is distinct by the slender pappus bristles that
darken distally. Except for the pale and densely
lepidote abaxial surfaces of the leaves of Diaz &
Pena 8754, the four specimens included here are
similar in every' respect, and they are treated as
conspecific.

At the time that the description was prepared,
there was some suspicion the unpublished name P.
klugii of Gerald Smith might apply. This has sub¬
sequently been confirmed by Smith, who agrees to
have it validated here. The Smith name has pre¬

viously been used on annotations and it appears in
print as a nomen nudum in the Catalogue of the
Flowering Plants and Gymnosperms of Peru (Brako
& Zarruchi, 1993).

The habitat is consistently cited as primary for¬
est. In one form it was cited from “bosque primario
sobre pendientes de 30%, arboles cubiertos de
musgos, suelos lentizoles.”

Paratypes. ECUADOR. Morona-Santiago: Guala-
i|uiza Canton, Cordillera del Condor, ridge top above Ban¬
deras, near disputed Ecuador— Peru border, 03°28'S,
78°15'W, 1350 m, 17 July 1993, A. Gentry 79959 (MO,
US). PERU. Amazonas: Euya Prov., Camporedondo, Tul-
lanya, base Cerro Huiesocunga, La Laguna, 06°06'33"S,
78°20'55"W, 2080 m, 1 Dec. 1996, C. Diaz A Pena 8754
(MO, US) (form with leaves discolorous and densely lep¬
idote below); Bagua Prov., Distrito Imaza, Tayu Mujaji,
eomunidad de Wawas, 05°15,25”S, 78°21,41,,W, 800 in.
23 Oct. 1997, A'. Rojas. Pena. Anag & Yagkuag 408 (MO,
US).

Piptocnrpliu vasquezii H. Robinson, sp. nov.
TYPE: Peru. Amazonas: Bagua Prov., Distrito
Imaza, eomunidad de Yamayakat, ()5°03'24"S,
78°20'17"W, 600 m, 9 July 1997. I\. Vdsquez,
Pena & Chavez 23981 (holotype, US; isotype,
MO). Figure 2.

A speciebus pluribus in inflorescentiis paniculatis et in
capitulis majoribus urceolatis in bracteis interioribus
apice late rotundatis et in appendicibus antherarum glan-
duliferis differt.

Coarse scandent shrubs or vines, with branches
spreading at 55-75° angles; stems and branches
brown, terete, closely shallowly striated, covered
with dense granular pubescence of nearly sessile,
peltately attached, shallowly stel lately lobed cells.
Leaves alternate, petioles 1—2 cm long; blades
broadly elliptical to obovate, mostly 12—17 X 4—9
cm, base broadly acute, usually somewhat unequal,
margins entire, apex shortly and sharply acuminate,
surfaces concolorous, sparsely pitted or lepidote
with minute scales or stellately lobed cells, adaxial
surface shinier than abaxial surface; venation pin¬
nate, with 8 or 9 secondary veins on each side,
spreading at 65—75° angles. Inflorescences terminal
panic les with paniculate branches, branchlets in
corymbiform clusters of few' heads; peduncles 2-6
mm long, w ith densely granular pubescence. Heads
10—1 I mm high; involucre cylindrical below, slight¬
ly constricted distally, terminating with somewhat
spreading expanded tips of longest bracts; bracts in
ca. 7 imbricated series, ovate to oblong, 1.5— 7.0 X
1.5— 2.0 mm, with granular pubescence on lowest
bracts and on distal dark areas of median bracts,
inner longest bracts glabrous or nearly glabrous,
castaneous, distally scarious with broadly rounded

Volume 12, Number 3
2002

Robinson

Piptocarpha from Ecuador and Peru

397

R . VAsquez ,

A. Pena £. E. Chavez . 239B1

MISSOURI BOTANICAL GARDEN HERBARIUM (MO)

Figure 2. Piptocarpha vasquezii H. Robinson, sp. nov. Holotype, Peru: Amazonas, Vdsquez et al. 23981 (US).

Distrito Imaza, Comunidad de
Yamayakat. Bosque primario.

05°03 ' 2*+ "S 78°20 ' 17 "W 600 m

3402050

398

Novon

apices. Florets 6 or 7 in a head; corollas white, ca.
7 mm long, mostly glabrous, with few glandular
dots on upper tube and toward tips of lobes, basal
tube ca. 3.5 mm long, lower narrow part ca. 1.5
mm long, upper 2 mm broadened, funnelform,
throat ca. 0.8 mm long, between insertion of stout
filaments and sinuses, with many transverse un¬
dulations, lobes linear, ca. 3 mm long, 0.5 mm
wide; anther thecae 3.3-3. 5 mm long, including
basal whitish, sclerified, sharply pointed tail ca. 0.5
mm long; apical appendages pale, triangular-ovate,
ca. 0.5 mm long, with a number of stalked glands
outside near base and middle. Achenes ca. 3 mm
long, cylindrical when immature, becoming some¬
what obcompressed, with many glandular dots when
young; pappus white, ca. 6 mm long, with ca. 80
slender bristles, broadened distally, with outer se¬
ries of some shorter bristles or narrow scales ca.
0.8 mm long. Pollen grains ca. 40 pm diam.

Piptocarpha vasquezii is a remarkably distinctive
new species presently known only from the two col¬
lections from primary forest in the Imaza District
of Amazonas, Peru. The species belongs to the
group with heads in terminal panicles, but the
heads are much larger than other such species and
the involucral bracts form a vase-like structure con¬
stricted above and then broadened apieally with en¬
larged rounded tips on the longer bracts. The apical
appendages of the anthers have a number of glands,
a characteristic found in many members of the sub¬
tribe Vernoniinae, but very rare in the Piptocar-
phinae. Single glands have been seen previously on
anther thecae beside apical appendages in one Ve¬
nezuelan specimen of Piptocarpha triflora (Aublet)
Bennett ex Baker ( Wurdack & Adderley 43490, US).

A wider sample may show that the characteristic is
more common. The new species is distinct among
those named here by the insertion of the anther
filaments ca. 0.8 mm below the bases of the corolla
sinuses instead of immediately below the sinuses.
The area between, the throat, has marked trans¬
verse undulations.

Gerald L. Smith has examined the type of the
species and places it in Piptocarpha subg. Ooce-
phalus.

Paratype. PERU. Bagua I'rov., Distrito Imaza, Yama-
yakat. 05°03'20"S, 78°20'23"W, 350 m, 7 June 1996, R.
Vdsquez & A. Vasquez 21064 (MO, US).

Acknowledgments. Susan Hunter is thanked for
the preparation of the scans of the types, Katherine
Rankin is thanked especially for her perseverance
in mounting the specimens of Piptocarpha vasquezii
with their extremely fragile heads, and Marjorie
Knowles is thanked for many other details regard¬
ing the preparation of the paper.

Literature Cited

Brako, I,. & J. L. Zarruchi. 1993. Catalogue of the f low¬
ering Plants and Gymnosperms of Peru. Monogr. Syst.
Bot. Missouri Bot. Card. 45.

Jones, S. B. 1980. Vernonieae. In: J. F. Macbride and
collaborators. Flora of Peru, family Compositae: Part 1.
Fieldiana, Bot. n.s. 5: 22—73.

Smith, G. F. 1981. New taxa in Piptocarpha R. Br. (Ver¬
nonieae: Compositae). Ann. Missouri Bot. Card. 68:
661-667.

- . 1982. Taxonomic considerations of Piptocarpha

(Compositae: Vernonieae) and new taxa in Brazil. Brit-
tonia 34: 210-218.

Steyermark, J. A. 1967. Flora del Auyan-Tepui. Acta Bot.
Venez. 2: 5—370.

- & C. Brewer-Carias. 1976. Fa vegetacidn de la

Cima del Macizo de Jaua. Bol. Soc. Venez. Ci. Nat.
22(132-133): 179-405.

New Taxa and a New Combination in Tidestromia (Amaranthaceae)

from North America

Ivonne Sanchez-del Pino and Hilda Flores Olvera
Departamento de Botanica, Instituto de Biologfa, UNAM, Apartado Postal 70-233, 04510
Mexico, D.F. Mexico. mahilda@servidor.unam.mx

ABSTRACT. A new species and a newT subspecies
of Tidestromia are described and illustrated. Tides¬
tromia valdesiana, restricted to Coahuila, Mexico,
is characterized by the absence of an involucre and
tepals lanceolate, aristate at the apex with a prom¬
inent midnerve and subulate trichomes. Tidestrom¬
ia lanuginosa subsp. eliassoniana is characterized
by having trichomes with the terminal cell projec¬
tions either irregular (as those of the subsp. lanu¬
ginosa) or spreading, and microspinulose pollen,
which is unique within (lie genus. A distribution
map for the two recognized subspecies of T. lanu¬
ginosa is provided. The new combination 7. suffru-
ticosa var. oblongifolia is proposed, and T. oblon-
gifolia subsp. cryptantha is reduced to synonymy
under the former; both names are lectotypified.

Resumen. Se describen e ilustran una especie y
una subespecie nuevas de Tidestromia. Tidestromia
valdesiana, endemica de Coahuila, Mexico, se ea-
racteriza por los tepalos lanceolados de apice aris-
tado con la vena media prominente y tricomas su-
bulados, asf como por no tener involucro.
Tidestromia lanuginosa subsp. eliassoniana se ca-
racteriza por tener tricomas con proyecciones de las
celulas terminales irregulares (como en la subsp.
lanuginosa) o patentes, asi como polen microespi-
nuloso que es unieo en el genero. Se incluye un
mapa de distribucion de las dos subespecies re-
conocidas de T. lanuginosa. Ademas se propone la
combi nacion nueva T. stiff rut icosa var. oblongifolia,
redueiendose como sinonimo T. oblongifolia subsp.
cryptantha ; ambos nombres son lectotipifieados.

Key words: Amaranthaceae, Froelichiinae,
Gomphreneae, North America, pollen, Tidestromia.

Tidestromia Standley is a small genus of six spe¬
cies in the Amaranthaceae, subfamily Gomphre-
noideae, tribe Gomphreneae, subtribe Froelichiinae
(Schinz, 1893), occurring in North American de¬
serts (Standley, 1916; Robertson, 1981; Eliasson,
1988; Henrickson, 1993), frequently on salty, gyp¬
seous, and calcareous soils. Tidestromia has been
circumscribed by having the flowers arranged in
small axillary glomerules and leaves that subtend

the inflorescence becoming hard and more or less
connate in age, thus forming an involucre (Standley,
1916; Eliasson, 1988). The pollen in Tidestromia
has been considered unique among Amaranthaceae
(Eliasson, 1988; Borsch, 1998). It differs from that
of other genera by being melareticulate, with me-
soporia continuously narrowed distallv (triangular
in cross section), tectate, with perforations concen¬
trated close to the apertures (Borsch, 1998). Al¬
though psilate pollen has been considered diagnos¬
tic of Tidestromia, Martin and Drew (1970)
described ornamented pollen of T. lanuginosa (Nut-
tall) Standley, which was never taken into consid¬
eration for the taxonomy of the genus.

Tidestromia has trichomes with a single row of
cells (uniseriate) with the upper cells usually with
sinuous interlocking junctions, thick cellulose
walls, and usually with projections. As was indi¬
cated by Carolin (1983), one subtype of this uni¬
seriate trichome has projections, if they occur, ap¬
parently at random, and the other subtype, which
we are calling candelabriform in this paper, is char¬
acterized by large projections arranged in a whorl
in the center of the cells. In Tidestromia, while the
size of the projections is characteristic of a partic¬
ular trichome, there are trichomes whose cells have
projections of differing size. We found also notable
differences in the apical cells of the trichomes.

In an ongoing revision of Tidestromia, observa¬
tions on the habit, the nature of the pubescence,
characters of the trichomes, tepals anti involucre,
as well as pollen micromorphology led us to rec¬
ognize an undescribed species and to propose a
new subspecies of T. lanuginosa. We also propose
a new combination for T. sujfruticosa (Torrey)
Standley, and we designate lectotypes for two
names.

Tidestromia valdesiana Sanchez-del Pino X Flo¬
res Olvera, sp. nov. TA PE: Mexico. Coahuila:
Mpio. Parras, ejido el Durazno, xerophytic
scrub on limestone mesa, 25°27'20"N,

1 02°06'36"W, 1635 m, 15 Oct. 2000, /. Sdn-
chez-del Pino 93, H. Flores Olvera & C. Go¬
mez- Hinostrosa (holotype, MEXU; isotypes,
ANSM. MO, NY. TEX).” Figures 1 , 2 A, 3 A. 3C.

Novon 12: 399-407. 2002.

400

Novon

Figure 1. Tidestromia valdesiana Sanchez-del Pino & Flores Olvera. — A. Habit. — B. Flowering branch. — C. Leaf
and involucral leaves surrounding the inflorescence. — I). Inner tepals, abaxial and lateral views. — E. Outer tepals,
abaxial view. — F. Bract. — G. Bracteoles. — H. Sexual verticils. — I. Pistil. — J. Androecium. — K. I.eaf and involucral
leaves in fruit. — L. Inflorescence. — M. Utricle. — N. Seed. Drawn from the holotype ( Sdnchez-del Pino 93 et a /.,
MEXU).

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Sanchez-del Pino & Flores Olvera
Tidestromia from North America

401

Herba perennis rhizomatosa, caulibus ascendentibus
vel decumbentibus, plerumque base gemmis lanatis,
dense lanuginosa, trichomatibus omnino vel partim can-
delabriformibus; inflorescentia plerumque per duo folia
involucralia inclusa, his ad maturitatem non induratis nee
connatis, involucrum non formantibus, vel raro inflores¬
centia foliis involucralibus carens ad nodos ubi duo vel
plures rami orientes; tepala lanceolata, apice aristata, tri¬
chomatibus barbatis subulatisque, nervo mediano cons-
picuo.

Herbaceous perennial from rhizome, up to 22 cm
tall, canescent, richly lanuginose, trichomes com¬
pletely or partially candelabriform. Stems grav-
green or red, ascending or decumbent, usually
bearing wooly to conspicuously wooly gemmae at
the base. Leaves alternate, gray-green, sometimes
red-shaded, ovate to widely ovate, 0.8— 2.2 X 0.7—
2.0 cm. thick, veins prominent beneath: apex acute
or obtuse, base cuneate, cordate or attenuate; pet¬
ioles up to I mm long. Inflorescence reduced to a
few, mostly 3 flowers; surrounded by two involucral
leaves that do not become indurate or connate with
age, thus not forming an involucre, or rarely invo¬
lucral leaves absent; involucral leaves gray-green,
sometimes red, ovate, broadly ovate, or lanceolate,
0. 1-0.6 X 0. 1-0.7 mm, thick, veins prominent be¬
neath, apex acute or obtuse, base cuneate or atten¬
uate; petiole up to 0.3 cm long, densely lanuginose.
Flowers 4. 5-5.5 mm long; bracts widely depressed
ovate, 2.5 X 1 .5 mm, lanuginose above to glabrous,
apex obtuse or rounded; braoteoles ovate, 1.9— 2.2
X 1.2— 1.5 mm, lanuginose above to glabrous, apex
acute; tepals yellow-tinged, lanceolate, 4-4.5 X I —
2 mm, coriaceous, apex aristate, midrib prominent,
thicker at the base, densely lanuginose; trichomes
subulate, barbed; staminal cup 0.6— 0.9 mm long,
filaments 1 mm long, anthers 0.8-1. 2 mm long,
pseudostaminodia short or absent; ovary 0.8— 1.2
mm long and wide, style 2-3 mm long, stigma bifid,
deltoid, dark brown, 0.3— 0.4 mm long. Utricle 2.1
X 1.4— 1.6 mm. Seeds red-brown, 1.5— 1.7 X 1.4
mm. Pollen psilate (Fig. 2A).

Distribution and habitat. This is a narrowly en¬
demic species of the Chihuahuan Desert Region,
known only from the type locality in Coahuila, Mex¬
ico, at 1635—1700 m. It is locally common restrict¬
ed to calcareous sites. Tidestromia valdesiana is re¬
stricted to the limestone mesa where it grows with
Agave lechuguilla Torrey, Dasylirion sp., Fouquie-
ria sp., and Larrea sp. Flowers have been observed
from September to November. Tidestromia suffruti-
cosa is sympatric with T. valdesiana.

Etymology. Tidestromia valdesiana is named in
honor of Javier Valdes Gutierrez, w hose support and
enthusiasm motivated our interest in the study of
the xerophytic flora of Mexico.

Discussion. Tidestromia valdesiana is the only
species in the genus with lanceolate tepals having
an aristate apex, prominent midnerve, and subulate
trichomes (fig. 3A). These trichomes have small
projections mostly reflexed with a barbed appear¬
ance. I his last character is also present in T. tenella
I. M. Johnston, restricted to Coahuila, Mexico, but
its trichomes are round-tipped (Fig. 3B). Tidestro¬
mia valdesiana is unique in the genus in that the
two leaves surrounding the inflorescence do not be¬
come indurate and connate in age and do not form
an involucre. An extreme condition in this species
is the absence of leaves surrounding some inflores¬
cences borne at nodes where two or more branches
originate (Fig. 1L). Tidestromia valdesiana is simi¬
lar to T. gemmata 1. M. Johnston, which is consid¬
ered nonspecific with T. suffruticosa (Torrey) Stand-
ley (Sdnehez-del Pino, 2001). Although both
species are perennial herbs, usually with gemmae
at the base, the gemmae densely wooly, and hirsute
vestiture due to the abundance of trichomes par¬
tially candelabriform with the terminal cells lacking
projections (Fig. 3C). they differ by other vegetative
and reproductive characters. Tidestromia suffruti¬
cosa is suffrutescent, without a rhizome; it has ovate
tepals with an acute or obtuse apex, midnerve not
prominent, in addition to completely or partially
candelabriform trichomes (Fig. 3D), and with pro¬
jections at random (Fig. 3E). The inflorescence in
this species is always surrounded by two involucral
leaves that become indurate and connate with age
forming an involucre.

Paratypes. MEXICO. Coahuila: Mpio. Parras, Ejido
Mesa del Durazno, Capd 1978 (ANSM), Capo 1979
(ANSM), /. Sdnchez-del Pino 94 et al. (CAS, MEXU,
TEX).

Tidestromia lanuginosa subsp. eliassoniana

Sanchez-del Pino & Flores Olvera, subsp. nov.
TYPE: Mexico. Sonora: Mpio. Guaymas, 15
km N of Guaymas on the road to Hermosillo,

1 Nov. 1985, B. Bartholomew 3654, L. R. Lan¬
drum , //. W. Li & T. S. Ying (holotype, MEXU;
isotypes, ASU, CAS, GH). Figures 2B, 2C,
3G— I, 4, 5.

A subspecie lanuginosa differt pollinis granis exina mi-
crospinulosa, cellula terminali trichomatum caulium, fo-
liorum et foliorum involucralium duarum classium: com-
muni, irregulariter ramificanti, et atypica, ramis
patentibus.

Stems light brown to gray-green, generally red;
stems, leaves, and involucral leaves covered by
completely candelabriform trichomes, with the ter¬
minal cell projections either irregular, as those of
the subspecies lanuginosa, or spreading. Leaves

402

Novon

Figure 2. SEM photomicrographs of pollen grains. — A. Tidestromia valdesiana (Capo 1978, ANSM). — B, C. Tides-
tromia lanuginosa subsp. eliassoniana ( Bohrer 1189, AKIZ). — I). Tidestromia suffruticosa var. suffruticosa (Sdnchez-
del Pino 77 el al., MEXU).

widely obovate or generally ovate-trullate or lan¬
ceolate; apex acute or rarely obtuse. Pollen mi-
crospinulose (Fig. 211, C).

Distribution and habitat. Tidestromia lanugi¬
nosa subsp. eliassoniana is known from the south¬
western United States in Utah, California, Arizona,
and New Mexico, to northwestern Mexico in Sonora,
Chihuahua, Baja California Sue, and Sinaloa (Fig.
5). It occurs in primary and disturbed vegetation in
grassland, xerophytic scrub, coastal thorn scrub,
thorn forest, and tropical deciduous forest, dwelling
on roadsides or in cultivated fields, from sea level

to 1463 m. The soils throughout its distributional
range are alluvial, sandy to clayey, gravelly, lime¬
stone, silty, igneous, salty, or gypseous.

Etymology. Tidestromia lanuginosa subsp.
eliassoniana is named in honor of Uno H. Eliasson
in recognition of his contributions to the taxonomy
of Amaranthaceae, including pollen morphology.

Discussion. Tidestromia lanuginosa is the most
widely distributed species of the genus. It is char¬
acterized by having, on the secondary branches, an
involucre formed by involucral leaves in which the
petiole becomes indurate with age and connate with

Figure 3. SEM photomicrographs of trichomes. — A. Subulate barbed trichome ol Tidestromia valdesiana (Capo 1978,
ANSM). — B. Bound-tipped barbed trichome of T. tenella ( Villarreal 4459 et al., ANSM). — C. Partially candelabriform
trichome of T. valdesiana ( Capo 1978, ANSM). — D. Partially candelabriform trichome of T. suffruticosa ( Chiang 9277,
NY). — E. Trichome with projections at random of T. suffruticosa ( Johnston 10674, MEXU). — F. Trichome with the
terminal cell projections irregular of T. lanuginosa subsp. lanuginosa ( Ekman 9861, GH). — G. Trichome with the
terminal cell projections spreading of T. lanuginosa subsp. eliassoniana ( Bartholomew 8654 et al., CAS). — H. Unis-
eriate macroform osteolate conical trichome of T. lanuginosa subsp. eliassoniana (Clark 8499, A BIZ). — 1. Uniseriate
macroform osteolate T-shaped trichome of T. lanuginosa subsp. eliassoniana (Clark 8499, A BIZ).

Volume 12, Number 3
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Tidestromia from North America

403

404

Novon

and lateral views. — F. Bract. — G. Bracteoles. — H. Androecium. — I. Pistil. — J. Leaf and involucral leaves in fruit.
B— J, drawn from the holotype ( Bartholomew 3654 et «/., MEXU).

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Sanchez-del Pino & Flores Olvera
Tidestromia from North America

405

the stem or with the leaf petiole and the stem. This
is an annual speeies, with leaves and involucral
leaves chartaceous or less commonly fleshy, stems
usually gray-green or red to yellow-green in the
field, usually richly lanuginose to glabrate, and
completely candelabriform trichomes. The tepals
present this type of trichome as well as trichomes
with small projections that occur in some cells. Due
to the wide morphological variation of the species,
some authors (Steyermark, 1932; Cory, 1936; Cor-
rell & Johnston, 1970; Robertson, 1981) recognized
two varieties: T. lanuginosa var. lanuginosa and va¬
riety carnosa (Steyermark) Cory. Nevertheless, we
agree with Johnston (1943), who recognized T. car¬
nosa as a distinctive species. This species has the
involucre on the secondary branches formed by the
connation of involucral leaf bases with or without
the leaf base, but never with the stem. In addition,
the young parts and the tips of the leaves are gla¬
brous or slightly pubescent; the stems, leaves, and
involucral leaves are fleshy and yellow-green in the
field.

Tidestromia lanuginosa is herein proposed to he
separated into two subspecies on the basis of pollen
and trichome features. Tidestromia lanuginosa
subsp. lanuginosa has the characteristic psilale
pollen of the genus as described by previous au¬
thors (Eliasson, 1988; Borsch, 1998), whereas T.
lanuginosa subsp. eliassoniana is characterized by
the occurrence of microspines on the tectum (Fig.
2B, C), which is unique in the genus but not in the
tribe. Tidestromia lanuginosa subsp. eliassoniana
also differs from the other subspecies by having
stems, leaves, and involucral leaves with trichomes
that have the terminal cell projections either irreg¬
ular, as in those of the subspecies lanuginosa (Fig.
3F), or spreading (Fig. 3G). The collection Clark
8499 (ARIZ) from Arizona has a different type of
trichome with cuticle granules and without projec¬
tions that were described by Reddy et al. (1998) as
uniseriate macroform osteolate. Among these, some
are conical (Fig. 3H) and others are T-shaped (Fig.
31), resembling trichomes sometimes present in T.
carnosa and T. tenella. Because this specimen also
has microspin ulose pollen, the trichomes are con¬
sidered an extreme variation for subspecies elias¬
soniana.

Based on the known morphological variation of
pollen present within genera of the Amaranthaceae,
the microspinulose pollen in T. lanuginosa subsp.
eliassoniana, unique within the genus, is of high
significance. Preliminary studies indicate that there
might be intermediate stages between psilate and
microspinulose pollen within Tidestromia. We ob¬
served parts of the tectum with nanoverrucae of

some samples of T. sujfruticosa var. sujfruticosa
(Fig. 2D), suggesting the need to do exhaustive pal-
ynological studies in the genus. Shifts between
completely smooth to sculptured tecta are extreme¬
ly rare within genera of Amaranthaceae and as cur¬
rently known otherwise only occur in the genus
Cyathula (Borsch, 1998).

Because the pollen of all other genera of Gomph-
reneae is generally microspinulose, it would appear
that this is a plesiomorphic condition within the
genus Tidestromia, but phylogenetic studies are
needed to test this. Pollination studies are also
needed to test whether the sculptured pollen is re¬
lated to a shift in pollen dispersal mechanisms po¬
tentially related to an increase in the amount of
pollen kit.

Tidestromia lanuginosa subsp. lanuginosa oc¬
curs from the central United States to northern
Mexico and the West Indies (Antilles), from Illinois
to the Dominican Republic, whereas subspecies
eliassoniana ranges from the southwestern United
States to northwestern Mexico (Sanchez-del Pino,
2001) (Fig. 5).

Paratypes. U.S.A. California: Riverside Co., 21 mi.
W of Tonopah along I- 10, Sep. 1991. Atwood 1 7084 (NY).
Utah: Washington Co., 3 mi. S of Saint George, Sep.
1941, Gould Mid (ARIZ, DS, GH. NY. UC). Arizona:
Cochise Co., Escala San Simon Valley, Aug. 1912, Gooding
1290 (ARIZ, NY); Coconino Co., along Grand Canyon,
airport road, 100 yards W of junction with Arizona 64-
180, ca. 8 mi. S of Grand Canyon Village, Sep. 1981,
Sauleda 6495 & Sauleda (ASU); Gila Co., near Canyon
of the Sally may. Sierra Ancha foothills, Sep. 1946, Gould
3883 (ARIZ, NY, UC); Graham Co., hwy. 666. 0.2 mi. E
of junction with hwy. 70, Sep. 1968, Pinkavu 13425 et al.
(ASU, NY); Greenlee Co., along U.S. hwy. 70. 8 6/10 mi.
E of Solmonville on road to Duncan, Oct. 1942, Wolf
11414 & Everett (UC); Maricopa Co., Tempe, on rocky
slopes of Double Butte, Sep. 1964, Fryxell 101 (ENCB,
NY); Mohave Co., E of Blythe, Sep. 1929. Jones 24819
(CAS, GH, MO, NY); Pima Co., Tucson, Oct. 1894, Tou¬
rney 1894 (ARIZ, CAS, GH. NY. UC); Pinal Co., along
hwy. 10 between Phoenix and Tucson. Sacaton rest stop,
Oct. 1982, Neese 12529 (ASU, NY); Santa Cruz Co., be¬
tween Sopori Scool and Arivaca, July 1966, Tate 278
(ASU); Yuma Co., SE of Alamo Lake, May 1980, Butter-
wick 6322 & Hillyard (ARIZ, ASU). New Mexico: Hi¬
dalgo Co., 1 mi. N of Rodeo, Aug. 1966, Cazier 407
(ASU). MEXICO. Baja California Sur: Mpio. MulegG
Bajada 6 mi. S\\ of Mulege, Oct. 1962, Wiggins 17964
& Wiggins (DS, US). Sonora: Mpio. Agua Prieta, Colonia
Morelos, region of the Rio de Bavispe, Aug. 1941, White
4145 (ARIZ. GH. NY): Mpio. Alamos, near Tepistate N
of Alamos, Nov. 1939, Gentry 4838 (ARIZ, NY); Mpio.
Altar, Hacienda Oquitoa (Cutting’s Ranch) 6 mi. E of Al¬
tar, Oct. 1932, Wiggins 5965 (DS, US); Mpio. Arizpe,
across the Rio Bacanuchi from Tauichopa, Tauichopa is
2.7 mi. by road N of Arizpe, Aug. 1958, Turner 183 &
Lowe (ARIZ); Mpio. Bacoachi, along wash about 14 mi. S
of Divisaderos, Sep. 1934, Wiggins 7474 (DS, GH, US);
Mpio. Benjamin Hill, Santa Ana km 145 Hermosillo— San-

406

Novon

Figure 5. Distribution of Tidestromia lanuginosa subsp. lanuginosa (line outlining range) and T. lanuginosa subsp.
eliassoniana (•).

ta Ana, 13.7 mi. (21.9 km) N of Benjamin Hill, Aug.
1985, Cowan 5524 et al. (NY); Mpio. Carbrt, Potrero 7
rancho La Granada CIPES, Sep. 1984, Miranda RLG-068
(ANSM. SLPM); Mpio. Cajeme, about 2.2 mi. NE of liwy.
15 (toward a microwave tower), about 6.9 mi. SE of Ciudad
Obregon, Sep. 1973, Stevens 2067 & Fairhurst (1)S,
ENCB, GH, MO, NY); Mpio. Guaymas, ca. 15 mi. SE of
Guaymas, June 1979, Webster 17001 & lynch (GH,
MFXU, MO); Mpio. Hermosillo. 27 mi. W of Hermosillo
on the road to Kino Bay, Aug. 1941, Wiggins 131 & Rollins
(AR1Z, DS, GH, MO, NY, UC); Mpio. Huatabampo, Nes-
cotahueca vicinity 5 km (by air) N-NE of Camahuiroa, 6.7
km N on Bachoco road from Die/ <le Abril road, 1.9 km
S-SW (by air) Nescotahueca, Jan. 1995, Friedman 179-
95 & Zittere (ASU); Mpio. La Colorada, La Colorada, May
1905, Clokey 1915 (UC); Mpio. Magdalena, bank of Kio
Sasobe, 10 mi. E of Magdalena, Sep. 1934. Wiggins 7110
(DS, GH); Mpio. Navojoa, turnoff to microwave road on
Cerro Prieto, 17 km E of Navojoa on road to Alamos, Mar.
1993, Van Devender 93-248 et at. (ARIZ); Mpio. Onavas,
Onavas, graphite mine NE of pueblo, Oct. 1986, Rea 1248
(ARIZ); Mpio. Pitiquito, Pitiquito, May 1925, Kennedy
7069 (CAS, UC); Mpio. Puerto Periasco, ca. 7 km N of
Puerto Penasco, at ca. 2 km N of airport, June 1985, Fel-
ger 85-790 (ARIZ. MEXU); Mpio. Sovopa, NE side of Rio
Yaqui bridge on Mex. 16, just S of Tonichi, Sep. 1996,
Van Devender 96-350 et al. (ARIZ); Mpio. Ures, along
hwy. 14 leading NE out of Hermosillo, ca. 37.5 km SW
of Ures, Aug. 1995, Snow 6593 & Frinzie (MEXU). Chi¬
huahua: along New Mexico— Chihuahua border 0.9 mi. W
of monument #3, July 1986, Worthington 14398 (NY). Si¬
naloa: Mpio. Culiacan, Culiacdn, Oct. 1904, Brandegee
s.n. (GH, UC); Mpio. El Fuerte, Valle del Carrizo, street
8(X), near San Francisco, Aug. 1975, Rodriguez 1607
(ARIZ, CAS, ENCB, MEXU, MO); Mpio. Salvador Alva¬
rado, on Cerro Tasirogojo, 9 km S Estaci6n Luis, 1 km W

(by air) Francisco Sarabia, 5.5 km (by air) SE Melchor
Ocampo, Aug. 1994, Friedman 210-94 A Espinosa (ASU);
Mpio. Sinaloa, Sinaloa de Leyva ca. I km N of Lagunilla,
to Santa Fe, Nov. 1990, Bojorquez 758 (MEXU).

Tidestromia suffruticosa var. oblongifolia (S.

Watson) Sanchez-del Pino & Flores Olvera,
comb, et stat. nov. Basionym: Cladothrix
oblongifolia S. Watson, Proc. Amer. Acad. Arts
17: 376. 1882. Tidestromia oblongifolia (S.
Watson) Standley, J. Wash. Acad. Sci. 6: 70.
1916. TYPE: U.S.A. Arizona: Yuma Co.,
Yuma, 25 June 1881. Pringle 1881 (lectotype,
here designated, GH; isolectotypes, G, K, NY,
US).

Tidestromia oblongifolia subsp. cryptantha (S. Watson)
Wiggins, Contr. Dudley Herb. 4: 16. 1950. Ba¬
sionym: Cladothrix cryptantha S. Watson, Proc.
Amer. Acad. Arts 26: 125. 1891. TYPE: U.S.A. Cal¬
ifornia: San Diego Co., Canso Creek, Nov. 1890, ()r-
cutt 2186 (lectotype, here designated, GH; isolec-
totype, K).

Tidestromia oblongifolia was recognized as a
separate species (e.g., Standley, 1917; Correll &
Johnston, 1970) based on minor differences in sta-
minodia and leaf characters, which are variable.
Also, two subspecies were proposed for T. oblon¬
gifolia (Wiggins, 1950) based on characters of leaf
form and size, and depth of the involucre. We rec¬
ognize T. oblongifolia as nonspecific with T. suffru-

Volume 12, Number 3
2002

Sanchez-del Pino & Flores Olvera
Tidestromia from North America

407

ticosa and propose two varieties based on charac¬
ters of leaves and internodes of the secondary
branches, which subtend fruiting involucres. Tides¬
tromia sujfruticosa var. sujfruticosa has leaves usu¬
ally ovate, rarely lanceolate, very broadly ovate, or
reniform, and internodes usually well defined, not
crowded. Tidestromia suffruticosa var. oblongifolia
has leaves usually oblong, rarely ovate-oblong, cir¬
cular, lanceolate or ovate, and internodes usually
reduced, crowded. These varieties are geographi¬
cally isolated: T. suffruticosa var. sujfruticosa ranges
through New Mexico, Texas, Chihuahua, Durango,
and Coahuila, whereas 7. sujfruticosa var. oblongi¬
folia ranges through California, Baja California,
Utah, Nevada, Arizona, and Sonora.

In the protologue of Cladothrix oblongifolia and
C. cryptantha, Watson cited syntypes. For the for¬
mer name Watson cited: “on the banks of Colorado,
near Chimney Peak (Dr. Newberry) and at Yuma
(C. G. Pringle) and in the Mohave Desert (S. B. &
W. F. Parish).” Specimens cited for C. cryptantha
are: Tarry 274 collected in 1881 at Colton, Cali¬
fornia, and Orcutt 2186 collected in 1890 at Canso
Creek. San Diego County. We chose the best-pre¬
served specimens at GH to lectotypify both these
names.

Acknowledgments. We are grateful to Fernando
Chiang for reviewing the manuscript and for the
Latin descriptions, and Helga Ochoterena for crit¬
ical comments. We thank the curators and staff of
ANSM, ARIZ, ASU, CAS, DS, ENCB. GH, K,
MEXU, MO, NY, and UC for the loan of specimens.
We appreciate the helpful assistance of Sara Fuentes
with the SEM photomicrographs. We thank Carlos
Gomez-Hinostrosa for his company in the field
studies. Albino Luna for the preparation of the
plant illustrations, and Samuel Aguilar Ogarrio,
Helga Ochoterena, and Alfredo Wong for the prep¬
aration of the figures. Finally, we thank Thomas

Borsch and Steve Clemants for useful comments

that highly improved the content of the manuscript.

Literature Cited

Borsch, T. 1998. Pollen types in the Amaranthaceae. Mor¬
phology and evolutionary significance. Grana 37: 129—
142.

Carolin. R. C. 1983. The trichomes of the Chenopodiaceae
and Amaranthaceae. Bot. Jahrh. Syst. 103: 451-466.

Correll, I). S. & M. C. Johnston. 1970. Pp. 551-574 in
Manual of the Vascular Plants of Texas. Texas Research
Foundation, Renner, Texas.

Cory, V. L. 1936. New names and combinations for Texas
plants. Rhodora 38: 404—408.

Eliasson, U. 1988. Floral morphology and taxonomic re¬
lations among the genera of Amaranthaceae in the New
World and the Hawaiian Islands. J. Linn. Soc., Bot. 96:
235-283.

Henrickson, J. 1993. Amaranthaceae. Pp. 130-134 in J.
C. Hickman (editor). The Jepson Manual. Higher Plants
of California. Univ. California Press, Berkeley.

Johnston, I. M. 1943. Noteworthy species from Mexico and
adjacent United States. 1. J. Arnold Arbor. 24: 227-
236.

Martin, S. P. & C. M. Drew. 1970. Additional scanning
electron photomicrographs of southwestern pollen
grains. J. Arizona Acad. Sci. 6: 140-161.

Reddy, M. H.. S. N. B. Reddy & R. R. V. Raju. 1998.
Structure, distribution and taxonomic importance of tri¬
chomes in the tribe Gomphreneae (Amaranthaceae). J.
Indian Bot. Soc. 77: 235-236.

Robertson, K. R. 1981. The genera of Amaranthaceae in
the southeastern United States. J. Arnold Arbor. 62:
267-314.

Sanchez-del Pino, I. 2001. Sistemdtica del Genero Tides¬
tromia Standi. (Amaranthaceae). Tesis Maestro en Cien-
cias, Universidad Nacional Autonoma de Mexico, Mex¬
ico.

Schinz, 11. 1893. Amaranthaceae. Pp. 91—118 in A. En-
gler & K. Prantl (editors). Die naturlichen Pflanzenfa-
milien, 1st ed., vol. Ill, la. W. Engelmann, Leipzig.

Standley, P C. 1916. Tidestromia, a new generic name. J.
Wash. Acad. Sci. 6: 69-70.

- . 1917. Amaranthaceae. North American Flora

21(2): 95-169. New York Botanical Garden, Bronx,
New York.

Steyermark, J. A. 1932. Some new spermatophytes from
Texas. Ann. Missouri Bot. Gard. 19: 389—395.

Wiggins, I. L. 1950. Taxonomic notes on plants from the
Sonoran Desert. Contr. Dudley Herb. 4(2): 15-17.

A New Tassadia (Apocynaceae, Asclepiadoideae) from Bolivia

and Brazil

W. I). Stevens

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

douglas.stevens@mobot.org

ABSTRACT. Tassadia capitata, from Santa Cruz.
Bolivia, and Para, Brazil, is described and illus¬
trated. This new species is most closely related to
T. guianensis Decaisne, and the two species are
somewhat intermediate between Tassadia and Sten-
omeria.

Key words: Apocynaceae, Asclepiadoideae, Bo¬
livia, Brazil, Stenomeria, Tassadia.

Tassadia Decaisne is a genus of about 25 species
of tropical and subtropical South America, with one
species, T. obovata Decaisne, ranging through Cen¬
tral America to Veracruz, Mexico. Tassadia belongs
to the tribe Asclepiadeae and is most closely re¬
lated to the genus Stenomeria Turez. The two gen¬
era share, in particular, an unusual arrangement of
the inflorescences. The true inflorescence is an ex¬
tra-axillary cyme, which is the normal inflorescence
of the Asclepiadoideae. However, these sessile or
nearly sessile, congested cymes are mostly ar¬
ranged along leafless axillary branches; cymes as¬
sociated with leafy nodes are also occasionally pro¬
duced. These inflorescences have been described
as thyrses or pleiothyrses (Fontella, 1977; Morillo,
1990), but those terms refer to mixed inflorescences
in which the primary axes are paniculate anti the
ultimate axes cymose (Jackson, 192B). I.iede and
Weberling (1995) also considered the leafless
branches to be inflorescences and erroneously com¬
pared them with the true, extra-axillary inflores¬
cences of Tylophora, which are superficially similar.
I.iede and Meve (2001) repeated the error in con¬
sidering the inflorescences of Tassadia to be “lax."
Stenomeria differs from Tassadia in having rostrate
and glabrous style apices, strongly contorted corolla
lobes, and slender and symmetrical follicles. In
Tassadia, only this new species and T. guianensis
have elongate style apices and in both cases they
are penicillate, the corolla lobes are slightly to not
at all contorted, and the follicles, when known, tend
to be broad and/or asymmetrical.

Tassadia capitata W. 1). Stevens, sp. nov. TYPE:
Bolivia. Santa Cruz. Velasco Prov.: Parque Na-
cional Noel Kempff Mercado, Arroyo Las Lon-
dras, a 15 km al N del ernpalme hacia el ase-
rradero Chore, bosque inundado temporalmente,
siempreverde, 14°24'18"S, 61°08'40”W, 150 m,
10 Oct. 1996, A. M. Carri/m, R. Guillen, M.
Garvizu, P. Soliz, J. Huffman & L Saucedo 360
(holotype, MO; isotype, USZ). Figure 1 A-D.

A Tassadia guianensi Decaisne apice styli capitato dig-
noscenda.

Twining (to right) vine, lower parts unknown,
stems densely puberulent with curved, multicellu¬
lar, red-tipped hairs 0.2— 0.4 mm long; sap white.
Leaves opposite, without pseudostipules, blade
3. 6-7. 5 X 1.7— 4.2 cm, elliptic, apex abruptly acu¬
minate, base acute to obtuse or nearly rounded, gla¬
brous above except with a line of hairs on midrib,
densely and uniformly puberulent below, lateral
veins 4 to 7, colleters 2; petiole 4—8 mm long, pu¬
berulent. Inflorescence extra-axillary, I per node,
congested-cymose, arranged on leafless or nearly
leafless axillary or terminal branches with the ap¬
pearance of inflorescences or occasionally with iso¬
lated inflorescences at leafy nodes, peduncle absent
to 1 mm long, pedicel 0.5-1. 2 mm long, bracts 0.6-
0.7 X 0.4— 0.7 mm, ovate; calyx divided to base,
with 0-2 colleters per sinus, lobes ovate to lance¬
olate with acute to obtuse tips, 0.8-1. 4 X 0.5— 0.9
mm, apparently green, glabrous to sparsely puber¬
ulent along midrib outside, glabrous inside; corolla
campanulate at base, aestivation weakly contorted
and dextrorse, white with a yellow or green tint,
tube 0.7-1. 5 mm long, glabrous outside, barbate
inside with glassy, unicellular hairs ea. 0.1 mm
long, lobes narrowly lanceolate with acute tips,
erect, 2.6— 3.6 mm long, 0.8— 0.9 mm wide at base,
glabrous outside, inside with margins densely pu¬
berulent with white, unicellular hairs ca. 0.05 mm
long; androecium and gynoecium entirely fused and
forming a short-stipitate gynostegium; corona gy-
nostegial, adnate to gynostegium at base, of 5 free
lobes opposite the anthers, these rounded and with

Novon 12: 408-410. 2002.

Volume 12, Number 3
2002

Stevens

Tassadia captitata from Bolivia and Brazil

409

Figure 1. A-D, Tassadia capitata W. D. Stevens i Carr ion et al. 360, MO). — A. Flowering branch. — B. Flower with
corolla partially removed. — C. Corona and gynostegium. — D. Pollinarium. — E. Tassadia guianensis ( Kappler 1410,
Suriname: Para, Feb. 1844. MO). Corona and gynostegium.

2 small lateral teeth, 0.6— 0.9 mm long, 0.5— 0.8 mm
wide at base; anthers quadrate, ca. 0.5 mm long
anti wide, 2-celled, terminal appendages ovate, ap-
pressed to base of style apex, guide rails linear, ca.
0.5 X 0.05 mm; corpusculum 0.12—0.13 X 0.06—
0.09 mm, ellipsoid, pale brown, translators 0.05—
0.08 mm long, terete, somewhat sigmoid, pollinia

pendent, 0.15—0.18 X 0.05—0.08 mm, ellipsoid;
style apex capitate, somewhat bilobed, 0.7— 0.8 mm
long, 0.5— 0.8 mm wide at tip, dilated apex densely
penicillate with projections ca. 0.1 mm long. Fruits
and seeds unknown.

This new species is similar to Tassadia guianen-

410

Novon

sis (Fig. IE) and probably closely related. The two
species share an elongate and penicillate style
apex, otherwise unknown in the genus. Tassadia
capitata differs from T. guianensis in many details,
including having the corolla lobes longer and nar¬
rower with shorter, more appressed hairs, having
the corolla tube barbate within, as opposed to gla¬
brous, having larger corona lobes that exceed in
length and cover the anthers, having quadrate rath¬
er than radially trapezoidal anthers and conse¬
quently the guide rails are vertical instead of flar¬
ing, and by a style apex that is distinctly capitate
and prominently penicillate, as opposed to subcy-
lindrical to conical and obscurely penic illate. Fon-
tella (1977) revised the genus and cited specimens
of T. guianensis from the Guianas, Amazonian Peru,
and the Brazilian state of Para, and it has since
been discovered in Amazonian Ecuador, while this
new species is known from Bolivia and Para, Brazil.

Both Fontella (1977) and Morillo (1990) have
pointed out the close relationship of Tassadia and
Stenomeria, with T. guianensis being the most in¬
termediate. This new species is likewise interme¬
diate but again the preponderance of characters
suggests keeping the genera separate and placing
the new species in Tassadia.

Paratypes. BOLIVIA. Santa Cruz: Velasco Province,
Parque Nacional Noel Kempff Mercado, Arroyo Las Lon-
dras, bosque de sartenejal, estacionalmente inundado,
14°24'09"S, 61°08'35"W, 150 m, 10 Oct. 1996, V/. Gar-
vizti, A. M. Carrion & R. Guillen 1R4 (MO, USZ). BRAZIL.
Para: Parque Nacional do Tapajos, Km 60 da estrada
Itaituba— Jacareeanga, illia cm frente ao Pimental, mata de
varzea, 26 Nov. 1978, M. G. Silva A C. Rosario .1976
(MO).

Acknowledgments. Alba Arbelaez (MO) pre¬
pared the illustration. The helpful comments of Vic¬
toria Hollowed (MO), Steven Lynch (LSUS), and
Eric Sundell (UAM) stimulated improvements in
the manuscript.

Literature Cited

Fontella P, J. 1977. Revisao taxonomica do genero Tas¬
sadia Decaisne (Asclepiadaceae). Arch. Jard. Bot. Rio
de Janeiro 21: 235—392.

Jackson, B. I). 1928. A Glossary ot Botanic Terms. Gerald
Duckworth. London.

Liede, S. & U. Meve. 2001. Taxonomic changes in Amer¬
ican Metastelminae (Apocynaceae— Asclepiadoideae).
Novon II: 171-182.

- & F. Weberling. 1995. On the inflorescence struc¬
ture of Asclepiadaceae. PI. Syst. Evol. 197: 99—109.
Morillo, G. 1990. Revision sinoptiea de Stenomeria Turcz.
(Asclepiadaceae). Acta Bot. Venez. 16: 79—91.

Una Especie Nueva y Notas sobre una Especie Notable de
Dioscorea (Dioscoreaceae) para Mexico

Oswaldo Tellez Valdes

Laboratorio de Recursos naturales UBIPRO, Faeultad de Estudios Superiores Iztacala
UNAM, Av. de Los Barrios 1, Los Reyes Iztacala, C.P. 54090 Tlalnepantla,

Edo. de Mexico, Mexico

Bernice G. Schubert

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

Resum EN. Dioscorea omiltemensis, una nueva es¬
pecie del estado de Guerrero, Mexico, es descrita.
Se ubica en la seccion Hyperocarpa, esta especie
nueva se distingue de D. oreodoxa de Mexico, asf
como de D. niederleinii y D. grisebachii de Suda-
merica dentro de la misma seccion por su morfol-
ogfa, distribucion geografica y preferencias ecolo-
gicas distintas. Ademas, en este trabajo, se
describen e ilustran las flores estaminadas de D.
tacanensis por primera vez, de ejemplares de la lo-
calidad tipo. Se discuten sus relaciones con D. cy-
phocarpa, se brindan datos de su distribucion geo¬
grafica y preferencias ecologicas.

ABSTRACT. Dioscorea omiltemensis , a new species
from the State of Guerrero, Mexico, is described.
Assigned to section Hyperocarpa, the new species
is distinguished from D. oreodoxa of Mexico, as
well as D. niederleinii and I). grisebachii of South
America within the same section, by its different
morphology, its geographic distribution, and its eco¬
logical preferences. In addition, in this paper the
staminate flowers of D. tacanensis are described
and illustrated for the first time from the collections
from the type locality; its relationship with D. cy-
phocarpa is discussed, as well as its geographic dis¬
tribution and ecologic preferences.

Key words: Dioscorea, Dioscoreaceae, Mexico.

Dioscorea omiltemensis 0. Tellez, sp. nov.
TIPO: Mexico. Guerrero: Mpio. Chilpancingo
de Los Bravo, Omiltemi, barranca Agua Frfa,
1 7°33'30"N, 99°4'52"W. alt. 2250 m, bosque
mesofilo de montana, Canada con corriente
permanente de agua, asociada a Croton, Mo-
nnina, Carpinus y Ostrya, 19 die. 1993, C.
Gonzalez 300 9 (holotipo, MEXU; isdtipo,
FCME). Figura 1.

Herba caule sinistrorsum volubili. Folia (3.5— )4. 0—6. 5

cm longa, (2.0— )3. 0-4. 5 cm lata, ovato-cordata, 7- vel 9-
nervata. I nflorescentia staminata incognita. Inflorescentia
pistillata 3.0— 5.2 cm longa, pauciflora. Flos pistillatus vi-
ridis; perianthi 1.5 mm longo, viridi; tepalis 1.0— 1.5 mm
longis, 0.5— 1.0 mm latis, ovatis vel oblongo-lanceolatis;
staminodis 3; stylis filiformibus. Capsula 0.6— 0.8 cm lon¬
ga, 0.35—0.45 cm lata, oblongoidea vel oblanceoloidea,
viridis, base acuta, apice rotundato; seminibus immaturis
brevi-alatis.

Rizomas desconocidos. Herbacea trepadora de-
licada, glabra; tallos levovolubles; pecfolos 2.3— 3.6
cm de largo, sulcados, glabros. Hojas (3.5— )4— 6.5
X (2— )3— 4.5 cm de ancho, alternas, ovado-corda-
das, membranaceas, glabras, 7- a 9-nervadas, ner-
vaduras no prominentes, apice agudo a acuminado,
base cordada, seno basal campanulado. Inflores-
cencias estaminadas desconoeidas. Inflorescencias
pistiladas 1 por axila, de 3-5.2 cm de largo, con 1
6 2 flores por inflorescencia; pedicelos 2^1 cm de
largo; bracteas 1—1.5 X 0.3— 0.4 mm, lanceoladas;
flores verdes; perianto en dos series, una interna y
otra externa, tepalos 6, 1—1.5 X 0.5 mm, ovado a
oblongo-lanceolados; estaminodios 3, filamentos ca.
1 mm de largo, restos de las anteras extrorsas per-
sistentes; estilos 3, fibres desde la base. Capsulas
inmaduras 0.6— 0.8 X 0.35—0.45 cm, verdes oblon-
gas a oblanceoladas, base aguda, apice redondeado;
semillas inmaduras casi totalmente carentes de un
ala, colapsadas.

Distribucion. Dioscorea omiltemensis es ende-
mica del estado de Guerrero al suroeste de Mexico.
Habita el bosque mesofilo de montana, a una alti-
tud de ca. 2250 m. La fructification ocurre durante
diciembre.

Dioscorea omiltemensis pertenece a la seccion
Hyperocarpa Uline, subgenero Helmia (Kunth) Ben-
tham, por poseer 3 estambres, los lrutos oblanceo-
lados, largo pedicelados, y las semillas practica-
mente carentes de un ala, caracterfsticas solo

Novon 12: 411-414. 2002.

412

Novon

Figura 1. Dioscorea omiltemensis 0. Tellez. — A. Habito de la planta. — IT Flor pistilada mostrando los estaminodios,
los restos de las anteras y las ramas del estilo (de C. Gonzalez 300, MFXU).

Volume 12, Number 3
2002

Tellez Valdes

Dioscorea omiltemensis de Mexico

413

compartidas por otras 3 especies conocidas en la
seceion (D. grisebachii Kunth, D. oreodoxa Schu¬
bert y D. niederleinii Knuth). Dos de estas especies
se distribuyen en Sudamerica (de Brasil al norte de
Argentina), y una mas en el occidente de Mexico.
Estas distribuciones ampliamente discontinuas no
son raras, ya que algunas secciones como Apodos-
temon Uline, Macrogynodium Uline, Strutantha
Uline y Trigonobasis Uline (Burk ill. I960; Knuth.
1924) comparten un importante numero de espe¬
cies entre Mexico y Sudamerica.

Dioscorea omiltemensis esta cercanamente rela-
cionada con D. oreodoxa, la cual crece en el cer-
cano estado de Colima, lambien en el occidente de
Mexico. Sin embargo, ambas especies difieren en
algunas caracterfstieas, D. omiltemensis tiene hojas
7- a 9-nervadas, peciolos lisos, flores verdes, y las
inflorescencias pistiladas son de 3—5.2 cm de largo,
con 1 a 2 flores por inflorescencia. En contraste,
D. oreodoxa tiene hojas 7-nervadas, peciolos ser-
rulados, flores purpureas, y las inflorescencias pis¬
tiladas son de 10—25 cm de largo, con 5 a 10 flores
por inflorescencia. Ademas, D. omiltemensis habita
en el bosque mesofilo de montana a una altitud de
2250 m, y florece y fructifica en diciembre; mien-
tras que I). oreodoxa habita en el bosque tropical
caducifolio a una altitud de 500 m, y florece y fruc¬
tifica de agosto a septiembre.

Ademas, en este mismo trabajo se deseriben las
caracterfstieas de las plantas estaminadas de Dios¬
corea tacanensis Lundell, una especie end6mica,
muy rara y notable por sus probables relaeiones.

Dioscorea tacanensis ha sido comunmente con-
fundida y hasta sinonimizada bajo D. cyphocarpa
Robinson por Matuda (1953), la especie mas cer¬
canamente relacionada a esta, y probablemente la
unica. Sin embargo, Schubert (1968) y Burkill
(1960) la consideran como una especie diferente.
Aetualmente, es faeil distinguir a cada una de estas
especies, D. tacanensis tiene flores purpureas, es-
tambres sesiles, las flores pistiladas, y los frutos son
puberulentos, mientras que D. cyphocarpa tiene
flores verdes, los estambres forman una columna, y
las floras pistiladas y los frutos son glabros. Ade¬
mas. D. tacanensis habita en el bosque mesofilo de
montana a una altitud entre 1500 y 2300 m, y es
endemica del estado de Chiapas, mientras que D.
cyphocarpa habita en el bosque tropical caducifolio
y los matorrales espinosos, a una altitud entre 1000
y 1390 m, y se distribuye en los estados de Gue¬
rrero y Oaxaca exclusivamente.

Por lo anterior, parece pertinente reu bicar a D.
tacanensis nuevamente como un taxon valido, ya
que definitivamente esta especie representa un ta¬
xon distinto a D. cyphocarpa.

Dioscorea tacanensis Lundell. Lloydia 2: 78.
1939. TIPO: Mexico. Chiapas: Union Juarez,
la falda sur del Volcan Tacana, Chiquihuites,
a la orilla de bosque mixto, alt. 1500 m, E.
Matuda 2416 9 (holotipo, MICH; isotipo,
MEXU). Figura 2.

Rizomas deconocidos. Tallos levovolubles, sul-
cados, esparcidamente puberulentos a glabrescen-
tes. Hojas 8.0—15.5 X 5.0—10.5 cm. alternas, am¬
pliamente ovadas a suborbiculares, 11- a
15-nervadas, la base profundamente cordada, apice
abruptamente caudado-acuminado, glabro; peciolos
5.0— 7.5 cm de largo, esparcidamente puberulentos
a glabrescentes, sulcados. Inflorescencias estami¬
nadas 16 2 racimos, 2.0— 5.0 cm de largo; raquis
ligeramente fractiflexo (en zigzag); flores 1 por bro-
te; pedicelos 1.0— 1.5 mm de largo; bracteas 1, 1.0—
1.5 mm de largo, linear-lanceoladas; perianto ca.
1.0 mm de largo, purpureo; tepalos 0.5— 0.6 mm de
largo, lanceolados, patentes; estambres 3, sesiles,
anteras extrorsas; pistilodio ausente. Inflorescen¬
cias pistiladas 1 6 2 racimos por axila, 11.0—15.0
cm de largo, raquis glandular-puberulento; flores
una por brote, pedicelos ca. 1.0 mm de largo; pe¬
rianto 1.5— 2.0 mm de largo; tepalos ca. 1.0 mm de
largo, insertos muy cerca de la base de la columna
estilar, columna estilar ca. 0.8 mm de largo, estilos
simples, teretes, delgados; estigmas 3; pedicelos
acrescentes en fruto, de 7.0—10.0 mm de largo.
Capsulas samaroides ca. 1.4 cm de largo, ca. 1.1
cm de ancho, con dos de los loculos abortados, el
tercero acrescente, membranaceo, oblongo a sub-
orbicular, castano-claro, puberulento; semilla 1,
6.0— 7.0 mm tie largo, ca. 5.0 mm de ancho, oblonga
a reniforme, castano-claro, lisa; practicamente no
alada o con escasa reminiscencias de lo que apa-
rentemente fue un ala.

Colecciones examinadas. MEXICO. Chiapas: Union
Juarez, Volcan Tacana, subiendo por Talquian, bosque rae-
sdfilo de montana. alt. 1700-2300 m, 8 julio 1991, L. M.
Gonzdlez V., R. Ramirez D., R. Gonzalez T. y R. Gonzalez
A. 4138 6 (IBUG).

Distribucidn. Esta especie esta restringida al
Volcan Tacana, en los Ionites entre el estado de
Chiapas y Guatemala. Habita en el bosque mesofilo
de montana, a una altitud entre 1500 y 2300 m. Se
encuentra comunmente asociada con Quercus , Al-
nus, Clethra, Fuchsia, Peperomia, Bomarea, Bego¬
nia, Gaultheria y Cuphea. La floracion y la fructi-
ficacion ocurren durante julio y agosto.

Estas dos especies D. tacanensis y I), cyphocarpa
que estan muy estrechamente relacionadas por com-
partir el mismo tipo de flores pistiladas, pedicelos lar¬
gos acrescentes, un fruto modificado con dos de las

414

Novon

Figura 2. Dioscorea tacanensis. — a. Rama con frutos. — b. Inflorescenoia estaminada. — c. Flor estaminada. — d.
Flor pistilada. — e. Fruto. (a y d de E. Matuda 2416, MEXU; bye de L. M. Gonzalez V. et al. 4128. IBUG.)

tres cavidades abortadas, y las semillas no aladas,
deben conformar un grupo diferente, al menos al nivel
de seceion dentro de Dioscorea. Por tales caractens-
tieas, Burkill (1960) y Schubert (1968) las ban con-
siderado como un posible estado intermedio y una liga
entre los generos Dioscorea y Rajania, debido a que
el fruto es sumamente semejante al del genero Ra¬
jania, principalmente por los pedicelos largos, las
cavidades abortadas, y la semilla no alada.

Agradecimientos. Agradezco a Elvia Esparza el
dibujo de D. omiltemensis , y a Regina O'Huges el
de D. tacanensis, en los dos casos excelentes. A
Barry Hammel por fungir como revisor de este

manuscrito, asf como a un revisor anonimo. Al her-
bario IBUG del Institute de Botanica, de la Univ-
ersidad de Guadalajara por facilitar colecciones
para el desarrollo de este trabajo.

Fiteratura Citada

Burkill, l. H. 1960. Organography and evolution of Dios¬
corea. J. Finn. Soc., Bot. 56(367): 319—412.

Knuth, R. 1924. Dioscoreaceae. In: Engler, Das Pflanzenr.
IV. 43 (Heft 87): 1-387.

Matuda, E. 1953. Fas Dioscoreas de Mdxico. Anales Inst.

Biol. Univ. Nac. Mexico, Ser. Bot. 24(2): 279—390.
Schubert, B. G. 1968. Aspects of taxonomy in the genus
l)i oscorea. Publ. Especial 8. Instituto Nacional de In-
vestigaciones Forestales. Mexico, D.F.

Arnaldoa argentea (Barnadesioideae: Asteraceae), a New Species
and a New Generic Record for Ecuador

Carmen Ulloa Ulloa and Peter M. Jorgensen
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
carmen.ulloa@mobot.org; peter.jorgensen@mobot.org

Michael 0. Dillon

Department of Botany, The Field Museum, Chicago, Illinois 60605, U.S.A.

dillon@fieldmuseum.org

ABSTRACT. A new species of Asteraceae, Arnal¬
doa argentea C. Ulloa, P. Jprgensen & M. 0. Dillon,
from southern Ecuador is described and illustrated.
It is characterized by its cream-white to light or¬
ange corollas and red-brown phyllaries covered by
a dense silvery pubescence, especially on the ad-
axial surface. The genus was previously known only
from northern Peru. A key to the species of Arnal¬
doa is presented.

Resumen. Se describe e ilustra una nueva espe-
cie de Asteraceae, Arnaldoa argentea C. Ulloa, P.
Jprgensen & M. O. Dillon, del sur de Ecuador. Esta
especie se caracteriza por las corolas de color blan-
co-crema a anaranjado palido y las filarias cafe-
rojizas cubiertas por un denso tomento argenteo es-
pecialmente en la superficie adaxial. El genero
anteriormente se conocfa solo del norte de Peru. Se
presenta una clave para todas las especies de Ar¬
naldoa.

Key words: Andes, Arnaldoa, Asteraceae, Bar¬
nadesioideae, Ecuador.

Recent collecting efforts in the Andean region of
southern Ecuador have revealed the presence of the
genus Arnaldoa Cabrera in the country (Ulloa Ulloa
& Jprgensen, in press). The genus was previously
known only from northern Peru (Ferreyra, 1995;
Stuessy & Sagastegui Alva, 1993). Arnaldoa is a
small genus of three species of shrubs, with long
axillary spines, solitary, terminal, homogamous, dis¬
coid capitula, and pseudobilabiate corollas. It be¬
longs to the South American subfamily Barnade¬
sioideae (formerly included as Barnadesiinae in the
Mutisieae, see Cabrera, 1977; Darling, 1991). The
members of this monophyletic subfamily, sister
group to the rest of the Asteraceae, are character¬
ized by a unique indumentum of barnadesioid vil¬
lous hairs (a long filiform cell attached to a swollen
cell, which is held by a basal epidermal cell, see

Erbar & Leins. 2000) on the corollas, achenes, and
pappus of most species (Bremer, 1994). Most mem¬
bers also have peculiar spines in pairs, sometimes
solitary or three to five together between the sub¬
tending leaf and the axillary bud (Bremer, 1994).
Various affinities with other, mainly Andean, genera
of this subfamily have been suggested for Arnaldoa.
Morphologically the genus has been placed be¬
tween Chuquiraga and Dasyphyllum (Cabrera,
1962, 1977), and an intergeneric hybridization be¬
tween Barnadesia and Chuquiraga has been sug¬
gested (Stuessy et al., 1996). A recent molecular
phylogeny (Gustafsson et al.. 2001) reveals a well-
supported Arnaldoa clade also comprising Fulealdea
and Dasyphyllum subg. Archidasyphyllum. Although
this group is morphologically heterogeneous, the
placement of Arnaldoa based on morphological char¬
acters is inconclusive and neither strongly contra-
die ts nor supports the molecular results (Gustafsson
et al., 2001).

The species of Arnaldoa have narrow distribu¬
tions and grow on wooded or shrubby slopes in
more or less xerophytic habitats, between 1370 and
3000 m. The species known in Peru have showy
bright orange-red or purple corollas, whereas the
specimens from Ecuador have cream-white or light
orange ones. Further comparison revealed that the
latter represent a new species.

Arnaldoa argentea C. Ulloa. P. j0rgensen & M.
0. Dillon, sp. nov. T'l PE: Ecuador. Loja: Ama-
luza, road to the antennas above town, at the
top, 5 Nov. 2000. 2400 m. 79°26'26"W.
4°33'58"S, P. M. Jorgensen, C. Ulloa, J. Ca-
ranqui, J. Madsen & 0. Sanchez 2234 (holo-
type, QCNE; isotypes, F, LOJA, MO, QCA,
USM, WU). Figures 1, 2.

Haec species habitu Arnaldoae weherbaueri, (ilamentis
pubescentibus A. macbrideanae similis, sed a hac capitulo
multo majore, ab ilia filamentis pubescentibus alque co-

Novon 12: 415-419. 2002.

416

Novon

Figure 1.
removed.
2234.

Arnaldoa argentea C. Ulloa, P. Jprgensen & M. 0. Dillon. — A. Habit. — B. Floret with some pappus bristles
— C. Stamen. Drawing based on photographs taken at the type locality and the isotypes of j0rgensen el al.

Volume 12, Number 3
2002

Ulloa Ulloa et al.

Arnaldoa argentea from Ecuador

417

Figure 2. Arnaldoa argentea. Flowering branch (left). Habit. Photographs by Jens Madsen taken at the type locality,
2001.

rolla ex pappo vix exserta, ab arnbabus corollae colore
distinguitur.

Tall, slender shrubs, up to 3.5 m tall, the branch¬
es striate, densely rufous hirsute-tomentose, gla-
brescent with age, with paired axillary spines, the
longest to (17— )27 mm long, projecting at about 90°
at the base of the petiole, generally with two ad¬
ditional pairs of shorter ones, 4—1 1 mm long.
Leaves simple, alternate, decreasing in size toward
the base of the capitula; petioles 3—6 mm long,
densely tomentose, with long tan hairs, glabrescent
with age; blades elliptic-oblong to ovate, (2.0— )4.3 —
7.8 X 1.2— 4.1 cm, coriaceous, apex obtuse, spiny-
mucronate, mucro 1.5—3 mm long, base unequal,
entire, tomentose to sparsely tomentose at the base
of the main nerve and becoming shiny adaxially.

densely tomentose abaxially and velvety to the
touch, with long pale yellow' to argenteous hairs es¬
pecially along the veins and margins, glabrescent
with age, 3-veined from near the base, tertiary ve¬
nation finely reticulate. Capitulescence of solitary,
terminal, sessile capitula, these homogamous, dis¬
coid, 40-58 X 35—50 mm, the involucre campan-
ulate; phyllaries 1 1— 1 5-seriate, slightly recurved at
apex and spreading with age, the outer series dark
green and progressively becoming red- to orange-
brown toward the median and inner series, all
densely argenteous tomentose in the 1/2— 2/3 upper
portion adaxially, and along the margins, glabrous
at the base, sparsely tomentose to glabrescent on
the abaxial surface, the outer phyllaries ovate to
triangular, 6—12 X 5—6 mm, acuminate to cuspi-

418

Novon

date, with a spiny apex 3—5 mm long, the median
phyllaries ovate, 15-18 X 3-5 mm, long-acuminate
with a spiny apex 4-6 mm long, the margins slight¬
ly reflexed, the inner phyllaries lanceolate, 35—38
X 2—3 mm, the margins reflexed. Florets 55 to 70,
the corollas 25—28.5 mm long, cream-white to light
orange, pseudobilabiate, with a 4-lobed expanded
limb and a single adaxial slender lobe, the tube 9—
10.5 X 2.5 mm, glabrous, the limb 16—17 mm long,
the lobes 4—8 mm long and densely white villous
outside, the adaxial lobe filiform, as long as the
limb, densely white-villous on the apical 5—6 mm;
filaments free, white-villous, the anthers 10-13 mm
long, caudate, the tails 1.5—2 mm long; style 30—
44 mm long, the stigmas cream becoming dark or¬
ange-brown. Achenes cylindrical, 6—7.5 X 2 mm,
densely white-villous; pappus of 21-26 villous bris¬
tles, 22-25 mm long.

Paratypes. ECUADOR. Loja: vicinity of Rellavista. at
“Las Antenas” (6—7 km from Amaluza), 31 May 2001,
2200-2300 m, 79°27'024"W, 4°33'421"S, J. E. Madsen &
C. Rosales HI 75 (LOJA not seen, MO), 14 Aug. 2001.
2400 m, J. E. Madsen et al. 8341 (LOJA not seen. MO).

Distribution, habitat, and conservation. This
species has only been collected on the top of a
small mountain just outside the town of Amaluza,
in a dry inter-Andean valley in the Province of
Loja, between 2200 and 2400 m. It was collected
in flower in May and June, less so in November,
and in fruit in August and November (Madsen,
pers. comm.). The area is heavily disturbed with
only small patches of natural vegetation on the hill¬
sides. The long spines may protect these plants
from the cattle and from being cut for fuel-wood;
however, they may not survive fires. These factors
combined with a so far very restricted distribution
make this species extremely vulnerable.

Etymology. The specific epithet refers to the
pubescence of the phyllaries and of the pappus,
which gives the capitula an overall silvery appear¬
ance.

This new species can be readily recognized from
the other species of Arnaldoa by its large silvery
capitula, with red- to orange-brown phyllaries, the
corollas barely exserted from the pappus, and the
villous filaments. It resembles A. weberbaueri
(Muschler) Ferreyra (including A. coccinasantha
(Muschler) Ferreyra) by the large capitula and
densely pubescent stems and leaves, but it differs
by having broader phyllaries (outer ones 5—6 mm
vs. 2—3 mm wide) that are more densely pubescent
on the adaxial surface (vs. the abaxial), villous fil¬
aments (vs. glabrous), and the corollas just slightly
exserted from the pappus (vs. much exserted). From

A. macbrideana Ferreyra it may be distinguished
by the pubescent leaves and stems (vs. glabrous)
and much larger capitula and florets. The corollas
of this new species are very similar to, but larger
than, those of A. macbrideana in their deeply lobed
limb with villous lobes, while the rest of the corolla
is glabrous both inside and outside; in A. weber¬
baueri the limb is only shortly 4-toothed at the apex
and the corolla is densely villous on its upper half
on the outside, and the tube is villous within. Al¬
though the corollas and styles of this new species
turn light orange to dark brown with age, they can¬
not compare with the bright red-orange to purple
showy corollas and styles of the other two species.
The pubescence of the filaments was suggested as
a species character (Erbar & Leins, 2000) to sep¬
arate A. macbrideana Ferreyra from A. weberbaueri
with glabrous ones. The species of this genus may
be separated by the following key.

Key to the Species of Arnaldoa

la. Filaments glabrous; corollas 32—50 mm long,
much exserted form the pappus, orange (rarely
purple), the tube villous inside, the limb shortly
4-toothed and longer than the adaxial slender
lobe; Peru (Ancash, Amazonas, Cajamarca, and

La Libertad) . A. weberbaueri

lb. Filaments densely villous; corolla slightly ex¬
serted from the pappus, the tube glabrous inside,
the limb deeply 4-lobed and as long as the ad¬
axial slender lobe.

2a. Corollas 15-19 mm long, purple; capitula
2. 5^4. 5 cm long; Peru (Lambayeque and

Piura) . A. macbrideana

2b. Corollas 25—28.5 mm long, cream-white to
light orange; capitula 4.0-5. 8 cm long; Ec¬
uador (Loja) . A. argentea

Acknowledgments. The National Geographic
Society (grant number 6327-98 to P. M. Jprgensen)
is thanked for supporting the field trips to southern
Ecuador, and Ecuador’s Ministerio del Ambiente
for the collecting permits. In Loja, we thank the
staff at the herbarium and Jens Madsen for his hos¬
pitality and for making the photographs available
to us. Roy Gereau provided the Latin diagnosis. We
thank John Pruski for his useful comments and dis¬
cussion and two anonymous reviewers.

Literature Cited

Bremer, K. 1994. Asteraceae: Cladisties & Classification.

Timber Press, Portland, Oregon.

Cabrera, A. L. 1962. Compuestas andinas nuevas. Bob
Soc. Argentina But. 10: 2 1 — 45.

- . 1977. Mutisieae — Systematic review. Pp. 1039—

1066 in V. H. Heywood, J. B. Harborne & B. L. Turner
(editors). The Biology and Chemistry of the Compositae.
Academic Press, London.

Erbar, C. & P. Leins. 2000. Some interesting features in

Volume 12, Number 3
2002

Ulloa Ulloa et al.

Arnaldoa argentea from Ecuador

419

the capitulum and flower of Arnaldoa macbrideana Fe-
rreyra (Asteraceae, Bamadesioideae). Bot. Jahrb. Svst.
122: 517-537.

Ferreyra, R. 1995. Family Asteraceae: Bart VI. In: J. F.
Maobride and collaborators (editors). Flora of Peru.
Fieldiana, Bot. (n.s.) 35: v, 1—101.

Gustafsson, M. H. G., A. S.-R. Pepper, V. A. Albert & M.
Kallersjo. 2001. Molecular phylogeny of the Barnade-
sioideae (Asteraceae). Nordic J. Bot. 21: 149—160.
Harling, G. 1991. Compositae— Mutisieae. In: G. Harling
& L. Andersson (editors), FI. Ecuador 42: 1-105.
Stuessy, T. F. & A. Sagastegui Alva. 1993. Revision de
Arnaldoa (Compositae, Bamadesioideae), g^nero ende-
mico del norte del Peru. Arnaldoa 1(4): 9—21.

- . T. Sang & M. 1,. DeVore. 1996. Phylogeny and

biogeography of the subfamily Bamadesioideae with im¬
plications for early evolution of the Compositae. Pp.
463-490 in D. J. N. Hind (editor), Compositae: System-
atics. Proceedings of the International Compositae Con¬
ference, Kew, 1994. Vol. 1. Royal Botanic Gardens,
Kew.

Ulloa Ulloa, C. & P. M. Jprgensen. In press. Sobre la
presencia del g6nero Arnaldoa (Asteraceae) en el Ec¬
uador. In: Z. Aguirre et al. (editors), Botanica Austroe-
cuatoriana — Estudios sobre los recursos vegetales en
las provincias de El Oro, Loja y Zamora-Chinchipe.
Ediciones Abya-yala. Quito, Ecuador.

Two New Species of Spiradiclis (Rubiaceae) from China

Wang Ruijiang

South China Institute of Botany, Chinese Academy of Sciences, Guangzhou 510650,

People’s Republic of China

ABSTRACT. Two new species, Spiradiclis loanu R.
J. Wang and S. chuniana R. J. Wang, from Guangxi,
China, are newly described and illustrated. Spirad¬
iclis loana is perhaps related to S. spathalata X. X.
Chen & C. C. Huang by its oblanceolate, elliptic,
or obovate and rosette-like leaves, but distin¬
guished by its long petioles, fewer secondary veins,
and white coronas. Spiradiclis chuniana is probably
related to S. tomentosa D. Fang & I). II. Oin by its
hairy leaves but distinguished by its truncate leaf
bases, long internodes and bracts, short pedicels,
and a ring of hair inside the middle of long-styled
flowers.

Key words: China, Rubiaceae, Spiradiclis.

I’he genus Spiradiclis Blumea (Rubiaceae— Hed-
yotideae) comprises 31 annual or perennial her¬
baceous species native to southern and southwest¬
ern China and northern India (Lo, 1999; Deb &
Rout, 1989) and mainly grows in mountainous, es¬
pecially limestone, areas. It is characterized by a
eineinnus cyme, heterostylous and 5-merous flow¬
ers, and globose, ovoid or linear-oblong dehiscent
capsules with four twisted or untwisted valves. Two
subgenera, subgenus Sinospiradiclis, characterized
by subglobose capsules and untwisted valves when
matured, and subgenus Spiradiclis. characterized
by ovate or linear-oblong and twisted valves, were
recommended (Lo. 1998). An expedition searching
for the Spiradiclis species in the Guangxi limestone
area was held in 2000 in order to discover more
about its biodiversity.

In the tribe Hedyotideae, the genus Spiradiclis
is very similar and closely related to such Chinese
genera as Hedyotis L. (s.l.) and Ophiorrhiza L. in
morphological characters, but differs from the for¬
mer by its 5-merous heterostylous not 4-merous
homostylous flowers, and from the latter by its
subglobose not obcordate or obconical capsules.

Spiradiclis loana R. J. Viang, sp. nov. TYPE: Chi¬
na. Guangxi: Nonggang Nature Reserve, alt.
280 m. 15 Aug. 2000 (fl & fr), R. ./. Wang 390
(holotype, I BSC; isotype, MO). Figure 1.

Species af finis S. spathalatae X. X. Chen & C. C.

Huang, sed petiolis 1—4.5 cm longis, nervis secundariis
utrinque ca. 10 paribus, corollarum tubis ca. 5 mm longis,
albis differt.

Herbs to 15 cm in height; stems erect, simple,
stout, pubescent. Leaves rosette-like, petiolate, the
blade 6—12 X 1.2—3 cm, oblanceolate, elliptic or
obovate, acute at apex, narrowed at base, spiny pi¬
lose adaxially, glaucous abaxially; secondary veins
ca. 10 pairs, slender, alternate or subopposite, con¬
spicuous beneath; petiole 1-4.5 cm long, slender,
pubescent; stipules persistent, interpetiolar, 1—2
mm wide at base, bilobed or not at apex, lobes
linear, 0.5—1 cm long, pubescent; internodes ca. 2
mm long. Inflorescence a terminal cyme; peduncle
6—10 cm long, pubescent; bracts ca. 1 mm long,
pubescent. Flowers heterostylous, corollas white;
long-styled flowers: calyx tubes ca. 1 mm long, ob-
conic, pubescent, calyx lobes ± as long as the tube,
triangular, acute, pubescent; corolla tubes 5-10
mm long, enlarged a little at middle, puberulous
outside, glabrous inside except with a pubescent
ring of long hairs at the middle and sparse simple
hairs above; lobes ca. I mm long, acute, pubescent
adaxially; stamens ca. 3.5 mm long, adnate a little
higher than the base of the corolla; anthers ca. 1.2
mm long, linear, situated near the ring of hairs;
ovaries ca. 0.6 mm long, bilocular; styles slender,
ca. 7 mm long; stigmas 2-lobed, ca. 0.8 mm; short-
styled flowers: calyxes as in long-styled flower; co¬
rolla tubes ca. 5.5 mm long, pubescent inside but
without a ring of long hairs, lobes ca. 1.2 mm long;
stamens adnate near the middle of the tube, ca. 2.5
mm long, anthers situated near the throat of the
tul je; styles ca. 3 mm long. Capsules subglobose,
ca. 2 mm diam., septicidal and loculicidal dehis¬
cence when mature. Seeds ca. 25. angled cylindric,
ca. 0.6 mm long, brown.

This species belongs to Spiradiclis subg. Sinos¬
piradiclis Lo because it has globose capsules and
four untwisted valves when the capsules mature. Its
oblanceolate, elliptic, or obovate and rosette-like
leaves make it look like S. spathalata X. X. Chen
& C. C. Huang, which has 5—8 mm long petioles,
15—25 secondary veins, and purplish red coronas,
but it differs from S. spathalata by its 1—4.5 cm

Novon 12: 420-423. 2002.

Volume 12, Number 3
2002

Wang

Spiradiclis from China

421

Figure 1. Spiradiclis loana R. J. Wang. — A. Habit. — B. Long-styled flower split and opened to show floral parts.
— C. Short-styled flower split to show floral parts. — D. Capsule. — E. Stipule. — F. Seeds. (Drawn from the type.)

422

No von

Figure 2. Spiradiclis chuniana R. J. Wang. — A. Habit. — lb Short-styled flower opened to show floral parts. — C.
Long-styled flower split to show floral parts. — I). Articulated hairs on the surface of the leaf. — E. Stipule. (Drawn
from the type.)

BDDDCDGDOOp

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2002

Wang

Spiradiclis from China

423

long petioles, 10 secondary veins, and white coro¬
nas.

Distribution and ecology. This species, collect¬
ed from Nonggang Nature Reserve, Guangxi, China,
grows under the secondary limestone forests at an
altitude of 200—280 m, where it is very common.
At the time it was found, the plant bore both white
flowers and green fruits. Therefore, the flowering
season might be from late July or early August to
the end of August or early September, and the fruit¬
ing season might be half a month later.

Notes. This species is named after Lo Hsien-
shui (IBSC), a Chinese taxonomist studying the Ru-
biaceae family.

Paratype. CHINA. Guangxi: Minqiangshe village,
Shanglong Xiang, Longzhou county, in densely mountain¬
ous forest, on stone, alt. 200 m, 4 Sep. 1958, Zhao-Qian
Zhang 11831 (IBSC).

Spiradiclis chuniana R. .). Wang, sp. nov. TYPE:
China. Guangxi: Nonggang Nature Reserve,
alt. 390 m, 15 Aug. 2000 (fl & young fr), R.
J. Wang 392 (holotype, IBSC; isotype. MO).
Figure 2.

Species nova S. tomentosae I). Fang & D. H. Qin, sim-
ilis, sed foliis ovatis, basi truncatis, bracteis brevioribus
1.2—2 mm longis, inflorescentiis densioribus, tubo corollae
floris longstylosi intus ad medium annulo piloso instructo
differt.

Annual herbs to 8 cm in height: stems simple,
stout, pubescent. Leaves ovate, 3—6.5 X 1.6—3 cm,
mucronate at apex, truncate at base, oblique,
sparsely covered with articulate villose hairs abax-
ially and adaxially, secondary veins 10 to 12 pairs,
distinct abaxially; petiole 1—3 cm long, pubescent;
stipules persistent, interpetiolar, 2- to 5-lobed,
lobes linear, 0.5-1 cm long, pubescent; internodes
1—1.5 cm long. Inflorescence a terminal cyme; pe¬
duncle 2.5—6 cm long, pubescent; pedicel 0—5 mm
long; bracts 1.2—2 mm long. Flowers heterostylous,
corollas white; long-styled (lowers: calyx tubes ca.
1 mm, obconic, pubescent; calyx lobes ± as long
as the tube, acute, pubescent; corolla tubes ca. 7.5
mm long, enlarged a little at the middle, puberu-
lous outside, glabrous inside except bearing a ring
of ca. 1 mm long white hairs at the middle and
sparsely pubescent above; lobes ca. 1 .5 mm long,
pubescent inside; stamens ca. 3 mm long, adnate a
little higher than the base of the corollas; anthers
ca. 1.2 mm long, linear, situated adjacent to the

ring of hairs; ovaries bilocular; styles ca. 5 mm
long, slender; stigmas bilobed. Short-styled flowers:
calyx tubes as in long-styled flowers; corolla tubes
pubescent inside but without a ring of long hairs;
stamens ca. 4 mm long, adnate nearly at the middle
of the tube, anthers situated lower than the throat;
styles ca. 2.5 mm long. Gapsules immature.

This species might belong to Spiradiclis subg.
Sinospiradiclis Lo because its immature capsules
look subglobose. Only after the capsules mature
can its position be assigned properly. In its habit
and leaf indument it is closely related to S. tomen-
tosa D. Fang & D. H. Qin. which has obovate or
oblanceolate leaves with an attenuate base, ca. 2
mm long internodes, 3—7 mm long bracts, and a
sparse inflorescence with distinct pedicels; the new
species is distinguished from the latter by its trun¬
cate leaf bases, 1—1.5 cm long internodes, 1.2-2
mm long bracts, the crowded inflorescence with in¬
distinct pedicels, and the corolla tube in long-
styled flowers bearing a hairy ring at the middle
inside. It differs from S. loana by its ovate leaves
with a truncate base and bearing articulate villose
hairs abaxially and adaxially.

Distribution and ecology. This species, collect¬
ed from Nonggang Nature Reserve, Guangxi, China,
grows in the shade of rocky and humid slopes under
secondary limestone forests at ca. 390 m altitude,
where it is apparently restricted. Based on field ob¬
servations, its flowering and fruiting season is main¬
ly from August to September.

Notes. This species is named after Chun Woon-
young, a former Academician of the Chinese Acad¬
emy of Sciences (CAS) and one of the pioneers of
plant taxonomy in China.

Acknowledgments. This work was supported by
a grant for Systematics and Evolutionary Biology
from the Chinese Academy of Sciences. 1 am grate¬
ful to Hu Chi-ming, the curator of the South China
Institute of Botany (IBSC), and anonymous review¬
ers for their suggestions.

Literature Cited

Deb, D. B. & R. C. Rout. 1989. Two new species of the
genus Spiradiclis (Rubiaceae) from India. Candollea 44:
225-229.

Lo, Hsienshui. 1998. Materials for Chinese Rubiaceae
(IV). Bull. Bot. Res. 18(3): 275-283.

- . 1999. Spiradiclis Bl. (Rubiaceae). In: Lo Hsien¬
shui, FI. Reipubl. Popularis Sin. 71(1): 86—110.

Nomenclatural Novelties in Chinese Elymus (Poaceae, Triticeae)

Guanghua Zhu

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

Shouliang Chen

Institute of Botany, Jiangsu Province and Chinese Academy of Sciences, Nanjing,
Jiangsu 210014, People’s Republic of China

Abstract. In the treatment of the Triticeae (Po¬
aceae) for the Flora of China, Roegneria k. Koch
is recognized as a synonym of Elymus L. This de¬
cision led to the following 18 nomenclatural nov¬
elties: E. abolinii (Drobow) Tzvelev var. nudiusculus
(L. B. Cai) S. L. Chen & G. Zhu, comb, et stat.
nov.; E. angustispiculatus S. L. Chen & G. Zhu,
nom. nov.; E. caianus S. L. Chen & G. Zhu, nom.
nov.; E. cheniae (L. B. Cai) G. Zhu, comb, nov.; E.
curtiaristatus (L. B. Cai) S. L. Chen & G. Zhu,
comb, nov.; E. debilis (I- B. Cai) S. L. Chen & G.
Zhu, comb, nov.; E. gmelinii (Ledebour) Tzvelev
var. macrantherus (Ohwi) S. L. Chen & G. Zhu,
comb, nov.; E. hongyuanensis (L. B. Cai) S. L. Chen
& G. Zhu, comb, nov.; E. kamoji (Ohwi) S. L. Chen
var. macerrimus G. Zhu, var. nov.; E. laxinodis (L.
B. Cai) S. L. Chen & G. Zhu, comb, nov.; E. mag-
nipodus (I,. B. Cai) S. I.. Chen & G. Zhu. comb,
nov.; E. serpentinus (L. B. Cai) S. L. Chen & G.
Zhu, comb, nov.; E. shouliangiae (L. B. Cai) G.
Zhu, comb, nov.; E. sinicus (Keng) S. L. Chen var.
medius (Keng) S. L. Chen & G. Zhu, comb, nov.;
E. sinoflexuosus S. L. Chen & G. Zhu, nom. nov.;
E. strictus (Keng) A. Love var. crassus (I.. B. Cai)
S. L. Chen & G. Zhu, comb, et stat. nov.; E. tri-
chospicula (L. B. Cai) S. L. Chen & G. Zhu, comb,
nov.; and E. yushuensis (L. B. Cai) S. L. Chen &
G. Zhu, comb. nov.

Key Words: China, Elymus, Poaceae. Roegner¬
ia, Triticeae.

The grass tribe Triticeae is complex for its tax¬
onomy at the generic level mainly because ol the
existence of many natural and artificial intergeneric
hybrids (Stebbins, 1956; Dewey, 1982; Barkworth
& Dewey, 1985; Kellogg, 1989). It has been sug¬
gested by different authors that the tribe consists
of a single genus or up to 38 genera (Love, 1984).
The generic limits of Roegneria K. Koch and Ely¬
mus L. are among the well-known taxonomic prob¬
lems in the tribe (Dewey, 1982, 1983. 1984; Love,
1984; Kellogg, 1989; Zhu et al., 1990; Zhu & Yang,

1990; Baum et ah. 1991; Cai. 1997; Yang et ah,

2001).

Linnaeus (1753) included five species within
Elymus, which was lectotypified by Hitchcock (in
Hitchcock & Green, 1929) on E. sibiricus L. Since
1753, the generic name has been adopted by most
authors, but with very different taxonomic concepts.
Traditionally, this genus consists of grass species
with more than one (often two) spikelets at each
node. Based on such a concept, Hitchcock (1935)
included Leymus Hochstetter and Psathrostachys
Nevski in his delimitation of Elymus. Nevski (1934)
placed these traditional Elymus species under Cli-
nelymus (Grisebach) Nevski. and included some
unrelated rhizomatous species under Elymus. This
treatment was accepted by Keng (1959). In addition
to these traditional species, Tzvelev (1976) also in¬
cluded in Elymus those self-pollinated species with
clustered habit and always one spikelet per node.

The name Roegneria K. Koch was published in
1848. based on R. caucasica K. Koch, and was
mostly ignored for almost a century afterward. Its
species, characterized by a single spikelet per
node, were mostly treated in Agropyron Gaertner
sensu lato (Bentham & Hooker. 1880; Hitchcock,
1935; Bor, 1960). When redefining the generic lim¬
its within the Triticeae, Nevski (1934) limited the
generic concept of Agropyron to only the typical
wheatgrass species and revived the name Roegner¬
ia. Agropyron sensu stricto has been widely ac¬
cepted since then. The genus Roegneria has been
accepted principally by Chinese and Japanese au¬
thors (Ohwi, 1941, 1942; Keng, 1959; Keng &
Chen. 1963; Yang, 1980, 1987; Zhu et al., 1990;
Zhu & Yang, 1990; Zhang et al.. 1991; Baum,
1991; Cai. 1997), but has been treated in the syn¬
onymy of Elymus by others (Melderis, 1980; Dewey,
1982; Love, 1984; Clayton & Renvoize, 1986:
Chen, 1997). In the treatment of the Triticeae for
the Flora Reipublicae Popularis Sinicae, Yang
(1987) included only those species with two spike-
lets per node under Elymus and included all those

Novon 12: 424-429. 2002.

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2002

Zhu & Chen

Chinese Elymus (Poaceae)

425

with one spikelet per node in Roegneria. Yang’s
concepts are widely accepted in China. The genus
Roegneria is recognized in all present Chinese local
floras, and new species are continually being de¬
scribed under this name (Zhu et ah, 1990; Zhu &
Yang, 1990; Cai, 1994, 1996a, 1996b, 1997; Cai
& Wang, 2001).

The aim of this article is not to justify the inclu¬
sion of Roegneria within Elymus. One may argue
both ways due to the taxonomic complexity ol the
whole tribe. As coauthors of the treatment of tribe
Triticeae for the English-language Flora of China
(Chen & Zhu, in prep.), we have decided to treat
Roegneria as a synonym of Elymus. This requires
the following 18 new names or new combinations
to be made available for the Flora.

1. Elymus abolinii (Drobow) Tzvelev var. mi-
diusculus (L. B. Cai) S. L. Chen & G. Zhu,
comb, et stat. nov. Basionym: Roegneria nu-
diuscula L. B. Cai. Acta Phytotax. Sin. 35:
171. 1997. TYPE: China. Xinjiang: Nileke,
along bank of stream, alt. 1650 m, 6 July
1976. Exped. Xinjiang 1746 (holotype, XJBI).

Only two collections are known of this variety. A
second specimen cited in the protologue, l\. R. Cui
82987 , was collected from Xiyuan County, Xin¬
jiang, but was not seen by the present authors. This
specimen is presumably deposited in Xinjiang Nor¬
mal University herbarium (XJNU), where the col¬
lector was employed. The holotype can be easily
identified as Elymus abolinii and differs only in
having spikelets with 4 or 5 florets each and an¬
thers ca. 5 mm long. Elymus abolinii is a variable
species. Other characters discussed in the proto¬
logue of Roegneria nudiuscula , such as spikes rel¬
atively short, glumes lanceolate and acute, lemma
densely pubescent, and the length of lemma awns,
all fall within the range of variation of E. abolinii.
Therefore, the former taxon can only be accepted
at the varietal level under the latter.

The basionym of Elymus abolinii is Agropyron
abolinii Drobow, the epithet of which was spelled
“ abolini ” in the protologue (Drobow, 1925). The
species was named after the type collector, Robert
Ivanovich Abolin, so according to Article 60.11 of
the ICBN (Greuter et ah, 2000), the epithet was
correctly changed to “ abolinii ” when Tzvelev made
the new combination under Elymus.

2. Elymus angustispiculatus S. L. Chen & G.

Zhu, nom. nov. Replaced synonym: Roegneria
angusta L. B. Cai, Acta Phytotax. Sin. 34: 332,
fig. 3(1-9). 1996. non Trinius ex Ledebour, FI.
Altaic. 1: 119. 1829. TYPE: China. Qinghai:
Xunhua, on mountain slope, alt. 2200 m, 8
July 1984, G. X. Lei 841810 (holotype,
HNWP).

A new name for this species is proposed here
because the epithet “ angustus ” is not available for
use in the genus Elymus. Elymus angustispiculatus
is compared in its protologue for unknown reasons
with E. barbicallus (Ohwi) S. L. Chen, which is not
closely related. The latter is easily distinguishable
by its awnless glumes. Elymus angustispiculatus is
in fact closest to E. tibeticus (Melderis) G. Singh in
having spikelets sessile, glumes shortly aristate,
lemma with long awns, and palea nearly as long as
the lemma. Elymus angustispiculatus differs from
E. tibeticus in having spikes 8—10 cm long (vs. 10—
16 cm long) and lemma sparsely pubescent (vs. gla¬
brous) with awns strongly recurved (vs. straight or
slightly curved).

3, Elymus caianus S. L. Chen & G. Zhu. nom.

nov. Replaced synonym: Roegneria gracilis L.
B. Cai. Acta Phytotax. Sin. 34: 328-330, fig.
1(9—15). 1996. non Philippi. I.innaea 33: 301.
1865. TYPE: China. Xizang [Tibet]: Gongbo-
gyamda, along bank of river, alt. 3970 m, 21
Sep. 1988, J. L. Yang et al. 880788 (holotype,
SAUT1).

A new name for this species is proposed here
because the epithet “ gracilis ” is not available for
use in the genus Elymus. It is named in honor of
Lianbing Cai, who first described this species. Ely¬
mus caianus was compared with E. puberulus
(Keng) A. Love in its protologue. However, little
similarity can be found between these two species.
Elymus caianus is readily distinguishable from the
sparsely tufted E. puberulus by its single-culmed
habit. In addition, E. caianus has spikelets purple
with 3 or 4 florets each and the lower glume much
less than half as long as the lemma, whereas E.
puberulus has spikelets green with 2 or 3 florets
each and the lower glume about half as long as the
lemma. Elymus caianus is similar to E. hongyu-
anensis L. B. Cai in the above characters, but dif¬
fers in having spikes more or less pendulous, palea
spiny but glabrous between the keels, ami anthers
yellow.

426

No von

4. Elymus cheniae (L. B. Cai) G. Zhu, comb. nov.

Basionym: Roegneria cheniae L. B. Cai, Acta
Phytotax. 34: 333, fig. 3(10-18). 1996. TYPE:
China. Xinjiang: Zhaosu, alt. 2300 m, 3 Aug.
1978, K. Tuo 780875 (holotype, XJBI).

Although without an explicit indication, Roeg¬
neria cheniae was originally named in honor of
Shouliang Chen. Elymus cheniae was compared
with E. sylvaticus (Keng & S. L. Chen) S. L. Chen
in the protologue. However, these two species share
little similarity except that their glumes are rela¬
tively narrow for the genus, and the lower glume is
more than half as long as the lemma. These char¬
acters are shared by many other species in the ge¬
nus. Elymus sylvaticus is easily distinguished from
the former as a much larger plant with shortly
awned glumes. Elymus cheniae is similar to E.
magnicaespes (D. F. Cui) L. B. Cai in having spikes
narrow, 5—13 cm long, glumes narrow and awnless,
and lemma blunt or slightly mucronate at the apex,
but differs in having 2-4 florets per spikelet vs. 4—
6 in E. magnicaespes.

5. Elymus curtiaristatus (L. B. Cai) S. L. Chen

& G. Zhu, comb. nov. Basionym: Roegneria
curtiaristata L. B. Cai, Guihaia 16: 200, fig.
1(10-15). 1996. TYPE: China. Xizang [Tibet]:
Changdu, Xishan, alt. 3400 m, 22 Aug. 1973,
Exped. Xizang 1988 (holotype, HNWP).

Elymus curtiaristatus is compared in its proto¬
logue with E. alashanicus (Keng) S. E. Chen, as
both have lower glumes half as long as the lemma.
However, E. curtiaristatus is easily distinguishable
by the awn of the lemma, whereas E. alashanicus
has lemmas nearly aw n less. Elymus curtiaristatus
is closer to E. calcicola Keng in having spikes
erect, glumes submuticate, lower glume half as long
as the lemma, lemmas with conspicuous awns, and
spikelets nearly sessile. Elymus curtiaristatus dif¬
fers in having lemmas with awns erect and short
(5—10 mm long) and culms 3-noded, whereas E.
alashanicus has lemmas with awns strongly re¬
curved and long (15-28 mm long) and culms often
5-noded.

6. Elymus debilis (L. B. Cai) S. L. Chen & G.

Zhu, comb. nov. Basionym: Roegneria debilis
L. B. Cai, Acta Phytotax. Sin. 34: 327, fig.
1(1-8). 1996. TYPE: China. Gansu: Sunan. in
forest, alt. 2350 m, 29 July 1991, T. N. He
2989 (holotype, HNWP).

Elymus debilis is only remotely similar to E. cal-
cicolus (Keng) A. Love, with which it is compared

in the protologue, in the slender appearance of the
culms. Elymus calcicolus is a much larger plant (ca.

I m tall), with spikes 12—20 cm long, whereas E.
debilis is smaller (up to 60 cm tall) with spikes 9-

II cm long. Elymus debdis is closest to E. schren-
hianus (Fischer & C. A. Meyer) Tzvelev in having
glumes shortly aristate, less than half as long as the
lemma, and lemma awns mostly straight or only
slightly curved. Elymus debilis differs from E.
schrenkianus in having culms slender with 5—7
nodes (vs. robust with 1-3 nodes), lemma 7-10 mm
long (vs. 15-25 mm long), and palea longer than
lemma (vs. as long as lemma).

7. Elymus gmelinii (Ledebour) Tzvelev var. ma-

cranllieriis (Ohwi) S. L. Chen & G. Zhu,
comb. nov. Basionym: Agropyron turczaninovii
Drobow var. macrantherum Ohwi, Acta Phy¬
totax. Geobot. 10: 98. 1941. TYPE: China.
“Mongolia interior [Nei Mongol) boreali-orien-
talis (Kochito Orientalis),” /. Hirayoshi 10277
(holotype, KYO).

Agropyron turczaninovii is regarded as a synonym
of Elymus gmelinii. 'Hie latter species is character¬
ized by having erect spikes, lemma awns divergent,
palea nearly as long as the lemma, and anthers very
short. Variety macrantherus differs from variety
gmelinii in having much taller culms (up to 100 cm
tall), longer spikes (5-25 cm long), broader glumes,
and lemma awns 10-14 mm long.

8. Elyinus hongyuanensis (E. B. Cai) S. E. Chen

& G. Zhu. comb. nov. Basionym: Roegneria
hongyuanensis L. B. Cai, Acta Phytotax. Sin.
35: 157, pi. 1. figs. 1-9. 1997. TYPE: China.
Sichuan: Hongyuan, in meadow, alt. 3400 m,
23 Sep. 1979, W. /. Xie 005 (holotype,
HNWP).

Elymus hongyuanensis is similar to E. caianus in
having glumes awnless, lower glume less than half
as long as the lemma, and lemma awns straight and
long, but differs in having spikes arching, palea
spiculose between the keels, and anthers black-yel¬
low.

9. Elymus kainoji (Ohwi) S. E. Chen var. ina-

cerrimus G. Zhu, var. nov. Roegneria kamoji
Ohwi var. macerrima Keng, in Keng & S. L.
Chen, J. Nanjing Univ. (Biol.) 3: 17. 1963,
nom. inval. (Art. 43.1). Roegneria kamoji
subsp. macerrima (Keng) N. R. Cui, Claves PI.
Xinjiang. 1: 153. 1982, comb, inval. TYPE:
Ch ina. Cuangxi: “Hsing-an |Xing'an], along
Li-Kiang [Lijiang],” 15 July 1937, H. Eung
21054 (holotype, N).

Volume 12, Number 3
2002

Zhu & Chen

Chinese Elymus (Poaceae)

427

Elymus hamoji var. macerrimus is validated by
reference to the diagnosis of Roegneria kamoji var.
rnacerrima. The latter is an invalid name under Ar¬
ticle 43.1 of the ICBN (Greuter et al., 2000) be¬
cause R. kamoji is itself invalid under Article
34.1(c), having been published merely as a syno¬
nym in the protologue of Agropyron kamoji Ohwi
(Ohwi, 1942: 179). Several authors have mistakenly
regarded R. kamoji as a valid name (keng. 1959:
Cui. 1982; Yang, 1987; Chen et al., 1987). Chen
(in Chen et al., 1987) published the new combi¬
nation Elymus kamoji , citing R. kamoji as the bas-
ionym with a full reference to its author and place
of publication. This can be regarded as a biblio¬
graphic error of citation lor A. kamoji under Article
33.4. and it does not invalidate the publication of
the new combination.

Elymus kamoji is characterized by having lower
glumes conspicuously shorter than the lemma, both
glumes and lemma with broad, membranous mar¬
gins, palea keels conspicuously winged, and lemma
not ciliate. Variety macerrimus differs from variety
kamoji in having leaf blades often recurved and ca.
2 mm wide (vs. 3-13 mm wide), spikes pale green
or grassy yellow (vs. green tinged with purple) and
2-6 cm long (vs. 7—20 cm long), and spikelets with

2- 5 (vs. 3—10) florets each.

10. Elymus laxinodis (L. B. Cai) S. L. Chen &
G. Zhu, comb. nov. Basionym: Roegneria lax¬
inodis L. B. Cai, Guihaia 16: 199, fig. 1(1—9).
1996. TYPE: China. Sichuan: Kangding, on
mountain slopes, alt. 3700 m, 18 Sep. 1973,
Exped. Xizang 2599 (holotype, HNWP).

Elymus laxinodis is similar to E. parviglumis
(Keng) A. I iive in having awnless glumes conspic¬
uously shorter than the lemma and lemma sparsely
hirtellous with awns longer than the lemma body.
Elymus laxinodis differs from E. parviglumis in
having spikelets each with 2-5 (vs. 5—9) florets, the
palea conspicuously shorter than the lemma (vs.
nearly as long as the lemma), and the glumes 2- or

3- veined (vs. 3— 5-veined).

1 1. Elymus magnipodus (E. B. Cai) S. L. Chen
& G. Zhu, comb. nov. Basionym: Roegneria
rnagnipoda L. B. Cai, Acta Phvtotax. Sin. 35:
164, pi. 1, figs. 18-26. 1997. TYPE: China.
Qinghai: Golmud, in gravelly places along riv¬
er banks, alt. 3160 m, 19 June 1963, Exped.
Abandoned Land 001 (holotype, HNWP).

Elymus magnipodus is similar to E. puberulus
(Keng) A. Love, but differs in having scapes slen¬
der. spikelets sessile, glumes submuticate, and

lemma aristate, with awns straight or slightly
curved. Elymus magnipodus differs from E. puber¬
ulus in having leaf blades recurved (vs. not re¬
curved), spikes erect (vs. nodding or recurved),
spikelets each with 6—8 (vs. 2—3) florets, and lem¬
ma awns about as long as (vs. twice as long as)
lemma body.

12. Elymus serpentinus (L. B. Cai) S. L. Chen
& G. Zhu. comb. nov. Basionym: Roegneria
serpentina L. B. Cai, Acta Phytotax. Sin. 35:
167, pi. 2, figs. 1-9. 1997. TYPE: China. He¬
bei: Yuxian, along bank of river, all. 2010 m,
10 July 1957, Exped. Shanxi 10147 (holotype,
HNWP).

Elymus serpentinus is similar to E. serotinus
(Keng) A. Love in having lax, narrow spikes, ob¬
long-lanceolate glumes, the lower glume aristate,
and lemma with a strongly recurved awn. However,
E. serpentinus differs from E. serotinus in having
leaf blades sparsely pilose on the adaxial surface,
glumes dentate on at least one side, the upper
glume also aristate, the lemma glabrous on the ab-
axial surface, with an awn 14-18 mm long, and the
palea much shorter than the lemma. Elymus ser¬
pentinus is also similar to E. sinoflexuosus S. L.
Chen & G. Zhu in spike appearance and the shapes
of the lemma and glumes, but both glumes are aris¬
tate in the former species and never dentate in the
latter.

13. Elymus shouliangiae (L. B. Cai) G. Zhu,
comb. nov. Basionym: Roegneria shouliangiae
L. B. Gai, Acta Phytotax. Sin. 35: 161, pi. 1,
figs. 10—17. 1997. TYPE: China. Xizang [Ti¬
bet]: Gyirong, river banks, alt. 2800 m, 6 July
1975, C. Y. Wu et al. 678 (holotype, PE).

Roegneria shouliangiae was named in honor of
Chen Shouliang. Elymus shouliangiae is similar to
E. brevipes (Keng) A. Love in having spikelets sub-
sessile, loosely arranged on the spike, glumes awn¬
less, lanceolate or oblong-lanceolate, and lemma
awns recurved. Elymus shouliangiae differs from E.
brevipes in having much longer spikelets (2.6— 3.2
cm long), more florets (8—10) in each spikelet,
much longer glumes (7—9 mm long), the upper
glume shortly aristate, and anthers brown, 4-5 mm
long.

14. Elymus sinicus (Keng) S. L. Chen var. med¬
ium (Keng) S. L. Chen & G. Zhu, comb. nov.
Basionym: Roegneria sinica var. media Keng,
in Keng & S. L. Chen, J. Nanjing. IJniv. (Biol.)
1: 35. 1963. TYPE: China. Shanxi: Mt. Wutai,
roadside, alt. 3500-4000 m, 11 July 1929. T.
Tang 1045 (holotype, PE).

428

Novon

Elymus sinicus is characterized by having culms
with 2 or 3 nodes, spikelets each with 4 or 5 florets,
the lower glume only slightly shorter than the lem¬
ma, and the lemma sparsely pubescent, with the
awn much longer than the lemma body. Variety
medius differs from variety sinicus in having leaf
blades up to 7 mm wide (vs. 3—4 mm wide) and
glumes shortly aristate with awns 1—3 mm long (vs.
1—1.8 cm long).

15. Elymus sinoflexuosus S. L. Chen & G. Zhu,
norm nov. Replaced synonym: Roegneria flex-
uosa L. B. Cai, Acta Phytotax. Sin. 34: 330,
fig. 2(1—9). 1996. non Tausch, Flora 20: 120.
1837. TYPE: China. Gansu: Zhangye, on
mountain slope, alt. 1750 m. 1 Aug. 1957, X.
Z. Zhang 203 (holotype, WIJK).

A new name for this species is proposed here
because the epithet ‘‘ fiexuosus ” is not available for
use in Elymus. The species is similar to E. serotinus
(Keng) A. Love in having lax. narrow spikes, ob¬
long-lanceolate glumes, and the lemma with a
strongly recurved awn. However, E. sinoflexuosus
differs from E. serotinus in having glabrous leaf
sheaths and glumes, glumes with 5-7 veins, the
lower glume aristate, and the palea glabrous be¬
tween the keels.

16. Elymus strictus (Keng) A. Love var. crassus
(I,. B. Cai) S. L. Chen & G. Zhu, comb, et stat.
nov. Basionym: Roegneria crassa L. B. Cai,
Acta Phytotax. Sin. 34: 332, fig. 2(10-16).
1996. TYPE: China. Ningxia: \anehi. on
mountain slope, alt. 1800 m, 21 July 1977, Z.

Y. Zhang & 11. J. Wang 140 (holotype, WUK).

Variety strictus is characterized by having lem¬
mas glabrous in the middle of the abaxial surface,
more or less puberulous elsewhere, and paleas pu-
berulous between the upper keels. Variety crassus
differs from variety strictus in having lemmas abax-
ially glabrous or hirsute along the veins, the callus
hirsute, and paleas glabrous between the keels.

17. Elymus trichospicula (L. B. Cai) S. L. Chen
& G. Zhu. comb. nov. Basionym: Roegneria
trichospicula L. B. Cai. Bull. Bot. Res., Harbin
14: 340, fig. 2. 1994. TYPE: China. Qinghai:
Yushu, near forest, alt. 3500 m, 17 Aug. 1981,

Z. I). Wei 22414 (holotype, NWBI).

Elymus trichospicula is characterized by having
glumes awnless with 3 keels, the lower glume about
half as long as the lemma, and the lemma densely
villous with an awn 6—12 mm long. In the proto-

logue, E. trichospicula was compared with E. sini¬
cus (Keng) S. L. Chen, but the latter has a much
larger lower glume, which is much more than half
as long as the lemma and can be easily distin¬
guished. Elymus trichospicula is similar to E. dol-
ichaterum (Keng) A. Love and E. leiotropis (Keng)
A. Love in the above characters, but the latter two
species have lemmas with much longer awns (15-
40 cm long) and glumes with 3-5 keels.

18. Elymus yusliuensis (L. B. Cai) S. L. Chen X
G. Zhu, comb. nov. Basionym: Roegneria yu-
shuensis L. B. Cai. Bull. Bot. Res., Harbin 14:
338, fig. 1. 1994. TYPE: China. Qinghai: Yu¬
shu, along roadside, alt. 3750 m, 24 Aug.
1980, Z. D. Wei 22105 (holotype, NWBI).

Elymus yushuensis and E. alashanicus (Keng) S.
L. Chen are similar and distinctive in the genus in
having spikes short (5—11 cm long), narrow, and
lax, glumes lanceolate or oblong-lanceolate, nearly
awnless, and lemmas slightly mucronate. Elymus
yushuensis differs from E. alashanicus in having
spikes pendent (vs. erect), purple (vs. pale yellow),
and hirsute (vs. glabrous).

Acknowledgments. We thank Nick Turland
(MO), Gerrit Davidse (MO), Sylvia Phillips (K), Vic¬
toria Hollowell (MO), and Roy Gereau (MO) for
their comments. Special thanks are due to Xilin
Yang from Inner Mongolia Teachers’ University for
his guidance to the senior author. We thank the
curators of the following herbaria for the loans of
specimens or other assistance with this study:
HNWP, PE, SAUTI (Institute of Wheat Research,
Sichuan Agricultural University), and XJBI (Xin¬
jiang Institute of Biology, Pedology and Desert Re¬
search. Chinese Academy of Sciences).

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study on the esterase and proxidase isoenzyme of Roeg¬
nerin K. Koch and Elymus Linnaeus. Acta Bot. Boreal. -
Occid. Sin. 10: 43-53.

A New Species of Oxytropis (Leguminosae) from Xizang (Tibet)

in China

Xiang-yun Zhu and Yu-fen Du

Institute of Botany, the Chinese Academy of Sciences, Bei jing 100093,

People’s Republic of China, xiangyunzhu@ns.ibcas.ac.cn

// iroyosh i Oh ash i

The Botanical Garden, Graduate School of Science, Tohoku University,

Sendai 980-0862, Japan

Abstract. A new species of Oxytropis (Legumi¬
nosae) from Xizang (Tibet) in China, O. qamdoensis
X. Y. Zhu, Y. F. Du & H. Ohashi, is described and
illustrated. The new species is very similar to 0.
ochrantha (sect. Baicalia Steller ex Bunge) and
shares with it verticillate and opposite leaflets and
glandular hairs, but differs in having leaflets with
9—1 1 whorls per leaf, the standard bilobed at the
apex, wings emarginate at the apex, the beak of the
keel-petals 1.0— 1.5 mm long, and pods with white-
tinged hairs.

Key words: China, Leguminosae, Oxytropis.

Oxytropis DC., a genus belonging to the tribe
Galegeae of Papilionoideae in the Leguminosae,
comprises about 300 species occurring in cold
mountainous regions of Europe, Asia, and North
America, and is most numerous in Central Asia
(Polhill, 1981). Bunge (1874) classified 181 species
of Oxytropis into 4 subgenera and 19 sections. For
Chinese Oxytropis, 125 species, 4 varieties, and 4
forms were grouped into 3 subgenera and 20 sec¬
tions (Zhu & Ohashi, 2000). Section Baicalia Stell¬
er ex Bunge now contains 16 species (including the
one described here), which are a [tart of 1 19 spe¬
cies in the subgenus Oxytropis in China. The sec¬
tion can be distinguished from the other sections
by having verticillate and opposite leaflets and
plants with glands (Zhu & Ohashi, 2000).

Oxytropis qainilocnsis X. Y. Zhu, Y. F. Du & 11.
Ohashi, sp. nov. TYPE: China. Xizang (Tibet):
Qamdo Xian, Karoo Zhen, Karoo Village, Jimo
Shan, within shrubs on dry and stony slope,
3215—3250 m. 16 Aug. 2000, X. Y. Zliu <X Y.
F. Du 20074 (holotype, PF: isotypes, PE,
TUS). Figure 1.

Haec species ah O. ochrantha foliolis 9—1 I verticillis,
vexillo bilobo ad apicem. alis emarginatis ad apieem, ros-

tris carinae 1.0— 1.5 mm longis, legumine albo-pubescenti
differt.

Aeaulescent perennial herb, 7—20 cm tall.
Leaves with white hairs, 5—8 cm long; leaflets ver¬
ticillate, or opposite at the upper part ol leaf axis,
acuminate or acute at apex, 9 to I 1 whorls per leaf,
4(6) per whorl, ovate or oblong-ovate, with white
hairs on both surfaces. 4—7 mm long, 1.5— 5.0 mm
wide, young leaflets with densely gray-white hairs;
stipules with white hairs, herbaceous, narrowly tri¬
angular, 3.5— 8.8 mm long, 2.0— 3.0 mm wide, con¬
nate w ith petiole at base. Racemes 5—13.5 cm long,
longer than leaves at fruiting time, shorter than
leaves at flowering time; bracts with white hairs,
ovate, 4. 5-7. 5 mm long, 2.0— 3.0 mm wide. Calyx
with white hairs, campanulate, 10-12 mm long, 5-
lobed, lobes lanceolate, 4.0— 6.0 mm long, ca. 0.1
mm wide, tube ca. 6.0 mm long. Corolla white;
standard broadly obovate, 16-19 mm long, 6.0— 7.0
mm wide, bilobed at apex; lamina narrowed to base;
wings clawed, lamina obovate, ca. 8 mm long, ca.
3 mm w ide, emarginate at apex, tapering to a claw',
claw ca. 7 mm long, ca. 0.5 mm wide, auriculate
at base, auricle ca. 3 mm long, ca. 1.5 mm wide;
keel-petals clawed, 11.5-17 mm long, lamina ob¬
long, 4.5— 8.0 mm long, 2.5— 3.0 mm wide, claw
7.0— 9.0 mm long, ca. 1.0 mm wide, auriculate at
base, auricle 1-1.5 mm long, 1—1.5 mm wide, beak
1.0— 1.5 mm long. Androecia diadelphous, ca. 11
mm long. Ovary pubescent, tubular, ca. 3 mm long,
ca. 0.5 mm wide, style incurved, ca. 7.5 mm long.
Pods with white hairs, ovoid, membranous, ca. 16.0
mm long, ca. 7.0 mm wide, with beak at the top of
pod, ca. 6.5 mm long, seeds cordiform, 1.78—2.20
mm long, 1.50—1.68 mm wide. Flowering and fruit¬
ing July-August.

Distribution. Known only from Qamdo Xian,
with shrubs on dry and stony slopes, Xizang, China;
3215-3250 m.

Novon 12: 430-432. 2002.

Volume 12, Number 3
2002

Zhu et al.

Oxytropis qamdoensis from Xizang

431

Figure 1. Holotype of Oxytropis qamdoensis X. Y. Zhu, Y. F. Du & H. Ohashi. — a. Calyx. — b. Bract (view from
outside). — c. Pod. — d. Androecium (view from outside). — e. Standard (view from inside). — f. Gynoecium. — g. Whole
of the type specimen. — h. Keel-petal (view from outside). — i. Wing (view from inside). Scale bars = 5 mm in a— f, h,
i; 10 mm in g. Drawn from the holotype, X. Y. Zhu <& Y. F. Du 20074.

Oxytropis qamdoensis differs from O. ochrantha
by leaflets with 9—1 1 whorls per leaf (vs. 13—19
whorls per leaf), standard bilobed at apex (vs.
rounded at apex), wings emarginate at apex (vs.
rounded at apex), beak of keel-petals 1.0— 1.5 mm
long (vs. 1 .5—2.0 mm long), and pods with white-
tinged hairs (vs. yellow hairs); it is distinguished

from O. ochrolongibracteata by plants small, less
than 0.2 m tall (vs. more than 0.4 m tall), flowers
white (vs. yellow), and most leaflets verticillate (vs.
opposite). This species is also different from O. bi¬
color in having white flowers (vs. blue-purple or
pale yellow), calyx lobes as long as its tube (vs.
shorter), and pods membranous (vs. coriaceous).

432

Novon

The key to Oxytropis qamdoensis and related spe¬
cies is given below.

Main Differences between Oxytropis qamdoensis and
Its Close Relatives

la. Stipules membranous; pods coriaceous . . 0. bicolor

lb. Stipules herbaceous; pods membranous.

2a. Plants large, more than 0.4 m tall; most leaf¬
lets opposite . 0. ochrolongibracteala

2b. Plants small, less than 0.2 m tall; most leaf¬
lets vertieillate.

3a. Leaflets 13—19 whorls per leaf; standard
rounded at apex; wings rounded at apex;
beak of keel-petals 1.5— 2.0 mm long;
pods with yellow hairs .... O. ochrantha
3b. Leaflets 9—11 whorls per leaf; standard
bilobed at apex; wings emarginate at
apex; beak of keel-petals 1.0- 1.5 mm

long; pods with white hairs .

. 0. qamdoensis

In August of 2000, Zhu and Du traveled to Xiz-
ang and collected this new species. Its population
occupied a small place with more or less 20 indi¬
viduals. which were growing among shrubs on dry

and stony slopes between 3215 and 3250 m and
distributed in Qamdo Xian of Xizang. Its distribu¬
tion in Xizang represents the western limit for this
section within China. The species should be pro¬
tected because of its small population and locality
close to a village.

Acknowledgments. This work was supported by
the National Nature Science Foundation of China
(grant number 30170072) and the Chinese Acad¬
emy of Sciences (On the Floristie Region and Dis¬
tribution Patterns of the Qinghai-Xizang Plateau
and Adjacent Areas [KSCX2-1 -06B]). We thank Y.
B. Sun for the drawing of the holotype specimen.

Literature Cited

Bunge, A. 1874. Species generis Oxytropis DC. Mem.

Acad. Petersh. VII, ser.. 22(1): 1-166.

Polhill, 11. M. 1981. Tribe 16. Galegeae (Broun) Torrey &
Gray (1838). Pp. 357-363 in R. M. Polhill & P. H.
Raven (editors). Advances in Legume Systematics, Part
1 . Royal Botanic Gardens, Kew.

Zhu X. Y. & H. Ohashi. 2000, Systematics of Chinese
Oxytropis DC. (Leguminosae). Cathaya 11-12: 1-218.

Volume 12, Number 3, pp. 309—432 of NOVON was published on 25 September 2002.

Volume 12
Number 4
2002

NOVON

Diplopanax vietnamensis, a New Species of Nyssaceae from
Vietnam — One More Living Representative of the Tertiary Flora

of Eurasia

Leonid V. Averyanov

Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov str.. 2,
St. -Peters burg, 197376, Russia, av@herb.bin.ras.spb.ru

Nguyen Tien Hiep

Institute of Ecology and Biological Resources oi the National Center for Natural Sciences
and Technology of Vietnam, Nghia Do, Cau Giay, Hanoi, Vietnam

Abstract. A new species of Diplopanax , I). viet¬
namensis (Nyssaceae, Comaceae s.l.), was discovered
in the mountains of southern Vietnam. It differs from
the single known species, Diplopanax stachyanthus,
by short, simple, densely pubescent inflorescences
and large yellow-white flowers with five prominent
bosses on the flower disc. This genus (with two spe¬
cies, D. stachyanthus and I), vietnamensis) is now re¬
garded as congeneric with Mastixicarpum, represen¬
tatives of which were an integral component of the
Mastixia- like paratropical broad-leaved evergreen
vegetation that covered much of the Northern Hemi¬
sphere f rom the uppermost Cretaceous to the late Mio¬
cene, about 65 to 7 million years ago.

Key words: Diplopanax, Nyssaceae, Tertiary
flora of Eurasia, Vietnam.

The montane flora within the eastern part of the
Indochinese Peninsula is still relatively unknown.
Without a doubt it is diverse and conceals a great
number of exciting discoveries. Floristically, the
highland forests of this area represent relietual
remnants of a primitive, tropical flora that occupied
w ide tropical and subtropical areas of Europe, Asia,
and North America during the early Tertiary, ap¬
proximately 40-70 million years ago (Axelrod et
al., 1998: Kubitzki & krutzseh. 1998; Zhang & Lu.
1998). Due to subsequent climate cooling and in¬
creased aridity, the European and North Asian rep¬

resentatives of this highland tropical flora and its
corresponding climatic zone have disappeared al¬
most completely. Small relietual and depauperate
elements of it can still be found in Portugal, the
Colchis, and in a few other orographieally favored
continental regions. The largest remaining humid
warm Tertiary flora was shifted to Southeast Asia.

One of the richest assemblages of ancient, tropical
Tertiary taxa may now be observed in highland trop¬
ical areas of Southeast Asia. Primary mountain for¬
ests of this region include a surprisingly large pro¬
portion of archaic genera well represented in the
fossil record of the Upper Cretaceous and bower Ter¬
tiary recorded from Europe, East Asia, and North
America (Kubitzki & Krutzseh, 1998; Martinetto,
1998). Among them are numerous Actinidiaceae,
Annonaceae, Berberidaceae, Chloranthaceae,
Daphniphyllaceae, Fagaeeae, Hamamelidaceae. Jug-
landaceae, Lardizabalaceae, Lauraeeae, Magnoli-
aceae, Menispermaceae, Nyssaceae. IVntaphylaea-
ceae, Platanaceae, Ranunculaceae, Rhoipteleaceae,
Sabiaceae, Saururaeeae, Theaceae, as well as a num¬
ber of gymnosperm genera such as Amentotaxus Pil-
ger (Cephalotaxaceae), Cephalotaxus Siebold & Zuc-
earini ex Endlicher (Cephalotaxaceae), Glyptostrobus
Endlicher (Taxodiaceae), Keteleeria Carriere (Pina-
ceae), Pseudolarix Gordon (Pinaceae), and a great
number of others (Axelrod et ah, 1998: Martinetto,
1998; Nguyen. 1998: Nguyen & Nguyen. 1998; Wu

Novon 12: 433-436. 2002.

434

Novon

Figure I. Diplopanax vietnamensis Averyanov & T. H. Nguyen. — a. Flowering leafy branchlet. — b. Portion of inflo¬
rescence with mature (lower buds just before opening. — r. Open (lower. — d. Adaxial surface of petal. — e. Calyx,
style, and bosses on the disc. (All drawn from an isotype, L Averyanov & T. H. Nguyen VH 596 , LE.)

Volume 12, Number 4
2002

Averyanov & Nguyen

Diplopanax vietnamensis (Nyssaceae)

435

& Wu, 1998). These plants were widely distributed
in the Northern Hemisphere during the Eocene and
Miocene. Most of these plants from these areas ex¬
isted during the past 5-10 million years including
such famous “living fossils” as Cathaya Chun &
Kuang (Pinaceae), Metasequoia Miki ex Hu & W. C.
Cheng (Taxodiaceae), and Pseudolarix Gordon (Pin¬
aceae) (Axelrod et al., 1998). Species of the genus
Diplopanax Handel-Mazzetti are typical representa¬
tives of this flora.

Diplopanax (with a single species, D. stachyan-
thus Handel-Mazzetti) was described in 1933 from
southern China and was initially placed in the Ar-
aliaceae (Handel-Mazzetti, 1933). Later, based on
the study of fruit structure, it was suggested that
the genus has affinities with Mastixia (Nyssaceae)
and especially with some fossil Mastixiaeeae (Hoo
& Tseng. 1978; Tseng, 1983). Comparison of extant
Diplopanax fruit pyrenes and fossil fruit remnants
of Miocene species of Mastixicarpum M. E. J.
Chandler gave evidence that the two genera may
he congeneric (Eyde & Xiang, 1990). It is remark¬
able that species of Mastixicarpum were wide¬
spread and integral components of Tertiary Euro¬
pean (loras especially during the Miocene. Fruits
of these plants are the most common fossils in lig¬
nite beds of Europe from the uppermost Cretaceous
to the late Miocene, roughly 65 to 7 million years
ago (Eyde & Xiang, 1990).

Woody fruits of Diplopanax anti Mastixicarpum
have a certain similarity and obviously represent
the same woody-fruited evolutionary line in the
family Nyssaceae (Cornaceae s.l.). It was long sup¬
posed that this line of woody-fruited relatives had
died out in the later Miocene about 5—10 million
years ago (Eyde & Xiang, 1990). With the descrip¬
tion of the new species, at least two modern species
still exist in Southeast Asia. The second extant spe¬
cies, Diplopanax vietnamensis, was recently discov¬
ered in southern Vietnam (Kon Turn Province). It
was observed as an occasional co-dominant of the
first forest stratum in primary, wet, evergreen,
broad-leaved tropical mountain forests. Other com¬
mon trees in these forests are also relictual genera
of Tertiary age from such families as Betulaceae,
Fagaceae, Hamamelidaceae, Eauraceae, Magnoli-
aceae, and Sabiaceae. This unique region repre¬
sents a model for the study ol the flora, vegetation,
and climate reconstructions from now-extinct
Northern Hemisphere forest floras of Tertiary age
(Martinetto, 1998). This area in Southeast Asia may
be the birthplace and cradle of the Holarctic and
even Paleotropic flora (Wu & Wu, 1998).

According to available data, Diplopanax includes
two extant and four extinct species. Distribution of

extant species comprises southern China and Viet¬
nam. Diplopanax stachyanthus is the type species of
the genus. The following dichotomous key is provid¬
ed for the identification of Diplopanax species.

Key to the Modekn Species of Diplopanax

la. Tree, about 8-12(15) m tall; inflorescence a ra¬

ceme or panicle branched at the base, 15—30 cm
long; rachis of inflorescence glabrous or sparsely
pubescent in basal portion; mature flower buds
before opening about 3 mm wide; flowers egg-
yellow, about 8 mm diam.; disc of the flower
without prominent bosses and ± smooth; style
glabrous . 1 . D. stachyanthus

lb. Tree, 15-25 m tall; inflorescence not a branched

spike, 5-8 cm long; rachis of inflorescence
densely pubescent in basal portion; mature flow¬
er buds before opening 4—5 mm wide; flowers
yellow-white, about 10-12 mm diam.; disc of the
flower with 5 prominent bilobed bosses sur¬
rounding the base of the style; style sparsely pu¬
bescent . 2. L). vietnamensis

1 . Diplopanax stachyanthus Handel-Mazzetti,

Sinensia 3(8): 197—198. 1933. TAPE: China.
Guangxi: Kwangsi, N. Luchen, Miu-shan, Bin-
long, 1200 m, 14 June 1928, K. C. Chin g 5969
(holotype, LU; isotype, NY).

This species is distributed in southern China (Prov¬
inces Hunan, Guangdong, Yunnan, and Guangxi
Zhuang Autonomous Region) and northern Vietnam
(Provinces Cao Bang and Vinh Phuc). Diplopanax
stachyanthus grows in wet evergreen broad -leaved
montane primary and open secondary forests and
shrubs, wood sides, at elevations oi 900-1600 m on
soils developed predominantly on silicate sands, clay,
granite, and rhyolites. It flowers in April— June.

Specimens examined. CHINA. Kwangsi: Sliap Man
Taai Shan, near Iu Shan village, SE Shang-sze, Kwangtung
Border (Shang-sze District). 23 May 1933, W. T. Tsang
22361 (LE). VIETNAM. Cao Bang Prov.: Le A pass at
elevation 1600 m a.s.l., 24 Nov. 1976, Khoi, Nhan & \e
71 (LE). Vinh Phuc Prov. : Tam Dao 2, 29 May 1977,
T. B. Nguyen 121 (LE).

2. Diplopanax vietnamensis Averyanov & 1. H.

Nguyen, sp. nov. TYPE: Vietnam. Prov. Kontum:
cloud evergreen primary forest on N slope of
Ngoe Linh mountain system, 2380 m, 9 Mar.
1995, L. Averyanov, N. T. Hiep VH 596' (holo¬
type, HN; isotypes, AUU, LE, MO, P). Figure 1.

1 Index VII (abbreviation of words Vietnamese High¬
lands) is used in the numbering of herbarium collections
that were made during fieldwork associated with the U.S.
National Geographic Society exploration program “1 lora
of Highlands of South Viet Nam.'

436

Novon

Inflorescentia brevis, simplex (eramosa), spiciformis, 5-
8 cm; inflorescentiae rhachis crassa, basi dense pilosa;
flores albido-flavides, magni, 10—12 mm in diametro, dis¬
cus florum cum 5 crassis excrescentiis; stylus disperse
pilosus — species nova a I). stachyantho (sola antea cog-
nita species generis) dignoscor.

Tree 15—25 m tall. Branehlets glabrous, apically
with numerous spirally arranged leaves appearing
verticillate within shortened internodes of the
branch. Leaves simple, entire, petioled, coriaceous,
bright glossy green and shiny, glabrous. Petioles
1-5-3 cm long and 2-3 mm thick. Leaf blade nar¬
rowly obovate, elliptic or oblong-lanceolate, 8—18
cm long, 3—6 cm wide, base narrowly cuneate, apex
shortly attenuate and rounded. Inflorescences axil¬
lary (usually appearing as subterminal or terminal),
spike 5—8 cm long, rachis 3—5 mm thick, densely
finely pubescent throughout its length with short
appressed white hairs. Flowers articulated, sessile,
actinomorphic, 5-merous, with 5 free petals, 5 ca¬
lyx teeth, and 10 stamens, 10—12 mm diam., yel¬
low-white, fragrant. Calyx broad, with 5-dentate
limb about 6 mm diam.; calyx lobes tooth-like,
short, with broad base and acute tip. Petals rigid
to coriaceous, with indexed lips in bud and in open
flowers, ovate with broad base, narrowed toward the
acute apex, about 5 mm long, 2.5-3 mm wide, with
prominent broad ovate hairy callus at basal half on
adaxial surface. Stamens shorter than petals, with
thick filaments; anthers with 2 longitudinally de¬
hiscent cells. Ovary inferior, 1 -celled, broadly ob¬
ovate, about 4 mm long, densely pubescent outside
with fine appressed white hairs. Style 1, undivided,
conical, with broad pubescent base, narrowing to¬
ward punctiform stigma, surrounded at the base
with 5 thick bilobed glabrous bosses. Fruits drupe¬
like, about 3—4 cm long, 1.6—2 cm wide, with very
hard, durable, woody endocarp of pyrene.

I he new species differs from l). stachyanthus by
its short, simple, spike-like inflorescence 5—8 cm
long, a densely pubescent basal portion of the in¬
florescence rachis, large yellow-white flowers about
10-12 mm in diameter, the presence of 5 promi¬
nent bosses on the flower disc, and a sparsely pu¬
bescent style. Diplopanax vietnamensis is known
from the mountainous regions of Kon Turn Province
in southern Vietnam, where it grows in wet ever¬
green broad-leaved montane closed primary forests
along ridges and on mountain summits at elevations
of 1000—2500 m on soils developed predominantly
on silicate sandstones, granite, and gneiss. Some¬
times this species was observed as a significant

component of the secondary forest stratum. This
tree flowers in March-May.

Paratypes. VIETNAM. Prov. Kontuni: cloud ever¬
green primary forest on N slope of Ngoc Linh mountain
system at 2400-2450 m, 6 Mar. 1095, L. Averyanov & /V.
7.' Hiep VH 561 (AUU, HN, EE, MO, P); Kon I’long Distr.,
Ifieu Municipality, Mang La forest enterprise (I4°30'N,
I08°25'E), 15 Apr. 2(MX), L. Averyanov , P. K. hoc, P. H.
Hoang, I). X. Du & N. T. Vinh VH 5111 (AUU, HN, LE,
MO).

Acknowledgments. Field studies in Vietnam,
the results of which are presented in this paper,
were funded by grants from the U.S. National Geo¬
graphic Society (grants 5094-93, 5803-96, 6383-
98 “Flora of Highlands of South Viet Nam”). We
thank I). K. Harder for comments and kind help in
improving the English.

Literature Cited

Axelrod, I). I., I. Al-Shehbaz & P. H. Raven. 1998. His¬
tory of the modern flora of China. Pp. 43-55 in Floristic
Characteristics and Diversity of East Asian Plants. Chi¬
na Higher Education Press, Beijing, and Springer- Ver-
lag, New York.

Eyde, R. H. & Q-N. Xiang. 1990. Fossil mastixioid (Cor-
naceae) alive in eastern Asia. Amer. J. Bot. 77: 689-
692.

Handel-Mazzetti, 11. 1933. Plantae novae Chingianae.

Pars. 3. Sinensia 3(8): 18.5—198.

Hoo, G. & C. Iseng. 1978. Araliaceae. Id. Reipubl. Pop¬
ular. Sin. Vol. 54.

Kubitzki, K. & Vi. Krutzseh. 1998. Origin of east and
south east Asian plant diversity. Pp. 56—70 in Floristic
Characteristics and Diversity of East Asian Plants. Chi¬
na Higher Education Press, Beijing, and Springer- Ver-
lag. New York.

Martinetto, E. 1998. East Asian elements in the Plio-
Pleistocene floras of Italy. Pp. 71-87 in Floristic Char¬
acteristics and Diversity of East Asian Plants. China
Higher Education Press, Beijing, and Springer- Verlag,
New York.

Nguyen, N. T. 1998. The Fansipan flora in relationship to
the Sino-Japanese floristic region. Pp. 111-122 in I).
E. Boufford N II. Obha (editors), Sino-Japanese Flora:
Its Characteristics and Diversification. University of To¬
kyo, Tokyo.

& T. L Nguyen. 1998. Diversity of vascular plants
of high mountain area: Sa Pa-Phan Si Pan. Hanoi. Nha
Xuat Ban Dai Hoc Quoc Cia Ha Noi. Jin Vietnamese. |
Iseng, C. 1983. The systematic position of Diplopanax
Hand.-Mazz. Acta Phytotax. Sin. 21(2): 151-152.

\\u. Z-Y. & S-G. Wu. 1998. A proposal fora new floristic
kingdom (realm) — I he E. Asiatic Kingdom, its delin¬
eation and characteristics. Pp. 3—42 in Floristic Char¬
acteristics and Diversity of East Asian Plants. China
Higher Education Press, Beijing, and Springer- Verlag,
New York.

Zhang. Z-Y. & A-M. Lu. 1998. On the origin, differenti¬
ation and dispersal of the Hamamelidaceae. Pp. 137—
153 in Floristic Characteristics and Diversity of East
Asian Plants. China Higher Education Press. Beijing,
and Springer- Verlag, New York.

Packera lyallii , a Corrected Name for Packera contermina
(Asteraceae: Senecioneae)

John F. Bain

Department of Biological Sciences, University of Lethbridge, Lethbridge, Alberta,
Canada T1K 3M4. Bain@uleth.ca

Abstract. the new combination Packera lyallii
(Klatt) J. F. Bain is proposed, based on Senecio lyal¬
lii Klatt.

Key words: Asteraceae, nomenclature, Packera,
Senecio, Senecioneae.

The segregate genus Packera A. Love & 1). Love
(= the aureoid Senecio complex) will be recognized
in the upcoming treatment of the Asteraceae for the
Flora of North America (North of Mexico). I his
change in treatment has necessitated a number of
recent nomenclatural transfers (e.g., Bain, 1999;
Track & Barkley, 1998, 1999). The name change
proposed herein is necessitated by a recent change
in the rules of nomenclature contained in the In¬
ternational Code of Botanical Nomenclature (St.
Louis Code) (1CBN) (Greuter et al., 2000).

The publication of the latest ICBN (2000), the
St. Louis Code, introduced a new rule (Article 53.6
Note 3, Ex. 20) that states that a rejected homonym
“remains legitimate and takes precedence over a
later synonym of the same rank should a transfer
to another genus (or species) be effected. ’ This
change has rendered Packera contermina, the re¬
cent new combination proposed by Bain (1999), su¬
perfluous. The basionym, Senecio lyallii, is legiti¬
mate in Packera according to the new
aforementioned rule. I herefore, the following new
combination is proposed:

Packera lyallii (Klatt) J. F. Bain. comb. nov. Bas-
ionym: Senecio lyallii Klatt, Ann. K. K. Na-
turhist. Hofmus. 9: 365. 1894, not Senecio
lyallii Hooker f. Senecio conterminus Green-
man, Ann. Missouri Bot. Card. 3: 101. 1916.
TYPE: Canada. Summit of the Rocky Moun¬
tains, 1861, Dr. Lyall s.n. (holotype, B? not
seen; isotype, GH).

Following the same rule, the taxon commonly
treated as Packera streptanthifolia (Greene) Weber
& Love (based on Senecio streptanthifolius Greene,
a name replacing the later homonym Senecio cym-
halarioides Nuttall) should hereafter be treated as
Packera cymbalarioides (Nuttall) Weber & Love, a
combination published in 1981.

Literature Cited

Bain, J. F. 1999. New combinations in Packera (Astera¬
ceae) and lectotypification of Packera ovina. Novon 9:
457-159.

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 & I). L. Ilawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Trock, D. K. & T. M. Barkley. 1998. Seven new nomen¬
clatural combinations and a new name in Packera (As¬
teraceae: Senecioneae). Sida 18: 385—387.

- Si - . 1999. Packera subnuda comb, nov., a

corrected name for Packera buekii (Asteraceae: Sene¬
cioneae). Sida 18: 635.

Novon 12: 437. 2002.

Additional Notes on Chinese Listera (Orchidaceae)

Singchi Chen (Xinqi Chen) and Yibo Luo

Laboratory of Systematic & Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, Nanxinchun 20, Xiangshan, Beijing 100093, Peoples Republic of China

Guanghua Zhu

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

Abstract. Two new species from China. Listera
microphylla S. C. Chen & Y. B. Luo and L. fangii
'Iang & Wang ex S. C. Chen & G. Zhu. are de-
scri bed and illustrated. Listera megalochila (S. C.
Chen) S. C. Chen & G. Zhu is elevated in status
from varietal to species level based on characters
of the floral lip. Their affinities with other related
species of Listera are briefly discussed.

Key words: China, Listera, Orchidaceae.

Listera It. Brown is a genus of ca. 35 species,
mainly distributed in boreal and temperate regions
of the Northern Hemisphere, with a few species
extending to Taiwan and Vietnam to the south. The
genus is easily distinguishable by the single pair of
simple, opposite, leaves with secondary venation
reticulate near the middle of the plant and a flower
with a bilobed lip that is commonly much larger
than the sepals and petals. Twenty-one species and
four varieties have been previously reported from
China (Chen & Luo, 1999).

In preparing a revised manuscript of this genus
for the Flora of China (English edition), the spec¬
imens preserved in PE have been carefully re-ex¬
amined, and the following additions and changes
are made:

1. Listera microphylla S. C. Chen & Y. B. Luo,
sp. nov. TYPE: China. NW Yunnan: Gong
Shan Co., 2500 m, Oct. 1935, C. IF. Wang
67264 (holotype, PE). Figure 1A, B.

Species nova insignis, L. mucronatae subsimilis, a qua
bene differt planta parvula 5-1 1 cm alta, caule aerio brev-
issimo (6—20 mm longo), foliis multo minoribus 6—3 X 7—

9 mm, fioribus minoribus.

Terrestrial herb, 5-11 cm tall; rootstock short,
with several ± fleshy roots. Aerial stem 6—20 mm
long below the single pair of leaves, glabrous.
Leaves 2, opposite, sessile, orbicular-ovate, 6—8 X
7-9 mm, obtuse-acute at apex, slightly fleshy, sec¬
ondary venation densely reticulate, covered with
minutely glandular hairs on both sides, the margin

Novon 12; 438-440. 2002.

nearly entire or slightly undulate. Peduncle 2.6-6
cm long, glabrous, sometimes with a sterile bract
in upper part; raceme 1.6-3 cm long, glabrescenl
or loosely glandular-hairy on rachis, 3- or 4-flow-
ered; bracts subovate or ovate-lanceolate, 3-4 X 2
mm; pedicel 1.5-2 mm long, slender, glabrous; ova¬
ry almost as long as the pedicel, glabrous. Flowers
greenish white, glabrous; dorsal sepal oblong-ovate,
ca. 3.5 X 1.5 mm, I -nerved, obtuse at apex; lateral
sepals lanceolate-ovate, slightly oblique, similar in
size to dorsal one, carinate distall v; petals similar
to dorsal sepal, but a little shorter and narrower;
lip obovate-cuneate in outline, ca. 5.2 mm long, 3.5
mm wide in upper part, 1 mm wide near the base,
bilobed at apex, with an indistinct keel from the
base terminating in a short mucro in the sinus;
lobes broadly oblong, ca. 1.4 X 1.6 mm, 3-nerved,
the margin nearly entire; column suberect, 1.5-2
mm long; anther ca. 1 mm long. Capsules ellipsoid,
erect, ca. 3 mm long, 2.5 mm thick.

Distribution. I his species is known only from
the type collection at an altitude of 2500 m on a
grassy slope.

I his species is similar to Listera mucronata Pan-
igrahi & J. ,). Wood, but differs in having a much
shorter stem and much smaller leaves. It is close
to L. brevicaulis King & Pantling, but differs in hav¬
ing a slightly pubescent scape and small auricles
at the base of the lip. Listera micrantha may also
be close to the newly described species but has a
short, trilobed lip and pubescent scape.

2. Listera fangii Tang & Wang ex S. C. Chen &
G. Zhu, sp. nov. 1 V PE: China. Sichuan: Guang
Xiang [Dujiangyanj, 800-1000 m, 14 July
1928, W. F. Fang 2204 (holotype, PE). Figure
1C, D.

Species L. nipponicae affinis, a qua imprimis differt pe-
dunculo et racemo fere usque ad 20 cm longo quam caule
multo longiore, labello infra medium unguiculato, auri-
culis late ovato-deltoideis.

Volume 12, Number 4
2002

Chen et al.
Chinese Listera

439

Figure 1. A. B. Listera mkrophylla S. C. Chen & Y. B. Luo. -A. Plant. B. Flower. C I). Listera fangii Tang &
Wang ex S. C. Chen & G. Zhu. — C. Plant. —I). Flower. — F. Listera megalochila (S. C. Chen) S. C. Chen & G. Zhu,

flower.

10 mm

440

Novon

Terrestrial herb, ea. 29 cm tall; rootstock short,
with several somewhat fleshy roots. Aerial stem
10.5 cm long below the single pair of leaves, gla¬
brous, enclosed in basal part by a tubular sheath
over 2 cm long. Leaves 2, opposite, subsessile,
ovate-orbicular, 2.1-2.8 X 2.8-3.0 cm, rounded-
acute at apex, broadly cuneate-subcordate at base,
minutely puberulous on upper surface, entire-mar¬
gined. Peduncle up to 15.3 cm long, covered with
glandular hairs, usually with a sterile bract in lower
part; raceme ca. 4.5 cm long (not completely ma¬
ture), glandular-hairy on rachis, 1 I -flowered; bracts
subovate-oblong, 1. 5-2.5 X 0.8 mm, ± glandular-
hairy; pedicel 3. 5-4.5 mm long, glandular-hairy;
ovary ca. 2 mm long, loosely granular-hairy or gla-
brescent. Flowers glabrous; sepals ovate-oblong, ca.
2 X 0.8 mm, obtuse at apex, 1 -nerved, lateral ones
slightly oblique; petals similar in outline to dorsal
sepal but slightly narrower; lip ca. 5 mm long, be¬
low the middle contracted into a claw ca. 2.8 X
0.8 mm, and on either side with a broadly ovate-
deltoid auricle ca. 0.8 mm long; the upper part it
fan-shaped, ca. 2X3 mm, bilobed deeply to half¬
way; column suberect, ca. 2 mm long.

Distribution. Known only from the type locality.

1 his new species is similar to Listera nipponica
Makino hut is easily distinguishable by its unusual
floral lip, which is composed of a basal claw and
an apical limb, spl it flower from peduncle, the pe¬
duncle elongate and much longer than either aerial

stem or raceme, and leaf upper surface minutely
puberulous.

3. Listera megalochila (S. C. Chen) S. C. Chen
& ( j . Zhu, stat. nov. Basionym: Listera gran-
diflora Rolfe var. megalochila S. C. (then, Acta
Phytotax. Sin. 25(6): 473. 1987. TYPE: China.
Sichuan: Da jin Xian, 2800 m, 27 June 1958,
X. Li 77925 (holotype, PE). Figure 1 E.

Listera megalochila is similar to L. grandiflora
Rolfe in having large flowers, lips ca. 1.6-1. 7 cm
long and deeply bifid, sinus entire, and columns 5—
t mm long, ami was first published as a variety of
the latter species. However, the floral lip obovate-
oblong, 5 mm wide at base, more than half of lip
width, and presence of a basal V-shaped nectary in
L. megalochila qualifies its specific status in the
genus; in L. grandiflora the floral lip is obovate-
cuneate, 2—3.5 mm wide at base, less than half of
lip width, and without nectar.

Distribution. 1 his species is known from for¬
ested slopes in Dajin in Kangding Xian in western
Sichuan. It is usually found above 2800 m.

Acknowledgments. Ilns study was supported by
the State Key Basic Research and Development
Project ((7200001680 1 ). We thank Cai Shuqin for
the I ine drawings.

Literature Cited

Chen, S. (.. & M. Luo. 1996. Listera. In K. Y. Lang
(editor). Flora Reipublicae Popularis Sinicae vol. 17.

Miscellaneous Notes on Sphagnum — 12

Howard Crum)

University of Michigan Herbarium, 3600 Varsity Drive,
Ann Arbor, Michigan 48108-2287, U.S.A.

ABSTRACT. Based on unique combinations of char¬
acters, five new species. Sphagnum [sect. Sphag¬
num | matogrossense. Sphagnum [sect. Sphagnum \
hertelianum. Sphagnum [sect. Sphagnum \ troll d ,
Sphagnum [sect. Suhsecunda] uruguayense, and
Sphagnum |sect. Acutifolia\ amazoneme, are added
to the rich Sphagnum flora of South America.

Key words: South America, Sphagnaceae,
Sphagnum.

For many years I have given special attention to
the genus Sphagnum in South America. I find it
frustrating that the genus is so abundantly repre¬
sented there, particularly in Brazil, and so poorly
known. I have taken a cautious approach to the
genus in North America, where the flora and its
ecological relationships are so well known. The
species are not very numerous, and most of them
can, in fact, be recognized in the field by aspect
and ecological niche. But I have found it necessary
to describe many new species from South America,
partly because collections are relatively few and
partly because I do not know the multiplicity of
habitats available to the genus or phytogeographic
relationships between the floras, hven in the case
of common species, every specimen has to be stud¬
ied in minute detail before deciding whether it has
been described before, and it is not possible to de¬
termine the range of variability from the relatively
few collections available. The species differ mainly
in microscopic structures that may indeed vary with
the habitat, but the kind of information given on
the labels is virtually meaningless. Even though 1
consider these five new species (and the many other
novelties that I have offered in description) fully
justified, further collection and study will be nec¬
essary to evaluate them.

Sphagnum [sect. Sphagnum | matogrossense
Crum. sp. nov. TYPE: Brazil. Ponte da Pedra:
Matto Grosso, June 1909. F. C. Hoehne 2147
(holotype, M). Figures 1-6.

Epidermis caulina stratis 3, subtiliter fibrosis vel efi-
brosis, parietes exteriores cellularum superficial! um fora-

mine uuo vel duo (raro tres); cylindrus lignosus atro-rufus.
Kolia caulina ad 1.4 mm longa, ad apicem fibrosa; cellulae
hyalinae superficie exteriore lacunis rotundis vel oblongis,
interiore lacunis nullis sed aliquando poris parvis in cel¬
lularum angulis. Kami duo; cellulae epidermidis subtiliter
fibrosae vel efibrosae, poris unis. Folia ramulina ad 3 mm
longa; cellulae hyalinae superficie exteriore 1—3 poris in
cellularum angulis conjunctis, interiore poris nullis; cel¬
lulae chlorophylliferae sectione transversali elliptieae vel
fusiformae, utroque latere foliorum liberae.

Pale, lawny plants up to about 6 cm high. Stem
cortex 3-layered, the epidermal cells short, delicate¬
ly fibrillose or not at all, 1— 2(— 3)-porose; wood cyl¬
inder very dark, blackish brown. Stem leaves pale
brown, ca. 1.4 mm long, lingulate and rounded at a
fringed apex but shortly concave-pointed; hyaline
cells undivided, fibrillose on both surfaces toward
the apex, sometimes as much as V4—V2 the leal length,
on the outer surface with 3-5 large, rounded, oblong,
or irregular gaps, on the inner surface without gaps
but occasionally with a small, ringed comer pore.
Branches in fascicles of 2 ( I spreading); cells of cor¬
tex delicately or not at all fibrillose, with I large
pore. Branch leaves somewhat spreading, 2.8—3 mm
long, broadly ovate-acute, deeply concave; hyaline
cells fibrillose on both surfaces, on the outer surface
with 1-3 end and corner pores, grouped in threes at
adjacent comers, commonly with membrane pleats,
on the inner surface with pores none; green cells
central and elliptic, exposed on both surfaces but
with thickened cell ends.

The plants resemble Sphagnum ouropretense C.
Muller & Warnstorf in having stems with a very
dark, blackish brown wood cylinder, but the stem
leaves have hyaline cells fibrillose only toward the
apex, branches in fascicles of two, and hyaline cells
of branch leaves commonly membrane-pleated.

Sphagnum [sect. Sphagnum] hertelianum Crum,
sp. nov. TYPE: Venezuela. Estado Bolivar: Kwei-
kin-ima Tepuy (= Ceiro Guaiquinima), 05°47'N,
63°51'W, 1500-1550 m, blanket bog with scat¬
tered rock outcrops and mossy dwarf forest, sub¬
merged, 13-15 Feb. 1990, //. Hertel 36,606 (ho¬
lotype, M; isotype. MICH). Figures 7—11.

+ Deceased 30 April 2002. I)r. Crum had essentially completed the manuscript for this article. With help from Steven
P. Churchill (MO), Crums manuscript was lightly edited in June 2002 by Marshall R. Crosby (MO).

Novon 12; 441-445. 2002.

442

Novon

Plantae parvae, columniformes, pallide incarnato-fus-
cae. Epidermis caulina et ramulina subtiliter vel baud fi-
brillosa. Folia caulina el ramulina similia; cellulae hyali-
nae superficie exteriore poris magnis ternis in angulis
conjunctis instructa, interiore poris et pseudoporis parvis
commissuralibus numerosis. Rami 2-fasciculati, subae-
t|uales. Folia ramulina cellulis chlorophylliferis sectione
transversal! ± triangulis, superficie interiore expositis.

Small, columnar, pale pinkish brown plants, up
to 4 cm high. Stems dark red-brown; cortical cells

in 2 layers, short, uniporose, delicately fibrillose or
non-fibrillose; wood cylinder blackish brown. Stem
leaves 2.5 mm long, oblong-1 ingulate, concave,
broad at the apex but concave-pointed, bordered
by a resorption furrow; hyaline cells non-septate,
fibrillose throughout, on the outer surface with
large, ringed pores in threes at adjacent angles and
also singly at side corners, on the inner surface with
ringed elliptic pores at corners and numerous

Volume 12, Number 4
2002

Crum

Notes on Sphagnum

443

smaller pores and pseudopores at commissures, of¬
ten in series of 2-5. Branches paired, subequal,
about 5 mm long, both directed upward and crowd¬
ed; epidermal cells uniporose, delicately fibrillose.
Branch leaves 2.5-3 mm long, oblong-ovate, ta¬
pered to the apex, concave, bordered by a resorp¬
tion furrow; hyaline cells similar to those of stem
leaves; green cells triangular, exposed on the inner
surface (and sometimes narrowly exposed on the
outer), the hyaline cells convex on the inner sur¬
face, bulging on the outer.

The plants have an interesting columnar form
owing to short branches crowded together and di¬
rected upward. The branches are in fascicles of two
and similar in size and form (neither pendent). The
cortical cells of stems and branches are weakly fi¬
brillose, if at all. The stem and branch leaves are
similar. They have, on the outer surface, large,
ringed, elliptic pores conspicuously in threes at ad¬
jacent angles and, on the inner surface, a few rather
large corner pores in addition to numerous small
pores and pseudopores at the commissures, often
in series of 2 to 5.

Sphagnum [sect. Sphagnum] trollii Crum, sp. nov.
TYPE: Colombia. Paramo de Boqueron bei Bo¬
gota, 3900 m, C. Troll (holotype, M). Figures
12-16.

Plantae ca. 4 cm altitudine, fulvae. Caules subfusci;
cellulae epidermidis caulinae fibrillosae, 1— 3-porosae; cy-
lindrus lignosus obscure rufo-brunneus. folia caulina eis
ramulinis similia sed plus lingulata, 1.5— 1.8 mm longa;
cellulae hyalinae fibrillosae, non septatae, superficie ex-
teriore poris commissuralibus elliptic i s (sursum ca. 5,
deorsum 8-9), sub apicibus laeunis magnis, interiore pler-
umque poris nullis. Folia ramulina 2—2.2 mm longa, ovata;
cellulae ehlorophylliferae orciformes, superficie utroque
anguste expositae.

Plants tawny, up to 4 cm high. Stems brown; cor¬
tex 3-layered, the epidermal cells fibrillose, 1-3-
porose; wood cylinder dark red-brown. Stem leaves
much like branch leaves in structure, though small¬
er. 1.5 mm long and more nearly lingulate, bor¬
dered by a resorption furrow; hyaline cells fibril¬
lose, undivided, on the outer surface with elliptic
commissural pores, about 5 toward the apex and 8
or 9 below, near the apex with large gaps, on the
inner surface with scarcely any pores. Branches in
fascicles of 3 (2 short and spreading); cells of the
epidermis uniporose, efibrillose or some few cells
fibrillose. Branch leaves 2-2.2 mm long, ovate; hy¬
aline cells with fewer pores but otherwise similar
to stem leaves; green cells orciform, narrowly ex¬
posed on both surfaces, more so on the outer.

The dark wood cylinder of stems and similar

stem and branch leaves are especially interesting.
Sphagnum lapazense Crum also has a dark wood
cylinder and similar stem and branch leaves, hut
the stem cortex is only 1-layered and lacks pores
and fibrils, and the leaves have only a few pseu¬
dopores on the outer surface hut numerous true
pores on the inner.

Sphagnum [sect. Subsecunda] uruguayense
Crum, sp. nov. TYPE: Uruguay. Depto. Mon¬
tevideo, Carrasco, in ripariis aren.-arg., 25
Oct. 1934, W. G. Herter 1432a (holotype, M).
Figures 17—24.

Hyalodermis caulis strato uno, cellulis parvis, indis-
tincte diversis; cylindrus lignosus brunneus. Folia caulina
1.5-1. 8 mm longa, oblongo-ovata, rotundo-obtusa; cellu¬
lae hyalinae superficie exteriore numerosae, interiore pau-
cae. Fasciculus ramorum ramis 3 (2 patentibus). Folia ra¬
mulina 2 mm longa, oblongo-ovata vel oblongo-lanceolata,
concavo-acuminata; cellulae hyalinae superficie exteriore
poris commissuralibus numerosis, interiore poris paucis;
cellulae ehlorophylliferae anguste triangulo-trapezoideae,
sectione transversali superficie exteriore anguste exposi-
lae.

Plants rather small, yellowish green. Cortical
cells of stem small and inconspicuous, in I layer;
wood cylinder brown. Stem leaves 1.5— 1.8 mm
long, oblong-ovate, rounded-obtuse, somewhat con¬
cave above, narrowly bordered; hyaline cells fibril¬
lose in the upper V2—2A, occasionally once- or even
twice-divided, on the outer surface with as many as
8 commissural pores, on the inner surface with
pores few, small, and rounded. Branches in fasci¬
cles of 3 (2 spreading). Branch leaves not at all
secund, 2-2.1 mm long, oblong-ovate or oblong-
lanceolate, concave; hyaline cells not or rarely di¬
vided, on the outer surface with numerous small,
rounded-elliptic commissural pores in nearly con¬
tinuous rows, on the inner surface with few small,
rounded pores; green cells in section narrowly tri¬
angular-trapezoidal. narrowly exposed on both sur¬
faces, more so on the outer.

Herter distributed this material as S. medium
Limpricht var. affine, mini, inval., and in aspect it
indeed resembles the section Sphagnum to which
that species belongs. However, it has none of the
diagnostic features of that section, and the small
pores of both stem and branch leaves are commis¬
sural in arrangement and numerous, as in most of
section Subsecunda. Notable features include the
unusually small cortical cells of the stem as well
as the relatively few small, rounded pores on the
inner surface of both stem and branch leaves and
more numerous, small, rounded-elliptic pores on
the outer surface, those of the branch leaves in
nearly continuous commissural rows.

444

Novon

Figures 12-23. Sphagnum trollii and .S', uruguayense. 12-16. S. trollii. —12. Branch leaves, X30. —13. Upper cells

of branch leaf, outer surface, X320. — 14. Portion of branch leaf in section, X320. — 15. Stem leaves, X30. _ 16.

Upper cells of stem leaf, outer surface, X320. 17-23. Sphagnum uruguayense. — 17. Branch leaves, X30. — 18. Upper
cells of branch leaf, outer surface, X320. —19. Upper cells of branch leaf, inner surface, X320. —20. Portion of

branch leaf in section, X320. — 21. Stem leaves, X30. — 22. Upper cells of stem leaf, outer surface, X320. _ 23.

Upper cells of stem leaf, inner surface, X320.

Sphagnum [sect. Acutifolia ] amazonense Crum,
si), nov. TYPE: Peru. Amazonas: Chachapoyas
District, Las Palmas zw. Balsas u. Leimemba,
06°45'S, 77°49'W, 3000 m, 31 Aug. 1973, P.
& E. Hegewald 6996 (holotype, MO). Figures
24-28.

Plantae fulvellae, ea. 6 cm altae. Cellulae epidermidis
eaulinae sine poris; cylindrus lignosus rutilus. Folia cau-
lina 1 .2 mm longa, subtriangularia, parce concava, valde
limbata, deorsum ± dilatata; cellulae hyalinae efibrillo-
sae, plerumque I (-2)-septatae, utroque superficie lacuna
magna perforata. Folia ramulina 1. 1-1.2 mm longa, an-
guste ovata, concava, ca. 3 seriebus cellularum elongatar-

Volume 12, Number 4
2002

Crum

Notes on Sphagnum

445

Figures 24—28. Sphagnum amazonense. — 24. Branch leaves, X30. — 25. Upper cells ol branch leaf, outer surface,
X320. —26. Upper cells of branch leaf, inner surface, X320. —27. Stem leaves, X30. —28. Upper cells of stem
leaf, inner surface, X320.

um limbata; cellulae hyalinae superficie exteriore poris
annulatis ellipticis numerosis ad commissuras, interiore
poris 1-3, rotundis, non annulatis; cellulae chlorophyli-
ferae sectione transversali triangulae. dorso foliorum ex-
positae; cellulae hyalinae utroque superficie valde con-
vexae.

Plants tawny, about 5 or 6 cm high. Stem cortex
without pores; wood cylinder red-yellow. Stem
leaves 1.2 mm long, ovate-triangular, acute, slightly
concave, heavily bordered by 4—5 rows of cells, the
border somewhat expanded toward the base; hya¬
line cells 1— 2-divided, efibrillose, with a large gap
on both surfaces. Branches in fascicles of 3 (1 pen¬
dent). Branch leaves 1. 1-1.2 mm long, narrowly
ovate, concave, bordered by about 3 rows of linear
cells; hyaline cells on the outer surface with nu¬

merous (up to 9) ringed, elliptic commissural pores,
on the inner surface with 1—3 rounded, unringed
pores; green cells triangular, exposed on the inner
surface, the hyaline cells bulging on both surfaces.

The plants have stem leaves that are more or less
triangular, acutely pointed, and very slightly con¬
cave; the hyaline cells are efibrillose and once or
twice divided and have large membrane gaps on
both surfaces. The branch leaves have rather nu¬
merous elliptic commissural pores on the outer sur¬
face and as many as three, rather large, rounded,
thin-margined pores (or gaps) on the inner surface;
the green cells are exposed on the inner surface,
and the hyaline cells are bulging on both surfaces.

Two New Species of Oxypetalum (Asclepiadoideae, Apocynaceae)

from Brazil

Maria Ana Farinaccio

Departamento de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, C.P. 11461,
05422-970, Sao Paulo, SP, Brazil, mafarinaccio@hotmail.com

Abstract. Oxypetalum habragynum and 0. he¬
llos, two new species of Asclepiadoideae, Apocy¬
naceae, are described and illustrated. Oxypetalum
habragynum has the membrane of the caudicles of
the pollinarium concave and not convex or plane
as in other species. Oxypetalum helios has the seg¬
ment of the corona 3-lobed, lacking an adaxial ap¬
pendage, and the gynostegium apex is filiform and
bifid at its distal third. I liese species are endemic
to the National Park of the Serra da Canastra in
the state of Minas Gerais, Brazil.

Key words: Apocynaceae, Asclepiadoideae,
Brazil, Oxypetalum.

Oxypetalum R. Brown (Asclepiadoideae, Apo-
cynaceae) is a Neotropical genus comprising about
100 species, occurring in a variety of habitats, prin¬
cipally in open areas like savannah, grassland, and
forest edges. Brazil is the center of diversity for the
genus, with a majority of species found within the
state of Minas Gerais. During the preparation of the
treatment for the flora of Serra da Canastra National
Park, Minas Gerais, in southeastern Brazil (Fig. 1),
two new species of Oxypetalum , 0. habragynum
and 0. helios, were recognized and are here de¬
scribed and illustrated.

Oxypetalum habrogynunt Farinaccio, sp. nov.

I \ PE: Brazil. Minas Gerais: Sao Roque de
Minas, Parque Nacional da Serra da Canastra,
estrada Sacramento— Sao Roque de Minas, ea-
poes apos cerradao, 21 Mar. 1998 (fl), M. A.
Farinaccio, P. T. Sana, A. C. Araujo, D. V. Arce
& P. Fiaschi 143 (holotype, SPF; isotypes, F,
MO). Figure 2.

Ah omnibus speciebus generis membrana pollinarii
concava uptime distinctum. Species quam maxime all in is
O. pachygyno, quae tamem eharacteribus inflorescentiae
florumque differ).

Vines, stems tomentose. Leaves opposite, patent;
petiole 1—2.2 cm long, somewhat sulcate, tomen¬
tose; blades 3.5-8(-l 1) X 2-3.2(-6) cm, oblong to
ovate-oblong, discolorous, puberulous to pubescent,
chartaceous, brochidodromous, apex rounded to

No VON 12: 446-450. 2002.

acute, mucronate, base cordate, with 2 to 5 colle-
ters at base ol adaxial side. Partial inflorescence
extra-axillary, alternate, 6- to 1 1-flowered. erect;
peduncles 0.5— 1.7 cm long, tomentose, bracts 1.5
X 0.5 mm, lanceolate, adaxial surface hirsute, ab¬
axial surface glabrous, with colleters at the base;
pedicels 2—6 mm long, pubescent to tomentose. Ca¬
lyx green to brown, divided almost to base, lobes
5—4-3 X 1-1.5 mm, lanceolate, apex acute, abaxial
surface pubescent to tomentose, adaxial glabrous,
l or 2 colleters below each sinus. Corolla abaxial
surface dark purple to green, pubescent, adaxial
white, cream or lightly pink, glabrous to puberu¬
lous, tube campanulate, 2-2.5 mm long, lobes 3.6-
5.5 X 2—2.1 mm, lanceolate, reflexed, margins hy¬
aline, apex aeule. Corona white, lobes 2.4— 3.2 X
1-1.7 mm, oblong to obovate, imbricate, the adaxial
surface with carunculate excrescences and a cen¬
tral tooth-like appendage, apex subtruncate to trun¬
cate, crenate. Gynostegium rostrate, white to lightly
pink, 1.5 mm long, 1.1 mm diam., sessile, apex
2. 5-4. 3 mm long, conical, bifid from the distal
third. Anthers 0.7— 1.5 X 0.6-1 mm, rectangular to
subquadrangular, terminal appendage 0.8-1 X
0.8-0. 9 mm, obscured by corona, ovate, apex acute,
wings longer than or equal to dorsum, straight. Cor-
pusculum 0.43-0.53 mm long, 0.1 1-0.13 mm wide,
lanceolate, thick-gibbous, apex acute, caudicles
0.13-0.19 mm long, flattened, broad, translucent,
with horny teeth 0.21-0.29 mm, pollinia 0.38-0.42
X 0.1 1-0.14 mm, oblong. Follicle black when se¬
nescent, 6 X 0.5 cm. Seeds unknown.

Oxypetalum habragynum is distinct from the oth¬
er species in the genus by the membrane of the
caudicles ol the pollinarium, which is concave and
not convex or plane as in other species. It is very
closely related to 0. pachygynum Decaisne, mainly
by vegetative characteristics, but may be distin¬
guished by characteristics of the partial inflores¬
cence and of the flowers (Table 1).

This species is endemic to the Serra da Canastra
National Park. It occurs in humid soil of grasslands,
around the borders of small forest tracts (“capdes”)
and gallery forests. It has been collected in flower

Volume 12, Number 4
2002

Farinaccio

Oxypetalum from Brazil

447

Figure 1. Map showing the Serra da Canastra National Park, Minas Gerais State, Brazil.

in March. July, and October, and with senescent
fruits in July. The specific epithet is an allusion to
its gynostegium, more delicate in comparison to the
gynostegium of O. pachygynum.

Paratypes. BRAZIL. Minas Gerais: Sao Roque <le
Minas, Parque Naeional da Serra da Canastra, estrada
para Sacramento, pr6ximo ao Corrego dos Passageiros, 20
Mar. 1995 (fl), J. /V. A 'akajima et al. 892 (HUFU); Corrego
do Barbara, 19 Oet. 1997 (H), J. N. Nakajima et al. 2972
(HUFU); estrada Saeramento-Sao Roque de Minas, Tres
Matinhas, 8 July 1999 (fl, fr), M. A. Farinaccio et al. 352
(SPF).

Oxypetalum helios Farinaccio, sp. nov. TYPE:
Brazil. Minas Gerais: Sao Roque de Minas,
Parque Naeional da Serra da Canastra, Cor¬
rego do Quilombo, Tres Matinhas, 8 July 1999
(fl, fr), M. A. Farinaccio , A. A. Araujo & F. P.
Gomes 351 (holotype, SPF; isotypes, HRCB, K,
MBM, MO, NY, RB). Figure 3.

Inter affines segmentis coronae 3-lobatis adaxiale ex-
appendiculatis et apiee gynostegii filiformi bifidoque diag-
noscitur.

Vines, stems pubescent. Leaves opposite, patent;
petiole 0.7— 1.3 cm, smooth, pubescent; blades 4.5—
8.5 X 1.7-3 cm, elliptic to lanceolate, sometimes
ovate, discolorous, subglabrate, minutely hairy
along the midvein and margins, chartaceous, with
brochidodromous venation, apex acute to obtuse,
mucronate to apiculate, base cordate with 2 colle-
ters at base of adaxial side. Partial inflorescence
extra-axillary, alternate, 2- to 9- (frequently 3)-flow-
ered, erect; peduncle 1.6— 2.2 mm long, pubescent,
bracts 2-3.3 X 0.5-0.7 mm, linear or oblong, pu-
berulous; pedicels 1-1.8 cm long, pubescent. Calyx
yellow to green, dark purple in the proximal region,
divided almost to base, lobes 2.5—6 X 0.5—1 mm,
lanceolate, apex acute, puberulous, 2 or 3 colleters

below each sinus. Corolla cream to yellow, cam-
panulate, tube 2.5-3 cm long, puberulous, lobes 1-
1.3 X 0.1-0. 2 cm, linear to narrowly oblong, patent
to reflexed, twisted, abaxial surface subglabrate,
adaxial surface pubescent, margins hyaline, apex
acute. Corona light green to cream, lobes 2—3 X

1.5- 1. 8 mm, oblong, 3-lobed, apex truncate,
curved. Gynostegium cream, 2.5 mm long, 0.8—1
mm diam., sessile, apex cream to lightly pink, 3—4
mm long, filiform, bifid from distal third. Anthers
0.72-1 X 0.5-0.72 mm. rectangular, terminal ap¬
pendage 1.1-1. 5 X 0.5-0.72 mm, oblong, apex
acute, emarginate, wings straight, longer than the
dorsum. Corpusculum 0.96-1.12 X 0.14—0.18 mm,
oblong, laminar, apex truncate, caudicles 0.11-
0.16 mm long, flattened, broad, translucent, with
horny teeth 0.19-0.24 mm long, pollinia 0.27-0.32
X 0.11-0.13 mm, oblong. Follicle green to brown,

5. 5- 8. 5 X 1.5-2. 5 cm, ovate, striate, puberulous.
Seeds 6-7 X 3-3.5 mm, ovate, verrueose.

Oxypetalum helios may be included in the 0.
insigne (Decaisne) Malme complex, principally on
the characteristics of habit (climbing plants) and
morphology of the pollinarium, which has the lam¬
inar corpusculum longer than pollinia. It is very
closely related to (). glahrum (Decaisne) Malme.
Both plants are sparsely pubescent, and they share
a similar leaf morphology and flower size. However,
0. helios is immediately distinguished from all spe¬
cies of that complex by the segment of the corona
3-lobed, lacking an adaxial appendage, and prin¬
cipally by the gynostegium apex, which is filiform
and bifid at its distal third.

This species is endemic to the Serra da Canastra
National Park. It occurs in humid soil around the
borders of small forest tracts (“capoes ). It has been
collected in flower and fruit in July, September, and

448

No von

Figure 2. Oxypetalum habrogynum Farinaccio. —A. Flowering branch. — B. Flower. — C. Adaxial surface of part of
the calyx showing colleters. — D. Adaxial surface of corona lobe. — E. Gynostegium. — F. Pollinarium: (a) corpusculum,
(b) caudicles, (c) pollinium; (a) + (b) = translator. — G. Senescent fruit. (A-F from the holotype, Farinaccio et al. 143 ;
G from farinaccio et al. 352.)

3 mm

Volume 12, Number 4
2002

Farinaccio

Oxypetalum from Brazil

449

Figure 3. Oxypetalum hellos Farinaccio. — A. Flowering branch. — B. Flower. — C. Adaxial surface of part of the
calyx showing colleters. — D. Adaxial surface of corona lobe. — E. Corona with lobe removed showing gynostegium.
— F. Pollinarium. — G. Fruit. — H. Seed. (From the holotype, Farinaccio et al. 351.)

450

Novon

Table 1. Differential characters between 0. habrogynum and 0. pachygynum.

Species

Characters

0. habrogynum

0. pachygynum

2-5

Rower number in the partial
inflorescence

Number of colleters in the calyx sinus

Color of the corolla

Tube length

Corolla lobe length

Abaxial face of the corolla lobes

Corona color

Gynostegium color

Gynostegium size

Gynostegium apical apex length

Anther wings

Terminal anther appendage

6-11

1-2

white, cream, or slightly pink

2—2.5 mm

3.6— 5.5 mm

glabrous to puberulous

white

white to slightly pink
1.5 X 1.1 mm
2. 5-4.3 mm
entire

0.8—1 mm, shorter than the corona

4

green to dark purple

2.5- 4 mm

6.5— 10.2 mm
gray-sericeous
green cream
dark purple

2—3 X 1.7-2 mm
4—6 mm
dentate

2.2 mm, exceeding the corona

October. The specific epithet is an allusion to its
yellow (lowers, which with their very conspicuous,
extremely narrow twisted lobes resemble the sun.

Paratypes. BRAZIL. Minas Gerais: Sao Roque de
Minas, Parque Nacional da Serra da Canastra, Corrego do
Quilombo, 26 Sep. 1995 (fl, fr), R. Romero et al. 2803
(HUFU, SPF); 20°10'17"S, 46°39'52"W, 14 July 1997 (fl,
fr), J. A. Lombardi 1862 (BHCB); Corrego do Quilombo,
lies Matinhas, 15 Oct. 1997 (fl, fr), J. l\l. Nahajima et al.
2869 (HUFU, SPF).

Acknowledgments. This work is part of a mas¬

ter’s thesis prepared under the guidance of Renato
Mello-Silva, at the Universidade de Sao Paulo
(USP). It was supported by Fundayao de Amparo it
Pesquisa do Estado de Sao Paulo (FAPESP) and
Conselho Nacional de Desenvolvimento Cientffico
e Tecnologieo (CNPq). I am grateful to Renato Mel¬
lo-Silva for the Latin diagnoses and helpful sug¬
gestions, Alexandre Carnier Nunes da Silva for pre¬
paring the map, and Henrique Graciano Muracho
for correcting the orthography of Oxypetalum helios.
The line drawings were prepared by Rogerio Lupo.

Two New Species of Ainsliaea (Asteraceae, Mutisieae) from China

and Vietnam

Susana E. Freire

Division Plantas Vasculares, Museo de La Plata, Paseo del Bosque, 1900 La Plata,
Argentina. freire@museo.fcnym.unlp.edu.ar

Abstract. Two new species, Ainsliaea pentaflora
from Tonkin, Vietnam, and Ainsliaea qianiana from
Yunnan. China, are described and illustrated. Ain¬
sliaea pentaflora is distinguished by its 5-flowered
capitula, rounded anther appendages, and rosulate
leaves with practically wingless petioles. Ainsliaea
qianiana is distinguished by its apiculate anther
appendages, rosulate leaves with petioles winged,
and 3-flowered capitula. These species are com¬
pared with closely related species. Included are
keys to those Ainsliaea with apiculate anther ap¬
pendages and 5-flowered capitula.

Key words: Ainsliaea, Asteraceae, China, Com-
positae. Mutisieae, Vietnam.

The genus Ainsliaea DC. (including Diaspanan-
thus Miquel) contains about 50 species distributed
in southeast Asia. All are perennial herbs with ro¬
sulate leaves or leaves arranged at the median part
of the stem, few-flowered capitula in spikes or pan¬
icles, slightly zygomorphic, deeply 5-lobed corollas
with unilateral lobes, and plumose pappus bristles.
The most closely related genera, Macroclinidium
Maximowicz, Myripnois Bunge, and Pertya Schultz
Bipontinus, usually differ by their scabrid, non-plu-
mose pappus bristles (Bremer, 1994).

During the course of a taxonomic revision of the
genus Ainsliaea by the author, the following two
species were recognized as undescribed.

Ainsliaea pentaflora S. E. Freire, sp. nov. TYPE:
Vietnam. Tonkin: Massif du Tom Dav, 900 m,
Dec. 1921. A. Petelot s.n. (holotype, C). Fig¬
ure 1.

Herba perennis, scaposa. Folia rosulata, longipetiolata;
lamina ovata, basi rotundata, leviter decurrentis, apice
acuta, in margine mucronato-denticulata, membranacea,
supra seabridula parce villosa vel glaberrima, subtus vil¬
losa, pinnativenia; petioli non alati, villosi. Folia inter in-
florescentiam pauca, elliptica. Capitula pentaflora, sessi-
lia, solitaria vel 2-fasciculata in panicula disposita.
Involucrum cylindrici-campanulatum. Phyllaria 6— 7-ser-
iata, coriacea, apice villosa. Flores isomorphi, omnes her-
maphroditi; corollae tubuliformes, 5-lobatae, cum lobis
unilateralibus; antherae apice rotundatae, basi sagittatae,

breviter laciniatae; styli rami obtusi. Achaenia villosa;
pappi setae uniseriatae, plumosae.

Plants perennial, herbaceous, ca. 0.35 m tall.
Stems erect, unbranched, villous. Leaves basally
clustered, rosulate, long-petiolate; petioles wing¬
less, 6.5—9 cm long, villous; leaf blades charta-
ceous, ovate, 7.5— 8.5 X 3.5—4 cm, apex acute, base
rounded, then cuneately narrowed into the petiole,
margins denticulate-mucronate; adaxial surface
slightly scabrous, sparsely villous to glabrous, tri-
chomes multicellular-unbranched (consisting ofun-
iseriate pedestal and long apical cells oblique sep¬
tate), ca. 0.7—1.25 mm long, abaxial surface villous,
veins somewhat more densely pubescent than rest
of abaxial surface; venation pinnate, secondary
veins 2 or 3 on each side of midrib, arcuate, the
secondary veins and midrib prominent. Cauline
leaves few, much reduced, ovate-elliptic. Capitula
sessile, solitary or 2, disposed in panicles, nodding
in maturity. Involucre 6- or 7-seriate, narrowly
campanulate, 13—15 X 12—14 mm; phyllaries co¬
riaceous. apically villous; outer phyllaries oblong-
ovate, 3-4 X 1.5 mm, the apices acute; inner phyl¬
laries linear-elliptic, 10-10.5 X 1.5— 1.7 mm. the
apices acute. Florets 5, hermaphroditic, corollas
glabrous, apically papillose, tubular, 4.5— 5.5 mm
long, 5-lobed. lobes ca. 1 mm long, with one deeper
slit ca. 1.5 mm, the corolla then pseudoligulate but
without an expanded limb, tube ca. 3.5 mm long.
Anthers 5, ca. 3 mm long, apical appendages
rounded, ca. 0.4 mm long, sagittate; tails 0.8—1 mm
long, shortly pilose. Style branches short, obtuse,
ca. 1.2 mm long. Achenes oblong, 7.5—8 mm long,
pilose. Pappus bristles uniseriate, ca. 39, plumose,
7.5—8 mm long, white.

The new species is known only from the type
collection from Tonkin, Vietnam, at approximately
900 m.

Ainsliaea pentaflora most closely resembles A.
latifolia (D. Don) Schultz Bipontinus. Both have
ovate leaves arranged at the base of the stem and
rounded apical anther appendages. However. A.
pentaflora differs from A. latifolia by its 5(vs. 3)-

Novon 12: 451-455. 2002.

452

Novon

Figure 1. Ainsliaea pentajiora S. E. Freire. — A. Habit. — B. Foliar trichome. — C. Phyllaries. — D. Capitula. — E.
Corolla. — F. Floret. — G. Style branches. — H. Stamen. Scale bars: A - 3 cm; B = 0.2 mm; C = 3.5 mm; 1) & F =
5 mm; E = 1.5 mm; G = 2.5 mm; H = 1 mm. Drawn from the holotype: Petelol s.n. (C).

Volume 12, Number 4
2002

Freire

Ainsliaea from China and Vietnam

453

flowered capitula, and its practically wingless (vs.
winged) petioles.

The new species epithet pentaflora refers to the
5-flowered capitula. Two other species of the genus
are also 5-flowered: A. brandisiana Kurz from
Myanmar (= Burma) and A. spanocephala Y. C.
Tseng from Thailand.

Kky to tup: Ainsliaea Species with 5-Flowered
Capitula

la. Leaf bases cuneate; anther appendages apically

rounded . A. pentaflora S. E. Freire

lb. Leaf bases rounded or cordate; anther append¬
ages apically acuminate.

2a. Leaf bases rounded . A. brandisiana Kurz

2b. Leaf bases cordate .

. A. spanocephala Y. C. Tseng

Ainsliaea qianiana S. E. Ereire, sp. nov. TYPE:
China. NE Yunnan: Colline de kin-tchong-
chan, 2990 m, E. E. Maire 2529 (holotype, W;
isotypes, NY. UC). Figure 2.

Herba perennis, scaposa. Folia rosulata, longipetiolata,
cum petiolo late alato; lamina elliptica vel ovala, in petiolo
subito angustata, apice acuta, in margine mucronato-den-
ticulata et villosa, crasse, utrinque glabra, subtus albida,
pinnativenia. Folia inter inflorescentiam pauca, lineari-el-
liptica. Capitula triflora, numerossisima, sessilia, 3-5-fas-
ciculato-spicata, horizontalia vel pendula, unilateralia,
spica simplex vel breviter ramosa. Involucrum cylindri-
cum. Phyllaria 6— 7-seriata, papyracea, omnes glabra. Flo¬
res isomorphi, omnes hermaphroditi; corollae rubescentes,
5-lobatae, cum lobis unilateralibus, linearibus, partitis
subaeque; antherae apice apiculatae. basi sagittatae, brev¬
iter laciniatae; styli rami breves, rotundati. Achaenia vil¬
losa; pappi setae uniseriatae, plumosae.

Plants perennial, herbaceous, 0.50-0.60 m tall.
Stems erect, unbranched, moderately villous.
Leaves basally clustered, rosulate, long-petiolate;
petioles winged, 10-11 cm long; leaf blades sub-
carnose, elliptic to elliptic-ovate, 7.5— 8.5 X 3.5—
3.8 cm, apex acute to obtuse, base abruptly round¬
ed and decurrent into the petiole, margins
denticulate-mucronate and villous, trichomes mul-
ticellular-unbranched (consisting of a uniseriate

pedestal and long apical cells obliquely septate),
ca. 0.6—1 mm long; upper and lower surfaces gla¬
brous, the surface whitish below; venation pinnate,
secondary veins 2 or 3 on each side ol midrib, ar¬
cuate, secondary veins and midrib prominent. Cau-
line leaves few, much reduced, linear-elliptic. Ca¬
pitula sessile, 3- to 5-fasciculate, rarely 1 or 2.
spicately disposed, nodding in anthesis, spike uni¬
lateral, single- or few-branched. Involucre 6- or 7-
seriate, cylindrical, 12-14.5 X 3—4 mm in anthesis;
phyllaries papyraceous, glabrous; outer phyllaries
linear-ovate, 2.8—3 X 0.8—1 mm, the apices acu¬
minate; inner phyllaries linear-ovate to I inear-el I ip-
tic, 11—13 X 1.2—2 mm, the apices acuminate to
acute. Florets 3, hermaphroditic, corollas light
pink, glabrous, apically papillose, 5-lobed, lobes
unilateral, oblong, ca. 3.5 mm long, with one deep¬
er slit ca. 5.5 mm long, the corolla then slightly
ligulate, tube ca. 4.5 mm long. Anthers 5, ca. 6 mm
long, apical appendages shortly apiculate, 1—1.5
mm long (apiculum ca. 0.2 mm long), sagittate; tails
ca. 1 .3 mm long, shortly pilose. Style branches
shortly bilobed. apically rounded, ca. 0.25 mm
long. Achenes oblong, 3—4 mm long, densely pi¬
lose. Pappus bristles uniseriate, ca. 39, plumose,
6—7 mm long, white.

Ainsliaea qianiana at present is known only from
the type collection from northeastern Yunnan, Chi¬
na, at approximately 2990 m.

The species is named to honor Qian Yi-yong, a
dedicated botanist and collector of the Chinese flo¬
ra.

The new species is characterized by its apiculate
anther appendages, rosulate leaves with petioles
winged, and capitula 3-flowered. Ainsliaea qiani¬
ana exhibits some likeness to A. macrocephala in
habit and capitulescence. That species is amply
distinguished by its apically rounded anther ap¬
pendages and its pilose phyllaries and leaves.
Twelve other species of the genus also have apic¬
ulate anther appendages.

Key to the Ainsliaea Species with Apiculate Anther Appendages

la. Feaves clustered at the median part of the stem.

2a. Leaves 3-parted . A. dissect a Franchet & Savatier

2h. Leaves lobed.

3a. Blades palmately lobed.

4a. Capitula 1-flowered . A. lancangensis Y. Y. Qian

4b. Capitula 3-flowered . A. acerifolia Schultz Bipontinus

3b. Blades pinnately lobed . A. cordifolia Franchet & Savatier

lb. Leaves ± rosulate.

5a. Leaves palmativeined; capitula I -flowered . A. uniflora Schultz Bipontinus

5b. Leaves pinnativeined; capitula 3- or 5-flowered.

6a. Capitula 5-flowered.

454

Novon

Figure 2. Ainsliaea qianiana S. E. Freire. — A. Habit. — B. Foliar trichome. — C. Phyllaries. — 1). Capitula. — E.
Floret without ovary. — F. Achene with pappus. — G. Stamen. — H. Style branches. Scale bars: A = 3 cm; B = 0.1
mm; C & F = 2 mm; 1) = 5 mm; E = 2.5 mm; G & H = I mm. Drawn from the holotype: Maire 2529 (W).

Volume 12, Number 4
2002

Freire

Ainsliaea from China and Vietnam

455

7a. Leaf bases rounded . 4. spanocephala Y. C. Tseng

7b. Leaf bases cordate . 4. brandisiano Kurz

6b. Capitula 3-flowered.

da. Petioles winged . 4. qianiana S. E. Freire

fib. Petioles wingless.

9a. Blades elliptic, rounded at the base.

10a. Blades 8-12 cm long . 4. pingbianensis Y. C. Tseng

l()b. Blades 4.5—5 cm long . 4. angustata C. C. Chang

9b. Blades oblong, cordate at the base.

11a. Phyllaries apically pilose . 4. cavaleriei H. L^veille

lib. Phyllaries glabrous.

12a. Blades slightly concolorous . 4. ramasa Hemsley

12b. Blades with lower surface whitish, veins reddish . . 4. rubrinervis C. C. Chang

Acknowledgments. I thank John Pru.ski for his
helpful comments and one anonymous reviewer for
critically reading the manuscript. Appreciation is
expressed to the curators of herbaria mentioned for
the loan of specimens that made this study possi¬
ble. 1 am also grateful to Emilio F. Rollie, Area de
Latin, Facultad de Humanidades, Universidad Na-
cional de La Plata, for correcting the Latin diag¬
noses and Victor H. Calvetti for inking the illustra¬

tions. Finally, I thank Victoria Hollowell for her
efficient editorial work. This study was supported
by the Consejo Nacional de Investigaciones Cien-
tfficas y Tecnieas (CONICET), Argentina.

Literature Cited

Bremer, K. 1994. Asteraceae: Cladistics & Classification.
Timber Press, Portland, Oregon.

Estudios en el Genero Schwartzia Vellozo (Marcgraviaceae): Una
Nueva Especie de la Cordillera Central Andina de Colombia

Diego Giraldo- (Janas

Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495,
Santafe de Bogota, D.C., Colombia. giraldoc@ciencias. unal.edu. co

Resumen. Se describe e ilustra una nueva espe-
cie de Schwartzia , S. parrae. La nueva espeeie se
conoce unicamente de la Cordillera Central Andina
de Colombia (Departamento de Antioquia). Se pre-
senta una clave para distinguir las especies col-
ombianas de Schwartzia.

ABSTRACT. A new species of Schwartzia, S. par¬
rae, is described and illustrated. The new species
is known only from the Andean Central Cordillera
of Colombia (Departamento de Antioquia). A key
for the Colombian species of Schwartzia is provid¬
ed.

Key words: Andes, Colombia, Marcgraviaceae,
Schwartzia.

El genero Schwartzia Vellozo, constituido por ca.
15 especies, se distribuye desde Costa Rica hasta
el sur de Brasil y se caracteriza por presentar in-
florescencias racemosas y flores largamente pedi-
celadas con un nectario (braetea nectarffera) ubi-
cado en la porcion medial a proximal del pedicelo
(Giraldo-Canas, 2001a, 2001b, 2001c). Este genero
fue considerado por mucho tiempo dentro del ge¬
nero Norantea Auhlet (Wittmack. 1878; Szyszylow-
icz, 1893; Gilg & Werdermann, 1925; De Boon,
1970; Eerreira, 1995), hasta que fue revalidado re-
cientemente por Bedell (1989).

Ambos generos presentan inflorescencias race¬
mosas, dilereneiandose basieamente por la longitud
del pedicelo, siendo en Schwartzia las llores lar¬
gamente pediceladas (pedicelo de 16—80 mm de
longitud), mientras que en Norantea son cortamente
pediceladas (pedicelo de 3—7 mm de longitud). Por
otra parte, en Schwartzia los nectarios reprodueti-
vos, cortamente peciolados (pecfolo de 1—7 mm de
longitud), se ubican en la poreibn medial a proxi¬
mal del pedicelo y en Norantea se localizan en el
tercio medio o hacia su porcion distal, presentando
un pecfolo de 5-20 mm de longitud. Norantea s.l.
ha sido dividido en 4 generos: Norantea s. str.,
Schwartzia, Sarcopera Bedell y Marcgraviastrum
(Wittmack ex Szyszylowicz) De Boon & S. Dressier
(De Boon & Dressier, 1997).

Como resultado del esludio del genero Schwart¬
zia y de la familia Marcgraviaceae (Giraldo-Canas,
1999, 2001a, 2001b, 2001c; Giraldo-Canas & Pic-
ca, 1999; Picca & Giraldo-Canas, 1999) se propone
una nueva especie, la que se describe e ilustra a
continuation:

Schwartzia parrae Giraldo-Canas, sp. nov. TIPO:
Colombia. Antioquia: Mun. Yolombd, vereda
El Bote, sitio Alto del Bote, 8.1 km SO de
Yolombd, bosque perturbado, 75°01'0,
6°32'N, 1550 m, I jul. 1989, II Callejas J.
Betancur & 0. Escobar 7957 (holotipo, HUA;
isdtipos, MO. NY). Figura 1.

Frutex foliis spiraliter alternantibus, peliolis 2-7 mm
longis, laminis (5-)9-22 cm longis et (1.3-)2-5.2 cm latis,
28—56 hidatodiis donatis. Racemus multiflorus. Flores
20—32, pedicellis 4—8 cm longis; stamina ca. 48, antheris
basi subcordatis, filamentis applanatis, verruculosis; ova-
rio conico; stigma mammiforme; braetea floralis cvmbifor-
mis, (7-)9-l8 mm longa et (3-)4— ft mm lata.

Arbustos escandentes, tallos cilfndricos a irre-
gularmente aplanados, fuertemente estriados; cor-
teza castano, ferrugfnea a cobriza. Hojas espirala-
das, subsesiles a cortamente pecioladas; pecfolo
ligeramente estriado, recurvado, canaliculado (el
canal muy angosto, ca. 1 mm de ancho), castano
claro a negruzco, de 2—7 mm tie longitud; ldminas
oblongas a obovadas, de (5— )9— 22 X (1.3— )2— 5.2
cm, de base redondeada a ligeramente cordata, ac-
uminadas, el apice prolongandose en un apfculo
negruzco de 1-1.5 mm de longitud, enteras, con
margenes ligeramente revolutos o involutos y en-
grosados, coriaceas, glabras; nervio medial conspi-
cuo en ambas earas, levemente hundido en la haz
y ligeramente abultado en el ernes, liso, de 1—3
mm de ancho en la porcion proximal tie la lamina
foliar, desvaneciendose hacia la porcion distal de
la lamina foliar; venacion secundaria suprimida o
apenas perceptible en la haz. debil en el enves,
entonces esta broquidodroma; 14—28 pares de glan-
dulas laminales en la superfirie abaxial, localizadas
a 1.5—1 I mm del margen, elfpticas, de 1 X 0.5

Novon 12: 456-459. 2002.

Volume 12, Number 4
2002

Giraldo-Canas

Schwartzia (Marcgraviaceae)

457

Figura 1. Schwartzia parrae Giraldo-Cafias. — A. Rama florffera. — B. Flor, pedicelo y braetea nectarifera. — C.
Ovario. (I)e Callejas e t al. 7957, holotipo, HU A.)

458

Novon

mm, con su orificio negruzco tie 0.2 X 0.15 mm,
distanciadas entre si 1—5 cm. Inflorescencia ter¬
minal, racemosa, multiflora (20—32 flores), densa;
pedunculo de 12-19 cm de longitud; pedicelos de
4—8 cm de longitud, cilmdrieos a moderadamente
aplanados, rectos a levemente flexuosos, lisos a lig-
eramente estriados, un poco mds gruesos en su por-
cidn distal, cobrizos, cada uno con una brdctea nec-
tarffera (nectario). Nectarios (bracteas nectarfferas)
cimbiformes, levemente rostrados, pendulos, verdes
a purpureos, ligeramente estriados longitudinal-
mente, pecfolos de los nectarios 1 .5—3 mm de lon¬
gitud, urnas de (7— )9— 18 X (3— )4— 8 mm, los orifi-
cios dispuestos hacia el eje de la inflorescencia y
ocupando una position transversal respecto a su
eje, los nectarios insertados por debajo del V3—Vb
proximal del pedicelo. Flores dispuestas en forma
espiralada, insercibn pedicelo-flor horizontal a
moderadamente oblfcua; bractbolas 2, sepaloides,
opuestas, orbiculares, verrueulosas, carnosas, 5—7
mm de longitud, margenes translucidos; sbpalos 5,
imbricados en dos series, orbiculares, verruculosos,
carnosos, 7—8 mm de longitud, margenes translu¬
cidos; petalos 5, connatos basalmente (antes de la
antesis) por ca. % de su longitud, obovado-elipticos,
de 8.5—1 1 X ca. 3 mm, carnosos, verruculosos; es-
tambres ca. 48, filamentos ca. 1.5 mm de longitud
(antes de la antesis), aplanados, verruculosos, ad-
natos a la base de los petalos, anteras introrsas,
elfptico-oblongas, subcordatas basalmente, de 3—4
mm de longitud; ovario supero, conico, ligeramente
estriado, ca. 3.5 X ca. 3 mm; estigma mamiforme.
Fruto desconocido.

Schwartzia parrae esta dedicada a mi amigo y
colega Carlos Parra (COL), estudioso de las familias
Brassicaceae, Myricaceae y Myrtaceae, quien a pe-
sar de su juventud ha contribuido significativa-
mente al conocimiento de la Flora de Colombia.
Esta especie sblo es conocida de la Cordillera Cen¬
tral Andina (Departamento de Antioquia) y crece
entre los 1 100 y 2000 m. Dado que sblo se conocen
de esta especie cuatro ejemplares eolombianos, se
inhere que bsta se encuentra en vfas de extincion,
dada la acelerada fragmentacibn y/o deslrueeibn de
los bosques en donde ella vive (observaciones per-
sonales).

Schwartzia parrae es facilmente distinguible de
las demas especies colombianas por sus nectarios
reproductivos pedicelares. los euales son cimbifor¬
mes. mientras que en las demds especies estos son
sacciformes, tubulares o globosos. Las cuatro es¬
pecies colombianas descritas pueden diferenciarse
fdcilmente por la siguiente clave:

Clave para Distinguir i.as Especies de Schwartzia
Presentes en Colombia

la. Nectarios reproductivos pedicelares cimbifor¬

mes; 14—28 pares de glbndulas laminales por
hoja; bractbolas de 5—7 mm de long.; sbpalos de
7—8 mm de long . S. parrae Giraldo-Canas

lb. Nectarios reproductivos pedicelares sacciformes,
tubulares a globosos; 0-6 pares de glbndulas
laminales por hoja; bractbolas de 1.8— 4.7 mm de
long.; sbpalos de 2—7 mm de long.

2a. Nectarios reproductivos pedicelares nacien-
do por debajo de la articulaeibn de los ped¬
icelos al eje de la inflorescencia; estambres
< 18; glbndulas laminales ubicadas a 6—10

mm del margen .

. S. diaz-piedrahitae Giraldo-Canas

2b. Nectarios reproductivos pedicelares nacien-
do en el tercio proximal del pedicelo; estam¬
bres 22—50; glandulas laminales ubicadas a
0.5— 4.5 mm del margen.

3a. Estambres 22—26; bractbolas ca. 2 mm
de long.; sbpalos de 2.2— 2.8 mm de
long.; ovario ca. 2.5 mm de long.; urnas
de los nectarios reproductivos pedice¬
lares de 9—17 mm de long .

. S. lozaniana Giraldo-Canas

3b. Estambres 50; bractbolas de 2.5—5 min
de long.; sbpalos de 5—7 mm de long.;
ovario de 3—8 mm de long.; urnas de los
nectarios reproductivos pedicelares de

15—33 mm de long .

. S. chocoensis Giraldo-Canas

Pardtipos. COLOMBIA. Antioquia: Mini. Amalfi, 8—
27 km NE de Amalfi, en la via Vetilla-Eraguas, sitios Sa¬
lazar v Marengo, I 150—1450 m, 7 die. 1989, Callejas et
al. 9102 (MCA); Mun. Guatapb, sector Santa Rita, zonas
aledanas a la carretera, 75°05'0, 6°10'N, 1800— 2000 m,
22 jun. 1997, Alzate et al. 247 (I I LI A); Mun. San Luis,
vereda Manizales, finca de Rambn Jaramillo, 1460—1760
m, 1 1 oct. 1981, Orozco et al. 549 (COL).

Agradecimientos. Quiero manifestar mi profun-
do reconocimiento al Instituto de Ciencias Natur-
ales de la Universidad Naeional de Colombia por
todas las facilidades que me brinda para llevar a
cabo los estudios botanicos. A los curadores de los
l.erbarios COAH. COL. CIL I ILIA, JAUM, MKDEL,
MO, NY. l’SO, SI, U IS. US y YEN por los presta-
mos enviados y/o por las facilidades brindadas dur¬
ante la visita a sus instalaciones. A S. Dressier (FR)
y V. Hollowell (MO) por su valiosa colaboracidn. A
H. Arellano (COL) por la elaboration de la ilustra-
ci6n. Esta contribution es derivada del proyecto
“Estudios sistematicos en el complejo Norantea
Aubl. (Marcgraviaceae)”, flnanciado por la Division
de Investigation DIB de la Universidad National
de Colombia, sede Bogota.

Literatura Citada

Bedell, H. G. 1989. Marcgraviaceae. En: R. A. Howard
(editor). Flora of the Lesser Antilles 5: 300-310.

De Boon, A. C. 1970. Family 121. Marcgraviaceae. En:

Volume 12, Number 4
2002

Giraldo-Canas

Schwartzia (Marcgraviaceae)

459

R. E. Woodson & R. W. Sehery (editors). Flora of Pan¬
ama, Part 4. Ann. Missouri Bot. Card. 57: 29—50.

- & S. Dressier. 1997. New taxa of A orantea Aubl.

s.l. (Marcgraviaceae) from Central America and adja¬
cent South America. Bot. Jahrb. Syst. 119: 327—335.

Ferreira, G. L. 1995. Estudo taxonomico das especies
brasileiras do genero Norantea Aublet (Marcgravi¬
aceae). Arch. Jard. Bot. Rio de Janeiro 33: 9—53.

Gilg, E. & E. Werdermann. 1925. Marcgraviaceae. En: A.
Fingler & K. Prantl (editors). Die nattirlichen Pflanzen-
familien, ed. 2, 21: 94-106.

Giraldo-Canas, D. 1999. Las Marcgraviaceae de la Ama¬
zonia colombiana: Flstudio preliminar sobre su diversi-
dad, distribucidn y fitogeograffa. Darwiniana 37: 15-24.

- . 2001a. Schwartzia brasiliensis (Marcgraviaceae),

nueva combinacion. 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: 477-482.

- & P. Picca. 1999. Tres nuevos registros de Marc-

gravia (Marcgraviaceae) para la flora mexicana. Hick-
enia 2(62): 289-292.

Picca, P. & D. Giraldo-Canas. 1999. Una nueva especie
de Marcgravia (Marcgraviaceae) de la Amazonia col¬
ombiana. Hickenia 2(63): 293-297.

Szyszylowicz, I. 1893. Marcgraviaceae. En: A. Engler &
K. Prantl (editors). Die natiirlichen Pflanzenfamilien
3(6): 157-164.

Wittmack, L. 1878. Marcgraviaceae. En: C. F. P. von Mar¬
tins (editor). Flora Brasiliensis 12(1): 213—258. Miin-
chen, Wien & Leipzig.

Rediscovered after 200 Years, Moraea ovata Thunberg Is a Rare
Ferraria (Iridaceae: Iridoideae) from Namaqualand, South Africa

Peter Goldblatt

It. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299,
St. Louis, Missouri 63166-0299, U.S.A. peter.goldblatt@mobot.org

John C. Manning

National Botanical Institute, P. Bag X7, Claremont 7735, South Africa.
manning@nbiet.nbi.ac.za

Abstract. Rediscovery in the late 1990s of the
mysterious Moraea ovata Thunberg, described in
1800 and first collected in 1793 in early fruit,
shows that the species belongs to tbe related genus
Ferraria. Flowers, only found in June 2001, show
that it has parallel anther locules and a spindle-
shaped ovary without a sterile beak, features con¬
sistent with section Ferraria of the genus. The new
combination F. ovata is provided for the species,
which we speculate is most closely related to F
densepunctulata, native to the west roast of Western
Cape Province, South Africa. The range of F. ovata
is restricted to low to middle elevations in central
Namaqualand in Northern Cape Province, South
Africa.

key words: Ferraria, Iridaceae, Moraea, phy¬
togeography, southern Africa.

Phe southern African Moraea ovata Thunberg
lias been a puzzling plant to botanists ever since it
was described just 200 years ago. It was named by
Carl Peter Thunberg in 1800 in bis preliminary ac¬
count of the Cape flora, Prodromus plantarum ca-
pensium. This work included plants from all of
southern Africa, but effectively only dealt with t lit'
winter-rainfall belt of the subcontinent, which ex¬
tends north and west of Cape Town along the coast
and near interior. Summer- rainfall southern Africa
had barely been explored botanically at the time.
Moraea ovata was only known to Thunberg from
three fruiting specimens, all of which lacked a
corm, the underground storage organ of most Iri¬
daceae, and of tt*ii a feature of taxonomic impor¬
tance. In fact, the reason the species was placed in
Moraea was most likely because it clearly had an
inferior ovary. Thunberg's account is succinct, but
accurately describes the short stem, round in sec¬
tion, bearing broadly ovate leaves. Later, in his Flo¬
ra capensis published in 1820, Thunberg men¬
tioned that he received his specimens from tin1

English plant collector Francis Masson, and that
they grew “in Namaquas juxta koks fontein” (in
Namaqualand near Koksfontein), the locality con¬
firmed by the annotation on the type sheet of M.
ovata, “Cocksfonteyn.”

So confusing was Moraea ovata that the English
botanist J. G. Baker, in his account of the Iridaceae
in Flora capensis (1897), treated the species as a
synonym of the western southern African Lapeirou-
sia pyramulalis (Lamarck) Goldblatt (which Baker
called L. fissifolia (Jacquin) Ker Gawler). The as¬
signment was made because L. pyramidalis has an
inflorescence with unusually broad bracts that re¬
semble in a crude way the leaves of M. ovata. I he
leaves of the latter are, however, true leaves, i.e.,
they do not subtend a flower nor are they associated
with a smaller inner bract as they would be in Lap-
eirousia il they were floral bracts surrounding a
llower. In 1928 a second English botanist, N. E.
Brown (1928). critically examined the specimens in
the Thunberg Herbarium housed at Uppsala, Swe¬
den, and concluded that the specimens were not a
hipeirousia, but he made no further comment. In a
revision ol the Gape species of Lapeironsia, Gold¬
blatt (1972) was not so certain, and M. ovata was
provisionally regarded as a somewhat divergent
form ol k. pyramidalis.

From the sparse literature relating to Masson’s
travels during his second visit to the Cape (1786—
1795), we assume it was on his 1793 visit to Na¬
maqualand that Masson collected these plants
(Gunn & Codd, 1981). It has been established that
Masson traveled as far as the Kamiesberg in that
year, since he collected certain species that only
occur there (Forbes, 1965), but his exact itinerary
is not known. There is no Koksfontein recorded in
the place names in tin* Kamiesberg. but Kookfon-
tein is the name given to the original farm on which
the present-day settlement ol Soebatslontein lies.

Novon 12: 460-464. 2002.

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Ferraria ovata from South Africa

461

Could Masson have passed though the area that lies
to the west of the Kamiesberg on his trip to or from
the mountains? This seemed likely, for in 1995 the
conservation biologist Anelise le Roux collected a
sterile specimen that exactly matched Masson’s
tvpe collection on the farm Doornfontein, which lies
on the eastern boundary of Kookfontein. We re¬
turned there in 1997 but failed to locate more
plants.

In 1999, during a multidisciplinary expedition to
Namaqualand, the biologists Nick Helme and Phil-
ip Desmet discovered a handful of plants that close¬
ly matched Masson’s specimens in the Knersvlakte
south of Kliprand on the farm Steenbokskraal (not
at all close to Soebatsfontein). Although the plants
were in bud. they did not look as if they would
flower, so three specimens were carefully dug up,
later to be replanted. These plants differed slightly
from those collected previously by Masson and le
Roux in having short, narrow basal leaves that were
unifacial in (he upper half as are those ol Ferraria ,
a genus of 10 species of semi-arid habitats in
southern and south tropical Africa (de Vos, 1979).
The corms were also exactly like those of a Ferraria
in their compressed, discoid shape, multiple inter¬
nodes, and lack of long-lived, fibrous tunics. The
habitat was not especially distinctive, for plants
were found growing on a gentle, sandy slope iu veld
dominated by dwarf succulents. In August 2000 we
discovered yet another population of the species,
again in sterile condition, west of Kliprand.

In early June 2001, this botanical mystery was
finally solved when the population that we found
the previous year was revisited. Several plants in
full bloom were found and the identity of Moraea
ovata could finally be established. The (lowers, like
the corm, match exactly those of Ferraria (de Vos,
1979), having subequal, clawed, lanceolate tepals,
with crisped margins. Like other species of Ferrar¬
ia. , the filaments are united in a slender column for
most of their length, and the style divides at the
apex of the filament column to form three flattened,
somewhat petaloid style branches that are deeply
forked, with an abaxial stigmatic surface, and with
fringed distal margins that extend over the stigma
lobes.

Ferraria ovata (Thunberg) Goldblatt N J. C. Man¬
ning, comb. nov. Basionym: Moraea ovata
Thunberg, Prod. pi. cap. 186. 1800. TYPE:
South Africa. Northern Cape (Namaqualand
near Koksfontein), without date, Masson in herb.
Thunberg 1225 (holotype, UPS). Figure 1.

Plants 4—20 cm high, mostly unbranched. Corm

discoid, ca. 2 cm diam., the old corms persisting
beneath the current one. Stem erect, terete. Basal
leaves linear, channeled below, to 4.5 cm long, the
upper half unifacial, ca. 2 mm wide, upper leaves
amplexicaul, broadly ovate and concave with nar¬
row reddish membranous margins, 1.5— 1.8 cm long
and up to 1.5 cm wide, with a short, unifacial tip;
leaves of immature plants unifacial, linear, up to
15 cm long, ca. 2 mm wide. Inflorescence a rhipi-
dium of 1 or 2 flowers, spathes green, leathery,
ovate with narrow reddish membranous margins,
the outer 10—12 mm long, the inner 12—15 mm
long. Flowers actinomorphic, lightly acrid-scented,
lasting 1 day, cream streaked with maroon, the te-
pal limbs spreading or lightly reflexed, yellow
marked with a few brown spots and with undulate
and minutely crisped chocolate brown margins; te¬
pals Iree, lanceolate, subequal, the inner slightly
smaller than the outer, the claws ascending, form¬
ing a cup ca. 7 mm deep and ca. 8 mm diam. at
the rim, the inner 15—18 X 5—6 mm with claws 6—
7 mm long, the outer 15—20 X 7—9 mm with claws
6—7 mm long; nectaries bipartite with two concave
hollows, 2X2 mm, situated between the lower
third and middle of the tepal claws. Filaments unit¬
ed in a smooth column, 7—8 mm long; anthers di¬
verging, appressed to the style branches, 2 mm
long, the anther lobes parallel. Ovary included in
the spathes, ovoid, 6 mm long; style dividing at the
apex of the filament column, the branches diverg¬
ing, deeply divided, terminating in feathery crests,
the stigma lobes abaxial, below the crests. Mature
capsules and seeds unknown.

Flowering time. June and possibly July.

Distribution. Ferraria ovata is apparently re¬
stricted to south-central Namaqualand, where it oc¬
curs on stony, granite-derived soil at middle ele¬
vations. Despite the few collections known, it may
be more common than the record indicates. Plants
are inconspicuous except in flower, and flowering
occurs early in the season when little plant explo¬
ration is undertaken to Namaqualand, where the
main flowering occurs in August and September.

Relationships. In her account of Ferraria, de
Vos (1979) recognized two main clusters of species
plus the monotypic section Glutinosae for the south
tropical African F. glutinosa (Baker) Rendle. Sec¬
tions Ferraria and Glutinosae comprise Ferraria
species with the apparently plesiomorphic feature
of parallel anther lobes. While section Glutinosae
has flowers with a plesiomorphic ovoid-truncate
ovary and capsule, species ol section Ferraria have
an elongate, spindle-shaped ovary lacking a sterile
beak (rostrum) except in F. schaeferi. Species of

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E

E

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Figure 1. Ferraria ovata (Thunberg) Goldblatt & J. C. Manning. — A. Entire plant with corm. — B. Inner tepal. —
C. Outer tepal. — I). Diagrammatic section of flower showing spindle-shaped ovary, filament column, and orientation
of the style branches and anthers. — E. Top view of style branches showing divided style branches w ith fringed margins
and position of the anthers (shaded). — F. Detail of the tip of one style branch and anther with parallel lobes. Drawn
by J. C. Manning from Manning 2.'i50 (NBG).

Volume 12, Number 4
2002

Goldblatt & Manning
Ferraria ovata from South Africa

463

Ferraria sect. Macroscyphae have an apomorphic
rostrate ovary with a sterile, somewhat tubular beak
at least 8 mm long, and anther lobes that either
diverge from the apex or are initially parallel but
later diverge toward the base (only F. ferrariola
(Jacquin) Willdenow, which thus seems somewhat
ambiguously placed in the section).

Ferraria ovata is clearly a member of section
Ferraria and resembles F. densepunctulata M. P. de
Vos of the Cape west coast in several aspects, par¬
ticularly its marked heterophylly, with the blades
of the lower leaves linear and up to 5 mm wide,
small stature, and especially significantly in the po¬
sition of the nectaries. In all other species of Fer¬
raria the nectaries are situated at the base of the
tepals, but in F. densepunctulata and F. ovata they
are situated near the middle of the tepal claws. The
two species resemble one another also in their early
flowering, in May and June. They differ most sig¬
nificantly in the shape of the cauline leaves and in
the color and longevity of the flowers. In F. dense¬
punctulata the cauline leaves are lanceolate, unlike

the ovate, cucullate cauline leaves of F. ovata, and
the pale gray or greenish flowers are finely marked
with maroon or purple spots, bear conspicuous yel¬
low-green nectaries, and remain open for two days,
unlike the yellow, boldly marked flowers of F. ovata
that last just one day and have inconspicuous
cream nectaries streaked with maroon.

Species of section Ferraria are confined to the
west coast and near interior of South Africa and
southwestern Namibia, an area that lies entirely
within the southern African winter-rainfall zone.

Biology. The slightly fetid-smelling, dull-col¬
ored, brown-speckled flowers are typical of Ferrar¬
ia. most species of which are visited by muscid and
calliphorid flies (unpublished data), and which we
assume are fly-pollinated, as did Vogel (1954). Fer¬
raria ovata may likewise be considered to share
this pollination strategy, for we observed several
muscid flies crawling on the flowers and carrying
loads of the orange-colored pollen of the species.
Flies were not captured for identification. Flies visit
the flowers to feed on nectar produced from the
nectaries located on the tepals.

A Revised Key to Section Ferraria
discussion

We include Ferrariola ferrariola in the key below, for although it was included in section Macroscyphae by de Vos,
its sectional position is ambiguous. It shares with section Ferraria the nearly parallel anther lobes (they diverge from
the base with age), but it has the rostrate ovary characteristic of section Macroscyphae, also, however, present in F.
schaeferi Dinter of section Ferraria. In particular, F. ferrariola shares the heterophyllous condition with F densepunc¬
tulata and F. ovata. Whatever its correct sectional position, it seems helpful to include F. ferrariola in the key to
section Ferraria.

la. Stem slender with internodes partly exposed; lower leaves linear to falcate, with blades usually less than 5
mm wide; nectaries situated at the base or near the middle of the tepals.

2a. basal leaf sheaths conspicuously reddish or red-spotted; nectaries situated at the base of the tepal claws;

ovary rostrate; stigmas large and flap-like, partially obscuring the anthers . F. ferrariola

2b. basal leaf sheaths at most faintly flushed reddish; nectaries situated near the middle of the tepal claws;
ovary rostrate; stigmas minute, at the tip of curving, horn-like branches.

3a. Cauline leaves ovate-cucullate; flowers yellow with brown margins and a few conspicuous spots;

nectaries cream streaked maroon . F. ovata

3b. Cauline leaves lanceolate-attenuate; flowers gray or greenish with minute brown or maroon speckling;

nectaries yellowish green . F. densepunctulata

lb. Stem stout, mostly covered by leaf bases; leaves sword-shaped to falcate, with blades more than 5 mm wide;
nectaries situated at the base of the tepals.

4a. Ovary witli a short tapering rostrum ea. 8 mm long; tepals 22— 25( — 30) mm long, yellow with dark brown

margins and blotches, often coalescing in the outer quarter; flowers sweetly scented . F. schaeferi

4b. Ovary without a tapering rostrum; tepals 25—35 mm long, flowers variously colored, usually dark maroon
or purple with paler margins or cream to pale yellow' and variously striped and blotched; flowers with an
unpleasant fetid scent.

5a. Leaves with a thickened zone in the middle and a strong pseudomidrib, 2-ranked . . . F. crispa burman
5b. Lea ves with numerous veins of equal size and without a pseudomidrib, often spirally 2-ranked . . .

. F. foliosa G. J. Lewis

Additional specimens (cited according to the geograph¬
ical quarter-degree square system for indicating latitude
and longitude in use in southern Africa). SOUTH AFRI¬
CA. Northern Cape: 30.17 (Hondeklipbaai) Farm
Doomfontein, Steenkamp Kraal (BA), 2 Sep. 1995 (ster¬
ile), Le Roux 4653 (JON K). Western Cape: 30.18 (Ka-
miesberg) Farm Gannabos on road from bitterfontein to

Kliprand (CD), 10 Aug. 2000 (sterile), Goldblatt & Man¬
ning 1 137.3 (MO, NRG). 6 June 2001, Manning 2350
(NbG); Knersvlakte, Farm Steenbokskraal, 20 Aug. 1999,
National Geographic- IRC Expedition 115 (NbG).

Acknowledgments. Support for our study by

464

Novon

grants 6704-00 and 7103-01 from the National
Geographic Society is gratefnlly recognized. We
thank Anelise Le Roux and Philip Desmid for
their assistance in relocating the elusive Ferraria
ovata.

Literature Cited

Baker, J. G. 1897. Iridaceae. Pp. 7-71 in W. T. Thiselton-
Dver (editor), f lora Capensis 6. Reeve, Ashford, Kent.
Brown, N. K. 1928. The Iridaceae of Thunberg's herbari¬
um. .1. Linn. Soc., But. 48: 34—42.

Lorbes, V. 1965. Pioneer Travellers in South Africa. Bal-
kema. Cape Town.

Goldblatt, P. 1972. A revision of the genera Lapeirousia
Pourret and Anornalheca Ker in the winter rainfall re¬
gion of South Africa. Contr. Bolus Herb. 4: 1—1 I I.
Gunn, M. & L. E. Codd. 1981. Botanical Exploration of
Southern Africa. A. A. Balkema, Cape Town.
Thunberg, C. P. 1800. Prodromus plantarum capensium.
J. Edman, Uppsala.

- . 1820. f lora capensis I. Bonnier, Copenhagen.

Vogel, S. 1954. Bliitenbiologische Typen als Elemente der
Sippengliederung. Bol. Stud. I: 1—338.

Vos, M. P. de. 1979. The African genus Ferraria. J. S.
African Bot. 45: 295—375.

One New Combination and One New Species in Leiomitra
(Trichocoleaceae, Hepatophyta) from Southern South America

Gabriela G. Hassel de Menendez

Museo Argentine de Ciencias Naturales “Bernardino Rivadavia,” Avda. Angel Gallardo 470,
1405 l)JR Buenos Aires, Argentina. hassel@muanbe.gov.ar

Abstract. Trichocolea elegans Lehmann is trans¬
ferred to the genus Leiomitra Lindherg, because gy-
noeeia have been found in several specimens with
a coelocaule, perianth, and thin calyptra. hnomitra
smaragdina Hassel sp. nov. also belongs to this ge¬
nus as confirmed through female specimens. Both
species, present in Chile and in Argentina, are de¬
scribed and illustrated, with the distinguishing
characters emphasized.

Rksumen. Trichocolea elegans Lehmann se trans-
tiere al genero Leiomitra Lindherg porque se en-
contraron ginoecios en varios espeefmenes con cel-
ocaule, periantio y una delgada caliptra. leiomitra
smaragdina lliissel sp. nov. pertenece tambien a
este genero como fue confirmado a traves de es-
peefmenes femeninos. Se describen e ilustran am-
bas especies, presentes en Chile y en Argentina,
con enfasis en sus caracteres distintivos.

Key words: Argentina, Chile, Hepatophyta,
Leiomitra, Trichocoleaceae.

The characters or circumscribing concepts for
the genera Trichocolea Dumortier eorr. Nees 1838,
noin. conserv., and Leiomitra Lindherg 1875 were
discussed by Hatcher (1957, 1959), Engel (1999),
and Schuster (1980, 1984, 2000). Trichocolea is
considered to he distinguished by a pinnate growth
form, the female organs terminal on main shoots,
and sporophytes surrounded by a hirsute calyptra,
covered with paraphyllia (see Hatcher, 1959).
Leiomitra does not present a regular pinnate growth
form, female organs are developed on short lateral
branches, and the calyptra is smooth and free, in¬
cluded in a coelocaule that ends in a perianth (see
Hatcher, 1959), corresponding to Trichocolea to-
mentosa (Swartz) Gottsehe in Trichocolea sect. Lae-
viflora Gottsehe (see Schuster, 1984, fig. 52).

Only Trichocolea was reported from southern
South America, with two species: T. elegans Leh¬
mann. by Hatcher (1957), Fulford (1963), Engel
(1978), and Schuster (2000), present along the
southwest humid side of the continent: and T. el-
liottii Stephani, a Caribbean species, disjunct and
restricted to the Juan Fernandez Islands by Hatcher

(1957) and Fulford (1963). Schuster (2000) con¬
cluded that some species in his tentative key of
Trichocolea (which includes T. elegans and T. el-
liottii) may prove to belong to the genus Leiomitra,
as the specimens he had seen were sterile.

The discovery of a third species, in a recent ex¬
cursion. induced me to check all my putative col¬
lections of Trichocolea sensu R. Hatcher as well as
those of some other institutions in the hope of re¬
vealing additional specimens. Instead I found some
female “ Trichocolea elegans ” plants and several
male plants. Androecia, gynoecia, and sporophytes
were unknown to Hatcher (1957) and to Fulford
(1963). Engel (1978: 78) cited a fertile specimen
from Chile, Brunswick Peninsula, B. Fortescue “c.
sporo.” hut gave no further details.

Close observation of the gynoecia found on what
was known as “ Trichocolea elegans " has shown that
all these specimens belong to Leiomitra. Therefore,
a nomenclatural change is necessary, along with the
description of the characters that support these
changes.

Leiomitra elegans (Lehmann) Hassel, comb. nov.
Basionym: Trichocolea elegans Lehmann, Nov.
Minus Cog. Stirp. Pug. 10: 8. 1857. TYPE:
Chile. Valdivia, Lechler s.n. (holotype, not
seen; isotype, NY). Figure I.

Trichocolea verticillala Stephani, Bill. Kongl. Svenska Ve-
tensk.-Akad. Ilandl. 20 (III. 6): 57, 1900. TYPE:
Chile. “Valdivia,” 1887. I)r. Hahn s.n. hl>. Jack (9
cum capsula) (lectotype, designated here. M); syn-
types: “Ensenada,” P. Dusen s.n. ((f); “Aysen,” P.
Dusen s.n. ((f): "Pto. Bueno,” P. Dusen s.n. ((f); “Ghi-
loe," I1. Dusen s.n. (G).

Trichocolea decrescens Stephani, Bill. Kongl. Svenska Ve-
tensk.-Akad. Ilandl. 46(9): 77, fig. 30b— c, 1911.
TYPE: Chile. “Caleta Kayo,” C. Skottsberg 557 (lee-
totype, designated by Engel (1978), G); “Juan Fer¬
nandez Islands. El Yunque,” C. Skottsberg s.n. (syn-
type).

Detailed descriptions. B. Hatcher (1957: 16,
ligs. 98-1 10), Fulford (1963: 48. fig. 6a-e).

Description of additional characters. Branching
sometimes l(or 2)-pinnate, frequently irregular, the

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Hassel de Menendez 467

Leiomitra (Trichocoleaceae, Hepatophyta)

main shoot not distinguishable; the distance oi leaf
insertion quite variable. Oil bodies ovoid to fusi¬
form, refringent pale yellowish, (6)8 to 1 I per leaf
cell. 5-7 X 2. 5-3. 5 gm. Male plants irregularly 1-
pinnate. Androecia inconspicuous with 3 to 4 pairs
of bracts on main shoot or branches, successive but
intercalary between long sets of normal leaf cycles.
Bracts of the same si/e and armature as vegetative
leaves, dorsal portion of lamina enlarged, covering
antheridia. Antheridia globose, 150—200 gin diam.,
1 (or 2) per bract, with biseriate stalk. Gynoecia
inconspicuous, terminal on main stem or enlarged
branches, but through development of adjacent ter¬
minal branch becoming delayed as short lateral
branches. Archegonia covered by bracts and brac-
teole similar to the leaves and amphigastria, al¬
though with an enlarged lamina, 0.4— 1.5 mm high
(including segments), the segments at their base 12
to 15 cells wide, the dorsal ones being larger. Ma¬
ture gynoecium 2.5—3 mm long, coeloeaule wall 8
to 9 cells thick, covered with 3 to 4 cycles of leaves
that are identical to the vegetative leaves. The uni-
stratose perianth, not surpassing the bracts, smooth
at its base, covered above with isolated but crowded
short, curved, simple, or furcate to opposed ciliate
structures, reminiscent of leaf eilia; mouth lacini-
ate-ciliate, each lacinia similar to bract segments,
8 to 10 cells wide at their base, with several op¬
posed pairs of cilia, final uniseriate filament rather
straight, 4 to 6 cells long. Calyptra free, thin, with
sterile archegonia on its base. Foot as deep as the
3 or 4 pairs of leaves. Seta 2.5— 3.5 mm long. Cap¬
sule brow n, ellipsoid. Capsule wall 4-stratose, outer
cells prismatic with thin walls and occasional
brown thickenings, inner cells fusiform, irregularly
arranged with semiannular brown thickenings.
Spores red-brown, hispid, 23 gm diam. Klaters
nearly straight, 205—250 fxm long, 5-1 1 ptm wide,
with 2 (to 3) helicoidal brown thickenings.

The distinguishing characters of Leiomitra ele-
gans (Lehmann) Hassel are: the pale grayish green
color; the spongy consistency; shoots ca. 1.2 mm
wide; leaves with 5 opposite ciliate arching seg¬
ments trifid at their bases.

The completely developed gynoecia of this taxon,
here described and illustrated for the first time and
verified from several samples, confirm that it be¬

longs to Leiomitra. The general aspect ol the gy¬
noecium of Leiomitra elegant is also reminiscent of
that of Leiomitra hirticaulis H. M. Schuster, (see
Schuster, 1984, fig. 52).

Examination of approximately 200 samples ol
Leiomitra elegans reveals that its growth form is
usually pendulous, loosely or compactly interwoven
with other hepatics. It occurs in dripping carpets,
not only on old trunks in the temperate forests, but
also on fallen logs, on humid rock walls, on crev¬
ices above waterfalls, wet banks by streams, and
the door of Nothofagus forests. Its presence has
been confirmed in Chile and Argentina, from Vi 1-
lariea in Chile to Isla de los Estados (Staten I.) at
the tip of Tierra del Fuego in Argentina, as well as
in the islands that surround Chiloe and the Channel
Islands, like Piazzi Island, Rennel Island, Vidal
Gormaz Island, Virtudes Island, and Diego de Al-
magro Island.

Selected specimens. CHILE. Corral, (d), R. A. Pdrez
Moreau s.n. (BA 2427). Region X: cerca de Termas El
Amarillo, (d). G. Hassel de Menendez (G.H.M.*) 11241
(BA). Region XI: 20 km W of I, a Tapera, Mina Sta. Te¬
resa, (d, 9), Hyvonen 5156 (BA); Parque Nacional (v)ueu-
lat, G.H.M. & M. Rubies 12661 (BA); Clarence Island, (d,
9). Hyvonen 2951 (BA); Seno Ultima Esperanza, Anedn
Sin Salida, Seno Union, (9), S. IV. Greene, Transecta Bo-
tdnica de la Patagonia Austral- Bryophytes (TBPA-R**)
878 b (BA); Piazzi Island, Caleta Ocasion. (9), G.H.M.,
TRPA-B 899 (BA); Rennel Island, (9), G.H.M., TRPA-B
1251 (BA); Vidal Gormaz Island, Caleta Nantuel. (9),
G.H.M., TRPA-R 1412j (BA), G.H.M., TRPA-R 1449 (BA);
Virtudes Island, Pto. Virtudes, (9), G.H.M. TRPA-R 1913
(BA). ARGENTINA. Neuquen: Brazo Blest, Arroyo Bra¬
vo, O. Ktihnemann (9, cum capsula) (BA 2161b). Rio
Negro: Pto. Blest, camino a Los Cantaros, G.H.M. 8921
(d) (BA), G.H.M. 9785 (d, 9) (BA); Rfo Negro, Lago
Frias, cerca de la casa del guardaparque, G.H.M. 177 (d,
9 cum capsula) (BA, MO), R. A. Perez Moreau (d) (BA
4612a), camino al lfmite con Chile, G.H.M. 364 (9 c.
caps) (BA). Chubut: Lago Mendndez, cireuito Rfo Cisnes,
G.H.M. 6475 (9 juv.) (BA). Tierra del Fuego: Isla de
los Estados. B. Hoppner, G.H.M. 4681 (9 juv.) (BA).

♦G.H.M. = G. Hassel de Menendez-, **' TBPA-B = Tran¬
secta Rotdnica de la Patagonia Austral- Bryophytes.

Leiomitra smaragdina Hassel, sp. nov. TYPE:
Chile. Parque Nacional Queulat, sendero de la
cascada, cuesta 615 msm, (<d, 9, cum cap¬
sula) 22 Mar. 1 999, G. Hassel de Menendez &
M. Rubies 12642 (holotype, BA; isotype, MO).
Figure 2.

Figure 1. Leiomitra elegans (Lehmann) Hassel. — A. Bracts, perianth (light shading), and capsule (darker shading).
— B. Longitudinal section of gynoecium. — C. Apical portion of perianth with superficial cilia and cuticle shaded on
left. — D. Apical portion of bract. — E. Leaf. — F. Amphigastrium. — G. Transverse section of capsule wall. (A, B, D
from G. Hassel de Menendez, Transecta Rotdnica de la Patagonia Austral- Bryophytes 1913 [BA]; C, G from G.H.M. 177
[BA]; E, F from G.H.M. & M. Rubies 12661 [BA].)

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Hassel de Menendez 469

Leiomitra (Trichocoleaceae, Hepatophyta)

Plantae smaragdinae minutae, facie hirsuta. Folia lam¬
ina deminuta 5-segmentata. Amphigastria 4-segmentata.
Segmenta attenuata ciliata; cilia rigida paribus oppositis,
0-2 furcata. Dioica. Gynoecium coelocaule bracteis tec to
ad apicem perianthio segmentato ciliato.

Plants minute, solitary or in groups, flat and
tightly adherent to other small plants, emerald
green, appearing hirsute, shoots including stem and
leaves 0.49—0.65 mm wide, regularly to irregularly
1- or 2-pinnate; branches alternate, most of them
lateral terminal of the Frullania type, of variable
length, very occasionally ventral intercalary
branches present. Stem cylindric, 97—146 gm
diam.. 5 to 7 (to 10) cells diam., with 14 to 26
peripheral cells, medullar cells 15-30 gun diam.,
epidermal cells 12—15 gun diam.; cortex undiffer¬
entiated. cuticle coarsely striolate. I’he main stem
with uniseriate Y- or I-shaped paraphyllia, 2 or 3
(to 5) cells long. Leaves spreading, with oblique
insertion, asymmetric, divided into 5 ciliate seg¬
ments. Leaves including segments 0.40-0.60 mm
long. Lamina (1)2 to 4 cells high (cells 35—47 gun
high. 12-23 gun wide, and 12-14 gun thick), some¬
times with small lateral teeth. Segments at base
(2)4 cells wide, distally 2 cells wide, these cells
35—47 gun long, segment apex tapered, 1 -seriate 3
to 4 cells long, the tip cell 75-130 gun long, central
segments with 4 to 5 pairs of opposite cilia, lateral
segments with 3 to 4 pairs ol cilia; proximal cilia

1 or 2 furcate; cilia consisting of rather still and
straight cells, borne on the abaxial side of the seg¬
ments. with furcate branching and swept toward the
tip of the shoot; whole surface of leaves coarsely
striolate. Amphigastria up to 0.99 mm long includ¬
ing segments. Undivided portion of the lamina I to

2 cells high; segments 4, the central sinus deepest;
cilia opposite as in leaves. Rhizoids, when present,
at base of amphigastria. Dioicous. Androecia fusi¬
form. conspicuous, on stem and branches with 5 to
6 or more (?) pairs ol saccate perigonial bracts, the
dors ally convex lamina 5 to 6 cells high, with seg¬
ments and cilia similar to the leaves; I or 2 an-
theridia per bract; globose, 160 gun diam., stalk
biseriate. Gynoecia inconspicuous, terminal on
stem, becoming displaced when unfertilized. Bracts
and braeteoles in 2 series surrounding the coelo¬
caule; bracts up to 2 mm long (including segments
and cilia), lamina higher than in leaves, with su¬

perficial branched cilia; the 5 segments ciliate as
in leaves; bracteole similar to amphigastria but
more shallowly segmented. All lips of segments and
cilia swept toward shoot tip. Perianth not emerging,
2-2.5 mm high, ending in segments and cilia sim¬
ilar to those of the leaves but with more delicate
cell walls. Calyptra thin; sterile archegonia at hot-
tom in contact with perianth base. Capsule dark
brown, spherical 3- or 4-stratose. Spores brown,
hispid, 18—23 gun diam. Klaters brown, 190—260
gun long, 9-12 gun diam. in the middle, attenuate
lips 5 gun diam., with 2 to 3 helicoidal thickenings.

Distinguishing characters of Leiomitra smarag-
dina, a species from southern South America, are:
the emerald green color when alive (hence the spe¬
cific epithet), the small size relative to other Leiom-
itra species, the reduced foliar lamina with the stiff
segments and cilia spreading and directed toward
lift of shoot, which give it a hirsute effect.

Other paraphyllose species of Leiomitra are pre¬
sent in tropical South America, as indicated by
Schuster (2000: 147) in a key: L. paraphyllina
Spruce, L. hirticaulis R. VI. Schuster, and /.. robusta
(Stephani) R. M. Schuster. In these taxa the leal
lamina is higher than in L. smaragdina, more than
4 cells, and the perianth is absent, low, or vestigial.

leiomitra smaragdina grows on bark of different
tree species, in the same habitat as Leiomitra ele¬
gant, but the smaller plants are less exuberant than
this species, and the individuals are often attached
to other hepatics, like Porella subsquarrosa (Nees
& Montagne) Trevisan, Chiloscyphus striatellus C.
Massalongo, C. ctenophyllus (Schiffner) Hassel, C.
infiexispinus (Hooker I. & Taylor) J. J. Engel & R.
VI. Schuster, or grow with Riccardia spinulifera C.
Massalongo, Plagiochila lophocoleoides Montagne,
P. bispinosa Lindenberg ex Gottsche, Acrobolbus
ochrophyllus (Hooker I. & Taylor) R. M. Schuster,
Pigafettoa crenulata G. Massalongo, Leiomitra (de-
go ns (Lehmann) Hassel, Tylimanthus limbatus Ste¬
phani, Jamesoniella colorata (Lehmann) Spruce ex
Schiffner, and Apometzgeria frontipilis (Lindberg)
Kuwahara & J. J. Engel. Thus far. the new species
has been found only in the XI Chilean Region,
where areas of the temperate rain forest of Notho-
fagus species still exist. In Argentina Leiomitra
smaragdina has been found on trunks of Nothofa-

Figure 2. Leiomitra smaragdina lliissel. — A. Female plant. — If. Male plant. — G. Perianth witli capsule (shading).
— I). Longitudinal section of gynoecium. — E. Apical portion of perianth with superficial cilia, cuticle indicated by
shading below. — F. Leaf. — (7 Amphigastrium. — II. I. Perigonial bracts. — J. Apical portion ol bract. — K. Iransverse
section of capsule wall. — L. Transverse section of stem with paraphyllia. (A— L from holotype, & M. Rubies

12642 | BA].) Note: C. D, E, .1 have a different magnification from similar details in Figure I for L elegant.

470

Novon

gas dombeyii, together with other bryophytes like
Plagiochila bispinosa Lirulenberg ex Gottsche, P.
heterodonta (Hooker f. & Taylor) Gottsche et al.,
and Leiomitra elegans in the Nahuel Huapi and Los
Alerees National Parks, located in the provinces
Neuquen, Rfo Negro, and Chubut. Only one spec¬
imen, the type, has been found with a single cap¬
sule.

Specimens examined. CHILE. XI Region: Lago Risso
Patrbn, I 15 msm, G.H.M. A- M. Rubies 1289} (RA); Car-
retera Austral, Quebradita, G.H.M. & M. Rubies 13094
(BA). ARGENTINA. Ncii<|iien: Parque National Nahuel
Huapi, Villa Angostura, camino al lfmite con Chile, a 500
m de la Aduana, sobre vara de Chusquea G.H.M. 8299 (9
juvenile sterilis) (BA). Rio Negro: Parque Nacional Na¬
huel Huapi, Lago Enas, camino al Tronador, G.H.M. 162
(9 juv. ster.) (BA 22013); Lago Nahuel Huapi, Pto. Blest,
G.H.M. 87 (9 juv. ster.) (BA 0471), G.H.M. 102 (9 juv.
sler.) (RA 0486). Gliuhut: Parque Nacional Los Alerees,
Lago Men^ndez, 0. Kiihnemann (RA 4946 a).

Acknowledgments. I thank the Consejo Nacion¬
al de Investigaciones Cientffieas y Teenicas de Ar¬
gentina (CONICET) for providing funds for travel
expenses in Argentina and Marta Rubies, who as¬
sisted me during the trip in southern Chile (1999)

and helped with literature citations. I am also in¬
debted to Matilde Gonsalves Carralves, who with

much skill illustrated the laxa presented here.

Literature Cited

Engel. J. J. 1078. A taxonomic and phytogeographic study
of Brunswick Peninsula (Strait of Magellan) Hepaticae
and Anthocerotae. Fieldiana, Bot. 41: 1—319.

- . 1999. Austral Hepaticae. 26. The identity, taxo¬
nomic position, and ecology of Trichocolea julacea
Hatcher (Trichocoleaceae). Novon 9: 25-28.

Fulford, M. 1963. Manual of the leafy Hepaticae of Latin
America. Part I. Mem. New York Bot. Card. I I: 1 — 172,
pi. 1-37 (unnumbered).

Hatcher, R. E. 1057. The genus Trichocolea in North,
Central, and South America (Hepaticae). Lloydia 20(3):
139-185.

- . 1959. The structure of the female inflorescence

and its taxonomic value in the genus Trichocolea (He¬
paticae). Lloydia 22(3): 208—214.

Schuster. R. M. 1080. New combinations and taxa of He¬
paticae, 1. Phytologia 45: 415—437.

- . 1984. Evolution, phylogeny and classification of

the Hepaticae. Pp. 892-107 in R. M. Schuster (editor),
New Manual of Bryology, Vol. 2. Hattori Botanical Lab¬
oratory, Nichinan.

- . 20(H). Austral Hepaticae. Part I. Beih. Nova Hed-

wigia 118: l-VIl, 1-524.

Two New Species of Pectis (Asteraceae: Tageteae) from

South America

David J. Kcil

Biological Sciences Department, California Polytechnic State University, San Luis Obispo,
California 93407, U.S.A. dkeil@calpoly.edu

Abstract. Pectis hassleri and P. pumila arc new
species, the former from the Gran Chaco area of
Paraguay and the latter from southwestern Ecuador
and northwestern Peru. Pectis Imssleri differs from
P. odorata by leaves that are glandular-punctate on
the adaxial as well as the ahaxial surfaces, by lon¬
ger and wider ligules of the ray florets, and by ray
pappi of awns and shorter bristles. Pectis pumila
differs from the closely related P. arida by its wider
leaves and by its sessile or subsessile capitula with
campanulate involucres and obovate phyllaries. A
hexaploid chromosome count of 2 n — 36It is newly
reported for P pumila.

Key words: Asteraceae, Compositae. Ecuador.
Paraguay, Pectis , Peru, South America, Tageteae.

As part of monographic studies of Pectis L. (As¬
teraceae: Tageteae), I have discovered two previ¬
ously undeseribed South American species.

Pectis hassleri keil, sp. nov. TYPE: Paraguay.
Gran Chaco: Loma Clavel, 23°20,S, Nov.
1903, K. Hassler 2491 (holotype, G: isotypes,
BM. K. GH, k. LIL. MICH, MO. NY. P. S, UC,
US).

P. odoratae affinis serf glandihus pellucidis numerosis
in lamina adaxiali. ligulis flosculorum radioruin longiori-
bus (5. 5-6.5 mm vs. 3-4 mm) et latioribus, et pappis
acheniorum radiorum 2—3 aristarum 2.5—4 mm longum et
ca. 10 setarum breviorum versus 30—40 setarum usque ad
7 mm longum diversa.

Annual or sometimes apparently perennial; herb¬
age probably scented, but nature of odor unknown.
Stems spreading to erect or ascending, 10—35 cm
long, several-branehed from near the base, few-
branched above, moderately to densely leafy, pur¬
ple-brown, glabrous or sparsely to densely puber-
ulent. Leaves opposite, lance-linear, often falcate,
2-5 cm long, 2—4 mm wide, acute, mucronate to
bristle-tipped, proximally eiliate with 2 to 5 pairs
of bristles 2—3 mm long, glabrous or sparsely dis-
tallv scaberulous on the margins, densely dotted on
both surfaces with round, pellucid glands 0.1— 0.2
mm diam., the glands of the adaxial surface smaller

and more numerous than those of the ahaxial sur¬
face. minutely ciliolate near the base, otherwise
glabrous. Capitula long-peduncled, solitary or in
open, few-headed cymes, ea. 38- to 75-flowered;
peduncles 3-11 cm long, slightly elavate, hearing
3 to 7 linear-acuminate scale-like bractlets 3—6 mm
long or the proximal bract larger and leaflike. In¬
volucres campanulate; phyllaries 8 (to 10). oblan-
eeolate to obovate, 6-8.5 X 2-3.5 mm, broadly
overlapping, subacute, weakly convex, proximally
rounded and gibbous, indurate-keeled in the prox¬
imal V2-V3. narrowly hyaline- or purple-margined in
the proximal %, densely dotted or streaked with
elliptical to linear glands, distally ciliolate. Ray flo¬
rets 8 (to 10): corolla yellow or reddening abaxially,
9-10 mm long, the tube 3-3.5 mm long, the lignle
narrowly ovate, 5.5— 6.5 mm long, glabrous. Disk
florets 30 to 65; corolla yellow, 4-6 mm long, weak¬
ly 2-lipped, slightly exceeding the pappus, the lube
and throat about equal, the ahaxial lip ca. 1 mm
long. 2-3 X as long as the 4 lobes of the adaxial
lip; anthers ca. 2 mm long; style exserted. Achenes
3.5—4 mm long, sparsely appressed-puberulent with
trie homes 0.2— 0.4 mm long; ray pappus of 2 or 3
stiff awns 2.5—4 mm long and ca. 10 shorter bristles
1-2 mm long; disk pappus of 20 to 30 slender bris¬
tles 2-5 mm long. Chromosome number unknown.

Distribution. Endemic to the Gran Chaco area
of Paraguay, this species is represented by only a
handful of collections with rather sparse data. I
have not been able to determine a precise location
for any of the collections examined. Elevation data
are unavailable. Essentially nothing is known about
the ecology of the species, known flowering dates
range from November to June.

Relationships. Pectis hassleri is one of the very-
few species of Pectis with glands on the adaxial leaf
surface. In most other species the glands are con¬
fined to the ahaxial surface of the leaf or to the
margins. The significance, if any, of these extra
glands is unknown. The glands are similar in size
and shape to those of the related P. odorata A. H.
R. Grisebach and P. substriata H. H. Rushy, both

Novon 12: 471-473. 2002.

472

Novon

of which are strongly scented. Presumably P. has-
sleri is scented as well.

I have named the species in honor of Erich Hass-
ler, a pioneer in botanical studies of Paraguay.
Ilassler recognized the type collection of P. hassleri
as a distinct taxon and considered it to be a variety
of P. odorata. He assigned it an unpublished vari¬
etal epithet alluding to broad ligules.

Paratypes. PARAGUAY. Chaco septentrionalis,
Puerto Tolanoera, 1907, A. Fiebrig 1330 (G); Chaco, 21°S
lat., 1906. K. Fiebrig 1459 (G, Z); Gran Chaco, sal., A.
Pride s .ii. (K); in region crossed by the lower portion of
the River Pilcomayo, June 1906, T. Rojas 257 (BAK, BM.
G, K. P).

Pedis piiniila I). J. Keil, sp. nov. TYPE: Peru.
Eambayeque: along Pan American Hwy. ea. 30
km NNW of Piura, 7 km S of Puente Sullana
(over Rfo Chira), 30 Mar. 1987, I). J. Keil et
al. 19917 (holotype, F; isotypes, ASU, BM,
CPUN, F, (;, (ill. HUT, K. MFXU, MO. NY,
OBI, OS, BSA. S, TEX, IJC, US, USM).

P. aridae I). J. Keil affinis sed (oliis saepe latioribus,
eapitulis sessilibus vel subsessilibus, involueris campan-
ulalis, et phyllariis obovatis diversa.

Erect or diffusely branched, often strongly scent¬
ed, tap-rooted annuals. Stems I to several from the
base, erect or diffusely spreading, 3—50 cm long,
cymosely branched distally, straw-colored to dark
purple-brown, puberulent throughout or in lines ex¬
tending from the connate flange and midribs of the
leal bases with soil, multicellular trichomes 0.1-
0.2 mm long. Leaves opposite, linear-oblaneeolate
or lanceolate, 2—4 cm long, 2-0 mm wide, obtuse
to subacute, mucronate or bristle-tipped, with the
base narrowed or more often conspicuously ex¬
panded. entire and eiliate to beyond the middle
with slender bristles 2—4 mm long or serrate with
bristle-tipped teeth, punctate abaxially with scat¬
tered. round, pellucid glands 0.2-0. 3 mm diam..
scaberulous along the margins, sparsely hirtellous
proximally along the margin and midrib with tri¬
chomes 0.1— 0.2 mm long, otherwise glabrous. Ca-
pitula solitary, sessile or subsessile in the forks of
the stems, subtended by 1 or 2 scarious, lance-
acuminate bractlets, 24- to 31 -flowered. Involucres
campanulate; phyllaries 5 or 0. obovate, broadly
overlapping, 5—7 X 2.5— 4 mm. obtuse to subacute,
mucronulate, broadly searious-margined, broadly
convex and inconspicuously round-keeled, distally
sparsely punctate with I to several elliptic glands
0.2— 0.3 mm long, sometimes streaked proximally
with linear glands or occasionally glandless, erose-
eiliolate at the apex, otherwise glabrous. Ray florets
5 or 6; corolla yellow' or becoming pink, 4—5 mm

long, the tube 1.5—2 mm long, the ligule slender,
2.5—3 mm long, glabrous. Disk florets 19 to 25;
corolla yellow, weakly 2-lipped, glabrous or glan-
dular-puberulent, 2.5— 3.5 mm long, the tube 0.8-
I mm long, the throat 0.6-1. 2 mm long, the abaxial
lip 1 — 1.3 mm long. 2— 3X as long as the 4 lobes of
the adaxial lip; anthers ca. I mm long; style about
as long as the corolla, included. Aehenes 2.7— 3.8
mm long, strigillose with 2-celled hairs 0.2— 0.3 mm
long. Ray pappus of I or 2 paleaceous-based, an-
trorsely barbed bristles 2—2.5 mm long or some¬
times reduced to scales 0.5 mm long or shorter,
sometimes with 1 or 2 additional scales between
the bristles. Disk pappus ol 15 to 25 unequal pa¬
leaceous-based, antrorsely barbed bristles 1.5—4
mm long. Chromosome number: 2 n = 36M.

Distribution. Pedis pumila occurs only in the
coastal lowlands ol southern Ecuador and northern
Peru at 0—250 m. Habitats include arid scrublands,
roadsides, and beaches. It often occurs in mixed
populations with the closely related Pedis arida
and with P. linifolia L. var. linifolia. Some herbar¬
ium specimens have mixed gatherings of P. arida
and P. pumila. In beach habitats P. pumila grows
in proximity to P. multiflosculosa (DC.) C. H.
Schultz Bipontinus.

Relationships. Pedis pumila is apparently most
closely related to P. arula. Both are much-branched
annuals with leaves densely gland-dotted abaxially.
The sessile or subsessile campanulate eapitula with
broad phyllaries differentiate P. pumila from P. ar¬
ida. which has peduncled eapitula with cylindrical
involucres and narrower phyllaries.

This species has been collected for many years,
but it either has gone undetermined in herbaria or
has been misidentified. Until recently, I believed it
to be the taxon described by II. Cassini (1819) as
Cryptopet alon ciliare (Pedis eryptopetala C. H.
Schultz Bipontinus). I now know that the type (P)
for that name is nonspecific with the type of Pedis
sessiliflora (Lessing) C. II. Schultz Bipontinus. Be¬
cause the specific epithet used by Cassini when he
described the taxon as a member of the genus Cryp-
topetalon is the same as that of a Linnaean species
of Pedis, many botanists have misidentified P. pum¬
ila as Pedis ciliaris L., a very different taxon not
known from western South America.

Chromosome number. A meiotic chromosome
number of 2 n = 36n (diakinesis) was determined
from Keil et al. 19917-D (OBI) and is herein newly
reported. 1 he well-established base number for
Pedis is .v = 12 (Keil. 1977; Keil el al., 1988).
Pedis pumila is therefore a hexaploid taxon.

Volume 12, Number 4
2002

Keil

Pectis from South America

473

Paratypes. ECUADOR. Guavas: Salinas, 7 Apr.
1939, E. Asplund 5620 (S); Salinas, Punt ilia, 1 May 1956,
E. Asplund 20369 (S); along Pacific Ocean betw. Santa
Elena & San Pablo, 17 Mar. 1973, L. Holm-Nielsen et at.
2032 (S); Chanduy, 18 Mar. 1973, /.. Holm-Nielsen et al.
2113 (COL, F, NY, S); Punta de Santa Elena, 21 Mar.
1973, /.. Holm-Nielsen et al. 2306 (F, MO, NY, S. U), 2309
(MO. NY, S), 2337 (NY, S), 2356 (E, MO, NY, S, U);
Anci'in Distr., Santa Elena Peninsula, 7 June 1932, Shep¬
pard s.n. (K): Chanduy, R. Spruce 6476 (RM, E. G, k. EE.
OXF), 6477 (RM, G, OXF. P); Salinas, Puntilla, 1-2 Mar.
1941, H. K. Svenson 11073 (US), 11096 (GH, US); Sali¬
nas, Puntilla, 21 Feb. 1941. II. K. Svenson 1 1 124 p.p.
[mixed with P. arida ] (US); Salinas. 15 Mar. 1941, //. K.
Svenson 11203 (GH, NY. US); Punta Centinela, 6 Apr.
1941, //. K. Svenson 11267 (NY); Punta Carnero. Penin¬
sula de Santa Elena, May 1978. E M. Valverde 2569 (MO).
PERU. Lambayeque: near Motupe, 18 Apr. 1953, R.
Eerreyra 9034 (MO, OBI, USM); 24 km S of Motupe, 29
Mar. 1987, I). J. Keil et al. 1991 1 (F, MO, OBI, UC).
Piura: El Alto, 14 May 1957, H. Ellenberg 1333 (U);
Piura, betw. Amotape & Talara, 19 Apr. 1949, R. Eerreyra
5935 p.p. [mixed with P. arida ] (US); Quebrada Mogollon,
Amotape hills, 29 Mar. 1941, O. Haught & H. K. Svenson
11520 p.p. | mixed with P. arida] (NY); E of La Brea, 25
km SE of Talara, 3 Mar. 1939, 0. B. Horton 1 1531 (G,
GH, UC); 39 km NW of Mari a Vellica, 30 Mar. 1987, D.
J. Keil et al. 19920 (CPUN, E, HUT, MO, NY. OBI, OS,
TEX, UC, USM); betw. Talara & Los Organos, 30 Mar.
1987, D. ,/. Keil et al. 19923 (OBI); Talara. s.d.. E. Smith
15 (MO, USM, OBI); Huancabamba, Serran, 2 Apr. 1939,
11. E. Stork 11363 p.p. [mixed with P. arida] (GH). Tuin-

bes: N of Tumbes, 7 May 1957, //. Ellenberg 1276 (U);
Caleta Cruz betw. Zorritos & Tumbes, 27 May 1957, R.
Eerreyra 12220 (MO); Caleta Cruz, 25 Apr. 1955, R. Eer¬
reyra et al. 10694 p.p. [mixed with P. arida] (OBI, USM);
beach just S of Cancas, 30 Mar. 1987, I). ./. Keil et al.
19925 (ASU, BM. CPUN, F, G, GH, HUT, k. MO, NY,
OBI, RSA, S, UC, US, USM); beach 2 km N of Cancas,
30 Mar. 1987, I). J. Keil et al. 19929 (CPUN, E. HUT,
MO, OBI, USM).

Acknowledgments. Research was supported by
National Science Foundation Grant DEB 81-04683
and a minigrant from California Polytechnic State
University. I am grateful to the curators of the cited
herbaria for the loan of material and/or hospitality
during herbarium visits. I am indebted to Charles
and Eizbeth Burandt with whom I shared Peruvian
adventures.

I .iterature Cited

Cassini, H. 1819. Cryptopetale, Cryptopetalon. P. 123 in
F. Cuvier, Dictionnaire des Sciences Naturelles, 2nd
ed., vol. 12. Paris.

keil. D. J. 1977. Chromosome studies in North and Cen-
tral American species of Pectis (Compositae: Tageteae).
Rhodora 79: 79-94.

- , M. A. Luckow Si I). J. Pinkava. 1988. Chromo¬
some studies in Asteraceae from the United States,
Mexico, the West Indies, and South America. Amer. J.
But. 75: 650-666.

A New Species of Anthurium (Araceae) from Southern Ecuador
and a Revision of the Anthurium oxybelium Schott Complex

Half M. Leimbeck

Department ol Systematic Botany, Institute of Biological Sciences, University of Aarhus,
IMordlandsvej 68, 8240 Risskov, Denmark. Ralf.Leimheck@biology.au.dk

Thomas B. Croat

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

tliomas.croat@mobot.org

ABSTRACT. Anthurium obpyriforme from Loja
province in southern Ecuador is described as new.
This species has been confused with A. oxybelium
Schott hut shows considerable morphological and
ecological differences. Anthurium oxybelium is in¬
troduced as the oldest name of a widespread upland
species, and its relevant synonymy is included. A
key is provided to distinguish Anthurium obpyrifor¬
me from the latter and other related high-altitude
species of Anthurium sect. Belolonchium in Ecua¬
dor.

Key words: Andes, Anthurium, Araceae, Ecua¬
dor.

The exclusively Neotropical Anthurium, with ap¬
proximately 1000 species, is the largest genus in
Araceae (Croat, 1999b). At least 227 named spe¬
cies of Anthurium are known from Ecuador (Croat,
1999a), and many Andean species remain unde¬
scribed, especially those with large cordate leaves
(Croat, 1999b). High-altitude Andean habitats arc
well collected and have relatively few Anthurium
species compared to forests at lower elevations.
Nevertheless, during fieldwork in the Cajanuma
area in the Podocarpus National Park in southern
Ecuador, it was recognized that besides Anthurium
oxybelium, which is the dominant Anthurium above
2500 m, another species was exceedingly abundant
at this altitude. Older collections showed that this
new species had formerly been misidentified and
confused with the widespread A. oxybelium. How¬
ever, there were no difficulties in discriminating be¬
tween the two species in the field, and field obser¬
vations on more than 1000 individuals of both
species and subsequent herbarium studies left no
doubt that the material represents a separate spe¬
cies.

Like Anthurium oxybelium, many other high An¬
dean species of Anthurium arc widely distributed.
Due to the large amount of variability in any spe-

Novon 12: 474-480. 2002.

t ies of Araceae in these montane habitats, some of
them have been described several times, especially
those from mesic upland sites that were the most
readily available habitats to the early collectors
(e.g., Andre, Holton, Lehmann, Purdie). In this
manner A. oxybelium has proven to have many syn¬
onyms, most of which were published by Louis So-
diro. who worked in Ecuador around the turn of the
last century. Anthurium oxybelium is the oldest
name in this complex, and it is related to Anthur¬
ium obpyriforme, the newly described species. Ow¬
ing to this relatedness, the taxonomic complexity of
this common widespread species, the inadequacy
of the original description, the marked change in
its status, and the extensive new synonymy, A. ox¬
ybelium is redescribed later in this paper.

Anthurium obpyriforme Leimbeck, sp. nov.
TYPE: Ecuador. Loja: Podocarpus National
Park, along trail from Cajanuma Visitors Cen¬
ter to Mirador, wet montane forest, 04°05'S,
79°10'W, 2750-3000 m, 29 Oct. 2000, R.
Leimbeck, J. Madsen & R. Windeballe 319 (ho-
lotype, AAL; isotypes, LOJA, M 0-5309365-
67, OCNA). Figure 1.

Planta epiphytica; intemodia brevia, 1.5—3 cm crassa;
caudex ad 5—30 cm longus; cataphylla mox in libras re-
soluta, 9—1 7 cm longa: petioles 25—96 cm longus, 1—9
mm crassus cum geniculo 1 .5-2.5 cm longo; lamina ob-
pyriforma, basi cordata, coriacea, 25—77 cm longa, 1 7—46
cm lata; peduneulus quam petioles duplo brevior, 14-43
cm longus, 2-5 mm crassus; spatha erccta, lanceolata vel
ovata, subviridis vel albida; spadix stipite 541 mm longo
suffultus, 7—22 cm longus, 6—14 cm crassus, viridi-griseus
vel rubineus.

Epiphytic, rarely terrestrial; internodes short,
1.5—3 cm diam.; stem short, generally 5— 30(-85)
cm long; eataphylls 9—17 cm long, drying light
brown to red-brown, usually weathering to a dense
mass ol light yellow fibers; leaves erect-spreading;

Volume 12, Number 4
2002

Leimbeck & Croat
Anthurium from Ecuador

475

Figure 1. Anthurium obpyriforme Leimbeck. — A. Leaf with cataphyll enclosing the petiole base. — B. Spathe and
spadix. — C. Spadix details. — I). Habit. (A from I si m beck & Windeballe 3.39; I?. C from the holotype, Lsimbeck, Madsen
& Windeballe .319.)

petioles 25—96 cm long, 4—9 mm diam., rigid, te¬
rete, esuleate to obtusely silicate adaxially, matte,
light green, drying brown to red-brown; sheath to 5
cm long; geniculum 1.5— 2.5 cm long, slightly thick¬

er than petiole, often red; blade obpyriform, deeply
cordate at base, acute to acuminate at apex, (16-)
25—77 cm long, (6— )17— 46 cm wide, ca. 1.6 times
longer than wide, broadest at petiole insertion, co-

476

Novon

riaceous, matte to semiglossy on both surfaces,
epunctate, deep green on upper surface, paler be¬
low, drying green to grayish green, margin undu¬
late; anterior lobe constricted at middle of blade,
with concave lateral margins, ea. 3 times longer
than posterior lobes; posterior lobes curved inward;
midrib eonvexly raised above, ± sharply V-shaped
below, light green, drying brown; primary lateral
veins 8 to 13 per side, departing midrib at 35^15°
angle, usually oidy lowermost primary lateral veins
prominent and raised in valleys (like a ridge rising
up longitudinally in the center of a groove) on up¬
per surface, the remainder sunken above, all pri¬
mary lateral veins narrowly raised on lower surface,
moderately straight or weakly curved to the margin,
light green, usually drying brown; interprimary
veins usually present, sometimes only near apex;
secondary veins sunken above, raised below, lesser
veins distinct in dry condition; collective vein usu¬
ally arising from the first basal vein, higher order
basal veins also collecting, but ultimately merging
with the margin, 1—3 mm from margin, sunken
above, raised below; basal veins 6 to 10 per side,
the first pair usually free to base, the 5th and higher
order basal veins coalesced to 5—9 cm, eonvexly
raised above, moderately acute below, light green;
posterior rib naked for most of its length along the
sinus, to 0 mm wide, ± flat, light green, at petiole
insertion often red; sinus parabolic to hippocrepi-
form with rounded to truncate apex; inflorescence
erect-spreading; peduncle 14-43 cm long, ca. half
as long as petiole, 2—5 mm diam., slender, terete,
matte, light green; spathe erect, directed at ca. 180°
angle with the peduncle, 6— 22 cm long, 1.5— 8.5 cm
wide, lanceolate to ovate, sometimes naviculiform,
coriaceous, light green to white at anthesis, pluri-
nerved, ± acuminate at apex, cordate at base, in¬
serted at ea. 45° angle on peduncle; stipe stout with
small bulge at base and apex, 5— 8(— 15) mm long,
same color as spathe; spadix spreading to arcuate-
pendent. curved at ca. 90° angle to peduncle, rigid,
tapered, 7—22 cm long, 6-14 mm diam., green-gray,
turning dark ruby; flowers rhombic, 2. 5-3.3 mm
long, 2.2— 2.5 mm wide (dry), 5 to 10 flowers in
principal spiral, 6 to 15 flowers in alternate spiral;
stamens yellow-white, exserted ca. 1 mm above te-
pals when dry; pollen yellow-white. Infruetescenee
not seen.

Anthurium obpyriforme is so far known only from
the western slopes of the Nudo de Sabanilla in Loja
province, at an altitude of 2700-3050 m, which is
near the tree line. The habitat is covered by upper
montane rain forest with ca. 3000 mm precipitation
per year, an annual mean temperature of I 1°C, and

an exceptionally high tree species richness (Mad¬
sen cV 011gaard, 1994) and epiphyte richness
(flpgh. 1992). Due to low canopy height (± 10 m)
and frequent treefalls the forest is quite open lo¬
cally. However, dense bamboo thickets of Chiisquea
sometimes form an almost impenetrable lower un¬
derstory. fhe dominant tree species is Weinrnannia
glabra I,, f. (Cunoniaceae), and the majority of trees
belong to Melastomataceae (Madsen & 011gaard.
1994). Anthurium obpyriforme grows mostly epi¬
phytically on the lower parts of the phorophyte, 0—
3 m above the ground. Among all encountered in¬
dividuals, only 14% were terrestrial, but most of
the terrestrial individuals were small and none of
them were fertile. With more than 750 individuals/
ha A. obpyriforme is very common, and after A. ox-
ybelium it is the second most abundant Anthurium
in di(‘ area. Only three other Araceae species occur
in the same habitat: A. nigrescens Engler, A. lon-
gegeniculatum Engler, and a third, yet undeter¬
mined Anthurium.

Anthurium obpyriforme has been confused with
A. oxybelium. Both species belong to Anthurium
sect. Belolonchium Schott emend. Engler, which is
characterized by coriaceous, cordate leaves with
conspicuous minor veins and cataphylls that usu¬
ally weather into a mass of fibers (Croat & Sheffer,
1983). However, A. obpyriforme differs in being epi¬
phytic with short internodes, while A. oxybelium is
primarily a scandent terrestrial with longer inter¬
nodes. Furthermore, A. obpyriforme has generally
larger leaf blades (25—77 cm long vs. rarely longer
than 30 cm) with strongly constricted anterior lobes
(vs. anterior lobes with ± straight margins), esul-
cate to obtusely silicate petioles (vs. sharply sili¬
cate), cataphylls always weathering into fibers (vs.
cataphylls at upper nodes remaining ± intact),
shorter peduncles (half as long as petioles vs. about
equal), and a shorter stipe (5-8 mm vs. 10-30 mm
long).

Anthurium obpyriforme is also similar to “A. me-
lampyi Croat ined.,” a yet unpublished species
from Colombia and Carchi province in northern Ec¬
uador. That species differs in having long, ribbed
petioles (100—140 cm long), large blades that often
are more than 1 m long, many primary lateral veins
(25 to 30 per side), and the collective vein arising
near the base, from one of the lowermost basal
veins.

Several other related cordate-leaved Anthurium
occur at high elevations in the Ecuadorian Andes,
most of which were placed in section Belolonchium
by Engler (1905). However, the section is poorly
defined and may represent an assemblage of spe¬
cies that could not be assigned to any of the other

Volume 12, Number 4
2002

Leimbeck & Croat
Anthurium from Ecuador

477

sections with cordate leaves (Croat & Sheffer,
1983). Species of Anthurium sect. Belolonchium
sensu Engler (1903) occurring above 2500 m
(Croat, 1999a) are presented in a key at the end of
this paper, to distinguish them from Anthurium oh -
pyriforme. The key is based on herbarium material
at AAU and species descriptions in Engler (1905),
Sodiro (1903b), and Croat and Rodriguez de Sal¬
vador (1995).

Paratypes. ECUADOR. Loja: Podocarpus National
Park, 1-ha sample plot N of Cajanuma Visitors Center,
wet montane forest, 04°05’S, 79°I0'W, 2000 m, 1 Nov.
2000, R. Leimbeck A- B. Windeballe 325 (AAU, LOJA,
QCNA), 4 Nov. 2000. R. leimbeck & B. Windeballe 339
(AAU); E of Nudo de Cajanuma, N of Centro de Infor-
maeidn. 04°05'S, 79°10'W, 2900 m. 20 Sep. 1989, 4.
Bpgh 47838 (AAU, MO). A. Bpgh 47840 (AAU), 26 Oct.
1988, J. Madsen 75508 (AAU); above Cajanuma, trail
from Centro de Intormacion towards Lagunas de Compa-
dre, just below forest limit, wet montane forest, 04°05'S,
79°10'W, 2900-3050 m, 16 Jan. 1989, ./. Madsen 85523
(AAU).

Anthurium oxybelium Schott, Oesterr. Hot.
Wochenbl. 7(39): 310. 1857. TYPE: Colombia.
Nueva Granada: Rfo Hacha. exact location un¬
known. Purdie s.n. (holotype, K).

Anthurium lividispica Sodiro, Anales Univ. Centr. Ecuador
15(108): 14. 1901. Syn. nov. TYPE: Ecuador. Pi-
chincha: El Corazdn, 2800 m. Mar. 1900, Sodiro s.n.
(specimen lost). Ecuador. Cotopaxi: El Corazon, Jan.
1901. Sodiro s.n. (holoneotype, designated here, G;
isoneotype, QPLS).

Anthurium luteolum Sodiro, Anales Univ. Centr. Ecuador
15(108): 13. 1901. Syn. nov. TYPE: Ecuador. Pi-
chincha: Oyacachi, ca. 2800 m, Sodiro s.n. (holo¬
type, Q).

Anthurium patulum Sodiro, Anales Univ. Centr. Ecuador
15(108): 14. 1901. Syn. nov. TYPE: Ecuador. Napo:
Oyacachi, Jan. 1900, Sodiro s.n. (holotype, B; iso¬
types, G, QPLS).

Anthurium psdurum Sodiro. Anales Univ. Centr. Ecuador
15(108): 14. 1901. Syn. nov. TYPE: Ecuador. Pi-
chineha: Oyacachi, ca. 2800 in, Sodiro s.n. (holo-
type, B: isotypes, G. QPLS).

Anthurium puelanum Sodiro, Anales Univ. Centr. Ecuador.
15(108): 17. 1901. Syn. nov. TYPE: Ecuador. Chim¬
borazo: Volean Tungurahua, Puela, Sodiro s.n. (ho¬
lotype, B; isotype. G).

Anthurium sclerophyllum Sodiro, Anales Univ. Centr. Ec¬
uador 15(108): 14. 1901. Syn. nov. TYPE: Ecuador.
Napo: Papallacta— Cuyujua, Feb. 1901, Sodiro s.n.
(holotype, QPLS; isotypes, (7. Q).

Anthurium stuns Sodiro, Anales Univ. Centr. Ecuador
15(108): 17. 1901. Syn. nov. TYPE: Ecuador. Pi-
chincha: Volean Pasochoa, 3000— 120(1 m, Sodiro s.n.
(holotype, B).

Anthurium albaretii J. E. Macbride, Candollea 5: 348.

1934. Syn. nov. TYPE: Ecuador. Piehincha: Volean

Pasochoa, Sodiro s.n. [28] (holotype, B).

Terrestrial or sometimes epiphytic, 0.3-l(-1.5)
m tall: internodes mostly 4—6 cm long, 1.5—2 cm
diam., except much shorter, usually about 1 cm
long near apex, sometimes much longer on older
stems, to 16 cm long; cataphylls 8-13 cm long,
persisting red-brown, intact to semi-intact toward
apex, becoming red-fibrous, eventually deciduous
at lower nodes; petioles subterete, sharply silicate,
19-64 cm long, 4—5 mm diam.. often red near apex;
blades narrowly ovate to triangular or weakly pan-
duriform, (12— )20— 30 cm long, (4— )1 0 — 26 cm w ide,

( 1 .3— )2.3 — 4 times longer than w ide, usually deeply
cordate at base, moderately coriaceous, semiglossy,
slightly bicolorous, usually drying red-brown on
both surfaces, sometimes gray-green: posterior
lobes (2— )4.5— 1 5(— 20) cm long, 2.7— 8(— 1 3) cm
wide, directed toward the base or slightly outward,
sometimes markedly curved inward and overlap¬
ping; margins of the anterior lobes convex to
straight or concave; midrib ± concolorous, narrow¬
ly raised in valley above, convex and paler below;
primary lateral veins (5)8 to 1 I pairs, etched-sunk-
en above, convex below, spreading at 50—75° angle;
tertiary veins etched above, slightly darker than
lower surface; collective veins arising from 3rd bas¬
al veins, sometimes from the first basal veins, 3-4
mm from margin; basal veins (3)4 to 9 pairs, the
1st and sometimes 2nd free to the base, the 3rd
and 4th coalesced to (0.5-)2-4 cm; posterior rib
naked for 1—3.5 cm along the sinus; sinus oblong
to closed to hippocrepiform, rarely parabolic; inflo¬
rescence erect to erect-spreading, held at about the
level of the leaves; peduncle 10—50 cm long; spathe
usuallv held horizontally and hooding spadix, 6-16
cm long, 1-3.5 cm wide, green, sometimes tinged
with red to greenish purple, rarely reddish or red.
long-acuminate to acicular (the acicular portion to
1.5 cm long); stipe 1—3 cm long; spadix (4— )8— 17
cm long, 7-15 mm diam., (5.7— )9— 1 7 times longer
than wide, often curved downward, green, olive-
green, greenish purple, dark purple, purplish
green, greenish brown, brown in age; pistils early;
tepals 3.5-4 mm long (diameter in longitudinal di¬
rection); stamens are exserted at anthesis.

Anthurium oxybelium ranges from Colombia (and
probably Venezuela) to Ecuador (Sueumbios, Napo,
Morona-Santiago, Zamora-Chinchipe, Cart hi. Im-
babura, Piehincha, Cotopaxi, Iungurahua, Canar,
Azuay, Loja) and Peru (Amazonas, Cajamarca, Pas¬
co, Cuzco) at 1400— 4300 m elevation. It is highly
variable, even within a single population, as is in-

478

Novon

dieated by the fact that Sodiro described so many
species from a single locality at Oyacachi. Despite
high variability, these plants share certain things in
common. Typically the intemodes are longer than
broad, though larger or older plants appear to have
thicker stems with shorter intemodes. Most cata-
phylls are persistent, and at least the upper ones
are usually intact, but they are mainly fibrous at
the base. Petioles are subterete and obtusely and
narrowly sulcate. I lie moderately coriaceous blades
vary greatly in shape, being primarily narrowly tri¬
angular to narrowly ovate with a usually obovate or
rarely hippocrepiform sinus and an acuminate
apex. I he margins of the anterior lobe are variable,
most commonly straight or broadly convex, though
sometimes concave. Basal vein numbers are mostly
4 to 6 per side, the first or sometimes the second
pair is free to the base, and several of the remain¬
der are coalesced to 4 cm. file posterior rib is na¬
ked along much of its length. I he inflorescences
are long-pedunculate, and while the peduncles av¬
erage about as long as the petioles, they may be
shorter, as long as, or longer than the petioles.
Spathes are variable, usually lanceolate to narrowly
elliptic, sometimes elliptic, usually green or green
tinged with purple, sometimes purple with green
veins. I he spadix is usually green, but sometimes
purple or red tinged, typically short and weakly
tapered to the apex, generally about as long as the
spathe or shorter. The spadix is commonly 10—17
times longer than wide, but on the stubbier forms
it may range up to little more than 5 times longer
than wide. Flowers art' large for the genus, usually
3.5—4 mm diam. with the stamens exserted at an-
thesis.

For many years this species has been called An-
thurium pulchrum Engler, an illegitimate name
since that name was already published by N. E.
Brown for another species. That illegitimate name
was published by Engler (1898: 449) based on a
Sodiro (Sodiro s.n. [28]) collection from Volcan Pa-
sochoa in Piehincha province. Synonymized here
under .4. oxybeliurn, A. albaretii was intended as a
new name for A. pulchrum by Maebride (1934).

As is the case of many Sodiro collections, no
specimens for A. luteolum were listed in the origi¬
nal description, w hich consisted only of a key. 1 1 is
first mention of any collection is in his first 1903
publication (Sodiro, 1903a: 281). where he stated
that A. luteolum was collected between Papallacta
and Cuyujua. Only a single collection exists with

this information, a collection at the Universidad
Central in Quito (Q).

A neotype was necessary for A. lividispica Sodiro
because no specimens exist from El Corazon dated
March 1900, as was listed by Sodiro. Instead, a
collection from the same locality but collected in
January 1901 has been designated here as the neo¬
type.

Anthurium macrourum was treated as a synonym
of A. patulum (now A. oxybeliurn) in the Catalogue
of the Vascular Plants of Ecuador (Croat, 1999a),
but owing to the lack of a type it appears best to
continue to assume that the species is distinct.
Most existing material of that species, which was
determined by Sodiro, have spadices longer and
proportionately more slender than those of A. oxy-
belium.

Most of the taxa synonymized in this treatment
were described from a few areas in Ecuador, all
very similar in life zone characteristics. Many spe¬
cies were described from a narrow region on the
eastern flanks of the Andes east of Quito, between
Papallacta, Cuyujua, and Oyacachi in Piehincha
province, along the road between Quito and Baeza.

I he remaining species were described from Volcan
Pasochoa or at Puela on Volcan I’ungurahua. Con¬
sidering the variability in this species complex,
none of the types listed above represent anything
out of the ordinary. Sodiro separated A. luteolum by
its having a yellow, rather than green spathe as in
A. oxybeliurn. Anthurium oxybeliurn is apparently
further distinguished in having the blades sagittate-
hastate and somewhat constricted above the petiole
attachment.

Anthurium lividispica and A. psilurum are distin-
quished by having the posterior lobes retrorse with
an ovate sinus, furthermore, A. lividispica has con¬
vex anterior lobes and a thick spadix, 1.3 times as
long as the spathe (vs. straight blade margins and
a slender spadix, twice as long as the spathe in A.
psilurum). All the characters used by Sodiro are
either variable ones or characters that change with
age, so that all the material easily fils into the com¬
plex as it is known today. Throughout its range and
even on a local populational level, A. oxybeliurn is
one of the most variable species in the genus. Spe¬
cies like A. oxybeliurn. A. dombeyanum Brongniart,
and A. nigrescens that range throughout much of the
Andes at high elevations occur in relatively isolated
populations on the tops of mountains, which may
account for the high variability from population to
population.

Volume 12, Number 4
2002

Leimbeck & Croat
Anthurium from Ecuador

479

Key to Hioii Andean Species of Anthvrwm sect. Remlonchium (sensu Enui.er, 1905) from Ecuador

la. Peduncle clearly shorter than petiole; internodes short, rarely 3 cm long.

2a. Flowering spadix sessile or short-stipitate (stipe 0—3 mm long); petioles to 173 cm long.

3a. Epiphyte; petioles many-ribbed; leaf blades prominently constricted above base; spathe white to

light green (northern Ecuador) . 4. jimenae Croat

3b. Terrestrial; petioles sulcate; leal blades ovate-cordate, not constricted above base; spathe red-

tinged, naviculiform (Colombia, Ecuador) . 4. cupulispathum Croat & Rodriguez

2l>. Flowering spadix long-stipitate (stipe 5—20 mm long); petioles to 100 cm long.

4a. Spathe broadly ovate, enclosing the spadix at base, but narrowly open at front; internodes i— 8

cm diam.; geniculum 5—7 cm long; leaves 45—90 cm wide (northern Ecuador) .

. A. gualeanum Engler

4b. Spathe lanceolate to ovate, not enclosing the spadix; internodes to 3 cm diam.; geniculum to 3
cm long; leaves less than 50 cm wide.

5a. Cataphylls usually less than 9 cm long; leaves subelliptic to oblong-sagittate, less than 15
cm wide; spathe purple on both sides with green-yellow linear nerves, longer than spadix

(eastern Andean slopes, Ecuador and Peru) . 4. variegatum Sodiro

5h. Cataphylls more than 9 cm long; leaves ovate to ohpyriform, more than 17 cm wide; spathe
solid green, purple, roseate or white, not purple with green-yellow nerves; shorter than
spadix.

6a. Cataphylls 25-30 cm long; petioles muriculate; leaf blades 60-120 cm long; major
veins with scaly, wing-like appendages on lower surface; spathe purple; stipe 15—20

mm long (Ecuador) . 4. scabrirwrve Sodiro

6b. Cataphy lls 9-20 cm long; petioles glabrous; leaf blades 40-77 cm long; veins without
appendages; spathe green to white or roseate; stipe less than 15 mm long.

7a. Leaves obpyriform, constricted above base, coriaceous; stipe ca. 5—8 mm long
with basal and apical bulge; spathe light green to white (southern Ecuador) . .

. A. obpyriforme Leimbeck

7b. Leaves ovate, subeoriaceous; stipe 12-15 mm long without bulge; spathe roseate

(Colombia to Bolivia) . 4. incurvatum Engler

Ih. Peduncle about equal to or longer than petiole; internodes more than 2 cm long.

8a. Leaf blades 40—70 cm long, 20-35 cm wide; spadix 12—50 cm long.

9a. Stipe 15—30 mm long; petioles plurisuleate; lamina thick, anterior lobe ca. 3 times as long as

posterior lobes; spathe wide lanceolate, ca. I 1 cm long, 3.5 cm wide (Ecuador) .

. A. rigidifolium Engler

9h. Stipe 5—10 mm long; petioles unisulcate; lamina herbaceous to cartilaginous, anterior lobe ca. 4
times as long as posterior lobes; spathe linear to lanceolate, 15—23 cm long, 1—3.5 cm wide.

10a. Lamina cartilaginous; spathe linear, 1 — 1.5 cm wide, purple; spadix 18—20 cm long (northern

Ecuador) . 4. tremulum Sodiro

10b. Lamina herbaceous; spathe lanceolate, 1.5— 3.5 cm wide, green-yellow; spadix 25—50 cm

long (northeastern Andean slope, only known from the type) . 4. vomeriforme Sodiro

8b. Leaf blades 12-35 cm long. 8-26 cm wide; spadix 1—17 cm long.

I la. Peduncle and petioles usually more than 25 cm long; cataphylls 8—13 cm long (Colombia to Peru)

. A. oxybelium Schott

lib. Peduncle and petioles usually less than 25 cm long; cataphylls less than 8 cm long.

12a. Lamina triangular-sagittate, 12—14 cm long, 8—10 cm wide, anterior lobe ca. 4 times as long
as posterior lobes, sinus between basal lobes arcuate to parabolic (southern Ecuador, only

known from the type) . 4. coerulescem Engler

12h. Lamina ovate-sagittate, ca. 25 cm long. 13-15 cm wide, anterior lobe 2-3 times as long as

posterior lobes; sinus between basal lobes hippocrepiform (Venezuela to Bolivia) .

. A. nigrescent Engler

Acknowledgments. We thank Henrik Balslev for
comments on earlier drafts, Lucy T. Smith for her
beautiful illustrations, Jens E. Madsen and Birgille
S. Windeballe for their help in the field, and Zhofre
Aguirre from the Loja herbarium for logistical sup¬
port. We are grateful for support from “Brorsons
Rejselegat” and the Danish Natural Science Ke-
searc h Council (grant #1 1-0390 lo Henrik Balslev).

Literature Cited

Bpgh, A. 1992. Composition and distribution of the vas¬
cular epiphyte flora of an Ecuadorian montane rain for¬
est. Selbyana 13: 25—34.

Croat, T. B. 1999a. Araceae. Pp. 227-246 in P. M.
Jprgensen & S. Leon-Yanez (editors). Catalogue of the
Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri
But. Card. Vol. 75.

- . 1999b. The Araceae of Peru — Distribution, spe¬
cies diversity and centers of endemism. Arnaldoa 6:
45-80.

- & J. Rodriguez de Salvador. 1995. Contributions

to the Araceae flora in northwestern Pichincha Prov¬
ince, Ecuador. Part 1: Anthurium of ENDESA Reserve.
Aroideana 18: 46—148.

- & R. D. Sheffer. 1983. The sectional groupings of

Anthurium (Araceae). Aroideana 6: 85—123.

Engler, A. 1898. Beitrage zur Kenntnis der Araceae. VIII.
Bot. Jahrb. Syst. 25: 352—476.

480

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- . 1905. Das Pflanzenreich. Regni vegetabilis con¬
spectus, Vol. IV. 23 H. Araceae— Pothoideae. Verlag von
Wilhelm Engelmann, Leipzig.

Macbride, J. K 1934. Renamed spermatophytes mostly
Peruvian. Candollea 5: 346-402.

Madsen, J. K. & I?. 0llgaard. 1994. Holistic composition.

structure, and dynamics of an upper montane rain forest
in southern Ecuador. Nordic J. Pot. 14: 403-423.
Sodiro, L. 1903a. Anturios Eeuatorianos. Anales Univ.
Centr. Ecuador 16: 267-282.

- . 1903b. Anturios Eeuatorianos. Monografia 11. Con-

tribuciones al conocimiento de la Flora Fcualoriana, Quito.

A New Micro gramma Subgenus Solanopteris (Polypodiaceae) from
Peru and a New Combination in the Subgenus

Blanca Leon

Museo de Historia Natural tie la Universidad Nacional Mayor de San Marcos, Av. Arenales
1256, Apartado 14-0434, Lima 14, Peru, and Herbarium, Plant Resources Center, Bio Labs
311, University of Texas at Austin, Austin, Texas 78712, U.S.A.
blanca. leon@mail.utexas.edu

Hamilton Beltran

Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Av. Arenales

1256, Apartado 14-0434, Lima 14, Peru

ABSTRACT. Based on a collection from north-cen¬
tral Peru, a new tuberous species of Polypodiaceae
is recognized and described as Microgramma fas¬
ten. It differs from other coenosoric species by the
presence of laminar scales intermixed among the
sori and slender paraphyses with elongate apical
cells possessing clear lumina and thin walls. A key
for species identification in subgenus Solanopteris ,
to which the new species belongs, is provided. In
addition, a new combination is made for a species
previously recognized in the genus Solanopteris.

Key words: Microgramma subg. Solanopteris.
Neotropics, Peru, Polypodiaceae.

In the Neotropics, modified stems resembling tu¬
bers generally inhabitated by ants are known only
in the genus Microgramma subg. Solanopteris (Co¬
peland) Lellinger (Wagner, 1972; Rauh, 1973; Go¬
mez, 1974; Lellinger, 1977; Moran, 1992). This
subgenus currently includes four species: M. bif-
rons (Hooker) Lellinger, M. bismarckii (Rauh) B.
Leon, comb, nov., M. brunei (Werckle ex Christ)
Lellinger, and M. tuberosa (Maxon) Lellinger, all
with dimorphic fronds. In Peru, two species have
been reported: M. bifrons from the Departments of
Amazonas, Cusco, Loreto, Madre de Dios, Pasco,
San Martin, and Ucayali, growing below 1000 m
elevation; and M. bismarckii from Cusco, Pasco,
and San Martin from altitudes above 1000 m (Tryon
& Stolze. 1993). Based on a collection made by the
second author in the Department of Ucayali at 1220
m, a new species is recognized and described here.

Microgramma fosteri B. Leon & II. Beltran, sp.
nov. TYPE: Peru. Ucayali: Cordillera Azul del
Biabo, cabeceras del no Pisqui, 8°28'45.6"S,
75°43'5.21”W. 1220 m, 12 Sep. 2000, //. Bel¬
tran, R. B. Foster & W. Alverson 3643 (holo-
type, USM). Kigures 1, 2A, B.

A Microgramma bismarckii et M. tuberosa squamis lam-
inaribus inter soros positis et paraphysibus gracilibus, cel-
lulis apicalibus elongatis luminibus hyalinis praeditis re-
cedit.

Epiphyte (Fig. 1A). Rhizomes long-creeping,
1.5—2 mm wide, branched, glaucous to dark brown,
apices and young parts covered with broadly ellip¬
tic non-clathrate peltate scales 0.5—0.75 mm long,
white or brown with white margins (Fig. 1B-E); tu¬
berous rhizome hollow, 1.5—2 cm wide, covered
with long acuminate scales from a round peltate
base, 1.4-1. 5 mm long (Eig. IE), shiny dark-brown
in the center. Phyllopodia short, inconspicuous.
Leaves dimorphic, 0.2— 0.5 cm apart, subcoria-
ceous, simple with entire margins, subsessile to
short -pet iolate, petiole 1—2 mm long. Trophophylls
(Fig. 1C) obovate to broadly elliptic, apex obtuse,
1.3—2 X 0.8— 1.2 cm, scattered scales on both sur¬
faces; scales linear, 1.1-1. 4 mm long (Eig. 1 H, 1);
costa prominent abaxially % of leaf length, primary
veins slightly prominulous. Sporophylls (lig. 1J. K),
oblong, apex retuse. 1.2—2 X 0.7— 0.8 cm, adaxially
with scattered dark brown scales, persistent along
margins, sporangia and paraphyses borne on a 2
mm broad receptacular line completely covering
the abaxial surface at maturity, with dark brown
laminar scales (Eig. 1L, M) 1 X 0.5 mm, base pel¬
tate with 0.1 mm long stalk, intermixed with spo¬
rangia and paraphyses. Paraphyses hairlike, 0.8—1
mm in length. 8—10 cells long. I cell wide, apical
cell hyaline, 40-60 pun long (Eig. 2 A, B). Spores
65—75 ^un long, with spines 8-13 /am long.

The specific epithet honors Robin B. Foster for
his work in Peru and his efforts in neotropical con¬
servation. Microgramma fosteri is characterized by
its leaves with entire margins, scattered persistent
abaxial scales, acrostichoid coenosori, I mm long

Novon 12: 481-485. 2002.

482

Novon

laminar scales intermixed with the sporangia, 1
cell-wide paraphyses with elongate apical cells
having clear lumina, and spores with long spines
8—13 pun. It differs from 17. bifroris and M. Brunei
by its trophophylls with entire margins, acrosti-
ehoid coenosori, and paraphyses with elongate api¬
cal cells. It can he distinguished from M. bismarckii
and M. tuberosa, both of which also have coenosori,
by its trophophyll with a retuse apex, completely
covered abaxially with sporangia when mature (vs.
having a nonsoriferous expanded apex), the pres¬
ence of shiny dark brown laminar scales among the
sporangia (vs. none or inconspicuous), and the pa¬
raphyses with elongate cells, clear lumina, and thin
apical cell walls (vs. having blunt apical cells with
dark lumina). Microgramma bismarckii and M. tub¬
erosa, which are known from few collections, are
poorly understood taxonomieally and may represent
different phases of a single species.

Wagner (1986) studied soral patterns in pleopel-
tid ferns and commented on the parallel situation
in Solanopteris and Microgramma with regard to
soral series ranging from discrete to confluent sori
to coenosori. The soral pattern in this newly de¬
scribed species links coenosoric development with
a reduction of the fertile blade. The presumed clos¬
est relative. M. bismarckii, better represented in
herbaria, presents laminar expansions beyond the
proximal and distal border of the sori but no con¬
current frond reduction. In the overall distribution
ol Microgramma subg. Solanopteris , coenosoric
species occupy altitudes above 1000 m.

I axonomic characters used for species recogni¬
tion in subgenus Solanopteris have included leaf
morphology, venation patterns, and soral shape.
However, with the exception of Wagner (1972). lit¬
tle attention has been paid to other characters such
as those found in paraphyses. Paraphyses in Solan¬
opteris can be described as one cell w ide for most
of their length. Differences from species to species
appear in the distal portions, where variation occurs
in cell number, dimensions, shape, and coloration
of the apical cell lumina (Fig. 2A-E). The distinc¬
tive acrostichoid coenosori in M.fosteri mostly con¬
tain thin paraphyses with elongate cells, and the
most distal cells have a clear lumina and thin walls

(fig. 2A. 8). The paraphyses in this new species
do not resemble those found in the discrete-sori
species (fig. 2C, I)), where the most distal cells are
broader than long and the conformed apex has one
or two blunt cells. They differ from those other coe¬
nosoric species (Fig. 2F), in which the distal cells
are also broader than long, and the terminal cell
has the lumina dark yellow-brown filled with tannin
at maturity. This is similar to the situation in Po¬
lypodium virginianum L. (Peterson & Kott, 1974)
in which the marginal cells are similarly darkened.

Affinities between subgenus Solanopteris and
some species of subgenus Microgramma are man¬
ifest in the paraphyses (e.g., Sota, 1986). For M.
baldwinii Brade and M. ulei (Ule) Stolze, both also
found in Peru, paraphyses are one cell wide. In M.
baldwinii, the three to four of the most distal cells
are broader than long. In some cases, the third row
is bicellular, as in M. bifroris and M. Brunei, and
the apical cell is hooked, with the lumina filled with
tannin. In M. ulei, the apical cell is blunt and dark,
more like those seen in M. bismarckii. Further re¬
search on paraphyses development and character¬
istics for these and other species of Microgramma
may contribute to a better understanding of these
affinities.

Nkw Combination

Microgramma bismarckii (Rauh) B. Leon. comb,
nov. Basionym: Solanopteris bismarckii Rauh.
Abh. Akad. W iss. Abh. Lit. Mainz, Math.-Na-
turwiss. Kl. 5: 232. 1973. TYPE: Peru. Pasco:
Oxapampa, 1800-2000 m, Ceja de la Mon¬
tana. July 1971, Werner Rauh 25790 (holotype,
HEID).

The following key distinguishes all currently rec¬
ognized species of Microgramma subg. Solanopter¬
is.

kiT to i ni. Si’kciks oi Microgramma si no. Soiaaorteris

1. Sporophylls with discrete sori; leaves charta-
ceous to subeoriaceous; paraphyses witli blunt

apical cells . 2

L. Sporophylls only with coenosori, occasionally
intermixed discrete sori present; leaves eoria-

Figure 1. Microgramma fosteri 8. Lertn & H. Beltrdn. — A. Habit. — B-E. Rhizome scales. — I). Central basal portion
of rhizome scale. — F. Tuber scale. — G. Abaxial view of trophophyll. —If. Adaxial trophophyll scale. — 1. Abaxial
trophophyll scale. J. Abaxial view ol sporophyll. — k. Adaxial view of sporophyll. — L. Adaxial sporophyll scale. —
M. Abaxial sporophyll scale. Drawn from Beltran. Foster & Alverson 5824 (USM). Stippled areas represent dark lumina
cells; striped areas represent insertion. The 0.2 mm bar applies to B, C, E, F, H, I, L, and M. The 0.8 cm bar applies
to G, J, and k.

Volume 12, Number 4
2002

Leon & Beltran

Microgramma fosteri from Peru

483

0.8 cm

B

200 iim

130 fim

Volume 12, Number 4
2002

Leon & Beltran

Microgramma fosteri from Peru

485

ceous; paraphyses with elongate apieal cells or
broader than long apical cells with lumina filled

with tannin . 3

2(1). Trophophyll margins lobed; leaves chartaceous

. M. hi from

2'. Trophophyll margins entire or rarely slightly re-

pand; leaves subcoriaceous . M. brunei

3(1). Sporophylls with retuse apex; sori acrostichoid;
dark brown laminar scales dispersed on both
surfaces; paraphyses with elongate apical cells

and terminal cell without tannin . M. fosteri

3'. Sporophylls with mucronate to acute apex; sori
elongate to discrete; laminar scales absent or
inconspicuous; paraphyses with distal cells
broader than long; terminal cell brown filled

with tannin . 4

4(3). Trophophyll apex mucronate . M. bismarckii

4'. Trophophyll apex acute or obtuse . . M. tuberosa

Acknowledgments. We thank the Environmental
Conservation Programs of The Field Museum, Chi¬
cago, for inviting the second author to participate
in the field expedition to Cordillera Azul. We also
thank Peter Jprgensen, John M. Mendenhall, Car¬
men Ulloa Ulloa, Tyana Wachter, and Kenneth
Young for their help and support during the prep¬
aration of the manuscript. For providing insightful
comments, suggestions, and literature we thank
George Yatskievych. We thank llenk van dec Werff
for his comments and help with the Latin diagnosis.
We thank David Dellinger for providing information
on paraphyses characters of Microgramma tuberosa

and Alan R. Smith for providing early comments
and photocopies of Solanopteris at UC. We also
thank Robbin C. Moran and an anonymous review¬
er for their comments.

Literature Cited

Gomez, L. 1). 1974. Biology of the potato-fern Solanopteris
brunei. Brenesia 4: 37—61.

Lellinger, D. B. 1977. Nomenclatural notes on some ferns
of Costa Rica, Panama, and Colombia. Amer. Fern J.
67: 58-60.

Moran. R. C. 1992. The potato fern. Fiddlehead Forum
19; 18-20.

Peterson, R. L. & L. S. Kotl. 1974. The sorus of Polypo¬
dium virginianum : Some aspects of the development
and structure of paraphyses and sporangia. Canad. J.
Rot. 52: 2283-2288.

Ranh. W. 1973. Solanopteris bismarckii Rauh. Abb. Akad.
Wiss. Abb. Lit. Mainz, Matli.-Naturwiss. Kl. 5: 223-
256.

Sota, F. 1986. Sobre la posicion sistematica tie Polypo¬
dium fuscopunetatum Hook, y Polypodium percussum
Cav. (Polypodiaceae s. str., Pteridophyta). Physis (Buen¬
os Aires) 44: 19-28.

Tryon, R. M. & R. G. Stolze. 1993. Pteridophyta of Peru.

Part V. Solanopteris. Fieldiana Bot. n.s. 32: 178-183.
Wagner, W. H., Jr. 1972. Solanopteris brunei, a little-
known fern epiphyte with dimorphic stems. Amer. Fern
J. 62: 33-43.

- . 1986. The New World fern genus Marginariopsis

(Polypodiaceae): An example of leaf dimorphy and coe-
nosory in generic delimitation. Bull. Torrey Bot. Club
113: 159-167.

Figure 2. Paraphyses. A, B. Paraphyses of Microgramma fosteri. Drawn from Beltran , hosier & Alverson 3824 (U5M).
— A. Paraphyses. — B. Detail of distal portion. — C. Paraphyses of Solanopteris bifrons. Drawn from Dick 270 (MO).

_ I). Paraphyses of Solanopteris brunei . drawn from Forero et al. 6022 (MO). — F. Paraphyses of Solanopteris bismarckii,

drawn from Rauh 35685 (LISM). — F. Paraphyses of Solanopteris bismarckii. two left figures drawn from Ceron et al.
5790 (MO), two right figures from G entry & Smith 45184 (MO). The 200 pm bar applies to A, C, I), and E. The 130
gm bar applies to B and F.

Two New Taxa of Maianthemum (Convallariaceae) from
Northwestern Yunnan, China

Li Rang and Li Heng

Kunming Institute of Botany, I he Chinese Academy of Sciences, Kunming, Yunnan 650204,
People’s Republic of China, lirong_mail@yahoo.com, liheng@public.km.yn.cn

ABSTRACT. As a result of expeditions to the Gao-
ligong Mountains in western Yunnan, China, the
authors discovered two new taxa. Maianthemum
dulongense H. Li var. coriaceum R. Li & H. Li and
M.fuscum (Wallich) LaFrankie var. cordatum R. Li
& H. Li are described, and their differences from
the type varieties are discussed.

Key wards: China, Convallariaceae, Maianthe¬
mum.

Maianthemum 0. Weber ex Wiggers (including
Smilacina Desfontaines) is a genus of about 35 spe¬
cies distributed widely in the northern temperate
region, the Himalayas, as well as subtropical mon¬
tane Asia and Central America (Mabberley, 1697;
Chen & Kawano, 2000). In the past (Wang & Tang.
1978; Dahlgren, 1980), it was separated into two
genera, Maianthemum and Smilacina , based on
whether the (lowers are dimerous (4 tepals, 4 sta¬
mens, 2 carpels) or trimerous (6 tepals, 6 stamens,
3 carpels). However, because of their overall sim¬
ilarity, the two genera have been continuously com¬
bined (Pursh, 1814; Link, 1821; Greene, 1888).

I herman (1956) challenged the separation of the
two genera on the basis of their uniform karyotype.
LaFrankie (1986a, 1986b) studied the New World
species of Maianthemum and concluded that Smi-
lai inn and Maianthemum should be combined. In
his note, LaFrankie summarized the evidence for
this combination and transferred the species of
Smilacina to Maianthemum (1986b). Along with Li
(1990) and Chen and Kawano (2000), the present
authors agree with LaFrankie’s transfers.

During the summer of 2000, a botanical expe¬
dition was carried out in Gongshan County and l)u-
longjiang Valley, Northwest Yunnan Province, Chi¬
na, to study the flora of the Gaoligong Mountains.
In addition to the two new varieties (Maianthemum
dulongense var. coriaceum and M. fuscum var. cor¬
datum) herein described, eight other species of
Maianthemum were found in this region: M. atro-
purpureum (Franchet) LaFrankie, M. dulongense H.
Li, M. fuscum (Wallich) LaFrankie, M. gongsha-
nense (S. Y. Liang) H. Li, M. henryi (Baker) La-

Novon 12: 486-491. 2002.

Frankie. M. oleraceum (Baker) LaFrankie, M. pur-
pureum (Wallich) LaFrankie, and M. tatsienense
(Franchet) LaFrankie.

LaFrankie (1986b) transferred all the species of
Smilacina to Maianthemum ; however, in his note
he did not present an infrageneric system for
Maianthemum. Hara (1987) studied the Asiatic
species of Smilacina (Hiroshi Hara passed away in
September 1986 and had not seen LaF rankie’s pa¬
lter about the transfer of this genus) and gave a
synopsis for the infrageneric system. In his note
(Hara, 1987). Smilacina can be divided into four
sections based on the combination of several sig¬
nificant features, such as corolla shape, insertion of
stamens, sexuality of flowers, and the surface struc¬
ture of pollen grains. Haras (1987) infrageneric
classification of the genus Smilacina is as follows:
Sect. I. Smilacina, Subsect. la. Smilacina ; Sub¬
sect. lb. Dioica Hara; Sect. 2. Tubifera Hara; Sect.
3. Oligobotrya (Baker) Hara; Sect. 4. Medora
(Kunth) Hara. Li (1990) (the second author) agreed
w ith LaFrankie s taxonomic treatment of Smilacina
and studied all the species of Maianthemum in the
world. In her note (Li, 1990), according to the
shape of the rhizomes, the number of foliage leaves,
the branching pattern of the inflorescence, the ba¬
sic number and the color of the flowers, a new in¬
frageneric system for Maianthemum was presented,
which divided the genus into two subgenera and
five sections. Also, all the species of Maianthemum
were recombined and rearranged as a new classi¬
fication system. Li’s infrageneric system of Maian¬
themum, based on Li (1990), is as follows:

I. Maianthemum subg. Medora (Kunth) H. Li
Individual rhizome units swollen and tuberous;
flowers trimerous; foliage leaves numerous.

I. Section Medora

Individual rhizome units spherical or sub-
spherical, ovoid; corolla cyathiform, purple,
or ashy pale, rarely white.

a. Subsection Medora
Inflorescence a panicle.

b. Subsection Dulongensis 11. Li
Inflorescence a raceme.

Volume 12, Number 4
2002

Li & Li

New Taxa of Maianthemum

487

2. Section Oligobotrya (Baker) H. Li
Individual rhizome units ovoid; com 11a tub-
iform; inflorescence racemose or paniculate
with a few branches, main axis with a single
flower at each node or with 2 to 7 flowers
clustered at each node; stamens inserted the
throat of the corolla.

3. Section Tatsienensis U. Li

Individual rhizome units claviform or hori¬
zontal cylindrical; corolla cyathiform.

II. Maianthemum subgenus Maianthemum

Individual rhizome units extended and stolon-

iferous; flowers dimerous or trimerous, snow

white; foliage leaves 2 to 4.

1. Section Smilacina (Desfontaines) H. Li
Flowers trimerous.

2. Section Maianthemum
Flowers dimerous.

According to Li’s infrageneric system, the new
variety Maianthemum dulongense var. coriaceum,
with suhspherical individual rhizome units, numer¬
ous leaves, racemose inflorescence, trimerous flow¬
ers, cyathiform corolla, and maroon perianth, be¬
longs to subsection Dulongensis H. Li of section
Medora. The new variety Maianthemum juscum var.
cordatum, with moniliform individual rhizome
units, numerous leaves, paniculate inflorescence,
trimerous flowers, cyathiform corolla, and purple
perianth, belongs to subsection Medora of section
Medora.

\f aiantheniuni dulongense H. Li var. cori-
aceum R. Li & H. Li, var. nov. TYPE: China.
Yunnan: Gongshan Xian, E side of Gaoligong
Mountains, on the trail from Qiqi to Dongshao
Fang, wet sloping meadow, 27°41,23"N,
98°28'26"E. 3400-3600 m, 17 July 2000, Li
Heng with Bruce Bartholomew, Philip Thomas,
Peter Fritsch, Dao Zhi-lin, Wang Zhong-lan &
Li Bong 12721 (holotype, KUN). Figure 1A— E.

A Maianthemo dulongensi var. dulongensi folio cori-
aceo, cordato-oblongo, nervis 21, nervillis conspicuis, flo-
ribus 24, perianth! marronini tepalis interioribus exterior-
ibus longioribus differt.

Terrestrial herb, 22—28 cm tall. Roots uniform.
10 to 15 per rhizome unit, at nodes and internodes,
8-13 cm X 0.5—1 mm. Rhizome a sympodium, 3—
4 cm long, the individual units suhspherical,
densely connected, 4-8 mm diam., stem scar on
node conspicuous, orbicular, 3—4 mm diam., inter¬
nodes very short. Leafy stem upright, 16-18 cm
long, dark purple, densely pubescent; foliage leaves
5; internodes 1-2 cm long, shorter apically. Leaf
sessile or subsessile; blade deep green, coriaceous.

above cordate to oblong, glabrous, apex short-acute,
base cordate, 3.6— 4.3 X 2.3-3 cm. veins 21. dense-
ranked, conspicuous, venulae between veins slight¬
ly conspicuous. Inflorescence 10 cm long, a cylin-
drie raceme with 24 flowers, fertile axis erect or
arching upward, densely pubescent. Flowers trim¬
erous; pedicel 2-5 mm long, densely pubescent,
with a broad triangular bract; perianth maroon, cu-
puliform, ca. 4 mm diam.; tepals oblong, cuspidate
apex, outer ca. 3X2 nun, inner longer, ca. 4 X 2
mm; stamens 6. white, filaments short, ca. 1 mm
long, anthers ovate, ca. I mm long; ovary conical,
pubescent, 3 locules, style inconspicuous, stigma
3-lobed. Flowering in July. Fruit not seen.

Distribution and habitat. Known only from the
type locality, where it has been collected in pri¬
marily evergreen broadleaf forest at 3400—3600 m.
This new variety is common in wet sloping mead¬
ows by rivers.

Maianthemum dulongense var. coriaceum differs
from variety dulongense in having a coriaceous,
cordate-oblong leaf blade with 21 veins and con¬
spicuous venulae, an inflorescence with 24 flowers,
a maroon perianth, and inner tepals longer than
outer tepals. By contrast, M. dulongense var. dulon¬
gense has a papyraceous, ovate leaf blade with 15
veins and inconspicuous venulae, an inflorescence
with 6 to 15 flowers, and a light purple perianth.

Maianthemum dulongense var. coriaceum is re¬
stricted to the east side of the Gaoligong Mountains
and occurs in a geographical area allopatric to the
typic variety, which occurs on the west side of the
Gaoligong Mountains and extends to southeastern
Xizang (11. Li. 1990, 1997). In our opinion, the
shape of the rhizomes, the number of foliage leaves,
the branching pattern of the inflorescence, and the
basic number and color of the flowers are usually
important diagnostic features for evaluating species
limits throughout the genus. However, the variety
coriaceum is distinguished from the typic variety by
the shape of the leaf blade, the number of veins,
and the number of flowers. Other features are the
same, and hence we recognize it as a variety of M.
dulongense.

Maianthemum dulongense var. dulongense was
first described in 1990 in Acta Botanica Yunnanica,
Suppl. 3. In Li’s (1990) infrageneric system of
Maianthemum, it was included in section Medora
subsect. Dulongensis because of its moniliform in¬
dividual rhizome units, numerous leaves, racemose
inflorescence, trimerous flowers, and cyathiform co¬
rolla.

488

Novon

2cm

2 mm

2 cm

Figure 1. A-F. Maianthemum dulongense H. Fi var. coriaceum R. Li & H. Li. — A. Rhizome, leafy shoot, and
infructescence. B. I.eaf. Fepals and stamens. — I). Gynoecium. — E. Ovary, longitudinal section. F. Maianthe¬
mum fuscum (Wallich) LaFrankic var. cordatum R. I.i & H. Li. — F. Leaf. (Drawn by Wang Ling from the holotypes.)

Volume 12, Number 4
2002

Li & Li

New Taxa of Maianthemum

489

Figure 2. Maianthemum fuscum var. cordatum (Diilong Jiang Expedition 6854, holotype, KUN).

Maianthemum fuscum (Wallich) LaFrankic var.
cordatum R. Li & H. Li, var. nov. I 't PI .:
China. Yunnan: Gongshan Xian, Dulnng Jiang
Valley, W side of Gaoligong Mountains, under
forest, 2200 rn, 16 May 1991, Dulong Jittng
Expedition 6854 (holotype, KUN; isotype,
MO). Figures 1 F, 2.

A Maianthemo fusco var .fusco folio cordato, floribus 20.
pedicello tenui differt.

Terrestrial herb, 20-40 cm tall. Roots uniform,
8 to 13 per rhizome unit, at nodes and internodes
7—15 cm long. Rhizome a sympodium, 3—8 cm

long, the individual units moniliform or subcylindr-
ical, densely connected, 1—1.5 cm diam., stem scar
on node conspicuous, discoidal, 3.4 mm diam., iu-
ternodes very short. Leafy stem arcuately ascend¬
ing. 15-30 cm long, green, cylindrical, glabrous;
foliage leaves 3—7; internodes 1.5—3 cm long. Leaf
petiolate, 2-3 cm long; blade green, chartaceous,
glabrous, ovate-cordate, 7-12 X 4-6.5 cm, abrupt¬
ly attenuate with an acumen of 1.1— 1.8 cm at the
apex, deeply cordate at the base. Inflorescence 5—
10 cm long, a panicle with 20 flowers, raehis con¬
spicuously zigzagged, glabrous; shortly peduncu¬
late. 8—15 mm long, glabrous, bractless. Flowers

490

No von

Irimerous; pedicel tenuous, 4—9 mm long, glabrous,
with a ca. 1 mm long triangular bract at the base;
perianth purple, subrotate, ca. 5-8 mm diam., 6-
lobed nearly to the base, connate part ca. 0.5 mm
long; tepals ovate, obtuse at the apex, outer 3 lobes
much smaller, ca. 2X2 mm, inner 3 lobes larger,
ca. 3 X 3 mm wide; stamens 6. light yellow, fila¬
ments ovoid, earnose, white, ca. 0.5 mm long, in¬
serted to the base of the corolla lobes, anthers
small, roundish, yellow, ca. 0. 4-0.5 mm long, bas-
i fixed; ovary ovate, ca. 1 mm long, 3 locules, style
very short, pyramidal, ca. 0.4 mm long, stigma ob¬
scurely 3-lobed. Flowering in April-Mav. fruiting
in June-December.

Distribution and habitat. The new variety oc¬
curs in northwestern Yunnan and southeastern Xiz-
ang, China. The type specimen was collected in
Dulong Jiang Valley by the Dulong Jiang expedi¬
tion in 1991. The plant grows in evergreen broad-
leaf forest at 1800—3000 m.

Maianthemum fuscum var. cordatum differs from
variety fuscum in having a cordate leaf blade, an
inflorescence with 20 flowers, and a tenuous pedi¬
cel. By contrast, variety fuscum has a lanceolate
leaf blade, an inflorescence with 3 to 10 flowers,
and a filiform pedicel.

Maianthemum fuscum var. cordatum is restricted
to the northern Gaoligong Mountains and southern
Xizang and occurs in a geographical area sympatrie
with the typic variety, which occurs in western Yun¬
nan, southern Xizang, and extends to eastern Him¬
alaya (including Nepal, Sikkim, Bhutan, northeast¬
ern India, and northern Burma) (Noltie, 1994; H.
Li, 1997). As mentioned earlier, we consider the
shape of the rhizomes, the number of foliage leaves,
the branching pattern of the inflorescence, and I he
basic number and the color ol the flowers usually
to be the important diagnostic features for evalu¬
ating species limits throughout the genus. However,
tin* shape of the blade and the number of flowers
are quite variable for variety fuscum. With the ex¬
ception of the minor differences mentioned above,
other morphological features between the new taxon
and variety fuscum are the same, and hence we
recognize this taxon as a variety of M. fuscum.

Maianthemum fuscum var. cordatum resembles
M. fuscum var. pilosum (Hara) S. Karthikeyan but
grows allopatrie to it (the latter occurs in Nepal and
Bhutan [Hara, 1987]), and it can be separated by
a few constant characters. Variety cordatum is dis¬
tinguished from variety pilosum in having a gla¬
brous leaf blade and the rachis of the inflorescence
conspicuously zigzagged and glabrous. By contrast,
variety pilosum is ciliate on the margin of tin1 leaf
blade, and the rachis of the inflorescence is almost
straight, sometimes with still spreading hairs.

In Hara s (1987) opinion, Maianthemum fuscum
var. fuscum belonged to Smilacina sect. Medora ,
but it was subsequently transferred to Maianthe¬
mum by LaFrankie (1986b). In Li's (1990) infra¬
generic system of Maianthemum, it was rearranged
in section Medora subsect. Medora because of its
moniliform individual rhizome units, numerous
leaves, paniculate inflorescence, trimerous flowers,
cyathiform corolla, and purple perianth. Therefore,
both M. fuscum and M. dulongense belong to the
same subgenus and section, but in different sub¬
sections.

Paratypes. CHINA. Yunnan: Congslian Xian, Dulong
Jiang Valley, W side of Gaoligong Mountains, in evergreen
forest, 2200 in. 20 May 1991, Didong Jiang Expedition
6939 (KUN): Zhiyenandai, in broad leaf evergreen forest,
22(H) in. 4 Se|>. 1982, Qizang Expedition 9918 (KUN);
Fugong Xian. Ouanmugulu-.VIudeng, 2500-3000 m, 31
July 1979, Nujiang Expedition 791658 (KUN). Xizang:
Dingjie Xian, Chengtangqi, in broad leaf forest, 2300 m. 6
June 1975, Qizang Expedition 5545 (KUN); Medog, Han-
mi, under forest, 2300 m, 26 Oct. 1992, Expedition to
Medog 0571 (KUN). 2100 in, 27 Oct. 1992, Expedition to
Medog 0846 (KUN), 1900 m, 28 Oct. 1992, Expedition to
Medog 0938 (KUN); Beiben, the back hill of Xirah, under
forest, 2200 m, 8 Dec. 1992, Expedition to Medog 2074
(KUN); I’angxin. Pangguo, in forest, 2100 m, 28 Feb.
1993, Expedition to Medog 4090 (KUN); Damn, under for¬
est. 2000 m, 6 Mar. 1993, Expedition to Medog 4234
(KUN); Rengqiangpeng, in forest. 2000 m. 22 Apr. 1993,
Expedition to Medog 5691 (KUN); Denxin. Wenlang, in
forest, 18(H) m, 29 Apr. 1993, Expedition to Metlog 6069
(KUN).

I he following key to the species of Maianthe¬
mum in the Gaoligong Mountains includes the two
new varieties described herein.

Key to the Species of M.manthemi \ t in the Gaoligong Mot ntains

la. Corolla distinct or connate at the base.

2a. Inflorescence a raceme.

3a. Leaf papyraceous, ovate; blade with 15 veins and inconspicuous venulae; inflorescence with 6 to 15

flowers; perianth light purple . W. dulongense var. dulongense

31). Leaf coriaceous, cordate-oblong; blade willi 21 veins and conspicuous venulae; inflorescence with 24

flowers; perianth maroon . ,W. dulongense var. coriaceum

21). Inflorescence a panicle.

Volume 12, Number 4
2002

Li & Li

New Taxa of Maianthemum

491

4a. Plant pubescent.

5a. Foliage leaves more than 4; inflorescence with more than 10 flowers.

6a. Leaf elliptic-ovate to broadly lanceolate, 12-21 cm long, long-cuspidate at the apex; style

conspicuous, 2—2.5 mm long . M. oleraceum

6h. Leaf elliptic to oblong, 7-13 cm long, acuminate at the apex; style short, 1.2 mm long . . .

. M. purpureum

5b. Foliage leaves 2; inflorescence with I to 4 flowers; plant 5—20 cm tall; leaf ovate to elliptic-

ovate, pubescent, 2—5 cm long, 1 .5—3 cm wide . M. gongshanense

4b. Plant glabrous.

7a. Petioles 1—4 cm long; tepals ovate, 3—4 mm long; stigma obscurely 3-lobed.

8a. Blade lanceolate; inflorescence with 3 to 10 flowers, pedicel filiform . . M. fuscum var. fuseum

8b. Blade cordate; inflorescence with 20 flowers, pedicel tenuous . M. fuscum var. cordatum

7b. Petioles short, less than l cm long; tepals narrowly lanceolate, 2-2.5 mm long; stigma deeply 3-

lobed . M. tatsienense

lb. Corolla conspicuously connate.

9a. Plant 30-80 cm tall; rhizome moniliform; flowers salver-shaped; corolla tube cylindrie, 6—7 mm long;

style ca. 2 mm long, stigma 3-lobed; berries green with purple spots . M. henryi

9b. Plant 80-150 cm tall; rhizome nodose; (lowers broadly eampanulate or rotate; corolla tube cupulate, 1-2

mm long; style ca. 0.5 mm long, stigma obscurely 3-lobed; berries red . M. alropupureum

Acknowledgments. 'Fite study is supported by
the National Nature Science Foundation of China
(grant no. 39670086), the National Geographic So¬
ciety (U.S.A., grant #6578-99), and the National
Science Foundation (U.S.A., award no. DEB-
0103795). We are gratefid to Wang Ling for pre¬
paring the drawing.

Literature Cited

Chen, S. C. & S. Kawano. 2000. Two new combinations
in Maianthemum (Convallariaceae). Novon 10: 113—

1 14.

Dahlgren, B. M. T. 1980. A revised system of classifica¬
tion of the angiosperms. Bot. J. Linn. Soc. 80: 91-124.
Greene, E. L. 1888. Bibliographic notes on well-known
plants. IX. Bull. Torrey Bot. Club 15: 285—287.

Hara, H. 1987. Notes towards a revision of the Asiatic
species of the genus Smilacina. J. Fac. Sei. Univ. Tokyo,
Sect. 3, Bot. 14: 137-159.

LaFrankie, J. V., Jr. 1986a. Morphology and taxonomy of

the New World species of Maianthemum (Liliaceae). J.
Arnold Arbor. 67: 371—439.

- . 1986b. Transfer of the species of Smilacina to

Maianthemum (Liliaceae). Taxon 35: 584—589.

Li, 11. 1990. Infrageneric system of the genus Maianthe¬
mum. Acta Bot. Yunnan., Suppl. 3: 1-12.

- . 1997. Liliaceae. In: C. Y. Wu (editor). Flora Yun-

nanica 7: 749. Science Press, Beijing.

Link, H. F. 1821. Enum. PI. hort. Kegii Bot. Berol., alt.
Pars. 1. Berol. Reimer.

Mabberley, I). J. 1997. The Plant-Book: A Portable Dic¬
tionary of the Vascular Plants, 2nd ed. Cambridge Univ.
Press, Cambridge.

Noltie, H. J. 1994. Convallariaceae. In: A. .1. C. Grierson
& 1). G. Long, Flora of Bhutan 3(1): 47. Royal Botanic
Garden Edinburgh.

Pursh, F. T. 1814. Flora Americae Septentrionalis. White,
Cochrane, London.

Therman, E. 1956. Cytotaxonomy of the tribe Polygona-
teae. Amer. J. Bot. 43: 134—142.

Wang, F. T. & T. Tang. 1978. Liliaceae. In: Inst. Bot. Acad.
Sin. (editor). Flora Reipublicae Popularis Sinicae 15:
26-40. Science Press, Beijing.

A New Species of Chirita (Gesneriaceae) from Guangxi, China

Li /lien- Yu

Laboratory of Systematic and Evolutionary botany. Institute of Botany, Chinese Academy of

Sciences, Beijing, 100093, China

Abstract. A new species from northern Guangxi.
China, Cliirilu longii Z. Y. Li, is described.

Key words: China, Chirita, Gesneriaceae.

An early-flowering plant of Chirita Buchanan-
Hamilton ex D. Don found in South China is here
recognized as a new species. Most Chinese species
in this genus bloom in spring, summer, or autumn;
only C. atropurpurea W. T. Wang blooms in winter
(February) (Wang et ah. 1998). The new species
was first collected and photographed by Long
Guang-Ri in January 1992 in the karst region of
Longan County of northern Guangxi. When he re¬
turned to this locality in February 1993 for further
exploration, he found that the habitat had been en¬
tirely destroyed due to human activities. Unfortu¬
nately, this plant has not yet been rediscovered in
the type locality or in adjacent regions (Long
Guang-Ri, pers. comm.. 1993. 1994, 1995).

Cliirilu longii Z. Y. Li, sp. nov. TYPE: China. Cu-
angxi: Longan Co., Qiaohan, Qinzhan, on cliff,
alt. 350 m, 12 Jan. 1992, Ding Guang-Ri
91004 (holotype, PE). Figure 1.

A C. tenuifolia W. T. Wang foliis minoribus, laminis
ohlanceolatis, pedunculis brevioribus 1.2— 2.3 cm longis,
calyre 8—12 mm longo, corolla majore ca. 4.3 cm longa,
stigmatis labio antico 2-fido differt.

Perennials, stemless. Rhizome ohconic, to 25 X
14 mm, internodes inconspicuous. Leaves 3 to 8,
crowded at apex of rhizome, opposite, petiolate;
blades oblaneeolate, 1.5— 2.5 X 0.4— 0.9 cm, her¬
baceous, upper surface dark green, paler beneath,
both surfaces appressed white-puberulous, base de¬
current, margin remotely undulate or undulate-cre-
nate, apex acute to obtuse; lateral veins 3 pairs,
inconspicuous; petioles 5—10 X 1-2.5 mm, velutin-
ous. Inflorescence of I or 2 cymes from the axils
of the crowded leaves, each with 1 flower; pedun¬
cles 12—23 mm, densely pubescent; bract 1, linear,
3—3.5 X 0.5 mm, both surfaces densely pubescent;
pedicels 3—4 mm, densely pubescent. Galyx 5-part-
ed from base, segments linear, unequal, 8—12 X
0.8— 1.2 mm, outside densely pubescent, inside
sparsely puberulent, margin entire, apex aeumi-

Novon 12: 492-494. 2002.

nate. Corolla lilac purple, ca. 4.3 cm, outside
sparsely puberulous, inside glabrous; tube ca. 2.7
cm, ca. I cm diam. at mouth; upper lip ca. 9 mm,

2- lobed, lobes suborbicular; lower lip ca. 16 mm,

3- lobed, each lobe oblong. Stamens 2; filaments
narrowly linear, adnate for ca. 12 mm to corolla
base, ca. 12 mm. geniculate above base, to 0.5 nun
wide, 0.3 mm wide apieally, upper part puberulent,
lower glabrous; anthers elliptic, ca. 2.5 mm, gla¬
brous. Staminodes 2. narrowly linear, adnate for ca.

I f mm to corolla base, ca. 2.5 mm, glabrous. Disc
annular, ca. 0.6 mm. margin lobed, glabrous. Pistil
ca. 23 mm, densely puberulous, ovary linear, ca. 8
X 0.8 mm; lower lobe of stigma obtrapezoid, 1.2
mm, 2-parted, each lobe oblong-lanceolate, ca. 0.9
mm. Fruit unknown.

Habitat. Growing on a limestone cliff at 350 in.
flowering occurs at least in January.

Distribution. Known only from the type locality
at Zhan Oin, Qiaohan, the karst region ol Longan
County, Guangxi, China.

Etymology. This species is named in honor of
Long Guang-Ri (1952—), a dendrologist at the f or¬
estry Bureau of Liuzhou Prefecture of Guangxi.

The genus Chirita consists of about 140 species,
distributed in subtropical and tropical Asia, rang¬
ing from northwestern India (Gujarat and Simla)
eastward to eastern China (Taiwan, Taidong) and
from central China (Gansu, Wenxian) southward to
southern Indonesia (Lessersunda Islands), at 90—
3200 m altitude (Wood, 1974; Wang et al., 1998).
There are 102 species in China, 54 in Guangxi
(Wang el ah. 1998; Fang el ah. 1999; present pa¬
per); most species discovered from the karst regions
were found at the lower elevation. Following
Clarkes subdivision (Clarke, 1883; Wood, 1974;
Wang et ah, 1990), C. longii Z. Y. Li is referred to
Chirita sect. Gibhosaccus Clarke. It is closely allied
to C. tenuifolia W. T. Wang (Wang, 1985; Wang et
ah. 1998), but differs from the latter in its smaller
and thicker leaves, oblaneeolate lamina (vs. elliptic
or broadly elliptic), shorter peduncles (1.2— 2.3 cm
vs. ca. 5 cm), longer calyx (8-12 mm vs. ca. 4.8
mm), larger corolla (ca. 4.3 cm vs. ca. 2.8 cm).

Volume 12, Number 4
2002

Li

Chirita longii (Gesneriaceae)

493

1 mm

Figure J. Chirita longii Z. Y. Li. — A. Flowering plant (with hairs enlarged). — B. Bract. — C. Opened corolla showing
stamens and staminodes. — D. Anther. — E. Calyx and pistil (with stigma enlarged at right). — F. Disc and cross section
of ovary. (Drawn from the holotype.)

494

Novon

obtrapezoid and 2-parted lower lip of stigmas (vs.
ligulate, apex rounded, never divided), and dow¬
ering in January (vs. August).

Acknowledgments. I thank Long Guang-Ri for
His assistance in the field; Wang Wen-Tsai for
checking the Latin description; Hong De-Yuan. Hu
Yu-Xi, Yang Qin-Er, and Zhu Guang-Hua for crit¬
ical comments; and Sun Ying-Bao for the drawing.
I he author is grateful lor detailed reviews of the
manuscript by Victoria C. Hollowed and Eric Roal-
son. This work was supported by the State Key Ba¬
sic Research and Development Plan (No.
G200004680] -1) and the National Natural Science
Foundation of China (No. 39870056).

Literature Cited

Clarke, C. It. 1883. Cyrtandreae. Pp. 1-303 in A. & C.
De Candolle (editors). Monograph iae Phanerogamarum,
Vol. 5(1). Sumptibus G. Masson, Parisiis.

Fang, I)., I). H. Din & Y. G. Wei. 1999. New plants of
the Gesneriaeeae from Guangxi of China (III). Acta
Phytotax. Sin. 37(6): 591-594.

Viang. Vi. T. 1985. Notulae de Gesneriaceis Sinensibus
(II). Bull. Hot. Res. 1(4): 35-75.

- . K. Y. Pan & Z. Y. Li. 1990. Gesneriaeeae. Pp.

125—581 in W. T. Wang (editor). Flora Reipublicae Po-
pularis Sinicae, Tonius 69. Science Press, Beijing.

- , - . - . A. I,. Weitzman & L. E. Skog.

1998. Gesneriaeeae. Pp. 241—101 in C. Y. Wu & P. H.
Raven (editors). Flora of China, Vol. 18. Science Press,
Beijing, and Missouri Botanical Garden, St. Louis.
Wood, I). 1974. A revision of Chirita (Gesneriaeeae).
Notes Roy. Rot. Gard. Edinburgh 33: 123-205.

New Combinations in Chinese Cotoneaster (Rosaceae)

Lu Lingti

Laboratory of Systematic and Evolutionary Biology, Institute of Botany, Chinese Academy of
Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China

Anthony R. Brack

Missouri Botanical Garden, % Harvard University Herbaria, 22 Divinity Avenue,
Cambridge, Massachusetts 02138-2094, U.S.A. brach@oeb.harvard.edu

ABSTRACT. During preparation of the account ol
Cotoneaster Medikus for the Flora of China, volume
9, it was found that some taxa required the follow¬
ing new combinations, namely Cotoneaster glauco-
phyllus Franchet var. serotinus (Hutchinson) L. I.
Lu & A. R. Brach, C. bullatus Bois \ar.floribundus
(Stapf) 1.. T. Lu & A. R. Brach, and C. gracilis
Rehder & E. H. Wilson var. difficilis (G. Klotz) L.
T. Lu.

Key words: China, Cotoneaster, Maloideae, Ro¬
saceae.

In the broad sense, the genus Cotoneaster Me-
dikus consists of about 90 species and is widely
distributed in temperate regions of northern Africa,
continental Asia, Europe, and North America
(mainly in Mexico). It is most abundant in south¬
western China, with about 58 species recorded from
t licit country which have been variously placed into
about seven (to more than 20) series. Their taxon¬
omy is complicated by hybridization and apomixis.
Further studies could help elucidate relationships
between and within species. While preparing an
account of Cotoneaster for the Flora of China, vol¬
ume 9 (Lu & Brach, in press), tin* present authors
noted that the following new combinations are nec¬
essary.

Cotoneaster glaucophyllus Franchet (1890: 222)
is a semi-evergreen shrub endemic to southwestern
China (Guangxi, Guizhou. Sichuan, and Yunnan
Provinces), with elliptic to ovate, glaucous leaves,
compact, compound, to more than 50-flowered cor¬
ymbs, spreading, white petals, and reddish yellow,
ovoid to obovoid fruit.

In addition to its typical variety, two other named
varieties occur in China: Cotoneaster glaucophyllus
var. vestitus W. W. Smith (1917: 21) and variety
meiophyllus W. W. Smith (loc. fit.). A form of this
species, f. serotinus (Hutchinson) Stapf, which was
originally described at specific rank (G. serotinus

Hutchinson), is best treated as a fourth variety; thus
a new combination is required.

Cotoneaster glaucophyllus Franchet var. sero¬
tinus (Hutchinson) L. T. Lu & A. R. Brach,
comb. nov. Basionym: Cotoneaster serotinus
Hutchinson. Bot. Mag. 146: I. 8854. 1920. Co¬
toneaster glaucophyllus Franchet f. serotinus
(Hutchinson) Stapf. Bot. Mag. 153: t. 91 (I.
1929. TYPE: plant cultivated in the garden of
Mr. G. H. Wollaston, Flaxley Cottage, Flax
Bourton, United Kingdom, 1919 (fr). grown
from seeds collected in China. Yunnan: G.
Forrest 6754 (holotype, K).

This fourth variety occurs from 1900 to 3000 m
in mountainous regions of western Yunnan Prov¬
ince. It is characterized by having leaf blades 4—6
cm. abaxially pubescent when young, gradually gla-
brescent, and inflorescences 50- to 60-flowered,
white tomentose initially, later glabrescent. I he
four varieties of Cotoneaster glaucophyllus are dis¬
tinguished as follows:

Ki a to Varieties ok Cotoneaster oiavcophyllus

la. Leaf blade 3.5-6 cm.

2a. Inflorescences to 40-ilowered, with soft yel¬
low hairs . var. glaucophyllus

2b. Inflorescences 50- to 60-flowered, white to¬
mentose initially, later glabrescent ....
. var. serotinus

lb. Leaf blade 1.5—3 cm.

3a. Leaf blade abaxially densely yellow tomen¬
tose when young, later glabrescent; inflores¬
cences to 40-flowered, larger, densely white

tomentose when young . var. vestitus

3b. Leaf blade abaxially slightly pubescent or
subglabrous; inflorescences 6- to 12-flow-
ered, smaller, with soft hairs . . var. meiophyllus

Cotoneaster bullatus Bois (in Vilmorin & Bois,
1904: 1 19) is a deciduous shrub endemic to south¬
western China (Hubei, Sichuan, Xizang, and Yun¬
nan Provinces), with conspicuously rugose, bullate

No von 12: 495-496. 2002.

496

No von

leaves, lax, 5- to 31 -flowered corymbs, erect, pink
petals, and red to dark purple, globose or ohovoid
fruit. In addition to its typical variety, another
named variety occurs in China: C. bullatus var. ma-
crophyllus Rehder & K. II. Wilson (in Sargent,
1912: 164). A form of this species, f . floribundus
(Stapf) Rehder & E. H. Wilson (originally described
as a form of C. moupinensis I’ ranchet), is best treat¬
ed as a third variety of C. bullatus ; thus a new
combination is required.

Cotoneaster hull ulus Bois var. florihuudiis

(Stapf) L. T. Lu & A. R. Brach, comb. nov.
Basionym: Cotoneaster moupinensis Franchet 1.
floribundus Stapf, Bot. Mag. 135: t. 8284.
1909. Cotoneaster bullatus f. floribundus
(Stapf) Rehder & E. H. Wilson, in Sargent, PI.
Wilson. I: 165. 1912. TYPE: plant cultivated
at the Royal Botanic Gardens, Kew, United
Kingdom, from seed collected in China. Sich¬
uan: “near Taehienlu" [Kangding], M. Vilmorin
2 1 2d (holotype, K).

Thi s second variety occurs in mountain forests
from 900 to 2100 m in western Sichuan Province.
It is characterized by having petioles 1.5(— 2.5) mm,
leaf blades ovate or obovate, less than 5 cm, and
inflorescences 4—6 cm in diameter, 15- to 31 -flow¬
ered.

The three varieties of Cotoneaster bullatus are
distinguished as follows:

Kky to Vahiktiks ok Cotoneaster hi liati s

la. Petiole 3—6 mm; inflorescences 5- to 13-flowered

. var. bullatus

Ih. Petiole 1.5— 2.5 mm; inflorescences II- to 31-
flowered.

2a. Leaf blade 5—15 X 2.5-3 cm. petiole ea. 2
mm; inflorescences 5—8 cm diam., II- or

more flowered . var. mucroph yllus

2b. Leal blade usually less than 5 cm, petiole
usually less than 2 mm; inflorescences 4—6

cm diam.. 15- or more flowered .

. var. floribundus

Cotoneaster gracilis Rehder & E. H. Wilson (in
Sargent, 1912: 167) is a deciduous shrub endemic
to central China (Gansu. Hubei, Shaanxi, and Sich¬
uan Provinces), with ovate to broadly elliptic
leaves, lax, l- to 7-flowered corymbs, erect, red
petals and red, obovoid, puberulous fruit. In addi¬
tion to its typical variety, another species, Cotone¬
aster difficilis G. Klotz, is very similar to it. but
differs mainly by its smaller leaf blades and few-
flowered inflorescences. This latter species is best

treated at varietal rank under C. gracilis ; thus a
new combination is required.

Cotoneaster gracilis Rehder & E. II. Wilson var.
difficilis (G. Klotz) L. T. Lu, comb. nov. Bas¬
ionym: Cotoneaster difficilis G. Klotz, Wiss. Z.
Eriedrich-Schiller-Univ. Jena, Math.-Natur-
wiss. Reihe 21(5-6): 1017. 1972. TYPE: Chi¬
na. Sichuan: “valley of Hsao chin ho |Xiaojin
River] near Mon kong ling. 7-10.000 ft.,” June
1908, E. II. Wilson 2169 (holotype. A; isotype,
BM).

This variety occurs from 1800 to 3000 m in
mountain regions of southeastern Gansu and west¬
ern Sichuan Provinces. Cotoneaster gracilis var. dif¬
ficilis is characterized by having leaf blades ovate-
elliptic to broadly elliptic, 7-14 X 5.5—8 mm,
abaxially tomentose initially, and inflorescences I-
to 3(to 6)-f lowered. It is distinguished from the typ¬
ical variety as follows:

key to Vahiktiks ok Cotoneaster cracii. is

la. Leaf blade ovate to oblong-ovate, 20—35 X 10-

20 mm, densely tomentose abaxially; inflores¬
cences 3- to 7-flowered . var. gracilis

lb. Leaf blade ovate-elliptic to broadly elliptic, 7—

14 X 5.5—8 mm, initially tomentose abaxially;
inflorescences 1- to 3(to 6)-flowered . . var. difficilis

Acknowledgments. We thank Nicholas J. Tur-
land (MO) for nomenclatural advice and editorial
help. We are grateful to two anonymous reviewers
for their comments.

Literature Cited

Franchet, A. 1889-1890. Plantae del avayanae. Plantes de
Chine recueillis au Yun-nan par I'Abbi* Delavay et de-
erites par A. Franchet. Paul Klincksieek, Paris.

Lu. Lingdi [Ling-ti] & Brach, A. R. In press. Cotoneaster
Medikus. In: Zhengyi Wu & P. H. Raven (editors). Flora
of China, Vol. 9. Science Press, Reijing, and Missouri
Botanical Garden Press, St. Louis.

Sargent, C. (editor). 191 1 — 1917. Plantae Wilsonianae. An
Enumeration of the Woody Plants Collected in Western
China for the Arnold Arboretum of Harvard University
During the Years 1907, 1908. and 1910 by E. II. Wil¬
son. Harvard Univ. Press. Cambridge, Massachusetts.
Smith. W. W. 191 i. Diagnoses speeierum novarum iu her-
bario Horti Regii Rotaniei Edinburgensis cognitarum
(species asiaticae). Notes Roy. Bot. Card. Edinburgh
10: 1-78.

Vilmorin. M. de & I). G. J. M. Bois. 1904. Frulicetuin
Vilmorinianum. Catalogus primarius. Catalogue des ar-
bustes existant en 1904 dans la collection de VI. Mau¬
rice LeviS | ue de Vilmorin avee la description d'espLees
nouvelles et d introduction reeenle. Librairie Agricole,
Paris.

Stylosanthes seabrana (Leguminosae: Papilionoideae), a New
Species from Bahia, Brazil

Brigitte L Maass

Institute for Crop and Animal Production in the Tropics, Georg- August-University, D-3707 t

Gottingen, Germany, bmaass@gwdg.de

Leendert 7 Mannetje

Herbarium Vadense (WAG), Wageningen University, The Netherlands, ltmannet@bos.nl

Abstract. During agronomic evaluation of intro¬
duced accessions of Stylosanthes Swartz in Colom¬
bia anil Queensland, Australia, a genotype differing
in morphology, chromosome number, and genetic
markers was discovered. The new species Stylosan¬
thes seabrana is closely related to S. scabra Vogel
and has been collected in Bahia, Brazil. Two cul-
tivars, ‘Primar’ and ‘Unica,' of this species now
newly recognized were released in Queensland,
Australia, in 1996 for use as pasture legumes.

Key words: Bahia, diploid, genetic resources,
germplasm, Stylosanthes seabrana , tropical pasture
legume.

Within a large germplasm collection of Stylosan¬
thes scabra Vogel at the Centro Internacional de
Agricultura Tropical (Cl AT), Colombia, Maass
(1989) found plants that were morphologically, ag-
ronomieally, and biochemically very' different from
S. scabra , while not belonging to any other known
species within this genus. They were referred to as
“cf. scabra-type” (Maass, 1989) and were also in¬
troduced to Queensland, Australia (Jansen & Kdve.
1996), where they were called "aff. scabra’ and
became commonly known as “Caatinga stylo. Two
cultivars, ’Primar' and ’Unica," were released in
Australia in 1996 for use as pasture legumes
(Anonymous, 1996).

In the morphological study of Jansen and Edve
(1996), the diploid accessions of "aff. scabra” were
compared to S. hamata Taubert ‘Verano’ and S. sca¬
bra ‘Seca.’ which are both tetraploid (Oram. 1990;
Stace & Cameron, 1987). However, these acces¬
sions from an undescribed taxon could morpholog¬
ically be distinguished from S. hamata by their
bristly leaflets (vs. non-bristly in S. hamata) and
from S. scabra by their narrower and glabrous leaf¬
lets (vs. elliptical to obovate and pubescent in S.
scabra ) except for prominent viscid bristles on the
lower surface and margins; prominent bristles on
stem nodes and stipules, stipule teeth with long lat¬

eral bristles in combination with an absent or very'
short terminal bristle, bristles often on part of the
internodes and on the inflorescence (vs. overall pu¬
bescence with occasional appearance of short, vis¬
cid bristles in S. scabra, but not as prominent); rel¬
atively longer pods and longer stylar remnants
(beaks) (vs. shorter pods and beaks in S. scabra )
(Jansen & Edve, 1996).

Additional morphological, rhizobial, cytological,
and molecular evidence supports the conclusion
that this is a separate, as yet unnamed, diploid (2n
= 20) (Liu, 1997; Liu & Musial, 1997) species
(Table 1), which belongs to Stylosanthes sect. Sty-
posanthes. From genetic evidence, Liu et al. (1996)
and Liu and Musial (1997) postulated that this un¬
known species and S. viscosa Swartz might be the
putative diploid progenitors of the allotetraploid S.
scabra ('2n = 40). Unlike S. scabra , the unnamed
species demonstrated a highly specific requirement
for effective nitrogen-fixing strains of Bradyrhizo-
bium (Date et ah, 1996).

KhA TO Sm.OSA.\THES seabrana, S. hamata, and S. scabra

la. Beak equal to or exceeding the upper article,

leaflets without bristles . S. hamata

1 b. Beak shorter than the upper article, leaflets with
bristles.

2a. Leaflets narrowly elliptical, glabrous except
for long bristles on the margins and midrib
and prominently raised veins on the lower

surface . S. seabrana

2b. Leaflets elliptical to obovate, pubescent with
bristles at least underneath or on the mar¬
gins without prominently raised veins on the
lower surface . .S’, scabra

Stylosanthes seabrana B. L. Maass & L. ‘t Man¬
netje, sp. nov. TYPE: Brazil. Bahia; State Road
BA052 from Irece 80 km to Xique-Xique,
10°56'S, 42°29'W, 29 June 1983, L Coradin,
R. Baker, F. B. de Souza, R. M. Harley A' S.
Linington LC 6261 (holotype, CEN; isotypes,
k. M, PA MG, RB, WAG).

Nov on 12: 497-500. 2002.

lable 1. Herbarium specimens, germplasm accessions, and ploidy status of Stylosanthes seabrana.

498

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Volume 12, Number 4
2002

Maass & ‘t Mannetje
Stylosanthes seabrana

499

S. scabrae affinis sed diploidea vice tetraploidea, rudi-
mento axis multo longiore (> 6 mm), folic*] is angustioribus
et glabrescentibus infra et ad marginen setis longioribus
vulgo viscidis munitis et infra venis manifest? prominen-
tibus albidis munitis, dentibus stipularum setis lateralibus
longis munitis, seta terminali nulla vel brevissima differt.

Perennial, woody at base: stems pilose on one
side, except at base, viscid bristles (> 1 nun) near
the nodes; stipules pilose, very bristly, long lateral
bristles present on st i pule teeth, but rarely with a
terminal bristle, if so this very short. Leaflets lan¬
ceolate to elliptic-acute, upper surface glabrous,
lower surface glabrous, except for long often viscid
bristles on the midrib below and on the margins,
veins raised, prominent below, whitish. Spikes cap¬
itate, small with several papilionaceous flowers,
with obovoid sulphur-yellow standard. Each flower
surrounded by a trifoliolate outer bract and two in¬
ner bracteoles, ciliate at the apex. Outer bract bris¬
tly, pilose on the margins. 3 pairs of veins. Axis
rudiment reaching a height halfway up the upper
article (7—8 mm long), pilose. Loment with 2 arti¬
cles; the upper article appressed pilose to woolly,
sometimes glabrous or slightly hairy on the inside
of the beak (stylar remnant), with a clear central
vein and weak lateral veins. Beak uncinate, pilose
to woolly, half the length of the upper article (ca.
2—3 mm from base to top of the bend). I lie lower
article densely pilose. Seed cream-colored. Chro¬
mosome number 2 n = 20.

Etymology. Specimens and several germplasm
accessions from the region around the town of Sea-
bra. in the Brazilian state of Bahia were among the
earliest collections of the species.

Distribution. Endemic to Bahia, Brazil, be¬
tween 9— 13°S and 40— 46°W.

Ecology. Stylosanthes seabrana occurs in areas
with low rainfall, long dry seasons, mainly on me¬
dium to heavy textured red or yellow soils with Caa-
tinga vegetation (Jones et ah. 1996). It is often
found growing sympatrically with S. scabra. The re¬
leased cultivars in Queensland, Australia, are frost
and anthracnose tolerant.

Paratypes. BRAZIL. Bahia: Muu. Barreiras, 28 Sep.
1978, L. Coradin et al. LC 1172 (CEN, PAMG); Mun.
Barreiras. 28 Sep. 1978, L. Coradin et al. LC 5186
(ILCA); Mun. Seabra, 29 Sep. 1978, L. Coradin et al. LC
5208 (ILCA); Mun. Seabra, 29 Sep. 1978, L. Coradin et
al. LC 5221 (ILCA); Mun. Seabra. 4 Oct. 1978, L Coradin
et al. LC P234 (CEN, PAMG); Rd. Irece-Xique Xique 3
kin NW of Irece, 18 feb. 1981, Bastos /5 (CEPEC, k);
Rd. Ilh6us— Itabuna km 22, 1 July 1981, Huge & Brito
1017 (CEPEC, K); Mun. Boninal 52 km S from Rd. BR
242 to Piata, 20 Aug. 1981, L. Coradin et al. LC 4.351
(CEN, HUEFS, k. NY); Mun. Palmeiras, 22 Aug. 1981,

L. Coradin et al. LC 4447 (CEN, ILCA); Rd. Jacobina—
Lage do Batata km 15, 28 June 1983, L Coradin et al.
LC 6171 (CEN, k, PAMG); Palmeiras ea. 250 km on Rd.
BR 242, 19 Mar. 1990. de Carvalho & Saunders 2962
(CEPEC. K).

Germplasm accessions. The largest part of the
herbarium specimens listed here was collected dur¬
ing different germplasm collecting missions direct¬
ed by L. Coradin from the Brazilian Centro Na-
cional de Recursos Geneticos e Biotecnologia
(CENARGEN) in collaboration with scientists from
Cl AT (Coradin & Schultze-Kraft, 1990) or the Roy¬
al Botanic Gardens, Kew. For purposes of conser¬
vation. research, and breeding, seed of most of the
collections is consequently stored in genebanks of
the respective institutions as well as in other rele¬
vant genetic resource collections, such as the Com¬
monwealth Scientific and Industrial Research Or¬
ganisation (CS1RO) in Australia or the International
Livestock Research Institute (1LRI) in Ethiopia (Ia-
ble 1). A number of additional germplasm acces¬
sions that belong to the same diploid species were
mentioned by Jansen and Edye (1996) and Liu
(1997).

Acknowledgments. We are grateful to G. P. Lew¬
is of the Royal Botanic Gardens, Kew. for critical
comments on the manuscript; to R. Lemmens of
Wageningen Herbarium for the Latin species de¬
scription; T. Barbosa Cavalcanti (CENARGEN), A.

M. Torres (CIAT), and J. Bone (MSB-Kew) for pro¬
viding comprehensive passport information; the
herbaria CEN. K, M. and WAG for making speci¬
mens available. BLM washes to acknowledge the
support of ILRI for carrying out part of this re¬
search.

Literature Cited

Anonymous. 1996. Caatinga stvlo Stylosanthes — “Primar
and “Unica.” PI. Var. J. 9(3): 19-20.

Coradin, L. & R. Schultze-Kraft. 1990. Germplasm col¬
lection of tropical pasture legumes in Brazil. Trop.
Agric. (Trinidad) 67; 98-100.

Date, R. A., L. A. Edye & C. J. Liu. 1996. Stylosanthes
sp. aff. scabra — A potential new forage plant for north¬
ern Australia. Trop. Grasslands 30: 133.

Jansen. P. I. & L. A. Edye. 1996. Variation within Stylo¬
santhes sp. aff. S. scabra and comparison with its closest
allies, S. scabra and S. hamata. Austral. .1. Agric. Res.
47: 985-996.

Jones, P. G„ R. Rebgetz, B. L. Maass & P. C. Kerridge.
1996. Genetic diversity in Stylosanthes species: A GIS
mapping approach. Pp. 186—189 in R. C. Pereira & L.
C. B. Nasser (editors). Biodiversidade e produyao sus-
tentavel de alimentos e fibras nos Cerrados: Proceed¬
ings of the First International Symposium on Tropical
Savannas, 24-29 March 1996. Empresa Brasileira de
Pesquisa Agropeeuaria (EMBRAPA), Centro de Pes-

500

Novon

quisu Agropecudria dos Cerrados (CPAC), Planaltina,
DF, Brazil.

I -iu, C. J. 1997. Verification of and genetic variation be¬
tween Stylosanthes sp. aff. S. scabra accessions. In:
Proc. Will International Grassland Congress, 8-19
June 1997, Winnipeg and Saskatoon, Canada, vol.
1(12): 7-8.

- & J. M. Musial. 1997. Stylosanthe s sp. all. scabra:

A diploid putative progenitor of Stylosanthes scabra. PI.
Syst. Evol. 208: 99-105.

- - & 1. W. Smith. 1996. Evidence for a low

level of genomic specificity of sequence-tagged-sites in
Stylosanthes. Theor. Appl. Genet. 93: 864-868.

Maass. B. L. 1989. Die tropische Weideleguminose Sty¬
losanthes scabra Vog. — Variabilitat, Leistungsstand und
Mogliehkeiten ziichterischer Verbesserung. Landbaufor-
schung Volkenrode, Sonderheft 97: 1-189.

Oram, R. N. 1990. Pp. 256—259 in Register of Australian
Herbage Plant Cultivars. 3rd ed. CSIRO, Canberra,
Australia.

Stace. H. M. & I). F. Cameron. 1987. Cytogenetic review
of taxa in Stylosanthes hamata sensu lato. Trop. Grass¬
lands 21: 182-188.

New Combinations and Synonyms in the Moss
Family Diphysciaceae

Zacharia L. K. Magombo

National Herbarium and Botanic Gardens of Malawi, P.O. Box 528, Zomba, Malawi

zlkmagombo@hotmail.com

Abstract. Phylogenetic analyses of sequence
data of the chloroplast protein coding genes rbc b
and r/w4, and the chloroplast encoded trn L (UAA)
intron suggest that morphological characters tradi¬
tionally used in delimiting the genera Diphyscium
Mohr, Muscoflorschuetzia Crosby, and Theriotia
Cardot in the moss family Diphysciaceae are ho¬
moplastic; Diphyscium is paraphyletic, and Theri¬
otia and Muscoflorschuetzia are nested in Diphys¬
cium. Therefore, only Diphyscium Mohr should be
recognized in Diphysciaceae, and Muscoflorschuet¬
zia Crosby and Theriotia Cardot should be made
synonyms of Diphyscium. Consequently, three com¬
binations are newly made: Diphyscium kashmirense
(H. Robinson) Magombo, Diphyscium lorifolium
(Cardot) Magombo, and Diphyscium pilmaiquen
(Crosby) Magombo.

Key words: Diphysciaceae, Diphyscium, moss,
Muscoflorsch uetzia, Theriotia.

The Diphysciaceae are a small family of mosses
distinguished by the unique combination of short se¬
tae. immersed capsules, collared axillary hairs, dis¬
tinctly differentiated perichaetial leaves that have
crenulate, dissected, laciniate or ciliate margins at
apex, and arthrodontous peristomes that have pleat¬
ed endostomes and exostomes that are rudimentary
or lacking. The Diphysciaceae grow on rocks (lime¬
stone or sandstone) or in rock crevices, on soil,
sometimes on rotten wood, and rarely on tree trunks.
They are found in forests, along trails or footpaths,
along streams and riverbanks, and sometimes close
to waterfalls, on wet or frequently watered substrates,
in fully or partially shaded places. Occasionally, the
Diphysciaceae occur fully submerged in water. The
Diphysciaceae grow at an altitudinal range from
close to sea level to as high as 3400 m.

Most species in the Diphysciaceae are restricted
in their distributions, although a few are wide¬
spread and exhibit disjunct distribution patterns
(Robinson, 1965; Allen, 1996; Crosby, 1977; De-
guchi, 1975, 1984a, 1984b: Norris, 1981; Noguchi
& Iwatsuki, 1987; Deguchi et ah, 1997; Tan, 1998).
Twelve species occur in the Old World, and seven

of these are restricted to Asia (Diphyscium fasci-
culatum Mitten. D. satoi Tuzibe, D. perminutum
Takaki, D. suzukii Z. Iwatsuki, D. fulvifolium Mil-
ten, Theriotia lorifolia Cardot, and T. kashmirensis
11. Robinson). The Old and the New World share
five species (D. foliosum (Hedwig) Mohr, D. mu-
cronifolium Mitten. D. longifolium Griffith. D. pocsii
(Bizot) R. H. Zander, and D. chiapense D. H. Nor¬
ris). Three species, D. fendleri Muller Halle, D.
domingense (Bridel) W. R. Buck & Steere, and
Muscoflorschuetzia pilmaiquen (Crosby) Crosby, are
restricted to the New World. Diphyscium foliosum
is the most widely distributed species of all Di¬
physciaceae. It is known from Central and North
America, throughout Europe (including the Arctic
regions), and Asia.

The Diphysciaceae traditionally consist of three
genera: Diphyscium Mohr (12—15 species), 7 heri-
otia Cardot (2 species), and Muscoflorschuetzia
Crosby (1 species). The first taxon now placed in
Diphyscium was originally described as Buxbaumia
foliosa Hedwig. Hedwig (1801) used the genus
Buxbaumia to accommodate two species, Buxbau¬
mia aphylla Hedwig and Buxbaumia foliosa Hed¬
wig, which he characterized as dioieous species
with terminal inflorescences and double peristomes
that have truncate exostomes and plicate endosto¬
mes. Based on differences in gametophyte mor¬
phology, Mohr (1803) established the genus Di¬
physcium, with a single species Diphyscium
foliosum (Hedwig) Mohr. Schwaegrichen (1830) re¬
turned D. foliosum to Buxbaumia, choosing not to
recognize Diphyscium Mohr because of its similar
peristome structure. However, Fleischer (1919)
agreed with Mohr and established the family Di¬
physciaceae for D. foliosum.

The genus Theriotia was erected by Cardot
(1904) based on a sterile collection ( Faurie 136)
from Ouen-San (Wonsan) in North Korea, which he
named Theriotia lorifolia Cardot. Lamina thickness
has traditionally been the main feature for distin¬
guishing Theriotia (3 to 14 cells thick) from Di¬
physcium (1 to 2 cells thick). Cardot (1904) origi-
nally placed Theriotia in the Syrrhopodontaceae,

Novon 12; 501-503. 2002.

502

Novon

because its leal form resembles that of Syrrhopodon
Schwaegriehen subg. Calymperidium Dozy & Molk-
enboer. I'he multistratose leaf structure of Theriotia
was also compared to that of Exodictyon Cardot
(Leucobryaceae), a genus that Cardot (1904) con¬
sidered intermediate between Syrrhopodontaceae
and Leucobryaceae. Brotherus (1925) was the hrst
to place T. lorifolia in the Diphysciaceae, because
its peristome is similar to that of Diphyscium. A
second species was added to Theriotia when Rob¬
inson (1965) described T. kashmirensis H. Robin¬
son with a leaf structure similar to that of T. lori¬
folia.

Crosby (1977, 1978) added a third genus to the
Diphysciaceae when he described Muscoflorschuet-
zia. He placed this genus in the Diphysciaceae,
because of its short setae, collared axillary hairs,
and inner perichaetial leaves that have ciliate mar¬
gins at the apex. Muscoflorschuetzia differs from
other members of the Diphysciaceae in its lack of
a peristome, long and narrow capsule shape, and
consistently unistratose lamina (Crosby, 1977,
1978).

Variation in morphological characters, particu¬
larly plant size, leaf morphology, lamina cell struc¬
ture, capsule morphology, exothecial cell morphol¬
ogy, stomata, peristome, annulus, and sexuality has
led to debate on character evolution and has af¬
fected ideas on how genera and species might be
related in the Diphysciaceae (Crosby, 1977, 1978;
Deguchi, 1975. 1984b: Allen. 1996; Deguchi et al.,
1997; Norris. 1981; Robinson, 1965). On the basis
of molecular data the Diphysciaceae are considered
monophyletic (Goffinet et al., 2001; Magombo.
2002) and are sister to haplolepideous and diplo-
lepideous mosses (Beckert et al., 1999, 2001; New¬
ton et al.. 2000: Goffinet et al., 2001; Magombo.
2002). However, phylogenetic analyses of sequence
data of the chloroplast protein coding genes rhc\.
and r/«4, and the chloroplast encoded trnL (UAA)
intron (see Magombo. 2002) suggest that the mor¬
phological characters traditionally used in delim¬
iting Diphyscium. Muscoflorschuetzia. and Theriotia
are homoplastic; recognition of Theriotia and Mus¬
coflorschuetzia makes Diphyscium paraphyletie
since both genera are nested in Diphyscium. There-
fore, only Diphyscium should be recognized in the
Diphysciaceae; Theriotia and Muscoflorschuetzia
are synonyms of Diphyscium. Consequently, three
combinations are newly made in Diphyscium. A de¬
tailed taxonomic account of the Diphysciaceae is
presented in the forthcoming revision of the family
in the Journal of the Hattori Botanical Laboratory.

Diphyscium Mohr. Observ. Bot. 34. 1803. TYPE:

Buxbaumia foliosa Hedwig (= Diphyscium fo-
liosum (Hedwig) Mohr).

Theriotia Cardot, Beih. Bot. Centralbl. 17: 8. 1904. Syn.
nov. TYPE: Theriotia lorifolia Cardot.

Muscoflorschuetzia Crosby, Bryologist 81: 338. 1978. Syn.
nov. Replacement name for Florschuetzia Crosby,
horn, illeg., non Florschuetzia Hopping & Muller.
TV PE: Florschuetzia pilmaiquen Crosby (= Muscof¬
lorschuetzia pilmaiquen (Crosby) Crosby).

J, Diphyscium kashmirense (H. Robinson) Ma¬
gombo, comb. nov. Basionym: Theriotia kash¬
mirensis H. Robinson. Bryologist 68: 314.
1965. TYPE: [Pakistan] Kashmir. Karakorum
Range, upper Hushe Valley, Atosar Valley, 17
July 1955, G. L. Webster & E. Nasir 6173a
(holotype, US).

Robinson (1965) used the name Theriotia kash¬
mirensis when he first described this species be¬
cause its leaf structure (3 to 14 cells thick) is sim¬
ilar to that of Theriotia lorifolia. The two taxa have
indeed been considered closely related and have
traditionally been separated from members of Di¬
physcium, which have lamina of one or two cells
thick. However, Theriotia kashmirensis is similar to
members of Diphyscium in peristome structure, col¬
lared axillary hairs, and inner perichaetial leaves
with ciliate margins. Furthermore, phylogenetic
analysis of molecular evidence (Magombo. 2002)
shows Theriotia kashmirensis nested in Diphyscium.
Therefore, a new combination is made.

2. Diphyscium lorifoliuui (Cardot) Magombo,
comb. nov. Basionym: Theriotia lorifolia Car-
dot, Beih. Bot. Centralbl. 17: 8. 1904. TYPE:
Coree [North Korea]. Ouen-San, October 1901,
B. 11. ,/. Faurie 136 (holotype. PC).

Like D. kashmirense , I). lorifolium is different
from other Diphysciaceae because of the lamina
structure (3 to 14 cells thick). However, it is similar
to other Diphyscium species in its peristome struc¬
ture, collared axillary hairs, as well as inner peri¬
chaetial leaves with ciliate margins. Furthermore,
placement of this species in a separate genus, Ther¬
iotia. is not supported by molecular evidence (Ma¬
gombo. 2002). Consequently, a new combination is
made. Both I). kashmirense and 1). lorifolium occur
in the Old World (Asia).

Volume 12, Number 4
2002

Magombo

Moss Family Diphysciaceae

503

3. Diphyscium pilmaiquen (Crosby) Magombo,
comb. nov. Basionym: Florschuetzia pilmai¬
quen Crosby, Bryologist 80: 149. 1977. Mus-
coflorschuetzia pilmaiquen (Crosby) Crosby,
Bryologist 81. 338. 1978. TYPE: Chile. Prov¬
ince Valdivia/Prov. Osorno: Forest Reserve at
Planta Hvdroelectrica Pilmaiquen. along Rio
Pilmaiquen. 29 Jan. 1976. M. R. Crosby 12235
(holotvpe, MO).

Diphyscium pilmaiquen is the most distinctive
species in Diphysciaceae because of its lack of
peristome, lamina consistently unistratose, and a
long and gradually narrow capsule. Its placement
in Diphyscium , and therefore the need lor a new
combination, is supported by collared axillary
hairs, immersed capsule, perichaetial leaves with
ciliate margins, and molecular evidence (see Cros¬
by, 1977. 1978: Magombo, 2002). Diphyscium pil¬
maiquen is rare, known only from the type locality
in southern Chile.

Acknowledgments. I am grateful to R. E. Ma-
gill, M. R. Crosby, and B. Allen for making valu¬
able comments on this paper. 1 thank W. Buck and
V. Hollowed for reviewing the manuscript. 1 am also
grateful to the curators at PC and US for loans of
relevant tvpe specimens. This work was made pos¬
sible with financial support from the National Sci¬
ence Foundation (NSF) PEET grant DEB-9522034
to the Missouri Botanical Garden, International
Center for Tropical Ecology (ICTE) of the Univer-
sitv of Missouri St. Louis (UMSL), Des Lee Labo¬
ratory of Molecular Systematics (UMSL), NSF grant
DEB-9870231, and National Geographic Society
grant no. 6733—00.

Literature Cited

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aceae) an African species new to Honduras. Nova Hed-
wigia 62: 371—375.

Beckert. S., S. Steihauser, H. Muhle & V. Knoop. 1999.
A molecular phylogeny of bryophytes based on nucle¬
otide sequences of the mitochondrial nad5 gene. PL
Svst. Evol. 218: 179-192.

- . H. Muhle. I). Pruchner & V. Knoop. 2001. The

mitochondrial nad2 gene as a novel marker locus for
phylogenetic analysis of early land plants: A compara¬

tive analysis in mosses. Molec. Phylogenet. Evol. 18:

1 17-126.’

Brotherus, V. A. 1925. Musci. In Engler. Nat. Pflanzen-
fam. 2, 10: 1—542.

Cardot, J. 1904. Premiere contribution a la flore brvolo-
gique de la Cor6e. Beih. Bot. Centralbl. 17: 1—44.

Crosby, M. R. 1977. Florschuetzia, a new genus of Bux-
baumiaceae (Musci) from southern Chile. Bryologist 80:
149-152.

- . 1978. Muscoflorschuetzia (Musci), a new name for

Florschuetzia Crosby. Bryologist 81: 338.

Deguchi, H. 1975. Male inflorescence of Theriotia lorifol-
ia and its Diphyscium type paraphyses. Proc. Bryol.
Soc. Japan I: 120-122.

- . 1984a. Study on Theriotia kashmirensis (Diphys¬
ciaceae, Musci). Bull. Natl. Sci. Mus., Tokyo, B 10:
143-152.

- . 1984b. Diphyscium unipapillosum, sp. nov. (I)i-

physciaceae, Musci) from Japan. J. Jap. Bot. 59: 97—
103.

- . J. Ueno & T. Yamaguchi. 1997. Taxonomic notes

on Diphyscium species with unipapillose leaf cells. J.
Hattori Bot. Lab. 82: 99—104.

Fleischer, M. 1919. Natiirliches System der Bryales. Pp.
89-90 in A. Engler & E. Gilg, Syllabus der Pflanzen-
familien. Ed. 7. Gebriider Borntraeger. Berlin.

Goffinet, B., J. C. Cox, A. J. Shaw & T. A. Hedderson
2001. The Bryophyta (Mosses): Systematic and evolu¬
tionary inferences from rps4 gene (cpDNA) phylogeny.
Ann. Bot. 87: 191-208.

Hedwig, J. 1801. Species Muscorum Erondosorum. Lip-
siae.

Magombo, Z. L. K. 2002. The phylogeny of basal peris-
tomate mosses: Evidence from cpDNA, and implica¬
tions for peristome evolution. Syst. Bot. (in press).

Mohr, D. M. H. 1803. Observationes Botanicae, pp. 34-
35, Kiliae.

Newton, A. E., C. J. Cox, J. G. Duckett. J. A. Wheeler,
B. Goffinet, A. J. Hedderson & B. D. Mishler. 20(H).
Evolution of the major moss lineages: Phylogenetic
analysis based on multiple gene sequences and mor¬
phology. Bryologist 103: 187-211.

Noguchi, A. & Z. Iwatsuki. 1987. Illustrated Moss Flora
of Japan, Part I. Hattori Botanical Laboratory, Nichin-
an.

Norris, D. H. 1981. Diphyscium chiapense, a new species
from southern Mexico. Bryologist 84: 375—378.

Robinson, 11. 1965. A small collection of bryophytes from
Kashmir. Bryologist 68: 313-320.

Schwaegrichen, E. 1830. Species Muscorum Frondosorum
Part 2. Pp. 1-108 in C. L. Wildenow (editor). Species
Plantarum Vol. 5. Berolini Impensis G. C. Nauck, Ber¬
lin.

Tan, B. 1998. Noteworthy disjunct patterns of Malesian
mosses. Pp. 235-241 in R. Hall & J. D. Holloway (ed¬
itors), Biogeography and Biological Evolution of SE
Asia. Backhuys Publishers, Leiden.

Isoetes toximontana (Isoetaceae), a New Quillwort with Green
Megaspores from the Northern Cape of South Africa

Lytton John Musselman

Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23529-0266,

U.S.A. lmusselm@odu.edu

,/. R. Roux

Compton Herbarium, National Botanical Institute, Private Bag X7, Claremont 7735,
South Africa. Roux@nbict.nbi.ac.za

Abstract. Isoetes toximontana is only the second
quillwort reported from the Northern Cape region
in South Africa. It occurs in shallow water on the
Oifberg. Megaspores are uniformly tubereulate on
the proximal surface; their distal surfaces may have
occasional rugi along with bullae. They are olive
green when dry. Microspores are brown and acu¬
leate. Preliminary field studies suggest that an un¬
expected diversity of quillworts is found in this re¬
gion.

hey words: Gifberg, Isoetaceae, Isoetes, North¬
ern Cape Province, Quillwort, South Africa.

Burrows (1990) recorded six species of Isoetes L.
from South Africa, with only two from the Cape
region (Cape Agulhus to the Namibian border): /.
capensis Duthie and /. stellenbossiensis Duthie.

I heir presence as the only quillworts in the flora is
confirmed in a recent comprehensive treatment of
the ferns and fern allies of the Cape (Roux, 2000).
Our report here of a new species is the first quill-
wort documented across this large geographical
area.

Isoetes toximontana L. J. Musselman & J. R
Roux, sp. nov. TYPE: South Africa. Northern
Cape Province: Gifberg, shallow water along
Gifberg Road, I km S of Oubergpad,
31°47'09"S, 18°45'7I"E, ca. 22 km SE of Van-
rhynsdorp, 565 m. 20 Sep. 2001, Lytton ./.
Musselman & Elizabeth R. Musselman 2001-
35 (holotype, NBG; isotype, MO). Figures 1,2.

Planta amphibia, emergens. Excaudex trigonus et rad-
icibus simplicibus. Folia usque ad 10. obscure viridia,
rigida. spiralia, usque ad 42 mm longa, apicibus obtusa.
in medio longitudinis ca. 0.5 mm lata, in sectione trans-
versali elliptica, cellulis epidermalibus omnibus proces-
sibus tubereulatis ornatis, prope basim folii tantum, cav-
ernulis aeriis quatuor etiam seriebus stomatum ad as
parallelis praeditis, littis peripheralibus absentibus. Me-
gasporophylla microsporophyllaque in eadem planta por-

Novon 12: 504-507. 2002.

lata. Velum absente; parietes sporangii pellucidos. Megas-
pora<‘ paucae (usque ad .30 per sporangium), olivaeeae,
275-320 gin in diam., in parietibus proximalibus distal-
ibusque, aeipie tuberculatae, tuberculis altitudine varian-
tilms, cingulo angusto sub crista aequatoria posito. Mi-
crosporae brunneae usque ad 25 gm longae, aculeatae
stelis remotis.

Plants amphibious, emergent. Rootstock three-
sided. Roots simple. Leaves 3 to 10 per plant, dull
green, stiff, spirally arranged, up to 42 mm long,
obtuse, ca. 0.5 mm wide at mid length, elliptic in
cross section. Leaf epidermal cells with tubereulate
outgrowths, with four air chambers near the base,
but these absent toward the leaf apex, with four
rows of stomata parallel to the air chambers, pe¬
ripheral strands lacking. Scales present. Megaspo-
rophylls and microsporophylls on the same plant.
Velum absent. Sporangium wall clear. Megaspores
few (up to 36 per sporangium), gray-green in color,
275—320 gun diam., uniformly tubereulate on the
proximal surface, the distal surface with occasional
rugi. Tubereules of vary ing heights, with a narrow
girdle below the equatorial flange. Microspores
brown en masse, to 25 gun long, aculeate with re¬
mote stelae. The name is derived from Gifberg, Af¬
rikaans for “poison mountain,” where the tvpe was
collected.

Spore color, ornamentation, and size. Isoetes
toximontana has green megaspores, a character first
reported in the genus by Duthie (1929) for /. stel¬
lenbossiensis (Fig. I B, I)). It can be distinguished
from other Southern African quillworts by mega¬
spore ornamentation and leaf features. Isoetes stel¬
lenbossiensis has reticulate megaspore ornamenta¬
tion; /. capensis megaspores are boldly rugate on
the distal surface. (Terminology for spore ornamen¬
tation follows Lellinger & Taylor, 1997.) Based on
the figures in Burrows (1990), there are four quill¬
worts in southern Africa with tubereulate mega-

Volume 12, Number 4
2002

Musselman & Roux

Isoetes toximontana from South Africa

505

Figure 1A-E. — A. Roadside ditch with shrubby restio. Type location. — B. Megaspores. Brown microspores are

evident on the surface. — C. I socles toximontana. Note the size of the plants. — D. Mixture of megaspores of I. toxi-
montana (green) and Musselman A' Musselman 21)1)1 -.13 (white). — I’.. Representative plants of I. toximontana. 1 he coin
is ca. 2 cm in diameter.

spores: /. giessi Launert, /. transvaalensis Jermy &
Schelpe, /. welwitschii A. Braun ex Kuhn, and /.
alstonii Heed & Verdcourt. Of these, only /. giessi
and /. welwitschii lack a velum. Isoetes welwitschii
is more tropical in its distribution and in South
Africa is known from only a few collections in the
eastern part of the country (Burrows, 1990). Quill-
worts of Namibia are poorly understood, including
the taxon known as /. giessi from central Namibia,
which does have tuberculate megaspores. However,
ornamentation is sparse and megaspores are white.
The size of megaspores and microspores of /. toxi¬
montana is similar to that of /. stellenhossiensis and
/. capensis, both diploids (B. I). Bray, unpublished),
suggesting that /. toximontana is also diploid. Mi¬

crospores are brown (Fig. IB) and have aculeate
macro ornamentation (Fig. 2C). Megasporophylls
and microsporophylls were found in mid September
on the same plant.

Leaves. Unlike /. stellenhossiensis, the Gifberg
(juillwort has stomata in rows on the adaxial surface
(Fig. 2B, F, G). Feaf epidermal cells have a dis¬
tinctive ornamentation consisting of knobby out¬
growths (Fdg. 2F. G). No peripheral strands (collen-
chyma-like cells) were found (Fig. 2D).

Rootstock and roots. The rootstock is three-sid¬
ed (Fig. 2H). No branched roots, characteristic of
most Isoetes species, were evident in /. toximon¬
tana.

506

Novon

Figure 2A-G. fsoetes toximontana. — A. SEM of megaspores. Scale = 200 gm. — B. SEM of leaf tip. Note the rows
of stomata on the adaxial side. Scale = 100 gm. — C. SEM of microspores. Scale = 10 gm. — I). SEM of cross section
of leaf at midpoint. Arrow indicates stomatal apparatus. Scale = 200 gm. — E. Megaspores of Mussel man & Musselman
2001-33 with rugulate ornamentation. — E. Cross section of leaf of Musselman Musselman 2001-33. Note the four
air chambers. — G. Iwo stomata of /. toximontana. — 1 1. Cross sections of rootstocks of /. toximontana.

Volume 12, Number 4
2002

Musselman & Roux

Isoetes toximontana from South Africa

507

Habitat. This diminutive species (Fig. 1C, E)
was found only as a submergent in a temporary
pond dominated by a semi-shrubby restio (Restion-
aceae) (Fig. 1A). Our preliminary fieldwork in the
Gifberg, the southwestern-most escarpment ol
sandstone of the Cape System, indicates that sev¬
eral taxa of quillworts may be present. One of these
quillworts has white megaspores with rugate orna¬
mentation (Fig. ID. 2F). At least one putative hy¬
brid was found, distinguished by aborted spores,
another indication of the diversity of Isoetes taxa
previously unknown in this part ol southern Africa.

Preliminary molecular work using ITS sequences
indicates that I. toximontana is a distinct species
with nine unique substitutions located in a clade
with /. capensis (W. C. Taylor, pers. comm.).

Acknowledgments. We thank Denise Wadsworth
for preparing materials for scanning electron mi¬

croscopy and Alan Savitzky for light microscopy.
Discussions with Carl Taylor were helpful and en¬
couraging. We thank him for sharing preliminary
data from his phylogenetic work on South African
quillworts. Research was supported in part by the
Mary Payne Hogan fund.

Literature Cited

Burrows, J. E. 1990. Southern African Ferns and Fern
Allies. Frandsen Publishers, Sandton.

Duthie, A. V. 1929. The species of Isoetes found in the
Union of South Africa. Trans. Roy. Soc. S. Afr. 17: 321-
330.

Lellinger, l). B. & W. C. Taylor. 1997. A classification of
spore ornamentation in the Pteridophyta. Pp. 33-42 in
B. J. Johns (editor), Holttum Memorial Volume. Royal
Botanic Gardens, Kew.

Roux, J. P. 2000. Pteridophytes. Ferns and Fern Allies.
Pp. 37-50 in P. Goldblatt & J. Manning, Cape Plants.
A Conspectus of the Cape Flora of South Africa. Stre-
litzia 9: 1—743.

Seven New Speeies and One New Combination in Carex
(Cyperaeeae) from North America

Robert F. C. Naczi

Claude E. Phillips Herbarium, Department of Agriculture & Natural Resources, Delaware
State University, Dover, Delaware 19901-2277, U.S.A. rnaczi@dsc.edu

Charles T. Bryson

USDA-ARS, Southern Weed Science Research Unit, P.0. Box 350, Stoneville,
Mississippi 38776, U.S.A. cbryson@ars.usda.gov

Theodore S. Cochrane

Herbarium, Department of Botany, University of Wisconsin-Madison, Madison,
Wisconsin 53706-1381, U.S.A. tscochra@facstaff.wisc.edu

ABSTRACT. Seven species in four sections of Car-
ex are described as new from North America: C.
acidicola Naczi, C. calcifugens Naczi, C. paenin-
sulae Naczi, E. L. Bridges & Orzell, and C. thornei
Naczi (all sect. Griseae ); C. kraliana Naczi & Bry¬
son (sect. Laxiflorae ); C. gholsonii Naczi & Coch¬
rane (sect. Granulares); and C. infirminervia Naczi
(sect. Deweyanae). Carex acidicola , C. calcifugens,
C. paeninsulae, and C. thornei . all members of the
C. oligocarpa complex, have distichous perigynia
and purple-red shoot bases. Carex acidicola , from
a few sites in Alabama and Georgia, is distinctive
in having the bases of the proximal bract blades
whitish. Carex calcifugens occurs in a few popula¬
tions in Florida, Georgia, North Carolina, and South
Carolina. It is characterized by having the purple-
red coloration at the plant base less extensive than
in other members of the C. oligocarpa complex, the
vegetative shoots exceeding the reproductive ones,
and by bearing the proximal-most spikes at the ba¬
ses of the shoots. Carex paeninsulae is restricted to
a few sites in Florida. It is unique in the C. oli¬
gocarpa complex in having relatively long rhi¬
zomes. Carex thornei occurs in a few sites in Ala¬
bama, Florida, and Georgia. It has short bract
blades and terminal spikes with long peduncles.
(uirex kraliana is widespread and frequent through¬
out most of the southeastern United States. It is
distinctive in its lanceolate or narrowly lanceolate,
wide bract blades. Carex gliolsonii occurs in Ala¬
bama. Florida, Georgia. North Carolina, and South
Carolina. It is rare in all portions of its range except
for parts of Florida. It is distinctive in being loosely
caespitose, having short ligules, short bract blades,
long and w ide terminal spikes, and long perigynia.

Novon 12: 508-532. 2002.

The chromosome number of C. gholsonii, deter¬
mined from two populations in Florida, is 16 II +
I IV. Carex infirminervia is widespread but infre¬
quent throughout much of the western United States
and western Canada. It is unique in its combination
of papillose culms, long ligules, perigynium beaks
occupying a high percentage of the perigynium
length, and toothless or short-toothed perigynium
apices. The new combination C. digitalis var. flor-
idana (F. II. Bailey) Naczi Bryson ( Carex sect.
Careyanae) is proposed to replace C. digitalis var.
asymmetrica Fernald because of the priority of Bai¬
ley’s epithet.

Key words: Carex, Carex sect. Careyanae. Car¬
ex sect. Deweyanae, Carex sect. Granulares, Carex
sect. Criseae, Carex sect. Laxiflorae, Cyperaeeae,
North America.

Recent studies of North American sedges have
resulted in the description of many new species,
particularly for the southeastern United States and
western North America. The southeastern U.S.
hosts a great diversity of Carex (Cyperaeeae) in a
wide range of habitats. Tucker (1987) estimated
165 species of Carex for the region. Western North
America also possesses a diverse array of Carex
species. In this paper, we increase the number of
known species of North American Carex by de¬
scribing seven new species, six from the Southeast
and one from the West. Four of these new species
belong to Carex sect. Criseae (F. H. Bailey) Kiiken-
thal, one belongs to Carex sect. Laxiflorae Kunth,
one is a member of Carex sect. Granulares (().
Fang) Mackenzie, and one belongs to Carex sect.
Deweyanae (Tuckerman ex Mackenzie) Mackenzie.

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

509

We also propose one new combination in Carex
sect. Careyanae.

Four New Species of Carex sect. Griseae

Carex sect. Griseae is a group of 2 I species en¬
demic to eastern North America. The center of di¬
versity for the section is the southeastern United
States, where 20 species occur. Most of diem in¬
habit mesic deciduous forests.

Apomorphies diagnosing Carex sect. Griseae are
the large number (usually 40 or more) of nerves
(sclerenchyma traces) that are impressed in the sur¬
face of living and dried perigynia. Additional dis¬
tinctive characters of the section, but not necessar¬
ily autapomorphic, are long-sheathing proximal
bracts, usually unisexual spikes with only the ter¬
minal one staminate, and glabrous perigynia that
are obtusely triangular or suborbicular in cross sec¬
tion and have entire orifices.

Carex oligocarpa Willdenow, C. bulbostylis Mac¬
kenzie, C. edwardsiana E. L. Bridges & Orzell, and
C. planispicata Naczi constitute a morphologically

distinctive lineage within Carex sect. Griseae. As
identified by Naczi (1999a). members of this C. oli¬
gocarpa complex have distichously imbricate peri¬
gynia and purple-red coloration on the proximal
(2. 7— )3.0— 1 2 cm of the shoot bases. Other members
of Carex sect. Griseae have spirally imbricate peri¬
gynia and brown shoot bases or, if purple-red, the
coloration is usually less than 3.7 cm high. Recent
herbarium, field, and laboratory work by Naczi has
resulted in the discovery of four new species within
the C. oligocarpa complex, which are described be¬
low as C. acidicola, C. calcifugens, C. paeninsulae,
and C. thornei. With these new species, the C. oli¬
gocarpa complex now contains eight species and
becomes the largest species complex within Carex
sect. Griseae. Members of this complex are also the
most difficult to identify in Carex sect. Griseae.
Most previous determinations of specimens of these
new species were as C. oligocarpa. Each of these
new species appears to be rare and quite limited
in geographic distribution. The new species are dis¬
tinguished from other members of the C. oligocarpa
complex in the following key.

Key to Members of the Carex oligocarpa Complex (Members of Carex sect. Griseae with Distichous Perigynia
and Purple-red Shoot Bases)

This key is designed for identification of complete and ample specimens bearing mature perigynia. Because collectors
often fail to gather rhizomes of C. paeninsulae , this species is keyed twice to permit identification of incomplete
specimens of it.

la. Perigynia much inflated, orbicular or suborbicular in cross section, (1.8— )2.0-2.5(— 2.8) mm wide, 1 .6-2.0(-2. 1)

times as long as wide . C. bulbostylis

lb. Perigynia tightly enveloping achene or slightly inflated, obtusely triangular in cross section, 1 .4— 1 .9(— 2.3) mm
wide, (1.8— )2.1— 3.3 times as long as wide.

2a. Loosely caespitose, longer rhizomes 10—58 mm long between shoots or branches of the rhizomes .

. C. paeninsulae sp. nov.

2b. Densely caespitose, longer rhizomes 0.2— 6(— 8) mm long between shoots or branches of the rhizomes.

3a. Proximal bracts with bases of blades white between veins (most easdy seen on abaxial surfaces of

blades) . C. acidicola sp. nov.

3b. Proximal bracts with bases of blades uniformly green.

4a. Perigynia (1.8— )2. 1-2. 6 times as long as wide, with bodies usually abruptly contracted near
apices to beaks (0.4— )0.5-1.0 mm long; longest lateral spike with 4 to 8(10) perigynia (including
undeveloped or aborted ones).

5a. Proximal-most spike usually considerably above base of shoot, inflorescence 46— 94(— 99)%
of culm height; vegetative shoots exceeded by culms or slightly exceeding culms, tallest
vegetative shoot 0.88-1.4 times as tall as tallest culm; hyaline band of sheaths of proximal
bracts apically convex and elongated (0.8— )1. 1—4.0 mm beyond sheath apex ... C. oligocarpa
5b. Proximal-most spike usually at base of shoot, inflorescence (88— )94— 99% of culm height;
vegetative shoots usually greatly exceeding culms, tallest vegetative shoot (0.80-)1.4— 4.4
times as tall as tallest culm; hyaline band of sheaths of proximal bracts apically concave or
truncate.

6a. Purple-red coloration at shoot base 2.7— 5. 2(— 5.6) cm high; bodies of pistillate scales

(2.2-)2. 5—3.0 mm long; achene beaks (0. 10-)0.20-0.30 mm long C. calcifugens sp. nov.
6b. Purple-red coloration at shoot base (3.7— )4. 7—1 1.7 cm high; bodies of pistillate scales

1.5— 2.1 (—2.5) mm long; achene beaks 0.05— 0.1 0(— 0.15) mm long . C. edwardsiana

4b. Perigynia (2.4— )2. 5-3.3 times as long as wide, with bodies gradually tapering to apices and thus
beakless or w ith beaks 0.1— 0.4 mm long; longest lateral spike with (5)< to 14 perigynia (including
undeveloped or aborted ones).

7a. Perigynia (1 .9-)2.0-2.3 times as long as achene bodies; achene beaks (0.3— )0. 4-0. 7 mm

long . C. planispicata

7b. Perigynia 1. 7-2.0 times as long as achene bodies; achene beaks 0. 1-0. 3(— 0.4) mm long.

510

Novon

8a. Longest (per plant) peduncle of terminal spike (2.2-)5.1-8.9(-10.4) cm long (for this
measurement, include the portion ol the peduncle enclosed in the sheath of the distal-
most lateral spike); 2 distal-most lateral spikes usually widely separate, longest (per
plant) internode between distal lateral spikes (5.3-)8.8-15.2 cm; vegetative shoots ex¬
ceeded bv culms or slightly exceeding culms, tallest vegetative shoot 0.5-1. 3 times as
tall as tallest culm . C. thomei sp. nov.

8b. Longest (per plant) peduncle of terminal spike 0.9-3. 2(-6.8) cm long (for this measure¬
ment, include the portion of the peduncle enclosed in the sheath of the distal-most
lateral spike); 2 distal-most lateral spikes usually approximate, longest (per plant) in¬
ternode between distal lateral spikes 0.9— 7. 4(— 9.8) cm; vegetative shoots greatly ex¬
ceeding culms, tallest vegetative shoot 1.4-2. 2 times as tall as tallest culm .

. C. paeninsulae sp. nov.

Carex acidicola Naezi, sp. nov. TYPE: U.S.A.
Georgia: Clarke Co., ca. 5 air mi. SSE of cen¬
ter of Athens, ea. 0.5 mi. S of end of Rock and
Shoals Rd.. 20 May 1995, K. F C. Naezi 4741
(holotvpe. DOV; isotvpes, GA, MICIL NCU,
NY, US, VDB, herb. Bryson). Figure 1.

A ceteris speciebus Carici oligocarpae affinibus inter¬
venes albis ad bases laminarum bractearum proximalium
differt.

Perennial, densely caespitose. Rhizomes 0.2-6
mm long between shoots or branches of the rhi¬
zomes. Shoot bases purple-red to (3.2-)4. 1-7.4 cm
high. Culms 1 1-41 cm tall. Leaves of reproductive
shoots with widest blade per plant (3.7— )4.3— 5.5
mm wide, deep green except proximal ones proxi-
mally whitish. Vegetative shoots 26—42 cm tall.
0.72—1.3 times as tall as culms. Infructescences
8.8—36 cm long, 68—95% of culm height, with the
spikes widely separate or the distal-most 2 spikes
overlapping, the internode between the distal lat¬
eral spikes 1.4—13 cm long, the longest (per plant)
internode between the distal lateral spikes 3.1-13
cm long; proximal bract blades proximally white
between veins, hyaline band of adaxial face of
sheath with apex truncate or concave or slightly
convex and elongated to 1.3 mm beyond sheath
apex; bract blade of distal-most lateral spike usu¬
ally exceeding terminal spike but occasionally
shorter than terminal spike, longest (per plant)
bract blade of distal-most lateral spike 4.9-12 cm
long. Spikes 2 to 5; terminal spike 12-49 mm long,

( 1.3— )2. 1—4.0 mm wide, the longest per plant
(2 1 — )26 — 49 mm long, usually much exceeding dis¬
tal-most lateral spike, on peduncle 1 4 — 98(— 141)
mm long, the longest peduncle per plant 18-98
(-141) mm long; lateral spikes — 2 1 X 3.2— 5.7
mm. entirely pistillate. (2)4- to 7(1 l)-flowered, the
longest per plant 5- to 7(1 l)-flowered, the perigynia
distichously imbricate, the internode between the
proximal-most scales in the proximal-most spike
2. 4-3.0 mm long, perigynia overlapping, with ratio
of length of longest lateral spike per plant (iti mm) :
number of perigynia = 2.0-2. 4. Staminate scales

1.2-6.5 X 1 .6— 2.4 mm, obtuse or acute, awnless.
Pistillate scales 2.8— 4.8 X 1.6-2. 1 mm; body
(2.0— )2. 3— 2.9 mm long, with midrib prolonged as
awn 0.6-2.8 mm long. Perigynia (3.7-)4.2-4.9 mm
long, 1.6-1. 9 mm wide, 2.3-2. 7(-2.9) times as long
as w ide, 1 .6— 1.8(— 1 .9) times as long as achene bod¬
ies, ascending, obtusely triangular in cross section,
nerves deeply impressed and 49 to 64, narrowly
obovate or obovate or narrowly elliptic or elliptic
in outline, gradually tapered from widest point to
base, gradually tapered or somewhat abruptly ta¬
pered from widest point to apex, beakless or with
straight beak; beaks 0— 0.7(— 0.9) mm long. 0-17%
of perigynium length, vertical. Achenes 3. 1-3.3 X
1.5-1. 8 mm, faces tightly enveloped by perigynia,
proximally abruptly contracted to stipe, distally
abruptly contracted to minute beak; stipe 0.3-0. 6
mm long, usually vertical; body 2. 4-2. 7 mm long;
beak 0.20—0.30 mm long, vertical.

Carex acidicola is unique in the C. oligocarpa
complex in having the proximal bracts with the ba¬
ses of the blades white between the veins. This fea¬
ture is seen most easily on the abaxial surface of
the blades. All other members of the complex have
bract blades that are uniformly green. The region
of the blades that is whitish is limited to a small
area at the very base of the bract blade (immedi¬
ately distal to the sheath summit). In this region,
the veins and tissues immediately flanking veins
are green, whereas the intervenal spaces are con¬
trastingly white. Phis whitish region occurs quite
consistently on the proximal bracts of all specimens
seen of C. acidicola , though it may be difficult to
discern the contrast of white and green on older,
faded specimens. Because the whitish region is
limited in extent and consistent in appearance, it
does not appear to be teratologic. Rather, it appears
to be a regular, predictable character state of C.
acidicola.

Carex acidicola is also distinctive in its relatively
wide leaves (widest leaf blade per plant [3.7— ]4. 3—
5.5 mm wide). All other members of the C. oligo¬
carpa complex have the widest leaf blade 1 .8-4.0

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

511

f igure 1. Carex acidicola Naczi. Distal portion of culm
(X 0.9); abaxial view of perigynium in long section to
reveal achene (upper), and abaxial view of whole perigyn¬
ium (lower) (both X 17.5).

mm wide, except for C. oligocarpa (widest leaf
blade 1 .8-3.9[-4.6] mm w ide), C. paeninsulae (3.0-
4.6 mm wide), and C. planispicata ([3.0— ]3. 5-6. 5
mm wide). Several features distinguish C. acidicola
and C. oligocarpa. Besides different proximal bract
blade coloration, C. acidicola lias a shorter hyaline
band of the adaxial face of the sheaths of the prox¬
imal-most bracts. In C. acidicola, the apex of the
hyaline band is usually truncate or concave, but
rarely elongated as much as 1.3 mm beyond the
sheath apex. In C. oligocarpa , the apex of the hy¬
aline band is convex and elongated (0.8-) 1 .1—4.0
mm beyond the sheath apex. Also, C. acidicola has
perigynia with a higher length: width ratio (peri-
gynia 2.3— 2.9 times as long as wide in C. acidicola
vs. 2. 0-2. 6 times as long as wide in C. oligocarpa).
Differences between C. acidicola and C. paenin¬
sulae include habit and rhizome length (densely
caespitose. with rhizomes 0.2—6 mm long in C. aci¬
dicola vs. loosely caespitose and rhizomes 7-58
mm long in C. paeninsulae), and length of vegeta¬
tive shoots relative to culms (vegetative shoots
0.72-1.3 times as tall as culms in C. acidicola vs.
1.4-2. 2 times as tall as culms in C. paeninsulae).
Carex acidicola differs from C. planispicata in three
ways. First. C. acidicola has longer terminal spikes.
The longest terminal spike per plant is (2. 1 — )2.6 —
4.9 cm long in C. acidicola , whereas it is 1 .3—
2.6(-3.0) cm long in C. planispicata. Second, peri¬
gynia of C. acidicola are 1.6— 1.8(— 1.9) times as
long as the achene bodies versus (1.9-)2.0-2.3 in
C. planispicata. Third, C. acidicola has the achene
beaks 0.2-0. 3 mm long versus (0.3— )0.4— 0.7 mm
long in C. planispicata.

Carex acidicola occurs disjunctly in three areas:
central Alabama, central Georgia, and near the
Chattahoochee River in easternmost Alabama and
southwestern Georgia. It occurs in the Piedmont,
Appalachian Plateaus, Ridge and Valley, and
Coastal Plain. Carex acidicola is a very rare and
local species, collected from only ten populations.
The type and paratypes are all the specimens seen
of C. acidicola. Few individuals occur at six of the
seven populations studied by Naczi. Only at the
type locality are plants relatively common.

Carex acidicola inhabits lightly shaded, acidic,
humic, sandy loams in mesic forests. Vascular
plants frequently associated with C. acidicola are
Aesculus sylvatica Bartram, C. superata Naczi, Rez-
nicek & B. A. Ford. Carya spp.. Cercis canadensis
L., Hexastylis arifolia (Michaux) Small. Juniperus
virginiana L., and Ostrya virginiana (Miller) k.
Koch. Unlike most of the other members of the Car¬
ex oligocarpa complex, notably the widespread and
frequent C. oligocarpa, C. acidicola grows in acidic

512

Novon

soils rather than eireumneutral ones. Whereas an¬
alyzed soils of eight populations of C. oligocarpa
have ,,H values of 6.3-7. 6. those of four popula¬
tions of C. acidicola have pH values of 5.2— 5.8.
This apparent requirement for acidic substrates
makes “ acidicola " (literally, “acid-dweller") an ap¬
propriate epithet for this ecologically distinctive
species.

Paratypes. U.S.A. Alabama: Bibb Co.. 3.7 mi. NW
of town of Sixmile, N of Little Cahaba River, 31 May 1997,
A 'aczi 6346 N MacDonald (DOV); I ,ee Co., ca. 1.5 mi. N
of Smiths Station, 24 Apr. 2002, Naczi 9136 (DOV, MICH,
MO, NY, VI)B, herb. Bryson); Tuscaloosa Co., valley of
Rocky Branch, 19 Apr. 1936, Harper 3487 ( CH, MO, NY.
US); N side of Croton bluff on Warrior River, 16 Apr.
1959, Harper 443 1 (CA, UNA); NK from Tuscaloosa,
along Black Warrior River, It) Apr. 1992. McKinney 4925
N Hillman (DOV); ca. 1.5 mi. NW of Belt M’son, at Black
Warrior River, 15 May 1995, Naczi 4624 (DOV, MICH);
ca. 9.5 mi. NK of center of Tuscaloosa, Rocky Branch
Public Use Area. 29 Apr. 1996. Naczi 5075 (DOV), 22
May 1996, Naczi 5437 & Bryson (DOV), 22 Apr. 1999,
Bryson 17073 & MacDonald (DOV, MICH. VDB, W IN,
herb. Bryson); ca. 6.5 mi. NK of center of Tuscaloosa, near
Hurricane Creek, I June 1997, Naczi 6368 (DOV). Geor¬
gia: Clarke Co., type locality, 29 Apr. 1981, Manhart 234
(DOV, herb. Bryson), 10 May 1986. Naczi 1065 N Man-
hart (DOV, MICH), 24 May 1988. Naczi 1963 (FKAS,
fSU, CA. DOV, MICH, UNA, herb. Bryson); Decatur Co.,
above Lake Seminole, 27 May 1984, Saucier s.n. (herb.
Bryson); Dekalb Co., Stone Mountain, 27 Apr. 1929, Mill¬
er 275 & Maguire (BH, CU); Early Co., W of Hilton, bluff
along Chattahoochee River, 14 May 1947, Thorne 3886
(CU); opposite Columbia, Alabama, bluffs of Chattahoo¬
chee, 11 Apr. 1977, Krai 59607 (DOV, VDB); Walton Co.,
ca. 1 mi. E of Walnut Grove, 24 May 1988, Naczi 1967
(MICH), 20 May 1995, Naczi 4745 (DOV, MICH).

Carex calcifugens Naczi, sp. nov. TYPE: U.S.A.
Georgia: Screven Co.. Blue Springs, Blue
Springs Landing on Savannah River, 2 May
1991, K. F. C. Naczi 2840 (holotype, DOV;
isotypes. FLAS, FSU, GA, GH. MICH, MO,
NCU, NY, PH, TENN, UNA, US. USCH. VDB,
A PI, VSC, Vt IN, herb. Bryson), f igure 2.

A Carice oligocarpa spica infima plerumque e basi eul-
mi portata, foliis surculorum vegetativorum culmos pler¬
umque superantibus, vilta hyalina adaxiali vaginae brae-
teae infimae ad apicem truncata vel concava differt; a
Carice edwardsiana basibus surculorum atrovinosis usque
2.7— 5.2(— 5.6) cm tantum altis, corporibus squamarum pis¬
til lataru m longioribus, rostris acheniorum longioribus dif¬
fert.

Perennial, densely eaespitose. Rhizomes 0.2—7
mm long between shoots or branches of the rhi¬
zomes. Shoot bases sparingly purple-red to 2.7-
5.2(-5.6) cm high. Culms 11-37 cm tall. Leaves of
reproductive shoots with widest blade pier plant
2.6-AA) mm wide, green. Vegetative shoots 23-45
cm tall, (0.8— ) 1.4— 1.6 times as tall as culms. In-

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Carex from North America

513

fructescences 19-34 cm long, (88-)94— 99% of
culm height, with the spikes separate or t lie distal-
most 2 to 3 spikes overlapping, the internode be¬
tween the distal lateral spikes (1.0— )2. 6— 12 cm
long, the longest (per plant) internode between the
distal lateral spikes 5.2-12 cm long; proximal bract
blades uniformly green, hyaline band ol adaxial
face of sheath with apex truncate or concave; bract
blade of distal-most lateral spike exceeding termi¬
nal spike, longest (per plant) bract blade of distal-
most lateral spike 6.1-15 cm long. Spikes 4 to 5;
terminal spike 8-35 X 1.4-2. 5 mm, the longest per
plant 14—35 mm long, exceeding distal-most lateral
spike, on peduncle 3.1-54(-73) mm long, the lon¬
gest peduncle per plant 17— 54(— 73) mm long; lat¬
eral spikes 11-17 X 4.5— 8.5 mm, entirely pistil¬
late. 2- to 8-flowered. the longest per plant 5- to
8-flowered, the perigynia distichously imbricate,
the internode between the proximal-most scales in
the proximal-most spike 3. 0—4. 3 mm long, perigyn¬
ia overlapping, with ratio of length of longest lateral
spike per plant (in mm) : number of perigynia =
2.2— 2.4. Staminate scales 4.3— 5.8 X 1.4— 1.9 mm.
usually acute but occasionally obtuse, awnless. Pis¬
tillate scales 3.6-5. 7 mm long, 1 .8-2. 1 (-2.4) mm
wide; body (2.2-)2. 5-3.0 mm long, with midrib pro¬
longed as awn 1 .0—3.2 mm long. Perigynia (3.7—)
3.9-5.0 mm long. 1 .6— 2.0(— 2. 1 ) mm wide, 2.1-
2.5(— 2.7) times as long as wide, 1.5— 2.0 times as
long as aehene bodies, ascending, obtusely trian¬
gular in cross section, nerves deeply impressed and
48 to 63. obovate or narrowly obovate in outline,
gradually tapered from w idest point to base, abrupt¬
ly tapered from widest point to straight or slightly
excurvet I beak; beaks (0.4— )0.6- 1 .0 mm long, 8.3-
21% of perigynium length, vertical or bent 10-30°
from vertical. Achenes 2.6— 3.4 X 1.4— 1.9 mm, lac¬
es tightly enveloped by perigynia, proximally
abruptly contracted to stipe, distal ly abruptly con¬
tracted to minute beak; stipe 0.2— 0.5(— 0.7) mm
long, vertical; body 2. 0-2. 6 mm long; beak
(0.10-)0. 20-0.30 mm long, vertical.

Carex calcifugens is morphologically most simi¬
lar to C. eduiardsiana and C. oligocarpa. All three
of these species have a densely caespitose habit,
the widest leaf blade per plant 1 .8-4.0 mm wide,
the longest pistillate spike 4- to 8(10)-flowered,
perigynia usually 2.1— 2.6 times as long as wide,
perigynium bodies abruptly contracted to beaks
(0.4— )0.5-l.0 mm long, and perigynia 1. 5-2.0
times as long as the aehene bodies. As indicated
in the key, C. calcifugens shares several features
with C. edwardsiana that distinguish both of these
species from C. oligocarpa: the infructescence with

the proximal-most spike usually at the base of the
plant, vegetative shoots usually greatly exceeding
culms, and hyaline bands of sheaths of proximal-
most bracts apieally truncate or concave. Carex cal¬
cifugens differs from C. edwardsiana in having the
purple-red coloration of the shoot bases less exten¬
sive, longer bodies ol the pistillate scales, and lon¬
ger aehene beaks.

Carex calcifugens ranges from eastern North Car¬
olina south to northern Florida, strictly on the
Coastal Plain. It is rare and very local. The type
and paratypes are all the specimens examined ol
C. calcifugens. The total number of documented
populations is about 18. Most populations seen
consisted of fewer than 20 individuals. At the type
locality, however, the plants were relatively numer¬
ous. I’he plants usually occur on slopes above
streams in mesic deciduous forests with diverse
herbaceous and woody angiosperm communities.
Specimen labels indicate C. oligocarpa also occurs
in maritime woodlands, frequent vascular plant as¬
sociates of C. calcifugens are Aesculus pavia L..
Carex abscondita Mackenzie, C. striatula Michaux,
Cercis canadensis , Cornus florida I... Fagus gran-
difolia Ehrhart, Hexastylis arifolia, Polystichum ac-
rostichoides (Michaux) Schott, and Sanguinaria
canadensis L. The substrates for four studied pop¬
ulations are well-drained sandy loams, loamy
sands, and sands. Analyses of the soils in the im¬
mediate vicinity of the roots of 6. calcifugens at
these sites reveal low levels ol calcium (calcium
concentration at four sites = 473-920 ppm). Since
several of the populations occur over shell mounds
and deposits of fossil shells, it is likely the paucity
of calcium is due to leaching. The epithet ^ calci¬
fugens" (“calcium-fleeing”) is fitting for this new
species because of its affinity for calcium-poor
soils, iu contrast to its close relatives C. oligocarpa
and C. edwardsiana. which are ealciphiles (calcium
concentration at eight sites of C. oligocarpa = 840—
5937 ppm, calcium concentration at four sites of 6.
edwardsiana = 5937—6190 ppm, Naczi, unpub¬
lished data).

Paratypes. U.S.A. Florida; Leon Co., near Iallahas-
see, 8 Apr. 1910, Harper 65 (GH, MIN, NV); ca. 2 mi.
ESE of Tallahassee, 26 Apr. 1925, Harper 50 (811, GH,
ILL, NY, PH. US); Liberty Go., Allen [Alum] Bluff of Ap¬
alachicola River, 1 I Apr. 1931, Palmer 58557 ((41. MO.
NY); ca. 6 mi. NE of Bristol, II May 1989. Orzell &
Bridges 9716 (DOV, MICH), 1 Apr. 1990, Orzell & Bridges
15061 (DOV, MICH). 14 Apr. 1990, Orzell & Bridges
15155 (DOV, MICH). Georgia: Baker Co., E side of
Ichauwaynochaway Creek, Ichauway Plantation, 1 1 Apr.
1986, Gholson 1 1599 (FLAS, herb. Bryson); Burke Co., I
of McBean, Shell Bluff Landing, 10 Apr. 1938, Clausen
& Trapido 5589 (BH, CU, OKL), 10 Apr. 1938, Pyron &
McVaugh 2502 (FSU, GA); 7 mi. NE of Shell Bluff, near

514

Novon

Savannah River, 18 May 1989, Naczi 2191 (IH)V, GA,
MICH, MO, NY. VI)B, herb. Bryson), 5 June 1990, Naczi
2522 (MICH), 25 May 1999, Morris 4525 & Patrick (HOY,
herb. Bryson); ea. 3 mi. E ol Waynesboro, along Brier
Creek, 18 May 1989, Naczi 2195 (I )(>V); Early Co., N of
Blakely, Colomokee Mounds Slate Park, 4 Apr. 1948,
Tliornc 7971 & Muenscher (CU, GA, GEO, GH); Effing¬
ham Co., N of Clyo, along the Savannah River, 21 Apr.
1902, Ahles 56549 el al. (NCU), 2 May 1991, Naczi 2552
(l)OV). North Carolina: Brunswick Co., Smith Island,
May 1894, Ashe s.n. (NCU); Carteret Co., near Lenoxville,
1 I Apr. 1898, Ashe s.n. (ESU. MICH. NCU); near Shac¬
kleford Pt., 1 I Apr. 1898. Ashe s.n. (NCU); 1.2 km from
the W end ol Shackleford Island, 20 Mav 1990, Peel 6794
& Ixirke (NCU); Dare Co., near Cape Hatteras, 10 June
1898, Ashe s.n. (NCU); Buxton Woods, 23 May 1954, Rad¬
ford 5116 (NCU). South Carolina: Aiken Co., Savannah
River Site, Compartment 48, Stand 37, 13 May 1993,
Stanford 152 (USCH); Savannah River Site, slope of Up¬
per Three Runs Creek, 10 June 1994, Hyatt 6055 (DOV,
MICH); Savannah River Plant, compartment 31, near Ty¬
ler Rd., along linker (.reek, 28 Apr. 2000, Pittman
1)425000 1 & Darr 213 (DOV. MO, USCH, herb. Bryson);
Allendale Co., Savannah River Operations Area of the
Atomic Energy Commission, 6 Apr. 1953, Batson & Kelley
s.n. (US(.H); Barnwell Co., Savannah River Plant, along
Lower Three Runs Creek. 28 Apr. 2000, Pittman
04250021 & Darr 227 (DOV, USCH); Charleston Co.,
Edisto State Park, 9 Apr. 1994, Pittman 04099405
(MICH, USCH); Dorchester Co., SW of Summerville, near
Ashley River, 2i May 1957, Aides 26-11 1 & Haesloop
(NCU), 19 Apr. 1962, Radford 44505 (FSU, GA, (ill,
NCU, NY, VDB), 25 May 1988, Naczi 1970 (DOV, FLAS,
MICH. MO, NY, USCH. VDB, WIN, herb. Bryson), 6 June
1990, Naczi 2524 (MICH); Orangeburg Co., S end of San¬
tee State Park, 19 May 1997, Pittman 05199702 (DOV
MICH. USCH).

Carex paeninsulae Naczi, E. 1,. Bridges & Orzell,
sp. nov. TYPE: U.S.A. Florida: Clay Co., ca. 1
mi. N of Green Cove Springs, Magnolia
Springs, 0.4 mi. N of rte. 17. W of Haven Av¬
enue and S of stream. 20 Apr. 1991, R. F. C.
Naczi 2770 (holotype, DOV; isotypes, FLAS,
FSU, GA, GH, MICH, MO. NY, VDB, WIN,
herb. Bryson). Figure 3.

Carex grisea var. angustifolia Boott, Illustr. Genus Carex
34. 1858. TYPE: U.S.A. Florida: |no additional lo¬
cality data], [no date). Chapman s.n. (lectotype, des¬
ignated here, k: isolectotype, k).

A ceteris speciebus Carici oligocarpae affinibus rhizo-
matibus longioribus differt.

Perennial, loosely eaespitose. Rhizomes 7—58
mm long between shoots or branches of the rhi¬
zomes. Shoot bases purple-red to 4.8-1 1.4 cm high.
Culms 8.9—39 cm tall. Leaves of reproductive
shoots with widest blade per plant 3.0-4.6 mm
wide, green. Vegetative shoots 33—59 cm tall. 1.4—
2.2 times as tall as culms. Infructescences 8.1—29
cm long, 73-95% of culm height, with llie distal-
most 2 to 3 spikes overlapping or rarely all spikes

separate, the internode between the distal lateral
spikes 0.4-7. 4(-9.8) cm long, the longest (per
plant) internode between the distal lateral spikes
0.9-7. 4(-9. 8) cm long; proximal bract blades uni-
formly green, hyaline band of adaxial face of sheath
with apex truncate or concave; bract blade of distal-
most lateral spike exceeding terminal spike, longest
(per plant) bract blade of distal-most lateral spike
3. 2-9.5 t -m lung. Spikes 4 to 6; terminal spike 14—
38 X 1.2— 2. 1 mm, the longest per plant 17—38 mm
long, exceeding distal-most lateral spike, on pe¬
duncle 3.0— 32(— 68) mm long, the longest peduncle
per plant 8.7-32(-68) mm long; lateral spikes 7-
20 X 3. 5-4. 8 mm, entirely pistillate, 2- to 12-flow-
ered, the longest per plant (5)7- to 12-flowered, the
perigynia distichously imbricate, the internode be¬
tween the proximal-most scales in the proximal-
most spike 2.4— 3.5 mm long, perigynia overlap¬
ping, with ratio of length of longest lateral spike
per plant (in mm): number of perigynia = 1.8-2. 5.
Staminate scales 3.9— 4.8 X 1.2— 1.8 mm, acute,
awnless or awned with awn up to 2.2 mm long.
Pistillate scales 2.5 — 4.2 X 1 .4—2.2 mm; body 1 .5—
2.4 mm long, with midrib prolonged as awn 0.4-
2.4 mm long. Perigynia 4. 1-4.7 X 1. 4-1.6 mm,
2.7— 3.1 limes as long as wide, 1. 8-1.9 times as
long as achene bodies, ascending, obtusely trian¬
gular in cross section, nerves deeply impressed and
49 to 66, narrowly obovate in outline, gradually ta¬
pered from widest point to base, gradually tapered
from widest point to apex, beakless or with straight
beak; beaks 0—0.4 mm long, 0—10% ol perigynium
length, vertical. Aehenes 2.0-3.4 X 1.3- 1.4 mm,
faces tightly enveloped by perigynia, proximally
abruptly contracted to stipe, distally abruptly con¬
tracted to minute beak; stipe 0.5— 0.6 mm long, ver¬
tical; body 2. 2-2. 6 mm long; beak 0.10-0.30 mm
long, vertical.

Carex paeninsulae is unique in the C. oligocarpa
complex in its loosely eaespitose habit as a resull
of relatively long rhizomes (longer rhizomes 10—58
mm long between shoots or branches of the rhizome
in C. paeninsulae). Other members of the complex
are densely eaespitose, with rhizomes short (0.2—8
mm long between shoots or branches of the rhi¬
zome). Carex paeninsulae differs from C. oligocarpa
in having vegetative shoots greatly exceeding the
culms (vegetative shoots 1. 4-2.2 times as tall as
culms in C. paeninsulae vs. 0.88-1.4 times as tall
as culms in C. oligocarpa), hyaline bands of adaxial
faces ol sheaths of proximal-most bracts truncate or
concave at their apices (convex and elongated 0.8—
4.0 mm beyond the sheath apex in C. oligocarpa),
and perigynia with a higher length: width ratio

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

515

Figure 3. Curex paeninsulae Naczi, E. E. Bridges &
0.9); and perigynium, lateral view (right; X 17.5).

Orzell. Hahit (left; X 0.45); distal portion of culm (middle; X

516

No von

(perigynia 2.7— 3. 1 times as long as wide in C. paen-
insulae vs. 2.0— 2.6 times as long as w ide in C. oli-
gocarpa). Carex paeninsulae is similar to C. calci-
Jugens and C. edwardsiana in having the vegetative
shoots exceeding t he culms, but differs from those
species in perigynia with a higher length: width ra¬
tio (perigynia 2.7— 3.1 times as long as wide in C.
paeninsulae versus 1.9-2.5[-2.7] times as long as
wide in C. calcifugens and C. edwardsiana). In ad¬
dition, C. paeninsulae has the purple-red coloration
extending higher on the bases of the plants than in
C. calcifugens : to 4.8—1 1.4 cm high in C. paenin¬
sulae versus to 2.7-5.2(-5.6) cm high in C. calci¬
fugens. Furthermore, C. paeninsulae has shorter
bodies of the pistillate scales than C. calcifugens:

1.5— 2.4 mm long in C. paeninsulae versus (2.2-)

2.5— 3.0 mm long in C. calcifugens. Relative to C.
edwardsiana, C. paeninsulae has longer achene
beaks. In C. paeninsulae, these beaks are 0.10—
0.30 mm long versus 0.05-0. 10(-0. 15) mm long in
C. edwardsiana.

Carex paeninsulae occurs in the northern half of
peninsular Florida, strictly on the Coastal Plain. It
is rare and local, collected from only about 10 pop¬
ulations. The type and paratypes represent all the
collections we have examined of this species.
Plants grow in sandy loams in mesie deciduous for¬
ests in hardwood hammocks. Frequent vascular
plant associates of C. paeninsulae include Arisaema
dracontium (1,.) Schott, Carpinus caroliniana Wal¬
ter. LiquiAambar styracijlua I,., Magnolia grandi-
flora I... (Juercus virgin iana Miller, Ruellia carol i-
n lens is (J. 1. G me I in) Steudel, Sabal minor
(Jacquin) Persoon, and .S', palmetto Schultes &
Schultes. Because of its apparent restriction to the
peninsula of Florida, "paeninsulae" is an appropri¬
ate epithet for this new species.

Edwin Bridges and Steve Orzell supplied nu¬
merous specimens and habitat notes to Robert Nac¬
zi for his description of Carex paeninsulae. In re¬
turn. Naczi has complied with their request to be
included as authors of the description.

Carex grisea var. angustifolia, as lectotypified
here, is a synonym of C. paeninsulae. Though frag¬
mentary, the lectotype possesses several features
that are diagnostic of C. paeninsulae: a relatively
long portion of a rhizome, a vegetative shoot dial
exceeds its associated culm, the longest peduncle
of the terminal spike 24 mm long, the longest lat¬
eral spike with 8 perigynia, and perigynia with a
length : width ratio of 2.8. Bootl s syntypes of Carex
grisea var. angustifolia are a mixture of C. paen¬
insulae, C. bulbostylis, and C. corrugata Fernald.
For the lectotype. Naczi has chosen the specimen

that best matches Rood’s description (Rood, 1858:
35), especially “. . . perigyniis minus turgidis, apice
saepe suhrostellatis . . . ,” which are more charac¬
teristic of C. paeninsulae than C. bulbostylis or C.
corrugata. I he lectotype is also the syntype that
Rood illustrated for C. grisea var. angustifolia
(Rood. 1858, pi. 87).

Paratypes. II. S. A. Florida: [county unknown |. “Flor¬
ida” [no additional data], 1859, Chapman s.n. ((ill);
[Bradford] (.<>., Mount Vernon, 1838, Chapman H (NY);
(.lay Co., type locality, 5 May 1990, Naczi 2373 (DOV,
MICH); [Duval) Co., Island of F oil George, 1817. Baldwin
s.n. (NY); Si. Johns, [no date, but most likely 1817 — see
Darlington, 1843], Baldwin s.n. (PH); Gilchrist [“bevy” on
label] Co., I mi. N of Trenton, 31 Mar. 1957, Krai 4385
(FLAS, FSU, GH, MSC, VDB); Hernando Co., near Pi-
neola, fern grottoes, 12 Apr. 1923, Small 10835 A* Mosier
(NY); Annutteliga Hammock, 26 Mar. 1958, Cooley 5807
& Baton (FSU, Cl), USF); Pineola Grotto, N of Brooksville
near Istachatta, 8 Mar. 1977, Wunderlin 5751 (USF); ea.
5 mi. N of Brooksville on rte. 41, 14 Apr. 1994, Libby,
Wears' A: Abbott s.n. (DOV, MICH); ca. 6 air mi. NF of
Brooksville, W side US 41. 12 Apr. 1994, Orzell A Bridg¬
es 22034 (1 IG. DOV, USF); bevy Co.. Waccasassa Bay
State Preserve, along boundary N of Turtle Creek. 12 Apr.
1996, Abbot i 8382 (DOV, FbAS); Marion Co., 4 mi. W of
Anthony, 2 Apr. 1950, Blake s.n. A Lewis (FbAS); E side
of Silver Springs, 13 Mar. 1976, Krai 57220 (DOV, VDB);
N side of US 27, ca. 1.6 mi. SF of junction with rte. 326
at Blichton, 28 Mar. 1993, Orzell A Bridges 21348 (DOV),
8 May 1993. Orzell A Bridges 21001 (FI G, DOV, MICH,
US. USF ); N side of LIS 27, ea. 1.8 mi. SF of junction
with rte. 326 at Blichton, 28 Mar. 1993. Orzell & Bridges
21352 (DOV, FTG, MICH, NY, US, USF. herb. Bryson);
ca. 3.1 air mi. N of Flemington, 17 Apr. 1993, Orzell A-
Bridges 21003 (US, FTG, DOV, MICH, USF); ea. 0.8 air
mi. NNW of junction of rte. 318 and US 301 in Citra. 17
Apr. 1993, Orzell A Bridges 21024 (FTG, DOV); 3.2 air
mi. SW of Belleview, 18 Apr. 1993, Orzell A' Bridges
21020 (FbAS, FSU, FTG, DOV, MICH. NY. US. herb.
Bryson); Seminole Co., near Oviedo, Black Hammock. 12
Apr. 1975, Wunderlin 5450 A Poppleton (USF); Sumter
Co., ca. 2.2 mi. FNF of l.inden, 15 Mar. 1992, Orzell A
Bridges 10107 (DOV, FbAS, FSU, FTG, CA. MICH, MO,
NCU, NY. US, herb. Bryson); Suwannee Co., ca. 10.3 air
mi. NW of bive Oak, on Suwannee River across from
mouth of Alapaha River, 21 Mar. 1991, Orzell A Bridges
10145 (FTG, MICH).

Carex lliurnei Naczi, sp. nov. TYPE: U.S.A. Al¬
abama: Russel] Co., 2.0 road mi. SW of Holy
Irinity, along E side of rte. 165, along S side
of Bluff Creek. 3 May 1996. R. F. C. Naczi
5214 (holotype, DOV; isotypes, MICH. MO,
NY, US, W IN. herb. Bryson). Figure 4.

A ceteris speciebus Carici oligocarpae affinibus laminis
braetearum brevibus, pedunculo spieae temiinalis longo,
perigyniis (2.4 — )2.5 — 3. 1 plo longioribus quam latioribus
differ! .

Perennial, densely caespitose. Rhizomes 0.8— (>.4
mm long between shoots or branches of the rhi-

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Naczi et al.

Carex from North America

517

Figure 4. Carex thornei Naczi. Distal portion of culm (X
0.9); lateral view of perigynium in long section to reveal
achene (X 17.5).

zomes. Shoot bases purple-red to 4.8-8. 1 cm high.
Culms 24—53 cm tall. Leaves of reproductive shoots
with widest blade per plant 2. 1 — 4.0 mm with;,
green. Vegetative shoots 28—49 cm tall. 0.54—1.3
times as tall as culms. Infructescences 13—38 cm
long, 77—96% of culm height, with the spikes usu¬
ally separate or occasionally the distal-most 2(— 3)
spikes overlapping, the internode between the dis¬
tal lateral spikes (1.6-)3.4-15.2 cm long, the lon¬
gest (per plant) internode between the distal lateral
spikes (5.3-)8.8-15.2 cm long; proximal bract
blades uniformly green, hyaline band of adaxial
face of sheath with apex usually truncate or con¬
cave but occasionally convex and elongated to
2.2(— 5.0) mm from sheath apex: bract blade of dis¬
tal-most lateral spike slightly exceeding terminal
spike or exceeded by terminal spike, longest (per
plant) bract blade of distal-most lateral spike 3.9—

1 1 cm long. Spikes 4 to 5; terminal spike 16—36
X 1.9— 2.5 mm. the longest per plant 17—36 mm
long, exceeding distal-most lateral spike, on pe¬
duncle 3.4— 89(-104) mm long, the longest pedun¬
cle per plant (22-)51-89(-104) mm long; lateral
spikes 5-21 X 3.2-4. 1 mm, entirely pistillate, 3-
to 1 2-flowered, the longest per plant 6- to 1 2-flow -
ered, the perigynia distiehously imbricate, the in¬
ternode between the proximal-most scales in the
proximal-most spike 1.7— 2.9 mm long, perigynia
overlapping, with ratio of length of longest lateral
spike per plant (in mm): number of perigynia =
1.8— 2.1. Staminate scales 4.3— 5.0 X 1.4— 1.8 mm,
usually acute but occasionally obtuse, awnless. Pis¬
tillate scales 3. 0-4.6 X 1. 6-2.0 mm; body 1.6-2. 6
mm long, with midrib prolonged as awn 1.1— 2.5
mm long. Perigynia 3.8—4.9 X 1.4— 1.7 mm,
(2.4— )2. 5-3.1 times as long as wide, 1. 7-2.0 times
as long as achene bodies, ascending, obtusely tri¬
angular in cross section, nerves deeply impressed
and 53 to 64, narrowly obovate or narrowly oblong
in outline, gradually tapered from w'idest point to
base, gradually tapered from widest point to apex,
beakless or with straight beak; beaks 0—0.4 mm
long, 0-11% of perigynium length, vertical.
Aehenes 2.6-3. 5 X 1.3-1. 6 mm, faces tightly en¬
veloped by perigynia, proximally abruptly contract¬
ed to stipe, distally abruptly contracted to minute
beak; stipe 0.2— 0.6 mm long, vertical; body 2.1-
2.7 mm long; beak 0. l-0.4(-0.5) mm long, vertical.

Carex thornei is unique in the C. oligocarpa com¬
plex in its combination of short bract blades (lon¬
gest distal-most bract blade per plant 3.9—1 I cm
long), long peduncles of the terminal spikes (lon¬
gest peduncle of terminal spike per plant [ 22 — ]5 1 —
89[ — 104] mm long), and high length: width ratio of

518

No von

perigynia (perigynia [2.4— ]2. 5-3.1 times as long as
wide). Morphologically, il is most similar to C. oli-
gocarpa and C. planispicata. Carex thornei differs
from C. oligocarpa in having shorter hract blades
(longest distal-most bract blade per plant 1 4.9— |
9.2-14 cm long in C. oligocarpa). longer peduncles
of tbe terminal spikes (9.B — 49[ — 1 04 1 mm long in
C. oligocarpa). perigynia with a higher length:
width ratio (perigynia 2.0-2. 6 times as long as wide
in C. oligocarpa). and perigynia beakless or indis¬
tinctly beaked with beaks to 0.4 mm long (C. oli¬
gocarpa has distinctly beaked perigynia with beaks
[0.4— ]0. 5— 1.0 mm long). From C. planispicata, C.
thornei differs by having longer peduncles ol the
terminal spikes (longest peduncle of terminal spike
per plant 2.4-49 mm long in C. planispicata). the
perigynia shorter relative to the achene bodies
(perigynia 1. 7-2.0 times as long as achene bodies
in C. thornei vs. 1 1.9— ]2.0— 2.3 times as long as tbe
achene bodies in C. planispicata), and shorter
achene beaks (0.1— 0.4[— 0.5] mm long in C. thornei
vs. [0.3— ]0.4— 0.7 mm long in C. planispicata).

Carex thornei is a narrow endemic of the lower
Chattahoochee Kiver-lower Flint River— upper Ap¬
alachicola River drainage in the region near the
junction of the borders of Alabama, Florida, and
Georgia. It occurs on the Coastal Plain, from the
Fall Line south to Liberty County, Florida, a dis¬
tance of only about 225 km (140 mi.). Within this
region, it is sometimes frequent, but local. The
paratypes cited below represent most of the known
populations ol the species. Carex thornei occurs in
mesic deciduous forests, usually on the upper por¬
tions of floodplains and adjacent slopes, in sandy
loams. Frequent associates include Carex abscon-
dita, C. basiantha Steudel. Cercis canadensis. Fa-
gas grandifolia, and Hexastylis arifolia.

Carex thornei has an unusual pattern of ende¬
mism, but not unique. Other species of flowering
plants that are endemic to the region of the lower
Chattahoochee, lower Flint, and upper Apalachi¬
cola rivers include Rhododendron prunifolium
(Small) Millais, Torreya taxifolia Arnott, and Taxits
floridana Chapman.

Carex thornei is named for Robert F. Thorne.
Long ago (Thorne, 1951). lie suggested what was
being called C. oligocarpa from southwestern Geor¬
gia likely represented a new taxon. Actually, each
of the three vouchers he cited as “C oligocarpa ”
is a different new species, C. thornei ( Thorne 3100
et al.), C. acidicola (Thorne 3366), and C. calcifug-
ens ( Thorne 7071 & Muenscher). It is fitting to rec¬
ognize the taxonomic acumen of one of the earliest
collectors of this species by naming it for him.

Paratypes. U.S.A. Alabama: Henry Co., vicinity of
Shorterville, along Chattahoochee River S from Farmers
Landing, 16 Apr. 2000, MacDonald 13815 (DOV); Lee
Co., 1.5 mi. N of Smith's Station, 7 Apr. 1949, Duncan
9205 A Smith (GA); Russell Co., ca. 2.5 mi. NW of Cot-
tonton, along Sand Branch, 3 May 1996, Naczi 5201
(DOV, MICH); type locality, 24 May 1997, Naczi 6170
(DOV, herb. Bryson). Florida: Gadsden Co., Chattahoo¬
chee, between Main Street & Morgan Avenue, 27 Apr.
1974, Gholson 809 (FLAS); Apalachicola River, Flat
Creek Boat Landing, 13 Apr. 1982, Gholson 9533 & Man-
hart (f I . AS), 13 Apr. 1982, Manhart 407 A Gholson
(DOV, MICH); Chattahoochee, along Apalachicola River
on River Landing Rd., 8 May 1995, MacDonald 8645
(DOV, MICH, YDB. herb. Bryson); W of Chattahoochee,
overlooking Apalachicola River, S of hwy. 90, 6 Apr.
2000, Bryson 17850 A Usnick (DOV, MICH, MO. USCH.
V DB. V I’l. WIN, hcrl). Bryson); Liberty Co., Torreya State
Park, along Apalachicola River, 8 Apr. 1972, Godfrey

1328 (1SU). Georgia: Baker Co.. W bank of Flint River,
It hauway Plantation, 4 Apr. 1986. Gholson 11593 (FLAS);
Chattahoochee Co., ca. 9 mi. W of Cusseta, along Oswi-
chee Creek. 23 May 1997, Naczi 6135 (DOV); ca. 10.5
mi. NW ol Cusseta, along Upatoi Creek, 24 May 1997,
Naczi 6148 (DOV); Decatur Co., I mi. N of Chattahoochee
(Florida), bluff along Flint River, 14 Apr. 1947, Thorne
3100 et al. (CU, GA, GEO, GH), 22 Mar. 1949, Thorne
9080 A Muenscher (GEO, MICH, MO, PENN, US); Lake
Seminole, East Bank Public Use Area. 13 Apr. 1982,
Manhart 401 (DOV, GA, GH, MICH, MO, NCU, NY,
PENN, USCH, Y DB. W IN, herb. Bryson), 20 Apr. 1985,
Gholson 11283 (FLAS), 12 May 1986, Naczi 1072 A- Ghol¬
son (DOV, MICH). 18 May 1995, Naczi 4710 (DOV,
MICH); Early Co.. W' ol Saffold railroad station, 26 Mar.
1949, Thorne 9207 A Muenscher (F); ca. 1.5 air mi. NW
of Hilton, along Chattahoochee River, 18 Mav 1995 , Naczi
4715 (DOV, MICH).

Carex krauana, a New Species of Carex sect.

Lax i FLORAE

Carex sect. Laxiflorae is a group of about 25 spe¬
cies endemic to eastern North America, western
North America, Central America, and Furasia
(Naczi, 1992). All but 2 of tin1 16 species that occur
in the Western Hemisphere inhabit mesic decidu¬
ous forests of the southeastern United States, the
center of diversity for the section.

Morphologic apomorphies that diagnose Carex
sect. Laxiflorae are culms with acute angles and
culms with epidermal cells conspicuously larger
than underlying cells (Naczi. 1992). Additional
morphologic features that characterize the species,
but are probably not apomorphic of section Ixixi/lo-
rae. include long-sheathing proximal bracts, usu¬
ally unisexual spikes with only the terminal one
staminate, glabrous perigynia that are obtusely tri¬
angular in cross section and have more or less en¬
tire orifices, and perigynia usually with 25 to 39
nerves (sclerenehyma traces) that are raised above
the surface in living and dried material.

Past authors frequently placed members of Carex

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2002

Naczi et al.

Carex from North America

519

sect. Laxiflorae in the same section as members of
Carex sect. Careyanae Tuckerman ex Ktikenthal
(e.g., Kiikenthal, 1909; Mackenzie, 1935; Fernald,
1950; Bryson, 1980; Manhart, 1986; Gleason &
Cronquist, 1991). Bryson (1980) and Manhart
(1986), based on morphologic and biochemical ev¬
idence, respectively, found section Laxiflorae s.l. to
consist of two subgroups that correspond to section
Careyanae and section Laxiflorae s. str. Phyloge¬
netic analysis of morphologic data reveals that Car¬
ex sect. Laxiflorae s. str. and section Careyanae are
not each other's sister groups, however, and uniting
them would create a biphyletic group (Naczi, 1992).
Analyses of DNA sequences also support treating

Carex sect. Laxiflorae and section Careyanae as
separate sections (Starr et al., 1999). For these rea¬
sons, we circumscribe Carex sect. Laxiflorae in the
strict sense to exclude members of section Carey¬
anae.

In the course of their ongoing research on the
systematics of Carex sect. Laxiflorae, Naczi and
Bryson have detected a previously undescribed
species in the section. This new species, here
named Carex kraliana, is widespread and frequent
throughout much of the southeastern United Stales.
In the following key, Carex kraliana is distin¬
guished from all other members of section Laxiflo¬
rae that occur in the Western Hemisphere.

C. leptonei

Key to Species ok Carex sect. Laxiflorae Native to the Western Hemisphere

Complete, ample specimens bearing mature perigynia are necessary for identification. Magnification of 10X and
bright illumination should be used when examining shoot liases for presence or absence of purplish coloration, since
the coloration is often limited to small areas at the very bases of the plants. When measuring perigynium width, care
should be taken to measure a perigynium at its widest point, since one face of a perigynium is usually slightly wider
than the other two. When measuring length of the peduncles of lateral spikes, include the portion of the peduncle
enclosed in the bract sheath.

la. Bract blades of distal lateral spikes lanceolate or narrowly lanceolate, wider than spikes anil concealing them
when viewed from abaxial surface, widest bract blade (per plant) of distal-most lateral spike (3.0— )3. 2— 8.3
mm wide.

2a. Widest leaf or bract blade 1 ,3-3.8(-5.0) cm wide; pistillate scales from proximal portions ol spikes

awnless (sometimes mucronate), with truncate bodies; perigynia (1.6-)1.7-1 .9 mm wide . . . 6. albursina
2b. Widest leaf or bract blade 0.5-1. 1 cm wide; pistillate scales from proximal portions of spikes awned,

with acute bodies; perigynia 1 .3—1 ,7(— 1 .8) mm wide . . C. kraliana sp. nov.

lb. Bract blades of distal lateral spikes linear, narrower than spikes and not concealing them when viewed from
abaxial surface, widest bract blade (per plant) ol distal-most lateral spike 0.5-3.4 mm wide.

3a. Perigynia 8- to 18-nerved .

3b. Perigynia (22)25- to 32-nerved.

4a. Shoot bases purplish, with purplish coloration ranging from slight tingeing of brown background
in basal 5 mm of shoots to strong staining that obscures brown background and extends 14 cm
high.

5a. Perigynia 2.4— 3.3 mm long.

6a. Perigynia closely overlapping, 1.8— 2.7 times as long as wide, with beaks 0.2— 0.8 nun
long; internode between proximal-most scales in proximal-most spike 1.1— 8)

mm long; longest lateral spike I 1-21 mm long; angles of bract sheaths denticulate

. C. gracilescens

6b. Perigynia loosely overlapping or separate, 1.7— 2.1 times as long as wide, with beaks
0. 1-0.3 mm long; internode between proximal-most scales in proximal-most spike 3.3-
14 mm long; longest lateral spike (15— )19— 33 mm long; angles of bract sheaths entire

or minutely papillate . C- ormostachya

5b. Perigynia 3.4— 4.5 mm long.

7a. Terminal spike exceeding bract blade of distal-most lateral spike, longest peduncle
(per plant) of terminal spike (31-)46-157 mm long; vegetative shoots with widest leal
blade 1.9-2. 8 times as wide as widest leaf or bract blade of reproductive shoots; blades
of overwintered leaves (often dying or dead at time ol collection and thus partially or
completely brown) with abaxial surfaces densely and minutely papillate . . C. purpurifera

7b. Terminal spike usually exceeded by bract blade of distal-most lateral spike, longest
peduncle (per plant) of terminal spike 1 2— 32(— 53) mm long; vegetative shoots with
widest leaf blade 1 . 1 - 1 .8(-2.4) times as wide as w idest leaf or bract blade of repro¬
ductive shoots; blades of overwintered leaves (often dying or dead at time of collection
and thus partially or completely brown) with abaxial surfaces smooth . . . . C. manhartii
4b. Shoot bases brownish or whitish, with purplish coloration completely absent.

8a. Perigynia closely overlapping, ratio of length of longest lateral spike (in mm): number of

perigynia = 0.8-1. 7.

9a. Perigynia 2.5-3.8(-4.1) mm long, 1 ,5-1.9(-2.2) times as long as achene bodies; peri¬
gynium beaks 0.2— 0.6 mm long.

10a. Most culms with 2 distal lateral spikes overlapping (occasionally most culms

520

Novon

91).

with distal lateral spikes separate, and only rarely with 3 to 4 distal lateral spikes

overlapping) . . . . (; blanda

10b. Most culms with 3 to 4 distal lateral spikes overlapping . C. conge stiflora

I erigynia (3.3— )3.9— 5.4 mm long, 1.9— 2.. 1 times as long as achene bodies; perigyniuin
beaks 0.5— 1.7 mm long.

1 la. Distal lateral spikes overlapping; terminal spike exceeded by distal-most lateral
spike or the 2 spikes subequal in height; perigynia ascending; most culms with

peduncle ol proximal-most spike arising in distal half of culm . C. crebriflora

1 I h. Distal lateral spikes separate; terminal spike clearly exceeding distal-most lateral
spike; perigynia spreading; most culms with peduncle of proximal-most spike
arising in proximal third of culm.

12a. Densely caespitose, with short rhizomes . C. styloflexa

12b. Loosely caespitose, with long rhizomes . C. chapmanii

Oh. Perigynia loosely overlapping or separate, ratio of length of longest lateral spike (in mm):
number of perigynia = 1.8— 3.4.

13a. Pistillate scales 1.9-2. 3 mm wide . hendersonii

13b. Pistillate scales 1.1 — 1.8 mm wide.

14a. Longer peduncles of proximal lateral spike
subtend; perigynia spreading.

15a. Densely caespitose, with short rhizomes . C. styloflexa

15b. Loosely caespitose, with long rhizomes . C. chapmanii

14b. Longer peduncles of proximal lateral spikes 1.4-3. 3(— 5.3) times as long as spikes
they subtend; perigynia ascending.

16a. Longest bract blade 5.0— 6.7(— 8.0) cm long; blades of overwintered leaves
(often dying or dead at time ol collection and thus partially or completely
brown) with abaxial surfaces densely and minutely papillate . . . C. radfordii
16b. Longest bract blade (4.5 — )6.5 — 1 5 cm long; blades of overwintered leaves
(often dying or dead at time of collection and thus part i al ly or completely
brown) with abaxial surfaces smooth or rarely sparsely and minutely pa¬
pillate.

17a. Achene body 1.8-2.2 mm long; perigynia (2.6-)3.2— 4.1(-4.6) mm

long; longest terminal spike 1 2— 2 4(— 34) mm long . C. laxiflora

17b. Achene body 2. 2-2. 8 mm long; perigynia (3.4— )3.9-5.1 mm long;

longest terminal spike 20-32(-36) mm long . C. striatula

4.6—14 times as long as spikes they

Larex kruliunu Naczi & Bryson, sp. nov. TYPE:
U.S.A. Mississippi: Winston Co., ca. 5 mi. NW
ol Louisville, W of Noxubee Crest Natural
Area, T16N, K12E, NW 1/4 of sect. 31, II
Apr. 2000, R. F. C. Naczi 8339, Bryson , Case,
Smith, MacDonald A: Goodlett (holotype, l)()V;
isotypes. MICH, MO, NY, WIN. herb. Bryson).
Eigure 5.

A Carice albursina laminis foliorum angustioribus,
squamis pistillatis aeutis vel aristatis, perigyniis angus¬
tioribus differt; a ceteris speeiebus sectionis Laxiflorarum
laminis bractearum lanceolatis vel anguste lanceolatis,
laminis bractearum latioribus differt.

Perennial, densely caespitose. Rhizomes 0.1— 0.8
cm long between shoots or branches of the rhi¬
zomes. Shoot bases dark brown to light brown. Re¬
productive shoots erect to spreading; culms 9.2—56
cm tall. Leaves of reproductive shoots 2 to 4; blades
L2— 21 cm X 2.6—1 1 mm, the widest per plant 5.3—

1 1 mm wide, dark green. Vegetative shoots 26-34
cm tall; leaves 4 to 6, similar to those of reproduc¬
tive shoots except blades 0.9-32 cm long; pseu¬
doculms 2.S-6.9 cm tall. Infructescences (1.6-)
2.6-18(-27) cm long, 9.3-47(-77)% of culm
height, with the distal-most 2 to 3 spikes overlap¬

ping, the internode between the distal lateral spikes
0.7—19 cm long, the internode between the proxi¬
mal spike 2.8—19 cm long; proximal-most bract
with blade 6.2—17 cm long and 19— 61% of culm
height; bract blade of distal-most lateral spike lan¬
ceolate or narrowly lanceolate, ( 1 .9-)3.4— 8.3 cm
long, the widest per plant (3.0— )3.2— 7. 1 mm wide,
wider than and concealing lateral spike when
viewed from abaxial surface, much exceeding ter¬
minal spike. Spikes 2 to 4, erect; terminal spike
5.8— 13(— 18) mm long, the longest per plant 7.6—
1 3( — 1 8) mm long, 0.9— 2.1 mm wide, the widest per
plant 1. 1-2.1 mm wide, entirely staminate, on pe¬
duncle 0.8-7. 3(— 27) mm long, usually exceeded by
distal-most lateral spike but sometimes subequal to
it; lateral spikes 5.3— 18(— 23) mm long, the longest
per plant 12— 18(— 23) mm long, 3.7— 4.6 mm wide,
entirely pistillate, (4)6- to 16-flowered, the perigyn¬
ia spirally imbricate, the internode between the
proximal-most scales in proximal-most spike 1.1—
3.2(— 3.8) mm long, perigynia closely overlapping,
with ratio of length of longest lateral spike per plant
(in mm): number of perigynia = 0.97-1. 6(- 1.9).
Staminate scales 1.9— 3.8 X 0.8— 1.5 mm, obtuse or
acute, awnless. Pistillate scales I.1-4.2 X 1 .4-1.8

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Carex from North America

521

Figure 5. Carex kraliana Naczi & Bryson. Perigynium, lateral view (left; X 20); distal portion of culm (middle; X 2);
and pistillate scale (riglit; X 20).

mm; body l.l-2.2(-2.4) mm long, ovate, acute,
usually awned when in proximal portions of spikes,
usually awnless when in distal portions ol spikes,
margins usually whitish; awn 0—2.2 mm long. Peri-
gynia 2.7— 4.2(— 5.0) mm long, 1.3— 1.7(— 1 .8) mm
wide, 1 .8-2.4(-2.9) times as long as wide, 1. 4-2.0
times as long as achene bodies, ascending, obtusely
triangular in cross section, laces convex, nerves
raised and 29 to 34, glabrous, yellow-green or pale
green or yellow-brown, obovate in outline, apex ex-
curved to beak, gradually tapered from widest point
to narrow base, abruptly or gradually tapered from
widest point to beak; beaks 0.2— 0.6(— 1 .0) mm long,
9.0—21% of perigynium length, smooth, entire, de¬
flected 30—45° from long axis of perigynium.
Aehenes 1 .9-2.5(-2.9) mm long, 1 .2— 1.6(— 1 .7) mm
wide, faces tightly enveloped by perigynia, obovoid
or oblong, obtusely trigonous, faces flat or slightly
convex, brown, basally abruptly contracted to stipe,
apically abruptly contracted to minute beak; stipe
0. 1-0.2 mm long, bent 10—30° from long axis of
achene or vertical; body 1 .8— 2.2(-2.4) mm long;
beak 0. 1-0.2 mm long, bent (10-)45-60° from long
axis of achene.

The most distinctive morphologic feature of Car¬
ex kraliana is its lanceolate or narrowly lanceolate,
relatively wide bract blades. The width of the
blades is greater than the width of the lateral
spikes. The bract blades are quite spathe-like,
since they obscure the lateral spikes when viewed
from the abaxial surface of the bracts, and much

exceed the spikes. Nearly all other members of
Carex sect. Laxiflorae have linear bract blades that
are narrower than or as wide as the lateral spikes.
Only C. albursina Sheldon has wide and spathe-
like bract blades similar to those of C. kraliana.

Carex kraliana differs from C. albursina in sev¬
eral ways. First, C. kraliana has narrower leaves
than C. albursina (widest leaf blade per plant 5.3-
II mm wide for C. kraliana vs. 13—50 mm wide
for C. albursina). Second, the pisti Hate scales of C.
kraliana are acute to awned, but truncate and awn¬
less in C. albursina. Third, the perigynia of C. kral¬
iana are narrower than those of C. albursina. Carex
kraliana has perigynia 1 .3-1 .7(-l .8) mm wide,
while those of C. albursina are (1.6— )1. 7— 1.9 mm
wide.

Carex kraliana is somewhat similar to several
other species of section Laxiflorae, particularly C.
blanda Dewey, C. congestiflora Reznicek & S. Gon¬
zalez, C. crebriflora Wiegand, C. laxiflora Lamarck,
and C. leptonervia (Fernald) Fernald. These species
share the following features with C. kraliana: brown
shoot bases, the distal-most lateral spike often over¬
lapping the terminal spike, and ascending, rela¬
tively short perigynia (perigynia often less than 4.3
mm long). Unlike C. kraliana, however, all these
other species have linear and narrow bract blades.
In C. blanda, C. congestiflora, C. crebriflora, C. lax¬
iflora, and C. leptonervia, the widest (per plant)
bract blade of the distal-most lateral spike is 0.9-
3.2 mm wide (rarely up to 4.2 mm wide in C. lax-

522

No von

i flora), whereas it is (3.0— )3. 2— 7.1 mm wide in C.
kraliana.

Additional features distinguish C. kraliana from
C. blanda, C. congest iflora, C. crebriflora, C. laxi-
jlora, and C. leptonervia. Carex kraliana has the
terminal spike exceeded by or subequal to the dis¬
tal-most lateral spike, unlike C. blanda , which has
tht‘ terminal spike usually clearly exceeding the
distal-most lateral spike. Carex kraliana has the
distal-most lateral spikes separate or no more than
2 of them overlapping, whereas C. congestiflora
usually has the 3 or 4 distal-most lateral spikes
overlapping. Also. C. kraliana has 1 to 3 lateral
spikes versus 2 to 5 for C. congestiflora. Relative
to C. crebriflora, C. kraliana has shorter pistillate
scale bodies: 1 . 1 — 2.2(— 2.4) mm long for C. kraliana
versus (2.2— )2. 5— 3.5 mm long for C. crebriflora.
Also, C. kraliana has narrower perigynia than C.
crebriflora: 1 .3 — 1.7 ( — 1 .8) mm wide (or C. kraliana
versus ( 1 .6— )1 .7— 2.2 mm wide for C. crebriflora.
Carex kraliana differs Irom C. laxiflora by its short¬
er terminal spikes: longest terminal spike per plant
7.6— 1 3( — 1 8) mm long in C. kraliana versus
(12— )19— 28(— 34) mm long in C. laxiflora. In addi¬
tion, C. kraliana has denser lateral spikes than C.
laxiflora: internode between proximal-most scales
of proximal-most spike 1 . 1 — 3.2( — 3.8) mm long in
C. kraliana versus 3. 3-8.2 mm long in C. laxiflora.
(,arex leptonervia has 8- to 18-nerved perigynia
(29- to 34-nerved in C. kraliana) and relatively
loosely overlapping perigynia: internode between
proximal-most scales of proximal-most spike 3.0-
6.9 mm long in C. leptonervia versus 1.1— 3.2
(—3.8) mm long in C. kraliana.

Probably because of its similarity to so many oth¬
er species, Carex kraliana remained undescribed
for so long. Older specimens of C. kraliana bear
identifications most often of C. blanda, C. crebriflo¬
ra, and C. laxiflora var. serrulata F. J. Hermann.
Naczi and Bryson, too, had thought the name C.
laxiflora var. serrulata applied to C. Indiana and
annotated many specimens with this misidentifiea-
tion. Recent examination of the type of C. laxiflora
var. serrulata (U.S.A. Indiana: Clark Co., ca. 1.5
mi. NW of Henryville, 25 May 1910, Dearn 6458,
holotype, CH; isotype, NY) reveals it is C. laxiflora,
however, and not C. kraliana. Because of confusion
of C. laxiflora var. serrulata with C. kraliana, re¬
ports of the former, particularly from the south¬
eastern U.S. (e.g.. Ream, 1940: Hermann, 1954,
1974; Braun, 1967; Hyatt, 1998; Reznicek & Gon-
zdlez-Elizondo, 1999), should be regarded as pos¬
sibly C. Indiana and should be re-evaluated.

Plants ol C. kraliana grow in mesic deciduous
forests, usually in loams, sandy loams, and silt

loams. Frequent associates of C. kraliana include
C. amphibola Steudel, C. basiantha, C. planispi-
cata, C. rosea Willdenow, Hexastylis arifolia, and
Podophyllum peltatum I.. Carex kraliana is fre¬
quent and widespread in the southeastern United
Stales, ranging in several physiographic provinces
from southern Maryland south to northern Florida
and west to eastern Arkansas and eastern Texas.
The paratypes are only a small fraction of the col¬
lections we have studied ol C. kraliana.

Naczi and Bryson name this species for our
friend and fellow student of Cyperaceae, Robert
Krai, in recognition ol his contributions to the
knowledge ol the flora of the southeastern United
States, particularly its great diversity ol sedges.

Paratypes. U.S.A. Alabama: Bibb Co., 6 mi. SE of
Centreville, 1 May I9B7, Bryson 5606 & krai (herb. Bry¬
son); Cleburne Co., ca. 10 mi. S of Borden Springs, 23
Apr. 1990, krai 77868 (VDB); Conecuh Co., ca. 1 mi. E
of Jay Villa. 2 May 1996, Naczi 5164 (DOV, MICH. MO,
NY, WIN, herb. Bryson); Coosa Co., ca. 3.5 mi. SW of
Unity. 14 Apr. 2000, Naczi 8874 & Case (DOV); Crenshaw
Co., ca. 2 mi. S of Uuverne, 17 Mar. 1993, Naczi 2974 <V
Welpton (DOV); Dekalb Co., ca. 0.5 mi. N of Portersville,
2 June 1997, Naczi 6422 (DOV); Elmore Co., ca. 1.8 mi.
SE of Titus, 23 Apr. 2002, Naczi 9115 (DOV, MICH,
VDB, WIN, herb. Bryson); Jackson Co., ca. 2.7 mi. N of
center ol Skyline, 22 Apr. 2002, Naczi 9049 (DOV, VDB);
Lawrence Co., ca. 8 mi. S of Ml. Hope, 21 May 1996,
Naczi 5408 & Bryson (DOV, MICH, herb. Bryson); Lee
Co., ca. 1.5 mi. N ol Smiths Station, 24 Apr. 2002, Naczi
9188 (DOV, VDB); Lowndes Co., 4.5 mi. E of Braggs, 6
Apr. 1993, kr<d 81989 (herb. Bryson); Madison Co.,
Huntsville, Monte Sano State Park, 6 May 1991, Naczi
2878 (DOV, MICH); Marion Co., N side of Hamilton, 17
May 1973, krai 50065 (herb. Bryson); Russell Co., ca.
2.5 mi. NW of Cottonton, 3 Mav 1996, Naczi 5189 (DOV,
MICH, MO, NY, IISCH. WIN, herb. Bryson); Tuscaloosa
Co., near mouth of Hurricane Creek. 28 Mar. 1911, Harper
141 (F. ILL); Winston Co., 5.5 mi. NE of Double Springs,
15 May 1992, Bryson 11652 (DOV, herb. Bryson). Ar¬
kansas: Cross Co., W of Birdeye, along S side of rte. 42,

20 May 1990, Naczi 2442 (DOV, herb. Bryson). Florida:
Gadsden Co., S side of Chattahoochee, bluffs of Apalach¬
icola River, 18 Mar. 1976, krai 57851 (DOV, MICH).
Georgia: Early Co., W of Saffold railroad station, SW
corner of county, 26 Mai. 1949, Thorne 9208 & Muenscher
(MICH); Harris Co., ca. 6 mi. W ol Mulberrv Grove, along
Mulberry Creek, 24 May 1997, Naczi 6152 (DOV); Jasper
Co., ca. 14 mi. SW ol Monticello, ca. 0.8 mi. NE of Oc-
mulgee River crossing by rte. 83, 19 May 1997, Naczi
6057 (DOV); Monroe Co., ca. 2.5 mi. SW of Russellville,
25 Apr. 2000, Naczi 8897 & Ford (DOV, herb. Bryson);
Stephens Co., ca. 3 mi. N ofToccoa, 3 May 1991, Naczi
2854 (DOV, herb. Bryson); Upson Co., ca. 1.4 mi. SSW
of Roland, 25 Apr. 2002, Naczi 9178 (DOV, VDB, herb.
Bryson). Indiana: Crawford Co., 1.7 mi. E of Birdseye,

2 1 May 1990, Naczi 2498 (DOV, herb. Bryson). Ken¬
tucky: Barren Co., ca. 3 mi. SSW of Haywood, Brigadoon
State Nature Preserve, 14 June 1997, Naczi 6459 & Heeg
(DOV); Clinton Co., ca. 3.3 mi. Vi NW ol Albany, 23 May
1998, Naczi < 187 & Ford (DOV, MICH, herb. Bryson);
Fulton Co., ca. 5 mi. SSW of Hickman. 27 June 1995,

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

523

Naczi 4853 & Reznicek (DOV, MICH); Robertson Co., ca.
4 mi. SE of Piqua, 28 May 1998, Naczi 7356 & Ford
(DOV); Rowan Co., ca. 7 mi. S of center of Morehead, I
June 1996, Naczi 5597 & Trauth (DOV, KNK. MICH,
herb. Bryson). Louisiana: Morehouse Parish, W o( Jones,

30 Apr. 1992, Thomas 128379, Bryson & Newton (VDB).
\1 aryland: Charles Co., Thomas Stone National Historic
Site (W of Rose Hill Rd., 2 mi. S of Marshalls Comer),

31 May 2(H) 1 . lx-a 2569 (DOV). Mississippi : Forrest Co.,
Ragland Hills, 11 Apr. 1991, Bryson 10640 Rosso
(DOV, herb. Bryson); Grenada Co., Camp McCain, 5 May
1995, MacDonald 8593 (DOV, herb. Bryson); Holmes Co.,
6.2 mi. E of Thornton, 14 Apr. 1992, Bryson 11223 &
Newton (DOV, herb. Bryson); Marion Co., 17.5—18 mi.
SSE of Columbia, 26 Apr. 1991, Bryson 10725 et al.
(DOV, herb. Bryson); Panola Co., 1.0 mi. W o( Pleasant
Grove, 19 May 1990, Naczi 2437 (DOV); Tishomingo Co.,
ca. 1.3 mi. S of Mingo, 17 May 1992, Bryson 11704
Warren (DOV, herb. Bryson). North Carolina: Montgom¬
ery Co., NW of Uwharrie, along West Branch of McLeans
Creek, 7 June 1990, Naczi 2540 (DOV); Rockingham Co.,
ca. 7 km W of Price, along South Mayo River, 2 May
1994, Wieboldt 8873 (DOV, VPI). Ohio: Lawrence Co., N
side of Sharp’s Creek Rd. (C-19), 2 mi. W of Symmes
Creek, 22 May 1997, McCormac 5796 (MICH). South
Carolina: Edgefield Co., ca. 8 mi. SW of Edgefield, 2
Apr. 1997, Nelson 18040 (DOV, USCH); Lancaster Co¬
ca. 2 mi. E of Taxahaw, 17 May 1996, Naczi 5319 (DOV);
York Co., ca. 4 mi. NW of Bethany, 17 May 1996, Naczi
5304 (DOV). Tennessee: Dyer Co., ca. 6 mi. NW of Dy-
ersburg, 27 June 1995, Naczi 4844 & Reznicek (DOV);
Grundy Co., ca. 10 mi. SW of Altamont, 29 May 1990,
Naczi 2517, Krai <& Raveill (DOV); Henderson Co., ca. I I
mi. W' of center of Lexington, Blue Goose, 26 June 1995,
Naczi 4798 <£: Reznicek (DOV); Jefferson Co.. Jefferson
City, Apr. 1844, Rugel s.n. (MICH); Perry Co., by Blue
Creek Rd., 4 mi. S of 1-40, 14 May 1996. Krai 85847
(DOV, MICH); Scott Co., ca. 1 mi. W of Fairview, 22 June
1993, Naczi 3195 & Reznicek (DOV, MICH). Texas: Mar¬
ion Co., E of N arm of Caddo Lake, 29 Mar. 1987, Orzell
& Bridges 4940 (MICH); Sabine Co., ca. 4.5 mi. N of
Milam, along upper reaches of Mason Creek, 12 Apr.
1988, Orzell & Bridges 6180 (MICH); Tyler Co., above B.
A. Steinhagen Lake, 4 Apr. 1989, Orzell & Bridges 8901
(MICH). Virginia: Prince William Co., just NE of Prince
William Forest Park, 21 May 1995, Lea 959 (GMUF);
Scott Co., ca. 6 mi. NE of Gate City, W of rte. 669 crossing
of Copper Creek, 21 June 1993, Naczi 3172, Reznicek &
Wieboldt (DOV, VPI); Southampton Co.. NW of Courlland,
near Davis School, along Nottoway River, 8 May 1940,
Fernald & Long 11784 (US); Sussex Co., 1.3 mi. NW ol
Fields Crossroads, 2 May 1997, Fleming 12821 (GMUF);
Westmoreland Co., Littlefields farm, end of SR 672, 22
June 1999, Dodge 1483 (GMUF).

Carex gholsonii, a New Species of Carex skct.
Granulares

Carex sect. Granulares is a group of six species
of North America and Central America (Naczi,
1992): C. atractodes F. J. Hermann, C. crawei Dew¬
ey, C. granularis Willdenow, C. microdonta Torrey
& Hooker, C. quichensis F. J. Hermann, and C.
gholsonii, described here. Four of these species (C.
crawei, C. gholsonii, C. granularis, and C. micro¬

donta) occur in flic southeastern United States, the
center of diversity of the section. Carex atractodes
occurs in Mexico, and C. quichensis ranges in Gua¬
temala. Members of this section inhabit mesic for¬
ests, moist meadows, moist prairies, and moist lake
shores.

Three apomorphies diagnose Carex sect. Gran¬
ulares: red-brown cells scattered in the epidermis
of perigynia, 25 or more perigynia per well-devel¬
oped lateral spike, and perigynia loosely envelop¬
ing the achenes (Naezi, 1992, 1997). Additional
distinctive morphologic features that characterize
the species, but are probably not apomorphic of
Carex sect. Granulares, include long-sheathing
proximal bracts, usually unisexual spikes with only
the terminal one staminate, glabrous perigynia that
are suborbicular or obtusely triangular in cross sec¬
tion and have more or less entire orifices, and peri¬
gynia usually with 25 to 39 nerves (sclerenehyma
traces) that are raised above the surface in living
and dried material.

Through their independent research on the sys-
tematics of Carex sect. Granulares, Naczi and
Cochrane have discovered a previously unde¬
scribed species of the southeastern U.S. This new'
species, here named C. gholsonii, is infrequent to
rare in its relatively small geographic range. In the
following key, C. gholsonii is distinguished from
other members of Carex sect. Granulares.

Kky to Carex sect. Granulares

la. Shoots scattered along long-creeping rhizomes;
terminal spike and distal-most lateral spike (un¬
less staminate) usually separate, longest (per
plant) peduncle ol terminal spike 2.2—15 cm
long; proximal-most spike usually arising from
proximal 30% of culm; perigynia with beaks 0.1-
0.9 mm long.

2a. Staminate scale bodies obtuse, awnless; wid¬
est leaf 1 .8-2.8(^4.4) mm wide; perigynia
2.2— 3.2 mm long, with beaks 0.1— 0.3 mm

long . C. crawei

2b. Staminate scale bodies acute, often awned;
widest leaf 2. 8-8.3 mm wide; perigynia 2.8—

4.2 mm long, with beaks 0.3— 0.9 mm long
. C. microdonta

lb. Shoots clumped on very short rhizomes; terminal
spike and distal-most lateral spike usually im¬
bricate, longest (per plant) peduncle of terminal
spike 0.2— 1 -6(— 1 4) cm long; proximal-most spike
usually arising from distal 70% of culm; peri¬
gynia with beaks 0.1— 0.3(— 0.5) mm long.

3a. Foliage usually glaucous; longest (per plant)
bract blade of distal-most lateral spike 4.1-
15.8 cm long; ligule of proximal-most bract
(3.6-)4.0-26 nun long; perigynia 2.0-
3.5(— 3.9) mm long, 1 .4—2. 2(— 2.4) times as

long as wide; densely caespitose .

. C. granularis

3b. Foliage usually green; longest (per plant)

524

Novon

bract blade of distal-most lateral spike 1.6—
4.6(— 7.1) cm long; ligule of proximal-most
bract (>.6-5.9(-6.5) mm long; perigynia 2.7—

4.0 mm long, (1.6— ) 1.9— 3.0 times as long as
wide; loosely caespitose.

4a. Longest (per plant) terminal spike 8—19
mm long; longest (per plant) peduncle
of terminal spike 0.4— 3.0 mm long . . .

. C. at r act odes

4b. Longest (per plant) terminal spike
( 1 7— ) 1 9— 38(—4 1 ) mm long; longest (per
plant) peduncle of terminal spike 3.5—

16(— 140) mm long.

5a. Widest leaf blade (5.7 — )6.4 — 1 1.3
mm wide; widest (per plant) termi¬
nal spike (2.4— )2.6— 3.4(— 4.0) mm
wide; perigynia 2.9— 4.0 mm long

. C. gholsonii sp. nov.

5b. Widest leaf blade 5.7— 6.2 mm
wide; widest (per plant) terminal
spike 2.2— 2.6 mm wide; perigynia
2.7—3. 1 mm long . C. quichensis

Carex gholsonii Naczi & Cochrane, sp. nov.
TYPE: U.S. A. Florida: Citrus Co., ea. 3 mi. S
of city of Crystal River, along S side of rte.
494, 1.4 mi. W of rte. 98, 24 Apr. 1991, R. F.
C. Naczi 2787 (holotype, I )() V; isotypes,
FI, AS, MICH. MO. NY, WIS, herb. Bryson).
Figure 6.

A Carice granulari liabitu laxe caespitoso, foliis viri-
dibus, ligulis brevioribus, laminis bractearum brevioribus,
perigyniis longioribus, perigyniis (1.6-)1.9— 3.0plo lon-
gioribus quam latioribus dilfert; a Carice atractode spiea
terminali longiore, peduneulo longiori insidente differt; a
Carice quichensi laminis foliorum latioribus, spiea termin¬
ali latiore, perigyniis longioribus differt.

Perennial, loosely caespitose. Rhizomes 0.1-4
cm long between shoots or branches of the rhi¬
zomes. Shoot bases light brown to dark brown. Re¬
productive shoots erect to spreading; culms 20—75
cm tall. Leaves of reproductive shoots 1 to 5; blades

2.3—26 cm X 2.2—11.3 mm, the widest per plaid
(5.7-)6.4-l 1.3 mm wide, green. Vegetative shoots
18—51 cm tall; leaves 3 to 6, similar to those of
reproductive shoots except blades 0.6—43 cm long;
pseudoculms 3.1— 9.0 cm tall. Infructescenees 24—
60 cm long, 51—93% ol culm height, with the dis¬
tal-most 2 to 3 spikes overlapping or rarely all
spikes separate, the internode between the distal
lateral spikes 0.9—17 cm long, the internode be¬
tween the proximal spikes 5.9—31 cm long; proxi¬
mal-most bract with blade 5.0—27 cm long and 17—
89% of culm height, ligule 0.6— 5. 9(— 6.5) mm long;
bract blade of distal-most lateral spike linear, 0.9—
4.6(— 7. 1 ) cm long, the longest per plant 1. 6-4.6
(—7.1) cm long, the widest per plant 1 .2— 2.4 mm
wide, not concealing lateral spike when vieweil
from abaxial surface, greatly exceeded by terminal

spike to slightly exceeding terminal spike. Spikes
4 to 6. erect; terminal spike 8.8— 34(— 41) mm long,
the longest per plant (17 — ) 1 9 — 34(-41) mm long,
1. 7-4.0 mm wide, the widest per plant (2.4— )2.6—
3.4(— 4.0) mm wide, entirely staminate, overlapping
and exceeding distal-most lateral spike or rarely
separate from distal-most lateral spike; peduncle of
terminal spike 1.4— 16(— 140) mm long, the longest
per plant 3.5— 16(— 140) mm long; lateral spikes

5.3— 25 X 4.2— 6.6 mm, the longest per plant 14—
25 mm long, entirely pistillate, 4- to 39-flowered,
the perigynia spirally imbricate, the internode be¬
tween the proximal-most scales in proximal-most
spike 0.9-2. 4 mm long, perigynia closely overlap¬
ping, with ratio of length of longest lateral spike
per plant (in mm) : number of perigynia = 0.58-
1.1. Staminate scales 3.7— 5.6 X 1.0— 1.7 mm, usu¬
ally acute but occasionally obtuse, awnless. Pistil¬
late scales (2.0— )2.2— 2.5( — 3.1) mm long, 1.0— 1.4
mm w ide; body 1.4— 2.2 mm long, ovate, acute and
awnless or short-awned, margins whitish to ferru¬
ginous and often reddish-streaked and speckled;
awn 0—1.0 mm long. Perigynia 2.9-4. 0 mm long,

1.3— 1.8(— 2.1) mm w ide, (1 .6— ) 1 .9— 3.0 times as long
as wide, 1.8— 2.0 times as long as aehene bodies,
ascending, suborbicular to obtusely triangular in
cross section (often flattened during specimen prep¬
aration). faces convex, nerves raised and 26 to 32,
glabrous, olive to brownish green, narrowly ovate to
ovate in outline, gradually tapered from widest
point to broad base, abruptly tapered from widest
point to exeurved or straight beak; beaks 0.1— 0.3
(-0.5) mm long, 3.0—17% of perigynium length,
smooth, entire, bent 0-30° from long axis of peri¬
gynium. Achenes ( 1 .9 — )2.2 — 2.7 mm long, 1 .2-1.6
mm w ide, faces loosely enveloped by perigynia, ob-
ovoid, obtusely trigonous, faces slightly concave or
flat, brown, basally abruptly contracted to stipe,
apically abruptly contracted to minute beak; stipe
0.2— 0.4 mm long, bent about 30° from long axis of
aehene; body 1 .7-2.0 mm long; beak 0.14). 5 mm
long, bent (10— )30— 90° from long axis of aehene or
rarely vertical.

Vt it h its caespitose habit, distal-most lateral
spike usually overlapping the terminal spike, ter¬
minal spikes on relatively short peduncles, and rel¬
atively short-beaked perigynia, Carex gholsonii is
most similar to C. atractodes, C. granularis, and C.
quichensis. In fact, most specimens of C. gholsonii
were originally identified as C. granularis. As in¬
dicated in the key, C. gholsonii differs from C.
granularis by having a loosely caespitose habit,
green foliage, shorter ligules, shorter bract blades,
longer perigynia, and perigynia with a higher

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

525

\\

\

\

Figure 6. Carex gholsonii Naczi & Cochrane. — A. Habit. — B. Distal portion of culm. — C. Ligule of proximal-most
bract, with (left) and without (right) included culm and peduncle of lateral spike. — I). Perigynium, lateral view (left)
and adaxial view (right). — E. Pistillate scales. — I . Ac hone, whole (left), cross sectioned (middle), and showing apex,
section, and base (right).

526

Novon

length : width ratio. Relative to C. atractodes, C.
gholsonii has longer terminal spikes and longer pe¬
duncles of the terminal spikes. Carex gholsonii dif¬
fers from C. quichensis by having wider leaf blades,
wider terminal spikes, and longer perigynia.

The number and arrangement of chromosomes in
two plants of Carex gholsonii at metaphase 1 are t6
II + I IV (methods as in Naezi, 1999b). One plant
was from Nassau County. Florida ( Naezi 2742 ,
DO V, MICH; greenhouse-grown voucher originally
from same population as Naezi 2269, cited below),
and the second was from Wakulla County, Florida
( Naezi 2741, DO V. MICH; greenhouse-grown
voucher originally from same population as Naezi
2383, cited below). The chromosome number of C.
gholsonii overlaps that of C. granularis (Naezi,
1999b). In addition, C. granularis frequently has 1
to 3 quadrivalents per cell at metaphase I, like C.
gholsonii. The chromosome numbers of C. atrac¬
todes and C. quichensis are unknown.

Carex gholsonii ranges from southeastern North
Carolina south to central peninsular Florida and
west to southeasternmost Alabama. It occurs only
on the Coastal Plain. It is quite rare in North Car¬
olina, South Carolina, Georgia, and Alabama. Only
in Florida is it relatively frequent, but even there
it is local. The paratypes cited below are most of
the specimens examined of C. gholsonii. No doubt
its low frequency is due to its requirement for a
specialized habitat, calcareous mucks and sandy
loams. In these substrates, it grows along streams,
in swampy forests, in hammocks, on floodplains,
and in open, grassy areas such as roadsides. Fre¬
quent associates of C. gholsonii include C. bromo-
ides Willdenow, C. chapmanii Steudel, C. godfreyi
Naezi, C. leptalea Wahlenberg, Rhynchospora mil-
iacea (Lamarck) A. Gray, Ruellia caroliniensis , Sa¬
ha l minor, and S. palmetto.

Although Carex granularis is present in the
southeastern United States, it is rare within the geo¬
graphic range of C. gholsonii. No syntopie occur¬
rences of these two species are known. Carex ghol¬
sonii is allopatric with the Central American C.
atractodes and C. quichensis.

Naezi and Cochrane name this species for Angus
K. Gholson, Jr., in gratitude for his assistance with
fieldwork (including showing Naezi the first plants
he saw of C. gholsonii), loans of specimens, and
avid documentation of the flora of the southeastern
United States. It is particularly appropriate that this
species be named for Gholson since in the field it
so often accompanies C. godfreyi, the namesake of
Mr. Gholson s longtime friend and field companion.

Paratypes. U.S.A. Alabama: Houston Co., ca. 2 mi.

S of Columbia, near George Andrews Lock & Dam, 3 May
1997, MacDonald 10309 (I)()V, FSU, MICH, UNA, VDB,
WIS, herb. Bryson). Florida: [no locality beyond state],
1842—1849, Rugel 239 (l S); [Alachua] Co., Gainesville,
12 Apr. 1897, Crawford s.n. (PH); Brevard Co., Merritt
Island National Wildlife Refuge, ca. 3 mi. N of Vehicle
Assembly Bldg., 16 Apr. 1994, Hyatt 5302 (DOV, MICH);
Citrus Co., Chassahowitzka, along Chassahowitzka River
just W of county park, 24 Apr. 1991, Naezi 2792 (DOV,
MICH); Columbia Co., 3 mi. W of Lula, 2 May 1965,
McDaniel 0035 (LI, AS, LSU, MO, NY); Dixie Co., ca. 2.5
mi. NF, of Jena, 24 Apr. 1991, Naezi 2794 (DOV, FI, AS,
GA. GH, MICH. MO. NCU, NY, USCII. VDB, VP1. WIN,
WIS, herb. Bryson); Gadsden Co., Chattahoochee, Lincoln
Drive, 18 Apr. 1974, Gholson 1030 (FI, AS); Hillsborough
Co., along Hillsborough River, next to Morris Bridge Rd.,
19 Apr. 1978, Arena 754 & Wunderlin (LISF); Jefferson
Co., W of Aucilla River, along S side of rte. 98, 13 May
1986, Naezi 1 100 & Godfrey (DOV); Lake Co., Astor Park.
0.2 mi. F, of rles. 40 & 445A junction, 22 Apr. 1991,
Naezi 2732 (DOV, Ft.AS, FSU, GH. MICH, MO, NY,
TENN, UNA, USCH, VDB, herb. Bryson); Leon Co., W
of Tallahassee, near Ocklockonee River, 5 Apr. 1925,
Small 1 1009 A Wherry (NY); ca. 0.5 mi. S of Ocklocko¬
nee, 12 Apr. 1925, Harper 10 (GH, NY. PH, US); ca. 6
mi. W of Tallahassee, 5 May 1960, Godfrey 59510 (FI, AS,
FSU, NCU, NY, SMU, US, USF); Levy Co., ca. 1.2 mi. W
of Otter Creek, along rte. 24, 9 Apr. 1977. Gholson 0031
A Godfrey (FLAS); Waccasassa Bay State Preserve, near
Jack’s Creek. 4 Mar. 1997, Abbott ' 10029 (FLAS, DOV);
Nassau Co., 1.8 mi. N of Callahan, along F side of rte. 1,

4 May 1990, Naezi 2309 (DOV, MICH); Polk Co., E of
rte. 60 crossing of Alafia River, 26 Mar. 1961, Ixikela
23900 (FLAS, GH, MIN, US, USF); St. Johns Co., N edge
ol St. Augustine, between rte. 1 and railroad, 15 Apr.
1982, Gorrell 53034 A Popenoe (NY); Taylor Co., ca. 2
mi. W of Hampton Springs, near Fenholloway River, low
hammock, 29 Mar. 1910. Harper 02 (CM. GH, NY); vi¬
cinity of Aucilla River, along rte. 98, 21 Apr. 1979, Ghol¬
son 7661 & Godfrey (FLAS); Volusia Co., Daytona, 14
Mar. 1904, Deam 1305 (MICH); High Trail fish camp, I I
Apr. 1978, Thorne s.n. (USF); Wakulla Co., St. Marks
Wildlife Refuge, 24 Apr. 1971. Godfrey 70197 (FSU. GH.
KNK, MSC. NY, PH, USF); ca. 0.7 mi. N of Newport, just

5 of Newport Spring, 6 May 1990, Naezi 2333 (DOV,
MICH). Georgia: Early Co., along Kirkland Creek, near
Howard’s Mill. I I Apr. 1938, Harper 3032 (F. GH, MICH,
MO, NY, PH, US); Jefferson Co., ca. 10 mi. SE of Lou¬
isville, 8 Apr. 1904, Harper 2104 (F. GH, MO. NY. US);
Lee Co., Fowlton Creek near Armena, 20 Mar. 1949, Thor¬
ne 9026 & Muenscher (GA). North Carolina: Jones Co.,
6.5 mi. E of Pollocksville, Island Creek, 20 Apr. 1952,
Godfrey 52245 & Radford (NY, TENN); New Hanover Co.,
Wilmington, Delgado, 21 Apr. 1923, Churchill 132 (GH);
Pender Co., 7—13 June 1899, Ashe s.n. (NCU); ca. 11.5
mi. NNW of Holly Ridge, ca. 0.3 mi. W of Onslow Co.
line, 20 May 1992, Reznicek 3943 & Reznicek (DOV,
MICH); ca. I mi. W of Onslow Co. line, 20 May 1992,
Reznicek 3952 A Reznicek (MICH). South Carolina: Aik¬
en Co., Savannah River Site, along Tinker Creek above
confluence with Upper Three Runs, 17 May 1993, Antieau
135 (USCII); Mlendale Co.. Watcheall Creek. W side ol
SC 3, 13 Apr. 1996, Nelson 17099 & Pittman (DOV,
MICH, USCH, VDB); Horry Co., 3 mi. N of Conway, 21
Apr. 1932, Weatherby 10431 A Grlscom (NY); Orangeburg
Co., Santee State Park, near Chapel Hill Baptist Church
NE of rte. 6, I f June 1988, Hill 19574 (CLEMS, MICH).

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

527

Carex in firminervia, a New Species or- Carex
SECT. DeWEYANAE

Carex sect. Deweyanae is a group of six to eight
species of North America and eastern Asia. Most
of the species occur in western North America,
where as many as five species are sympatric in por¬
tions of British Columbia, Montana, and Washing¬
ton. Members of Carex sect. Deweyanae usually in¬
habit mesic to wet-mesic woodlands.

Plants belonging to Carex sect. Deweyanae are
lax sedges with setaceous proximal bracts, spikes
that are usually gynecandrous or pistillate, distig-
matic pistillate flowers, and spongy-based, beaked,
thick-margined, non-puncticulate perigynia. The
section contains C. bolanderi Olney, C. bromoides
Willdenow subsp. bromoides, C. bromoides subs]).
montana Naczi, C. deweyana Schweinitz var. driv¬
er aim. C. deweyana var. collectanea Fernald, C.
leptopoda Mackenzie, C. senanensis Ohwi, and C.
infirminervia, described here. Reznicek and Ball
(1980) referred C. laeviculmis Meinshausen to sec¬
tion Deweyanae. It and the very similar (possibly
nonspecific) C. kreczetoviczii T. V. Egorova may be¬
long to another section since they have shorter
spikes, perigynia with lower length: width ratios,
shorter perigynium beaks, and perigynia that are

more spreading than is typical for section Dewey¬
anae. All of the members of Carex sect. Deweyanae
are native to North America, except C. kreczetoviczii
(northeastern Russia) and C. senanensis (Japan).

Taxonomy within Carex sect. Deweyanae is con¬
troversial, particularly for C. deweyana and its
western North American allies. In this group, some
authors recognize as many as three species, C. bo¬
landeri, C. deweyana, and C. leptopoda (Mackenzie,
1931; Hermann, 1970; Scoggan, 1978). Most recent
authors, however, lump the three species, recogniz¬
ing only C. deweyana with no infraspecific taxa
(e.g., Hitchcock et al., 1969; Cronquist et al., 1977;
Taylor, 1983; Klinka et al., 1989; Hurd et al.,
1998). A few authors follow an intermediate course,
treating C. leptopoda as C. deweyana subsp. lepto¬
poda (Mackenzie) Calder & R. L. Taylor (Calder &
Taylor, 1965, 1968; Mastrogiuseppe, 1993).

Detection of a new species of Carex sect. Dew¬
eyanae from western North America results from
ongoing revisionary studies of the section by Naczi.
Bast failure to recognize this new species probably
contributed to confusion among the species in the
section. The new species, here named C. infirmi¬
nervia, is distinguished from other members of the
section in the following key.

Key to Carex sect. Deweyanae

I his key works best with complete and ample spec imens hearing mature perigynia; immature or incomplete speci¬
mens are difficult or impossible to identify. Since perigynia from the midregion of the spikes are most characteristic of
the taxa, these perigynia should he studied in preference to perigynia from the proximal or distal portions of spikes.

la. Longest (per plant) proximal spike 5.5— 9.8 mm long; perigynium beaks 0.4— 1 .1(— 1.3) mm long (beaks mea¬
sured from aehene apex to perigynium apex); widest leaf blade 1 .3— 2.0(-2.3) mm wide.

2a. Infruetescences (16— )28— 58(— 87) mm long . C. laeviculmis

2b. Infruetescences 15—25 mm long . C. kreczetoviczii

lb. Longest (per plant) proximal spike 8.7—27 mm long; perigynium beaks 0. 9-2.8 mm long (beaks measured
from aehene apex to perigynium apex); widest leaf blade 1.3— 5.9 mm wide.

3a. Perigynia (3.3— )4. 1-6.7 times as long as wide, abaxially with (3)4 to 8 complete nerves (those that extend
unbroken from perigynium base to base of perigynium beak); achenes 1.9— 2.9 times as long as wide;
w idest leaf blade 1.3-4. 4 mm wide.

4a. Culms 0.5— 1.1 mm w ide at mid-height; w idest leaf blade 1.3— 2. 9(— 3.1) mm wide; perigynium beaks

30-42% of perigynium length . C. bromoides subsp. bromoides

4b. Culms 1.0— 1.6 mm wide at mid-height; w idest leaf blade (2.3— )2. 8— 4.4 mm wide; perigynium beaks

36-48% of perigynium length . C. bromoides subsp. montana

3b. Perigynia 2.3— 3.9(-4.2) times as long as wide, abaxially with 0 to 7 complete nerves (those that extend
unbroken from perigynium base to base of perigynium beak); achenes 1.2-1. 8 times as long as wide;
widest leaf blade (2.2— )2. 4— 5.9 mm wide.

5a. Ligule of distal leaf on culm 0.9-2. 2 mm long; pistillate scale bodies 2. 8-4. 2 mm long; ac henes
(1 .8—) 1 .9— 2.2 mm long; longest infructeseenc-e with 2 to 5 spikes; longest proximal spike with 5—12
perigynia (including undeveloped or aborted ones).

6a. Longest infructescence 35—56 turn long, longest internode between proximal spikes (11 — ) 1 3—34

mm; longest proximal bract 15—49 mm long . C. deweyana var. deweyana

6b. Longest infructescence 12—32 mm long, longest internode between proximal spikes 6-10 mm;

longest proximal bract 9 — 1 2 ( — 1 8) mm long . C. deweyana var. collectanea

5b. Ligule of distal leaf on culm (2. l-).3. 1-8. 1 mm long; pistillate scale bodies 2. l-3.3(-3.8) mm long;
achenes 1.4— 1.9(— 2.2) mm long; longest inlructescence with (4)5 to 9 spikes; longest proximal spike
with (9)12 to 37 perigynia (including undeveloped or aborted ones).

7a. Perigynium beaks 0.9— 1.5(— 1.7) mm long, 28—38% of perigynium length; pistillate scale bodies
2. 1 — 2.9( — 3. 1 ) mm long . C. leptopoda

528

No von

71). Perigynium beaks (1 ,4-)l .6-2.7 mm long, 38-50% of perigynium length; pistillate scale bodies

(2.3— )2. 7— 3.8 mm long.

8a. Perigynium apices with teeth 0-0.2(— 0.4) mm long, teeth 0—8% of perigynium length (avoid
perigynia with teeth lengthened by tearing of sinus between teeth); longest infructescence
with (4)5 to 6(7) spike's; culm at mid-height papillose (best seen with 20X magnification on
young culms) . C. infirminervia sp. nov.

8b. Perigynium apices with teeth (0.2-)0.3— 0.6(— 1.0) mm long, teeth (6— )9— 20% of perigynium
length (avoid perigynia with teeth lengthened by tearing of sinus between teeth); longest
infructescence with (5)6 to 9 spikes; culm at mid-height smooth, serrulate, or scabridulous
(check with 20X magnification).

9a. Perigynia abaxially with (2)3 to 7 complete nerves; perigynium margins denticulate for

34—51% of perigynium length . C. bolanderi

9b. Perigynia abaxially with 0 to 2(3) complete nerves; perigynium margins denticulate for

49—64% of perigynium length . C. senanensis

Carex infirminervia Naczi, sp. nov. TYPE: [Can¬
ada. British Columbia]: Nelson, Silver King
Mine Rd., roadside at about 5000 It., 31 July
1943, ./■ 1C East hum 10949 (holotype, UBC;
isotype, DAO). Figure 7.

A Carice bolandero et Carice senanensi culmis papil-
losis. infniotescentiis spieis paucioribus instructis, apici-
bus perigyniorum edentatis vel brevidentatis differt.

Perennial, densely caespitose. Culms 10— 83 cm
tall, 0.8-1. 3 mm wide at mid-height, covered with
minute papillae at mid-height. Leaves with blades
1.4— 3.8 mm wide, the widest blades 2.3— 3.8 mm
wide; ligules of distal leaves on culm (2.1— )2.6— 6.8
mm long. Infructescences 27—68 mm long, with the
proximal-most 2 spikes overlapping or separate, in¬

i' igure 7. Carex infirminervia Naczi. Perigynium, abaxial
view (X 20).

ternode between proximal-most spikes 7.3—28 mm
long; proximal-most bract I 1-61 mm long. Spikes
(4)5 to 6(7), usually gynecandrous but also some¬
times pistillate or staminate; terminal spike 9.6-16
X 3.4— 6.2 mm, usually with 1 to 3 staminate and
14 to 21 pistillate flowers, sessile or on peduncle
up to 1.8 mm long; proximal-most spike 9.7—18 X
3. 7-6.6 mm, the longest per plant 12-18 mm long,
usually with 1 to 3 staminate and 11 to 22 pistillate
(lowers, the longest per plant with 12 to 22 pistillate
(lowers. Pistillate scales 2.9— 4.4 mm long, acumi¬
nate to short-awned with awn up to l.l mm long;
bodies 2.9— 3.8 X 1.4— 1.8 mm, whitish to easta-
neous except for green midrib. Staminate scales
3.1— 4.3 X 1.0— 1.5 mm, acute to short-awned, mar¬
gins whitish to castaneous. Perigynia 3. 7-5.3 mm
long, 1.1— 1.3(— 1.4) mm wide, erect to appressed-
erect, narrowly lanceolate in outline, 3.2— 3. 9(— 4.2)
times as long as wide, green to brown, unnerved
adaxially, unnerved or weakly I- to 3(4)-nerved
abaxially, gradually narrowed to a beak; beaks 1.5—
2.2 mm long, 39-49% of total perigynium length,
denticulate on margins for distal 49—60% of length
of perigynium, the apex toothless or bidentulate
with teeth 0. 1— 0.2(— 0.4) mm long, teeth 0—8% of
perigynium length. Achenes 1.6—2. 2 X l.l— 1.3
mm, 1.5— 1.8 times as long as wide. Stigmas 1.8—
2.5 mm long. Anthers 1.3— 1.8 mm long.

Carex infirminervia most closely resembles C.
bolanderi and C. senanensis. Carex infirminervia
possesses relatively long ligules (s 2.1 mm long),
relatively long bodies of the pistillate scales (> 2.9
mm long), perigynia 3.2— 3.9(— 4.2) times as long as
wide, and perigynium beaks occupying a relatively
high percentage of the perigynium length (^ 39%).
Only C. bolanderi and C. senanensis also have this
combination of features; other species in section
Deweyanae have shorter ligules (C. deweya.no),
shorter pistillate scale bodies (C. leptopoda ), peri¬
gynia with either a lower ( C . leptopoda) or a higher
(C. bromoides) length: width ratio, and perigynium

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

529

beaks occupying a lower percentage of the perigyn-
ium length ( (.. deweyana, C. leptopoda).

Carex infirminervia differs from both C. bolanderi
and C. senanensis in at least three ways. First, at
mid-height, the culms of C. infirminervia are cov¬
ered with minute papillae (best seen with 20X
magnification), whereas those of C. bolanderi and
C. senanensis are smooth, serrulate, or seabridu-
lous. The papillae on culms of C. infirminervia have
their long axes perpendicular to the culm surface,
unlike the serrulations and minute, antrorse teeth
that are sometimes present in the mid-region ol die
culms of C. bolanderi and C. senanensis. The pa¬
pillae are delicate, and many wear off the culms
with age. By the time most perigynia have shed
from a culm of C. infirminervia, the papillae may
be patchily distributed or nearly absent. Thus,
when studying the morphology of culms of C. bo¬
landeri, C. infirminervia, and C. senanensis, one
should examine more than one culm per specimen
and include young culms in this examination. With¬
in Carex sect. Deweyanae, papillose culms are not
unique to C. infirminervia. Regularly, C. leptopoda
has papillose culms, and C. deweyana occasionally
has them. However, within the group of C. bolan¬
deri, C. infirminervia, and C. senanensis, papillose
culms are restricted to C. infirminervia.

A second feature that distinguishes C. infirmi¬
nervia from C. bolanderi and C. senanensis is its
fewer spikes per infructescence. The longest in-
fructescence per plant in C. infirminervia bears (4)5
to 6(7) spikes, whereas C. bolanderi has (5)6 to 9
spikes and C. senanensis has 7 to 9 spikes. A third
diagnostic feature of C. infirminervia is the tooth¬
less or relatively short-toothed perigynium apex.
Whereas apices of perigynia of C. infirminervia
have teeth 0— 0.2(— 0.4) mm long, those of C. bolan¬
deri and C. senanensis bear teeth (0.2— )(). 3—
0.6(— 1.0) mm long.

Additional features distinguish C. infirminervia
and C. bolanderi. The perigynium margins of C.
infirminervia are denticulate for the distal 49—60%
of the perigynium length, whereas in C. bolanderi.
the margins are denticulate for 34— 51% of the peri¬
gynium length. Also, abaxial surfaces of perigynia
of C. infirminervia usually have fewer and fainter
nerves than those of C. bolanderi. In C. infirminer¬
via, 0 to 3(4) fine nerves extend unbroken from the
perigynium base to the base of the perigynium
beak, whereas perigynia of C. bolanderi have (2)3
to 7 nerves, which are usually relatively thick. The
epithet ‘"infirminervia” (“weak-nerved”) is appro¬
priate for the new species since its relatively few
and faint perigynium nerves distinguish it from C.

bolanderi , the species to which it is morphologically
and geographically most similar.

Carex infirminervia ranges from southwestern Al¬
berta and central British Columbia south to north¬
ern Colorado and west to central California. Por¬
tions of the geographic range of Carex infirminervia
overlap parts of the ranges of C. deweyana and C.
leptopoda Carex infirminervia is allopatric with the
eastern North American C. bromoides and the Jap¬
anese C. senanensis. Carex infirminervia is sympat-
rie with C. bolanderi over much of the ranges of
the two species. Herbarium specimens document at
least two locations for syntopy of C. infirminervia
and C. bolanderi : one in British Columbia (Nelson,
Six Mile Lakes, common on plateau, 4500 ft., 27
June 1940, Eastham 7578 |UBC] — C. infirminer¬
via, and Eastham 7577 [UBC| — C. bolanderi) and
the other in California (El Dorado Co., Strawberry
Creek, 5900 ft., 20 July 1 397. Brainerd 179 [VT] —
C. infirminervia, anti Brainerd 180 [VT] — C. bolan¬
deri). In both cases, the collectors apparently no¬
ticed a difference between the two species, since
they assigned different collection numbers to the
specimens of each. Co-occurrence of C. infirminer¬
via and C. bolanderi, with both apparently main¬
taining their distinctions, is further evidence in
support of their status as two species.

Though widespread, C. infirminervia is infre¬
quent and its populations are scattered. Judging
from herbarium labels, it usually occurs in wood¬
lands and their edges, often on slopes above
streams. A few collectors have indicated C. infir¬
minervia also grows in more open habitats, such as
grassy slopes and roadsides. Perhaps these open
habitats are near woodlands.

Most collections of C. infirminervia bear original
identificati ins of C. bolanderi, C. leptopoda, or C.
deweyana. Carex infirminervia shares some features
with C. bolanderi (discussed above), C. deweyana
(e.g., relatively long pistillate scale bodies, peri-
gynium apex toothless or short-toothed, few and
faint perigynium nerves), and C. leptopoda (e.g.,
relatively long ligules, perigynium apex toothless or
short-toothed, few and faint perigynium nerves).
Such shared characteristics probably account for
much of the conflicting classification within tin-
section. Recognition of the diagnostic features of C.
infirminervia permits precise circumscription of it
and similar species, with resultant clarification of
the taxonomy of section Deweyanae.

Paratypes. CANADA. Alberta: Lake Louise, 22 July
1904, M acorn 6 4142 (CH, NY); Carbondale River, 16
July 1944, Carmack 328 (ALI A, US); Waterloo Lakes Na¬
tional Park, above Waterton Lake, 13 July 1953, Breitung
15880 (NY. IJS); Waterton Lakes National Park. L shore

530

Novon

of Cameron Cake, I Aug. 1956, Hermann 13049 (ALTA,
US). British Columbia: Selkirk Mountains, Glacier,
4300 ft.. 10 Aug. 1897, Brainerd s.n. (VT); Selkirk Moun¬
tains, Glacier— Avalanche Mountain, 23 July 1908, Hallers
398 A" Holway (GH, 11 M); Nelson, Six Mile Lakes, 27 June
1940, Eastham 7578 (UBC); ca. I I mi. W of Revelstoke,
near Victor Lake, 3 July 1941, Hitchcock 7547 A Marlin
(WTU); Rossland, Little Shee|i Creek Trail. 5 July 1942,
Eastham 9978 (UBC); Fernie, Fairy Creek, 5 July 1947,
Eastham 15652 (DAO, UBC); ca. 9 mi. NF of Nelson,
Siteum Creek Delta. 19 June 195.3, Colder 9385 & Savile
(DAO, UBC, UC); NNK of Naramata, 1.5 road mi. SW of
Chute Lake, 8 July 1953, Calder 10242 A Savile (DAO,
DS); Manning Park, 12 mi. NW of ranger station, along
Hope— Princeton Hwy., 3100 ft., 16 July 1953, Calder
10606 A' Savile (DAO); just NW o( Azouzetla Lake, Pine
Pass, along Hart Hwy., 2900 It., 1,3 July I960, Calder
26902 A- Kukkonen (DAO); Galena, at Hill Creek, 21 June
1962, Taylor A Szczawinski 513 (UBC); Azouzetla Lake,
Hart Hwy., 650 m, 5 July 1963, Taylor A Szczawinski 6 36
(UBC); 20 km NK of Vernon, Silver Star Mountain, 5 July
1982. Botham 2948 (DAO. MICH), Botham 2962 (DAO,
MICH); ca. 8 km F of Yahk, provincial rest area on N
side of hwy. .3, 27 June 2000, Eord 00125 A Saarela
(WIN); Manning Provincial Park, trail between Strike and
Flash Lake, 29 June 2000. Eord 00139 A Saarela (DOV.
WIN). U.S.A. California: FI Dorado Co.. Strawberry
Creek, 5900 ft.. 20 July 1897, Brainerd 179 (VT); Lake
Tahoe, Emerald Bay, 21 July 1941. Rose 41334 (CAS);
Fresno Co., 2 mi. N of General Grant, National Park
Headquarters, 29 June 1940, Rose 40686 (CAS); Glenn
Co., Canyon of Snow Basin Creek, 5 Aug. 1943, Howell
19106 (CAS, US); Humboldt Co., Trinity Summit, lower
end of Brett Hole, 23 July 1935, Tracy 14162 (CAS. DS.
NCU, UC); Mariposa Co., Lake Tenaya Trail, 14 Aug.
1894, Congdon s.n. (DS); Nevada Co., between Donner
Pass and West Lake, 23 July 1943, Howell 18721 (CAS);
Placer Co., Lake Tahoe, Chambers Lodge, 25 July 1944,
Rose 44245 (CAS); Plumas Co., 8 mi. SW of Johnsville,
McRae Meadows, 28 June 1951, Rose 51029 (COLO);
Lassen Volcanic National Park, Boiling Springs Lake. 20
July 1960, Howell 35759 ((LAS); Siskiyou Co., Spirit Lake,
Marble Mountains, 3 Aug. 1939, Howell 14989 (CAS,
UC); SF end of Long Gulch Lake. Klamath National For¬
est, T39N, R9W, SF 1/4 sect. 32. 12 Aug. 1987, Tallent
901 (MICH); Trinity Co., N Fork Coffee Creek, Trinity
River, 19 Aug. 1916, Goldsmith 19a (JFPS); [Tuolumne]
Co. [originally labeled as FI Dorado Co.], Jones Hill,
1879, Shockley 605 (JFPS). Colorado: Boulder Co., Mar¬
ysville, F of Eldora, N-facing slope Tennessee Mountain,
Shickley property, 23 July 1971. Colson 71-23 (COLO);
Gunnison Co., Gunnison National Forest, West Flk Wil-
derness, along Cascade Creek. 0.25 mi. upstream from
Coal Creek confluence, 7200 It., 19 July 1994, Rondeau
94-203 A DeCoursey (COLO); Routt Co., Routt National
Forest, lower Silver Creek, ea. 8100 ft., 13 Aug. 1993,
Huff 435 A Kettler (COLO). Idaho: Bonner Co., 20 mi.
N of Sandpoint, 3.5 mi. NF of Jeru Peak, Homstead Creek
Watershed, Selkirk Mts., 14 Aug. 1969, Stickney 1954
(RM); Elmore Co., ca. 6 mi. from Atlanta on road to Fealh-
erville, 22 July 1944, Hitchcock 10203 A Muhlick (CAS,
Gil, NY, RM, UC. LIS, WTU); Latah Co., NF of Moscow,
Thatuna Hills, 12 July 1939, Baker 1428 (WTU); Valley
Co., 3 mi. NW of Payette Lake, 25 June 1946, Hitchcock
13949 A Muhlick (CAS. MO. NY. KM. UC, UTC, WTU);
12 mi. N of McCall, Brush Creek Rd., Payette National
Forest, 13 Aug. 1971. lewis 2144 (UTC); upper six mile

creek. Middle Fork Payette, Boise National Forest, 14 July
1972. lewis 2451 (UTC). Montana: [Flathead] Co.,
Nyack, 5 Aug. 1894, Williams 466 (US): Flathead Co..
Upper Whitefish Lake, above campground, Whitefish
Range, 2 July 1972, Lackschewitz 3718 (RM); Glacier Co.,
Glacier National Park, along trail from Many Glacier Hotel
to Swiftcurrent Pass, 19 July 1919 , Stand ley 1601 1 (US);
Ravalli Co., 10 mi. S of Alta, head of Beaver Creek, 2
July 1946, Hitchcock 14362 A Muhlick (CAS, NY. WTU).
Nevada: Washoe Co., Carson Range, Little Valley, 26 July
1975, Tiehm 1640 (NY. RFNO). Oregon: Baker Co., HI
mi. above Richland, Eagle Creek. 28 June 1938, Peck
19969 (CAS, WILLU); [Hood River] Co., Mount Hood,
near White River, 23 July 1928, Thompson 5006 (MO.
WTU); Klamath Co., Crater Lake National Park. Vidae
Ridge, 15 Aug. 1950, Baker 7244 (CAS); N end of Lake
of Woods. Peck 16589 (Vi 1LLU); Lane Co., Lookout Ridge,
along Lookout Ridge Rd., Andrews Exp. Area, 25 July
1968. Eranklin 772 (OSC, RM); Morrow Co., Umatilla Na¬
tional Forest, Alder Creek, along FS Road 21, 10 July
1993, Wilson 6 153 et al. (MICH); [Umatilla] Co., I.aka.
Blue Mountains, 12 June 1886. Henderson 1784 (GH);
Wallowa Co., Eagle Cap Wilderness, S of S end of road in
Lostine Canyon, 8 Aug. 1994, Wilson 7392 et al. (MICH).
I tali: [Salt Lake] Co., Alta, Wasatch Mountains, 8 Aug.
1879, Jones 1218 (MICH, NY, UTC); Salt Lake Co., Little
Cottonwood Canyon, W hite Pine Fork Trail, 13 Aug. 1983,
Arnow 6149 (NY). Washington: Asotin Co., SW of An-
atone, head of Cottonwood Creek. T7N, R44F, sect. 6, 9
July 1949. Cronquist 5944 A Jones (COLO, GH, MICH.
NY, UC, LIS. UTC, WTU); [Chelan] Co., Stehekin, June
1902, Griffiths 198 A Cotton (GH); Cascade Tunnel, 14
July 191 1 . Jones s.n. (DS); Stevens Pass. June 192}). Grant
s.n. (RM); Chelan Co., Stehekin River, below Cottonwood
Camp, 31 Aug. 1956, Woven 10300 (CAS); Columbia Co.,
Blue Mountains, I I Aug. 1897, Horner R485B524 (GH);
Pierce Co., Mount Rainier National Park, trail to Glacier
Basin, White River Campground, 2 Aug. 1977, Colson 77-
60 (COLO); Whatcom Co., S of Colonial Campground,
Thunder Creek Trail, 14 June 1991, Castaner 10723
(DOV). Wyoming: Lincoln Co., Skull Creek al junction
of Little Greys River. 28 June 1980. Tuhy 66 (NY); Park
Co., ca. 1.5 mi. N of rtes. 14, 16, and 20, along Grinnell
Creek Trail, 9 Aug. 1980, Evert 2343 (COLO, NY. RM);
along Mormon Creek, ca. 2-3 mi. N of Hwy. 14, 19 July
1985, Evert 8535 (NY); Teton Co., 1.3 mi. F of Driggs,
Idaho, 2 mi. E of Treasure Mountain Scout Camp. Teton
Forest Camp, 16 July 1956, Anderson 507 (NY. UTC).

A New Combination for a Variety of Carex
digitalis (Carex sect, Careyaixae)

Carex digitalis Willdenow is widespread in for¬
ests of eastern North America and common
throughout most of its range. Fernald (1950) rec¬
ognized two non-autonymic varieties for this spe¬
cies, C. digitalis var. asymmetrica Fernald and C.
digitalis var. macropoda Fernald. In the forthcom¬
ing treatment of Carex sect. Careyanae for the Flo¬
ra of North America, Bryson and Naczi (in press)
recognize three varieties for C. digitalis, using the
same circumscriptions as Fernald. However, the
nomenclature of one of these varieties must change.
Over half a century prior to Fernald, Bailey (1889)

Volume 12, Number 4
2002

Naczi et al.

Carex from North America

531

described C. laxiculmis var. jloridana, which is the
same taxon as C. digitalis var. asymmetrica. In or¬
der to use Bailey's prior epithet and align it with
C. digitalis, it must be transferred, as follows.

Carex digitalis var. floridaua (L. H. Bailey) Naczi
& Bryson, comb. nov. Basionym: Carex laxi-
culmis var. Jloridana L. H. Bailey. Mem. Torrey
Bot. Club 1: 47. 1889. TYPE: U.S.A. Florida:
[no additional locality information, no date].
Chapman s.n. (lectotype, designated here,
BH).

Carex digitalis var. asymmetrica Femald, Bhodora 43:
544. 1941. Syn. nov. TYPE: U.S.A. Virginia: South¬
ampton Co., NW of Applewhite Church, near Three
Creek, 8 May 1940, Fernald & Long 1 1791 (holo-
type, GH; isotypes, Gil. PH).

Carex digitalis var. Jloridana is diagnosed by rel¬
atively long perigynia (3.2— 4.2 mm long) with con¬
spicuously excurved apices. Specimens ol the two
other varieties, C. digitalis var. digitalis and C. dig¬
italis var. macropoda, have perigynia 2.4— 3.4 mm
long with barely excurved apices. Examination of
the types of C. digitalis var. asymmetrica and C.
laxiculmis var. Jloridana reveal that both have long
perigynia with excurved apices. In addition, Naczi
and Bryson discern no taxonomic differences be¬
tween these type specimens. Hence, we conclude
they are the same taxon.

Bailey cited two syntypes for his C. laxiculmis
var. Jloridana, Chapman s.n. and Curtiss s.n., both
from Florida. We choose the Chapman specimen as
the lectotype because it more closely matches the
description, and it bears mature and ample peri¬
gynia, whereas the other specimen ( Curtiss s.n.) has
immature and few perigynia.

Plants of C. digitalis var. Jloridana inhabit mesic,
well-drained (usually sandy) soils of deciduous and
deciduous-pine forests from southern Maryland
south and west to eastern Texas, mostly on the
Coastal Plain.

Acknowledgments. We thank Peter Ball and
Anton A. Reznicek for reviewing this paper and
providing many helpful comments. Victoria C. Hol¬
lowed made numerous insightful suggestions for
improving the manuscript. We also thank Frederick
W. Case. Jr., Angus K. Gholson, Jr., the late Robert
K. Godfrey, John R. MacDonald, James B. Manhart,
Anton Reznicek, and William I). Welpton for as¬
sistance with fieldwork. For sending their collec¬
tions for our study, we are grateful to .1. Richard
Abbott, Angus Gholson, Bruce A. Ford, John Mac¬
Donald, Albert B. Pittman, Christopher Eea, and
Randy Mears. Chris Eea brought our attention to

collections of Carex kraliana at CMUF and US. Su¬
san A. Reznicek drew Figures 1 to 5 and 7. Kandis
E. Elliot drewr Figure 6. The Kentucky State Nature
Preserves Commission granted permission to col¬
lect at some of its preserves. We also thank curators
of the cited herbaria for loans ol specimens or as¬
sistance during our visits. National Science Foun¬
dation Doctoral Dissertation Improvement Grant
BSR-9001260, the University of Michigan, Dela¬
ware State University, and the* Kentucky State Na¬
ture Preserv es Commission provided financial sup¬
port.

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of the genus Carex. Mem. Torrey Bot. Club. 1: 1—85.
Boott, F. 1858. Illustrations of the Genus Carex , Vol. 1.
William Pamplin, London.

Braun, E. E. 1967. The Monocotyledoneae: Cat-tails to
Orchids. Ohio State Univ. Press, Columbus.

Bryson, C. T. 1980. A Revision of the North American
Carex sect. Laxiflorae (Cyperaceae). Ph.D. Dissertation.
Mississippi State University, Mississippi Slate.

- 5 K. F. C. Naczi. In press. Carex sect. Careyanae.

In Floia of North America North of Mexico, Vol. 23.
Oxford Univ. Press, New York.

Calder, J. A. & B. E. Taylor. 1965. New taxa and nomen-
elatura changes with respect to the llora of the Queen
Charlotte Islands, British Columbia. Canad. J. Bot. 43:
1387-1400.

- (S B. L. Taylor. 1968. Flora of the Queen Char¬
lotte Is ands. Part 1: Systematics of the Vascular Plants.
Monogi. 4(1). Canada Dept. Agriculture, Research
Branch, Ottawa.

Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. E.
Reveal & P K. Holmgren. 1977. Intermountain Flora.
Vol. 6. Columbia Univ. Press, New York.

Darlington, W. 1843. Beliquiae Baldwinianae: Selections
from the Correspondence of the Eate William Baldwin.
M.D. Kimber & Sharpless, Philadelphia.

Deam, C. C. 1940. Flora of Indiana. Indiana Dept, of Con¬
servation, Indianapolis.

Fernald, M. E. 1950. Gray’s Manual ol Botany, 8th ed.
American Book, New York.

Gleason, H. A. & A. Cronquist. 1991. Manual of Vascular
Plants >f Northeastern United Sates and Adjacent Can¬
ada. 2nd ed. New York Botanical Garden, Bronx, New
York.

Hermann F. J. 1954. Addenda to North American earices.

Amer. Midi. Naturalist 51: 265-286.

- . 970. Manual of the Carices of the Rocky Moun¬
tains and Colorado Basin. Agric. Handbook 374. Forest
Service, U.S.D.A., Washington.

- . 974. Manual of the Genus Carex in Mexico and

Central America. Agric. Handbook 467, Forest Service,
U.S.D.A., Washington.

Hitchcock, C. L., A. Cronquist & M. Owtibey. 1969. Vas¬
cular Plants of the Pacific Northwest. Part 1: Vascular
Cryptogams, Gymnosperms, and Monocotyledons. Univ.
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Hurd, E. G., N. E. Shaw, J. Mastrogiuseppe, E. C. Smith-
man & S. Goodrich. 1998. field Guide to Intermountain
Sedges Gen. Tech. Bep. BMBS-GTR-10. U.S.D.A.,

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Forest Service, Rocky Mountain Research Station, Og¬
den, Utah.

Hyatt, I*. K. 1998. Arkansas Carex (Cyperaceae): A briefly
annotated list. Sida 18: 535—554.

klinka, k., V. J. krajina, A. Ceska & A. M. Scagel. 1989.
Indicator Plants of Coastal Rritish Columbia. Univ. Brit¬
ish Columbia Press, Vancouver.

kilkenthal, G. 1909. Cyperaceae— Caricoideae. In: A. En-
gler (editor). Das Pflanzenreich, IV. 20, Heft 38: 1—824.
Wilhelm Englemann, l^ip/.ig.

Mackenzie, k. k. 1931. Cyperaceae — Cariceae. N. Amer¬
ican Flora 18: 1-168.

- . 1935. Cyperaceae— Cariceae. N. American Flora

18: 169-478.

Manhart, J. R. 1986. Foliar flavonoids of the North Amer¬
ican members of Carex sect. Laxiflorae (Cyperaceae).
Bioehem. Syst. Ecol. 14: 85—90.

Mastrogiuseppe, J. 1993. Carex. Pp. I 107-1 138 in ,1. (5
Hickman (editor). The Jepson Manual: Higher Plants of
California. Univ. California Press, Berkeley.

Nacz.i, R. h. (.. 1992. Systematics of Carex sect. Griseae
(Cyperaceae). Ph.l). Dissertation, University of Michi¬
gan, Ann Arbor.

- . 1997. Phylogeny reconstruction in Carex sections

Careyanae and Granulares (Cyperaceae). Amer. J. Bot.
[Suppl.] 84: 218-219.

- . 1999a. < arex planispicata, a widespread and fre¬

quent new species ol Carex sect. Griseae (Cyperaceae)
from the eastern United States of America. J. kentucky
Acad. Sci. 60: 37-44.

- . 1999b. Chromosome numbers of some eastern

North American species of Carex and Eleocharis (Cy¬
peraceae). Contr. Univ. Michigan Herb. 22: 105-119.

Reznieek, A. A. & P. W. Ball. 1980. The taxonomy of
Carex sect. Stellulatae in North America north of Mex¬
ico. Contr. Univ. Michigan Herb. 14: 153—203.

- & S. Gonzalez-Flizondo. 1999. New species of

Carex (Cyperaceae) from Chiapas, Mexico. Contr. Univ.
Michigan Herb. 22: 121-130.

Seoggan. II. .1. 1978. The Flora of Canada. Part 2: Pteri-
dophyta, Gymnospermae, Monoeotyledoneae. Publ. in
Botany 7(2). National Museums of Canada, Ottawa.

Starr, J. R„ II. J. Bayer & B. A. Ford. 1999. The phylo¬
genetic position of Carex sect. Phyllostachys and its im¬
plications for phylogeny and subgenerie circumscription
in Carex (Cyperaceae). Amer. J. Bot. 86: 563—577.

Taylor, I. M. C. 1983. The Sedge Family (Cyperaceae).
Handbook 43. British Columbia Provincial Museum,
Victoria.

I home, R. F. 1951. Vascular plants previously unreported
f rom Georgia. Castanea 16: 29-48.

lucker, G. C. 1987. The genera (if Cyperaceae in the
southeastern United States. J. Arnold Arbor. 68: 361—
445.

Mikania oreophila (Asteraceae, Eupatorieae), a New Species from

Southern Brazil

Mara Rejane Ritter and Silvia Teresi iha Sfoggia Miotto
Depto. de Botanica — IB/UFRGS, Campus do Vale, Porto Alegre, Rio Grande do Sul,
Brazil 91509-900. mrritter@lerra.com.br

Abstract. Mikania oreophila, a new species ol
Asteraceae (Eupatorieae) from the southern Brazil¬
ian highlands, is described and illustrated. The new
species is related to M. paranensis, from which it
differs by the thinly chartaceous and somewhat ba-
sally attenuate leaves and subinvolucral bracts
placed at the base of the peduncles.

Rksumen. Se describe e ilustra Mikania oreophi¬
la, una nueva especie de Asteraceae (Eupatorieae)
de las montanas del sur de Brasil. La nueva especie
esta emparentada con 17. paranensis, de la cual se
diferencia por sus hojas finamente cartaceas y algo
atenuadas en la base y por las bracteas subinvol-
ucrales dispuestas en la base de los peduneulos.

Key words: Asteraceae, Brazil, Eupatorieae,
Mikania.

Mikania Willdenow is a genus in Asteraceae
with about 430 species distributed mainly in the
tropical regions, mostly in South America (Holmes,
1995). There are about 171 species recorded lor
Brazil (King & Robinson, 1987). As part ol a study
of the genus in southern Brazil, an undescribed
species came to our attention and is described
herein as new.

IVlikaniu oreophila M. R. Ritter & Miotto, sp. nov.
TYPE: Brazil. Rio Grande do Sul: Sao Eran-
cisco de Paula, Alpes de Sao Francisco, 18
Mar. 2000, 47. R. Ritter & S. L. de Carvalho
twite 1110 (holotype, ICN; isotypes, FLOR,
MBM, MO). Figure 1.

Suffrutices volubiles, breviter puberuli vel pubescentes;
folia opposita, sine stipulis, aliquando linea trichomatum
interpetiolari, hastala, decurrentia, superficiebus arnbabus
cum trichomatibus et glandulis sparsis, trinervibus. Cap-
ituli peduneulati, pilosi, compositi in inflorescentiis pan-
iculiformibus. Bractea subinvoluerata lineari-lanceolata
posila in base pedunculi et rare paulo superiore. Bracteae
involucratae elliptico-lanceolatae. Pappus cum circiter40
setis. Cypselae glabrae.

Twining lianas; stems terete, fistulose with age,
striate, |uiberulent to pubescent with simple pluri-
cellular uniseriate trichomes up to 1.5 mm long.

Leaves simple, opposite, petiolate, estipulate; pet¬
ioles 1.3— 2.5 cm long, occasionally with a row of
trichomes between them; blade thinly chartaceous,
triplinerved, (6)6.5-9 cm long, (2. 5)3. 5-6(6. 3) cm
wide, base attenuate, apex long-acuminate, margins
with up to six pairs of teeth, < • i I i ate*, both surfaces
with trichomes like those on the stems and sparse
sessile glands, more densely pubescent abaxially,
especially so on veins. Capitulescence terminal or
axillary, paniculate, branches terete. Capitula up to
7.5 mm tall, peduncles 1—3.5 mm long, densely
pubescent; subinvolucral bract linear-lanceolate,
1.5— 2(2.5) mm long, 0.4— 0.7 mm wide, located at
the base of the peduncle or occasionally a little
more distal in capitula borne on short peduncles,
margins entire, eiliate, apex long-acuminate, abax¬
ially puberulent, sometimes sparsely glandular; in-
volucral bracts elliptic-lanceolate, (2.5)3— 4 mm
long, 0.8-1 mm wide, margins entire, apically acu¬
minate, eiliate, pubescent to puberulent, sometimes
glandular. Corollas discoid, tube ea. 0.5— 0.8 mm
long, limb ca. 1.8— 2.5 mm long, lobes ca. 0.4— 0.5
mm long, glandular. Cypselae brownish, 2.5— 3.5
mm long glabrous, weakly glandular. Pappus with
ca. 40 biislles, 3—4 mm long, with short stiff cells,
whitish, sometimes pink-tinged.

Habitat and distribution. Mikania oreophila
thrives in borders of woods in the highlands from
the State of Rio Grande do Sul to the Serra do Mar
in Parana, reaching elevations of up to 1300 m. It
was also collected in the State of Sao Paulo at el¬
evations of 750—870 m, but with typical climatic
characteristics of the highland “campos," with hu¬
mid wooc s with frequent logs and lower night tem¬
peratures.

Phenology. Flowering and fruiting plants were
collected from February lo May.

Ktymology. The epithet is derived from the
Greek words oros (mountain) and philos (loving), in
allusion to the montane habitat of the species.

Mikania oreophila is similar to Mikania para¬
nensis, from which it is set apart by the pilose, thin-

Novon 12; 533-535. 2002.

534

Novon

Figure I. Mikania oreophila M. R. Ritter & Miotto. — A. Flowering branch. — B. Leaf. — C. Detail of stem node
showing row of trichomes. — I). Capitula. — E. Floret. — F. Subinvolucral bract. — G. Involucral bract. (A, C-G: M. R.
Hiller & .S’. I.. tie Carvalho /site H10, from holotype in ICN; B: I). II. Falkenberg 7 1 68, from paratype in FLOR.)

ly chartaceous, and basally attenuate leaves, and
by the subinvolucral bracts located at the base of
the peduncles, or sometimes more distal, and the
corolla tube uniformly widened from base to apex.
In M. paranensis the leaves are glabrous, charta¬
ceous to coriaceous, the blades are triangular and

not basally attenuate, the subinvolucral bracts arc
placed at the base ol the capitula, and the corolla
is clearly divided into a narrow proximal half and
a wider distal one. Mikania oreophila may also be
confused with M. chlorolepis Baker, but in the latter
species the leaves are longer, up to 16 cm long,

Volume 12, Number 4
2002

Ritter <k Miotto

Mikania oreophila from Brazil

535

and the subinvolucral bracts are located at the base
of the capitula; moreover, M. chlorolepis is not
known to occur in montane habitats in southern
Brazil.

According to the sectional classification pro¬
posed by Holmes (1996), Mikania oreophila should
be assigned to section Summikania due to its basal
subinvolucral bracts, although the two species with
which it is compared are better placed in section
Mikania since they bear apical subinvolucral
bracts; another character used by Holmes, the di¬
rection of the capitulescence maturation, could not
be stated confidently in either species — indeed, in
one specimen of M. paranensis where maturation
was clearly verifiable (Ritter IGOR. ICN 115651) it
occurred racemosely, in a way contrary to the pat¬
tern of section Mikania and expected to be found
in section Summikania. So, if further investigation
proves such overlap of characters, it will be nec¬
essary to look for some additional ones to reinforce
the useful sectional classification of Holmes.

Paratypes. BRAZIL Parana: Guaratuha, serra tie
Arayatuha. morro dos Perdidos, 18 Feb. 1998, II. M. her-
nandes & F. P. Santos 69 (ICN), 12 Mar. 1999, I,. C.
Cdndido A M. Hassegawa 12 (UPCB), 8 Apr. 2000. /'..
Harbosu A' 0. S. Ribas 419 (MBM), 8 Apr. 2000. E. Bar¬
bosa & 0. .S'. Ribas 432 (MBM); Nova Galicia, 18 Feb.
1916, P. Ihtsen 17701 (NY, S). Rio Grande do Sul:
Cam hard do Sul, em dire§ao a Sao Jose dos Ausentes, a
19 km de entroncamento para Retiro, 9 May 2000. M. It.
Ritter 1162 (ICN); a 17 km de Fortaleza dos Aparados

para Cambard do Sul, 28 Mar. 1999, M. R. Ritter 1007
(ICN). M. R. Ritter 1008 (ICN): fazenda Velha, Celulose
Carnbara, 5 Apr. 1994, A. Silveira A C. Mansan 11924
(HAS, ICN): Ouro Verde, 10 Apr. 1995, T. Lewinson, PIC
95176 (ICN). Santa Catarina: Urubici, alto da serra do
Corvo Branco, acesso na SC 439, junto ao eorte no morro,
24 Feb. 1995, I). B. Falkenberg 7168 (FLOR); trilha para
o topo da serra do Corvo Branco, no alto da SC 439, junto
ao eorte no morro, 28 Mar. 1995, I). B. b alkenberg 7375
(FLOR) Sao Paulo: Sao Paulo, distrito de Marsilao.
Parque Estadual da Serra do Mar, Nueleo Curucutu, 27
Mar. 1996, It. J. F. Garcia. G. M. P. Ferreira & C. M. A.
Pimentel 772 (PMSP, SPF).

Acknowledgments. We thank Marcia Vignoli da
Silva for her accurate drawing of this new species,
Marcos Sobral for stimulus and suggestions, Ely
Fumagalli Horta for correcting tin* Latin diagnosis,
and Nelson Ivo Matzenbacher for the valuable help
with the English version. The curators of the cited
herbaria are thanked for their help and availability.

Literature Cited

Holmes W. C. 1995. A review preparatory to an infrage¬
neric classification of Mikania (tribe: Eupatorieae). Pp.
239-254 in I). J. N. Hind, C. Jeffrey & G. V. Pope
(ed ite rs), Advances in Compositae Systematics, Vol. 1.
Roya Botanic Gardens, Kew.

- 1996. A proposed sectional classification of Mi¬
kania (Eupatorieae). Pp. 621-626 in I). J. N. Hind &
H. J. Beentje (editors), Compositae: Systematics. Pro¬
ceedings of the International Compositae Conference,
Kew, 1994, Vol. 1. Royal Botanic Gardens, Kew.

King, R. M. & 11. Robinson. 1987. The Genera of the
Eupa orieae (Asteraceae). Monogr. Syst. Rot. Missouri
Bot. Gard. Vol. 22.

A New Species of Faramea (Rubiaceae) from Costa Rica

Alexander Rodriguez G.

Instituto Nacional tie Biodiversidad (INBio), Apto. 22-3100, Santo Domingo de Heredia,

Costa Hica. arodrig@inbio.ac.cr

Abstract. Faramea zamorensis, a new species
from Costa Rica, is described and illustrated. Its
affinity to other species is discussed. The species
is easily distinguished from congeners by its calyx
lobes 5-7 mm long, its funnel-shaped corolla, and
its costate hypanthium and fruits.

Key words: Costa Rica, Faramea. Rubiaceae.

Faramea Aublet is a large genus of about 130
species, ranging from Mexico, Central America,
and the West Indies throughout tropical South
America (Burger & Taylor, 1993). During a field
study in the northern zone of Costa Rica, a new
species having subsessile, funnel-shaped flowers
was found and is described here. Including this
new species, there are 17 species of Faramea re¬
ported for Costa Rica, mainly in lowland rainfo¬
rests.

Faramea zamorensis Al. Rodriguez, sp. nov.
TYPE: ( msta Rica. Alajuela: San Carlos, Boca
Tapada, Cuenca del Rfo San Carlos, Finca San
Jorge, 10°43'20"N, 84°1 1'50"W, 100 m, 10
Feb. 1996, Al. Rodriguez & A. Estrada 1 067
(holotype, 1NB; isotvpes, CR, MO). Figure 1.

Species facile distinguenda of) calyce lobulis 5—7 mm
longis, corolla infundibuliformi, hypanthio costato a con-
generis diversa.

Shrub 3 m tall, glabrous throughout; stem later¬
ally flattened and remaining so with age. Leaves
sessile or with petioles up to I mm long; blade
elliptic to lance-elliptic, 10.5-18 X 3. 3-8. 5 cm,
apex acuminate with slender tips 10-17 mm long,
base cordate to subcordate, drying chartaceous,
grayish green or blackish green above and below;
secondary veins 10 to 16 pier side, spreading wide¬
ly, tertiary venation reticulated, thickened to slight¬
ly raised adaxially, slightly raised abaxially. Stip¬
ules interpetiolar, caducous, triangular, 5-1 I mm
long including arista, aristae 4—6 mm long. Flowers
terminal, 2 or 3 per fascicle, peduncles 0—1.5 mm
long, bracts ca. 7 mm long, linear-lanceolate, easily
deciduous; pedicels 1.6-2 mm long, ebracteate, hy¬
panthium turbinate, ribbed, 9-1 I mm long; calyx
lobes 5-7 mm long, linear, glabrous. Corolla fun¬
nel-shaped, violet and drying blackish, glabrous
externally, tube ca. 18 mm long, slightly widening
at mouth, ca. I mm diam. near the base and 2—3
mm diam. in the wider part, 4-lobed, with lobes 6-
7 mm long, linear-triangular, acute and ascending.
Ovary unilocular. Fruits drupaceous, leathery exo-
earp, subglobose, ca. 18 mm diam., ribs prominent
when dried, glabrous, maturing bluish black, drying
grayish black or grayish yellow, with persistent ca¬
lyx tube and lobes, tube ea. 10 mm long and 3-4
mm diam.; seed 1 per fruit, globose, ca. 8 mm
diam.

Fable I. Comparison of some characters of Faramea zamorensis , F. correae, F. sanblasensis, and F. permaenifolia.

F. zamorensis

F. correae

F sanblasensis

F. permagnifolia

Petiole (mm)

0-1

0

0 or subsessile

1-5

Leaf base

subcordate-cordate

rounded-subeordate

rounded-subeordate

rounded-truncate

Inflorescence

2 or 3 fasciculate flow-

2 or 8 fasciculate flow-

many flowers in a con-

3 or 6 fasciculate flow-

ers. terminal

ers. terminal

gested cyme, termi¬
nal

ers, terminal

Hypanthium

ribbed

terete

terete

terete

Calyx lobes

5—7 mm

truncate

truncate

truncate-denticulate

Corolla

funnel-shaped

salver-shaped

salver-shape

salver-shaped

Lobe

ascending

divergent

divergent

divergent

Fruit

ribbed

terete

terete

terete

Distribution

northern Costa Rica
and southern Nica¬
ragua

central Panama

eastern Panama

southern Costa Rica

Novon 12: 536-538. 2002.

Volume 12, Number 4
2002

Rodriguez

Faramea zamorensis from Costa Rica

537

Figure 1. Faramea zamorensis Al. Rodriguez ( Rodriguez & Estrada 1067). — a. Leaves and fruit. — b. Abaxial view
of leaf showing tertiary venation. — c. Stipule. — d. Leaves and flowers. — e. Stem.

Phenology. Flowering in January and February;
collected in fruit in both months.

Distribution. Known from wet forest in northern
Costa Rica and in southern Nicaragua at 1—100 m.
Faramea zamorensis has the narrowly triangular.

aristate stipules, unilocular ovary, and one-seeded
fruit of most Faramea species. It is distinguished
by its subsessile leaves, which are cordate or sub-
cordate at the base, subsessile fasciculate flowers
with a ribbed hypanthium, calyx lobes 5—7 mm

538

Novon

long, funnel-shaped corolla, and ribbed fruit. Be¬
cause of the slightly petiolate to sessile, basally cor¬
date to subcordate leaves and congested inflores¬
cences, this species is similar to F. correae C. M.
Taylor, F. sanblasensis C. M. Taylor, and F permag-
nifolia Dwyer ex C. M. Taylor. From all those it
differs by its much longer calyx lobes, funnel-
shaped rather than salverform corolla with ascend¬
ing rather than divergent lobes, and the ribbed
rather than terete hypanthium and fruit (see Table

I).

The specific epithet honors Nelson Zamora, a
Costa Rican botanist and colleague, who also col¬
lected this species, and whose extensive work has

greatly advanced the knowledge of the country’s flo¬
ra.

Paratype. COSTA RICA. Alajuela: San Carlos, Boca
Tapada, Cuenca del Rfo San Carlos. Finca San Jorge, 50
m. 2 Jan. 1997, Zamora 2549 (CR. INB, MO).

I .iterature Cited

Dwyer, .1. 1). 1980. Rubiaceae. In: R. K. Woodson. Jr. &
R. W. Schery et al. (editors), Flora of Panama. Ann.
Missouri Bot. Card. 67: 1—256.

Burger, W. & C. M. Taylor. 1993. Rubiaceae. In: W. Bur¬
ger (editor). Flora Costaricensis. Fieldiana, Bot. n.s. 33:
1-333.

Taylor, C. M. 2001. Rubiaceae. Pp. 2206— 2284 in: W. I).
Stevens, C. Ulfoa Ulloa, A. Pool & 0. M. Montiel (ed¬
itors), Flora de Nicaragua, Tomo 111. Monogr. Syst. Bot.
Missouri Bot. Card. Vol. 85.

Sladenia integrifolia (Sladeniaceae), a New Species from China

Shui Yu- Min, Z hang Guang-Jie, Zhou Zhe-Kun
Kunming Institute of Botany, the Chinese Academy of Sciences, Kunming 650204,

P. R. China, shuiyumin@hotmail.com

Mo Ming-Zhong

Jinping f orestry Bureau, The Forestry Department of \unnan Province, Yunnan,

Jinping 661500, P. R. China

ABSTRACT. A new species of Sladenia (Sladeni¬
aceae) from China, Sladenia integrifolia Y. M. Shui,
is described and illustrated. The relationship to and
characters distinguishing the new species from
Sladenia celastrifolia Kurz are discussed. I he sys¬
tematic position of the genus is also reviewed.

Key words: China, Sladenia , Sladeniaceae.

Between 15 and 26 October 1999, the authors
made an expedition to the Xilongshan range of
Jingping County, southeastern Yunnan, China, ad¬
jacent to southwestern Vietnam. During the expe¬
dition, the specimen Zhou Zhe-Kun el at. EXLS-
0039 (fr) was identified as Sladenia celastrifolia
Kurz. However, further study showed that the entire
margin of t he leaves differed from the serrated mar¬
gin of those oi Sladenia celastrifolia. On 15 May
2001, another specimen. Mo Ming-Zhong, Mao
Rong-Hua <V Yu Zhi-Yong 05 (fl), was collected,
and more differences were found. As a result, a new
species of Sladenia Kurz (Sladeniaceae) is here de¬
scribed.

Sladenia integrifolia Y. M. Shui, sp. nov. 1 't Ph:
China. Yunnan: Jinping Co., Zhemi commu¬
nity, Citongba to Liangzi, second dry evergreen
forests, 1100-1300 m, 15 May 2001 (11), Mo
Ming-Zhong, Mao Rong-Hua & Yu Zhi-Yong
05 (holotype, KUN 0735701; isotypes, MO.
PE). Figure 1.

Species Sladeniae celastrifoliae similis, seel foliis iute-
gris, floribus lateralibus pedicellis 2.54 mm longis, se-
palis late ovatis, 2.0 mm longis, 1.5 mm latis, filamentis
base non connatis, fructibus 3 mm longis et crustaceis,
costis non conspicuis, rlitfert.

Evergreen trees, 13—15 m tall; branehlets green,
terete, turgid, glabrous throughout; buds broadly
ovate, glabrous. Leaves spiral, papery, glabrous,
ovate or lanceolate, 5-1 I cm long, 2. 5-4.0 cm
wide, adaxially deep green, abaxially greenish;
base broadly cuneate, slightly decurrent on the pet¬

iole; margin entire, apex acuminate or caudate,
with obtuse tip: midrib shallowly canaliculate on
adaxial surface, raised on abaxial, lateral veins in
7 to 9 pairs, raised on both surfaces, secondary
nerves sparse, slightly prominent on both surfaces;
petiole 0. 7-0.9 cm long, glabrous. Dichotomous
cymes axillary, usually 4- or 5-forked, congested,
with 14 to 17 llowers; peduncle glabrous, 0.4— 0.8
cm; terminal flowers with pedicels 0.2— 0.3 mm
long, laieral Mowers with pedicels 2.5— 4.0 mm long;
bracts narrowly ovate, ca. 2 mm long; bracteoles
ovate, ca. 0.8 mm long, glabrous, caducous. Sepals
broadly ovate, imbricate, ca. 2 mm long, 1.5 mm
wide, obtuse at apex, with eiliate margin, not
sprawling-reverse in fruit. Corolla white, urn¬
shaped, glabrous, ca. 3 mm long, ca. 3 mm diam.
at base, 5-fid at apex; corolla lube 0.7— 0.8 mm
long; lobes 5, involute, concave, oblong, 2.2-2. 3
mm long, 1.5— 1.7 mm wide. Stamens 8 to 10, in¬
serted at base of corolla; filaments free, ca. 0.8 mm
long, Cc . 0.4 mm wide, acute at apex; anthers in¬
dexed, ovate, ca. 0.8 mm long, apex 2-dehiscent
with 2 hairs, base sagittate with 2 hairs on each
side; ovary 3-locular, conoid, ca. 2 mm long, ca.
0.9 mm diam. at base, glabrous, apex continuous
with style, tip 3-fid. Eruit ovoid, crustaceous, slight¬
ly striate, ca. 3 mm long, 1. 0-1.1 mm diam. Seeds
trigonal, winged, ca. 1.5 mm long, ca. 0.3 cm wide.

Habit at. In secondary evergreen forests, 1000-
1300 m.

Phenology. Flowering March to June; fruiting
July to December.

Distribution. China (SE Yunnan Province).

The Mower and fruit characteristics of Zhou Zhe-
Kun et al. EXLS-0039 clearly belong to those of the
genus Sladenia, as elaborated on by Ming (1997).
However, the entire leaves show that it is not iden¬
tical to Sladenia celastrifolia. The new species fur¬
ther dilfers from Sladenia celastrifolia in its gla¬
brous twig, broadly ovate sepals, 2.5— 4.0 mm long

Novon 12: 539-542. 2002.

540

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2 mm

Volume 12, Number 4
2002

Shui el al.

Sladenia integrifolia from China

541

Table 1 . Differe

■nces between Sladenia celastrifolia and S. integrifolia.

Characteristics

Sladenia integrifolia Y. M. Shui

Sladenia celastrifolia Kurz

Leaf margin

entire

serrate, rarely entire

Lateral pedicel

2.5— 4 mm

7—10 mm

Cymes

4- or 5-forked, congested

2- or 3-forked, spananthus

Sepal

broadly ovate; not erect in mature fruit; 2.0
mm long, 1.5 mm wide

oblong; sprawling-reverse in mature fruit; 5—
6 mm long, 2—3 mm wide

Corolla

2.9— 3.1 mm long, connate at base

5.3— 6.4 mm long, almost distinct at base

Corolla tube

0.7— 0.8 mm long, ca. *4 as long as corolla

0.3— 0.4 mm long, ca. 1/17 as long as corolla

Corolla lobe

broadly oblong; 2.2— 2.3 mm long, 1.5— 1.7
mm wide

oblong; 5—6 mm long, 2—3 mm wide

Filament

0.8 mm long, 0.4 mm wide, distinct at base

1.5 mm long. 1 mm wide, connate at most
weakly

Stamens

8 to 10

10 to 13

Anther

ovate, 0.8 mm long

sagittate, 2 mm long

Ovary

conoid, obtuse at apex

pyramidal, acuminate at apex

Fruit

ovoid, slightly striate, crustaceous; 3 mm
long, 1.0— 1.1 mm diam.

bottle-shaped, ribbed, woody; 7-8 mm long.
3—4 mm diam.

Seed

1 .5 mm long, 0.3 cm wide

3 mm long, 1 mm wide

lateral pedicels, and 3 mm long, slightly striate
fruits (Table I ).

fhe formerly monotypic genus Sladenia is allied
with Dilleniaceae (Gilg, 1893), Linaceae (Haller.
1923), Sladeniaceae (Airy Shaw, 1964). Aetinidi-
aceae (Gilg & Werdermann. 1925; Hutchinson.
1969), and Theaceae (Kurz, 1873; keng, 1962;
Cronquist, 1981; Takhtajan, 1996; Thorne, 2000).
Airy Shaw (1964) considered it to be a monotypic
family, Sladeniaceae. In Sladenia the basic number
of chromosomes (hi. 2002), the character of wood
anatomy (Deng & Baas, 1990, 1991), the apical
pore ol the anthers (Kobuski, 1951; keng, 1962),
palynology (Wei et al.. 1997), and embryology (l.i
et al., 2002) are all considerably different from the
family Theaceae. Molecular l)NA allies Sladenia
with the East African Ficalhoa Hiern. the two form¬
ing the sister taxon to Ternstroemiaceae (Anderberg
et al., 2002). It is thus reasonable that the group is
considered to be a monotypic family. Currently, ma¬
terials of testa anatomy and chemistry are poorly
known. A detailed systematic study of the charac¬
teristics and relationships in light of the new spe¬
cies will be significant to understanding the posi¬
tion of the genus Sladenia in angiosperms.

Paratypes. CHINA. Yunnan: Jinping Co., Zhemi
community, Citongba to biangzi, in secondary evergreen

forests, 1290 in, 17 Oct. 1999 (fr), Zhou Zhe-Knn , Fei
Yong, Shvi Yu-Min, Zhang Guang-Jie & Yang Jian-Kun
EXLS-Oa.V (KUN, MO, PE), 25 June 2001 (fl). Hu Yun-
Qan A- Deng Ling 22 (KUN), I July 2001 (yng. fr). I.i Lu
et Kong L'ong-Rui 2001-7-1 (KUN,' MO. PE).

Acknowledgments. This research was funded by
the National Natural Science Foundation of China
(grant no. 39930020), the National Natural Science
Foundation of Yunnan Province, China (grant no.
2001C22Q), and the Taxonomy and Flora Special
Foundation, Chinese Academy of Sciences (C AS),
fhe authors are also indebted to Wu Zheng-yi.
Ming Tian-lu, and Peng Hua (kUN) for their guid¬
ance. hi Lu was insightful in modifying the errors
in the figure. Additionally, Wu Xi-Lin is thanked
for his excellent drawing. Finally, Mao Yong-hua
and Yu Zhi-yong were very helpful during field¬
work.

Literature Cited

Airy Shaw H. K. 1964. Diagnoses of new families, new
names, etc. for the seventh edition of W illis’s dictionary.
Kew Bu I. 18: 249-273.

Anderberg A., C. Rydin & M. Kallersjo. 2002. Phyloge¬
netic relationships in the order Ericales s.l.: Analyses
of molecular data from five genes from the plastid and
mitochondrial genomes. Amer. J. Bot. 89: 677—687.
Cronquist, A. 1981. P. 322 in An Integrated System of

Figure 1. Sladenia integrifolia Y. M. Shui. — a. Habit (in flower). — b. Habit (in fruit). — e. flower. — d. Adaxial
view of stamen. — e. Lateral view of stamen. — f. Opened corolla and stamens. — g. Fruit with persistent calyx. — h.
Seed. — i. Transverse section of fruit, a, c— f from the holotype (Mo Ming-Zhong, Mao Rong-Hua & Yu Zhi-Yong 05);
b, g— i from a paratype (Zhou Zhe-Kun, Fei Yong, Shut Yu- Min, Zhang Guang-Jie & Yang Jian-Kun EXLS-0039).
Drawn by Wu Xi-lin.

542

No von

Classification of Flowering Plants. Columbia Univ.
Press, New York.

Deng, L. & P. Haas. 1990. Wood anatomy of trees and
shrubs from China II. Theaeeae. I.A.W.A. Bull., N.S.
I 1 : 337-378.

- & - . 1991. The wood anatomy of the Thea-

ceao. I.A.W.A. Bull., N.S. 12: 333-353.

Gilg, E. 1893. Dilleniaeeae. In: A. Engler & k. Prantl,
Die NatUrliehen Pflanzenfamilien: 100-128. Leipzig,
Germany.

- & E. Werdermann. 1925. Actinidiaceae. In: A.

Engler & k. Prantl, Die NatUrliehen Pflanzenfamilien.
2nd ed., 21: 36—47. Leipzig, Germany.

Haller, II. 1923. Be it rage zur kenntnis der Linaeeae. Du-
mort. Beih. Bot. Centralbl. 39: 1 — 178.

Hutchinson, J. 1969. P. 289 in Evolution and Phylogeny
of Flowering Plants. Academic Press, London & New
York.

keng, H. 1962. Comparative morphological studies in
Theaeeae. Univ. Calif. Pub]. Bot. 33: 355—356.

kobuski, C. E. 1951. Studies in the Theaeeae, XXIV. The
genus Sladenia. .1. Arnold Arbor. 32: 403-408.

Kurz, S. 1873. On a few new plants from Yunnan [sic]. .1.
Bot. II: 193-196, t. 133.

Li, L. 2001. Chromosome number of Sladenia celastrifolia.
Acta Bot. Yunnan. 23: 223—224.

- , 11. X. Liang, II. Peng & S. C. Tucker. 2002.

Embryology of Sladenia and its systematic affinities.
Amer. J. Bot. (in press).

Ming, T. L. 1997. Sladeniaceae. In: Wu (5 Y. (editor).
Flora Yunnanica. Lomus 8. 383-385, lig. 98: 6-1 I. Sci¬
ence Press. Beijing.

I'akhtajan, A. 1996. R 1 64 in Diversity and Classification
of Flowering Plants. Columbia Univ. Press, Newr York.
Thorne, R. F. 2000. The classification and geography of
the (lowering plants: Dicotyledons of the Class Angios-
permae, Dilleniidae, Rosidae, Asteridae and Lamidae.
Bot. Rev. 66(4): 491.

Wei, Z. X.. I). Z. Li. X. k Fan & X. L. Zhang. 1997.
Pollen ultrastructure of Pentaphyllaceae and Sladeni¬
aceae and their relationships to the family Theaeeae.
Acta Bot. Yunnan. 21(2): 202—206

Drosera peruensis (Droseraceae), a New Species from Peru

Tania Regina dos Santos Silva

Depto. de Ciencias Biologicas, Universidade Estadual de Feira de Santana, BR 116, km 3,
CEP 44031-460, Eeira de Santana, Bahia, Brazil. taniasilva@mailbr.com.br

Mireya D. Correa A.

Herbario de la Universidad de Panama and Instituto Smithsonian de Investigaciones
Tropicales, Apartado 2072, Balboa, \ncon, Panama

Abstract. During a revision of the genus Drosera
for Flora Neo tropica, a new species was found.
Drosera peruensis is described as new from Peru for
the neotropics and is placed in D. subg. Drosera in
section Drosera. It is characterized by the presence
of elongated stems, leaves that become patent when
old, obovate-oblong blades with distinct petioles,
filamentous trichomes on the abaxial leaf surface,
and inflorescence with filamentous trichomes.

Key words: Drosera, Droseraceae, Peru.

Drosera peruensis T. Silva & M. D. Correa, sp.
nov. TYPE: Peru. Pasco: Oxapampa, Cordi¬
llera Yanachaga, Cerro Pajonal, “chacos” 12
km SE ol Oxapampa, 75°20'W, 20°35'S,
2700-2800 m. valley of Rio Yamaquizu, above
Chacos to a microwave construction site, a
footpath toward a crest behind the site toward
a small lagoon, 7 Oct. 1982, R. B. Foster 9066
(holotype, PM A; isotypes, F, MO). Figures 1. 2.

Herba 4—11.5 cm alta. Caules elongati. 2-t cm longi.
Folia rosulata, vinacea, matura pendentia, laminae oblon-
gae, 4—7 mm longae, ad 1 mm latae. Inflorescentiae 1—2,
vinaceae, 4—7 cm longae, flores 2-4; ovaria 3-carpel I aria,
styli 3, profunde 2-partiti; stigmata 6, clavata. fructus
capsularis, valvis 3; semina oblonga, reticulata.

Herb, 4—1 1.5 cm tall; stem 2^4 cm long. Leaves
reddish, patent when old, 10-12 cm long, blades
obovate-oblong, 4-7 X 1 mm, adaxial surface vil¬
lous, glandular trichomes, abaxial surface densely
villous, filamentous trichomes to 1 mm long, pale
gold; petioles 6—7 X 1 mm, adaxial surface gla¬
brous, abaxial surface villous, filamentous tri¬
chomes, 2—2.5 mm long, pale gold; stipules rect¬
angular, membranaceous, ca. 7 mm long, laeiniate,
laciniae ca. 6 mm long. Inflorescences 1 or 2. red-
tinged, flowers 2 or 4, 4-7 cm long, villous, with
filamentous trichomes, peduncle 3.5—6 cm long,
pedicel reddish. Flowers with reddish sepals, 4 X
1.5 mm, fused in lower Ft, abaxial surface villous,
with reddish filamentous trichomes to 1 mm long,

Figure 1. Drosera peruensis T. Silva & M. D. Correa. —
A. Habit. — B. Stipule. — C. Leaf. — D. Seed. ( Foster
9066, PM A.)

Novon 12: 543-545. 2002.

544

Novon

Figure 2. SKM photographs of Drosera peruensis. — A. Leaf, abaxial surface detail. — B Seed. — C. Seed, surface
detail. — I). Pollen, proximal view. — E. Pollen, pore detail. (Foster 9066 , PM A.)

Volume 12, Number 4
2002

Silva & Correa

Drosera oeruensis from Peru

545

lobes oblong; petals white or red; ovary 3-carpel-
late, styles 3 bipartite to the base, stigmas 6, cla-
vate. Fruit a 3-valvate capsule; seed oblong, testa
reticulate.

The genus Drosera L. is distributed throughout
Australia, America, and Africa, with a few species
in Asia and Europe. Twenty species are found in
the neotropics, concentrated in northern South
America (Venezuela, Guyana, Surinam, and French
Guiana) and eastern Brazil. Two species are found
in Peru: D. chrysolepis Taubert and D. peruensis T.
Silva & M. 1). Correa.

Drosera peruensis is distinguished from all other
Drosera species by the presence of elongated stems,
patent leaves when old, obovate-oblong blades with
distinct petioles and densely covered by filamen¬
tous trichomes on the abaxial leaf surface, and the
inflorescence with filamentous trichomes. The area
where it was found was a shrubland on white sand¬
stone, with spongy Sphagnum humus up to 2 m
deep, except where burned. The species was colo¬
nizing a sandy wet landslide on a steep slope, form¬
ing small pillars with old leaves and stems. It is
only known from the type collection.

Drosera peruensis is distinguished from D. chry¬
solepis by having leaves with oblong-obovate blades
4—7 mm ong, a petiole 6—7 mm long, and rectan¬
gular stipules, whereas D. chrysolepis has narrowly
oblong blades 1-3 mm long, a petiole 2-5 cm long,
and triangular stipules. The blades of Drosera per¬
uensis are similar in shape to the obovate-oblong
ones in D. arenicola Steyermark. The new species
has filamentous trichomes on the inflorescence,
while D. arenicola has glandular trichomes.

Drosera peruensis is placed in Drosera subg.
Drosera 1 ased on its subterraneous, non-tubercu-
late root system and in section Drosera because it
is stipulated and has three styles divided to the
base (Seine & Barthlott, 1994).

Acknowledgments. We thank Robin Foster,
Field Museum of Natural History, Chicago, for
specimens from Peru and the Office of Fellowships
and Grants, Smithsonian Institution, Washington.
D.C., for T. Silva’s financial support.

Literature Cited

Seine, IT A W. Barthlott. 1994. Some proposals on the
infrageneric classification on Drosera L. Taxon 43: 583-
589.

Caobangia , a New Genus and Species of Polypodiaceae

from Vietnam

Alan R. Smith

University Herbarium, 1001 Valley Life Sciences Bldg. #2465, University of California,
Berkeley, California 94720-2465, U.S.A. arsmith@uclink4.berkeley.edu

Zhang Xian- Chun

Institute of Botany, Academia Sinica, 20 Nanxincun, Xiangshan, 100093 Beijing,
People’s Republic of China, xianchunzhang@yahoo.com.cn

ABSTRACT. We describe and illustrate the mono-
typic new genus and species Caolmngia squamata
(Polypodiaceae), endemic to limestone ridges in
northern Vietnam. Caobangia is superficially sim¬
ilar to the predominantly Neotropical genera Micro-
gramma and Pleopeltis, but its affinities appear to
he with Lepidogrammitis, comprising about eight
species in China and southeastern Asia. Caobangia
is characterized by long-creeping filiform rhizomes,
clathrate rhizome scales, simple, monomorphic to
subdimorphic, densely acicular-scaly blades, and a
single row of sori on each side of the midrib.

Key words: Caobangia, Polypodiaceae, Viet¬
nam.

Discovery of a new genus of ferns, based solely
on previously undescribed species of an avowedly
non-hybrid nature, is an uncommon event. Quite
recently, a new genus and species ( Mankyua che-
juense B.-Y. Sun et al.) of the eusporangiate fern
family Ophioglossaceae was described from Cheju
Island, Korea (Sun et al., 2001), but previous to
that, the last such new genera of which we are
aware were the description of Podosorus angustatns
Holttum (I960), Polypodiaceae, from the Philip¬
pines, Cystoathyrium chinense Clung (1906), Athy-
riaceae, from China, and A ’ephopleris maxonii Del¬
linger (1966), Pteridaceae, from Colombia.

Recent pteridophyte floras of areas within east¬
ern Asia (ail to provide clues as to the identity of
our species, and one quickly comes to “blind al¬
leys" in available keys. In the recent Flora Male-
siana treatment for Polypodiaceae (Hovenkamp et
al., 1998), our undescribed taxon will key to Lem-
nuiphyllum s.l. (including lepidogrammitis), in the
sense that name is applied by Hovenkamp, but his
description of Lemmaphyllum is at variance with
Caobangia in several important respects (see dis¬
cussion under the new genus below). In an effort
to identify the plant we describe herein, we have

consulted, without success, keys to genera and spe¬
cies from the following areas: Vietnam (Tardieu-
Blot & Christensen, 1941; Pham-lloang, 1991,
1999); Thailand (Smitinand X Larsen, 1989); Chi¬
na (Lin, 2000); Malaysia (Holttum, 1954); Malesia
(Hovenkamp et al., 1998); and Taiwan (Huang.
1994). We have also shown a specimen of the plant
in question to knowledgeable colleagues in Asia
(especially China and Japan), Europe, and North
America, and no one has seen anything remotely
resembling the plant described below. Without hes¬
itation, therefore, we name this taxon as a new ge¬
nus and species.

Caobangia A. R. Smith X X. C. Zhang, gen. nov.

TY PE: Caobangia squamata A. R. Smith & X.

C. Zhang. Figure 1.

A lepisoro et Lepidogrammitide I am in is dense squa-
matis, squamis fere laminis occultantibus, rubro-brunneis,
plemmque 5-6 mm longis, base peltatis clathratis mar-
ginibus dentatis, apice acicularibus non-elathratis dentatis
differt; a Lemmaphyllo s. str. sori mtundatis diserelis ca.
2 mm diametro, laminis monomorphis vel subdimorphis
squamis numerosis acicularibus differt.

Rhizomes long-creeping, filiform, sparingly
branched, approximately terete, dorsiventral with
two rows of dorsal fronds, rather sparsely set with
roots, not ant-inhabited, densely scaly; scales lack¬
ing hairs at the bases (non-comose), acicular and
nearly entire from peltately attached, clathrate,
strongly and irregularly toothed bases. Fronds
short-stipitate to nearly sessile; stipes scaly, lacking
obvious articulation lines or swellings at bases;
blades herbaceous to chartaceous, entire, elliptic to
ovate, monomorphic to subdimorphic in size and
shape, dark green-brown when dried, stellate hairs
lacking, densely scaly on both sides with persistent
red-brown, hairlike scales; blade scales with pel¬
tate, clathrate, laciniate-margined bases and very
long, acicular, uniformly red-brown, non-clathrate,

Novon 12: 546-550. 2002.

Volume 12, Number 4
2002

Smith & ;!hang
Caobang a from Vietnam

547

Figure 1. Caobangia squamata A. It. Smith & X. C. Zhang, based on Loc et al. CHL 1698 (PE, UC). A. I lant

habit. B. Abaxial blade surface, with sorus. — C. Adaxial blade surface. — D. Rhizome scale. — E. Seale from abaxial

surface of blade. — F. Scale from adaxial surface of blade. — G. Scale near sorus. — H. Sporangium. — I. Spores. — J.
Venation in a portion of fertile leaf, with scales removed; stippled areas indicate soral receptacles, with sporangia
removed.

548

Novon

dentate to laciniate tips. Venation: midribs distinct,
sclerenchymatous (darkened, hut hidden by red-
brown scales), main lateral veins indistinct or dis¬
tinct oidy in the basal 1/5 or less in cleared fronds,
10—13 per side, anastomosing and forming irregular
areoles (1 row of large areoles adjacent to midrib,
0- 2 additional rows of smaller areoles toward mar¬
gins), areoles with occasional free veinlets, these
simple (unforked), almost always recurrent, im¬
mersed, not or only faintly visible adaxially, veins
toward blade margins often ending freelv, hyda-
thodes lacking. Sori in single' rows within the larger
areoles, midway between midrib and each margin,
round, not confluent, ± obscured by acicular
scales. Sporangia stalked, lacking paraphyses, w ith
16 to 19 indurated annulus cells, mixed with acic¬
ular scales; spores monolete, whitish, papillate to
rugose, fully mature spores not seen.

riie' name Caobangia is chosen to acknowledge
the area where this most distinctive fern grows. It
is the name of the province, as well as the name of
the capital city of the province. “Cao,” in Vietnam¬
ese, means high; “Bang” refers to a locality with
more or less flat terrain. The province of Cao Bang
is in a limestone area, carved with many often nar¬
row' valleys. I he city ol Cao Bang lies in the largest
valley of this province, surrounded by high moun¬
tains. The Cao Bang limestone area is one of the
ric best areas for plant diversity in Vietnam, and
many endemic or relatively restricted plants are
known from the region (Averyanov, 1998; Loc et ah,
1999a, 1999b; Hiep et ah, 2000). Recent collec¬
tions from this area from diverse families of ferns
and seed plants have proven to be first records of
species from Vietnam, e.g., Cyrtomium hemionitis
II. Christ (Dryopteridaceae) (see (http://mobot.
mobot.nrg/\X 3 I /Search/vast. html)), Loxogratnme
grammitoides (Baker) C. Christensen (Polypodi-
aceae), Pteris actiniopteroides C. Christensen (Pter-
idaceae), Pseudotsuga brevifolia W. C. Cheng & L.
k. [• u (Pinaeeae), Campylotropis henryi (Schindler)
Schindler (Leguminosae), Luculia yunnanensis S. Y.
Hu (Rubiaeeae), Myrsine kwangsiensis (E. II. Walk¬
er) I’ipoly & C. Chen (Myrsinaceae). Paphiopedil-
urn diant hum T. Tang & F. T. Wang and Paphio-
pedilum micranthum T. Tang & F. T. Wang
(Orchidaceae), and Swertia nervosa (Wallich ex G.
Don) C. B. Clarke (Gentianaceae) (see Foe et ah,
1999b, for more examples).

The habitat of Caobangia is on limestone ridges
in closed tropical seasonal submontane conifer for¬
est dominated by Pseudotsuga brevifolia W. C.
Cheng «S' L. k. fu mixed with Pin us kwangtungen-
sis Chun ex Isiang, sometimes with Calocedrus ma-

crolepis kurz and Amentotaxus argotaenia (Hance)
Pilger. The common angiosperm found in the over¬
story is Quercus. In the understory (7-15 m tall)
are the conifers Nageia fleuryi (Nickel) de Lauben-
fels, and Podocarpus brevifolius (Stapf) Foxworthy,
and angiosperms Platycarya strobilacea Siebold &
Zuccarini, Pistacea weinmannifolia J. Poisson ex
Franehet, and species of Acer, Carpinus , Elaeocar-
pus, Eriobotryu , Sorbus, Schefflera, and Ulmus.
Shrubs and herbs abound; among the latter are nu¬
merous species of Orchidaceae, terrestrial as well
as epiphytic. Ferns and bryophytes are similarly
abundant, both as lithophytes and epiphytes. The
limestone ridges on which this forest community
occurs are extremely eroded, composed of resistant,
marble-like rock outcrops interspersed with thin
soil pockets. The climate is tropical, with cool win¬
ters (5 months, from November to March, with
mean monthly temperature below 17° Celsius) and
summer rains. There is no dry period: all months
have rainfall over 25 mm (see Van et ah. 2000).

Affinities of this distinctive new genus are ob¬
scure, on initial inspection, other than being an un¬
doubted member of the Polypodiaceae. Character¬
istics placing it firmly in Polypodiaceae include the
long-creeping, sparingly branched, clathrate-sealy
rhizomes; articulate fronds; simple blades; areolate
venation with included recurrent veinlets; lack of
hairs on the blades; round, dorsal, exindusiate sori;
and monolete, bilateral whitish spores. Moreover, it
grows on rocks, a habitat favored by many members
of this family. W ithin Polypodiaceae, only Platycer-
ium and Pyrrosia in the Paleotropics and subtropics
be. ir such dense scales on the blade, and the scale
type found in Caobangia (peltate-acicular and elath-
rate at the base) is strongly at variance with the stel¬
late scales found in platycerioid genera. We suggest
that Caobangia is probably a member of the lepi-
soroid clade, comprising the genera Drymotaenium ,
Lepisorus, Lepidogrammitis, Lemmaphyllum, and,
more remotely, Belvisia (tribe Lepisoreae of Hennip-
man et ah, 1990). From all of these, Caobangia dif¬
fers in the densely scaly blades. Additionally, from
Lepisorus it differs in lacking peltate scales from the
receptacle. Caobangia is more distantly related to
the microsoroid genera, which include Microsorurn .
Leptochilus, Neocheiropteris, Colysis , and Phymato-
sorus (Nooteboom, 1997, 1999), and it differs from
these by the single regular row of sori between mid¬
rib and blade margin and the abundant peltate-acic¬
ular scales on the blades.

I hi1 closest living relative of Caobangia may be
the genus Lepidogrammitis Clung, sometimes treat¬
ed as a synonym of Lemmaphyllum (e.g., by Hen-
niprnan et ah, 1990). Lepidogrammitis (Cliing,

Volume 12, Number 4
2002

Smith & Zhang
Caobangia from Vietnam

549

1940), which we prefer to recognize at generic
rank, comprises perhaps eight species in China and
southeastern Asia (Lin, 2000). It is characterized
by long-creeping, filiform rhizomes, clathrate rhi¬
zome seales, fronds with simple, monomorphic to
subdimorphic or dimorphic blades, and, on each
fertile blade, a single row of sori on each side of
the midrib. Most of the species are restricted to
China. Lepidogrammitis differs from Caobangia in
having the rhizome scales decidedly clathrate
throughout their entire length (including tips), gla¬
brous or nearly glabrous, subcoriaceous or fleshy
blades, strongly articulate stipe bases, and pres¬
ence of stalked peltate scales within the sori. The
venation in at least some species of lepidogram¬
mitis (see, e.g.. Lin, 2000, fig. 19.8, lepidogram-
mitis drymoglossoides (Baker) Ching) is similar to
that of Caobangia (Fig. I J ) exeept that in Caoban¬
gia there is a less continuous intramarginal union
of the vein tips around the periphery of the blade,
and veins of Caobangia often end freely near the
margin.

If one compares Lemmaphyllum sensu Hoven-
kamp (in Hovenkamp et al., 1998) with Caobangia.
one notes the following differences: rhizome scales
peltately attached at the base (never basifixed, as
described for some Lemmaphyllum ), seales also
acicular and essentially entire beyond the base (vs.
dentate to lacerate in Lemmaphyllum); blades her¬
baceous to chartaeeous, densely scaly on both sur¬
faces (vs. more or less succulent and glabrous or
essentially so in Lemmaphyllum); veins forming one
large and an incomplete smaller second row of ar-
eoles between midrib and margin (vs. 2 or 3 rows
of areoles in Lemmaphyllum; see, e.g.. fig. 19.2 in
Lin, 2000); and sori round (vs. round to elliptic to
longitudinally coenosoriate in Lemmaphyllum), sori
lacking round-peltate paraphyses (vs. paraphyses
round-peltate in Lemmaphyllum). Fronds of Lem-
maphyllum s. str. (2 or 3 species) are often strongly
dimorphic, and the sori appear linear and subacros-
tichoid (confluent) at maturity. Lemmaphyllum car-
nosum (Hooker) C. Brest bears a few scattered
seales on the lamina abaxially (e.g., van der WerJ]
et al. 14216. UC, from Vietnam), but these are
round-peltate to ovate, much less strikingly acicu¬
lar than those in Caobangia , only about I mm long,
and strongly clathrate throughout. lemmaphyllum
accedens (Blume) Donk, sometimes placed in the
segregate genus Weatherbya, is widespread in Ma-
lesia to Polynesia and is yet more distantly related.
Both lemmaphyllum s. str. and lepidogrammitis
are relatively common epiphytes in low-elevation
forests in southern China and southeast Asia.

There is no close relationship that we see to any

known genus of New World Polypodiaceae. From
Pleopeltis, perhaps the most superficially similar
genus, Caobangia differs in having acicular scales
from a peltate base on all parts of the blades (in¬
stead of round-peltate scales). The rhizome scales
of Caobangia are non-comose anti uniformly col¬
ored, contrasting with the comose, bicolored scales
found in many species of Pleopeltis. Microgramma,
also with simple blades, has non-clathrate rhizome
scales lacking acicular tips; il is more remotely re¬
lated, although still somewhat superficially similar.
Unpublished molecular work by Haufler et al. place
Pleopeltis and Microgramma solidly within a large
clade of beotropical Polypodiaceae, and separate
from nearly all Old World genera of Polypodiaceae,
which form their own large clade (Schneider et al.,
ms. in pre 5.).

Caobangia squamata A. IL Smith & X. C. Zhang,
sp. ncv. TYPE: Vietnam. Cao Bang: Ha Lang,
Thang Loi, vicinity of Lung Sam village,
22°45'N, 1 06°42'E, 550-600 m, 24 Apr.
1999, P K. Loc, P. H. Hoang & /.. Averyanov
CBL ’69H (holotype. HN not seen; isotypes.
LE net seen, KUN not seen. MO 5291090 not
seen, PE, UC). Figure 1.

A Lepidogrammitide rostrata (Beddome) Ching laminis
dense squaniatis, squamis fere laminis occultantibus, rub-
ro-brunneis, plerumque 5-6 mm longis, base peltatis
clathratis, marginibus dentatis vel laciniatis, apice aci-
cularibus dentatis non-clathratis differt.

Rhizomes ca. I mm diam., 2-4 cm between stipe
liases, densely covered with numerous ascending
and outwardly curved, overlapping scales; scales
dark red-brown, ca. 5 mm long, peltate bases ca.
0. 4—1.2 mm wide (including teeth), also with a few
short teeth to 0.2 mm long in proximal third of the
long acicular scale tips, distal portion of scales un-
iseriate for 1—2 mm. Fronds to 7.5 X 1.8 cm; stipes
mostly 1—10 mm long, with scales similar to those
of blades; sterile blades ovate to broadly elliptic or
short-oblong, often shorter and more rounded than
the fertile at their bases but equaling the fertile in
width (transitional shapes appear to occur), mostly
ca. half the length of the fertile fronds or less,
rounded at tips, densely covered (but some laminar
tissue visible between overlapping scales) on both
sides with persistent scales mostly 5—6 mm long,
scales acic jlar from peltate, clathrate, dentate-mar-
gined or laeiniate bases, the acicular tips uniformly
red-brown and non-clathrate, bearing up to ca. 19
teeth per side, teeth up to 0.3 mm long, scales
curved and ascending or spreading, extending up
to 5 mm beyond blade margins. Sori up to ca. 9 on

550

Novon

each side of midrib, up to 2 mm diam., difficult to
see because of the dense overlayer of scales.

Plants rare, epipetric on vertical shaded lime¬
stone cliffs of northern exposure, in secondary wet
evergreen broadleaved forest with remnants ol pri¬
mary elements on sleep slopes and bluffs of karst
remnant limestone ridge. File type was distributed
under the name Irpisorus, and is the only known
collection.

Acknowledgments. We especially thank Phan
Ke Loe, Hanoi University, one of the collectors of
this plant, for comments on the habitat and vege¬
tation of this area. We thank also Leonid Averyanov,
Komarov Botanical Institute, St. Petersburg, for the
suggestion ol the generic name, Hans Nooteboom,
Leiden, and W. M. Chu. Yunnan University, for ex¬
amining and commenting on plants of Caobangia,
and two anonymous reviewers for their constructive
and helpful comments. Harald Schneider, Duke
University and Univ. Gottingen, has kindly made
available a photograph of the venation of cleared
leaves. Finally, we thank the artist. Ji Chao-Zhen,
lor rendering tin- illustration, and Linda Vorobik for
final alterations in the plate. The expedition that
collected Caobangia for the first time was funded
by a grant Irom the U. S. National Geographic So¬
ciety to L. V. Averyanov (grant no. 6300-98).

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vae et rarae in flora Vietnami. Novosti Sist. Vyssh. Hast.
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Ching, R. C. 1940. On natural classification of the family
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- . 1966. Three new fern genera. Acta Phytotax. Sin.

II: 17-29.

Hennipman, E., P. Veldhoen & K. U. Kramer. 1990. Po¬
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dophytes and Gymnosperms. Springer- Verlag, Berlin.
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new species for the flora of Vietnam collected from the

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- . 1966. New ferns from Malesia. Kew Bull. 20:

455-460.

Hovenkamp, P. et al. 1998. Polypodiaceae. Pp. 1-234 in
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Huang, f.-C. (editor-in-chief). 1994. Flora of Taiwan, 2nd
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Lellinger, I). B. 1966. Nephopteris , a new genus of ferns
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Lin, \. 2000. Pteridophyta: Dipteridaceae, Polypodiaceae,
Platyceriaceae, Loxogrammaceae, Salviniaceae, Chei-
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Sinicae 6(2): 1-404.

Loe, P. k., N. T. Iliep & L. Averyanov. 1999a. Some
threatened plant species on limestone of Caobang prov.
needed to be protected in priority. Vietnam Forest. Rev.
12: 35—36 [in Vietnamese, summary in English],

- . - & - . 1999b. Are there new plant

records from Caobang limestone? In Conservation and
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stitute, Hanoi [in Vietnamese].

Nooteboom, H. P. 1997. The mierosoroid ferns. Blumea
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- . 1999. The mierosoroid Polypodiaceae: The gen¬
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X.-C. Zhang & K.-H. Shing (editors), Ching Memorial
Volume. China Forestry Publishing House, Beijing.

Pham-Hoang. lid. 1991. QSyco Vidt Nam = An Illustrated
Flora of Vietnam, Vol. 1(1). Pham- Ho&ng Ho, Montreal.

- . 1999. CSyco Vidt Nam = An Illustrated Flora of

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ridophytes. Vol. 3(4): 481-640.

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Mankyua (Ophioglossaceae): A new fern genus from
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H. Lecomte (editor), Flore Cdndrale de l’Indo-Chine.
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Bioclimatic Diagrams of Vietnam. Vietnam National
University Publishing House, Hanoi.

A New Species of Gonolobus (Apocynaceae, Asclepiadoideae) from

Mesoamerica

W. I). Stevens and Olga Martha Montiel

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
stevens@mobot.org; olgamartha.mont iel@ruobot.org

Abstract. Gonolobus taylorianus, a Mesoameri-
can endemic, is described and illustrated.

Key words: Apocynaceae, Asclepiadoideae,
Gonolobus, Mesoamerica.

The genus Gonolobus Michaux has about 150
species ranging from temperate North America to
temperate South America. The greatest diversity is
found in southern Mexico and northern Central
America. The genus is most clearly distinguished
from the other genera of the Gonolobinae by an
extra anther appendage that is always found on the
distal dorsal edge of the anther and is usually
fleshy, of a contrasting color, and more or less pat¬
ent. Additionally, the corona is always a simple,
entire or 5-lobed, fleshy ring at the junction of the
corolla and the gynostegium, there is frequently a
well-developed coralline annulus, the corolla lobes
are frequently strongly overlapping and twisted in
bud, the indumentum is usually comprised of hairs
of one length, and the follicles usually have five
complete wings.

This new species came to light in the process of
preparing floristic accounts for Flora de Nicaragua
and Flora Mesoamericana, and is here described.

Gonolobus taylorianus W. I). Stevens & Montiel.
sp. nov. TYPE: Nicaragua. Matagalpa: along
road from Muy Muy to Esquipulas, ca. 19 km
W of Esquipulas, 12°42'N, 85°45'W, elev. 400
m, 30 July 1978, IE D. Stevens 9563 (holotype,
MO). Figure 1.

Gonolobus taylorianus W. I). Stevens N Montiel; ex af-
finitate G. nigri et speeierum affinium floribus majoribus
et indumento annuli limbique eodem distinguendus.

Vine, twining (to right), woody below, corky bark
absent; latex white, roots fibrous; young stems pu¬
bescent at nodes and in 2 lines on internodes, in¬
dumentum mixed, long hairs multicellular, pale
yellow, reflexed, 0.15—0.4 mm long, short hairs
multicellular, capitate-glandular, pale brown, erect,
0.05—0.1 mm long; internodes 2-16 cm long.
Leaves opposite, without pseudostipules, blades
ovate to elliptic, 5.9-12.5 X 2.2-7. 1 cm, apex acu¬

minate to attenuate, base lobate, lobes divergent to
convergent, sinus 0.3— 1.7 cm deep, glabrous or
with scattered long (0.3-0. 7 mm) and/or glandular
hairs, lateral veins pinnate, 4 to 6 pairs, middle
veins 35—45° to midrib, colleters 2 or 4; petiole
2.2-7 cm long, glabrous or with mixed indumentum
on adaxial side. Inflorescence extra-axillary. 1 per
node, eongested-racemiform, 4- to 18-flowered,
with indumentum of stem or rarely glabrous, sessile
or peduncle l-6(-l3) mm long, axis 1-10 mm long,
pedicel 4—11 mm long, bracts 0.6-2.6 X 0. 1-0.5
mm, linear: calyx tube 0.5-1. 2 mm long, with 1 to
2 colleters per sinus, lobes lanceolate to debate,
with acute tips, reflexed, 2.3-6. 2 X 1.3-2 mm,
green, glabrous or with a few long hairs on margin
and base o’ abaxial side; corolla rotate, aestivation
imbricate and dextrorse, brown, green, or purple,
tube 1.7—2 7 mm long, glabrous or with mixed in¬
dumentum outside, long hairs to 0.1 mm long, in¬
side densely pubescent with glassy, erect or curly,
unicellular hairs 0. 1-0.3 mm long, faucal annulus
tubular, 5-lobed, 0.5-1. 1 mm tall at lobes, 0.3-0.6
mm tall between lobes, outside with same indu¬
mentum as adjacent corolla tube, corolla lobes del¬
taic to lanceolate with acute tips, with conspicuous
to inconspicuous auricles at base, 6.6—1 1.7 X 3.2—
4.7 mm. re lexed, glabrous or with mixed indumen¬
tum outside, inside glabrous to densely pubescent
with glassy erect or curly, unicellular hairs 0.1— 0.3
mm long; gynostegium with stipe 0.3— 0.6 X 0.7—1
mm, 5-ribbed, corona — hidden by faucal annulus,
a 5-lobed disk, adnate to corolla and gynostegium
stipe, 0. 3-0.6 mm thick, 0.6-1. 2 mm wide between
lobes, 0.9- 1.5 mm wide at lobes, yellow, anther
appendages deflexed to nearly patent, spathulate,
0.8-1. 3 mm long, 0.9-1. 3 mm wide at tip. fleshy,
smooth to papillate, orange, terminal appendages
tightly appressed to margin of style apex, 0.2— 0.5
X 1.7-2. 1 mm, green, guide rails straight, parallel,

0.6— 1 .2 X 0.1— 0.3 mm; corpusculum 0.2-0.28 X
0.07-0.13 mm, sagittate, pale brown to black,
translators 0.25-0.33 X 0.1-0.18 mm, broadly
winged, pcllinia 0.8—1.03 X 0.37—0.53 mm, ob-
ovate to pyriform, somewhat asymmetrical, exea-

Novon 12: 551-554. 2002.

552

Novon

Volume 12, Number 4
2002

Stevens & Montiel 553

Gonolobus taylorianus from Mesoamerica

vated, sterile and translucid at attachment; style
apex 2.4— 3.4 mm wide, star-shaped, shallowly con¬
cave, smooth, red-brown. Follicles single, fusiform
with narrow asymmetric base, 11—19 X 4.5—7 cm,
glabrous, spongy, mottled white and [tale green,
smooth or irregularly winged (4 complete wings +

I incomplete distal wing, 2 complete + 2 incom¬
plete wings, or I to several incomplete wings),
wings 2-8 mm wide; seeds obovate, 7—9.3 X 3.7—
5.5 mm, flat, dark gray to gray-brown, verrucose-
reticulate, margin 0.5—1 mm wide, irregularly cren-
ulate on distal 3A-Vi, coma 2.5-5 cm long, white.

This species has most often been annotated as
Gonolobus salvinii Hemsley, but also as G. niger
(Cavanilles) R. Brown ex Schultes, G. edulis Hem¬
sley, G. fraternus Schlechtendahl, G. stenanthus
(Standley) Woodson, G. roeanus L. 0. Williams, as
well as a few generic misidentifications. One col¬
lection, Standley 26259, was even annotated as the
type of a new species of Gonolobus. but the name
was never published. The new species was de¬
scribed in the Flora of Guatemala (Standley & Wil¬
liams, 1969) under the name Gonolobus salvinii,
but examination of the type of that name reveals
that it corresponds to a rare Guatemalan endemic,
quite a different species that is described in that
same publication under the name G. longipetiolatus
Woodson. Gonolobus longipetiolatus will be consid¬
ered a synonym of G. salvinii, while the more com¬
mon and widespread species is here described. The
new species Gonolobus taylorianus was treated as
"Gonolobus sp. D in Flora de Nicaragua (Stevens
et al.. 2001).

Gonolobus taylorianus is related to G. niger, but
that species has long peduncles, much smaller
flowers, purple-black corollas, a low faucal annulus
that is, along with the rest of the corolla, completely
glabrous, and larger follicles that arc completely
smooth or have only a few small fragments of wings,
appearing to be prickles on the distal part of the
follicle; G. niger ranges from northwestern Mexico
to Guatemala. Gonolobus roeanus is also related and
has brown corollas, but has smaller flowers and a
low annulus that has only a terminal fringe of hairs,
while the rest of the annulus and adaxial corolla
surface is glabrous; G. roeanus seems to be known
only from the flowering type collection from Gua¬
temala. This small group of Gonolobus, which in¬
cludes other undescribed species, is unusual in
having a mixed indumentum and follicles that are
not uniformly 5-winged. These characters suggest
a relationship with Matelea, but the dorsal anther
appendages are well developed and there is no

doubt that he group is best accommodated within
Gonolobus.

This new species is variable in a number of char¬
acters, especially corolla color (green, brown, or
purple), flower size, although always much larger
than the related species, distribution of the indu¬
mentum on the corolla (tending to be restricted to
the annulut and limb on southern collections, but
extending onto the lobes on the northern collec¬
tions), the fruit wings (from smooth to various com¬
binations of complete and incomplete wings), and
in ecological requirements (moist, wet. or cloud for¬
ests between near sea level and 1300 m). Gonolobus
taylorianus flowers mostly from July to September,
and fruits mature from January to March. The un¬
usual morp lological variability is perhaps related
to the fact that the species is often cultivated for
its fruits. The developing fruits are eaten as a veg¬
etable, raw when young, cooked when more mature,
or boiled with sugar at any stage, and are available
in markets in Guatemala and El Salvador during
tin; fruiting season (Standley & Williams, 1969).
“Cuchamper” is the most common local name for
the species, and “Siguamper,” “Chinchayote,” and
“Pol la” are also recorded from herbarium labels. It
seems to be primarily cultivated as a dooryard
plant, but long cultivation and selection over a rel¬
atively broad geographic range could be accentu¬
ating the apparent variability of the species.

This species is named in honor of Jack Crawford
Taylor (14 \pril 1922-). influential patron of the
arts and sciences. It is most fitting that this new
species, at once beautiful and useful, is named for
a man whose dedication to preserving and under¬
standing our environment is manifest.

Paratypes. GUATEMALA. Jalapa: along Hwy. 19
about 6.3 mi. SE of Jalapa, 16 July 1971. Stevens 1247
(MO). Guatemala: camino a Humitos, Km 34 Amatitlan,
31 Aug. 1997, MacVean & MacVean 7096 (MO, UVAL).
Santa Rosa: Cerro Gordo, Sep. 1892. Heyde A' Lux ex
Smith 4007 (MO). HONDURAS. Colon: base of Cerro
Piedra Blanca, along Bonito Oriental— Limon road, ca. 5
km NE of Be nito Oriental, ca. 50 m SIS of Rio Piedra
Blanca, 5 Feb. 1993, Evans 1079 (MO); old road to Cas¬
tilla, E of airport to Jericho, 12 Jan. 1981, Saunders 072
(MO, NY). Francisco Morazan: Villa San Roque, 1 Aug.
1950, Standley 26259 (MO); Sabana Grande, 9 Sep. 1945,
Valerio 3205 (MO); Quebrada La Orejona, NE de la
UNAH. 17 A ig. 1978, Zelaya 04 (MO). Olaucho: mar-
genes del Rfo Talgua. 6 km SE de Catacamas, 16 Mar.
1987, Ortega 203 (MO). EL SALVADOR. La Libertad:
San Salvador, Jardfn BoUinico La Laguna, 3 Sep. 1984,
Berendsohn & Berendsohn 197 (MO); San Salvador, Jardfn
Botdnico La Laguna, 20 Sep. 1988, Villacorta 101 (MO);
San Salvador, Jardfn Botdnico La Laguna, 25 July 1989,
Villacorta 343 (LAGU, MO); Mun. Antiguo Cuscutlan,
laderas de La Laguna, casa W de vigilante, 28 Sep. 1989.
Villacorta et al. 450 (B. LAGU, MO). La Paz: Hacienda

554

No von

Santa Marfa, Canton La Lucha, Tecoluca, 13 Oct. 1988,
Herendsohn & R. Calderdn 1 1 <17 (MO). Santa Ana: Santa
Ana, 1928, S. Calderdn 2409 (MO); Santa Ana, 7 mi. NK
of Metapdn on road to Cerro Monte Cristo, 30 July 1977,
Croat 42280 (MO). San Salvador: San Bartolo, llopango,
12 Aug. 1973, Flores 520 (MO). Sonsonate: Cantbn La
Calcra. nacimiento del Bfo Grande de Sonsonate, 21 July
1989, Villacorta et al. 424 (LAGU. MO). NICARAGUA.
Boaeo: summit and upper NW slope of Cerro Mombach-
ito. 18 Jan. 1981. Stevens 18969 (MO). Cliontales: 2.8
km N of Cuapa, 21 Jan. 1978, Stevens 6149 (MO). Estcli:
Salto de Kstanzuela. 13 Aug. I97(>. Hall 7687 (MO); I
km al S de Santa Cruz, eamino a Cuajiniquil, 6 Mar. 1982,
Moreno 15712 (MO); Hacienda Varela, 17 km de Estelf
eamino a El Sauce, 17 Sep. 1982. Moreno 1741 1 (MO).
Granada: NE del Volcdn Mombacho, empalme de los
(‘aminos a Santa Isabel y a Cutirre, 16 Sep. 1980, Moreno
2642 (MO); NE del Volcdn Mombacho, Hacienda Las De-
lieias, 8-9 km sobre la carretera a Cutirre, 16 Sep. 1980,
Moreno 2695 (MO); Volean Mombacho, empalme de los
caminos a Las Delicias y a La Calera, 16 Sep. 1980, Mo¬
reno 2700 (MO); Volcdn Mombacho, La Esmeralda, 19
Mar. 1982, Moreno 15945 (MO). Vladriz: Cerro Volean
Somoto (Tepesomoto), lado Oeste, Finca San Martin, 25
Sep. 1980, Moreno 2964 (MO). Matagalpa: Ouebrada
Malacal. Hacienda La Bonanza, 20 km de Matagalpa, 21
Jan. 1982, Castro 2405A (MO); along Hwy. 5, 24.6 km

from Hwy. 3 intersection, 8.6 km SW of Bfo Tuma bridge,
26 Jan. 1982, Stevens A- Montiel 21458 (MO). Rivas: lsla
Ometepe, Volcdn Concepcion, Los Hatillos, eamino a Las
Delicias, 9 Aug. 1984, Robleto 990 (MO). Zelaya: be¬
tween Cerro Saslaya and San Josb del Hormiguero, 13
Mar. 1978, Stevens 7068 (MO); vicinity of Wan i. Bfo Uli,
15 Mar. 1978, Stevens 7271 (MO). COSTA BICA. Guan-
aeaste: Santa Bosa National Park, 6 Jan. 1980, Janzen
11874 (MO). Puntarenas: Guru, 2 Sep. 1995, Sanders et
al. 17751 (MO, UCB).

Acknowledgments. Michael Gilbert kindly
loaned his expertise in the Asclepiadaceae in ex¬
amining the type of Gonolobus salvinii. Alba Ar-
belaez prepared the drawing of this complicated
species with her usual skills. Victoria Hollowell
and Amy Pool suggested key improvements in the
language.

I .iterature Cited

Standley, P. C. & L. O. Williams. 1969. Flora of Guate¬
mala. Asclepiadaceae. Fieldiana, Bot. 24(8, part 4);
407-472.

Stevens, W. I)., C. Ulloa U„ A. Pool & O. M. Montiel.
2001. Flora de Nicaragua. Monogr. Syst. Bot. Missouri
Bot. Card. 85: i-xlii, 1—2666.

Rubiacearum Americanarum Magna Hama Pars IX. New Species
and a New Combination in Hippotis and Pentagonia (Hippotideae)
from Central and Western South America

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166. U.S.A.
Charlotte.Taylor@mobot.org

ABSTRACT. The following new species are de¬
scribed: Hippotis stellata C. M. Taylor N Rova.
found from eastern Panama to northwestern Ecua¬
dor and distinguished by its closely grouped or pos¬
sibly stellate trichomes; Pentagonia angustifolia C.
M. Taylor, found in western Panama and formerly
confused with P. nuciformis Dwyer; P. dwyeriana C.
M. T aylcr, found in western Panama and formerly
confused with P. macrophylla Bentham; Pentagonia
monocaulis C. M. Taylor, found in eastern Costa
Rica and formerly confused with P. donnell-smithii
(Standley) Standley; and Pentagonia sanblasensis
C. M. Taylor, found in central Panama and also
formerly confused with P. macrophylla. The new
combination H. panamensis (Dwyer) C. M. Taylor is
based on Duroia panamensis Dwyer, and a descrip¬
tion is presented for this poorly known species,
which is actually the most commonly collected spe¬
cies of Hippotis in Central America and adjacent
northwestern Colombia.

Key words: Hippotideae, Hippotis, neotropics,
Pentagonia , Rubiaceae.

The Neotropical genus Hippotis Ruiz & Pavon
(Hippotideae; Robbrecht, 1993; Rova & Anders-
son. 1995) comprises about 11 species of shrubs
and small trees found in wet lowland forests from
Nicaragua to Bolivia. This genus is distinguished
in the Rubiaceae by the combination of its lack of
raphides; its interpetiolar stipules that are convo¬
lute or induplicate-valvate with the resulting struc¬
ture often twisted again; its leaves with the higher
order venation striolate or lineolate (i.e., very close¬
ly set and parallel within the areoles; Rova & An-
dersson, 1995); its axillary, usually solitary flowers;
its well-developed calyx limbs that are spathaceous
(i.e., completely fused in bud forming a closed
structure that is split along one side or sometimes
irregularly by the elongating corolla); its funnelform
corollas with five valvate or reduplicate-valvate
lobes; its baccate fleshy fruits with the large calyx
limb usually persistent; and its seeds that are an¬
gled. generally smooth, and numerous on placentas

that are axile at the base and parietal above (Rova
& Andersson, 1995). The corollas are often rather
zygomorph c with the tube broadly curved. Hippotis
was last studied as a whole by Steyermark (1965);
this work is now somewhat dated.

The Neotropical genus Pentagonia Bentham
(Uip| lotideae; Robbrecht, 1993; Rova & Anders¬
son. 1995) comprises about 25 to 30 species of
shrubs and small trees found in wet lowland to pre-
montane forests from Guatemala to Peru, in both
primary ard secondary vegetation. This genus is
distinguished in the Rubiaceae by the combination
of its fleshy, relatively large vegetative and repro¬
ductive structures; its lack of raphides; its inter¬
petiolar coivolute stipules; its leaves with the ab-
axial and often also the adaxial surfaces finely
striate (described and illustrated in detail by Rova
& Andersson, 1995); its axillary, cymose to sub-
capitate or glomerulate, sessile to usually shortly
pedunculate inflorescences; its calyx limb that is
usually five-lobed with the lobes often well devel¬
oped and imbricated in bud, or alternatively this
limb is sometimes spathaceous; its corollas with
well-develcped tubes and five valvate lobes; its five
included stamens with the filaments pubescent on
their bases, its baccate, coriaceous to rather woody,
usually lenticellate, generally subglobose fruits;
and its seeds that are angled, smooth on the sides,
and numerous on fleshy placentas that are axile at
the base aid parietal above (Rova & Andersson,
1995). Plants of Pentagonia usually contain gen¬
erous amounts of mucilaginous latex or sap. which
is visible in dried specimens as elastic filaments
that stretch between broken edges of most tissues.
The flowers are often slightly zygomorphic in their
broadly curved tubes and the arrangement of their
lobes and anthers, with the anthers usually grouped
on the lower side of the flower and imbricated end-
to-end because of the twisting and thus relative
shortening of the filaments from the upper side of
the flower.

Several morphological structures found in some

Novon 12; 555—562. 2002.

556

Novon

species of Pentagonia are unusual in the Rubi-
aceae. The most striking of these is pinnatifid leaf
shape (e.g., P. tinajita Seemann, P. lobata C. M.
laylor). Pinnatifid leaves may be lobed shallowly
(e.g., some plants of P. tinajita ) to essentially com¬
pletely, with the blade tissue between the lobes
sometimes reduced to a ridge of tissue along the
costa (e.g., some plants of P. alba Dwyer and I1,
pinnatifida Seemann). Pinnatifid leaves are infre¬
quent in the Rubiaceae, and are otherwise known
in the neotropics only in occasional plants of some
species of Simira. Pinnatifid Pentagonia species
are most numerous in Panama and Costa Rica.
Some species of Pentagonia have a “trash bucket”
habit, with sessile or subsessile leaves that are
rounded to auriculate at the base. These leaf bases
form a cup or platform around the stem where leaf
litter accumulates (e.g., P. wendlandii Hooker f.).
Occasional herbarium specimens have small ad¬
ventitious roots arising from the stems just above
the nodes; these probably grew into the decaying
accumulated litter. A number of species of Penta¬
gonia are striking also in their unbranched or mon-
ocaulous habit (e.g., P. nuicrophylla Rentham).

I hese plants may be as much as 3.5 m tall, and
this monocaulous habit is loosely correlated with
leaves that are relatively large. The leaves of these
monocaulous species may be as much as 1.2 m long
(e.g.. P. magnifica K. Krause, P. gigantifolia
Ducke). This aspect of the habit is here considered
a generally consistent character for species of Pen¬
tagonia; some taxonomists have considered it var¬
iable within a species. Species of Pentagonia also
show a notable variability in the development of
their inflorescence bracts: in general, these are ei¬
ther relatively reduced to absent (to I mm long;
e.g., P. parvifolia Steyermark), or quite well devel¬
oped (5—30 mm long, e.g., P. macrophylla, P. wen¬
dlandii). Degree of development of the inflores¬
cence bracts is also here considered a generally
consistent character within species of Pentagonia.

Standley (1914b) presented a taxonomic history
of this genus and noted that the name Pentagonia
Rentham which was applied to these plants was
illegitimate because it had been used twice previ¬
ously, and that Watsonamra Kuntze was the first
valid and legitimate name for them. At the same
time Standley (1914a) published the genus name
Nothophlelna Standley for a Costa Rican species
that he distinguished from Watsonamra by its eam-
panulate, only shallowly lobed calyx limb. Subse¬
quently, the name Pentagonia Rentham was con¬
served for this group, and Nothophlelna was
reduced to synonymy of Pentagonia (Burger & Tay¬
lor, 1993) as the relatively wide variation in this

group’s calyx morphology became evident. Penta¬
gonia has not been studied as a whole since Stan-
dley’s (1914b) article and is relatively poorly
known. The large fleshy plants are difficult to pre-
pan* as good herbarium specimens. Most species of
Pentagonia are locally uncommon in Central Amer¬
ica and Colombia (pers. obs.; of. Ernst, 1989). Eittle
is known about population-level morphological var¬
iation. though study of this would be useful for the
characters considered taxonomically informative
(e.g., pubescence, corolla, and fruit sizes). Freeman
and Stiles (1990), Stiles and Freeman (1993), and
Janzen (1971) noted that Pentagonia flowers pro¬
duce significant nectar, and in a lowland forest in
Costa Rica are visited by hermit hummingbirds,
large bees, and lepidopterans. The floral biology of
Pentagonia macrophylla was studied by McDade
(1986), who found the flowers to be protandrous,
sequentially unisexual in function, and strongly
synchronized among the plants of a single popula¬
tion.

The new species of Hippotis and Pentagonia de¬
scribed below were discovered during preparation
of the Rubiaceae treatment for the Flora Mesoam-
ericana.

New Spkcies and a New Combination

Hippotis pannmcnsis (Dwyer) C. M. Taylor, comb,
nov. Rasionym: Duroia panamensis Dwyer,
Ann. Missouri Rot. Card. 55: 138. 1968.
TYPE: I ’anama. Bocas del loro: Duwebdulup
Peak, N of R. Terebe across from W. Huron
(behind chiefs house), 300-900 m, 13 Apr.
1968. ,/. II. Kirkbride, Jr. & ,/. A. Duke 571
(holotype, MO-1968346; isotypes, MO-
1968345, PM A).

'frees or treelets to 17 m tall; stems pilose to
hirsute, h'aves elliptic to usually obovate, 12-35 X
5.5-18 cm, at apex acuminate, at base cuneate to
subtruncate, drying papyraceous to chartaceous, on
both surfaces pilose to hirsute; secondary veins 7
to 10 pairs, not looping to interconnect, without
domatia; petioles 1—3.5 cm long; stipules elliptic: to
lanceolate, 1.5—3 cm long, obtuse to acute. Flowers
solitary, ebracteate; peduncles 1-10 mm long; hy-
panthium ellipsoid, ca. 5 mm long, densely seri¬
ceous; calyx limb spathaceous, 2.5-4 cm long, hir¬
sute*. apex entire and acute* to irregularly 2- or
3-lobed, leebes acute; corolla funnelform, white to
cream, externally sericeous to hirsute, internally
glabrems. tube 35—45 mm long, lobes 5, triangular,
5—8 mm long, aeute to obtuse; anthers ea. 3 mm
long, filaments hirsute at base; stigmas ea. 2 mm

Volume 12, Number 4
2002

Taylor

Hippotis and Pentagonia

557

long. Fruits ellipsoid, 3—4 X 1—3 cm, hirsute to
pilose; seeds ea. 2 mm long.

This species is found in wet forests at 0—900 m
elevation from Nicaragua to northeastern Colombia;
it has been collected with flowers most frequently
in February and also in March, April, August
through October, and December, and in fruit in
February and April through October.

Dwyer (1980) described this species in Duroia
Auhlet probably based mainly on its general as¬
pect, the hirsute to pilose pubescence of all its veg¬
etative organs, its externally sericeous corollas, and
its usually solitary, rather large (i.e., several cm in
diameter), baccate fruits. However, his species
clearly belongs instead to Hippotis as shown by its
interpetiolar, induplicate-valvate stipules (vs. fully
fused into a circumscissile calyptrate structure in
Duroia ), its lineolate higher-order leaf venation (vs.
irregularly reticulated in Duroia), its bisexual flow¬
ers (vs. unisexual on dioecious plants in Duroia),
its spathaceous calyx limb (vs. tubular and truncate
to regularly lobed in Duroia). and its five corolla
lobes (vs. six to nine in Duroia).

Dwyer (1980) treated this species together with
the following, Hippotis stellata, under the name
Hippotis albiflora H. Karsten. However, //. albiflora
is a species restricted to Colombia and Venezuela
(Steyermark. 1965). Because of this confusion and
also the limited usefulness of the original descrip¬
tion of Duroia panamensis, a complete description
and a list of representative specimens an* presented
here. Hippotis panamensis is distinguished from the
other species in this genus by the combination of
its spreading pubescence on the vegetative organs
and fruits, its stipules 1.5—3 cm long, its petioles
1—3.5 cm long, its solitary flowers with peduncles
1 — 10 mm long, its relatively long calyx limbs, and
its relatively long white corollas. Hippotis albiflora
differs in its appressed pubescence, its (lowers 1 to
3 per leaf axil on pedicels 7-1 1 mm long, its calyx
limbs lf>— 23 mm long, and its corollas 35—42 mm
long.

Representative specimens. COLOMBIA. Choco:
Barque National <le Utrfa, en la falta de la serranfa que
bordea el Rfo San Pichf, F. Garcia G. & Agualimpia 470
(MO). COSTA RICA. Cartago: 24 kin NE of Turrialba on
highway to Limon, then E at Tres Equis on jeep road 1.5
km, Liesner et al. 15379 (MO). Heredia: Finea La Selva,
the OTS field station on the Rio Puerto Viejo just E of its
junction with the Rio Sarapiquf, along W Boundary Trail,
2900 m line, Grayum 2352 (MO). Limon: Cerro Coronel,
E of Laguna Danto, Stevens & Montiel 24363 (MO, PTBG).
NICARAGUA. Kio San Juan: sobre el Rfo Sabalo. R. R.
Moreno A Roblelo 26007 (MO). Zelaya: along road to Co-
Ionia Yolaina, Colonia La Esperanza, etc. [sic], ca. 1.3 km
SE of intersection with road between Nueva Cuinea and

Colonia Verdun, immediately upriver from bridge over
Cano Sardira, Stevens 6306 (MO). PANAMA. Boras del
Toro: near highway to Chiriquf Grande, 10 road-mi. from
continental divide and about 2 road-mi. along road E of
highway, MiPherson 11822 (MO).

Hippotis stellata C. M. Taylor & J. 11. F. Rova,
sp. nov. TYPE: Panama. Darien: Parque Na-
cional del Darien, ridge between N and S
branches of Rfo Pucuro, in forest N of old vil¬
lage of Tacarcuna, ca. 18 km N of Pucuro,
8°05'N, 77°16'W, 600-800 m, 24 Oct. 1987.
/>. Hummel, G. de Nevers, H. Cuadros & //.
Herrera 16473 (holotype, PMA; isotypes, MO-
3607390, PTBG-7273). Figure 1.

Haec spe des a congeneris foliis calyce fnictuque pu-
bescentia plerumque adpressa vestitis, limbo foliari sub-
tus secus venationem tertiariam trichomatibus ramosis ex
ramulo centrali elongato etiam ramulis basalibus brevibus
numerosis constantibus induto, limbo calycino 30—10 nun
longo atque eorollae albae tubo ca. 50 mm longo ac lobulis
10—12 mm longis distinguitur.

Shrubs < r small trees to 15 m tall; stems densely
strigillose o sericeous, leaves elliptic to obovate,
13—35 X 6-18 cm. at apex acuminate, at base ob¬
tuse to rounded, drying papyraceous to eharta-
ceous, adaxially strigillose and scabridulous, abax-
ially strigil ose and hirtellous with trichomes mixed
simple and sessile-stellate; secondary veins 8 to 10
pairs, not to weakly looping to interconnect, without
domatia or rarely with crypt-type domatia; petioles
1—3 cm lor g; stipules lanceolate-oblong, 9-16 mm
long, obtuse. Flowers solitary; peduncles 4—6 mm
long, densely strigillose to sericeous; hypanthium
turbinate, (> — 7 mm long, densely velutinous; calyx
limb 30—40 mm long, densely strigillose, spatha¬
ceous and acute, irregularly and shortly 1- to 4-
lobed; corolla funnelform, white, externally mod¬
erately to densely sericeous, internally glabrous,
tube ca. 50 mm long, lobes ligulate, 10—12 mm
long, obtuse to rounded; anthers and stigmas not
seen. Fruits ellipsoid, ca. 3X2 cm, densely stri¬
gillose, brown.

Distribution, habitat, and phenology. In wet
forest at 350—1500 m, eastern Panama through
western coastal Colombia to northwestern Ecuador;
collected with flowers in February, April, June, Oc¬
tober, and November, with fruits in February, April,
May, July, and September.

This nev/ species is distinguished from other
Hippotis by the unusual trichomes found on its leaf
undersides, which are unlike the trichomes known
from other Hippotis species: these are apparently
stellate or multiradiate with one large central arm
and 3 to 20 shorter arms arranged generally in a

558

Novon

Figure I. Hippotis stellata C. M. Taylor & Rova. — A. Fruiting branch. — I?. Flower. — C. 1), E. Branched trichomes
from abaxial leaf surface. A. based on Orozco et al. 2347', I?, C. I). E, based on Hummel et al. 16473. A. B to 5-cm
scale; C. 1). E to 1-mm scale.

basal rosette. Each of the arms is unicellular, 'flic
species epithet refers to these trichomes. Individual
plants of some other species of Hippotis (e.g., H.
paruimensis) have a mixture of short and long tri¬
chomes on their leal undersides, and these are oc¬
casionally grouped to almost fasciculate. It seems
likely that the apparently stellate trichomes found
in H. stellata are formed from the fusion among
these grouped unequal trichomes rather than by
branching of individual trichomes. Such stellate or
closely grouped trichomes are otherwise unreported
in Rubiaceae (Robbrecht, 1988).

This new species is also distinguished by its ca¬
lyx limbs 30-40 mm long and its white corollas
with tubes ca. 50 mm long and lobes 10—12 mm
long. Dwyer (1980) included this species together
with Hippotis panamensis in his circumscription of
H. albijlora, as discussed above under //. pana¬
mensis. However. //. albijlora differs from this new
species in its pedicels 7-1 I mm long, its calyx limb
16-23 mm long, and its corollas 35—42 mm long.
Also similar to H. stellata is II. grandiflora Sley-

ermark from Pacific coastal Colombia; //. grandi¬
flora differs from this new species in its stipules
30—35 mm long, its petioles 6—10 mm long, its
bracteoles 30 X 10-15 mm long, its calyx limb
with the tube 43—47 mm long and lobes 6—10 mm
long, and its corolla ca. 80 mm long.

Paratypes. COLOMBIA. Antioquia: mpio. Frontino,
correg. La Blanquita, region de Murrf, via Nutibara-La
Blanquita, 14.5 km 0 de Nutibara, 15—16 km del Alto de
Cuevas-La Blanquita, Callejas et al. 6762 (HIJA, MO);
Parque Nacional Natural “Las Orqu ideas,” Sector Vena-
dos, Cogollo et id. 2865 (JAUM, MO); Murrf, La Blan¬
quita, Rfo Murrf, hills above village, Gentry et id. 75739
(MO); mpio. Coconut. vereda La Pinuela, carretera a San
Francisco, Giraldo-CaHas 916 (HUA, MO); mpio. Fronti¬
no, vereda Venados, Parque Nacional Natural “Las Or-
qufdeas,” Quebrada Las Manzanares, Pipoly el al. 1821 1
(JAUM, MO). Choco: mpio. Nuquf, correg. Termales,
Quebrada Piedra Piedra, Acevedo- Rodriguez et al. 6783
(MO. US); NW of Alto Curiche, Duke 11258 (MO). Kis-
ars dda : mpio. de Mistrato, correg. de Puerto de Oro, ver¬
eda Chirrineha, Finca La Cilia, G. I. Orozco et al. 2347
(COL, MO). ECUADOR. Carchi: canton Tulcan, Reserva
Etnica Awa, parroquia El Chical, Centro Gualpi Medio,

Volume 12, Number 4
2002

Taylor

Hippotis .and Pentagonia

559

Rio Cabumi, A. Grijalva et al. 591 (MO); canton lulcan,
around encampment in Gualpi Chico area of Awa Reser¬
vation, NW and SE, Hoover et al. 5695 (MO); canton Tul-
can, Reserva Etnica Awa, parroquia El Chical, Centro San
Marcos, P. Mdrulez et al. .373 (MO); canton Tulcan, Re¬
serva Indfgena Awa, comunidad San Marcos, 25 km al
NW de El Chical, parroquia Maldonado, I). Rubio et al.
1051 (MO). PANAMA. Sail Bias: Pemasky, Sendero Ina
I gar. R. Peralta 60H (MO).

Pentagonia angustifolia C. M. Taylor, sp. nov.
TYPE: Panama. Veraguas: on Caribbean slope
above Rfo Primero Brazo 5 mi. NW ol Santa
Fe, 700-1200 m, 18-19 Mar. 1973, R. L. Lies-
ner 993 (holotype, MO-2257090). Figure 2C.

Haec species a Pentagonia nuciformi habitu monocauli
breviore, a Pentagonia monocauli foliis angustis 10-16
cm latis atque limbo calycino glabrescente 17-18 mm lon-
go distinguitur.

Slender trees to 4 m tall, unbranched; stems gla-
brescent. Leaves entire, narrowly oblanceolate to
narrowly ligulate or elliptic-oblong, 35-50 X 10—
16 cm, at apex acute to acuminate, at base acute
to attenuate or obtuse to rounded, drying charta-
ceous, adaxially glabrous, abaxially glabrescent or
sometimes strigillose on principal veins; secondary
veins I I to 13 pairs; petioles 5-8.5 cm; stipules
lanceolate to ovate, 2. 5-4.8 cm long, acute to acu¬
minate, smooth, abaxially and adaxially glabrous.
Inflorescences congested-cymose, puberulous to gla¬
brescent; peduncles 5—7 cm long; bracts reduced
or absent; pedicels 2-3 mm long; flowers ca. 7, ped¬
icellate; hypanthium turbinate, ca. 5 mm long, gla¬
brescent; calyx limb 17-18 mm long, glabrescent,
color not noted (probably green), lobed for
lobes 5. elliptic, ciliolate, obtuse to rounded, some¬
what cucullate; corolla in bud tubular, color not
noted, externally glabrescent, internally not seen,
tube to 22 mm long, lobes 5, debate, to 5 mm long,
acute; anthers and stigmas not seen, bruits several,
subglobose, 25—30 mm diam., densely lenticellate,
yellow, glabrescent; seeds not seen.

Habitat, distribution, and phenology. In wet
forest at 450-1200 m in western Panama; collected
in flower in March and August, in fruit in May and
October.

This new' species differs from Pentagonia nuci-
f or mis Dwyer by its relatively short monocaulous
habit, its pedicellate flowers, and its longer calyx
limbs; plants of P. nuciformis branch and frequently
are up to 15 m tall, w ith sessile or subsessile flow¬
ers and calyx limbs 5-8 mm long. Pentagonia an¬
gustifolia is similar in habit to P. monocaulis (de¬
scribed below), but P. monocaulis can be quickly
distinguished by its broader leaves, 26— 36 cm

wide. The specific epithet of this new species refers
to its relatively narrow leaves. The flowers of the
type collection may be mature, but more likely
these are well-developed buds that partially and
premature lv opened after collection.

Paratypts. PANAMA. Veraguas: valley of Rfo Dos
Bocas along road between Escuela Agricola Alto Piedra
and Calov<4)ora, 15.6 km NW of Santa F'e, Croat 27561
(MO); Cerro Tute, W of Santa Ee, beyond Alto de Piedra,
McPherson 7187 (MO); NW of Santa Ee. 8.8 km from
Escuela Agricola Alto de Piedra, Pacific slope, Mori <&
Kallunki 6 184 (MO).

Pentagonia dwyeriana C. M. Taylor, sp. nov.
TYPE; Panama. Code: bills above road 18 km
past Sardinilla on way to Nombre de Dios
(road not finished), 150—300 m, 2 Aug. 1974,
V H. Croat 26123 (holotype, MO-2205421).
Figuie 2A, B.

Haec species a Pentagonia macrophylla bracleis acutis
anguste triangularibus usque lingulatis distinguitur.

Small trees to 5 m tall, branching pattern not
noted; steins densely pilosulous or hirtellous some¬
times becoming glabrescent. Leaves entire, oblan¬
ceolate to narrowly elliptic-oblong, 35—65 X 12—
25 cm, at apex acute to somewhat acuminate, at
base obtuse to rounded, drying chartaceous, adax¬
ially and ; baxially moderately to densely pilosulous
or hirtellous; secondary veins 13 to 18 pairs; peti¬
oles 6.5—10.5 cm long; stipules narrowly triangular,
32-40 mm long, acute, smooth, adaxially sericeous
to glabrescent, abaxially densely sericeous. Inflo¬
rescences glomerate to congested-cymose, densely
strigillose: peduncles 0-10 mm long; bracts nar¬
row lv triangular to lanceolate, 1.5—3 X 0.3— 0.8 cm,
acute to long-acute; flowers sessile or subsessile;
hypanthium turbinate, ca. 6 mm long, densely se¬
riceous; cilyx limb ca. 15 mm long, densely stri¬
gillose to sericeous, lobed for ca. Vi, perhaps yellow,
lobes 5, elliptic to ovate, obtuse to rounded; corolla
tubular, p ile yellow, externally densely velutinous
except glabrous in basal V4-V3, internally not seen,
tube ca. 30 mm long, lobes 5, narrowly triangular
to ligulate, ca. 6 mm long, acute; anthers and stig¬
mas not seen. Fruits several, subglobose to ellip¬
soid, 2—3 X 2-2.5 cm, yellow, sparsely lenticellate,
glabrescent, with bracts persisting and pink; seeds
ca. 4 mm long.

Habitat distribution, and phenology. In wet
forest at 150-700 m in western Panama; collected
in flower in August, in fruit in January and August.

I'liis new species is similar to Pentagonia ma¬
crophylla Bentham, but it is easily separated from
that by ils narrower, acute inflorescence bracts.

560

Novon

figure 2. A, B. Pentagonia dwyeriuna (.. M. laylor. based on (.rout 26123. — A. Part of stem with infrurteseenee.
— B. Flower partly dissected with (left to right) bract, calyx, and corolla. — C. Pentagonia angustifolia C. M. Taylor,
based on Liesner 993. I), K. Pentagonia sanblasensis C. M. Taylor, based on Sytsrna & Andersson 4500. — I). Part of
stern with inflorescence. E. Corolla, partly dissected, h G, H. Pentagonia monocaulis C. M. Taylor, based on Croat
35720. — F. Part of stem with infructescence and broken stipule. — G. Part of stem with inflorescence. — H. Flower.
A, C, I), G to 5-cm stale; B, E, H to 1-cm scale.

Pentagonia macrophylla lias obtuse to rounded
bracts (>-15 mm wide, and also differs from this
new species in its usually fewer secondary leaf
veins (10 to 14 pairs) and its calyx limb lobed for
% or more of its length. The epithet of this new
species commemorates John I). Dwyer, who pre¬

pared the Rubiaceae treatment for the Flora of
Panama (Dwyer. 1980).

Paratypes. PANAMA. Colon: N of Diamante, ridge
NW of abandoned mine on Quebrada dc la Mina. II. W.
Churchill A- de Nevers 4213 (MO), 4214 (MO, PM A).

Volume 12, Number 4
2002

Taylor

Hippotis and Pentagonia

561

Pcnlugoiiiu monocaulis C. M. Taylor, sp. nov.
TYPE: Costa Rica. Heredia: near Puerto Viejo
along road near Rio Sucio, 20 m. 27 May
1976, T. B. Croat 35720 (holotype, MO-
2381 703). Figure 2K. G, H.

Haec species a Pentagonia donnell-smithii habitu sim-
plici, limbo calycino dense strigilloso atque corolla cre-
mosa luteave extus dense velutino-tomentulosa distingui-
tur.

Shrubs or small trees to 3.5 m tall, unbranched
or perhaps sparsely branched; stems pilosulous or
puberulous to glabrescent. Leaves entire, obovate to
broadly elliptic, 30-63 X 26-36 cm, at apex acute
to obtuse or somewhat acuminate, at base obtuse to
rounded or truncate, drying chartaceous to subco-
riaeeous, adaxially glabrescent, abaxially strigillose
to glabrescent on the lamina and moderately to
densely strigillose on the principal veins; secondary
veins 12 to 16 pairs; petioles 3-15 cm long; stip¬
ules ligulate to lanceolate, 3.5— 7.5 cm long, acute,
adaxially glabrous, abaxially moderately to densely
strigillose to glabrescent. Inflorescences subcapitate
to eongested-eymose, strigillose to glabrescent; pe¬
duncles 0.5-2 cm long; axes conspicuously lenti-
cellate; bracts none or reduced, up to I mm long,
rounded; pedicels 0-3 mm long; flowers sessile to
shortly pedicellate; hypanthium cylindrical to ellip¬
soid. 4—5 mm long, densely strigillose; calyx limb
10—15 mm long, densely strigillose, green, lobed
for V3-V2. lobes 5, elliptic to ligulate. obtuse to
rounded, entire to sparsely eiliolate, frequently
somewhat cucullate; corolla tubular, cream to usu¬
ally yellow, externally densely velutinous-tomentu-
lose except glabrous near base, internally glabrous
except densely hirtellous just above stamen inser¬
tion. tube 28-34 mm long, lobes 5, narrowly tri¬
angular or lanceolate, 8—10 mm long, acute: an¬
thers ca. 3.5 mm long, positioned variously % of
length of corolla tube above base to just below co¬
rolla throat, filaments ca. 12 mm long, villous on
bases; stigmas ca. 3.5 mm long, positioned near
anthers. Fruits several, subglobose to ellipsoid. 20—
25 mm diam., glabrescent, lentieellate, brown to
orange; seeds ca. 2.3 nun long.

Habitat, distribution , and phenology. In wet
forests at 0—900 m. Caribbean slopes of Costa Rica;
collected in (lower in January, March through July,
and October, in fruit in January, July, September,
and November.

This new species is similar to Pentagonia don¬
nell-smithii (Standley) Standley (Burger & Taylor,
1993; Taylor, 2001); it differs from P. donnell-smi¬
thii in its low unbranched habit, its densely stri¬

gillose calyx limbs, and its externally densely ve-
lutinous-tomentulose, cream to yellow corollas. In
contrast. P. donnell-smithii has a branched habit up
to 15 m tall, glabrous calyx limbs, and externally
glabrous, white to pink corollas. The specific epi¬
thet of P. monocaulis refers to its habit.

Paratypes. COSTA RICA. Heredia: Barque Nat ional
Braulio Carrillo, estacion Magsasay, G. Carballo 91 (INB.
MO); cant6n de Sarapiquf, Rio Sarapiquf near Puerto Vie¬
jo. Estacidn Biologica La Selva, A. bait re 6 (INB. MO),
Gentry A- Ortiz 78595 (MO), /.. ,/. Poveda 697 (MO), I).
Smith 538 DUKE, MO); canton de Sarapiquf, cuenca del
Sarapiquf, La Virgen, margen del Rfo San Ramon, earre-
terea al Tirimbina, A. Rodriguez & Fernandez 1413 (INB.
MO). Linicn: cantdn de Pococi, Refugio Nat ional Barra
del Colorado, Llanura de Tortuguero, Sardinas, F Araya
et al. 236 ilNB, MO); between Rfo Chirripocito and Rfo
Sardina (“Sardinal”), Grayum 9784 (CR, MO); Guapiles,
tin road from Guapiles to Rfo Chirripo. Barringer & Go-
rnez-Lauritc 2369 (MO); hills 3.4 airline km S of Islas
Buena Vista in the Rfo Colorado, 16 airline km SW of
Barra del C olorado, Davidse & Herrera 31280 (MO): path
from Rfo Sacio to Gonzalez farm. Braulio Carrillo, /.. D.
Gomez et al. 22736 (MO); Cerro Coronel, E of Laguna
Danto, Stevens 23702 (MO); E of Rfo Zapote, Stevens
23944 (MO), 24300 (MO), Stevens et al. 24677 (MO); Bar¬
ra tlel Colorado, N side, between town anti ocean beach,
Stevens 24099 (MO). NICARAGUA. Rio San Juan: enlre
el Pueblo t e San Juan tlel Norte Nuevo y la easa tie Ra¬
mon Castillo, viajando por el cano San Juanillo, Rueda et
al. 1862 (MO).

Pentagonia sanblasensis C. M. Taylor, sp. nov.
TYPE: Panama. San Bias: El Elano— Cartf
roatl, 12 mi. from Pan American Hwy., 350—
400 m, 10 May 1981. K. Sytsma & L. Anders-
son 4500 (holotype, MO-2929899), figure
21). F.

Haec species a congeneris pubescentia dense velutina,
foliis satis amplis, inflorescentia subsessili glomerata, ca-
Ivcis limbo profunde lobato 12-20 mm longo ac lobulis
aeutis atqus fructu luteo 22-25 mm in diametro tlistin-
guitur.

Small trees to 5 111 tall, possibly mouocaulous;
stems glabrescent, a little to markedly quadrangu¬
lar anti sometimes strongly channeled on sides.
halves nairowly elliptic-oblong to elliptic or oblan-
ceolate, 30-82 X 12—32 cm, at apex obtuse to
acute, at base obtuse to rounded or truncate, drying
chartaceous to subcoriaceous, adaxially glabres¬
cent. abaxially densely velutinous; secondary veins
12 to 21 pairs; petioles 3.5-9 cm long; stipules
narrowly triangular to lanceolate, 1.5—5 cm long,
acute, smooth or sometimes with midrib thickened,
adaxially glabrous, abaxially densely strigillose. In¬
florescences glomerate, velutinous or pilosulous to
glabrescei t; peduncles 0—8 mm long; bracts absent
or minute; flowers sessile" or subsessile; hypanthium

562

Novon

turbinate, 5— 6 mm long, densely velutinous; ealyx
limb ca. 12 mm long, densely velutinous, lobed for
% to nearly completely, green, lobes 5, lanceolate
to ovate, acute; corolla tubular-funnelform, white to
yellow, externally densely velutinous except gla¬
brous in basal VS, internally glabrous except pu-
berulous in basal VS. tube ca. 43 mm long, lobes 5,
narrowly triangular, ca. 10 mm long, acute; anthers
ca. 4 mm long, positioned ca. % of length of corolla
tube above its base, filaments 15—17 mm long, vil¬
lous on bases; stigmas ca. 2 mm long, situated near
anthers. Fruits several, subglobose to ovoid, 22-25
mm diam., velutinous to glabrescent, yellow; ma¬
ture seeds not seen.

Habitat, distribution, and phenology. In wet
forests at 0—400 m in central Panama; collected in
flower in May and June, in fruit in March, April,
and September.

This new species is distinguished by its combi¬
nation of densely velutinous pubescence, relatively
large leaves (even for Pentagonia, i.e., to I m or
more long), glomerate subsessile inflorescences,
deeply lobed calyx limbs 12-20 mm long, acute
calyx lobes, and yellow fruits 22-25 mm in diam¬
eter. I he specific epithet refers to the only region
from which this new species is known. Similar spe¬
cies are Pentagonia macrophylla and P. duyeriana,
which both differ from P. sanblasensis in their well-
developed red inflorescence bracts 5—20 mm long,
and P. sprucei Standley, which differs in its sub-
truncate to shallowly lobed calyx limbs 10-12 mm
long with the lobes obtuse to rounded.

Paratypes. PANAMA. San Bias: Kl Llano— Cartf road,
19 kin from Interamerican Hwy., de Nevers & Herman
3830 (MO, PMA); Km 27.6, Rfo Pingandi, de Nevers el
al. 5027 (MO); Km 26.5, along Rfo Cartf Chico, de Nevers
et al. 5544 (MO, I’M A), 5831 (MO sterile, PMA), 7830
(MO); Cangandf, de Nevers 5698 (MO sterile); Rfo Can-
gandf, camino entre el pueblo Cangandf y el aeropuerto
de Mandinga, II. Herrera 107 (MO. PMA); frente a la Isla
Miria Ubigandup, camino Sangandi, H. Herrera & Harris
321 (MO); 34—38 km from Pan-American Hwy. on Kl Lla-
no-Cartf road, Knapp & Schmalzel 5466 (MO. PMA).

Acknowledgments. I thank the curators of

DUKE, F, and PMA for access to specimens and
information, and in particular Mireya Correa; Johan
H. E. Rova for generously providing much infor¬
mation and many helpful comments; Roy E. Gereau
for preparation of the Latin diagnoses; L. Anders-
son for much helpful advice and providing addi¬
tional data; 0. M. Montiel and R. Magill for signif¬
icant encouragement of this work; and Robin
foster, Tyana Wachter, and a grant from the Andrew
Mellon Foundation to the Field Museum of Natural
History for support for travel to consult that collec¬
tion.

Literature Cited

Burger, \\. C. & C. M. Taylor. 1993. Flora Costaricensis:
Family #202. Rubiaceae. Fieldiana, Bot. n.s. 33: 1 —
333.

Dwyer, J. I). 1980. Flora of Panama Part IX. Family 179.

Rubiaceae. Ann. Missouri Bot. Card. 67: 1-522.

Lrnst, K. A. 1989. Insect herbivory on a tropical under-
storv tree: Effects of leaf age and habitat. Biotropica
21(3): 1 94— 1 99.

Freeman, C. E. & F. G. Stiles. 1990. [Abstract 171] Floral
nectar sugar compositions of Costa Rican hummingbird-
visited taxa. Amer. J. Bot. 77(6): 67.

Janzen, I). H. 1971. Euglossine bees as long distance pol¬
linators of tropical plants. Science 171: 203-205.
McDade, L. C. 1986. Protandry, synchronized flowering
and sequential phenotypic unisexuality iu Neotropical
Pentagonia macrophylla (Rubiaceae). Oecologia 68(2):
218-223.

Robbreeht, L. 1988. Iropical woody Rubiaceae. Opera
Bot. Belg. I: 1-271.

- . 1993. Supplement to the 1988 outline of the clas¬
sification of the Rubiaceae, Index to genera. Opera Bot.
Belg. 6: 1 73- 1 96.

Rova, J. H. E. & l„ Andersson. 1995. A reevaluation of
the tribes Hippotideae and Tammsieae (Rubiaceae).
Nordic J. Bot. 15: 269-284.

Standley, P. C. 1914a. Nothophlebia, a new genus of Ru¬
biaceae from Costa Rica. Contr. U.S. Natl. Herb. 17:
437-438.

- . 1914b. A revision of the genus Watsonamra.

Contr. U.S. Natl. Herb. 17: 439-444.

Steyermark, J. A. 1965. GOiero Hippotis. Acta Bot. Venez.
1: 85-102.

Stiles, F. G. & C. E. Freeman. 1993. Patterns in floral
nectar characteristics of some bird-visited plant species
from Costa Rica. Biotropica 25: 191—205.

I'aylor, C. M. 2001. Rubiaceae. In: W. I). Stevens et al.,
Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot.
Card. Vol. 85, Tomo 3: 2206—2284.

Rubiacearum Americanarum Magna Hama Pars X. New Species
and a New Subspecies of Faramea (Coussareae) from Central and

South America

Charlotte 17. Taylor

Missour i Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Charlotte,' 1 aylor@mobot.org

ABSTRACT. Descriptions and illustrations are pre¬
sented for the new species Faramea accumulans C.
M. Taylor and F. areolata C. M. Taylor of the San
Bias region of eastern Panama, F guaramacalensis
C. M. Taylor from cloud forests in northwestern Ve¬
nezuela. and F. longistipula C. M. Taylor from mon¬
tane forests in northwestern Colombia. The new
subspecies F tamberlikiana subsp. sessifolia (P. H.
Allen) C. M. Taylor from southern Costa Rica rep¬
resents the northernmost range extension of this
species.

Key words: Coussareae, Faramea, neotropics,
Rubiaceae.

During study of recent collections of Faramea
Aublet from Central and South America for the Flo¬
ra Mesoamericana and other projects, the following
undescribed taxa were discovered. A summary of
the morphology of Faramea has recently been pre¬
sented (Taylor, 1999). This Neotropical genus of
about 130 species is characterized by indehiscent.
often leathery, blue to black fruits with a single
rather thin-walled seed; the ovaries 1 -locular or in¬
completely 2-locular with one or two basal ovules;
and internally glabrous corollas with four valvate
lobes. Faramea is similar to Coussarea Aublet;
Coussarea is distinguished by its ovules inserted at
the same point (vs. separately) and its vertical (vs.
horizontal) seeds, and additionally usually by its
inflorescences white (vs. green to blue) and its
fruits white and spongy. Faramea has not been
studied as a whole, but several floristic treatments
have been presented, notably by Mueller (1881),
Steyermark (1967, 1974), Burger and Taylor
(1993), and Taylor (1999). Mueller recognized four
sections within the genus, but subsequent authors
have not used or expanded the infrageneric clas¬
sification of Faramea.

Faramea is particularly well represented in wet
lowland forests and montane forests, but it is found
throughout a wide range of Neotropical habitats in¬
cluding pyrrhophvtic vegetation (e.g., F. crassifolia
Bentham), gallery forests in savanna regions (e.g..

F. sessilifolia (Kunth) DC.), seasonal Antillean for¬
ests (e.g., F. occidental^ (L.) A. Richard), and At¬
lantic forests from Brazil (e.g., F percyanea Zahl-
bruckner) through Paraguay (e.g., F cyanea
Mueller Argoviensis). The reproductive biology of
Faramea has not been studied in much detail. I he
fruits of Faramea species are fleshy and presum¬
ably dispersed by animals. The flowers of several
species of Faramea are visited by butterflies (Bawa
& Beach, 1983), and are fragrant and white or blue
w ith the color usually consistent in a particular spe¬
cies. As in Coussarea, the corollas of some species
apparenth elongate markedly and rapidly just be¬
fore anthesis, so measurements of flower buds are
often not informative about size of the flowers at
anthesis. The white flowers of several species (e.g..
F. occidentals, pers. obs., Puerto Rico) have an
odor similar to that of gardenia, w hile the blue flow¬
ers (e.g.. F. multiflora A. Richard ex DC., pers.
obs., Costa Rica) have a sweeter and lighter fra¬
grance. Blue flowers are uncommon in the Rubi¬
aceae and not known in Coussarea. They have been
considered derived in the family (Robbrecht. 1988)
and thus are likely derived within Faramea. The
flowers of many species of Faramea are distylous,
but sometimes do not show the reciprocal place¬
ment of anthers and stigmas found in most Rubi¬
aceae (e.g., Faivre & McDade, 2001): in several
species (e.g., F. guaramacalensis, described below),
the anthers occupy nearly the same position in the
corolla tube in all flowers seen while the styles are
markedly dimorphic, so that the stigmas of the
short-styled flowers are positioned below1 the an¬
thers and included (e.g.. Fig. 2C), while those of
the long-styled (lowers are positioned at or just
above the corolla throat.

Some morphological features of Faramea are un¬
usual in the Rubiaceae and have been used by pre¬
vious authors to characterize the genus, but are ac¬
tually not found in all species and thus are
distinctive but not diagnostic. These features in¬
clude blue flowers (discussed above), stipule form,

Novon 12: 563-570. 2002.

564

Novon

leal arrangement, leal venation pattern, fruit shape,
and calyx limb length.

I Ik1 stipules are olten described for Faramea
(e.g., Steyermark, 1 9 < I) as interpetiolar, triangular,
and aristate. I he majority of species ol Faramea
do have this feature, and it allows recognition of
this genus because the aristas ol the stipules on the
terminal bud typically cross each other at their ba¬
ses and diverge at their apices in a manner found
only in Faramea. However, some species of Fara¬
mea lack aristas on their triangular stipules (e.g.,
F flavicans (Roemer & Schultes) Standley), while
other species have the stipules completely fused
into a ealyptrate structure that may have two short
projections at the apex (e.g., F. longistipula, de¬
scribed below), may terminate in a single short aris¬
ta (e.g., h. calyptrata C. M. Taylor), or may lack any
terminal projection (e.g., F. parvibractea Steyer¬
mark).

Another distinctive vegetative feature of Fara¬
mea is a distichous leal arrangement; in the major¬
ity of Rubiaeeae, including Coussarea, the leaves
are decussate. However, the entire genus has not
been surveyed in the field, and this character is not
always evident on herbarium specimens. The leaves
of many Faramea species have a distinctive vena¬
tion pattern with one or more well-developed sub¬
marginal veins (e.g., F. suerrensis Donnell Smith, F
exemplaris Standley); however, the submarginal
veins are only weakly developed in other species
(e.g.. F occidental is).

Similarly distinctive but not diagnostic are the
strongly oblate (i.e., depressed-globose), laterally
flattened fruits found in many Faramea species
(e.g., F suerrensis, F. multiflora), though the fruits
ol several Faramea species are ellipsoid to globose
(e.g., F occidental is). Additionally, the calyx limb
is usually rather short, truncate, and denticulate,
but also may be rather well developed and even
spathaeeous to irregularly lobed (e.g., F. cyathea
Standley, F. macrocalyx Mueller Argoviensis).

Other features that are particularly variable in
Faramea are inflorescence position and arrange¬
ment. Inflorescence position varies from consis¬
tently terminal (e.g.. F. suerrensis, F multiflora), to
variably axillary and/or terminal (e.g.. F. occiden¬
tals, F. torquata Mueller Argoviensis), to consis¬
tently axillary (e.g., F. egregia Sandwith, F. ma-
guirei Steyermark. F. guaramacalensis). The
inflorescences of some species are even supraaxil-
lary, inserted I to several mm above the junction
ol the leaf and the stem (e.g., F. spathacea Mueller
Argoviensis ex Standley, F guaramacalensis); the
developmental origin of this is not clear. As to in¬
florescence arrangement, Faramea shows nearly

the lull range found in the Rubiaeeae: from flowers
solitary (e.g., F capulifolia Dwyer), to few (e.g.. F.
quinquejlora Poeppig & Endlieher), to numerous
(e.g., F. multiflora); from flowers sessile or subses-
sile (e.g., F maguirei). to borne on well-developed
peduncles and/or pedicels (e.g., F. egregia, F. guar¬
amacalensis), to presenting all these conditions in
a single species (e.g.. F. Occident a l is); from inflo¬
rescence bracts reduced (e.g., F. guaramacalensis),
to w'oll developed (e.g., F. guianensis Aublet) and
sometimes even petaloid (e.g.. F. anisocalyx Poep¬
pig & Endlieher; Classen-Bockhoff, 1996); and
Irom thyrsiform with several orders ol branching
(e.g., F. multiflora, F. suerrensis) to densely capitate
(e.g.. F papirifolia (Steyermark) C. M. Taylor). Sim¬
ilarly extensive variation in inflorescence arrange¬
ment is found in several other genera of Rubiaeeae,
including Coussarea.

Faramea accumulans C. M. Taylor, sp. nov.
TYPE: Panama. San Bias: vecindad de Bio
Diablo, frente a la Isla Nargana. a unos 15 km
de la costa, 9°22'N, 78°35'W, 100-180 m. 2
July 1992, II. Herrera, II Obladia, R. (Madia
& N. Blanco I I9B (holotype, PM A; isotype,
MO-520402 1 ). Eigure II).

Haec species a congeneris foliis elliptico-oblongis 28-
14 cm longis basi cordulatis amplexicaulibus. inflorescen-
lia cymosa ax i I lari, pedicellis 5—12 mm longis, limbo ca-
lycino truncato ea. 3 mm longo atque fructu laevi oblato
ea. 10 X 14 mm distinguitur.

Small trees to 5 m tall, glabrous. Leaves elliptic-
oblong, 28-44 X 13—24 cm, at apex acute to short¬
ly acuminate, at base shortly cordate and amplex-
icaul, drying chartaceous, adaxially shiny, abaxially
matte; secondary veins I I to 16 pairs, looping to
interconnect forming an undulating submarginal
vein, adaxially costa prominent and remaining ve¬
nation plane, abaxially costa prominent and re¬
maining venation plane to prominulous; petioles 0-
3 mm long; stipules interpetiolar, persistent, main
body deltoid. 9—12 mm long, abaxially smooth, at
apex aristate with arista 5—6 mm long. Inflorescenc¬
es axillary, cymose, 3—3.5 cm long, glabrous, ebrac-
teale; peduncles I or 2 per axil; pedicels 5-12 nun
long; flowers with hypanthium cylindrical to turbi¬
nate, ca. 1 mm long; calyx limb ca. 3 mm long,
truncate, corolla in young bud glabrous externally,
not seen at maturity; anthers and stigmas not seen.
Infructescences similar to inflorescences; fruits ob¬
late, ca. 10 X 14 mm, smooth, purple.

Distribution, habitat, and phenology. Wet for¬
ests at 50-180 m in northeastern Panama; collected

Volume 12, Number 4
2002

Taylor

Faramea

565

566

Novon

with flowers in hud in July, with fruits in July and
August.

This species apparently accumulates detritus in
the clasping leaf bases that surround die stems, and
the specific epithet refers to this habit. Faramea
accumulans is similar to F. correae C. M. Taylor of
central Panama; F correae differs in its smaller
leaves, 8—18 X 2.5—7 cm, and its fewer, sessile,
terminal flowers. Faramea accumulans is also sim¬
ilar to F permagnifolia Dwyer ex C. M. Taylor of
southeastern Costa Rica; F. permagnifolia differs in
its oblanceolate to obovate leaves that are rounded
to truncate at the base and its fasciculate terminal
flowers. Faramea accumulans is also similar to an
unnamed species from northern Costa Rica and ad¬
jacent Nicaragua, which differs in its carinate ca¬
lyces and fruits; and F. accumulans is similar to F.
cyathocalyx Standley of northwestern Colombia,
which differs in its fasciculate (lowers and petiolate
leaves that are acute to cuneatc at the base.

Paratypes. PANAMA. Sail Bias: Cordillera frenle a
la Isla Nargand, Ribera de Rfo Diablo, C. Galdames el at.
1480 (MO. I’M A); Rfo Diablo y veeindad de Duque Sui,
a linos 10 km de la costa frente a la Isla de Nargana, ruta
haeia Cerro Ibedon. //. Herrera el al. 1178 (MO, I’M A);
veeindad del Rio Diablo, 8—9 km de la costa, II. Herrera
el al. 1728 (MO. PMA).

Faramea areoluta C. !M. Taylor, sp. nov. TYPE:
Panama. San Bias: Nusagandi and road to Car-
tf, 9°18'N, 78°58'W, 400 m, 18 July 1980, ./.
F. McDonagh, H. A. Lewis, A'. J. Gumpel & .4.
./. Plumptre 172 (holotype, I’M A; isotypes,
BM-614523, MO-3622277). Figure I A-C.

Haec species Farameae eurycarpae similis. sed ab ea
foliis rigidioribus in sicco plumbeo-viridibus, infloreseen-
tiae ax i bus secundariis verticillatis atque limbo calycino
0.5— 0.8 mm fongo distinguitur.

Shrubs or trees to 5 m tall, glabrous. Leaves el¬
liptic-oblong to lanceolate, 8-19 X 2-8 cm. at apex
acuminate with tips often well developed, at base
cuneate to usually rounded, truncate, or occasion¬
ally shortly cordate, drying chartaceous, adaxially
matte, usually shiny abaxially; secondary veins 8
to 13 pairs, looping to interconnect forming an un¬
dulate-angled, usually reticulating submarginal
vein, adaxially and abaxially costa prominent and
remaining venation reticulated and prominulous;
petioles 3—7 mm long; stipules shortly to completely
united around stem into a continuous sheath, ca¬
ducous, main body of interpetiolar portion 5-7 mm
long, abaxially smooth, at apex obtuse to broadly
so, aristate with arista 2-5 mm long, inserted at or
a little below apex. Inflorescences terminal, cymose,
blue to white, glabrous; peduncles 2.5-0 cm long;

branched portion corymbiform, 5-11 X 5—9 cm;
secondary axes 3 to 8, verticillate to subverticillate;
bracts triangular, to 1.5 mm long; pedicels 2-7 mm
long; flowers distylous; hypanthium ellipsoid to cy¬
lindrical, 1-1.5 mm long; calyx limb 0.5-0. 8 mm
long, denticulate; corolla salverform, blue, exter¬
nally glabrous, tube 7—8 mm long, lobes narrowly
triangular, 3—5 mm long, acute; anthers and stigmas
not seen in good condition. Infructescences similar
to inflorescences; fruits oblate, ca. 5X11 mm, lat¬
erally somewhat flattened, smooth, blue.

Distribution , habitat, and phenology. In wet
forests and cloud forests at 200—800 m in central
and eastern Panama; collected with flowers in Jan¬
uary, March through September, and December,
with fruits in January, April, June, July, and Octo¬
ber through December.

I his species has previously been confused with
Faramea eurycarpa Donnell Smith (e.g., Dwyer,
1980), but F. eurycarpa differs in its leaves that dry
papyraceous and usually yellow-green and its calyx
limbs 1-3 mm long. The leaves of F. areoluta in
particular are notable in being shiny abaxially and
having the submarginal veins rather angled and the
higher order venation of both surfaces raised and
arranged in distinctive angled areoles (Fig. 1C); the
specific epithet refers to this distinctive venation.
Faramea areoluta is similar to F. ampla Standley
of northwestern Colombia; F. ampla differs in its
leaves that dry papyraceous, are cuneate to obtuse
at the base, and have submarginal veins more
smoothly undulating and little or not at all reticu¬
lated.

Due to the poor preservation of the flowers on
the specimens seen, the exact position of the an¬
thers al anthesis is not clear, but it appears to be
generally the same in both short-styled and long-
styled flowers while the stigma position differs.

Paralypes. PANAMA. Colon: ridge top leading N
from Rio Escandaloso towards Cerro liurja [sic], Hummel
2t 10 (MO); Santa Rita Ridge Road, 26— 22 km from Tran-
sisthmica Highway, Sytsma 1325 (MO); ridge between Rfo
Piedras and Rfo Gatlin waterbeds, along trail from end of
Santa Rita Ridge Road, ca. 5— 8 km SW of Cerro Bruja,
Sytsma el al. 4292 (MO). Panama: Plano— Cartf road, I
mi. past sawmill on dirt road, T. Antonio 2533 (MO): along
newly cut road from El Plano to Cartf-Tupile, 12 mi.
above Pan-Ant Highway, Croat 22874 (MO), Liesner 1120
(MO); El Plano to Cartf Road, 14 km N of Panamerican
Highway, ./. Folsom £■ Kauke 1411 (MO), 14.9 km N of
Panamerican Highway, Folsom el al. 1449 (MO), 8.2 (on
new road, 8.6 on old road) mi. from Pan-American High¬
way on die PI Llano— Cartf Road, Knapp & llufl 4421
(MO); high point ol ridges S of I pet f , 5-6 hrs. walk from
Choc6 village, Serranfa de Maje. Knapp el al. 4528 (MO);
along new El Llano-Cartf road. 8-12 km N of El Plano,
Nee et al. 8769 (MO); El Plano-Cartf Road, 16 km N of

Volume 12, Number 4
2002

Taylor

Faramea

567

Pan Am. Highway at El Llano, Nee & Dressier 9356 (MO),
16-18.5 km by road. Nee & Tyson 10968 (MO); El Llano-
Cartf Road. Km 10-15, P. Maas et al. 2829 (MO); El
Llano-Cartf road, 10.6 mi. from the Pan American High¬
way, J. S. Miller et al. 875 (MO); El Llano-Cartf road, just

5 of San Bias border, J. S. Miller et al. 101 1 (MO); El
Llano-Cartf road, 10 km from Inter- American Highway,
Mori & Kallunki 2299 (MO), 2305 (MO), 9.6-1 1 km, Mori

6 Kallunki 3521 (MO), 21.1 km, Mori & Kallunki 5112
(MO), 11-12 km. Mori et al. 6885 (MO), 8-1 1 km. Mori
7707 (MO); El Llano-Cartf Road, ea. 18 km from Pan Am
Highway, Stein 1029 (MO); El Llano-Cartf Road, 15 km
above Pan-American Highway, Systma A Sytsma 3140
(MO). San Bias: El Llano-Cartf Road, vicinity Nusagan-
df, sendero Nusagandf down to the Atlantic coast, Croat
69256 (MO); Nusagandi, along the continental divide on
the El Llano-Cartf Road, de Nevers & Rerez 3581 (MO);
Nusagandi, summit of El Llano-Cartf Road, along the first
stretch of the trocha S of camp, de Nevers & Gonzalez 3641
(MO); El Llano-Cartf Road, Km 19.1. de Nevers et al.
5010 (MO). 5876 (MO), 7343 (MO), Km 27, de Nevers el
al. 5094 (MO), Km 16.7, de Nevers & Charnley 5903
(MO); 23-29 km from Pan-American Highway on El Lla¬
no— Cartf Road, Knapp 1856 (MO).

Faramea guaramacalensis C. M. Taylor, sp. nov.
TYPE: Venezuela. Lara: distrito Moran, car-
retera desde Humacaro Alto hacia Guaito,
2200 tn, 14 Nov. 1984, H. van der Werff & R.
Rivero 7890 (holotype, MO-5204020; isotypes,
PORT, US, VEN). Eigure 21), E.

Haec species a Faramea larensi inflorescentia axi I lari,
pedicellis longioribus atque limbo calycino breviore; a F
spathacea limbo calycino pedicellis et lobulis corollinis
brevioribus distinguitur.

Shrubs and small trees to 6 m tall; stems gla¬
brous. Leaves elliptic to obovate or elliptic-oblong,
4-14 X 1.5—6 cm, at apex acute with tips 5-20
mm long and usually quite slender, at base cuneate
to rounded, drying papyraceous to chartaceous, gla¬
brous on both surfaces; secondary veins 6 to 9
pairs, looping to interconnect forming a curving to
rather angled submarginal vein, without domatia,
adaxially venation plane or costa prominulous,
abaxially costa prominent, secondary veins promi¬
nulous, and remaining venation plane to usually
thickened; petioles 3-5 mm long; stipules glabrous,
persistent, shortly united around stem, main body
interpetiolar portion deltoid to broadly triangular.
3—5 mm long, abaxially smooth in basal half and
costate in upper part, al apex acute, aristate, arista
linear, 1-3 mm long, apparently glandular at its
apex. Inflorescences shortly supraaxillary, produced
at first subterminal node of stems, inserted 1-2 mm
above node, glabrous; peduncles 1 per axil, 16-28
nun long, terminating in a single 3-flowered cyme;
bracts 0.5-2 mm long, narrowly triangular to linear;
pedicels 3-8 mm long; flowers distylous, fragrant;
hypanthium narrowly turbinate to cylindrical, ca.

1.5 mm long, glabrous; calyx limb 0.8—1 mm long,
sinuate tc truncate, with 4 linear teeth ca. 0.3 mm
long; corollas salverform to narrowly funnelform,
white, glabrous externally, tubes 13—18 mm long,
lobes 4, narrowly triangular to lanceolate, 7-8 mm
long, acuie; anthers ca. 4.5 mm long, included, in
short-styled form situated just below corolla throat,
in long-styled form situated just above middle of
corolla tube; stigmas 2, linear, in short-styled form
ca. 3.5 mm long, included, situated near middle of
corolla tube, in long-styled form ca. 2.5 mm long,
partially exserted. Infructescences similar to inflo¬
rescences or sometimes displaced to second sub-
terminal node by subsequent stem grow th; fruits ob¬
late, ca. 6X8 mm long, glabrous, color not noted.

Habitat, distribution, and phenology. In cloud
forests at 1950-2200 m in Lara, Portuguesa, and
probably adjacent Trujillo states of northwestern
Venezuela; collected in flower in June and Novem¬
ber. in immature fruit in May, June, and November.

This new species is distinguished by the com¬
bination of its shortly petiolate leaves, persistent
aristate stipules, axillary, few-flowered inflorescenc¬
es with well-developed peduncles and pedicels, rel¬
atively short denticulate calyx limb, white corollas
with well-developed tubes, oblate (but not laterally
flattened) fruits, and cloud forest habitat. 'Ellis spe¬
cies was first noticed as undescribed by J. A. Stey-
ermark, who passed the type collection on to J. I).
Dwyer, who never advanced work on it and later
passed this same specimen to me (J. I). Dwyer,
pers. comm.). Rut this species has been best doc¬
umented by the work of Cuello (1996, 1999) in
Guaramacal National Park, and the specific epithet
refers to ts occurrence here. Faramea guarama-
calensis was treated as “ Faramea aff. larensis ” in
the recent flora of this park (Dorr et ah, 2000).
Cuello’s focused floristic sampling showed strong
altitudinal stratification in the vegetation of this
park and the distribution of many of its plant spe¬
cies, including F. guaramacalensis.

Faramea larensis Steyermark is also found in
cloud forest in northwestern Venezuela, but differs
from this new1 species in its terminal inflorescences,
stipule aristas ca. 4 mm long, pedicels 1.5— 3.5 mm
long, and 'ulyx limbs 1—1.5 mm long. Faramea lar¬
ensis is k lown only from a single collection w ith
immature flowers, so the measurements provided by
Steyermark represent only a minimum flower size.

In the treatment of the Rubiaeeae of Venezuela
(Steyermark, 1974), Faramea guaramacalensis will
key to two different species, F. spathacea and F.
egregia. Faramea spathacea differs from I'. guara¬
macalensis in its peduncles 0—4 mm long, pedicels

568

Novon

5 cm

Kigurt> 2. A-C. Faramea longistipula C. M. Taylor. — A. Flowering branch. — B. Flower bud. — C. Flower shortly
alter corolla has fallen off. I). E. Faramea guaramaealensis C. M. Taylor. — I). Flowering branch. — E. Flower, partially
dissected. A, I) to 5-cm scale. A. based on Cogollo el al. 7839 ; B. C, based on Pipoly el at. 17263: D, E, based on
van der Werff & Rivero 7890.

Volume 12, Number 4
2002

Taylor

Faramea

569

10—30 mm long, calyx limbs 6—8 mm long, corolla
lobes 13-20 mm long, and distribution in Venezue¬
la at 700-1400 m; F. egregia differs from F guar-
amacalensis in its solitary flowers, calyx limbs ca.
6 mm long, corolla lobes 16-30 mm long, and dis¬
tribution in Guyana and adjacent southeastern Ve¬
nezuela at 500-1000 m.

Paratypes. VENEZUELA. Porluguesa: mpio. Sucre,
Parque Nacional Guaramacal, sector El Alto, 1 unite con
el Estado Trujillo, Camino Real Paramito— Batatal, 9° 19—
20'N, 70°05'W, A. Cuello et al. 1623 (PORT), 1639
(PORT), 1699 (PORT).

Faramea lougisti|mlu C. M. Taylor, sp. nov. TYPE:
Colombia. Antioquia: mpio. Urrao, vereda Calles,
Rio Calles, Parque Nacional Natural “l,as Or-
quideas,” 4°36'N, 77°05'W, 1400-1500 m, 3
May 1995, R. Fonnegra & Grupo de Palinologia
Semestre 1-95 5496 (holotype, HUA; isotype,
MO-5204022). Figure 2A-C.

Haec species a congeneris stipulis spathaceis breviter
biaristatis 3.6— 6.5 cm longis distinguitur.

Shrubs and small trees to 6 m tall; stems gla¬
brous. Leaves elliptic-oblong to narrowly elliptic, 8—
19 X 2.5—6 cm, at apex acuminate with tips 12—
25 mm long and usually slender, at base cuneate
to obtuse, drying papyraceous to chartaceous and
often discolorous, glabrous on both surfaces; sec¬
ondary veins 10 to 22 pairs, looping to interconnect
forming an angled to undulate submarginal vein,
without domatia, adaxially costa prominulous and
remaining venation plane, abaxially costa promi¬
nent and secondary veins and reticulated higher
order venation thickened to prominulous; petioles
5—12 mm long; stipules glabrous, united into a lat¬
erally flattened sheath, main body of sheath 3.5—
6.5 cm long, at apex acute to obtuse, with aristas
2, closely set, 3.5—5 mm long, as new leaves ex¬
pand sheath splitting from apex for up to V3 forming
a spathaceous structure, eventually deciduous by
fragmentation. Inflorescences terminal, corymbi-
form-rounded, 3.5—5 X 4.5—6 cm, branched to sev¬
eral orders, white, glabrous, subsessile and tripar¬
tite with subtending leaves often deciduous,
principal axes terminating in dichotomous cymules
of 3 to 5 flowers; bracts none or deltoid, to 0.5 mm
long; pedicels 3—12 mm long; flowers with hypan-
thium ellipsoid to subglobose, 1 — 1.2 mm long, gla¬
brous; calyx limb 0.3— 0.5 mm long, truncate, entire
to 4-dentieulate, glabrous; corolla salverform, white
to blue, externally glabrous, tube 7-9 mm long,
lobes 4, narrowly triangular to narrowly lanceolate,
3-4 mm long, acute; anthers ca. 5 mm long, in¬
cluded, situated just above middle of corolla tube;
stigmas 2, linear, ca. 1 mm long, included, situated

near base of corolla tube on style ca. 1 mm long.
Infructescences to 5 X 8 cm, their axes appearing
jointed due to internodes often becoming thickened
and nodes contracted; fruits oblate, ca. 8X15 mm,
laterally rather flattened, glabrous, smooth, leath¬
ery, purple.

Habitat, distribution, and phenology. In wet
montane forest at 1390—1750 m in the northwestern
Cordillera Occidental of the Andes; collected in
flower in May, November, and December, in im¬
mature fruit in March, May, and December.

This new species is distinguished by its unusu¬
ally well developed, calyptrate to spathaceous stip¬
ules; it is also distinctive in its combination of cor-
ymbiform, well-branched inflorescences, slender
white to blue corollas, oblate laterally flattened
fruits, and montane forest habitat. The relatively
large stipules separate this species from the other
Andean species of Faramea except F calyptrata C.
M. Taylor of montane southwestern Colombia and
northern Ecuador; the specific epithet refers to
these stipules. Faramea calyptrata differs from F.
longistipula in its stipules that are apieally entire
or bear only 1 arista, its 5 to 10 fasciculate pedun¬
cles each producing a single flower, and its corollas
with tubes 16—25 mm long. Faramea longistipula
is one of several species of Faramea from western
Colombia that Standley annotated with the unpub¬
lished epithet “cclata."

Paratypes. COLOMBIA. Antioquia: mpio. Urrao,
vereda Calles, Parque Nacional Natural “Las Orqufdeas,”
margen derecha del Rio Calles, Cogollo et al. 2564
(JAUM, MO), 3915 (JAUM, MO), 7435 (JAUM, MO),
7839 (JAUM. MO), Pipoly et al. 17263 (JAUM. MO); Alto
de Palmitas, Pipoly et al. 17637 (JAUM, MO); Quebrada
La Agudelo, D. Cardenas et al. 3180 (JAUM. MO).

Faramea tamberlikiana Mueller Argoviensis subsp.
sessifolia (P. H. Allen) C. M. Taylor, comb, et
stat. nov. Basionym: Faramea sessifolia P. H. Al¬
len, Rain Forests of Golfo Dulce 409. 1956.
TYPE: Costa Rica. Puntarenas: Esquinas Forest,
area between the Rio Esquinas and Palamar, 2(H)
ft., 22 May 1950, P. H. Allen 5539 (holotype,
US; isotypes, F- 1716255, F- 1 509883).

As circumscribed here, Faramea tamberlikiana
Mueller Argoviensis is found in the western Ama¬
zon Basin of South America, throughout wet low¬
land Colombia including along its Pacific coast,
and north into Panama and southern Costa Rica.
Reproductive features are similar in plants from
throughout this region, but there is notable varia¬
tion in vegetative features. In particular F. tamber-

570

No von

likiana subsp. tamberlikiana, found in the Amazon
basin and central and eastern Colombia, has peti¬
oles 3-13 mm long, leaves with the bases acute to
rounded or truncate, and calyx limbs ca. 0.2 mm
long, while the allopatric subspecies sessifolia,
found from southern Costa Rica to western Colom¬
bia, has petioles 1-5 mm long, leaves that are
rounded to truncate or cordate at the base, and ca¬
lyx limbs 0.5— 1.5 mm long. The leaves of vegetative
stems of subspecies sessifolia are subsessile, rela¬
tively larger, cordate at the base, and markedly am-
plexicaul, while those of reproductive stems are
usually shortly petiolate, relatively smaller, and
truncate to rounded at the base, though occasional
leaves ol reproductive stems are similar to those of
vegetative stems. In contrast, subspecies tamberli¬
kiana has similar leaves on both vegetative and re¬
productive stems. As a species F. sessifolia was
considered restricted to southern Costa Rica (e.g..
Burger & Taylor, 1993), but as circumscribed here
its range is wider.

Specimens examined. COLOMBIA. Antioquia: mpio.
Cdceres, Kms 10—15 SK de Cdceras, sobre La Truncal de
La Paz, Callejas et al. 5354 (HUA, MO); mpio. Tarazd,
eorreg. Kl 12. vereda Banablanco, 210 km NE <le Medel-
lfn, Troncal del Caribe, en la vfa El 12-Barroblanco, cuen-
ca del rfo Pubi, Callejas et al. 10376 (HUA. MO); mpio.
Puerto Mare, Ifmites con la vereda El Prodigio (mpio. Sail
Luis), vereda Serranfas. I). Cardenas et al. 3017 (JAUM,
MO). Chord: mpio. Nuquf, eorreg. Termales, N of Que-
brada Piedra Piedra, Acevedo- Rodriguez et al. 6365 (MO,
US); Serranfa de Baudo, along road between Las Animas
and Palo on Rfo Palo, ca. 5 km SW of Pato, Croat 56036
(MO); carretera Panamericana (en construceidn), adelante
del Rfo Paid, K. Forero et <d. 5662 (COL, MO); Guayabal
(just N of Quibdd), Juncosa et id. 700 (NY). COSTA RICA.
Puntarenas: Reserva Eorestal Golfo Duke, Los Mogos,
Golfito, R. Aguilar 546 (CR, MO); along road from Pana-
merican Highway al Piedras Blancas to Rincdn (on Osa
Peninsula), 3.7 mi. Vi of Panamerican Highway, Croat
67636 (MO); Parque Nacional Corcovado, Sirena, on trail
toward Laguna Corcovado (“The Swamp”), Delprete 5202
(CR, MO); fila before Rancho Ouemado. near Rincdn, Osa
Peninsula. Gentry et id. 73652 (MO). 73634 (MO sterile);
ridge between Rfo Riyito (valley of Laguna Chocuaco) and
Ouebrada Banegas, S of Cerro Rancho Ouemado (ca. 7
km W of Rincdn de Osa), Grayum et al. 7566 (MO); Re¬
serva Eorestal Golfo Dulce, Osa Peninsula, Trocha de La
Tarde rd. 10 km SVt of La Plana, S of Rincon de Osa, E
of the Rfo Rincon valley, Hummel cV Robleto 16730 (CR.
MO); Osa Peninsula, trail from Rincdn de Osa to Rancho
Ouemado. //. Kennedy 1940 (MO); Corcovado National
Park, trail from base of hills to Los Chiles, Liesner 3075
(MO); Corcovado National Park, Llorona Plateau, along
trail above Llorona station, Neill 5045 (MO); along the
Cam i no al Paeffico, W of Rincdn de Osa, Osa Peninsula,
near Mile 12, Raven 21502 (E, MO); R.N.V.S. Golfito, Gol¬
fito, Valle de Coto Colorado, f ila La Gamba, Al. Segura
31 (INB, MO); on road to radio and telecommunications
tower, 6 km N of Golfito, Utley & Utley 4337 (DUKE,

MO); Estacidn Biologica Marenco, parte NO de la Esta-
cidn, sendero camino publico, Peninsula Osa, Zamora et
al. 1200 (CR, MO). PANAMA. Code: on the Atlantic
side ca. 5 hr. walk from sawmill at El Cope, along slopes
above Norte Rfo Blanco near small village of Cano Susio,
T. Antonio 3623 (MO); Penonom6— Coclecito, 5.6 mi. N of
Llano Grande, along Rfo Cascajal, 1.4 mi. N of continental
divide, Croat 67473 (MO); area between Cano Blanco del
Norte, Cano Sucio and Chorro del Rfo Tife, Davidse &
Hamilton 23633 (MO); 9 km N off Llano Grande on road
to Coclesito, Hummel 1307 (MO). Colon: along Portobe-
lo— Nombre de Dios road, 10 km W of Nombre de Dios,
Knapp & Mallet 5707 (MO).

Acknowledgments. I thank R. E. Gereau for prep¬
aration of the Latin diagnoses; the curators of COL,
F, HUA, JAUM, PM A, PORT, and US for access to
or information about specimens, and in particular M.
Correa; R. Foster, T. Wachter, and the Andrew Mellon
Foundation through a grant to the Field Museum of
Natural History for support for travel to that institu¬
tion; O. M. Montiel and R. Magill for significant en¬
couragement of this work; and A. Cogollo, Q. Jimenez,
J. Sanchez, L. Dorr, N. Cuello, G. Aymard, and ,1.
Dwyer for providing information.

Literature (filed

Bawa, K. S. & J. H. Beach. 1983. Self-incompatibility
systems in the Rubiaceae of a tropical lowland wet for¬
est. Amer. J. Rot. 70: 1281-1288.

Burger, W. C. & C. M. Taylor. 1993. Flora Coslarieensis:
Family #202. Rubiaceae. Eieldiana, Bot. n.s. 33: 1-
333.

Classen-Boekhoff, R. 1996. A survey of flower-like inflo¬
rescences in the Rubiaceae. Opera Bot. Belg. 7: 329—
367.

Cuello A., N. L. 1996. Floristic Diversity and Structure of
the Montane Forests of Cruz Carrillo National Park in
the Venezuelan Andes. M.S. Thesis, University of Mis-
souri-St. Louis.

- (editor). 1999. Parque Nacional Guaramacal.

UNELLEZ-Fundacion Polar, Caracas.

Dorr, L. J., B. Stergios, A. R. Smith & N. Cuello A. 2000.
Catalogue of the vascular plants of Guaramacal National
Park. Portuguesa and Trujillo States, Venezuela. Contr.
U.S. Natl. Herb. 40: 1-155.

Dwyer, J. I). 1980. Flora of Panama Part IX. Family 179.

Rubiaceae. Ann. Missouri Bot. Card. 67: 1-522.

Faivre, A. E. & L. A. McDade. 2001. Population-level
variation in the expression of heterostyly in three spe¬
cies of Rubiaceae: Does reciprocal placement of anthers
and stigmas characterize heterostyly? Amer. J. Bot. 88:
841-853.

Mueller, J. 1881. Faramea. El. bras. 6(5): 105—162.
Robbrecht, E. 1 988. Tropical Woody Rubiaceae. Opera
Bot. Belg. 1: 1-271.

Steyermark, J. A. 1967. Faramea. In: B. Maguire & Col¬
laborators. The Botany of the Guyana Highland — Part
VII. Mem. New \ork Bot. Gard. 17: 371—396.

- . 1974. Faramea. In: T. Lasser (editor). Flora de

Venezuela 9(1): 935—967.

Taylor, C. M. 1999. Rubiaceae— Coussareae. In: G. Hailing
& L. Andersson (editors), FI. Ecuador 62: 238—307.

Rul )iacearum Americanarum Magna Hama Pars XI. A New
Species of Alseis (Calycophylleae) from Central America and Notes
on the Morphology of this Neotropical Genus

Charlotte M. Taylor

Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Charlotte.Taylor@mobot.org

Abstract. The Neotropical tree genus Alseis is
unusual in t tie Rubiaceae in having a typically de¬
ciduous habit and protogynous flowers that are pro¬
duced just before or with the new leaves. Alseis
costaricensis C. M. Taylor shares characteristics
w ith both A. hondurensis Standley and A. blachiana
llemsley of Panama and adjacent Colombia, but is
allopatrie to and differs morphologically from both
of these species.

Key words: Alseis, Calycophylleae, neotropics,
Rubiaceae.

The genus Alseis Schott includes about 15 spe¬
cies of Neotropical trees found in lowland wet. sea¬
sonal, and dry forests and also in savanna habitats
from southern Mexico to southern Brazil. Alseis is
distinguished by its spiciform inflorescences; its
relatively short corollas with the reduced lobes nar¬
rowly imbricate or sometimes apparently open in
bud; its densely pubescent stamen filaments; its nu¬
merous ovules on pendulous placentas that are at¬
tached to the apical part of the septum; its narrow
capsular fruits; and its flattened w inged seeds (An-
dersson, 1994). Alseis was long classified in the
tribe Cinchoneae (e.g., Steyermark, 1974; Rob-
brecht, 1988), but careful study by Andersson and
Persson (1991) showed that this tribe was rather
heterogeneous. Consequently they as well as later
authors (e.g., Robbrecht, 1993) classified Alseis in
the tribe Calycophylleae L. Andersson & Persson,
which comprises several genera of Neotropical
trees.

Alseis is well circumscribed as a genus but not
well documented by most authors, and is poorly
understood at the species level. Several features of
Alseis that are unusual in the Rubiaceae have been
documented repeatedly and independently by re¬
cent collectors, notably its usually deciduous habit
and its protogynous flowers. Below 1 discuss these
unusual features and describe a new species that
was discovered during preparation of the Rubi¬
aceae treatment for tbe Flora Mesoamericana.

Alseis trees are typically relatively large in both

height and diameter, and often reach the subcanopy
or canopy of the forest. Some specimen labels re¬
port heights up to 35 m, although most describe the
trees as 15-20 m tall. Collection of fertile speci¬
mens from trees this size is often difficult, and Al¬
seis is unsurprisingly represented by relatively few
specimens from most regions. Additionally, many
species of Alseis are reported by specimen labels
and field observations to be at least facultatively
deciduous and to produce their flowers usually just
before or with the flushing of the new leaves (A.
Jardim, pers. comm.; Andersson, 1994), an uncom¬
mon habit among Neotropical Rubiaceae. Thus,
flowers of Alseis are nearly always collected sepa¬
rately from the leaves and fruits, so correlating
these structures is difficult. And, the stipules of
Alseis are usually quickly caducous as the new
leaves flush, so they cannot be used to help cor¬
relate flowering and fruiting collections as can be
done with many Rubiaceae.

The floral biology of Alseis is also unusual in
being protogynous (Andersson, 1994; Burger &
Taylor, 1993). This condition is found in several
other Neotropical genera of Rubiaceae (e.g., War-
szewiczia Klotzsch, Simira Aublet), but the flowers
of Alseis show more marked developmental differ¬
ences than in other genera, in particular in corolla
size and form. Typically the corollas of Alseis are
tubular and apparently pale green in the first, pis¬
tillate stage (Fig. 1 B), but then they elongate, wid¬
en, and become white and campanulate in the sec¬
ond, staminate phase (Fig. ID). The corolla in the
staminate phase may be two to three times as large
as it was in the pistillate phase, and the pubescent
filaments that enlarge in the staminate phase pro¬
duce the appearance of a densely barbate corolla
throat in this phase. This complicated floral biology
has also contributed to the poor taxonomic knowl¬
edge of the genus, because species descriptions fre¬
quently do not specify which stage of the corolla is
being described. Clearly comparisons of corollas in
different stages are not useful for species distinc¬
tions.

Novon 12: 571-574. 2002.

572

Novon

figure 1. Alseis costaricensis (.. M. laylor. — A. Stem with old inflorescences (corollas fallen). — B. Detail of inflo¬
rescence showing two flowers in pistillate stage. — C. One flower between tin1 pistillate stage and anthesis. — I). Detail
of inflorescence showing one flower at anthesis. — E. Detail of inflorescence showing one old flower, after the corolla
has fallen. A, based on Jimenez et al. 670, to 5-cm scale; B, C, D, E, all based on Aguilar 927, to 5-mm scale.

The capsules of Alseis species are all quite sim¬
ilar. about 1—2 cm long, relatively narrow (1—5 mm
wide), widest at the top (obeonic, obovoid, turbi¬
nate, clavate), chartaceous to woody, septicidal from
the apex, and two-valved (Steyermark, 1074: lf>2,
fig. 20). After dehiscence the capsule valves typi¬
cally remain attached to the old infructeseence axes
for some time. The orientation of the capsules and
the old valves, ascending versus reflexed, has been
considered useful to distinguish among species
(e.g., Standley & Williams, 1075) but actually
seems to be quite variable. For example, all ori¬
entations from strongly ascending to completely re¬
flexed are found among the specimens of A. black-
iana Hemsley from Barro Colorado Island, a
relatively small site. The individual capsule valves
sometimes show a partial secondary splitting from

the top, but this feature is variable and apparently
not taxonomically significant.

Alseis costaricensis C. M. laylor, sp. nov. TYPE:
Costa Rica. Puntarenas: Reserva Forestal Gol-
fo Dulee, Aguabuena, sector cuenca Norte,
8°42'N, 83°32'W, 50-150 m, 13 Feb. 1992,
R. Aguilar 927 (holotype, CR; isotypes, F-
2142377, MO-4362115). Figure 1.

Haec species Alseidi blackianae Hemsley et A. hondu-
rensi Standley similis, sed a hac lobulis calycinis brevior-
ibtis (0.5—1 mm longis) atque floribus sessilibus, ab ilia
inflorescentiis terminalibus axillaribusque plerumque ra-
mosis distinguitur.

Trees to 30 m tall, deciduous; young twigs stri-
gillose to glabrescent. Leaves oblanceolate to ob-
ovate, 7—22 X 3.5— 9.5 cm, at apex acute to some-

Volume 12, Number 4
2002

Taylor

Alseis

573

what acuminate, at base acute, papyraceous, above
and below sparsely strigillose rapidly becoming gla-
brescent except persistently puberulous to strigil¬
lose on costa and secondary veins; secondary veins
10 to 19 pairs, weakly looping to interconnect near
margins, abaxially with hirtellous to pilosulous
domatia in axils, adaxially costa, secondary veins,
and higher-order venation prominulous, abaxially
costa prominulous to prominent and secondary
veins and higher order venation prominulous; pet¬
ioles 10-25 mm long, puberulous to glabrescent;
stipules 5—6 mm long, narrowly triangular, acute to
acuminate. Inflorescences terminal and also in axils
of uppermost leaves, produced immediately before
new leaves, 15—25 X 10—20 cm, branched; sec¬
ondary axes 1 to 2 pairs, densely strigillose to to-
mentellous, with trichomes generally appressed;
peduncles 1-2 cm long; bracts narrowly triangular
to linear, 0.8— 1.5 mm long, acute. Flowers sessile,
protogynous; hypanthium 1—1.5 mm long, densely
tomentellous to strigillose; calyx limb divided near¬
ly to base, lobes 5, 0.5—1 mm long, lanceolate to
ovate; corolla in staminate phase campanulate,
white to cream, 2-2.5 mm long, externally glabrous
except often shortly puberulous near base, inter¬
nally glabrous, lobes 5, reduced (i.e.. 0.2— 0.3 mm
long), rounded, in bud narrowly imbricate to ap¬
parently open; anthers ca. I mm long, in staminate
phase exserted. filaments 5—7 mm long, glabrescent
in upper third and densely villous below; mature
stigmas 2—2.5 mm long in pistillate phase, in sta¬
minate phase becoming twisted. Capsules narrowly
turbinate, 9-12 X 1-1.5 mm. chartaceous to some¬
what woody, brown, glabrescent; seeds linear-fusi¬
form, 5—8 X 0.5— 0.8 mm, papery.

Habitat, distribution, and phenology . Seasonal
forests at 50—700 m in south-central Costa Rica;
collected in flower in February, March, and June,
in fruit in May. July, and September.

This species was treated hy Burger and Taylor
(1993) as “ Alseis sp. aff. A. hondurensis Standi.,”
but with further study of this genus in Central
America, its status is now clearer. Alseis costaricen-
sis has been previously confused with both A. hon¬
durensis Standley and A. blacfciana. Alseis blacki-
ana is found in central Panama and adjacent
northwestern Colombia, and is quite similar to this
new species in its floral morphology and also ap¬
parently in producing its flowers before its new
leaves. However, A. blackiana differs from A. cos-
taricensis in having unbranched, usually axillary in¬
florescences. Alseis hondurensis is found from
southern Mexico through Honduras and is similar

to A. costaricensis in having branched terminal in¬
florescences. However, A. hondurensis differs from
A. costaricensis in its longer calyx lobes 1-2.5 mm
long, and apparently in producing its flowers after
the new leaves have flushed. These three species
all appear to be allopatric, with A. costaricensis oc¬
cupying a range between but isolated from the other
tw o. The intermediate morphology of A. costaricen¬
sis. which shares some features with each of these
other species, together with its intermediate range
suggest the possibilities that it may be of hybrid
origin, or that these three species are not complete¬
ly isolated, or that these three species may have
arisen from a formerly continuous population, per¬
haps with some clinal variation. At present, none
of these possibilities can be excluded.

Paratypes. COSTA RICA. Alajuela: San Carlos, Ha¬
ber et al. 1804 (MO), Haber et al. 1825 (F, MO). Guan-
acaste: Parque National Guanacaste, Estaci6n Pitilla,
Sendero El Mismo. Einca La Pasmompa, P. Rios 229 (CR,
MO sterile). Puntarenas: Drake-Osa, 3 km al E de Bahia
de Drake, camino a Agujitas, Q. Jimenez et al. 670 (CR,
F, MO); canton de Garabito, Reserva Bioldgica Carara.
cuenca del Rio Grande tie Tarcoles, camino a Coopeear-
ara, Q. Jimenez et al. 1366 (CR, F, MO); canton de Osa,
Rancho Quemado, sector norte, camino a Estero Guerra,
./. Marin 412 (CR. F, MO); cantdn de Garabito, Reserva
Biologiea Carara, cuenca del Rio Grande de TArcoles, Es-
taeidn Quebrada Bonita, E. Rojas 50 (CR, MO); canton de
Osa, Peninsula de Osa, Aguas Buenas Norte, Sendero Los
Guapinoles, Zamora et al. 1891 (CR, MO sterile); Reserva
Biologiea Carara, Sector Quebrada Bonita, Sitio Quebrada
Bonita, R. Zuniga 252 (CR, E, MO). San Jose: Carara
National Park, Rfo Carara near Carara Guard Post, Gentry
et al. 79559 (CR, MO sterile); Reserva Eorestal El Can-
grejo Mastatal de Puriscal, along road between Puriscal
and Quepos, forest patches along Rio Negro at base of
Cerro El Cangrejo, Hammel et al. 17120 (CR, F, MO).

Acknowledgments. I thank R. E. Gereau for
preparation of the Latin diagnosis; the curators of
CR and F for access to specimens; A. Jardim for
generously providing information; I .. Andersson
and M. V. L. Pereira-Moura for helpful review of
this manuscript; R. Foster, T. Wachter. and the An¬
drew Mellon Foundation through a grant to the
Field Museum of Natural History for logistical help
and support for travel to this institution; and 0. M.
Montiel and R. Magill for significant encourage¬
ment and help with this work.

Literature Cited

Andersson, L. 1994. Alseis. FI. Ecuador 50: 85—89.
- & C. Persson. 1991. Circumscription of the tribe

Cinchoneae (Rubiaceae) — A cladistic approach. PI.

Syst. Evol. 178: 65-94.

Burger, W. C. & C. M. Taylor. 1993. Flora Costaricensis,

Family #202, Rubiaceae. Fieldiana, Bot. n.s. 33: 1 —

333.

574

Novon

Robbrecht, E. 1988. Tropical Woody Rubiaceae. Opera
Rot. Belg. 1: 1-271.

- . 1993. Supplement to the 1988 outline of the clas¬
sification of the Rubiaceae. Index to Genera. Opera Hot.
Belg. 6: 173—196.

Standley, I’. C. & L. O. Williams. 1975. Flora of Guate¬

mala — Part XI (Rubiaceae). Fieldiana, Rot. 24(11): 1—
274.

Steyermark, J. A. 1974. Rubiaceae. In: T. Fasser (editor),
Flora de Venezuela 9: 1—2070. Instituto Botanieo, I)i-
receidn de Recursos Naturales Renovables, Ministerio
de Agricultura y Cria, Caracas.

A Synopsis of Persea (Lauraceae) in Central America

Henk van der Werff

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

henk.vanderwerff@mobot.org

Abstract. During preparation of a treatment of
Persea for the Flora Mesoamericana three new spe¬
cies were recognized. A description of these species
(Persea albiramea van der Werff, P. brevipetiolata
van der Werff. and P. laevifolia van der Werff) is
presented, as well as a key to all the known species
in the Flora Mesoamericana region including brief
rrotes giving synonyms, distribution, and noteworthy
features for these species.

Key words: Alseodaphne, Central America,
Lauraceae, Persea.

fhe genus Persea was described by Miller in his
Gardeners Dictionary, ed. 4, in 1754; the type spe¬
cies. P americana Miller, was described in the 8th
edition of the Gardener's Dictionary (1768). Since
then, a rather large number of species have been
placed in Persea, from both (sub)tropical Asia and
America. The genus has been variously defined and
is consequently quite variable in flower and fruit
characters. The American species were last revised
by Kopp (1966), who recognized two subgenera in
the neotropics. Persea subg. Persea is characterized
by having equal or subequal tepals, these reflexed
and usually deciduous in fruit, in having quadri-
locular anthers, stipitate glands at the base of the
stamens of the 3rd whorl, and a pubescent pistil;
the subgenus Eriodaphne Nees by its unequal te¬
pals (the inner 3 clearly longer than the outer 3),
the tepals persistent and patent in fruit, by its an¬
thers with a variable number of loeules (0. 2. or 4),
sessile glands at the base of the filaments of the
inner three stamens, and the pistil glabrous or pu¬
bescent. The main differences between the two sub¬
genera (equal tepals, these reflexed or usually de¬
ciduous in fruit vs. unequal tepals patent and
persistent in fruit) are easy to observe, and most
species can be placed in a subgenus without any
hesitation. However, in Kopp’s (1966) treatment she
also included a few anomalous species, which had
equal tepals, persistent and patent in fruit. Some
of these species are known from Central America.
Finally, there is one species from Suriname, P. ju-
lianae van der Werff, with strongly unequal tepals
that are deciduous in fruit. The seemingly clear di¬

vision of Neotropical Persea species in two subgen¬
era can no longer be maintained.

Kostermans (1993) added a further complication
by describing the new genus Mutisiopersea, which
differed from Persea solely by its persistent and
indurate tepals in the fruiting stage, whereas the
tepals in Persea s. str. are persistent or deciduous,
but never indurate. I do not accept Mutisiopersea
because 1 find the distinction between indurate and
non-indurate tepals vague; in some species the te¬
pals are clearly hardened and swollen in fruit, in
others slightly so or not at all. A few Central Amer¬
ican species of Persea were transferred by Koster¬
mans (1993) to Mutisiopersea.

Two of the species described in this contribution
(P. albiramea and P. laevifolia ) differ in several
characters from the other Persea species and war¬
rant further comment. Both species have the leaves
clearly clustered at the tips of the branches, their
leaves have a glaucous lower surface, and the te¬
pals are small, not exceeding 3 mm, and deciduous
or persistent in fruit. If persistent (in P. albiramea ),
the tepals are not enlarged and are visible as small
bracts at the base of the fruit. If deciduous (in P.
laevifolia), the tepals are united at the base and
dehisce as a unit, together with the stamens, unlike
the situation in P. americana, where the tepals ini¬
tially persist in young fruits and later break off in¬
dividually. Mature fruits of P. albiramea are seated
on a swollen pedicel, and fruits of P. laevifolia are
not yet known. This combination of characters is
also found in Alseodaphne Nees, an Asian genus
closely related to Persea (Chanderbali et ah, 2001),
and these two species would very likely be includ¬
ed iu Alseodaphne had they been collected in trop¬
ical Asia (van der Werff. 2001). Because of the
great variation in floral characters present among
the Neotropical species placed in Persea, it would
be very difficult to separate the new species from
Persea and I prefer to include them in Persea while
realizing that a modern study might well radically
change generic concepts in Persea s.l.

Type and density of indument are important
characters in the species delimitation of Lauraceae.
While density of pubescence is in general fairly

Novon 12: 575-586. 2002.

576

Novon

Table 1 . Spe<

i'ies groups of Persea in i

Central America.

Tepals of whorl

1 and 2

Tepals in fruit

Length of tepals

Position of leaves

Group 1

equal

deciduous

4—6 mm

evenly distributed

Group 2

unequal

persistent

3—5 mm

evenly distributed

Group 3

equal

persistent

2.5—5 mm

clustered

Group 4

(sub)equal

persistent or
deciduous

2-3 mm

clustered

constant within a species, I have found that some¬
times this character is much more variable in Per-
sea species than in species of other genera of Laur-
aceae. One example is the case of Persea bullata
Kopp. This species is only known from southern
Ecuador and was described as having densely pu¬
bescent twigs and pilose leaves. Subsequent col¬
lections have shown that this species can be en¬
tirely glabrous, and 1 have found glabrous plants
growing almost side by side with pubescent ones.
A comparable situation exists with P. boldufolia
Mez. Kopp (1966) cited only the type, collected
near Molinapampa, Dept. Amazonas, Peru. Around
Molinapampa large areas occur covered by a shrub¬
by vegetation growing on poor, sandstone-derived
soils. In this scrub I found not only plants exactly
matching the type, which is glabrous, but also
plants that are moderately pubescent and plants
that are densely pubescent. The densely pubescent
plants are similar to / \ ferruginea HBK. Plants
ranging from glabrous to densely pubescent grow
less than 100 m apart in the same vegetation, and
thus it seems reasonable to consider P. boldufolia
as a glabrous form of P. ferruginea. These examples
led me to identify specimens from Costa Rica with
a moderate to sparse, erect indument as the much
more densely pubescent P. donnell-smithii Mez; de¬
scribing a new species based only or largely on the
density of indument in this case seemed unwise.

Persea Miller, Card. Diet. ed. 4, 1754. TYPE: Per¬
sea americana Miller.

Mutisiopersea Kostermans, Rheedea 3: 133. 1993. Syn.
nov. TYPK: Mutisiopersea mutisii (HBK) Kostermans.

Trees and shrubs. Leaves alternate, evenly dis¬
tributed along the twigs or clustered, pinnately
veined, axillary tufts of hairs or domatia lacking.
Inflorescences in the axils of leaves or cataphylls,
panic ulate-cymose, lateral flowers of a cyme always
strictly opposite, blowers bisexual; tepals 6, equal
or unequal; if unequal, outer 3 tepals smaller than
the inner 3; tepals deciduous in fruit or persistent;

if persistent, usually remaining entire, rarely with
the tips of the inner tepals breaking off; stamens 9
or 6; if 6. stamens of whorl Ill staminodial; anthers
usually 4-eelled, rarely 2-celled; inner 3 stamens
with 2 glands at the base; staminodia of whorl IV
present, each with a sagittate or triangular apex,
f ruit with or without persistent tepals at the base,
but never with a cupule. 80—90 species in the neo¬
tropics, I species on the Canary Islands, ca. 100
species in tropical and subtropical Asia.

The altitudinal range given under the species is
taken from Central American specimens and may
vary in other parts of its distribution.

The Central American species of Persea can be
placed in the following species groups (Table 1):

• Species with equal tepals, these ultimately de¬
ciduous in fruit; tepals rather large (4-6 mm
long); subgenus Persea with P americana and P.
schiedeana Nees. Restricted to Central America,
but P. americana widely cultivated.

• Species with unequal tepals, these persistent in
fruit (part of the longer tepals sometimes break¬
ing off and tepals appearing equal in fruit); te-
pals usually 3—5 mm long; subgenus Kriodaphne
(including species placed in Mutisiopersea) with
12 species. An additional 50—60 species in
South America.

• Species with equal tepals, these persistent in
fruit; tepals 2.5—5 mm long. No name has been
proposed for this group; it includes 4 species, P
albida Kostermans with leaves evenly distribut¬
ed, and P. brevipetiolata, P. rigens C. K. Allen,
and P. silvatica van der Werff. with leaves clus¬
tered at the swollen tips of the branches. Also a
few species from South America and P. indica
(L.) Sprengel from the Canary Islands.

• Species with small (2—3 mm long), equal or sub¬
equal tepals, these persistent or deciduous in
fruit; pedicel swollen in fruit; leaves clustered,
glaucous below. Similar to Asian Alseodaphne,
including P. albiramea, P. laevifolia, and possi¬
bly P. perglauca Lundell. Also in South America

Volume 12, Number 4
2002

van der Werff

Persea in Central America

577

(P sphaerocarpa (Winkler) Kostermans from Bo¬
livia and an undescribed species from Ecuador).

Key to the Species of Persea in Central America

1 . Tepals equal or nearly so . 2

1'. Tepals unequal, the outer 3 clearly shorter
(not more than 2A the length of) than the inner

tepals . 11

2(1). Pistil pubescent; pedicels 5-15 mm long ... 3

2'. Pistil glabrous; pedicels to 5 mm long .... 4

3(2). Pedicels 5-8 mm long; bracts surrounding ter¬
minal buds uniformly pubescent; pistil sparse¬
ly to densely pubescent . P. americana

3'. Pedicels 10-25 mm long; bracts surrounding
the terminal buds with a scarious margin; pis¬
til densely pubescent . P schiedeana

4(2). Lower leaf surfaces glaucous; tepals 2-3 mm

long . 5

4'. Lower leaf surfaces not glaucous; tepals 3 mm
or longer (P. brevipetiolata with tepals 2.5 mm

long) . . 7

5(4). Leaves 15—30 cm long; petioles 15-40 mm

long . P. albiramea

5'. Leaves to 15 cm long, petioles to 16 mm long

. 6

6(5). Bark of twigs whitish; midrib impressed and
reticulation raised on the upper leaf surface;
terminal buds sparsely to moderately ap-

pressed pubescent . P. perglauca

6'. Bark of twigs dark brown; midrib and reticu¬
lation immersed on the upper leaf surface; ter¬
minal buds densely white pubescent ....

. P. laevifolia

7(4). Leaves whorled, clustered near the tips of the
branches; branches with thickened nodes from

which multiple branches emerge . 8

T . Leaves evenly distributed; tips of branches not

swollen, without multiple branches ... P albida
8(7). Inflorescences and flowers densely white pu¬
bescent, the surface of the flowers completely

covered . P rigens

8'. Inflorescences and flowers sparsely pubescent
or if densely pubescent, the indument brown

. 9

9(8). Petioles to 8 mm long; tertiary venation raised
on the upper leaf surface, forming a coarse
reticulum; flowers sparsely pubescent to

subglabrous . P. brevipetiolata

9'. Petioles 8-25 mm long; tertiary venation im¬
mersed or raised on the upper leaf surface; if
raised, forming a fine reticulum; flowers mod¬
erately to sparsely pubescent . 10

10(9). Leaves 3-6 cm wide, the base slightly decur¬
rent on the petiole; flowers moderately pubes¬
cent, the hairs ascending to erect . . . P rufescens
10'. Leaves 7-12 cm wide, the base not decurrent
on the petiole; flowers sparsely pubescent; the

hairs appressed . P- silvatica

11(1). Flowers with nine 2-celled stamens P. cuneata
11'. Flowers with six or nine 4-celled stamens 12
12(11). Flowers with six 4-celled stamens ....

. P. pseudofasciculata

12'. Flowers with nine 4-celled stamens .... 13

13(12). Outer tepals glabrous or nearly so on the out¬
er surface; inner tepals moderately to dense¬

ly pubescent on the outer surface, the differ¬
ence in indument between inner and outer

surface readily visible . P. povedae

13'. Indument, if any, on inner and outer tepals

similar . 14

14(13). Leaves sessile, the base cordate or rounded

. P. sessilis

14'. Leaves petiolate, the base never cordate 15
15(14). Twigs densely tomentose, the surface com¬
pletely covered; lower leaf surface moderate¬
ly to densely tomentose, the indument dis-

cernable to the touch . P. donnell-smithii

15'. Twigs glabrous or variously pubescent, but
not densely tomentose; lower leaf surface
usually glabrous or appressed pubescent; if
hairs ascending, then straight, not curly or

crisped . 16

16(15). Lower leaf surface covered by a short, dense,
felt-like indument, this sometimes difficult to
discern; individual hairs not or scarcely vis¬
ible . P. vesticula

16'. Lower leaf surface glabrous or with ap¬
pressed or ascending hairs; if pubescent, in¬
dument not felt-like and individual hairs vis¬
ible . 17

17(16). Leaves, twigs, and terminal buds glabrous;

leaves longer than 10 cm . P. standleyi

17'. Plants vegetatively with some indument (oc¬
casionally specimens of P. obtusifolia may be
glabrous, but this species has leaves gener¬
ally less than 10 cm long) . 18

18(17). Leaves and twigs initially pubescent, the
hairs erect or ascending, but soon becoming
glabrous; leaves coriaceous, rarely exceeding

10 cm in length . P obtusifolia

18'. Indument of twigs and leaves when present,
strictly appressed; leaves coriaceous or char-
taceous, usually exceeding 10 cm (to 10 cm

long in P. brenesii) . 19

19(18). Leaves elliptic, to 10 X 3.5 cm, petioles 1-

1.5 cm; indument on twigs and lower leaf

surface dense, consisting of coarse, ap¬
pressed hairs . P brenesii

19'. Leaves elliptic to broadly elliptic or ovate,
generally longer and/or broader than 10 X

3.5 cm; indument variable; if dense, then

hairs fine . 20

20(19). Inner tepals 3-4 times longer than the outer
tepals; flowers with a distinct, slender pedi¬
cel; distal part of the inner tepals breaking
off in fruit, thus fruit seemingly subtended

by 6 equal tepals . P caerulea

20'. Inner tepals twice as long as the outer ones;

flowers sessile or nearly so, without slender
pedicel; distal part of inner tepals not break¬
ing off in fruit, thus fruit subtended by 6

unequal tepals . 21

21(20). Inflorescences clustered at the base of sea¬
sonal growth or near the tip of the twigs, in
the axils of bracts or leaves, about as long as

the leaves . P veraguasensis

21'. Inflorescences ± evenly distributed along
the seasonal growth, mostly in the axils of
leaves, rarely more than half as long as the
subtending leaves . P liebmannii

578

Novon

Persea ulhidu Kostermans, Reinwardtia 7: 511.
1969. Replaced name: Persea pallida Mez <K
Piltier, Bull. Herb. Boissier, ser. 2, 3: 231.
1903, non (Nees) Oliver, Hooker’s Icon. PI. 14:
11. 1880. TYPE: Costa Rica. Valle de Goto,
Piltier l! I! I (isotype, G).

Persea guatemalensis Lundell, Wrightia 5: 37. 1974. Syn.
nov. TYPK: (Guatemala. Contreras 10943 (isotype,
MO).

Distribution. Guatemala, Honduras, Costa
Rica: possibly Chiapas (Breedlove 31300, CAS, a
fruiting specimen), from 1200 to 2100 m altitude.

Rest recognized by its densely pubescent flowers,
equal tepals, and evenly distributed (not clustered)
leaves. Differs from P americana in its glabrous
(not pubescent) pistil and shorter tepals (to 4.5 mm
long in P. albida, to 6 mm long in P. americana).

Persea nlhirumeu van der Werff, sp. nov. TYPE:
Costa Rica. Puntarenas: Canton de Osa, Ran¬
cho Quemado, Aug. 1991, Marin P2B (ho-
lotype, MO; isotype, INB). Figure 1.

A congeneris foliis magnis, subtus glaueis, ad apices
ramulorum aggregatis, fforibis |)arvis, glabris, tepalis su-
baequalibus diversa est.

Irees. to 30 m. Twigs thick, ridged or angular,
with pale gray bark, glabrous or with a few ap-
pressed hairs near the apex, grouped toward the tip
of leafless shoots, clusters ol bract scars lacking;
terminal buds densely pubescent, the hairs brown¬
ish. ascending, completely covering the surface, not
protected by bracts. Leaves 15-30 X 5-10 cm, el¬
liptic, chartaceous, grouped along the distal parts
of seasonal shoots, tin* base and apex acute, the
margin thickened, glabrous on both surfaces, or mi¬
nutely puberulous along the major veins on the low¬
er surface, the lower surface glaucous, midrib im¬
mersed or slightly raised, lateral veins and tertiary
venation immersed on the upper surface, midrib
and lateral veins raised, tertiary venation immersed
on the lower surface; lateral veins 10—16 on each
side; petioles 1.5-4 cm, ridged, glabrous, dark
brown and contrasting in color with the light-col¬
ored twigs. Inflorescences 7—15 cm, paniculate-cy-
mose, sparsely puberulous, in the axils of bracts
immediately below the tip of seasonal flushes.
Flowers 2.5-3 mm diam.. white, pedicels slightly
longer than the flowers. Tepals ca. 2 mm, the outer
3 slightly shorter than (7, to y. the length of) the

inner 3, erect at anthesis, sparsely puberulous or
glabrous on the outer surface, puberulous on the
inner surface; stamens 9, 4-celled, ca. 1.5 mm, the
filaments pubescent, about as long as the anthers,
slightly narrower than the anthers; outer 6 anthers
with the cells introrse. inner 3 with the cells ex-
trorse, filaments of the inner 3 stamens with 2 glo¬
bose glands at the base; staminodia 0.8 mm, broad¬
ly triangular, with a cordate apex, abaxially
pubescent, adaxially pubescent near tin* base only;
pistil 1.5 mm, glabrous, the style distinct and about
as long as the ovary; receptacle cup-shaped, dense¬
ly pubescent inside. Fruits 1.4 X 1.9 cm, wider
than long, tepals persisting at the base of the fruits,
reflexed, not enlarged or thickened; pedicel thick¬
ened toward the fruit, at the base 4 mm diam., near
the tip 8 mm diam.

Distribution. Costa Rica, Panama, from 200 to
600 m.

Persea albiramea is a verv distinctive species
and is easily recognized bv the combination of
large, glaucous leaves grouped near the tips of the
branches, the thick, pale twigs, and the1 long peti¬
oles. A s is the case with P. laevifolia, it does not
fit well in any of the proposed infrageneric taxa of
Persea due to its small flowers with subequal tepals
and the persistent, but not enlarged tepals in fruit.
It strongly resembles Asiatic species placed in Al-
seodaphne or Dehaasia and is likely closely related
to these species. An unusual feature, shared with
P. laevifolia, is the impressed tertiary venation on
the upper leaf surface. Most Persea species have
the tertiary venation raised and forming a fine re¬
ticulum. A related species from Bolivia is P. sphaer-
ocarpa (Winkler) Kostermans with large, clustered
leaves, these glaucous below, and thick twigs with
pale bark; it dilfers from P. albiramea in its erect
indument on the lower leaf surface and along the
major veins. Persea sphaerocarpa is only known
from the fruiting type collection. Several collections
from Ecuador also show the combination of clus¬
tered. glaucous leaves and thick twigs with pale
bark. These collections are either sterile or in bud,
and additional collections are needed for their
identification.

I lie Panamanian specimen included here differs
from the Costa Rican specimens in its glabrous
flower buds; vegetatively, it is a good match with
tin1 Costa Rican specimens.

— »

figure I (left). Holotype of Persea albiramea van der Werff.
figure 2 (right). Holotype of Persea brevipetiolata van der Werff.

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2002

van der Werff

Persea in Central America

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580

Novon

Paratypes. COSTA NIC A. Punlart ‘lias: Canton dr
Osa. G. Harm, A ./. Marin 1980 (1NB, MO). />'. Hummel
A' J. Mann 1831. >{ INB. MO), G. Herrera 5019 ( INB. MO),
K. Thomsen 401 (MO). PANAMA. Darien: Rio Tuquesa,
T. Croat 27170 (MO).

Persea americana Miller, Gard. Diet. ed. 8. 1768.
1 ai urns persea L. TYPE: Plate 222, fig. 2 in
Sloane (1 i25; lectotype, designated here).

Persea amerirana var. angustifolia Miranda, Anal. Inst.
Biol. Mexico 17: 126. 1946. TYPE: Mexico. Puebla,
Miranda 3482 (UNA not seen).

Persea drymifolia Schlechtendal A Chamisso, l.innaea (>:
365. 1831. Persea amerirana var. drymifolia (Schle¬
chtendal & Chamisso) Blake, J. Wash. Acad. Sri. 10:
15. 1920. Persea gratissima var. drymifolia (Schle¬
chtendal & Chamisso) Mez, Jahrb. Konigl. Bot. Cart.
Berlin 5: 147. 1889. TYPE: Mexico. Srhiede A- Dep-
pe 1 140 (isotype, MO).

Persea edulis Rafinesque, Sylva Tell. 134. 1838. Nom. su-
perfl. (or Persea amerirana Miller.

Persea Jlorrosa Mez, Jahrb. Konigl. Bot. Cart. Berlin 5:
148. 1889. I V PE: Mexico. Liebmann 85 (holotype,
C).

Persea gigantea I.. O. Williams. Ceiba 4: 39. 1953. TYPE:
Honduras. Williams A' Molina 12029 (holotype. EAP
not seen).

Persea gratissima Gaertner, Fruct. Sem. PI. 3: 222. 1807.
Nom. superfl. for Persea amerirana Miller.

Persea gratissima var. oblonga Meissner, DC. Prodr. 15(1):
53. 1864. TYPE: Mexico. Aschenborn 110 (syntype,
not seen), Galeotti 7010 (syntype, not seen); Peru.
Ruiz A Pavon s.n. (syntype, not seen); Brazil. Riedel
106 (syntype, not seen). Martins s.n. (syntype, not
seen): Java, Zollinger 457 A 3101 ex parte (syntype,
not seen).

Persea gratissima var. praerox Nees, Syst. Laurin. 129.
1836. Persea gratissima var. maerophylla Meissner,
DC. Prodr. 15(1): 53. 1864. TYPE: Peru. Poeppig
2446 (not seen).

Persea leiogyne Blake, J. Wash. Acad. Sci. 10: 19. 1920.
TYPE: U.S.A. Florida: Popenoe 219 (holotype, US
not seen).

Persea nubigena Williams, Ceiba 1: 55. 1950. Persea
amerirana var. nubigena (Williams) Kopp, Mem.
New York Bot. Card. 14: 19. 1966. TYPE: Guate¬
mala. Williams A Molina 16833 (isotype, MO).

Persea paucilriplinervia Lundell, Wrightia 5: 146. 1975
TYPE: G uatemala. Lundell A Contreras 19177 (iso¬
type, MO).

Persea steyermarkii Allen, J. Arnold Arbor. 26: 286. 1945.
TYPE: Guatemala. Steyermark 37061 (holotype, F).

Distribution. Widely cultivated in tropical and
subtropical countries for its edible fruits.

Persea amerirana is morphologically a variable
species and is closely related to P. schiedeana. It
can be separated from this species by its shorter
pedicels (to 8 mm long), the uniformly pubescent
bracts protecting the terminal buds, and its narrow¬
er leaves. Smith (1966) reported that Persea arner-
icana cotyledons were found in cave deposits ap¬
proximately 10.000 years old; thus die species has
been cultivated for a long period. Much of the var¬
iation in fruit shape and size as well as variation
in indument and leaf shape can be attributed to the
process of cultivation. A classification based on
tin *se characters (for instance, the one proposed by
L. 0. Williams in 1977) will only lead to the rec¬
ognition of cultivars; I prefer to accept Persea amer-
icana in a wide sense and ignore the cultivated
races of this species. It seems likely that P. atner-
icana originated in the highlands of Central Amer¬
ica (Kopp. 1966). However, 1 would not know how
to recognize the "wild" P. amerirana and include
all specimens matching the description given above
in P. amerirana. regardless of whether they come
from obviously cultivated or seemingly native trees.

I he type of P. leiogyne is from a cultivated tree;

I assume that the types of P americana, P. gratis¬
sima , and P. edulis also come from cultivated trees,
based on their origin outside the Central American
highlands or their specific epithets.

Iwo invalidly published names, Persea tolima-
nensis (Zentmyer A Schieber, 1990) and P. zent-
myerii (Schieber A Berg, 1987). have been applied
to P. amerirana s.l. These names were published
without a Latin diagnosis, and no type specimens
were designated.

Kopp (1966) designated the description in Clu-
sius (1601: 2) as the lectotype ol Laurus persea, but
because descriptions cannot serve as lectotypes ac¬
cording to Article 8.1 of the St. Louis Code (Greuter
et al„ 2000), a new one had to be chosen. The
illustration in Sloane (1725) is quite adequate, has
a brief description as well, and is backed by a spec¬
imen (no. 965) in the Sloane herbarium at BM.
rherefore, I designate here the illustration in Slo¬
ane ( I 1 25) as the lectotype of Laurus persea L. and
therefore of Persea amerirana Miller.

Persea hrenesii Standley, Publ. Field Mus. Nat.
Hist., Bot. Ser. 18: 458. 1937. Mutisiopersea
hrenesii (Standley) Kostermans. Bheedea 3:
134. 1993. TYPE: Costa Rica. La Palma de
San Ramon, Brenes 4451 (holotype, F; isotype,
CR not seen).

Distribution. Costa Rica, from 900 to 1200 m
altitude.

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2002

van der Werff

Persea in Central America

581

A poorly known species, best recognized by its
dense indument, consisting of coarse, appressed
hairs on twigs and lower leaf surface.

Persea hrevipetiolata van der Werff, sp. nov.
TYPE: Mexico. Vera Cruz: Mpio. Jesus Car¬
ranza, Apr. 1985, T. Wendt, ./. Garcui, W.
Thomas, ./. Grimes & A. Montero 4794 (holo-
type, CHAPA; isotype, MO). Figure 2.

Perseae rufescenlis siniilis, sed floribus inflorescenti-
isque glabris vel subglabris, reticulatione subtus elevata
et petiolis brevioribus recedit.

Small trees, to 8 rn. Twigs smooth, terete, solid,
glabrous, thickened at nodes, from which multiple
branches may emerge, a few bracts or bract scars
present at the base of seasonal growth; terminal
buds small, glabrous, protected by a few bracts.
Leaves 10-16 X 2.5—5 cm, elliptic, clustered near
the tips of seasonal growth, coriaceous; the base
acute or somewhat decurrent on the petiole, apex
gradually acute, glabrous on both surfaces, midrib,
lateral veins, and tertiary venation weakly raised
on tbe upper surface, the tertiary venation forming
a coarse reticulum, midrib, lateral veins, and ter¬
tiary venation raised on the lower surface; lateral
veins 7-10 pairs; petioles 5-8 mm. glabrous, flat
above, with the same color as the twigs. Inflores¬
cences 10-15 cm, paniculate-cymose, glabrous or
nearly so, often with a few bracts or bract scars at
the very base, in the axils of leaves or bracts at the
nodes. Flowers 4 mm diam., green-yellow, pedicels
5 mm. Tepals equal or subequal, with the outer
tepals slightly shorter, 2.5 mm, broadly elliptic,
erect or weakly spreading at anthesis, the outer sur¬
face glabrous or very sparsely and minutely pubes¬
cent, the inner surface glabrous or nearly so, sta¬
mens 9. 4-celled. ca. 2 mm, the anthers as long as
the filaments, anthers glabrous, filaments pubes¬
cent. filaments half as wide as the anthers; outer 6
stamens with introrse cells, inner 3 with extrose-
lateral cells, filaments of the inner stamens with 2
globose glands near the base; staminodia present,
1.3 mm, broadly triangular, abaxiallv pubescent;
pistil 2 mm, glabrous, the ovary 1.2 mm, distinct
from the narrow style; receptacle shallowly cup¬
shaped, glabrous inside. Fruits globose, ca. 2 cm
diam., tepals persistent in fruit, but not enlarged,
spreading, the pedicel a little thickened and warty.

Distribution. Persea hrevipetiolata is only
known from three collections from Mexican lowland
forests (200—250 m altitude) on the Isthmus of Te¬
huantepec. It is closely related to P rufescens Fun¬
ded (Fig. 3), which occurs in montane forests be¬

tween 1500 and 2500 m in Chiapas and Oaxaca,
Mexico; it differs from this species in its glabrous
(or nearly so) flowers and inflorescences, in the
raised reticulation on the lower leaf surface, and in
its shorter (to 8 mm long vs. 15-25 mm in P. ru¬
fescens) petioles. Persea hrevipetiolata is also close
to the Costa Rican-Panamanian P. rigens C. K. Al¬
len and P. silvatica van der Werff. The former can
be recognized by its dense, white indument on flow¬
ers and inflorescences; the latter differs in its larger
and wider leaves and longer inflorescences. All four
species in this group are rarely collected, and more
collections are needed for a better understanding
of this group.

Para types. MEXICO. Oaxaca: Mpio. Mati'as Romero,
Hernandez & Sanchez U-16 (CHAPA). Vera Cruz: Mpio.
Hidalgotitlan, T. Wendt & I. Navarrete 3252 (MO).

Persea caerulea (Ruiz & Pavon) Mez, Jahrb. kon-
igl. Rot. Cart. Berlin 5: 171. 1889. Lauras ca¬
erulea Ruiz & Pavon. FI. Peruv. 4: t. 350.
1802. TYPE: Peru. Ruiz & Pavon s.n. (M A not
seen).

Persea petiolaris HBk. Nov. Gen. Sp. 2: 159. 181 < . 1 'i IT.:
Colombia. Humboldt & Bonpland s.n. (B not seen).

Persea laevigata HBK. Nov. Gen. Sp. 2: 157. 181 7. IV PE:
Colombia. Humboldt s.n. (B not seen).

Persea lignilepala Lasser. Bol. Soe. Venez. Ci. Nat. 9: 177.

1944. TYPE: Venezuela. Pittier 7541 (holotype,
VEN not seen).

Persea skutchii C. K. Allen, .1. Arnold Arbor. 24: 289.

1945. TYPE: Costa Rica. Skutch 4812 (holotype, GH
not seen).

Distribution. Honduras, Nicaragua, Costa Rica,
Panama, South America, from 500 to 1800 m alti¬
tude.

A common species characterized by its strongly
unequal tepals, with the outer ones Ys to Va the
length of the inner ones and small (1 cm diam.),
greenish blue fruits. Vegetatively, it resembles P.
americana, which has equal tepals and much larger
fruits.

Persea cuneata Meissner. DC. Prodr. 15(1): 46.
1864. Beilschmiedia cuneata (Meissner) Kos-
termans, Bol. Teen. Inst. Agron. N. 28: 59.
1959. Mutisiopersea cuneata (Meissner) kos-
termans, Rheedea 3: 134. 1993. TYPE: Co¬
lombia. Jervise (K not seen).

Distribution. Costa Rica, Panama, northern
South America, from 1000 to 1900 m altitude.

The only species of Persea with nine 2-celled
stamens in Central America. Vegetatively very' sim-

582

Novon

ilar to P. povedae Burger, but this species has larger
leaves, (lowers (with 4-celled stamens), and fruits.

Persea doiinell-smilhii Me/, Arbeiten Konigl.
Hot. Gart. Breslau 1: 113. 1892. Mutisiopersea
donnell-smithii (Mez) Kostermans, Bheedea 3:
134. 1993. TV PE: Guatemala. Alta Verapaz,
J. Donnell Smith 1718 (syntype, MO); Mexico.
Liebmann Laurac. 1 1 (syntype, C not seen).

Distribution. Chiapas, Guatemala, Honduras,
Nicaragua, Costa Rica, from 900 to 2400 m alti¬
tude.

A rather common, montane species characterized
by its unequal tepals and dense, erect indument on
twigs and lower leaf surface. Collections from Costa
Rica placed here have a sparser indument than the
collections from Nicaragua northward. It is similar
to Persea charnissonis Mez from Mexico in indu¬
ment, but the latter has narrower leaves. It is also
possible that I1, donnell-smithii is nonspecific with
P. charnissonis ; the fact that a syntype of P. donnell-
smithii was collected in Veracruz, Mexico, lends
support to this idea.

Person luevifolia van der Werff, sp. nov. TYPE:
Costa Rica. Heredia: Chilamate de Sarapiqui,
May 1989, H. Hummel & M. Grayum 17334
(holotype, MO; isotypes, BM, BO, CANB, E,
HBG, INB, KLIN. L, EE. MEXU, MO, NY. P.
PMA, TI, US, XAE). Eigure 4.

Perseae perglaucae similis, sed ramulis fuscis, gemmis
terminalibus dense pubescentibus costisque immersis re-
cedit.

frees, to 20 m. Twigs slightly ridged, glabrous or
nearly so, dark brown, the branching not clustered,
clusters of bract scars lacking; terminal buds
densely white pubescent, not protected by bracts.
Eeaves 7—15 X 2-4 cm, chartaeeous, elliptic to
slightly obovate, alternate and clustered near the
tips of the branches; base narrowly euneate and
somewhat decurrent on the petiole, the margin
thickened, the apex acute or shortly acuminate, gla¬
brous on both surfaces, midrib immersed or weakly
raised, lateral veins and tertiary venation immersed
on the upper surface, midrib raised, lateral veins
and tertiary venation immersed on the lower sur¬
face, lower surface glaucous; lateral veins 7-10 on
each side, poorly visible; petioles 7-14 mm, gla¬
brous, canaliculate. Inflorescences 4—12 cm, panic¬

ulate-cymose, sparsely pubescent when immature,
glabrous in young fruiting stage, near the base of
seasonal flushes. Flowers unknown; description of
floral parts based on remnants at the base of young
fruits. Tepals 1.7 mm, broadly elliptic, equal, the
outer surface sparsely to moderately, the inner sur¬
face moderately to densely pubescent; stamens 9,
all 4-cel led, but often only 2 or 3 cells opening,
the inner 3 stamens sometimes with the upper 2
cells vestigial, the outer six I mm, the anther clear¬
ly wider than the filament, the cells introrse, anther
about as long as the basally pubescent filament;
inner 3 stamens as long as the outer 6, anther
slightly wider than the filament, filament as long as
the anther, densely pubescent, with large glands
near the base, staminodia 0.8 mm, broadly trian¬
gular, pubescent on both surfaces, pistil glabrous.
Young fruits globose, the largest seen 6 mm diam.,
tepals in very young stage persisting, but ultimately
dehiscing as a unit with the stamens attached or,
less frequently, individually.

Distribution. Costa Rica, only known from the
lower Atlantic slope. 100—200 rn.

Persea laevifolia is an unusual Persea species
because of its small, equal tepals, which are de¬
hiscent in the fruiting stage. Thus, it does not fit
well in the previously recognized infragenerie taxa
of Persea. Most Neotropical species of Persea have
unequal tepals or equal, persisting tepals or equal,
deciduous tepals that are not basally united. Nearly
all have clearly larger flowers. Persea laevifolia re¬
sembles P. perglauca from Guatemala vegetatively,
but differs in having dark brown (not very pale to
whitish) twigs, a densely pubescent (not sparsely
pubescent) terminal bud, immersed (not raised or
slightly raised) lateral and tertiary veins on both
surfaces, the leaf bases slightly decurrent on the
petioles and the midrib immersed (not impressed)
on the upper leaf surface. These differences are
slight, but constant, and I expect that when good
flowering collections become available floral differ¬
ences will be found as well.

Paralypes. COSTA BICA. Heredia: Chilamate, II.
Hammel A- /. Chacon 16055 (INB, MO). II. Hammel cl al.
14135 (MO), C. Herrera 4987 (INB, MO); Sar apiqui, Kan-
cho El Bejuco, II. van der Werff el al. 14045 (INB. MO).

Eigure 3 (left). Persea rufescens.

figure I (right). Holotype of Persea laevifolia van der Werff.

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van der Werff

Persea in Central America

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584

Novon

Persea liebmannii Mez, Jahrb. Konigl. Bol. Cart.
Berlin 5: 166. 1886. Mutisiopersea liebmannii
(Mez) Kostermans, Rheedea 3: 135. 1993.
1 YPE: Mexico. Oaxaca: Liebmann 115 (syn-
type, C not seen). Liebmann 1 16 (syntype, C
not seen, photo MO).

Persea flavifolia Lundell, Contr. Univ. Michigan Herb. 6:
17. 1941. TYPE: Mexico. Mahnla 1821 (isotype,
MO).

Persea petenensis Lundell. Wrightia 4: 109. 1969. Syn.
nov. TYPE: Ouatemala. Contreras 902 (isotype, MO).

Persea podadenia Blake, Contr. Gray Herb. 52: 62. 1917.
TYPE: Mexico. Palmer II 9 (isotype, MO).

Distribution. Mexico. Guatemala, from 2000 to
2500 m.

Can he recognized by its rather large (to 16 cm
long and 8 cm wide) leaves, inflorescences evenly
distributed along the twigs, and appressed indu-
ment. Vegetatively similar to P. veraguasensis See-
mann from Panama and Costa Rica, which has lon¬
ger inflorescences clustered near the tips of the
branches.

Persea ohtusifolia Kopp, Mem. New York Rot.
Card. 14: 81. 1966. TYPE: Panama. Chiriqui:
Cerro Copete, Allen 1888 (isotype, MO).

Distribution. Costa Rica, Panama, from 2500 to
3300 m.

Best recognized by its rather small (to 10 cm
long), coriaceous leaves, which have, when young,
an erect or ascending indument, but which become
quickly glabrous with age.

Persea perglauca Lundell, Wrightia 5: 147. 1975.
TYPE: Guatemala. Baja Verapaz, Lundell di-
Contreras 19217 (isotype, MO).

Distribution. Guatemala; altitudinal distribu¬
tion not known.

Typical are the whitish twigs contrasting with the
dark petioles, clustered leaves, which are glaucous
on the lower surface, the impressed midrib, and the
small flowers with tepals 2 mm long. Similar to P.
laevifolia, which has darker twigs and an immersed
midrib. It is possible that P. perglauca is only a
small-flowered variety of P. americana.

Persea povedae Burger, Eieldiana Bot., n.s. 23:
105. 1990. TYPE: Costa Rica. Alajuela: Po-
veda 740 (isotype, MO).

Distribution. Costa Rica, Panama, Ecuador, Ve¬
nezuela, from 700 to I 100 m.

Readily recognized by the indument on the te¬

pals (inner 3 tepals much more densely pubescent
than the outer 3) and its 4-celled stamens. Similar
to P. cuneata, which has 2 -celled stamens.

Persea pscudofaseirulata Kopp, Mem. New York
Bot. Card. 14: 85. 1966. TYPE: Bolivia. La
Paz, Krukoff 11288 (isotype, MO).

Distribution. Costa Rica (I collection). South
America, at 500 m.

The only species of Persea with six fertile sta¬
mens and whorl III staminodial in Central America.
The inflorescences are short, branched, and dense¬
ly pubescent.

Pt -rsea rigens Allen, J. Arnold Arbor. 26: 297.
1945. I \ IT,: Panama. Chiriqui: Little 6058
(isolectotype, MO).

Distribution. Costa Rica, Panama, probably Co¬
lombia, from 1360 to 1600 m.

A poorly known species with clustered leaves,
thickened branch tips, and densely white-pubes¬
cent flowers. Similar to P. silvatica, which has less
pubescent flowers. Kopp (1966) discussed the
problems with the type designated by Allen. She
concluded that the holotype ( Little 6075 , E) was a
mixed collection consisting of a sterile tw ig of Little
6075 and a flowering twig of Little 6058 and se¬
lected the flowering twig of Little 6058 in E as the
leetotype. On labels of duplicates of Little 6075 (for
instance, in MO) it is clearly mentioned that no
flowers or fruits were in evidence, while labels of
Little 6058 state that immature flowers were pre¬
sent. Allen (1945) described flowers of P. rigens
and thus must have seen material of Little 6058.
Both Little 6058 and 6075 were collected in Pan¬
ama. Allen’s statement that Little 6075 was col¬
lected in Costa Rica is incorrect.

Persea rufescens Lundell, Wrightia 5: 38. 1974.

I YPE: Mexico. Chiapas: Matilda 5394 (iso¬
type, MO).

Distribution. Oaxaca, Chiapas, from 1500 to
2550 m.

A poorly known species with clustered leaves,
thickened branch tips, and moderately brown-pu¬
bescent flowers, with spreading to ascending hairs.
See under P. brevipetiolata for differences with that
species.

Persea schiedeana Nees, Sy st. Laurin. 130. 1836.
Persea gratissima var. schiedeana (Nees) Meis¬
sner. DC. Prodr. 15(1): 53. 1864. TYPE: Mex¬
ico. Misantla, Schiede 1141 (holotype, B not
seen).

Volume 12, Number 4
2002

van der Werff

Persea in Central America

585

Persea pittieri Mez, But. Jahrb. Syst. 30, Beibl. 67: 15.
1901. TYPE: Costa Rica. Pitlier 1156 (holotype, BR
not seen).

Distribution. From Mexico to Panama, at 800-
2500 altitude.

Readily identifiable by its large flowers (tepals
6-8 mm long) and long pedicels (10-25 mm long).
Its 1 eaves are somewhat clustered and to 20 cm
broad.

Persea sessilis Standley & Steyermark, Publ.
Field Mus. Nat. Hist., Bot. Sen 23: 1 15. 1944.
TYPE: Guatemala. Zacapa, Steyermark 424H7
(holotype, F).

Distribution. Guatemala, at 2100—2400 m.
Rarely collected, but easily identified by its ses¬
sile leaves. Flowers and fruits poorly known, but
definitely a Persea species. Only known from die
type collection.

Persea silvatica van der Werff, Fieldiana Bot.. n.s.
23: 107. 1990. TYPE: Costa Rica. Heredia:
Schatz & Young 964 (holotype, MO).

Distribution. Costa Rica, at 80—300 m.

Best recognized by its clustered leaves, swollen
branch tips, and sparsely pubescent flowers. Close
to P. rigens , which has densely white-pubescent
flowers.

Persea stamllcyi Allen. J. Arnold Arbor. 26: 301.
1945. Mutisiopersea standleyi (Allen) Koster-
mans, Rheedea 3: 135. 1993. TYPE: Guate¬
mala. Solola: Steyermark 47 ISO (holotype, F).

Distribution. Guatemala, Honduras, from 1500
to 2 1 00 m.

Poorly known and best recognized by its glabrous
leaves, twigs, and vegetative buds.

Persea veraguasensis Seemann, Bot. Voy. Herald
193. 1854. Mutisiopersea veraguasensis (See¬
mann) Kostermans, Rheedea 3: 135. 1993.
TYPE: Panama. Veraguas: Volcan Chiriquf,
Seemann I57P (RM, photo MO).

Distribution. Costa Rica. Panama, from 1200 to
2000 m.

Similar to P. liebmannii, but differs in having the
inflorescences about as long as the leaves and clus¬
tered at the base of seasonal growth. Meissner
(1864) erroneously used the name P. veraguensis for
this species.

Persea vesticula Standley & Steyermark, Publ.
Field Mus. Nat. Hist., Bot. Ser. 23: 1 16. 1944.
Mutisiopersea vesticula (Standley & Steyer¬
mark) Kostermans, Rheedea 3: 135. 1993.
TYPE: Guatemala. San Marcos, Steyermark
36207 (holotype, F).

Persea chiapensis Lundell, Wrightia 1: 150. 1946. TYPE:
Mexico. Chiapas: Matuda 549H (isotype, MO).

Persea popenoei L. O. Williams, Ceiba 1: 57. 1950. T\ PE:
Honduras. Williams & Molina 15692 (isotype, MO).

Distribution. Chiapas, Guatemala. Honduras,
El Salvador, from 1800 to 2500 m.

Characterized by a short, dense, felt-like indu-
ment on the lower leaf surface, which may be dif¬
ficult to discern. Specimens from Costa Rica pre¬
viously identified as P. vesticula are here placed in
P. obtusifolia. These specimens lack the dense, felt¬
like indument on the lower leaf surface and instead
have longer, straighler hairs, which may wear off
with age.

Exci.i dki) Species

Persea primatogena Williams & Molina = Beilsch-
miedia riparia Miranda (Nishida, 1999).

Acknowledgments. 1 thank T. Wendt for making
the holotype of Persea brevipetiolata available and
Fred Keusenkothen for scanning the herbarium
specimens from which the illustrations were made.

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inary survey of the Mexican and Central American spe¬
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Clusius, C. 1601. Rariorum plantarum historia. Antwer¬
pen, pp. 1 —364.

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Nomenclature (Saint Louis Code). Regnum Veg. 138.
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in the Western Hemisphere (Persea— Lauraceae). Mem.
New York Bot. Card. 14: 1 — 117.

Kostermans, A. J. G. H. 1993. Mutisiopersea Kostermans,
a new genus in Lauraceae. Rheedia 3: 132—135.
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itor), Prodromus systematis naturalis regni vegetabilis
15(1): 1-260.

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in the Neotropics. Ann. Missouri Bot. Card. 86: 657—
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Schieber, E. t!4 B. Bergh. 1987. Persea zentmyerii, a new
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van der Werff, II. 2001. An annotated key to the genera
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A new species for Central America. Acta llort. 275:
383-386.

Isoetes yunguiensis (Isoetaceae), a New Basic Diploid Quillwort

from China

Wang Qing-Feng and Liu Xing

Herbarium, College of Life Sciences, Wuhan University, Wuhan 430072, People’s Republic
of China, wangqf97@public.wh.hb.cn and evotaxon@whu.edu.cn

W. Carl Taylor

Department of Botany, Milwaukee Public Museum, 800 West Wells Street, Milwaukee,
Wisconsin 53233-1478, U.S.A. ct@mpm.edu

He Zhao- Hong

Herbarium, Laboratory of Pteridophyta, Institute of Ecology & Geobotany, Yunnan
University, Kunming, Yunnan 650091, People’s Republic of China.
zhaorong_he@yahoo.com.cn

ABSTRACT. Isoetes yunguiensis, a basic diploid
species with a chromosome number of 2 n = 22. is
described and illustrated. This species occurs on
the Yunnan— Guizhou Plateau in southwest China.
It is similar to /. japonica in its appearance and
emergent aquatic habit. Herbarium specimens of /.
yunguiensis previously have been misidentified as
/. japonica. Isoetes yunguiensis differs from /. ja¬
ponica in megaspore and microspore morphology
and chromosome number.

Key words: China, East Asia, Isoetaceae. Isoe¬
tes, quillwort, Yunnan-Guizhou Plateau.

Two basic diploid species (2 n = 22), Isoetes hyp-
sophila Handel-Mazzetti and /. taiwanensis DeVol,
one tetraploid species (2 n = 44), Isoetes sinensis
Palmer, and one hexaploid species (2re = 66), /.
japonica A. Braun, have been reported from China
(Diao, 1990: DeVol, 1972). Over the last four years,
field, herbarium, and laboratory studies have been
conducted on t lie genus Isoetes in the vast area of
mainland China, where quillworts are increasingly
threatened with extirpation due to ongoing anthro¬
pogenic changes. During the course of these stud¬
ies, specimens collected from the Yunnan-Guizhou
Plateau in southwest China that superficially re¬
sembled and had been identified as I. japonica were
found in herbaria. Study of these herbarium spec¬
imens and live specimens from the field revealed
that they were different from I. japonica in spore
morphology and chromosome number. These spec¬
imens represent a previously undescribed species
of Isoetes.

Isoetes yunguiensis Wang Q. K & W. C. Taylor,
sp. nov. TYPE: China. Guizhou: Pingba Coun¬
ty, Shashi Valley, plants in J uncus bufolius—
Eriocaulon schochianum marsh, along a stream
originating from the Shashi Valley Reservoir,
ca. 4 km E of Pingba Town, 9 Aug. 2001,
Wang Q. F., Liu X. & Yang X. L. WH2001 166
(holotype, WH). Figures 1. 2.

Planta aquatica, submersa vel emergens ex caudice tri-
lobo. Folia 20 ad 70, viridia, 10-52 cm longa, patentis-
sima, basi dilatata et complanata, marginibus membran-
aeea; ligula triangulata, cuspidata, 1.0— 2.5 mm longa,

1 .5— 3.0 mm lata. Sporangium basale, ovale, 2—7 mm lon-
gum, 1.5— 4.5 mm latum; velum brevissimum, fibris peri-
phericis destitutum. Megasporae albae, 340-430 gm dia-
metro, cristate-reticulatae. Microsporae pallide canae in
inassa dispositae, ellipticae, 20—25 gm longae, laeves vel
granulatae. Chromosomatum numerus 2 n — 22 (Fig. 21).

Plants aquatic, submersed or emergent. Root-
stock 3-lobed. Leaves spirally arranged, widely
spreading, 1 0—52 cm long, ca. 2.5 mm wide at mid
length, in tufts of 20 to 70, flattened on upper side,
rounded on lower side (Fig. 1C), base expanded
with membranous margins, peripheral fibers absent.
Ligule triangular, cuspidate, 1-2.5 X 1.5— 3.0 mm.
Velum rudimentary, covering only the distal edge
of the sporangium. Sporangia basal, oval, 2—7 X

1 .5- 4.5 mm (F ig. I B). Megaspores gray when wet,
white when dry, 340-430 gun diam. (mean = 390
gun), with cristate-reticulate ornamentation (Fig.
2A-E). Microspores gray in mass, elliptic, 20—25
gun long (mean = 23 gun), laevigate to granulate
(Fig. 2F-H). Some sporangia contain both mega¬
spores and microspores. The megaspores in these
mixed sporangia are slightly smaller than those

Novon 12: 587-591. 2002.

588

Novon

Figure 1. Isoetes yunguiensis Wang U. F. & W. C. Taylor. — A. Habit. — 15. Adaxial view of leaf base. — C. Transsection
of leaf at mid length. Scale bar: A = 2 cm; H = I cm; C = I mm.

found in normal megasporangia. Chromosomes: 2 n

= 22.

Distribution. In China, Kunming City and Xim-
dian County of Yunnan Province; Pingba County
and Guiyang City of Guizhou Province.

The specific epithet for this species is derived
from die names Yunnan and Guizhou, the provinces
from which the new species lias been collected.
These two provinces constitute t he main section of

the Yunnan— Guizhou Plateau in the upper reaches
of the Yangtze River in southwest China.

Spore morphology. Spores of /. yunguiensis
were obtained from specimens collected at the
Shashi Valley and Shuangshao Village localities.
Samples were prepared for scanning electron mi¬
croscopy following the methods of Huang et al.
(1992). Observations, measurements, and photo¬
micrographs were made using a Hitachi S-800

Volume 12, Number 4
2002

Wang et al.

Isoetes yunguiensis from China

589

Figure 2. Isoetes yunguiensis Wang Q. F. & W. C. Taylor. — A. Megaspores. — B. Proximal view of megaspore. — C.
Distal view of megaspore. — D. Equatorial view of megaspore. — E. Megaspore surface. — F. Microspores. — G. Micro-
spore. — II. Microspore surface. — I. Somatic chromosomes in mitotic root lip squash (Chu IF M. & He Z. R. 29306
(MIL)). Scale bar: A = 200 gin; B & I) = 75 gm; C = 50 gun; E & F = 15 gm; G = 4 gun; H = 1 gun; I
1 1 ) gun.

scanning electron microscope anil with Olympus
BX60 and SZX12 research microscopes. Mega¬
spores of /. yunguiensis have a cristate, irregularly
reticulate texture with the crests of the ridges un¬
even (Fig. 2 A— E). These megaspores are 360—450
gun diam. (x = 390 gun diani., N = 30). In contrast,
the megaspores ol I. japonica have a more regularly
reticulate texture with the crests of t lie ridges more
uniform (Takamiya, 1999: 124, fig. I OB). The

megaspores of /. japonica are larger, 390—550 gun
diam. (x = 500 gun diam., N = 30).

The cristate-reticulate megaspores of /. yun¬
guiensis clearly differ from the other basic diploid
species in East Asia. Isoetes asiatica (Makino) Ma-
kino has echinate megaspores (Takamiya et al.,
1997), /. hypsophila has laevigate megaspores
(Palmer, 1927), and /. taiwanensis has rugulate-re-
ticulate megaspores (Huang et al., 1992; Britton X

590

Novon

Brunton, 1991). The tetraploid /. sinensis has mega¬
spores with a cristate texture (Takamiya et al.„
1997: Takamiya, 1999).

Microspores ol /. yunguiensis have a laevigate to
verrucate texture (Fig. 2K. G). At higher magnifi¬
cations, using the scanning electron microscope,
the microspores are seen to have a granulate texture
(Fig. 211). In texture, the microspores of /. yun¬
guiensis are similar to l. japonica (Takamiya, 1999:
124, fig. 10E), but /. yunguiensis microspores are
smaller than /. japonica microspores. The micro¬
spores of /. yunguiensis are 20-25 gm long and
average 23 gun long. In contrast, the microspores
of /. japonica are 28-34 gun long and average 31
gun long.

Although the megaspores and microspores of /.
yunguiensis differ in size from those of /. japonica,
I. yunguiensis is similar to /. japonica in its ap¬
pearance and emergent aquatic habit. They are
much alike in eorm, leaf, sporangium, and velum
characters. Both species occur in marshes, ponds,
and streams. Early descriptions of Isoetes species
were based on habit and habitat, and this could be
the reason all of the /. yunguiensis plants deposited
in different herbaria in China were identified as /.
japonica.

Cytology. Plant materials for chromosome
counts were collected from the Shashi Valley and
Shuangshao Village localities and cultured in pots
at the Botanical Garden, College of Fife Sciences,
Wuhan University, and in the Botany Department
at the Milwaukee Public Museum. Bool tips were
prepared for observation of somatic chromosomes
following the methods ol Takamiya et al. ( 1994) and
Taylor and Fuebke (1988). Chromosome counts
from 70 cells in root tip squashes from 14 individ¬
uals showed that /. yunguiensis is a basic diploid
with the chromosome number of 2 n = 22 (Fig. 21).
The discovery of this basic diploid species brings
the total of basic diploids in East Asia to four, in¬
cluding /. asiatica, I. hypsophila, and /. taiwanen-
sis, out of a total of about eight species.

Isoetes yunguiensis is the only East Asian, basic
diploid species with a vegetative morphology sim¬
ilar to the hexaploid /. japonica. Therefore, one
could hypothesize that /. yunguiensis is involved in
the evolution of /. japonica. It is possible that /.
japonica is an autohexaploid, arising through au-
t()| )olvploidy in the basic diploid /. yunguiensis. Al¬
ternatively, it is possible that /. japonica is an al-
lohexaploid arising through allopolyploidy
involving a cross between /. yunguiensis and a tet¬
raploid species to produce a sterile triploid hybrid
that doubled its chromosome number. Relation¬

ships among the basic diploid and polyploid Isoetes
species distributed in East Asia, such as those be¬
tween the basic diploid /. yunguiensis and the hexa¬
ploid /. japonica, need to be investigated by eyto-
logical and molecular studies.

Ecology. The streams, marshes, and ponds in
which /. yunguiensis grows belong to the river sys¬
tems of the Yangtze River on the Yunnan— Guizhou
Plateau at an altitude of 1200-1900 m. Water pH
value here is about 6.3. As with most quillworts, /.
yunguiensis is easy to overlook, for it blends in with
associated species such as Juncus bufonius L., Leer-
sia hexandra Swartz, Rotala rotundifolia (Buchan-
an-Hamilton) koehne, Mariscus umbellatus Vald.
HippurL s' vulgaris L., Eriocaulon schochianum Han-
del-Mazzetti, and Hypericum japonicum Thunberg
ex Murray.

Megaspores of /. yunguiensis mature from the
middle of June to the end of September. Micro¬
spores develop from the end of June to the middle
of October. By the end of October most sporophylls
have decayed, and only a few small leaves that lack
sporangia remain green during the winter. A similar
overwintering condition is found in /. sinensis.

Like all other quillworts in China, /. yunguiensis
is a rare and endangered species. To date it has
been found at only four sites. Field investigations
have revealed that two of these populations have
been decimated. Of the remaining two sites, the
Shuangshao population covers about 100 square
meters and contains less than 100 plants, and the
Pingba population covers about 400 square meters
and holds 200 to 300 individuals. Pollution and
increasing urbanization in China continue to raise
the risk of extinction for this rare species. Searches
for more field localities as well as conservation
studies on this species are urgently needed.

Paratypes. CHINA. Guizhou: Pingba County, Shashi
Valley, Wang P S. 75043, 75425 (COB I . HGAS); Guiyang
City, Wangwu, Peng //., Wang P. S. & Zhao P. 70060
(HGAS). Yunnan: Kunming City, Heilongtan, Chit Hi M.
3636 (PY II. W 11); Xundian County, Tianshengqiao, Song
Z. II. 80902 (PY U. \\ H); Kunming City, Shuangshao Vil¬
lage, swamp near reservoir, Chu W. M. & He Z. I\. 29306
(MIL. PYU).

Acknowledgments. We are grateful for the as¬
sistance and cooperation of the curators of CDBI,
HGAS. kUN, NAS. PE, and PYll. We also thank
Guo You-Hao, College of Life Sciences, Wuhan
University, for the Latin description of the species.
We have benefited from discussions with Chu Wei-
Ming. Yunnan University. We acknowledge Wang
Pei-Shan of Guizhou Academy of Sciences for his
kind and keen-eyed assistance in the field. Wang
Yong and Chen Bao-lian helped us with the line

Volume 12, Number 4
2002

Wang et al.

Isoetes yunguiensis from China

591

drawings. Neil T. Luebke, Botany Department, Mil¬
waukee Public Museum, produced the chromosome
figure. The State Key Basic Research and Devel¬
opment Plan of China (G2000046805) supported
this research.

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Palmer, T. C. 1927. A Chinese Isoetes. Amer. Fern J. 17:
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Takamiya, M. 1999. Natural history and taxonomy of Is¬
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- , M. Watanabe & K. Ono. 1994. Biosystematie

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- . - & - . 1997. Biosystematie studies

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Taylor, W. C. & N. T. Luebke. 1988. Isoetes X hickeyi: A
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Volume 12, Number l, pp. 1 — 154 of NOVON was published on 2 April 2002.
Volume 12. Number 2, pp. 155—308 of NOVON was published on 8 July 2002.
Volume 12, Number 3. pp. 309-432 of NOVON was published on 25 September 2002.
Volume 12, Number 4, pp. 433-591 of NOVON was published on 26 December 2002.