Volume 13
Number 1
2003
NOVON
A New Species of Doliocarpus and a New Species of Tetracera
(Dilleniaceae) from Brazil
Gerardo Ay mard C.
UNELLEZ-Guanare, Programa de R. N. R., Herbario Universitario (PORT), Mesa de
Cavacas, Estado Portuguesa, Venezuela 2333
ABSTRACT. During the course of a taxonomic
study of the Neotropical species of Dilleniaceae,
two new species from Brazil were discovered: Do¬
liocarpus lombardii from the “Mata Atlantica’’ and
Tetracera boomii from the “Mata de Restringa”
and “Cerrado” vegetation dominated by Curatella
americana L. The two new species are described,
illustrated, and compared to closely related spe¬
cies. By its sessile inflorescence Doliocarpus lom¬
bardii is closely related to D. sessiliflorus Martius,
but differs from that species in its longer leaf
blades (9-22 cm long), densely pubescent on the
lower surface; sepals 5, 6—1 1 mm long, ovate or
elliptic, adpressed sericeous in midrib internally;
and fruit densely hispid. Tetracera boomii is
known only from two places in Bahia and Sergipe
states, Brazil, and is similar to T. lasiocarpa Ei-
chler in that both bear follicles 1—1.5 cm long,
densely hispid at the apex; however, T. boomii dif¬
fers from that species in its leaves rigid-coria¬
ceous, elliptic, aerolate below and margins den¬
tate, inflorescences 1-2 cm long, bracteoles ca. 6
mm long, sepals 11—14, filaments 4-5 mm long,
and aril completely covering the seed.
Key words: Brazil, Dilleniaceae, Doliocarpus,
Tetracera.
Species of Doliocarpus Rolander are mostly lia¬
nas (rarely shrubs), with inflorescences that are
ramiflorous, fasciculate, or glomerate, the ovary un-
icarpellate, 1-celled, the fruit a berry, and seeds
covered with white arils. Tetracera L. is the only
genus in the Dilleniaceae with either unisexual or
bisexual flowers; all Neotropical species are andro-
dioecious (Dickinson, 1968; Kubitzki, 1970). Do¬
liocarpus includes about 45 species and Tetracera
ca. 20; Brazil is the center of diversity for both
genera.
Tetracera boomii G. Aymard C., sp. nov. TYPE:
Brazil. Bahia: Entre Rios, 23 km from Subau-
ma on road to Entre Rios, 29 May 1981, S. A.
Mori & B. M. Boom 14196 (holotype, CEPEC
not seen; isotype, NY). Eigure 1.
Haec species T. lasiocarpae affinis, sed ab ea foliis rig-
ido-coriaceis ellipticis subtus areolatis margine dentatis,
inflorescentia 1-2 cm longa dense adpresso-pubescente;
sepalis 11 ad 14, petalis 4 intus dense sericeis, lilamentis
4—5 mm longis atque arillo semen omnino tegente.
Liana, bark brown, flaking; branches and
branchlets sparsely adpressed pubescent to gla-
brescent when mature. Leaves elliptic, rigid-coria¬
ceous, 4—22 X 3—13 cm, base cuneate, apex rotun-
date, scabrous and sparsely pilose on the upper
surface, papillate and glabrescent on the lower sur¬
face, except for the midrib and secondary nerves,
these sparsely adpressed pubescent, with stellate
and simple trichomes, margins dentate mostly in
the upper half, the veins elevated, areolate below,
sunken above, with 6 to 15 parallel nerves on each
side of mid-nerve, craspedodromous venation; pet¬
ioles 0.5-2.2 cm long. 1-4 mm wide, subwinged,
canaliculate, adpressed pubescent. Inflorescence
thyrsoid, axillary or terminal, 1—2 cm long, rachis
densely adpressed pubescent, bracteoles lanceo¬
late, ca. 6 mm long, sparsely adpressed pubescent
externally, sericeous internally mostly in the mid¬
dle, pedicel 0.5—5 mm long, 1 to 3 flowers per
Novon 13: 1-4. 2003.
2
Novon
lcm
Figure 1. Tetracera boomii G. Aymard C. —A. Branch with leaves and inflorescences. —B. Upper leaf surface. —C.
Upper part of the inflorescence. —D. Lower leaf surface. —E. Cross section of stem with vascular tissue arranged in
bands separated by abundant parenchyma. (Drawn from the isotype, Mori & Boom 14196, NY.)
cyme. Flowers androdioecious, bisexual; sepals 11-
14, unequal, the outer 4—10 mm long, broadly
ovate, the inner 10—12 mm long, broadly ovate,
sparsely adpressed sericeous externally, densely se¬
riceous internally, except at the margins; petals 4,
obovate to oblanceolate, 8—12 mm long, glabrous
externally, glabrous internally except in the mid¬
point, which is densely adpressed sericeous. Sta¬
mens 100—120, filaments 4—5 mm long, glabrous,
anthers oblong, ca. 0.75 mm long; staminate flowers
not seen. Caqiels 7 to 8, ca. 5 mm long, free,
densely hispid sericeous, style ca. 2 mm long, gla¬
brous. Follicles 1 — 1.5 cm long, divergent, densely
hispid at the apex, with large yellow trichomes;
Volume 13, Number 1
2003
Aymard
Doliocarpus and Tetracera
3
Table 1. Comparison of the distinguishing characters of Tetracera boomii and T. lasiocarpa.
Tetracera boomii
Tetracera lasiocarpa
Leaves
rigid-coriaceous, elliptic, areolate below; mar¬
gins dentate
subcoriaceous, obovate or elliptic, without are-
oles; margins mucronate-dentate
Inflorescence
1—2 cm long, densely adpressed pubescent;
bracteoles ca. 6 mm long
5-10 cm long, slrigose pubescent; bracteoles
2—4 mm long
Sepals
11-14
7-9
Petals
4, densely sericeous at midpoint internally
5, glabrous internally
Filaments
4—5 mm long
ca. 3 mm long
Aril
completely covering the seed
covering half the seed
style persistent, 1—2 mm long; seeds 1 per carpel,
4—5 mm long, black, aril laciniate, completely cov¬
ering the seed.
Distribution and habitat. Tetracera boomii is
known only from two collections, in “Mata da Res-
tringa” and “Cerrado” areas dominated by Cura-
tella americana in the Brazilian states of Bahia and
Sergipe.
Because of its scabrous leaves and many-flowered
cymes, Tetracera boomii belongs to the section Te¬
tracera (Kubitzki, 1970). Based on the size of the
follicles (1—1.5 cm long), which are densely hispid
at the apex with large yellow trichomes, this new
species is related to T. lasiocarpa from Bahia, Brazil.
However, T. boomii can be distinguished from T. la¬
siocarpa by the characters indicated in Table 1.
This species is named in honor of Brian M.
Boom, whose collections and fieldwork have added
much to our knowledge of the Bahian flora.
Paratypes. BRAZIL. Sergipe: Santa Luzia do Itanhi,
500 m S of Distrito de Castro, 29 Nov. 1993, S. C. de
Sant'Ana et al. 480 (CEPEC, NY).
Doliocarpus lombardii G. Aymard C., sp. nov.
TYPE: Brazil. Minas Gerais: Parque Estadual
do Rio Doce, Marlieria-Salao Dourado, 15 June
1995, ./. A. Lombardi ).
A New Alpine Species of Rhododendron (Ericaceae) from
New Guinea
Gillian A. Brown
Australian National Herbarium, Centre for Plant Biodiversity Research, CSIRO Plant
Industry, GPO Box 1600, Canberra, ACT. 2601, Australia, and School of Bot any. The
University of Melbourne, Victoria, 3010, Australia. GiIlian.Brown@csiro.au
Lyn A. Craven
Australian National Herbarium, Centre for Plant Biodiversity Research, CSIRO Plant
Industry, GPO Box 1600, Canberra, AC I, 2601, Australia. Lyn.Craven@csiro.au
ABSTRACT. A new species of Rhododendron sect.
Vireya from Mount Jaya, West Papua (Irian Jaya).
Indonesia, is described and illustrated. The new
species. Rhododendron xenium Gillian Brown &
Craven, is related to R. womersleyi and R. rubinei-
florum, of subsection Euvireya, based on scale and
corolla features and leaf size. An identification key
to R. xenium and related species in subsection Eu¬
vireya is provided.
Key words: Ericaceae, New Guinea, Rhododen¬
dron.
During the Carstensz Glaciers Expedition of
1971-1972 to New ( Hiinea. the expedition biolo¬
gist, G. S. Hope, made two collections of Rhodo¬
dendron that subsequently were identified as R. an¬
agalliflorum Wernham (L. Craven, in herb, sched.
1972, and H. Sleumer in herb, sched. 1973). Fol¬
lowing the realization by Craven in 1979 that the
name R. anagalliflorum was commonly being mis¬
applied to an undescribed species, subsequently
described as R. rubineifiorum Craven (Craven,
1980), his investigation of the Mount Java (Car¬
stensz Mountains) specimens led to the conclusion
that these probably represented a third species.
The Mount Jaya specimens key out between Rho¬
dodendron womersleyi Sleumer and R. anagalliflo¬
rum in Sleumers (1966: 568) classic treatment of
section Vireya for Malesia. Differences between R.
anagalliflorum and R. rubineifiorum are given in
Craven (1980); in Sleumers treatment specimens of
the latter species key directly to R. anagalliflorum.
Closer examinations of the two Mount Jaya speci¬
mens, and also of specimens of related species,
have now confirmed that the former specimens rep¬
resent a new species of the genus, described below
as Rhododendron xenium Gillian Brown A Craven.
In the account of the Carstensz Glaciers Expedition
Novon 13: 26-29. 2003.
(Hope, 1976), the name R. anagalliflorum refers to
the present new species.
Rhododendron xenium is placed in section Vireya
subsect. Euvireya ser. Linnaeoidea Sleumer based
on scale and corolla features and leaf size. It has
the typical Euvireya scales, sessile, moderately
lobed with a small variously colored center, and a
corolla that is not salver-shaped; it is placed in
series Linnaeoidea because of its small leaves, less
than 1 cm by 0.6 cm (Sleumer, 1966). So far this
species has only been recorded from Mount Jaya in
the Indonesian province of West Papua (Irian Java)
on the island of New Guinea. It appears to be most
closely related to R. womersleyi and R. rubineifio¬
rum: however, it differs from these two in numerous
characters, outlined m Iable 1. The most obvious
differing characters are the length of the pedicel
and style at anthesis, and the arrangement of the
leaves. Other notable differences between the three
species occur in the leaf apex shape and size, co¬
rolla shape, pedicel indumentum, perulae indu¬
mentum, stamen length, and staminal filament in¬
dumentum (Table 1).
The species epithet is derived from the Greek.
xenion, gift to a guest, and is in reference to the
long pedicel that presents the flower well above the
foliage and thus facilitates feeding (and pollination)
by nectarivorous birds that are presumed to visit
the flowers of this species.
Rhododendron xenium Gillian Brown & Craven,
sp. nov. T^ PE: Indonesia. West Papua (Irian
Jaya): limestone hill at S end of Carstensz
Meadow, Mount Jaya (Carstensz Mountains.),
alt. 3540 m, 13 Dec. 1971, G. Hope, Carstensz
Glaciers Expedition. ANU 10847 (holotvpe.
CANB; isotype, BO). Figure 1.
A R. womersleyi pedicello longiore ((15.2—)]0.5—31
Volume 13, Number 1
2003
Brown & Craven
Rhododendron xenium from New Guinea
27
Table 1. Species comparison table. Characters that
rubineiflorum are shown and described.
differ among Rhododendron
xenium, R. womersleyi, and R.
Character
R. xenium
R. womersleyi
R. rubineiflorum
Leaf arrangement
scattered throughout a sea¬
sonal growth unit, not
pseudowhorled
rather densely set along
the distal part of the
branchlet
in 3—5-merous pseudo¬
whorls, rarely opposite
Leaf apex
short-acuminate
short-acuminate, subacute
acute to subacuminate
Leaf blade (mm)
(2-)3-8 X 2-4.5
5—8(-10, rarely 15) X 4—
6(—7, rarely 10)
4.2-10 X 2-5
Pedicel
—indumentum
scales and hairs
scales and hairs
scales only
—length at anthesis
(15.2-) 19.5-31
8— 12(— 15)
6-6.5
(mm)
Perulae
abaxial surface with scales
(very sparse)
abaxial surface glabrous
abaxial surface glabrous
Calyx indumentum
scales and hairs
scales and hairs
scales
Calvx lobe lengths
0.7-1.5
0.6-1.3
1.3-1.5
(mm)
Corolla curvature
slightly curved
slightly curved
straight
Corolla shape
campanulate
tubular
campanulate, broadly fun¬
nel-shaped campanulate
Corolla length (mm)
(15-) 17-20
21-25.7
13-24
Corolla tube length
(9-) 11-13.5
15.3-19.5
10-14
(mm)
Corolla lobes
circular, circular-obovate
obovate-circular
obovate-wide oblong
Stamens
slightly longer than corolla
tube
slightly longer than corolla
tube
shorter than the corolla
tube
Li laments
± hairy at the very base,
glabrous above
glabrous, sometimes hairy
in the proximal region
glabrous
Ovary dimensions
(2.7-)3-4 X ca. 2
4 X 1.5
2.2-3.5 X 1.5
(mm)
Style length at anthe-
4.3-84
10.5-12
1.3-3
sis (mm)
mm) et corolla longiore per anthesin, corolla campanulata
et breviore ((15-)17-20 mm), et stylo breviore (4.3-S.4
mm) et R. rubinefloro loliis dispersis et lamina apice brev-
iacuminata, perulis lepidotis persparsim, pedicello long¬
iore et lepidoto pilosoque, corolla parum curvata, filamen-
tis staminium pilosis, et stylo longiore differt.
Terrestrial shrub to ea. 20 cm tall. Branchlets
moderately to densely lepidote (the scales in part
persistent), terete to subterete. Leaves dispersed
throughout a seasonal growth unit, not pseudo-
whorled; leaf blade ovate to ovate-elliptic (2—)3-8
X 2—4.5 mm, the apex shortly acuminate, the base
broadly cuneate, the margin entire, slightly revo¬
lute, the midrib impressed above in lower 1/2—2/3,
inconspicuous to just visible at the base of the up¬
per surface, the veins inconspicuous, dark green
above with the margin becoming brown, light brown
beneath, initially lepidote on both surfaces becom¬
ing glabrous above with age (with visible impres¬
sions where the scales once were), persistently sub-
densely to laxly lepidote beneath (the scales brown,
irregularly lobed and sessile); petiole 1—1.6 X 0.5
mm, flattened, lepidote on both sutfaces. FloraI bud
unopened 4.5-6.5 X 2.5 mm; outer perulae ovate,
acuminate at the apex, the abaxial surface very
sparselv lepidote, the scales on the margin obscure¬
ly stalked; inner perulae the same as the outer per¬
ulae but narrower and shortly acuminate. Pedicels
densely lepidote and hairy, (15.2—) 19.5—31 X 0.6—
0.9 mm at anthesis, 34-41 mm long in fruit. Flow¬
ers solitary, held erect, the perulae sometimes per¬
sistent. Calyx 5-lobed, ca. 2 mm diant.. obtuse to
rounded-obtuse, lepidote and hairy; lobes 0.7-1.5
X ca. 0.8 mm, the lobe margin with stalked scales.
Corolla slightly curved campanulate, (15—) 1 <-20 X
13-15.5 mm, bright red to red, waxy, moderately
lepidote outside (extending over the entire tube and
onto the lower middle part of the lobes), glabrous
inside; tube (9—)11—13.5 X 3.5-5 mm (wide at the
base) and 5.3-8 mm (wide at the throat); lobes cir¬
cular to eircular-obovate, 5.5—8.5 X 6.5—8.5 mm,
overlapping at the base, rounded or retuse at the
apex. Stamens 10, appearing unequal in length, ex-
28
Novon
D
5 mm
5 mm
5 mm
Figure I. Rhododendron xenium Gillian Brown & Craven. —A. Habit, flower orientation, and fruit dehiscence. —B.
Inner perula (detail of the margin enlarged; stalks of scales not apparent). —C. Stamen. —1). Gynoecium. Drawn from
G. Hope ANU 10847 (CANB).
serted slightly from the corolla tube, stamens ap¬
pear to be arranged equally around the mouth; fil¬
aments linear, 10—11 mm long, glabrous except for
a few hairs at the very base; anthers broadly oblong,
1.2-1.6 X ca. 1 mm. Disc 10-lobed, ca. 0.6 mm
high, 2—2.5 mm diam., with dense erect white hairs
at the apex. Ovary subcylindrieal, (2.7-)3^t X ca.
2 mm, broadly tapering to the style, ribbed, densely
covered in short white hairs over a subdense to lax
layer of scales; style 4.3—8.4 X ca. 0.7 mm, hairy
in the proximal 1/4—1/2, glabrous for the remainder,
slightly shorter than the corolla tube; stigma capi¬
tate, 5-lobed. Capsule fusiform; immature fruits 14—
14.5 X ca. 3.5 mm, purplish when dry, both hairy
and lepidote (less dense than on the ovary); mature
fruits ca. 20 mm long (excluding the style), brown,
with no visible indumentum, the style persistent,
when dehiscing the valves peeling back with only
slight twisting at the base. Seeds not seen.
N.B. Floral measurements were determined by
the method given in Argent et al. (1988).
Distribution and habitat. Known only from
Carstensz Meadow at Mount Jaya in West Papua
(Irian Jaya), New Guinea. Recorded as growing in
crevices in peat in very griked limestone slopes
[trenched limestone slopes due to solution along a
Volume 13, Number 1
2003
Brown & Craven
Rhododendron xenium from New Guinea
29
joint], and on rocky slopes on the forest margin.
Altitude 3520-3540 m.
Paratype. INDONESIA. West Papua (Irian Java):
limestone hill at S end of Carstensz Meadow, Mount Jaya
(Carstensz Mountains.), alt. 3520 m, 13 Dec. 1971, C.
Hope, C.G.E., ANll 10840A (CANB).
Key to Rhododendron xenium and Related Species
Couplet 2 of subsection Euvireya ser. Linnaeoidea in
Sleumer (1966: 568) can be replaced with the following
couplets that will permit specimens of Rhododendron an -
agalliflorum, R. rubineiflorum, R. womersleyi, and R. xen¬
ium to be identified.
2. Style at anthesis relatively short, i.e., approxi¬
mately as long as the ovary or shorter. Calyx ex¬
clusively lepidote. Corolla straight, never curved.
2a. Corolla funnel-shaped with elliptic- lobes,
white, flushed with purple from the base.
Anthers 0.6 mm long. R. anagalliflorum
2a. Corolla campanulate to broadly funnel-
shaped campanulate with orbicular lobes,
red to pink. Anthers 1.2-1.5 mm long . . .
. R. rubineiflorum
2. Style at anthesis at least two times as long as the
ovary. Calyx both lepidote and hairy. Corolla
slightly curved.
2b. Pedicel shorter than the total length of the
corolla at anthesis. Corolla tubular. Abaxial
surface of perulae glabrous .... R. womersleyi
2b. Pedicel longer than the total length of the
corolla at anthesis. Corolla campanulate.
Abaxial surface of perulae ± sparsely lepi¬
dote . /?. xenium
Acknowledgments. The curator of CANB is
thanked for the opportunity to study specimens de¬
posited therein. The first author is the recipient of
a Melbourne University Research Scholarship. The
Baker Foundation is thanked for its financial sup¬
port of the research project of which the present
contribution forms a part. Sharyn Wragg is thanked
for preparing the illustration, and the interest of
John L. Rouse in our research is gratefully ac¬
knowledged.
Literature Cited
Argent, G., A. Lamb, A. Phillipps & S. Collenette. 1988.
Rhododendrons of Sabah. Sabah Parks Publication no.
8. Sabah Parks Trustees, Kota Kinabalu.
Craven, L. A. 1980. A new Rhododendron (Ericaceae)
from New Guinea. Notes Roy. Rot. Gard. Edinburgh 38:
141-144.
Hope, G. S. 1976. Vegetation. Pp. 113-172 in G. S. Hope,
J. A. Peterson, U. Radok & 1. Allison (editors). The
Equatorial Glaciers of New Guinea: Results of the
1971—1973 Australian Universities Expedition to Irian
Jaya: Survey, Glaciology, Meteorology, Biology and Pa-
laeoenvironments. Balkema, Rotterdam.
Sleumer, H. 1966. Rhododendron. Pp. 469—668 in C. G.
G. J. van Steenis (editor). Flora Malesiana ser. I, Vol.
6. Wolters-Noordhoff, Groningen.
A New Species of Casearia (A lacourtiaceae) from Mexico
Gonzalo Castillo-Ca mpos
Departamento de Sistematica Vegetal. Institute de Ecologfa, A.C., km 2.5 Carretera Antigua
a Coatepec No. 351. Congregacion El Haya, 91070, Xalapa, Veracruz, Mexico. Author for
correspondence: castillo@ecologia.edu.inx
Ma. Elena Medina Abreo
Departamento de Diagnostico Regional, Instituto de Ecologfa, A.C., km 2.5 Carretera
Antigua a Coatepec No. 351, Congregacion El Haya, 91070, Xalapa, Veracruz, Mexico.
inedi nama@ecologia.edu. mx
Abstract. Casearia guevarana, a new species of
Flacourtiaceae from the coastal dunes of the Nat¬
ural Reserve "El Mono de la Mancha,” from the
states of Veracruz and Tabasco, Mexico, is de¬
scribed and illustrated. Its affinities with the closely
related species C. aculeata Jacquin and C. obovata
Schlechtendal are discussed. This new species dif¬
fers from its congeners by having broadly obtuse
leal apices, large, six-channeled fruits, and larger,
more numerous seeds.
Resumen. Se describe como nueva a Casearia
guevarana (Flacourtiaceae) de las dunas costeras
de la Reserva Natural FI Morro de La Mancha,
Veracruz y Tabasco, Mexico. Esta nueva especie
esta cercanamente relacionada con Casearia acu¬
leata Jacquin y C. obovata Schlechtendal, de las
que se distingue por presentar el apice de las hojas
ampliamente obtuso, frutos 6-acanalados y de may¬
or tamano, con semillas mas grandes y inas nu-
merosas.
Key words: Casearia, coastal, dunes, Flacour¬
tiaceae, La Mancha, Mexico, Tabasco, Veracruz.
In 1998, during an inventory of trees and shrubs
associated with an evergreen seasonal rainforest at
the Nature Reserve and Center for Coastal Re¬
search at La Mancha (C1C0LMA) in Veracruz,
Mexico, a tree rarely seen in the middle stratum of
the forest was sampled and reported. At the time,
this tree was totally lacking reproductive structures,
so that it was only possible to classify it with any
certainty to the genus Casearia (Flacourtiaceae).
However, during two years subsequent to this initial
find, flowers (2000) and fruits (2001) were obtained.
After having examined and compared vouchers of
C. aculeata and C. obovata with those recently col¬
lected from the Natural Reserve of La Mancha, it
was found that they corresponded to an unde¬
scribed species. Based upon this additional infor¬
mation, this specimen was considered to represent
a new species. Its detailed description is reported
below.
Casearia Jacquin is a pantropical genus of about
180 species of trees and shrubs. It is most diverse
in (sub)tropical Central and South America, where
some 75 species occur, including 8 in Veracruz.
I lie genus is subdivided into six sections, the larg¬
est, and only one to occur outside of the Neotropics,
is section Casearia. Three of the sections are rep¬
resented in Veracruz: sect. Crateria, sect. Piparea,
and sect. Casearia. Section Casearia is further di¬
vided into six informal groups, three of which occur
in Veracruz, i.e. Nitidae, Aculeatae, and Decan-
drae.
The newly described Casearia guevarana has an
undivided style, free stamens, and free interstam-
inal disc lobes. These characters place the species
in Casearia sect. Casearia. The following charac¬
ters place the species in section Casearia, group
Aculeatae; an undivided style, at times obsolete,
and capitate stigma; plants generally with spines,
at times not present in herbarium vouchers; and
stamens generally 8, flowers more or less tubular,
base obtuse-truncate often slightly swollen just be¬
fore flowering. Two other species of the Aculeatae
group occur in Veracruz, C. aculeata and C. obov¬
ata.
Casearia guevarana Castillo-Campos & Medina,
sp. nov. TYPE: Mexico. Veracruz: Nature Re¬
serve “LI Morro de la Mancha,” Mun. Ado-
pan, approx. 10 m. 24 May 2001, C. Castillo
C. 20801 (holotype, XAL; isotypes, ENCB,
MEXU). Figure I.
Arbor 3-6 ni alta erect a spinis rectis munita vel mer-
mis; foliis oppositis vel altemis, ellipticis vel ovatis, pe-
No von 13: 30-33. 2003.
Volume 13, Number 1
2003
Castillo-Campos & Medina
Casearia guevarana from Mexico
31
Figure I. Casearia guevarana Caslillo-C. & Medina. —a. Branch with infruetescence. —b. Inflorescence in bud.
—c. Detail of the inflorescence with bracts. —d. Detad of an open flower in natural state. e. Immature fruit. f.
Detail of fmit apex. —g. Mature fruit. —h. Open fruit. —i. Valve. —j. Seed. (Based on the paratype Castillo 19902.)
32
Novon
tiolatis, 2.0—8.8 cm longis, 1.4—4.2 cm latis. Flores albi
perfecti pedicellati, pedicellis ad dimidium articulatis; se-
palis 5 ovatis, 5-6 mm longis, 2.2-2.5 mm latis; stami-
nibus 8. Fructus capsularis subglobosus, juventute 3-cos¬
tatus, maturitate 6-canaliculatus.
Perennial tree or shrub; 3-6 rn high; trunk about
20 cm diam.; bark smooth; older branches brown;
prickles brown to ash-gray, darker at prickle apex;
in young branches, frequently ending in a prickle,
or prickle-less, pubescent, lentieellate; prickle
straight (when present), 0.7-1.7 cm long, 1-2.5 mm
wide at base, basal half lentieellate, apex acute.
Leaves alternate, distichous, simple, petiolate, ob-
ovate-elliptic, 2—8.8 cm long, 1.4—4.2 cm wide, co¬
riaceous; base attenuate, euneate, apex broadly ob¬
tuse, frequently shallow-emarginate; margin
crenate with 3 to 10 teeth per side, upper and lower
surface lustrous with a few scattered hyaline hairs
on the underside, glabrescent; venation brochidod-
romous, central vein pubescent on both surfaces,
more densely so on the upper surface, lateral veins
3 to 7, tertiary venation reticulate. Petiole 2-6 X
0.5—0.8 mm, pubescent on upper and underside.
Flowers bisexual, 1 to 11 (frequently 3) in axillary
peduneled fascicles, white. Bracts ovate, 0.5-1.1 X
0.5—1.0 mm, pubescent, chartaceous, with acute
apex and entire margin, yellowish when young,
turning to light green and then light brown when
mature. Pedicels 5-11 X 0.5 mm. articulate near
middle, glabrescent. Sepals 5, imbricate, 5-6 X
2.2-2.5 mm, spreading, ovate, with obtuse apex
and entire margin, outside glabrescent. inside gla¬
brous, deciduous although frequently persistent in
immature fruits. Stamens 8, filaments 3.5 mm long,
flattened, glabrous; anthers 0.5 X 0.5 mm. Disc
lobes alternate with stamens, villous. Ovary ovoid,
villous; style undivided; stigma capitate, 0.6-0.8
mm long. Fruit capsular with septicidal dehiscence,
subglobose, 3-ribbed when immature, 6-channeled
when mature (3 prominent channels, 3 shallow), 9-
14 X 7-10 mm, green when young and yellowish
when mature, purple at base and along the more
prominent channels, splitting into 3 valves, re-
trorse, 5—10 X 5—7 mm; endocarp red at maturity,
the inner surface of each valve with 1-7 small
white spots. Seeds 16—21, yellow to orange at ma¬
turity, brilliant, usually ovate, angular, 3-6 X 2-
4.5 mm, base truncate; seed coat reticulate, com¬
pletely covered by aril, aril mucilaginous, orange
with small white spots.
Ph enology. Casearia guevarana flowers anti
fruits between May and November, and its flowering
and fruiting periods are brief. Flowers and fruits
are fragile, detaching easily as branches are moved
by the wind. Young fruits are ribbed, but at matu¬
rity these are lost anti substituted by channels or
longitudinal depressions. These characters are lost
in dried specimens because well-developed fruits
open during the drying process.
Habitat. Casearia guevarana is found in ever¬
green seasonal rainforest along the coastal dunes of
El Morro tie la Mancha in central Veracruz, Mexico.
A rare tree forming part of the middle stratum of
the tropical deciduous forest, it is associated with
Brosimum alicastrum Swartz, Bursera simaruba (L.)
Sargent, Cedrela odorata L., Ehretia tinifolia L.,
Enterolobium cyclocarpum (Jacquin) Grisebach, Ex¬
ostem a mexicanum A. Gray, Ficus cotinifolia
Knntli, Ginoria nudiflora (Hemsley) Koehne, Gyr-
ocarpus jatrophifolius Domin, Tabebuia rosea (Ber-
toloni) DC., and Nectandra salicifolia (Kunth) Nees.
Distribution. Casearia guevarana has a limited
distribution, being found in just two localities: one
in the state of Veracruz and the other in the state
of Tabasco. Both sites occur on the coastal plain of
the Gulf of Mexico in evergreen seasonal rainforest
at above sea level.
Etymology. The specific epithet alludes to Ser¬
gio Guevara Sada, paying particular homage to his
research and enthusiasm in consolidating and pro¬
moting the Nature Reserve and Center for Coastal
Research at La Mancha (CICOLMA), Veracruz,
Mexico.
Discussion. Casearia guevarana is closely re¬
lated to C. aculeata and C. obovata, with which it
shares certain morphological characteristics, espe¬
cially the presence of prickles, leaf form, petiole
length, pedicel articulation, number of stamens, pu¬
bescence of disc lobules, and ovary and (lower col¬
or. Casearia guevarana is distinguished from C.
aculeata and C. obovata by its glabrous, lustrous
leaves; broadly obtuse leaf apices; larger, six-chan¬
neled fruit; and larger, more numerous seeds (Table
1). Casearia aculeata and C. obovata are similar,
differing only in leaf form, apex, and pubescence
(Nee, 1999). Casearia obovata and C. guevarana
are endemic to Mexico, whereas C. aculeata has a
much wider distribution, from Sinaloa and Tamau-
lipas in northern Mexico to South America and the
Antilles (Sleumer, 1980; Nee, 1999).
Paratypes. MEXICO. Tabasco: km 21 along the high¬
way that runs from Parafso to La Barra de Tupilco, Mun.
Parafso, 47. A. Magana A. 482 (XAL); km 1.7 along the
highway from Nicolas Bravo-Mecoacan to Aqufles Serdan,
Man. Parafso, (xnvan .3205 (FNCB). Veracruz: nature re¬
serve at El Morro de La Mancha, Mun. Actopan, G. Cas¬
tillo C. 19902. 19238 (ENCB. MEXU, XAL).
Acknowledgments. Our sincerest thanks are ex¬
tended to Jerzy Rzedowski for the Latin description
Volume 13, Number 1
2003
Castillo-Campos & Medina
Casearia guevarana from Mexico
33
Table 1. Similarities and differences between Casearia guevarana , C. aculeata, and C. obovata (Sleumer, 1980;
Nee, 1999).
Casearia guevarana
Casearia aculeata
Casearia obovata
Leaf
Shape
obovate, elliptic
oblong, ovate-oblong, obovate obovate
Pubescence
glabrous, except on the central vein
pubescent to glabrous
pubescent
Length (cm)
2.0-8.8
5-8
3.5-9.0
Width (cm)
1.4-4.2
3-4
2-5
Apex
broadly obtuse, frequently emarginate
abruptly attenuate
abruptly acuminate
Base
attenuate, cuneate
acute to obtuse
acute to obtuse
Petiole
Length (mm)
2-6
2-6
3-10
Flower
Number
1 to 11, frequently 3
5 to 10
few
Color
white
yellow-green, white
green-white
Stamens
8
8
8
Filament
glabrous
glabrous
glabrous
Disc lohed
villous
villous
villous
Ovary
villous
villous
pubescent
Pedicel
Length (mm)
5-11
4-6
3-8
Articulation
present
present
present
Fruit
Shape
subglobose, 6-channeled
subglobose, obtusely triangu- subglobose
l.,.-
Length (mm)
9-14
5-6
6-8
W idth (mm)
7-10
5-8
6-8
Seed
Number
16-21
3-10
few
Length (mm)
3-6
4
2.5
Shape
ovoid to angular
ovoid to angular
ovoid to angular
of Casearia guevarana and for his comments on the
manuscript. Thanks are also given to Francisco Ko¬
rea H. and Andres F. Vovides for their sound sug¬
gestions on tiie manuscript, Edmundo Saavedra for
preparing the illustration, to the herbaria MEXU
and ENCB for loan of specimens used in this re¬
search, and to Dan Bennack for translating the orig¬
inal article from Spanish to English. This research
was funded by CONABIO (L-228) and Institute de
Ecologfa, A. C. (904—14).
Literature Cited
Nee, M. 1999. Flacourtiaceae. In: Flora de Veracruz. Fasc.
Ill: 1-79.
Sleumer, 0. H. 1980. Flacourtiaceae. FI. Neotrop. Monogr.
22: 1-499.
A New Species of Tovomita (Clusiaceae) from Amazonian
Venezuela and Peru
Nidia L. Cuello A.
Herbario Universitario (PORT), Programs R.N.R., UNELLEZ-Guanare, Mesa us
(non elevatis), pedicellis 9-1 l(—20) mm longis, gemmis flor-
alibus masculis mucronatis, sepalis 4 exterioribus orbicu-
latis, petalis interioribus apice breviter difatatis, staminum
filamentis crassiusculis quadratis atque fructu pyriformi in
stylum 1 mm vix attingentem desinente differt.
Presumed monoecious treelet. 3—8 m tall.
Branches terete, glabrous. Latex whitish, wood with
sweet aroma. Leaves opposite, decussate, glabrous
on both surfaces, clustered toward upper parts of
branches with internodes (3—)4—8(—11) mm long;
the petioles plane, 3—7(—12) mm long; the blades
membranous, lanceolate to elliptic-lanceolate, (3—)
4—10 X (0.5—) 1-2.5 cm, attenuate at base, acute-
acuminate to caudate at apex, the apex 5—15 mm
long, base 1.5—2 mm wide, margin thinly revolute,
main nerve elevated in upper surface, impressed
beneath, lateral veins inconspicuous in mature
leaves, slightly visible in younger leaves but not
prominent, less than 3 mm apart. Inflorescences cy-
moid, terminal. 1.5—3 cm long, usually divided into
3 branches near the base, but secondary flowers of
lateral branches caducous, leaving each branch 1-
(lowered and appearing as a simple dichasium. pe¬
duncle 1-6 mm long, terminal internode 6-20 mm
long, internodes of lateral branches 3.5—20 mm
long; pedicels slender, 9—1 1 (—20) mm long in [tri-
mar)' flowers. Bracts caducous, not seen. Flower
buds ovoid, 4-5 X 2.5—3 mm, the staminate buds
mucronulate, apex ca. 0.5—1 mm long, the pistillate
buds slightly shorter and broader. Sepals 4, the 2
external sepals green, coriaceous, fused up to the
No VON 13: 34-36. 2003.
Volume 13, Number 1
2003
Cuello
Tovomita stergiosii (Clusiaceae)
35
Figure 1. Tovomita stergiosii Cuello. Habit with details of flowers and leaves. —A. Habit and staminate flowering
branch. —B. Staminate flower bud. —C. External sepals. —D. Internal sepals. — E. External petals. —E. Internal
petals. —G. Stamens. —H. Pistillode. —I. Pistillate flower bud. —.). External sepals. —K. Internal sepals. —L.
External petals. —M. Internal petals. —N. Staminodes. —O. Pistil. —P. Branch bearing young fruits. —Q. Detail of
leaf margin from abaxial view. —B-V. Leaf shapes and sizes. (A based on Colella et al. 2016, PORT; B—H and V
based on Stergios et al. 13161. PORT; l-R based on Rimachi 7616. MO; S based on McDaniel & Rimachi 20466. MO;
T based on Aymard & Delgado 8423, POR I; and U based on Castillo & Camaripano 8362. PORT.)
36
Novon
middle, 4—5 X 3—4 mm, broadly oblong to orbic¬
ular, cymbiform, truncated at base, tbe outer of the
two mucronulate in staminate flowers; tbe 2 internal
sepals petaloid, imbricate, broadly oblong, 3—5 X
2-3 mm, the inner one bonded, narrower than tbe
outer, apiculate in staminate flower, apex ca. I mm
long. Petals 4, varied in shape, free and distinct in
bud, white, 3—5 X 1.5—2 mm, decussate, the 2 ex¬
ternal petals oblong, rounded to truncate or sinu-
olate at apex, the 2 internal petals contracted at
1/3 the distance from the apex anti shortly widened
at apex, the claw 2.5—3 mm long, 1.5—2 mm wide,
the blade 1—1.5 mm long, 3—4 mm wide. Stamens
ca. 20, filaments moderately thick, tetragonous, ca.
0.5 mm witle, 2.5—3.5 mm long, thecae obliquely
attached, 0.5—1 mm long. Staminodes ca. 20, 3.5—
4 mm long. Pistil in female buds 2—2.5 mm long,
shortly urceolate, wider and slightly angulate at
base, ovary 4-celled, styles 4, distinct, 0.5—0.6 mm
long, stigmas cordate to rounded, 0.5—0.7 mm
diam. Pistillode in staminate buds very small, ca.
0.5 mm long. Fruit a fleshy septicidal capsule, pyr¬
iform when still closed, 1.7—2.2 X 0.7—1.4 cm,
green externally, pink within, estipitate, with 4
short, persistent, crowned styles 0.8—1 mm long,
stigmas rounded in fruit, 0.5—0.7 mm diam. Seeds
reniform, arillate, 7.5—9 mm long, 4—5 mm wide.
Distribution and ecology. Tovomita stergiosii is
a small tree known from shrubby white-sand sa¬
vannas and edges of inundated lowland forest along
blackwater rivers, between 90 and 250 m elevation
in Venezuelan and Peruvian Amazonia. The new'
species is known in Peruvian highland Quechua as
“chullachaqui caspi” (. Rimachi 7615, MO).
Tovomita stergiosii is most similar to T. brasilien-
sis (Martius) Walpers from Brazil, French Guiana,
and Colombia (Martius, 1826; Walpers, 1842; En-
gler, 1858) with regard to habit and leaf features.
Both are small trees with terete branches, and both
have small, membranous leaves on short inter¬
nodes. However, T. stergiosii differs in having lan¬
ceolate leaves rather than oblong-elliptic leaves,
with an attenuate rather than acute base and an
acute-acuminate to caudate rather than obtuse
apex. Additionally, in T. stergiosii the main leaf
nerve is impressed and not prominent, and lateral
nerves are slightly visible on the undersurface. In
contrast, T. brasiliensis has leaves with a prominent
main nerve and its lateral nerves are slightly raised
beneath. Furthermore, the flower buds in the new'
species are mucronulate in staminate flowers, with
four sepals, the outer sepal orbicular-mucronulate,
the two inner petals shortly widened at the apex.
In T. brasiliensis, the flower buds are rounded at
the apex, not mucronulate, with only two ovate-ob¬
tuse sepals, and petals obovate-oblong and obtuse.
Also, tbe new species differs by its longer pedicels,
9-11 (—20) mm long, less numerous stamens (ca.
20) with moderately thick and quadrangular fila¬
ments, and a pyriform fruit with short styles ca. 1
mm long. Tovomita brasiliensis has shorter pedicels,
6—8 mm long, ca. 50 stamens with flattened, filiform
filaments, and an obovate fruit with longer, persis¬
tent styles up to 3 mm long.
Among specimens examined of Tovomita stergio¬
sii, a variation in the sizes and shapes of leaves was
noted, especially in the shape of the leaf apices,
which can vary from acute to acuminate in Vene¬
zuelan and Peruvian specimens and even to cau¬
date in some of the Peruvian specimens.
This species is named in honor of Basil Stergios
(1940-) from PORT, in recognition of bis enthusi¬
asm for botanical exploration in Venezuela and for
collecting the type specimen.
Paratypes. VENEZUELA. Territorio Federal Ama¬
zonas: Atabapo, Alto Rib Orinoco, Cano Tama Tama. 2
km al E de Tama Tama, 3 Mar. 1990 (fr), G. Aymard A
L. Delgado 8423 (MO. PORT, YEN not seen); Casiquiare.
Rio Casiquiare, Laguna de Paciva y sus alrededores, 8
Feb. 1991 (fl, fr), M. Colella, ,/. Veldzco, E. Guayamare &
S. Camico 201 6 (NY. PORT); Rio Sipapo, cerca de Cerro
Pelota, 20 Eeb. 2001 (fr), A. Castillo & B. Camaripano
8362 (PORT, YEN not seen). PERU. Loreto: Maynas,
Iquitos, Rio Nanay, Quebrada de Momdn, un poco arriba
del caserio de San Jose, Aug. 1984 (11, fr), M. Rimachi
7616 (MO); Rio Nanay, two bends below entrance to Mapa
Cocha, 14 Jan. 1976 (11). S. McDaniel & M. Rimachi
20466 (MO); vicinity of Iquitos (collection data lost). 1977
(fl). ,/. Revilla 3610 (MO); Rio Mamdn near Rio Nanay, 1
Sep. 1972 (11), 7.’ Croat 19887 (MO).
Acknowledgments. 1 thank the Missouri Botan¬
ical Garden for making its facilities available for
research. I thank Gerardo Aymard (PORT), Basil
Stergios (PORT), Barrv Hammel (MO), and an
anonymous reviewer for their comments and sug¬
gestions on the manuscript. I thank Bruno Manara
(VEN) and Roy Gereau (MO) for their expertise
with the Latin diagnosis, and Angelina Licata
(PORT) for preparing the illustration.
Literature Cited
Cuello, N. 1998. Tovomita (Clusiaceae). Pp. 313—322 in
P. E. Berry, B. K. Holst & K. Yatskievyeh (editors).
Flora ol the Venezuelan Guayana, Vol. 4. Missouri Bo¬
tanical Garden Press, St. Louis.
Engler. A. 1858. Guttiferae. Pp. 382—474 in C. F. P. von
Martius (editor), f lora Brasiliensis Vol. XII (1).
Martius, C. F. P. 1826. Marialvaea. Nova Genera et Spe¬
cies Plantarum II: 83.
Walpers, G. 1842. Tovomita. Repert. But. Sist. 1: 392.
New and Reconsidered Mexican Acanthaceae X. Flora del
Bajio Region
Thomas F. Daniel
Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco,
California 94118, U.S.A. tdaniel@calacademy.org
Abstract. Three new species (Carlowrightia
venturae, Jnsticia zamudioi, and Tetramerium car-
ranzae) and a new combination (Mirandea hyssopus
based on Rhytiglossa hyssopus Nees) art' proposed
for the acanthaceous flora of the region in central
Mexico covered by the Flora del Bajio y de Re-
giones Adyacentes. Morphological and palynological
characters are used to distinguish these taxa from
their Mexican congeners. Studies of Acanthaceae
in this region reveal the presence of 59 species (5
endemic) in 18 genera there.
Rest MEN. Tres especies nuevas ( Carlowrightia ven¬
turae, Justicia zamudioi, y Tetramerium carranzae)
y una combinacion nueva [Mirandea hyssopus bas-
ada en Rhytiglossa hyssopus Nees) se proponen
para la flora de acantaceas de la region de Mexico
central cubierta por la Flora del Bajio y de Re¬
gia nes Adyacentes. Los earacteres morfologieos y
palinologicos se utilizan para distinguir estos taxa
de sus congeneres mexicanos. Los estudios de
Acanthaceae en esta region revelan la presencia de
59 especies (5 endemicas) en 18 generos allf.
Key words: Acanthaceae, Carlowrightia, Justi¬
cia, Mexico, Mirandea, pollen. Tetramerium.
file relatively high and flat region located in the
southern extremity of central Mexico's Altiplano,
comprising portions of the states of Guanajuato, Mi¬
choacan, and Queretaro, is commonly referred to as
“el Bajio” (Rzedowski & Calderon de Rzedowski,
1987). An amplification of this region, including all
of Guanajuato and Queretaro as well as much of
northern Michoacan, is the subject of the Flora del
Bajio y de Regiones Adyacentes, a project of the
Instituto de Ecologfa in Patzeuaro, Michoacan
(Calderon de Rzedowski & Rzedowski, 1991). This
amplified region comprises about 50,000 square ki¬
lometers, contains considerable topographic relief,
and is estimated to have more than 5500 species
of plants (Calderon de Rzedowski & Rzedowski,
1991). Portions of the amplified region, especially
in northern Guanajuato and adjacent Queretaro, re¬
mained botanically unexplored until the late 1980s
and 1990s. Extensive field activities associated
with the ongoing floristic project have resulted in
approximately 55,000 plant collections since 1985
when the project was initiated (J. Rzedowski, pers.
comm.). Studies of these collections have revealed
numerous undescribed species and distributional
range extensions in various taxa (e.g., Calderon de
Rzedowski, 1991: Calderon de Rzedowski & Rze¬
dowski. 1997; Rzedowski & Calderon de Rzedows¬
ki, 1995. 1997, 1998a, 1998b; Daniel, 1999).
In this report three new species and a new com¬
bination are proposed for the acanthaceous flora of
the amplified Bajio region. Studies of Acanthaceae
in this region to date reveal the presence of 59
species in 18 genera (Daniel, unpublished). Sixteen
of these genera are native to the region and five
species (including those treated here) appear to be
endemic there. An account of the family in the Ba¬
jio region is being prepared for the flora noted
above.
Carlowrightia venturae T. F. Daniel, sp. nov.
TYPE: Mexico. Guanajuato: Mpio. Victoria,
Cerro de la Luz, camino a Xichu, 1800 m, 26
July 1991, F. Ventura V. & E. Lopez P. 9336
(holotype, IEB; isotype, CAS). Figure 1.
Herbae perennes decumbentes usque ad f .5 dm altae.
Caules juniores bisulcati. pubescentes trichomatibus
eglandulosis. Folia petiolata, petioli usque ad 3 mm longi,
laminae lanceolato-ovatae vel ovatae, 10—22 X 2.8—8 mm,
2.1— 3.8-plo longiores quam latiores. Dichasia redacta, 1-
fiora, sessilia in axillis bractearum. Calyces 3.7—6 mm
longi, extus glandulosi. Corollae subactinomorphae, azur-
ae, 7-9.5 mm longae. Capsulae 9-10 mm longae, extus
pubescentes. Semina 3.5—3.8 X 3.3—3.5 mm, testae tub-
ereulatae ad margines dentatae.
Decumbent perennial herbs to 1.5 dm tall from
stout woody caudices. Young stems bisulcate, ± bi-
fariously pubescent with retrorse eglandular tri-
chomes 0.05-0.1 mm long, trichomes concentrated
in (or restricted to) 2 lines. Leaves petiolate, peti¬
oles to 3 mm long, blades lance-ovate to ovate, 10—
22 X 2.8—8 mm, 2.1-3.8 times longer than wide,
truncate to rounded at base, acuminate at apex,
surfaces with several orders of venation prominent.
Novon 13: 37-48. 2003.
38
Novon
Figure I. Carlowrightia venturae T. k Daniel. —a. Habit. —b. Knlargement of section of young stem. —c. Leaf.
—d. Inflorescence node with flower. —e. Distal portion of stamen. —f. Distal portion of style. —g. Capsule and calyx.
—h. Seed. —i. Enlargement ol seminal tubercles, a—c from Ventura • Myoxanthus ruschii Fraga & L. Kollmann. —A. Habit and inflorescence. —15. Flower. —C. Ovary, column,
and lip, side view in normal position. —I). Ovary and column, from below. —F. Dorsal sepal. —F. Petal, from above'
—G. Petal, from below. — H. Lateral sepal, from above. —I. Lip, from above. —J. Lip, from below. —K. Anther, from
above. L. Anther, from below. —M. Pollinia. Drawn from (he type specimen (C. N. Fraga 7HO), by C. N. Fraga.
Volume 13, Number 1
2003
Fraga & Kollman
Myoxanthus ruschii from Brazil
51
lobe elliptic, obtuse, somewhat verrucose with rev¬
olute margins, thick, the lower half with low, erect,
rounded margins, the disc with a broad, smooth,
slightlv concave area, the base subtruncate, with a
pair of minute lateral lobes reflexed on the end, 3-
veined; column 1-1.5 X 0.5 mm, yellow, flecked
with red, stout incurved, 3 mm long, with rounded
wings above the middle, the column-foot stout. 2
mm long, concave, with a pair of thick calli near
the apex, the upper margin of the anther fimbriate.
2 pollinia-pairs, yellow, obovoid. Capsule unknown.
Etymology. The name of the new species pays
homage to Augusto Ruschi, a naturalist involved in
the conservation of natural areas in Espirito Santo,
especially through the Santa Lucia Biological Sta¬
tion. which was where he started his botanical stud¬
ies. particularly with Orchidaceae.
The new species is apparently related to Myox¬
anthus punctatus (Barbosa Rodrigues) Luer, M. lon-
chophyllus (Barbosa Rodrigues) Luer. and M. sei-
delii (Pabst) Luer. from which it is distinguished by
its elliptic leaves, labellum thick and non-deflexed
with revolute margins, and fimbriate anther.
The presence of an elliptical labellum with the
terminal lobule having a verrucose margin, also ob¬
served in Myoxanthus punctatus and M. sei delii ,
distinguishes M. ruschii from M. lonchophyllus,
which has an ovate labellum, somewhat hispid-pa¬
pillose apically.
The yellow coloring with red spots, non-clavate
petals, and general elliptic form of the labellum
with subtruncate base with a pair of lateral lobes
reflexed on the end. found in Myoxanthus ruschii ,
are characters absent in M. punctatus and M. sei-
delii.
Paratype. BRAZIL. Espirito Santo: Santa Teresa, Val-
sugana Velha, Santa Lucia Biological Station, dry trail, ca.
19°57' 10”-19°59'00"S, 40°31'30"-40°32'25"W, 700 in.
Atlantic forest. 28 Apr. 2000 (fl), /.. ./. C. Kallmann 2899,
C. N. Fraga, V. G. Demuner. E. M. C. heme & II. R. Silva
(MBML).
Acknowledgments. We acknowledge Conselho
Nacional de Desenvolvimento Cientifico e Tecnol-
ogico (CNPq) for financial support, Helio de Quei-
roz Boudet Fernandes, Director of the Mello Leitao
Biological Museum and MBML herbarium curator,
for assistance during fieldwork in Santa Teresa. Jor¬
ge Fontella Pereira for the Latin diagnosis, and Fa-
bio de Barros, Marcos Sobral, and two reviewers
and the editor for suggestions and help with the
English.
Literature Cited
Luer, C. A. 1982. A reevaluation of the genus Myoxanthus
(Orchidaceae). Selbyana 7: 34—54.
-. 1992. leones Pleurothallidinarum IX: Systemat-
ics of Myoxanthus. Addenda to Platystele, Pleurothallis
subgenus Scapula and Scaphosepalurn. Monogr. Syst.
Bot. Missouri But. Card. 44: 1-128.
-. 1997. leones Pleurothallidinarum XV: System-
atics of Trichosalpinx. Addenda to Dracula, Masdeval-
lia, Myoxanthus and Scaphosepalurn. Corrigenda to Le-
panthes of Ecuador. Monogr. Syst. Bot. Missouri Bot.
Card. 64: 1—136.
Mendes, S. L. & M. P. Padovan. 2000. A Estayao Biolo-
gica de Santa Lucia, Santa Teresa. Espirito Santo. Bob
Mus. Biol. Mello Leitao (n. ser.) 11/12: 7-34.
Myers. N. B. A., C. G. Mittermeier. G. A. B. Fonseca &
J. Kent. 2000. Biodiversity hot spots for conservation
priorities. Nature 403: 853—858.
New Species of Cyclanthaceae from Southern Central America and
Northern South America
Burry E. Hammel
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.
barry.hamrnel@mobot.org
Abstract. Four new species of Asplundia and
one of Dicranopygium are described and illustrat¬
ed. Asplundia allenii is endemic to Panama. D. ta-
tica is endemic to Costa Rica. .4. brunneistigma is
known from Costa Rica and Panama, A. ceci from
Costa Rica to northern Colombia, and A. albicarpa
from Costa Rica to Ecuador. Asplundia stenophylla,
described originally from Costa Rica from sterile
material and now known from Costa Rica to Ec¬
uador, is here provided with a full description in¬
cluding staminate and pistillate flowers, as well as
fruits.
Key words: Asplundia, Central America. Cy¬
clanthaceae. Dicran opygium.
Over the course of the last 20+ years, collecting
for the Flora of Panama, Flora Mesoamericana,
and the Manual de Plantas de Costa Rica projects,
as well as systematic collecting of large-leaved
monocots by particular investigators, has given rise
to the accumulation of numerous undescribed spe¬
cies of Cyclanthaceae from the region. Principally
to make names available for the last-mentioned flo-
ristic project, the formal description of these spe¬
cies is here initiated.
All of the species of Asplundia Hailing, de¬
scribed below, pertain to the subgenus Asplundia,
characterized by asymmetrical and flat-topped
staminate flowers, rather than symmetrical and
concave as in subgenus Choanopsis (Marling,
1958). Although the great majority of species in
subgenus Asplundia have leaves that are subtri¬
costate to distinctly tricostate, three of the five
species here described are unusual in having es¬
sentially unicostate leaves. Their obvious mor¬
phological similarity and certain relation to par¬
ticular tricostate-leaved species is discussed.
One species of Dicranopygium Marling subg. Di¬
cranopygium is described. It is unusual (though
not unique) among members of the genus for its
very short and broad anthers.
Asplundia albicarpa Hammel. sp. nov. TYPE:
Costa Rica. San Jose: Parque Nacional Braulio
Carrillo, Estacion Carrillo, 400 m, 28 July
1985 (fJ. fr). R. Hammel A J. Trainer 14253
(holotype, MO |2 sheets, MO-5609384 & MO-
5609385]; isotypes, COT, CR. GIT 1NB. PM A,
QCNE, US). Figure 1.
Asplundiae stenophyllae affinis sed difieri laminis fo-
liorum menoribus profunde bifidis cum segmentis latiori-
bus menoribus profunde plicatis, fforibus staminiferis ma-
joribus cum staminibus pluribus et majoribus, fforibus
pistillatis cum tepalis in fructu longioribus, et stigmatibus
in fructu conduplicatis.
Long-stemmed (1-2 m) climbing epiphyte and
also often scandent terrestrial plant with conspic¬
uous adventitious roots. Petiole (1 ()—) 19-52 cm
long, the base dull green or often purple, the epi¬
dermis not brittle, with scant brown scales through¬
out, the sheath a thin, light brown and slight I v fi¬
brous soon caducous ribbon, the apex ± terete.
Leaf blades 30—52(—65) cm long, bifid from just
over 1/3—2/3 of their length, decurrent on the pet -
iole, unicostate, the lower surface with very scant
light brown scales; leaf segments 4-1 I cm wide,
lanceolate, acuminate, shallowly plicate, the veins
of the plicae not adaxially elevated. Peduncle dur¬
ing anthesis ca. 5 cm long, up to 7(—8) cm long in
later stages, the epidermis like that of the petiole
base. Spathes 3 or 4, the lowest attached below the
middle, the others ± evenly spaced above, the out¬
er ones green to brown or “maroon,” the inner ones
white, soon fibrous and caducous. Spadix at anthe¬
sis ca. 2.5—4(—5.5) X 1.5 cm, cylindrical. Staminate
flowers ca. 4 mm long, asymmetrical; receptacle ca.
2.5 mm wide, flat; perianth lobes 3 to 5, ca. 1-1.2
mm long, reaching to just above the base of the
anthers; stamens ca. 40-50, the anthers 0.8-1 X
0.5 mm, from conspicuous basal bulbs ca. 0.2 mm
long. Pistillate flowers ca. 4 mm wide at anthesis,
to ca. 10 mm wide in young fruiting stage; tepals
1-1.5 mm long and connate (at the base) at anthe¬
sis, 5—6 mm long and connate in young fruit (in
dried mature fruit to 8 mm long and connate below
the middle), broadly acute to rounded truncate.
Novon 13: 52-63. 2003.
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
53
Figure 1 Asplundia albicarpa Hammel. —A. Habit with detail of leaf blade (upper right), lower surface. — B Sta¬
gnate flower. —C. Pistillate flower at anthesis. —D. Pistillate flower in fruit. (A-C from Hammel & Trainer 14253 ; D
from Croat & Grayum 60250.)
equal to slightly surpassing the stigmas; styles lack¬
ing, the stigmas broadly ovoid to nearly quadrate,
with the pubescent fertile part surrounded by ster¬
ile tissue that becomes conduplicately folded and
turned upward in fruit, wholly surrounded by the
tepals. Fruiting spadix 4-7 X 2-2.5 cm, cylindri¬
cal; fruits green-white, sometimes tinged purple,
the seeds 1—1.3 X 0.6 mm wide, flattened, orange.
54
Novon
Distribution. Very wet primary forest of central
Costa Rica to Colombia and Ecuador, 200—1800 m.
Etymology. I lie epithet of this new species re¬
fers to the often greenish white color of the fruiting
spadix, a somewhat unusual color for species of ,4.s-
plundia, which are more often yellow-orange or
dark green.
Discussion. This new species appears to lie
closely related to Asplundia stenophylla, which lias
leaves that are deeply and prominently plicate
throughout, as well as narrower and more deeply
bifid (A. albicarpa leaves are usually bifid only to
about the middle and deeply plicate only at the
base). The staminate flowers in A. albicarpa are
somewhat larger with usually more and larger sta¬
mens, and the pistillate (lowers have tepals that be¬
come longer and stigmas with a conduplicately
folded sterile part in fruit. As indicated by the ep¬
ithet. the fruits of this species are often white or
greenish to yellowish white at maturity, whereas
those of A. stenophylla appear to remain green or
to become greenish yellow. Mature fruits of most
Central American species of Asplundia known to
the author are bright yellow, orange, or dull red at
maturity. Asplundia albicarpa and ,4. stenophylla
sometimes occur more or less sympatrically. as for
example in Parque Nacional Braulio Carrillo in
Costa Rica, in Chiriqui Province of western Pana¬
ma, and at Ea Planada Reserve in southern Colom¬
bia.
In size and shape of the leaves, Asplundia albi¬
carpa somewhat resembles A. polymera (Handel-
Mazetti) Hailing subsp. reitzii Hurling of south¬
eastern Brazil. That species, however, has more
deeply bifid leaf blades and the stamens are about
half as long and half in number, in addition to the
pistillate flowers having tepals reduced to low
wales.
Additional specimens (H = flower, fr = fruit, otherwise
sterile). COSTA RICA. Alajuela: Junto al Rio Sarapiquf
cerca del puente, camino a Colonia La Virgen del Socorro,
Hammel el al. 17743 (CR, INB, MO). Cartago: 6 km N
of La Suiza on the road to Paeayitas, Solomon 19227{ INB.
MO). Heredia: Parque Nacional Braulio Carrillo, 8 km K
of San Ramon, (fr), Loiselle 115 (MO); Kstacidn El Ceibo.
(fr), Aguilar A Hammel 29 (CR. INB. MO), Boyle 3108
(INB); Rara Avis, (fl), Vargas A- Frazee 48 (INB); Atlantic
slope of Voledri Barba, (fr), ('.rayitm et al. 7776 (CR). I.i-
"ion: N Hank of Lila de Matama, (fr). Grayum 10975.
11054 (INB); Parque Nacional Braulio Carrillo, Estacidn
Ouehrada Gonzalez a Sendero Botarama, (fr), Hammel et
al. 19090 (CR, INB. MO), Hurtado 93 (INB), (fr), A. Rod¬
riguez et al. 1497 (CR, INB. MO), (fl). M. M. Chavarria
el al. 206 (CR); Teleferico del Bosque, (fr), Hammel 19941
(INB). Hammel 19942 (CR. INB. MO). San Jose: Parque
Nacional Braulio Carrillo, Lstacion Carillo. (fr). L. I). Go¬
mez et al. 22912 (MO); sendero La Montura, (fr), Hammel
et al. 10839 ( CR. INB). PANAMA. Bocas del Toro: Lor-
tuna Dam area, in forest along road to Chiriqin Grande,
(fl. fr), Ci urcliill 5532. 5866. 5878 (MO), (fr). Groat A
Grayum 63250 (MO), (fr), Hammel A McPherson 14449.
14468 (MO). Hammel et al. 14676. 14693 (MO), (fr). Mc¬
Pherson A’ Aranda 10056 (MO). Chiriqui: along road to
Lortuna dam site from Gualaea, (fr). Antonio 5097 (MO.
PM A): Ouehrada los Chorros, (fr), Churchill A Churchill
6071 (MO), (fr), Churchill A Churchill 6072 (MO. PM A,
COL), (fr), Churchill & Churchill 6112. 6146 (MO); vicin¬
ity of Lortuna Dam, (fr). Groat 66568 (F, MO. PMA). (fr),
Croat 67853 (GB, MO. PMA, US), (fr), Croat A Zhu
76315 (MO. PMA), (fr), Croat & Zhu 76343 (MO), (fr),
McPherson 1 282 (MO); slope NW ol confluence of Rio
Horninto and Rio Chiriqui (fr), Stevens 18350 (MO), (fr).
Sytsma A Stevens 2203. 2225 (MO), (fr), van der Werff &
van Hardeveld 6653 (MO), (fr), van der Werff & van llar-
develd 6869 (MO); Cerro Colorado mine area. Hammel A
Trainer 14926 (MO). Code: above LI Cope (fr), Croat
67507 (MO, PMA); on continental divide above town,
Hammel 13648 (MO). Darien: Cerro Pine. (fr). Duke A
Elias 13799 (MO); slopes of Cerro Tacareuna, (fr), Ham¬
mel et at. 16535 (MO). Panama: vicinity of Cerro Jefe,
(fr), Croat 35912 (MO). Hammel A Trainer 15029 (MO),
(fr). Mori 7988 (MO), (fr), Witherspoon et al. 8284 (GB,
MO); Cerro Brewster region, (fr). Hammel A de Severs
13549. 13554 (MO), (fr), McPherson 7573 (MO). San
Bias (Comarea): Cerro Obu, (fl). de Severs et al. 8048
(MO), (fl). de Severs et al. 8072. 8085 (MO). Veraguas:
vicinity of Escuela Alto Piedra, (fr), Antonio 2965 (MO,
L), (fl. fr), Groat 27605. 49018. 66994 (MO), (fr), Ham¬
ilton A Kroger 3951-A (MO), (fr). Mori et al. 3916 (MO).
COLOMBIA. Chord: along road between Bolivar and
Ouibdo, near km marker 147, (fr), Croat 52095 (MO);
Municipio de San Jose del Palmar, hoyo del Rio Torito,
(fr). Torero et al. 7335 (MO). Nariiio: Reserva Natural La
Planada, (fr), Benavides 8968 (MO), (fr). Gentry et al.
35176 (MO), (fl). Gentry et al. 55100 (MO). ECUADOR.
Carchi: Cerro Golondrinas, Boyle et al. 1550. 1764 (MO).
Esmeraldas: Reserva Etnica Awd, (fr), Aulestia et al. 315
(MO); Mataje village, (fr), Neil! et at. 11773 (MO); Ouin-
inde, Bilsa Biological Reserve, (fr). Pitman A Bass 794
(MO. (JCNE).
Several collections, all from the Chiriqui Prov¬
ince of Panama, have leaves of a length and fruits
ol a shape more like those of A. albicarpa. but w idth
of the leaf segments and depth of the division more
like A. stenophylla. These possible hybrids between
those two species are cited below:
PANAMA. Chiriqui: vicinity of Lortuna dam, Croat
66580 (MO, PMA), Croat 67823 (MO), McPherson 7281
(MO).
Another possible hybrid with A. albicarpa in¬
volves. remarkably, the tricostate species A. eurys-
patha Harling. See also discussion under .4. sten¬
ophylla. A population from near La Suiza,
Turrialba, in Costa Rica seems to demonstrate a
gradation in leaf and fruit morphology between .4.
albicarpa and A. euryspathir. an apparently inter¬
mediate specimen is cited below:
COSTA RICA. Cartago: de La Suiza de ' I’urrialba rum-
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
55
bo a Pacayitas, (fr), Hammel & Kennedy 21121 (CR. INB.
MO).
Asplundia allenii Hammel, sp. nov. I V PE: Pana¬
ma. Code: El Valle de Anton, vicinity Finca
Tomas Arias, 600 m, 5 Aug. 1946 (fl & fr), P.
Allen 3623 (holotype, G; isotype, F). Figure 2.
Asplundiae sleeperae valde affinis, ob tepalis pistilatis
3-lobatis et forma stigmatum, serl differt laminis foliis for-
mose palmatipartitis (vice simpliciter I >i part it is) cum plu-
ri Gns segment is veris, non laesione fort u i ta. et antheris
longioribus.
Terrestrial plant, acaulescent or with a short,
erect stem up to 50 cm long. Petiole 112-150 cm
long, the base dull green in life, drying slightly to
conspicuously reddish brown, heal blades 61—75
cm long, split at the middle ca. 2/3 their length but
overall split into 25-30 divisions that reach up to
3 cm from the base, tricostate, the lateral costae
running in the margin at the base, the surface lack¬
ing scales; leaf segments 2.5-5 cm wide, linear,
narrowly acute to acuminate. Peduncle during an-
thesis ca. 10 cm long, at least 14 cm long in later
stages. Spathes 4, the lowest one attached at about
the middle, the others somewhat clustered apieally,
fibrous and soon caducous. Spadix at anthesis 3 X
1.5 cm, ellipsoid. Staminate flowers ca. 4 mm long,
asymmetrical; receptacle 3—4 mm wide, flat; peri¬
anth lobes 5 to 7, ca. 1 mm long; stamens 30-40,
the anthers 1—1.4 X 0.5 mm, from rather large bas¬
al bulbs. Pistillate flowers 3-5 mm w ide at anthesis,
6-8 mm wide in young fruiting stage; lepals at an¬
thesis ca. 1 X 2 mm, free, 3-lobed nearly to base,
the lobes ca. 3 mm long in fruit, acute, free and
shorter than the stigmas; styles ca. 1 mm long, free
at anthesis, ca. 2 mm long and connate in the lower
1/2 in young fruit, the stigmas broadly ovate, not
flanked by sterile tissue, slightly projecting be¬
tween the tepals, distinctly grooved in young fruit.
Fruiting spadix ca. 5 X 1.5 cm, cylindrical; fruits
(immature) greenish white, the seeds ca. 1.5 X 0.8
mm, flattened, orange.
Distribution. This rare species is known from
lowland forest (50—270 m) from four sites in middle
to eastern Panama. It has been collected fertile only
once from the region where it is most common (Rio
Boquerdn and Rio Guanche); it is lound there in
more or less open areas, in sandy soil along rivers
and also on slopes in forest.
Etymology. This species is named for Paul H.
Allen (191 1-1963), longtime resident and botanist
of the American tropics, collector of the first known
specimen and, to date, the only known flowering
collection of the species.
Discussion. Except for the very strikingly pal-
mate-divided leaves with numerous, linear divi¬
sions, this species matches Asplundia sleeperae
Grayum & Hammel quite closely, differing from it
in fertile characters principally by the slightly lon¬
ger anthers (0.7-1.2 mm in A. sleeperae vs. 1-1.4
mm in A. allenii). Asplundia sleeperae is now known
from wet forests of Nicaragua to central Panama,
from near sea level up to about 1200 m.
By virtue of its similarly 3-lobed pistillate tepals,
as well as the shape of the stigmas, the much small¬
er A. pittieri (Woodson) Harling is also obviously
related to the present new' species. Asplundia pit¬
tieri is itself unique in the genus for having fruits
that do not dehisce at maturity. In all other species
known to the author, the stigmas and tepals form a
unit that dehisces at the base, leaving the succulent
mass of seeds on the spadix rachis. In A. pittieri
these fruit “caps” remain fused, and the multiple
“fruit” (the spadix) becomes engorged and nearly
smooth at maturity, releasing the juicy, seed-laden
contents more or less explosively when punctured,
somewhat similar to species of Dicranopygium. As¬
plundia pittieri has a narrowly disjunct distribution,
known in Costa Rica only from the wet forests of
the southern Pacific slope (especially the Osa Pen¬
insula) and then again in Panama from Code, Pan¬
ama. and San Bias Provinces. Some material from
northwestern Colombia has also been tentatively
identified as A. pittieri and A. sleeperae.
Numerous species of Asplundia have been de¬
scribed as having the lateral segments “secondarily
split” into numerous lobes (cf. Harling, 1958,
1973), but the wording of descriptions and the
dried specimens themselves have left considerable
doubt as to whether or not these splits are actual
lobes or merely random lacerations by physical
damage to very broad-leaved species. Harling, in
fact, implied that the latter is the case: “The leaf
of the schizophylla type is, however, frayed and sec¬
ondarily deeply cleft into many irregular lobes.
Such lobation of old leaf blades is nothing remark¬
able, but a characteristic feature of many Asplundia
spec ies” (Harling, 1958: 235). With this new spe¬
cies I mean to leave no doubt: the divisions are
developmental, perhaps brought about by differ¬
ential growth, not lacerations from physical damage
to leaves that would otherwise be only bifid. In fact,
the identity of the species is dependent, in large
part, upon this distinction. In southern Central
America only one other species of Asplundia has
similarly divided leaves, the epiphytic A. aurantia-
ca Harling, most certainly related to A. dominguen-
sis Harling and A. gamotepala Harling, all quite
56
Novon
Figure 2. Asplundia allenii Hammel. —A. Habit. —B. Detail of leaf blade, lower surface. —C. Staminate flower will,
det ail of stamen. —1). Pistillate flower at anthesis, top view, side view. (A, I? from Hammel & Trainer 14773 ; C I)
from Allen 3623.)
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
57
distinct from A. allenii (see also discussion under
A. ceci).
Additional specimens. PANAMA. Colon: Rfo Guanche,
ca. 5 km upstream from road to Portobello, Hammel &
Trainer 14773 (MO). Darien: Parque Nacional del Dari¬
en, Estacion Rancho Frio. Hammel et al. 16148 (MO), (fr).
Hammel et al. 16160 (MO). Panama: Rio Boqueron, (fr).
Dressier 4696 (GB), Hammel & Kress 1.3405 (DI KE).
Hammel 16092 (MO).
Aspluntlia hrunneistigma Hammel. sp. nov.
TYPE: Costa Rica. Heredia: Parque Nacional
Braulio Carrillo, primary forest between Rfo
Pejo and Rfo Sardinalito, 700—800 m, 3 Apr.
1986 (fr), M. Grayum 6709 (holotype, MO; iso-
tvpes, CR. GB, US). Figure 3.
Asplundiae peruvianae valde affinis, ob stigmatibus late
rotundatis ad aliquantum quadratis, epidermide petiolo-
rum flava et rimosa, et laminis foliorum infra cum squamis
brunneis, sed ab ea et speciebus affinibus differt habitu
praecipue terrestri, foliis unicostalibus et vadosis bifidis.
Terrestrial or shortly climbing caulescent plant,
the stem 1 —2(—3) m long. Petiole 25-40 cm long,
the base with yellowish tan, shiny and brittle epi¬
dermis, elsewhere with brown scales. Leaf blades
40—75 cm long, bifid 1/3—1/2(—2/3) their length,
unicostate, narrowly acute to decurrent on the pet¬
iole, the surface (especially below along veins) with
conspicuous brown scales; leaf segments 6—14 cm
wide, broadly lanceolate, shortly acuminate. Pe¬
duncle during anthesis ca. 10 cm long, up to 15
cm long in later stages, the base with epidermis
like that of the petiole base. Spathes 3 or 4, the
lowest one attached below r the middle, the others ±
evenly spaced above, their epidermis somewhat
shiny and tan as in the petiole bases, but soon fi¬
brous and caducous. Spadix at anthesis 3—5 X 1.5—
2 cm, cylindrical. Staminate flowers 5-7 mm long,
asymmetrical; receptacle 3-4 mm wide, flat; peri¬
anth lobes 5-7, 1.5-2.5 mm long; stamens 25-35?,
the anthers 1.0-1.5 X 0.5 mm, from conspicuous
basal bulbs ca. 0.5 mm long. Pistillate flowers ca.
7 mm wide at anthesis, to ca. 10 mm wide in young
fruiting stage; tepals ca. 5 mm long and free to the
base at anthesis, 6-7 mm long and free or connate
to just above the middle, broadly acute to rounded,
about the same height as the stigmas in young fruit;
styles ca. 3 mm long and free at anthesis, ca. 7 mm
long and connate to just above the middle in young
fruit, the stigmas broadly ovate, wholly surrounded
by the tepals, the sterile part conduplicately folded
and turned upward, above the conspicuously red¬
dish brown sulcus. Fruiting spadix 5-8 X 2.5 cm,
cylindrical; fruits orange, the seeds 1-1.2 mm long,
0.5 mm wide, flattened, orange.
Distribution. Asplundia hrunneistigma is known
only from mid-elevation (400—1000 m), very wet,
primary forests of central Costa Rica to central Pan¬
ama.
Etymology. The epithet "hrunneistigma” refers
to the brown fertile tissue (contrasting with the
green sterile tissue) of the stigmas as seen in live
material. Although that may not be particularly dis¬
tinctive among Asplundia species, when I was first
collecting these plants it seemed so and the name
has stuck.
Discussion. Asplundia hrunneistigma is easy to
distinguish by its relatively long-stemmed, usually
terrestrial to sometimes climbing habit, and the
narrow, shallowly divided, unicostate leaves with
yellow, cracking epidermis throughout the petiole
bases. Although this species might seem to resem¬
ble A. guianensis Harling, as per Harling’s key
(1958), examination of type material suggests oth¬
erwise. Asplundia guianensis does have a petiole
with scant patches of shiny yellow and brittle epi¬
dermis, but it is otherw ise quite unlike the present
species (see discussion under A. ceci). Especially
the structure of the stigmas in A. guianensis, being
somewhat laterally flattened and without surround¬
ing sterile tissue, is very unlike that of A. hrun¬
neistigma, which instead needs to be compared
with A. peruviana Harling, A. krukoffii Harling, A.
venezuelensis Harling, and A. xiphophylla Harling.
All of these have the very same petiolar epidermis,
bright yellow and cracking throughout (even in the
sheath), brown scales abundant on the lower leaf
surface and often also on the petiole and sheaths,
as well as stigmas that are quadrate to broadly
rounded, rather than laterally flattened at anthesis.
From all. A. hrunneistigma is distinct by the less
deeply divided and always strictly unicostate leaf
blades, as well as by its usually terrestrial (though
long-stemmed) habit.
Asplundia hrunneistigma is regularly infected by
a gall-producing fly in the family Cecidomyiidae
(Uiptera). Their bright orange larvae, which affect
the aerial roots, give rise to conspicuous globular
galls (to ca. 1.5 cm), not seen on other Cyclantha¬
ceae nearby. This and similar galls on the petioles
of A. euryspatha are the only know n records of galls
on Cyclanthaceae (Paul Hanson, pers. comm.).
Most of the fertile material known for this species
is in fruit; the scant flowering material available is
old, and thus it is difficult to determine the number
of stamens.
Additional specimens. COSTA RICA. Heredia: Parque
Nacional Braulio Carrillo, puesto el Ceibo, Boyle 2883
(CR. 1NB. MO), Boyle 3034 (INB. MO); between Rfo Pejo
and Rfo Sardinalito, (fr), Grayum & G. Herrera 7907 (CR,
58
Novon
Figure 3. Asplundia brunneistignw Hammel. —A. Habit, —lb Staminate flower (stamens fallen) with detail of stamen
—C. Pistillate flower, young fruiting stage. (A from Hammel 22380: 15. C from Hammel & Kress 13639.)
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
59
MO, F); S\A of Horquetas on road to Plastico (Haras Avis),
Hummel 16710 (CH. INB). Linidn: Quebrada Molinete,
propiedad do Rainforest Aerial Tramway, (fr), Hammel &
Perry 19726 (CR, MO), (fr). Hammel 19945 (CR, IMi.
MO), Hammel 22580 (INB). PANAMA. Code: near con¬
tinental divide above El Cope, (lr), Hammel 1060 (MO),
(fl, fr), Hammel & Kress 11335, 13444, 13462 (DUKE),
(fr), Hammel & Kress 13447, 13639 (DUKE, MO). Colon:
Santa Rita ridge, Hammel et al. 14381 (MO). Panama:
Cerro Jefe, trail from end of road past Los Altos de Pacora,
(11). Hammel & (le Nevers 13572 (MO, PMA); in Clusia
and Colpolhrinax forest near radio tower, (fr), Hammel &
Trainer 15030 (MO).
Asplundia ceci Hammel, sp. nov. TYPE: Costa
Rica. San Jose: Parque Nacional Braulio Car¬
rillo, Estacion Carrillo, 600 m. 28 July 1985
(11 & fr), H. Hammel & ./. Trainer 14260 (ho-
lotype, MO; isotypes, COE, CR. CR, PMA,
IJS). Figure 4.
Asplundiae sanctae-ritae et aliis speciebus valde affmis,
ob floribus pistillatis dense crebris, stigmatibus lateraliter
complanatis, et squamis badiis basium petiolorum, sed all
iis differt laminis foliorum parvioribus convexis, profun-
dioribus bipartitis, cum costis lateralibus debilibus mar-
ginalibus, et praecipue pedunculo et infructescentia bre-
vioribus.
Profusely branching and climbing plant (liana)
or rarely terrestrial, the stem to 15 m or longer.
Petiole 25—40 cm long, the base dull, not cracking,
reddish brown (with dense scales), elsewhere with
scattered reddish brown scales. Leal blades 25—50
cm long, bifid 3/4—4/5 of their length, subtricostate,
the lateral costae in the margin, broadly acute to
the base, the surface (especially below along veins)
with scant reddish brown scales; leaf segments 6—
10 cm wide, elliptic to oblanceolate, markedly con¬
vex above and cuspidate. Peduncle during anthesis
ca. 3 cm long, 4-5(-6) cm long in fruit, also with
reddish brown scales. Spathes mostly 4. ± evenly
spaced above the middle, dark reddish brown, rig¬
idly fibrous to almost woody, tardily caducous. Spa¬
dix at anthesis ca. 3-4 X 1.5 cm, ellipsoidal. Sta-
minate flowers ca. 5 mm long, asymmetrical;
receptacle 2.5—3 mm w ide, flat; perianth lobes 4 to
6, 0.8—1.2 mm long; stamens 30—35, the anthers
1—1.4 mm long, from small basal bulbs ca. 0.2 mm
long. Pistillate flowers ca. 4 mm wide at anthesis,
7—9 mm wide in fruit; tepals ca. 2 mm long and
free to the base at anthesis, 2.5—3 mm long and
connate to just above the middle, broadly rounded
to truncate and sometimes emarginate, same height
as or slightly lower than the stigmas in fruit: styles
ca. 2 mm long and fused at anthesis, 2.5—3 mm
long in fruit, the stigmas narrowly elliptic, laterally
compressed, slightly projecting between the tepals
at anthesis, later laterally swollen with the fertile
part protruding. Fruiting spadix 4—6 X 1.5—2.5 cm,
ellipsoid; fruits green, the seeds 1-1.7 mm long,
0.6—0.7 mm wide, flattened, orange.
Distribution, This somewhat rare species is
known from scattered localities in very wet, mid¬
elevation (500—900 m) primary forests from central
Costa Rica to northwestern Colombia.
Etymology. I am pleased to honor Cecilia
(“Ceci”) Herrera Chacon by naming this species
after her. She came to us for an interview for the
position of assistant to the Manual Project back in
1987, at the suggestion of her cousin, renowned
Costa Rican plantsman. Gerardo Herrera. She has
been with us, loyal, hard-working, and consistent,
ever since. Thank you, Ceci. The epithet “ceci" is
used as a noun in apposition with no Latin termi¬
nal ion. as sanctioned by the Saint Louis Code, Ar¬
ticle 23.1 (Greuter et al.. 2000), and therefore not
to be corrected (Art. 60.11).
Discussion. This copiously branching climber
is easily recognized by its dark reddish brown
stems, petiole bases, and spathes, and its small in¬
distinctly subtricostate leaves, which, in life, are
distinctly domed (convex), with the margin and tip
curved down. With respect to the reddish brown
stems and characters of the inflorescences (fibrous
and leathery spathes) and flowers (especially the
somewhat laterally compressed stigmas), among
Costa Rican species it is most similar (and probably
related) to A. aurantiaca and A. sanctae-ritae Gal-
eano & Bernal. From those two, as well as from
other related South American species, Asplundia
ceci is distinct for its relatively small leaves with
the lateral costae weakly developed and entirely in
the margin, high climbing habit, and for the short
peduncles and fruiting spadices. Overall this spe¬
cies looks most like A. guianensis, which also is
said to be a rather high-climbing (to 7 m) liana.
This latter species has similarly but even more
weakly subtricostate leaves, traces of the reddish
brown scales on various structures, and very similar
pistillate flowers. The new species differs from A.
guianensis by its somewhat smaller leaf blades with
more w idelv divergent divisions, the shorter pedun¬
cles, and by the pistillate tepals being slightly
shorter than, rather than surpassing, the stigmas in
fruit. The spathe(s) of A. guianensis was said by the
collector to be “whitish" while those of A. ceci are
reddish brown. Asplundia cabererae Harling and A.
gamotepala Harling may also share relationship by
virtue of the reddish brown scales and the structure
of the pistillate flowers, but both have much larger
leaves, and longer peduncles (30—35 cm in the lat¬
ter species). Asplundia cabrerae also differs from A.
60
Novon
Figure 4. Asplundia ceci Hammel. —A. Habit. —B. Detail of inflorescences with spathes. —C. Staminate flower with
detail of stamens. —D. Pistillate flower at anthesis. (All from Hammel 22378.)
ceci by virtue of the tepals of the pistillate flowers
being Iree even in fruiting material. In summary,
by reason of the reddish brown petiole bases and
the structure of the pistillate flowers, A. brunneis-
tigma is most similar to A. cabrerae, A. gamotepala,
A. guianensis , A. pycnantha, and A. sanctae-ritae,
but dillers from all of them by its much smaller and
more deeply divided leaves with weak lateral costae
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
61
(wholly in the margin), as well as by the much
shorter peduncle and infructescence. In addition, it
can be separated from A. pycnantha by the lused
styles, from A. cabrerae by the fused tepals, and
from A. guianensis by the stigmas surpassing the
tepals.
Additional specimens. COSTA RICA. Heredia: Parque
Nacional Braulio Carrillo, puesto el Ceibo, Hoyle 2908
(1NB, MO); between Rfo Peje and Rio Sardinalito, (fl. Ir),
Grayum 6697 (CR. F, MO, GB, US); Horquetas to Finca
Plastico, (fl, fr), Hammel A: Robles 16692 (CR). Limon:
Quebrada Molinete, propiedad de Rainforest Aerial I rain¬
way, (fl. fr), Hammel 22378 (INB): Parque Nacional Brau¬
lio Carrillo, Sendero Botarrama, (fl. fr), Hammel 19082
(CR, INB), (fr), Hammel & Troyo 19953 (CR. INB, MO),
Lepiz 178 (INB); Rfo Sukut por el Rfo Uren, (fl), Hammel
et al. 17593 (CR, INB). San Jose: Parque Nacional Brau¬
lio Carrillo, Estacidn Carrillo, (fl. fr). Hammel & Trainer
14250 (CR. F, MO, K), (fl, fr), Hammel & Chacon 16053
(CR, MO). PANAMA. Code: El Valle de Anton, vicinity
of Ea Mesa, (fl. fr), Croat 67243 (MO. PM A, CAS). CO-
LOMBIA. Antioquia: Municipio de San Luis, por la au-
topista Medellfn-Bogota, quebrada “La Mariola,” (fr), Co-
gollo et al. 4947 (MO. JAUM), (11), Hoy a & Hernandez
975 (MO, JAUM).
Because fertile material of Asplundia stenophylla
has never been described, and because one of the
new species here proposed (A. albicarpa) appears
to be closely related to it, I here provide a complete
description based on the abundant fertile material
now known.
Asplundia stenophylla (Standley) Hailing, Acta
Horti Berg. 17: 43. 1954. Carludovica steno¬
phylla Standley, Publ. Field Mus. Nat. Hist.
Bot. Ser. 18: 130. 1937. TYPE: Costa Rica.
Cartago: El Muneco, Standley 33829 (holo-
type, US).
Long-stemmed (up to 5 m) climbing epiphyte, or
rarely terrestrial plant. Petiole 20—44 cm long, the
base dull green or often purple, the epidermis not
brittle, sometimes with scant brown scales through¬
out. the apex deeply grooved. Leal blades (44— )50—
77 cm long, bifid from 5/6—9/10 of their length,
decurrent on the petiole, unicostate to faintly sub¬
tricostate, the lateral costae well in from the mar¬
gin; leaf segments 2.5—6 cm wide, linear, long acu¬
minate, deeply and permanently plicate throughout,
the veins of the plicae adaxially raised. Peduncle
during anthesis 4-7 cm long, up to 11 cm long in
later stages. Spathes 3 or 4. the lowermost at about
the middle, the others ± clustered above, cream-
colored, turning brown and soon caducous. Spadix
at anthesis 3^1.5 X 1-1.5 cm, cylindrical. Stami-
nate (lowers 3-3.5 mm long, asymmetrical; recep¬
tacle 2.5-3 mm wide, flat; perianth lobes 3-5, 0.5-
1.1 mm long, reaching to the base of the anthers
or just above; stamens 30—42, the anthers 0.5—1
mm long, ca. 0.5 mm wide, from basal bulbs ca.
0.2 mm long. Pistillate flowers 4—5 mm wide at
anthesis, to 11 mm wide in young fruiting stage;
tepals 2-2.5 mm long and connate (at the base) at
anthesis, 3.5-4 mm long and connate in young fruit
(in mature fruit no longer, and connate to above the
middle), acute to broadly so al anthesis, rounded
truncate in later stages, equal to or slightly exceed¬
ed by (in later stages) the stigmas; styles essentially
lacking (1 mm long at most), the stigmas broadly
elliptic to ovoid quadrate with the pubescent fertile
part flanked by sterile tissue that is only slightly
raised in later stages, slightly protruding between
the tepals or surrounded by (in later stages) the
tepals. Fruiting spadix 5-9(-12 fide Murphy 1172)
X 1.5-3.5 cm, cylindrical; fruits green, sometimes
tinged purple or brown, the seeds 1.2-1.5 mm long,
0.6-0.7 mm wide, flattened, orange.
Distribution. This species is quite common in
very wet cloud forest from 1300 to 2200 m from
Costa Rica to Ecuador.
Discussion. Asplundia stenophylla, until now
described from only sterile material and distinctive
because of its long, narrow, very deeply divided and
deeply corrugated leaves, belongs to the subgenus
Asplundia, where it appears to be closely related to
A. albicarpa, described herein. It appears to be fur¬
ther distinct from A. albicarpa by virtue of its pet¬
ioles being deeply grooved all the way to its apex,
whereas those of the latter are nearly terete at the
apex. Asplundia stenophylla is a perplexing species
because in spite of having very narrow leaf blades,
they are often indistinctly subtricostate, a situation
usually found only in much broader-leaved species.
Specimens examined. COSTA RICA. Alajuela: Monte
Verde Reserve, E slope of Cerros Centinelas, (fr), Grayum
5366 (MO); 15 km NW of San Ramon, Cerro Azahar, (fr),
Liesner et al. 15586 (MO). Cartago: ca. 1-2 km above El
Muneco, (fl), Luteyn 3235 (DUKE). Limon: Cordillera de
Talamanca, Rfo Sinf, (fr), Davidse & Herrera 29111 (MO);
Alto Lari, (fl). Herrera 5197 (MO). Puntarenas: Cordillera
de Talamanca, between Cerro Frantzius and Cerro Pittier,
(fr), Davidse et al. 28410 (MO); Rfo Bella Vista, NIX' of
Las Alturas, (fl), Davidse 24342 (MO); Monte Verde Re¬
serve, Cerros Centinelas, (fr), Grayum & Sleeper 3850
(MO); Sendero Pantanoso, (fr). Haber ex Hello 5203 (MO);
Sendero Rfo, (fl). Hammel & Trainer 13783 (MO); Sendero
Nuboso, (fl. fr), Hammel 13854 (MO); Sendero Brillante,
(fl), Hammel & Zuchowski 13888 (MO); Las Alturas, (fr),
Murphy 1172 (DUKE, MO). San Jose: about 5 km NE of
San Isidro de Heredia, (fl), John & Kathy Utley 5081
(DUKE). PANAMA. Bocas del Toro: Cerro Colorado
mine area, (fl). Hammel & Trainer 14902 (MO). Chiriquf:
Cerro Horqueta, (fr), Blum & Dwyer 2669 (MO); near Cer¬
ro Pate Macho NE of Boquete, Hammel et al. 11395 (MO);
Boquete, Palo Alto area, (fl), Hammel et al. 14388 (MO);
62
Novon
I" inca Ojo de Agua, (fl), Knapp 1589 (MO); Boquete, Cerro
Horqueta, (fr), Dwyer & Hayden 7729 (MO); Distr. Bug-
aba. Santa Clara. Hartmann’s Finca, (fl), van der Werff
Herrera 7125 (MO). COLOMBIA. Narino: Reserva Nat¬
ural La Planada, (fr), Benavides 8918 (MO), Hammel A
Bernal 15800 (MO). ECUADOR. El Oro: Montana Schi-
cay, near Cachii aran, (fr), Steyermark 54116 (F). Pichin-
eha: Canton Quito, Chiriboga, carretera vieja Quito-Sto.
Domingo, (fr), Cerdri & Iguago 8515 (MO), (fr). Hammel
& Trainer 15812 (MO). Santo Domingo to Quito, Tandapi.
(fr). Hammel & Trainer 15892 (MO); Reserva Orquideo-
logica El Pahumma, (fl), Suin et al. 897 (MO).
At several sites in Costa Rica this species ap¬
pears to grade into something with very broad, con¬
spicuously tricostate leaves. In these areas, fertile
individuals of typical A. stenophylla can be easily
found, but also common are fertile individuals, in¬
termediate between that and distinctly tricostate,
broad-leaved individuals. On close examination,
tbe intermediately broad-leaved individuals are
slightly different (mostly in terms of the staminate
flowers) from both A. stenophylla and what appears
to be tbe other parent (A. euryspatha) in a complex
hybrid zone. Collections of the narrower-leaved, but
distinctly tricostate, intermediate forms are cited
here below and will be labeled “A. stenophylla
prob. X A. euryspatha.”
Although cytology has been helpful in suspected
cases of hybridity, e.g., in ferns and grasses, and
one might suggest such a line of inquiry to pursue
further these cases in Asplundia, my own attempts
to gel chromosome counts for Cyclanthaceae have
led only to frustration. It is not for lack of trying
that the family is still very poorly known cytologi-
cally (see, e.g., Eriksson, 1995). Presently, so little
is known that nothing can be said even about pos¬
sible chromosome evolution at the generic level, let
alone comparisons be made between species sus¬
pected of hybridization.
Probable hybrids with A. euryspatha. COSTA RICA.
Cartage*: Tapantf, tributary of Quebrada Casa Blanca, (fr),
Grayum 8612 (MO); between Parafso and Turrialba, (fr),
Solomon 19206 (CR. MO); Canton de El Guarco. Carretera
Interamericana, entre Cangreja y Casamata, Hammel
19811 (INB). Heredia; Parque Nacional Braulio Carrillo,
Puesto El Ceibo, Boyle 1881 (CR, MO). Puntarenas:
Monte Verde, along Rio Guacimal below lecheria, (fl. fr),
Hammel & Trainer 18799 (CR. F. GB. MO. US); sendero
brillante, (fr), Hammel & Zuchowski 18889 (MO). San
Jose: Canton de Perez Zeleddn, km marker 105—1 16 Car¬
retera Interamericana, (fr), Hammel 18557 (CR, MO); km
marker I 17, 1750 m, (fr). Hammel 19885 (CR. INB).
Dicranopygium tatica Hammel, sp. nov. TYPE:
Costa Rica. Limon: Rio Bananito, along small
stream on Selva Bananito Lodge property. 300
m, 24 IVlar. 2001 (fl & fr), H. Hammel & ,/. F.
Morales 22857 (holotype, INB; isotypes. COL,
CR, MO), f igure 5.
Dicranopygio venezuelano similis, ob antheris parvis
plus minusve sessilibus connectivis late triangularis in-
sidentibus, sed differ! floribus staminiferis asymmetricis
cum tepalis et staminibus paucioribus, et segmentis fo-
liorum multo angustioribus.
Short-stemmed terrestrial plant, often on rock
cliffs along streams and in forest. Petiole 20—57 cm
long, the base often dull purple, tbe sheaths falling
± entire, not as fibers. Leaf blades 35—53 cm long,
bifid to about 4/5 of their length, unicostate; leaf
segments 1.5—2.5 cm wide, linear, long acuminate,
deeply and closely plicate with ea. 5 ridges per cm
(dried material). Peduncle during anlhesis 10-13
cm long, up to 22 cm long in later stages. Spathes
3. 4-6.5 cm long, long acuminate, tinged dull pur¬
ple. Spadix at anthesis 1-1.5 X 0.8-1 cm. ovoid
elliptic. Staminate flowers ea. 2 mm long, asym¬
metrical: receptacle 2 mm w ide, flat: perianth lobes
5. 0.5-0.6 mm long, reaching to about the middle
of the anthers; stamens I 1 — 14, the anthers 0.5—0.8
nun long and about as wide, essentially without
basal bulbs but the connective broadly triangular
from about the middle down. Pistillate flowers ea.
2.5 mm wide at anthesis, 4—4.5 mm wide in young
fruiting stage, the tepals ea. I nun long, free at the
base at anthesis, ea. 2 mm long and free in young
hint, truncate, equal to the stigmas in height; styles
ca. 0.5 mm long, the stigmas broadly ovate elliptic
to rounded, wholly surrounded by the tepals. Ma¬
ture fruiting spadix not known, but judging from
the immature ones more than 2 cm long and ca.
1.5 cm wide, ovoid to cylindrical.
Distribution. So far this species is known only
from the Caribbean lowlands of Costa Rica from the
two nearby watersheds of Rfo Barbilla and Rib
Bananito at about 200—700 m elevation.
Etymology. Iatiea is one of the many nick¬
names by which one of this species’ early collec¬
tors, Gerardo Herrera, is fondly known among his
colleagues and friends. Gerardo, who was also one
of the Manual Project’s early collectors, is re¬
nowned among those who know his collections, and
is the stuff ol legend among those who have been
in the field with him. The epithet ’’tatica” is used
as a noun in apposition.
Discussion. Dicranopygium tatica is unusual
among congeners of southern Central America, not
only for the long, linear leaf segments, but also for
its very small anthers with basally broadened con¬
nectives. Ry way of the anther size, shape, and
number, as w r ell as characters ol the pistillate flow¬
ers, the species seems to be related to I). Venezue¬
la num Hailing even though that species has sym¬
metrical staminate flowers (with perianth lobes on
Volume 13, Number 1
2003
Hammel
New Species of Cyclanthaceae
63
Figure 5. Dicranopygium tatica Hammel. Habit (in frame). Details, left to right: infructescence; inflorescence; sta-
minate flower; pistillate flower (at anthesis). (All from Hammel 22.157.)
all sides of the receptacle). The only other species,
al least in Central America, with similar anthers is
I). harlingii, which, however, has minute, nearly
invisible staminate perianth lobes and only 5 to 8
stamens. The present new species is strikingly dif¬
ferent vegetatively, for its long and narrow leaf seg¬
ments, from all others with anthers of similar size
and shape, and different also in details of fhe flow¬
ers.
Additional specimens. COSTA KICA. Limbn: Cordillera
. Female inflorescence with 2 mature (mils. —c. Young male
inflorescence. —d. Older male inflorescence. Drawn from Marling & Stahl 26727 (S).
86
Novon
the midvein flanked by two major basal veins. Both
male and female inflorescence types with very
short, simple peduncle 1-2 mm, followed by one
fertile internode. Male inflorescence < 1.5 cm long,
flowers to 50 per fertile bract, regularly triseriate.
the terminal, exhausted part of the fertile internode
becoming rod-like. Female inflorescence < I cm
long, flowers mostly 3 per fertile bract, the fertile
internode swollen to 6 mm in fruit, and investing
the bases of at least the young fruits. Fruits about
1 cm in width, longitudinally compressed, in shape
a thick, rounded disk, white, with small, recurved
petals.
Dendrophthora harlingii is a species closely re¬
lated to D. dalstroemii kuijt. but with some sharp
differences. Dendrophthora dalstroemii has irregu¬
larly uniseriate flowers, the inflorescence peduncle
making up about half of the length of the entire
inflorescence, which mostly has 2 fertile internodes;
it lacks basal cataphylls; and its leaves are consis¬
tently ovate rather than obovate. Finally, its fruiting
internode does not show the distinctive swelling
that characterizes I). harlingii.
Dendrophthora harlingii is perhaps more closely
related to D. cuneifolia kuijt from Venezuela (kuijt.
1990), which, in general appearance, resembles it
strikingly. Dendrophthora cuneifolia is monoecious,
however, with unisexual inflorescences of both
types occurring even on the same twig. Addition¬
ally. that species bears to 30 or more female flowers
per fertile internode, and the flower seriation of the
male inflorescence soon becomes irregular, while it
remains strictly triseriate in D. harlingii , even on
old inflorescences. The fruit of I). harlingii also
appears much larger, and longitudinally com¬
pressed rather than more or less globular, but the
shape of fruits is difficult to ascertain from herbar¬
ium material, f inally, inflorescence peduncles in I).
cuneifolia may be up to 1 cm long, while those of
D. harlingii rarely exceed I mm; and basal cala-
phvlls are absent in the former, present in the latter.
Yet another species that may well be related is
Dendrophthora virgata (Trelease) kuijt as illustrat¬
ed in kuijt (1980), which differs in having male
and female inflorescence peduncles to 6 mm long,
to 10 flowers per female fertile bract, and much
smaller (< 4 mm) leaves.
Paratype. As the type. Hurling & Stahl 267.17 (S).
46. Dendrophthora meridana kuijt. sp. nov.
TYPE: Venezuela. Trujillo, border with Lara:
Parque Nacional Dinira, Paramo de Jabon, fila
cerca de la cumbre del Pico Jabon. shrubby
paramo vegetation between boulders and rocky
escarpments, 9°34'24"N, 70°07'03"W, 3100
m. 1 1 Aug. 1999, R. Riina, R. Duno. R. Ghin-
aglia & R. Gonto 62.3 (holotype, VEN
301604). Figure 1 I.
Planta monoecia, glabra, ereeta, sat carnosa, caulibus
teretibus; cataphylla basalia nulla; folia usque ad 13 X 3
mm, oblanceolata. lnfiorescentia mascula in axilla pro-
phy 1 li posita, fforihus triseriatis usque ad 18 unaquaque
bractea fertili subtends. lnfiorescentia feminea in axilla
folii posita, floribus 4(6) triseriatis unaquaque bractea fer¬
tili subtends.
Rigidly erect, glabrous, monoecious plants ca. 30
cm in height, densely branched, internodes to 2.5
cm long, terete, basal cataphylls absent, basal phyl-
lotaxy transverse. Leaves to 13 X 3 mm, oblaneeo-
late. fleshy, apex obtuse or nearly so. base cuneate,
petiole indistinct; venation obscure. Inflorescences
with separate sexes, to ca. 12 mm long, triseriate,
peduncles 4 to 5 mm, simple, fertile internode one.
Male inflorescences in secondary positions or low¬
est on lateral branches, to 10 mm long, flowers to
about 18 per fertile bract. Female inflorescence
mostly in primary axillary positions, flowers 4(6)
per fertile bract. Fruit 2X2 mm. globose, white,
petals spreading.
I he small-flowered species of Dendrophthora ,
especially in the Bolivian and Venezuelan Andes,
are in great need of revision, and the present spe¬
cies resembles a number of those that have already
been described. The known and similar species dif¬
fer from D. meridana in characteristics involving
basal cataphylls, number of flowers per fertile
bract, the number of fertile internodes per inflores¬
cence, in being short-tomentose, and in their male
and female flower distribution and leaf morphology.
I have scrutinized the species known so far (kuijt,
2000) and cannot match the present species with
any known one. A similar but not quite identical
distributional pattern of male and female inflores¬
cences has been noted for another Venezuelan spe¬
cies. I). cuneifolia. which may be closely related
but has a very different leaf shape and size, and
nearly twice as many female flowers per fertile
bract (kuijt. 1990).
The specific epithet refers to the Cordillera de
Merida, where both known specimens were col¬
lected. The holotype is a unieate.
Paratype. VENEZUELA. Merida: Sierra Nevada, near
Alto del Aguada, 3(X)-3500 m, /,. Bemardi et al. 171II
(LEA).
Volume 13, Number 1
2003
Kuijt
Mistletoe Notes
87
Figure 11. Dendrophthora meridana Kuijt. —a. Flabit. lateral branch with female inflorescences flanked by two male
inflorescences in prophyllar positions. —b. Young infructescence. Drawn from Riina et al. 623 (VEN).
88
Novon
47. Dcndrophtliora warmiiigfii (Eichler) Kuijt,
comb. nov. Basionym: Phoradendron warmin¬
gii Eichler, in Warming, Vidensk. Meddel.
Dansk. Naturhist. Foren. Kjoebenhavn. 209.
1870. TYPE: Brazil. Lagoa Santa, on Vochysia
elliptica, 14 Jan. 1865, E. Warming 383 (ho-
lotype, C).
Dendrophthora tepuiana (Steyermark) Kuijt, Proc. Kon.
Ned. Akad. Wetensch. 93: 138-139. 1990. Venezuela. Bo¬
livar: Ptari-tepui, 16(H) m. Steyermark 59688 (holotype, F;
isotype, US).
I have found, in the holotype of Dendrophthora
warmingii, two instances of an apparently female
flower with a small anther on one of the petals, as
happens very occasionally in Phoradendron and
Dendrophthora. 1 hese anthers, although underde¬
veloped, were almost certainly unilocular and, on
the basis of this character, I feel confident about
the assignment of this species to Dendrophthora. It
should be noted that Phoradendron rugulosum Ur¬
ban, which Rizzini (1991) recombined to Phora¬
dendron warmingii Eichler var. rugulosum (Urban)
Rizzini, is a synonym of P. pteroneuron Eichler
(Kuijt, 2003).
Acknowledgment. The work here reported was
done with financial support from the National Sci¬
ence and Engineering Research Council of Canada.
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-. 1991. N ovos taxons do Brasil nas Loranthaceae
e Rutaceae. Rev. Brasil. Biol. 51: 455-462.
Van Iieghem, P. 1895. Sur le groupement des espfeces en
genres dans les Loranthacdes k calice dialysdpale et
anthferes oscillantes ou Struthanth^es. Bull. Soc. Bot.
France 42: 161-180.
Venturelli, M. 1981. Embriologia de Struthanthus vulgaris
(Loranthaeeae-Loranthoideae). Kurtziana 14: 73-1 (H).
Dischidia cleistantha (Apocynaceae, Asclepiadoideae): A New
Philippine Endemic
Tatyana Livshultz
L.H. Bailey Hortorium, Department of Plant Sciences, 228 Plant Science Building, Cornell
University, Ithaca. New York 14853-5908, U.S.A. tl44@cornell.edu
ABSTRACT. Dischidia cleistantha , a new species
endemic to the Philippines, is described and illus¬
trated. It is distinguished from three other species
of Dischidia sect. Conchophyllum native to the Phil¬
ippines by the character combination of corolla
adaxially glabrous, 7—7.5 mm long, with enrol line
corona lobes opposite the corolla lobes, fleshy tri¬
angular dorsal staminal appendages, and unstalked
staminal corona lobes.
Key words: Apocynaceae, Asclepiadaceae, As¬
clepiadoideae, Conchophyllum , Dischidia, Philip¬
pines.
Ongoing studies of the genus Dischidia R. Brown
have revealed that a plant known in cultivation as
Dischidia “Thailand Red" belongs to a new species
endemic to the Philippines.
Materials and Methods
Specimens were observed with light and scan¬
ning electron microscopy (SEM). For SEM. flowers
that had been preserved in FA A and stored in 70%
EtOH were put through a dehydration series and
critical-point dried; anther margins and pollinaria
were removed from fresh flowers (vouchered by Liv¬
shultz 911— 6 . RH) and air-dried. All specimens ob¬
served with SEM were mounted on aluminum stubs
with two-sided tape and dissected. Stubs were coat¬
ed with gold-palladium and viewed with a Zeiss
960 SEM. All photography is digital. Figure 2C was
retouched to remove a small nick in the corolla
tube, 2D to remove a charge line, and Figure 2E
to remove cracks in the background; retouching did
not alter the morphology or micromorphology de¬
picted in the images. Distribution of sclerenchyma
in the stamens was determined by staining spirit-
preserved flowers with phloroglucinol-HCl.
Dischidia cleistantha Livshultz, sp. nov. T\ PE:
Philippines. Zamboanga Province: Mindanao.
Sep. 1922, E. I). Merrill 11639 (holotype, K;
isotypes. P. US). Figures 1, 2, 3A, C—D.
Species foliis patelliformibus (convexo-concavis) in-
structa, a speciebus aliis omnibus distincta tubo corollae
intus omnino glabro, lobis corollinis coronae lobis corollae
oppositis, corona staminali ex lobis sessilibus ancorifor-
rnibus sub antheras insertis formata, appendicibus stam-
inalibus dorsalibus ex lobis earnosis triangularibus con-
stantibus.
Plants succulent, herbaceous epiphytic vines,
root climbers and twiners, with white latex in all
parts (Fig. 1A), glabrous except for sparse hairs
near the margins of some younger leaves, lacking
wax chimneys on stems and leaves (Fig. 3A). Roots
on mature plants adventitious, always paired below
each node, additional roots occasionally produced
along internodes (Fig. 1 R). Sterns terete; diameter
1.5—3 mm: internodes 3.5—7.6 cm long. Stipular
colleters a minute pair, triangular or ovate, to 0.5
mm long. Leaves always opposite, decussate at first,
becoming distichous, peltate at maturity; petiole 1-
3 X 1—2 mm; lamina with indexed margins at first,
at maturity the adaxial surface convex, green, the
abaxial surface concave, purple (Fig. 1A. R), 1.3—
3.6 X 1.6—5 cm. transverse-ovate to reniform. apex
acute to obtuse, base truncate to slightly cordate,
bearing a small protuberance with 1 to 3 colleters
on the adaxial side (Fig. 1C); venation obscurely
raised adaxially, invisible abaxially, brochidodrom-
ous, with 3-4 pairs of secondaries (Fig. 1C). Inflo¬
rescence (Fig. ID) extra-axillary', umbelliform or
compound-umbelliform, persistent; peduncles 0.4—
2.0 X 0.15—0.2 cm, bearing 1—3(—4) stout rachises
at their apices; rachises 3—4 mm diam.. up to 1.4
cm long, with extremely short internodes, each ra-
chis bearing 6 to 12 flowers and buds at its tip
when actively growing; lower portions of rachises
covered with transversely elliptical scars where
flowers have fallen. Each flower subtended by two
membranous triangular to lunate bracts ca. 0.5 X
0.5 mm; the bracts becoming chartaceous, persis¬
tent after flowers fall, but wearing off from the old¬
est parts of the rachises; pedicels 3-4 X 1 mm. pale
green; sepals quincuncial. 1.5 X 1.5 mm, with a
minute ovate colleter within each sinus, ovate, apex
obtuse, pale green with hyaline margins, margins
ciliate or not; corolla tube 7—7.5 X 3.5 mm, suc¬
culent, urceolate, 5-ribbed, the rounded, shallow
Novon 13: 89-96. 2003.
90
Novon
DtiJjldv&i ol ‘
Volume 13, Number 1
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Livshultz
Dischidia cleistantha from the Philippines
91
ribs separated by valleys opposite the corolla lobe
sinuses (Fig. IE, F); corolla lobes valvate, 2 X i
mm, tips cucullate (Figs. 2A. 3C), connivent at an-
thesis, adaxial epidermal cells flattish tabular, or¬
namented with longitudinal striations (Pig. 2H) (ter¬
minology from Christensen & Hansen, 1998);
corolline corona of 5 prominent lunate wedges in¬
serted adaxially. 2.5 mm from the base of the co¬
rolla, opposite the lobes, inner surface of corolla
tube rugose below the corolline corona (Pig. 2C);
corolla at anthesis white suffused with purple, the
pigment most prominent on the valleys and on the
margins of the corolla lobes, pigment actually in
the subepidermal cells of the adaxial corolla sur¬
face, on the margins and tips of the corolla lobes,
the valleys between ridges (Fig. 3C), the corolline
corona lobes and the tube below the corolline co¬
rona lobes, entire flower becoming pale pink with
age. Gynostegium 2.5 X 2 mm, conic in outline,
almost sessile, basal (filament) tube 0.2 mm long
(P ig. 1G); anthers triangular, 2X1 mm; connec¬
tives dark purple with triangular, white, hyaline ap¬
pendages covering the style-head apex; each anther
margin with an inner and an outer edge (P ig. 2E);
the outer edges form an open “mouth" at the base
of the anther margins, revealing the inner edges
(Fig. 2D); staminal corona of five unstalked lobes
inserted at the bases of the anthers, each lobe with
two recurved arms at its apex, white (Fig. 21)); dor¬
sal staminal appendages of five fleshy, triangular
lobes inserted on the abaxial surface of the anthers
(Pigs. 1G, 2D, 3C), each lobe suffused with pale
purple pigment; sclerenchyma restricted to the an¬
ther margins, lateral margins of the dorsal staminal
appendages, and the distal and outer edges of the
arms of staminal corona lobes; pollinarium (Pigs.
2F, 3D) with pollinia acropetal to the corpusculum
when in situ, corpusculum narrowly elliptic, 230—
240 X 60—80 pun, apex obtuse, rounded to trun¬
cate; caudicles flattened, with a thin region forming
a channel along the outer edge (Fig. 2F), triangular
with a long, narrow “thumb" extending from the
apex of the outer edge, 300—330 (including thumb)
X 70—90 gm; pollinium dorsiventrally flattened,
narrow-obovate to elliptic, apex slightly truncate,
480—510 X 100—110 pirn; carpels 1.2 X 0.4 mm
diam., obclavate (Figs. IG, 3C), style-head obtur-
binate, 5-angled in cross section with 5 reflexed
lobes alternating with the stamens, 1.4 X 0.8 mm;
apex acuminate, 0.8 X ca. 0.2 mm (Figs. IG, 3C).
Fruit a single follicle developing from each flower,
pedicels 5—6 X ca. 1 mm, follicle 5.0—5.4 X 0.3—
0.5 cm diam., reflexed relative to the pedicel, lan¬
ceolate in outline, widest 3.5 cm from the apex,
gradually tapering toward both ends, apex obtuse,
base rounded, cross section terete or half-terete
(distorted by pressing), coriaceous and glabrous;
seeds attached to a white, papery placenta, about
20 per fruit, 3X1 mm, ovoid, obscurely winged
on the margin, testa smooth, glabrous; coma 2-2.5
cm long, white, silky.
Flowering specimens collected in January, Feb¬
ruary, August, September, November, and Decem¬
ber from wild populations; continuously flowering
under greenhouse conditions in Ithaca, New York.
Common names. The Tagalog name is reported
as “pag-ong-pag-ongan" ( Fox 9073). In cultivation,
this species is called '“'Dischidia ‘Thailand Red"' or
“J9. Thailand Blush"’.
Etymology. The specific epithet is derived from
the closed appearance at anthesis of flowers from
living plants.
Discussion
Dischidia cleistantha is one of several previously
undescribed shell-leaved species from the Philip¬
pines. Specimens of D. cleistantha have been iden¬
tified as D. purpurea Merrill, D. platyphylla Schle-
chter, and D. imbricata (Blume) Steudel on
herbarium labels. Merrill 8319 was listed among
the exsiecatae of D. platyphylla in the Enumeration
of Philippine Flowering Plants (Merrill. 1923). The
first two of these species are endemic to the Phil¬
ippines (Merrill, 1923), whereas D. imbricata is
widespread in Malesia (Rintz. 1980) but collected
only from Palawan in the Philippines (Eivshultz,
pers. obs.). Table 1 lists characters that can be used
to distinguish them. The glabrous flowers of D.
cleistantha easily distinguish it from D. platyphylla
and D. imbricata, which have dense hairs on the
bases (D. platyphylla) or tips and margins (D. im¬
bricata) of the adaxial side of the corolla lobes. Dis¬
chidia cleistantha has much larger flowers than D.
ed and buttressed at maturity, the
walls essentially straight when immature, forming a
wavy configuration at maturity; style base essen¬
tially absent, at most a depressed-pyramidal rem¬
nant of tissue or a low, 3-sided rim at achene apex,
0.1 mm long or less, 0.1—0.2 mm wide, brown; bris¬
tles 2—4, subulate, reddish, often represented only
by short rudiments that rarely reach to middle of
achene, the longer often with antrorse to divergent,
blunt- or round-tipped barbs distally, these often
reduced to small nubs proximally; embryo of the
Carex- type.
Distribution. I he genus contains two species
endemic to Mexico.
Etymology. Cypringlea is named in honor of
botanical explorer Cyrus Guernsey Pringle (1838—
1911), the first collector of C. analecta , who col¬
lected plants in Mexico between the years ± 1882
and 1907.
Key to the Genera of the Tribe Scirpeae
1. Perianth of scale-like tepals, persistent on the rachilla . Oreobolopsis T. Koyama & Guaglianone
1 . I erianth of bristles, often persistent at base of achene . 2
2(1). Bristles 10 to many, greatly elongating after anthesis and often forming white cottony masses around
o’ spikelets . ...... . Eriophorum I..
Z . Bristles 3 to 6 or rudimentary, short or reduced, rarely elongating and forming white or brown wooly
masses around spikelets. '
3(2). Bristles with spine-like, retrorsely set barbs. 4
Volume 13, Number 1
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Strong
New Genus of Cyperaceae
125
3 '. Bristles with anlrorsely or divergently set hairs or blunt barbs, rarely smooth, sometimes rudimen¬
tary .. • ^
4(3). Inflorescence compound, anthelate w ith elongate rays; spikelets numerous. Scirpus L.
4'. Inflorescence a simple head of 1 to few spikelets at the summit of the culm . 5
5(4). Inflorescence pseudolateral; style 2-branched. Amphiscirpus Oteng-Yeboah
5'. Inflorescence capitate; style 3-branched. Phylloscirpus C. B. Clarke
6(3). Leaf blades reduced, often with rounded or thickened tips; inflorescence typically
of a single spikelet; achenes beaked or shortly so; bristles well-developed ....
. Trichophorum Persoon
6 '. Leaf blades well-developed, attenuate to triquetrous apex; inflorescences anthelate
or paniculate; achenes unbeaked; bristles rudimentary. Cypringlea
Key to the Species of Cypringlea
1 . Leaf blades linear, (2-)3-7.2 mm wide, flattened
distally, often V-shaped or folded proximally, es¬
sentially straight, slightly curving only toward
apex; inflorescence a compound, umbel-like cor¬
ymb, 3.5-11 X 2-7 cm, the spikelets in small
fascicles or sometimes solitary at tips of shorter
secondary rays .1. C. analecta
I'. Leaf blades linear-filiform, wiry, 0.3-0.8 mm
wide, triangular-channeled, wavy and curling; in¬
florescence a single dense fascicle or cymose
panicle of 2—3 subcontiguous fascicles of spike-
lets, 1-3 X 1-2 cm. 2. C. coahuilensis
Description of the Species
I. Cypringlea analecta (Beetle) M. I. Strong,
comb. nov. Basionym: Scirpus analecta Beetle,
Brittonia 5(2): 148. 1944 (as “analecti ).
TYPE: Mexico. San Luis Potosf: Minas de San
Rafael. May 1911 (July 1911, UC #157273),
C. A. Purpus 5454 (holotype, NY; isotypes, F,
GH. MO, UC, US). Figures 2, 4.
Scirpus analecta var. evadens C. I). Adams, Ann. Mis¬
souri. Bot. Card. 78: 254. 1991. Syn. nov. TYPE: Mexico.
Chiapas: steep slope with Quercus and Pinus, 3 mi. S of
Aguacatenango along road to Pinola Las Rosas, Municipio
of Venustiano Carranza. 56(H) ft., 25 June 1965, I). E.
Breedlove 10570 (holotype, F #1679132).
Culms solitary’, often forming basal offshoots. 30—
70 cm tall, 1—2.3 mm wide, erect to ascending,
slender, triquetrous, firm but flexuous, coarsely
ribbed, green to pale green, the angles finely and
closely antrorsely scabrous, often smooth proximal¬
ly, the sheath-clad bases (4—)7—12 mm wide.
Leaves 5-14, 16-80 cm long, ascending, approxi¬
mate at or just above the base, often appearing sub-
verticillate, ecauline; sheaths short, herbaceous,
finely and distinctly veined, light green or often
stramineous, old sheath bases brown, often persis¬
tent and becoming fibrillose, the inner band with a
truncate to deeply U-shaped orifice; ligule a narrow,
thickened band of tissue; blades linear, (2—)3—7.2
mm wide, flattened distallv, often V-shaped or fold¬
ed proximally, finely veined and green both adaxi-
allv and abaxiallv, margins and abaxial midvein
finely and closely antrorsely scabrous, long-atten¬
uate to triquetrous apex. Inflorescence a compound,
umbel-like corymb, terminal, 3.5—11 X 2—7 cm;
involucral bracts several, the lowermost 1—2 leaf¬
like, elongated and often exceeding the inflores¬
cence, the uppermost linear-setaceous; rays as¬
cending to curving with age, flexuous, flattened or
compressed-trigonous, antrorsely scabrous on mar¬
gins distallv, smooth proximally. the spikelets often
fascicled or sometimes solitary at tips of shorter
secondary rays; spikelets 13—90 per inflorescence,
ovoid-ellipsoid or sometimes ellipsoid-obovoid
when immature, 5-7(-8) X 2-3 mm. with spread¬
ing scales, cuneate at base, acuminate at apex;
scales broadly rounded in cross section, dorsally
truncate or slightly furrowed longitudinally along
midcosta, thinly herbaceous, light yellow-brown to
yellow-brown, sometimes with brown sides, margins
narrowly scarious proximally, midcosta 3-nerved,
light green, thickening distally, prolonged beyond
the obtuse apex as a short mucro. lateral nerves
indistinct; fertile scales 10—20, ovate-elliptic to ob¬
long-ovate. 1.8-3 X 1-1.3 mm; sterile scales 2-4
at base (excluding 2-nerved prophyll), ovate, short¬
er than the fertile. Flowers loosely spirally arranged
or appearing 4-ranked, the rachilla slightly elon¬
gated between the flowers; anthers 1.5—2 mm long,
apiculate at apex, rounded with minute papillae at
base; style branches reddish, scaly-fimbriate,
equaling to 1/3 longer than unbranched portion.
Achene obovate to elliptic-obovate, 1.5-1.8 X 0.8-
l.l mm, rounded at apex, margins sometimes dis¬
tinctly costate proximally, indistinctly so distally.
Distribution. Endemic to Mexico. Occurs on
the western side of the Sierra Madre Oriental in the
states of San Luis Potosf and Hidalgo, south to
Puebla state in the Cerro de Paxtle, and southeast
to a disjunct population in the Sierra Madre de
Chiapas in Chiapas state.
Etymology. “Analecta” or “analects” is a plu¬
ral noun sometimes used in titles meaning extracts
or collections, typically from a particular classical
author. It also has a slightly pejorative sense of
126
Novon
Figure 2. Achenes of Cypringlea analecta (Beetle) M. T. Strong. —A, B. Achenes. —C. Detail of epidermal cells.
I). Detail of distal end ol bristle. —E. Apex showing detail of style base. From Pringle 3175 (US).
Volume 13, Number 1
2003
Strong
New Genus of Cyperaceae
127
Figure 3. Achenes of Cypringlea coahuilensis (Svenson) M. T. Strong. —A, B. Achenes. C. Detail of epidermal
cells. _D. Detail of distal ends of Bristles. —E. Apex showing detail of style base. From Johnston 8763 (LL).
inluuliml
128
Novon
ISOTYPE of: Sclrpus analecri Beetle
Brittonia 5(2): 148. 1944.
determined by Mark T. Strong, 2000
UNITED STATES NATIONAL HERBARIUM
^SiumotT^
I 157272 J
''v'JS/Ty 0 e
PLANTS MEXICAN*; PURPUSIAN*
SAN LUIS POTOSI
BR/vMDt.‘i£E HEfUkAKiUfc
<2 ft V 9W
Figure 4. Herbarium type specimen, Purpux 5454 (UC), isotype of Cypringlea analecta (Beetle) M. T. Strong.
Volume 13, Number 1
2003
Strong
New Genus of Cyperaceae
129
“crumbs” or “leftovers.” There seems to be no jus¬
tification for retaining “analecti” (as originally pub¬
lished by Beetle), and the usual form analecta
(noun in nominative plural) is here restored.
Cypringlea analecta was first named Scirpus po-
tosinus by Britton (1892: 77) in a checklist of North
American Cyperaceae. It was based on a Pringle
collection (3175) from San Luis Potosf, Mexico.
Britton gave the author as C. B. Clarke and cited
it as ined. It was later listed by Clarke (1908: 89)
in a manuscript published alter his death, as Scir¬
pus pot.asien.sis C. B. Clarke, citing the earlier ref¬
erence of Britton. Beetle (1944), noting that both
these proposed names were nomina nuda, proposed
the name Scirpus analecti and designated Purpus
5454 as the holotype. Beetle also indicated that two
other specimens he did not see belonged to this
species. Parry Palmer 905 and Arsene 2636,
brought to his attention by Hugh O’Neill, curator
of the Langlois Herbarium, Catholic University
(CU). However, these two specimens (both at US)
belong to another species, Karinia mexicana (Brit¬
ton) Reznicek & McVaugh. Cypringlea analecta has
continued to be treated as a species of Scirpus, most
recently by Adams (1994) in Flora Mesoamericana
and Espejo Serna and Lopez Ferrari (1997).
I have found no taxonomic significance to the
inflorescence of Scirpus analecta var. evadens C. D.
Adams in which the spikelets are solitary at ray
tips. The specimen Purpus 2889 (GH. US), from
Puebla, is intermediate between the fascicled
spikelets of the type material and that of S. var.
evadens in having both solitary and fascicled spike¬
lets at ray tips.
Representative specimens. MEXICO. Hidalgo: Jacala,
July 19,' 16. E. Lyonnet 1318 (US). Puebla: vicinity of San
Luis Tultitlanapa near Oaxaca, Cerro de Paxtle, rocky
slopes, July 1907, C. A. Purpus 2889 ((ill, US). San Luis
Potosf: moist banks of ravines, 15 Aug. 1890, C. G. Prin¬
gle 3175 (F, ISC. MU, NY. US).
2. Cypringlea coahuilensis (Svenson) M. T.
Strong, comb. nov. Basionym: Scirpus coahu¬
ilensis Svenson, J. Arnold Arbor. 25: 46. 1944.
TY PE: Mexico. Western Coahuila: high lime¬
stone ridge with a forest of Pinus pinceana ,
abundant on rocky pine-clad slopes, SW end
of the Sierra de la Fragua, 1-2 km N of Puerto
Colorado, 2 Sep. 1941. /. M. Johnston 8763
(holotype, Gil not seen; isotype, TEX—2).
Figures 3, 5.
Culms caespitose, 30—50 cm tall. 0.6—1 mm
wide, ascending, obtusely trigonous, wiry, stiff,
coarsely ribbed, smooth, pale green, the sheath-
clad bases 5—10 mm wide. Leaves 3 to 5, basal and
lower eauline, 10-35 cm long, ascending; sheaths
herbaceous to thickly herbaceous proximally, mem¬
branous distally, coarsely veined, brown proximally,
pale distally, the inner band with a deeply V-
shaped orifice, auriculate at junction with blade;
ligule a narrow band of lacerate tissue formed near
base of the auricles; blades linear-filiform, wiry,
0.3—0.8 mm wide, triangular-channeled, wavy and
curling, coarsely veined abaxially, smooth and V-
channeled adaxially, pale green both adaxially and
abaxially, margins closely antrorsely scabrous, tri¬
quetrous at apex. Inflorescence a single fascicle or
a cymose panicle of 2—3 subcontiguous fascicles of
spikelets at the summit of the culm, 1-3 X 1-2
cm; involucral bracts 2—3, linear-setaceous, 0.3—
0.5 mm wide, the lowermost 1—2 greatly exceeding
the inflorescence, 2—16 cm long; rays 1-2, short,
ascending, flexuous, obtusely trigonous, crescenti-
form or subterete in cross section, smooth on mar¬
gins; spikelets 3-9 per inflorescence, sublaterally
compressed, ovate-elliptic to elliptic, 7—10 X 2—3
mm, acute to acuminate on both ends, the scales
spreading with developing achenes; scales obtuse
to rounded or broadly rounded in cross section,
thinly herbaceous to submembranous, finely longi¬
tudinally cellular-striate, whitish- or stramineous-
translucent, glabrous, margins broadly scarious,
midcosta 3-nerved, pale green, prolonged beyond
the obtuse to subrounded apex as a short mucro,
lateral nerves indistinct; fertile scales 20-30, ovate
to widely ovate or ovate-elliptic, 2.8—4 X 1.6—2.6
mm, the prophyll at base of spikelet well-devel¬
oped, 2-nerved, about same size as fertile scales;
sterile scales 0-1 at base (excluding 2-nerved pro¬
phyll), ovate, slightly shorter than the fertile. Flow¬
ers spirally arranged or sometimes subdistichous
proximally on rachilla; anthers 1.7—2.2 mm long,
rounded-apiculate with minute papillae at apex,
short-sagittate with minute papillae at tips of lobes;
style branches scaly-fimbriate, equaling to slightly
shorter than unbranched portion of style. At hene
obovate to broadly obovate, 1.6—2 X 1.1—1.5 mm,
obtuse to subrounded at apex, margins indistinctly
costate.
Distribution. Endemic to Mexico and known
only from the type collection from a high limestone
ridge in the southwestern end of the Sierra de la
Fragua, western Coahuila.
Discussion
These two plants representing the new genus,
first described as Scirpus L., and like Scirpus (today
treated by many in a strict sense with the conserved
130
Novon
Scirpus coahuilensis Svenson
Jour. Arnold Arb. 25: 46. 1944.
ISOTYPE
MEXICO: western Coahuila
I. M. Johnston, no. 8?63 Sept. S, 1941
Scirpus coahuilsnsis Svenson, n. sp.
Abundant on rocky pine-clad slopes.
Coarse tufts 1-3 ft. tall*
Southwestern end qf the Sierra de la Fragva, 1-2
km. north of Puerto Colorado; a high limestone
ridge with a forest of Pinus Pinceana.
Figure 5. Herbarium type specimen, Johnston 8763 (LL), isotype of Cypringlea coahuilensis (Svenson) M. T. Strong.
Volume 13, Number 1
2003
Strong
New Genus of Cyperaceae
131
type, Scirpus sylvaticus L.), are members of the sub¬
family Cyperoideae Sussengutli following the most
recent Cyperaceae classification of Goetghebeur
(1998). The spirally arranged scales, embryo type,
morphological and epidermal structure of the
achene, and presence of persistent perianth bristles
places them in the tribe Scirpeae Kunth ex Du-
mortier. Scirpus sensu stricto is characterized by
having anthelate inflorescences, a Fimbristylis- type
embryo, spikelets with all scales fertile, beaked
achenes, and generally well-developed bristles with
sharply tipped retrorse barbs. Cypringlea differs
from it in having a Carex- type embryo (Fig. 1), 1
to 3 of the basal scales of the spikelet sterile (emp¬
ty), unbeaked achenes, and rudimentary bristles
that bear what appear to be blunt, rudimentary an-
trorsely or divergently set hairs or barbs at their
tips (Figs. 2D. 3D).
The embryos of Cypringlea (Fig. 1) are of the
Carex- type (Van der Veken. 1965; Goetghebeur,
1986) and are closest to its nearest relative, Tri-
chophorurn Persoon. Cypringlea differs from Tricho-
phorum in having well-developed leaf blades and
inflorescence bracts that are attenuate to a trique¬
trous apex; anthelate or paniculate inflorescences
with capitate or fascicled spikelets; achenes (Figs.
2A, 28. 2E; 3A, 3B. 3E) that are unbeaked, the
disarticulation of the style occurring at the apex of
the achene, bordered at most by a low, marginal
rim (Figs. 2F, 3E); and poorly developed bristles
(Figs. 2D. 3D) that bear rudimentary antrorse or
divergent hairs or barbs distally that are blunt or
rounded at their tips, or reduced to tiny nubs prox-
imallv. On the other hand. Trichophorum has re¬
duced leaf blades and inflorescence bracts that are
generally blunt and obtuse or rounded and often
thickened at their apices; inflorescences of a single
spikelet. rarely anthelate and never with spikelets
fascicled or capitate; beaked or shortly beaked
achenes, the style disarticulating at the apex of the
beak; and well-developed perianth bristles that are
132
Novon
densely beset with antrorse or divergent fine hairs
or harhs.
The achene pericarp cells of Cypringlea are iso-
diametric and 6-sided (Figs. 2C, 3C). When im¬
mature, they have a low central nodule and the
walls are essentially non-wavy and markedly raised
above the surface, forming a reticulation. At ma¬
turity, the walls are of a wavy configuration and
each has one central conical body with well-devel¬
oped buttresses between the walls and the body. In
species of Trichophorum, the cell walls of the peri¬
carp are generally non-wavy and lack buttresses
between the walls and apical nodules (Schuyler,
1971).
Species of Trichophorum are holarctic anti gen¬
erally occur in wet or inundated alpine, tundra, or
acidic bog habitats, rarely in damp or somewhat
dry, mesic habitats in woodlands where, for exam¬
ple, the North American species T. planifolium
(Sprengel) Palla occurs. Cypringlea species occur
on dry, steep, rocky slopes and ravines, often on
limestone substrate with pine and oak. 'Hie com¬
bination of a strikingly different habitat, well-de¬
veloped leaf blades that are attenuate at their api¬
ces, anthelate or paniculate inflorescences,
spikelets often fascicled or in capitate clusters, and
rudimentary bristles, separates Cypringlea from Tri¬
chophorum.
Cypringlea is endemic to Mexico. It occurs from
the Sierra de la Fragua (western side of the Sierra
Madre Oriental mountain range) in western Coa-
luiila state (type locality of C. coahuilensis ) in the
north, south to the western side of the Sierra Madre
Oriental in the states of San Luis Potosf, Hidalgo,
and Puebla (Cerro de Paxtle), and southeast to
Chiapas state in the easternmost extent of the Sierra
Madre de Chiapas (Fig. 6). Both species occur on
steep, rocky slopes and ravines, often on limestone
substrate with pine and oak.
Acknowledgments. I thank Dan Nicolson (US)
for reviewing the manuscript and providing the in¬
formation on the etymology of the epithet analecta ;
the National Museum of Natural History Scanning
Electron Microscopy Lab for use of their facilities;
and curators at F, GH. ISC, LL, UC, VI. and W for
searching their collections and loaning specimens.
Literature Cited
Adams, C. 1). 1994. Scirpus (Cyperaeeae). Pp. 448-449,
in: G. Davidse, M. Sousa S. & A. O. Chater (editors),
Flora Mesoamericana, Vol. 6, Alismataceae a Cypera-
ceae. Universidad Nacional Autdnoma de Mexico D.F.;
Missouri Botanical Garden, St. Louis; The Natural His¬
tory Museum, London.
Beetle, A. A. 1944. A known but undescribed Mexican
Scirpus. Brittonia 5(2): 148.
Britton, N. L. 1892. A list of species of the genera Scirpus
and Rhynchospora occurring in North America. Trans.
New York Acad. Sci. II: 74—93.
Clarke, C. B. 1908. New genera and species of Cypera-
ceae. Bull. Misc. Inform. Addit. Ser. 8: 1-196.
Espejo Serna, A. & A. B. Lopez Ferrari. 1997. Las Mon-
ocotiledbneas Mexicanas. Una Sinopsis Florfstica. 1.
Lista de Referencia, Parte V. Cyperaeeae. Consejo Na¬
cional de la Flora de Mexico, A.C., Universidad Autdn-
oma Metropolitana, y Consejo Nacional para el Cono-
cimiento y Uso de la Biodiversidad. Mexico, D.F.
Goetghebeur, P. 1986. Genera Cyperacearum. Fen bijdr-
age tot de Kennis van de Morphologie, Systematiek en
Fylogenese van de Cyperaceae-Genera. Unpublished
Dr. Sci. Thesis, State University, Gent, Belgium.
-. 1998. Cyperaeeae. Pp. 141-190 in: K. Kubitzki
(editor). The Families and Genera of Flowering Plants,
Vol. 4. Springer-Verlag, Berlin, Heidelberg.
Parker, K. J. 1976. Freezing in the preparation of cleared
plant material. Stain Technol. 51: 61—62.
Schuyler, A. E. 1971. Scanning electron microscopy of
achene epidermis in species of Scirpus (Cyperaeeae)
and related genera. Proc. Acad. Nat. Sci. Philadelphia
123: 29-52.
Van der Veken, P. 1965. Contribution a l’embryographie
systematique des Cyperaeeae—Cyperoideae. Bull. Jard.
Bot. Flat 35: 285-354.
Wilson, K. L. 1989. Proposal to conserve 468 Scirpus L.
(Cyperaeeae) with S. syivaticus L. as type. Taxon 38:
316-320.
Two New Winged Species of Symbolanthus
(Gentianaceae: Helieae) from Colombia
Lena Struwe
Department of Ecology, Evolution, and Natural Resources, Rutgers University—Cook College,
237 Foran Hall, 59 Dudley Road, New Brunswick, New Jersey 08901—8551, U.S.A.
struwe@aesop.rutgers.edu
Abstract. Two new r Colombian species were
identified in the Neotropical genus Symbolanthus
(Gentianaceae: Helieae) during an ongoing revi¬
sionary study. Symbolanthus pterocalyx Struwe is
characterized by its green to yellow corollas, strong¬
ly winged stems, and long-acuminate calyx lobes
with a long, thin dorsal keel on each lobe. The
other new’ species, Symbolanthus tetrapterus Stru¬
we, also has strongly winged stems, but corollas
variously colored in green, white, pink, to red-violet
and calyx lobes rounded and non-keeled. Both spe¬
cies are endemic to small areas in the Andes of
western Colombia. The western outlier Cerro del
Torra of the Cordillera Occidental in the province
of Choco (Colombia) is the only known locality of
Symbolanthus tetrapterus. Symbolanthus pterocalyx
occurs in high-elevation cloud forests, subparamo,
and paramo in a small area around Medellin in the
province of Antioquia (Colombia).
Rksumen. Se describen dos especies nuevas del
genero neotropical Symbolanthus (Gentianaceae:
Hel ieae). Symbolanthus pterocalyx Struwe se car-
acteriza por las corollas verdes o amarillas, los tal-
los fuertamente alados, y por los lobulos del caliz
largos, acuminados y con una quilla dorsal larga y
delgada sobre cada lobulo. La otra especie nueva,
Symbolanthus tetrapterus Struwe, tiene tambien tal-
los fuertemente alados y las corolas varfan desde
verdes o blancas hasta rosadas o rojo-violetas, pero
los lobulos del caliz son redondos y no quillados.
Las dos especies son endemicas de areas restrin-
gidas en los Andes occidentales de Colombia. Sym¬
bolanthus tetrapterus se ha registrado solamente en
el Cerro del Torra, saliente de la Cordillera Occi¬
dental en el departamento del Choco (Colombia).
Symbolanthus pterocalyx se eneuentra en bosques
nublados de elevaciones altas, subparamos, y pa¬
ramos cerca de Medellin, en el departamento de
Antioquia (Colombia).
Key words: Colombia, Gentianaceae, Helieae,
Neotropics, Sym bo la nth us.
Symbolanthus G. Don is one of the most colorful
and large-flowered genera of tropical American
Gentianaceae. It is a member of tribe Helieae, a
group that also includes Chelonanthus (Grisebach)
Gilg, Irlbachia Martius, Macrocarpaea (Grisebach)
Gilg, and Tachia Aublet (Struwe et al., 2002). Sym¬
bolanthus has the broadly bilamellate stigmas,
curved stamens, pollen in tetrads, slightly zygo-
morphic flowers, and woody habit that are charac¬
teristic ol this tribe. Its position in the Helieae is
also supported by molecular data from several
sources.
The plants of Symbolanthus are shrubby, some¬
times small trees, and less often herbaceous. The
flowers are 4—10 cm long, and two main types of
corollas occur in the genus. The first type are sal¬
ver-shaped and brightly pink, red, or purple corol¬
las, often with white nectar guides on the lobes.
The second type is more broadly shaped, with co¬
rollas that are basically white, yellow, to green. In
most recent Andean floristic treatments one poly¬
morphic species, Symbolanthus calygonus (Ruiz &
Pavon) Grisebach, has been recognized (e.g., Mon-
tufar, 2000; Pringle, 1995). During the last de¬
cades, dozens of previously described species (e.g.,
Grisebach, 1849; Gilg, 1906) have been recognized
as synonyms under Symbolanthus calygonus follow¬
ing an unpublished revision by Noor van Heusden
(Utrecht University). For her revision, she annotat¬
ed all investigated Symbolanthus herbarium mate¬
rial with the name Symbolanthus calygonus (except
for two highly distinct and narrowly endemic spe¬
cies). In herbaria and Andean floras, this view has
been followed until now. Such a broadly circum¬
scribed Symbolanthus calygonus included widely
diverse calyx and corolla morphology traits. Im¬
mediately apparent is the corolla color, which in
Symbolanthus calygonus sensu lato varies from
deep magenta and red through pink to white, yel¬
low, and green. Using this broad circumscription,
Symbolanthus calygonus also had a distribution
that covered all of the Andes from Bolivia in the
south to Costa Rica in the north.
Novon 13: 133-140. 2003.
134
Novon
Van Heusden included the handful of Symbolan¬
thus species of the Guayana Highlands in her
broadly circumscribed species concept, but this
view lias never been followed in florist it* works of
that area (cf. Maguire & Boom, 1989; Struwe et al.,
1999). Similarly, the Central American material has
been consistently treated as a distinct species, e.g.,
Symbolanthus pulcherrimus Gilg (Standley, 1938;
Elias & Robyns, 1975).
An ongoing revision of Symbolanthus by the au¬
thor reveals that Symbolanthus calygonus is a spe¬
cies that is restricted to the Huanuco region in Peru
and is characterized by having a thick “hump-like”
dorsal keel on each calyx lobe. This suggests that
many Andean species face resurrection, e.g., Sym¬
bolanthus daturoides (Grisebach ex Hooker) Gilg.
S. magnificus Gilg, and 5. vasculosus (Grisebach)
Gilg. all of which have distinct morphological char¬
acters that can be used for field recognition. These
investigations also indicate several species remain¬
ing to be described, especially from Bolivia and
Peru. The number of species estimated in Symbo¬
lanthus totals at least 30, including two species of
the genus Wurdackanthus Maguire, which is now
synonymized under Symbolanthus (Gould & Stru¬
we, in prep.; Struwe & Gould, in prep.).
This paper describes two new species from Go-
lombia that are distinctly different morphologically
from Symbolanthus calygonus. Significantly, these
more narrowly circumscribed species not only dif¬
fer in leaf and floral morphologies from Symbolan¬
thus calygonus sensu stricto, but the color scheme
of the corolla is also more consistent within each
species (i.e., each species has either the red-pink
scheme or the white-yellow-green scheme). Some
individuals have a mixture of corolla colors, such
as a white tube on a red corolla or a pink tinge on
a white corolla. As discussed below, Symbolanthus
tetrapterus is exceptional because it has corollas
varying in color from red-purple, through pink, to
white anti green. In contrast, Symbolanthus ptero-
calyx has a light green to yellow corolla (rarely pale
pink or with pink markings).
The two new species described here have in
common strongly quadrangular and 4-winged
stems. In Symbolanthus tetrapterus, the calyx lobes
are acute anti non-alate. Symbolanthus pterocalyx
differs in having long-acuminate calyx lobes with a
long and thin hyaline keel, traits that are unique
to this latter species. Several non-winged species
of Symbolanthus are previously known from Colom¬
bia ami are currently being revised (Struwe, in
prep.).
Material and Methods
Herbarium material from HE A, K, MO, NY, U,
and US was studied. Selected flowers from herbar¬
ium material were rehydrated in a solution ol 70%
ethanol, glycerol, and 5% Aerosol OT (9:1:1) for at
least 24 hours.
TaXONOM tc Tn E A I M ENT
Symbolanthus pterocalyx Struwe, sp. nov.
TYPE: Colombia. Antioquia: Santa Elena,
1500-2000 m, shrub 15 ft., fl. pale yellow-
green, 1 Jan. 1930, IT. ,4. Archer 1287 (holo-
type, NY; isotypes, K, US). Eigure 1.
Haec species Symbolantho calygono siniilis, sed ah ea
caulibus manifeste alatis, lobis ealycis longioribus longe-
apieulatis manifeste alatis, tubo corollae ealyce 1.0—1.25-
plo longiore, et corollis albis, flavis vel viridis differt.
Shrub, branched, 0.6—4 m tall. Branches qua¬
drangular, with interpetiolar lines; flowering stems
3—11 mm diam., internodes 1.2-4.8 cm long,
winged; wing 1—3 mm wide. Leaves widely elliptic,
(1.5—)2.0-2.5X longer than wide, broadest at the
middle to slightly above, short- to long-petiolate,
membranaceous when dry; lamina 5.5—16.2 X 1.6-
6.5 cm; leaf apex acute to apiculate; leaf margin
flat; leaf base long-attenuate; venation visible, 2
prominent pairs of basally divergent secondary
veins; petiole 0.5—1.5(—2.0) cm long. Inflorescence
I - to 4(to 9)-flowered; subtending bracts and brac-
teoles linear to narrowly triangular, acute to acu¬
minate, with dorsal keel, 9—12 X 2—3 mm at base;
pedicel 26-42 mm long at anthesis, upper part of
pedicel often keeled or 5-angular by decurrent ca¬
lyx wings, pedicel 2-4 mm diam., transition from
pedicel to flower gradual; flowers erect at anthesis.
Calyx divided to ca. 0.8 of its length, at anthesis
(2.6—)3.2-5.0 X (1.5—)2.0-2.9 cm; lobes narrowly
ovate, (2.3-)2.8-4.0 X 1.1-1.7 cm, erect, but not
tightly appressed to the corolla base, with a large,
long, thin, often hyaline keel, 3—5 mm high, usually
decurrent into the peduncle; apex long-apiculate
(rarely acute); margin broadly hyaline; colleters
(fingershaped, multicellular glands) present on
large area on the inside of the base of each calyx
lobe. Corolla broadly funnel-shaped, slightly zygo-
morphic with two lower lobes recurved, (5.0—)5.5—
6.8(—7.8) cm long, basic color greenish white, pale
yellow, yellow, to green (rarely pale pink), tube ex¬
ternally sometimes with reddish or pink lines, thin
when dry; tube broadly widened toward mouth,
straight, 1.0— 1.25X longer than calyx, 3.4—5.0 cm
long, 0.8—1.5 cm w ide below insertion of stamens.
Volume 13, Number 1
2003
Struwe
Symbolanthus from Colombia
135
Figure 1. Symbolanthus pterocalyx Struwe (Archer 1287, K holotype, NY isotype: A—C; Zarucchi & Cardenas 4322,
NY paratype: D—G). —A. Habit of flowering stem. —B. Close-up of winged stem and interpetiolar lines.
—C. Flower bud showing calyx lobes with dorsal ridges decurrent onto the pedicel and comparatively large, linear
bracts. —D. Opened calyx with colleters (glands) on internal surface. —E. Opened corolla (partial) with stamens and
corona. —F. Gynoecium with glandular disk at base of ovary. —G. Inside of corolla tube with corona and base of
capsule, erect (rarely slightly nodding), broadly el¬
liptic, 2.4-3.1 X 1.4-2.1 mm, same length or short¬
er than calyx, green, dehiscent, persistent basal
part of style 8-23 mm long; calyx lobes 26—50 mm
long in fruit; seeds angular, brown, ca. 1 mm.
Vernacular names. Suspiro {Lehmann 5449), Ta-
baquillo pequeno (Giraldo et al. 1.399).
Phenology. Flowering specimens have been
collected in all months of the year except October
and December; fruiting specimens have been found
during January to April and August to October.
Habitat and distribution. Symbolanthus ptero¬
calyx only occurs in high-altitude cloud forests,
subparamo, and paramo habitats in the province of
filaments.
(1.8—)2.6—3.0 cm wide at mouth; lobes broadly
ovate, 2.0-3.0 X 2.3—2.8 cm, (1.0—) 1.2—1.5 X lon¬
ger than wide, spreading to slightly reflexed at un¬
thesis, apex abruptly (long-)apiculate; corolla apex
in bud tapering. Stamens inserted ca. 7—15 mm
from the base of the corolla; corona well-developed,
ca. 10 mm tall, as a coronate ring with 5 plicae,
basally fused with filaments, free part ca. 5—6 mm
tall, with small pockets behind corona, plicae rect¬
angular and 5—6 X ca. 4 mm, upper edge fimbriate;
filaments inserted into anther at ca. 1/3—1/4 from
base of anther; anthers white to cream (young),
brown (older), ca. 6—9 mm long. Ovary ca. 6—7 mm
long; style 28—35 mm long; stigma lobes oblong,
green to white, 3-4 mm long. Fruit a fleshy-walled
136
Novon
80° W 70° W
Figure 2. Distribution map for Symbolanthus pterocalyx
and S. tetrapterus.
Antioquia, Colombia (Fig. 2). It has been found at
Cerro del Padre Amaya, Paramo “El Mono,” Pa¬
ramo Sabanazo, Cerro San F elix, and Cerro Marcon
in the Santa Helena area at altitudes between 1500
and 3200 m. Several collectors noted on herbarium
labels that this is a common plant in these areas.
It also grows in disturbed areas and is noted to
recover degraded soils (vidi Giraldo et al. 1399 :
“USO: Pro[te]ctor de fuentes de agua, recuperador
tie suelos”).
Etymology. This species is named pterocalyx
(= winged calyx) after its prominent wings on the
calyx.
Paratypes. COLOMBIA. Antioquia: Mun. Bello, cor-
regimiento de San Felix, 5 June 1989, Echeverri & Con¬
treras 86 (HUA); Corregimiento San Felix (Torres de Te-
lecomunicaciones), 22 May 1991, Fonnegra & Curso
Taxononua Vegetal 3733 (HUA, U); Cerro San Felix, 11
Mar. 1992, Fonnegra & Grupo Tax. Plan. Vase. I Sern/92
3979 (HUA); Mun. de Caldas, Vereda La Corrala, FINCA
"La Zarza.” 28 Apr. 1986, de Escobar & Giraldo 6649
(HUA); Mun. de Belmira, Pdramo “El Morro,” Proyeeto
Heserva Natural “El Pdramo de Belmira,” 22 Apr. 1993,
Fonnegra & Tuberquia 4637 (HUA, MO, U); Paramo Sa¬
banazo, Estacion Ecoldgica “El Refugio,” 14 May 1996,
Fonnegra et al. 5984 (HUA); Mun. Buriticd, Corregimento
El Cuarco, Bosque Andino, 12 Mar. 2000, Molina et al.
431 (HUA); Mun. de Envigado, borde de carretera y di-
vorcio de aguas, 16 Apr. 1996, Correa et al. 334 (HUA,
JAUM not seen; det. here as cf. pterocalyx), 18 Apr. 1996,
Correa et al. 515 (HUA, J AUM not seen); recorrido Ionite
Envigado, Caldas y El Retiro haeia el Oriente siguiendo
el divorcio de aguas, 25 Apr. 1996, Correa et al. 712
(HUA, JAUM not seen); Mun. Frontino, Corregimiento
Nutibara, Vereda Alto de Cuevas, Quebrada del Oso, 23
Apr. 1991, Fonnegra & Curso Taxonomta Vegetal 3536
(HUA, U); euenea alia del Rfo Cuevas, 13 Jan. 1987,
Sanchez et al. 894 (HUA, MEDEL not seen), 13 July
1987, Sanchez et al. 1366 (HUA. MEDEL not seen): Mun.
de La Estrella, vereda El Romeral, sitio La Laguna, 17
Aug. 1992, Giraldo-Canas et al. 1085 (HUA); Mun. La
Union, Corregimiento San Miguel, carretera La Union-
Sonson, borde de Canada, cerea al Rio Pietlras, 5 July
1987. Marulanda et al. 304 (HUA); Mun. Medellin. Bo-
querdn, Cerro del Padre Amaya, Feb. 1984, Agudelo &
Betancur 32 (HUA); Corregimiento San Cristobal, Ver. La
Palma, 1 Apr. 1997, Giraldo et al. 1399 (HUA, JAUM not
seen); bosque de subparamo, antena de INRAVISION, 2
Sep. 1981. Hoyos et al. 112 (HUA); ca. 19 km W of Me¬
dellin (rd. to Antioquia), cloud forests 5—10 km from high¬
way along road to TV towers, 25 Mar. 1979, Luteyn &
Lebrdn-Luteyn 7081 (HUA, NY); Mililar, 15 May 1982,
Moreno & Buitrago 21 (HUA), 4 Aug. 1979, Palacios et
al. 85 (HUA); 11 km from Medellin—Santa Fe de Antio¬
quia highway on road to summit, 18 Mar. 1987, Zarucchi
& Madrigal 4853 (HUA, MO, U); along road to Cerro del
Padre Amaya, 10 km from Medellin—Santa Fe de Antio¬
quia road, 15 Sep. 1987, Zarucchi A Brant 5341 (HUA,
MO), 4 Jan. 1969, Uribe Uribe 6225 (US); Mun. de Santa
Rosa, carretera haeia Aragon 5 km, 2 Oct. 1982, Curso
Fcologia Vegetal, Universidad de Antioquia s.ri. (HUA);
Mun. de Yarumal, Wachst in dichten laubwaldern um Ya-
rumal, Nov. 1891, Lehmann 5449 (K); N of Yarumal on
road to Cerro Marconi (telecommunication towers for
town), S slope near peak, 29 Nov. 1986, Zarucchi & Car¬
denas 4322 (HUA, MO, NY, U); unknown municipality,
Santa Helena, forest bond., 9 Jan. 1880, Kalbreyer 1304
(K); Rionegro, 24 Jan. 1992, Vargas 790 (HUA).
Symholanthiis tetrapterus Struwe, sp. nov.
TYPE: Colombia. Choeo: Mun. San Jose del
Palmar, Cerro del Torra, cumbre del Torra
Grande, 2770 m. 13 Jan. 1984, F. A. Silver-
stone-Sopkin, TV. Paz, A. Duque & H. Bayona
1803 (holotype, U; isotypes, CUVC, MO). Fig¬
ure 3.
Haec species Symbolantho calygono similis, sed ab ea
caulibus manifeste alatis, foliis obovatis vel obovatis-ob-
longis sessilis (raro subsessilis), et calycibus non alatis
differt.
Shrub, branched, 1—4 m tall. Branches quadran¬
gular, with interpetiolar lines; flowering stems 5—8
mm diam.; internodes 11—39 mm long, winged,
wing broader toward upper part of each internode,
often with a wavy to undulating edge; wing (0.5—)
1.5—3.0 mm wide. Leaves obovate or obovate-ob-
long, 2.0—2.5X longer than wide, broadest above
middle, sessile (seldom subsessile), membrana¬
ceous to slightly coriaceous when dry; lamina 3.2—
12.9 X 1.3—6.3 cm; leaf apex rounding to acute
apex; leaf margin flat; leaf base cuneate; venation
visible, 2 pairs of basally divergent secondary
veins; petiole 0(—5) mm long. Inflorescence 1- to 4-
flowered; subtending bracts and bracteoles trian¬
gular. acute, membranaceous, 2—3 mm long, ca. 3
mm wide at base; pedicel 18-29 mm long at an-
thesis, terete, 2—3 mm diam.. transition from ped¬
icel to flower gradual; flowers erect at anthesis. Ca¬
lyx divided to ca. 0.9 of its length, at anthesis 2.2—
2.8 X 1.3—1.5 cm; lobes narrowly ovate. 1.3—2.5 X
0.8—1.0 cm, appressed (but not tightly) to the co¬
rolla base, without dorsal keel, but dorsallv thick-
Volume 13, Number 1
2003
Struwe
Symbolanthus from Colombia
137
Figure 3. Symbolanthus tetrapterus Struwe (, Silverstone-Sopkin et al. 1803 , MO isotype: A, B, F; Silverstone-Sopkin et
al. 4140, U paratype: C-E). —A. Habit of flowering stem, with close-up of erase corolla lobe apex at right. —B. Close-
up of winged stem and interpetiolar lines. —C. Flower before anthesis showing non-keeled calyx lobes. —D. Inside of
corolla with corona and anthers. —E. Gynoecium (note glandular base of ovary). —F. Close-up of branch with young
fruits.
ened; apex acute, mucronate; margin very broadly
hyaline; colleters present as a glandular colleter
area on the inside of the hase of each calyx lobe.
Corolla narrowly funnel-shaped, actinomorphic,
6.2-6.8 cm long, basic color white, green, pink
(less often dark pink to red-purple), sometimes also
the corolla tube white internally, green externally,
and corolla lobes showy dark pink (vidi Silverstone-
Sopkin et al. 4140), thin when dry; tube cylindric
to slightly widened toward mouth, straight. 2.0-
2.5X longer than calyx, 4.8—5.2 cm long, ca. 0.7
cm wide below insertion of stamens, 1.2-2.2 cm
wide at mouth; lobes broadly ovate, 1.5—2.0 X 1.5—
2.0 cm, 1.0—1.3 X times longer than wide, erect to
spreading at anthesis, apex obtuse, with a small
apiculate tip; corolla apex in bud tapering. Stamens
inserted ca. 10-15 mm from the base of the corolla;
corona well-developed, ca. 10 mm tall, as a coro¬
nate ring with fused upper edge, with small pockets
between corona and corolla, fused with filaments
for 5 mm, upper portion (5-6 mm tall) free from
the filaments, corona edge uneven, papillate; fila¬
ments inserted into anther ca. 1/4 from base of an¬
ther; anthers white, 6—7 mm long. Ovary ca. 6—7
mm long; style ca. 2 cm long; stigma lobes oblong,
ca. 3 mm long. Fruit a fleshy-walled capsule, nod¬
ding, globose to elliptic, 2.0—3.4 X 1.8—2.4 cm
wide, same length as calyx, green, probably dehis¬
cent, persistent basal part of style 2-6(-22) mm
long; calyx lobes 2.1-3.0 cm long in fruit; seeds
not seen.
Phenology. Flowering and fruiting specimens
have been collected in January and August.
138
Novon
Table 1. Comparison
of the two new species Symbolanthus
pterocalyx and .S’, tetrapterus
with S. calygonus.
S. calygonus
S. pterocalyx
S. tetrapterus
W ings on stem
not or barely winged
prominently winged
prominently winged
Leaf shape
elliptic-lanceolate
widely elliptic
obovate to obovate-ob-
long
Leaves petiolate vs. ses¬
sile
distinctly petiolate leaves
distinctly petiolate
leaves
sessile (rarely subses-
sile)
Leaf base
cuneate
cuneate
strongly attenuate
Bract length
4—10 mm
10—12 mm
2—3 mm
Calyx lobe length
1.5—2.4 cm
2.8—4.0 cm
1.3-2.5 cm
Calyx lobe apex
obtuse to slightly acute
long-apiculate
acute, mucronate
Calyx wings
short “hump-like” keel
long thin wing on the
back of each calyx
lobe, often decurrent
wing absent
Corolla tube length vs.
calyx length
2—3X longer
I.0-1.25X longer
2.0—2.5X longer
Corolla color
pink to red
white, yellow, to green
(rarely pale pink)
green, white, pink, or
red-violet corollas
Distribution
Peru; Huanuco
Colombia: Antioquia
Colombia: Choco
Habitat and distribution. Symbolanthus tetrap-
terus is known only from Cerro del Torra in south¬
ern Choc6 in Colombia (Fig. 2). It grows in several
habitats on this mountain at ea. 1900—2800 m al¬
titude, from disturbed shrubby vegetation, primary
forests, to open paramo vegetation summit domi¬
nated by A 'eurolepis Meissner, Paepalanthus Kunth,
Blechnum P. Browne, and Rubiaceae and Melas-
tomataceae shrubs anti very few dwarf trees, but
devoid of Espeletia Mutis ex Bonpland (Silverstone-
Sopkin & Ramos-Perez, 1995). This type of para¬
mo-like vegetation usually occurs at a higher alti¬
tude in the Andes, and its occurrence on Cerro del
Torra is probably due to wind exposure (Silver-
stone-Sopkin & Ramos-Perez, 1995).
Etymology. This species epithet is derived
from tetrapterus (or 4 wings) after its four prominent
and decurrent wings along the stems.
Paratypes. COLOMBIA. Choco: Municipio San Just'*
del Palmar, Cerro del Torra, vertiente nordeste, 8 Aug.
1982, Silverstone-Sopkin 1260 (CIJVC, MO); rumble, !!
Aug. 1982, Silverstone-Sophin 1268 (CUVC, MO); vertien¬
te nororiental, arriba del helipuerto, 6 Jan. 1984. Silver-
stone-Sopkin et al. 1587 (CUVC, MO. U); cumbre del Tor¬
ra Grande, 13 Jan. 1984, Silverstone-Sopkin et al. 1804
(CUVC, MO); vertiente oriental, helipuerto, 6 Aug. 1988.
Silverstone-Sopkin et al. 4140 (CUVC 2 sheets, NY, U):
filo de cumbre, 15 Aug. 1988. Silverstone-Sopkin et al.
4421 (CUVC, U), 15 Aug. 1988, Silverstone-Sopkin et al.
4422 (CUVC, U), 21 Aug. 1988, Silverstone-Sopkin et al.
4664 (CUVC, U).
Discussion
Symbolanthus pterocalyx differs from S. calygon-
us in several notable characters (Table 1). These
include prominently winged stems (vs. not or barely
winged stems), longer bracts (10—12 mm vs. 4—10
mm), longer and long-apiculate calyx lobes (28-40
mm vs. 15-24 mm and obtuse to slightly acute)
with a long thin wing on the back of each calyx
lobe (vs. a short “hump-like” keel), corolla tube
1.0—1.25 times longer than calyx (vs. 2—3 times),
and white, yellow, to green corollas (vs. pink to
red). Only one known collection of Symbolanthus
pterocalyx has corollas that are recorded as pale
pink (Zarucchi & Cardenas 4322) as opposed to
yellow 1 and green, so corollas of the pink-red color
scheme are very uncommon in this species. In a
few specimens the calyx keels are not strongly de¬
veloped toward the apex of the calyx lobes, but the
keels are always present at the very upper portion
of the pedicel to the base of the calyx lobes. Usu¬
ally the same specimens that have less well-devel¬
oped wings on the stems will always have the long
and acuminate calyx lobes (at least somewhat
keeled) and greenish yellowish corollas (e.g., Fon-
negra et al. 5984). The calyx wings are often de¬
current as ridges down onto the pedicel (e.g., in
Marulanda et al. 304, where the wings from the
two outer calyx lobes reach down to the base of the
pedicel and cause it to be distinctly flattened in
cross section).
Gentians are rarely eaten by insects and other
herbivores, but several collections of Symbolanthus
pterocalyx have circular to 5 mm large holes in
their leaves (e.g., Marulanda et al. 304, Vargas
790). Nearly all gentians contain large amounts of
extremely bitter chemical compounds (seeo-iri-
Volume 13, Number 1
2003
Struwe
Symbolanthus from Colombia
139
doids; Samuelsson, 1992: 153; Jensen & Shripse-
ma, 2002), but it is not known if these reduce her-
bivory. Furthermore, according to herbarium label
information from Agudelo & Betancur 32, in Sym¬
bolanthus pterocalyx, wasps (Vespidae) are associ¬
ated with the hollow stems and beetles (Coleoptera)
are found in the (presumably floral) receptacles.
Hummingbirds have been recorded as visitors to a
Symbolanthus tetrapterus plant with white to green-
colored corollas ( Silverstone-Sopkin et al. 4140).
Symbolanthus tetrapterus differs from S. calygon-
us in its prominently winged stems (vs. not or barely
winged stems), obovate to obovate-oblong and ses¬
sile (rarely subsessile) leaves (vs. elliptic-lanceo¬
late and distinctly petiolate leaves) with a strongly
attenuate base (vs. cuneate), shorter bracts (2—3
mm vs. 4— 10 mm), non-alate calyces (vs. a dorsal
keel on each calyx lobe), and green, white, pink,
or red-violet corollas (vs. always pink to red: Table
ll .
The mountain Cerro del Torra is an isolated out¬
lier west of the Cordillera Occidental of the Andes
and reaches about 2800 m elevation. It is the only
known locality of Symbolanthus tetrapterus, and
over 25 other endemic plant species have been de¬
scribed from this mountain (in Ericaceae, Melas-
tomataceae, Orchidaceae, and Solanaceae). In fact,
of the three gentians so far found on Cerro del Tor¬
ra. possibly two are endemic to this mountain (Sym¬
bolanthus tetrapterus and Macrocarpaea sp. nov.. J.
R. Grant, unpublished). The floristic study of Cerro
del Torra by Silverstone-Sopkin and Ramos-Perez
(1995) showed an overwhelming floristic affinity be¬
tween Cerro del Torra and the Cordillera Occiden¬
tal. This result suggested that this mountain is not
a phytogeographical entity distinct from the Cor¬
dillera Occidental.
The phylogenetic relationships of Symbolanthus
pterocalyx and .S', tetrapterus are uncertain at pres¬
ent, and the closest relatives to each cannot be dis¬
cerned based on simple comparison of morpholog¬
ical traits. Molecular studies including other
species of Symbolanthus have shown that sequence
variation in the fast-evolving 5S-NTS nrDNA region
is very low and does not resolve species relation¬
ships in Andean Symbolanthus very well (Gould &
Struwe, in prep.). However, molecular data suggest
that there is support for the hypothesis that green-
white vs. red-pink-flowered species do not consti¬
tute major monophyletic groupings. These two new
species are not very similar to each other apart from
their winged stems, and other morphological fea¬
tures do not suggest a close relationship to each
other. Symbolanthus pterocalyx from Antioquia
shows more similarities with the eastern Andean
species S. vasculosus in its long-petiolate leaves
and large, campanulate corolla with wide corolla
lobes, inferring possible bat pollination. The more
southern Symbolanthus tetrapterus has sessile
leaves and salver-shaped flowers indicating hum¬
mingbirds as possible pollinators. Both species
have winged stems but this character occurs in oth¬
er species as well, e.g., Symbolanthus vasculosus.
The presence of winged stems in these species is
probably due to parallel evolution of this trait in
geographically separated populations, rather than a
result of common ancestry. Further phylogenetic
studies are needed, and until suitable DNA mark¬
ers become available, hypotheses about species re¬
lationships will have to be based on selected mor¬
phological characteristics alone.
Acknowledgments. The author gratefully thanks
Jason R. Grant for translations, Latin diagnoses,
and unpublished information: Philip Silverstone-
Sopkin for information regarding Cerro del Torra
and comments on the manuscript; Jim Luteyn for
information on S. pterocalyx; Rocfo Cortes for Span¬
ish translations; Francisco Javier Roldan for infor¬
mation on S. pterocalyx; Max Weigend and Rodrigo
Bernal for information on Colombian localities; as
well as the late Rupert Barneby for advice on Fatin
names. The herbaria CUVC. HUA. k, MO. NY, U.
and US are also gratefully acknowledged for loans
or information about their herbarium material. Bob-
bi Angell skillfully prepared the plant illustrations.
This work was funded by the Lewis B. and Dorothy
Cullman Program for Molecular Systematics Stud¬
ies, The New York Botanical Garden, and Rutgers
University—Cook College. The 1988 expedition to
Cerro del Torra, during which some of the Symbo¬
lanthus tetrapterus specimens were collected and
photographed, was financed by a National Geo¬
graphic Society grant (#3884—88) to Philip Silver¬
stone-Sopkin.
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aceae—Systematics and Natural History. Cambridge
Univ. Press, Cambridge.
Aculeatae , a New Series in Bauhinia Section Pauletia
(Leguniinosae, Caesalpinioideae, Cercideae)
Angela Maria S. da F. Vaz
Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Rua Pacheco Leao 915, Rio de
Janeiro, RJ, 22460-030, Brazil, amvaz@jbrj.gov.br
Ana Maria G. de Azevedo Tozzi
Universidade Estadual de Campinas, Instituto de Biologia, Botanica, Caixa Postal 6109,
Campinas, SP, 13083—970, Brazil
Abstract. Bauhinia ser. Aculeatae Vaz & A. M.
G. Azevedo, a new series in section Pauletia, is
established as separate from Bauhinia ser. Cansen-
ia (Rafinesque) Wunderlin, k. Larsen & S. S. Lar¬
sen. A description of the new series is presented
together with a description of series Cansenia newly
circumscribed.
Key words: Bauhinia sect. Pauletia, Brazil,
Caesalpinioideae, Cercideae, Leguminosae.
Bauhinia L. is a pantropical genus of tribe Cer¬
cideae (Leguminosae, Caesalpinioideae). Wunder-
lin’s (1976) and Wunderlin et al.’s (1981, 1987)
studies led to a classification of the Cercideae, in¬
cluding 4 subgenera, 22 sections, and 30 series for
Bauhinia. Bauhinia sect. Pauletia (Cavanilles) DC.
is characterized by the presence of 10 fertile sta¬
mens or try 5 fertile stamens alternating with 5 sta¬
mens with reduced anthers or, rarely, with 6 to 8
fertile stamens and 2 to 4 staminodes. In all cases,
filaments are connate at the base and generally
have pollen grains with a reticulated sexine and
with a supratectal gemoid process. Section Pauletia
encompasses ca. 70 species in tropical America
and 2 species in southern Asia, South China, and
the Malesian Area, according to Wunderlin et al.
(1987). In tropical America, its distribution ranges
from Mexico to Argentina, encompassing 53 spe¬
cies of Bauhinia in Brazil (Vaz, 2001).
Bentham (1870) divided Bauhinia sect. Pauletia
into three informal groups of species based on the
attributes of petals and stamens: (1) narrow petals,
10 perfect stamens, without prickles; (2) narrow
petals, alternate (antepetalous) filaments with
smaller or deficient anthers or anantheriferous,
prickles robust-growing; (3) obtuse petals, 10 per¬
fect stamens, prickles robust-growing. Wunderlin et
al. (1987) divided Bauhinia sect. Pauletia into five
series, including also features of the calyx, pollen,
and fruits: Bauhinia ser. Cansenia (Rafinesque)
Wunderlin, K. Larsen & S. S. Larsen. B. ser. Acu-
minatae Wunderlin, K. Larsen & S. S. Larsen, B.
ser. Perlebia (Martius) Wunderlin, k. Larsen & S.
S. Larsen, B. ser. Pentandrae Wunderlin, k. Larsen
& S. S. Larsen, and B. ser. Ariaria (Cuervo Mar¬
quez) Wunderlin, K. Larsen & S. S. Larsen. Here,
the first and third groups proposed by Bentham
(1870) whose species have 10 perfect anthers were
merged into series Cansenia. Nevertheless, Wun-
derlin’s Cansenia comprises two distinct groups in
Bauhinia and should therefore be divided into se¬
ries corresponding roughly to the first and third in¬
formal groups proposed by Bentham (1870).
This paper, which is part of a more extensive
taxonomic study of Bauhinia sect. Pauletia (Vaz,
2001), emends the taxonomic circumscription of its
inclusive series Cansenia and proposes a new sep¬
arate series. To circumscribe both series, we have
added the following new diagnostic characters: in¬
florescence type, the presence/absence of prickles,
and the presence/absence of the extrafloral nectar¬
ies. Ihe inflorescences in Neotropical Bauhinia
sect. Pauletia are terminal pseudoracemes, which
always have successively geminate pedicels. The
sympodium is leafy at base, or the leaves are re¬
placed by foliaceous bracts (series Cansenia and
Ariaria). In another group of species (series Acu¬
leatae, Pentandrae, Perlebia ), flowers are arranged
in 2(3-6) leaf-opposed clusters (cymoids). These
inflorescences are of terminal origin, although re¬
ferred to as axillary or supra-axillary by most au¬
thors. Both types of inflorescence (pseudoracemes
and cymoids) have already been recognized in Bau¬
hinia by Urban (1885). Structures described by
Bentham (1870) as “aculei” ( = prickles) are in
fact prickles in some species and extrafloral nec¬
taries in others. The extrafloral nectaries, as rec¬
ognized here, were first detected for B. curvula
Bentham (Besende et al., 1994, see fig. 1), although
Novon 13: 141-144. 2003.
142
Novon
a “nectar secreting thread'" has already been re¬
corded by van der Pijl (1952: 292) for an American
species not identified. The extrafloral nectaries are
smaller, not pungent, intrastipular projections. 1-2
mm long, subulate or ovoid in shape. They are also
presented in pseudoracemes as paired projections
from foliaceous bracts. In some species extrafloral
nectaries may be rudimentary or obsolete. Adpe-
ciolar prickles are always pungent, straight or hook¬
shaped, 1-10 mm long. They are positioned below
or beside the adjacent stipules. In some species,
the alternate prickle may be rudimentary or obso¬
lete.
Buuliinia ser. Cansenia (Rafinesque) Wunderlin,
K. Larsen & S. S. Larsen, Biol. Skr. 28: 12.
1987. hie emendavit. Basionym: Cansenia Raf¬
inesque, Sylva Tellur. 122. 1838. TYPE: Can¬
senia ungulata (L.) Rafinesque (lectotype, des¬
ignated by Wunderlin (1976)).
Plant with terminal branches ascendent, prickles
absent; extrafloral nectaries present or rudimentary.
Inflorescence a terminal pseudoraceme, leafless,
with foliaceous bracts or sometimes with scattered
reduced leaves toward the apex, partial inflores¬
cence adnate to the main axis by geminate pedi¬
cels, (1— )2(—3)-flowered; calyx at anthesis irregu¬
larly connate at the apex and splitting into 2 to 5
lobes; petals narrow, acute or acuminate or obtuse
(only in B. cheilantha ); 10 fertile stamens, connate
at the inner internal face, but without ligular ap¬
pendix.
Suffrutescent. about 0.5—2 m high, when growing
in "campo open formations or Brazilian savannas,
with a conspicuous xylopodium—a subterranean
organ adapted for vegetative propagation during
rainy periods, after a long-lasting dry season and
periodical burning of the aerial portion of plant
cover (Rizzini & Heringer, 1961, 1962). Also found
growing as trees or treelets, 4—15 m tall in forest,
including gallery forests. The multi-stemmed
shrubby habit is found in Brazilian savannas
(burned periodically with scattered trees and a her¬
baceous ground cover), secondary forests, and dis¬
turbed areas in Brazil.
Pollen triangular-shaped in polar view, angulap-
erturate 3(4) colpi short, covered with a colpus
membrane (figs. 10-14 in Ferguson & Pearce,
1986; Melhem & Labouriau, 1963; Vaz, 2001, part
1, fig. 16D).
Ibis series, here newly circumscribed, corre¬
sponds to the first group of Bauhinia sect. Pauletia
(Bentham, 1870), except for Bauhinia marginata
(Bongard) Steudel and Bauhinia hirsuta (Bongard)
Vogel ( — B. hirsutijlora Vaz), both species belong¬
ing to series Pentandrae.
Species Included
1. Bauhinia acuruana Moricand, PL Nouv. Amer.
6, [). 77, tab. 5. 1840.
2. Bauhinia aureopunctata Ducke, Arch. Jard.
Bot. Rio de Janeiro 4: 53. 1925.
3. Bauhinia bombaciflora Ducke, Arch. Jard. Bot.
Rio de Janeiro 3: 104, fig. 5. 1922.
4. Bauhinia brevipes Vogel. Linnaea 13: 307.
1839.
5. Bauhinia burchellii Bentham. in Martius, FI.
Bras. 15(2): 186. 1870.
6. Bauhinia caloneura Malme, Bih. Kongl. Sven-
ska Vetensk.-Akad. Handl. 25 (3. n° 11): 29.
pi. 2. 1900.
7. Bauhinia campestris Malme, Ark. Bot. 5(5): 10.
1905.
8. Bauhinia candelabriformis R. S. Cowan, Contr.
Sci. Los Angeles County Mus. 13: 4. fig. 1.
1957.
9. Bauhinia cheilantha (Bongard) Steudel, Nom.
Bot., ed. 2. v. 1. p. 191 sphalm. 291. 1840.
10. Bauhinia cinnamomea DC., Prodr. p. 517.
1825.
11. Bauhinia canwayi Rushy. Bull. New York Bot.
Card. 8(28): 92. 1912.
12. Bauhinia cupulata Bentham, in Marlins, FI.
Bras. 15(2): 188. 1870.
13. Bauhinia curvula Bentham, in Martius, FI.
Bras. 15(2): 194. 1870.
14. Bauhinia dubia G. Don, Gen. Syst. 2, p. 463.
1832, non B. dubia Vogel.
15. Bauhinia dumosa Bentham, in Martius, FI.
Bras. 15(2): 194. 1870.
16. Bauhinia fusconervis (Bongard) Steudel, Nom.
Bot. ed. 2, v. 1. p 1. p. 191 sphalm. 291. 1840.
17. Bauhinia gardneri Bentham, in Martius, FI.
Bras. 15(2): 186. 1870.
18. Bauhinia goyazensis Harms, Bot. Jalirb. Syst.
33(72): 21. 1903.
19. Bauhinia grandifolia (Bongard) Steudel, Nom.
Bot. ed. 2, v. 1. p. 191 sphalm. 291. 1840.
20. Bauhinia holophylla (Bongard) Steudel. Nom.
Bot. ed. 2. v. 1. p. 191 sphalm. 291. 1840.
21. Bauhinia leptantha Malme, Ark. Bot. 5(5): 1 1.
1905.
22. Bauhinia longicuspis Bentham, in Martius. FI.
Bras. 15(2): 185. 1870.
23. Bauhinia longifolia (Bongard) Steudel, Nom.
Bot. ed. 2. v. 1, p. 191 sphalm. 291. 1840.
24. Bauhinia longipedicellata Ducke, Arch. Jard.
Bot. Rio de Janeiro 4: 105. 1922.
Volume 13, Number 1
2003
Vaz & Azevedo
Bauhinia ser. Aculeatae
143
25. Bauhinia malacotricha Harms, Hot. Jahrb. Syst.
33(72): 22. 1903.
26. Bauhinia malacotrichoides Cowan, Contr. Sci.
Los Angeles County Mus. 13: 8, fig. 3. 1957.
27. Bauhinia membranacea Bentham, in Martius,
FI. Bras. 15(2): 187. 1870.
28. Bauhinia pulchella Bentham, in Martius, FI.
Bras. 15(2): 190. tab. 49. 1870.
29. Bauhinia rufa (Bongard) Steudel, Nom. But.
ed. 2, v. 1, p. 192. 1840.
30. Bauhinia smilacifolia Bentham, in Martius, FI.
Bras. 15(2): 183. 1870.
31. Bauhinia subclavata Bentham, in Martius, 11.
Bras. 15(2): 188. 1870.
32. Bauhinia tenella Bentham, in Martius, FI.
Bras. 15(2): 195. 1870.
33. Bauhinia ungulata L., Sp. PI. ed. 1, p. 374.
1753.
34. Bauhinia urocalyx Harms, Notizbl. Konigl.
Bot. Cart. Berlin 6(59): 308. 1915.
Bauhinia ser. Aruleatac Vaz & A. M. G. Azevedo,
ser. nov. TYPE: Bauhinia aculeata L.
Rami terniinales vulgo subhorizontales, aculei gemini
vel subsolitarii; nectaria extrafloralia nulla. Inflorescentiae
foliosae, partiales unilaterales, 2(3-6)-florae; calyx per an-
thesin in spatham deflexam bine fissus ad basim; petala
lata, oblanceolata vel spathulata; stamina 10, omnia fer-
tilia, base interne appendice ligulari praedita.
Plant with terminal branches subhorizontal,
prickles paired or subsolitary; extrafloral nectaries
absent. Inflorescence leafy, partial inflorescences
unilateral, 2(3—6)-flowered, calyx spathaceous at
anthesis and then splitting to the base, petals
broad, oblanceolate or spathulate; stamens 10, all
fertile, at inner base with ligular appendix.
Trees or treelets, ca. 4—15 m tall, found in forest
formations in tropical America, including gallery
forests of Brazilian cerrado, but tiever suffrutescent.
although they may survive as multi-stemmed
shrubs in secondary forest, including disturbed ad¬
jacent sites. In high-altitude grassland (“campos de
altitude”) in southeastern Brazil, in Rio de Janeiro
and Sao Paulo states, they may occur as treelets.
Pollen (4—)5—6-zonocolpate with colpi wide and
short ( = type 3 subtype b. in Gamerro & Fortun-
ato, 2001, figs. 5-7; Vaz, 2001. part 1, fig. 16B;
see also Barth & Bousada, 1964, figs. 1, 2).
This series corresponds to the third group of
Bauhinia sect. Pauletia (Bentham, 1870), exclud¬
ing Bauhinia corniculata Bentham, which belongs
to series Pentandrae and B. cheilantha cited above,
under series Cansenia.
Species Included
1. Bauhinia acreana Harms, Notizbl. Konigl. Bot.
Gart. Berlin 6: 307. 1915.
2. Bauhinia aculeata L., Sp. PI. ed. 1, 374. 1753.
3. Bauhinia affinis Vogel, Linnaea 10: 594. 1836.
4. Bauhinia albicans Vogel. Linnaea 13: 304.
1839.
5. Bauhinia caatingae Harms, Bot. Jahrb. Syst.
42: 209. 1908. (from caatinga, a Brazilian veg¬
etation type; sphalm. catingae).
6. Bauhinia forficata Link, Enum. Hort. Berol. 1:
404. 1821.
7. Bauhinia integerrima Bentham, in Martius, FI.
Bras. 15(2): 197. 1870.
8. Bauhinia mollis (Bongard) I). Dietrich, Syn. PI.
2: 1475. 1840.
9. Bauhinia ovata (Bongard) Vogel. Linnaea 13:
300. 1839.
10. Bauhinia platypetala Bentham, in Martius, FI.
Bras. 15(2): 198. 1870.
Acknowledgments. The authors thank Dorothy
I). Araujo (UFRJ. Rio de Janeiro) for the English
translation and Tarciso S. Filgueiras (IBGE, Bra¬
silia) for his review and improvement of the Latin
diagnosis. We appreciate the helpful comments of
Dan H. Nicolson (US), Victoria C. Hollowell (MO),
and R. P. Wunderlin (Tampa, Florida). Thanks to
the Conselho Nacional de Desenvolvimento Cien-
tffico e Teenologico (CNPq) for a Ph.D. fellowship
at Unicamp University given to Angela S. da F. Vaz.
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dos pdlens das plantas arboreas do Brasil meridional.
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Bentham, G. 1870. Leguminosae II. Svvartzieae et Caesal-
pinieae. Pp. 179-254 in (7 P. F. von Martius & A. G.
Fichler (editors), Flora Brasiliensis, Vol. 15 (2). Frid.
Fleischer, Leipzig.
Ferguson. I. K. & K. J. Pearce. 1986. Observations on the
pollen morphology of the genus Bauhinia L. (Legumi-
nosae, Caesalpinioideae) in the Neotropics. Pp. 283—
296 in S. Blackmore & 1. K. Ferguson (editors), Pollen
and Spores: Forms and Function. The Linnean Society
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Gamerro, J. C. & R. H. Fortunato. 2001. Morfologi'a del
polen de las especies Argentinas de Bauhinia (Cerei-
deae, Caesalpinioideae, Fabaeeae). Ann. Missouri Bot.
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Melhem, T. S. & M. L. S. Labouriau. 1963. Pollen grains
of plants of the “cerrado”—V: Leguminosae, Caesalpi¬
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Neerl. 1(2): 287-309.
Rezende, M. H„ L. A. Cardoso & A. L. Vannucci. 1994.
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gans of trees and shrubs from some southern Brazilian
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de Biologia, Universidade Estadual de Campinas, Cam¬
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Wunderlin, B. P. 1976. Enumeration and typification of
genera in the tribe Cercideae. Rhodora 78: 750—760.
-, K. Uarsen & S. S. Uarsen. 1981. Cercideae. Pp.
107—116 in B. M. Polhill & P. II. Raven (editors). Ad¬
vances in Legume Systematics, Vol. 1. Royal Botanic
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-.-& -. 1987. Reorganization of the
Cercideae (Fabaceae: Caesalpinioideae). Biol. Skr. 28:
1-40.
An Overview of the Subspecies of Paullinia obovata
(Sapindaceae-Paullinieae) in Peru
Caroline S. Weckerle
Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008 Zurich,
Switzerland, weckerle@systbot.unizh.ch
Carlos Reynel
Facultad de Ciencias Forestales/Herbario Forestal, Universidad Nacional Agraria La Molina,
Ap. Postal 451, Lima 1, Peru, reynel@lamolina.edu.pe
ABSTRACT. An overview of the subspecies of
Paullinia obovata in Peru is presented, recognizing
five subspecies. One new subspecies, P. obovata
subsp. flava Weckerle & Igersheim. is described.
Paullinia brentberlinei is transferred to the rank of
subspecies of P. obovata. Three varieties of P. obov¬
ata, i.e., var. obovata, var. polymorpha, var. subro¬
tunda, are recognized at the subspecies level. All
subspecies are illustrated, and their comparative
morphological features are compared in a table.
If KSUMF.N. Se presenta una revision de las subes-
pecies de Paullinia obovata en Peru. Son recono-
cidas cinco subespecies. Se describe una nueva
subespecie, P. obovata subsp. flava Weckerle & Ig¬
ersheim. Paullinia brentberlinei es reducida a una
subespecie de P. obovata. Ires variedades de P.
obovata (var. obovata, var. polymorpha, var. subro¬
tunda) son reconocidas como subespecies. Se pro-
vee una tabla resumen eon los caracteres morfolo-
gicos para la comparacidn de los taxa, asf como
ilustraciones para cada uno de ellos.
Key words: Paullinia, Paullinieae, Peru, Sap-
indaceae.
The tribe Paullinieae (Sapindaceae) mainly oc¬
curs in tropical America; exceptions are a few spe¬
cies of Cardiospermum L. with a pantropical distri¬
bution and Paullinia pinnata L., which is also
found in tropical Africa and Madagascar (Radlko-
fer, 1933). The tribe consists of lianas (plants grow¬
ing in open areas may be non-climbing treelets)
and vines with tendrils, a growth form not very
common to the Sapindaceae and found in the Paul¬
linieae sensu Radlkofer (1933) only. The genera of
this tribe have very similar flowers, characterized
by zygomorphy and four petals with basal append¬
ages. The taxa are separated mainly by fruit and
seed characters.
Paullinia L. is the second largest genus of the
tribe Paullinieae with about 200 species. Most of
them occur in the rain forests of Amazonia. About
70 species are found in Peru (Simpson, 1976; Ger-
eau. 1993), chiefly in the lowland rain forest and
the rain forest of the eastern Andean slopes. The
genus Paullinia is polymorphic with regard to veg¬
etative features as well as fruit and seed morphol¬
ogy. Cross sections of the stems have a single stele
or less commonly numerous steles, which means
that a central stele can be surrounded by one, two,
or three peripheral steles. The leaves are alternate
and often 5-foliolate pinnate but may vary from 3-
foliolate to biternate or multifoliolately ternate-pin-
nate, with margined or winged petioles and raehis-
es. The 3-valved capsules are stipitate, subsessile
to sessile, 3-locular, and uniovulate. Paullinia can
be divided into three main species groups, differing
conspicuously in macromorphological fruit charac¬
ters. (1) The majority ol taxa (± 2/3) are charac¬
terized by globose, subglobose, fusiform, obovate,
or elavate fruits; (2) several species (± 1/3) have
fruits with dorsal wing-like outgrowths; (3) five spe¬
cies have echinate fruits (Radlkofer, 1933). The
mature capsules are often reddish, more rarely yel¬
lowish or greenish. The seeds are globose, ovate,
ellipsoid, or compressed ellipsoid with a black or
brown exotesta. They are partly or totally enclosed
with a white aril (actually an arilloid sensu van der
Pijl (1957)) of different texture.
Paullinia obovata (Ruiz & Pavon) Persoon is re¬
stricted to western Amazonia (Ecuador, Peru, Bo¬
livia, and adjacent regions of Colombia and Brazil)
with its main distribution in Peru (Simpson, 1976).
The stem has a single stele, the leaves are 5-fo-
liolate pinnate with unwinged or occasionally mar¬
gined petioles and rachises, and the capsules are
stipitate or subsessile, 2.5-9.0 cm long, with thick
and fibrous valves. The usually three seeds are at
least half enclosed by a sweet aril.
Novon 13: 145-152. 2003.
146
Novon
Figure I. Geographic distribution of the subspecies of
Paullinia obovata in Peru, based on specimen locations.
The new subspecies Paullinia obovata subsp.
flava Weckerle & Igersheim was found in the low¬
lands of Peru in the vicinity of Jenaro Herrera, de¬
partment Loreto. Examination of herbarium mate¬
rial (USM) showed that this subspecies also occurs
further south in the department Madre de Dios (Fig.
1). It differs from the other subspecies mainly by
its yellow instead of red fruits (Table 1, Fig. 3A, B)
and its distribution restricted to the lowlands (100—
400 m).
Paullinia brentberlinei Croat is known from the
tropical lowlands (200-300 rn) of northern Peru
(Fig. 1). Examination of the type material as well
as other collections reveals that this species differs
from P. obovata subsp. polymorpha only by having
larger fruits (8—9 cm in subsp. brentberlinei vs. 4—
7 cm in subsp. polymorpha, cf. Table 1). Therefore,
this species is transferred to P. obovata. It is prob¬
ably best to accept it as a subspecies of P. obovata,
as suggested by Beck (1991) in his Ph.D. thesis.
In his partial revision Simpson (1976) recognized
in P. obovata three varieties: var. obovata, var. po¬
lymorpha I). R. Simpson, and var. subrotunda (Ruiz
& Pavon) D. R. Simpson. They differ mainly in
their size and shape of the fruits (Fig. 2A-C) and
some vegetative characters (cf. Table 1). The vari¬
eties of P. obovata as delimited by Simpson (1976)
show a geographical correlation with the main mor¬
phological characters and are therefore treated as
subspecies (Hamilton & Reichard, 1992). Subspe¬
cies obovata and subspecies polymorpha, both with
fusiform and stipitate fruits, differ in fruit size (2.5-
4.5 cm in subsp. obovata vs. 4—7 cm in subsp. po¬
lymorpha), serratioji type and indument of the leaf¬
lets (conspicuous, beginning below or above
middle; mostly glabrous in subsp. obovata vs. in¬
conspicuous, beginning above middle; pubescent
beneath in subsp. polymorpha, cf. Table 1). Their
distribution area is overlapping. Subspecies obov¬
ata has its main area of distribution between 700
and 1000 m, but may also be found in the lowlands
((200)-400 m), whereas subspecies polymorpha is
mainly found in the lowlands but reaches elevations
of up to 800 m. Subspecies subrotunda has obovate,
(sub)sessile fruits and is restricted to middle ele¬
vations ((700)—1200—1800 m) of the central eastern
Andean slopes. Subspecific rank for these taxa also
provides more consistency of the infraspecific de¬
limitation within Paullinia. Intermediate forms of
subspecies of P. obovata are occasionally found
(e.g., leaf characters pointing to subsp. obovata
combined with fruit characters typically found in
subsp. polymorpha).
Paullinia obovata (Ruiz & Pavon) Persoon, Syn.
PI. 1:443. 1805. Basionym: Semarillaria obov¬
ata Ruiz & Pavon, FI. Peruv. Prodr. 54. 1794;
FI. Peruv. IV: 156-157. 1954. TYPE: Peru.
Pozuzo [not preserved or destroyed?]. FEC-
TOTYPE (designated here): “ Semarillaria
obovata ” [icon no. 338 in] Ruiz & Pavon, FI.
Peruv. IV: 189. 1954.
In the Flora Peruvianae, et Chilensis Prodromus
54, Ruiz and Pavon (1794) mentioned the species
name Semarillaria obovata for the first time. Only
a short sentence is given, and neither specimens
nor localities are cited. The same authors present
in the FI. Peruv. IV (1954), a broader description
stating “[ Semarillaria obovata ] habitat in Peruviae
Andium nemoribus Pozuzo,” but still without a ci¬
tation of specimens. In his revision Simpson (1976)
designated “Ruiz & Pavon s.n. from Pozuzo” as a
“fictive type” of P. obovata. Photographs (negatives)
of lectoparatypes are cited, i.e., F 53208 [53204]
and F 23655 (ex G). These photographs show ster¬
ile specimens, which contain no original label or
note from Ruiz and Pavon, and the corresponding
specimen to F 23655 could not be found in Geneva
(G). Therefore, these two specimens cannot be
linked to the information provided in the proto-
logue. Since Ruiz noted in his journal Semarillaria
Table 1. Comparative morphological features of the subspecies of Paullinia obovata.
Volume 13, Number 1
2003
Weckerle & Reynel
Paullinia obovata in Peru
147
148
Novon
Figure 2. Typical fruit shapes in subspecies of Paullinia obovata. —A. Paullinia obovata subsp. obovata, Weckerle
& Igersheim 000330-2/1 (MOL). —ft. Paullinia obovata subsp . polymorpha, Weckerle & Igersheim 010202-1/2 (MOL).
—C. Paullinia obovata subsp. subrotunda, Weckerle & Igersheim 000329-1/1 (MOL). Scale bar 2 cm.
obovata among those species he “succeeded in de¬
scribing in lluanuco, of those that were worked on
in Macora and that were burned in that fire” (as
cited in Ruiz, [1777-1788] 1940: 203), it can be
expected that the set of collections made by Ruiz
and Pavon and used lor their description and illus¬
tration prepared in Peru (Lack, 1979) was de¬
stroyed. There is no doubt about the good quality
of the illustration that accompanies Ruiz and Pa-
vdn’s (1954) description of the species in FI. Peruv.
IV. It agrees unambiguously with the taxon as un¬
derstood by us and shows clearly, besides leaf char¬
acters, the pertinent fruit characters. The illustra¬
tion given by Ruiz and Pavon (1954) is therefore
designated herewith as the lectotype of Paullinia
obovata (Ruiz & Pavdn) Persoon.
1. Paullinia obovata (Ruiz & Pavon) Persoon
subsp. obovata
Distribution and habitat. Paullinia obovata
subsp. obovata is a rather widespread taxon and
occurs mainly in the forests of the eastern Andean
slopes between 700 and 1000 m, but may also be
found in the lowlands ((200)-400 m) of Junfn, Pas¬
co, San Martin, and Ucayali, along rivers, edges of
secondary forests, and along roadsides (Fig. 1).
Phenology. Flowering specimens were collect¬
ed during October and January and specimens with
mature and immature fruits during January, March,
April, May, July, and October.
Discussion. Vegetatively, Paullinia obovata
subsp. obovata is very similar to P. obovata subsp.
flava, and both taxa seem to be closely related. In
both subspecies the serration of the margins of the
leaflets begins below or above the middle, and the
lower surface of the leaflets is mostly glabrous (Ta¬
ble I). The two subspecies differ markedly in the
color of the capsules at maturity (red in subsp.
obovata vs. yellow in subsp. flava, cf. Fig. 3A, R
and C-E), and in contrast to subspecies flava,
which is restricted to the lowlands, subspecies
obovata occurs mainly in the eastern Andean slopes
(Fig. 1).
Specimens examined. PERU. No locality given: Pavdn
644 (G), Pavdn 1821 (G), Pavdn s.n. ((f). San Martin:
Mariscal Cdceres, Tocache Nuevo, Fundo Miramar, “en
bosque alto." 28 Mar. 1970, Schunke 3879 (G, MO, MOL);
“al horde del Rio en bosque alto,” I July 1974, Schunke
7091 (G, MO, USM); Fundo Miramar, “en bosque alto,”
Volume 13, Number 1
2003
Weckerle & Reynel
Paullinia obovata in Peru
149
10 Oct. 1980, Schunke 12338 (USM, Z), Schunke 13388
(USM, Z). Ucayali: Coronell Portillo, Pucallpa, Rfo Pach-
itea, 30 Jan. 1988, Ayala 6002 (MO); Leoncio Prado (Yar-
inacocha), 15 May 1984, Vdsquez 5004 (MO); Previsto, on
the bank of stream, 10 Oct. 1962, Woytkowski 7573 (MO).
Junui: Puente Paucartambo, 30 Mar. 2000, Weckerle &
Igersheim 000330-2/1 (MOL); Perene, road between Per¬
ene and La Esperanza, 4 Apr. 2000, Weckerle & Igersheim
000404-1/5 (MOL); riverside of Rfo Paucartambo, near
Puente Colgante along road to Perene, 19 July 2000,
Weckerle & Igersheim 000719—1/2 (MOL); La Merced,
road between Puente Colorado and Casa Rlanca, 5 Apr.
2000. Weckerle & Igersheim 000405—1/4 (MOL), 710 m,
3 July 2000, Weckerle & Igersheim 000703-2/1 (MOL);
road between Puente Colorado and La Alianza, 3 July
2000, Weckerle <£: Igersheim 000703—1/2 (MOL); road
along Quebrada Pescarola, 10 July 2000, Weckerle & Ig¬
ersheim 000710-1/6 (MOL); Puente Quimiri, riverside of
Rfo Chanchamayo near confluence with Quebrada Sheila,
18 July 2000, Weckerle & Igersheim 000718-2/1 (MOL);
Villa Rica, road between Cedro Pampa and Purus, river¬
side of Quebrada Aqua Blanca, ca. 7 July 2000, Weckerle
A Igersheim 000707-2/1 (MOL); San Ramon, road be¬
tween Puente Victoria and Lourdes, along Quebrada Ox-
abamba, 11 July 2000. Weckerle & Igersheim 000711-1/
2 (MOL); road between Puente Victoria and Lourdes,
along Quebrada Oxabamba, 11 July 2000, Weckerle & Ig¬
ersheim 000711-1/6 (MOL); riverside between Puente
Victoria and Puente Almandra, 15 Jan. 2001, Weckerle &
Igersheim 010115—1/2 (MOL).
2. Paullinia obovata subsp. Hava Weckerle & Ig¬
ersheim, subsp. nov. TYPE; Peat. Ixireto: May-
nas, Requena, ca. 7 km W of Jenaro Herrera,
vicinity of Laguna Vainilla. inundated primary
forest (Braga), 04°54'46"S, 73°44'20"W, ca. 1 10
m, 27 May 2(X)0, A. Vdsquez, C. S. Weckerle &
A. Igersheim 000527-1/4 (holotype, MOL; iso¬
type, Z). Figure 3A, B.
Ab aliis subspeciebus capsulis llavis cum lineis dehis-
centiae rubris differt.
Liana, stems sparsely pubescent to mostly gla¬
brous, stem with a single stele, up to 8-9 cm diam.
Leaves pinnately 5-foliolate, petiole and rachis
wingless, petiole 4-14 cm, rachis 2-5 cm, stipules
subulate, deciduous. Petiolule 0.3—0.8 cm, leaflets
chartaceous, ovate, elliptic to obovate, 7.5-21.5 cm
long (including the petiolule), 4—9.5 cm wide, the
apex acuminate, the base acute, obtuse to rounded,
margins with serration beginning above or below
middle, never at base, mostly glabrous except for
the hair tufts in the axils between primary and sec¬
ondary nerves, some secondary nerves bifurcating
near the margin. Infructescence 11 — 19 cm long, ra¬
chis rusty tomentose, bracts deciduous. Fruits stip-
itate or subsessile, fusiform to obovate, 3—4 cm long
(including the 0.3-0.5 cm long stipe), 2.5-3.5 cm
wide, yellow when mature with three red dehis¬
cence lines, the valves fibrous, with fibers running
longitudinally, the three septa usually persistent;
seeds dark brown or black, characteristically cov¬
ered by a white aril.
Distribution and habitat. Paullinia obovata
subsp. flava is restricted to lowland rain forests
along river margins or seasonally inundated areas
at elevations from 100 to 400 m (Fig. 1).
Phenology. The two specimens examined with
buds were collected in October and November. The
remaining specimens show young and mature fruits.
They have been collected in January, March. May,
and June, respectively.
Discussion. Herbarium material of Paullinia
obovata subsp. flava is very similar to that of sub¬
species obovata (fruit size and shape, serration type
and indument of leaflets) and therefore sometimes
difficult to distinguish. The diagnostic feature for
subspecies flava, the yellow color of the capsules
with the three red dehiscence lines, cannot be seen
directly on the dried herbarium specimen. How¬
ever, after drying, the capsules of subspecies flava
appear to be more orange-colored than the capsules
of all other subspecies of Paullinia obovata, which
are brown. Thus, notes on the labels concerning the
color of mature fruits at their time of collection are
important for identification.
Paratypes. PERL. Loreto: Maynas, Requena, along
Quebrada Cano Iricahua near the confluence with Rio
Ucayali, 31 May 2000, Vdsquez, Weckerle & Igersheim
000531-1/6 (MOL); Reserva Nacional Pacaya Samiria, Si-
taraco, 29 Mar. 1993, del Carpio 1977 (USM); Campa-
mento La Huaca, 9 June 1993, del Carpio 2448 (USM);
Alto Amazonas, Rfo Pastaza, “una hora arriba de la boca
del Lago Rimachi,” 25 Jan. 1979, Diaz & Ruiz 914 (MO).
Madre de Dios: Manu, vicinity of Cocha Cashu Station,
7 Mar. 1977, Bell & Foster 6192 (USM); moist floodplain
forest, lake edge, 2 Oct. 1991. Burnham RJB691 (USM);
Cocha Juarez, Rfo Manu, 3—5 May 1987, Nunez, Lees &
Wright 8027 (USM); vicinity of Cocha Cashu Station, lake
margin, 3 Nov. 1976, Terborgh 5185 (USM).
3. Paullinia obovata subsp. hrentberlinei
(Croat) Weckerle & H. T. Beck, stat. nov. Bas-
ionym: Paullinia hrentberlinei Croat, Phytolo-
gia 38: 73-74. 1977. TYPE: Peru. Amazonas:
Quebrada Huampami, 10 km from its mouth
at the Rfo Cenepa, 700 ft. [ca. 200 m], 23 Nov.
1972. B. Berlin 336 (holotype, MO; isotypes,
USM. US not seen).
Distribution and habitat. Paullinia obovata
subsp. hrentberlinei is only known from the lowland
forest of northern Peru (200-300 m). It occurs
mainly in the department Amazonas along river¬
sides and in inundated areas (Fig. 1).
Phenology. No flowering specimens were ex-
150
Novon
Figure 3. Mature fruits of subspecies of Paullinia obovata. A, B. Paullinia obovata subsp. flava. —A. Yellow fruits
from the type collection plant. —B. Fruit revealing seeds with aril. C—E. Paullinia obovata subsp. obovata. —C. Bed
fruits. —D. Fruit revealing seeds with aril. —F.. |Inset| Persistent septa with arillate seeds. F', G. Paullinia obovata
subsp. subrotunda. —K. Red fruits. —G. | Inset] Fruit revealing seeds w ith aril. A. B are from Vdsquez, Weckerle &
Volume 13, Number 1
2003
Weckerle & Reynel
Paullinia obovata in Peru
151
amined. Fruiting specimens were collected during
November, January, March, and May.
Discussion. Examination of the type material of
Paullinia brentberlinei as well as other collections
reveals that this species only differs from P. obovata
subsp. polymorpha by having larger fruits (8—9 cm
in subsp. brentberlinei vs. 4—7 cm in subsp. poly¬
morpha). The vegetative features are the same as
in subspecies polymorpha , i.e., the inconspicuous
serration of the leaflets begins above the middle
and the leaflets are pubescent beneath with incon¬
spicuous hair tufts (Table 1).
Specimens examined. PERU. Amazonas: Quebrada
Huampami, Rio Cenepa. 26 May 1973, Kayap 818 (USM);
Quebrada Huampami, monte al lado de Huampami, 20
Jan. 1973. Kayap 189 (USM): Yamayakat, bosque de ri-
bera, 13 Mar. 1996. Jaramillo. Jaramillo & Chamit 1388
(USM); Bagua, Imaza-Chipe, bosque de ribera del Mara-
non, 28 May 1996, Vdsquez et al. 21026 (USM).
4. Paullinia obovata subsp. polymorpha (I). R.
Simpson) Weckerle, stat. nov. Basionym: Paul¬
linia obovata var. polymorpha I). R. Simpson,
Fieldiana 36: 155. 1976. TYPE: Peru. San
Martin: Mariseal Caeeres, Tocache Nuevo,
“bosque alto,” Quebrada de Tananta, 1 7 Dec.
1970, J. Schunke 4580 (holotype, F not seen;
isotypes, G, MOL).
Distribution and habitat. Paullinia obovata
subsp. polymorpha is found in the tropical lowland
and the foothills of the Andes at 400—800 m (Fig.
1 ).
Phenology. Flowering specimens were collect¬
ed during February and specimens with mature and
immature fruits during December, February, and
April.
Discussion. Simpson (1976) chose Schunke
4580 as the type specimen for P. obovata subsp.
polymorpha. However, this specimen does not rep¬
resent the typical character set of subspecies po¬
lymorpha. The leaves point much more to subspe¬
cies obovata (serration conspicuous, beginning
below middle) than to subspecies polymorpha, and
the fruits of the specimen, which bear the most
important characters for infraspecific delimitation,
are immature. More representative material of sub¬
species polymorpha can be seen on Schunke 4574,
for the vegetative, and Smith 1234, for the fruit
characters.
Specimens examined. PERU. San Marlin: Mariseal Ca¬
eeres, Tocache Nuevo, “bosque alto,” Quebrada de Tan¬
anta. 17 Dec. 1970. Schunke 4574 (G). Huanuco: Pach-
itea, Comunidad Nativas Santa Marta, about 50 km by
river from Puerto Inca, on bank of Sungaruyacu, 11 Apr.
1982, Smith 1234 (AMAZ, MO. MOL, USM); Pozuzo, road
between Pozuzo and Santa Rosa, 2 Feb. 2001, Weckerle <£
Igersheim 010202-1/1 (MOL), Weckerle & Igersheim
010202-1/2 (MOL).
5. Paullinia obovata subsp. subrotunda (Ruiz &
I’avon) Weckerle, stat. nov. Basionym: Semar-
illaria subrotunda Ruiz & Pa von, FI. Peruv.
Prodr. 54. I 794. Paullinia subrotunda (Ruiz &
Pavon) Persoon, Syn. PI. 1: 443, no. 15. 1805.
Paullinia obovata var. subrotunda (Ruiz & Pa¬
von) D. R. Simpson, Fieldiana 36(12): 153-
154. 1976. TYPE: Peru. No locality given: H.
Ruiz s.n. \J. Tafalla .s./j.) (lectotype, designated
by Simpson (1976). MA not seen; isolectotype,
US).
Distribution and habitat. It is ordy known from
the central eastern Andean slopes of Peru, occur¬
ring between (700-)1200 and 1800 m in the de¬
partments of Junfn, Pasco, and Huanuco. in sec¬
ondary forests, along roads and riversides (fig. 1).
Phenology. Flowering specimens were collect¬
ed during December and January and specimens
with mature and immature fruits during March and
April.
Discussion. The type collection for Paullinia
obovata subsp. subrotunda (Ruiz s.n. [Tafalla s.n. |)
was probably collected by Tafalla and sent to Ruiz.
Details supporting this were discussed by Simpson
(1976: 154).
Paullinia obovata subsp. subrotunda differs from
all other subspecies by its broader fruits, somewhat
resembling walnuts when green, and by the broader
seeds and thicker fruit walls. Unlike the lectotype,
which bears no fruits, the icon 336 in FI. Peruv. I\
is a very good reference for this subspecies.
Specimens examined. PERU. Pasco: Oxapampa, Villa
Rica-Yesu road, 28-29 Dec. 1983, Smith. Brack & Meza
5427 (MO); Oxapampa, roadside between Pusapno and
Oxapampa, 18 Mar. 2000. Weckerle & Igersheim 000318-
3/1 (MOL); Oxapampa, Pusapno, 15 Mar. 2000. Weckerle
& Igersheim 000315-2/1 (MOL), 14 Apr. 2000, Weckerle
& Igersheim 000414-1/4 (MOL), 28 Jan. 2001, Weckerle
& Igersheim 010128-1/1 (MOL); Pusapno, along Rio Pus¬
apno, 16 Mar. 2000, Weckerle & Igersheim 000316-2/1
(MOL). Junin: Villa Rica, along road to Ocanal, edge of
Igersheim 000527—1/4 (MOL, Z); C and E from Weckerle & Igersheim 000405—1/4 (MOL); I) from Weckerle A' Igersheim
000703-2/1 (MOL); F from Weckerle & Igersheim 000315-2/1 (MOL); and G from Weckerle & Igersheim 000318-3/1
(MOL). Scale bar 2 cm. Photographs by G. Weckerle & A. Igersheim.
152
Novon
“Laguna," 29 Mar. 2000. Weckerle & Igersheim 000329—
I/I (MOL), 7 July 2000. Weckerle & Igersheim 000707-
/// (MOL). Huanuco: Tingo Maria, 22 June 2000. Weck¬
erle & Igersheim 000622-1/2 (MOL).
Acknowledgments. We thank the Institute) Na-
cional de Recursos Naturales (INRENA) for the
permission (N° 006-2000-INRENA-DGF-DTCF) to
collect plants, the Instituto de Investigaciones de
la Amazonia Peruana and the staff of the Jenaro
Herrera research station for logistical and facilities
support, and the curators and staffs of the following
herbaria for loans or assistance during study v isits:
AMAZ, G, M, MO. MOL, US, USM. We also thank
R. Rutishauser, R. Nyffeler, and C. D. K. Cook for
critical comments on the manuscript, F. Stauffer for
the Spanish translation of the abstract, and A. Ig¬
ersheim for help in many aspects of the held ex¬
pedition. This project has been partly founded by
the Splinter Legal and the “Georges-und-Antoine-
Claraz-Schenkung.”
Literature Cited
Heck, H. T. 1991. The Taxonomy and Economic Botany
of the Cultivated Guarand and Its Wild Relatives and
the Generic Limits within the Paullinieae (Sapinda-
eeae). Ph.D. Thesis, City University of New York. Dis¬
sertation Abstracts AAT 9207049,
Gereau, R. E. 1993. Sapindaceae. In: L. Brako & J. L.
Zarucchi (editors). Catalogue of the Flowering Plants
and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri
Bot. Card. 45: 1059-1068.
Hamilton, C. W. & S. H. Reichard. 1992. Current practice
in the use of subspecies, variety and forma in the clas¬
sification of wild plants. Taxon 41: 485—498.
Lack, H. W. 1979. Die sudamerikanischen Sammlungen
von II. Ruiz und Mitarbeitern im Botanischen Museum
Berlin-Dahlem. Willdenowia 9: 177—198.
Pijl. van der L. 1957. On the seeds of Nephelium, Eupho¬
ria, Litchi and Aesculus, and the seeds of Sapindaceae
in general. Acta Bot. Neerl. 6: 618-641.
Radlkofer, L. 1933. Sapindaceae. In: A. Engler (editor).
Das Pflanzenreich IV 165. 1: 1-1018. Wilhelm Engel-
mann, Leipzig.
Ruiz, H. 11 777—1788] 1940. Travels of Ruiz, Pav6n, and
Dombey in Peru and Chile (1777-17815) [translated by
B. E. Dahlgren]. Field Mus. Nat. Hist., Bot. Ser. 21: 1 —
372.
- & J. Pavon. 1794. Flora Peruvianae et Chilensis
Prodromus. Madrid.
- & -. 1954. Flora Peruviana et Chilensis,
Vol. IV. Anales Inst. Bot. Cavanilles 12: 113-195.
Simpson, I). R. 1976. A partial revision of Paullinia (Sap¬
indaceae) for Ecuador, Peru and Bolivia. Fieldiana 36:
125-163.
A New Species of Ephedra (Ephedraceae) from China
Yong Yang and Dezhi Fu
Herbarium, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.
ephedra@ns.ibcas.ac.cn
Guanghua Zhu
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166—0299, U. S. A.
guanghua.zhu@mobot.org
Abstract. A new species. Ephedra rituensis Y.
Yang, 1). Z. Fu & G. Zhu, is described and illus¬
trated. This species is similar to Ephedra interme¬
dia Schrenk & C. A. Meyer in having hi- or tri-
ovulate cones, and to E. saxatilis Royle ex Florin
in having strong and well-developed woody stems
and branchlets. It differs from E. intermedia mainly
by the shorter micropylar tube straight or slightly
curved and the less connate bracts of the upper¬
most whorl of female cones, and from E. saxatilis
bv the longer micropylar tube, the bi- or tri-ovulate
female cones, more pairs or whorls of bracts, and
the smaller seeds. The new species should be as¬
cribed to Ephedra sect. Pseudobaccatae Stapf since
its female cones bear fleshy bracts at maturity.
Key words : China, Ephedra, Ephedra sect.
Pseudobaccatae, Ephedraceae.
Ephedra rituensis Y. Yang, I). Z. Fu & G. Zhu,
sp. nov. TYPE: China. Xizang: Rutog Xian,
4600 m, (9) 30 July 1974, Xizang Exped.
3676 (holotype, HNWP; isotypes, PE, XJRI).
Figure 1.
Haec species ad Ephedram intermediam et E. saxatilern
maxime accedit, sed a hac strobilo foemineo hi- vel tri-
ovulato bractearum paribus verticillisve multioribus mun-
ito, seminibus minoribus atque micropyles tubo longiore,
ab ilia micropyles tubo breviore recto vel leviter curvata
distinguitur.
Dioecious subshrub, erect, to 50 cm tall, major
stems lignified and woody, erect or procumbent,
fawn-colored, sometimes ash-gray, up to 6 mm
diam., ascendingly branched; branchlets stout,
erect or suberect, yellowish green, flat at beginning,
slightly warty when elderly, solitary, opposite or fas¬
ciculate, 1.8-2.9 mm diam.. purplish brown rned-
ullated; internodes longitudinally striate, 1.4—3.9
cm long, 1.3-2.9 mm diam. Leaves white, mem¬
branous, 3-4 mm long, 2 or 3 at nodes, with 2
purplish brown veins in middle part, connate for
2/3-4/S of total length, fused sheath cylindric, api¬
ces triangular, acute or acuminate, not recurved.
Male cones sessile, solitary or opposite, 4-6 mm
long, ovoid; bracts in 3 to 4 pairs, ovate, obtuse;
synangia 6 to 8, sessile, densely aggregated on top
of column. Female cones sessile, solitary, opposite
or clustered, 5—7 mm long, 2.8-4 mm diam.; bracts
in 4 or 5 pairs or whorls, those of uppermost whorl
broadly ovate, apex obtuse, connate for 1/3—1/2
their length, red and fleshy at maturity, other whorls
of bracts smaller and distinct from each other.
Seeds 2 or 3, concealed by bracts, purplish brown,
ovoid, ca. 5 mm long, ca. 3 mm wide, smooth, dor-
sally convex and ventrallv plane, or carinate; mi¬
cropylar tube straight or slightly curved, ca. 2.2 mm
long, exserted.
Etymology. The specific epithet is derived from
Chinese ping-yin of the county where the holotype
of Ephedra rituensis was collected.
Distribution. Ephedra rituensis is found in some
inaccessible localities of the Tibetan Plateau. Most
specimens examined were collected from Xizang,
China, but a few were found in Yecheng of southern
Xinjiang and Qinghaihu of Qinghai, China.
Habitat. Newly described Ephedra rituensis
grows in slightly divergent habitats at 3200—4600
m. The holotype was found in a dried river beach
at 4600 m; other examined specimens were discov¬
ered in crevices of rocks, sand dunes, or dried grav¬
el river beaches at 3200-4270 m. This species may
live in a relatively moist condition since most spec¬
imens were collected near a river or lake.
Relationships. This new species is similar to
Ephedra intermedia in having bi- or tri-ovulate
cones, and to E. saxatilis in having strong and well-
developed woody stems and branchlets. It differs
from E. intermedia mainly by its shorter micropylar
tube straight or slightly curved, the less connate
bracts of the uppermost whorl, and the narrower
female cones, and from E. saxatilis by its longer
micropylar tube, bi- or tri-ovulate female cones,
Novon 13: 153-155. 2003.
154
Novon
Figure 1. Ephedra rituensis Y. Yang, D. Z. Fu & G. Zhu (holotype). —A. Habit, female plant, X], —B. Seed, X6.
—C. The uppermost whorl of bracts to show the degree of fusion, X2. —D. Female cone, X3.
Volume 13, Number 1
2003
Yang et al.
Ephedra rituensis from China
155
smaller seeds, and more pairs or whorls of bracts
of female cones (Meyer, 1846; Stapf, 1889; Florin,
1933; Fu et al., 1999).
The most widely used sectional classification of
Ephedra was made by Stapf (1889) based on the
nature of bracts of female cones, although there are
different opinions (Soskov, 1968; Pachomova, 1971;
Mussayev, 1978). Bracts of female cones in Ephe¬
dra at maturity may be fleshy, coriaceous, or mem¬
branous, representing section Pseudohaccatae,
section Asarca, and section Alatae, respectively
(Stapf, 1889). This newly described species clearly
belongs to Ephedra sect. Pseudohaccatae.
Paratypes. CHINA. Xizaug: Burang Xian, (9) 18 Aug.
1974, Tibet Exped. 3999 (HNWP, XJ8I); Burang Xian,
(3), Vegetation Group , Qianghai-Xizang Exped. 13006
(PE); Rutog Xian. (9) 14 Sep. 1976, Vegetation Group ,
Qinghai-Xizang Exped. 13665 (PE); Gyirong Xian, (9) 30
July 1975, Wu C. Y., Chen S. K., Du Q. & Lu S. I, 75-
493 (PE); Zhag'yaf) Xian, (9) I Sep. 1976. Qingzang Ex¬
ped. 12981 (PE). Xinjiang: Yeeheng, (9) 23 Aug. 1987,
Wu Y. //. 1172 (PE). Qinghai: Qinghaihu, 1 July 1958.
Tsoong Pu-Chiu 8329 (PE).
Acknowledgments. We thank Helmut Freitag,
Boy Gereau. and Maria Maier-Stolte for their valu¬
able comments, ami Yingbao Sun for preparing the
illustration. We are grateful to the curators of
HNWP, PE, and XJB1 for the loans of specimens.
This paper is supported by Project Ksxc2-sw-108
(Chinese Academy of Sciences).
l.iterature Cited
Florin, R. 1933. fiber einiger neue oder wenig bekannte
asiatische Ephedra- Arten der Sect. Pseudohaccatae
Stapf. Kongl. Svenska Vetensk. Acad. Handl. 12: 1—44.
Fu, L. K.. Y. F. Yu & H. Riedl. 1999. Ephedraceae. Pp.
97-101 in C. Y. Wu & P. Raven (editors). Flora of Chi¬
na, Vol. 4. Science Press, Beijing, and Missouri Botan¬
ical Garden Press, St. Louis.
Meyer, C. A. 1846. Versuch einer Monographic der Gat-
tung Ephedra. Mem. Acad. Imp. Sci. St. Petersbourg,
Ser. 6 (Sci. Nat.) 5: 225—298.
Mussayev, I. F. 1978. On geography and phylogeny of
some representatives of the genus Ephedra L. Bot.
Zhurn. (Moscow & Leningrad) 63: 523—543.
Pachomova, M. G. 1971. Ephedraceae. Pp. 25—33 in V. I.
Gluhov, M. G. Matzenko & M. G. Pachomova (editors),
Plantae Asiae Centralis, Vol. 6. Academia Scientiarum
LRSS Institutum Botanicum nomine V. L. Komarovii,
Leningrad.
Soskov, L. I). 1968. Three lines of development within the
section Ephedra of the genus Ephedra L. in the U.S.S.R.
Bot. Zhurn. (Moscow & Leningrad) 53: 85—91.
Stapf, 0. 1889. Die Arten der Gattung Ephedra (Mono¬
graph). Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-
Naturwiss. Kl. 56(2): 1-112.
Rhododendron maxiongense (Ericaceae), a New Species from
Yunnan, China
Zhang Chang-Qin
Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming, Yunnan
650204, People’s Republic of China, zhangchangqin@mail.kib.ac.cn
D. Paterson
Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland,
United Kingdom. D.Paterson@rbge.org.uk
Abstract. Rhododendron maxiongense C.-Q.
/hang & D. Paterson (Ericaceae), from Yunnan,
China, is described as new. It is similar to R. aber-
conwayi Cowan, which has elliptical thick and co¬
riaceous leaves witli recurved margins that are not
undulate and have red punctate hair. The calyx of
R. aberconwayi is sparsely hairy and glandular-cil-
iate; the corolla is open-campanulate, white to pale
rose, with purple flecks; the pedicels are sparsely
hairy and stipitate-glandular; and there is pilus on
the base of the stamen. Rhododendron maxiongense
differs from R. aberconwayi in its habitat as a dwarf
shrub 0.6 to 0.9 m in height; leaves with undulated
recurved margins without red punctate hair; inflo¬
rescence 8- to 13-flowered; corolla white, without
(leeks, apex pale green or pale rose, and lobe apex
acute; stamens glabrous; calyx densely hairy; and
pedicels densely hairy.
Key words: China, Ericaceae, Rhododendron,
Yunnan.
Rhododendron I,, is a large genus of the Erica¬
ceae containing approximately 960 species world¬
wide, of which about 542 species (no! including
subspecies arrd varieties) are distributed in tire wild
in western China (Fang, 1999), with 227 species in
Yunnan (Fang, 1999) (Cullen, 1980; Chamberlain,
1982; Mei & Fang, 1986). H. Sleurner divided the
genus into 8 subgenera, but J. Cullen and 1). E.
Chamberlain divided it into 5 subgenera.
When investigating plants of Rhododendron in
Maxiongshan, northeast Yunnan, southwest China,
in May 2000, sorrre specimens were collected and
all but one was identified. After studying the un¬
identified specimen and literature, and checking
other specimens deposited in KUN, we believe it
represents a new species within subsection Irror-
ata of subgenus Hymenanthes. Subsection Irrorata
contains 21 species in China, of which 16 are
No VON 13: 156-158. 2003.
found in Yunnan (Hu & Fang, 1994; Chamberlain,
1982).
Rhododendron maxiongense C.-Q. Zhang & I).
Paterson, sp. nov. TYPE: China. Yunnan:
Zhanyi Maxiongshan, 25°35'N, 103°50'E,
2440 m, 6 May 2001, C.-Q. Zhang 2001-01
(holotype, KUN). Figure 1.
Species affinis R. aberconwayi, sed pumilo frutice 0.6-
0.9 m, foliis rubris puncticulosis trichomatibus carentibus,
inflorescentiis 8-13-fforis, purpureis punctis carentibus,
apice viridi-albo vel rubello, cafyce velato confertim pilis,
staminibus glabrescentibus, differt.
Dwarf shrub 0.6-0.9 m high; young shoots
sparsely tomentose or glandular, bark glabrous,
lea ves clustering at the end of shoots; leaf blades
elliptic or ovate-elliptic, thick and coriaceous, 3-6
X 1.2-2.5 cm, apex acute or mucronate, base near¬
ly rounded, margin undulate and recurved; upper
blade surface green, glabrous, lower blade surface
pale green; midrib strongly sunken above and
raised beneath, lateral veins in 10 to 13 pairs,
sunken on upper surface and raised beneath; blade
petioles 0.7—1 cm, grooved above, with densely
glandular hair; racemose umbellate inflorescences
8- to 13-flowered, rachis ca. 1-2 cm with white
trichomes; pedicels 0.8—2 cm, with densely white
glandular trichomes; calyx lobes 5, triangular, 1—2
mm, densely hairy, glandular-ciliate. Corolla open-
campanulate, 1.5—2 cm long, diameter at mouth 2—
4.5 cm, lacking nectar pouches, glabrous within,
white, without dots, apex pale green or pale rose;
corolla lobes 5, apex acute, 1-1.2 X 1.4-1.6 cm;
stamens 10, glabrous, anthers long-rounded, ca. 2
mm long, ovary conical, ca. 3.5 X 3 mm, densely
glandular, style shorter than corolla, 1.3—1.7 cm
long, glandular throughout, stigma pale green or
pale rose, capsule unseen.
Distribution. Rhododendron maxiongense is
Volume 13, Number 1
2003
Zhang & Paterson
New Rhododendron from China
157
Figure 1. Rhododendron maxiongense C.-Q. Zhang & I). Paterson. —A. Inflorescence. — B. Petiole showing setose
glandular hair. — C. Acute apex of corolla lobes. — D. Pedicel and calyx hair. — D-l. Pedicel glandular hair. E.
Stamen. —F. Gynoecium. All drawn front the type collection, C. Q. Zhang 2001-01 , by \ang jian-kun.
known only from Zhanyi County, northeast Yunnan.
China. This species grows on slopes of open Firms
yunnanensis Franchet var. pygmaea (J. R. Xue) J.
R. Xue forests at 2440 m above sea level.
Rhododendron maxiongense resembles R. aber-
conwayi : both are evergreen shrubs with open-cam-
panulate flowers, from northwest Yunnan, China.
However, the latter species differs in its larger habit
(1.5 to 2.5 m high); inflorescence with 6 to 12 flow¬
ers, and its corolla with purple flecks inside and
emarginated lobes; the base of the stamen has pi-
lus.
Morphologically, Rhododendron maxiongense, R.
aberconwayi, and 17 other species including R. ir-
rorala Franchet and R. agastum Balfour f. & W. W.
Smith all belong to subsection Irrorata. These are
shrubs or small trees; bark rough; young shoots
stipitate-glandular; leaves ovate to elliptic or ob¬
long, lower surface usually glabrous when mature
but with persistent punctate base with a thin veil
of dendroid hairs embedded in a surface film; in¬
florescence lax or dense, 4- to 20-flowered; rhachis
5—10(—35) mm, calyx minute or cupular, with broad
lobes ami up to 6 mm long. Corolla 5- to 7-lobed,
tubular to open-campanulate, with or without nectar
pouches, white (rarely yellow) to mauve or deep
158
Novon
crimson, usually with darker flecks and stipitate-
glandular; style glabrous or glandular to tip (Sleu-
mer, 1949). This subsection comprises about 21
species in China, 16 in Yunnan (Hu & Fang, 1994;
Chamberlain, 1982).
Acknowledgments. The study is supported by the
Chinese Academy of Sciences (the Knowledge In¬
novation Project and grant number KZI-12) and the
Nature Science Foundation of Yunnan (number
98co90M). Special thank s are due to T. L Mei, who
made constructive suggestions. Thanks also to Yang
jian-kun, who prepared the illustration.
Literature Cited
Chamberlain, I). K 1982. A revision of Rhododendron II.
Subgenus Hymenanthes. Notes Roy. Hot. Card. Edin¬
burgh 39(2): 209-486.
Cullen, J. 1980. A revision of Rhododendron subgenus
Rhododendron sections Rhododendron & Pogonanthum.
Notes Roy. Hot. Gard. Edinburgh 39(1): 1—207.
fang, Rh. Ch. 1999. Flora Reipublicae Popularis Sinicae,
Tomus 57(1). Science Press, Beijing.
Hu, L. Ch.& M. Y. Fang. 1994. Flora Reipubl ieae Popu¬
laris Sinicae, Tomus 57(2). Science Press, Beijing.
Mei, T. I. & Fang Ruieheng. 1986. Flora Y unnaniea, To¬
mus 4. Science Press, Beijing.
Sleumer, M. 1949. Fin System derGattung Rhododendron.
Bot. Jahrb. 74: 511-533.
Erratum
Crandall-Stotler, B. J., R. E. Stotler & C. H. Ford.
2002. Contributions toward a monograph of
Petalophyllum (Marchantiophyta). Novon 12:
334-337.
We thank R. Grolle (JE) for calling to our
attention the misspelling of Petalophyllum
hodgsonii sp. nov. (p. 336). In accordance
with Article 60.11 of ICBN, the termination of
the epithet “ hodgsonii ” should be corrected
to “ hodgsoniae .”
Novon 13: 159. 2003.
Volume 13, Number 1, pp. 1-160 of NOVON was published on 25 March 2003.
Volume 13
Number 2
2003
NOVON
Chromosome Cytology and Taxonomy of the Red Goblet-Flowered
Species of Clidemia (Melastomataceae: Miconieae) in
Central and South America
Frank Almeda
Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco,
California 94118-4599, U.S.A. falmeda@calacademy.org
Abstract. Clidemia clandestina (Melastomata-
ceae) of Costa Rica and Panama is newly described
as part of a review of the goblet-flowered species
from tropical America. This natural group of three
species, including C. garciabarrigae and C. utley-
ana, is characterized by 5-merous flowers that are
goblet-shaped in profile view with erect red petals
that are connivent and cucullate at anthesis, minute
anther thecae with wide ventrally inclined apical
pores, and a completely inferior 3-locular ovary. An
original chromosome count of n = 17 is reported
for C. utleyana . and a count of n = 23 is reported
for C. clandestina and C. garciabarrigae. The sig¬
nificance of these chromosome numbers is evalu¬
ated in the context of previous reports for the tribe
Miconieae together with a diagnostic key, descrip¬
tions, illustrations, new distributional information,
phenological notes, and citations of representative
specimens.
Key words: Central America, chromosome cy¬
tology, Clidemia, Melastomataceae, Miconieae, red
goblet-flowered species, South America.
Clidemia I). Don sensu lato, a berry-fruited ge¬
nus of Melastomataceae with over 170 species, is
diverse and widespread in the American tropics but
still little-known and undercollected in several
parts of its range. The three species treated here
are collectively distributed from Costa Rica and
Panama disjunctly south to Colombia and north¬
western Ecuador where they occur in some of the
wettest environments of the two contiguous biodi¬
versity hot spots in the region (Mittermeier et al.,
1999). They constitute a distinctive assemblage
characterized by their scandent or reclining (some¬
times epiphytic) habit, completely inferior 3-locular
ovary, 5-merous flowers that are goblet-shaped in
profile view (Figs. 21), 3D) with unusual red petals
that are erect, connivent, and conspicuously con¬
cave at anthesis (Figs. 2E, 3H). In addition, the
anther thecae are minute, incurved, and have com¬
paratively wide ventrally inclined apical pores. A
taxonomic account of this species group is pre¬
sented to facilitate the description of C. clandestina
from Costa Rica and Panama and to provide the
first chromosome information for each of the three
species.
Chromosome Cytology
The first gametic chromosome number reports
are presented here for one population each of C.
clandestina, C. garciabarrigae, and C. utleyana. Al¬
though some pollen mother cells of both C. clan¬
destina and C. garciabarrigae showed some lagging
chromosomes, meiosis for the most part appeared
to be regular with n = 23 for the few r cells exam¬
ined (Fig. 1 A—1C). In C. utleyana meiosis was con¬
sistently regular with n = 17 in all cells examined
(Fig. ID). Bud material for these counts was col¬
lected from natural populations in the field, fixed
in modified Carnoy’s solution (4 chloroform, 3 eth¬
anol, 1 glacial acetic acid, v/v/v, Bradley, 1948),
transferred to 70% ethanol, anti stored under re¬
frigeration. All counts were made with a Zeiss light
Novon 13: 161-169. 2003.
162
No von
A
C
B
D
i_i
20^m
F igure 1. Camera luciila drawings of meiotic chromosome figures of Clidemia. —A, B. C. clandestina, n = 23, meta¬
phase I (A l me da et al. 5907). —C. C. garciabarrigae , n = 23, metaphase 1 (Almeda el al. 7504). —D. C. utleyana , n
= 17, metaphase I (Almeda et al. 6864).
microscope with phase contrast and a 100X oil im¬
mersion objective. Drawings of meiotic figures were
made with camera lucida at a magnification of
2I00X. Voucher collections for the counts reported
here are marked with an asterisk among the spec¬
imens cited.
The count of n = 17, reported here for C. utley¬
ana , is the most commonly reported gametic chro¬
mosome number in the tribe Miconieae and appears
to he the base number for Clidemia and several
related genera (Almeda & Chuang, 1992; Suit &
Wurdack, 1980). Ten of the 14 genera in the Mi¬
conieae investigated to date have x = I 7 and the
other four genera have n = 15, 20, 28. and other
numbers that can best be interpreted as dysploid
derivatives of polyploid numbers (Almeda, 1997a).
Almeda and Chuang (1992) speculated that n = IT
may have arisen as a dysploid derivative of an au-
totetraploid based on x = 9 or by ancient hybrid¬
ization of species with 7 and 10 or 8 and 9 haploid
chromosome numbers. Because chromosome num¬
bers ranging from x — 9 through x = 12 are known
for several genera in the Melastomataceae and sev¬
eral families in the Myrtales (Almeda, 1997a,
1997b), it is likely that some number in this range
is the original base chromosome number in the
Volume 13, Number 2
2003
Almeda
Red Goblet-Flowered Clidemia
163
family. Thus, a polyploid origin for the Miconieae
is clearly indicated, but unlike other tribes in the
family, the Miconieae appear to have no extant taxa
with lower base numbers that could qualify as an¬
cestral base numbers (Almeda. 1997a).
Goldenberg and Shepherd (1998) demonstrated
a close association between high chromosome num¬
ber (above n = 17), low but highly variable pollen
viability, and apomixis. Gametic chromosome
counts of n — 23 and 24 have been reported for
several species of Clidemia and Miconia (Almeda,
1997a; Almeda & Chuang, 1992; Suit & Wurdack,
1980), and those species with these higher numbers
that have been investigated experimentally have
been shown to be apomictic (Goldenberg & Shep¬
herd, 1998). Apomixis has been reported or sus¬
pected in several species of Melastomataceae, a
majority of which are concentrated in the tribe Mi¬
conieae (Almeda & Chuang, 1992; Goldenberg &
Shepherd, 1998; Renner, 1989).
Access to good anthers with pollen was limited
for most goblet-flowered species of Clidemia ex¬
amined for this study. Pollen viability estimated for
one individual of each species (based on a sample
of 200 grains stained with aniline blue in lacto-
phenol for 24 hrs.) was 51% for C. utleyana ( Al¬
meda 6864), 0% for C. clandestina (Croat 67238),
and 5% for C. garciabarrigae (0llgaard 37666).
Judging from the extremely low pollen viability for
the latter two species coupled with their higher
chromosome number (n = 23), one may logically
predict that they are also apomictic. Study of ad¬
ditional populations and emasculation experiments
are needed, however, for all of the taxa treated here
in order to confirm the occurrence of apomixis and
to better evaluate any morphological, cytological, or
environmental correlates with this breeding system.
Key to the Red Goih.et-Feowered Species of
Clidemia
la. Leaf blades glabrous on the adaxial (upper) sur¬
face; inflorescence a multibranched panieuliform
dichasium; bracts and bracteoles of the inflores¬
cence widest at the base.
2a. Leaf blades essentially Hat on the adaxial
surface; abaxial (lower) surface with spread¬
ing smooth hairs restricted to the blade base
where the primary veins diverge from the
median vein; hypanthia minutely and decid-
uously brown-lepidote; petals copiously cov¬
ered with minute papillae on both surfaces
. C. clandestina
2b. Leaf blades bullate on the adaxial surface;
abaxial leaf surface with smooth spreading
hairs covering all the elevated veins and
veinlets; hypanthia sparsely to moderately
covered with smooth spreading hairs and a
ground layer of scattered minute glandular-
punctate hairs; petals completely smooth on
both surfaces. C. utleyana
lb. Leaf blades pubescent on the adaxial surface;
inflorescence a solitary or trifurcate collection of
racemes; bracts and bracteoles of the inflores¬
cence widest above the base . ... C. garciabarrigae
1. Clidemia clandestina Almeda. sp. nov. TYPE:
Panama. Cocl6: Cerro Gaital, E slope and ridg¬
es leading to the summit with montane rain¬
forest and elfin woodland, 8°40'N, 80°07'W,
elev. 1050-1100 m, 24 Feb. 1988, F Almeda
et al. 5907* (holotype, CAS; isotypes, AAU,
CR. INB, MEXL, MO, NY, PM A). Figure 2.
Frutex terrestris vel epiphyticus. Hamuli primum quad-
rangulati demum teretes glabri. Lamina 5.7—14 X 3—7.3
cm elliptica vel elliptico-ovata apice acuminato vel cau-
dato-acuminato basi obtusa vel rotundata, 3—5-plinervata,
supra glabra, subtus in venarum primariarum axillis (et
supra axillis) modice setosa pilis simplicibus 0.5—1 mm
longis persistentibus. Inflorescentia 3—8.5 cm multiflora;
flores 5-meri; calyeis tubus 0.1—0.25 mm alius, lobis in-
terioribus 0.25 X 0.5 mm triangularibus, dentibus exter-
ioribus subulatis 0.25—0.5 mm eminentibus. Stamina iso-
morphica glabra; antherarum thecae 0.25 X 0.25 mm
oblongae exappendiculatae, poro ventraliter inclinato.
Ovarium 3-loculare et omnino inferum apice glabro.
Erect or scandent terrestrial shrub 1-2.5 m tall
or dangling epiphyte. Internodes ± quadrate, be¬
coming rounded with age, and sometimes bearing
adventitious roots but otherwise essentially gla¬
brous. Young vegetative buds, pedicels, and hypan¬
thia minutely and caducously brown lepidote.
Leaves of a pair ± equal in length or somewhat
unequal but never markedly so; petioles 0.6—1.5
cm long; blades ± coriaceous when dry, 5.7—14 X
3—7.3 cm, elliptic sometimes varying to elliptic-
ovate, apex acuminate to caudate-acuminate, base
obtuse to rounded, margin entire to inconspicuously
ciliate-serrulate, 3- to 5-plinerved with the inner¬
most pair of primary veins diverging from the me¬
dian vein 0.2-0.8 cm above the blade base, gla¬
brous adaxially and moderately lepidote abaxially
on the actual surface with hair-tuft domatia bearing
hairs 0.5—1 mm long along the basal portion of the
primary veins and where the innermost primaries
diverge from the median veins at the blade base.
Inflorescence a terminal modified dichasium 3—8.5
cm long becoming pseudolateral with elongation of
lateral branches, sometimes divaricately branched
from the base; bracts and bracteoles subulate to
narrowly triangular, 0.5—1 mm long and about 0.25
mm wide at the base. Pedicels 0.5—1 mm long. Hy¬
panthia (at anthesis) campanulate, ca. 1 mm long
from the base to the vascular ring. Calyx tube 0.1—
0.25 mm long, the calyx lobes 0.25 X 0.5 mm,
bluntly triangular, hyaline and typically obscured
164
Novon
Figure 2. Clidemia clandestina Almeda. —A. Habit. —B. Representative leaf (abaxial surface) at left and leaf base
(abaxial surface) at lower right. —C. Flower bud showing pedicel and subtending bracteoles. —I). Flower at anthesis
(profile view) with one petal removed. —E. Petal (3/4 profile view) on left and enlargement of abaxial surface on right.
—F. Stamens, ventral view on left and profile view on right. —G. Berries. —H. Seeds. (A—II from Almeda el al. 5907.)
Volume 13, Number 2
2003
Almeda
Red Goblet-Flowered Clidemia
165
by the exterior teeth; ealyx teeth 5, linear to su¬
bulate, 0.25—0.5 mm long. Petals 5, copiously cov¬
ered with minute papillae on both surfaces, dark
red but drying maroon-black, obovate, erect and
concave adaxially at anthesis. 1 mm long and 1 mm
wide. Stamens 10, isomorphic and forming an in¬
curved circle around the style; filaments glabrous,
1 mm long, gradually tapered from the base to the
apex, pink but drying red; anthers 0.25 mm long
and 0.25 mm wide, pale yellow flushed with crim¬
son-maroon basally, ± oblong, rounded at the apex
with a ventrally inclined pore; connective crimson-
maroon but drying purple-black, thickened at the
base of the anther sac but not conspicuously pro¬
longed or appendaged. Ovary completely inferior,
3-locular, apex glabrous. Style straight, glabrous,
1.25 mm long; stigma blunt and rounded to ± trun¬
cate. Berry maroon when immature but turning pur¬
ple-black when ripe, 4-5 X 4-5 mm when dry.
Seeds ovoid to obovoid in outline, I mm long,
bluntly and complexly tuberculate on the convex
(ace.
Phenology. Collected in flower and fruit during
February, March, July, August, November, and De¬
cember.
Distribution. Local and uncommon in rain for¬
ests, cloud forests, and elfin woodland from north¬
ern and central Costa Rica disjunctly to western
and central Panama at 700—1400 m.
Although collections of this species were first
made in 1979, followed by several more in the
1980s, none had intact floral parts for critical anal¬
ysis. Like so many Melastomataceae, the floral
parts of this species are especially fugacious and
unless care is taken to press flowers separately or
preserve them in alcohol they are commonly absent
on herbarium specimens. Details of floral morphol¬
ogy presented here were discernable only after I
had the opportunity to study and collect this spe¬
cies in Panama in 1996.
Clidemia clandestina is readily distinguished
from C. garciabarrigae and C. utleyana by its gla¬
brous internodes and leaf blades that are glabrous
adaxially and moderately lepidote abaxiallv on the
actual surface with tufts of smooth spreading hairs
along the basal portion of the primary veins and
where the innermost primaries diverge from the me¬
dian veins at the blade base. In addition, the young
vegetative buds, pedicels, and hvpanthia are mi¬
nutely and caducously brown lepidote and the pet¬
als are distinctly papillose on both surfaces. The
flowers of C. clandestina are small, dark red in col¬
or, and inconspicuous. The inflorescences are typ¬
ically shorter than subtending leaves and often as¬
sume a position hidden under the blades. The
specific epithet for this species, which is derived
from the Latin word clandestinus , secret or hidden,
emphasizes these features.
Paratypes. COSTA RICA. Alajuela: Canton de San
Ramon, Bosque Eterno de Los Ninos, Cordillera de Tilar-
an, Valle del Rfo Penas Blancas, 7 Oct. 1993, Bello &
Cruz 5364 (INB); Upala, Bijagua, El Pi Ion. Cabeceras del
Rfo Celeste, 10°49'N, 84°27'W. 13 Nov. 1987. Herrera
1263 (CAS, CR, MO); along road from San Ramon north¬
ward through Balsa, ca. 16.7 km N of bridge over Que-
brada Volio and ca. 7.5 km N of bridge over Rfo Balsa,
10° 10— 15'N, 84°30-35'W, 29 Aug. 1979, Stevens 13890
(CAS, CR, MO). Lirndn: Canton de Talamanca, Rratsi,
Amubri, Alto Lari. Kivat, bajando haeia el Rfo Dapari,
margen derecha, 9°24'20"N, 83°05'35"W, 10 Mar. 1992,
Herrera 5280 (CAS, INB, MO); Cantrtn de Talamanca,
Rratsi, Amubri, Alto Lari, Kivut, fila mayor entre los Rfos
Dapari y Lari, desvidndose hacia el este por fifa secun¬
daria, cuenca del Rfo Lari. 9°23'50"N, 83°05'10"W, 17
Mar. 1992, Herrera 5383 (INB); Canton de Talamanca, lila
de exploracion minera entre Rfo Sukut y Rfo Carbri, Cerro
Schtomat, 9°22'40"N, 82°56'30"W, 13 July 1989, Solis et
td. 21 (INB). PANAMA. Chiriqui: Lortuna Dam area
about 3.7 km S of the Sitio de Presa offices, trail through
Ouebrada Aleman, 9 Feb. 1996, Almeda et al. 7565 (CAS,
MO. PMA); Lortuna Dam area. North Quebrada de Arena
along river, 8°46'N. 82°12'W, 9 Feb. 1984, Churchill et
al. 4709 (MO). Code: vicinity of La Mesa, N of LI Valle
de Anton, along E edge of Cerro Gaital on hogback ridge
leading to summit, 8°37’N, 80°08’W, 13 July 1987. Croat
67238 (CAS); Cerro Gaital, elfin forest, 8°37’N, 80°6’W,
26 Nov. 1984, de Nevers & Charnley 4379 (CAS); slopes
and summit of Cerro Gaital, N of LI Valle, 8°40'N,
80°07'W, 10 July 1982, Knapp et al. 6008 (CAS); Divide
SW of La Mesa at end of logging road, 80°5‘W. 8°35’N,
26 Dee. 1982, Stein & Hamilton 998 (CAS, MO).
2. Clidemia garciabarrigae Wurdack, Phytolo-
gia 7: 239. I960. TYPE: Colombia. Narino:
between Altaquer and Ricuarte along the road
to Barbacoas, 1140—1300 m, 3—5 Aug. 1948,
H. Garcia-Barriga 13116 (holotype, US; iso¬
type, COL). Figure 3.
Terrestrial or epiphytic weakly branched shrub
with pendent or arching branches 1—2 m long typ¬
ically requiring the support of adjacent vegetation.
Internodes ± terete and covered with smooth,
eglandular, spreading hairs (0.5—)1—3 mm long like
the inflorescence raehis and both foliar surfaces;
the nodes ± swollen. Leaves of a pair dimorphic
in size and shape; blades subcoriaceous when dry,
the larger one at each node, 6-10.2 X 2-5.4 cm,
ovate-elliptic to ovate-oblong, apex abruptly acu¬
minate, base broadly rounded to subcordate, mar¬
gin entire, 5-nerved or 5-plinerved, the innermost
pair of primaries diverging from the median vein
2—3 mm above the blade base; petiole 3—5 mm
long; the smaller blade 1.7—4.4 X 1.5—3.6 cm,
broadly ovate, apex short-apiculate, base cordate,
166
Novon
2 mm
1 mm
Figure 3. Clidemia garciabarrigae Wurdack. —A. Habit. —B. Representative leaves at a node, larger leaf showing
abaxial surface (left) and smaller leaf showing abaxial surface (right). —C. Floral bracteoles (abaxial surfaces). —D.
Flower at anthesis (profile view) with one petal removed. —E. Immature berry (profile view). —F. Stamens, ventral
view on left and profile view on right. —G. Seeds. —H. Petal (3/4 profile view). (A, G from Hampshire & Whitefoord
738; B—F, H from Almeda el al. 7504.)
Volume 13, Number 2
2003
Almeda
Red Goblet-Flowered Clidemia
167
margin entire, 3-5-nerved, petiole barely prolonged
and subsessile or up to 1 mm long. Inflorescence a
trifurcate raceme or occasionally a solitary raceme
3.5— 5 cm long that is at first terminal but becomes
pseudolateral with elongation of lateral branches;
bracteoles elliptic to linear-oblong or somewhat ob-
ovate, tardily deciduous, 2—4 X 0.25—2.5 mm. gla¬
brous adaxially, sparsely to moderately covered
abaxiallv with smooth spreading hairs and a sparse
ground layer of small glands. Pedicels (at anthesis)
0.25—1 mm long. Hypanthia (at anthesis) campan-
ulate to cupulate, 1—1.5 mm long from the base to
the vascular ring, sparsely covered with smooth
spreading hairs 0.5—1.5 mm long and a ground lay¬
er of scattered rusty-brown glandular hairs. Calyx
tube obsolete, the calyx lobes 0.5—0.8 X 0.75 mm,
rounded-triangular, somewhat hyaline and mostly
obscured by the exterior teeth; calyx teeth 5, nar¬
rowly triangular, 1 X 0.75 mm. Petals 5. glabrous,
dark red but drying maroon-black, suborbicular,
erect, connivent and concave adaxially at anthesis,
1.5— 2 X 1.5—2 mm. Stamens 10, isomorphic and
forming an incurved circle around the style; fila¬
ments glabrous, 1—1.5 X 0.25 mm, gradually ta¬
pered from the base to the apex; anthers 0.5 X 0.5
mm, pale yellow apically but deep red toward the
base, ± oblong to somewhat rectangular in outline,
± rounded apically with a ventrally inclined pore;
connective crimson-maroon but drying purple-
black, thickened and ± truncate ventrally at the
base of the anther sac but not prolonged or appen-
daged. Ovary completely inferior, 3-locular, apex
glabrous and 10-costate. Style straight, glabrous, 2—
2.5 mm long; stigma truncate to ± rounded. Berry
changing through green, red, blue, and ultimately
black when ripe, 4-5 X 4—5 mm when dry. Seeds
± obovoid in outline, 1 mm long, tuberculate on
the broader distal end.
Phenology. Collected in flower and fruit from
February through April and in September and No¬
vember.
Distribution. Local anti uncommon in open ar¬
eas in cloud forests and lake margins from disjunct
localities in western and central Panama to south¬
ern Colombia and northern Ecuador at 700—1300
m. fhe label on a collection from Carchi, Ecuador
(Tipaz et al. 1035), gives the elevation as 3200 m.
This is extraordinarily high for this species and
should be regarded with suspicion until confirmed.
In the protologue, this species was reported only
from Colombia. It was first collected in Ecuador in
1979. Although it wasn't included in Wurdack’s
(1980) melastome treatment for Ecuador, it was re¬
ported from that country shortly thereafter (Wur-
dack, 1982: 61). The range of C. garciabarrigae is
here extended north to Panama, making it the most
widespread member of the globlet-flowered species
ol Clidemia.
Clidemia garciabarrigae differs from the other
two species treated here in having an indument ol
smooth, spreading hairs on both foliar surfaces, leaf
blades that are conspicuously dimorphic in size and
shape at each node, and an inflorescence that typ¬
ically consists of trifurcate (sometimes varying to
solitary) racemes. It also differs from both conge¬
ners by its floral bracteoles that are widest well
above the base.
Representative specimens examined. ECUADOR. Car-
chi: El Pailon, ca. 45 km below Maldonado along a foot¬
path to Tobar Donoso, 2 Nov. 1979, Madison & Hesse 7093
(US); Cantdn Parroquia Chieal, sector Gualpi medio, Re-
serva Indfgena Awa, sendero a San Marcos al norte de la
oasa comunal. 78°16'W, 01°02'N, 23—27 May 1992. Tipaz
et (d. 1035 (CAS, MO). Esmeraldas: San Lorenzo Canton
Reserva Indfgena Awa, canon del Rfo Mira, 10 km al
oeste de Alto Tambo, 78°26'W, 01°02'N, 16—26 Mar.
1991, Rubio et al. 1227 (US). Pirliinrlia: Road El Par-
afso—Saguangal, I I km from El Parafso, 78°46'W, 0°]2'N,
2 May 1982, 0llgaard et al. 37666 (CAS). PANAMA.
Chiriqui: Edwin Eabrega Dam and Reserve in Fortuna,
Quebrada Bonita ca. 7.5 km N of Sitio de Presa office
enroute to Chiriqui Grande, 5 Feb. 1996, Almeda et al.
7504* (BM. CAS, INB. MO, MEXU, PM A); Gualaca-Chi-
riquf Grande, 4—8 mi. beyond IRHE facilities at Dam, 4
mi. N of bridge over Bayano Lake, 8°46'N, 82°16'W, 23
Sep. 1987, Croat 6H0I9 (CAS, MO, PMA): Fortuna Lake,
cleared forest on banks of lake, 8°45'N, 82°12'W, 20 Mar.
1985. Hampshire & Whitefoord 733 ((IAS. MO): Fortuna.
camino de Quebrada Bonita llegando por el embalse hacia
el norte, 8 Apr. 1987, Valdespino et al. 596 (CAS. PMA).
Code: forested slopes above El Cope along abandoned
road leading to the Continental Divide, 25 Feb. 1988,
Almeda et al. 5944 (CAS).
3. Clidemia utleyana Almeda, Bull. Torrey But.
Club 106: 189. 1979. TYPE: Costa Rica. Car¬
tage: continuation of C.K. #224 on the prop¬
erty of ICE hydroelectric plant (now Tapantf
National Park), about 4.4 km beyond main
gate to property and 17 km beyond the Church
in Orosf. elev. 1500-1800 m, 13 Dec. 1975,
F. Almeda & J. Utley 2666 (holotype. CAS;
isotypes, CR, F, MO, US).
Laxly branched erect to reclining terrestrial or
epiphytic shrub mostly 0.5—1.5 m tall. Internodes
and uppermost branchlets terete, moderately to co¬
piously beset with smooth, eglandular, spreading
hairs mostly 1—2 mm long like the inflorescence
rachis. Leaves ol a pair somewhat unequal in size,
otherwise identical in shape and indument details,
blades chartaceous to subcoriaceous when dry, en¬
tire but conspicuously revolute, narrowly ovate to
168
Novon
± cordate, 5- to 7-nerved, glabrous and bullate
adaxially, the abaxial surface beset with a mixture
of smooth spreading hairs and sessile glandular
hairs largely confined to the elevated primary and
secondary veins; petioles 1.5—3 mm long or the
blades occasionally subsessile and ± clasping.
Larger leaf at each node 4—10 X 1.5-4.5 cm.
Smaller leaf (2— )3—6 X 1 —4 cm. Inflorescence a
terminal modified dichasium 1.5—2.5 cm long be¬
coming pseudolateral with elongation of lateral
branches; bracteoles linear-lanceolate to subulate,
persistent, 2-4 X 0.5 mm, beset with spreading
simple hairs and sessile glandular-punctate hairs
abaxially or essentially glabrous and then tipped
with an apical hair. Pedicels (at anthesis) 0.5—1 mm
long. Hypanthia (at anthesis) campanulate, 1—1.5
mm long from the base to the vascular ring, sparse¬
ly covered with smooth spreading hairs and a
ground layer of scattered glandular-punctate hairs.
Calyx tube 0.25 mm long, the calyx lobes l X 1
mm, rounded-triangular, somewhat hyaline and ob¬
scured by the exterior teeth; calyx teeth 5, lance-
triangular to ± setiform, 0.25—0.5 X 0.25 mm. Pet¬
als 5, glabrous, dark red. ovate to narrowly obovate,
minutely glandular-eiliolate, erect, connivent and
concave adaxially at anthesis, 1.5—2 X 1 — 1.5 mm.
Stamens 10, isomorphic and forming an incurved
circle around the style; filaments glabrous, 1—1.5
X 0.5 mm, subulate; anthers 0.5 X 0.25 mm, pale
yellow flushed with red along the connective dor-
sally and basally at the filament insertion, oblong
to obovoid in outline, ± truncate distally with a
ventrally inclined pore; connective thickened dor-
so-basally but not dilated or prolonged below the
thecae. Ovary completely inferior, 3-locular, apex
glabrous and 10-costate. Style straight, glabrous, 2
mm long; stigma truncate, berry dark purple at ma¬
turity, 4—4.5 X 4—5 mm. Seeds narrowly pyriform,
1 mm long, ± papillate on the broader distal end.
Phenology. Collected in flower and fruit from
November through February and from June through
August.
Distribution. Rare and local in cloud forests
and ridgetop elfin vegetation in seepage areas, par¬
tially shaded vertical banks, and secondary thickets
from central Costa Rica disjunctly south to western
Panama with one known population in western Co¬
lombia from 1090 to 1800 in.
Among the three goblet-flowered species, C.
utleyana is most similar to C. garciabarrigae but
differs most notably in its leaf blades that are gla¬
brous and bullate adaxially. It also differs in its
dichasial inflorescence and its floral bracteoles that
are broadest at the base.
For an illustration of this species see Almeda
(1979: 190).
Representative specimens examined. COLOMBIA.
Choc *6: Carretera Ansermanuevo/San Jose del Palmar,
8.4 km del Alto del Galapago, 19 Feb. 1977. Forera et al.
3015 (COL). COSTA BICA. Cartago: Refugio Nat ional
de Vida Silvestre Tapantf, Sendero Oropendofa and vicin¬
ity, approximately 1.2—2.7 km from the reserve entrance,
1 Mar. 1991, Almeda et al. 6864* (AAU, BM, CAS, CR.
INB, MEXU, MO, NY, I’M A); Tapantf Hydroelectric Re¬
serve along Rio Grande de Orosf, 4.5 km beyond small
bridge which crosses river inside the preserve, along road
to diversion dam, 28 June 1976, Croat 36084 (CAS). He¬
redia: Parque Nacional Braulio Carrillo between Bfo Peje
and Rio Sardinal. Atlantic slope of Volcdn Barva,
10°15.5'N, 84°05'W, 13 Nov. 1986, Crayurn & Herrera
78 77 (CAS, INB, MO). PANAMA. Boras del Toro: Ed¬
win Fabrega Dam and Reserve in Fortuna along the Con¬
tinental Divide Trail, 12.9 km N of Sitio de Presa offices
above the Dam, 08°45'04"N, 82°15'04"W, 7 Feb. 1996,
Almeda et al. 7534 ((IAS. MO. PMA). Chiriqui: Edwin
Fabrega Dam and Reserve in Fortuna, Quebrada Bonita
ca. 7.5 km N of Sitio de Presa offices enroute to Chiriqui
Grande, 5 Feb. 1996, Almeda et al. 7503 (CAS, PMA).
Veraguas: summit of Cerro Jute with low windswept fog-
shrouded elfin vegetation, 18 Feb. 1996, Almeda et al.
7629 (CAS, PMA).
Acknowledgments. 1 thank Lawrence M. Kelly
for examining and photographing the isotype of C.
garciabarrigae at COL; the staffs of the Museo Na¬
cional de Costa Rica, Institute de Biodiversidad,
Costa Rica, Universidad de Panama, Missouri Bo¬
tanical Garden, and the Smithsonian Tropical Re¬
search Institute for logistical support in the field;
Fei Mei Chuang for assistance with the cytological
work; Ellen del Valle for preparing the species il¬
lustrations; and the curators and staffs of the fol¬
lowing herbaria for loans, gifts of specimens, and/
or assistance during study visits: BM, BR, CAS,
CR, DS, DUKE, F, INB. K, MEXU, MO, NY. P,
PMA, SCZ, US. Fieldwork for this study was sup¬
ported by National Science Foundation Grants DEB
78—25620 and BSR 8614880 (Flora Mesoameri-
cana), the G. Lindsay Field Research Fund, and
the Nathan Jay and Virginia Friedman Fund of the
California Academy of Sciences.
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308.
& T. I. Chuang. 1992. Chromosome numbers and
Volume 13, Number 2
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Almeda
Red Goblet-Flowered Clidemia
169
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Arabidopsis bactriana Belongs to Dielsiocharis (Brassicaceae)
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Sabir ,/. Junussov
Institute of Botany, Tajik Academy of Sciences, Karamova Street 27,
Dushanbe 734017, Tajikistan
ABSTRACT. The Tajikistan endemic Arabidopsis
bactriana is transferred to Dielsiocharis, a genus
previously recognized as monotypic. Detailed de¬
scription of A. bactriana is provided, and a key
distinguishing the two species of Dielsiocharis is
given.
Key words: Arabidopsis, Brassicaceae, Dielsi¬
ocharis, Taj i k i stan.
During a visit to Dushanbe (Tajikistan) in 2002,
the first author had the chance to examine many
species of Brassicaceae at TAD. One of the species,
previously placed in Arabidopsis (DC.) Heynhold,
A. bactriana Ovczinnikov & Junussov, was studied
critically, anti the present paper deals with its prop¬
er generic placement.
Arabidopsis bactriana cannot be maintained in
Arabidopsis because it is a pulvinate, scapose, ca-
nescent perennial with almost exclusively dendritic
trichomes, leafless stems, pale yellow flowers, and
non-torulose fruits. As defined by O’Kane and Al-
Shehbaz (1997), Arabidopsis consists of annual, bi¬
ennial, or non-pulvinate, non-scapose, non-canes-
cent perennials with simple and forked (never
dendritic) trichomes, leafy stems, white to lavender
or rarely purple flowers, and torulose fruits.
Al-Shehbaz et al. (1999) excluded Arabidopsis
bactriana from Arabidopsis and, prior to the exam¬
ination of the type material, suggested that the spe¬
cies might belong to Crucihimalaya Al-Shehbaz et
al. However, we now believe that the species should
be assigned to Dielsiocharis 0. E. Schulz, a genus
previously recognized by Schulz (1924. 1936) and
Hedge (1968) as monotypic.
In nearly all aspects of Arabidopsis bactriana, the
species is more closely related to Dielsiocharis kot-
schyi (Boissier) 0. E. Schulz (northern Iran and ad¬
jacent southern Turkmenistan) than to any other
species of the Brassicaceae. Both are densely caes-
pitose, pulvinate, tomentose, canescent, scapose
perennials with almost exclusively dendritic tri¬
chomes (some simple trichomes at the petiolar
Novon 13: 170-171. 2003.
base); entire, oblong, oblanceolate, or obovate, ro-
sulale basal leaves attenuating to the petiole; leaf¬
less stems; ebracteate, few- to several-flowered ra¬
cemes elongated in fruit; bright or pale yellow
petals 2.5-5 mm long; linear to oblong-linear, often
recurved, non-torulose, dehiscent, terete fruits; mi¬
nute, entire stigmas; and oblong, uniseriate seeds
with incumbent cotyledons (Hedge, 1968; Junus¬
sov, 1978).
Dielsiocharis bactriana (Ovczinnikov A Junus¬
sov) Al-Shehbaz & Junussov, comb. nov. Bas-
ionym: Arabidopsis bactriana Ovczinnikov &
Junussov, FI. Tajikistan 5: 626. 1975. TYPE:
Tajikistan. Mt. kugi-Frusch, calcareous
ground, 2800 m, 4 July 1956, D. N. Ovczin¬
nikov A M. S. Lazareva 4079 (holotype, TAD;
isotype, TAD).
Perennial herbs to 4 cm tall, pulvinate, scapose;
caudex woody, compact, covered with persistent
leaves of previous years. Leaves oblong-oblanceo-
late to obovate, 0.3—2 cm long, 1—3.5 mm wide,
canescent, densely tomentose with stalked, den¬
dritic trichomes, these with much longer stalks and
simple along petiolar base, base attenuate or cu-
neate, margin entire, apex obtuse. Cauline leaves
absent. Raceme ebracteate, 3- to 6-flowered, elon¬
gated in fruit. Sepals oblong, to 2 mm long, tomen¬
tose outside. Petals pale yellow, narrowly oblanceo¬
late, 2.5—3 mm long, apex rounded. Fruiting
pedicel slender, filiform, 5—10 mm long, sparsely
tomentose. Ovules 34 to 40 per ovary'. Fruit linear,
terete, curved, not torulose, 7-15 X 0.7-1 mm;
valves glabrous, midvein absent or obscure; style
0.1-0.5 mm long; stigma minute, entire. Seeds un¬
iseriate, oblong, brown, ca. 0.7 X 0.3 mm long;
cotyledons incumbent.
Di elsiocharis bactriana is known thus far only
from the type collection. It is easily distinguished
from D. kotschyi by having fewer-flowered racemes.
Volume 13, Number 2
2003
Al-Shehbaz & Junussov
Arabidopsis bactriana transferred
171
glabrous fruit valves, and more ovules per ovary
(see key).
Key to the Species of Dielsiocharis
la. Fruit valves glabrous; racemes 3- to 6-flowered;
ovules 34 to 40 per ovary. D. bactriana
lb. Fruit valves pubescent; racemes 10- to 20-flow-
ered; ovules 6 to 14 per ovary . D. kotschyi
Acknowledgments. We are most grateful to Ac¬
ademician Hikmat Hisoriev, Director of the Insti¬
tute of Botany of the Tajik Academy of Sciences,
for his help during the 2002 visit of Al-Shehbaz to
TAD. We thank the National Geographic Society
(Grant N712I-01) for financial support. Victoria C.
Hollowell and Steve L. O'Kane, Jr., are thanked for
their comments on the manuscript.
Literature Cited
Al-Shehbaz, I. A., S. I,. O'Kane, Jr. & R. A. Price. 1999.
Generic placement of species excluded from Arabidopsis
(Brassicaceae). Novon 9: 296—307.
Hedge, i. C. 1968. Sisymbrieae. In: K. H. Bechinger (ed¬
itor), FI. Iranica 57: 309-342. Akademische Druck-u.
Verlagsanslalt, Graz-Austria.
Junussov, S. 1978. Arabidopsis. In: P. N. Ovczinnikov (ed¬
itor), FI. Tajikistan 5: 44—51. USSR Academy of Sci¬
ences, Leningrad.
O’Kane, S. L., Jr. & I. A. Al-Shehbaz. 1997. A synopsis
of Arabidopsis (Brassicaceae). Novon 7: 323—327.
Schulz, O. F. 1924. Cruciferae-Sisymbrieae. In: A. Engler
(editor). Pflanzenreich IV. 105(Heft 86): 1—388. Verlag
von Wilhelm Engelmann, Leipzig.
-. 1936. Crueiferae. In: A. Engler & K. Prantl (ed¬
itors), Nat. Pflanzenfam., ed. 2, 17B: 227—658. Verlag
von Wilhelm Engelmann, Leipzig.
Drabopsis Is United with Draba (Brassieaceae)
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ihsan.al-shehbaz@mobot.org
Marcus Koch
Institute of Botany, University for Agricultural Sciences Vienna, Gregor-Mendel-Str. 33,
A-1180 Vienna, Austria. koch@edv 1 .boku.ac.at
ABSTRACT. Drabopsis nuda is transferred to Dra¬
ba, and (he two genera are united. The distinguish¬
ing characters separating these two genera and Ar-
abidopsis are discussed.
Key words: Arabidopsis, Brassieaceae, Crucifer-
ae, Draba. Drabopsis.
The systematic position of Drabopsis K. Koch has
been the source of continuous controversy. The ge¬
nus has long been considered to be monotypic.
Naqshi and Javied (1984) added another species,
D. brevisiliqua Naqshi & Javied, that was said to
differ from the highly variable I), nuda (Belanger)
Stapf by having shorter fruits (ca. 1.2 cm) and non-
Hexuous scapes. In our opinion, these differences
are trivial and D. brevisiliqua is reduced to synon¬
ymy of D. nuda (see below).
Drabopsis nuda is distributed from western Xin¬
jiang (China), central Asia, Kashmir, and Afghan¬
istan westward into Iran, Iraq, Turkey, Caucasus,
and Crete (Gustavsson, 1977). The nomenclature of
the species has been dealt with by Leonard (1977),
and the species should be recognized as D. nuda.
instead of the later-published D. verna K. Koch, as
was done by Cullen (1965), Hedge (1968), Jafri
(1973), and Gustavsson (1977).
Drabopsis nuda does not fit appropriately in any
genus. It was originally described as Arabis b., and
it resembles Arabis auriculata Lamarck in having
linear, somewhat latiseptate fruits, stout fruiting
pedicels nearly as thick as fruit, and accumbent
cotyledons. However, the species cannot be accom¬
modated m Arabis because it has yellow flowers and
leafless scapes. The presence in D. nuda of accum¬
bent instead of incumbent cotyledons and leafless
instead of leafy stems strongly supports its exclu¬
sion from Sisymbrium L., where it was placed by
Boissier (1867).
Busch (1909) and Bornmuller (1911) transferred
Drabopsis verna anti I), nuda It) Arabidopsis (DC.)
Heynhold, respectively. No reasons were given to
Novon 13: 172-173. 2003.
support such transfer, and Jafri (1973) recommend¬
ed that the two genera be united, though he main¬
tained both. 1 he lack in Drabopsis nuda of any cau-
line leaves and the presence of stout fruiting
pedicels nearly as thick as the fruit, yellow flowers,
and stalked, almost exclusively stellate trichomes
clearly do not support its placement in Arabidopsis.
All species of Arabidopsis have cauline leaves,
fruiting pedicels narrower than fruits, white to lav¬
ender flowers, and a mixture of simple and forked
trichomes. Furthermore, molecular data (O'Kane &
Al-Shehbaz, in prep.) demonstrate that Arabidopsis
and Drabopsis are unrelated. Al-Shehbaz et al.
(1999) excluded the species from Arabidopsis but
maintained it in Drabopsis.
I’he recognition of Drabopsis as a monotypic ge¬
nus does not solve its affinity and relationship. In
every aspect of its scapose habit, leafless scapes,
annual duration, flower color, accumbent cotyle¬
dons, and trichome morphology, Drabopsis nuda is
more at home in Draba. In fact, Boissier (1867) was
the first to point out the resemblance of the species
to Draba L. (as Erophila DC.). The only morpho¬
logical oddity brought about by the placement of
Drabopsis nuda in Draba is the relative length/
width ratio of the fruit. No other species in Draba
has fruits as many as 45 times longer than broad,
though this extreme length/width ratio of the fruits
in Drabopsis nuda is the maximum, and according
to Zhou et al. (2001), the species has fruits (10—)
17—33(—45) X 0.8—1 mm. Other species of Draba
with linear fruits as long as 30 times the width are
found in the Himalayan and central Asian I), au-
cheri Boissier and the Tibetan D. nylamensis Al-
Shehbaz (Al-Shehbaz, 2002).
Schulz (1936) placed Drabopsis in the tribe Ar-
abideae and Draba in the tribe Drabeae. His tribal
assignment of the two genera is not supported by
morphological or molecular data, and it has been
shown that the closest relatives of Draba s.l. are
Volume 13, Number 2
2003
Al-Shehbaz & Koch
Drabopsis and Draba United
173
Eurasian Arabis and Aubrieta Adanson (Koch & Al-
Shehbaz, 2002).
Molecular studies (Koch & Al-Shehbaz, unpub¬
lished) clearly show that Drabopsis nuda is close to
the annual species of Draba (e.g., D. nemorosa L.,
D. muralis L., D. verna L). Phylogenetic analysis
using DNA sequence variation of the internal tran¬
scribed spacers ITS-1 and ITS-2 of nuclear ribo-
somal DNA demonstrated that Draba (including Er-
ophila) forms a monophyletic assemblage. The vast
majority of perennial species of Draba are com¬
bined to one large group, and all annual taxa, in¬
cluding Drabopsis nuda, are positioned basal to
them.
Draba nuda (Belanger) Al-Shehbaz & M. Koch,
comb. nov. Basionym: Arabis nuda Belanger,
Voy. Indes Or.. Bot. t. 15a. 1834. TYPE: illus¬
tration t. 15a.
Drabopsis brevisiliqua Naqshi & Javied, J. Econ.
lax. Bot. 5: 966. 1984. Syn. nov. TYPE: Kash¬
mir. Harwan, A. R. Naqshi 3784 (KASH).
Draba nuda is easily distinguished from all of
the other annual, yellow-flowered species of Draba
by having leafless scapes, stout fruiting pedicels
nearly as thick as the fruit, slightly flexuous raehis,
and slightlv compressed, slender, linear fruits (1-)
1.7-3.3(-4.5) cm X 0.8—1.2(—1.6) mm.
Acknowledgments. We are grateful to Victoria
C. Hollowed and Steve L. O'Kane, Jr., for their
comments on the manuscript.
Literature Cited
Al-Shehbaz, I. A. 2002. Six new species of Draba (Bras-
sicaceae) from the Himalayas. Novon 12: 314—318.
-. S. L. O’Kane, Jr. & R. A. Price. 1999. Generic
placement of species excluded from Arabidopsis (Bras-
sicaeeae). Novon 9: 296—307.
Boissier, E. 1867. Flora orientalis sive enumeratio plan-
tarum in Oriente a Graecia et Aegypto ad Indiae fines
hucusque observatarum. Vol. 1. H. Georg, Basel & Ge¬
neve.
Bornmiiller, J. 1911. Colleetiones Staussianae novae.
Beih. Bot. Centralbl. 28(2): 458—535.
Busch, N. 1909. Arabidopsis. In: N. Kusnezow, N. Busch
& A. Fomin (editors). Flora Caucasica Critica 3(4):
456—467.
Cullen, J. 1965. Drabopsis. In: P. H. Davis (editor). Flora
of Turkey 1: 444. University Press, Edinburgh.
Gustavsson, L.-A. 1977. Drabopsis verna C. Koch (Bras-
sicaceae) new to Europe. Bot. Not. 130: 213—214.
Hedge. I. C. 1968. Arabideae. In: K. H. Rechinger (edi¬
tor), Flora Iranica 57: 193—218. Akademische Druck-
u. Verlagsanstalt, Graz-Austria.
Jafri, S. M. H. 1973. Rrassicaceae. In: E. Nasir & S. I.
AH (editors), Flora of Pakistan, Vol. 55. Ferozsons, Ka¬
rachi.
Koch, M & I. A. Al-Shehbaz. 2002. Molecular data in¬
dicate complex intra- and intercontinental differentia¬
tion of American Draba (Brassicaceae). Ann. Missouri
Bot. Card. 89: 88-109.
Leonard, J. 1977. Le genre irano-touranien Drabopsis et
son unique espfece I), nuda (Cruciferae). Publ. Cairo
Univ. Herb. 7-8: 295-297.
Naqshi, A. R. & G. N. Javied. 1984. Drabopsis brevisiliqua
(Cruciferae)—A new species from Kashmir. J. Econ.
Taxon. Bot. 5: 963-966.
Schulz, <). F. 1936. Cruciferae. In: A. Fngler & K. Prantl
(editors), Nat. Pflanzenfam., ed. 2, 17B: 227—658. Ver-
lag von Wilhelm Engelmann, Leipzig.
Zhou, T. Y.. L. L. Lu, G. Yang & I. A. Al-Shehbaz. 2001.
Brassicaceae (Cruciferae). Pp. 1-193 in Z. Y. Wu & P.
H. Raven (editors). Flora of China, Vol. 8. Science
Press, Beijing, and Missouri Botanical Garden Press,
St. Louis.
Three New Species of Biophytum (Oxalidaceae) from the
Venezuelan Guayana
Gerardo Aymard C.
UNELLEZ-Guanare, Programa ruary to May.
Conservation status—IUCN Red List Catego¬
ry. Least Concern (LG). This is a relatively
widespread species, although low collecting fre¬
quency infers that further work is required to assess
local vulnerability.
Additional material examined. PAPUA NEW GUIN¬
EA. Central Province: Tovobada Hills, ca. 9 mi. [15
km] NNW of Port Moresby, ca. 500 ft. [152 m], 14 May
1967, Pullen 6980 (A, BO, Bill, CANB, K. L, LAE. US);
near SE side of Little Mount Lawes, ca. 16 mi. |26 km]
N of Port Moresby, ca. 150 ft. [46 m], 25 Apr. 1967, Pul¬
len 6814 (A. BRI, CANB, K, L, LAE); Kanosia, ca. 50 ft.
115 m], 9 Eeh. 1935, Carr 11220 (K); Port Moresby sub¬
district, below Vetorogu Hill, Tovobada Peaks, 3 km N of
Barune, 60 m, 24 May 1974, Frodin & Field Botany Class
UPNG 4323 (K). AUSTRALIA. Torres Strait Is.: Sun¬
day Island, June 1821, Cunningham 7 (K); Sunday Island,
s.c. s.n. (K). Queensland: Cook District, 1 km S of Siei-
sia. 21 Nov. 1999, Wannan NSW 443387 (BRI, NSW ).
Notes. Psilanthus brassii is similar to P. mabe-
sae (Elmer) J.-F. Leroy from the Philippines, but
differs in that its leaves, flowers (corolla), and fruits
are glabrous (not puberulous to pubescent as in P.
mabesae). Herbarium and literature studies indi¬
cate that P. brassii is the only species of Psilanthus
occurring in Papua New Guinea and Australia.
Acknowledgments. The author thanks K. Chal-
lis. B. k. Brummitt, D. Bridson, and G. Zijlstra for
careful guidance with nomenclatural aspects of this
paper. The directors and staff at the following her¬
baria are acknowledged for their assistance with
access to herbarium collections: BO, CANB, L, and
NSW 7 .
Literature Cited
Bridson, D. M. 1987. Nomenclatural notes on Psilanthus,
including Coffea sect. Paracoffea (Rubiaceae tribe Cof-
feeae). Kew Bull. 42: 453-460.
-. 1988a. Coffea. Pp. 415-474 in R. M. Polhill (ed¬
itor), D. M. Bridson & B. Verdcourt, Flora of Tropical
East Africa, Rubiaceae, Pt. 2. Balkema, Rotterdam/
Brookfield.
-. 1988b. Classification. Pp. 61—75 in G. Wrigley,
Coffee. Longmans, Tropical Agriculture Series, New
York.
Chevalier, A. 1947. Les cafeiers du globe, fasc. 3: Sys-
tematique des cafeiers et faux-cafeiers maladies et in-
sectes nuisibles. Encycl. Biol. 28: 1—352.
Davis, A. P. & F. Rakotonasolo. 2000. Three new species
of Coffea L. (Rubiaceae) from Madagascar. Kew" Bull.
55: 405^06.
Greuter, W., J. McNeill, E. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Eilgueiras, 1). H. Nicolson, P C. Silva.
J. E. Skog, P. Trehane, N. J. Turland & D. I.. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Leroy, J.-F. 1967. Diagnose differentielle du genre Para¬
coffea Leroy. J. Agric. Trop. Bot. Appl. 14: 276.
-. 1968a. Un Cafeier du genre Paracoffea en Nou-
velle-Guinee. J. Agric. Trop. Bot. Appl. 14: 598-600.
|1967, publ. 1968.]
-. 1968b. La loi de correlation de croissance chez
les Cafeiers: Cas d’une espece d’Indonesie (Paracoffea
floresiana). J. Agric. Trop. Bot. Appl. 14: 601—608.
11967, publ. 1968.]
-. 1972. Note sur un cafeier du Sud-East Asiatique:
Paracoffea merguensis (Ridl.) Leroy. Adansonia, s6r. 2,
11: 603-614. [1971, publ. 1972.]
-. 1980a. Les grandes lignees de Cafeiers. Ass. Sci.
Internat. Cafe (ASIC). 9th Colloque: 473-477.
-. 1980b. Evolution et taxogenese chez les Cafeiers
(Coffea L., Psilanthus Hook.f. et Nostolachma Durand).
Hypothese sur leur origine. Compt. Rend. Acad. Sci.
Paris, ser. D, 29: 593—596.
-. 1981. Les cafeiers du genre Psilanthus (Rubi-
acees) en Afrique orientale et en Asie et Ties du Paci-
fique. Bull. Mus. Natl. Hist. Nat.. B, Adansonia 3: 251 —
258.
Lobreau-Callen, D. & J.-F’. Leroy. 1980. Quelques don-
rfees palynologiques sur le genre Coffea et autres genres
du eercle des Cafeiers. ASIC, 9th Colloque: 479-506.
Moens, P. 1962. La place systematique du Coffea lebru-
niana Germain et Kesler. Bull. Jard. Bot. Brux. 32: 131.
Robl irecht, E. 1981. Studies in tropical African Rubi¬
aceae (II): 5. A survey of Argocoffeopsis. (6) A revision
of Calycosiphonia. Bull. Jard. Bot. Nat. Belg. 51(3/4):
359-378.
Sivarajan, V. V., S. I). Biju & P. Mathew. 1992. Revision
of the genus Psilanthus Hook.f. (Rubiaceae tribe Cof-
feeae) in India. Bot. Bull. Acad. Sin. (Taipei) 33: 209—
224.
Stoffelen, P. 1998. Coffea and Psilanthus in Tropical Af¬
rica: A Systematic and Palynological Study, Including
a Revision of the West and Central African Species.
Ph.D. Thesis, Katholieke Universiteit Leuven.
-, E. Robbrecht & E. Smets. 1997. Pollen mor¬
phology of Coffea and Psilanthus (Rubiaceae), mainly
from Africa. Grana 36: 313—327.
Wong, K. M. 1989. Rubiaceae. Pp. 324-425 in E. S. P.
Ng, Tree Flora of Malaya 4. Longman, Selangor.
184
No von
Appendix 1. Outline classification of Coffea and Psilan-
thus.
Coffea L„ Sp. PL: 172. 1753.
Cuff. •a subg. Coffea I . TYPE: Coffea arabica L.
Around 90 species. Africa, Madagascar, Mascarenes.
Coffea subg. Psilanthopsis (A. Chevalier) J.-F. Leroy, Ass.
Sci. Internat. Calc, (ASIC) 9th Colloque: 475. 1980.
Psilanthopsis A. Chevalier, J. Agric. Trop. Bot. Appl.
19: 404. 1939. TYPE: Coffea kapakata (A. Cheva¬
lier) Bridson.
Monotypic. Africa (Angola).
Paolia Chiovenda, Result. Sci. Miss. Stenfan. -Pauli So-
mal. Ital. 1: 93. 1916. TYPE: Paolia jasminoides
Chiovenda (= Coffea rhamnifolia (Chiovenda) Brid¬
son).
Monotypic. Africa (Somalia and Kenya).
Coffea subg. Baracoffea (J.-F. Leroy) J.-F. Leroy, Ass.
Sci. Internat. Cafe, (ASIC) 9th Colloque: 475. 1980.
Coffea sect. Baracoffea J.-F Leroy, Comp. Rend.
Acad. Sci. Paris 252: 2287. 196L TYPE: Coffea
humbertii.
Seven species (including four undescribed). Madagas¬
car.
|Paracoffea subg. Insulanoparacoffea J.-F. Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), noin. nud.]
Psilanilius Hooker L, Gen. PL: 115. 1873.
Psilanlliiis subg. Psilanilius Hooker f. TYPE: Psilanthns
mannii Hooker f.
Two species. Africa (central and western).
Psilanthns subg. Afrocoffea (Moens) Bridson, Kew Bull.
42: 454. 1987. Coffea subg. Afrocoffea Moens, Bull.
Jard. Bot. Brux. 32: 131. 1962. TYPE: Psilanthus
lebrunianus (Germain & Kesler) Bridson.
About 17 species. Africa, Asia, Australasia.
Coffea sect. Paracoffea Miquel, FI. Ind. Batavae: 308.
1856. Psilanthus subg. Paracoffea (Miquel) J.-F.
Leroy, Ass. Sci. Internat. Cafe, (ASIC) 9th Colloque:
475. 1980. TY PE: Coffea horsfieldiana Miquel.
Paracoffea J.-F. Leroy, J. Agric. Trop. Bot. Appl. 14: 276.
1967. TYPE: Coffea horsfieldiana Miquel.
|Paracoffea subg. Afroparacoffea J.-F. Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
|Paracoffea subg. Melanoparacoffea J.-F. Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
Two New Species of Streptocarpus (Gesneriaceae) from
South Africa
T. ,/. Edwards
School of Botany and Zoology, University of Natal. Private Bag X01, Scottsville 3209,
South Africa
Abstract. A new species of the Streptocarpus
rexii aggregate is described from I tala Game Re¬
serve in KwaZulu-Natal, South Africa. The geo¬
graphic isolation of S. kunhardtii, its pale methyl
violet (lowers, and long fruits distinguish it from S.
roseoalbus, its closest relative. A second new spe¬
cies, S. hilburtianus, is described from two high-
altitude localities in Mapumalanga, South Africa.
The alliances of this species are unclear, although
corolla coloration and leaf vestiture approach S.
galpinii ; however, it differs in its cylindrical corolla
tube, its much shorter pedicels, and its distinctive
pollen.
Key words: Gesneriaceae, South Africa, Strep¬
tocarpus.
The recognition of species limits within Strepto¬
carpus Lindlev is complicated by the lack ol fertil¬
ity barriers between species ol the typical subge-
nus. This feature has considerable impact on
species concepts for the genus and heightens the
importance of allopatry in the speciation process.
For the most part, Streptocarpus species are limited
to forest patches that have shrunk and expanded,
with climatic vicissitudes, through geological his¬
tory. Over the last 2 million years, southern Africa
has experienced 20 climatic cycles, each lasting ca.
100,000 years (Deacon, 1983; Tyson. 1986; Deacon
& Lancaster, 1988). The resulting fragmentation ol
populations produces a powerful driving force for
allopatric speciation, which in my opinion has been
fundamental in the divergence ol Streptocarpus (see
Weigend & Edwards, 1994a, 1994b).
1. Streptocarpus kunhardtii I. J. Edwards, sp.
nov. TYPE: South Africa. KwaZulu-Natal: Itala
Nature Reserve. Mpophomeni Gorge, S-facing
cliffs, 14.12.1996, C. Carbutt 35 (holotype,
NU: isotvpes, E, MO, PRE).
Haec species ad gregem Streptocarpi rexii pertinens
quoad corollae formam notasque ad S. roseoalbum maxime
accedit, sed ah eo foliis majoribus plerumque suberectis.
fructu majore atque lobulis corollinis inferioribus pallide
malvicoloribus distinguitur.
Perennial rosulate, rhizome poorly developed.
Leaves suberect, up to 120-250 X 40-55 mm, mar¬
gin eremite, base cuneate, decurrent on the petiole;
petiolode 10-15 X 6-8 mm, red-brown to green,
pilose. Inflorescence 1- or 2(to 4)-flowered; pilose
with glandular and filiform trichomes throughout;
peduncle up to 100—240 mm, maroon to green;
bracts lanceolate, 2-3 X 0.75 mm; pedicel 9-15
mm. Calyx segments linear, 5.5—6.5 mm, glandular
pilose. Corolla broadly infundibuliform, tube white,
24—29 X 3 mm broadening to 10 mm in the throat,
bent in the lower third; pilose with glandular and
filiform trichomes on the outside; yellow stripe on
the tube floor sometimes broadened in the basal
third: lower limb 15—20 mm long, lobes 10—15 X
10-12 mm. pale methyl violet (Wilson. 1941) with
darker central stripes; upper lobes 7—13 X 10—1 I
mm. Stamens 2; filaments 8—9 mm, white, with
stalked glandular trichomes; anthers connate; stam-
i nodes 2—3 X 0.5 mm, glabrous. Pollen prolate,
17-20 X 9—10 /xm, tricolpate; mesocolpium dif¬
fusely perforate, scabrate (Fig. 1). Ovary 10—14 mm
long, densely pubescent, green; style 8—12 mm
long, glandular, white. Fruit 80—110 X 2.5 mm.
Seeds 0.5-0.7 mm long, sculpturing reticulate.
Distribution. Chris Kunhardt collected this ro¬
sulate Streptocarpus from Itala Game Reseve in the
1980s, shortly before his death. Vague provenance
details of the population were kept in his living
collection, and relocation of the population proved
difficult. Recently the species was rediscovered at
Itala Nature Reserve ( Carbutt 35) in sparse popu¬
lations. deeply shaded on south-facing cliffs, usu¬
ally near streams of the Mpophomeni Gorge.
Phenology. The species flowers through De¬
cember and January, fruiting between December
and February 7 .
Etymology. The species epithet honors its dis¬
coverer. the late Chris Kunhardt, an avid amateur
botanist and Streptocarpus enthusiast.
Corolla morphology and color patterning suggest
an alliance with S. roseoalbus Weigend & T. J. Ed¬
wards, which occurs in Lowveld vegetation
(Acocks, 1988) around Barberton (Mpumalanga). In
Novon 13: 185-188. 2003.
186
Novon
Figure 1. Pollen micrographs. —A. Streptocarpus kun-
hardtii (Carbult 35). Scale bar 6 gm. —B. Streptocarpus
hilburtianus (Edwards 682). Scale bar 5 gun.
S. roseoalbus the corolla is 34—45 mm long witli a
short tube of 25—30 mm; its proximal width is 3
mm broadening to 9-10 mm in the throat. Corolla
color is, however, distinctive: S. roseoalbus is al¬
ways pink and S. kunhardtii is always methyl violet.
I he leaves of S. roseoalbus are small and more or
less prostrate, while in S. kunhardtii leaves are usu¬
ally suberect. In addition, the mature fruits of S.
roseoalbus are significantly shorter than those of the
new species, never exceeding 55 mm.
Streptocarpus kunhardtii further plugs the dis¬
junctions between the closely allied rosulate Strep¬
tocarpus species ol the eastern seaboard of South
Alrica: its morphology closely approximates these
species, and its pollen is indistinguishable (Wei-
gend & Edwards, 1996).
Paratypes. SOUTH AFRICA. KwaZulu-Natal: 2731
(Louwsburg) 11 a I a Game Reserve (CB), 14 Dec. 1996. R.
Scott Shaw 8597 (MO).
2. Streptocarpus liilhurtiaiius T. J. Edwards, sp.
nov. PE: South Africa. Mpumalanga: Long
Tom Pass, Die Geut, 09.12.1992, T. J. Ed¬
wards 682 (holotype, NU). Figure 2.
Haec species Streptocarpo galpinii similis, sell ah eo
corollae tubo elongato cylindricoque labio inferiore ad
basim macula alba luteave carente atque pedieellis bre-
vioribus distinguitur.
Plards perennial, leaves I to 3, prostrate, lamina
50-100 X 30—70 mm, base broadly cuneate,
densely pilose, sericeous. Inflorescence cymose 2-
to 10-flowered, lower flowers often sub-opposite;
peduncle 50—140 mm long, densely glandular pi¬
lose; pedicels 10—14 mm long, densely glandular
pilose; bracts spathulate, 2.5-3 X 0.5 mm. Calyx
segments deltoid, 2.5-3 X 1.5 mm, slightly spread¬
ing. densely glandular pilose. Corolla 28—30 mm
long, sparsely glandular pilose outside, glabreseent
inside but sparingly pubescent on the floor; tube
cylindric 18—23 mm, basal width 3—4 mm, mouth
width 4.5—5 mm, deep mauve outside, white inside;
lower lip 10 mm long, three-lobed, lobes 6X9
mm, pale mauve diluting to white in the throat, 2
mauve sinus chevrons sometimes present; upper lip
i mm long, 2-lobed. Stamens 2, filaments arising
in the lower third of the corolla tube, filaments 4—
5 mm long, connective sparsely glandular pilose,
anthers connate, 2 mm long; staminodes 2. 2-3 mm
long, glabrous. Pollen spheroidal. 10—12 X 10—12
/am, trieolpate, colpi weakly verrucose, mesocolpi
scabrate, apparently imperforate, poles apocolpate.
Gynoecium 10-12 mm long; ovary 5-6 mm long,
densely sericeous; style 5—6 mm long, pilose; stig¬
ma stomatomorphic. Capsule 25—30 X 2.5-3 mm.
pilose.
Distribution. This species has only been col¬
lected twice, in Mpumalanga, South Afric a. Popu¬
lations occur at high altitudes (ca. 2000 m) on lith-
osols above cliffs in afromontane grasslands. The
plants grow r in very exposed positions, sparingly
shaded by rock ledges and associated scrub.
Phenology. Flowering and fruiting collections
of this species were made in November and De¬
cember.
Etymology. The epithet is a combination of two
names in recognition of the combined contributions
made by 0. M. Hilliard and B. E. Burtt to the sys-
tematics of Streptocarpus.
The niche and vegetative form of .S', hilburtianus
is similar to S. galpinii J. I). Hooker, but they are
distinguished by pedicel length and corolla mor-
Volume 13, Number 2
2003
Edwards
Streptocarpus from South Africa
187
Figure 2. Streptocarpus hilburtianus (Edwards 682). —A. Flowering plant. Scale Far 20 mm. —B. Floral dissection.
Scale bar 5 mm. —C. Gynoecium. Scale bar 2 mm. Drawn by A. J. Beaumont.
phology. In S. hilburtianus tin* pedicels are consis¬
tently shorter. 10—14 mm long, compared to 25—90
mm long in S. galpinii. The corolla of S. galpinii
is distinctive for the genus, being short and open
with a violet purple limb and a white patch, often
colored with yellow, at the base of the lower lip. In
contrast the new species has a cylindrical corolla
tube usually with two chevrons of deep mauve
flanking the medial lobe of the lower lip. The pollen
of S. hilburtianus is unique in the genus (fig. 1).
The tectal ornamentation is very similar to that of
the 5. rexii aggregate but the grains are spheroidal
and not prolate.
Paratype. SOUTH AFRICA. Mpumalanga: 2530
(Lydenburg) Hartebeesvlakte../. Kluge 2026 (PRE).
Acknowledgments. Jaco Truter is thanked for
discussions on habitat and for tracking down the
second population of S. hilburtianus at Hartebeesv¬
lakte. Angela Beaumont contributed the line draw-
188
Novon
ing of S. hillmrtianus. Clinton Carbutt and Hob
Scott-Shaw are thanked for relocating the popula¬
tions of 5. kunhardtii. Mike Lambert is acknowl¬
edged for his continued assistance with Latin di¬
agnoses, and I valued the improvements suggested
by E. Hennessy. The research was funded by the
National Research Foundation of South Africa and
the University of Natal Research Fund.
Literature Cited
Acocks, J. I’. H. 1988. Veld types of South Africa, ed. 3.
Mem. Hot. Surv. S. Africa. No. 57.
Deacon, H. J. 1983. Another look at Pleistocene climates
of South Africa. S. African J. Sci. 79: 325-328.
- & N. Lancaster. 1988. Late (Quaternary Palaeoen-
vironments of Southern Africa. Clarendon Press, Ox¬
ford.
Tyson, I’. I). 1986. Climatic Change and Variability in
Southern Africa. Oxford Univ. Press, Cape Town.
Weigend, M. & T. J. Edwards. 1994a. Notes on Strepto-
carpus primulifolius (Gesneriaceae). S. African J. Bot.
60: 168-169.
-&-. 1994b. Notes on Streptocarpus cyaneus
and -S', parviflorus. Sendtnera 2: 365-376.
&-• 1996. flie palynology of Streptocarpus
and the other African and Malagasy Gesneriaceae and
its systematical implications. Bot. Jahrb. Syst. I 18: 59-
80.
Wilson, IL F. 1941. Horticultural Colour Chart 39. British
Colour Council in collaboration with the Royal Horti¬
cultural Society. Henry Stone & Sons, Banbury.
Two New Species of Rhododendron (Ericaceae) from China
Gao Lian-ming and Li De-zhu
Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, Yunnan,
China, gaolm@mail.kib.ac.cn; dzl@mail.kib.ac.cn
ABSTRACT. Two new species. Rhododendron day-
aoshanense L. M. Gao & I). Z. Li from Guangxi.
southern China, and Rhododendron longdobum L.
M. Gao & D. Z. Li from Yunnan, southwestern Chi¬
na, are described and illustrated here. Rhododen¬
dron dayaoshanense, known only from Dayaoshan
Mountain, Guangxi. is distinguished by the follow¬
ing characters: the leaves narrowly oblanceolate or
oblong-lanceolate, short setose on undersurface and
midrib, petiole densely glandular setose, pedicel
densely gray-puberulent and glandular-pilose as
well as ovary densely flavescent-tomentose. Rho¬
dodendron longilobum is collected only from Daw-
eishan Mountain, southeast Yunnan, and is distin¬
guished from its close relatives by the characters
of the leaf blades with prominent lateral primary
veins on both surfaces, calyx lobes membrana¬
ceous, linear, linear-lingulate or narrowly triangu¬
lar, and 0.5—2 cm long; ovary densely flavescent
tomentose and truncate at the apex.
Key words: China, Ericaceae, Rhododendron.
The two new species in this paper belong to Rho¬
dodendron subg. Azaleastrum, seel. Choniastrum,
which is recognized by the following characters: ev¬
ergreen shrubs or small trees; inflorescence buds
lateral at the end of branches, 1- to several-flow¬
ered; corolla narrowly funnel-shaped, 5-lobed; sta¬
mens 10; ovary glabrous or pilose, cylindric, (5)6-
locular. Section Choniastrum is comprised of about
14 species in China (He et ah. 1994). Species of
section Choniastrum are mainly distributed in
southern China; only R. taiense Hutchinson is en¬
demic to Thailand, and R. moulmainense Hooker f.
extends southward to the Malay Peninsula and
westward to Assam in India (Philipson & Philipson,
1986; Fang & Ming, 1995).
Kliododemlron dayaoshanense L. M. Gao & 1).
Z. Li, sp. nov. TYPE: China. East Guangxi:
Jinxiu Xian. Dayaoshan, alt. I 180 m. 2 May
2000 (ft). L. M. Gao 20059 (holotype, KUN;
isotype, KUN). Figure 1.
Species valde affinis R. henryi Hance, sed laminis fo-
liorum oblanceofatis vel oblongo-lanceolatis, ft—IT cm lon-
gis, 1.8—2.5 cm latis, pedicellis dense cinereo-pubescen-
tibus et sparsim glanduloso-pilosis; ovario dense
flavescenti- tomentoso bene distincta.
Small tree, 3—4 m high; young branches slender,
terete, glandular setose; mature branches brown,
glabrous. Leaves clustering at the ends of branches,
pseudo-verticillate; blade narrowly oblanceolate or
oblong-lanceolate, 8—13 X 1.8—2.5 cm; apex acu¬
minate, base cuneate, margin slightly revolute,
midrib slightly depressed above and raised be¬
neath. lateral veins 14 to 18 pairs, prominent on
both surfaces, upper surface green, glabrous, un¬
dersurface pale green, short setae scattered on un¬
dersurface, dense on midrib. Petiole 0.8—1.5 cm
long, grooved above, glandular setose. Inflores¬
cence lateral, 4- to 5-flowered, umbellate; rachis
brown, glabrous, 0.5 cm long; pedicels 2.5—3.5 cm
long, densely gray pubescent and sparsely glan¬
dular pilose. Calyx rim undulate, lobes 5, triangu¬
lar, ea. I mm long, glabrous. Corolla pale rose, nar¬
rowly funnelform, 5-5.5 cm long, glabrous outside;
lobes 5, oblong-obovate or elliptic, 3.5—4 X 2.5—3
cm, tube 1.5—2 cm long, upper lobe with yellow
blotch inside. Stamens 10, shortly exserted. fila¬
ments 4—4.5 cm long, puberulous in basal half but
glabrate at base; anthers ovate, ca. 2 mm long, pur¬
ple. Ovary oblong-cylindric, 7 mm long, densely
flavescent tomentose. Style glabrous, 4—4.5 cm
long; stigma green, capitate. Capsule not seen.
Distribution and habitat. The new species is
known only from the type collection. It was found
about 8 km east of Dayaoshan Mountain, Jinxiu
county, east Guangxi Province, growing on a slope
near the roadside, on the margin of mixed forest at
ea. I 180 m altitude. Little plant collection has been
done in the area. Thus it is not surprising that the
species had not been collected before, and it is
likely that it is not really rare on Dayaoshan Moun¬
tain.
Rhododendron dayaoshanense resembles R. hen¬
ryi Hance in sharing 4 to 5 flowers per inflores¬
cence, pink corolla, cylindric ovary, and tapering
ovary apex. However, R. dayaoshanense differs from
R. henryi in the leaves oblanceolate or oblong-lan¬
ceolate, calyces minute, glabrous, pedicel densely
gray-puberulent and glandular-pilose, as well as
Novon 13: 189-192. 2003.
190
Novon
Figure 1. Rhododendron dayaoshunen.se I . M. (;ao & I). /. I.i. —A. Flowering branch. —B. Pistil. —C. Basal portion
of abaxial leaf surlace. —I). Stamen. Drawn from the holotype ( L. M. Gao 20059).
Volume 13, Number 2
2003
Gao & Li
New Rhododendron from China
191
Figure 2. Rhododendron longilobum L M. Gao & I). /. Li. —A. Young fruit branch. —B. Pistil. —C. Pistil with
calyx lobes. —I). Stamen. Drawn from the holotype {Rhododendron Picture Group 83—0357).
1cm
192
Novon
ovary densely flavescent-tomentose. Rhododendron
dayaoshanense is also similar to R. cavaleriei in
having 4 to 5 flowers per inflorescence, pedicel
densely gray-pubescent, ovary densely flavescent-
tomentose, and ovary apex tapering. However, the
later species differs in the young branches gla¬
brous, the leaves glabrous, the lateral primary veins
inconspicuous, and petiole glabrous.
The new species is possibly a hybrid between R.
cavaleriei Leveille and R. championae Hooker f. be¬
cause R. dayaoshanense is distributed together with
R. cavaleriei and R. championae in this region and
some characters of R. dayaoshanense were shared
by R. cavaleriei and R. championae. The sequences
of ITS and TrnL-F of R. dayaoshanense are tin-
same as those of R. cavaleriei and R. championae.
Rhododendron longilobum L. M. Gao & I). Z.
Li. sp. nov. TYPE: China. Southeast Yunnan:
Pingbian Xian, Daweishan, near Shiuwei-
cheng, alt. 1950 m, 9 May 1983, Rhododen¬
dron Picture Group 83-0357 (holotype, KUN;
isotype, KUN). Figure 2.
Species valde affinis R. cavaleriei sed laminis foliorum
oblongis vel oblongo-ellipticis, nervis lateralibus foliorum
utrinque conspicuis; lobis calycis membranaceis, 5—20
mm longis, 1 mm latis, linearibus vel lineari-lingulatis vel
anguste triangularibus, ovario dense flavescenti-tomento-
so. apice truncate differt.
Small trees, 6—8 m high: young branches terete,
gray, glabrous. Leaves clustering at the ends of
branches, pseudo-verticillate; blades elliptic or
narrowly elliptic, ca. 7-10.5 X 1.8-2.5 cm. apex
long acuminate, base cuneate, margin slightly rev-
olute; upper surface green, under surface pale
green, glabrous, midrib depressed above and raised
beneath, lateral veins 12 to 16 pairs, prominent on
both surfaces. Petioles 0.8-1.5 cm long, grooved
above, glabrous. Inflorescences lateral, 4- to 5-flow-
ered. umbrella clustering near the end of leafy
shoot; outer bud scale gray pubescent, margin and
apex minutely ciliate; rachis brown, ca. 0.5 cm
long, glabrous; pedicels 2-2.5 cm long, glabrous.
Calyx lobes 5, membranaceous, linear, linear-lin-
gulate, or narrowly triangular, 5—20 X 1 mm, mar¬
gins sometimes minutely ciliate and apical rim hav¬
ing tufts of long weak hairs. Corolla pale rose,
narrowly funnelform, glabrous outside; lobes 5, 3—
3.5 cm long, tube ca. 1 cm long. Stamens 10. short¬
ly exserted, filaments 3—3.5 cm long, with puber¬
al. -nee on lower half; anthers ovate, ca. 2 mm long.
Ovary narrowly cylindric, 7-12 mm long, densely
flavescent tomentose, apex truncate; style glabrous,
3—3.5 cm long. Capsule not seen.
Distribution and habitat. The new species is
probably rare as it is known only from the type
collection, which occurs in evergreen broad-leaved
forest, near the top of Daweishan Mountain at ca.
1950 m altitude, Pingbian County, southeast Yun¬
nan province. This species occurs together with R.
tutcherae Hemsley & E. H. Wilson, R. moulmai-
nense Hooker f.. and R. hancockii Hemsley of sec¬
tion Choniastrum. The habitat of the new species
is very humid, which does not benefit growth and
dispersion of seeds. It may be one of the reasons
why this new' species is rare at the type locality.
I he species is easily distinguished from other
species of section Choniastrum by the leaf blades
with prominent lateral primary veins on both sur¬
faces, the calyx lobes membranaceous, up to 2 cm
long, and the ovary densely flavescent tomentose
and truncate at the 1 apex. Rhododendron longilobum
is most similar to R. cavaleriei in having glabrous
leaves, several flowers per inflorescence, and
densely tomentose ovary. However, it differs from
R. cavaleriei in having prominent lateral primary
veins on both surfaces, the calyx lobes membra¬
naceous and up to 2 cm long, and the ovary trun¬
cate at the apex. Rhododendron kaliense Fang & M.
Y. He, R. linearicupulare Tam, R. huguangense
Tam, and R. longilobum all have well-developed
calyces, and the calyx lobes are variable in size.
However, R. longilobum is easily distinguished
from R. kaliense, R. linearicupulare, and R. hu¬
guangense by the characters of the ovarv densely
flavescent tomentose and truncate at the apex. The
ovary is glabrous and tapering at the apex of R.
kaliense, R. linearicupulare, and R. huguangense,
which is different from R. longilobum.
Acknowledgments. We thank Fang Rhui-cheng
(KUN) and Li Xi-wen (KUN) for their help review¬
ing this manuscript. We are thankful to Tan Hai-
ming for his assistance during our fieldwork. We
also thank Wu Xi-lin and Yang Jian-kun for their
drawings. I his work was funded by a keynote pro¬
ject of biology of the knowledge innovation program
of the Chinese Academy of Sciences (KSCX 2-1-
068).
Literature Cited
Fang. R. C. & I. 1.. Ming. 1995. The floristic study on the
genus Rhododendron. Acta Bot. Yunnan. 17(4): 359-
379.
He, M. Y„ M. Y. Fang, W. G. Hu & L Z. Hu. 1994.
Ericaceae. In Flora Reipublicae Popularis Sinicae
57(1): 340-366.
Philipson. W. R. X M. N. Philipson. 1986. A revision of
Rhododendron III. Subgenera Azaleas!rum. Mumeaza-
lea, Candidastrum and Therorhodion. Notes Roy. Bot.
Card. Edinburgh 44(1): 1-23.
Primula calyptrata , a New Species in Section Carolinella
(Primulaceae) from Yunnan, China
Gong Xun and Fang Rhui-cheng
Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China.
gongxun@mail.kih.ac.cn
Abstract. Primula calyptrata X. Gong X R. C.
Fang (Primulaceae) is described and illustrated.
Analysis of its habitat, distribution, and capsule
morphology indicates that this species is a member
of Primula sect. Carolinella (Helmslev) Pax, which
is endemic to humid subtropical regions of Yunnan,
China, northern Vietnam, and Thailand, and char¬
acterized by an unusual calyptrate capsule.
Key words: China, Primula, Primulaceae, sec¬
tion Carolinella . Yunnan.
During an investigation of plants in southeastern
Yunnan, China, in April 2001. some specimens of
Primula were collected and recognized as a new
species. After further studies, the species was be¬
lieved to belong to section Carolinella. Living
plants were introduced into Kunming Botanical
Garden for cultivation and observation. Specimens
were also collected by Shui Yu-min et al. from
Zhongzai in Malipo County, Yunnan Province, in
2002.
Members of section Carolinella are unique in
Primula for having a calyptrate capsule with a cir-
cumscissile lid, a feature also shared by the genera
Pomatosace Maximowicz, Anagallis I... Soldanella
I... and Bryocarpum Hooker f. & Thomson in Pri¬
mulaceae (Pax & Knuth, 1905). file evolutionary
trend of the fruits in Primulaceae is believed to be
from indehiscent capsule to valvate capsule to ca¬
lyptrate capsule with a circumscissile lid (Hu Q-
M.. 1994; Chen F-H. & Hu, 1990). This feature in
section Carolinella is a link between Primula and
the other genera; however, other morphological
characters and distribution clearly separate them.
The phylogenetic relationships among other sec¬
tions in Primula and other genera of Primulaceae
are still unclear because studies are lacking in
chromosome number, pollen type, and seed mor¬
phology. The phenomenon that the calyptrate cap¬
sule occurs in the genera Pomatosace, Anagallis,
Soldanella, and Bryocarpum in Primulaceae may be
a result of convergent evolution (Kong X Liu,
1999).
Section Carolinella is believed to be a primitive
group in Primula (Hu Q-M., 1994); this new taxon
is considered to be primitive for having alternate
and long-petioled leaves, 2—2.5 cm long stems, and
homostylic flowers. In addition. Primula sect. Car¬
olinella is a typical endemic of East Asia and is
limited to southwestern and southern China and ad¬
jacent North Vietnam and Thailand. Eleven species
were recognized in this section: almost all are sten-
oehoric and have very small populations, some of
which are in imminent danger. All of the species
in this section occur in the warm-moist southern
subtropical evergreen broadleaf lorest oi low alti¬
tudes within the limestone region.
Primula calyptrata X. Gong X R. C. Fang, sp.
nov. TYPE: China. SE Yunnan: Malipo County,
Xiajinchang, 1870 m. 16 Apr. 2001, X. Cong
02815 (holotvpe, KUN). Figure 1.
Species in hac sectione caulibus brevibus, foliis subtus
purpuratis valde differt. Planta P. intanoensi e Thailandia
valde proxima, sed caulibus 2—2.5 cm longis, foliis alter-
nantibus, corollis purpureis, tubis 1.7 cm longis, 1 imbis
1.5— 1.8 cm in diam., pedicellis 2^1 mm longis differt.
Herbs perennial, completely efarinose, stems
and both surfaces of leaf blade as well as petioles
with dense articulated hairs. Rhizome elongated, 2
cm long, with numerous fibrous roots, producing 1
or 2 leaf-bearing stems above. Stem 2—2.5 cm long
before or in anthesis, purple. Leaves alternate, leaf
blade suborbicular, 2.8—4 X 2.6—3.8 cm, apex
rounded, base cordate, margin irregularly undulate,
thick-papyraceous when dry, grayish green above,
purple below, midrib and 4 pairs of lateral veins
obscure above and plane but distinct below; petiole
2.5— 5.5 cm long. Scape 1 or 2. 12—18 cm high,
sparsely white villous, with 1 to 2 superimposed
umbels with 2 to 5 slightly seeund flowers in each;
pedicels 2-4 mm long, with dense articulated hairs;
bracts ovate-lanceolate, 4—6 mm long, prominently
1-nerved, with dense articulated hairs; calyx cam-
panulate, 5—6 mm long, with dense articulated
hairs, divided to 1/2—1/3 into ovate-triangular
lobes, lobe with 3 distinct nerves; corolla purple
with a yellow eye, salverform. pubescent outside,
Novon 13: 193-195. 2003.
194
Novon
Figure 1. Primula calyptrata X. Gong & R. C. Fang. —A. Habit. —I!. Opened corolla, internal view showing the
position of the stamens and hairs. —C. Pistil. —I). Opened calyx, external view—E. Capsule, showing dehiscent
manner. Drawn by Yang Jian-kun from the holotype, X. Gong 02815 (RUN).
Volume 13, Number 2
2003
Gong & Fang
Primula calyptrata from Yunnan
195
tube 1.7 cm long with a hairy annulus inside, limb
1.5—1.8 cm diam., lobes oblong-obovate, deeply
emarginate; homostylous, stamens inserted in
throat, anthers below annulus; style ea. 1.7 cm,
subequal to corolla tube. Capsule enclosed in calyx
tube, dehiscing by means of a ealyptra.
Distribution. Known only from the type and
paratype specimens collected from both Xiajin-
chang and Zhongzai forests of Malipo County of
southeastern Yunnan, China.
Habitat and phenology. Understory species in
an evergreen broad-leaf forest composed of mem¬
bers of the Magnoliaceae, Fagaeeae, and Laura-
ceae, growing on limestone with moss; alt. 1700—
1870 m; (lowering March-April, fruiting in August.
Primula calyptrata is distinguished from the oth¬
er species of Primula sect. Carolinella by having
short stems and a leaf blade that is purple on the
abaxial surface. This species is closely related to
Primula intanoensis T. Yainazaki collected from
Thailand, but can be distinguished by stems 2-2.5
cm long, leaves alternate, corollas purple, tubes 1.7
cm long, limbs 1.5—1.8 cm in diameter, and pedi¬
cels 2-4 mm long. Primula intanoensis is stemless,
with leaves in a rosette, corollas white, tubes a little
longer than to hall again as long as the calyxes,
limbs 0.8 cm in diameter, and pedicels 5—7 mm
long.
Paratype. CHINA. SE Yunnan: M alipo County,
Zhongzai, 1700 m, 23 Mar. 2002, Yumin Shui el al. 20336
(KUN).
With the inclusion of the species described here,
11 species are known in Primula sect. Carolinella.
Key to the Species of Primula section Carouneua
la. Inflorescences umbellate, umbels 1 or 2 super¬
imposed, 2- to 6(9)-flowered; leaf blade pubes¬
cent.
2a. Leaf blade suborbieular or ovate-elliptic,
base distinctly cordate, both surfaces
markedly pubescent; flowers homostylous.
3a. Stem 2—2.5 cm long, leaves alter¬
nate; corolla purple, tube 1.7 cm
long, 2—3 times as long as calyx,
limb 1.5—1.8 cm diam.; China . . .
. . P. calyptrata X. Gong & R. C. Fang
3b. Stem absent; leaves a rosette; corolla
white, tube a little longer than to
half as long again as the calyx, limb
ca. 0.8 cm diam.; Thailand . . .
. P. intanoensis T. Yamazaki
2b. Leaf blade oblong, elliptic, broadly ovate
or broadly elliptic, base broadly cuneate
or cordate, both surfaces pubescent or
adaxially glabrous; flowers heterostylous.
4a. Pedicel and calyx glabrous;
leaves densely pilose on both
surfaces; China.
P. levicalyx C. M. Hu & Z. R. Xu
4b. Pedicel and calyx fulvous pi¬
lose; leaves pubescent on mar¬
gin and abaxial surface.
5a. Leaf blade subcordate or
rounded at base; calyx
4.5—7.5 mm. parted to
middle; China.
... 8 wangii F. H. Chen &
C. M. Hu
5b. Leaf blade cuneate or
rounded at base.
6a. Calyx 3-4 mm,
parted to 2/3;
China .
. ... P. kweichouensis
W. W. Smith
6b. Calyx 6—7 mm,
parted to 1/3;
China .
. . P. kwangtungensis
W. W. Smith
lb. Inflorescences abbreviated or almost umbellate,
(6)12- to 28-flowered; leal blade glabrous or
glabrescent; flowers heterostylous.
7a. Leaf blade acute or acuminate at apex.
8a. Leaf blade cuneate or rounded at
base; calyx ca. 3 mm, inconspicu¬
ously 5-veined; China, Vietnam . . .
. P. henryi (Hemsley) Pax
8b. Leaf blade cordate at base; calyx 7—
10 mm, prominently 10-veined; Chi¬
na, Vietnam . . P. chapaensis Gagnepain
7b. Leaf blade obtuse or rounded at apex.
9a. Leaf blade cordate at base.
10a. Leaf blade elliptic-ovate to
broadly ovate, 4.5-6 X 3.5—5.5
cm; petiole 2—5 cm long; Viet¬
nam .
. . P. cardioeides \\. W. Smith & H.
R. Fletcher
10b. Leaf blade oblong to suborbic-
ular or ovate-rounded, 7—18 X
5—15 cm; petiole 7-26 cm
long; China . . P. partschiana Pax
9b. Leaf blade broadly cuneate to sub¬
rounded at base; China.
. P. rugosa N. P. Balakrishnan
Acknowledgments. The authors are grateful to
Pan Yue-zhi for help with fieldwork and Yang zhi-
Yun for cultivating and observing the plants.
Literature Cited
Chen F-H. & Hu Q-M. 1990. Primulaceae. In: Flora Rei-
publicae Popularis Sinicae 59(2): 59-66. Science Press,
Beijing.
Hu Q-M. 1994. On the geographical distribution of the
Primulaceae. J. Trop. Subtrop. Bot. 2(4): 1-14.
Kong H-Z. & Liu J-Q. 1999. Karyomorphology of the ge¬
nus Pomatosace Maxin. (Primulaceae). Act. Phytotax.
Sin. 37(5); 445-450.
Pax, F. & R. Knuth. 1905. Primulaceae. In: A. Engler,
Das Pflanzenreich. Vol. 27: 1—15. Verlag von Wilhelm
Engelmann, Leipzig.
Two New Species of Primula (Primulaceae) from China
Hu Chi Ming
South China Institute of Botany, Academia Siniea, Guangzhou 510650,
People’s Republic of China, huqm@scih.ac.cn
Ceng Yu Ying
West China Subalpine Botanical Garden, Institute of Botany, Academia Siniea,
Dujiangyan 61 1830, China
ABSTRACT. Two new species of Primula from
western China are described, illustrated, and com¬
pared with putatively related species. Both new
species belong to section Petiolares subsect. Davi-
dii (Balfour I.) Craib, which is characterized bv the
capsule crumbling irregularly at maturity, the basal
buds always having a covering of paleaceous scales,
and the leaves being more or less coriaceous, per¬
sisting into the following spring, with veins often
prominently raised and alveolate abaxially and im¬
pressed adaxially. In this subsection there are about
18 species all distributed in Sichuan or Yunnan,
with only one species extending to Guizhou, Hubei.
Hunan, and Jiangxi.
Key words: China, Primula , Primulaceae.
Since the publication of the Primulaceae treat¬
ment in the Flora Reipublicae Popularis Sinicae
(Hu, 1990), the authors carried out a project of in¬
troducing wild species of Primula for cultivation in
the West China Subalpine Botanical Garden in Du¬
jiangyan City in Sichuan Province. In the process
we rediscovered P. davidii Franchet anti P. filchner-
ae knuth, both of which were previously known
only from the type material collected nearly a cen¬
tury ago. In addition, we discovered two new spe¬
cies belonging to section Petiolares subsect. Davi¬
dii, characterized by the capsule crumbling
irregularly at maturity, the basal buds always hav¬
ing a covering of paleaceous scales, and the leaves
being more or less coriaceous, persisting into the
following spring, with veins often prominently
raised and alveolate abaxially and impressed adax¬
ially. In this subsection there are about 18 species
all distributed in Sichuan or Yunnan, with only one
species extending to Guizhou, Hubei, Hunan, and
Jiangxi. The new species are described here.
Primula bergenioides C. M. Hu A Y. Y. Geug,
sp. nov. TYPE: China. Sichuan: Emei Shan,
Jing Shui, alt. 1105 m, 17 Mar. 1994, Y. Y.
Ceng l )4—()()d (holotype, I BSC). Figure 1.
Species affinis P. hylobiae et P. klaverianae, a quibus
foliis scapisque glabrissimis, calyce tantum ail tertian]
partem in lobos ovatos fisso differl; ab altera etiam floribus
heterogeneis differl.
Perennial herbs, efarinose, with a short compar¬
atively stout rhizome, basally with brown palea¬
ceous bud-scales at flowering time. Leaves forming
a spreading rosette, to 30 cm diam.; leaf blade ob¬
long-elliptic to elliptic-obovate, 12—20 X 6—9 cm.
apex rounded, tapering from the middle into a very
short or almost obsolete winged petiole, irregularly
crenate-dentate, papery when dry, ± bullate on the
margin and covered with minute glandular hairs
when just unfolded from the resting buds, soon gla-
brate, smooth and quite glabrous on both surfaces
when mature; lateral veins 11 to 13 pairs, plane
adaxially, raised abaxially, subparallel, angled to
midrib ca. 65 Q , secondary veins obscure on both
surfaces. Scapes 13—20 cm, I to 2 per plant, gla¬
brous. carrying a terminal umbel of 5 to 1 I flowers;
bracts lanceolate, 5—8 mm long, brown, paleaceous,
glabrous; pedicels 1 — 1.5 cm, elongated to 2.5 cm
in fruit, sparsely glandular-puberulent. Flowers
heteromorphie; calyx narrowly campanulate, 10—12
mm long, glandular-puberulent, 5-veined, split to a
third of its length; lobes ovate, ca. 3.5 mm wide,
apex acute; corolla funnel-shaped, lavender-blue,
exannulate. Thrum flower: corolla tube ca. 13 mm
long, cylindrical, ampliate above insertion of sta¬
mens; limb 1.8— 2.5 cm across; lobes obovate, 9 X
8 mm, apex emarginate; stamens inserted at apex
of corolla tube; filaments ca. 1.5 mm; anthers lin¬
ear, ca. 2 mm long; style 5—6 mm long; stigma cap¬
itate. Capsule subglobose, included in calyx tube.
Distribution and habitat. known only from the
type locality, growing in fissures on dripping wet
sandstone cliffs.
Primula bergenioides is most closely related to
P. hylobia W. W. Smith and P. klaveriana Forrest.
From both it can be easily distinguished by its gla¬
brous leaves and scapes and by its calyx splitting
No VON 13: 196-199. 2003.
Volume 13, Number 2
2003
Hu & Geng
Chinese Primula
197
198
No von
Figure 2. Primula tenuituba C. M. Hu & Y. Y. Geng. —A. Habit. —B. Calyx opened up to show pistil. —C. Short-
styled flower: portion of opened corolla showing stamens. (Drawn from the holotype.)
Volume 13, Number 2
2003
Hu & Geng
Chinese Primula
199
only to a third of its length (not to the middle as in
the above-mentioned species). Moreover, it stands
apart from P. klaveriana by its heterogeneous flow¬
er.
The specific epithet refers to the comparatively
large and dark green leaves, which look somewhat
like those of Bergenia.
Paratype. CHINA. Sichuan: Kmei Shan, .ling Shui, 7
May 1990, C. M. Hu 9067 (IliSC).
Primula tenuitulm C. M. Hu & Y. Y. Geng, sp.
nov. TYPE: China. Sichuan: Ying Jing Xian.
Hua Tan Xiang, Li Jia Yan, alt. 800 m, 14 May
1992. V. E Geng, Z. L Zhao et al. 92001 (ho-
lotype, IBSC). Figure 2.
Species valde affinis P. epilosae Craib, sed (loribus mi-
noribus, tubis corollarum tenuioribus, foliis scapis inflo-
rescentiisque glabris, rete venularum invisibili differt.
Perennial herbs with a comparatively stout rhi¬
zome from which arise numerous fibrous roots and
clothed at base by withered old leaves. Leaves
forming a rosette to 15 cm diam.; leaf blade 5—8 X
1.5—3 cm at flowering time, 7-11 X 2.5—3.5 cm at
fruiting, oblong-obovate to oblanceolate, tapering
from a rounded apex into a very short or almost
obsolete winged petiole, irregularly crenate-dentate
at margin, thickly papery, adaxial surface scurfy-
glandular at flowering, glabrous when mature, ±
white farinose on abaxial surface; lateral veins 8 to
11 pairs, prominent on abaxial surface, angled to
the midrib ca. 60 e . ending in teeth, secondary veins
very obscure or invisible. Scapes 4.5—6 cm long, I
to 4 per plant, glabrous, carry ing a terminal umbel
of 3 to 7 flowers; bracts 3—4 mm long, acuminate
from the broad base, brown, paleaceous; pedicels
1.2-1.5 mm, lengthening to 2 cm in fruit, glabrous.
Flowers heteromorphic; calyx campanulate, ca. 7
mm long, 5-veined, glabrous, split almost to mid¬
dle; lobes ovate, acuminate; corolla violet-blue.
Thrum flower: corolla tube slender, ca. 12 mm long,
ampliate above insertion of stamens, limb ca. 1.8
cm diam.; lobes obovate, ca. 7 mm long, deeply
emarginate; stamens inserted near apex of corolla
tul >e; filaments very short; anthers ca. 1.5 mm long;
ovary glabrous; style ca. 5 mm long. Capsule glo¬
bose, slightly shorter than the persistent calyx.
Distribution and habitat. Known only from the
type locality in Sichuan, growing on exposed cliffs
near a waterfall; alt. ca. 800 m.
Primula tenuituba is most closely related to P
epilosa Craib, from which it differs by the smaller
flowers with a very slender corolla tube and by the
leaves being glabrous and not strongly reticulate
with a network of veinlets on the abaxial surface.
The specific epithet refers to the slender corolla
tube of the plant.
Paratype. CHINA. Sichuan: Ying Jing Xian, Hua
Tan Xiang. Li Jia Yan, alt. 800 m, 17 May 1990, Y. Y.
Geng, Z. /.. Zhao et al. 90068 (IBSC).
Acknowledgments. Phis study was supported by
a grant from the National Science Foundation of
China (39970053). We thank Ihsan A. Al-Shehbaz
for his review of the manuscript prior to submis¬
sion.
Literature Cited
Hu. C. M. 1990. Primulaceae (2). In: F. II. Chen & C. M.
Hu (editors). Flora Reipubl. Popularis Sin. 59(2): 1—
317. Science Press, Beijing.
A New Subspecies of Hippophae (Elaeagnaceae) from China
Lian Yongshan, Chen Xuelin, Sun Kun*, and Ma Ruijun
Institute of Botany, Northwest Normal University, Lanzhou 730070, Gansu, China.
* Author for correspondence: Ksun@nwnu.edu.cn.
ABSTRACT. In this paper, a new subspecies of
Hippophae (Elaeagnaceae), H. rhamnoides subsp.
wolongensis Y. S. Lian, K. Sun & X. L. Chen, from
China is described and illustrated. According to an
analysis of the distribution and main characters, it
is allied to H. rhamnoides subsp. sinensis Rousi but
is distinct in having rosulately arranged flower
buds, larger, alternate leaves with usually more or
less sinuous margins, longer petioles, and longer
peduncles. Hippophae rhamnoides subsp. wolon¬
gensis is rare and occurs only in Wenchuan and
Mao Counties of Sichuan Province, at 1660-1920
m. A key including five subspecies of Hippophae
rhamnoides in China is provided.
Key words: China, Elaeagnaceae, Hippophae,
Sichuan.
Hippophae L. is a small genus of Elaeagnaceae
in which 6 species and 10 subspecies have been
recognized (Lian, 2000). These species are distrib¬
uted widely but sparsely in Asia and Europe. Most
of them, however, are restricted to the Oinghai-
Tibetan plateau and adjacent areas, and only the
species Hippophae rhamnoides I,, occurs in both
Asia and Europe. All of the species and six of the
subspecies are found in China (Lian, 1900, 2000;
Lian & Chen. 1996).
'flic authors, investigating populations of Hippo¬
phae rhamnoides in Gansu, Qinghai, Sichuan, Yun¬
nan, and Xizang (Tibet) Provinces in 2000, found
several unusual populations from Wenchuan and
Mao Counties in Sichuan. These populations were
found to occur together with populations of //.
rhamnoides subsp. sinensis in the above locations
but at different elevations. After critical study
based on ample specimens gathered from the spe¬
cial populations, the specimens were found to rep¬
resent a new subspecies of H. rhamnoides. Follow¬
ing are the description and discussion of the new
subspecies. A key including five subspecies of Hip¬
pophae rhamnoides in China is also provided.
Hippophae rhamnoides L., Sp. PI. 1023. 1753.
Hippophae rhamnoides , which comprises nine
subspecies, is now widely distributed in temperate
Asia and Europe. Subspecies rhamnoides , subspe¬
cies fluviatilis V. Soest, and subspecies carpatica
Kousi are represented in Europe, and subspecies
caueasia Rousi, subspecies turkestanica Rousi.
subspecies mongolica Rousi, subspecies sinensis
Rousi, subspecies yunnanensis Rousi, and the new
subspecies wolongensis are represented in Asia.
Subspecies sinensis, subspecies yunnanensis, and
subspecies wolongensis are restricted to China.
Hippophae rhamnoides subsp. wolongensis Y.
S. Lian, K. Sun & X. L. Chen, subsp. nov.
TYPE: China. Sichuan: Wenchuan, Wolong,
1920 m, in the Sea Buckthorn shrub commu¬
nity on mountain slopes or river banks (fruit),
25 Sep. 2000. Y. S. Lian, X. L. Chen & K. Sun
nil (holotype [pistillate], NWTC). Figure I.
Subspecies haec margine foliorum plus minusve sinu-
ata, petiolis longioribus 2-4 mm longis, pedunculis 3-5
mm longis, gernmis florum masculorum 5—1 I. femineorum
5—7, rosulatim insertis, fructibus et seminibus conspicue
minoribus differt.
Shrubs or small trees, deciduous, 3—5 m high;
usually much branched, young branches red-
brown, striate, and covered with scales on the sur¬
face; old branches dark brown, constricted into a
spine tip. Leaves usually alternate, or sometimes
subopposite on the upper branchlets, elliptic-lan¬
ceolate, 50-85 X 6-16 mm, acute or acuminate at
the apex, base cuneate or broadly cuneate, blade
margins usually ± sinuous, upper blade surface
green, covered by scales, often early deciduous,
lower blade surface covered by dense, gray scales,
sometimes mixed with a few red-brown scales; cos¬
tae impressed in a conspicuous groove on the upper
surface; petioles 2—4' mm long. Staminate flower
buds 5-11, pistillate 5-7, rosulately arranged, with
red-brown scales on the outside. Fruits yellow,
subglobose, 4—5.5(6) X (4.5)5—6(7) mm, with
sparse red-brown scales; peduncle 3—5 mm long.
Seeds obovate to oblong, mueronate at the apex, 3—
4 X ca. 1.5 mm, seed coat easily separated from
endocarp. Flowers were only collected in bud; fruit¬
ing in September to October. This subspecies is
distributed in the shrub community on mountain
slopes at 1660-1920 m.
Nov on 13: 200-202. 2003.
Volume 13, Number 2
2003
Lian et al.
New Subspecies of Hippophae
201
Figure I. Hippophae rhamnoides subsp. wolongensis Y. S. Lian, K. Sun & X. I,. Chen. —A. Habit of pistillate plant
with fruits. —B. Pistillate flower bud. —C. Seed. —I). Habit of staminate plant with flower buds. —F,. Staminate flower
bud. —F. Basal leaf blade of staminate plant showing petiole and scales. —G. Close-up of scale trichomes. (A, B, C,
F & C based on E S. Lian, X. L. Chen & K. Sun wl 1 (pistillate, NWTC); I) X E based on E S. Lian, A". L. Chen &
K. Sun w21 (staminate, NWTC); drawn by Jianlu Bai.)
202
No von
This subspecies has an affinity with Hippophae
rhamnoides subsp. sinensis in several characters,
including the endocarp easily separating from the
seed coat, the length of the fruit usually less than
or equal to the width, and the leaves relatively larg¬
er. They dilfer in that the flower buds of subspecies
wolongensis are rosulately arranged while those of
subspecies sinensis are 4-angled, tower-shaped in
staminate form and cruciform in pistillate form;
larger, alternate leaves with usually more or less
sinuous margins in subspecies wolongensis vs.
smaller, opposite or subopposite leaves with entire
margins in subspecies sinensis ; and longer petioles
(2—4 mm long vs. 1.5—3 mm) and peduncles (3—5
mm vs. 1—2.5 mm) in subsp. wolongensis than in
subsp. sinensis. Subspecies wolongensis is confined
to Wenchuan and Mao Counties of Sichuan Prov¬
ince, but subspecies sinensis is widely distributed
in southwestern, northwestern, and northern China.
Hippophae rhamnoides subsp. wolongensis occurs
together with subspecies sinensis in Wenchuan and
Mao Counties; however, subspecies wolongensis oc¬
curs at 1660—1920 m while subspecies sinensis is
found at over 2100 m.
Paratype. CHINA. Sichuan: Mao County, Changhai,
1660 m. in the Sea Buckthorn shrub community on moun¬
tain slopes (flowers in buds), 26 Sep. 2000, Y. S. Lian, X.
L. Chen & K. Sun w21 (staminate, NWTC).
Key to Five Subspecies of Hippophae rhamnoides in
China
la. Flower buds rosulately arranged or alternatively
opposite or subopposite; length of fruits usually
less than or equal to breadth.
2a. Leaves elliptic-lanceolate, blade margins usu¬
ally more or less sinuous; flower buds rosula¬
tely arranged; peduncles 3—5 mm long . . .
...//. rhamnoides subsp. wolongensis Y. S. Lian,
K. Sun & X. L. Chen
2b. Leaves lanceolate or narrowly lanceolate,
blade margins entire; flower buds opposite
or subopposite; peduncles 1—2.5 mm long.
3a. Branehlets stiff; most leaves opposite or
subopposite, lower blade surface cov¬
ered with white scales or rarely mixed
with rusty-red scales, petioles 1.5—3 mm
long; staminate flower buds manifestly
4-angled, tower-shaped, pistillate flower
buds cruciform; most fruits tangerine,
endocarp separating easily from the
seed coat .
. H. rhamnoides subsp. sinensis Bousi
3b. Branehlets soft; most leaves alternate,
lower blade surface covered with rusty-
red scales, petioles 1—1.3(2) mm long;
staminate flower buds inconspicuous, 4-
angled, tower-shaped, pistillate flower
buds near cruciform or bifid ovate; most
fruits yellow, endocarp sometimes hard
to separate from the seed coat ....
. . //. rhamnoides subsp. yunnanensis Bousi
1 b. Flower buds spirally arranged; the length of
fruits usually longer than breadth.
4a. Branches silvery, usually with more-
branched thorns; leaf blades 2—4(5) mm
wide, upper surface silvery; peduncles 3—
4(7) mm long.
.//. rhamnoides subsp. turkestanica Bousi
4b. Branches brown, with less- and un-branched
thorns; leaf blades (3)4—8 mm wide, upper
surface often green: peduncles 1—4 mm long
. H. rhamnoides subsp. mongolica Bousi
Acknowledgments. We thank the reviewers for
providing useful remarks and suggestions. The pre¬
sent study was sponsored by the National Natural
Science Foundation of China (30270091) and the
Science and Knowledge Innovation Project of
Northwest Normal University (No. 02) in Lanzhou,
Gansu. China.
I .iterature Cited
Lian, Y. S. 1988. The new discoveries of the genus Hip¬
pophae. Acta Phytotax. Sin. 26: 235-237.
-(editor). 2000. The Plant Biology and Chemistry
of the Genus Hippophae. Gansu Science and Technology
Press, Lanzhou.
-& X. L. Chen. 1996. The systematic classification
of the genus Hippophae. Hippophae 9(1): 15—24.
A New Species of Cyrubidium (Orchiclaceae) from Tibet
(Xizang), China
Chunlin Long, Zhiling Dao, and Heng Li
Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming,
Yunnan 650204, People’s Republic of China, long@mail.kib.ac.cn
ABSTRACT. Cymbidium chawalongen.se C. L.
Long, H. Li & Z. L. Dao (Orchidaceae), collected
from southeast Xizang (Tibet). China, is described
as new. The new species is similar to C. floribun-
dum Lindley in morphology and habitat, hut differs
in having an erect scape, very widely spread flow¬
ers, pale green sepals and petals, and a white, non-
mottled lip.
Key words: China. Cymbidium, Orchidaceae,
Tibet, Yunnan.
There are about 48 species in the genus Cym¬
bidium Swartz, distributed iu tropical and subtrop¬
ical regions of Asia and Australia (Chen, 1999).
About 29 species are distributed in China, of which
28 are found in Yunnan Province (Chen, 1999).
This may suggest that the Chinese center of species
diversity of the genus is Yunnan. Because of the
particular significance of Cymbidium to horticulture
and traditional culture in China and its neighboring
countries, the discovery of a new member of the
genus will stimulate much interest among horticul¬
turists anil orchid enthusiasts (Yang et al.. 1993).
The new Cymbidium species was collected during
a recent biodiversity expedition and appears differ¬
ent from all other species in the genus.
The distribution of this species includes the type
locality in southeast Tibet and extends into north¬
west Yunnan. This area lies at the conjunc tion of
two of the world’s most biologically rich and most
threatened environments, i.e., the Indo-Burma and
South-Central China hotspots of biodiversity (Myers
et al.. 2000). Some 5000 higher plant species have
been recorded from the Gaoligong Mountains be¬
tween the great Salween and Irrawaddy Rivers (Li
et al., 2000) in this region. Among them, 266 spe¬
cies are orchids. At least 10 orchid species, in¬
cluding Cymbidium gongshanense H. Li & K. M.
Feng, Calanthe dulongense 11. Li, Paphiopedilum
markianum Fowlie, and Coelogyne tarongen.se Han-
del-Mazzetti, are considered endemic to this region
(Li et al., 2000). This new orchid species is also
endemic to the Gaoligong Mountains.
Gymhidiiun chawalongense C. I.. Long. II. Li &
Z. L. Dao, sp. nov. TYPE: China. Xizang [ Ti¬
bet |: Chayu Xian, Chawalong, 2300 m, in co¬
niferous forest, flowers pale greenish, 14 May
2001, Gaoligongshan Expedition 13727 (ho-
lotype, KUN). Figure 1.
Species nova Cymbidio floribundo affinis, sed scape er-
ecto 22 cm longo, Horibus patulis, sepalis et petalis chlo-
roticis, labello albo non maculato differt.
Plants terrestrial, to 43 cm tall, with a short rhi¬
zome. Pseudobulbs congested, cylindric, slightly
flattened, 1.5 cm long and thick, enclosed in cat-
aphylls and leaf bases. Cataphylls 3 or 4. widely
ovate and lanceolate, 2—5.5 X 2-2.5 cm at base,
acuminate at apex. Leaves usually 5 per pseudo¬
bull), dark green, thinly coriaceous, slightly arch¬
ing, 27—43 cm long (including sheath), 1.4 cm
broad, apex acute and unequally retuse, base
slightly narrowed, plicate, articulate, midrib de¬
pressed above and prominent below; sheath green,
2.5—5 cm long. Scape lateral from pseudobulb base,
erect, robust, to 22 cm tall; peduncle pale green, 9
cm long, 3—4 mm diam., base enclosed in 5 or 6
cataphylls; raceme green, 10 cm long, loosely 17-
flowered; pedicel and ovary green-tinged, cylindric,
3.3 cm long, shortened upward, the terminal one
2.5 cm long; bracts small, green, deltoid, 2—2.3 cm
long; flowers widely spread, to 4 cm across; sepals
and petals pale green, lip white, not mottled; mid¬
dle sepal oblong, 2 cm long, 3 mm broad, apex
acute, margins slightly recurved, 3-nerved; lateral
sepals similar to middle but falcately curved, 2.2
X 0.4 cm, 5-nerved; petals elliptic, 1.8 X 0.7 cm,
margins slightly recurved, 5-nerved; lip sub-ob-
long, to 1.8 cm, 3-lobed, lateral lobes erect, to ca.
1.5 mm; mid-lobe subovate, 1 X 0.5 cm, smooth,
recurved; disc with 2 yellow lamellae; column sem-
icylindric, arcuate, erect, to 1.5 cm long, to 4—5
mm broad al upper part. Anther yellow 7 , eucullate;
pollinia 2, waxy, ovoid, ca. 1 mm diam., stipe short,
viscid disc oblong; ovary ca. 2.5 mm diam., stigma
transversely oblong, deeply concave. Flowering in
May.
Novon 13: 203-205. 2003.
204
Novon
Volume 13, Number 2
2003
Long et al.
Cymbidium chawalongense from Tibet
205
Distribution. Endemic to China from Chayu in
southeastern Xizang (Tibet) and northwestern Yun¬
nan. Populations were occasionally seen in Gong-
shan Xian and Weixi Xian of northwestern Yunnan,
but specimens were not collected from these loca¬
tions to protect the natural populations. Two living
collections are being kept in the Kunming Botani¬
cal Garden. We saw the living plants in forests
three times and in the local markets two times. The
new species was found in coniferous forest, ever¬
green forest, and secondary successions, at alti¬
tudes from 1600 to 2300 m. Forests there mainly
consist of Pinus yunnanensis Franchet, Betula sp.,
Acer spp., Cinnamomum spp., Litsea spp., Lindera
spp., and Lithocarpus spp. Understory plants are
represented by Excoecaria acerifolia F Didrichsen,
Campylotropis spp., and Desmodium spp.
Notes. Both Cymbidium chawalongense and C.
floribundum are members of subgenus Cymbidium.
They are very similar in morphology by having
smaller flowers, lip and column separated, and two
pollinia. Their natural habitat is also similar. They
can, however, be easily distinguished because of
the very opened flowers, pale green sepals and pet¬
als. and the white, non-mottled lip of the new spe¬
cies.
Acknowledgments. We are grateful to Yitao Liu
for preparing the illustration. We also thank Li
Bong for his assistance in preparation of the man¬
uscript. The study is supported by the Natural Sci¬
ence Foundation of China (30170102), Yunnan
Province (2001C0058M & 2001PY0017), Ministry
of Science and Technology of China (2001 DEA
10009), and German Technical Cooperation Agen¬
cy (GTZ 00.2047.8-001.02) .
Literature Cited
Chen, S. C. [Chen, X. Q.] (editor). 1999. Flora Reipubli-
c-ae Popularis Sinicae (Tomus 18): 191—227. Science
Press, Beijing.
Li, II., H. J. Guo & Z. L. Dao (editors). 2000. Flora of
Gaoligong Mountains. Science Press, Beijing.
Myers, N., R. A. Mitternieier. C. G. Mittermeier, G. A. B.
da Fonseca & J. Kent. 2000. Biodiversity hotspots for
conservation priorities. Nature 403: 853—858.
Yang. Z. II., 0- T. Zhang, Z. Z. Feng, K. Y. Lang & H.
Li. 1993. The Orchids. China Fsperanto Press, Beijing.
A New Species of Posoqueria (Rubiaceae) from Bahia, Brazil
Leila Macias
Departamento de Botanica, Instituto de Biologia, Universidade Federal de Pelotas (UFPEL),
Caixa Postal 354, CEP 96010-900 Pelotas, RS. Brazil, lmacias@uol.com.br
Luiza Sumiko Kinoshita
Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas
(UNICAMP), Caixa Postal 6109, CEP 13083-970 Campinas, SP, Brazil.
luizakin@unicamp.br
ABSTRACT. Posoqueria bahiensis from southern
Bahia, Brazil, is described as a new species. It dif¬
fers most notably from all the other species of the
genus in being totally glabrous except at the corolla
fauce and anthers and in the densely multiflowered
inflorescence with ea. 40 flowers.
Key words: Brazil. Posoqueria, Rubiaceae.
During the revision of the genus Posoqueria Au-
blet, it was evident that specimens from southern
Bahia, Brazil, represent a new species. Posoqueria
bahiensis is herein described and illustrated.
Posoqueria is a Neotropical genus belonging to
the subfamily Ixoroideae, tribe Gardenieae, sub¬
tribe Gardeniinae (Robbrecht, 1988, 1993). The
Brazilian species were mainly treated by Schumann
(1889). The whole genus was treated by Macias
(1988). who recognized 14 species.
Posoqueria bahiensis L. Macias & L. S. Kinosh¬
ita, sp. nov. TYPE: Brazil. Bahia: Marau, res-
tinga, 6 Oct. 1965 (fl). R. P. Belem 1877 (ho-
lotype, UB; isotypes, CEPEG, IAN, NY, VEN).
Figure 1.
Posoqueriae latifoliae (Budge) Roemer & Schultes fo-
liorum textura tantum similis sed planta omnis glabra (fau-
ce et antheris pilosis), inflorescentiis densis ea. 40 floribus
differt.
Treelet up to 5 m high, entirely glabrous. Leaf
blades 13.8—32 X 6.8—23 cm, ovate, oblong to el¬
liptic, acute or acuminate at apex, entire or rarely
slightly undulate and slightly revolute at margin,
rounded at base, strongly coriaceous, dark green
and shining above, paler and slightly opaque be¬
neath, secondary nerves in 5 to 8 pairs, alternate
or subopposite; petioles 1.5—3 cm long; stipules in-
terpetiolar, 0.8-1.5 X 0.5—0.8 cm, deltoid, acute at
apex. Inflorescence almost spherical, ca. 30 cm
diarn.; 3-times branched eorymbiform cymes, pe¬
duncle 6—8 cm long; bracts linear, 5—8 nun long.
Novon 13: 206-208. 2003.
Flowers ca. 40, very fragrant, pedicels 0.6—1.3 cm
long; bracteoles linear, ca. 2 mm long. Hypanthium
ca. 2 mm long, obovoid, glabrous; calyx limb-tube
ca. 2 mm long with a few colleters inside at the
base; calyx lobes triangular, ca. I mm long, ciliate;
corolla hypocrateriform, 9-14 cm long, tube pale
green, 6—12 cm long, 1—2 mm wide at base, inter¬
nal proximal region minutely papillate, distal re¬
gion with conspicuous sparsely distributed round
papillae; fauce glabrous to sparsely villous; lobes
white, 1.5—1.8 X 0.3-0.5 cm, oblong, rounded at
apex. Filaments 1.8—2 mm long; anthers ca. 0.5 cm
long, linear-lanceolate, sparsely puberulous. Hy¬
panthium glabrous; style 4-4.9 cm long, stigma ca.
2 mm long. Fruits ovoid, coriaceous, 3.5—5.2 X
2.8—3.8 cm, apricot colored (in vivo), seeds black¬
ish, ca. 1.1 X 0.7 cm.
Distribution. In southern Bahia, Brazil, in dis¬
turbed rain forest.
Phenology. Flowering in October; fruit collect¬
ed in January to September.
Posoqueria bahiensis differs from other species
of the genus in the glabrescent corolla fauce and
anthers, the densely flowered inflorescence (ca. 40
flowers), and the strongly coriaceous leaves. Table
1 shows a character comparison among four species
of Posoqueria morphologically related to P. bahien¬
sis. Posoqueria bahiensis is similar to P. latifolia
(Budge) Roemer & Schultes in the texture of the
leaves, but differs by the dense inflorescence with
ca. 40 flowers. It is also similar to P. williamsii
Steyermark, but differs in leaf indumentum and
texture and number of flowers. The new species
also resembles P. maxima Standley, but this has
lax inflorescences with only 10 to 20 flowers. Po¬
soqueria bahiensis is known only from southern Ba¬
hia. Brazil.
Paratypes. BRAZIL. Itnhia: Parque Nacional de
Monte Pascoal, on NW side of Monte Paseoal at low al-
Volume 13, Number 2
2003
Macias & Kinoshita
Posoqueria bahiensis from Brazil
207
Figure 1. Posoqueria bahiensis L Macias & L. S. Kinoshita. —A. Flowering branch. —IF Hypanthiuni, external view.
—C. Ovary, longitudinal section showing the colleters alternating with the lobes. —D. Corolla fauce and anthers. —
E. Stamen, sparsely puberulous anthers. —F. Longitudinal section of the fruit. (A—E from Bel&m 1877, UB; F from
Harley et al. 17898, CEPEC.)
Iahle 1. Comparison of some morphological characters among five Posoqueria species.
208
Novon
titude, between IBDF field hut and gates of the Parque
Nacional. 39°25'W, 16°53'S, 12 Jan. 1977 (fr). R. M. Hur¬
ley et at. 1789H (CEPEC); Uruyuca, District of Serra
Grande, 7.3 km along road Serra Grande/1 tacarf*, Fazenda
Eagoa do Conjunto Fazenda Santa Cruz, 39°01'W,
14°25'S, 1 Fev. 1991 (fr), ./. C. Jardim el ul. 607 (CEPEC,
MO). I 1-21 Set. 1991, (fr), A. M. Carvalho et al. 3521
(CEPEC), 26 Oct. 1993 (H). U. W. Thomas el al. 10166
(CEPEC).
Acknowledgments. The authors thank Tarciso S.
Filgueiras for revision of the Latin diagnosis and
the curators of the following herbaria for making
their collections available for study: CEPEC, IAN,
NY. 1 11. VEN. They also thank CAPES for gran!
support for Leila Macias during studies for her
M.Sc. thesis.
Literature Cited
Macias, I.. 1988. Revisao Taxonomica do Genero Poso¬
queria Aubl. (Rubiaceae). Tese de Mestrado, Instituto
de Biologia. Universidade de Campinas, Campinas.
(Unpublished.]
Robbrecht, E. 1988. Tropical woody Rubiaceae. Charac¬
teristic features and progressions. Contributions to a
new subfamilial classification. Opera Rot. Belg. 1: 1-
271.
-. 1993. Supplement to tin* 1988 outline of the clas¬
sification of the Rubiaceae—Index to genera. In: E.
Robbrecht (editor), Advances in Rubiaceae Macrosys-
tematics. Opera Rot. Belg. 6: 173—196.
Schumann, K. 1889. Rubiaceae. Pp. 329—340 in C. F. P.
von IMartius, A. G. Fielder & I. Urban (editors). Flora
Brasiliensis. V. 6. Fleischer, Leipzig.
A New Species of Tillandsia (Bromeliaceae) from the Mexican
Yucatan Peninsula
Ivon M. Ramirez M. and German Carnevali F.C.
Herbario CICY, Centro de Investigation Cientifica de Yucatan, A.C., Calle 43 # 130,
Colonia Chuburna de Hidalgo, Merida 97200, Yucatan, Mexico.
ramirez@cicy.iTix; carneval@cicy.mx
ABSTRACT. A new species of Tillandsia is [im¬
posed. It is similar to Tillandsia brachycaulos
Schleehtendal and Tillandsia balbisiana Schultes
f., two sympatric species. The new species is most
similar to T. brachycaulos in flower and bract color,
but the elongated inflorescences resemble those of
T. balbisiana, although the branches are much
shorter.
Key words: Bromeliaceae, Mexico, Tillandsia,
Yucatan.
The Bromeliaceae of the Yucatan Peninsula have
recently been reviewed by Ramirez and Carnevali
(1999). Later, the same authors proposed the ad¬
ditional species, T. jaguactalensis I. Ramirez, Car¬
nevali & Chi (Ramirez et ah. 2000), bringing to 29
species the number of taxa in the genus that were
thus known to occur in the Mexican portions of the
Yucatan Peninsula. This makes Tillandsia the larg¬
est genus of epiphytes in the area, followed by the
orchid genera Epidendrum L. (12 species) and Tn-
chocentrum Poeppig & Endlicher s.l. (8 species)
(Carnevali et ah, 2001, and unpublished data). Fur¬
ther exploration and study of the area’s Bromeli¬
aceae will certainly continue to yield taxonomic
novelties.
The novelty here presented is noteworthy be¬
cause it seems intermediate between two sympatric
species, Tillandsia brachycaulos and Tillandsia
balbisiana, which are widespread and locally com¬
mon in most of the Yucatan Peninsula.
Tillandsia niaya I. Ramirez & Carnevali, sp. nov.
TYPE: Mexico. Yucatan: Municipio Merida,
Parque Nacional de Dzibilchaltun, vegetacion
de selva baja caducifolia, aprox. 89°35'49"W.
21°5'44"N. ago. 2001 |"planta epffita, flores
moradas, 1 6 2 abiertas simultaneamente, es-
tigma apenas saliendo por encima del tubo de
la corola, creciendo con Tillandsia brachycau¬
los Schltdh, T. balbisiana Schult. f., T. elon-
gata (Kunth.) var. subimbricata (Baker) I,. B.
Sm.. T. paucifolia Baker”]. /. Ramirez, G. Car¬
nevali, F. Chi-May & U. Gonzalez 9 29 (holo-
type, CICY). Figure 1.
Species liaec T. brachycaulos affinis, sed plant is major-
ibus, foliis reflexis paucioribus, inflorescentiis scaposis,
ramulis pedunculum adpressis diversa; T. balbisiana ra-
mis infloreseentiae brevioribus non-complanatis, floribus
(asciculatis recedit.
Medium-sized epiphytic herbs, 28—40 cm tall.
Rosettes erect, with 16—20 leaves. Foliar sheaths
densely lepidote on both surfaces, membranaceous,
green, (4-)5-7 cm long, (2.5-)3-4 cm wide, oblong,
attenuate toward the apex; lamina narrowly trian¬
gular. attenuate, apex recurved, twisted when dry,
green on both surfaces, minutely lepidote, (12—)30—
35 cm long, (1,5-)1.8-2(-3) cm wide, somewhat
concave. Inflorescence a panicle with short, ap-
pressed branches, 27-35 cm long measured from
the base of the rosette, erect, cylindric, many-flow¬
ered, as long as to much longer than the leaves;
peduncle either partially enveloped (1/2 of its
length) by the primary bracts or completely ex¬
posed; 14-18 cm long measured from the base of
the rosette to the base of the fertile portion of the
inflorescence; fertile portion of the inflorescence
19-15 cm long, with 7 to 9 polystichously arranged
appressed branches; primary bracts from broadly
elliptic to ovate-elliptic, 17—28 cm long, free por¬
tion of the primary bract 8—12 mm w ide, foliaceous,
abruptly tapering into a narrowly triangular acu¬
minate lamina, diverging and exposing the apex of
the branch, basally red, apically green, minutely
lepidote on both surfaces, nerved when dry.
Branches appressed to the main axis or slightly di¬
verging, with 3—5 flowers each, 3—3.5 cm long, 0.9—
1 cm wide. Floral bract ca. 2 cm long, ca. 0.7 cm
wide, narrowly elliptic, acute, densely lepidote in¬
side, green outside, apically red, keeled, t lowers
4—4.5 cm long, erect, 1 or 2 open per branch at
any given time, thus inflorescences display several
flowers open simultaneously, each one lasting one
day; sepals 12-15 mm long, 3-4 mm wide, poste¬
rior ones connate for 2/3 of their length, the anterior
one connate for 1/3 of its length, white, apically
reddish, sparsely lepidote; petals imbricate, making
a tube-like corolla, 32—34 mm long. 5—6 mm wide.
No von 13: 209-211. 2003.
210
Novon
2. mm
Figure 1. Tillandsia maya I. Kamfrez & Carnevali. —A. Habit with inflorescence, showing also an old inflorescence
from the previous rosette. —B. Flower with floral bract anti fascicle bracts. —C. Flower with sepals. —L). Floral bract
flattened. —E. Sepals flattened. —F. Petal flattened. —G. Stamen with dorsifixed anther. —II. Ovary, style, and stigma
with details of the stigrnatic region. (Drawn from holotype, Ramirez el at. 929, CICY.)
Volume 13, Number 2
2003
Ramirez & Carnevali
Tillandsia maya from Mexico
211
base narrowly oblong, blade elliptic, obtuse, pur¬
ple, constricted at 14-13 mm from the base, base
narrowly oblong; stamens in two series, white, fil¬
aments 4 and 7 cm long, pollen bright yellow; an¬
thers 3 mm long, 1 mm wide, versatile, dorsifixed,
dark brownish; style 32-35 mm long, stigma con-
duplicate-spiral, with three white lobes, 2 mm long
each, surpassing the anthers in anthesis, white,
green—tinged; ovary 5 mm long, 2 mm wide, locules
2.5 mm long, 30 to 33 ovules per locule.
Etymology. The specific epithet honors the
Maya, the first people to inhabit the area where the
plants of this new taxon were collected.
Tillandsia maya belongs in subgenus Tillandsia
and seems closely related to Tillandsia balbisiana.
With this species, T. maya shares a similar vege¬
tative habit (pseudobulbose rosette) with curled
leaves and an elongated inflorescence. The rosettes
of T. maya, however, are less conspicuously pseu-
dobulbose and the leaves are comparatively less
lepidote, looking greener than in T. balbisiana. The
inflorescences are proportionally shorter in T. maya
than in T. balbisiana.
The inflorescence of Tillandsia maya is similar
to that of the recently described Tillandsia maypatii
I. Ramirez & Carnevali (Ramirez & Carnevali.
1999). Roth taxa share a condensed panicle with
sessile, appressed branches. Tillandsia maypatii.
however, features larger (5—5.5 cm long), white
flowers, more numerous leaves (30 to 40) in a more
open rosette. The inflorescences in T. maypatii are
also proportionally shorter and denser.
The habitat where this new taxon occurs is a low
caducifolious forest, dominated by Gymnopodium
floribnndum Rolfe trees, a member of the Polygon-
aceae. Tillandsia maya grows as an epiphyte at rel¬
atively low elevations (0.5—3 m) from the floor and
among several other epiphytic species such as Til¬
landsia elongata var. subimbricata (Baker) L. B.
Smith, T. recurvata (I..) L, T. fasciculata Swartz, T.
schiedeana Steudel, T. paucifolia Baker. Aechmea
bracteata (Swartz) Grisebach, and epiphytic orchids
such as Trichocentrum cebolleta (Jacquin) M. W.
Chase & N. H. Williams and Harrisella porrecta
(Reichenbach f.) Fawcett & Rendle. Despite five
years of fieldwork in the park, only two plants of
this new taxon have been found, one of them under
cultivation at the Jardfn Botanico Regional C1CY.
Paratype. MEXICO. Yucatan: Municipio Merida.
Parque Nacional de Dzibilchaltiin, aprox. 89°35'49"W,
21°5'44"N, ago. 2002 [“planta epffita, flores moradas, 1 6
2 abiertas simultaneamente, estigma apenas saliendo por
encima del tubo de la eorola, creciendo con Tillandsia
brachycaulos Schltdl. y T. balbisiana Sehult. f.”], /. Ra¬
mirez & G. Carnevali 956 (MO).
Acknowledgments. The authors thank Francisco
Chi-May, Ulises Gonzalez, and Filogonio May Pat
for field assistance and specimen handling. We are
indebted to Ginette Farreny for the line drawing of
the new taxon. Rossana Marrufo drew the dissected
sepals of the new entity.
Literature Cited
Carnevali, G., J. L. Tapia-Munoz. R. Jimdnez-Maehorro,
I,. Sanchez-Saldana, L. Ibarra-Gonzalez. I. M. Ramirez
X M. P. Gomez-Judrez. 2001. Notes on the flora of the
Yucatan Peninsula If: A synopsis of the orchid flora of
the Mexican Yucatan Peninsula and a tentative check¬
list of the Orchidaceae of the Yucatan Peninsula Biotic
Province. Harvard Pap. Bot. 5: 383—466.
Ramirez, I. & G. Carnevali. 1999. A new taxon of Til¬
landsia, some new records, and a checklist of the Bro-
meliaceae from the Yucatan Peninsula. Harvard Pap.
Bot. 4(1): 18.5-194.
-, G. Carnevali & F. Chi-May. 2000. Portraits of
Bromeliaceae from the Mexican Yucatdn Peninsula—11:
A new species of Tillandsia. J. Bromeliad Soc. 50(2):
62-67.
New Status and Combinations for Japanese Taxa of
Ch ionograph is (M ela n I h i aceae)
Noriyuki Tanaka
Department of Education, School of Liberal Arts, Teikyo University, 359 Otsuka, Hachioji,
Tokyo 192-0395, Japan, ntanaka@main.teikyo-u.ac.jp
ABSTRACT. Chionographis japonica var. hisauchi-
ana Okuyama is raised to specific rank as C. his-
auchiana (Okuyama) N. Tanaka, since it differs
markedly from C. japonica (Willdenow) Maximowicz
var .japonica in several characters. Chionographis ja¬
ponica var. minoensis H. Hara and C. japonica var.
kurohimensis Ajima & Satomi are transferred to C.
hisauchiana at subspecific rank as C. hisauchiana
subsp. minoensis (H. Hara) N. Tanaka and C. his¬
auchiana subsp. kurohimensis (Ajima & Satomi) N.
Tanaka, respectively. A key to the four taxa accepted
here is presented.
Key words: Chionographis, Japan. Melanthi-
aceae.
The genus Chionographis Maximowicz is often
placed in the tribe Heloniadeae, together with Cha-
maelirium Willdenow of eastern North America and
a few other genera (Engler, 1888, tribe as Helon-
ieae; Hutchinson, 1934; Tamura, 1.998), under Me-
lanthiaceae (Maximowicz, 1867, as Melanthaeeae;
Tamura, 1998) or Liliaceae (Engler, 1888; Hutch¬
inson, 1934). It is apparently most closely allied to
Chamaelirium, from which it differs mainly by its
zygomorphic flowers (vs. actinomorphic), unequal
tepals (vs. equal), fewer ovules (2 vs. 6—12 ovules
per locule), polycentric chromosomes (vs. monocen¬
tric), and in the sexual attributes (plants usually
hermaphrodite or gynodioeeious vs. dioecious) (Ta¬
naka & Tanaka, 1977. 1979; Tanaka, 1985).
Chionographis is distributed in Japan, Korea,
and southern China (Hara, 1968; Chen, 1980; Ta¬
naka, 1985; Chen & Tamura, 2000). According to
Hara (1968), it comprises four species, of which
two, C. japonica (Figs. I A, 2 A) and C. koidzumiana
Ohwi, are distributed in Japan (the former also in
southern Korea). Only the former species, as tra¬
ditionally circumscribed, is considered here. Hara
11968) recognized three subspecies under C. japon¬
ica, i.e., subspecies japonica, subspecies hisauchi¬
ana (Okuyama) H. Hara (Figs. IB, 2B. under the
name C. hisauchiana subsp. hisauchiana), and sub¬
species minoensis (H. Hara) H. Hara (Fig. 2C, un¬
der the name C. hisauchiana subsp. minoensis).
Novon 13: 212-215. 2003.
Later, a new variety, C. japonica var. kurohimensis
Ajima & Satomi (Fig. 21). under the name C. his¬
auchiana subsp. kurohimensis), was described by
Ajima (1976).
Hara's Chionographis japonica subsp. hisauchiana
(figs. IB, 2B) differs markedly from C. japonica
subsp .japonica (Figs. 1 A. 2A) in several characters
such as floral, vegetative, chromosomal, sexual, and
breeding attributes (Tanaka, 1985, and kev below). It
seems, therefore, more appropriate to treat it at spe¬
cific rank. Haras C. japonica subsp. minoensis (Fig.
2C) and Ajima and Satomi s C. japonica var. kurohi¬
mensis (Fig. 2D) are, respectively, much closer in mor¬
phological, chromosomal, sexual, and breeding char¬
acters to C. hisauchiana than to C. japonica (Tanaka,
1.985, and key below). Chionographis japonica subsp.
minoensis differs from C. hisauchiana mainly in leaf
characters and has a disjunct distribution from the
latter. It is transferred here to C. hisauchiana without
any change in rank. Chionographis japonica var. ku¬
rohimensis differs from C. hisauchiana subsp. hisau¬
chiana and C. hisauchiana subsp. minoensis in chro¬
mosome number and some morphological characters
(Tanaka, 1985, and key below) and has a disjunct
distribution from the latter subspecies. It is therefore
treated here as a third subspecies of C. hisauchiana.
Chionographis hisauchiana as here circum¬
scribed (figs. IB, 2B—2D) is distributed in more
northern regions in Japan (central and northern
Honshu) than C. japonica (Figs. 1A. 2A; kviishu.
Shikoku and west to central Honshu) with a very
minor overlap in their ranges.
A more detailed account of the variation and tax¬
onomy of the genus Chionographis will be forth¬
coming.
Chionographis hisauchiana (Okuyama) N. Ta¬
naka, comb, et stat. nov. Basionym: Chiono¬
graphis japonica var. hisauchiana Okuyama, J.
Jap. Bot. 27: 268. 1952. Chionographis japon¬
ica subsp. hisauchiana (Okuyama) H. Hara. J.
Jap. Bot. 43: 263. 1968. TYPE: Japan. Hon¬
shu: Prov. Musashi [Tokyo-to], Kasumimura, 3
June 1951. T. Satow 7594 (holotype, TNS).
Figures I B, 2B.
Volume 13, Number 2
2003
Tanaka
Japanese Chionographis
213
5mm
A B
Figure 1. Flowers of Chionographis Irom Japan. —A. C. japonica (W'illdenow) Maximowicz from Osaka, Kochi
Pref., Shikoku (A'. Tanaka 1 (THU)). —IF C. hisauchiana (Okuyama) N. Tanaka subsp. hisauchiana from Naguri-
mura, Saitama Pref.. Honshu (A'. Tanaka la (THU))- Both flowers bisexual and at same scale. Drawn from living
plants.
Selected additional specimens. JAPAN. Honshu: Sai¬
tama Pref., Iruma-gun, Naguri-mura, 19 Oct. 1990, A. Ta¬
naka M32B (TEU). 4 June 1990, A'. Tanaka la (THU);
Saitama Pref., Iruma-gun, Mt. Ogami, 25 May 1952, M.
Mizushima s.n. (TI); Tokyo-to. Ome-shi, Osogi-mura, 23
Oct. 1949. T. Satow 7495 (TI).
Chionographis hisaiiehiana subsp. minoensis
(II. Hara) N. Tanaka, comb. nov. Basionym:
Chionographis japonica var. minoensis II.
Hara. .). Jap. Bot. 36: 239. 1961. Chionogra¬
phis japonica subsp. minoensis (H. Hara) H.
Hara. J. Jap. Bot. 43: 263. 1968. TYPE: Ja¬
pan. Honshu: Prov. Mino [Pref. Gifu). Yokok-
ura-mura, N of Kanbara, alt. ca. 300 in. 5 June
1961. //. Hara & S. Kurosawa s.n. (holotype,
TI). Figure 2C.
Selected additional specimens. JAPAN. Honshu: Gifu
Pref., Motosu-gun, Neo-mura, 26 Mar. 1987. A. Tanaka 3
(THU); Gifu Pref., Gujo-gun, Gujo-hachiman, Nabi, 17
Sep. 1967, K. Inami s.n. ( INS-184890); Gifu Pref., Mino,
Natsuzaka, 29 Mav 1968. II. Ilara & S. Kurosawa s.n.
(TI).
Chionographis hisauchiana subsp. kuroliinien-
sis (Ajima & Satomi) N. Tanaka, comb, et slat,
nov. Basionym: Chionographis japonica var.
kurohimensis Ajima & Satomi, J. Geobot. 24:
48. 1976. TYPE: Japan. Honshu: Pref. Niiga¬
ta, Ml. Kurohime, 21 June 1975, T. Ajima
4546 (holotype, KANA-83861). Figure 21).
Selected additional specimens. JAPAN. Honshu: Ni¬
igata Pref., Santo-gun, Koshiji-cho, Hakama, 13 May
1980. A. Tanaka 7 (TEU); Niigata Pref., Murakami-shi.
Kasfiio, 2 June 1963, Y. Ikegami 56443 (TNS-162516);
Niigata Pref.. kariwa-gun. Mt. Hachikoku. 30 May 1952,
.S’. Iwano s.n. (KANA-13338).
Selected specimens of Chionographis japonica. JAPAN.
Kyushu: Nagasaki Pref., Mt. Unzen. June 1934, T. Ma-
kino s.n. (MAK-1.37210); Kagoshima Pref.. koshiki Isl.,
Shimo-koshiki-mura, Nagahama, 31 July 1961. A. Nitta
s.n. (KYO). Shikoku: Kochi Pref.. Osaka. 4 June 1984.
A. Tanaka I (TEU); Kochi Pref., Sakawa, 26 May 1889.
T. Makino s.n. (MAK-137200). Honshu: Hiroshima Pref..
Taishakukyo, I June 1984. A. Tanaka I (TEU); Gifu Pref.,
Motosu-gun, Neo-mura, 25 May 1910, M. Matsui s.n.
(MAK-137171).
Kei to Four Taxa of Chionographis Occurring in J a¬
pan
la. Flowers with lower 2 tepals vestigial, with 1 or
both often absent (Fig. 1A); main veins of leaf
blades obscurely impressed and fine veins tend¬
ing to be slightly elevated adaxially (Fig. 2A);
leaf blades usually opaque adaxially; plants usu¬
ally hermaphrodite but self-incompatible; 2 n =
24. C. japonica
lb. Flowers with lower 2 tepals shorter than upper
ones but always present (Fig. 1B); main veins of
leaf blades (dearly impressed and fine veins not
elevated adaxially; leaf blades lustrous or nearly
opaque adaxially (Fig. 2B—2D); populations usu¬
ally gynodioecious, sometimes hermaphrodite
where hermaphroditic individuals self-eompati-
Iile; 2 n = 42 or 44.
2a. Leaf blades nearly opaque adaxially (Fig.
2B), comparatively thin; upper tepals 2.7—
5.7 mm long (Fig. IB); 2n = 42 .
. C. hisauchiana subsp. hisauchiana
2b. Leaf blades lustrous adaxially (Fig. 2C. 2D),
comparatively thick; upper tepals 3.6—12.0
mm long.
214
Novon
\f I III
2 3 4 - 5 —6 - 7 - 8 — 9 — 10 — 11 — 12 - 12
■■ I p
Figure 2. Leaves »i four taxa of Chiorwgraphis from Honshu. Japan. —A. C. japonica (Willdenow) Maximowicz from
laishakukyo, Hiroshima Pref. (N. Tanaka I (TEU)). —B. C. hisauchiana (Okuvama) N. Tanaka subsp. hisauchiana
from Naguri-mura, Saitama I’ref. (A. Tanaka M32B (TEU)). —C. C. hisauchiana subsp. minoensis (H. Ilara) N. Tanaka
from Neo-mura, Gifu Pref. (A. Tanaka 3 (TEU)). —D. C. hisauchiana subsp. kurohimensis (Ajima & Satomi) N. Tanaka
from llakama, Niigata Pref. ( l\, Tanaka 7 (TEU)). All from living [>lants in cultivation.
3a. Leaf blades usually narrowly elliptic or
narrowly oblong (Fig. 2C); main veins
usually deeply impressed adaxially; up¬
per tepals 3.6-6.9 mm long; '2n = 42
. C. hisauchiana subsp. minoenisis
3b. Leaf blades usually elliptic to oblong
(Fig. 2D); main veins moderately im¬
pressed adaxially; upper tepals 3.7-
12.0 mm long; 2 n = 44.
. C. hisauchiana subsp. kurohimensis
In tlie above key the ranges of variation in te-
pal length of the three subspecies of C. hisau¬
chiana are based on the following samples: sub¬
species hisauchiana, 33 individuals from I
locality; subspecies minoenisis, 35 from 1; sub¬
species kurohimensis, 446 from 13. The ranges
include measurements of both female and her¬
maphrodite plants.
Acknowledgments. I am most grateful to the
keepers of KANA, KYO, MAK. TI, and TNS for
permission to examine their specimens. I also ex¬
press my deep gratitude to Roy E. Gereau (MO).
R. J. F. Henderson (BR1), and Victoria C. Hol¬
lowed (MO) for their critical reviews of the man¬
uscript.
Literature Cited
Ajima, T. 1976. A new variety of Chiorwgraphis japonica
Maxim. J. Geobot. 24: 45=48.
Chen, S. C. 1980. Chionographis. Pp. 13—15 in F. T. Viang
& T. Tang (editors). Flora Reipublicae Popularis Sini-
eae, 14. Science Press, Beijing.
-& M. N. Tamura. 2000. Chionographis. P. 88 in
/.. \. Vfu & P. II. Raven (editors). Flora of China. Vol.
24. Science Press, Beijing, and Missouri Botanical Gar¬
den Press, St. Louis.
Engler, A. 1888. Melanthioideae—Helonieae. Pp. 21-22 in
A. Engler & K. Prantl (editors), Die natiirlichen Pflan-
zenfamilien, 2(5). Wilhelm Engelmann, Leipzig.
Hara, H. 1968. A revision of the genus Chionographis
(Liliaceae). J. Jap. Bot. 43: 257-288, pis. 17, 18.
Hutchinson, J. 1934. The Families of Flowering Plants II.
Monocotyledons. Macmillan, London.
Maximowicz, C. J. 1867. Diagnoses breves plantarum no-
varum Japoniae et Manshuriae 3. Bull. Acad. Imp. Sci.
Saint-Paters bourg.
Tamura, M. N. 1998. Melanthiaceae. Pp. 369—380 in K.
Kubitzki (editor), The Families and Genera of Vascular
Volume 13, Number 2
2003
Tanaka
Japanese Chionographis
215
Plants, 3. Flowering Plants. Monocotyledons: l.ilianae
(except Orchidaceae). Springer-Verlag, Berlin.
Tanaka, N. 1985. Sexual variation in the genus Chiono¬
graphis. Shuseibutsugaku-kenkyu 9: 11—19. [In Japa¬
nese.]
-(initials as N. Y.) & N. Tanaka. 1977. Chromo¬
some studies in Chionographis (Liliaceae) I. On the ho-
lokinetic nature of chromosomes in Chionographis ja-
ponica Maxim. Cytologia 42: 753—763.
-(initials as N. Y.) & -. 1979. Chromosome
studies in Chionographis (I.iliaceae) II. Morphological
characteristics of the somatic chromosomes of four Jap¬
anese members. Cytologia 44: 935—949.
Rubiacearum Americanarum Magna Hama Pars XII. A New
Species of Ixora (Ixoreae) from Panama
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. I .ouis, Missouri 63166, U.S.A.
Charlotte.Taylor@mobot.org
Abstract. The new species Ixora knappiae C. M.
Taylor (Rubiaeeae) from die San Bias region of Pan¬
ama, described and illustrated here, is notable for
its opposite, subsessile. basally rounded to cordate
leaves.
Key words: Ixora, Ixoreae, Mesoamerica, Pan¬
ama, Rubiaeeae.
fhe pantropical genus Ixora L. (Ixoreae; An-
dreason & Bremer, 2000) comprises about 300 to
400 species of shrubs and small trees found in
moist and wet vegetation (I)e Block, 1998). The
majority of Ixora species are found in Asia, and
this genus is also well represented in Africa, Mad¬
agascar, and the Neotropics. Ixora has perhaps 30
Neotropical species, where as noted bv De Block
(1998) it is quite poorly studied. Several Asian spe¬
cies are widely cultivated in tropical regions; a kev
to these was presented by Bridson (1988). Ixora has
never been reviewed as a whole for the Neotropics,
though it has been treated fioristically for some
Neotropical regions, notably by Steyermark (1967,
1974). Ixora was long included in the tribe Pavet-
leae (Robbrecht, 1993; De Block, 1998), but more
recent work (Andreason & Bremer, 2000) has
shown that it is better placed in a separate Ixoreae.
In the Neotropics Ixora is distinguished within
die Rubiaeeae by its stipules that are interpetiolar,
narrowly triangular, generally persistent, and often
also shortly united intrapetiolarly; its calyx limbs
that are four-lobed with the lobes often narrowly
triangular to linear; its salverform corollas with the
tubes generally glabrous internally; its corolla lobes
four, elliptic to ovate, and convolute in aestivation;
its stamens lour and usually inserted in the corolla
throat on short filaments with the anthers partially
exserted; its stigmas two, short, and exserted; its
ovary bilocular with the ovules solitary in each lot—
ule and apically attached; and its fleshy drupaceous
fruits with the pyrenes two, planoconvex, generally
smooth dorsally (i.e., abaxially), and usually rather
thin-walled. The leaves lack domatia, of either fo-
veolate or pilosulous form. The inflorescences are
generally corymbiform and bracteate, and may be
terminal, eauliflorous, or ramiflorous. The fragrant
flowers are homostylous and generally white, yel¬
low, or red. In her excellent revision of Ixora in
Africa, l)e Block (1998) also highlighted the petiole
bases, which are articulated from the stems (at least
in dried specimens) in Ixora, as a good character
for recognition of this genus. This character is
found in Neotropical Ixora species as well. Other
Neotropical Rubiaeeae genera have not yet been
widely surveyed for this feature; articulated petiole
bases are known in a few other paleotropical gen¬
era, of several different tribes. The inflorescence
axes of Ixora are also usually articulated, often giv¬
ing them a distinctive appearance when dry.
The flowers of Ixora exhibit an adaptation for
pollination that is more common in Rubiaeeae of
the Paleotropics than the Neotropics, called sec¬
ondary pollen presentation. The flowers of Ixora
species are protandrous, and the anthers in bud are
held closely around the style and stigmatic arms.
Rather than presenting their pollen directly to pol¬
linators, the anthers release and transfer some or
all of their pollen onto the upper part of the style
and the non-receptive abaxial surfaces of the stig-
matic arms. Then this transferred pollen is exposed
to pollinators for some period of time on these
structures; after this the stigmatic arms separate
and the stigmatic surfaces become receptive.
The inflorescences of Ixora are considered ter¬
minal, most often arising directly from the apices
of the main stems but sometimes borne on short
lateral stems produced in the leaf axils. These lat¬
eral stems are called short shoots bearing terminal
inflorescences by De Block (1998). In some Neo¬
tropical species tbe stem portion of these short
shoots may be reduced so that the inflorescences
appear to be axillary (e.g., /. yavitensis Steyermark).
The inflorescences of Ixora are usually regularly
branched with the higher order axes as well devel¬
oped as the axis from which they branch; this con¬
dition was called “trichotomous” by De Block
(1998). ’I'he inflorescences may vary markedly
among species of Ixora in their degree ol expan¬
sion, from strongly congested, with the axes only
Nov ON 13: 216-219. 2003.
Volume 13, Number 2
2003
Taylor
Ixora from Panama
217
shortly developed and the flowers sessile and sub-
sessile, to relatively very open with the axes and
pedicels all well developed. As in other Rubiaceae,
similar inflorescence arrangements have sometimes
been described in different ways. De Block (1998)
considered the inflorescences to be either pedun¬
culate or sessile; sessile inflorescences comprise
three main units or partial inflorescences, which
are the parts of the inflorescence borne from, re¬
spectively, the two lateral peduncles or basalmost
secondary axes, and the central peduncle or pri¬
mary axis above its junction with that basalmost
pair of secondary axes. This sessile arrangement,
with three axes or peduncles each bearing
branched higher order axes and flowers, was called
“trichotomous” by Steyermark (1974). The distinc¬
tion between sessile and pedunculate inflorescenc¬
es has been considered less significant in studies
of Neotropical Ixora, where inflorescence arrange¬
ment is considered more irregularly variable in
general and several species may have either con¬
dition with the peduncle length varying continu¬
ously (e.g., /. acuminatissima Miiller Argoviensis,
Steyermark, 1974; /. floribunda (A. Richard) Gri-
sebach, Dwyer, 1980).
A number of distinctive characters vary within
the genus. The leaves may be opposite, ternate, or
infrequently verticillate, and frequently have a dis¬
tinctive venation pattern. This pattern has the sec¬
ondary veins relatively weakly developed but the
intersecondary veins numerous and relatively well
marked, so these types of veins are difficult to dis¬
tinguish and the blade appears densely venose. The
presence vs. absence of pubescence on the external
surface of the corolla has been given significant
taxonomic weight (e.g.. Steyermark. 1967) though
it is known to vary within some species of Ixora
(e.g.. De Block. 1998). This corolla pubescence is
often very short, and should be examined with good
magnification. As noted above, several species
show continuous variation in inflorescence arrange¬
ment from sessile to subsessile or pedunculate with
the peduncles well developed.
The species described below was discovered
during study of Rubiaceae collections from Central
America for the Flora Mesoamericana project.
Ixora knuppiae C. M. Taylor, sp. nov. T\ RE: Pan¬
ama. Colon: end of Santa Rita Ridge Road. 21
km from Transisthmian Highway, 400—500 m,
9°25'N, 79°37'W, 22 May 1982. 5. Knapp &
R. Schmalzel 5234 (holotype, MO-5333614).
Figure 1.
Haeo species a congeneris foliis subsessilibus base ro-
tundatis cordulatisve, infloreseentia termiriali pedicellos
usque ad 3 mm longos gerente atque corollae albae extus
glabrae tubo 3.5—4 mm longo ac lobulis ca. 1.5 mm longis
distinguitur.
Shrubs to 3.5 m tall, stems glabrous. Leaves op¬
posite, elliptic to oblanceolate, 10—19 X 4—7.5 cm,
at apex acute to acuminate, at base rounded to
shortly cordate, drying chartaceous, glabrous on
both surfaces; secondary veins 6 to 13 pairs, retic¬
ulating or weakly looping to interconnect near mar¬
gins; petioles 1—2 mm long; stipules persistent, in-
terpetiolar, triangular, 2.5—3 mm long, glabrous on
both surfaces, abaxially costate, aristate with aristas
2— 3 mm long. Inflorescences borne at the apices of
the main stems, pyramidal in outline. 4—5 X 4—5
cm, subsessile, lax, puberulous; bracts subtending
axes triangular, 0.5-1 mm long, acute, those sub¬
tending flowers triangular to lanceolate, ovate, or
ligulate, 0.5—0.8 mm long, acute; pedicels 0—3 mm
long; flowers sessile and shortly pedicellate in 2- to
3- flowered dichasial cymules; hypanthium ellip¬
soid, 1-1.5 mm long, puberulous; calyx limb 0.5—
0.8 mm long, denticulate, puberulous; corolla sal-
verform, white, externally sparsely puberulous,
internally glabrous, tube 3.5-4 mm long, lobes el¬
liptic, ca. 1.5 mm long, obtuse to rounded; anthers
ca. 1.2 mm long, partially exserted; stigmas 2, lan¬
ceolate to spatulate, ca. 0.8 mm long, exserted.
Fruits subglobose, 6—7 mm diam., glabrescent, ma¬
ture color unknown.
Distribution , habitat, and phenology. In wet
forests at 250—500 m in eastern and central Pan¬
ama; collected in flower in May, in fruit in Oc¬
tober.
This new species is distinguished from other
Neotropical species of Ixora by its combination
of opposite leaves that are subsessile and round¬
ed to shortly cordate at the base; its inflores¬
cences borne at the apices of the main stems with
pedicels to 3 mm long; its white corollas that are
sparsely puberulous externally; and its corollas
with tubes 3.5—4 mm long and lobes ca. 1.5 mm
long. A few T other Neotropical species of Ixora
share with I. knappiae subsessile leaves with
rounded to cordate bases; their distinctions from
/. knappiae are outlined below. Ixora acumina¬
tissima of Amazonian South America can often
be distinguished from I. knappiae by its ternate
leaf arrangement, though occasional plants of
this species do have opposite leaves, and I. ac¬
uminatissima can reliably be separated by its
larger corollas, with the tube 6-10 mm long and
the lobes 3-4 mm long. Ixora intropilosa Stey¬
ermark is found on tepuis in Guyana and eastern
218
Novon
Figure l. Ixora knappiae C. M. Taylor. —A. Flowering branch; based on de Nevers & Charnley 5916. —B. Detail of
inflorescence showing eymule with two flowers at anthesis; based on Knapp & Schmalzel 5234.
Venezuela, and can also be distinguished from /.
knappiae by its larger corollas, with the tube 8—
12 mm long and the lobes ca. 3.5 mm long. Ixora
piresii Steyermark of northeastern Brazil and the
Guianas can be distinguished from /. knappiae
by its ternate leaves 29—33 cm long, its pedun¬
cles ca. 5.5 cm long, and its larger corollas with
the tube ca. 16 mm long and the lobes ca. 7 mm
long. Ixora potaroensis Steyermark of Guyana dif¬
fers from /. knappiae in its leaves 32-48 cm long
arranged in verticils of four and its peduncles
12.5—14 cm long. ( Ixora potaroensis was de¬
scribed based on a single fruiting specimen, and
may not be distinct from I. piresii .) And, some
plants of /. killipii Standley, a species of Ama¬
zonian Ecuador, Peru, Colombia, and adjacent
Brazil, also have very short petioles and abruptly
rounded to truncate leaf bases; /. killipii can be
distinguished from /. knappiae by its flowers all
borne on pedicels 2—10 mm long and its larger
corollas, with the tube 25—30 mm long and the
lobes 8—9 mm long.
The specific epithet honors Sandra Knapp, a
prolific taxonomist and collector of the Neotropical
flora.
Paratypes. PANAMA. San Bias: El Llano—Cartf
Road, Kin 16.7, trail W to waterfall 5 km from road,
9°19'N, 78°55'W, de Nevers A- Charnley 5916 (MO): Gam-
pamento Nusagandi, en ELC a 19.1 km de la vfa Pana-
mericana, Sendero Wedar Igar. 9°1 l'N, 78°15'W. H. Her¬
rera et al. 1263 (MO).
Acknowledgments. I thank the curators of F and
PMA, in particular M. Correa, for access to speci¬
mens and providing information; B. E. Gereau for
preparation of the Latin diagnosis; P. De Block for
an invaluable review; T. Wachter, R. Foster, and the
Andrew Mellon Foundation through a grant to the
Field Museum of Natural History for support for
travel to that institution; and R. Magill and 0. M.
Montiel for their significant facilitation of this work.
Literature Cited
Andreason, k. & B. Bremer. 2000. Combined phyloge¬
netic analysis in the Rubiaceae—Ixoroideae: Morpholo¬
gy, nuclear and chloroplast DNA data. Amer. J. Bot. 87:
1731-1748.
Bridson, 1). M. 1988. Ixora. Flora of Tropical East Africa,
Rubiaceae 2: 610-617. Royal Botanic Gardens, Kew.
De Block, P. 1998. The African species of Ixora (Rubi¬
aceae— Pavelteae). Opera Bot. Belg. 9: 1-218.
Dwyer, J. I). 1980. Flora of Panama Part IX. Family 179.
Rubiaceae. Ann. Missouri Bot. Card. 67: 1—522.
Volume 13, Number 2
2003
Taylor
Ixora from Panama
219
Robbrecht, E. 1993. Genera Rubiacearum. Opera Hot.
Relg. 6: 173-196.
Steyermark, J. A. 1967. Ixora. Mem. New York Rot. Card.
17: 341-354.
-. 1974. Ixora. In: T. Lasser (editor), Flora de Ve¬
nezuela 9(2): 888—901. Instituto Botanico, Direccion de
Recursos Naturales Renovables, Ministerio de Agricul-
lura v Cria, Caraeas.
Rubiacearum Americanarum Magna Hama Pars XML A New
Species of Schradera (Schradereae) from Mesoamerica
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
charlotte.taylor@mobot.org
ABSTRACT. The new species Schradera obtusifolia
C. M. Taylor (Rubi aceae) from Costa Rica and Pan¬
ama is described and illustrated. This has previ¬
ously been confused with S. costaricensis Dwyer,
which also has obtuse to rounded leal apices.
Key words: Costa Rica, Panama, Rubiaeeae,
Schradera.
Schradera Vahl (Schradereae; Robbrecht, 1993)
is a tropical genus of succulent, lianescent, epi¬
phytic shrubs that climb by adventitious roots. This
genus is distinguished by its habit, which is un¬
usual in the Rubiaeeae, together with its interpe-
tiolar, relatively large, ligulate to elliptic stipules
that are quickly caducous, its terminal capitate in¬
florescences with the heads usually surrounded by
a well-developed involucre, its truncate, usually tu¬
bular calyx limb, its succulent, salverform, white
corollas with the tubes often rather stout, its corolla
lobes five to six, valvate, triangular in cross section,
and apieally prolonged beyond the contact zone
into a terminal appendage, its bilocular ovaries with
axile placentas bearing numerous ovules, and its
fleshy fruits with numerous seeds embedded in the
pulp, file flowers are distylous and apparently noc¬
turnal with a sweet fragrance similar to that of gar¬
denias. At least some species have markedly dif¬
ferent leaf forms in the juvenile and reproductive
stages (e.g., S. vahlii Steyermark. pers. obs., Puerto
Rico).
The corolla lobes are unusual in form in Schrad¬
era. These corolla lobes have been referred to as
valvate plus, variously, “hooded” (Puff et ah, 1993),
with an appendage (Steyermark, 1974), or witli a
cucullate tooth (Steyermark, 1963). The corolla
lobes of Schradera are typically very fleshy and flat¬
tened to triangular in cross section. They are val¬
vate in bud with the margins of adjacent lobes in
contact along part or most of their length, but at
the apex they are typically prolonged into another
portion that is not in contact with the other lobes.
Enlargements or prolongations of the abaxial or api¬
cal portion of valvate corolla lobes are found in
some other Neotropical Rubiaeeae, in both diurnal
and nocturnal flowers (e.g.. Palicourea cornigera C.
\1. Taylor, Rudgea tayloriae Aymard, Dorr & Coel-
lo, Rsyclwiria cornigera Bentham, Faramea uricin-
ata C. M. Taylor). The function of these structures
is unknown.
Schradera was long treated as a Neotropical ge¬
nus of about 30 species and placed in the tribe
Schradereae together with the genus Lncinaea DC.
of southeastern Asian and the southwestern Pacific
islands. More recently Puff et al. (1993) reviewed
this tribe in detail and concluded that no morpho¬
logical differences separate Schradera from Lnci¬
naea , and therefore these are better combined. The
name Schradera Vahl was published in 1796 and
consequently has priority over Lncinaea DC., pub¬
lished in 1830. ( Schradera Vahl is conserved
against Schradera Heister ex Medicus, published
in 1791.) Puff el al. (1993) formally synonymized
Lncinaea under Schradera, but did not make com¬
binations in Schradera for any of species of Lnci¬
naea. Subsequently Puff et al. (1998) studied some
of the paleotropieal species of Schradera and pub¬
lished new combinations for these, but tin 1 rest of
the paleotropieal species and all of the Neotropical
species are poorly known. In the Neolropies
Schradera is found from Costa Rica to Peru,
through Venezuela to the Guianas and eastern Bra¬
zil. and in the Antilles from Hispaniola though tin*
Lesser Antilles to Trinidad. For the Neotropics
Schradera has only been treated floristically for a
few regions, notably by Steyermark (1963).
I he following new species was discovered during
preparation of the Rubiaeeae for Flora Mesoamer-
icana. Schradera is difficult to study in large part
because of its sporadic collection, as w ith most epi¬
phytes, due to its habit, with the flowers usually
produced in the forest subcanopy and canopy. Once
collected, specimens are usually difficult to intepret
because of their succulent tissues, which often
make poor specimens. Thus the range of variation
generally found within an individual species in
supposedly taxonomically informative characters
such as number of flowers per inflorescence head,
corolla size, or peduncle length is not vet clear.
Nov on 13: 220-222. 2003.
Volume 13, Number 2
2003
Taylor
Schradera obtusifolia from Mesoamerica
221
Figure 1. Schradera obtusifolia C. M. lav lor, based on G. Herrera 3325. —A. Flowering branch. —B. Flower at
anthesis.
Schradera obtusifolia C. M. Taylor, sp. nov.
TYPE: ( iosta Kira. Limon: canton Talamanea.
Alto Uren, siguiendo el camino entre la Ouc-
hrada Chaho y Crorina, 9°21'N. 82°59'W, TOO
m, 23 July 1989, G. Herrera 3325 (holotype,
CR; isotype, MO-3844960). Figure I.
Haee species a Schradera costaricensi Dwyer inflores-
centia 5- ad 10-flora, calycis limbo 5—6 mm longo atque
corollae tubo H— 10 mm longo ac lobulis 10—12 mm longis
distinguitur.
Glabrous shrubs or lianas. Leaves elliptic-oblong,
4— 8.5 X 1.5—3.5 cm, at apex obtuse to rounded, at
base obtuse, drying subcoriaceous; secondary and
higher-order venation not visible; petioles 5—20 mm
long; stipules interpetiolar, caducous, ligulale to el¬
liptic, ca. 15 mm long, obtuse to rounded. Inflores¬
cences terminal; peduncles 1 to 3, 1—4 cm long,
each bearing one head; heads hemispherical to
subglobose, 1.5—2.5 cm diam. (not including corol¬
las), 5- to 10-flowered; involucre continuous, 8—9
mm long, truncate though sometimes splitting along
the margins and appearing irregularly lobed; flowers
with hvpanthium ellipsoid to subglobose or turbi¬
nate, 5—6 mm long, glabrous; calyx limb tubular,
5— 6 mm long, truncate, glabrous; corollas salver-
form, white, ramose, glabrous externally and inter¬
nally, tube 8—10 mm long, lobes 5. linear, 10-12
mm long, at apex with prolongation 3—6 mm long
beyond contact zone, acute, triangular to somewhat
flattened in cross section; anthers ca. 3 mm long,
partially exserted; styles and stigmas not seen.
Fruits not seen.
Distribution , habitat, and phenology. In wet
forests at 250—800 m, Costa Rica and western Pan¬
ama; collected in flower in June, July, and October.
This new species is similar to and apparently
allopatric with Schradera costaricensis Dwyer. Both
of these species are notable for their obtuse to
rounded leaf apices; the species epithet of S. ob¬
tusifolia refers to this character. Dwyer, Burger, and
Taylor (Burger & Taylor, 1993) combined all of the
obtuse- to rounded-leaved plants into a single Costa
Rican species, but with more collections now avail¬
able two species can be distinguished among these
plants. The type of S. obtusifolia was in fact cited
there as a paratype of S. costaricensis. Schradera
costaricensis differs from 5. obtusifolia by its heads
with 20 to 40 flowers, its calyx limbs 3—5 mm long,
and its corollas with the tubes 8—9 mm long and
the lobes 4—5 mm long. Schradera costaricensis is
found in the Cordillera Tilaran of northern Costa
Rica, while S. obtusifolia is found in the Cordillera
Talamanea of southern Costa Rica through western
Panama. These mountain ranges are not continuous
and are floristically distinct (Haber, 2000).
Paratypes. COSTA RICA. Limon: camino de Fila Di-
mat (rasa de Hermogenes Pereira) hasta Soki pasando por
la quebrada, /.. I). Gomez et at. 23353 (F, MO). PANAMA.
Bocas del Toro: F 1.5 mi. off road from divide to Chi-
riquf Grande, D'Arcy 16332 (MO).
Acknowledgments. 1 thank the curators of CR
and F for access to specimens and information; R.
E. Cereau for preparation of the Latin diagnosis: J.
Sanchez (CR) for very helpful comments; E. Rob-
222
Novon
brecht and P. De Block for excellent reviews and
significant help improving the manuscript; T. Wach-
ter, R. Foster, and the Andrew Mellon Foundation
through a grant to the Field Museum of Natural
History for support for travel to that institution; and
R. Magill and O. M. Montiel for their significant
facilitation of this work.
Literature Cited
Burger, W. C. & C. M. Taylor. 1993. Flora Costaricensis:
Family #202. Rubiaceae. Fieldiana, Bot. n.s. 33: 1-
333.
Haber, W. A. 2000. Plants and vegetation. Pp. 39—70 in
N. M. Nadkarni & IN. T. Wheelwright (editors), Montev-
erde, Ecology and Conservation of a Tropical Cloud For¬
est. Oxford Univ. Press, New York.
Puff, C., L. Andersson, U. Rohrhofer & A. Igersheim.
1993. The tribe Schradereae (Rubiaceae) reexamined.
Bot. Jahrb. Syst. 114: 449-479.
-, J. Greimler & R. Buchner. 199B. Revision of
Schradera (Rubiaceae—Schradereae) in Malesia. Blu-
mea 43: 287—335.
Robbrecht, E. 1993. Genera Rubiacearum. Opera Bot.
Belg. 6: 173-196.
Steyermark, J. A. 1963. Schradera. Mem. New York Bot.
Gard. 10: 259-278.
-. 1974. Schradera. hi: T. Lasser (editor). Flora de
Venezuela 9(1): 429—443.
Rubiacearum Americanarum Magna Hama Pars XIV. New Species
of Malanea (Guettardeae) from Western South America
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
charlotte.taylor@mobot.org
Abstract. The new species Malanea campylo-
carpa from the western Amazon basin, M. cylindri-
ca from east-central Peru, and M. ecuadorensis from
east-central Ecuador are described and illustrated
here. These are all notable for their cylindrical
fruits, and all have previously been confused with
M. boliviano Standley. A key to Malanea in western
South America includes eight species.
Key words: Guettardeae, Malanea, Rubiaceae,
South America.
The Neotropical genus Malanea Aublet (Guet¬
tardeae; Robbrecht, 1993) comprises about 35 spe¬
cies of lianas, woody vines, and shrubs found in
moist and wet vegetation from southeastern Nica¬
ragua to eastern Brazil and northern Bolivia. The
genus has not been reviewed as a whole, and in
general has been rather poorly studied. The major¬
ity of its species are found in northeastern South
America, most of them in the Guayana region. Stey-
ermark (1965) reported 25 species, though several
of these were based on only one or two specimens
and distinguished only by the density and distri¬
bution of their pubescence; they deserve reevalu¬
ation when more specimens are available.
Malanea is distinguished by the combination of
its usually climbing habit; its leaves with lineolate
venation; its interpetiolar, generally caducous stip¬
ules; its axillary inflorescences; its corollas with the
tubes relatively short and the lobes valvate in bud;
and its fleshy fruits each with a single hard pyrene.
This distinctive lineolate venation pattern has the
tertiary veins closely parallel, though these veins
and thus this feature are not readily visible in many
species of Malanea. Such lineolate venation is
found in many species of the Guettardeae as well
as several other Rubiaceae tribes (e.g., Hippotis
Ruiz & Pavon, Hippotideae; Sommera Schlechten-
dal. Hippotideae; Joosia H. Karsten, Cinchoneae).
The vining or liana habit of many species of Ma¬
lanea is relatively uncommon among Neotropical
Rubiaceae. However, it is found in a few species
of other genera belonging to both the Guettardeae
(e.g., Chomelia stergiosii Steyermark, C. volubilis
(Standley) Steyermark) and other tribes (e.g., Un-
caria guianensis (Aublet) J. F. Gmelin, Naucleeae;
Randia altiscandens (Ducke) C. \1. Taylor, Garden-
ieae). The delimitation of the Neotropical genera of
Guettardeae is largely traditional and probably de¬
serves reevaluation, in particular the recognition of
Malanea based primarily on its climbing habit.
The flowers of Malanea are generally pale green
to white, yellow, or orange, and thus probably in¬
sect-pollinated. The corollas are usually densely
pubescent in the tube and throat, and often this
pubescence extends onto the corolla lobes. The
species of Malanea that have been studied have
homostylous flowers, although flowers are known for
only a few species so the occurrence of distyly in
this genus cannot yet be excluded. Distyly is pre¬
sent in some species of the related genera Chomelia
Jacquin and Guettarda L. The succulent fleshy
fruits of Malanea are presumed to be dispersed by
animals.
The species described below were discovered
during preparation of the Rubiaceae treatment for
the Flora oj the Venezuelan Guayana. A key is pre¬
sented here to all the Malanea species known from
western South America, in this case comprising Bo¬
livia. Peru, Ecuador, and Colombia.
Kky to the Species of Malanea in Western South
America
I. Inflorescences spiciform, unbranched or with 1 to
2 pairs of secondary axes, with the flowers borne
in sessile glomerules along the primary and sec¬
ondary axes; dried fruits cylindrical, similar in di¬
ameter throughout; pyrenes 1-locular.
2. Plants hirsute throughout, the trichomes long
and spreading; bracts 3—12 mm long ....
. M. ecuadorensis C. M. Taylor
2'. Plants strigose to strigillose throughout, the
trichomes appressed; bracts 1—3.5 mm long.
3. f ruits strongly curved; leaves 9—20 X 4—
10 cm; corolla pale green, with the tube
ca. 2 mm long .
. M. campylocarpa C. M. Taylor
3'. Fruits straight or slightly curved; leaves
4.5—13 X 2—7.5 cm; corolla yellow-or¬
ange, with the tube ca. 4 mm long ....
. M. cylindrica C. M. Taylor
Novon 13: 223-227. 2003.
224
Novon
on Coello 19. —E. Flower bud: based on Ceron .1922. F', iehiiiclia:
Cerro Pugsi, NW slope of Yolcan Pichincha, Bleiweiss
1136 (NY); Tandaeato, Heilborn 496 (S). Tungiirahiiu:
Cordillera de Ulanganates, valley of Rfo Sangarinas (De-
saguadero), Asplund 9792 (S).
I. 25. Notopleura zarucchiana C. M. Taylor,
sp. nov. TY PE: Colombia. Antioquia: Son son,
Los Cristos, sitios “Paramo de Sonson" y
“Monumento," sobre la cuchilla “Comadrejal,”
10 km SE de Sonson en la via al Narino,
05°38'N, 75°15'W, 2500-2740 m, 17 Aug.
1992, R. Callejas & F. J. Roldan 10511 (ho-
lotype, HUA-83070). Figure 2E-H.
Haee species a congeneris foliorum venis secundariis
cum marginibus incrassatis conjunctis ac venatione ter-
tiaria abaxialiter manifesta, inflorescentia cymosa eonges-
ta, calycis limbo 2—3.5 mm longo atque corollae extus
hirtellae tubo 10—12 mm longo distinguitur.
Succulent, terrestrial herbs or soft shrubs to 0.8
m tall, unbranched or apparently little-branched;
stems densely hirtellous. Leaves lanceolate to ovate,
3.5—7 X 1 .5-4 cm, at apex acute, al base cuneate
to obtuse or subtruncate, drying papyraceous to
chartaeeous, adaxially densely pilosulous to subfloc-
cose, abaxially paler and glabrous except strigillose
to hirtellous on principal veins; secondary veins 6 to
10 pairs, extending to unite with margins, adaxially
venation plane, abaxially costa prominent, secondary
veins prominulous, and reticulated tertiary venation
visible and often thickened; margins markedly thick¬
ened, narrowly winged; petioles 3-22 mm long,
densely hirtellous; stipules glabrous, interpetiolar or
shortly united around stem, broadly triangular, 2—3
mm long, succulent, persistent, at apex with 2—5 lin¬
ear glands 0.3—1 mm long, caducous. Inflorescences
pseudoaxillary, congested-cymose, hirtellous, as¬
cending; peduncles 1.5-4 cm long; branched portion
rounded-corymbiform to hemispherical, 1—1.5 X
0.8—2.5 cm (without corollas), with I pair of short
secondary axes; bracts narrowly triangular, 6—15 mm
long, acute, glabrous; pedicels 0-1 mm long; flowers
subsessile to shortly pedicellate in glomerules or
congested cymules; hypanthium turbinate, ca. 1 mm
long, glabrous; calyx limb glabrous, 2—3.5 mm long,
divided for 2/3—3/4, lobes 4 to 5. narrowly triangular,
acute; corolla infundibuliform, white, externally
Volume 13, Number 2
2003
Taylor 255
Notopleura from Central and South America
sparsely to moderately hirtellous, internally glabrous,
tube 10-12 mm long, lobes 4 or 5, 3.5—5 mm long,
ligulate, dorsallv smooth; anthers ea. 1.2 mm long,
situated just above middle of corolla tube; stigmas
ca. 0.8 mm long, situated in corolla throat. Infruc-
tescences and fruits not seen.
Habitat, distribution, and phenology . In wet
premontane and montane forests at 1700 to 3300
m in the Cordilleras Occidental, Central, and Ori¬
ental of Colombia; collected in flower February
through June and August.
Notopleura zarucchiana is distinguished by the
combination of its leaves with the secondary veins
uniting with the thickened margins and the tertiary 1
venation visible abaxially, its congested-cymose in¬
florescences, its calyx limbs 2—3.5 mm long, its ex¬
ternally hirtellous corollas, and its corolla tubes
10—12 mm long. The species epithet honors James
L. Zarucchi, an American botanist who among other
accomplishments has documented the Colombian
flora with excellent specimens. Several other mon¬
tane Colombian species of Notopleura also have
thickened leaf margins, notably N. macropodantha,
but only N. longiflora (described above) also has
similarly long corollas. Notopleura longiflora differs
from N. zarucchiana in its calyx limbs 5—7.5 mm
long and corollas externally glabrous to sparsely
puberulous with tubes 12.5—14 mm long; N. ma¬
cropodantha differs from N. zarucchiana in its
adaxially glabrous leaves with the secondary veins
not extending all the way to the margins, its stipules
with two well-developed lobes, its corolla tubes 3-
4 mm long, and its corolla lobes ca. 1.5 mm long.
Notopleura zarucchiana is included in the “Noto-
pleura macropodantha group” discussed in the in¬
troductory part of this article.
Paratypes. COLOMBIA. Antioquia: Sonson. San
Pablo, via Sonson-Narino. 13.3 km SE de Sonson. Callejas
et al. 6311 (HUA); along old camino "Paramo de Sonson”
along or near carretera between Sonson and Narino, Core
741 (US); 10 km al K de Sonson, Scolnik et id. l9An237
(US). Cundinamarca: subida al Alto de Tibre, F. (). /u-
loaga 4045 (COL). Kisaralda: Santuario, Las Colonias,
200 m abajo del “Reposo,” Maeizo de Tainana,./. II. Tor¬
res et al. 1574 (COL p.p., plant “B”).
II. Notopleura subg. Viscagoga (Baillon) C. M.
Taylor, Ann. Missouri Rot. Card. 88: 511.
2001. Uragoga sect. Viscagoga Baillon, Ad-
ansonia 12: 227. 1879. Psychotria ser. Visca¬
goga (Baillon) Steyermark. Mem. New York
Bot. Card. 23: 500. 1972. TYPE: Notopleura
guadalupensis (DC.) C. M. Taylor.
This subgenus comprises 15 species (including
the 5 described in this article), which are recog¬
nizable by their epiphytic, usually branched habit,
terminal and sometimes also pseudoaxillary inflo¬
rescences, and pyrenes 2 to 0. Plants of subgenus
Viscagoga have stipules with a truncate sheath and
one to several short glandular interpetiolar append¬
ages, or in a few species these are apparently lack¬
ing. Species of subgenus Viscagoga are found gen¬
erally throughout the range of Notopleura, which is
expanded here with the discovery of N. bahiensis
C. M. Taylor.
II. 1. Notopleura bahiensis C. M. Taylor, sp.
nov. TYPE: Brazil. Bahia: Uruyuca, Parque
Estadual Serra do Conduru, trail to summit of
Cerro do Conduru, Southern Bahian Wet For¬
est. I4°29'S, 39°06'W, 210 m. 28 Sep. 2001,
W. W. Thomas, J. Jardim, S. Sant'Ana & F.
Junchum 12166 (holotype, CEPEC; isotypes,
MO-5568186, NY).
Haec species a Notopleura multiramosa fructus pyrenis
4 ad 6 atque distributione disjuncta orientali-brasiliensi
distinguitur.
Succulent suffrutescent herbs, epiphytic, to 2 m
tall; stems glabrous. Leaves elliptic to lance-elliptic
or oblanceolate, 3—9.5 X 1-3.5 cm, at apex acute,
at base acute to obtuse, glabrous, when dry sub-
coriaceous; secondary and higher-order venation
not visible; petioles 3—7 mm long; stipules glabrous,
united around stem into a truncate tubular sheath
1.5—2.5 mm long, near base of interpetiolar portion
with a conical appendage 0.5—1 mm long, at apex
this glandular, sheath generally persistent, append¬
age generally persistent but becoming dry and in¬
durated. Inflorescences terminal, glabrous or with
higher-order axes sometimes puberulous, cymose,
color not noted; peduncles 0.8—1.2 cm long;
branched portion 1.5—4 X 2-6 cm, rounded-cor-
ymbiform, with secondary axes 1 to 2 pairs; bracts
deltoid, 0.3—1 mm long; pedicels 0.5—2 mm long;
flowers pedicellate in dichasial cymules of 3(5): hy-
panthium ellipsoid, ca. 1 mm long, densely puber¬
ulous; calyx limb densely puberulous, 0.3—0.5 mm
long, truncate; corolla in bud externally puberu¬
lous. mature corolla, anthers, and stigmas not seen.
Fruits ellipsoid to subglobose, ca. 4X4 mm, gla-
brescent, orange; pyrenes 4, dorsallv smooth.
Habitat, distribution, and phenology. In wet
forests in Bahia, eastern Brazil, at 500 to 760 m;
collected in flower bud in June, in fruit in March,
May, and September.
Notopleura bahiensis is apparently disjunct in its
distribution from other Notopleura species; the spe¬
cies epithet refers to its geographic distribution.
The species of Notopleura with the geographical
256
Novon
Figure 7. New species of Notopleura suhg. Viscagoga. A, B, A 'otopleura elegant C. M. Taylor. —A. Stem witli leaves
and inflorescence; based on Hoyle 928 (MO). —B. Flower; based on Dmt 2056 (MO). —C. Notopleura vargasiana C.
M. Taylor, flower; based on Werner 275 (MO). I)— F, Notopleura cocleensis C. M. Taylor. —D. Flower; based on Knapp
et al. 5999 (MO). —E. Stem node (same node as in F) with stipule and basal portions of petioles, interpetiolar view;
based on Croat 27446 (MO). —F. Same stem node as in E, intrapetiolar view with petiole removed. G, H, Notopleura
episcandens C. M. Taylor & 1). H. Lorence; based on Knapp el al. 4506 (MO). —G. Stem node with basal portions of
petioles. —H. Corolla. C, D, E, F, G, H to same scale.
range closest to this new species is N. multiramosa
of the Guianas, which is similar to N. bahiensis in
general aspect and in having truncate calyx limbs.
In addition to their geographical separation, N.
multiramosa differs morphologically from N. bah¬
iensis in its fruits with consistently two pyrenes.
There may be differences in floral morphology as
well, but the flowers of both species are poorly
known. The fruits are described on a few collec¬
tions as orange, but it is possible that these later
become red then black at maturity as in several
other epiphytic species of Notopleura.
Paratypes. BRAZIL. Bahia; Sao Jose da Vitoria, ca.
9 km na estrada S. Jose-lna. ramal a dereita ca. 3 km
adentro, A. M. tie Carvalho et al. 6499 (NY); Rod. Rio
Branco a Una, H. .S'. Pinheiro 1249 (US); estrada Sao Jose,
Una, Km 8, ramal a dereita a partir de Sao Jose, A. M.
Amorim et al. 1689 (MO); Una, riberao das Palmeiras,
Serra Javi. ramal com entrada no km I I da rodovia Sao
Jos£/Una, lado S a 9 km ate cima da Sena, 9 km por ar
SE Sao Jose, /,. A. Mottos da Silva & Judziewicz 4154
(US); Uruyuca, 7.4 km N of Serra Grande on road to Ita-
eare, Fazenda Lagoa do Conjunto Fazenda Santa Cruz, W.
W. Thomas et al. 10907 (MO. NY).
II. 2. Notopleura cocleensis C. M. Taylor, sp.
nov. TYPE: Panama. Code: slopes and summit
of Cerro Gaital, N of El Valle, 8°40 , N,
80°07'W. 1000-1400 m, 10 July 1982, S.
Knapp. J. Mallet & H. Dressier 5999 (holotype,
MO-3032258). Figure 7D-F.
Ilaee species a eongeneris foliorum elliptieorum 1.2—
4.5 X 0.5—2 cm venis secundariis non manifestis, pedun-
culo 0.5—1.2 cm longo, stipularum appendieibus glandu-
laribus pluribus 0.5—1 mm longis, calycis limbo 1.5—3 mm
longo, corollae tubo ca. 3 mm longo ac lobulis ca. 1.5 mm
longo atque fruclu nigro distinguitur.
Epiphytic, succulent suffrutescent herbs or
shrubs to 0.3 m tall; stems glabrous. Leaves el¬
liptic to elliptic-oblong, 1.2—4.5 X 0.5—2 cm, at
apex acute and shortly apiculate, at base acute
to cuneate, glabrous, when dry coriaceous, slight¬
ly to markedly discolorous; secondary veins not
or almost not evident; petioles 2—4 mm long,
winged for most or all of their length; stipules
glabrous, united around stem into a tubular trun¬
cate sheath 0.8—1.5 mm long, interpetiolar por¬
tion near base with a medial cartilaginous rib ter¬
minating at middle of sheath in a group of
Volume 13, Number 2
2003
Taylor 257
Notopleura from Central and South America
caducous, glandular appendages 0.5—1 mm long.
Inflorescences terminal and pseudoaxillary, gla¬
brous, paniculate; peduncles 0.5-1.2 cm long;
branched portion 1 X 1.5—2.5 cm, corymbiform,
with secondary axes 1(2) pair, bracts 1.5—7 mm
long, linear, acute, those subtending flowers 1.5—
3 mm long; pedicels 1.5—3 mm long; flowers ped¬
icellate in cymules of 3 to 5; hypanthium ellip¬
soid, ca. 0.8 mm long, glabrous; calyx limb 1.5—
3.5 mm long, glabrous, red, deeply lobed, lobes
4, ligulate, acute to acuminate; corolla funnel-
form. white to pink, externally glabrous, inter¬
nally glabrous except hirtellous in a ring situated
near middle of tube, tube ca. 3 mm long, lobes
4, triangular, ca. 1.5 mm long, dorsally smooth;
anthers ca. 1 mm long, partially exserted; stigmas
ca. 1 mm long, exserted. Fruits subglobose, ca.
4.5 mm diam., glabrous, red then black; pyrenes
4, dorsally smooth or usually with a weak central
ridge.
Habitat, distribution, and phenology. Western
Panama, in wet forest and cloud forest at 800 to
1400 m. collected in flower in March. July, and
December, in fruit in March and July.
Notopleura cocleensis is known only from Code
Province, Panama, and the specific epithet refers
to this distribution. It is similar to N. maxonii,
N. elegans (described below), N. guadalupensis,
and N. episcandens (described below). Notopleura
maxonii is sympatric with N. cocleensis, and can
be distinguished from it by its calyx limb 0.8—1
mm long and its leaves 0.7-2.5 X 0.2-0. 8 cm;
N. elegans is also sympatric with N. cocleensis,
and can be distinguished from it by its calyx
limbs 0.8—1.8 mm long and corolla tubes 6—10
mm long; N. episcandens from eastern Panama
can be distinguished from N. cocleensis by its
ovate to elliptic-lanceolate leaves and solitary
triangular interpetiolar appendages; and N.
guadalupensis is sympatric with N. cocleensis and
can be distinguished from it by its ovate to
lanceolate leaves and calyx limbs 0.5—0.8 mm
long.
Paratypes. PANAMA. Code: La Mesa, above El Val¬
le de Anton, ca. 2 km W of Cerro Pilon, Croat 37446
(MO); 3-mountain ridge above El Valle, C. Hamilton et al.
4148 (MO); foothills and summit of Cerro Caracol. near
La Mesa N of El Valle de Anton, Knapp 1114 (MO); Cerro
Caracol, Kirkbride 1108A (MO p.p., plant labeled B); di¬
vide SW of La Mesa at end of logging road, Stein & Ham¬
ilton 11)09 (MO); ridge from summit of Cerro Carocoral
| sic; Caracol] to summit of Cerro Gaital, N rim of El Valle,
Sytsma 3826 (MO).
II. 3. Notopleura elegans C. M. Taylor, sp. nov.
TYPE: Costa Rica. Heredia: Parque Nacional
Braulio Carillo, sendero de transecto, on flat
portion of ridge during descent from Cerro las
Martas to old road from San Rafael, Vara Blan¬
ca to Refugio at 2070 m. I0°10'07"N,
84°06'48"W, 2210-2240 m, 5 June 1992, B.
Boyle 928 (holotype, MO-3294927; isotypes,
CR, INB). Kigure 7A, B.
Haee species a Notopleura guadalupensi limbo calycino
0.8—1.8 mm longo atque tubo eorollino 6—10 mm longo
distinguitur.
Epiphytic, succulent shrubs and suffrutescent
herbs to 1 m tall; stems glabrous. Leaves elliptic,
1.2-5.2 X 0.7—2.8 cm, at apex acute to somewhat
acuminate, at base acute to cuneate, glabrous,
when dry chartaceous to subcoriaceous, slightly to
markedly discolorous; secondary veins not evident;
petioles 2—8 mm long; stipules glabrous, united
around the stem into a truncate, tubular sheath 1-
1.5 mm long, interpetiolar portion with 1 to 2 tri¬
angular, caducous, glandular appendices 0.5—1 mm
long. Inflorescences terminal and pseudoaxillary,
glabrous, paniculate, pink to red; peduncles (1.5)2—
4.5 cm long; branched portion 2-4 X 2-5 cm, cor¬
ymbiform to broadly triangular, with secondary axes
1 to 3 pairs; bracts linear, 1—6 mm, acute, those
subtending flowers 1—3 mm long; pedicels 1—5 mm
long; flowers pedicellate in cymules of 2 to 3; hy¬
panthium ellipsoid, ca. I mm long, glabrous; calyx
limb 0.8—1.8 mm long, glabrous, lobed for 1/2-
deeply, lobes 4, triangular, acute; corolla slenderly
funnel form, white sometimes tinged with blue, ex¬
ternally and internally glabrous, tube 6—10 mm
long, lobes 4, 1.5—2.5 mm long, triangular, dorsally
smooth; anthers ca. 1.2 mm long, partially exserted;
stigmas ca. 0.2 mm long, exserted. Fruits subglo¬
bose, ca. 4 mm diam., glabrous, orange to red; py¬
renes 4, dorsally smooth.
Habitat, distribution , and phenology. Northern
Costa Rica and western Panama, in wet forest, oak
forest, and cloud forest at 900 to 2482 m; collected
in flower in January, March, and May through Oc¬
tober, in fruit in January, March, April, June, July,
and September through November.
Notopleura elegans is similar to N. guadalupen¬
sis, N. cocleensis, N. pithecobia (Standley) C. M.
Taylor, and N. vargasiana (described below). No¬
topleura guadalupensis and N. pithecobia differ
from N. elegans in their shorter calyx limbs, 0.5—1
mm long, and shorter corolla tubes, ca. 3 mm long;
N. cocleensis differs from N. elegans in its calyx
limbs 1.5—3 mm long and its corolla tubes ca. 3
mm long. The distinctions between N. elegans and
258
Novon
N. vargasiana are discussed under tins latter spe¬
cies, below. The specific epithet of this new species
refers to the graceful inflorescences with relatively
well-developed pedicels and corollas.
Paratypes. COSTA RICA. Alajuela: near Quebrada
Desengario. 1 km K of Vara Blanca, Lent 2056 (MO). Car-
tago: near and along crest of ridge S of Alto Patillos (and
a continuation of the latter), ca. 6 km SE of Tapantf, Gra-
yum & Herrera 7717 (MO). Guanacaste: Estaeion Cacao,
Parque Nacional Guanacaste, R. Espinoza 77 (MO): trail
to Cerro Cacao from Estaeion Bioldgica Cacao, Short et
al. 85 (BM). Heredia: N end of Cerros has Marias, N
slope of Volcan Barva, Grayum 7282 (MO); headwaters of
Rio Santo Domingo, ca. 3 km E of San Rafael de Vara
Blanca, N slope of Volcan Barba, Grayum 7536 (BM). Ei-
indn: Talamanca, Quebrada Kuisa, trail up of camp [sic],
ca. 500 m, Cordillera de Talamanca, Bittner 1801 (MO).
Puntarenas: Monteverde Reserve, Nuboso Trail. Haber
510 (MO), Haber ex llello 2232 (MO); Monteverde Re¬
serve, Haber 533 (MO); Monteverde Reserve, Sendero San
Euis. Haber & Bello 2495 (MO); Monteverde Cloud Forest
Reserve, road to TV towers. Pacific slope, Haber ex Bello
& Lierheimer 4466 (MO); Monteverde, upper valley and
ridges of Veracruz river valley S of Monteverde Reserve,
Pacific drainage, Haber ex Bello & Lierheimer 4525 (MO);
Cerro Ojo de Agua, Monteverde, Haber ine also
resulted in few infections on western hemlock (3%),
but all of the infections produced aerial shoots
(Wass, unpublished). Inoculations on shore pine
produced many more viable infections (42%) and
all of the infections produced shoots. In these ex¬
periments, inoculations with seeds from the dwarf
mistletoe on western hemlock onto shore pine pro¬
duced no infections, while 29% of seeds inoculated
on western hemlock produced infections and 93%
of these produced aerial shoots (Wass, unpub¬
lished).
Hawksworth (1987) summarized the taxonomy of
hemlock dwarf mistletoe and separated this species
into three different races: a western hemlock race,
a shore pine race, and a mountain hemlock race.
Hawksworth et al. (1992) presented another inter¬
pretation for the classification of hemlock dwarf
mistletoe. They described the mountain hemlock
race proposed by Hawksworth (1987) as a subspe¬
cies ol hemlock dwarf mistletoe (mountain hemlock
dwarf mistletoe— Arceuthobium tsugense (Rosen-
< I a h I) G. N. Jones subsp. mertensianae Hawksworth
& Niekrent), but maintained the dwarf mistletoe
Novon 13: 268-276. 2003.
Volume 13, Number 2
2003
Wass & Mathiasen
Arceuthobium tsugense subsp. contortae
269
parasitizing shore pine as a race of hemlock dwarf
mistletoe. Nickrent and Stell (1990) reported that
their analysis of isozymes could not he used to dis¬
tinguish the dwarf mistletoe parasitizing shore pine
from the mistletoe parasitizing western hemlock
and Nickrent et al. (1994) reported that nuclear
ribosomal DNA internal transcribed spacer se¬
quences were not useful in distinguishing between
the subspecies or races of hemlock dwarf mistletoe.
Therefore, in a revision of their monograph on 4r-
ceuthobium, Hawksworth and Wiens (1996) main¬
tained the race designation for the dwarf mistletoe
on shore pine.
In 1997, we began intensive studies of the mor¬
phology and host range of hemlock dwarf mistletoe
throughout its geographic range. Based on the re¬
sults of our work, we provide data that support the
classification of the dwarf mistletoe parasitizing
shore pine in British Columbia and Washington as
a subspecies of hemlock dwarf mistletoe, thereby
eliminating the use of the race concept in Arceu¬
thobium.
Arceuthobium tsugense (Rosendahl) C. N. Jones
subsp. contortae Wass & Mathiasen. subsp.
nov. TYPE: Canada. British Columbia: Van¬
couver Island, Mount Wells, elevation ca. 335
m, 8 Aug. 2001, E. E Wfm 26096 (holotype,
DAVFP; isotypes, ASC, MO, US).
Plantae 3—11 (6) cm altae; antliesis mense Julio— ()<■-
tobri; fructus maturitas Augusto—Novemhri. In I bn us con¬
torta var. contorta parasiticae.
Plants 3-1 I cm in height (mean ca. 6 cm); basal
diameter of dominant plants 2—5 mm (mean 3 mm);
third internode length 5-16 mm (mean 9.95 mm)
and 1.75 mm wide; staminate and pistillate plants
primarily green-brown, but some yellow-green; sta¬
minate flowers 3- or 4-partite; mature fruit length
3.3—5.7 mm (mean 4.6 mm) and 2.1-4.2 mm wide
(mean 3.1 mm). Seed length 1.8—3.0 mm (mean 2.5
mm) and width 1.0—1.7 mm (mean 1.4 mm).
Phenology. Antliesis from mid July through
early October with peaks in late July to mid August;
seed dispersal from mid September to early Novem¬
ber with peaks in mid September to mid October.
Habit. Parasitic principally on Pinus contorta
var. contorta, but occasionally parasitic on Tsuga
heterophylla and rarely on Pinus monticola Douglas
ex D. Don.
Distribution. Shore pine dwarf mistletoe
(SPDM) occurs on Vancouver Island and other is¬
lands off die coast of British Columbia as far north
as Maurelle Island and on the mainland coast of
British Columbia from south of Sechelt to near
Powell River (Fig. I). There are also reports of
SPDM near Port Clements on the Queen Charlotte
Islands, on Malcolm Island, and near Terrace, Brit¬
ish Columbia (Smith & Wass. 1976; Wass, 1976;
Hawksworth. 1987). These represent possible pop¬
ulations of SPDM that are approximately 400—500
km north of the northernmost populations on Van¬
couver Island or the main coast of British Colum¬
bia. Examination of the dwarf mistletoe collection
records for the Port Clements and Malcolm Island
sites and a visit to the Terrace site indicated that
these sites are actually rare crossovers of western
hemlock dwarf mistletoe onto shore pine. There¬
fore, the northern range limit of shore pine dwarf
mistletoe is on Maurelle Island (Fig. 1). Elevational
range is from near sea level to 800 m.
Morphological Measijkkments
Ten SPDM populations scattered throughout its
principal range in British Columbia and on Orcas
Island were sampled (Fig. 1). Five western hemlock
dwarf mistletoe (WHDM) populations from within
the geographic range of SPDM and 14 other
W HDM populations were sampled for comparison
(Figs. 1. 2). From each population, 20 infections
(10 male and 10 female) were collected and the
dominant shoot from each infection was used for
morphological measurements. The infections were
collected, placed in a cooler to keep them fresh
during transport, and stored in a cold room until
measured. Plant measurements were made within
two days of collection. The dwarf mistletoe plants
were measured using a dissecting microscope with
a micrometer.
The dwarf mistletoe plant characters measured
were those used by Hawksworth and Wiens (1996)
for taxonomic classification. The following morpho¬
logical characters were recorded: (1) height, basal
diameter, third internode length and width, and col¬
or of the tallest male and female shoot from each
infection collected; (2) mature fruit length, width,
and color; (3) seed length, width, and color; (4) sta¬
minate flower diameter; (5) number, length, and
width of staminate perianth lobes; (6) anther dis¬
tance from the perianth lobe tip; (7) anther diam¬
eter; and (8) pre-flowering lateral staminate spike
length. Analysis of variance (ANOVA). using a gen¬
eral linear model procedure for unbalanced de¬
signs, was used to determine significant differences
between shore pine dwarf mistletoe and western
hemlock dwarf mistletoe morphological measure¬
ments (F < 0.05). All statistical analyses were per¬
formed with SAS computer programs (SAS Institute
Inc., 1985).
270
Novon
Volume 13, Number 2
2003
Wass & Mathiasen
Arceuthobium tsugense subsp. contortae
271
Male plants of shore pine dwarf mistletoe were
significantly shorter (mean 5.6 cm) than those of
western hemlock dwarf mistletoe (mean 7.8 cm).
Although the range in male plant height of the two
mistletoes overlaps somewhat, the tallest plants of
SPDM do not reach the maximum heights of
WHDM (Table I). Shoot third internode length was
significantly shorter and third internode width was
significantly wider for SPDM than for W HDM ( fa¬
ble 1). Staminate flower diameter, perianth length,
and perianth width were significantly larger for
SPDM than for W HDM. Anther distance from peri¬
anth tip was significantly greater for SPDM than for
WHDM (Table l). Another difference between
SPDM and WHDM is plant color. Male plants of
SPDM are predominantly green-brown whereas
male plants of WHDM are predominantly green-
yellow. For both mistletoes, the number of perianth
lobes on male flowers is three or four, but the pre¬
dominance of 3-partite or 4-partite flowers varies.
Female plants of shore pine dwarf mistletoe were
significantly shorter (mean 6.6 cm) than those of
western hemlock dwarf mistletoe (mean 8.0 cm)
(Table 2). Again, the range in heights of the dom¬
inant female plants overlaps for these mistletoes,
but the height of SPDM female plants does not
reach the size of those of WHDM (Table 2). Shoot
third internode length was significantly shorter for
SPDM than for WHDM. The fruit width and seed
width of SPDM were significantly wider than for
WHDM. The color of female plants of SPDM is
predominantly green-brown. Female plants of
WHDM are green-brown, green-yellow, or purplish.
Fruit and seed color is similar for both mistletoes,
as is seed length (Table 2). Table ,3 summarizes the
principal morphological differences between SPDM
and WHDM.
Host Susceptibility Based on Nati ral
Infection
In order to determine the susceptibility of shore
pine and western hemlock to shore pine dwarf mis¬
tletoe and western hemlock dwarf mistletoe based
on natural infection, 173 temporary plots 6 meters
in radius (0.04 ha) were established around large,
severely infected residual trees (western hemlock
or shore pine) at 10 locations (Fig. 1). In each plot,
trees greater than 1.37 meters in height were sam¬
pled and the species and dwarf mistletoe rating
(DMR, Hawksworth, 1977) was recorded for each
live tree in a plot.
Ninety-nine percent of the shore pine sampled
(765 trees) in the plots located in shore pine forests
were infected by shore pine dwarf mistletoe (Table
4), and these trees had a mean DMR of 5.0 on a
scale of 0—6. This high level of infection (dearly
demonstrates the high level of susceptibility of
shore pine to this mistletoe.
In forests composed of mixtures of shore pine
and western hemlock infested with shore pine dwarf
mistletoe, 95% of the shore pine was infected, and
these trees had a mean DMR of 3.7. Of the 802
western hemlocks sampled in these forests, only
21% were infected by shore pine dwarf mistletoe
(Table 4). These western hemlocks only had a mean
DMR of 0.3. These infection levels indicate that
shore pine is highly susceptible to infection by
SPDM while western hemlock is (dearly much less
susceptible.
In forests of mixed shore pine and western hem¬
lock infested with western hemlock dwarf mistletoe,
96% of the 138 western hemlock sampled were in¬
fected (Table 4). These trees were severely infected
because their average DMR was 4.3. In contrast,
WHDM infection of the shore pine sampled (77
trees) was only 1% with an average DMR of only
0.1. These infection incidences demonstrate the
high susceptibility of western hemlock to W HDM
(a principal host) and the very low susceptibility of
shore pine (a rare host) (Table 4).
Shore Pine Dwarf Mistletoe
Plants of shore pine dwarf mistletoe and western
hemlock dwarf mistletoe have morphological simi¬
larities, but they also have some consistent mor¬
phological differences (Table 3). Male and female
plants of SPDM are consistently smaller than
WHDM. In addition, the color of male SPDM plants
is frequently green-brown and occasionally yellow-
green, while the color ol W HDM male plants is
consistently yellow-green. Staminate flowers of
SPDM are consistently larger than the flowers of
Figure 1. Dashed line encloses the distribution of Arceuthobium tsugense subsp. contortae. Black circles indicate
known populations and numbered circles indicate populations sampled for morphological measurements and host sus¬
ceptibility: Washington: 1 —Orcas Island; British Columbia: 2 —Mount Wells, 3—Mount Helmcken, 4—Bluff Mountain,
5—Mount Work, 6—Trap Mountain, 7—Spider Lake, 8—Cortes Island, 9—Texada Island, 10—Sechelt. Black squares
indicate populations of Arceuthobium tsugense subsp. tsugense sampled for morphological measurements and host sus¬
ceptibility: British Columbia: 11 —Spider Lake, 12 —Texada Island, 13 —Bowser, 14 —Holt Creek, 15 —Caycuse Sum¬
mit.
272
Novon
Figure 2. Distribution of Arceuthobium tsugen.se subsp. tsugense in Washington, Oregon, and California (Hawksworth
& Wiens, 1996). Black circles indicate known populations and numbered circles indicate populations where mor-
Volume 13, Number 2
2003
Wass & Mathiasen
Arceuthobium tsugense subsp. contortae
273
Table 1. Morphological characteristics of male plants for shore pine and western hemlock dwarf mistletoes for
collections from British Columbia and the United States.
Shore pine dwarf mistletoe
Western hemlock dwarf mistletoe
Probability
Male plant characters*
Mean
Range
N/n**
Mean
Range
N/n
>F
Tallest shoot length
5.6
3.2-10.8
11/110
7.8
3.4-16.1
20/270
0.0001
Shoot basal diameter
2.8
1.7-4.7
1 1/110
2.6
1.3-5.0
20/269
0.2104
Shoot third internode
length
9.2
5.8-15.7
11/110
1 1.8
4.5-23.0
20/270
0.0001
Shoot third internode
width
1.8
1.1-2.5
11/110
1.6
0.8-3.0
20/270
0.0111
Third internode length/
width ratio
5.3
2.9-8.8
1 1/110
7.5
2.7-15.0
20/270
0.0001
Staminate spike length
12.6
5.0-22.0
1 1/110
10.8
5.0-27.4
16/260
0.1590
Staminate flower diame-
ter
4.3
2.8-5.0
1 1/110
3.6
2.2-0.4
16/260
0.0056
Anther diameter
0.7
0.4-1.2
11/110
0.7
0.3-1.3
16/260
0.7650
Staminate perianth
length
1.8
1.2-2.8
11/110
1.5
1.0-2.2
16/260
0.0015
Staminate perianth
width
1.4
1.0-2.3
1 1/110
1.2
0.8-2.2
16/260
0.0248
Anther distance from
perianth tip
0.9
0.4-1.5
11/110
0.5
0.2-1.2
16/260
0.0002
* Shoot length measured
in cm;
other characters
measured in
mm.
** N/n: number of populations sampled over number of individual measurements.
Vi 11 DM. Because we sampled several populations
of both dwarf mistletoes and completed at least 100
measurements of the morphological characters se¬
lected for analysis, we feel the morphological dif¬
ferences we found between SPDM and W 11 DM are
consistent and are taxonomically important discon¬
tinuities.
The principal host ol shore pine dwarf mistletoe
is clearly shore pine, and not western hemlock,
which we classify as an occasional host of SPDM
using the host susceptibility system of Hawksworth
and Wiens (1972, 1996). When exposed to western
hemlock dwarf mistletoe in mixed western hem¬
lock/shore pine forests, shore pine is a rare host of
this mistletoe. These large differences in suscepti¬
bility between these hosts represent distinct and
consistent physiological discontinuities between
these mistletoes and represent taxonomically im¬
portant characters distinguishing them. Artificial
inoculations also indicate that these large differ¬
ences in host preference exist between SPDM and
WHDM (Smith, 1974; Smith & Wass, 1979; Wass,
unpublished).
Hawksworth and Wiens (1972. 1996: 146) de¬
fined subspecies in Arceuthobium as “geographi¬
cally restricted populations delimited by a few rel¬
atively small but consistent variations.’' Although
the distributions of the shore pine dwarf mistletoe
and western hemlock dwarf mistletoe overlap, they
have a few consistent morphological differences be¬
tween them and clearly demonstrate different levels
of parasitism on shore pine and western hemlock.
Because of these morphological and physiological
differences, the SPDM should he recognized as a
subspecies of hemlock dwarf mistletoe based on
Hawksworth and Wiens' concept of subspecies in
Arceuthobium.
Furthermore, giving shore pine dwarf mistletoe
taxonomic status at the subspecific level is consis¬
tent with the Hawksworth et al. (1992) classification
of mountain hemlock dwarf mistletoe as a subspe¬
cies of hemlock dwarf mistletoe. Our field studies
in the Calapooya Mountains of south central Oregon
indicate that mountain hemlock dwarf mistletoe is
not geographically restricted from populations of
western hemlock dwarf mistletoe. In addition, only
<—
phological data was collected. Washington: I—Snoqualmie Pass, 2—Westport, 3—Huckleberry Creek, 4—Cortright
Creek, 5—Clearwater Creek, 6—Wind River Experimental Forest: Oregon: 7—Wapinitia Pass, 8—Desolation Sad¬
dle; 9—Huckleberry Creek, 10—Indigo Spring; II—Wall Creek, 12 —Calapooya Ridge, 13 —Union Creek. 14—
Iron Mountain.
274
Novon
Table 2. Morphological characteristics of female plants
collections from British Columbia and the United States.
for shore pine
and western
hemlock dwarf mistletoes for
Female plant
Shore pine dwarf mistletoe
Western hemlock dwarf mistletoe
characters*
Mean
Range
N/n**
Mean
Range
N/n**
>F
Tallest shoot length
6.6
4.0-9.5
1 1/1 10
8.0
3.8-13.7
20/265
0.0031
Shoot basal diameter
3.3
1.8-5.0
1 1/1 10
2.7
1.3-5.5
20/265
0.0214
Shoot third internode
length
10.7
5.3-16.4
1 I/I 10
12.3
6.0-22.0
20/265
0.0050
Shoot third internode
width
1.7
1.3-2.5
1 1/1 10
1.6
1.0-3.1
20/265
0.2584
Third internode length/
width ratio
6.2
3.6-0.4
1 1/110
7.8
3.3-16.0
20/265
0.0007
Fruit length
4.6
3.3-5.7
1 I/I 10
4.4
3.3-5.5
18/210
0.2058
Fruit width
3.1
2.1—4.2
1 I/I 10
2.9
2.2-3.5
18/210
0.0363
Seed length
2.5
1.8-3.0
10/100
2.6
1.8-3.5
17/200
0.1193
Seed width
1.4
1.0-1.7
10/100
1.1
0.8-1.4
17/200
0.0001
* Shoot length measured
in cm;
other characters
measured
in mm.
** N/n: number of populations sampled over number of individual measurements.
a lew morphological and physiological differences
distinguish these subspecies (Hawksworth et al.,
1992; Hawksworth & Wiens, 1996). Differences in
plant size, phenology, and host range are the prin¬
cipal characters used by Hawksworth et al. (1992)
to separate those subspecies, anti these are the
same characters that distinguish shore pine dwarf
mistletoe from western hemlock dwarf mistletoe
(Table 3). Therefore, the classification of shore pine
dwarf mistletoe as a subspecies of hemlock dwarf
mistletoe, instead of a race, is more consistent witli
the interpretation for hemlock dwarf mistletoe pro¬
posed by Hawksworth et al. (1992) and with the
taxonomic framework established in Hawksworth
and Wiens’ (1996) monograph of Arceuthobium.
Arceuthobium tsugense subsp. contortae. Paratypes. All
citations based on Pinus contorta var. contorta. U.S.A.
Washington: San Juan Co., summit of Mount Constitu¬
tion. Orcas Island, 1921, Beattie 5801 (FPF); summit of
Mount Constitution, 1965, Graham & Thompson s.n.
(FPF); Mount Constitution, near summit, 1972, Hawk¬
sworth et al. 1442 (FPF); Moran State Park, Orcas Island,
1987, Hawksworth 2211 (FPF); Little Summit, Orcas Is¬
land, 1997. Mathiasen 9702 (ASC); summit of Mount Con¬
stitution, 1997, Mathiasen 9702 (ASC); midway between
Table 3. Principal morphological and physiological differences between shore pine dwarf mistletoe (SPDM) and
western hemlock dwarf mistletoe (WHI)M).
Characters
SPDM
WHOM
Mean plant height:
male
5.6 cm
7.8 cm
female
6.6 cm
8.0 cm
Mean length of third internode:
male
9.2 mm
1 1.8 mm
female
10.7 mm
12.3 mm
Mean flower diameter
4.3 mm
3.6 mm
Mean seed width
1.4 mm
1.1 mm
Seed length to width ratio
1.8
2.4
Plant color:
male
green-brown
yellow-green
female
green-brown
yellow-green/purple
Host susceptibility:
shore pine
principal host
occasional host
western hemlock
rare host
principal host
Peak seed dispersal
one week earlier than W HDM
one week later than SPDM
Volume 13, Number 2
2003
Wass & Mathiasen
Arceuthobium tsugense subsp. contortae
275
Table 4. Infection of shore pine and western hemlock by shore
mistletoe.
pine dwarf mis
itletoe and western hemlock dwarf
Shore
pine
Western
hemlock
frees
Percent
frees
Percent
Dwarf mistletoe
Forest type
sampled
infected
sampled
infected
shore pine
pure shore pine
765
99
—
—
shore pine and western
81 1
95
802
21
hemlock mix
western hemlock
western hemlock and shore
77
1
138
96
pine mix
Little Summit and Mount Constitution, 1997, Mathiasen
9705 (ASC). CANADA. British Columbia: Vancouver
Island, Metchosin, 1954, Kuijt 602 (FPF); Vancouver Is¬
land, Home Lake, 1954, Kuijt s.n. (FPF); Home Lake,
1969, R. S. Smith s.n. (FPF); Home Lake, 1977, Hawk-
sworth 1814 (FPF); Horne Lake, 1987, Nick rent 2664
(FPF); Horne Lake, 1997, Vancouver Island, Courtenay,
1954, Foster s.n. (FPF); Vancouver Island, Coldstream,
near summit of Mount Finlayson, 1965, Ziller s.n. (FPF);
Vancouver Island, summit of Mount Finlayson, 1987,
Nickrent 2666 (FPF); Vancouver Island, Nanoose Bay,
1954, Kuijt s.n. (FPF); Savary Island, 1918, Davidson s.n.
(FPF); Vancouver Island, Spider Lake, 6 km S of Quali-
cum, 1987, Nickrent 2665 (FPF); Spider Lake, Lakeview
lid., Wass 26851 (3) and 26852 ( 9) (DAVFP); mainland
British Columbia, Sechelt, 1954, Kuijt s.n. (FPF); Sechelt,
1997 and 1998, 26861 (6) and 26862 (9) (DAVFP);
Cortes Island, Mansons Landing, 1997, Wass 26853 (6)
and 26854 ( 9) (DAVFP); Vancouver Island, summit of
Mount Helmcken, Victoria, 1997, Ho.s.s 26855 (d) and
26856 (9) (DAVFP); Vancouver Island, summit of Bluff
Mountain, Sooke, 1997, Wass 26857 (cJ) and 26858 (9)
(DAVFP); Vancouver Island, summit of Mount Work, Sid¬
ney, 1997. Wass 26859 (d) and 26860 (9) (DAVFP); Van¬
couver Island, summit of Trap Mountain, Sooke, 1998,
Wass 26865 (d) and 26866 (9) (DAVFP); Texada Island.
Mt. Pocahontas Rd., 1997 and 1998, Wass 26863 (d)and
26864 ( 9) (DAVFP).
Arceuthobium tsugense subsp. tsugense. Paratypes. All
citations based on Tsuga helerophyila. U.S.A. Washing¬
ton: Skamania Co., Wind River Experimental Forest,
1998 and 2000, Mathiasen 98125 ( 9) and 0028 (d)
(ASC); Clearwater Creek, 1998 and 2000, Mathiasen
98126 ( 9) and 0029 (d) (ASC); Lewis Co., Cortrighl
Creek, 1998 and 2000, Mathiasen 98127 (9) and 0030
(d) (ASC); Pierce Co., Huckleberry Creek. 1998 and
2000, Mathiasen 98128 ( 9) and 0031 (d) (ASC); Kittitas
Co.. Snoqualmie Pass, 1998 and 2000, Mathiasen 98129
(9) and 0032 (d) (ASC); Grays Harbor Co., Westport,
1998, Mathiasen 98130 (9) (ASC); Oregon: Jackson Co.,
NE of Union Creek. 1998 and 2000. Mathiasen 98113
(9) and 0034 (d) (ASC); Douglas Co., Calapooya Ridge,
1998, Mathiasen 98116 (9) (ASC); Lane Co., Indigo
Springs, 1998 and 2000, Mathiasen 98118 (9) and 0033
(d) (ASC); Wall Creek, 1998 and 2000, Mathiasen 98121
(9) and 0035 (d) (ASC); Huckleberry Creek. 1998 and
2000, Mathiasen 98122 ( 9) and 0036 (d) (ASC); Wasco
Co., Wapinitia Pass, 1998 and 2000. Mathiasen 98124
(9) and 0037 (d) (ASC); Lincoln Co.. Desolation Saddle,
1998, Mathiasen 98131 (9) (ASC); Coos Co., E of Iron
Mountain, 1998, Mathiasen 98132 ( 9) (ASC). CANADA.
British Columbia: Vancouver Island, Spider Lake, Quali-
cum Beach. 1997, Wy/.s.s- 26869 (d) and 26870 (9)
(DAVFP); Texada Island, Mt. Pocahontas Rd., 1997 and
1998, Wass 26871 (d) and 26872 (9) (DAVFP); Vancou¬
ver Island, Holt Creek, Duncan, 1998. Wa.M 26873 (d)
and 26874 (9) (DAVFP); Vancouver Island, Caycuse
Summit, Honeymoon Bay, 1997 and 1998. Wass 26875
(d) and 26876 (9) (DAVFP); Vancouver Island, 1.2 km
up Crosby Rd., Bowser, 1998, Wass 26877 (d) and 26878
(9) (DAVFP).
Acknowledgments. We thank Richard B. Smith,
Carolyn M. Daugherty, and Greg Filip for manu¬
script reviews. Technical assistance was provided
by Leo Unger and Jason Nazar.
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- & -. 1996. Dwarf mistletoes: Biology, pa¬
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Hunt, R. S. & R. B. Smith. 1978. Natural infection of two
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dence for genetic differentiation in two host races of
hemlock dwarf mistletoe ( Arceuthobium tsugense).
Biochem. Syst. Fcol. 18: 267—280.
- , K. P. Schuette & E. M. Starr. 1994. A molecular
phylogeny of Arceuthobium (Viscaceae) based on nuele-
276
Novon
ar ribosomal 1)!\A internal transcribed spacer sequenc¬
es. Amer. J. Bot. 81: 1149-1160.
SAS INSTITUTE INC. 1985. SAS user’s guide: Statistics.
Version 5 Ed. SAS Institute Inc., Cary, North Carolina.
Smith, R. B. 1974. Infection and development of dwarf
mistletoes on plantation-grown trees in British Colum¬
bia. Canad. Forest Serv., Pacific Forestry Centre, In¬
form. Rep. BC-X-97.
-& E. F. Wass. 1976. Field evaluation of ecological
differentiation of dwarf mistletoe on shore pine and
western hemlock. Canad. J. Forest. Res. 6: 225—228.
- & -. 1979. Infection trials with three dwarf
mistletoe species within and beyond their known ranges
in British Columbia. Canad. J. PI. Pathol. I: 47—57.
Wass, E. F. 1976. Ecology of shore pine stands infested
with dwarf mistletoe on southeastern Vancouver Island.
Canad. Forest Serv., Pacific Forestry Centre, Inform.
Rep. BC-X-142.
A New Species of the Genus Tulipa (Liliaceae) from China
Yi-Zhi Zliao
Faculty of Life Science, Inner Mongolia University, Hohhot 010021, China
Abstract. Tulipa mongolica Y. Z. Zhao (Lili¬
aceae) is described from Inner Mongolia, China.
The new species is distinguished by its filaments
2.5—3.0 times as long as die anthers. It is related
to T. heteropetala Ledebour on the basis of its fil¬
aments widened from the mid-point down, narrow¬
ing on both sides of the filament.
Key words: China, Inner Mongolia, Liliaceae,
Tulipa.
flic genus Tulipa L. contains about 150 species,
about 14 of which are found in China. Among these
there is only one species in Mongolia and Inner
Mongolia. Pile two species Tulipa heteropetala and
Tulipa uniflora (L.) Besser ex Baker have the clos¬
est relationship with this new species. The distri¬
bution of Tulipa heteropetala is in the north of Xin¬
jiang, Siberia, and Central Asia. Tulipa uniflora
occurs in Siberia and Central Asia.
The new species was referred to Tulipa hetero-
petala in Mao (1980) and Tulipa uniflora in Ninbu
(1994). After comparing the features among these
three species of Tulipa the author thought this spe¬
cies was new to science.
The new species was distributed solitarily over a
sandy crack of volcanic cone in the XilinGol Meng
of Inner Mongolia. The species is now extinct at
the type locality because of heavy grazing pressure.
Tulipa mongolica Y. Z. Zhao, sp. nov. TYPE: Chi¬
na. Inner Mongolia: XilinGol Meng. Baiyinxi-
le, 20 May 1979. Shu-run Uu 395 (holotype,
HIMC; isotypes. HIMC). Figure 1A-C.
Haec species T. heteropetalae Ledebour affinis, sed ab
ea perianthii segmentis intemis oblongo-obovatis, 2.5-4.5
<'in longis apice acutis; filamentis 2.5—3.0-plo longioribus
quam antheris atque pistillo longiore quam staminibus dif¬
fer!.
Perennial, 10—25 cm tall. Bulb ovate, 1—2 cm
diam.; skin of bulb papery, dark brown, pubescent
on inner surface; stem glabrous. Leaves 2, closely
alternate, narrowly lanceolate, 8-1 I cm long, 4-9
nun wide, often curved outward, glabrous. Flower
single, terminal; perianth segments 6, in two series
of 3, yellow, 2.5—4.5 cm long, 4—8 mm wide, apices
acute; outer perianth segments 3, narrowly oblan-
eeolate, externally green-purple; inner perianth 3,
oblong-obovate, same length as above, 8—15 mm
wide; stamens 6, in two series with 3 long, to 2.4
cm, and 3 short, to 2.2 cm long; filaments 2.5—3.0
times as long as anthers, yellow, glabrous, widened
below their mid-point, gradually narrowed toward
both sides; anther yellow, narrowly oblong, 4—0 mm
long; pistil exceeding the stamens; style 1 cm long.
Fruit and seed not found.
Habitat. On stony hillside fields.
Distribution. Inner Mongolia. XilinGol Meng,
Baiyinxile.
Paralype. CHINA. Inner Mongolia: same locality as
type collection, 8 May 1965, Zhi-jie Xu 413 (HIMC).
flic chief morphological differences between the
new species and two similar species are summa¬
rized in the following key couplet. The three spe¬
cies of Tulipa also share in common a yellow flower,
long style, and pubescence on the top of the inner
skin of the bulb.
Ke> to thk Three Tvupa Species in Inner Mongolia,
China
la. Only the base of the filaments of stamens wid¬
ened. narrowing to the apex .
. 1. T. uniflora (L.) Besser ex Baker
If). Filaments widened from the mid-point down,
then narrowing on both sides of die filament.
2a. Stamen filaments equal in length to anthers or
slightly longer; pistil shorter than stamens; in¬
ner perianth ovate to rhombic, 2—2.5 cm long,
apex acuminate; leaves 2. loosely separate
.2. 7. heteropetala Ledebour
2b. Stamen filaments 2.5—3.0 times as long as
anther; pistil longer than stamens; inner
perianth oblong to obovate, 2.5—4.5 cm long,
apex acute; leaves 2, closely alternate . . .
.3. T. mongolica Y. Z. Zhao
Paratype. CHINA. Inner Mongolia: XilinGol Meng,
Baiyinxile, 8 May 1965, Zhi-jie Xu 413 (HIMC).
Literature Cited
Mao, Zu-mei. 1980. Tulipa. Pp. 86-97 in Flora Reipub-
lieae Popularis Sinicae, Vol. 14. Science Press, Beijing.
Ninbu. 1994. Flora lntramongolica, 2nd ed. Typis lntra-
mongolicae Popularis Hohhot. 5: 469.
Novon 13: 277-278. 2003.
278
Novon
Figure 1. Tulipa mongolica Y. Z. Zhao. —A. Plant. —B. Skin of bulb. —C. Part of flower.
Note
Packera lyallii (Asteraceae: Senecioneae) Is a Superfluous Name
ABSTRACT. Packera contermina is (he correct
name for the taxon erroneously published as Pack-
era lyallii.
Key words: Asteraceae. Packera, Senecioneae.
My publication of the name Packera lyallii
(Klatt) Bain (2002) was done in error, based on
misinterpretation of Artiele 53.6 of (he ICBN
(Greuter et al., 2000).
Article 53.6 does not apply in the above case
because Packera lyallii is a later homonym and is
not a homonym of equal priority to which the article
refers. Packera contermina (Greenman) Bain is
therefore the correct name for this taxon, and P.
lyallii is a superfluous name.
My argument for adopting the name Packera
cymbalarioides (Nuttall) Weber and Love to replace
Packera streptanthifolia (Greene) Weber and Love
is based on a similar misinterpretation of Article
53.6 and is also incorrect.
Literature Cited
Bain, J. F. 2002. Packera lyallii, a corrected name for
Packera contermina (Asteraceae: Senecioneae). Novon
12: 437.
Greuter, W., J. McNeill, F. K. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & D. L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
—John F. Ruin. Biological Sciences Department, University
of Lethbridge, 4401 University Drive, Lethbridge, Alberta
TIL .'1M4. Canada, bain@uleth.can
Novon 13: 279. 2003.
Volume 13, Number 2. pp. 161-280 of NO VON was published on 24 June 2003.
Volume 13
Number 3
2003
NOVON
A New Hard Pine (Pinus, Pinaceae) from Taiwan
Roman Businsky
Silva Tarouca Research Institute for Landscape and Ornamental Gardening (RILOG),
252 43 Pruhonice, Czech Republic, businsky@vukoz.cz
Abstract. Finns fragilissima Businsky (Pina¬
ceae), a new species of Finns subg. Finns , is de¬
scribed from southeastern Taiwan. Comprised of
trees with very sparse crown and fragile, symmet¬
rical. 6—9 cm long cones with often flat apophyses,
it appears to be most closely related to F. luchuensis
Mayr, endemic to the Nansei Islands, and to F. tai-
wanensis Havata. The latter is circumscribed here
as a Taiwan endemic with the exclusion of super¬
ficially similar but probably unrelated mainland
Chinese pines. These three allied species are clas¬
sified here as the sole representatives of Finns
subg. Finns ser. Luchuenses E. Murray.
Key words: Pinaceae, Finns , Pinus subg. Finns
ser. Luchuenses, Taiwan.
The author has been studying natural popula¬
tions of Eurasian pines for more than ten years,
concentrating on East Asia. The main object of the
study is a taxonomic revision of Pinus in that re¬
gion, the Revision of Pines ol East Asia (REPEA)
Project. Within the framework of this project, the
author undertook a series of research expeditions
to East Asia: eight to the People’s Republic of Chi¬
na (1990—2001), two to Japan (1991 and 1997), two
to Vietnam (1994 and 1997), and one to Taiwan
(1991), Indonesia (1994). and the Philippines
(2000). The study has been designed to obtain good
field knowledge, above all of variability within and
between populations, and to collect representative
herbarium and photographic material of all Asian
species of the genus Firms, allowing a more uniform
approach to classification (see also Businsky,
1999).
Taxonomy
During an exploration in 1991 of forest stands in
southern Taiwan, on the eastern (Pacific) side of the
island's central mountain range, a remarkable pop¬
ulation of a hard pine (= Firms subg. Firms ) near
Wulu village in the northern part of Taitung County
was found. The only species known from Taiwan
showing certain resemblance in general tree habit,
external leaf characters, and some cone characters
to this population is Firms massoniana Lambert.
Critchfield and Little (1966), using unpublished
data at the Taiwan Forest Research Institute, re¬
ported P. massoniana only from northern Taiwan.
However, Liu (1966) and Li (1975) also reported F.
massoniana in the south, but only from the eastern
coastal hills along the border between Taitung and
Hualien Counties. Liu (1966: fig. 33) listed only
one species of hard pine, F. taiwanensis Hayata, in
the Wulu region. In a narrow taxonomic concept
(excluding Pinus hwangshanensis W. Y. Hsia, see
below), P. taiwanensis is confined to middle and
high altitudes of Taiwan, widespread mainly be¬
tween 2200 and 2900 m. However, the pine from
the vicinity of Wulu occurs near or below 900 m,
anti differs from P. taiwanensis in several morpho¬
logical characters (Table 1). No other indigenous
hard pine is known from Taiwan. Outside of Taiwan,
the Wulu population approaches F luchuensis Mayr
in some characters (leaf morphology and anatomy,
distinctly broad tree crown); this is a species en¬
demic to the nearby flat islands of the Nansei (Ryu¬
kyu) Archipelago, northeast of Taiwan (ef. Critch¬
field & Little, 1966; Yamazaki, 1995). The
character set found in the Wulu pine is different
from that of the three geographically closest spe-
Novon 13: 281-288. 2003.
282
Novon
eies: P. massoniana, P. taiwanensis, and P. luchuen-
sis. The Wulu pine also cannot he equated widi any
other pine indigenous to East Asia, either; so as
discussed below it is described here as a new spe¬
cies.
I'imis fragilissima Businsky, sp. nov. TYPE: Tai¬
wan. Taitung County: below the great bend of
Southern Cross-Island Highway on S slopes
about I km N of Wulu village, in mixed forest
on W declivity of a stony ridge descending S,
alt. 930 m. 23°10'40"N, 121°02'E, 18 Dec.
1991, B. Businsky 32172 (holotype, PR; iso¬
types, R, BM, C, G, MO, P, PE, TAI & Her¬
barium of the RILOG). Figure 1.
Arbor usque 30 m alta, corona eonspicue sparse dif¬
fusa, lata; folia (12—) 16—20(—22) cm longa, canalibus re-
siniferis 4 ad 6(7), plerumque medialibus et interdum
(sub)marginalibus; strobili ovuliferi hornotini lere 9—10
mm lali, pedunculis 5—10 mm longis, squamis mucronatis
praeter partem strobilorum basalem; strobili maturi (5—)6—
9(—10) cm longi, fragiles, patentes vel leviter reflexi;
apophyses plerumque prominentes, non tumidae. A Pinus
massoniana differt imprimis canalibus resiniferis foliorum
plerumque medialibus et umbonibus strobilorum maturo-
rum plerumque prominentibus, non planis et concavis; a
P. taiwanensi differt imprimis foliis longioribus, strobilis
maturis longioribus, fragilibus et coronis arborum sparse
diffusis; a P. luchuensi differt imprimis strobilis maturis
longioribus et fragilibus, non suberectis, apophysibus
plerumque pyramidal ibus, non rotundatis vel tumidis et
coronis arborum sparse diffusis.
Tall trees attaining a height of 30 m with con¬
spicuously sparse crowns 20 m or more wide,
branches spreading, remarkably frangible, with
leaves falling during the second to third year. Bark
developing late, initially irregularly scaly, on trunks
of old trees conspicuously thick, deeply longitudi¬
nally fissured anti forming ribs often around 10 cm
thick. Annual branches uninodal, relatively long;
shoots of the current year relatively thin (fertile
shoots with conelets 3-6 mm thick), yellow-brown,
initially slightly pruinose, glossy, with inconspicu¬
ous shallow and narrow grooves formed by the de¬
current bases of the primary bracts; areolae among
them low and relatively flat. Shoots of the previous
year gray-brown. Buds cylindrical, up to ea. 3 cm
long, with scales in the upper half loose to erecto-
patent. narrowly lanceolate, with inconspicuous
dorsal keel, brown in the middle, paler toward the
lacerate margins, with long white fringes; scales not
persistent at the bases of shoots. Leaves in fascicles
of 2, occasionally in 3s on fertile shoots, (12—) 16—
20(—22) cm long, (0.9-)1.0-1.2(-1.35) mm wide
(fresh), pale green, relatively fine and flexible,
straight or slightly bent: amphistomatic; edges ir¬
regularly and rather densely acute-serrate, with (35
to)45 to 65(to 80) teeth per cm in the middle part;
sheaths (8—) 10— 13(— 15) mm long in the first year.
Leaf resin duets 4 to 6(7); two ducts near leaf edges
always median, of largest diameter; 1 to 3(4) ducts
dorsal, usually of smaller diameter, median or 1 to
2 of them marginal, occasionally submarginal, or
near endodermis: usually one duct near the ventral
side of small diameter, median to marginal (rarely
larger and septal). Leaf hypodermis formed mostly
by one layer of cells with relatively thin walls,
sometimes a second layer in scattered patches on
the dorsal side. Pollen cones ca. 2.5 cm long. Ovu¬
late cones after the first growing season (conelets)
on 5—10 mm long peduncles, 13—18(-20) mm long
and ea. 9—10 mm wide, ovoid to elliptic cylindrical,
erect, subterminal, usually in whorls of 3 to 5. The
exposed part of the scales ol the basal quarter of
the conelet rounded, without a distinct keel or mu-
ero; the other scales often show a considerably el¬
evated area with a distinct transverse keel without
discernible apex, dorsally depressed, with short
mucro situated eccentrically on the ventral side,
beginning from the swollen decurrent base and ori¬
ented backward over the keel. Mature seed cones
spreading or slightly reflexed on slender 5-10 mm
long peduncles, symmetrical, ovoid to oblong con¬
ical. fragile, (5—)6—9(—10) cm long, 5—8 cm wide
when open; usually persistent for a few years, fall¬
ing mostly without peduncle. Cone scales thin,
densely arranged, 120 to 220. the largest 20—30
mm long and 12-15 mm wide, with length/width
ratio 1.5—2.5, often broader below the middle than
the apophysis width. Apophyses cinnamon brown, in
outline irregularly transverse obtrullate, 1 1-14 mm
wide, with rounded to broadly cuneate distal edge,
± broadly pyramidal (not rounded or tumid), with
conspicuous, sharp transverse keel and generally
concave, depressed proximal side (often with raised
proximal corner). Umbo slightly sunken, but usu¬
ally prominent, transverse, (2—)2.5-4(—4.5) mm
wide on seed scales, pyramidal or rooflike, keeled,
often depressed on the proximal side, with minute,
slender, ca. 0.2—0.5 mm long, erect or recurved
mucro. Seeds ca. 4—5.5 X 2.2—3 mm, with wings
I 1-25 X 4.5—7.5 mm.
Etymology. The specific epithet (denoting “fra-
gilest”) refers to the remarkably frangible branches
and branchlets, more so than for all other East
Asian pines, and to the fragile consistency of the
scales of mature seed cones.
Distkibution and Ecology
Pinus fragilissima was observed only in a rela¬
tively small valley system of the Hsinwulu River,
Volume 13, Number 3
2003
Businsky
Pinus fragilissima from Taiwan
283
figure 1 . Pinus fragilissima Businsky. —A. fertile branehlet in winter season (current shoot, with top of previous year's
shoot) with leaf fascicles, terminal bud and ovulate cones. —B. Mature dosed cone. —C. Open seed cone. —I). Cross
sections at the middle of leaves showing position of resin ducts. —E. Open seed cone ol another individual. A—D from R.
Businsky 32172. holo- and isotypes, and E from R. Businsky 32173, paratype. A, B, C. E: the same scale bar. All drawn 1>\
Ludmila Businska.
284
Novon
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Volume 13, Number 3
2003
Businsky
Pinus fragilissima from Taiwan
285
descending east-southeast from the Kuan Shan
massif (3666 m) within the Chungyang Shanmo
mountain region, in the vicinity of and below Wulu
village in the northern part of Taitung County, Tai¬
wan, on the eastern side of the island’s central
mountain range. The population of the new pine in
the main valley of this system (i.e., along the South¬
ern Cross-Island Highway) occurs in elevations be¬
tween 500 and 1000 m, but it cannot be excluded
that it reaches slightly higher altitudes in some of
the side valleys of the system. The valley is char¬
acterized by steep, rocky slopes predominantly cov¬
ered with broad-leaved woodlands. Pinus fragilis-
sima prefers slopes of southern to western
exposures, confined to rocky sites with reduced
competition from broad-leaved trees. Potentially,
tbe new pine may be found in suitable habitats in
other less easily accessible valley systems in the
Chungyang Shanmo mountains in Taitung and Hu-
alien Counties.
Another population of pines occurs in similar al¬
titudes eastward, opposite the slopes of the Chun¬
gyang Shanmo mountains, i.e., in a narrow belt ol
coastal hills (situated between ca. 23°00' and
23°25'N) called Haian Shanmo. This population
was identified as Pinus massoniana by bin (1966)
and Li (1975). Earlier on, Li (1963) cited a single
specimen, Tanaka 10453, from southern Taiwan, at
“Taitung,” as P. massoniana. Liu’s map (1966: fig.
31) of P massoniana in Taiwan includes an isolated
area in these coastal hills as a narrow strip about
60 km long following the main ridge of the range.
A single dot representing a herbarium specimen is
in the middle of this strip, in the vicinity of Mt.
Xingang Shan (1682 m, on the border of Taitung
and Hualien Counties, above the port of Cheng-
kung), about 20 km east of the observed site ol P.
fragilissima in the Hsinwulu valley. No herbarium
specimens are cited by Liu, but Li (1963) is cited,
and thus it is presumed here that the dot refers to
the specimen cited by Li (Tanaka 10453). A du¬
plicate of Tanaka 10453 (C), determined as P. mas¬
soniana by the collector, was examined and un¬
doubtedly is P. fragilissima ; consequently the whole
population of pines in the Haian Shanmo is con¬
sidered here belonging to this species. This moun¬
tain range reaches the highest elevation in Xingang
Shan, which is close to the Hsinwulu River system.
The region with P. fragilissima as a whole is char¬
acterized by dry winters and wet summers (Walter
& Lieth, 1964: part 2.4).
Paratypes. TAIWAN. Taitung County, below the great
bend of Southern Cross-Island Highway on S slopes about
1 km N of Wulu village, in sparse mixed forest on S stony
declivity, alt. 930 m, 23°10'40"N, 12T02'E, 18 Dec.
1991, R. Businsky 32173 (I’ll); Tai to-cho [Taitung],
Kaede, 7 Mar. 1931. T. Tanaka 10453 (C).
Discussion
Pinus fragilissima is similar to P. luchuensis, par¬
ticularly in leaf size and anatomy. However, they
otherwise differ morphologically and in their eco¬
logical requirements. The new species differs from
P. massoniana in several characters, notably in the
position of the resin ducts, a character usually giv¬
en as important in the classification and determi¬
nation of the Asian hard pines (cf. Gaussen, 1960;
kwei & Lee, 1963; Cheng et ah. 1975; Law et al.,
1978). On the other hand, both species occupy a
similar altitudinal range; according to Liu (1966)
and IVlirov (1967), P. massoniana in Taiwan is con¬
fined to 300—1300 m. The new species is distinct
from P. taiwanensis in general habit (sparse broad
crowns), but further differences exist in leaf length,
cone size, and scale fragility. Although both species
occur in the same mountain region, their altitudinal
ranges are different and barely overlap. The alti¬
tude of 750 m recorded as the lower limit for P.
taiwanensis by Li (1963: 51), Liu (1966: 17), and
Mirov (1967: 284) is in conflict with field obser¬
vations ol the present author in several regions.
Even isolated trees of P. taiwanensis below its main
zone of occurrence were above 1500 m in most of
these regions. The 750 m record may be a result
of the inclusion of P. fragilissima or P. massoniana
in the distribution of P. taiwanensis. Sheue et al.
(2000) analyzed variation of leaf anatomy attributed
to P taiwanensis in the Tachia river system (mainly
in Taichung County on the west side of northern
Taiwan) in five localities from an altitude of 700 m
(Kukuan) to 3100 m (Konankaun). Li (1963, 1975)
and Liu (1966) reported from this region also an
occurrence of P. massoniana. The present author
studied pine populations and collected samples of
trees in the Tachia river system in the mentioned
altitudinal range as well. The population of a hard
pine in the vicinity ol Kukuan (occurring between
500 and 1000 m) is, however, considered to be P.
massoniana, comprising also some individuals mor¬
phologically transitional to P. taiwanensis (probably
of hybrid origin). On the basis of a suite of mor¬
phological and anatomical characters, ranges of
variability, and the geographic distribution, the trio
of Pinus luchuensis, P. taiwanensis, and P. fragil¬
issima appears as a related group, accepted here as
the series Luchuenses E. Murray within the typical
section and subsection of the genus.
Some Chinese authors, e.g.. Law et al. (1978)
and Eu and Li in Fu et al. (1999), include mainland
populations described (and accepted here) as Pinus
286
Novon
hwangshanensis W. Y. Ilsia in P. taiwanensis. How¬
ever. P. hwangshanensis appears more closely re¬
lated to the Japanese P. thunbergii Parlatore (Wang
& Szmidt, 1993). Field and herbarium studies of P.
hwangshanensis (see Appendix 1) show it to be dif¬
ferent from P fragilissima in general habit, mor¬
phology, and anatomical characters, and also in
ecological requirements. While P. hwangshanensis
prefers exposed rocks on the tops and ridges of
continental mountains predominantly above 900 m,
P. fragilissima prefers sheltered rocky sites in val¬
leys of (sub)eoastal mountains predominantly below
900 m.
Key to the Species Morphologically Similar to
PlNUS FRACtUSStMA
1. Leaves with resin duets always marginal, usu¬
ally less than 1 mm wide and more than 12 cm
long; apophyses of seed cones relatively thin,
flat or slightly prominent, umbo flat and gen¬
erally concave as a whole, with indistinct or
depressed transversal keel . P. massoniana
1'. Leaves with resin ducts only or largely median,
often more than I mm wide or less than 14 cm
long; apophyses of seed cones usually broadly
pyramidal or roundly prominent to tumid, umbo
prominent (although sometimes sunken), gen¬
erally pyramidal and pointed or rooflike (some¬
times ± depressed on the proximal side or on
the top), with distinct and usually raised trans¬
versal keel . 2
2(1). Leaves of mature trees (7—) 12—20(—22) cm long;
seed cones generally symmetrical, early decid¬
uous or persistent for a lew years; branchlets
remarkably frangible; species of flat islands,
hills, and submontane valleys; only or predom¬
inantly below an altitude of 9(H) m . 3
2'. Leaves of mature trees (4—)5—14(—18) cm long;
seed cones generally asymmetrical, usually
firmly persistent for more years; branchlets rel¬
atively firm; species of mountain tops, ridges,
and slopes, predominantly above an altitude of
9(H) m ... 4
3(2). bud scales persistent at the bases of shoots;
mature seed cones suberect to spreading, 4—6
(—6.5) cm long, with apophyses ± roundly
raised, usually at first pruinose; usually low
trees with relatively dense crown . . . P. luchuensis
3'. Hud scales not persistent at the bases of shoots;
mature seed cones spreading or slightly re¬
flexed, (5—)6—9(—I()) cm long, with apophyses
± pyramidally prominent, not pruinose; tall
trees with conspicuously sparse crown . . .
. P. fragilissima
4(2). Apophyses of seed cones usually light brown,
not contrasting in color with indistinct sealing
band on scales, proximal side of apophyses
generally convex (sometimes convex only near
umbo and concave toward the proximal apoph¬
ysis corner), umbo ± sunken but not flat . . .
. P. hwangshanensis
4'. Apophyses of seed cones usually cinnamon
brown, contrasting in color with distinct, lighter
sealing band on scales, the proximal side of
apophyses generally concave (sloped almost
from the umbo, sometimes with raised proximal
apophysis corner), umbo usually not sunken
. P. taiwanensis
Note. If. R. Mill in the Flora of China (Fu et
al., 1999) gave the following diagnostic characters
of Pinus hwangshanensis and P. taiwanensis: length
of needle sheaths, number of teeth per cm in the
middle part of needle margins, color of pollen
cones, and umbo morphology of seed scales. These
characters, except for color of pollen cones, were
tested by the author in detail in 44 individuals of
the former species and in 32 of the latter, in both
cases from several regions (see Appendix 1 for her¬
barium material examined). Only the umbo mor¬
phology was found partly in correspondence with
Mill (in Fu et al., 1999)—i.e., umbo sunken (“de¬
pressed”) but not flat versus not sunken (“not flat”).
The difference in quality of mucro (“prickle”) re¬
ported there was not found. Moreover, both leaf
characters were found entirely insignificant for dis¬
crimination between these species. That is to say:
the reported number of teeth does not agree for P.
taiwanensis (it does not even correspond with both
the type specimens), with the found range of
(25—)32—55(—70), not “26—35(—39)”; the reported
length of sheaths does not agree, either, for P.
hwangshanensis , with the found range of (6—) 10—
14(—17) mm, not “0.5—1 cm” (sheaths longer than
12 mm were found in populations in the Dabie
Shan Mis. north of the Yangtze River).
Acknowledgments. The author is grateful to the
management of the Silva Tarouea Research Insti¬
tute for Landscape and Ornamental Gardening,
Pruhonice, Czech Republic, for administration of
the REPEA Project with the support of the Czech-
East Asian Research Fund (grant No. 32), and to
the staff of Herbarium PR, Botany Department. Na¬
tional Museum in Prague, for their help with the
distribution of herbarium specimens abroad. The
author wishes especially to thank his wife, Ludmila
Businska, for many years of her infinite help, above
all in the field during the above-mentioned research
expeditions to East Asia.
Literature Cited
Businsky, It. 1999. Taxonomic revision of Eurasian pines
(genus Pinus L.)—Survey ol species and infraspecific
taxa according to latest knowledge. Acta Pruhon. 68: 7—
86 .
Cheng. Y\. C., L. k. Fu & C. Y. Cheng. 1975. Gymnos-
perniae Sinicae. Acta Phytotax. Sin. 13(4): 56—90, pi.
I -66.
Critehlield. Vi. B. & L. L. Little. 1966. Geographic l)is-
Volume 13, Number 3
2003
Businsky
Pinus fragilissima from Taiwan
287
tribution of the Pines of the World. U.S.D.A., Forest
Serv., Misc. Publ. 991. Washington, D.C.
Fu, L. K„ IN. Li & R. R. Mill. 1999. Pinaceae. Pp. 11-
52 in '/. Y. Wu & P. H. Raven (editors), Flora of China,
Yol. 4. Science Press, Beijing, and Missouri Botanical
Garden Press, St. Louis.
Gaussen, H. I960. Les Gymnospermes actuelles et fossi-
les. Fasc. VI, Chap. XL Generalites, Genre Pinus. Trav.
Lab. Forest. Toulouse, I. II, sect. 1, vol. 1, part. 2: I -
272.
Kwei, Y. L. & C. I,. Lee. 1963. Anatomical studies oi the
leaf structure of Chinese pines. Acta Bot. Sin. 11: 44—
60, pi. I—VI.
Law, Y. W., W. C. Cheng, L. k. Fu, C. I). Chu & C. S.
Chao. 1978. Pinus. Pp. 204—281 in W. C. Cheng & L.
K. Fu (editors). Flora Reipublicae Popularis Sinicae
Vol. 7. Science Press, Beijing.
Li. H. L. 1963. Woody Flora of Taiwan. Livingston Pub¬
lishing, Narberth, Pennsylvania.
-. 1975. Pinaceae. Pp. 514—529 in H. L. Li, T. S.
Liu, T. C. Huang. T. Koyama & C. F. I)e Vol (editors),
Flora of Taiwan Vol. 1. Epoch Publishing, Taipei.
Liu. T. 1966. Study on the phytogeography of the conifers
and taxads of Taiwan. Bull. Taiwan Forest. Res. Inst.
122: 1-33.
Mirov, N. T. 1967. The Genus Pinus. Ronald Press, New
York.
Sheue, C. R„ Yang. Y. P„ Kuo-Huang, L. L. 2000. Struc¬
tural variation of the needles of Pinus taiwanensis Hay.
along an elevational gradient. Taiwan ,1. Forest. Sci. 15:
337-349.
Walter, H. & H. Lieth. 1964. Klimadiagramm-Weltatlas.
2. Lieferung. VEB G. Fischer Verlag, Jena.
Wang, X. R. & A. E. Szmidt. 1993. Chloroplast DNA-
based phylogeny of Asian Pinus species (Pinaceae). PI.
Syst. Evol. 188: 197-211.
Yamazaki, T. 1995. Pinaceae. Pp. 266—277 in K. Iwatsuki,
T. Yamazaki, 1). E. Boufford & H. Ohba (editors), Flora
of Japan Vol. 1. Kodansha, Tokyo.
Appendix 1. Herbarium Material Examined
HERBARIUM SAMPLES COLLECTED:
List of herbarium samples collected by the author is
ordered chronologically below according to localities
where natural populations of relevant species were studied
and representative trees sampled (month, year, and col¬
lection numbers are in brackets; R. B. is an abbreviation
of R. Businsky). Collections listed are stored in the her¬
barium of the Silva Tarouca Research Institute for Land¬
scape and Ornamental Gardening (RILOG), 252 43 Pru-
honice, Czech Republic.
Pinus taiwanensis Hayata:
• Central Cross-Island Highway SSL near Lishan, Nantou
Co., Taiwan; all. about 2050 m [Nov. 1991: R. R.
32137}
• Hsuehshan massif (3884 m), SE slopes, Taichung Co.,
Taiwan; alt. 2400—3400 m [Nov./Dec. 1991: R. R.
32140, 32141a—d, 32142, 32144a-i\
• Central Cross-Island Highway at Tayuling, Hualien Co.,
N part, Taiwan; alt. about 2600 m [Dec. 1991: R. R.
32146, 32147, 32148, 32158, 32159}
• Along the road from Tayuling to Wushe, Nantou & Hu¬
alien Co. boundary, N part, Taiwan; all. 2700—3200 m
[Dec. 1991: R. R. 32152, 32153, 32154, 32155 1
• Both sides of central ridge along the Southern Cross-
Island Highway, Kaohsiung & Taitung Co., Taiwan; alt.
2600-2800 m |Dec. 1991: R. R. 32164, 32165, 32166,
32167 1
Pinus luchuensis Mayr:
• Naha City suburbs, Okinawa Island, Nansei Islands,
Japan; alt. 20-30 m [Nov. 1991: R. R. 32120, 32121,
32122, 32123, 32124}
• Ishigaki town environs. Ishigaki Island, Nansei Islands,
Japan; alt. 80 m [Nov. 1991: R. R. 32125, 32126,
32127}
Pinus massoniana Lambert:
• Kukuan environs, Tachia valley, Taichung Co., Taiwan;
alt. 500-1000 m [Nov. 1991: R. R. 32131. 32132,
32133}
• Lechang Co., Guangdong, China; alt. 900—1200 m |Jan.
1994: R. R. 39121, 39124, 39125}
• Xingshan Co., W Hubei, China; alt. about 1400 m [July
1995: R. R. 42115}
• Tianzhu Shan massif (1488 m), Qianshan Co., SW An¬
hui, China; alt. 300—700 m [July 1995: R. R. 42116}
• Longming environs, Tiandeng Co., SW Guangxi, China;
alt. 500 m [Aug. 1998: R. R. 46121, 46122. 46123}
• Between Longming & Shangying, Tiandeng Co., SW
Guangxi, China; alt. 600 m [Aug. 1998: R. R. 46124}
• Darning Shan Mts., S slopes, Winning Co., Guangxi,
China; alt. 400-1000 m [Sep. 1998: R. R. 46126,
46127}
• Laibang environs, Dabie Shan Mts., Yuexi Co., Anhui,
China; all. 700 m [Sep. 1998: R. R. 46134}
Pinus hwangshanensis W. Y. Hsia:
• SW boundary of Rucheng Co. (Hunan) with Guangdong,
China; all. 1500 m [Jan. 1994: R. R. 39126, 39128.
39129}
• Tianzhu Shan massif (1488 m), Qianshan Co.. SW An¬
hui, China; alt. 600—1450 m |July 1995: R. R. 42117.
42118, 42119 1
• Miaodao Shan massif (1432 in). Yuexi Co.. SW Anhui.
China; alt. 800—1400 m [July 1995: R. R. 42120,
42121 |
• Mt. Tiantangzhai (1729 m), Yingshan Co., E Hubei,
China; alt. 700-1500 m [July 1995: R. R. 42124.
42125, 42126}
• Yaoluoping environs, Dabie Shan Mts., Yuexi Co., An¬
hui. China; alt. 900—1300 m [Sep. 1998: R. R. 46135,
46141, 46142, 46143}
• Between Laibang and Huo Shan massif. Dabie Shan
Mts., Yuexi Co., Anhui, China; alt. 1300 m |Sep. 1998:
R. R. 46144, 46145]
Pinus fragilissima Businsky—additional field label
information:
R. Businsky 32172 (holotype)—old tree with 270 cm trunk
circumference (at 1.3 m height), 23 m high, crown
about 20 m in diameter
R. Businsky 32173 (paratype)—old tree with 220 cm trunk
circumference, branches to 12 m long
OLDER SPECIMENS EXAMINED:
Pinus hwangshanensis W. Y. Hsia:
K. K. Tsoong 4495: A nhwei [= Anhui], Hwangshan;
9.10.1924. Det. as P hwangshanensis by W. Y. Hsia.
| PE]
M. Chen 703: Chekiang [= Zhejiang], Tianmu Shan;
1.7.1933. Det. as P. tabulaeformis Carr, by? W. C.
Cheng, and as P. hwangshanensis by W. Y. Hsia. [PE]
M. Chen 1267: Anhwei [= Anhui], Hwangshan, Shizi
288
Novon
Feng, 6200 ft.; 4.10.1933. Det. as P. tabulaeformis
Carr, by W. C. Cbeng, and as P. hwangshanensis by W.
Y. Hsia / H. H. Hu. [PE: two specimens]
II. II. Hu 2621: Kiangsi [= Jiangxi], [Jiujiang Cn.|, Eu
Shan, Ruling [= Guling], trees 30-40 m high; Aug.
1934. Del. as P. tabulaeformis Carr, by H. H. Hu, and
as P. hwangshanensis by W. Y. Hsia in 1936. “Para-
type." [PE]
P. C. Tsoong 3111: Anhwei [= Anhui], Hwangshan,
Hsi-hai-men; coll. N. T. Liou & P. C. Tsoong,
28.8.1935. Det. as P hwangshanensis by W. Y. Hsia.
| PE]
I*. C. Tsoong 3161: Anhwei [ = Anliuij, upper slope of
Mt. Tientu near summit; 13.6.1936. Det. as P. hwang¬
shanensis. | PE |
P. C. Tsoong 3919: Anhwei [= Anhui], on approaching
Shin-Jin, in wood; 9.7.1936. Det. as P. thunbergii, and
later, in 1956, as P. hwangshanensis. [PE]
X. Y. lie 30300: Zhej iang, Tianmu Shan; s. a. Det. as
P. hwangshanensis in 1964. [In Chinese; PE: specimens
ol five individuals]
P. taiwanensis Hayata:
T. Kuwukami & IJ. Mori 2097: Taiwan, Central Mis;
Nov. 1906. Lectotype of P. taiwanensis Hayata, des¬
ignated by A. Farjon, 4.3.1992. [TJ|
B. Hayata & U. Mori 7142: Taiwan, Randaizan;
9.8.1908. Syntype of P. taiwanensis Hayata. [Tl]
U. Mori, s. n.: Taiwan; s. a. Type of P. brevispica Hayata.
[TI]
A New Jacquinia (Theophrastaceae) from the Yucatan Peninsula
with a Synopsis of Pale-Flowered Species in Mesoanierica
German Carnevali Fernandez-Concha*, Silvia Hernandez-Aguilar, and Jose' Luis Tapia Munoz
Herbarium CIGY, Centro de Investigacion Cientifica de Yucatan, A.C., A.P. 87, Cordemex,
Merida 97310, Yucatan, Mexico, curneval@cicy.nix; silviaha@cicy.mx; jltapia@cicy.mx.
^Author to whom reprint requests should be addressed.
ABSTRACT. A synopsis of Mesoamerican Jacqui¬
nia (Theophrastaceae) species with white, yellow¬
ish, or cream-colored flowers is presented. A key
for their identification is provided. All of these spe¬
cies are restricted to northern Mesoameriea. Among
these taxa Jacquinia sak-lol Carnevali, Hernandez-
Aguilar & Tapia-Munoz, from the state of Quintana
Roo in the Mexican Yucatan Peninsula, is proposed
as new. The new entity is described and illustrated.
The proposed novelty is related to Jacquinia pal-
udicola Standley, but it is a shrub and features
smaller, more coriaceous leaves with a revolute
margin and larger flowers.
RESUMEN. Se presenta una sinopsis de las espe-
cies de Jacquinia (Theophrastaceae) de Mesoam-
erica con flores blaneas, amarillentas o color crema.
Se provee una clave para su identification. Estas
especies estan restringidas al norte de Mesoamer-
ica. Entre ellas, Jacquinia sak-lol Carnevali, Her¬
nandez-Aguilar & Tapia-Munoz del estado de
Quintana Roo en la Peninsula de Yucatan Mexi-
cana es propuesta como nueva. La nueva entidad
es descrita e ilustrada. Ea nueva espeeie es affn a
Jacquinia paludicola Standley pero es un arbusto,
tiene hojas de tamano menor y de textura mas eor-
iacea con el margen revoluto y flores mayores.
Key words: Jacquinia, Mexico, Theophrasta¬
ceae, Yucatan.
The genus Jacquinia E. (Theophrastaceae), com¬
prising 35 to 38 species, was recently revised by
Stahl (1989. 1995). A synopsis of Central American
taxa was published shortly thereafter (Stahl. 1989),
in which 11 species within Mesoameriea were rec¬
ognized. Eater, a somewhat aberrant species, Jac¬
quinia nemophila Pittier, restricted to Panama and
featuring larger, thinly textured green flowers and
large, wide leaves, was transferred to a new genus,
Votschia Stahl. This action reduced the number of
Central American Jacquinia to 10 (Stahl, 1993). An
additional species, J. morenoana Castillo-Campos
& E. Medina, was recently described from the state
of Veracruz in Mexico (Castillo-Campos & Medina,
1998).
Many species of Jacquinia are typically charac¬
terized by brightly colored red or orange flowers.
Among the Mesoamerican Jacquinia taxa treated by
Stahl (1989), five were described as having flowers
of other, paler colors, namely white or pale yellow.
As Stahl (1989) suggested, when discussing ,/. al-
Inflora Lundell this difference in flower color may
indicate a different pollinator fauna, although we
have no information on pollinators and floral biol¬
ogy of these species.
In Mesoameriea, the taxa of Jacquinia with white
or yellowish flowers are concentrated in the north¬
ern half. No Jacquinia species with pale-colored
flowers are known from Costa Rica or Panama, al¬
though there are several in the West Indies and one,
Jacquinia armillaris Jacquin, occurs in northern
South America (Stahl, 1995). Nicaragua, which is
the southernmost limit for pale-colored Jacquinia
species in Mesoameriea and where four Jacquinia
species are known to occur, harbors one pale-flow¬
ered taxon, J. longifolia Standley, the most wide¬
spread and variable of the taxa treated in this ac-
eount (Stahl. 2001). On the other hand, Belize,
where another set of four Jacquinia taxa have been
reported (Balick et ah, 2000), is home to three pale-
flowered species of the genus, suggesting an in¬
crease in species diversity with these floral colors
toward the north of Central America.
The center of diversity of these pale-flow 7 ered
Jacquinia in Mesoameriea is the Yucatan Peninsula
Biotic Province, where all the know n Mesoamerican
pale-flowered taxa of the genus occur. The Yucatan
Peninsula Biotic Province was defined in Duran et
al. (1998) and somewhat differently by Carnevali el
al. (2001) as including the three Mexican states of
Yucatan, Campeche, and Quintana Roo besides the
Peten Department of Guatemala, and the three
northernmost districts of Belize, namely, Belize, Co-
rozal, and Orange Walk. Here (Trejo-Torres A Her¬
nandez-Aguilar, pers. comm.) seven species of Jac¬
quinia occur (including the novelty proposed
Novon 13: 289-295. 2003.
290
Novon
herein), five of which feature blossoms in the hues
of white, pale yellow, or cream-color and are the
subject of this account.
The pale-flowered Jacquinia species in Mesoam-
erica include considerable variation in vegetative
architecture, leaf shape, inflorescence length, (low¬
er shape, and habitat preferences (see Stahl, 1989.
and comments below). For example, Jacquinia ar-
borea and ,/. longifolia have white flowers that are
a smaller version of the orange-red flowers ol ./.
macrocarpa Cavanilles with the free portions ol the
petals (“corolla lobes") diverging from the tube
(which in this case is totally hidden inside the ca¬
lyx) immediately above the calyx lobes. Further¬
more, the staminodes are petal-like and only some¬
what smaller than the corolla lobes, thus giving the
effect of a widely opening, 10-lobed corolla (or with
two series of perianth whorls, one larger than the
internal one). The stamens are exserted. Jacquinia
albiflora has flowers similar in shape to those of the
two taxa above, hut instead of having the obovate,
obtuse leaves found in them, it features narrowly
elliptic leaves, and the staminodes are more petal¬
like.
On the other hand, J. paludicola and J. sak-lol
have a short but definite corolla lube exserted be¬
yond the calyx apex. The stamens are inserted in
the corolla tube. The staminodes are much smaller
than the petals, anther-like, with the apices thick¬
ened, the bases cordate or subcordate, and a gen¬
eral anchor-like outline. They are either inserted in
the corolla mouth or barely protruding. Since there
is an exserted corolla tube and the staminodes are
relatively small and different from the petals, the
flowers have a rotate, 5-lobed corolla.
Thus, the Mesoamerican pale-flowered taxa ol
Jacquinia appear to belong in different species
groups, suggesting that the Moral color feature has
evolved independently several times in the genus.
A cladistic analysis o {Jacquinia would be required
to address the question of a single or multiple or¬
igins of the floral colors found in the genus.
The five Mesoamerican Jacquinia taxa with pale-
colored Ilowers can be distinguished with the fol¬
lowing key:
Key to Mesoamerican Jacqi /am Species with
Paee-Coeoked F powers
la. Leaves obovate or obovate-spathulate, apex
rounded, with or without a terminal spine (most
commonly this lacking).
2a. Young branches densely lepidote; leaves
thick, coriaceous, the margin revolute; inflo¬
rescences as long as to longer than tin*
leaves.2. ./. arborea
2b. Young branches glabrous or puberulent;
leaves thin, thickly membranous, the margin
flat; inflorescences much shorter than leaves
. 3. ./. longifolia
lb. Leaves elliptic or narrowly elliptic, acute, always
with well-developed terminal spines.
3a. Leaves less than 1.5 cm wide, narrowly el¬
liptic; inflorescences composed of (2)3- to
7(9)-flowered racemes; rachis conspicuous;
many branchlets flowering simultaneously;
corolla campanulate, up to 7.5 mm long; sta¬
mens exserted; staminodes petaloid . . .
. 1 . ./. albiflora
3b. Leaves 1.7—4 cm wide, elliptic; inflores¬
cences composed of short 1- to 3-flowered
racemes or several-flowered fascicles; rachis
inconspicuous, always shorter than the pe¬
duncle. few branchlets flowering at any giv¬
en time; corolla campanulate or tubular-ro¬
tate, usually longer than 8.5 mm long;
stamens inserted or barely protruding at co¬
rolla mouth; staminodes ± stamen-like, i.e.,
the apical portion thickened and differently
shaped than the basal portion.
4a. Sparsely branched shrub 0.8—3 m tall;
leaves thickly coriaceous, 4.5—8.5 cm
long, 1.7—3.3 cm wide, margins revo¬
lute; inflorescence composed of a short,
2- to 4-flowered raceme or pseudoum¬
bel; pedicels 7.9-8.1 mm long; corolla
deeply tubular, (7.5—)9—10 mm long;
stamens inserted within the tube ....
.5. J. sak-lol
4b. Much-branched trees 3—10 m tall:
leaves thinly coriaceous, 10—14 cm
long, 3.1-4.5 cm wide, margins flat; in¬
florescence composed of a 4- to 8-How-
ered raceme or pseudoumbel; pedicels
9—11 mm long; corolla campanulate or
shortly tubular, 6 mm long; stamens
barely protruding al corolla mouth . . .
.4. J. paludicola
Synopsis of the. Mesoamerican Jacqu,m\
Species with Pai.e-Coeored Flowers
Jacquinia 1,.. Diss. FI. Jam. App. 1759. TYPE:
Jacquinia ruscifolia L.
1. Jacquinia albiflora Lundell. Wrightia 2: 60.
I960. Jacquinia aurantiaca var. albiflora
(Lundell) Lundell, Wrightia 3: I 14. 1964.
TYPE: Guatemala. Peten: Tikal National Park,
Tikal, in low forest on top of Temple III, 29
July 1959, E. Contreras 45 (holotype, LL not
seen; isotypes, MO photocopy, S not seen, TEX
not seen, LIS).
Among other pale-flowered species in the area
covered, Jacquinia albiflora is easily recognized by
the combination of 3- to 7-flowered racemes pro¬
duced on relatively long peduncles (10-25 mm
long) and the relatively small leaves, these 2.2—
3.8(4.8) cm long (average: 3.26, standard deviation:
0.55, n = 55), 0.6—1.2(1.5) cm wide (average 0.99,
Volume 13, Number 3
2003
Carnevali et al.
Mesoamerican Jacquinia
291
standard deviation: 0.20, n = 55). The relatively
long peduncles allow for inflorescences that are
commonly about as long as to longer than the
leaves. The leaves are elliptic or narrowly elliptic
and are seldom wider than 1.2 cm (rarely up to 1.5
cm wide). The species is endemic to the Yucatan
Peninsula Biotic Province (as defined in Carnevali
et al.. 2001). The flowers are white, eventually fad¬
ing into pale yellow upon aging. Specimens of Jac¬
quinia albiflora are commonly mistaken for ./. ma-
crocarpa Cavanilles, but in ,/. macrocarpa flowers
are red or orange-red and the leaves are much larg¬
er and proportionally wider with a range of dimen¬
sions of (2.9)3.5—5.5(6.5) cm long (average: 4.14,
standard deviation: 0.98, n = 109) and (1.0)1.3-
1.8(2.0) cm wide (average: 1.52, standard devia¬
tion: 0.44. n = 109). Jacquinia albiflora. with its
relatively long inflorescences and many simulta¬
neously opening flowers that are similar to those of
./. macrocarpa. albeit smaller, is most probably re¬
lated to this taxon rather than to other pale-flowered
taxa.
The specimen C. Cowan .1009 (CICY, CSAT,
MEXU), from “Bajo de Moroeoy al K de Nicolas
Bravo, carretera Escarcega-Chetumal, Mexico." is
atypical in displaying larger leaves of a thinner tex¬
ture. The population represented by this collection
is tentatively referred to ,/. albiflora until a more
thorough sampling of its variation is available. It
may represent a different (perhaps undescribed)
species.
Jacquinia albiflora has the following floral di¬
mensions: calyx lobes 1.5—2 nun long, 1.5—2 mm
wide; corolla tube inserted within the calyx 1.5—2
mm long, the corolla lobes ca. 3 nun long, 2 mm
wide; the staminodes are 1.5 mm long and wide;
stamens 1.8—2 mm long, filaments 0.9—1 mm long,
anthers ca. 1 mm long.
Distribution and habitat notes. Jacquinia albi¬
flora has been reported from the Mexican states of
the Yucatdn Peninsula (Campeche, Quintana Boo,
and Yucatan), in the Guatemalan Peten, and in
northern Belize. It is usually a heavily branched
shrub 1—2.5 m tall. It grows in several kinds of
ecosystems, but most collections have been made
in low inundated forests or “tintales,” at elevations
of 0-250 m. Most flowering collections have been
made in April to July, while fruiting collections
have been made from May until November.
Representative specimens examined. GUATEMALA.
Peten: Tikal National Park. 12-15 Apr. 1931. H. H. Bart¬
lett 12594 (A). 24 Apr. 1971. R. Tun Ortiz 1724 (US); in
ramonal covering the mins, 20 Apr. 1968, E. Contreras
7/16 (US); Aguada Naranjal on Arroyo Corriental, 10 I eh.
1959. C. L. Lundell 15461 ((ill); on top of Temple III. 30
Jan. 1964. C. L. Landed 17613 (GH); 4 km NO de Uax-
aetun, 18 May 1973, R. Tun Ortiz 2588 (US); San Cle¬
mente to Dos Arroyos. 1 May 1931. H. H. Bartlett 12821
(A, US). MEXICO. Campe che: Municipio Campeche,
Crucero de P. Trueba y autopista, Ciudad de Campeche,
19°48 15”N, 90°31'30"W, 8 m, selva liaja caducifolia se¬
cundaria, 2 Apr. 2000. C. Gutierrez B. 6802 (CICY.
UCAM). 28 Oct. 2002. G. Carnevali et al. 6557 (CICY.
NY); Municipio Calakmul. 500 m al N del poblado 11 de
Mayo, 18°05'52''N, 89°27'42"W. 4 May 1998. E. Madrid
et al. 813 (CICY. MEXU): Municipio Calakmul, 59 km N
ol Xpujil, camino a Xcan-ha, 18°59'51"N, 89°18'04"YV. 9
Apr. 1998. E. Martinez et al. 30615 (MEXU); 2 km al NE
de Calakmul, 18°27'24"N, 89°47'25''W. 8 Aug. 1997. E.
Martinez et al. 28188 (MEXU); a 6 km al S de la entrada
a Calakmul, 18°26’39 W N, 89°26 , 39 ,, W. 120 msnm, 16 Oct.
1997, E. Martinez S. et al. 28851 (MEXU); a 8 km al NE
de Bel-ha, 18°57'38"N, 89°14'42"W, 9 Apr. 1998, E. Mar¬
tinez S. et al. 30588 (MEXU). Quintana Roo: Municipio
Othdn P. Blanco, Ejido Caobas, aprox. 18°47'18"N,
88°59'47"W, 28 Apr. 1999. G. Carnevali el al. 5433
(CICY, MEXU); Lake Chichankanab, Apr. 1917, F. Can¬
nier 23653 (GH. US). Yucatan: Municipio Valladolid,
parte Sur-este del ejido San Lorenzo perteneciente al
pueblo de Pixoy, 20°42'28"N, 88°15'41"W, 21 Apr. 1983.
E. Ucdn 2410 (CICY).
2. Jacquinia arborea Vahl. Ecol. Amer. 26: 26.
1796. Jacquinia armillaris var. arborea (Vahl)
Grisebach, FI. Brit. West.-bid. Isl. 397. 1861.
I Y PE: Lesser Antilles. Monserrat: J. Ryan s.n.
(lectotype, designated by Stahl (1992: 59). C
not seen).
Jacquinia arborea is a member of the Caribbean
complex of J. armillaris Jacquin, a group of three
closely related taxa (Stahl, 1992). Jacquinia arbo¬
rea is distinctive among pale-flowered taxa in Me-
soamerica due to the combination of broadly ob-
ovate to obovate leaves with revolute margins, and
elongated, racemose inflorescences with 7 to 25(40)
flowers.
Jacquinia arborea has the following floral dimen¬
sions: calyx lobes 2.2-3.2 mm long. 2.5—3.2 mm
wide; corolla tube 4-5 mm long, the corolla lobes
2.8— 4 mm long, 2.5—3.5 mm wide; the staminodes
1.8— 3 mm long, 2-3 mm wide; stamens 2.7—3.8 mm
long, filaments 1-1.8 mm long, anthers 1.7—2 mm
long (StShl, 1992).
Distribution and habitat notes. In Mesoamerica
Jacquinia arborea is only known from three local¬
ities. One of them is the Islas de la Bahia in Hon¬
duras. The second locality is Cozumel Island, in
Quintana Roo State, Mexico (Stahl, 1992). The
third is the mainland at the archaeological site of
Tulum, just across the narrow 7 strait that separates
Cozumel from mainland Quintana Roo (see collec¬
tions below). Outside of Mesoamerica, this species
is known from Jamaica, eastern Hispaniola, Puerto
292
Novon
Rico, the Lesser Antilles, Trinidad and Tobago, and
Curazao and Bonaire (Stahl, 1992).
In the area covered by this paper, Jacquinia ar-
borea has always been collected growing in coastal
dune shrubland, characterized by sclerophyllous
shrubs and with an abundance of succulent herbs.
Jacquinia arborea is usually a shrub or a small tree
3-6 tn tall. It has been collected in flower in No-
vember-December. while mature fruits have been
found in May—June.
Representative specimens studied. MEXICO. (Quin¬
tana Ron: Municipio Cozumel, Isla de Cozumel, 35 km
al S de San Miguel de Cozumel sobre el camino al faro y
la punta Sur, 19 June 1935, E. Cabrera & //. de Cabrera
8709 (MEXU, XAL); Municipio Solidaridad, Tulum Ar¬
cheological Zone, 20°12'48"N, 87°25'45"W, 24 Nov.
1980, ./. /. Calzada et al. 7206 (CICY, XAL); Xel-lla la¬
goon, 20°19'05"N, 87°21 '25"W, 22 Apr. 1985,./. S. Flares
10583 (CICY, XAL).
3. Jacquinia longifolia Standley, J. Wash. Acad.
Sci. 14: 241. 1924. TYPE: El Salvador. San
Vicente: vicinity ol San Vicente, alt. 350—500
m. 2-11 Mar. 1922. I>. C. Standley 21159 (ho-
lotype, US; isotypes, F, G not seen, GH not
seen, MO, NY not seen).
Jacquinia schippii Standley, Publ. Carnegie Inst. Wash.
461(4): 78. 1935. TYPE: Belize. Toledo: Jacinto
Hills, in forest, alt. 270 m, 11 Dec. 1932, 11. A.
Schipp 1222 (holotype, !: isotypes, A. BM, (>, (41.
K, MA. MICH, MO. NY, S, none of the isotypes
seen).
Jacquinia longifolia is easily recognized by the
spathulate, thinly textured leaves with a rounded to
broadly acute apex; terminal spines are present or
absent. The species seems to be extremely variable
regarding size and disposition of the leaves, and
the degree of development of the pubescence found
in the lower third ol the leaves, petioles, and inflo¬
rescences.
This species is similar to Jacquinia paludicola
Standley. Both laxa are shade-loving and have
leaves with flat margins that dry to a pale, dull
green-brown hue. file flowers of the two species are
similar in size and overall morphology and are sim¬
ilarly produced on short racemes or pseudo-umbels
of 4 to 8 flowers, which are much shorter than the
subtending leaves. The stain inodes of ./. paludicola
are, however, more anther-like than in J. longifolia
and are also inserted within the corolla mouth or
barely protruding from it. Among Mesoamerican
taxa of the genus with pale-colored (lowers, Jac¬
quinia longifolia may be closest phylogenetically to
J. arborea because of its obovate, apically rounded
or obtuse leaves and the overall structure of the
flowers. Jacquinia arborea, however, features much
longer inflorescences with many more flowers and
is a plant of coastal habitats.
Jacquinia longifolia has the following floral di¬
mensions: calyx lobes 2—2.5 mm long, 2—2./ mm
wide; corolla tube 2.5—3 mm long, the corolla lobes
2.5—3.2 mm long, 2-2.8 mm wide; staminodes 2—
2.2 mm long, 1.5-2 mm wide at apex; stamens 3—
3.7 mm long.
Distribution and habitat notes. Jacquinia lon¬
gifolia occurs in Mexico (Quintana Boo), Belize.
Guatemala (Peten), El Salvador, Honduras, and
Nicaragua (Stahl, 1989, 2001; Balick et al., 2000).
file species has been collected at elevations rang¬
ing from —5 to 950 m, apparently always in humid
vegetation associations such as tall evergreen for¬
ests. file single collection known from the Yucatan
Peninsula was gathered in a low inundated forest.
Flowering collections have been made in May
through December.
Representative specimens studied. EL SALVADOR.
San Vicente: vicinity of San Vicente, 7—14 Feb. 1947, P.
C. Standley & F. Padilla 2298 ((41. US), 2-1 1 Mar. 1922,
P. C. Standley■ 21742 (GH, US). MEXICO. Quintana
Ron: Municipio Carrillo Puerto, Reserva de la Bibsfera
de Sian Ka’an. 20°50'N, 89°00'W, 28 June 1984. R. Dur¬
an & I. Olmsted 1012 (CIQR, MO. XAL).
4. Jacquinia paludicola Standley, Field Mus.
Nat. Hist.. But. Ser. II: 138. 1932. TYPE: Be¬
lize. Belize: Forest Home, alt. 60 m. I 1 Sep.
1932, W. A. Schipp 1028 (holotype, F not seen;
isotypes, MO, NY, neither isotype seen).
Jacquinia paludicola is closely related to ./. lon¬
gifolia, as discussed above. Both tend to attain larg¬
er sizes than the other three pale-flowered Jacqui¬
nia species known from Mesoamerica, often being
up to 20 m tall in J. longifolia. Jacquinia paludi¬
cola may be collected as an understory shrub or
treelet up to 4 m tall, but some collections indicate
heights up to 7 m. As in ,/. longifolia , the leaves
are thin, but in J. paludicola the leaves are elliptic,
acute, and long acuminate, and of larger sizes, to
14 cm long and 4.5 cm wide. The flowers have been
described by collectors as yellow.
Distribution and habitat notes. Jacquinia pal¬
udicola is known only from the humid portions of
the southern Yucatan Peninsula Biotic Province
ranging into southern Belize and the Izabal De¬
partment of Guatemala (Stahl. 1989, 2001; Balick
et al.. 2000). It has not yet been collected in the
Mexican states of Quintana Ron and Campeche, blit
it is to be expected there. Jacquinia paludicola
grows in tall evergreen forests and has been col¬
lected in flower in June through September; fruiting
Volume 13, Number 3
2003
Carnevali et al.
Mesoamerican Jacquinia
293
material has been sampled in September to Janu¬
ary.
Jacquinia paludicola has the following lloral di¬
mensions: calyx lobes ca. 3 mm long, 2.3—2.5 mm
wide; corolla tube ca. 5 mm long, the corolla lobes
ca. 2.5 mm long. ca. 2 mm wide; staminodes ca. 2
mm long, ca. 1.5 mm wide at apex; stamens 3.4-
3.6 mm long, filaments 2 mm long, anthers 1.5 mm
long.
Representative specimens studied. BELIZE. Toledo:
Blue Creek. 16°2<>'N. 88°45'W, 12 June 1981. C. White-
foord .3217 (MEXU); Joe Taylor Creek. 30 Sep. 1952. R.
H. Gentle 7772 (GH, US). GUATEMALA. Alta Verapaz:
bordering Rio Chiyu, 3.5 km. 28 Sep. 1968. E. Contreras
7753 (US): 6 km from Chahal. 16 Oct. 1968. E. Contreras
7946 (US). Peten: Cerro Tzul. La (Timbre. C. L. Lundell
& E. Contreras 20176 (MEXU); Poptun. en camino a San
Luis, km 117. 7 Jan. 1972. R. Tun Ortiz 2160 (US), 7
Dec. 1970. R. Tun Ortiz 14R4 (US); Cadenas Road, km
142. 3 Nov. 1966. E. Contreras 6515 (US); San Luis, Rfo
Quebrada Seea. 11 July 1959. C. L. Lundell 16364 (till).
Izabal: along Rfo Frio and tributaries, 18 Dei'. 1941. ./.
,4. Steyermark 41537 , 41546 (US).
5. Jacquinia sak-lol Carnevali, Hernandez-Agui-
lar & Tapia-Munoz, sp. nov. T\ PE: Mexico.
Quintana Roo: Municipio Othon P. Blanco,
Ejido Caobas, Sabana del Jaguactal, un desvfo
de 9.5 km por earretera de terracerfa al oeste
de la earretera hacia Pres Garantfas. unos 21
km al sur de la earretera principal Xpuji I—
Chetumal. aprox. 18°18'()()"N. 89°07'00"W, 31
May 2002, F. Lara & R. Santos 6 (holotype,
CICY; isotype, MO). Figure I.
Species haec ./. paludicolae similis sed foliis coriaceis
brevioribus proportione latiore marginibus revolutis, llo-
ribus majoribus crassioribus, tubo proportione longiore
abhorret; J. longifoliae similis sed foliis ellipticis acutis,
marginibus revolutis, Horibus majoribus differt.
Shrubs 0.8— 1,5(—3) m tall; brancldets pubescent;
terminal buds densely covered by scales (perulate);
leaves 4.5—8.5 cm long, 1.7—3.3 cm wide, pseudo-
verticillate on or near branch apex, and at the apex
of next lower brachyblast, coriaceous, elliptic,
acute to (rarely) obtuse or (teratologically) subtrun¬
cate, ending in a stiff mucro 1-1.5 mm long, ba-
sally decurrent, margin revolute, upper surface gla¬
brous. punctate, underside lightly pubescent on the
basal third, particularly over the mid-vein, venation
brochidodromous; mid-vein channeled on the upper
side, prominent on the underside; petiole ca. 1 mm
long; inflorescence terminal or axillar. (1)2- to 4-
flowered, ca. 9—12 mm long; peduncles ca. 5 mm
long; pedicels 7.9—8.1 mm long, glabrous, the flow¬
ers 1—2 mm apart, thus inflorescence appearing
pseudo-umbellate; flowers horizontally patent to
sub-nutant, cream-white or pale yellow-white, ca. 1
cm long, glabrous; sepals 5 mm long, suborbicular
or broadly elliptic, imbricate, margins ciliate, thin;
corolla rotate, the tube 6—7 mm long, lobes ca. 4
mm long and wide; staminodes alternate to corolla
lobes, ca. 2 mm long and wide, transversely oblong
or broadly anchor-like in general outline, apex
broadly obtuse to rounded; stamens inserted within
the corolla tube. ca. 4 mm long, filaments 2 mm;
anthers extrorse, 3 mm long, 1.6 mm wide; stvle
0.5 mm long, persistent, stigma capitate; fruit glo¬
bose, ca. 1.5—2 cm diam., yellow at maturity, sur¬
face rugose; seeds brown, flat, 5.5 mm long. 4 mm
wide.
Etymology. From the Mayan “sak,” white, and
“lol,” flower, in allusion to the white (or nearly
white) flowers of this species. The epithet choice
sak-lol is allowed by Article 23.5 of the Code
(Greuter et al., 2000) as a noun in apposition. The
hyphen is deliberate and supported by Article 60.9.
ex. 17.
Phis distinctive new species is vegetatively sim¬
ilar to some forms of the variable Jacquinia macro-
carpa Cavanilles. A look at the flowers of both spe¬
cies, however, reveals immediately that such
similarity is superficial. Jacquinia rnacrocarpa fea¬
tures bright orange, campanulate flowers, while J.
sak-lol displays cream-white, tubular blooms.
The relationships of Jacquinia sak-lol seem to lie
with J. paludicola. In both species the elliptic
leaves are arranged in pseudo-whorls of 2 to 4
leaves along the stems but mostly close to the apex
of the branches. The pseudo-whorls are separateil
by 5—10 cm on the* branches (up to 15 cm long in
J. paludicola). Both species feature a tubular, rotate
corolla with the staminodes and the stamens in¬
serted within the corolla tube. The staminodes are
anther-like and conspicuously smaller than the co¬
rolla lobes. Both species also feature short-pedun-
cled inflorescences with relatively few flowers.
Jacquinia sak-lol, however, is a shrub (as op¬
posed to a tree) up to 2 m tall. The leaves are
smaller and of a thicker texture. The margins of the
leaf blade are revolute in ./. sak-lol while they are
flat in ./. paludicola. The flowers are larger in ./.
sak-lol with a corolla tube of 6—7 mm long (vs. ca.
5 mm long) and the corolla lobes ca. 4 mm long
and wide, as opposed to the 2.5 mm long, 2 mm
wide corolla lobes of,/. paludicola.
Distribution and habitat notes. Jacquinia sak-
lol occurs in shady spots inside low-inundated for¬
ests (locally known as “tintales” because of the
dominance of Haematoxylum campechianum L.) or
in a mosaic of tall evergreen forests and tintales.
294
Novon
1cm
Figure I. Jacquinia sak-lol Carnevali. Hernandez-Aguilar & Tapia-Munoz. —A. Flowering habit. —B. Flower, lateral
view. —C. Flower tube split lengthwise, showing staminodes, stamens, and ovary; because of some distortion that
occurred in the process of cutting and laying out the tube, the staminodes are not totally alternate with the corolla
lobes. Drawn by Rossana Marrufo based on G. Carnevali, F. May-Pat & M. Gomez 5063 (CICY).
Volume 13, Number 3
2003
Carnevali et al.
Mesoamerican Jacquinia
295
Elevations range from 50 to 250 m. It is a sparsely
branched shrub or small tree that appears to grow
al low population densities. The species has been
collected in flower only in May, while fruiting spec¬
imens have been gathered in August and January.
Paratypes. MEXICO. Quintana Roo: Municipio Oth-
<5n I*. Blanco, 5 km al N lie La Union, alrededores del
rancho Paso del Danto, 17°57'30"N, 88°53'00"W, aprox.
150 msmn. 27 Jan. 1999, P. May el al. 1386 (CICY), 2
Aug. 1992, ,/. Granados 58 (CICY); Ejido Caobas, 26 km
al S de la carretera Xpujil a Chetumal, por la via a Ties
Garantfas, sobre la brecha ca. 3 km al W de la carretera,
ca. 18°13'30”N, 89°04'20"W, 27 May 1998, G. Carnevali
el al. 5063 (CICY).
Acknowledgments. We thank the curators of the
herbaria who loaned us material or allowed access
to their collections. These herbaria include: CI( V )R,
F, MEXU, MO, NY. the Herbarium of the Univer-
sidad Autonoma de Yucatan. UC AM, US, and XAL.
Rossana Marrufo (CICY) elaborated the illustration
of Jacquinia sak-lol. Ginnete Farreny improved on
details of this illustration. Ivon M. Ramirez and Jor¬
ge Carlos Trejo Torres provided discussion and
comments. Filogonio May Pat helped with speci¬
men handling and fieldwork. We are deeply in¬
debted to two anonymous reviewers whose accurate
comments drastically helped improve this article.
The editor of the journal also contributed to the
improvement ol the manuscript, particularly in mat¬
ters of nomenclature. Most of the fieldwork carried
out during this research was funded by CONARIO
(Comision Nacional para el Estudio de la Biodiv-
ersidad grant no. K0.37 to the senior author for the
project “Flora Ilustrada de la Peninsula de Yucatan
Mexicana”) and by CICY (Centro de Investigacion
Cientffica de Yucatan).
Literature Cited
Balick. M. J.. M. H. Nee & D. E. Atha. 2000. Checklist
of the vascular plants of Belize. Mem. New York Bot.
Card. 85: 1-246.
Carnevali, G., ,|. L. Tapia-Munoz, B. Jimenez-Machorro,
E. Sanchez-Saldana, I,. Ibarra-Gonzdlez. I. M. Ramirez
& M. P. Gomez-Juarez. 2001. Notes on the flora of the
Yucatan Peninsula II: A synopsis of the orchid flora of
the Mexican Yucatan Peninsula and a tentative check¬
list of the Orchidaceae of the Yucatan Peninsula Biotic
Province. Harvard Pap. Bot. 5: 383^466.
Castillo-Campos, G. & M. E. Medina Abreo. 1998. A new
species of Jacquinia (Theophrastaceae) from Veracruz,
Mexico. Novon 8: 129—132.
Duran, R., .1. C. Trejo- 1 Torres & G. Ibarra-Manriquez.
1998. Endemic phytotaxa of the peninsula of Yucatan.
Harvard Pap. Bot. 3: 263—314.
Greuter, VC. J. McNeill. F. R. Barrie. H. M. Burdet, V.
Demoulin, T. S. Eilgueiras, D. H. Nicolson, P. C. Silva.
J. E. Skog. P Trehane, N. J. Turland X 1). E. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Stahl. B. 1989. A synopsis of Central American Theo¬
phrastaceae. Nordic J. Bot. 9: 15—30.
-. 1992. On the identity of Jacquinia armillaris
(Theophrastaceae) and related species. Brittonia 44:
54—60.
-. 1993. Votschia, a new genus of Theophrastaceae
from northeastern Panama. Brittonia 45: 204—207.
-. 1995. A synopsis of Jacquinia (Theophrastaceae)
in the Antilles and South America. Nordic J. Bot. 15:
493-511.
-. 2001. Theophrastaceae. In W. I). Stevens, C. Ll-
loa Llloa, A. Pool & (). M. Montiel (editors), Flora de
Nicaragua. Monogr. Syst. Bot. Missouri Bot. Card.
85(3): 2448-2450.
Acer yangbiense (Aceraceae), a New Species from Yunnan, China
Yousheng Chen and Qiner Yang
Laboratory of Systematic and Evolutionary botany. Institute of Botany, Chinese Academy
of Sciences, Xiangshan, Beijing 100093, People’s Republic of China.
maple@ns.ibcas.ac.cn; qeyang@ns.ibcas.ac.cn
Guanghua Zhu
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Guanghua.Zhu@mobot.org
Abstract. Acer yangbiense Y. S. Chen & Q. K.
Yang, a new species from Yangbi County, Yunnan
Province, is described and illustrated. This species
is similar to Acer leipoense Fang & Soong from west¬
ern Sichuan in having a pale gray abaxial surface
of the leaves, a much longer glabrous infruetes-
cence and fruiting pedicels, and convex nutlets, but
differs by its larger 5-lobed leaves with a cordate
base, abaxially densely pubescent veins and vein-
lets, and pubescent young branchlets and petioles.
Key words: Acer ; Aceraceae, China, Yunnan.
In the course of checking specimens of the Chi¬
nese Aceraceae in the herbarium of Sichuan Uni¬
versity (SZ) in 2061, a specimen from Yangbi Coun¬
ty, Yunnan, R. C. Ching 22525, caught our
attention. Fang Wen-Pei identified it as Acer kung¬
shanense Fang on the determinavit slip in 1959,
although this species was not formally published
by him and C. Y. Chang until 1966 (Fang, 1966).
At first glance this specimen is somewhat similar
to A. kungshanense in having leaves abaxially
densely hairy, but differs in having 5-lobed leaves,
a pale gray abaxial surface of the leaves, a glabrous
infructescence, and convex nutlets. In contrast, A.
kungshanense lias 3-lobed leaves, a yellow-brown
abaxial surface of the leaves, a pubescent infruc¬
tescence, and ovoid-globose nutlets. This specimen
is more similar to A. leipoense in having a pale gray
abaxial surface of the leaves, a glabrous infructesc¬
ence, and convex nutlets, but differs by its 5-lobed
leaves, which are abaxially densely hairy along the
veins and veinlets, and thus may represent an un¬
described species. Unfortunately, this specimen is
in poor condition, with the infructescence being in¬
complete, so in April 2002 the first author made an
expedition to Yangbi County in northwest Yunnan
to collect more specimens. A large tree bearing
young fruits was found. Although flowering speci¬
mens were unavailable, some fruits that were still
bearing perianths and anthers were gathered. This
confirmed to the authors that the plant really is a
new species.
Acer yangbiense Y. S. Chen & Q. E. Yang, sp.
nov. TYPE: China. Yunnan: Yangbi County,
Malutang, forests, alt. 2400 m, 24 Apr. 2002,
Y. S. Chen 2010 (holotype, PE; isotypes, MO,
PE). Figures 1, 2.
Species .4. leipoensi atfinis, sed differt foliis majoribus,
10—20 cm longis, I 1-25 cm latis, 5-lobatis, basi cordatis,
nervis primariis 5, nervis et venis subtus densissime pu-
bescentibus.
Deciduous tree, up to 20 m tall; trunk up to 20
cm diam.; branchlets of the present year greenish,
pubescent; branchlets of the previous year brown-
green, [tale gray pubescent; those more than two
years old light brown or dark brown, glabrescent.
with noticeable brown-yellow lenticels; winter buds
ovoid, dark brown, imbricate scales about 9 pairs,
tomentose outside, dropping off soon after the flow¬
ers or leaves have matured. Leaves chartaceous,
10-20 X 11—25 cm, usually broader than long,
shallowly 5-lobed, base cordate, deep green above,
glabrous, pale green beneath, abaxially very dense¬
ly pubescent along veins and veinlets; basal lobes
usually smaller, apex acuminate, or accidentally
absent: middle and lateral lobes triangular-ovate,
apex acuminate, entire or remotely toothed with a
few sinuous teeth; primary veins 5, impressed on
the upper surface, conspicuously prominent below;
central lobe with 7 to 9 pairs of parallel lateral
veins; petioles 4—17 cm long, pale gray pubescent.
Racemes glabrous, pendulous, arising from leafless
lateral buds of 2- or 3-year-old branchlets. Flowers
hermaphroditic, yellow-green; sepals 5, ovate-ob¬
long, ca. 4.5 X 4 mm, yellow-green, glabrous; pet¬
als 5, ovate, yellow-green, base narrow. Intrastam-
inal disk glabrous; stamens 8; filaments ca. 2-3 mm
long; anthers ovoid, ca. 1 mm long; styles 2, base
Nov ON 13: 296-299. 2003.
Volume 13, Number 3
2003
Chen et al.
Acer yangbiense from Yunnan
297
Figure 1. Acer yangbiense Y. S. Chen & Q. E. Yang. —A. Branch with infruetescence. —B. Sepal. —C. Petal. —D.
Bud scale. —E. Pubescence on the abaxial surface of leaves. —F. Details of young fruit with persistent stamens. Drawn
from E S. Chen 2010 (PE).
united, the free portion curved downward; stigmas
2, simple. Infruetescence pendulous, ca. 9—32 X 7
cm; fruits 9 to 17 per raceme, red-green when
young, brown-yellow when mature; wings together
with nutlets 4.7-5.5 X 1.4—1.7 cm, strongly veined,
spreading at acute or nearly right angles; fruiting
pedicels 2.7—3.4 cm long, glabrous; nutlets ca. 7
mm diam., middle convex, globose, villous.
298
Novon
Figure 2. Acer yangbien.se. —Above. Fruiting branch. —Below. Pubescence on the abaxial leaf surface. Photographs
by Yousheng Chen.
Distribution and habitat. This species seems
restricted in distribution. So far it is known only
from its type locality, a valley in the western slope
of Mt. Cangshan, Yangbi, northwestern Yunnan.
Just above this area grows a large forest of Rho¬
dodendron delavayi Franchet. Its population is
small, consisting of no more than 10 trees sparsely
scattered near a small village. It is therefore to be
considered a rare and endangered species. Accord¬
ing the IUCN red list categories and criteria, Ver-
Volume 13, Number 3
2003
Chen et al.
Acer yangbiense from Yunnan
299
sion 3.1 (IUCN. 2001), il should be categorized as
a critically endangered species.
Taxonomic remarks. According to Ogata
(1965), Acer sect. Lithocarpa Pax is characterized
by having dioecious flowers, racemose inflorescenc¬
es from leafless lateral buds, and buds with 8 to 12
pairs of imbricate scales. This section includes a
few species with ranges from the Himalayas to Ja¬
pan. Among them, A. leipoense subsp. leipoense is
found only in Leipo County in Sichuan Province,
and A. leipoense subsp. leucotrichum Fang only in
Baoxing and Tianquan Counties in Sichuan Prov¬
ince. The first author made an expedition to find
these two subspecies in western Sichuan in 2001,
but failed to find specimens of either one. We have
also examined specimens of Aceraceae in the major
herbaria in China and found just two specimens
each of A. leipoense subsp. leipoense and A. lei¬
poense subsp. leucotrichum. To our knowledge, the
two subspecies of A. leipoense are also critically
endangered. The two subspecies of A. leipoense and
A. yangbiense can be distinguished from the re¬
maining species in section Lithocarpa Pax by hav¬
ing a pale gray abaxial surface ol the leaves, convex
nutlets, and a much longer infruetescence and
fruiting pedicels.
Acer yangbiense is similar to A. leipoense subsp.
leipoense in having a pale gray abaxial surface of
the leaves, a long glabrous infruetescence and fruit¬
ing pedicels, and convex nutlets, but differs by its
larger 5-lobed leaves very densely pubescent abax-
ially along veins and veinlets, and pubescent young
branchlets and petioles. Acer leipoense subsp. leu¬
cotrichum is different from the new species by its
much smaller 3-lobed leaves (9—1 1 X 11—12 cm)
and nutlets (3.5-4.3 X 0.8—1.3 cm), and shorter
fruiting pedicels (1.8—3 cm). The leaf bases of both
subspecies of A. leipoense are rounded, while those
of A. yangbiense are cordate. The leaves of A. lei¬
poense subsp. leucotrichum are abaxially only
sparsely pubescent and the hairs cannot be felt
with fingers, while the leaves of Acer yangbiense are
abaxially so densely pubescent that the hairs can
easily be felt. Thus, A. yangbiense can be distin¬
guished from A. leipoense subsp. leucotrichum by
the pubescence character.
fhe circumscription of Acer sect. Lithocarpa
made by de Jong (1994) is almost the same as that
made by Ogata (1965) except that de Jong treated
A. macrophyllum Pursh in this section as a mono-
specific series, series Macrophylla Pojarkova ex
Momotani, while Ogata treated it as an independent
section, section Macrophylla (Pojarkova) Ogata.
Acer yangbiense is a member of Acer sect. Litho¬
carpa ser. Lithocarpa by having racemes arising
from leafless lateral buds and about nine pairs of
imbricate scales on the buds.
Paratype. CHINA. Yunnan: Yangbi, Ma-lu-tang, 3
May 1929. R. C. Ching 22525 (SZ).
Key to A. yanghikn.se, A. leipoense, and A. kuncshanense
la. Abaxial surface of leaves yellow-brown; infruc-
tescence 7-9 cm long, glabrous; fruits 4 to 7 per
raceme; nutlets globose. Acer kurigslumen.se
lb. Abaxial surface of the leaves pale gray; infruc-
tescence 10—32 cm long. ± pubescent; fruits 9
to 17 per raceme; nutlets convex.
2a. Leaves 9—1 1 X 11 — 12 cm, sparsely pubes¬
cent abaxially along veins and veinlets (hairs
cannot be felt with fingers); leaf blade 3-
lobed, base rounded; nutlets 3.5—4.3 X 0.8—
1.3 cm . Acer leipoense
2b. Leaves 10-20 X I 1-25 cm. densely pubes¬
cent (hairs can be easily felt with fingers);
leaf blade 5-lobed, base cordate; nutlets
4.7—5.5 X 1.4—1.7 cm. Acer yangbiense
Acknowledgments. We thank two anonymous
referees for their valuable comments, anti Sun
Yingbao for his illustration.
Literature Cited
Fang, W. P. 1966. Revisio taxorum Aceracearum sinica-
rum. Acta Phytotax. Sin. 11(2); 139—189.
IUCN. 2001. IUCN Red List Categories and Criteria: Vers.
3.1. IUCN Species Survival Commission. IUCN. Gland,
Switzerland, and Cambridge, U.K.
Jong, P. C. de. 1994. Taxonomy and reproductive biology
of maples. Pp. 69—103 in I). M. van Gelderen. P. C. de
Jong, H. J. Oterdoom & J. R. P. van Hoey Smith (edi¬
tors), Maples of the World. Timber Press, Portland, Or-
egeon.
Ogata, K. 1965. A systematic study of the genus Acer.
Bull. Tokyo Univ. Forest 63. 89—205.
Sterculiaceae (Byttnerieae) Nuevas de Peru
Carmen L. Cristobal
Instituto de Botanica del Nordeste, C.C. 209, 3400 Corrientes, Bep. Argentina.
ihone@agr.unne.edu.ar.
RESUMEN. Se describen e ilustran dos Sterculi-
aceae nuevas del norte de Peru: Ayenia peregrina
y Byttneria sagasteguii. Se comparan ambas nove-
dades con las espeeies afines. Ayenia peregrina cor-
responde a la seccion Leiayenia. Se llama la aten-
ci6n sobre el hallazgo de un representante de esta
seccion en Sudameriea, ya que el area conocida
abarcaba el centro sur de los Estados Unidos de
America, Centroamerica y las Anti lias Mayores,
Byttneria sagastegui pertenece a la seccion Inea-
sica.
Abstract. Ayenia peregrina and Byttneria sagas¬
teguii, two new species from the north of Peru, are
described, illustrated, and compared to related spe¬
cies. The remarkable finding of A. peregrina is dis¬
cussed because it belongs to Ayenia sect. Leiayenia,
known up to now from the south-central U.S.A.,
Central America, and the Greater Antilles. Bytt¬
neria sagasteguii belongs to section Incasica.
Key words: Ayenia, Byttneria, Peru. Sterculi¬
aceae.
Ayenia peregrina Cristobal, sp. nov. TIPO: Peru.
Amazonas: along roail Pedro Ruiz—Chachapo-
yas, in dry scrub, with cacti and bromeliads,
1300—1900 in, I I Mar. 1998. //. van der Werff,
B. Cray, R. Vdsquez & R. Rojas 14669 (liolo-
tipo, MO; isdtipos, CTES, F. MO. NY). Figu¬
re 1.
Suffrutex ca. 1 m altus pilis stellatis dense tectus. Folia
ovata-lanceolata dentata discolora subtus cinerea. Cimae
axillares breves triflorae. Petala longissime unguieulata
cueullo subtrapeziforme apice integro dentibus duobus in-
lus instrueta; lamina petalorum nu 1 la; androgynophorus
nullus; tubus staminalis urceolalus antheribus trilocula-
ribus staminodiis cucullatis. Fructus ca. 4 mm latus acu-
leolis brevissimis et pilis stellatis tectus.
Subarbusto de 0.90—1 m alt., talk) lenoso con
corteza grisdcea, delgada, finamente estriada con
lenticelas redondeadas, ramas con una alternancia
de porciones cicatricosas, desnudas y entrenudos
muy cortos, con porciones con entrenudos largos de
1—1.5 cm long., los extremos florfferos y hojosos
amarillo-tomentosos, cubiertos de una gruesa capa
de pelos estrellados, intrincados, con numerosos ra¬
dios, sesiles o largamente estipitados, estipite de
ca. 0.5 mm long., castano, el extremo fragil. Estf-
pulas linear-subuladas, caedizas, 1.5—3 mm long.,
tomentosas. Peciolo terete, 1—1.5 cm long., con el
mismo indumento del tallo. Lamina discolora, ova-
do-laneeolada, base cordada, apice agudo, 3—5 X
1.5-2.8 cm, margen dentado, en las hojas mas vie-
jas el apice de los dientes glabro, papiloso v oscuro.
epifilo gris-verdoso. tomentoso, hipofilo cinereo, in¬
dumento mas denso que en el epifilo y con pelos
mas largos, sobre las venas pelos estipitados, venas
basales 7—9. desde la vena media 3—4 pares de
venas laterales mas o menos rectas y sobresalientes
hasta el margen. Alabastros globosos. Flores reu-
nidas en cimas axilares abreviadas, 3-floras, pe-
dunculo y pedicelos 3—7 mm long., ambos mas lar¬
gos en el fruto, braetdolas 1—1.5 mm long.,
subuladas, tomentosas, agrupadas en el extremo del
pedunculo. Sepalos 2.7—3 X 1.5—2 mm, soldados
cerca de la base, ovales. abruptamente afinados en
el 1/3 apical, cara externa tomentosa, pelos estre¬
llados sesiles y brevemente estipitados, cara
interna con la vena media muy sobresaliente, pelos
glandulares breves disperses. Petalos rojizos, gla-
bros, opuestos a los estambres, largamente ungui-
culados, lamina nula, porcidn basal de la una lili-
forme de 5 mm long., capucha subtrapezoidal,
adosada al horde del tubo estaminal, ca. 1.5 X 2
mm, ldbulos con el margen distal eroso, cara ex¬
terna con la mitad apical ampollada, extremo prox¬
imal redoutleado con un diminuto mucrdn oscuro
en el centro, y con dos dientes incurvados que
abrazan al filamento estaminal, cara interna con
una leve concavidad que aloja a la antera. Andro-
gindforo nulo. Tubo estaminal urceolado, 1 mm alt.,
glabro, anteras tritecas en el extremo de una corta
porcidn libre de filamento, estaminodios cuculados,
margen externo con un pequeno apfculo medio.
Ovario mas o menos globoso, eubierto de mamilas,
estilo exerto, estigma redondeado. Gapsula globosa,
ca. 4 mm diam., a la madurez disgregada en 5 co¬
cos uniseminados dehiscentes a lo largo de la su-
tura y de parte de la vena media dorsal, mitades
curvas separadas entre sf, cara externa cubierta de
aculeolos cilfndricos, hasta 1/2 mm long., oscuros,
Novon 13: 300-304. 2003.
Volume 13, Number 3
2003
Cristobal
Sterculiaceae de Peru
301
Figura 1. Ayenia peregrina Cristobal. —A. Kama. —B. Capucha del petalo y parte de la una, eara externa. —C.
Capucha del petalo, eara interna. —D. Porcidn del tubo estaminal, visto desde arriba y gineeeo. —E. Hor: parte del
caliz sin el indumento, 1 petalo separado del tubo estaminal visto de frente, tubo estaminal y gineeeo. —F. Coco
debiscente. —G. Semilla, vista lateral. A. F, G: dibujado del tipo van der Werff el al. 14669 (CTFS). B—E: Vdsquez el
al. 25942 (CTFS).
302
Novon
con pelos estrellados, tambien pelos estrellados en¬
tre los aculeolos, cara interna lustrosa, diminutos
pelos glandulares rojizos dispersos y escasos pelos
largos, simples. Semillas angostamente ovoides, ap-
ice conico, 2.5 X 1 .5 mm, castanas o negras, caras
laterales rugosas mas o menos planas, dorso con-
vexo tambien rugoso. pequenas verrugas dispersas
en toda la superficie.
Ayenia peregrina pertenece a la seccion Ijeia-
yenia Grisebach caracterizada por los petalos sin
lamina, constituidos solo por la capucha con dos
dientes incurvados en el extremo proximal, que
abrazan la port ion libre del filamento estaminal.
Como se senald en trabajos anteriores (Cristobal,
I960, 1961, 1962), las especies de esta seccion
tlilieren notablemente entre sf en el aspecto vege-
tativo y en diversos caracteres florales, en contra-
posicion a las otras secciones del genero donde con
frecuencia las diferencias interespeefficas son muy
finas. A1 comparar las flores de A. peregrina con
las de las especies afines, vemos que, curiosa-
mente, las mas semejantes son las de A. micro-
phylla A. Gray, especie que ocupa el extremo norte
del area de la seccion. Se diferencian sin embargo
porque en A. peregrina la capucha de los petalos
es ampollada, de mayor tamano y con los lobulos
erosos. En A. microphylla la capucha posee en el
margen proximal una superficie plana prolongada
en un notable apfeulo retrorso hacia la cara exter¬
na. Otra diferencia es el androginoforo, nulo en A.
peregrina y aunque breve, [)resente en A. micro¬
phylla. Finalmente, el tamano de la hojas y el in¬
dumenta, que hace que las de A. peregrina sean
discoloras, las diferencian a simple vista.
Con este hallazgo ascienden a 12 las especies
de la seccion Leiayenia. En el hemisferio norte A.
microphylla vive en el centro-sur de los Estados
Unidos de America y NE de Mexico, cinco especies
son exclusivas de Mexico y cinco viven en las An-
t iI las Mayores. El hallazgo de un miembro de esta
seccion en los Andes del !N de Peru, es un ejemplo
particularmente interesante para el estudioso del
origen y de las relaciones de la flora peruana. Ya
Weberbauer (1945: 657). en su capftulo sobre la
“Genesis de la Flora Peruana”, llama la atencion
sobre la disyuncion del area de especies xeroffticas
comunes entre el sur de los Estados Unidos y Mex¬
ico y territories interandinos de Peru.
Ea seccion Leayenia era la unica del genero cuya
area no abarcaba Sudamerica, ya que A. rnagna I,.,
que se extiende desde el Caribe hasta el NW de
Paraguay, representa a la seccion Cyhiostigma (Tur-
czaninow) Grisebach. La seccion Ayenia esta difun-
dida en ambos hemisferios.
Pardtipo. PERU. Amazonas: Ragua Distrito, Pongo
de Rentema. Bosque del valle seco del Maranon, 05 Q 30’S,
78 U 30'W, 400 m, 4 Feb. 1999, A“. Vdsquez, C. Diaz. F. Sal¬
vador el al. 25942 (CTES, F, MO. NY).
Byttneria sagasteguii Cristobal, sp. nov. TIPO:
Peru. Cajamarca: Chota, Tucupampa (Llama-
Huambos), ladera, 2500 m s.m., sufrutice con
flores vinosas, 17 Mar. 1997, A. Sagastegui A.,
S. Leiva G. A' V Quipuscoa 15947 (holdtipo. F;
isotipos, CTES, HUT). Figura 2.
Frutex caulibus pentagonis in costis parvis aeuleis rec-
tis 0.5—2 mm longis. Lamina ovata-lanceolata vel lateova-
ta supra pilis setiformibus antrorsis subtus tomentosis pi-
lis simplicibus. Inflorescentia nuda ramosa 3—5 cm longa.
Unguis petalorum membranaceus alis rotundis; lamina
leviter pubescens in dimidio superiore incrassata ad api-
cem acuta; antherae biloculares; staminodia membranacea
in latere extemo prominentia acuta.
Arbusto de ca. 2 m alt. (Sagastegui A., comuni-
eacion personal), aparentemente tortuoso, setuloso,
pelos simples, largos, gruesos distribuidos en todos
sus organos. Tallo, cerca de los extremos flonferos,
8—13 mm diam., macizo o fistuloso en los entren-
udos, con 5 caras planas demarcadas por costillas
colenquimatosas sobresalientes, amarillentas, con
pequenos aguijones rectos, fragiles, densamente
dispuestos, no alineados entre sf, de 0.5—2 mm
long., punzantes y terminados en 1 pelo, cerca del
apice de las ramas setas dispersas, sobre las cos¬
tillas y escasas en las caras. Estfpulas fugaces, fi-
namente subuladas, ea. 4 mm long., cara interna
glabra, cara externa pilosa. Pecfolo 12-25 mm
long., inerme, con los mismos pelos del tallo. La¬
mina del extremo de las ramas ovado-lanceolada,
base redondeada o subcordada, apice agudo, en ra¬
mas mas viejas anchamente ovada, base eordada,
apice agudo, margen serrado, etitero cerca de la
base, epifilo uniformemente cubierto de setas an-
trorsas, que dejan ver la superficie de la epidermis,
hipofilo densamente cubierto de pelos setiformes
especialmente sobre las venas, mas densamente
dispuestas y desordenadas, venacion poco notable
en el epifilo v sobresaliente en el hipofilo, desde la
base parten 2—3 pares de venas laterales y desde
la vena media 5—6 pares de venas enfrentadas mas
o menos rectas y paralelas hasta el margen, nec-
tario ubieado en la base de la vena media, desta-
candose solamente por ser una porcion, mas oseura,
algo mas ancha y con escaso indumenta. Flores
reunidas en inflorescencias tirsoideas desnudas,
axilares y terminales de 3-5 cm long., el eje con
cerca 5 nudos origina en cada uno de ellos 3—5
eimas umbeliformes plurifloras de 1.5-2 cm long.,
en la base de cada cima un verticilo de bracteolas
rojizas, membranaceas, soldadas en la base, laci-
Volume 13, Number 3
2003
Cristobal
Sterculiaceae de Peru
303
Figura 2. Byttneria sagasteguii Cristobal. —A. Kama. —B. Porci6n de tallo. —C. Petalo cara interna. —D. Petalo
3/4 perfil. —E. Flor mostrando 1 sepalo, 1 petalo visto de frente con la capucha algo abierta. tubo estaminal y gineceo.
Dibujado del tipo Sagdstegui A. et al. 15947 (CTES).
niadas, pedicelo 4—5 mm long. Sepalos morados,
4—5 X 2.3—2.5 mm, soldados en el tercio basal,
lobulos ovados, cara interna con pelos glandulares
diminutos dispersos, cara externa con pelos sim¬
ples, largos, dispersos. Petalos 3—5 mm long., di-
vididos en dos porciones de aproximadamente el
mismo largo: la capucha amarillenta, glabra, mem-
branacea, con las alas muy pequenas, redondeadas,
curvadas levemente hacia arriba, y la lamina con
escasos pelos cortos dispersos, la mitad basal pla¬
na, membranacea, amarillenta. y la mitad apical,
morada, aneha, carnosa, algo aplastada, aguda.
Tubo estaminal campanulado, glabro, membrana-
ceo, 1.3—1.5 mm long., amarillento excepto una
mancha morada en la cara externa de los estami-
nodios, estambres con una portion muy breve de
304
Novon
filamento, anteras algo divergentes, ditecas; esta-
minodios membrandceos, margen con un apfculo
central y un pequeno lobulo redondeado y algo re-
curvado a cada lado, cara externa con una promi-
nencia aguda sobre la linea media. Gineceo ca. 1.5
mm long., sobresaliendo levemente del tubo esta-
minal, ovario papiloso, lobulos del estigma del mis-
mo ancho (|ue el eslilo, redondeados. Fruto no vis-
to.
Byttneria sagasteguii pertenece a la seccion In-
casica Cristobal (Cristobal, 1976), ya que los pe-
talos se apoyan en el borde del tubo estaminal, los
estaminodios son membrandceos con una promi-
nencia aguda en la cara externa y los tallos tienen
costillas sobresalientes con aguijones.
Dentro de la seccion, esta nueva especie se di-
ferencia por la siguiente eombinacion de caracter-
es: tallos con pequenos y numerosos aguijones rec¬
tos no alineados entre si, hojas densamente
cubiertas de largos pelos setiformes, lamina de los
petalos con la mitad distal abultada, carnosa, mo-
rada y con escasos pelos dispersos e inflorescencia
amplia, tirsoidea, desnuda, tanto axilar como ter¬
minal. Si nos esforzamos en encontrar alguna es¬
pecie afin se puede comparar con B. flexuosa Kil-
lip, especie de Ecuador, pero los caracteres arriba
mencionados las separan facilmente.
En la revision de Byttneria Loefling (Cristobal,
1976) se incluyeron I I especies en la seccion In-
casica. Posteriormente Dorr (1997, 1999), adiciona
una especie nueva de Ecuador: B. jaramilloana
Dorr y pasa a la sinonimia de B. parviflora Bentham
a B. glabrescens Bentham.
Agradecimientos. Gracias a los curadores de
los herbarios consultados por el envfo del material,
y al Dr. Abundio Sagastegui Alva, de la Universi-
dad Privada Anterior Orrego, de Trujillo, quien co-
lecciono la Byttneria que lleva su nombre, por da-
tos adicionales sobre la especie. Gracias tambien a
Daniel Cian, por dibujar las ramas de las dos fi-
guras y por pasar en tinta los detalles florales di-
liujados por mi misma.
Literature Citada
Cristobal, C. L. I960. Revisidn del genero Ayenia (Ster¬
euliaceae). Opera Lilloana 4: 1—230.
-. 1961. Nueva contribucidn al estudio del gdnero
Ayenia L. (Stereuliaceae). Anales lust. Biol. Univ. Nac.
Mexico 32(1-2): 191-200.
-. 1962. Notas sobre el genero Ayenia. Lilloa 31:
207-212.
-. 1976. Estudio taxondmico del genero Byttneria
Loefling (Stereuliaceae). Bonplandia 4: 1—428.
Dorr, L. J. 1997. A new species of Byttneria (Stereuli¬
aceae) from Ecuador. Biollania Ed. Esp. 6: 305—309.
-. 1999. Stereuliaceae. In P M. Jorgensen & S.
Ledn-Yanez (editors), Catalogue of the Vascular Plants
ol Ecuador. Monogr. Syst. Bot. Missouri Bot. Card. 75:
918-922.
Weberbauer, A. 1945. El Mundo Vegetal de los Andes
Peruanos. Ed. 2. Lima, Peru.
Eleutherococcus humillimus , a New Combination
in Ch inese Araliaceae
Deng Yunfei
South Cliina Institute of Botany, Chinese Academy of Sciences, Guangzhou,
Guangdong, 510650, People’s Republic of China, yfdeng@scib.ac.cn
ABSTRACT. A new combination, Eleutherococcus
humillimus (Y. S. Lian & X. L. Chen) Y. F. Deng,
is made.
Key words: Acanthopanax, Araliaceae. China.
Eleutherococcus.
Eleutherococcus Maximowicz (1859) is the cor¬
rect name for the genus previously widely referred
to as Acanthopanax (Decaisne & Planchon) Witte
(1861). Nomenclatural problems involving these
genera have been discussed by several authors over
the years (Hu. 1980; Ohashi, 1987; Hsu & Pan,
1993; Zhu et al., 1995). Eleutherococcus has about
25 species distributed mainly in northeastern Asia,
including China, Korea, and Japan, with some
members extending to southwestern Asia (Kim &
Sun, 2000).
Acanthopanax was recognized at the subgeneric
rank under Panax L. by Decaisne and Planchon
(1854), and was later raised by Witte (1861) to ge¬
neric rank. Harms (1897: 49) combined both under
one genus and adopted the name Acanthopanax,
which has been widely used since by many bota¬
nists (Harms. 1918; hi. 1942; Hoo & Tseng, 1978;
Philipson, 1979; Shang, 1985a, 1985b). However,
the name Acanthopanax does not have priority un¬
der the current ICON (Greuter et ah. 2000), and
several recent authors (e.g.. Hu. 1980; Stone, 1980;
Ohashi, 1987: Hsu & Pan. 1993; Zhu et ah, 1995)
have accordingly transferred taxa from Acanthopan¬
ax to Eleutherococcus.
In 1918 Harms classified the genus into seven
sections, namely, sect. Eleutherococcus, sect. Ce-
phalopanax, sect. Euacanthopanax, sect. Zanthox-
ylopanax, sect. Evodiopanax, sect. Sciadophyllo-
ides, and sect. Kalopanax. This system has been
accepted by many authors (Hoo & Tseng, 1978;
Shang, 1985a, 1985b; Ohashi, 1987), but section
Kalopanax, typified by K. pictus (Thunberg) Nakai,
is generally treated as a distinct genus. Several
years later, Nakai (1924) elevated section Evodi¬
opanax to generic rank, then Shang et ah (2000)
synonymized it with Gamblea C. 8. Clarke and
made four new' combinations. Shang and Huang
(1993) raised section Sciadophylloides to generic
rank as Chengiopanax C. B. Shang & .1. Y. Huang
with two species, C.fargesii (Franchet) C. B. Shang
& J. Y. Huang and C. sciadophylloides (Franchet &
Savatier) C. B. Shang & .1. Y. Huang. R ecognition
of the unarmed genera Gamblea and Chengiopanax
as distinct from Eleutherococcus, which has spines
or prickles, was recently confirmed using ITS se¬
quence data (Wen et ah, 2001).
Acanthopanax humillimus Y. S. Lian X X. L.
Chen was described by Lian and Chen (1994)
based on collections from Gansu Province, China,
but the necessary combination in Eleutherococcus
is made here.
Eleutherococcus huuiilliiiius (Y. S. Lian & X. L.
Chen) Y. F. Deng, comb. nov. Basionym: Acan¬
thopanax humillimus Y. S. Lian & X. L. Chen.
Acta Bot. Boreal.-Occid. Sin. 14(6): 76. 1994.
TYPE: China. Gansu: Lintan Xian, “Lianhua
mountain, in locis udis regionis in sylvis,”
2700 m, 9 July 1994, Y. S. Lian et al. 94065
(holotype, NWTC).
Distribution. A rare species endemic to south¬
ern Gansu and northern Sichuan Provinces of Chi¬
na, in alpine habitats at high elevations (2700-
3000 m).
Eleutherococcus humillimus most closely resem¬
bles E. giraldii (Harms) Nakai but differs in being
a subshrub, only 5-15 cm tall (vs. 1-3 m), with 3
or 4 (rarely 5) ovaries and styles (vs. 5 ovaries and
styles), the latter connate for ca. 1/5 their length
(rather than 1/4—1/2). Eleutherococcus humillimus
occurs at high elevations and may be an alpine
vicariant of E. giraldii.
Additional specimens examined. CHINA. Sichuan:
Hongyuan Xian, Shuajingsi, Yakou, alt. 3450 in. in ravine,
8 Oct. 1957, Li Xing 72410 (IBSC, SZ); Hongyuan Xian.
Shuajingsi, Yakou, all. 3400 m, 9 Oct. 1957, Li Xing
72444 (IBSC, SZ); Nanping Xian, Jiuzhaigou, 3200 in, iri
Picea forest, 25 Sep. 1986, Wu Zhengyi et al. HI I (FUN);
Nanping xian, Jiuzhaigou, Changhai, 2800 m. in Picea
forest, 26 Sep. 1986, Wu Zhengyi et al. 982 (KI N); Nanp¬
ing Xian, Jiuzhaigou, in forest, 25 Sep. 1986. Wu Zhengyi
el al. 1110 (KUN): Zoige (Ruoergai) Xian, Baxi gongshe.
Novon 13: 305-306. 2003.
306
Novon
Yangyugou, in Picea forest, alt. 3100 rn, 28 June 1975,
Chengdu Biological Institute 10063 (IBSC). Gansu:
Dangchang xian, Daheba, in subalpine shrubby, alt. 3000
m, 24 Sep. 1979, Lion Yongshan 790922 (NWTC); Tewo
xian, Lazikou Linchang, in the forest, on the rock, alt.
3000 m. 24 July 1980, Lion Yongshan et al. 300222
(NWTC).
Acknowledgments. I thank Nicholas I in land
(MO) for editorial help with the manuscript. I am
grateful to two anonymous reviewers for their com¬
ments. This work was partly supported by a grant
from the Director Foundation of the South China
Institute of Botany, CAS.
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Nijhoff and Dr. W. Junk Publishers, The Hague.
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Malaysian Forester 43: 395.
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Evolution of Araliaceae: A phylogenetic analysis based
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167.
W itte, H. 1861. Les Araliaeees des jardins Hollandais.
Ann. Hort. Bot. 4: 87-95.
Zhu, X. Y., S. Ge & D. Y. Hong. 1995. A critical note on
the scientific name Acanthopanax Miq|.| and several
new combinations. Bull. Bot. Res., Harbin 15: 441 —
443.
Transfer of Madagascan Glochidion to Phyllanthus
(Euphorbiaceae s.l. or Phyllanthaceae)
Petra Hoffmann
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 SAB,
United Kingdom, p.hoffmann@rbgkew.org.uk
Gordon McPherson
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.
gordon.mcpherson@mobot.org
Abstract. Seven endemic Madagascan species
previously assigned to the genus Glochidion J. H.
& G. Forster by Leandri are re-examined morpho¬
logically and found to belong to Phyllanthus L. (Eu¬
phorbiaceae s.l. or Phyllanthaceae). As a conse¬
quence, the following five new combinations are
made: Phyllanthus ankaratrae (Leandri) Petra
Hoffmann & McPherson, P. humbertii (Leandri) Pe¬
tra Hoffmann & Mc Pherson, P. lichenisilvae (Lean-
dri) Petra Hoffmann McPherson, P. marojejiensis
(Leandri) Petra Hoffmann & McPherson, and P.
perrieri (Leandri) Petra Hoffmann & McPherson.
Key words: Euphorbiaceae, Glochidion. Mada¬
gascar. Phyllanthus.
The Euphorbiaceae—Phyllanthoideae volume of
the Flore de Madagascar et des Comores (Leandri,
1938) treated seven species in the genus Glochi¬
dion J. R. & G. Forster, distinguishing them from
Phyllanthus L. by their reduced, or small and en¬
tire styles. Leandri described the first of these sev¬
en species, G. perrieri , in 1934 and placed it in
section Heteroglochidion Mtiller Argoviensis. This
section is typified by the New Caledonian species
G. baladensis (Baillon) Muller Argoviensis, which
has long, free styles and is today again considered
a species of Phyllanthus (Schmid & McPherson.
1991). In 1937, Leandri described G. lichenisilvae
and transferred two of his own Phyllanthus species,
P- oreichtitus and P. sambiranensis , to Glochidion.
Three further species, G. ankaratrae, G. humbertii.
and G. marojejiense, were described in 1937 (Lean-
dri. 1957).
Glochidion and Phyllanthus are both members of
suhtribe Flueggeinae (tribe Phyllantheae), the larg¬
est and most difficult group in subfamily Phyllan¬
thoideae. or family Phyllanthaceae sensu APG (e.g.,
Savolainen et al., 2000, Chase et ah. 2002). The
last complete treatment of this group at species lev¬
el was published by Muller Argoviensis (1866),
who combined Glochidion with Phyllanthus. Ben-
tham and Hooker (1880) followed Muller in this but
most later authors (e.g.. Hooker, 1887. Pax & Hoff¬
mann. 1931, Webster, 1994, Radcliffe-Smith, 2001)
have regarded Glochidion as a separate genus.
Phyllanthus is an extremely diverse pantropical
group of ea. 800 species that may not be mono-
phyletic (Webster, 1994, Wurdaek et ah, in prep.).
Molecular analyses are under way as a collabora¬
tion between the University of Vienna and the Roy¬
al Botanic Gardens, Kew. to shed light on the po¬
sition of Glochidion , which Webster (1994: 46)
considered to "represent a specialized offshoot from
ancestral laxa within Phyllanthus." Glochidion is
generally recognized as being restricted to Asia and
Australasia (Webster, 1994), sometimes with the ex¬
ception of the Madagascan species (Radcliffe-
Smith. 2001).
Webster (1994) and Radcliffe-Smith (2001) dis¬
tinguished the two genera mainly on the basis of
the following characters: floral disc (usually present
in Phyllanthus , totally absent in Glochidion) and
seed morphology (dry and not ventrally invaginated
in Phyllanthus. usually fleshy (with sarcotesta) and
ventrally invaginated in Glochidion). Stuppy (1996)
confirmed the* importance of seed characters in sep¬
arating the two taxa.
Examination of the material in Paris (including
all types) showed that all seven Madagascan spe¬
cies must be referred to Phyllanthus on the basis
of both disc and (where available) seed characters.
Brunei (198/) was the first to come to this conclu¬
sion when examining Glochidion oreichtitum and G.
sambiranense. He found the pollen of these two spe¬
cies to be tricolporate whereas Glochidion has ste-
phanocolporate pollen (Webster, 1994). Brunei sug¬
gested an affinity with Phyllanthus subg. Kirgatielia
for the species with free stamens, and with Phyl¬
lanthus subg. Gomphidium for the species with con-
Novon 13: 307-310. 2003.
308
Novon
nate androecia. He did not, however, study any of
the species without valid names in Phyllanthus.
While Glochidion oreichtitum and G. sambira-
nense were previously published as Phyllanthus
and have valid combinations in that genus, the re¬
maining five species have no valid names outside
Glochidion. We were first presented with this prob¬
lem when identifying Euphorbiaceae collections
(cited below under the respective species) made
during the recently published botanical inventory
(Gautier, 2002) of the Reserve Speciale de Man-
ongarivo by the Conservatoire et Jardin Botaniques
de la \ ille de Geneve (G). We again encountered
the problem while reviewing the Euphorbiaceae
chapter in the Generic Tree Flora of Madagascar
(Schatz, 2001), and when preparing an overview of
the Euphorbiaceae of Madagascar for the Natural
History of Madagascar (Hoffmann & McPherson,
2003).
To correct the false impression that the Asian
and Australasian genus Glochidion is present in
Madagascar, we propose the following new combi¬
nations. This paper is intended to clarify the ge¬
neric boundaries and to make information on the
species concerned available to researchers outside
the Paris Herbarium. Where necessary, lectotypi-
fication will he undertaken in the course of a post¬
graduate study currently in progress at the Univer¬
sity of Antananarivo and the Royal Botanic
Gardens, kew. that will result in a full taxonomic
revision of Madagascan Phyllanthus.
Phyllanthus ankaratrae (Leandri) Petra Hoff¬
mann & McPherson, comb. nov. Basionym:
Glochidion ankaratrae Leandri, Mem. Inst.
Sci. Madagascar. Ser. B, Biol. Veg. 8: 214.
1957. TYPE: Madagascar. Environs
d’Ambositra, foret de Ranomena, //. Humbert
& C. F. Swingle 4845 (MO, P).
known from the type only.
Pliyllanthus Iiiinihcrlii (Leandri) Petra Hoffmann
& McPherson, comb. nov. Basionym: Glochi¬
dion humbertii Leandri, Mem. Inst. Sci. Mad¬
agascar, Ser. B. Biol. Veg. 8: 215. 1957.
TYPE: Madagascar. Pentes orientales du Mas¬
sif du Marojejy (Nord-Est), a 1’ouest de la ri¬
viere Manantenina, affluent de la Lokoho, foret
ombrophile sur laterite de gneiss. ± 1450 m.
H. Humbert 22495 (MO, P).
known from the type only.
Phyllanthus lichenisilvae (Leandri) Petra Hoff¬
mann & McPherson, comb. nov. Basionym:
Glochidion lichenisilvae Leandri, Notul. Syst.
(Paris) 0: 29. 1937. TYPE: Madagascar. Mont
Tsaratanana, 2000 m, silve a lichens, //. Per¬
rier de la B&thie 16414 (P).
known from the type only.
PliyllanthuH inarojejiensis (Leandri) Petra Hoff¬
mann & McPherson, comb. nov. Basionym:
Glochidion marojejiense Leandri, Mem. Inst.
Sci. Madagascar, Ser. B. Biol. Veg. 8: 215.
1957. SYNTYPES: Madagascar. Sommet ori¬
ental du Massif de Marojejy (Nord-Est) a
l’ouest de la haute Manantenina, affluent de la
Lokoho, 1850-2137 m, gneiss el quartzite,
dans la vegetation ericoide, II. Humbert 22761
(MO, P); Vallee de la Lokoho (nord-est): mont
Beondroka an nord de Maroambihy, 1400—
1450 m, rochers de la crete (gneiss et quartz¬
ite), H. Humbert 25608 (P).
known from the types only.
Phvllanthus perrieri (Leandri) Petra Hoffmann N
McPherson, comb. nov. Basionym: Glochidion
perrieri Leandri, Bull. Soc. Bot. France 81:
606. 1934. TYPE: Madagascar. Massif de
Manongarivo, 800—1000 m, hois humides, H.
Perrier de la Bathie 9918 (P).
Selected further collections. MADAGASCAR. Aiitsi-
ranana Province: Reserve Speciale de Manongarivo,
Rekolosy, 14°02'S, 48°18'E, vallon en amont de la chute
de la riviere Rekolosy, I 100 m. Derleth 117 (G, k, MO);
Reserve Speciale de Manongarivo, Besinkara, I4°04’S,
48°17'E, Ambalafary, 500 in en-dessou du hameau, 300
in, Gautier & Chatelain 2426 (G, k, MO); Reserve Spe¬
ciale de Manongarivo, Besinkara, 14°04'S, 48°17'E, Am¬
balafary, 250 in. Gautier 4009 (G, MO); Reserve Speciale
de Manongarivo, above Ambodisakoana, 1100—1200 m.
McPherson A’ van der Werff 16407 (MO. TAN).
As mentioned above, two of the species listed
under Glochidion by Leandri (1958) do not need
new combinations because Leandri had initially
published them in Pliyllanthus. One of them was
first published under the illegitimate name P. mon-
ticola (1934) hut given a nomen novum, P. oreich-
titus, a year later. The correct citations are as fol¬
lows:
Volume 13, Number 3
2003
Hoffmann & McPherson
Madagascan Glochidion
309
Phyllanthus oreichtitus Leandri, Cat. PL Madag.,
Euphorb.: 24. 1935. Replaced name: Phyllan¬
thus monticola Leandri, Bull. Soc. Bot. France
81: 450. 1934. nom. illeg., non Phyllanthus
monticola Hutchinson & Dalziel. 1928. Glo¬
chidion oreichtitum (Leandri) Leandri, Not id.
Syst. (Paris) 6: 29. 1937. SYNTYPES: Centre:
Andringitra, foret a mousses et sous-bois her-
bace vers 1800 et 2000 m, H. Perrier de la
Bathie 13670 (P), same loeality and collector
14406 (P).
Selected further collections. MADAGASCAR. Antsi-
ranana Province: Reserve Speciale de Manongarivo,
Ambahatra, cours superieur, 13°59'S, 48°26'E, an bord
du bras de 1"Ambahatra, 900 m h l’Vt du point cote 1037,
700 m, Gautier et al. 3499 (G, K. MO). Antananarivo
Province: 30 km N of Ankazobe, 18°15'S, 47°09'K, rem¬
nant patches of plateau forest, 850 m. Miller & Phillipson
3717 (K, MO).
Phyllanthus sambiranensis Leandri, Bull. Bot.
Soc. France 81: 451. 1934. Glochidion sam-
biranense (Leandri) Leandri, Notul. Syst. (Par¬
is) 6: 29. 1937. SYNTYPES: Centre, Haul
Bassin du Sambi ratio, 500 m, quartzites, ro-
cailles, 41. Perrier de la Bathie 969B (P);
Sources d’Andraronga, versant NE, 1200 m,
rocailles, quartzites, H. Perrier de la Bathie
972B (P); Massif de Manongarivo, 1000 m.
bois sec, gres liasiques, 41. Perrier de la Bathie
9917 (P).
Selected further collections. MADAGASCAR. ‘"Chiefly
from North-West Madagascar,” Baron 5625 (K). Antsiran-
ana Province: Reserve Speciale de Manongarivo, Belo-
kosy, 14°02'S, 48°18'E, Crete 1 km du sommet, 1390 m,
Gautier & Chatelain 2714 (G, K, MO); Reserve Speciale
de Manongarivo, Belokosy, 14°02'S, 48°18'E, sommet du
Belokosy, 1490 m, Gautier & Derleth 2568 (G. K, MO):
Reserve Speciale de Manongarivo, Ambahatra, cours su¬
perieur. 13°59'S, 48°26'E, crete entre les deux bras de
l’Ambahatra, crete principale, 200 m au NNE du point
cdtt 5 1037, Gautier et al. 3489 (G. K, MO).
Evaluation of these seven taxa is made difficult
by the scarcity of material. Because very few spec¬
imens exist outside the Paris herbarium, the main
characters are briefly described here. They are
woody plants with triangular, non-auriculate cata-
phylls. All except for Phyllanthus oreichtitus are
completely glabrous. Leaf shape and size vary
greatly between the species, from large (up to 9 X
4 cm) ovate leaves in P. humhertii to small (ca. 0.5
X 0.15 cm), falcate leaves in P. marojejiertsis that
give the plant a distinctly ericoid habit. All seven
species are monoecious; the cymules are bisexual
(P. humhertii), or unisexual w ith staminate and pis¬
tillate flowers found on separate branches (P. sam¬
biranensis) or the same branch (remaining five spe¬
cies).
The number of sepals in African Phyllanthus is
usually stable and of diagnostic value (cf. Keay,
1958: 384). Brunei (1987) used this character to
define infrageneric groups (series) in Phyllanthus.
Of the seven species in question, Phyllanthus or¬
eichtitus always has six sepals, P. ankaratrae has
five sepals in staminate and six sepals in pistillate
flowers, and the remaining species have five sepals
in both sexes. The shapes of the staminate sepals
and disc are similar in all seven species; discrete
alternisepalous disc glands are always developed.
Androecium morphology divides the seven spe¬
cies under consideration in two groups. The first
group comprises Phyllanthus ankaratrae, P. hum-
bertii, P. lichenisilvae, and P. marojejiertsis with
completely connate stamens, and in this way re¬
sembles true Glochidion. Phyllanthus ankaratrae
and P. humhertii have three stamens with non-apic-
ulate anthers, whereas the androecium of P. lich¬
enisilvae consists of four stamens w ith a central ap-
iculum, and 4 J . marojejiensis has five stamens with
individual apicula. The remaining three species
have free stamens, numbering live in P. perrieri and
P. sambiranensis, and six in P. oreichtitus. Anthers
are much longer than wide in all species and al¬
ways dehisce longitudinally.
Pistillate pedicels of these seven species vary
greatly in length from 1 to 15 mm. The pistillate
disc is extremely reduced to lacking (Phyllanthus
ankaratrae and P. humhertii), distinct and annular-
pentagonal ( 4 J . lichenisilvae), irregularly dissected
(P. oreichtitus), or consists of discrete, minute al¬
ternisepalous lobes (P marojejiensis, P. perrieri, and
P. sambiranensis). The ovary is uniformly glabrous
and three-locular.
Great variation is found in styles and stigmas.
Phyllanthus perrieri stands out in having erect, te¬
rete, entire styles up to 1.5 mm long that appear to
be fused at least apieally. It was the first Madagas¬
can species to be described as Glochidion (Leandri,
1934), mainly because of the fused styles, as is the
case in true Glochidion. The styles and stigmas of
the remaining six species classified as Glochidion
by Leandri (1958) are free and no more than 0.5
mm long, and of two principal shapes. In the first
three species of this group, the short styles are more
or less terete and bifid. In 4 J . ankaratrae and P.
oreichtitus they are inserted centrally and are api-
cally reflexed, whereas the styles of P. lichenisilvae
arise off-center and lean inward. In the remaining
three species under consideration the styles seem
to be reduced and there are three rather large, en¬
tire, flat stigmas. In P humhertii and P. marojejien-
310
Novon
sis these arise centrally and are spread flat against
the ovary, whereas P sambiranensis has similar but
excentric, inward-leaning stigmas.
Fruits and seeds are unknown in Phyllanthus
ankaratrae and P. lichenisilvae, and no seed has
been seen of P. marojejiensis. Where known, fruits
are smooth capsules, and the seeds are uniformly
triquetrous with a distinct elliptical, usually slightly
impressed but not invaginated chalaza and a
smooth dry seed coat.
It is obvious from this morphological discussion
that the Madagascan species of Phyllanthus united
by Leandri as Glochidion show considerable diver¬
sity. They represent distinct species with unique
combinations of characters although the distinc¬
tiveness of the infraspecific entities G. oreichtitum
subsp. tsaratanense Leandri and G. samhiranense
var. trapezophyllum Leandri is questionable.
Acknowledgments. We thank the staff of the La-
boratoire de Phanerogamic, Museum National
d’Histoire Naturelle, Paris (P), for giving us access
to their collection, and Laurent Gautier and his
team (G) for first drawing our attention to the prob¬
lem with their collections from Manongarivo.
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ques genres voisins de la Tribu des Phyllantheae Du-
mort. (Euphorbiaeeae, Phyllantheae) en Afrique inter-
tropieale et it Madagascar. These Doctorale, Universite
L. Pasteur Strasbourg I.
Chase, M. W, S. Zmarzty, M. I). Lledo, k. J. Wurdack, S.
M. Swensen & M. E. Eay. 2002. Vi hen in doubt, put it
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la Reserve Speeiale de Manongarivo, Madagascar. In:
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tique el Faunistique de la Reserve Speeiale de Man¬
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Hoffmann. P. & G. McPherson. 2003. Euphorbiaeeae—
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Oversea Governments and Administrations, London.
Leandri, J. 1934. Sur la presence a Madagascar du genre
Glochidion (Euphorbiacees). Bull. Soc. Bot. France 81:
606-607.
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de Madagascar. Notul. Syst. (Paris) 6: 11—35.
-. 1957. Notes systemaliques sur les Euphorbi¬
acees—Phyllanthees de Madagascar. MOn. Inst. Sci.
Madagascar, Ser. It. Biol. VTg. 8: 205—261.
-. 1958. Euphorbiacees 1 (Phyllanthoideae). In: II.
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I lie famille. 1 — 199. Firmin-Didot, Paris.
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Savolainen, V., M. F. Fay, I). C. Albach, A. Backlund, M.
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Delphinium lihengianum (Ranunculaceae), a New Species from
Xizang, China
Luo Yan and Yang Qiner
Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy
of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.
luoyan@ns.ibcas.ac.cn; qeyang@ns.ibcas.ac.cn
Guanghua Zhu
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. Delphinium lihengianum Q. E. Yang
& Luo. a new species of the Ranunculaceae from
Yadong County, Xizang. is described and illustrat¬
ed. This new species is similar to D. caeruleum
Jacquemont, especially in the indumentum of the
stem and pedicels anil in the leaf divisions, hut
differs in having the staminodes bifid to the middle
ami three carpels.
Key words: China. Delphinium, Ranunculaceae,
Xizang.
During the second author’s visit to the Royal Bo¬
tanic Gardens. Kew, from December 2001 to June
2002, the type specimen of Delphinium norlonii
Dunn (1927). E. F. Norton 369, which was collect¬
ed from the sandy soil above Karnpa Dzong, 12
miles north of Sikkim. Xizang. was carefully ex¬
amined. file sheet includes six perfect, small plant
individuals, which have one to three large inflated-
globose flowers and black-brown staminodes. Be¬
cause of these two characters D. norlonii should be
a member of D. subg. Delphinastrum (DC.) Peter-
rnann sect. Elatopsis Hath. However, Wang (1962)
placed D. norlonii in D. subg. Delphinastrum sect.
Delphinastrum, citing only one specimen, P. C.
Tsoong 5950, from Chumbi, Yadong County, south¬
eastern Xizang, under this species and stating that
he did not see its type specimen. This treatment
was followed by Wang (1979) and Wang and War-
nock (2001). As the members in section Elatopsis
are different from those in section Delphinastrum
in the color of the staminodes, we suspected that
the plant referred to D. norlonii by Wang (1962),
P C. Tsoong 5950, probably represented a mis-
identification, and thus its identity should be re¬
determined. Close examination of this specimen in
the Herbarium of the Institute of Botany, Chinese
Academy of Sciences (PE), which includes two
plants mounted on one sheet, has shown that it rep¬
resents an undescribed species.
Delphinium lihengianum O. E. Yang & Y. Luo,
sp. nov. TYPE: China. Xizang: Yadong County,
Chumbi, meadow in forest, July 1953 (fl), P.
C. Tsoong 5950 (holotype, PE). Eigure 1.
Herba perennis; caule graeili, 12-22 cm alto, terete,
ea. 1 cm diametro, adpresse pubescente, simplici vel 1-
ramoso, 2—3-foliato. Folia eaulina inferiora longe petiolata;
petiolo 3.2—5 cm longo, adpresse pubescente, basi anguste
vaginato; lamina utrinque sparse sericea, ambitu orbicu-
lari-pentagona, ca. 3 cm lata, trisecta, partitione centrali
rhombica tripartita fere usque ad costam, lobulis lineari-
lanceolatis vel linearibus 1—2 mm latis, partitionibus la-
teralibus oblique tetragonis inaequaliter 4-partitis fere
usque ad basim, lobis inaequaliter bifidis. Folia eaulina
superiora minora. Flores singuli ad apicem eaulis vel ap-
icem rami, 3.3—3.7 cm longi; pedicellis adpresse pubes¬
cent i bus, infra medium 2-bracteolatis, bracteolis lineari¬
bus, 1—3.3 cm longis, sessilibus vel breviter stipitatis;
sepalo superiore caeruleo, elliptico vel elliptico-ovato,
1.6—1.8 cm longo, extus pubescente, calcari cylindrico-
subulato, 1.7—1.9 cm longo, recto vel apice leviter deor-
sum eurvato, sepalis lateralibus late elliptieis, ca. 2 cm
longis, ca. 10 mm latis, sepalis inferioribus elliptieis, ca.
2 cm longis, ca. 5 mm latis; petalis in sicco brunneo-
flavis, glabris, apice emarginatis; staminodiis caeruleis,
limbo paulo longiore quam ungue, elliptico-obovato, bifido
usque ad medium, margine sparse ciliato, adaxialiter flav-
ido-barbato; staminum (ilamentis sparse puberulis vel gla¬
bris; carpellis 3, dense brunneo-villosis.
Perennial herbs. Stem slender, 12-22 cm tall,
terete, ca. I cm diam., adpressed pubescent, simple
or 1-branched, 2- to 3-leaved. Lower cauline leaves
long-petiolate; petiole 3.2—5 cm long, adpressed
pubescent, base narrowly vaginate; leaf blade
sparsely sericeous on both surfaces, orbicular-pen¬
tagonal in outline, ca. 3 cm wide, trisect, central
lobe rhombic, tripartite nearly to midvein, lobules
linear-lanceolate or linear, 1—2 mm wide, lateral
lobes obliquely tetragonal, unequally 4-partite
nearly to base, lobules unequally bifid. Upper cau¬
line leaves smaller. Flowers singular at apex of
stem or of branch, 3.3—3.7 cm long; pedicel ad¬
pressed pubescent, 2-bracteolate below middle;
Novon 13: 311-312. 2003.
312
Novon
Figure 1. Delphinium lihengianum O. K. Yang & Y. Luo.
—A. Habit. —B. Staminode. —C. Carpels. Drawn from
the type, P. C. Tsoong 5950 (PE).
bracteoles linear, 1-3.3 cm long, sessile or shortly
stipitate. Upper sepal blue, elliptic or elliptic-ovate,
1.6—1.8 cm long, pubescent outside; spur cylindric-
subulate, 1.7—1.9 cm long, straight or apex slightly
downcurved; lateral sepals broadly elliptic, ca. 2 X
ca. I cm; lower sepals elliptic, ca. 2 cm X ca. 5
mm. Petals brown-yellow when dry, glabrous, apex
emarginate. Staminodes blue; limb slightly longer
than claw, elliptic-obovate, bifid to middle, margin
sparsely ciliate, ventrally yellow barbate. Stamen
filaments sparsely puberulous or glabrous. Carpels
3, densely brown villose.
Distribution and habitat. Delphinium lihen¬
gianum is only known from its type locality. Yadong
County in southeastern Xizang, China, a border
area between China, Bhutan, and Sikkim, which is
not easily accessible. According to the collector's
note, this species grows in meadows in lorest.
Fieldwork needs to be done to know more about
the habitat of this species.
Taxonomic remarks. Delphinium lihengianum
is possibly related to I). caeruleum in usually hav¬
ing a slender and adpressed pubescent stem, ad-
pressed pubescent pedicels, and 3-sect leaves with
linear or linear-lanceolate lobules, but < 1 iffers in
having staminodes bifid to the middle and three
carpels. Its systematic position is far from D. nor-
tonii, which is distinguishable by its one to three
large inflated-globose flowers and black-brown
staminodes. In the original description, Dunn did
not mention the color of the staminodes, an impor¬
tant character in the classification of the genus Del¬
phinium. Our new species, D. lihengianum, which
has 3-sect leaves with linear or linear-lanceolate
lobules, blue staminodes, and the flower solitary at
the apex of the main stem or branch, should be a
member of subgenus Delphinastrum sect. Delphi-
nastrum ser. Caerulea W. T. Wang.
This species is named in honor of Professor Li
Heng from the Kunming Institute of Botany, Chi¬
nese Academy of Sciences, who provided valuable
guidance for the second author’s professional ca¬
reer.
Acknowledgments. We are most grateful to
Wang Wentsai, professor at the Institute of Botany,
Chinese Academy of Sciences, for his direction in
writing this paper. The second author thanks the
Keepers of K and PE for permission to examine
specimens or for the loan of material. We thank
Nick Turland for his comments on the manuscript
and Sun Yingbao for the illustration. This work was
supported by the National Natural Science Foun¬
dation of China (Grant No. 30070057) and a Key
Project of Chinese Academy of Sciences (KSCXZ-
SW-101A).
I .iterature Cited
Dunn. S. T. 1927. Plantarum novarum in Herbario Horti
Regii conservatarum. Bull. Misc. Inform. Kew 1927:
247.
Wang, W. T. 1962. Critical review of tin* genus Delphinium
from the ranunculaceous flora ol China (continued II).
Acta Bot. Sin. 10: 261—283.
-. 1979. Delphinium. Up. 326—462 in Flora Kei-
publicae Popularis Sinicae, Vol. 27. Science Press, Bei¬
jing.
-& M. J. Warnock. 2001. Delphinium. Pp. 223—
274 in /. Y. Wu & P. H. Raven (editors). Flora of China.
Vol. 6. Science Press, Beijing, & Missouri Botanical
Garden Press, St. Louis.
Luisma, a New Genus of Grammitidaceae
(Pteridophyta) from Colombia
M. I Murillo
Instituto de Ciencias Naturales, Herbario Nat ional Colombiano, Universidad Nacional de
Colombia, Apartado 7495, Bogota D.C., Colombia. mariateresamurillo@yahoo.com.mx
Alan R. Smith
University Herbarium, 1001 Valley Life Sciences Bldg. #2465, University of California,
Berkeley, California, 94720-2465, U.S.A. arsmith@ut link4.berkeley.edu
ABSTRACT. We describe and illustrate a monotyp-
ic new genus and species of fern, Luisma bivascu-
laris M. f. Murillo & A. B. Smith (Grammitida-
ceae), endemic from die Colombian Andes. Luisma
is similar to narrow-leaved species of Campyloneu-
rum (Polypodiaceae). However, it agrees with gram-
mitid ferns in having green, trilete spores (vs. hy¬
aline or yellowish, monolete spores in nearly all
Polypodiaceae) anti free veins (vs. anastomosing in
Campyloneurum). From all Grammitidaceae, Luis¬
ma differs in having two petiolar bundles. Other
distinguishing characters include suberect, radially
symmetrical rhizomes, sparse hyaline stipe hairs,
adaxial hydathodes, and clathrate, setulose rhizome
scales.
Rksumkn. Nosotros describimos e ilustramos un
nuevo geriero y especie tie helecho monotfpico,
Luisma bivascularis M. T. Murillo & A. R. Smith
(Grammitidaceae), endemieo de los Andes colom-
liianos. Luisma es similar a especies tie Campylo¬
neurum (Polypodiaceae) con hojas angostas. Sin
embargo, concuerda con las gramitidaceas por te¬
ller esporas verdes, triletes (vs. esporas hialinas o
amarillentas en casi todas las Polypodiaceae) y
venas libres (vs. anastomosadas en Campyloneu¬
rum). De todas las Grammitidaceae Luisma tlifiere
por tener dos haces peciolares. Otros caracteres
que lo distinguen incluyen un rizoma radialmente
simetrico, con pelos hialinos esparcidos en el es-
tfpite. hidatodos adaxiales y eseamas del rizoma
clatradas, setulosas.
Key words: Colombia, Grammitidaceae, Luis¬
ma.
In this paper we describe a new genus of Gram¬
mitidaceae in Colombia. A number of recent papers
establish a generic framework for the family in the
Neotropics. These include revisions or synoptical
treatments of Ceradenia (Bishop, 1988). Cochli-
dium (Bishop, 1978), Enterosora (Bishop & Smith,
1992), Grammitis sensu stricto (Bishop, 1977). Lel-
lingeria (Smith et al., 1991), Melpomene (Smith &
Moran, 1992), Micropolypodium (Smith. 1992),
Terpsichore (Smith. 1993), and Zygophlebia (Bish¬
op. 1989). Prior to these studies, Copeland (1952a,
1952b. 1955) dealt with various elements of Neo¬
tropical Grammitidaceae in a synoptical way, and
Morton (1967) and Smith and Moran (in Moran &
Riba. 1995) treated grammitids from adjacent Ec¬
uador and from Mesoameriea, respectively.
Discovery of a new genus of ferns, based solely
on previously undescribed species, is an uncom¬
mon event, notwithstanding several exciting discov¬
eries in the last two years. Sun et al. (2001) recently
described a new genus and species (Mankyua che-
juense B.-Y. Sun et al.) of the eusporangiate fern
family Ophioglossaceae from Cheju Island. Korea.
Even more recently, Caohangia squamata (Poly¬
podiaceae) has been described as a new genus and
species from Vietnam (Smith & Zhang, 2002). In
the Neotropics, the last such monotype named was
Nephopteris maxonii Lellinger (1966), a poorly
known plant (still known only from the type) from
Colombia, a country generally acknowledged to be
among the richest for ferns (as well as vascular
plants) in the Neotropics, and also among the rich¬
est in the world (Cuatrecasas, 1958; Gentry. 1982;
Moran, 1996; Rangel-Ch., 2000; Hammen, 1995).
Below we describe another monotypic genus, Luis¬
ma, with its sole species L. bivascularis. I he genus
is known onlv from the type.
Luisma bivascularis M. T. Murillo & A. B. Smith,
gen. et sp. nov. TYPE: Colombia. Risaralda:
Municipio de Mistrato. entre los corregimien-
tos de Geguadas y Puerto de Oro, selva de
pisones, 1550 m. zona de bosque pluvial pre-
montano, bosque primario, 30 mar. 1992. ./. L.
Fernandez 9613 (holotype, COL 468054). Fig¬
ure 1.
Novon 13: 313-317. 2003.
314
Novon
Figure I. Luisma bivascularis M. T. Murillo & A. R. Smith, gen. et sp. nov., based on J. Fernandez 96/.i (COL).
—A. Plant habit. —15. Rhizome scale. —C. Petiole and blade base. —I). Petiole, in cross section, showing two vascular
bundles. —E. Venation of a portion of the blade. —F. Sporangium, with spores.
Volume 13, Number 3
2003
Murillo & Smith
Luisma from Colombia
315
Plantae epiphytical. Rhizomata radialiter symmetrica,
erecta vel suberecta, apice squamata, squamis fumeis, ni-
tentibus, clathratis, usque 2.2 X 0.25 mm. lanceolatis,
basi peltatis ad marginem ciliatis, ciliis 0.05 rum longis,
liyalinis. 5ti[)ites teretes, basi cum duo fascibus vascular-
ibus in sectione transversali, 16-25 X 0.3-0.4 mm, fusci,
trichomatibus hyalinis simplibus, phyllopodiis carentibus.
Laminae prasinae in statu sicco, anguste ellipticae, 9-20
X 0.3—0.4 cm, membranaceae, basi et apice longiattenu-
atae. margine integrae; costae venaeque glabrae utrinque;
nervi libri, 2—3-furcati; hydathodi adaxialiter prominentes
ellipticae. Sori rotundati, ca. I mm diam., confinis costae,
uniseriales in lateribus ambabus costae, paraphysibus ab-
sentibus, capsulis sporangiorum glabris. Sporae virides,
globosae, 30-36 gm diam. (in acido laeteo), triletae.
Plants epiphytic. Rhizomes radially symmetrical,
erect or suberect, scaly at the apex, scales smoky
dark gray, shining, clathrate, up to 2.2 X 0.25 mm,
lanceolate, peltate at the base, margins ciliate, cilia
0.05 mm long, hyaline. Stipes terete, darkened
(brownish), at the base with two vascular bundles
in cross section, 16—25 X 0.3—0.4 mm, trichomes
hyaline, simple; phyllopodia absent. Blades green¬
ish in the dried state, narrowly elliptic, 9—20 X
0.3-0.4 cm, membranaceous, at the base and apex
long-attenuate, at the margins entire, midribs and
veins glabrous on both sides; nerves free, forked;
hydathodes adaxially prominent, elliptical. Sori
round, ca. 1 mm diam., adjacent to (touching) the
midrib, uniseriate on each side of the costa, lacking
paraphyses; sporangial capsules glabrous. Spores
green, globose, 30-36 /cm diam. (in lactic acid),
trilete.
Etymology. The genus is dedicated to Luis Ma¬
ria Murillo (father of the first author), pioneer in
entomology in Colombia and author of many sci¬
entific works. He was one of the founders in 1929
of the Academia Colombiana de Ciencias Exactas,
Ftsicas y Naturales, and editor of the journal for 10
years (1952—1961). He belonged to the Soeiedad
Colombiana de Geografta. was an honorary member
of the Soeiedad Real Entomologica de Belgiea, and
founded tlie Instituto de Altos Estudios del Go-
bierno Naeional. After more than 40 years of con¬
tinuous service in the Ministry of Agriculture as
"Jefe de Sanidad Vegetal”, Colombia’s President
Alberto Lleras Camargo conferred the Cruz de Boy-
aca in 1962, in recognition of his outstanding work
and exceptional scientific merits. We cite some of
his many publications (Murillo. 1937. 1944, 1951.
1956).
rile species epithet refers to the petioles, which
in cross section have two vascular bundles; this is
the only genus known in Grammitidaceae that has
thi s vascular configuration. The implications of this
finding for the phylogenetic position of Luisma
within the Grammitidaceae remain intriguing but
unknown, at present.
Habitat and distribution. The type locality is a
subandean pluvial forest, where the environment is
favorable for the existence of diverse plant life, in¬
cluding numerous epiphytes, ferns, mosses, li¬
chens. aroids, and orchids. In this area there have
been found recently several new bird species, in
Yireo (Salaman & Stiles, 1996) and Glaucidium
(Robbins & Stiles, 1999); a new lepidopteran in¬
sect, Antirrhea (Andrade, 1993); a new tree species
in the genus Matisia, Bombacaceae (Eernandez-
Alonso, 2001); and a new Columnea, Gesneriaceae
(Amaya-Marquez, 2002).
I he species is known from only a single collec¬
tion. and is seemingly rare. However, the locality
is relatively inaccessible, and the great floristic di¬
versity of the region, as well as the paucity of col¬
lections from the area, may obscure the fact that it
could lx 1 locally common. More likely, it is of very
restricted occurrence.
At first glance, Luisma bivascularis looks like a
species of Campyloneurum (Polypodiaeeae), but in
studying the characters further, we conclude that it
has no close affinity with that genus. Clearly, the
species is a member of the Grammitidaceae be¬
cause of its chlorophyllous (green) trilete spores
and free veins (although a few Polypodiaeeae sensu
stricto also have free veins, e.g., some species of
Polypodium and also Pecluma , and a few grammi-
tids have anastomosing veins, e.g., Zygophlebia).
Also the rhizome scales, which are strongly clath¬
rate and with hyaline marginal setulae, are remi¬
niscent of Lellingeria (Grammitidaceae) and unlike
anything known in Polypodiaeeae. The rhizomes
appear to be radially symmetrical and suberect,
like some grammitid ferns but unlike almost all Po-
lypodiaceae. We are unable to place the specimen
in any known genus of Grammitidaceae in either
the Neotropics or Paleotropies. From all known
genera and species of Grammitidaceae, Luisma bi¬
vascularis differs in having two vascular bundles in
the petioles. Many species of grammitids have been
checked for this character (Parris, 1990; A. R.
Smith & B. S. Parris, unpublished data), including
some that are larger than Luisma. In addition, Luis¬
ma differs from the largely Neotropical genera Cer-
adenia. Enterosora, and Zygophlebia in having
veins ending in prominent hydathodes adaxially.
Luisma further differs from Ceradenia in having
clathrate rhizome scales and from Zygophlebia in
lacking distinct phyllopodia.
from Terpsichore, Melpomene, and Micropolypo¬
dium, other Neotropical grammitid genera, Luisma
differs in lacking long reddish or red-brown setae
316
Novon
on the blades and petioles; in fact, die fronds of
Luisma are nearly glabrous or with only sparse
whitish hairs on the stipes. Melpomene further dif¬
fers in having entire marginal rhizome scales and
pinnatifid to pinnatiseet leaves. Like Micropolypo¬
dium and some species of Terpsichore , Luisma has
hyaline marginal setae on the rhizome scales, but
it differs from all species in those genera in having
entire leaves. From Lellingeria, which lias similar
rhizome scales. Luisma differs in its simple, entire
blades, lack of forked hairs on the stipes and ra-
ehises, and generally quite glabrous blades.
Other simple-bladed species of Grammitidaceae
in the Neotropics include Cochlidium, the species
of which have dull, coneolorous, orange to tan, en¬
tire rhizome scales; Grammitis, with species having
black sclerenchymatous margins and lustrous, con¬
eolorous, orange to tan. entire rhizome scales; and
Iximaphlehia (Greater Antilles), with anastomosing
veins and rhizome scales similar to those of Coch¬
lidium and Grammitis. American species of Gram¬
mitis sensu stricto lacking a black margin occur in
southern South America, but these have orangish,
coneolorous, entire rhizome scales and are very
small-bladed species, generally less than ca. 5 cm
long. We do not believe that any American genus
of Grammitidaceae is closely related to Luisma. nor
can we demonstrate that any Old World genus of
Grammitidaceae has close relationships. Without
molecular data, the closest affinities of Luisma will
probably remain obscure, and we believe it likely
that Luisma is one of the most isolated species and
genera in the family, phylogeneticallv.
Artikiciai
la. Blade
2a.
21 ).
Blade
5a.
5b.
Key to Genera of Grammitidaceae in Colombia
s entire, deeply serrate, crenate, or broadly and shallowly lobed.
Petioles each with two vascular bundles; rhizome scales clathrate, with ciliate margins. Luisma
Petioles each with a single vascular bundle; rhizome scales non-clathrate, entire or at least not ciliate.
.5a. Blade margins sclerified, dark brown or black. Grammitis
3b. Blade margins not sclerified, greenish or yellowish.
4a. Rhizomes radially symmetrical; blades entire to deeply serrate, usually loss than 5 mm wide;
hydathodes evident or not adaxiallv. Cochlidium
4b. Rhizomes dorsiventral; blades entire, crenate, or broadly lobed, usually more than 5 mm
wide; hydathodes absent adaxially. Enterosora
■s pinnatifid, pinnatiseet, or 1-pinnate.
Hydathodes absent adaxially.
6a. Veins free; opaque whitish stalked glands borne in soil and sometimes also on blades . . Ceradenia
6b. Veins areolate; opaque whitish glands lacking in sori and on blades. Zygophlebia
Hydathodes present adaxially.
7a. Rhizome scales not clathrate.
8a. Blades less than 1 cm wide, with one sorus per segment; main veins in segments (pinnae)
simple or with a single acroscopic branch; rhizomes radially symmetric .... Micropolypodium
8b. Blades usually more than 1 cm wide, with more than one sorus per segment; main veins in
segments (pinnae) pinnately branched; rhizomes dorsiventral to radial. Terpsichore
7b. Rhizome scales usually strongly clathrate.
9a. Setae lacking on margins of scales, but sometimes with several minute papillae at scale tips
. Melpomene
9b. Setae on margins of scales hyaline to reddish.
10a. Cell luminae clear; blades lacking black elavate fungal fruiting bodies. Lellingeria
I Ob. Cell luminae darkened; blades often with black elavate fungal fruiting bodies, espe¬
cially along rachises, costae, and among sporangia. Terpsichore
Acknowledgments. We express our gratitude to
the Working Group on Grammitidaceae for their
comments: Tom Ranker, Barbara Parris, Chris
Haulier, and Harald Schneider. We also thank
members of the Instituto de Ciencias Naturales de
la Universidad Nacional de Colombia for facilities
ami suggestions on the manuscript. Lastly, we
thank Juan Carlos Pinzon for executing the illus¬
trations.
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Luisma from Colombia
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pygmy-owl (Strigidae: Glaucidium) from the Pacific
slope of the northern Andes. Auk I 16: 305—315.
Salaman, P. G. W. & F. G. Stiles. 1996. A distinctive new
species of vireo (Passeriformes: Vireonidae) from the
western Andes of Colombia. Ibis 138: 610-619.
Smith, A. R. 1992. A review of the fern genus .1 hcro-
polypodium (Grammitidaceae). Novon 2 : 419—425.
-. 1993. Terpsichore, a new genus of Grammitida¬
ceae (Pteridophyta). Novon 3: 478— 489.
-& R. C. Moran. 1992. Melpomene, a new genus
of Grammitidaceae (Pteridophyta). Novon 2: 426—432.
- & X.-C. Zhang. 2002. Caobangia, a new genus
and species of Polypodiaceae from Vietnam. Novon 12:
546—550.
-, R. C. Moran & E. E. Bishop. 1991. Lellingeria,
a new genus of Grammitidaceae. Amer. Fern J. 81: 76—
88 .
Sun, B. Y., M. H. Kim, C. II. Kim & C.-W. Park. 2001.
Mankuyu (Ophioglossaceae): A new fern genus from
Cheju Island, Korea. Taxon 50: 1019—1024.
Notes on Grasses (Poaceae) for I lie Flora of China, I: Deyeuxia
Sylvia M. Phillips
Herbarium, Royal Botanic Gardens, kew, Surrey TW9 3AB, United Kingdom.
s.phillips@kew.org
Wen-Li Clien
Institute of Botany, Chinese Academy of Sciences, Beijing, 100093, China.
chenwl@ns.ihcas.ac.cn
Abstract. Two problematical species from the
Himalaya that lie on the boundary between Agrostis
and Deyeuxia are discussed. One is transferred to De¬
yeuxia as Deyeuxia petelotii with the new synonym D.
continentalis. The name Deyeuxia abnormis J. D.
Hooker is retained for the second species, which is
confirmed as distinct from I), zenkeri. Two other spe¬
cies are transferred to Deyeuxia from Calamagrostis:
Deyeuxia sichuanemis and Deyeuxia korotkyi.
Key words: Aniselytron , Calamagrostis, China,
Deyeuxia, Himalaya, Poaceae.
I’he genera Agrostis L, Calamagrostis Adanson,
and Deyeuxia Clarion ex P. Beauvois form an in¬
tergrading complex of three incompletely separated
entities. Treatments of this complex vary consider¬
ably, even in recent works, so it has been necessary
to carefully consider the options for the treatment
for the Flora of China.
Agrostis has always been maintained separately,
while Calamagrostis and Deyeuxia are either main¬
tained separately or Deyeuxia is sunk into Cala¬
magrostis. Calamagrostis and Deyeuxia are kept
separate by S. b. Lu (1987) in Flora Reipublicae
Popularis Sinicae, but united by Koyama (1987) in
his treatment of the Japanese flora. They are united
by Tzvelev (1976) for the former Soviet Union, but
kept separate by Bor (1960) for the Indian subcon¬
tinent anil Veldkamp (1996) for Malesia. Simon
(1993) kept them separate in Australia and also
Rugolo de Agrasar (1978) in South America. There
is no consistency within Chinese provincial Floras.
For example, Deyeuxia was maintained in Flora
Qinghaiica by Lu (1999), but sunk in Flora Si-
chuanica by J. L. Yang (1988).
Although Agrostis has always been maintained
separately, the boundaries between Agrostis and
both Calamagrostis and Deyeuxia are just as ill-
defined as that between Calamagrostis and De¬
yeuxia. It therefore seems logical to us that either
one large genus with sections is recognized, or else
three segregate genera, but not a halfway position
of two genera. We prefer to recognize three genera
to avoid an unwieldy conglomerate, while acknowl¬
edging that some species will cause problems. This
also follows the treatment of the complex for China
by Lu (1987), and avoids the need for many new
combinations in Calamagrostis. Following this con¬
cept, most species of Calamagrostis sensu lato are
referable to Deyeuxia. The genus Calamagrostis
sensu stricto is confined to relatively few north tem¬
perate species.
The majority of species in the complex can be
placed in one of the three genera without difficulty,
according to the following key:
la. Glumes clearly shorter than floret.
2a. Lower glume up to 1/2 floret length, usually
much shorter or vestigial; raehilla extension
minute, glabrous. Aniselytron
2b. Lower glume 1/2 floret length or more; raeh¬
illa extension distinct, penicillate . . . Deyeuxia
lb. Glumes equaling or longer than floret.
3a. Spikelets usually less than 5 mm; callus gla¬
brous or shortly hairy; lemma hyaline; raeh¬
illa extension often absent . Agrostis
3b. Spikelets often more than 5 mm; callus
bearded, hairs 1/3 as long as to longer than
floret (if shorter, penicillate raehilla exten¬
sion present); lemma membranous to firm;
raehilla extension present or absent.
4a. Lemma at least 3/4 as long as glumes,
usually firm; callus hairs almost as long
as to clearly shorter than floret; raehilla
extension present, penicillate . . Deyeuxia
4b. Lemma 1/2-2/3 as long as glumes,
membranous; callus hairs often much
exceeding floret; raehilla extension ab¬
sent, or if present glabrous or shortly
hairy . Calamagrostis
There are a number of troublesome intermedi¬
ates, and it must be stressed that no character com¬
binations provide a definitive way for assigning all
tin 1 species in this group. Two such problematical
species, Deyeuxia petelotii (Hitchcock) S. \1. Phil¬
lips & W. L. Chen and I). abnormis J. I). Hooker,
Novon 13: 318-321. 2003.
Volume 13, Number 3
2003
Phillips & Chen
Deyeuxia (Poaceae)
319
which we have had to consider for the Flora of
China account, are discussed below. They lie on
the boundary between Agrostis and Deyeuxia , with
an open panicle of small spikelets as in Agrostis,
but a bearded callus and penicillate rachilla exten¬
sion as in Deyeuxia. Two other species are trans¬
ferred here from Calamagrostis to Deyeuxia.
A lew intermediate and outlying species from
this complex have been described in two other gen¬
era, Aniselytron Merrill and Aulacolepis Hackel.
The small Asian genus Aniselytron was revised by
Korthof and Veldkamp (1984 [1985]), and is dis¬
cussed below under I). petelotii. Aulacolepis Hackel
(1907) is illegitimate, as it is a later homonym of
Aulacolepis Ettingshausen (1893). Under our ge¬
neric concept, its species are divided between An¬
iselytron and Deyeuxia.
1). •yeuxia petelotii (Hitchcock) S. VI. Phillips &
W. E. Chen, comb. nov. Basionym: Aulacolepis
petelotii Hitchcock, J. Wash. Acad. Sci. 24:
291. 1934, gen. illegit. Aniselytron petelotii
(Hitchcock) Sojak, Cas. Nar. Mu/. Praze. Rada
Prir. 148: 202. 1979 [1980]. Neoaulacolepis
petelotii (Hitchcock) Rauschert, Taxon 31:
561. 1982. Agrostis petelotii (Hitchcock) Nol-
tie, Edinburgh J. Hot. 56: 386. 1999. Cala¬
magrostis petelotii (Hitchcock) Govaerts. World
Checkl. Seed PI. 3: 11. 1999. TYPE: Vietnam.
Tonkin: near Chapu, ca. 1900 m, Aug. 1933,
A. Petelot 4743 (holotype, US; isotype, P).
Agrostis continentalis Handel-Mazzetti, Symb. Sin. 7:
1297. 1. 40 f. 2. 1936. Syn. nov. Deyeuxia continen¬
talis (Handel-Mazzetti) I,. I.iou, Vase. PI. Hengduan
Mountains 2: 2240. 1994. TV PE: China. Yunnan:
between Yuanmou and Hailo, 1050-1350 m, 10 Oct.
1914. //. Handel-Mazzetti 5(118 (holotype. W).
Anisachne gracilis Keng, J. Wash. Acad. Sci. 48: 117.
1958. Aniselytron gracilis (Keng) N. X. Zhao. J. IYop.
& Subtrop. Rot. 3(2): 50. 1995. TV PE: China.
Guizhou: Pichieh, 1400 m, I June 1943. Hou Hsueh-
yuh 2148 (holotype, N not seen).
Anisachne gracilis var. multinodis V. V . Qian, Bull. Rot. Res..
Harbin 18: 398. 1998. TYPE: China. Yunnan: l,an-
eang, I'XMI m. 24 Nov. 1993. Qian Yi-yong 8154 (ho¬
lotype. Ill I RC: isotypes. NEFU. SMAO none seen).
Distribution. Northeastern India, Bhutan, Chi¬
na (Guizhou, Yunnan), North Vietnam.
Deyeuxia petelotii appears to he widespread from
northeastern India through southern China to north¬
ern Vietnam, at elevations of 1000—3400 m. It has
been described as new several times within this
area, and its anomalous generic position can be
seen from the synonymy above. We agree with l„
Lion (1994) that it is best placed in Deyeuxia be¬
cause of its bearded callus and penicillate rachilla
extension. The description of the rachilla as naked
in Hitchcocks (1934) protologue is a mistake. The
tifi is bare, but there are long hairs on its lower
half. I ,iou made the transfer based on Agrostis con¬
tinentalis Handel-Mazzetti, a name that has hitherto
been missed by authors outside China. However,
this is predated by Aulacolepis petelotii Hitchcock
from Vietnam.
In Deyeuxia petelotii the lemma usually slightly ex¬
ceeds the glumes, or at least the lower glume, as de¬
picted in the figure accompanying the protologue of
Anisachne gracilis Keng (1958: 118). A paratype
specimen, Liu Tchen-ngo 21012 (PE), has been seen.
Keng (1958) described his new genus Anisachne on
the basis of the glumes being a little shorter than the
lemma. He acknowledged that otherwise he would re¬
fer it to Deyeuxia, Although the glumes are usually
longer than the lemma in Deyeuxia, shorter glumes
are also seen in a few species from Australia and New'
Guinea. Vickery (1940) and Simon (1993) both placed
these short-glumed Australian species in Deyeuxia. A
further complication is that spikelets affected by nem¬
atode infection may have abnormally elongated lem¬
mas. as noted by Noltie (2000: 600) and seen by the
second author in I). holciformis (Jaubert & Spaeh) Bor
and I), scahrescens Grisebach in southwestern China.
These slightly short-glumed species of Deyeuxia
should not be equated with the situation in the
small genus Aniselytron. where the glumes are both
very much shorter than the floret, with the lower
glume sometimes vestigial. Aniselytron is further
distinguished by its short glabrous rachilla exten¬
sion. We agree with the concept of Aniselytron as
set out hv Korthof and Veldkamp (1984 [1985]),
comprising two species, A. treutleri ((). Kuntze) So¬
jak and A. agrostoides Merrill, both of which occur
in China. Aniselytron is placed in synonymy under
Calamagrostis by Clayton and Renvoize (1986:
135), and this treatment is followed by Noltie
(2000: 61 7), although in a note he implicitly agreed
with Korthof and Veldkamp that Aniselytron merits
separate generic status.
It has not been possible to see the type of Ani¬
sachne gracilis var. multinodis Y. V . Qian. The fig¬
ure in the protologue shows the floret longer than
the glumes, as in D. petelotii. but the habit is more
typical of D. abnormis.
Deyeuxia abnormis J. D. Hooker, El. Bril. India
7: 268. 1897 [1896]. Calamagrostis abnormis
(J. I). Hooker) U. Shukla, Grasses N.-East. In¬
dia: 45. 1996. TV PE: India. Khasia, Nonkreem,
Oct. 1850, D. Hooker & I. Thomson s.n. (lec-
totype, designated by Noltie (1999: 386, Agros¬
tis 12, right-hand specimen “B”), K).
320
Novon
Agrostis nagensis Bor, Kew Bull. 9: 497. 1954. Deyeuxia
nagensis (Bor) Veldkamp, J. Econ. Tax. Bol. 13: 74.
1989. TYPE: NE India. Nagaland, Japvo, 28 Sep.
1935, /V. L. Bor 6449 (holotype, K).
Distribution. Northeastern India, Bhutan, Chi¬
na (Yunnan).
Deyeuxia abnormis is very similar to D. petelotii,
as recognized by Keng when describing Anisachne.
The two have often been confused, especially in
northeastern India where both may grow together.
The confusion dates hack to Hooker (1896: 268),
although il seems likely that he did distinguish
them in the field as he assigned two species num¬
bers, Agrostis nos. 11 & 12 (these are not collection
numbers). All elements of the protologue of D. ab¬
normis at Kew, except two ol the three sheets of
Hooker & Thomson Agrostis 12, are indeed D. ab¬
normis. Two of the sheets of Agrostis 12 are Agrostis
petelotii , and the third is a mixture of both species,
with the right-hand specimen “B" being D. abnor¬
mis. Noltie (1999: 386) lectotypified D. abnormis
on this right-hand specimen “B” of the mixed
sheet, correcting a previous wrong leetotypifieation
by Bor. Bor (I960: 392) recognized only one spe¬
cies, D. abnormis (under the name Agrostis zenkeri ,
see below). However, close inspection clearly re¬
veals that two species are involved. Their distin¬
guishing features have been set out by Noltie
(1999: 387), and the main points are summarized
below:
Spikelets 1.9—2.5(—3) mm: glumes slightly shorter
than floret; callus hairs < 1 mm, not visible
between glumes; leaf sheaths smooth ... I). petelotii
Spikelets 2.5—3.5 mm; glumes equaling or longer
than floret; callus hairs 1.2-2 mm, visible between
glumes; leaf sheaths scabrous . D. abnormis
The name Deyeuxia abnormis has frequently
been treated as a synonym of Agrostis zenkeri Tri¬
llins (1841). Veldkamp transferred A. zenkeri to De¬
yeuxia as A. zenkeri (Trinius) Veldkamp (1985), and
Davidse placed il in Calamagrostis as C. zenkeri
(Trinius) Davidse (1994). However, we regard this
as a different, little-known species, probably from
southern India.
The type of Agrostis zenkeri (holotype, LE) con¬
sists of a single flowering culm without basal parts.
There is a photograph of the type at Kew and a full
description by Veldkamp (Korthof & Veldkamp,
1984 [1985]: 219). As pointed out by Noltie (1999:
384), much has already been written about the cor¬
rect identity of this taxon, but it is necessary to add
a further note here. Veldkamp was unable to equate
it with any species known to him, following J. D.
Hooker (1896: 258) who many years earlier listed
it as a doubtful species.
Bor (1954. 1960) thought the type specimen of
Agrostis zenkeri represented the same species as
specimens from northeastern India described as
Deyeuxia abnormis by Hooker. This view has been
generally followed and was accepted by Noltie
(1999. 2000). However, we find that the type ol A.
zenkeri differs from material of D. abnormis in sev¬
eral important respects as follows:
Pulea 2/3 as long as lemma; anthers ca. 0.8 mm;
callus hairs 1.2-1.8 mm. Deyeuxia abnormis
Palea as long as or slightly longer than lemma;
anthers ca 1.2 mm; callus hairs ca. 1 mm.
. Agrostis zenkeri
Therefore, we disagree with Bor and Noltie that
Deyeuxia abnormis should be treated as a synonym
of A. zenkeri, but instead agree with Veldkamp that
A. zenkeri is a species of uncertain application.
There has been much discussion about the prov¬
enance of the type of Agrostis zenkeri. The type
specimen at LE quite clearly bears a label stating
that it comes from the Nilgiris in southern India,
and Bor (1954) argued convincingly that it was col¬
lected by Bernard Schmid and then sent to Zenker.
As no further material of this species has been col¬
lected from the area, it has been supposed that a
mix-up of labels might have occurred (Bor, 1954;
Noltie, 1999). However, there is no real evidence
for this. Bor also erroneously described the lemma
of A. zenkeri as 3-nerved: in fact, it is 5-nerved as
correctly observed by Hooker (1896: 258). Both A.
zenkeri and D. abnormis have 5-nerved lemmas.
The protologue description of Calamagrostis sri-
lankensis Davidse (1994: 107) from montane grass¬
land in Sri Lanka agrees quite well with the de¬
scription of the type of Agrostis zenkeri by
Veldkamp. In this species the lemma midnerve is
variable, sometimes extending nearly to the lemma
lip (as in A. zenkeri) and sometimes excurrent from
near the base. Il has been transferred to Deyeuxia
by Veldkamp (1996: 410). Unfortunately, there is
no isotype specimen of C. srilankensis at Kew, al¬
though it is said to be present in the protologue.
We are not pursuing this further as it is not a Chi¬
nese problem, but it seems likely that the type of
A. zenkeri was indeed collected in southern India,
and may even be nonspecific with D. srilankensis.
Deyeuxia siehuanensis (J. L. Yang) S. M. Phillips
& W. L. Chen, comb. nov. Basionym: Cala¬
magrostis siehuanensis J. L. Yang, Acta Bot.
Yunnan. 5: 47. 1983. TYPE: China. Sichuan:
Wenchuan, Wolong. 2850 m, ,/. L. Yang B22I
(holotype, SAUT not seen: isotypes, CDB1 not
seen. PE).
Volume 13, Number 3
2003
Phillips & Chen
Deyeuxia (Poaceae)
321
Distribution. China (Gansu, Sichuan).
Deyeuxia sichuanensis has the scabrous lemma,
callus hairs only half as long as the lemma, and
penicillate rachilla extension of a typical Deyeuxia,
as nicely shown in the illustration accompanying
the protologue (1983: 48). It is similar to D. ne-
palensis Bor, but can he distinguished as follows:
Lemma awned from near middle; awn ca. 1.2 mm,
included in spikelet; anthers ca. 1 mm.
. D. sichuanensis
Lemma awned from near apex; awn 3—6 mm, exserted
from spikelet; anthers ca. 2 mm. I). nepalensis
Deyeuxia koroikyi (Litvinov) S. M. Phillips & W.
L. Chen, comb. nov. Basionym: Calamagrostis
korotkyi Litvinov, Schedae Herb. FI. Boss. 55:
no. 2750. 1918. TYPE: Korotky H.F.R. 2750
(holotype, LE not seen).
Calamagrostis turczaninowii Litvinov, Not. Syst. Herb.
Hort. Petrop. 2: 115. 1921. Calamagrostis korotkyi
subsp. turkczaninowii (Litvinov) Tzvelev, Zlaki
SSSR: 304. 1976. Deyeuxia turczaninowii (Litvinov)
Y. L. Chang ex S. L. Lu, FI. Reipubl. Popularis Sin.
9(3): 205. 1987. TYPE: Russia. Zaihaikal district, 6
Aug. 1909, I. Blagoveshchenskii & G. Poplavskya
1472 (lectotype, designated by Tzvelev (1976: 304),
LE not seen).
Distribution. Mongolia, Russia (E Siberia), Chi¬
na (N Heilongjiang. N Nei Mongol. NW Xinjiang).
Acknowledgments. The Chinese author was
supported by the National Natural Science Foun¬
dation of China (30270102) and the Chinese Acad¬
emy of Sciences (KSCX2-1—06B). Vie thank the
Director of the Herbarium at the Smithsonian In¬
stitution, Washington, D.C., for sending the type of
Aulacolepis petelotii on loan to Kew. and the Direc¬
tors of the Paris and Vienna herbaria for sending
types to Beijing.
Literature Cited
Bor, N. L. 1954. Notes on Asiatic Grasses: XVII. The
species Agrostis zenkeri Trim Kew Bull. 9: 441-442.
-. 1960. The Grasses of Burma, Ceylon, India and
Pakistan. Oxford, London, New York, Paris.
Clayton, W. 1). & S. A. Renvoize. 1986. Genera Gramin-
um. Kew Bull. Addit. Ser. XIII. Royal Botanic Gardens,
Kew.
Davidse, G. 1994. Calamagrostis. Pp. 106—108 in Flora
of Ceylon, Vol. 8. Amerind, New Delhi.
Hitchcock, A. S. 1934. New species of Aulacolepis and
other grasses. J. Wash. Acad. Sci. 24(7): 290-292.
Hooker, J. 1). 1896. Flora of British India, Vol. 7. Gra-
mineae. Published under authority of Secretary of State
for India in Council, London.
Keng, Y. L. 1958. Two new genera of grasses from China.
.1. Wash. Acad. Sci. 48: 117-118.
Korthof, H. M. & J. F. Veldkamp. 1984 [19851. A revision
of Aniselytron with some new combinations in Deyeuxia
in SE Asia (Gramineae). Gard. Bull. Singapore 37: 213—
223.
Koyama, T. 1987. Grasses of Japan and its Neighbouring
Regions. Kodansha, Tokyo.
Liou, L. 1994. Deyeuxia. Pp. 2232—2241 in Vascular
Plants of the Hengduan Mountains. Science Press, Bei¬
jing.
Lu, S. L. 1987. Anisachne, Calamagrostis & Deyeuxia. In
Flora Reipublicae Popularis Sinicae 9(3). Science
Press. Beijing.
-. 1999. Calamagrostis & Deyeuxia. Pp. 130-138
in Flora Qinghaiica, Vol. 4. Qinghai People’s Publishing
House, Xining.
Nollie, II. J. 1999. Notes relating to the Flora of Bhutan:
XXXIX. Gramineae II. Edinburgh J. Bot. 56: 381—404.
-. 2000. Flora of Bhutan, Vol. 3(2). The Grasses of
Bhutan. Royal Botanic Garden Edinburgh & Royal
Government ol Bhutan.
Rugolo de Agrasar, Z. E. 1978. Las especies australes del
genero Deyeuxia Clar. (Gramineae) de la Argentina y
de Chile. Darwiniana 21: 417—453.
Simon, B. 1993. A Key to Australian Grasses. Dept, of
Primary Industries, Brisbane.
Tzvelev, N. N. 1976. Zlaki SSSR. Nauka Publishers, Len¬
ingrad.
-. 1983. Grasses of the Soviet Union. Ed. Fedorov.
Oxonian Press Pvt. Ltd.. New Delhi & Calcutta (trans¬
lation of Tzvelev, 1976).
Veldkamp, J. F. 1996. Name changes in Gramineae. Blu-
mea 41: 407-411.
Vickery, J. 1940. A revision of Australian species of De¬
yeuxia, with notes on the status of the genera Cala¬
magrostis and Deyeuxia. Contr. New South Wales Nall.
Herb. 1: 43-82.
Yang, J. L. 1988. Calamagrostis. Pp. 75-113 in Flora Si-
chuanica, Vol. 5(2). Sichuan Sciences & Technology
Publishing House, Chengdu.
A New Species of Rothmannia (Rubiaceae) from Yunnan, China
Shui Yu-Min and Chen Wen-Hong
Kunming Institute of Botany, The Chinese Academy of Sciences, Kunming 650204, China.
shuiyumin@hotmail.com
Chen Wei-Chlu
South China Institute of Botany, The Chinese Academy of Sciences,
Guangzhou 510650, China
Abstract. A new species in Ruhiaceae (Garden-
ieae), Rothmannia daweishanensis Y. M. Shui & W.
H. Chen, is described from Yunnan, China. Its di¬
agnostic characters and close relationship with
Rothmannia vietnamensis Tirvengadum are dis¬
cussed.
Key words: China, Rothmannia , Rubiaceae,
Yunnan.
During a phytogeographic study in the Daweis-
han range, southeastern Yunnan, and the revision
of Rubiaceae for the Flora of Yunnan in 2001,
Chen Wei-Chiu found a collection of Rubiaceae in
young fruit, K M. Shui <£ W. H. Chen 12242, un¬
recorded for China (Chen, 1999). hater, another
collection in botli flower and fruit, K M. Shui, C.
X. Pu & Z. A. Nie 1449 6, confirmed that a new
species in Rubiaceae had been discovered.
In accordance with studies of the tribe Garden-
ieae (Ixoroideae) in southern and southeastern Asia
(Tirvengadum, 1983; Robbreeht & Puff, 1986) and
Africa (Bridson, 1983, 1988), we place the new
species in the genus Rothmannia Thunberg, albeit
provisionally because of the difficulty of delimita¬
tion of the genera of Gardenieae. This is thus the
first record of the genus for China and Yunnan.
Because this genus is known only from tropical
Asia and Africa, its discovery in Yunnan adds
weight to the opinion that the flora of the Daweis-
han range has a tropical element and is allied to
the flora of the Paleotropieal Region (Wu & Wu,
1998).
Rothmannia occurs in tropical and southern Af¬
rica, east into the Seychelles as well as in southern
and southeastern Asia, with 63 species recognized
as valid in the International Plant Names Index
((www.ipni.org)). According to Jarvie and Ermayanti
(1996), together with Tirvengadum (1983), the di¬
agnostic features of the genus are: shrubs or large
trees (generally 4—15 m tall), spines absent; bud ter¬
minal and nude; inflorescences terminal, paniculi-
form; flowers coming from a single point on the stem,
or on a branched axis; petals 5, joined together, not
fleshy, entire; corolla sometimes large, purple speck¬
led inside the tube; stamens 5. subsessile, not fused,
adherent to the perianth, equal in size; styles soli¬
tary, with 2-lobed stigmas; carpels connate, 2 (or 3),
with 2 (or 3) locules; fruit berry-like, globular or
spherical, fleshy, sometimes up to .5—6 cm in diam¬
eter; seeds flat, sunk in a pulpy matrix. Some species
have been cultivated for horticulture, such as Roth¬
mannia capensis Thunberg ((nature.jardin.free.fr/
arbre/ft_rothmannia.html)) and Rothmannia globosa
(I lochstetter) keay ((www.shop.sunshine-seeds.de/
d_l 1159_Rolhmannia_globosa34658.htm)).
Rolhiuannia daweishanensis \. M. Shui & W. H.
Chen, sp. nov. TYPE: China. Yunnan: Maguan
County, Culinqing community, Woody Station
nearby Nanxi Community <>1 Hekou County,
22°40'N. 103°53'E, 500-600 m. 20 Nov. 2000
(II & fr), Y. M. Shui, C. X. Pu & Z. 1. Nie
14496 (holotype, KEN; isotypes, KEN. MO).
Figure 1.
Species Rothmanniae vietnamensi si mi I is, seef calyois
lobis lineari-oblongis, 1.4— E6 cm longis (nec 0.7 cm) et
oorollis campanulatis 5—5.5 cm longis (nec tubaeformibus
ca. 2.2 cm) differt.
Tree 10—15 m, glabrous throughout; bark lenti-
cellate, gray-shaded; young twigs dichotomous.
Leaves: blades chartaeeous, ovate, 10—14 X 4—5
cm, attenuate slightly from the middle to narrowly
cuneate at the base, slightly decurrent on the pet¬
iole, apex distinctly acuminate, tip obtuse, both
surfaces green-brown and glossy, midrib shallowly
canaliculate at adaxial surface, prominent at ab-
axial surface, lateral nerves 5 or 6 pairs; secondary
nerves faintly visible adaxially, slightly prominent
abaxially, almost all in 32—35° angle to the midrib;
petiole 0.3 cm long; stipules broadly triangular-
ovate, 1.5—2.0 X 2—3 mm, distinct at base, acu¬
minate at apex. Inflorescence uniflorous, protruding
Novon 13: 322-324. 2003.
Volume 13, Number 3 Shui et al. 323
2003 Rothmannia daweishanensis from Yunnan
Figure 1. Rothmannia daweishanensis Y. VI. Shui & W. H. Chen. —A. Habit. —B. Corolla. —C. Calyx lobes and
pedicel. —D. Fruit. —E. Transverse section of fruit and seeds. Drawn by Liu Yi-Tao from the holotype (K M. Shui,
C. X. Pu A /.. I., Nie 14496).
324
Novon
from a very reduced branch above a pair of leaves,
superficially terminal or axillary; reduced branch
ca. 5 mm long; pedicels 2.0—2.5 cm long with 3
bracteoles; bracteoles alternate and triangular, 1.5
X 1.0 mm. glabrous outside, sericeous inside. Ca¬
lyx lobes linear-oblong, 14—16 X 0.7-0.9 mm. ob¬
tuse at apex, 1-nerved, deciduous. Calyx tube 2—
2.5 mm long, 5—6 mm diam., sparsely sericeous
outside, densely long-pubescent, and with short
colleters inside. Corolla campanulate, tubes gla¬
brous, 5—5.5 cm long, 3 cm diam. at mid-point,
white outside, striated, anil mottled purple inside,
abruptly contracted at base into a slender tube 6—
7 mm long and 5 mm diam.; corolla lobes 5, gla¬
brous, broadly ovate, 1.8—2.0 X 2.2—2.3 cm, obtuse
at apex, white outside, with purple mottling inside,
deciduous and not evident on fruit. Stamens 5, al¬
ternate to corolla lobes, subsessile, inserted on the
upper part of corolla tube; anthers linear, 1.9—2.0
X 0.1 cm, longitudinally dehiscent, tapered to 0.5
mm; ovary parietal, seemingly 2-locular, ovules nu¬
merous in each row on placentas, placentas amal¬
gamating at the center of the ovary. Style 5 cm long,
stigmatic surface up to 2.0 cm long, ca. 0.1 cm
diam., oblong-lanceolate. Fruit elongate when
young, ca. 3.5 X 1.6 cm, spherical at maturity, ca.
4—5 cm diam., slightly inflated and decurrent at
base. Seeds numerous, lenticular, 1.2 cm long, 0.8
cm wide. 0.4 cm thick, sunk in a pulpy matrix.
Habitat and distribution. Among the rocky
crevices in the limestone rainy forests, 300—600 m;
known only from Yunnan, China.
Phenology. Flowering from October to Decem¬
ber; fruiting from December throughout the follow¬
ing year until the next March.
Etymology. Rothmannia daweishanensis is
named after the Daweishan mountain range located
in southeastern Yunnan, which is one of the richest
floras noted for Yunnan as well as China (Shui,
2000 ).
Although it is difficult to delimit the genera of
Gardeniaceae, the solitary flower, slightly lobate co¬
rolla tubes to 5—5.5 cm long, as well as the obovoid
or obovoid-oblong fruits, are all similar to the char¬
acters noted for Rothmannia (Tirvengadum, 1983).
Within this genus, Rothmannia daweishanensis is
similar to Rothmannia vietnamensis Tirvengadum
in the shape of the fruits, but the latter differs
mainly in its wider, broadly triangular calyx lobes
to 0.7 cm and in its shorter corolla tubes to only
2.2 cm long.
Paratypes. CHINA. SE Yunnan: Maguan County,
Gulinqing, Arboretum-examining Station, in tropical sea¬
sonal rainy forest. 2 Oct. 2(M)2, Y. M. Slim. W. H. Chen &
S. Sheng .30098 (KI N. PE); Maguan, Gulinqing, Xin-
tian-cun, alt. 500—600 in. in subtropical montane forest of
limestone area. Hi Aug. 1994, .S’. K. flu. X. Cheng. Y. P.
Yang. ./. Murata, T. Sugawara, T. Kawahara & II. Na-
gamasu 2524 (KUN); llekou County, Nanxi, Huayudong
National Forestry Park, 300 m, 22 Apr. 2000. Y. \1. Slim
A- Hi II. Chen 12242 (KUN); Nanxi, 24 Nov. 1982, S. Q.
Xiao & P. X. He 3222 (SWFC).
Acknowledgments. This research is funded by
the National Natural Science Foundation of Yunnan
(2001C0022Q), the Taxonomy and Flora Special
Foundation. Chinese Academy of Sciences (CAS),
and the Key Laboratory Foundation of Yunnan
Province Forestry Academy, China. The authors are
also indebted to Wu Zhengyi (KUN) for his helpful
instruction and Fang Ruizheng for correcting the
manuscript. Additionally, Liu Yi-Tao was very help¬
ful in providing the drawing.
Literature Cited
Bridson, 1). M. 1983. Notes on Rothmannia fischeri and
allied species (Rubiaceae subfam. Cinchonoideae). Kew
Bull. 39: 67-72.
-. 1988. Rubiaceae. In I). M. Bridson & B. Verd-
court. F lora of Tropical Fiasl Africa: 511—519.
Chen, W. C. 1999. Trib. Gardenieae A. Rich, ex DC. In:
H. S. Lo (editor), FI. Reipubl. Popularis Sin. 71(1):
330—386. Science Press, Beijing.
Jarvie. J. K. & Ermayanti (1996 onward). “Tree Genera of
Borneo—Descriptions and Illustrations” (http://django.
harvard.edu/users/jjarvie/borneo.htm/)
Robbrecht, FT & C. Puff. 1986. A survey of Gardenieae
and related tribes (Rubiaceae). But. Jahrb. Syst. 108:
63-137.
Shui. Y. M. 2000. The Preliminary Study of Phytogeog¬
raphy of the Angiosperms in the Daweishan, SE Yun¬
nan, China. Ph.D. Thesis, Kunming Institute of Botany,
The Chinese Academy of Sciences.
Tirvengadum, D. I). 1983. New taxa and name changes in
tropical Asiatic Rubiaceae. Nordic J. Rot. 3: 155—469.
Wu, C. Y. N S. G. Wu. 1998. A proposal for a new floristic
Kingdom (Realm)—The E. Asiatic Kingdom, its delin¬
eation and characteristics. Pp. 3—43 in A. L. Zhang &
S. G. Wu (editors), Floristic Characteristics and Diver¬
sity of East Asian Plants. China Higher Education
Press, Beijing, Springer-Verlag, Berlin.
Mostuea muricata (Gelsemiaceae), a New Species from Brazil
Marcos Sobral
Departamento de Botanica UFMG, Av. Antonio Carlos 6627, 31270-110. Belo Horizonte,
MG, Brazil. sobral@icb.ufmg.br
Lucia Rossi
Herbario, Instituto de Botanica de Sao Paulo, Caixa Postal 4005, 01061-970, Sao Paulo,
SP, Brazil, lrossi@uol.com.br
Abstract. Mostuea muricata, an American spe¬
cies from a predominantly African genus, is de¬
scribed. illustrated, and compared with die African
M. brunonis and the American M. surinamensis,
from which it is distinguished mainly by its muri-
cate fruits and rugose, glabrous, and viscose seeds.
This species was collected in the Brazilian states
of Mato Grosso and Sao Paulo, and its finding rep¬
resents a southward extension of the range of Mos¬
tuea in South America, before now known only in
Suriname and northern Brazil.
Resumen. Se describe y ilustra Mostuea murica¬
ta, una especie americana de un genero principal-
mente afrieano. Se la compara con la especie af-
ricana M. brunonis y con la especie americana M.
surinamensis, de las cuales se difereneia especial-
mente por sus frutos muricados y sus semillas ru-
gosas, glabras y viscosas. Esta especie fue colec-
tada en los estados brasilenos de Mato Grosso y
Sao Paulo, y su hallazgo representa una extension
austral del area geografica del genero en Sudamer-
ica, que hasta ahora se conocia solo en Surinam y
el norte de Brasil.
Key words: Brazil. Gelsemiaceae, Mostuea.
Mostuea Didrichsen is a small genus with eight
recognized species (Leeuwenberg, 1961), seven in
Africa and one disjunct in northern South America.
It was formerly included in the Eoganiaceae, to¬
gether with Gelsemium, in the tribe Gelsemieae
(Leeuwenberg, 1961); nevertheless, recent phylo¬
genetic studies (Struwe et al.. 1994; APG, 1998;
Backlund et al., 2000) strongly support the recog¬
nition of tribe Gelsemieae as a separate family, Gel¬
semiaceae, mainly characterized by heterostylous
flowers with twice-divided stigmas, latrorse anthers,
imbricated corolla aestivation, and flattened seeds
(Struwe et al., 1994). In this paper we describe a
second South American species, collected in the
Brazilian states of Sao Paulo and Mato Grosso.
Mostuea muricata Sobral & Lc. Rossi, sp. nov.
TYPE: Brazil. Sao Paulo: Sao Carlos, Santa
Eudbxia, Nov. 1993. M. Sobral & J. R. Steh-
mann 8049 (holotype, SP; isotypes. 1CN,
MBM. MO). Figures I. 2.
Species liaec M. brunoni et M. surinamensi proxima, a
quibus fruetibus muricatis, seminibus viseosis recedit.
Undershrub to shrub 0.5—1.5(2) m high. Branch¬
es dichotomous, sparsely to densely whitish-tomen-
tose with hairs 0.1—0.2 mm long, eventually in un¬
equally developed pairs, always with 2 pairs of
leaves and bearing flowers, with 2 callose, elliptic,
and somewhat paler protuberances 1.5—2.0 mm
long by the insertion of the petioles. Leaves dis¬
colored in dried specimens, ovate, obovate, or el-
liptic-obovate, the proximal ones strongly unequal
in size, (3)10—40 X (2)5—25 mm, smaller than the
distal ones, these equal or subequal in size, asym¬
metrical. 30—75 X 17—40 nun. with pale hairs 0.1—
0.2 mm scattered adaxially and more dense and
velutinous abaxially, especially on nerves; apex
acute; base cuneate or rounded, frequently cordate
and then with the lobes concealing adaxial petioles;
central and secondary nerves sulcate adaxially and
salient abaxially. the secondary nerves 6 to 8(10)
pairs; petioles 1.0—1.5(2) X 1.0—1.3 mm, scarcely
visible from the adaxial side; stipules interpetiolar
triangular, 1.0—1.2 X 1.2—1.5 mm, 1 - to 5-laciniate,
the laeiniae 0.8—1.0 X 0.2 mm; colleters linear, up
to 10 X 1 mm, inside the stipules. Inflorescences
terminal diehasia or dichotomous cymes, smaller
than the leaves, 3- to 20-florous, rarely uniflorous;
peduncles 10—20 mm long; braeteoles linear, 1—3
X 0.1 mm. Flowers heterostylous, pentamerous,
rarely tetramerous; pedicels absent or to 3 X 0.2
mm; calyx 1.7-2.0 mm long, sparsely to densely
covered with hairs, with equal or slightly unequal
laeiniae 1—2 X 0.3—0.8 mm; corolla infundibuli-
form, white, glabrous or scarcely pilose externally
at the apex, 5.5—8.0 X 2.0—2.5 mm in the widest
Novon 13: 325-328. 2003.
326
Novon
2 mm
2 mm
2 mm
3cm
2 mm
2 mm
2 mm
2 mm
Figure 1. Mostuea muricata Sobral & Ic. Rossi. —A. bruiting branch. —B. I.ongistylous flower. —C. Longistylous
flower in longitudinal section, drawn from fresh material showing recurved corolla lobes. —I). Brevistylous flower in
longitudinal section. —E. Fruit. —F. Seed, adaxial view. —G. Seed, abaxial view. A, E-G: Jaccoud 20 ; B, C: Rossi et
al. 1700; I): Rossi et al. 1697.
Volume 13, Number 3
2003
Sobral & Rossi
Mostuea muricata from Brazil
327
(li I!), and Sampaio 5094 , (R)) were examined.
portion, the lobes 1.5—2.5 X 1.5—2.0 mm, occa¬
sionally slightly revolute in nature; stamens equal
in size, 5-7 mm long and with exserted anthers in
brevistylous flowers, included and somewhat un¬
equal ca. 4 mm long in the longistylous flowers;
styles twice-branched, included and ca. 4 mm long
in brevistylous flowers and ca. 6 mm long and ex¬
serted in longistylous ones; ovary 1.8—2.0 X 1.5—
1.8 mm, externally 4-lobed, sometimes slightly ver-
rucose, glabrous or with scattered white hairs about
328
Novon
0.1 mm long at the apex, bilocular with 2 ovules
with central-basal placentation by locule. Fruit an
obeordate capsule, 5-6 X 7-10 mm, strongly mu-
ricate, with excrescences 0.1—0.3(0.5) mm long
crowned with a tuft of 4 to 5 pale hairs about 0.1
mm long. Seeds 2 to 4, red, shining, and viscose
when fresh, plano-convex, ellipsoid, glabrous, ru¬
gose, with a visible raphe and hylum in the flat¬
tened face, 4.5—5.0 X 3.5 mm.
Vernacular name. Agarra-agarra (from Jaccoud
20). “Agarra” is the Portuguese word for “grasp,”
and it is probable that the name has derived from
the ability of the seeds to attach to clothes or skin.
Distribution, habitat, and phenology. Disjunct
African and American families and genera are rel¬
atively common (Gentry (1993) listed 1 18 genera
in 51 families). Mostuea is an example of such a
phenomenon, with seven of its eight recognized
species (Leeuwenberg, 1961) occurring in Africa
and one in South America. The discovery of a spe¬
cies of Mostuea in central and southeastern Brazil
expands southward the geographical distribution of
the genus in South America. As far as known, the
distribution of the two American species does not
overlap (M. surinamensis Bentham is recorded only
for Suriname and northern Brazil), and in overall
morphology they look less similar to each other
than M. muricata to the African M. brunonis l)i-
drichsen (see Affinities below). Thus, Mostuea as
presently defined (Leeuwenberg, 1961) comprises
nine species, seven in Africa anti two in South
America. Mostuea muricata grows in shady and
sandy sites in the interior of mesophyllous forests
(“cerradoes”), where it eventually occurs in great
densities in the understory. Flowers were collected
from September to January, and fruits from October
to February.
Affinities. Mostuea muricata appears related to
African M. brunonis and American M. surinamensis
(for descriptions see Leeuwenberg, 1961) but is
markedly set apart from both by fruit and seed mor¬
phology. The fruits in M. brunonis are smooth, gla¬
brous or appressed-pubescent, and the seeds mi¬
nutely reticulate, dull and densely appressed-pilose;
fruits in M. surinamensis are smooth, glabrous, deep¬
ly bilobed (2 to 4 times as wide as long vs. lobes at
most as wide as long in all other species), and seeds
are glabrous and shiny, apparently not viscose. Veg-
etatively, M. muricata may be hard to distinguish
from M. brunonis: this highly variable and wide¬
spread species (from Ghana to Madagascar; I.eeu-
wenberg, 1961) has a wide range of leaf variation in
size (6—280 X 3-125 mm) and shape (orbicular to
oblong-ovate or lanceolate); from M. surinamensis it
can be separated by the elliptic to oblong-elliptic
leaves in this species. We have also observed that
in M. muricata all branches have a limited size, bear
four leaves, and end in an inflorescence—that is,
apparently there are no vegetative branches, while
they may be found in the other two species; never¬
theless, it would be premature to take it as a diag¬
nostic feature.
Paratypes. BRAZIL. Mato Grosso: Sao Felix do Ar-
aguaia, beira do rio Araguaia, 11°38'11.7"S,
50°40'23.7"W, 17 Mar. 1997, Souza et al. 14231 (ESA.
5P). Sao Paulo: Anhembi, mala da fazenda Barreiro
Rico, 5 Oct. 1956, Kuhlmann 3971 (RB, SI’): Descalvado,
Fazenda Graciosa, 3 Feb. 1966, Bordo 58 (SP); estrada
em di reran a Santa Eudbxia, 8 Nov. 1995. Rossi et al.
1696. 1697 (SP). Rossi et al. 1700 (SP); Fazenda Jaguarao,
Nov. 1993. Sobral 6i Stehmann 8045 (ICN, MBM, SP).
8046 (ICN. SP), 8047 (ICN, MBM. SP), 8048 (ICN, SP),
21 Jan. 1997, Stehmann A' Sobral 2234 (BHCB, ICN); Sao
Carlos, Santa Eudoxia. proximo & Fazenda Figueira Bran¬
ca, 30 Sep. 1980, Semir et al. 11540 (UEC), 11541 (UEC);
Sao Simao, Bocaina, proximo ao ebrrego da Prata, 8 July
1961. Jaccoud 20 (RB, SP), 30 Nov. I960. Kuhlmann
5004 (SP): Fazenda Bocaina, 29 Nov. 1961, Mattos 8674
(RB, SP).
Acknowledgments. We thank Joao B. Stehmann,
Elofsa de Araujo Rodrigues, and Maria Lucia Ka¬
wasaki for their kind help during the collections of
Mostuea muricata, the curators of BR, ESA, MO,
SP, and UEC, and Rogerio Lupo, for the beautiful
drawing of the species.
Literature Cited
APG (The Angiosperm Phylogeny Group). 1998. An or¬
dinal classification for the flowering plants. Ann. Mis¬
souri Bot. Card. 85: 531—555.
Backlund, M.. B. Oxelman & B. Bremer. 2000. Phyloge¬
netic relationships within the Gentianales based on
ndhY and rbc I. sequences with particular reference to
the Loganiaceae. Amer. J. Bot. 87: 1029—1043.
Gentry, A. H. 1993. Diversity and Horistic composition of
lowland tropical forests in Africa and South America.
Pp. 500—547 in P. Goldblatt (editor), Biological Rela¬
tionships between Africa and South America. Yale Univ.
Press, New Haven.
Leeuwenberg, A. J. M. 1961. The Loganiaceae of Africa.
11. A revision of Mostuea. Meded. Landbouwhogesehool
61(4): 1-31.
Struwe, 1... V. A. Albert & B. Bremer. 1994. Cladistics
and family level classification of the Gentianales. Cla¬
distics 10: 175—206.
New Combinations in Rohdea (Convallariaceae)
Noriyuki Tanaka
Department of Education, School of Liberal Arts, Teikyo University, 359 Otsuka, Hachioji,
Tokyo 192-0395, Japan, ntanaka@main.teikyo-u.ac.jp
ABSTRACT. Comparative survey shows that Roh¬
dea differs from eight species of Tupistra, one spe¬
cies of Campylandra (C. aurantiaca), and one spe¬
cies of Gonioscypha (G. eucomoides) mainly by the
degree ol thickness of the perianth and in the shape
and length of the perianth lobes. As these differ¬
ences do not seem to deserve generic delimitation,
those species of the last three genera are here
transferred to Rohdea. As a consequence, the fol¬
lowing 10 new combinations are proposed: Rohdea
aurantiaca (Baker) N. Tanaka, Rohdea chinensis
(Baker) N. Tanaka. Rohdea delavayi (Franchet) N.
Tanaka. Rohdea emeiensis (Z. Y. Zhu) N. Tanaka,
Rohdea ensifolia (F. T. Wang & Ts. Tang) N. Tanaka,
Rohdea eucomoides (Baker) IN. Tanaka, Rohdea jin-
shanensis (/. L. Yang & X. G. Luo) N. Tanaka,
Rohdea longipedunculata (F. T. Wang & S. Yun
Liang) N. Tanaka, Rohdea tonkinensis (Baillon) N.
Tanaka, and Rohdea verrueulosa (Q). H. Chen) IN.
Tanaka.
Key words: Campylandra, Eastern Asia, Gon¬
ioscypha. Rohdea, Tupistra.
The genus Rohdea was first described by Roth
(1821) with R. japonica (Thunberg) Roth from Ja¬
pan. Later, five other specific names were originally
published in the genus (cf. Liang, 1978; Liang &
Tamura, 2000). Recent studies by Liang (1978) and
Liang and Tamura (2000) recognized only one spe¬
cies, R. japonica (Fig. 1), in this genus. The five
other names were either submerged in R. japonica
or treated in other related genera ( Tupistra, Cam¬
pylandra). In reviewing the generic circumscription
of Rohdea, a comparison with its allied genera Tup¬
istra, Campylandra, and Gonioscypha is inevitable.
Tupistra was described by Ker Gawler (1814) with
T. squalida from Ambon (Amboyna), Indonesia,
though its provenance is questioned (Jessop, 1979).
Campylandra was founded by Baker (1875) with
the Himalayan species C. aurantiaca Baker. Tup¬
istra and Campylandra were regarded as distinct
by some botanists (e.g.. Baker. 1875; Engler, 1888;
Hutchinson, 1934; Tamura et al., 2000), while
Campylandra was treated as congeneric with Tup¬
istra by others (e.g., Bentham, 1883; Hooker, 1892;
Baillon, 1894; Liang, 1978; Huang & Li, 1990).
Under the latter view, many species with the fea¬
tures of Campylandra were described as Tupistra
(cf. Liang & Tamura, 2000; Tamura et al., 2000).
All or many of the species of Tupistra dealt with in
many papers (e.g., Liang, 1978; Huang et al.. 1989,
1990; Ma & Hong, 1990; Yang & Zhu, 1990;
Huang & Hong, 1997; Li, 1997) are those corre¬
sponding with Campylandra.
Huang and Li (1990) subdivided Tupistra into
two subgenera, subg. Tupistra and subg. Campylan¬
dra, the former comprising 13 species and the latter
14. Regarding Campylandra as a distinct genus,
Tamura et al. (2000) transferred 12 species initially
described under Tupistra and 2 species of Rohdea
to it. Liang and Tamura (2000) recognized 16 spe¬
cies in all in Campylandra. Yamashita and Tamura
(2001) recently added 1 new species to Campylan¬
dra, bringing the total to 17. I agree with these
botanists in regarding Campylandra as generically
distinct from Tupistra. Campylandra differs from
Tupistra in having small trisected stigmas (Fig. 24,
C) (vs. usually moderately to largely peltate, fun-
gill iform. or trilobed stigmas), orange to scarlet
smooth pericarps (Liang, 1978; Li, 1997) (vs. dark,
somewhat purplish brown to almost black, often
muricate or warty pericarps) (cf. Wan, 1984; my
own observations on T. albiflora K. Larsen and T.
fungilliformis F. T. Wang & S. Yun Liang, of which
the specimens are cited below), unimodal karyo¬
types (vs. bimodal karyotypes) (Huang et al., 1989,
1990; Yang & Zhu, 1990; Yang, 1995; Huang &
Liu, 1996), and monosulcate ellipsoidal pollen (vs.
inaperturate spheroidal pollen) (Ma & Hong, 1990;
Huang & Hong, 1997).
Meanwhile, Campylandra was treated as distinct
from Rohdea by many botanists (e.g.. Baker, 1875;
Bentham, 1883; Engler, 1888; Hutchinson, 1934;
Conran & Tamura, 1998; Liang & Tamura, 2000).
According to Baker (1875). Campylandra differs
from Rohdea in having an elongate style (vs. almost
lacking), short filaments (vs. none), and anthers lo¬
cated beneath the throat of the perianth tube (vs.
at the throat of the tube). According to current
knowledge, however, these distinctions are insig¬
nificant or partially incorrect. An indistinct style is
Novon 13: 329-333. 2003.
330
Novon
2 mm
Figure 1. Flower of Rohdea japonica. —A. Flower in front view. —B. Flower longitudinally sectioned with pistil
removed. —C. Pistil in longitudinal view. —D. Stamen in longitudinal view 1 . A-C to same upper scale, 1) to lower
scale. Drawn by Noriyuki Tanaka from a living plant in cultivation (/V. Tanaka s.n., TFLJ).
also seen in some species ol Campylandra, sucli as
C. siamensis Yamashita & M. N. Tamura, C. ver-
ruculosa (Q. II. Chen) M. N. Tamura, S. Yun I.iang
& Turland, and C. kwangtungensis Dandy. Anthers
are located at or near the throat ol the tube in C.
urotepala (Handel-Mazzetti) M. N. Tamura, S. Yun
Liang & Turland, and C. longipedunculata (F. T.
Wang & S. Yun Liang) M. N. Tamura, S. Yun Liang
& Turland (Fig. 2A, B). In Rohdea japonica, con¬
trary to Baker’s statement, filaments are evidently
present, though short, as shown in Figure I (A. B,
and D).
Conran and Tamura (1998) reported that Cam¬
pylandra has one stigma while Rohdea has three.
In my examination, both genera share a trisected
stigma in common (Figs. 1A, C, 2A, C). According
to Chen et al. (2000), Rohdea differs from Cam¬
pylandra by its indexed, very short perianth lobes
(vs. spreading to incurved lobes, which are 1/3—1/2
as long as the tube). In their diagnosis the term
“indexed” may be used to express a more abrupt
or angulated incurvature than “incurved." Rohdea
certainly has comparatively short perianth lobes
(Fig. 1A, B), but as to their curvature, it cannot
simply be said, at least at the dowering peak, that
the lobes are indexed. The perianth ol R. japonica
is incrassate (Fig. 1 A, B), and the lobes are, on the
ventral side, incurved to nearly erect (Fig. IB), and
on the dorsal side incurved to strongly indexed
(Fig. 1A, B). In Campylandra, many species (e.g.,
C. longipedunculata and C. verruculosa ) also have
perianth lobes showing incurvature at least in their
Figure 2. Flower of Rohdea longipedunculata (Campylandra longipedunculata). —A. Front view. —B. Longitudinal
section with pistil removed. —C. Pistil in longitudinal view. A-C to same scale. Drawn by Noriyuki Tanaka from a
living plant in cultivation, which was raised from a seed of II. Li, N. Tanaka & T. ./. Xiao 356 (TEU).
Volume 13, Number 3
2003
Tanaka
Rohdea (Convallariaceae)
331
upper part (Fig. 2A. B). The degree of incurvature
of the perianth lobes seems, therefore, not to be
decisive for distinguishing the two genera.
In my observation, Rohdea {R. japonica) differs
from Campylandra mainly by the following fea¬
tures: (1) the perianth lobes are short (vs. moder¬
ately long) and truncate or subtruncate (vs. round
to acuminate) at the apex, and (2) the perianth is
very incrassate with the lobes (usually upper ones)
particularly thickened dorsally (vs. less incrassate
and less dorsally thickened) (Figs. 1A, B. 2A. B).
These differences, however, seem to be insufficient
for the purpose of generic separation. Among oth¬
ers, the second difference appears to be rather
quantitative. Besides the close similarity in various
organs or parts of the plants (e.g., basal two-ranked
foliage leaves, flowers in a compact spike, fleshy
usually greenish or yellowish perianths, small tri¬
sected stigmas, and orange to scarlet smooth peri¬
carps). the two genera share similar unimodal kar¬
yotypes with the basic chromosome number x — 19
(Yang & Zhu, 1990), and monosulcate, ellipsoidal
pollen with a perforate or reticulate exine (Ma . nov.
TYPE: French Guiana. Carbet Grillon, 80 m.
9 Mar. 1999. B. Dutreve & F. Crazier 7 (ho-
lotype, MO; isotypes, CAY, K, NY, P, US). Fig¬
ure 4.
A ceteris speciebus Licariae subgeneris Cannellae foliis
subtus rufo-tomentosis, obovatis et ramulis juvenalibus
rufo-tomentosis recedit.
Trees. 25 m. Twigs terete or ridged, seasonal
growth densely dark red-brown tomentose. the sur¬
face completely covered; older twigs becoming gla¬
brous; terminal buds densely tomentose. Leaves 8—
18 X 3.5—7 cm, elliptic-obovate to obovate, coria¬
ceous, grouped near the lips of the branches, al¬
ternate, the base obtuse or cordate, the margin often
bent downward, the apex shortly acuminate, the up¬
per surface glabrous except for some tomentose in-
dument on the basal part of the midrib, the lower
surface reddish tomentose, completely covered by
the indument, the indument darker along the mid¬
rib; midrib, lateral veins, and tertiary venation
slightly raised on the upper surface, midrib and
lateral veins raised, tertiary venation not visible on
the lower surface; lateral veins 7 to 10 on each side;
petioles T)— 10 mm, with a similar indument to the
twigs, ± terete. Inflorescences 2 cm long, panicu-
late-cymose, densely tomentose, the surface com¬
pletely covered by the indument, in the axils of
leaves or bracts. Flowers ea. 2 mm long, globose or
ellipsoid, densely tomentellous, the hairs shorter
than those on the inflorescence axes, pedicels as
long as or slightly longer than the (lowers; tepals
six, ea. 0.5 mm long, triangular or broadly trian¬
gular, equal or the outer 3 slightly larger, much
shorter than the receptacle, densely tomentellous
outside, glabrous inside, outer 6 stamens stamino-
dial, tepaloid, a little smaller than the tepals, fertile
stamens 3, representing whorl III. ca. I mm long,
the basal part pubescent, the distal part glabrous.
anther cells 2 longitudinal slits, introrse-Iatrorse,
not opening with valves, glands, if present, fused
with the broadened base of the stamens and diffi¬
cult to discern; inner staminodia (whorl IV) not
seen; pistil glabrous, 1.3 mm long, the ovary glo¬
bose, 0.4 mm diam., style slender, stigma not dis¬
tinct; receptacle deep, brown-pubescent inside. In-
fmetescences short, excluding fruit to 5 cm, cupule
2.5—3 cm long, 3—3.5 cm diam., 1.5 cm deep, the
outer margin spreading, fruit ellipsoid, ca. 3 cm
long.
Flowers and fruits in March.
Licaria rufotomentosa is easily recognized by the
ferruginous indument on the lower leaf surface,
young twigs, and inflorescences. The clustered
leaves, the introrse-lateral anther cells, and the
large (to 3.5 cm diam.) cupules place the new spe¬
cies in Licaria subg. Cannella. It is closely related
to L. cannella (Meissner) Kostermans, a variable
species with three varieties (Kurz. 2000). In leaf
characters (short petioles, leaf bases obtuse or
rounded) the new species resembles the variety ten-
uicarpa (Kostermans) Kurz, but the tomentose in¬
dument on leaves anti twigs is not known from any
species of the subgenus Cannella.
fhe basionym of Licaria cannella is Aydendron
cannella Meissner (Meissner, 1864). Later, Meis¬
sner (1866) used the spelling “ canella ” instead of
" cannella ” in the Flora Brasiliensis treatment, and
this spelling has been used by all later authors. The
spelling cannella in Meissner (1864) was probably
based on the common name (Bois cannelle) cited
in the original publication. Because cannella does
not seem to be a typographical or orthographical
error, article 60.1 of the Code (Greater et al., 2000)
requires that the original spelling be retained.
4, Licaria sessiliflora van der Vie iff, sp. nov.
TYPE; Venezuela. Estado Amazonas: Rio Pas-
imoni, 4 Nov. 1994, B. Stergios, M. Nino , /.
Sikoura, A. Piven, P. iMartinez & N. Friederich
16276 (holotype, PORT; isotype, MO). Fig¬
ure 5.
A congeneris floribus sessilibus tubulosisque, foliis re-
ticulatis differt.
Tree, 8 m. Twigs terete, glabrous; terminal buds
glabrous. Leaves 9—12 X 3—4 cm, elliptic or ob-
ovate-elliptie, glabrous on both surfaces, alternate
or the two distal leaves subopposite, coriaceous, the
—>
figures 5, 6. —5 (left). Isotype of Licaria sessiliflora van der Werff. —-6 (right). Holotype of Licaria subsessilis van
rler Werff.
Volume 13, Number 3
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van der Werff
Lauraceae from South America
343
344
Novon
base acute or cuneate, the apex bluntly acute or
obtuse, tertiary venation raised on both surfaces,
midrib and lateral veins slightly raised on both sur¬
faces; lateral veins 3 to 5 pairs; petioles 1.3—1.8
cm, strongly canaliculate. Inflorescences in the ax¬
ils of bracts near the terminal buds, to 6 cm long,
paniculate-cymose, toward the flowers minutely
brown-puberulous. Flowers pink, sessile, the per¬
sistent bracts clasping the base of the flowers, tu¬
bular. 3—4 mm long; tepals 6, erect, much shorter
than the floral tube, ca. 0.5 mm long; stamens 3,
2-celled, exserted at anthesis, the anther cells lat¬
eral or lateral-apical, opening sideways; staminodia
6, representing the outer 2 whorls, lanceolate, frag-
ile. 1 mm long or more, just visible inside the flower
and reaching the tips of the tepals; staminodia of
whorl IV and glands at the base of the filaments of
the stamens not seen; pistil glabrous, ca. 2 mm
long, receptacle glabrous inside. Fruit unknown.
Flowers in November.
Licaria sessiliflora is a distinct species due to its
sessile flowers, persistent bracts, and leaves with
raised reticulation. Kurz (2000) recognized three
subgenera in his revision of Licaria, primarily
based on the opening of llie anther cells. In the
subgenus Cannella Kurz the anther cells open by
slits and do not have flaps; in subgenus Armeniaca
Kurz the anther cells open by flaps attached at the
base of the anther cells, while in subgenus Licaria
the anther cells open by flaps attached at the top
of the anther cells. This new species does not fit in
any of his subgenera: the flaps are attached later¬
ally rather than at the top or bottom of the anther
cells. Only one other species has anther cells open¬
ing in the same way: L. debilis (Mez) Kostermans,
but this species has opposite, chartaceous leaves,
lacks the raised reticulation, and has globose to
ellipsoid flowers. The raised reticulation is shared
with L. puchury-major (Martins) Kostermans, but
the latter species has the anther cells opening by
slits and lacks the flaps present in nearly all other
species. Thus, the relationships of the new species
are not clear.
5. Licaria snhsessilis van der Werff, sp. nov.
TYPE: Peru. Depto. Cajamarea: Prov. San Ig¬
nacio, Distr. San Jose de Lourdes, Cerro Pi-
corana, 2250—2300 m. 2 Dec. 1998, ,/. Cam¬
pos, L. Zurita & M. Camizan 5H74 (holotype,
MO 5593919; isotypes, HBG, MO. NY, US).
Figure 6.
A ceteris speciebus subgeneris Cannellae inflorescen-
tiis floribusque tomentosis, foliis subsessilibus, basi ro-
tundatis recedit.
frees, 15 m. Twigs terete or angular, glabrous,
solid, terminal buds glabrous or nearly so. Leaves
5—17 X 3—6.5 cm, coriaceous, alternate and clus¬
tered near the tips of the twigs, broadly elliptic to
elliptic-obovate, the base rounded, rarely obtuse or
cordate, to attenuate, the margins flat or slightly
incurved, the apex obtuse or very shortly acumi¬
nate; glabrous on both surfaces, the lower surface
densely gland-dotted; midrib slightly raised, lateral
veins and tertiary venation immersed on tin* upper
surface, midrib, lateral veins, and tertiary venation
raised on the lower surface; lateral veins 6 to 10
on each side; petioles to 1 cm long, thick, flat
above, glabrous. Inflorescences 2-12 cm long, pa¬
niculate-cymose, in the axils of leaves, densely red-
brown tomentose, the surfaces completely covered.
Flowers globose or a little longer than wide, ca. 3
mm diam., pedicels at anthesis about as long as the
flowers or slightly shorter, the flowers often nod¬
ding; tepals 6, ca. 1.5 mm long, difficult to separate
from the receptacle because of the dense indument,
equal or nearly so. triangular, tomentose on the out¬
er surface, glabous on the inner surface, outer 6
stamens staminodial, tepaloid, slightly smaller than
the tepals; functional stamens 3, representing whorl
III. 1.5 mm long, the anther cells appearing as lon¬
gitudinal slits on the inner surface of the anthers,
glands of the inner stamens lacking or completely
fused with the filaments, staminodes of whorl IV
not seen; pistil glabrous, 2 mm long, the receptacle
deep, pubescent inside. Cupules with coarse warts,
ca. 3 cm high. 4 cm diam., the margin 8 mm thick,
fruits 3—4 X 3 cm.
Flowers in December and March; fruits in
March.
The anther cells opening as introrse slits and the
clustered leaves place Licaria subsessilis in subge¬
nus Cannella. It differs from the other species in
this subgenus in its tomentose inflorescences and
flowers and its glabrous leaves. The lower leaf sur¬
face is densely gland-dotted. Its high-altitude hab¬
itat is also uncommon in Licaria. Three collections
from southern Ecuador (Prov. Zamora Chinchipe,
Estacion Cienlifiea San Francisco, 2100 m alt.) dif¬
fer from typical L. subsessilis in having an attenuate
or acute leaf base, obovate-elliptic leaves, and pet¬
ioles 5—10 mm long. At first I considered these
specimens to represent a distinct species closely
related to L. subsessilis. However, a very recent col¬
lection from this locality fits typical L. subsessilis
perfectly. Because I think it highly unlikely that
two closely related species, differing only in leaf
characters, occur in the same locality, and because
L. cannella, a lowland species closely related to L.
Volume 13, Number 3
2003
van der Werff
Lauraceae from South America
345
subsessilis, is quite variable in leaf characters, I
provisionally place the specimens with attenuate
leaf bases, obovate-elliptic- leaves, and distinct pet¬
ioles in L. subsessilis. Additional collections may
well show that two species are involved, but as long
as these collections are not at hand, it seems better
to accept that L. subsessilis is variable in leaf char¬
acters.
Paratypes. ECUADOR. Zamora Chinchipe: Esta-
cidn Cientfflca San Francisco, 30 km E of Loja on road
to Zamora, 2100 m, (fl. fr), Chimbo & Chamba 95 (E()JA.
MO, QCNE). (fl). Homeier & Scheffer 303 (MO. OCNE).
(fr), Neill & QCNE Botany Interns 12619. 12620 (MO.
OCNE).
III. Mezilaurus Taubert. Hot. Centralbl. 50: 21.
1892.
Mezilaurus is a small Neotropical genus with ea.
20 species. It was revised by van der Werff ( 1987)
who accepted 18 species; however, 2 species were
later transferred to the new genus Williamoden-
clron (Kubitzki & Richter, 1987). The latter genus
differed from Mezilaurus in its three 4-celled rath¬
er than 2-celled stamens. The androecium of Me¬
zilaurus is thus similar to that of Licaria , but Me¬
zilaurus differs from that genus in its clustered
(not alternate) leaves and in its fruit, subtended
by a small disc and not seated in a deep cupule
as in Licaria.
1. Mezilaurus campaucola van der Werff, sp.
nov. TAPE: Peru. Depto. Amazonas: Prov.
Condorcanqui, Distr. El Cenepa, Cerro Tutino,
600 m, 24 June 1997. R. Vasquez, A. Pena, E.
Chavez, E. Quiaco & A. Ampam 24170 (holo-
type. MO 5593918; isotypes, HBG, K, NY,
US). Figure 7.
Foiiis apice obtusis, floribus glabris, pedieellatis Mezi-
lauri navalii similis, sed foiiis latioribus, staminibus
dense pubescentibus et floribus latioribus quam longiori-
bus recedit.
Trees to 20 m. Twigs terete, glabrous, with con¬
spicuous scars of fallen leaves near the tips, ter¬
minal buds appressed pubescent. Leaves 8—16 X
4—7 cm, obovate or obovate-elliptic, coriaceous,
clustered near the tips of the branches, the base
attenuate, gradually narrowed into the petiole, the
margin flat, the apex obtuse, rounded or emargin-
ate; glabrous on both surfaces; midrib, lateral veins,
and tertiary venation weakly raised on the upper
surface, midrib raised, lateral veins and tertiary ve¬
nation weakly raised on the lower surface; lateral
veins 7 to 10 on each side; petioles poorly distin¬
guishable due to the decurrent leaf bases, 1—2.5
cm long, glabrous, flat. Inflorescences 3-8 cm, in
the axils of bracts near the tips ol the branches,
paniculate, the flowers clustered at the tips of the
secondary axes or a few racemose along the sec¬
ondary axes; glabrous or with a few appressed
hairs. Flowers yellow, ca. 1.5 mm wide, I mm long
(excluding the exserted anthers), pedicels 2—3 mm
long, glabrous; tepals 6, broadly triangular, 0.5 mm
wide, wider than long, glabrous on the outer sur¬
face, pubescent on the inner surface, erect or
slightly incurved at anthesis; stamens 3, ca. 1 mm
long, lree. the densely pubescent filaments as wide
as tbe glabrous anthers, anthers 2-celled, the cells
extrorse, opening back to back, staminodia and
glands not seen, pistil glabrous, 1.5 mm, the ovary
ca. 1 mm, distinct from the slender style, stigma
inconspicuous; receptacle bowl-shaped, glabrous
inside. Fruits ± round, 2 cm diam., the tepals per¬
sisting and forming a disc with the receptacle, 5
mm diam.; pedicel not swollen in fruit.
Flowers in June, September, October, and No¬
vember; fruits in June and October.
In the most recent revision of Mezilaurus (van
der Werff, 1987). Mezilaurus campaucola would key
to M. navalium (Allemao) Taubert ex M ez because
of its glabrous leaves with rounded apices and its
pedicellate, glabrous flowers. Mezilaurus navalium
is only known from the Atlantic rain forests in
southern Brazil and differs from tbe new species in
its narrow' leaves (to 3.5 cm wide vs. 4—7 cm wide
in M. campaucola ), and the shape of its flowers
(longer than wide vs. wider than long in M. cam¬
paucola). It also has chartaceous leaves, while M.
campaucola has coriaceous leaves. The epithet
campaucola is derived from campau. the Aguaruna
name for the soil type to which this species is re¬
stricted. Campau indicates soils not suitable for
cultivation; these are mostly shallow soils on sand¬
stone, covered with a thick, spongy humus laver.
The vegetation on these soils is generally a low
forest. Several species found on sandstone outcrops
in the campau show a relationship with the flora of
the sandstone mountains in the Venezuelan Gua-
yana, such as Euceraea nitida Martius (Flacourti-
aceae) and Aratitiyopea Steyermark & P. E. Berry
(Xyridaceae).
The epithet is the deliberate choice of the author
(Art. 23.2. St. Louis Code, Greuter et al.. 2000) and
is not to be regarded as a correctable error.
Paratypes. PERU. Amazonas: Prov. Bagua, Comu-
nidad de Kampaenza, 25 Sep. 1994. Jaramillo et al. 502
(MO); Comunidad Agauruna de Kusu-Listra, Cerro Apag.
18 Nov. 1996, Diaz et al. B5I3 (MO): Comunidad de Wa-
was, Tayu Mujaji. 25 Oct. 1997, Rojas et al. 473. 496
(MO).
346
Novon
Volume 13, Number 3
2003
van der Werff
Lauraceae from South America
347
2. Mezilaurus manausensis van der Werff, sp.
nov. TYPE: Brazil. Amazonas: Mun. de Ma¬
naus, Reserva No. 2206 do projeto Dinamica
Biologica de Fragmentos Florestais, free no.
1745, Mackenzie e equipe s.n., INPA/WWF
2206.1745 (holotype, MO 5593904). Figure 8.
Mezilauro micranthae similis, sed foliis chartaceis, flo-
ribus glabris, supra basin constrictis et indumento gem-
marum terminalium ascendenti recedit.
Trees, height unknown; trunks to 18 m. Twigs
terete, corky, initially appressed or ascending pu¬
bescent. soon becoming glabrous, with conspicuous
scars of fallen leaves; terminal buds densely pu¬
bescent, the hairs ascending. Leaves 8—27 X 3—8.5
cm, elliptic to elliptic-obovate, chartaceous, clus¬
tered near the tips of the twigs, the base gradually
narrowed into the petiole, the petiole often slightly
winged, the margin flat, the apex acute; the upper
leal surface glabrous, the lower surface glabrous or
with some appressed hairs along the major veins;
midrib raised, lateral veins slightly raised and ter¬
tiary venation immersed on the upper surface, mid¬
rib. lateral veins, and tertiary venation raised on
the lower surface; lateral veins 7 to 10 on each side;
petioles poorly differentiated from the leaf base, 1-
1.5 cm long, thickened at the base, terete or flat on
the upper side. Inflorescences 3—7 cm. paniculate,
glabrous, in the axils of bracts at the tips of the
twigs, the flowers clustered at tips of the secondary
axes, bracts pubescent, present at the base of the
secondary axes and flowers. Flowers glabrous, 1.5
mm long, 2 mm wide, constricted above the base,
the tepals erect, pedicels 3—4 mm long. Tepals 6,
subequal, the outer 3 a little shorter than the inner
3, broadly triangular, the inner 3 ca. 0.5 mm long,
the outer three 0.3—0.4 mm long; stamens 3, 0.8
mm long, the filament as wide as the anther, pu¬
bescent on the inner surface, shortly exserted, the
anther cells visible above the shorter, outer tepals,
anthers 2-cel led, the cells extrorse, rather small,
opening back to back; staminodia and glands not
seen; pistil glabrous, 1.5 mm long, the slender style
as long as the ovary; receptacle deeply cup-shaped,
pubescent inside. Fruits ellipsoid, 2.5 X 1.3 cm,
tepals persisting in fruiting stage, in young fruits
clearly recognizable, in mature fruits transformed
into a lobed disc, 5 mm diam.
Flowers in May and September; fruits in Decem¬
ber and January.
Species of Mezilaurus are difficult to recognize
due to their similarity in vegetative characters and
small flowers. In the key to Mezilaurus species in
van der Werff (1987), M. manausense would key to
M. sprucei (Meissner) Taubert ex Mez, which differs
in its long-pedicellate flowers, or M. micrantha van
der Werff, which differs in its slightly pubescent
flowers that are not constricted above the base, its
clearly exserted anthers with large anther cells, and
the strictly appressed indument of the terminal
buds. Although the differences between M. micran¬
tha and the new species are not striking, I describe
M. manausensis as a new species because of its
indument and floral differences from M. micrantha.
The new species is only known from a number of
specimens collected on plots made for the Biolog¬
ical Dynamics of Forest Fragments Project north of
Manaus, Brazil.
Paratypes. BRAZIL. Amazonas: Muti. de Manaus,
Projeto Dinamica Biologica de Fragmentos Florestais,
(lowering specimens: Reserva 1301 , tree 3221; 3258;
3412; 3587; 4106; Reserva 2303, tree 1707 (all MO); fruit¬
ing specimens: Reserva 1113, tree 35; Reserva 1301, tree
5154; Reserva 3402, tree 971 (all MO).
IV. Nectandra Rolander ex Rottboell, Acta Lit.
Lniv. Hafn. 1: 279. 1778.
Nectandra is a rather large genus of about 115
species, recently revised by Rohwer (1993). No
flowering material of the species described below
was available during the preparation of his revision,
and the fruiting specimens that were available were
insufficient for a description. Recent flowering col¬
lections are the basis for the following description.
1. Nectandra tomentosa van der Werff, sp. nov.
T\ PE: Ecuador. Zamora-Chinchipe: Campa-
mento Miazi along Rio Nangaritza, 900 m, 19
Feb. 1994. //. van der Werff. II. Gray, E. Freire
& M. Tirado 13268 (holotype, QCNF 104809;
isotypes, AAU. HBG, MO, QRS). Figure 9.
A congeneris foliis cordatis, ramulis ferrugineo-tomen-
tosis recedit.
Trees, 25 m tall. Twigs angular, ferruginous-to-
mentose, terminal buds tomentose. Leaves 13—30
X 4—10 cm, alternate, firmly chartaceous, elliptic
to oblong, the apex sharply acute to acuminate,
acumen to 3 cm long, toward the base gradually
narrowed and at the base abruptly cordate; midrib
figures 7, B. —7 (left). Holotype of Mezilaurus campaucola van der Werff. —8 (right). Holotype of Mezilaurus
manausensis van der Werff.
N? 5593908
348
Novon
Volume 13, Number 3
2003
van der Werff
Lauraceae from South America
349
and lateral veins impressed on the upper surface,
prominently raised on the lower surface, tertiary ve¬
nation immersed on the upper, raised on the lower
surface; upper surface glabrous or with some ap-
pressed hairs along the major veins, lower surface
glabrous or nearly so, but midrib and lateral veins
minutely puberulous, the hairs appressed to as¬
cending; axillary tufts of hairs absent; lateral veins
7 to 10. Petioles 5—10 mm long, with a similar in-
dument to the twigs. Inflorescences in the axils of
normal leaves, paniculate-cymose, to 15 cm long,
the main axes tomentose, changing to appressed
pubescent on the higher-order axes. Flowers white
or green-yellow, 3—4 mm diam.; tepals 0, equal, ca.
1.6 mm long, ± spreading at anthesis, inside with
the base sparsely papillose, the upper half densely
so, outside sparsely appressed pubescent; base of
tepals united and tepals falling off in old flowers as
a unit. Stamens 9, 4-eelled, ca. 0.5 mm long, with
a distinct filament, a sterile tip lacking, the cells
arranged in a shallow arc witli the lower pair shill¬
ing toward a lateral position; staminodia 3, stipiti-
form. Pistil glabrous, ca. 1 mm long, the style about
as long as the ovary. Receptacle inside basally gla¬
brous, the upper part appressed pubescent. Fruit
ellipsoid, 1.8 X 1.2 cm, cupule shallowly cup¬
shaped, warty, toward the base narrowed into the
swollen pedicel.
Flowers in December, February, and March;
fruits in September, December, and January.
Nectandra tomentosa is only known from the val¬
ley of the Rio Nangaritza, Ecuador, and adjacent
Peru where it is fairly common along rivers and in
forests adjoining the rivers on alluvial soil. Diag¬
nostic for this new species is the combination of
tomentose twigs and leaves with a cordate base.
Floral characters such as stamens with distinct fil¬
aments, a relatively long style, and the absence of
a sterile tip of the anthers indicate a close rela¬
tionship to the N. membranacea complex of Rohwer
(1993). Within this complex it is obviously close to
N. cordata Rohwer, which has similar leaves with
a cordate base. However, N. cordata has sparsely
appressed pubescent twigs and inflorescences,
quite unlike the indument in N. tomentosa', it is
only known from the type, collected near Balsa-
puerto, Loreto, Peru. Fruiting specimens of N. to¬
mentosa have been annotated by me previously as
N. cordata and N. cuneato-cordata Mez and may
have been distributed under those names.
Paratypes. ECUADOR. Zainora-Chinchipe: Rio
Nangaritza, 12 Dec. 1990, Neill & Palacios 97I3A (MO,
QCNE), 6 Dec. 1990. Palacios & Neill 6576 (MO, QCNE),
7 Dec. 1990, Palacios 6602 (MO. QCNE), 7 Dec. 1990.
Neill 9577 (MO, QCNE), 28 July 1993, Gentry 30529
(MO), 20 Feb. 1994, van der Werff et al. 13333 (MO,
QCNE, QRS). PERU. Amazonas: Prov. Condorcanqui,
Rio Cenepa, Mamayaque, 20 Feb. 1997, Vdsquez et al.
22651 (MO); Prov. Bagua, Dtto. Imaza, Kusu-Eistra, 19
Sep. 1996, Diaz et al. 3297 (MO); along road Chiriaco—
Bagua, 20 Mar. 2001, van der Werff et al. 16242 (CANB.
MO); Yamayakat, 6 Nov. 1996, Vdsquez et al. 21553 (MO),
20 Jan. 1973, Kayap 192 (MO).
V. Ocoiea Aublet, Hist. PI. Guiane 2: 780. 1775.
Ocotea is the largest genus of Lauraceae in the
Neotropics, with at least 300 species. A few species
occur in Africa, and about 30 are known from Mad¬
agascar. Because of its size and lack of recent treat¬
ments, Ocotea species are difficult to identify or
recognize. Additional species will almost certainly
need to be described. Rohwer’s (1986) synopsis of
the genus has been helpful in our understanding ol
the species, but much work remains to be done,
especially on the South American species. The
Central American species have been treated by van
der Werff (2002). A recent DNA-based phylogeny
of Lauraceae (Chanderbali et al., 2001) has indi¬
cated that Ocotea is polyphyletic and that the spe¬
cies with bisexual flowers need to be separated
from the species with unisexual flowers.
1. Ocotea arenaria van der Werff, sp. nov.
TYPE; Peru. Amazonas: vicinity of Molina-
pampa, 14 Mar. 1998, //. van der Werff, B.
Cray, R. Vdsquez & R. Rojas 14941 (holotype,
MO 5593908; isotypes, B, HBG, K. NY, QRS,
TI, US). Figure 10.
Ocoteae sericeae similis, sed floribus minoribus, tepalis
intus dense pubescentibus, inflorescentiis late paniculatis
recedi t.
Small, dioecious trees, to 7 m. Twigs angular or
ridged, solid, densely brown-tomentellous, the sur¬
face completely covered by the indument, this turn¬
ing darker with age and finally wearing off; terminal
buds densely brown-tomentellous. Leaves 6—11 X
1.5—2.5 cm, stiffly chartaceous, alternate, lanceo¬
late or narrowly elliptic, the base sharply acute,
narrowly decurrent on the petiole, the margin plane
or slightly revolute near the base, the apex acute,
sometimes bluntly so; midrib, secondary' veins, and
tertiary venation raised on the upper surface, the
Figures 9, 10. —9 (left). Isotype (MO) of Nectandra tomentosa van der Werff. Insert: Cupule and fruit.
Ilolotype ol Ocotea arenaria van der Werff.
10 (right).
350
No von
tertiary venation forming a line reticulum, midrib
raised, lateral veins and tertiary' venation immersed
or weakly raised on the lower surface; upper sur¬
face glabrous and shiny, on young leaves some
short, curly hairs present near the base, lower leaf
surface densely appressed pubescent, the surface
completely covered, the indument not sericeous,
vernation lines generally visible on young leaves,
but not on older leaves; domatia absent; lateral
veins I 1 to 14 pairs, weakly developed and difficult
to count; petioles 7—13 mm, flat above, with a sim¬
ilar indument to the twigs, the leaf bases often nar¬
rowly decurrent on the petioles. Inflorescences 6—
14 cm. paniculate-cymose, densely tomentellous,
the surface not visible, in the axils of leaves, f low¬
ers green, 3—4 mm diam. when tepals erect, 5—6
mm diam. when tepals spreading; staminate flow¬
ers: tepals 2 mm long, broadly ovate, spreading at
anthesis, densely brown-tomentellous outside,
densely pubescent on the inner surface, both sur¬
faces covered by the indument, stamens 9,
4-celled, the outer 6 stamens 1.2 mm, the filament
pubescent, much shorter than the anther, anther
cells opening introrse, inner 3 stamens 1.5 mm, the
filament pubescent, about as long as the anther,
glands present at the base of the filament, stami-
nodia not seen, pistillode 1.5 mm, with a well-de¬
veloped style and a thickened base, receptacle
deep, pubescent near the upper rim, otherwise gla¬
brous; pistillate flowers: tepals as in staminate flow¬
ers, but erect or half-erect at anthesis, staminodia
9, 0.6 mm, glands present at the base of the inner
3 staminodia, pistil 1.5 mm, the style about as long
as the ovary, a stigma well-developed; receptacle
deep, glabrous inside. Fruits and cupule unknown.
f lowers in March.
The dense, more or less appressed indument on
the lower leaf surface, the vernation lines, anti the
much-branched inflorescences place 0 . arenaria in
the (). guianensis group of Rohwer (1986). Within
this group its closest relative is 0 . sericea, a high-
altitude species reported from Venezuela, Colom¬
bia, and Ecuador. This species differs from (). ar-
enaria in having wider leaves with a sericeous
indument and a sharp, almost mueronate apex, te¬
pals ea. 3 mm long anti nearly glabrous on the inner
surface, and larger stamens and pistil. Ocotea ar¬
enaria is presently only known from the vicinity of
Molinapampa, at an altitude of 2300 m, near
Chachapoyas, in the Department of Amazonas,
Peru, where it was found in scrub on sandy soil,
with several indicators of poor soils, such as Utri-
cularia L., Eriocaulaceae, Xyris L., Sphagnum E.,
Pterozonium brevifrons (A. S. Smith) Lellinger, and
Doryopteris crenulans (Fee) H. Christ. A few col¬
lections from Colombia, as yet unidentified, are
close to 0 . arenaria, but differ in details of the
indument. None of these collections have flowers,
but two have fruits seated in double-margined cu-
pules. Double-margined cupules are known to oc¬
cur in two lowland species of the 0 . guianensis
group, but had not been reported from Andean spe¬
cies at higher altitudes.
Paratype. PERU. Amazonas: a few kin from Moli¬
napampa in scrub on nutrient-poor sandstone, 13 Mar.
1998, ran der Weiff et al. 14838 (MO).
2. Ocotea brevipetiolata van der Werff, sp. nov.
TYPE: Ecuador. Prov. Zamora-Chinchipe:
along new road Loja-Zamora, 2500 m, 25 Apr.
1987, //. van der Werff & IF Palacios 9004
(holotype, QCNE 167077; isotypes, AAU,
HUG. K. MO. NY, US). Figure I 1.
Ex affinitate Ocoteae floribundae, sed petiolis perbrev-
ibus, foliis basi rotundatis vel obtusis et floribus dense
pubeseentibus recedit.
Dioecious shrubs or trees to 20 m. Twigs ridged
to clearly angular, glabrous. Terminal buds pro¬
tected by glabrous bracts, inner bracts sometimes
pubescent or with pubescent margins; bracts some¬
times persisting as small, leaflike structures, to 2
cm long; scars of fallen bracts not clustered. Leaves
6—10 X 2.5—4 cm. rarely 15 X 8 cm, broadly el¬
liptic to obovate-elliptic, alternate, frequently
grouped together near the tips of the branches, co¬
riaceous, glabrous on both surfaces, the base obtuse
to rounded, the apex bluntly acute or obtuse; mid¬
rib, lateral veins, and tertiary venation immersed
on the upper surface, midrib raised, lateral veins
immersed and tertiary venation slightly raised on
the lower surface; lateral veins 5 to 7 on each side;
petioles 1-2 mm long, glabrous, flat above, the
leaves nearly sessile. Inflorescences to 7 cm long,
paniculate-cymose or racemose when small, near
the base moderately or sparsely pubescent, the
hairs ascending to erect, toward the flowers becom¬
ing densely tomentellous, the indument largely or
entirely covering the surface: in the axils of leaves
or bracts near the tips of the branches, often ap¬
pearing terminal, but a terminal, vegetative bud al-
Figures II, 12. —II (left). Isotype (MO) of Ocotea brevipetiolata van der Werff. Insert: Cupule and fruit,
(right). Isotype (MO) of Ocotea hirtostyla van der Werff. Insert: Cupule and fruit.
12
5593910
Volume 13, Number 3
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van der Werff
Lauraceae from South America
351
352
Novon
ways present. Flowers unisexual. 6—7 mm diam.,
white, the pedicels 1—1.5 nun long; tepals 3 nun
long, rather densely pubescent on the outer surface,
moderately pubescent on the inner surface. Stami-
nate flowers with 9 stamens, all 4-celled, the outer
6 stamens 2 mm long, glabrous, the anthers as long
as the filaments, the locelli filling the entire anther
and opening introrse; inner 3 stamens as outer 6,
but the locelli opening extrorse-lateral and the fil¬
aments with 2 well-developed glands at the base;
pistillode 2—2.5 mm long, pubescent, the style well-
developed; receptacle shallow, pubescent inside.
Pistillate flowers with 9 staminodes, these 1.5 mm
long, with 4 rudimentary locelli each; the filaments
slightly longer than the anthers, inner 3 staminodes
with 2 glands at the base of the filaments; pistil
2.5—3 mm long, the glabrous ovary as long as the
pubescent style, tbe stigma conspicuous, discoid.
Fruits rounded, ca. 1 cm diam., the cupule very
shallowly bowl-shaped to plate-like, with a double
margin, the tepals persisting on the outer margin.
Flowers in March. April, and May; fruits in De¬
cember.
Ocotea brevipetiolata is part of the Ocotea flori-
bunda group (Rohwer, 1986) by virtue of its uni¬
sexual flowers, pubescent pistil, and plate-like cu¬
pule with a double margin and persistent tepals. It
differs from the other species of this group in its
short petioles, leaves with obtuse to rotund bases,
glabrous terminal buds (or with at least the outer
bracts glabrous), and the rather dense indument of
inflorescences and (lowers. Ocotea floribunda
(Swartz) Mez is a widespread and somewhat vari¬
able species; it rarely occurs over 1500 m altitude.
Rohwer (1986) included two Andean species in his
0. floribunda group. One is 0. architectorum Mez,
known only from the type collected in Cajamarca,
Peru, at 2700—2900 m; the other is (). caniflora
Mez. known only from the type collected near Cuz¬
co, Peru, at 1700 m. Both of these species have
clearly petiolate leaves and cannot be confused
with O. brevipetiolata. Our knowledge about the
distribution of these species is still poor, due to a
lack of good collections. Most collections of O. brev¬
ipetiolata are from southern Ecuador, but there are
single collections from Colombia and Peru. A re¬
cent collection from the Reserva Maquipueuna in
Ecuador belongs to O. caniflora, while the distri¬
bution of 0. architectorum is still in question for
lack of good specimens.
Paratypes. COFOMBfA. Depto. del Valle: Mpio.
Tulua, old pistillate ff, Devia 1989 (MO). ECUADOR.
Loja: Carretera Loja—Zamora, eerca al curnhre, 23 Dec.
1991 (fr), Rubio et al. 2265 (MO); Parque Nacional Po-
doearpus, along road from Yangana to Cerro Toledo, 29
Apr. 1987 (stam. fl). van tier Werff & Palacios 9192 (HBG,
MO. QCNE), 29 Apr. 1987 (pist. fl). van tier Werff & Pa¬
lacios 9194 (HBG. MO. QCNE); pass on the road Loja—
Zamora, 9 May 1987 (pist. II), van tier Werff 9452 (MO),
9 May 1987 (stam. fl). van der Werff 9452 (MO). Zamora
Oiinchipe: along new road Loja—Zamora, 25 Apr. 1987
(fr. buds), van der Werff & Palacios 9000 (MO. OCNE);
along old trail from Nudo de Sabanillo to Vallodolid, 6
May 1987 (pist. fl). van der Werff &: Palacios 9282 (HBG,
MO. OCNE). 6 May 1987 (pist. fl. fr). van der Werff &
Palacios 9286 (HBG, MO. QCNE). PERU. Huamieo:
HuAnuco—Tingo Maria road, km. 456, 13 Apr. 1977, Gen¬
try et al. 19204 (MO).
3. Ocotea liirtostyla van tier Werff. sp. nov.
TYPE: Ecuador. Prov. Napo: Canton Tena, Es-
tacion Bioldgica Jatun Sacha, 400 m, 13 May
1990. IE Palacios & E. Freire 5100 (holotype,
QCNE 38682; isotypes, AALJ. HBG. MO, NY,
US). Figure 12.
Ocoteae floribundae affinis sed foliis subtus pilosis, ra-
mulis tomentellis recedit.
Dioecious trees to 17 m. Twigs somewhat angu¬
lar. densely brown-tomentellous, the surface of
young twigs completely covered by the indument,
becoming glabrous with age. Terminal buds densely
brown tomentellous. Leaves 1 1—26 X 5-12 cm, al¬
ternate, broadly elliptic to broadly obovate, (irmly
chartaceous, the base acute, the apex acuminate,
acumen to 2 cm long, but frequently broken off.
upper surface glabrous or with some erect hairs
along the major veins, lower surface pilose, the
hairs soft to the touch, erect and scattered, but
denser and becoming tomentellous along the lateral
veins and midrib; lateral veins 6 to 9 pairs, vena¬
tion immersed on the upper surface, midrib and
lateral veins raised, tertiary venation weakly raised
on the lower surface. Petioles 8-19 mm long, with
a similar indument to the twigs. Inflorescences ax¬
illary, 5-15 cm long, densely brown tomentellous,
paniculate-cymose. Flowers green, ca. 7 mm diam.,
the tepals ca. 3 mm long, ovate-elliptic, spreading,
pubescent at both surfaces. Staminate flowers with
nine 4-celled stamens, the outer 6 stamens ca. 2.5
mm long, the filaments ca. 1.5 mm long, moderately
pubescent, the anther cells introrse, the upper 2
largely overlapping with the lower 2; inner 3 sta¬
mens of similar size, but anthers with 2 cells ex-
trorse and 2 cells lateral; base of tbe filaments of
the inner 3 stamens seemingly fused; 2 globose
glands present at the base of the inner stamens;
staminodia not seen; pistillode slender, ca. 2 mm
long, pubescent, stigma present. Pistillate flowers
with 9 staminodes, these ca. 1 mm long, the fila¬
ments sparsely pubescent or glabrous, glands pres¬
ent at the base of the inner 3 staminodes. pistil 3
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2003
van der Werff
Lauraceae from South America
353
mm long, the ovary glabrous, ca. 1.2 mm long, the
style in bud folded in an S shape, becoming straight
in open flowers, pubescent, receptacle glabrous in¬
side. Fruits round to broadly ellipsoid, ca. 1.3 X
1.2 cm, seated on a thick, flat, plate-like cupule
with a double margin.
Flowers in May; fruits in September—December.
Ocotea hirtostyla is closely related to the wide¬
spread O. floribunda and its allies, which are char¬
acterized by having pubescent styles and plate-like,
thick cupules with a double margin and often some¬
what persistent tepals on the cupules. Ocotea hir¬
tostyla differs from the other species in this group
by its tomentellous twigs anti the soft, erect hairs
on the lower surface of the leaves. It is currently
only known from the Napo province in Amazonian
Ecuador; all but one collection come from the Jatun
Sacha reserve where collecting has been relatively
intense. It can be expected elsewhere in Amazonia
at the foothills of the Andes.
Paratypes. ECUADOR. Napo: Estacion Biologica Ja¬
tun Sadia, 6 May 1990. Palacios et al. 4930 (MO). 13
May 1990 (pist. ff). Palacios & Freire 5106 (MO), ft May
1990 (pisl. 11), Palacios & Freire 5146 (MO), ft May 1990
(stam. fl). Palacios & Freire 5160 (MO), 8 Sep. 1992 (fr),
Palacios et al. 10402 (MO), 6 Dec. J988 (fr), Palacios
3286 (MO). Sep. 1988 (fr), Palacios 3003 (MO); Rio Ara-
juno. Sola Coclia, Oct. 1995 (fr), Neill et al. 6941 (MO):
Napo, Cabeceras de la Quebrada Apayacu, 15 Dec. 1992
(fr), Zuleta 63 (MO).
4. Ocotea multinervis van der Werff, sp. nov.
TYPE: Peru. Dept. Piura: Prov. Morropon,
Carretera entre Chalaco y San Miguel, 2000
m, 18 Oct. 1988. C. Diaz & R. Vazquez 3030
(holotype, MO 5593912; isotypes, HRG, k.
NY, US). Eigure 13.
A congeneris nervis lateralibus numerosis et superficie-
bus ambabus tepalorum dense tomentellis recedit.
Trees, to 35 m. Twigs angular or ridged, 6-10
mm diam. 5 cm below the apex, densely tomentel¬
lous, glabrescent with age; terminal buds densely
tomentellous. Leaves alternate or grouped near the
tips of the branches, 9-17 X 4.5—8 cm, coriaceous,
elliptic or broadly elliptic, the base cordate to ob¬
tuse, the apex obtuse, rarely somewhat acute, the
upper surface glabrous, the lower surface densely
tomentellous, the indument covering the surface
completely; midrib immersed, lateral veins (slight¬
ly) impressed and tertiary' venation not visible on
the upper surface, midrib and lateral veins promi¬
nently raised, tertiary venation inconspicuously
raised or not visible on the lower surface; domatia
lacking; lateral veins 14 to 21 on each side; peti¬
oles short, to 5 mm long, but leaves often appearing
sessile. Inflorescences in the axils of normal leaves
or eataphylls, 5-14 cm long, densely tomentellous,
the surface completely covered by the indument,
paniculate-cymose. Flowers perfect (or unisexual?),
1 cm diam.; tepals spreading or reflexed at anthe-
sis, elliptic, 3 mm long, densely tomentellous on
both surfaces, the surface not visible, stamens 9,
all 4-celled, the outer 6 ca. 1 mm long, sessile or
nearly so, the upper pair of cells introrse, the lower
pair almost latrorse, the anther dorsally pubescent,
inner 3 with a short (about half as long as the an¬
ther) filament, the lower pair of cells extrorse, the
upper lateral; glands at the base in the inner sta¬
mens sessile, globose; staminodia minute, stipiti-
form, with a few hairs; pistil glabrous, 1.5 mm long,
receptacle initially rather deep, with some ap-
pressed hairs inside, but soon becoming shallow.
Fruit ellipsoid, 2 X 1.5 cm, cupule very shallow,
almost plate-like, ca. 1.5 cm diam., the pedicel
slightly thickened and abruptly widened into the
cupule.
Flowers in May and October; fruits in September.
Ocotea multinervis stands out by its tomentellous
leaves with many pairs of lateral veins and by its
tepals, densely tomentellous on both surfaces. It is
a tall tree in montane forests between 2000 and
3000 m. Its relationships within Ocotea are not ob¬
vious. The two flowering collections (the type and
Cuamacas 81) have anthers with open anther cells,
but without any pollen grains remaining. Thus, I
am not convinced that the flowers are hermaphro¬
dite; it is possible that the flowers are unisexual,
pistillate in the two flowering collections and that
the staminodia are relatively well developed. Ad¬
ditional collections will show whether this is the
case. In some flowers the glands at the base of the
inner three stamens are rather strongly enlarged
and protrude somewhat between the outer stamens,
a character found in Pleurothyrium. However, all
Pleurothyrium species with spreading tepals also
have the glands fused in a disc and surrounding
the stamens. In the new species the glands are not
fused in a disc and therefore I prefer to place it in
Ocotea, an already variable genus. One of the para¬
types ( Cuamacas et al. 81) was previously identi¬
fied as Ocotea arnottiana (Nees) van der Werff and
distributed under that name. A collection from Co¬
lombia ( Gentry & Cuadros 79141, MO) resembles
O. multinervis in leaf characters, but the young
fruits and remnants of the tepals are nearly gla¬
brous. Possibly it represents a related species, but
more material is needed to solve its identity.
Paratypes. PERU. Piura: Montana de Cuyas, 8 kin
NE of Ayabaca, 26 Sep. 1991 (fr). Gentry et al. 75116
(MO). ECUADOR. Imbahura: Caserfo Tablachupa, Par-
N? 5593913
354
Novon
Volume 13, Number 3
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van der Werff
Lauraceae from South America
355
roquia Plaza Gutierrez, 13 May 1992 (fl), Cuamaeas el al.
81 (MO. QCNE). COLOMBIA. Quindio: Keserva del Alto
Quindio Acaime, 10 July 1990 (fr), Renjifo 143 (MO).
5. Ocolea scalariformis van der Werff, sp. nov.
TYPE: Ecuador. Napo: Orellana Canton, Ya-
suni National Park, I). Neill 10426 (holotype,
QCNE 10426; isotype, MO). Figure 14.
Ocoteae tabacifoliae similis, sed basi foliorum decur-
renti, fructibus ovoideis. venatione scalariforme et floribus
subsessilibus recedit.
Dioecious trees, to 25 m. Twigs roundly angular
to rounded, densely red-brown pubescent, the hairs
erect and completely covering the twigs, terminal
buds densely red-brown tomentose. Leaves 16—35
X 9—1 1 cm, alternate, (irmly ehartaceous, elliptic
or broadly elliptic, rarely slightly ovate, the base
acute and somewhat decurrent on the petiole, the
margin slightly revolute, the apex acuminate, the
acumen to 2 cm long, or appearing obtuse (but then
probably damaged), the upper surface glabrous but
with traces of pubescence on the major veins, the
lower leaf surface moderately densely pubescent,
the hairs erect, discernable to the touch, most of
the surface visible, the indument denser and to¬
mentose along the major veins, domatia lacking,
midrib, lateral veins, and tertiary venation im¬
pressed on the upper surface, prominently raised
on the lower surface, the tertiary veins connecting
the lateral veins under an angle of 90°, and forming
a regular, ladder-like pattern; lateral veins 12 to 15
on each side, petioles 1-1.5 cm, with a similar in¬
dument to the twigs, canaliculate. Inflorescences
20-30 cm, panieulate-cymose, red-brown tomen¬
tose. the surface completely covered, in the axils of
leaves. Flowers 4—5 mm diam., moderately to
densely puberulous outside, unisexual; pedicels 1—
1.5 mm, shorter than the (lowers, bracts subtending
the flowers often persistent at anthesis and some¬
times even after flowers have fallen off; tepals 6.
equal, 2 mm long, elliptic, moderately to densely
puberulous outside, ± spreading at anthesis, mod¬
erately pubescent on the inner surface. Staminate
flowers: stamens 9. all 4-celled, ca. I mm, the fil¬
ament ca. 0.1 mm, very short, pubescent, the anther
glabrous, rectangular, the cells arranged in 2 pairs,
introrse; inner 3 stamens 1.5 mm, the filament ca.
0.4 mm, pubescent, the anther glabrous, cells ±
lateral; base of the filament with 2 small, globose
glands; staminodia not seen; pistillode 1.2 mm, gla-
<—
brous, the base swollen and containing an ovule¬
like body, pistillode with a small stigma, reaching
in the flower as high as the base of the anthers;
receptacle bowl-shaped, glabrous inside. Pistillate
flowers: tepals as in staminate flowers; staminodia
9. ca. 0.5 mm, the short filament pubescent, oth¬
erwise glabrous, glands present at the base of the
inner 3 staminodes; pistil 1.5 mm, glabrous, stvle
ca. 0.6 mm, stigma platelike, a little wider than the
style; receptacle deeply cup-shaped, with some ap-
pressed hairs inside. Fruits 20 X 13 mm, ellipsoid
to ovoid, the cupule small, platelike, scarcely more
than an undulating margin of the thickened pedi¬
cel.
Flowers in June and July: fruits in December.
Ocolea scalariformis can be confused with (). ta-
bacifolia (Meissner) Rohwer. Both species have
large, pubescent leaves with erect hairs, tomentose
twigs, and have been reported from the Amazon
lowland rain forest. However, 0. scalariformis dif¬
fers in its leaf bases, which are decurrent on the
petiole (obtuse to rounded in O. tabacifolia), in its
impressed venation on the upper leaf surface, in
the prominently raised, scalariform venation on the
lower leaf surface, in the ellipsoid to ovoid fruits
(rounded in O. tabacifolia), and in the pistillode
with a distinctly swollen base (pistillode threadlike
in O. tabacifolia). The pistillode with a swollen
base and the shape of the fruits do not suggest a
close relationship with 0 . tabacifolia, but point to
the 0. minarum group of Rohwer (1986). The spe¬
cies of this group are vegetatively variable, but all
share clearly pistillate and seemingly bisexual (fer¬
tile stamens and a pistil or pistillode with a swollen
base) flowers and ellipsoid fruits seated on a small,
platelike cupule. None of the species in this group
resembles 0 . scalariformis vegetatively, but I feel
that, based on flower and fruit characters, the new
species belongs in the 0 . minarum group.
Paratypes. ECUADOR. Francisco tie Orellana: Yu-
suni Scientific Station, 2 Dec*. 1997 (fr). Romoleroux et al.
3028 (MO), 15 July 2000 (pist. fl). Villa 378 (MO).
VI. Pleurothyrium Nees, in Findley, Nat. Syst.
Bot. 442. 1836.
Pleurothyrium is a Neotropical genus recently
revised by van der Werff (1993) who included 40
species in the genus. It is easily recognized by the
enlarged glands at the base of the inner stamens.
f igures 13, 14. —13 (lelt). Holotype of Ocolea multinervis van der Werff.
van der Werff. Insert: Cupule and fruit.
14 (right). Isotype of Ocotea scalariformis
356
Novon
N? 5593914
Ydmay*k*tBo%que prinur10,
04'35'S 78*19'W 380 ».
Arbolito d» fe m. de alto, floras
11 Qctubr* 1995
N.Jaramillo y O.Chamik 850
MISSOURI BOTANICAL GARDEN HERBARIUM
Figure 15. Holotype of Pleurothyrium arcuatum van tier Werff.
Often the glands protrude between the outer sta¬
mens and form a disc enclosing the stamens. Sev¬
eral species, including the one described below,
have a pronounced brochidodromous venation.
1. Pleurothyrium arcuatum van der Werff. sp.
nov. TYPE: Peru. Depto. Amazonas: Prov. Bag-
ua, Yamayakat, 320 m, I I Oct. 1995, /V. Jar-
amillo & I). Chamih 850 (holotype, MO
5593914; isotypes, HBG, K, NY, US). Figure
15.
A congeneris foliis glabris, alternis, apice acuminatis,
venatione brochidodroma, ramulis solidis, juvenalibus
castaneo-tomentellis recedit.
Small tree. 6 m. Twigs terete, solid, the apical
part with the inflorescences densely brown tomen-
tellous, the older parts glabrous, with a pale bark;
terminal buds densely brown tomentellous. heaves
10—20 X 4.5—7 cm, elliptic, coriaceous or firmly
chartaceous, alternate, evenly distributed, the base
acute or obtuse, the margin flat, the apex acuminate
or acute, the acumen I cm long, glabrous on both
surfaces except for some remnants of the tomentel¬
lous indument along the midrib, midrib immersed
or slightly raised, lateral veins impressed, tertiary
venation immersed on the upper surface, midrib
and lateral veins raised, tertiary" venation slightly
raised on the lower surface; lateral veins conspic¬
uously loop-connected, 8 to 12 on each side; peti¬
oles 1.5—2 cm, with a similar indument to the twigs,
flat on the upper side. Inflorescences 4—10 cm.
Volume 13, Number 3
2003
van der Werff
Lauraceae from South America
357
moderately to densely brown-tomentellous. panie-
ulate-cymose. in the axils of bracts near the tips of
the branches. Flowers 7—8 mm diam., yellow, ped¬
icels 2—3 mm long; tepals 6, ca. 3 mm long, elliptic,
equal or the inner 3 slightly narrower than the outer
3, spreading in older flowers, the outer surface
brown tomentellous, distally becoming papillose,
inner surface papillose, stamens 9, 4-celled, the
outer 6 with the anthers bent inward and the cells
lateral, the inner 3 with the anther bent toward the
outside and the cells lateral; the 2 cells on each
side of the anther opening back to back; glands
enlarged and forming a ring surrounding the sta¬
mens and obscuring the filaments; staminodia not
seen; pistil ca. 2 mm, with a few hairs, the style
short, ca. 0.5 mm; receptacle deeply cup-shaped,
pubescent inside. Fruits and cupules not known.
Flowers in October.
In the key of van der Werff (1993), P. arcuatum
would key to the last four species, based on its
enlarged glands surrounding the stamens, glabrous,
alternate leaves, and the papillose inner surface ol
the tepals. The conspicuously loop-connected lat¬
eral veins immediately set the new species apart in
this group. Pleurothyrium arcuatum is only known
from the type collection, made in the vicinity of the
Maranon in northern Peru. The epithet arcuatum
refers to the arcuate, loop-connected lateral veins.
Acknowledgments. Fieldwork by the author was
supported by National Geographic Society grant
#3480-86 and by a grant from the Mellon Foun¬
dation; their support is gratefully acknowledged. I
also thank my colleagues who have sent me gifts
for identification during the past two decades. Fred
Keusenkothen kindly provided the images of the
new species.
Literature Cited
Chanderbali, A. S., H. van der Werff & S. Renner. 2001.
Phylogeny and historical biogeography of Lauraceae:
Evidence from chloroplast and nuclear genomes. Ann.
Missouri Rot. Card. 88: 104—134.
Greuter, W., J. McNeill, E. T. Rarrie, H. M. Rurdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
J. E. Skog. P. Trehane, N. J. Turland N I). E. Hawk-
sworth (editors). 2000. International Code of Rotanical
Nomenclature (Saint Louis Code). Regnurn Veg. 138.
Kubitzki, K. & H. (». Richter. 1987. Williamodendron ku-
bitzki & Richter, a new genus of Neotropical Lauraceae.
Rot. Jahrb. Svst. 109: 49-58.
kurz. H. 2000. Revision der Gattung Licaria (Lauraceae).
Milt. Inst. Allg. Rot. Hamburg 28/29: 89—221.
Lorea Hernandez, F. G. 1996. A Systematic Revision of
the Neotropical Species ol Cinnamomum Schaeffer
(Lauraceae). Ph.D. Thesis, University of Missouri, St.
Louis.
Meissner, C. F. 1864. Lauraceae. In: A. de Candolle (ed¬
itor), Prodromus systematis naturalis regni vegetabilis
15(1): 1-260.
-. 1866. Lauraceae. In: C. F. P. von Martius (editor).
Flora Rrasiliensis 5(2): 138—295.
Rohwer, J. G. 1986. Prodromus einer Monographic der
Gattung Ocotea Aubl. (Lauraceae) sensu lato. Mitt. Inst.
Allg. Rot. Hamburg 20: 1-278.
-. 1993. Nectandra. FI. Neotrop. 60: 1—333.
van der Werff, H. 1987. A revision ol Mezilaurus (Laura¬
ceae). Ann. Missouri Rot. Card. 74: 153—182.
-. 1993. A revision of the genus Pleurothyrium
(Lauraceae). Ann. Missouri Rot. Gard. 80: 39—119.
-. 2002. A synopsis of Ocotea (Lauraceae) in Cen¬
tral America and southern Mexico. Ann. Missouri Rot.
Gard. 89: 429-451.
Revised Nomenclature and a New North American Record for the
Villose Cliff Brake (Pellaea, Pteridophyta)
Michael I). Windham
Utah Museum of Natural History, University of Utah, Salt Uake City, Utah 84112-0050, U.S.A.
windham@umnh.utah.edu
George Yatskievych
Missouri Botanical Garden, P.O. Box 299, St. Uouis, Missouri 63166-0299, U.S.A.
george.yatskievych@mobot.org
Abstract. Originally described hy Fee in 1857.
the villose cliff brake was not validly published for
another 136 years. In 1993, this member of the
Pteridaceae (sen.su Try on & Tryon) was named Pel¬
laea ternifolia subsp. villosa Windham, acknowl¬
edging its overall similarity to the ternate cliff
brake. Although formally including it within P. ter¬
nifolia (Cavanilles) Link, Windham suggested that
the villose cliff brake was a probable allotetraploid
produced by hybridization between typical P. ter¬
nifolia and some other (then unidentified) taxon.
Recent work by Mendoza et al. has identified a new
species, P. ribae Mendoza & Windham, that ap¬
pears to be the "missing parent." Considering the
evidence that the villose cliff brake is a reproduc-
tively competent taxon that originated as a hybrid
between P. ternifolia subsp. ternifolia and P. ribae,
we now feel that it should be recognized as a dis¬
tinct species. Herein, we propose a new combina¬
tion, Pellaea villosa (Windham) Windham & Yat¬
skievych, for the villose cliff brake. We also report
a northern range extension to Arizona, the second
record for the area covered by the Flora of North
America.
Key words: Arizona, North America, Pellaea,
Pteridaceae.
While preparing a taxonomic treatment of Pel¬
laea for Volume 2 of the f lora of North America
Project (Windham. 1993b), the senior author de¬
termined that U.S. populations assigned to Pellaea
ternifolia (Cavanilles) Link (Pteridaceae: sublam.
Cheilanthoideae, sensu Tryon & Tryon, 1982) rep¬
resent "three genetically distinct taxa characterized
by differences in leaf morphology and chromosome
number" (Windham, 1993a: 42). The most distinc¬
tive element, a tetraploid with sparsely villous
fronds, was segregated as P ternifolia (Cavanilles)
Link subsp. villosa Windham, the villose cliff
brake. Reported from a single locality within the
boundaries of the Flora (Windham, 1993b), this
taxon is relatively widespread in Mexico and prob¬
ably originated there (Windham, 1993a). Consid¬
ering this distribution, it is not surprising that sub¬
sequent work on the pellaeas of Mexico (Mendoza
et al., 2001; Mickel, in press) has substantially im¬
proved our understanding of P. ternifolia subsp. vil¬
losa. These new data require a reconsideration of
both the taxonomy and the known distribution of
this taxon.
Pellaea ternifolia was first described (as Pteris
ternifolia Cavanilles) in 1802. It has the most ex¬
tensive distribution of any species in the genus,
ranging from the southwestern United States to
southern Chile, with disjunct populations in Hawaii
and Hispaniola (Tryon, 1957). Within this vast
range, there is considerable habitat diversity and
ample opportunity for geographic isolation. These
factors have combined to produce a puzzling array
of morphotypes and an equally complex synonymy
(Tryon, 1957: 150). Nearly all of the synonyms and
proposed segregates have essentially glabrous
leaves like the type material of P. ternifolia from
Peru (Nee s.n .; lectotype [designated by Christen¬
sen. 1937]: MA not seen; isoleetotype: F). The first
author to discuss the villose cliff brake per se was
fVe (1857), who provided an accurate description
and the provisional name “P. lanuginosa.” Although
impressed by the apparent distinctness of this tax¬
on, Fee merely discussed it under P. ternifolia and
stated that he wanted to see more material before
recognizing it formally. Thus, the name “P. lanu¬
ginosa" Fee was considered a nomen provisorium
by Tryon (1957), and has never been validated.
Since Fee’s initial recognition of the villose cliff
brake, its distinctive nature has become more evi¬
dent. Alice Tryon noted a correlation between chro¬
mosome number and leaf morphology in Pellaea
ternifolia , stating that "the tetraploid plants gen-
Novon 13: 358-362. 2003.
Volume 13, Number 3
2003
Windham & Yatskievych
Villose Cliff Brake
359
Table 1. Comparison of selected taxa belonging to the /
of P. villosa. The characters listed for P. ternifolia apply only
1 ellaea ternifolia comple
to the diploid cytotype i
x demonstrating the intermediacy
of subspecies ternifolia (see text).
Characters
I’ ternifolia
P. villosa
P. ribae
Rhizome scale shape
Linear-subulate
Linear
Linear-filiform
Rhizome scale length
5—7 mm
6—12 mm
8-15 mm
Rhizome scale coloration
Bicolored
Mostly hicolored
Concolored
Petioles and rachises
Glabrous or with a
few scattered tri -
chomes
± Villous
Densely villous
Distal portion of petioles
Grooved or flattened
adaxially
Rounded to slightly
flattened adaxially
Rounded adaxially
Pinnae
Ternately divided ex¬
cept at leaf apex
Usually entire for dis¬
tal Vi—Yi
Entire throughout
Ahaxial pinna surfaces
Glabrous
Sparsely villous
Densely villous
Apical mucro on pinnae
Well-developed
Weakly developed
Absent
Fargest ultimate segments
Usually <18 mm
long
Usually >18 mm
long
Usuallv <18 mm long
Mean spore size
39-45 gm
46—53 gm
53—60 gm
Chromosome number
n = 29 (diploid)
n = 58 (tetraploid)
Unknown
erallv resemble the diploids, but the pinnae tend
to be entire and the rachises more pubescent"
(Tryon, 1972: 230). Based on isozyme profiles and
additional chromosome studies, the senior author
suggested that the “tetraploids are segmental allo¬
polyploids produced by hybridization between typ¬
ical P. ternifolia and other (as yet unidentified) dip¬
loid elements within the P. ternifolia complex"
(Windham, 1993a: 42). With no specimens of the
“missing parent” at hand, Windham (1993a,
1993b) took a conservative approach to the taxon¬
omy of the group and recognized the villose cliff
brake as a subspecies of P. ternifolia.
Subsequent work on the cliff brakes of Mexico
(Mendoza et al.. 2001; Mickel. in press) indicates
that this classification needs revision. Central to our
change in thinking is the discovery of Pellaea ribae
Mendoza & Windham (Mendoza el al.. 2001), the
probable “missing parent" of the villose eliff brake.
As shown in Figure 1 and Table 1. subspecies vil-
losa is morphologically intermediate between dip¬
loid P. ternifolia subsp. ternifolia and P. ribae in
nearly every character examined. From minute de¬
tails of the rhizome scales to the more obvious fea¬
tures of frond dissection and pubescence, the vil¬
lose cliff brake exhibits clear intermediacy. The
only exception is in the length of the largest ulti¬
mate segments. These usually exceed Iff mm in
subspecies villosa and are usually less than 18 mm
in both diploid P. ternifolia subsp. ternifolia and P.
ribae (Table 1). Such increases in frond segment
size are common in hybrid ferns (e.g., Windham.
1988; Paler & Barrington, 1995) and may be at¬
tributable to heterosis (Grant, 1975).
Although morphological intermediacy is expect¬
ed in taxa of hybrid origin, it rarely extends to mean
spore size, as it does in the case of the villose cliff
brake (Table 1). In general, allotetraploid ferns pro¬
duce spores that are somewhat larger than either
parent due to increased DNA content. The lack of
a chromosome count for Pellaea ribae leaves open
two possibilities. If spore sizes are strictly correlat¬
ed with ploidy level in this group, then the mate¬
rials of P ribae currently at hand likely represent
a polyploid derivative of the “missing parent" of
subspecies villosa. Alternatively, P. ribae may ex¬
emplify a rare exception to the correlation between
chromosome number and spore size. The species
appears to be a strict gypsophile, and larger prop-
agules might provide an adaptive advantage in such
a harsh habitat. Notholaena bryopoda Maxon, an¬
other strict gypsophile that is known to be diploid,
also bas exceptionally large spores (Windham, un¬
published). Plants of P. ribae derived from spores
of the type collection currently are being grown at
the Missouri Botanical Garden in hopes of testing
these hypotheses.
Regardless of the actual chromosome number of
Pellaea ribae, the data at hand indicate that the
features distinguishing subspecies villosa from typ¬
ical P. ternifolia derive from a genome shared with
P. ribae. This genome is strikingly divergent from
that of diploid P. ternifolia, confirming the sugges¬
tion (Windham, 1993a) that the villose cliff brake
is an allotetraploid. Although subspecies villosa
was formed originally by interspecific hybridization
between P. ternifolia and P. ribae, we contend that
it is a reproductively competent allopolyploid taxon
360
Novon
Figure 1. A—C. Diploid P. ternifolia (Cav.) Link subsp. ternifolia drawn from Mickel 3849 (NY). —A. Leaf and rhizome.
—B. Racbis and pinna attachment (adaxial surface). —C. Rhizome scale. D—F. P. villosa (Windham) Windham &
Yatskievych drawn from Rzedowski 22894 (NY). — 1). Leaf and rhizome. —K. Rachis and pinna attachment (abaxial
surface). — F. Rhizome scale. G—I. P. ribae Mendoza & W indham drawn from Heil 5247 (BRY). —G. Leaf and rhizome.
—H. Rachis and pinna attachment (abaxial surface). —I. Rhizome scale. Scale bar = 5 cm (A, I). G), 1 mm (B, C,
E, F. H). Adapted from Mickel (in press).
Volume 13, Number 3
2003
Windham & Yatskievych
Villose Cliff Brake
361
lliat should be treated taxonomically as a separate
species. It is amply distinct morphologically from
both P ternifolia and P ribae (Table 1).
The appendix on nothotaxa in the International
Code of Botanical Nomenclature (Greuter et al..
2000) states in Article H.5.1. “The appropriate
rank of a nothotaxon is that of the postulated or
known parent taxa." However, Article 3.2 states, “If
the postulated or known parent taxa are of unequal
rank, the appropriate rank of the nothotaxon is the
lowest of these ranks.” Given that we postulate that
P villosa arose following hybridization between a
subspecies of P. ternifolia and the species P ribae,
these statements might be interpreted to require the
continued recognition of P. villosa at the subspecific
rather than specific level. We draw r attention to
problems of interpretation created by this section
of the Code, especially as applied to fern nomen¬
clature. The present practice among biosystem-
atists, including most pteridologists, is to distin¬
guish between primary hybrids and the
reproductively competent taxa (sensu Wagner,
1954) that evolve from them following polyploidi-
zation. Such allopolyploids are reproductively iso¬
lated from either parent and function as full spe¬
cies; therefore we believe they should not be
treated nomenclaturally as hybrids.
Although we considered validating Fee's nomen
provisorium for this taxon, we have not seen the
type (staff at P were unable to locate the specimen).
Thus we are uncertain as to the precise application
of the name. We therefore propose the following
new combination based on the epithet applied by
Windham (1993a):
Pellaea villosa (Windham) Windham & Yatskiev¬
ych, comb, et stat. nov. Basionym: Pellaea ter¬
nifolia (Cavanilles) Link subsp. villosa
Windham, Contr. LJniv. Michigan Herb. 19: 43.
1993. TYPE: Mexico. Hidalgo: rocky hills,
Lena Station, 8300 ft., 24 Aug. 1905, C. G.
Pringle 10025 (holotype, LL; isotypes, ARIZ,
BRY, CAS, COLO, DAO, ENCB, F, GH. INI),
KANG, LL, MSC. OKLA, SMU, TEX, UC, US,
VT, WIS).
This species, occurring on various rocky sub¬
strates, is widespread in the Sierra Madre Occiden¬
tal and Sierra Madre Oriental of northern and cen¬
tral Mexico. Mickel (in press) has identified
specimens from the states of Chihuahua, Coahuila,
Durango, Guanajuato, Hidalgo, Mexico. Michoacan,
Nuevo Leon, Puebla, San Luis Potosi, and Sonora.
In the United States, the villose cliff brake has been
reported from a single locality in the Davis Moun¬
tains of western Texas (Windham, 1993b). However,
recent botanical surveys of the Madrean “sky is¬
lands” of southern Arizona have identified a second
U.S. population:
U.S.A. Arizona: Cochise Co., Huachuca Mountains,
Lone Mountain Watershed, Sycamore Canyon, above Mud
Spring, 31 "24'N I 10'23Vi'W elev. 5700 ft.. E-facing slope
just above arroyo bottom in pine-oak woods with Juniperus
deppeana, Quercus emoryi, Eragrostis, Muhlenbergia emer-
sleyi, Cyperus, Bouteloua, Tradescantia, Ranunculus ari-
zonicus, Ipomoea, Calliandra, Bommeria, terrestrial; un¬
common, 1 1 Aug. 1996, M. Fishbein 2861, with K. Hooper
& R. Coleman (ARIZ).
This new population is situated approximately
300 km north to northwest of the closest known
localities in the Sierra Madre Occidental and 600
km west-northwest of the only other U.S. collection
site. It remains to be seen whether Pellaea villosa
will be found in the mountains of southern New
Mexico, which have fewer Madrean habitats. At the
moment, it belongs to a group of species (including
the ferns Cheilanthes lendigera (Cavanilles) Swartz,
Notholaena aschenborniana Klotzsch, N. neglecta
Maxon, and P. ternifolia (Cavanilles) Link subsp.
arizonica Windham) that are known from Arizona
and trans-Pecos Texas but not from intervening re¬
gions of New Mexico.
Acknowledgments. We are grateful to John
Mickel for sharing his extensive knowledge of the
taxa under consideration and for permission to use
illustrations (skillfully drawn by Haruto Fukuda)
from his upcoming book on the ferns and allied
plants ol Mexico. We thank Mark Fishbein for per¬
mission to publish his discovery of Pellaea villosa
in Arizona, and express our gratitude to the cura¬
tors of the cited herbaria for loans and access to
their collections. Our reviewers, R. James Hickey,
Gerald J. Gastony, Ruth Kirkpatrick, and Alan R.
Smith, provided insightful comments that improved
our discussion greatly. We also w ish to acknowledge
the Utah Museum of Natural History (UMNH) and
the New York Botanical Garden (NY) for providing
funding that made this work possible. Finally, we
thank Jill Schwartz for her help with scanning and
assembling the figure.
Literature Cited
Christensen, C. 1937. Taxonomic fern studies III. Revi¬
sion of the genera and species of ferns described by A.
J. Cavanilles. Dansk Rot. Ark. 9(3): 3—32, fig. 1. tab.
1-L
Fee, A. L. A. 1857. Iconographie des espfeces nouvelles
decrites on enumerees dans le genera filicum et revision
des publications ant6rieures relatives t) la famille des
fougeres, 111. Memoires sur ies Families de Fougeres 8:
67—138. Veuve Berger-Levault et fils, Strasbourg.
362
Novon
Grant. V. 1975. Genetics of Flowering Plants. Columbia
Univ. Press, New York.
Greuter, W.. J. McNeill. F. li. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). H. Nicolson, P. C. Silva.
J. K. Skog. P. Trehane, N. J. Turland & 1). I,. Hawk-
sworth. 2000. International Code of Botanical Nomen¬
clature (Saint Louis Code). Regnum Veg. 138.
Mendoza, A., M. Windham, B. Perez-Garcfa «S G. Yat-
skievych. 2001. Una nueva especie de Pellaea (Pteri-
daceae) del estado de San Luis Potosf, Mexico. Acta
Bot. Mex. 57: 15-21.
Mickel, J. T. In press. Pellaea. In J. T. Mickel & A. R.
Smith, Pteridophyte flora of Mexico. Mem. New York
Bot. Garth
Paler. M. H. & I). S. Barrington. 1995. The hybrid Cys-
topteris fragilis X C. tenuis (Dryopteridaceae) anti the
relationship between its tetraploid progenitors. Syst.
Bot. 20: 528-545.
Tryon, A. F. 1957. A revision of the fern genus Pellaea
section Pellaea. Ann. Missouri Bot. Garth 44: 125—193.
-. 1972. Spores, chromosomes anti relations of the
fern Pellaea atropurpurea. Rhodora 74: 220—241.
Tryon, R. M. & A. F. Tryon. 1982. Ferns anti Allietl
Plants, with Special Reference to Tropical America.
Springer-Verlag, New' York.
Wagner, W. H., Jr. 1954. Reticulate evolution in the Ap¬
palachian aspleniums. Evolution 8: 103—118.
Windham, M. I). 1988. The Origin and Genetic Diversi¬
fication of Polyploid Taxa in the Pellaea wrightiana
Complex (Adiantaceae). Ph.D. Dissertation, University
of Kansas.
-. 1993a. New taxa and nomenclatural changes in
the North American fern flora. Contr. Univ. Michigan
Herb. 19: 31-61.
-. 1993b. Pellaea. Pp. 175-186 in Flora of North
America Editorial Committee (editors). Flora of North Amer¬
ica North of Mexico. Vol. 2. Oxford Univ. Press, New York.
A New Species and a New Specific Synonym of Pedicularis
(Scrophulariaceae) from I lie Hengduan Mountains, China
Yang Fu-Sheng, Hong De-Yuan, and Wang Xiao-Quan
Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, Beijing 100093, China. Hongdy@ns.ibcas.ac.cn
ABSTRACT. Pedicularis inflexirostris F. S. Yang, I).
\. Hong & X. Q. Wang, a new species from the
Hengduan Mountains, China, is described. Pedi¬
cularis inflexirostris is different from P. tatarinowii
Maximowicz in having more slender stems, fewer
branches, a cucullate middle lobe of the lower co¬
rolla lip, and glabrous filaments. Pedicularis deqi-
nensis H. P. Yang is reduced to a synonym of P
ramosissima Bonati on the basis of their common
characters of long and dense branches, recurved
calyx teeth, pilose indumentum on the inner margin
of the calyx, and cucullate middle lobe of the lower
corolla lip.
key words: China. Hengduan Mountains, Pc fl¬
ic ula ris, Scrophu lariaceae.
Pedicularis 1... consisting of about 500 species,
is the largest genus in the Scrophulariaceae. The
genus, confined to the Northern Hemisphere, is a
member of the arctic-alpine flora with the majority
of species occurring in meadows. Recorded in Chi¬
na are about 352 species, of which 214 are con¬
centrated in the Hengduan Mountains (western
Sichuan, eastern Tibet, and western Yunnan),
where most Pedicularis are endemic (Hong, 1983;
Yang et al., 1998).
Due to the large number of species, high diver¬
sity of (lowers, and extensive parallel evolution of
floral characters in Pedicularis, it is difficult to re¬
construct a natural intrageneric classification sys¬
tem. Quite a few influential but controversial sys¬
tems have been proposed (Steven. 1823; Bunge,
1841; Maximowicz, 1888; Bonati. 1910; Limpricht,
1924: Li. 1948. 1949; Tsoong, 1955, 1963), but Li’s
(1948. 1949) and Tsoong’s (1955, 1963) systems
are the most outstanding. Li’s system includes 3
groups, 22 sections, 79 series, and 282 species,
while Isoong’s includes 13 groups, 21 subgroups,
112 series, and 329 species (Tsoong, 1963). De¬
spite so many classification systems proposed, tax¬
onomic research on Pedicularis in the eastern Him¬
alayas, the modern diversity center of Pedicularis ,
is insufficient compared to the abundant Pedicu¬
laris species in the region.
In the summers of 2000 and 2001 we conducted
an extensive field investigation on Pedicularis in
the Hengduan Mountains and collected a large
amount of Pedicularis specimens. Based on exam¬
ination and identification of the specimens, we
found a new species and nominated it as Pedicu¬
laris inflexirostris F. S. Yang, D. Y. Hong & X. Q.
Wang on the basis of the indexed beak. According
to Li’s system, the new species should be placed in
the group Cyclophyllum sect. Orthosiphonia ser.
Myriophyllae Maximowicz, while in Tsoong’s sys¬
tem. it belongs to the grex Orthosiphonia Tsoong
ser. Myriophyllae Maximowicz. The series Myrio¬
phyllae w as founded bv Maximowicz (1878) and re¬
defined by Maximowicz (1888), Brain (1890), and
Limpricht (1924). Li (1948) narrowed the circum¬
scription of this series, removing Pedicularis curv-
ituba Maximowicz and P anas Maximowicz with
curved anti deflexed corolla tubes to form a new
series, series Curvitubae Li. Therefore, the series
Myriophyllae in Li’s system includes four species,
P. myriophylla Pallas, P. alaschanica Maximowicz,
P. tatarinowii, and P. provoti Franchet, character¬
ized by a dedexed beak and more or less straight
corolla tube. Tsoong (1963) broadened the circum¬
scription of the series Myriophyllae. adding a new
species Pedicularis pseudocurvituba Tsoong, and
transferring here P. scolopax Maximowicz and P
cristatella Pennell & Li from other series. Thus the
series Myriophyllae, according to Tsoong (1963), in¬
cludes seven species having a dedexed to straight
and horizontal beak. Shown in the following key are
the circumscription of the series and the distinct¬
ness of our new r species.
Key to the Species oe Series Myriophyllm-:
la. Beak of corolla less than 1 mm long; lateral calyx
lobes entire. I>. myriophylla
lb. Beak of corolla 2—5 mm long; lateral calyx lobes
serrate, pinnatifid, or entire.
2a. Beak of corolla 3—5 mm long; galea crested
. P. cristatella
2b. Beak of corolla 2—3 mm long; galea not
crested.
3a. ( Corolla purple-red; beak of corolla
strongly indexed.
Novon 13: 363-367. 2003.
364
Novon
4a. Middle lobe of lower lip cucullate;
filaments all glabrous . . P. inflexirostris
4b. Middle lobe of lower lip elongated,
not cucullate; filaments all pubes¬
cent . P. tatarinowii
3b. Corolla yellow; beak of corolla slightly
curved.
5a. Bracts all or at least proximal ones
longer than flowers; filaments all
glabrous or only 2 glabrous.
6a. Corolla 2—2.5 cm long; anterior
filaments villous, posterior ones
glabrous. P. alaschanica
6b. Corolla ca. 1.5 cm long; fila¬
ments all glabrous . . . P. scolopax
5b. Bracts all shorter than flowers; fil¬
aments all pubescent.
7a. Basal leaves persistent; stems
herbaceous, unbranched above;
calyx at least Yi ( left at anterior
. P. pseudocurviluba
7b. Basal leaves caducous; stems
± woody, short-branched
throughout; calyx barely 16
cleft at anterior .... P. curvituba
Pedicularis inflexirostris F. S. 'fang. I). V. Hong
& X. Q. Wang, sp. nov. TYPE: China. Tibet:
Jomda County, Mt. Wangna, dry slope meadow,
ea. 3700 m. 13 Ang. 2001, F S. Yang Y0180
(holotype, designated here, PE). Eigure 1.
Species P tatarinowii affinis, a qua caulibus tenuiori-
bus medio 1—2.5 mm diametro, ramis paucioribus usque
ad 4, galeae parte ereela graciliore 1.5 mm diametro, labio
infero minore 8—9 mm lato, lobo medio cucullato, fila¬
ment i s omnibus glabris differt.
Annuals, 20-45 cm tall, ± black when dry.
Rootstocks vertical, woody, 4—8 cm long. Steins sol¬
itary or sometimes caespitose, 1—2.5 mm diam. at
the middle part, erect, with 4 lines ol hairs, and 0
to 4 branches at middle and upper parts ol stems.
Basal leaves opposite, middle and upper leaves in
whorls of 4, long crisp pilose along petioles and
nerves; petioles 1—2 cm long; laminae oblong-lan¬
ceolate. 15—30 X 8—15 mm, pinnatisect; dentate
segments 10- to 13-paired, linear-lanceolate. Inflo¬
rescences to 15 cm long, interrupted at lower part;
lower bracts similar to upper leaves in shape, top
bracts dilated at the base, ovate. Calyx tube 5—7
mm long, membranous, with 10 thick veins and 5
triangular to lanceolate teeth, gray-tomentose. Co¬
rolla 13—15 mm long, pale purple, tube straight, as
long as or slightly longer than the calyx; erect part
of the galea 4—5 mm long, 1.4—1.5 mm wide, hor¬
izontal part of the galea strongly dilated to 4 mm
wide; beak bent downward and backward, ca. 2 mm
long; lower lip 6 X 8—9 mm, middle lobe cucullate,
x h as wide as lateral lips. Filaments glabrous. Cap¬
sules lanceolate.
Pedicularis inflexirostris resembles P. tatarinowii
(ser. Myriophyllae) in having pinnatisect laminae, a
straight corolla tube, and recurved beak, but dillers
from the latter in having more slender stems, fewer
branches at the middle part, and thinner helmet
(Table 1). The cucullate middle lobe ol the lower
lip and glabrous filaments are diagnostic characters
of the new T species, by which it is readily distin¬
guished from P. tatarinowii. The lower lip ol P. in¬
flexirostris is also similar to that of the species in
series Longicaules Prain (P. dielsiana Bonati, P.
longicaulis Franchet) in shape, but the pinnatisect
laminae, racemose inflorescences, and indexed
beak of the corolla show that P. inflexirostris is re¬
lated to the grex Cyclocladus ser. Myriophyllae
Maximow icz.
Habitat and distribution. Pedicularis inflexiros¬
tris is found in Garze County in western Sichuan,
and Jomda and Qamdo Counties in eastern Tibet,
at altitudes of 3700—3900 m, in meadows on dry
slopes.
Paratypes. CHINA. Tibet: Qamdo County, Mt. Kajila.
dry slope meadows, ca. 3700 m, 7 Aug. 2001. h. S. Yang
Y0150 (PE). Sichuan: Garze County, Mt. Zhuodala. dry
slope meadows, ca. 3900 in, 2 Aug. 2001, F. S. Yang
Y013I (PE).
Pedicularis ramosissima was described by Bonati
(1908). I.i (1948) placed it in the group Cyclophyl-
lurn sect. Orthosiphonia ser. Pectinatae Prain. while
Tsoong (1903) treated it in the grex Cyclocladus
subgrex Cyclocladus ser. Cradles Maximow icz. The
series Pectinatae was founded by Prain (1890) and
revised by Limpricht (1924). Li (1948) narrowed
the circumscription of the series to include five spe¬
cies. Pedicularis scolopax, P. moupinensis Franchet,
P. tantalorhyncha Franchet, P. atuntsiensis Bonati,
and P. ramosissima, characterized by dilated and
serrated bracts at the upper part of inflorescences,
and more or less straight and horizontal beak.
Tsoong (1963) redefined the series Pectinatae Prain
to include only one new species, Pedicularis rhyn-
chotricha Tsoong, with a twisted and pilose beak to
distinguish it from the species in the series Pectin¬
atae Prain of previous systems, transferring P. ra¬
mosissima to the series Cradles. Thus the series
Cradles, according to Tsoong’s system (1963), in¬
cludes two species. Pedicularis ramosissima and P.
gracilis Wallieh. characterized by excessive
branches at the upper part of stems and flowers
with an elongated beak. The circumscription ol the
series and the distinctness of the two species are
shown in the following key.
Volume 13, Number 3
2003
Yang et al.
Pedicularis from China
365
Figure 1. A-F. Pedicularis inflexirostris F. S. Yang, I). Y. Hong & X. (,). Wang. (Based on the collection F S. Yang
Y0150, Ph.) —A, 15. Habit. —C. Flower. —D. Lower lip. —E. Calyx. —F. Corolla and stamens. G, H. Pedicularis
tatarinowii. (Redrawn from figure 48 in Tsoong, 1963.) —G. Flower. —H. Corolla and stamens.
366
Novon
Table 1.
Diagnostic characters distinguishing Pedicularis inflexirostris from P. tatarinowii.
Number
of branches
Diam. of stems
at middle
part (mm)
Width of
helmet at
erect part
(mm)
Width of lower
corolla lip (mm)
Shape of middle
lobe of lower
corolla lip
Filaments
P. inflexirostris
0-4
1-2.5
1.4-1.5
8-9
cucullate
glabrous
P. tatarinowii
0-26
1.5-5
2.5-3
13-15
elongated, not cucullate
pubescent
Key to Species of Pei net iaris in Series Gkacii.es
la. Calyx teeth recurved, pilose on inner surface;
middle lobe of lower corolla lip cucullate ....
. P. ramosissima
II). Calyx teeth straight, glabrous on inner surface;
middle lobe of lower corolla lip not cucullate
. P gracilis
Pedicularis ramosissima Bonati, Bull. Sue. But.
France. 55: 246. 1908. TYPE: China. Sichuan:
Yargong, Aug. 1904, R. P. Soulid 5283 (holo-
type, P not seen, photo PE).
Pedicularis deqinensis II. P. Yang, Acta Phytotax. Sin. 28:
137. 1990. Syn. nov. TYPE: China. Yunnan: Deqen
County, 2900—3400 m, 21 Aug. 1981. Inst. Bat.
Acad. Sin. Hengduanshan Exped. 3332 (holotype,
PE; isotype. PE).
The original description of l\ ramosissima was
based on a single specimen, Soulid 5283. With
more specimens examined, the description is re¬
vised here, adding some characters.
Stems 20—90 cm tall. Petioles 5—20 mm long,
laminae 25—60 X 8—30 mm, dentate segments 5-
to 12-paired. Calyx tube ca. 5 mm long with 5 teeth
recurved. Corolla pale purple, 15-18 mm long,
lube straight: erect part of galea 6-8 mm long; low¬
er lip 8X12 mm, middle lobe cucullate.
Pedicularis deqinensis was noted by Yang (1990)
to be close to P. cristatella, and was grouped, with¬
out further explanation, into series Myriophyllae, in
which the latter also nested. Probably the elongated
beak and the crest on the helmet were characters
that prompted the author to relate P. deqinensis to
P. cristatella and then to the grex Orthosiphonia ser.
Myriophyllae. But in Pedicularis, convergent and
parallel evolution of the corolla prevails, so that it
may be unjustifiable to circumscribe intrageneric
laxa based only on differences in the corolla (l.i.
1951; Tsoong, 1955). Based on our observation of
the specimens (Inst. Hot. Acad. Sin. Hengduanshan
Exped. 3332) of P. deqinensis, long and dense
branches in whorls of 4 at the middle and upper
parts of stems and sparse, opposite or 3-verticillate
flowers clearly indicate its position in the grex Cy-
cloctadus ser. Graciles. Furthermore, all important
characters of P. deqinensis, such as habit, arrange¬
ment of branches and flowers, shape of leaf and
corolla, and size of calyx and corolla, are perfectly
consistent with those of P. ramosissima. In partic¬
ular. the recurved calyx teeth, pilose indumentum
on the inner margin of the calyx, and cucullate
middle lobe of the lower lip further indicate that P.
deqinensis should be merged into P ramosissima,
and these characters are the principal ones to dis¬
tinguish P. ramosissima from its close species in
the series Graciles.
Habitat and distribution. Pedicularis ramosis¬
sima is found in Deqen County in western Yunnan
and Zogang County in eastern Tibet, at altitudes of
2900—3700 m, under shrubs anti Quercus forests.
Additional specimen examined. CHINA, libel: Zo-
gang County, under shrubs at dry gravelly slope, ca. 3650
m. 18 Aug. 2001, E. S. Yang Y0182 (PE).
Acknowledgments. This work was supported by
the State Key Basic Research and Development
Plan of China (G2000046804). Vie are grateful to
two anonymous reviewers anti scientific editor V. C.
Hollowed for their valuable comments and sugges¬
tions, to Wang Wen-Tsai in the Laboratory of Sys¬
tematic and Evolutionary Botany, Institute of Bot¬
any, the Chinese Academy of Sciences (Beijing), for
his help in Latin, to Cai Shu-Qin for the drawing,
and to Ma Li-Ming for her help in many aspects.
Literature Cited
Bonati, G. 1908. Sur quelques Pedieulaires chinoises nou-
velles. Bull. Soe. Bot. France 55: 246.
-. 1910. Contribution it lYtude du genre Pedicular¬
is. Bull. Soe. Bot. France 57. Mem. 18: 1—35.
Bunge, A. 1841. Ueber eine neue Art der Gattung Pedi¬
cularis. Bull. Acad. Imp. Sei. Saint-Petersbourg 8: 241-
253.
Hong, D. Y. 1983. The distribution of Scrophulariaeeae in
the Holarctic with special reference to the floristic re¬
lationships between eastern Asia and eastern North
America. Ann. Missouri Bot. Gard. 70: 701—712.
Li, H. L. 1948. A revision of the genus Pedicularis in
China I. Proc. Acad. Nat. Sci. Philadelphia l(M): 205-
378.
-. 1949. A revision of the genus Pedicularis in Chi¬
na II. Proc. Acad. Nat. Sei. Philadelphia 101: 1—214.
Volume 13, Number 3
2003
Yang et al.
Pedicularis from China
367
-. 1951. Evolution in the flowers ol Pedicularis.
Evolution 5: 158-164.
Eimpricht, W. 1924. Studien uber die Gattung Pedicularis.
Repert. Spec. Nov. Regni Veg. 20: 161—265.
Maximowicz, C. J. 1878. Diagnoses plantarum novarum
Asiaticarum (2). Bull. Acad. Imp. Sei. Saint-Peters-
bourg 24: 26—89.
-. 1888. Diagnoses plantarum novarum Asiaticarum
(12). Bull. Acad. Imp. Sci. Saint-Petersbourg 82: 477-
629.
Prain, I). 1890. The species of Pedicularis of the Indian
Empire and its frontiers. Ann. Roy. But. Card. (Calcut¬
ta) 3: 1-196.
Steven, C. 1823. Monographia Pedicularis. \1em. Soc.
Imp. Naturalistes Moscou 6: 1-60.
Tsoong, P. C. 1955. A new system for the genus Pedicu¬
laris. Acta Phytotax. Sin. 4: 71-147.
-. 1963. Scrophulariaceae (Para II). Pp. 1-378 in
S. S. Chien & W. Y. Chun (editors), Elora Reipublicae
Popularis Sinicae (68). Science Press, Beijing.
'tang. H. P. 1990. New taxa of Pedicularis from the lleng-
duan Mountain. Acta Phytotax. Sin. 28: 13—144.
-, N. II. Holmgren & R. R. Mill. 1998. Pedicularis
L. Pp. 97-209 in /. Y. Wu & P. H. Raven (editors).
Flora of China, Vol. 18. Science Press. Beijing, & Mis¬
souri Botanical Garden Press, St. Louis.
Taxonomic Novelties in Andropogon (Poaceae—Andropogoneae)
for Brazil
Ana Zanin
Departamento de Botanica, Universidade Federal de Santa Catarina, Irindade, Florianopolis,
Santa Catarina, 88040—900, Brazil, anazanin@terra.com.br
Hilda Maria Longhi-Wagner
Departamento de Botanica, Universidade Federal do Rio Grande do Sul, Avenida Paulo
Gama, s.n., Porto Alegre, Rio Grande do Sul, 90046-900, Brazil, hmlw@vant.com.br
Abstract. Arulropogon brasiliensis and A. mono-
cladus, as well as a new variety, A. ingratus var.
hirsutus from Brazil, are described. One new com¬
bination, A. bogotensis (Hackel) A. Zanin & Lon¬
ghi-Wagner, is proposed. Morphological descrip¬
tions, illustrations, data on habitat, and
geographical distribution are provided.
Resumo. Andropogon brasiliensis e A. monocla-
dus, bem como uma nova variedade, A. ingratus
var. hirsutus para o Brasil, sao descritas. Uma nova
combinagao, A. bogotensis (Hackel) A. Zanin &
Longhi-Wagner e proposta. Sao fornecidas descri-
goes rnorfologicas, ilustragoes. dados sobre habitat
e distribuigao.
Key words: Andropogoneae, Brazil, Gramineae,
Panicoideae, Poaceae.
The genus Andropogon L. includes ca. 100 spe¬
cies (Clayton & Renvoize, 1986) distributed
throughout the tropics, with centers of species di¬
versity in Africa and tropical America (Clayton &
Renvoize, 1982). Thirty species occur in Brazil, in¬
cluding A. gayanus Kunth, introduced from Africa
for cultivation. In Brazil, the greatest species di¬
versity is found in “eerrado" and “campos rupes-
tres” of the southeastern and central-western re¬
gions.
Andropogon brasiliensis A. Zanin & Longhi-
Wagner, sp. nov. TV PE: Brazil. Minas Gerais:
mun. Congonhas do Norte, Serra da Carapina,
18°52'S, 43°14'W, 2 Mar. 1998, R. C. Forzza,
./. R. Pirani, A. C. Marcato, M. C. Assis & A.
Rapini 694 (holotype, SPF; isotypes, FLOR,
k). Figure 1.
Flabitu A. inacrothrix affinis sed praecipue laminis fo-
liorum brevioribus cl angustioribus, spiculis pedicellatis
longioribus et ligula membranaceo-ciliata differt. Laminae
3.5—17 X 0.05—0.2(0.4) cm, ligula membranaceo-c iliata.
inflorescentiae graeiles, plerumque terminales tantum,
2—3(—5) ramis per spatheolam, spiculae pedieellatae (2—)
3.1—5 mm longae.
Perennial , caespitose; culms 34—64 cm tall, gla¬
brous; 3 to 5 glabrous nodes. Leaf sheaths usually
shorter than the internodes, glabrous; blades 3.5—
17 X 0.05—0.2(—0.4) cm, linear, usually condupli-
eate, margins convolute or involute, apex acute or
subobtuse, base straight, green on both faces, gla¬
brous on abaxial face, puberulous on adaxial face,
usually setose in the proximal portion, with mar¬
ginal or submarginal hairs 2-5 mm long, apex sca¬
brous on the margins; ligule 0.2—0.6 mm long,
membranous ciliate. Inflorescence 2.5-6 cm long,
lax, elongated, usually composed of terminal, or
both terminal and axillary inflorescence units, these
units with 2 to 3(to 5) non-branched flowering
branches, equal or subequal in length, conjugate or
subdigitate, exserted or partially enclosed by the
spatheole; peduncle of the inflorescence unit 8—19
cm long; spatheole 5.5—14 cm long. Pedicels and
rachis internodes 2—3(—4) mm long, linear, subequal
in length, with hairs shorter than the sessile spike-
let or as much as 1.5 times its length. Sessile spike-
let 4.5-6.5 mm long, monoclinous, stramineous-
green; lower glume 4.5—6.5 X 0.6—1 mm, slightly
concave, lanceolate, bidentate, chartaceous,
4-nerved, without intercarinal nerves, without
groove, scabrous on the upper third of the nerves,
margins glabrous; upper glume 4.1-5 X 0.8-1.4
mm, acute or apiculate, chartaceous, 3-nerved, sca¬
brous on the upper third of the central nerve, ciliate
on the upper half of the margins; lower lemma 3.9—
4.5 X 0.7-1 mm, hyaline or hyaline vinaceous,
2-nerved, ciliate on the upper half of the margins;
palea absent; upper lemma 3—1 X 0.4—0.9 mm, per¬
fect hyaline or stramineous hyaline, 3-nerved, gla¬
brous or sparsely ciliate on the margins; apex bifid
on the upper third or quarter, awned, awn 12—22
mm long; palea 1.2—2.8 X 0.3—0.7 mm, hyaline,
Nov ON 13: 368-375. 2003.
Volume 13, Number 3
2003
Zanin & Longhi-Wagner
Andropogon from Brazil
369
Figure 1. Andropogon brasiliensis A. Zanin & Longhi-Wagner. —A. Habit. —B. Ligule. —C. Terminal dispersal unit
of the flowering branches. —1). Dispersal unit of the mid portion of the flowering branches. — E. Raehis internode. —
F Upper lemma of the sessile spikelet. — G. Caryopsis, dorsal view. — H. Caryopsis, ventral view. Based on B. C.
Forzza et at. 694 (A), A. B. July et at. IBS I (B-F). and A. M. Giulietti CFSC 7329 (G, H).
370
Novon
nerveless, acute or irregularly denticulate, glabrous
or eiliate at the apex. Lodicules 2, ca. 0.7 mm long,
glabrous; stamens 3, anthers 0.7-1.2 mm long, yel¬
low. Caryopsis 1.5-1.8 X 2-3 mm. Pedicellate
spikelet neuter, (2-)3.1-5 X 0.1-0.2 mm, shorter
than or as long as the sessile spikelet, always nar¬
rower.
Andropogon brasiliensis occurs in “campos ru-
pestres” of the Espinhayo Range as isolated indi¬
viduals in humid, sandy, and rocky soils. I he new
species resembles A. macrothrix Trinius, but the
plants are more delicate, with shorter leaf blades,
up to 17 cm long and 2 mm wide. In A. macrothrix ,
the leaf blades are usually more than 2 mm wide
and extend up to 38 cm long. Furthermore. .4. ma¬
crothrix has terminal and axillary inflorescences
that are more robust, with 2 to 1 I flowering branch¬
es. and pedicellate spikelets 1.2 X 3.2 mm long,
usually shorter than those of A. brasiliensis. The
ligules of A. brasiliensis are membranaceous eiliate.
while in 4. macrothrix the ligules are membranous
eiliolate or membranous with an erose apex.
Paratypes. BRAZIL. Minas Gerais: Diamantina, re-
giao da Bandeira, 12 km de Diamantina, ao sul da rodovia
Diamantina—Belo Horizonte, 9—13 July 1977, A. G. Hur-
man 25 (SP): Jaboticatubas, km 141 ao longo da rodovia
Lagoa Santa-Coneeiyao do Mato Dentro— Diamantina, I .
Apr. 1972. A. H. Joly et al. 1851 (ICN, SI’. SPF); Lima
Duarte, Sena do Ibitipoca, Pico do Piao, 20 July 1998, .4.
Zimin et al. 740 (FLOR), 23 Mar. 1999, L. G. Rodela IC-
3 (FLOR); Santana do Riacho, km 137 ao longo da Ro¬
dovia Belo Horizonte—Concei^ao do Mato Dentro, s.d., A.
M. Giulietti et al. CFSC 7329 (SPF); Sao Tome das Letras,
II June 1987, .S'. C. Pereira & M. II. Lagoa s.n. (ESAL
7677, IBGF 20138); morro da entrada da eidade, direyao
Baependi—Sao Tome das Letras, 6 Dec. 1997, A. Zanin A'
//. M. Longhi-Wagner 673 (FLOR); Tiradentes, 26 June
1986, S. C. Pereira s.n. (ESAL 6537).
Andropogon inonocladus A. Zanin & Longhi-
Wagner. sp. nov. TAPE: Brazil. Brasilia, Re-
serva Ecoldgica do IBGE, 4 Aug. 1979. E. P.
Heringer et id. 1716 (holotype, IBGE; isoty pes.
CEN. ICN. SP. UEC). Figure 2.
A. laterali subsp. laterali alfinis, sed ramo florifero un¬
ion per spatheolam praedito et plantis robustis 170—250
cm longis distal.
Perennial, eaespitose; culms 170—250 cm tall,
glabrous; 9 to 10 glabrous nodes. Leaf sheaths lon¬
ger and/or shorter than the internodes, villous or
hirsute; blades (22-)50-70 X 0.5-0.7 cm, linear,
flat, apex obtuse or abruptly acute, base straight,
glaucous and villous or hirsute on both faces, less
frequently glabrous, margins scabrous; ligule 1—1.2
mm long, membranous eiliolate. Inflorescence 2.5—
5 cm long, lax. elongated, composed of terminal
and axillary inflorescence units, these units with
one non-branched flowering branch, usually par¬
tially enclosed by the spatheole; peduncle of the
inflorescence unit 1.5-6.5 cm long; spatheole 3.5—
7 cm long. Pedicels and rachis internodes 2.2-3.5
mm long, linear, subequal in length, with hairs
shorter than sessile spikelet. Sessile spikelet 4.5—5
mm long, monoclinous, castaneous, stramineous or
stramineous-vinaeeous; lower glume 4.5—5 X 1-1.1
mm long, slightly concave, lanceolate, chartaceous,
2-nerved, without intercarinal nerves, without
groove, scabrous on the upper third or half of the
nerves, margins glabrous, apex bidentate or erose;
upper glume 3.5—4 X 0.8—1 mm, apiculate, char¬
taceous, 3-nerved. scabrous on the upper third or
half of the central nerve, eiliate on the upper half
of the margins; lower lemma 3—4 X 0.7—1 mm, hy¬
aline, 2- to 3-nerved, eiliate on the upper half of
the margins; palea absent; upper lemma 3—3.5 X
0.3—0.6 mm, perfect stramineous or stramineous
hyaline, 1- to 3-nerved, glabrous on the margins;
apex bifid on the upper third or fifth, awned, awn
8—9 mm long; palea 0.8—1.2 X 0.6 mm. hyaline,
nerveless, irregularly dentate, glabrous. Lodicules
2, ca. 0.8 mm long, glabrous; stamens 3, anthers
0.6—1 mm long, yellow. Caryopsis 2—3 X 0.8 mm.
Pedicellate spikelet usually staminate, rarely neuter
in the same plant, neuter spikelet 3—3.5 X 0.2 mm;
staminate spikelets 4.5-5.5 mm long, somewhat
longer than the sessile spikelet, stramineous or
stramineous-vinaeeous, awnless; lower glume 4.5—
5.5 X 1-1.2 mm. apiculate or erose, chartaceous,
5-nerved, scabrous on the upper hall or third, es¬
pecially on the nerves, glabrous on the margins;
upper glume 4 X 1.5 mm, chartaceous, 3-nerved,
erose, sparsely scabrous, especially on the nerves,
glabrous or eiliate on the margins; lower lemma
3.5—4.2 X 1.2 mm, hyaline or hyaline-vinaceous,
I-nerved, eiliate on the upper half of the margins;
palea absent; upper lemma 2.8—3.5 X 0.6—1 mm,
hyaline, 1-nerved, eiliate on the upper half of the
margins; palea 0.6 X 0.5 mm, hyaline, nerveless,
irregularly dentate, glabrous. Lodicules 2, ca. 0.7
mm long, glabrous; stamens 3, anthers 2-2.5 mm
long, yellow.
Andropogon inonocladus occurs in marshes of the
“cerrado,” in central Brazil and less frequently in
the state of Bahia, also in marshes. The new species
resembles A. lateralis Nees subsp. lateralis due to
its well-developed and staminate pedicellate spike¬
lets with the lower glume of the sessile spikelet
slightly concave, without nerves or grooves between
the keels. Andropogon lateralis subsp. lateralis dif¬
fers by always having two or more flowering branch-
Volume 13, Number 3
2003
Zanin & Longhi-Wagner
Andropogon from Brazil
371
figure 2. Andropogon monocladus A. Zanin & Ix)nghi-Wagner. —A. Flowering culm, basal portion. —B. Flowering
culm, apical portion. —C. Terminal dispersal unit of the (lowering branches. —I). Dispersal unit of the mid portion of
the flowering branches. —E. Kaehis intemode. —F. Upper lemma of the sessile spikelet. —G. Caryopsis, dorsal view.
—H. Caryopsis, ventral view. Based on l\ .S'. Cdmara & T. S. Filgueiras 118.
372
No von
es in each inflorescence unit and by the plants be¬
ing smaller. Furthermore, its distribution is
primarily in southern Brazil, while A monocladus
occurs in central Brazil.
Paratypes. BRAZIL. Bahia: Correntina, Fazenda Ja-
tohiS, 8 Aug. 1992, M. A. Silva et al. 1606 (IBGE). Dis¬
trito Federal: Brasilia, Cristo Bedentor, 15 Jan. 1991,
P. S. Camara T. S. Filgueiras I !H (IBGE); Sobradinho,
27 Sep. 1965, //. .S'. Irwin et al. 0734 (US, foto). Goias:
Jataf, 30 July 1956, A. Macedo 4617 (I AC); Municipality
not specified, s.d.. Fazenda da Vargem, Glaziou 22366 (K,
W).
Amlropogou ingrains Hackel var. hirsutus A.
Zanin ■ 12. and
Macedo 4908, the awns were observed to be 2—10
mm long combined with awnless spikelets in the
same inflorescence. The muticous sessile spikelet
and the smaller pedicellate spikelets give the inflo¬
rescence of A. bogotensis a much more delicate as¬
pect than in ,4. lateralis subsp. lateralis.
Specimens examined. BRAZIL. Minas Gerais: Dia-
mantina, Sena de Santo Antonio, 27-30 Dec. 1929, A.
Chase 10443 (VIC). 19 Jan. 1984, T. S. Filgueiras cV A.
G. Barman 1086 (IBGE); 12 km de Diamantina, ao sul da
rodovia Diamantina—Belo Horizonte, 9—13 Job 1977. A.
G. Barman 41 (SP); estrada de Sousa ate Sao Joan da
Chapada, 1 Apr. 1980, A. G. Barman 549 (SP); estrada
Datas-Diamantina, BR 259, km 590, It) Dec. 1997, A.
Zanin <£■ A. C. Araujo 716 (ELOR, SPF); safda da cidade.
em diregao a Biribiri. 11 Dec. 1997, A. Zanin tV A. C.
Araujo 719 (KLOR. SPF); Lavras Novas, Serra de Itatiaia
e Serra de Lavras Novas, 6 Sep. 1978, A. G. Barman 392
(SP); Sao Joao Del Rei, proximo a Sao Joao Del Rei, Oct.
1896, A. Silveira 1842 (R); Uberlandia, Fstacao de Su-
cupira. 24 Jan. 1957, A. Macedo 4908 (IBGE, SP).
Acknowledgments. The authors are grateful to
Tarciso Filgueiras for his assistance in the elabo¬
ration of the Latin diagnoses and to Rogerio l.upo
for the illustrations. The first author thanks CAPES-
P1CDT, and the second author thanks CNPq (Re¬
search Brazilian Council) for grants to support their
work.
Literature Cited
Clayton, W. I). & S. A. Renvoize. 1982. Gramineae. Pp.
767—782 in R. M. Polhill (editor). Flora of Tropical East
Africa, Part 3. Balkema, Rotterdam.
-& -. 1986. Genera Graminum: Grasses of
the World. Her Majesty’s Stationery Office, London.
(Kew Bull. Addit. Ser., 13).
Hackel, E. 1889. Andropogoneae. In: A. De Candolle &
C. De Candolle (editors), Monogr. Phan. 6: 1—716.
Renvoize, S. 1998. Gramineas de Bolivia. Royal Botanic
Gardens, Kew.
Volume 13, Number 3, pp. 281-376 of NOVON was published on 10 September 2003.
Volume 13
Number 4
2003
NOVON
A New Species and a Nomenclatural Change in
Graptopetalum (Crassulaceae)
Haul Acevedo-Rosas
Posgrado en Sistematica, Instituto de Ecologfa, A.C. Apdo. Postal 63, Xalapa, Veracruz,
91000, Mexico, and Depto. de Geografia, CUCSH, Universidad de Guadalajara, Av. de los
Maestros y M. Barcena, Guadalajara, Jalisco, 44120, Mexico, acerosl2@yahoo.com
Miguel J. Chazaro
Departamento de Geografia, CUCSH, Universidad de Guadalajara, Av. I)e los Maestros y M.
Barcena, Guadalajara, Jalisco, 44120, Mexico, chazaro55@hotmail.com
ABSTRACT. Graptopetalum glassii from Colima,
Mexico, is described and illustrated. It is closely
related to G. pentandrum, differing in being smaller
and in having a herbaceous habit and a larger num¬
ber of oblanceolate pale green leaves. Graptopetal¬
um pentandrum subsp. superbum is raised to spe¬
cies rank. Characters that support the change of
status are a larger size of the plants, a squamose
surface of the stem, a pink-violet color of the ro¬
sette, oblong-obovate leaves, and a profusely
branched inflorescence.
RESUMEN. Graptopetalum glassii de Colima, Mex¬
ico, es descrita e ilustrada. Esta cercanamente re-
lacionada con G. pentandrum ; sin embargo, difiere
en poseer un menor tamano, un habito herbaceo
con un mayor numero de hojas oblanceoladas de
color verde mas palido. Graptopetalum pentandrum
subsp. superbum es elevada a nivel de especie. Ca-
racteres que apoyan el cambio de estatus son un
tamano mas grande, una superficie del tallo esca-
mosa, un color rosa-violeta de la roseta, hojas
oblongo-obovadas y un mayor numero de ramas en
la infloreseencia.
Key words: Colima, Crassulaceae, Graptopeta¬
lum, Mexico.
Graptopetalum Rose (Crassulaceae) includes ap¬
proximately 19 species from Arizona, U.S.A., to Oa¬
xaca, Mexico, found in inaccessible places in semi-
arid vegetation over rocky hills or walls of ravines,
from sea level to 7500 ft. Only a few species have
wide distribution, such as G. pachyphyllum Rose
and G. rusbyi (Greene) Rose, while most of the spe¬
cies are restricted to specific mountains or ravines.
For example, G. mendozae Glass & Chazaro grows
in the foothills of small mountains in the north of
Veracruz, Mexico (Glass & Chazaro, 1997), and G.
marginatum Kimnach & Moran is known only from
the north of Tepic in Nayarit, Mexico (Kimnach &
Moran, 2002).
Plants of Graptopetalum are characterized by
caulescent or acaulescent rosettes. The caulescent
species are pendent or bushy, and the leaves are
thick. The inflorescence is lateral and the flowers
usually have a fetid odor; petals are pale, erect, and
fused in the lower hall and rotate in the apex, where
they have red to brown dots often forming bands
across the petals. The stamens are initially erect;
after anthesis they become strongly recurved (Uhl,
1970).
In 1995 Charles Glass and Mario Mendoza found
a population of a Graptopetalum species restricted
to a gypsum wall on the Pacific slopes of Mexico
in the state of Colima. Due to its distinctive char¬
acters, we describe it here as G. glassii. This new
Novon 13: 377-380. 2003.
378
Novon
species and the new combination that follow are the
result of a phylogenetic study of the genus Grap-
topetalum, which is currently in preparation by the
first author. The names are here published sepa¬
rately to make them immediately available for use.
Graptopctaluiii glassii Acevedo-Rosas & Chaza-
ro, sp. nov. TYPE: Mexico. Colima: Municipio
de Ixtlahuacan, Carretera libre de Ixtlahuacan
a Colima, justo enfrente del senalamiento del
km 21 de la autopista Colima—Tecoman, sobre
una pared de suelo yesoso con exposicion NE,
410 m, 12 ene. 2000, R. Acevedo, M. Chdzaro
& J. A. Machuca 1724 (holotype, XAE; iso¬
types, MO, NY). Eigure 1.
Graptopetalo pentandro cognatum. Plantae perennes
caespitosae. Caules maximam partem decumbentes breves
vel plantae acaules. Rosula ejus G. pentandri minor sed
folds pluribus munita congestaque. Folia oblanceolata vel
late oblanceolata, albo-viridia. Inflorescentia sub cymae
compositae formam; flores pentandri; petala triangulari-
lanceolata-acuta; calyx cupuliformis; gynoecium obova-
tum.
Plants perennial, caespitose, forming compact
clusters; stem mostly decumbent, smooth, short or
the plants stemless; suberect at first, then pendent,
to 55 mm long, 3—5 mm thick; at first nearly white-
glaucous, later red-brown-glaucous; mostly caespi-
tosely branching by ascending (when young) and
pendent (with age) slender stems from below ro¬
sette. Rosettes 20—30(40) mm diam. when mature,
with 30 to 40(50) crowded leaves; leaves white-
green to pale blue-green to yellow-green with palest
rosy blush, especially in youngest leaves and with
exposure to more intense light, glabrous, slightly
glaucous, ascending-erect when young, later the
apical ones ascending-erect and the basal ones ex¬
panding, 13-16(20) X 8-12 mm, ea. 2-5 mm
thick, inner surface slightly concave, outer surface
markedly convex, vaguely keeled in upper 2—4 mm,
oblanceolate to broadly oblanceolate; apex some¬
what acute or slightly apiculate, light green. Inflo¬
rescence a compound cyme, generally 6—12 cm
high; unbranched or with 1 or 2(3) simple or bi¬
furcate branches; peduncle 2.5—8 cm tall, 2—3.5
mm thick, smooth, light green, pink or red-tinged,
with 16 to 18 bract leaves, soon deciduous, similar
to rosette leaves but ovate rather than oblanceolate,
diminishing markedly in size, 7-10 X 6—7 mm, 2.5
mm thick, bracteoles mostly elliptic to lanceolate,
1.5—3 X 1—1.5 mm; flower buds about 9 mm long,
typically 5-merous; calyx cup-shaped, 4 mm high,
sepals 5, gray-green, tips 3 mm long; corolla ca. 8—
9 mm long when unopened, 12—14 mm wide when
expanded, yellow-white marked with red, flecks
ami partial banding toward the tips, the tips nearly
solid red, petals 5 (rarely 6), 3.5 mm wide near
base, triangular-lanceolate, acute, united for ca. 2.7
mm, rotate in the distal half; stamens 5, antesepa-
lous, at first erect, after anthesis strongly reflexed
between the petals, 6—7 mm long, ca. 0.4 mm wide,
base adnate to corolla tube, filaments white, the
apical half pink or red dotted, anthers with grav-
yellow pollen; nectar glands 5, yellow, 0.3 mm tall,
0.5 mm wide, 0.3 mm thick, somewhat reniform,
the inner face strongly concave, the external con¬
vex; gynoecium obovate, green-yellow, reddened
apieally, ca. 6—6.5 mm long, 3—3.5 mm thick, the
styles 1 mm long or less.
Distribution and ecology. Known only from the
type locality at an elevation of 410 m, where the
plant occurs in tropical deciduous forest. Grapto-
petalum glassii grows on gypsum slopes of hills.
Phenology. Graptopetalum glassii flowers from
late February to April or early May. Flowers pro¬
duce a disagreeable fetid odor.
Etymology. The species is named alter Charles
Glass (1934—1998), who collected plants of the new
taxon for the first time. Mr. Glass worked as the
Curator of Plants for Cante, A.C. (a non-govern¬
mental organization), a small botanical garden lo¬
cated in San Miguel de Allende, Guanajuato, Mex¬
ico, where he studied succulent plants such as
Cactaceae and Crassulaceae. Mr. Glass carried out
many field trips to remote and interesting places of
Mexico, where he found and described new taxa of
these groups.
Graptopetalum glassii differs from G. pentandrum
in its shorter herbaceous habit, caespitose growth,
a smaller size of the stem, rosette, and leaves, and
a shorter inflorescence (Table l). Graptopetalum
glassii can be recognized in the field among other
known species of the genus by its beautiful little
light green and dense rosettes, its shorter and slen¬
der stems, and the numerous conglomerate leaves
(more than 40); this discovery also represents the
third record in the genus having haplostemonous
flowers.
Plants of Graptopetalum glassii are sold as G.
sotoi; M. Chdzaro and C. Glass were planning to
publish it. Eigure 1 depicts a cultivated plant,
which is more branched and robust than wild
plants. Table 1 summarizes the main differences
among Graptopetalum glassii and the closely relat¬
ed species G. pentandrum and G. superbum, and is
based on our own observations.
Paratype. MEXICO. Colima: near La Salada, approx.
25 km S of Colima on a steep and shaded gypsum slope,
29 Dec. 1995, C. Glass <£• M. Mendoza 8910 (Cante, .4.6'.).
Volume 13, Number 4
2003
Acevedo-Rosas & Chazaro
Graptopetalum (Crassulaceae)
379
Figure 1. Graptopetalum glassii Acevedo-Rosas & Chdzaro. —a. Habit. —b. Propagation of plant by leaf cutting. —
c. Front view of flower. —d. Lateral view of flower. —e. Petal and stamens. —f. Stamen. —g. Gynoeeium with nectary
scales. Based on living collection established at “Clavijero” botanical garden in Xalapa, Veracruz, Mexico (acc. no.
2003-16), holotype: H. Acevedo el al. 1724 (XAL).
380
No von
Table 1. Distinguishing characters of Graptopetalum glassii
and its closest relatives.
G. glassii
G. pentandrum
G. superbum
Habit
herbaceous
suffrutex
suffrutex
Plant
caespitose
caespitose-ramose
ramose
Size
16 cm
40 cm
80 cm
Stem surlace
smooth
smooth
squamose
Stem diameter
3—5 mm
3—6 mm
10-12 mm
Leaf form
oblaneeolate
obovate
oblong-obovate
Leaf color
blue-green to
blue-green to
gray-blue to
white-green
white-blue
pink-violet
Leal length
13—20 mm
20-40 mm
40—55 mm
Rosette diameter
40 mm
60 mm
‘K) mm
Number of leaves per rosette
30-40
15-30
12-20
Inflorescence length
6-12 cm
20—35 cm
30—40 cm
Number of branches per inflorescence
1-2
3-4
12-15
Gruploprtuliim supeibiim (Kimnach) Aeevedo-
Kosas, stat. nov. Basionym: Graptopetalum
pentandrum Moran subsp. superbum Kimnach,
Gael. Succ. J. (Los Angeles) 59: 142. 1987.
TYPE: Cultivated. Mexico. La Barca, near
Guadalajara, the native locality unknown, Avi-
na s.n., HBG acc. no. 49307 (holotype, HNT
sheet #6392).
Graptopetalum superbum was described as a sub¬
species of G. pentandrum by Kimnach (1987).
Graptopetalum superbum is distinguished from G.
pentandrum by its larger size, a squamose surface
of the stem, a pink-violet color of the rosette, leaves
oblong-obovate, a profusely branched inflorescence
(see Table I), and different chromosome number ( G.
superbum n = 64, G. pentandrum n = 32, accord¬
ing to Uhl (in Kimnach. 1987)).
Morphological cladistic analyses indicate that
Graptopetalum glassii, G. pentandrum, and G. su¬
perbum are closely related and belong to a small
group (Acevedo et al., in press) that occurs in the
area of Nueva Galicia in western Mexico. Among
the characters they share are live stamens in a sin¬
gle whorl (haplostemonous flowers) instead of ten
in two whorls, which the rest of the species in the
genus have. Among Crassulaceae, only some gen¬
era of Sedoideae and the representatives in the
Crassuloideae include haplostemonous species, and
all are from the Old World (Berger, 1930).
Acknowledgments. The authors gratefully ac¬
knowledge Victoria Sosa for her review and sug¬
gestions for the manuscript, Jerzy Bzedowski for his
writing of the Latin diagnosis. Edmundo Saavedra
for the illustration, and J. Antonio Machuca for his
help in the field. Fieldwork was possible through a
grant by CONACyT (225260-5-29378N). The first
author recognizes the support from the SEP-SESIC-
PROMEP Fellowship Program and the University
of Guadalajara, Mexico.
Literature Cited
Acevedo, R., V. Sosa & F. G. I.orea. In press. Phylogenetic
relationships and morphological patterns in Graptope¬
talum (Crassulaceae). Brittonia 56.
Berger, A. 1930. Crassulaceae. In A. Engler & k. Prantl,
Die naturlichen Pflanzenfamilien, 2nd ed. Wilhelm En-
gelmann. Leipzig (18a): 352-482.
Glass, C. & M. Chdzaro B. 1997. Ena nueva especie de
Graptopetalum (Crassulaceae) del norte de Veracruz.
Cact. Sue. Mex. 42: 79-82.
Kimnach, M. 1987. A new succulent from Mexico: Grap¬
topetalum pentandrum subsp. superbum. Cact. Succ. J.
(Iajs Angeles) 59: 140—143.
- & R. Moran. 2002. Graptopetalum marginatum, a
new species from Nayarit, Mexico. Cact. Succ. J. (Los
Angeles) 74: 196—198.
Uhl, C. II. 1970. Chromosomes of Graptopetalum and
Thompsonella (Crassulaceae). Amer. J. Bot. 57: 111 5—
1 121 .
Transfer of Most North American Species of Arabis to
Boechera (Brassicaceae)
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. Based on extensive molecular, cyto-
logical, and morphological studies, most of the
North American species of Arabis (Brassicaceae)
are recognized herein as members of the genus
Boechera. Morphological differences between the
two genera are discussed, and species formerly
placed in Arabis and not included here in Boechera
are assigned to other genera. Thirty-two new com¬
binations in Boechera are proposed: B. bodiensis
(Rollins) Al-Shehbaz, B. breweri (S. Watson) Al-
Shehbaz, B. canadensis (L.) Al-Shehbaz, B. con-
stancei (Rollins) Al-Shehbaz, B. cusickii (S. Watson)
Al-Shehbaz, B. dispar (M. E. Jones) Al-Shehbaz, B.
falcifructa (Rollins) Al-Shehbaz, B. fruticosa (A.
Nelson) Al-Shehbaz, B. glaucovalvula (M. E. Jones)
Al-Shehbaz, B. hastatula (Greene) Al-Shehbaz, B.
hirshbergiae (S. Boyd) Al-Shehbaz, B. hoffmanii
(Munz) Al-Shehbaz, B. inyoensis (Rollins) Al-Sheh-
baz, B. johnstonii (Munz) Al-Shehbaz, B. koehleri
(T. J. Howell) Al-Shehbaz, B. laevigata (Muhlen¬
berg ex Willdenow) Al-Shehbaz, B. missouriensis
(Greene) Al-Shehbaz, B. ophira (Rollins) Al-Sheh¬
baz, B. parishii (S. Watson) Al-Shehbaz, B. patens
(Sullivant) Al-Shehbaz, B. perstellata (E. Braun) Al-
Shehbaz, B. pinzlae (Rollins) Al-Shehbaz, B. pla-
tysperma (A. Gray) Al-Shehbaz, B. pygmaea (Rol¬
lins) Al-Shehbaz, B. rectissima (Greene)
Al-Shehbaz, B. repanda (S. Watson) Al-Shehbaz, B.
rigidissima (Rollins) Al-Shehbaz, B. rollei (Rollins)
Al-Shehbaz, B. shortii (Fernald) Al-Shehbaz, B.
stricta (Graham) Al-Shehbaz, B. subpinnatifida (S.
Watson) Al-Shehbaz, and B. tiehmii (Rollins) Al-
Shehbaz. Three recently described species, Arabis
calderi G. A. Mulligan, A. murrayi G. A. Mulligan,
and B. glareosa Dorn, are reduced to synonymy of
B. divaricarpa (A. Nelson) A Love & D. Love, B.
lyallii, and B. gunnisoniana (Rollins) W. A. Weber,
respectively.
Key words: Arabidopsis, Arabis , Boechera, Bras¬
sicaceae. Pennellia, Turritis.
As traditionally delimited by recent North Amer¬
ican botanists (e.g., Hopkins, 1937; Rollins, 1941,
1993b; Mulligan, 1996). the genus Arabis L. (Bras¬
sicaceae) was rather broadly circumscribed to in¬
clude species now placed in Arabidopsis Heynhold,
Boechera A. Love & D. Love, Pennellia Nieuwland,
and Turritis L. (see below). Several European bot¬
anists also adopted such a broad concept. For ex¬
ample, Busch (1939) included in Arabis species
now placed in Alyssopsis Boissier; Arabidopsis ; Cru-
cihimalaya Al-Shehbaz, O'Kane & R. A. Price;
Neotorularia Hedge & J. Leonard; Rhammatophyl-
lum 0. E. Schulz; and Turritis. By contrast, the re¬
cent treatment of Arabis by Jones and Akeroyd
(1993) included species now assigned to Arabidop¬
sis, Diplotaxis DC., Fourraea Greuter & Burdet,
and Turritis (author’s compilation). All of the above
authors delimited Arabis exclusively on a combi¬
nation of latiseptate siliques (fruits flattened par¬
allel to the septum), branched trichomes, and ac-
cumbent cotyledons. However, this combination of
characters evolved independently many times with¬
in the Brassicaceae (Appel & Al-Shehbaz, 2002),
and it characterizes some or all species of the ten
genera above (minus Diplotaxis ), as well as those
of Aplanodes Marais (South Africa), Aubrieta Adan-
son (Europe), Desideria Pampanini (central Asia
and Himalayas), Draba L. (several Asian species),
Eremobium Boissier (northern Africa, Middle East),
Erysimum L. (some Eurasian species), Eurycarpus
Botschantzev (Himalayas), Lepidostemon J. I).
Hooker & Thomson (Himalayas), Matthiola R.
Brown (Africa, Eurasia), Pachycladon J. D. Hooker
(New Zealand and Tasmania), Pennellia (some
North American members), Phoenicaulis Nuttall
(U.S.A.), Sibara Greene (North America), Sisym-
briopsis Botschantzev & Tzvelev (central Asia),
Smelowskia C. A. Meyer ex Ledebour (Far East rep¬
resentatives), and Stevenia Adams & Fischer (east¬
ern and central Asia).
Rollins (1993b) recognized 80 North American
species in Arabis, of which 24 were further divided
into 40 varieties, while Mulligan (1996) accepted
30 species and 6 varieties in Alaska, Canada, ami
Greenland. As shown below, extensive molecular
studies on members of North American Arabis
clearly indicate that the genus, as circumscribed
Novon 13: 381-391. 2003.
382
Novon
by Rollins (1993b) and Mulligan (1996), is poly-
phyletic and represents a heterogeneous assem¬
blage of four genera: Arabidopsis, Boechera, Pen-
nellia, and Turritis. first. Arabis lyrata L.,
including A. media N. Busch and A. hamchatica
(Kischer) Ledebour, has been shown by Al-Shehbaz
and O’Kane (2002), O’Kane and Al-Shehbaz (1997,
2003), Koch et al. (1999. 2000, 2001), Heenan et
al. (2002), and Miyashita et al. (1998) to belong to
Arabidopsis. Second, Arabis glabra (L.) Bernhardi
was shown by Koch et al. (1999, 2000, 2001) and
Heenan et al. (2002) to be unrelated to Arabis and
to belong to Turritis, as originally treated by Lin¬
naeus (1753) and as accepted in a number of mono¬
graphic, floristic, and systematic accounts of the
family (e.g., Torrey & Gray, 1838; Hayek, 1911;
Burdet, 1967; Schulz, 1936; Schultze-Motel, 1986;
Zhou et al., 2001; Appel & Al-Shehbaz, 2002).
Third, Arabis microsperma Rollins and A. tricornuta
Rollins belong to Pennellia (Bailey et al., 2002;
Price et al.. 2001; Fuentes-Soriano, 2002), a genus
most closely related to Halimolobos Tausch, Man-
coa Weddell, Boechera, and Sphaerocardamum
Shauer, and clearly unrelated to Arabis. Finally, the
bulk of North American Arabis is recognized here
as Boechera.
File genus Boechera was segregated from Arabis
by Love and Love (1976) and was said to dilfer by
having a base chromosome number of* = 7 instead
of x = 8. These authors did not provide any mor¬
phological characters that otherwise distinguished
the two genera. Rollins (in Rollins & Riidenberg,
1977) strongly disagreed with the splitting of Arabis
based on chromosome numbers, though he admit¬
ted (p. 102) that species with x — 7 have “a some¬
what different circle of close affinity” from those
with x = 8, which are related to the Eurasian mem¬
bers of the genus. Weber (1982: 369) concluded
that “The difference in basic chromosome number
[in Arabis] . . . does represent a divergent phylo¬
genetic line, and the genetic barrier that it presents
to interbreeding of the units is sufficient justifica¬
tion for thinking in terms of discrete genera.”
Several phylogenetic studies (Galloway et al.,
1998; Koch et ah. 1999, 2000, 2001, 2003; Heen¬
an et al., 2002; O'Kane & Al-Shehbaz, 2003) have
clearly demonstrated that Boechera is unrelated to
Arabis alpina L., the type species of Arabis, or to
the Eurasian and North American species with x =
8. Therefore, molecular, cytological, and morpho¬
logical data (see below) strongly support the rec¬
ognition of Boechera as a genus well-defined from
the rest of Arabis, hereafter Arabis s. str., with x =
8. Only about ten native North American species
belong to Arabis s. sir. (Mitchell-Olds, unpub¬
lished): A. aculeolata Greene, A. arenicola (Rich¬
ardson & Hooker) Gelert, A. blepharophylla Hooker
& Arnott, A. hirsuta L., A. macdonaldiana East-
wood. A. modesta Rollins (if indeed different from
the next species), A. oregana Rollins, A. crusisetosa
Constance & Rollins, A. furcata S. Watson, and A.
nuttallii B. L. Robinson. Chromosome counts based
on x — 8 are known for all except the last three
species, which are retained in this genus on the
basis of the nuclear ribosomal DNA ITS region
(Mitchell-Olds, pers. comm.), leaf indumentum,
and fruit morphology. Whether or not the western
North American A. eschscholtziana Andrzejowski ex
Ledebour should be recognized as a distinct spe¬
cies (Mulligan, 1996), as a variety of A. hirsuta
(Rollins, 1993b), or as a subspecies or synonym of
the latter is a matter of judgment and will be dealt
with in a separate paper. With the exclusion of the
ten North American species of Arabis s. sir., as well
as those transferred to Arabidopsis (A. lyrata ), Tur¬
ritis (A. glabra), and Pennellia (A. microsperma and
A. tricornuta), the remaining species assigned to
Arabis by Rollins (1993b) and Mulligan (1996,
2002) should be segregated as Boechera.
A critical comparison of the Eurasian and North
American species of Arabis s. str. and Boechera
reveals that there are a number of morphological
characters that readily distinguish the two. No
species of Arabis s. str. has falcate, secund, widely
spreading, pendent, or strongly reflexed fruits, but
the majority of Boechera species have at least one
ol these aspects of fruit morphology. Species of
Arabis s. str. have erect, ascending, or divaricate-
ascending, straight fruits. Furthermore, no species
of Arabis s. str. has dendritic, irregularly bifurcate,
or sessile trichomes on the leaves, and the vast
majority of Boechera species have at least one of
these trichome types. In Arabis s. str., the tri¬
chomes are stalked and either forked, 3-rayed, or
cruciform, and their rays are almost always un¬
branched. The present author and Sara Fuentes-
Soriano are preparing a detailed atlas ol trichome
morphology of Arabis s.l. A few Eurasian species
assigned to Arabis have dendritic trichomes, and
they are to be transferred to other genera (Al-
Shehbaz, unpublished). In species with glabrous
leaves, characters of the fruit, seed, or leaf mor¬
phology (see below) help in assigning them to ei¬
ther of the two genera.
A few other characters have some limited use
in the separation of Arabis s. str. and Boechera.
For example, the fruits in Arabis s. str. are gen¬
erally narrow (often less than 2 mm wide) and al¬
ways glabrous. Although many species of Boechera
have narrow and glabrous fruits, several others
Volume 13, Number 4
2003
Al-Shehbaz
Boechera
383
have fruits 3—8 mm wide or fruit valves sparsely
to moderately pubescent. The seeds in Arabis s.
str. are uniseriate and either narrowly winged or
wingless, just as in a number of Boechera species.
However, several species of Boechera have broadly
winged seeds (wing 1-3 mm wide) and/or biseriate
seeds.
Weber (1982) indicated that Arabis is separated
from Boechera by having loose, slender root systems
(instead of shortly clustered caudices) and large,
numerous, and often toothed (instead of smaller,
fewer, and entire) leaves. These differences were
not recognized by Al-Shehbaz (1988), Rollins
(1993b), and Mulligan (1996). A closer examina¬
tion of the caudex and leaves in Arabis s.l. world¬
wide further shows that the caudex, which is absent
in the annual and biennial species, is not useful
taxonomically and only the leaves are occasionally
helpful in the separation of the two genera.
To my knowledge, Arabis s. str. does not include
any species with both polyploidy and apomixis.
Apomixis in Arabis s.l. was first reported by Bother
(1947) in A. holboellii and subsequently confirmed
in other critical studies (Bot her. 1951, 1954, 1966.
1969). Mulligan (1996) suggested that sexual and
apomictic populations are found within A. Colum¬
biana Macoun, A. divaricarpa A. Nelson, A. drurn-
mondii A. (fray, A. exilis A. Nelson, A. lemmonii S.
Watson, A. lyallii S. Watson, A. microphylla Nuttall,
A. pinetorum Tidestrom, and A. sparsiflora Nuttall.
These species, recognized here as members of
Boechera, include triploid plants based on x = 7,
and they exhibit some meiotic irregularities, in¬
cluding univalents, multivalents, and B-chromo-
somes (Boeher, 1947, 1969; Johnson, 1970; Mul¬
ligan. 1964, 1996; Packer, 1964; Rollins, 1993b;
Rollins & Riidenberg, 1971; Roy, 1995; Vorobik,
1985).
Jones anil Akeroyd (1993) recognized three spe¬
cies in Arabis s.l. with strongly falcate and/or re-
llexed fruits: A. laxa Smith (eastern Europe and
southwestern Asia), A. turrita E. (central and east¬
ern Europe), and A. pendula L. (Eurasia). Arabis
laxa has long been recognized as Turritis laxa
(Smith) Hayek in several other floristic works (e.g.,
Cullen, 1965; Hedge, 1968; Meikle, 1977), and on
the basis of overall morphology (e.g., glaucous cau-
line leaves, creamy white or yellowish flowers,
slightly 4-angled fruits) and a chromosome number
of n — 6 (Maassoumi, 1980), it is perfectly at home
in Turritis. Arabis turrita, with 2 n = 16 (e.g.. Bur-
det, 1967; and numerous subsequent counts), is the
only species in the genus with bracteate racemes
and fruits to 9 cm long. Molecular studies (Koch et
ah, 2000. 2001) strongly support the separation of
this species to a distinct genus. In addition to its
chromosome number of 2 n — 21 (Berkutenko &
Gurzenkov. 1976; Berkutenko et ah, 1984), A. pen¬
dula is anomalous in Arabis s. str. because it has
sessile stellate trichomes with an erect central ray
much coarser than the other rays. One might be
tempted to place it in Boechera, but molecular data
(O'Kane & Al-Shehbaz, 2003) place it in a clade
well separated from both Arabis s. str. and Boech¬
era. It may well represent an independent genus,
though it is beyond the scope of this paper to deal
with it. Another anomalous European Arabis rec¬
ognized by Jones and Akeroyd (1993) is A. brassica
(Leers) Rauschert. a species with 2 n = 14 (Polat-
schek, 1966; Burdet, 1967; Titz, 1967; van Loon
et ah, 1971) often reported as A. paueiflora (Grimm)
Gracke. However, molecular data (Koch et ah,
2001) indicate that this species is unrelated to Ar¬
abis s. str., and a closer examination of morphology
supports Greuter and Burdet (1984) in transferring
it to the monotypic Fourraea Greuter & Burdet, as
F. alpina (L.) Greuter & Burdet.
The present account of Boechera was prepared
during work on the genus for an upcoming volume
of the Flora of North America, anil the new com¬
binations are proposed herein to make the species
names available for phylogenetic studies in pro¬
gress. Of the 40 additional varieties recognized by
Rollins in Arabis s.l., 35 fall within the present cir¬
cumscription of Boechera. Mulligan (1996) raised 4
of these 35 varieties to the species rank and re¬
duced 9 others to synonymy of other taxa. The sta¬
tus of these infraspecific taxa will be addressed in
a subsequent publication following the examination
of all types. A key to all species of Boechera will
lie presented when we have a better understanding
of the taxa involved, especially the representatives
from the Russian Far East.
Under each species of Boechera, its synonym in
Arabis is given regardless whether or not its bas-
ionym is in Arabis. This is important for comparing
the names in Boechera with those in Rollins
(1993b) and Mulligan (1996). Love and Love
(1976, 1982), Weber (1982. 1989. 1996), and Dorn
(2003) have already transferred several species of
North American Arabis to Boechera, and these are
listed below with full information on the type col¬
lections because this information was not given by
these authors. Types marked “not seen" were com¬
piled from Hopkins (1937), Rollins (1941), and
Mulligan (1996).
Boechera beckwithii (S. Watson) Dorn, Brittonia
55: 3. 2003. Basionym: Arabis beckwithii S.
Watson, Proc. Amer. Acad. Arts 22: 467.
384
Novon
1887. TYPE: U.S.A. Nevada: Quartz Mts., W
of (Ireal Sail Lake, Lieutenant Beckwith s.n.
(leetotype, designated by Rollins (1941: 452),
GH).
Watson (1887) cited four syntypes under the
original description of Arabis beckwithii. Rollins
(1941) listed Beckwith’s collection as the type, and
this was taken as the leetotype.
Boechera bodiensis (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis bodiensis Rollins, Contr.
Gray Herb. 212: 113. 1982. TYPE: U.S.A.
California: Mono Co., 2 mi. NW ol Masonic,
5.9 mi. from Bridgeport-Sweetwater highway,
loose soil of old mine, 8100 ft., 3 Aug. 1945,
/. L. Wiggins & R. C. Rollins 536 (holotype,
(ill; isotypes, DS, (ill, UC).
Boechera breweri (S. Watson) Al-Shehbaz, comb,
nov. Basionym: Arabis breweri S. Watson. Proe.
Amer. Acad. Arts II: 123. 1875. TYPE:
U.S.A. California: [Contra Costa Co.], rocky
summit of eastern peak of Mt. Diablo, 4 May
1862, W. //. Brewer I0B6 (leetotype, designat¬
ed by Rollins (1941: 408), GH; isotypes, l)S,
MO, UC [2], US).
Watson (1875) cited lour syntypes under the
original description ol Arabis breweri. Rollins
(1941) listed Brewer’s collection as the type, and
this was taken as the leetotype of the species.
Boechera canadensis (L.) Al-Shehbaz, comb,
nov. Basionym: Arabis canadensis L., Sp. PI.
2: 665. 1753. TYPE: leetotype, designated by
Reveal in Cafferty A Jarvis (2002: 531), ,/.
Clayton 400, left-hand specimen. Herb. Linn.
No. 842.12 (LINN; isotype, BM).
Boechera cobrensis (M. E. Jones) Dorn, Vase. PI.
Wyom. ed. 3: 375. 2001. Basionym: Arabis
cobrensis M. E. Jones, Contr. W. Bot. 12: 1.
1908. TYPE: U.S.A. Nevada: Elko Co., Cobre,
clay hills, 6000 ft., 16 June 1906, M. E. Jones
s.n. (holotype, POM; isotypes, BM, CAS. MO.
POM. US [2]).
Boechera constancei (Rollins) Al-Shehbaz,
comb. nov. Basionym: Arabis constancei Rol¬
lins, Contr. Gray Herb. 201: 5. 1971. TYPE:
U.S.A. California: Plumas Co., 7.6 mi. SE of
Quincy (at Spring Garden overpass) on road to
Blairsden, near rocks on open serpentine, W-
SW facing gentle slope, 1400 ft., 1 1 July 1969,
/,. Constance & T. I. Chuang 3875 (holotype,
GH; isotype, UC).
Boechera crandallii (B. L. Robinson) W'. A. We¬
ber. Phytologia 51: 369. 1982. Basionym: Ar-
abis crandallii B. L. Robinson, Bot. Gaz. 28:
135. 1899. TYPE: U.S.A. Colorado: Montrose
Co.. Cimarron, 7000 ft., 18 May 1898. C. S.
Crandall 6 (holotype, GH; isotype, NY not
seen).
Boechera cusiekii (S. Watson) Al-Shehbaz, comb,
nov. Basionym: Arabis cusiekii S. Watson, Proe.
Amer. Acad. Arts 17: 363. 1882. TYPE:
U.S.A. Oregon: Union Co., rocky ridges, early
spring 1879. W. C. Cusick 727 (holotype, GH;
isotvpe, US).
Boechera demissa (Greene) W. A. Weber, Phy¬
tologia 51: 370. 1982. Basionym: Arabis de¬
missa Greene, PI. Baker. 3: 8. 1901. TYPE:
U.S.A. Colorado: Gunnison Co., Cimarron,
stony riverbed, 4 June 1901, C. F. Baker 16
(holotype, NDG 5264).
Boechera dispar (M. E. Jones) Al-Shehbaz, comb,
nov. Basionym: Arabis dispar M. E. Jones,
Contr. W'. Bot. 8: 41. 1898. TYPE: U.S.A. Cal¬
ifornia: Inyo Co., Panamint Mts., Pleasant
Canyon, 5500 It., 6 May 1897, M. E. Jones
s.n. (holotype, POM; isotypes, MO, POM. US,
UTC not seen).
Boechera divaricarpa (A. Nelson) A. Love & 1).
Love, Bot. Not. 128: 513. 1976. Basionym: Ar¬
abis divaricarpa A. Nelson, Bot. Gaz. 30: 193.
1900. TYPE: U.S.A. Wyoming: Yellowstone
National Park, Yellowstone Lake, stony and
sandy banks ol lake, A. & E. Nelson 6622 (lee¬
totype, designated by Hopkins (1937: 132),
RM; isotype, GH).
Arabis calderi G. A. Mulligan, Rhodora 67: 1 11. 1996.
Syn. nov. TYPE: Canada. British Columbia: Indian
River at Mile 34 from Alaska Hwy. on A11 in road,
ca. 59°54'N, 133°48'W, open grassy flats on bench
above river, 9 June 1960, J. A. Colder & J. M. Gillett
25180 (holotype, DAO).
Nelson (1900) cited two collections (6352 and
6622) in his original description ol Arabis divari¬
carpa, but Rollins (1941) indicated that the type is
A. & E. Nelson 6332, apparently a typographical
error. However, Hopkins (1937) was the first to des¬
ignate a leetotype for the species.
An examination of the holotype of Arabis calderi
clearly shows that the plant is quite similar to those
cited by Mulligan (1996) as A. divaricarpa var. di¬
varicarpa, including the type collection of A. pra-
tincola Greene. There is no single character that
supports the maintenance of A. calderi as a distinct
species, and the trichomes of basal leaves are in¬
distinguishable in the two. As shown by Rollins
(1983), Boechera divaricarpa (as Arabis) is quite
Volume 13, Number 4
2003
Al-Shehbaz
Boechera
385
variable in leaf indumentum and fruit orientation,
and various populations of this species show mor¬
phological tendencies toward one of its closely re¬
lated species, B. stricta (as A. drummondii ) and B.
holboellii.
Boechera falcatoria (Hollins) Dorn. Brittonia 55:
3. 2003. Basionym: Arabis falcatoria Rollins,
Contr. (nay Herb. 212: 106. 1982. TYPE:
U.S.A. Utah: Box Elder Co., 3.8 mi. N of
Grouse Creek on road to Oakley, Idaho, marble
chip rock, 9 June 1981, K. C. & K. W. Rollins
81259 (holotype, GH; isotypes, GH. LE, RM,
RSA, UC).
Boechera falcif'ructa (Rollins) Al-Shehbaz,
comb. nov. Basionym: Arabis falcifructa Rol¬
lins, Contr. Gray Herb. 212: 112. 1982. TYPE:
U.S.A. Nevada: Elko Co., near U.S. Hwy. 93,
betw. Thousand Springs & Jackpot, 37 mi. S
of Jackpot, rock crevices, sagebrush area on
slope of a high ridge, 18 June 1979, R. C. &
K. W. Rollins 79267 (holotype, GH; isotype,
MO).
Boechera fecunda (Rollins) Dorn, Brittonia 55:
3. 2003. Basionym: Arabis fecunda Rollins,
Contr. Gray Herb. 214: 1. 1984. TYPE: U.S.A.
Montana: Ravalli Co., big game range E of
Corvallis, rocky terrain near sagebrush, 4000
m, 13 June 1976, Jaculyn Cory 1611 (holo¬
type. MONTU not seen; isotype, (ill).
Boechera fendleri (S. Watson) W. A. Weber, Phy-
tologia 51: 370. 1982. Basionym: Arabis hol¬
boellii var. fendleri S. Watson, in A. Gray, Syn-
op. FI. N. Amer. 1: 164. 1895. TYPE: U.S.A.
New Mexico: without locality, 1847,4. Fendler
27 (holotype, GH; isotypes, MO, NY not seen,
UC).
Boechera fernaldiana (Rollins) W. A. Weber,
Phytologia 51: 370. 1982. Basionym: Arabis
fernaldiana Rollins, Rhodora 43: 430. 1941.
TYPE: U.S.A. Nevada: Nye Co., Toiyabe Mts.,
E slope of Toiyabe Dome, among rocks &
around sagebrush, 10,000 ft., 13 July 1938,/?.
C. Rollins & T. S. Chambers 2520 (holotype,
GH; isotypes, l)S. MO. UC).
Arabis vivariensis S. L Welsh, Great Basin Naturalist 46:
263. 1986. Boechera vivariensis (S. I.. Welsh) W. A.
Weber. Phytologia 67: 425. 1989. TYPE: U.S.A.
Utah: Uintah Co., Jones Hole, National Fish Hatch¬
ery, T3S, R25E, Sec. 1, 6000 ft.. 16 May 1979, S.
L. Welsh A* E. Neese 18341 (holotype, BUY not seen;
isotypes, CAS, GH, BM, RSA, UC, US).
Welsh (1986) indicated that Arabis vivariensis
differs from A. fernaldiana in style length, fruit
width, and petal length, and Weber (1989) trans¬
ferred the species to Boechera. Although I have not
examined the holotype of A. vivariensis at BRY, the
study of six isotypes supports Rollins (1993b) in
reducing the species to synonymy of A. fernaldiana.
The differences on which 4. vivariensis was based
are rather minor and show continuous variation.
Boechera fruticosa (A. Nelson) Al-Shehbaz,
comb. nov. Basionym: Arabis fruticosa A. Nel¬
son, Bot. Gaz. 30: 190. 1900. TYPE: U.S.A.
Wyoming: Yellowstone National Park, Undine
Falls, dry roadsides, 6 July 1899, 4. & E. Nel¬
son 5681 (holotype, RM; isotypes, GH, MO,
NY not seen, US).
Rollins (1983, 1993b) suggested that this species
is most likely a variant and possibly synonym of
Arabis divaricarpa. The two taxa differ considerably
in habit as well as the trichomes of basal leaves
and merit their recognition as distinct species. The
base of Boechera fruticosa is somewhat woody and
with a few to several stems, and the trichomes are
dendritic on minute stalks. By contrast, B. divari¬
carpa is herbaceous and simple at the base, and
the trichomes are sessile and often with undivided
branches.
Boechera glaucovalvula (M. E. Jones) Al-Sheh¬
baz, comb. nov. Basionym: Arabis glaucoval¬
vula M. E. Jones, Contr. W. Bot. 8: 40. 1898.
TYPE: U.S.A. California: Inyo Co., Darwin
Mesa, Argus Mts., 5000 ft., 8 May 1897, Mar¬
cus E. Jones s.n. (holotype, POM; isotypes,
BM, DS, GH, MO, NY not seen, PH, POM [2],
RM, UC [3], US 12], UTC not seen).
Boechera gracilipes (Greene) Dorn, Brittonia 55:
3. 2003. Basionym: Arabis gracilipes Greene,
Pittonia 4: 193. 1900. TYPE: U.S.A. Arizona:
Coconino Co., Flagstaff, May 1893, N. C. Wil¬
son s.n. (holotype, NDG 4325).
Boechera guiiiiisoniana (Rollins) W. A. Weber,
Phytologia 51: 370. 1982. Basionym: Arabis
gunnisoniana Rollins, Rhodora 43: 434. 1941.
TYPE: U.S.A. Colorado: Gunnison Co., 6 mi.
E of Gunnison, near Saguache Creek, barren
rocky knoll, 8000 ft., 21 May 1938, R. C. Rol¬
lins 2090 (holotype, GH; isotypes, BM. DAO,
GH, RM, UC).
Boechera glareosa Dorn, Brittonia 55: 1. 2(X)3. Syn. nov.
TYPE: U.S.A. Utah: Uintah Go., ca. 14 mi. E of
Jensen, 40°21,077'N, l()9°04417'W, 67(H) ft. (2040
m), 12 May 2(M)2. R. 1). Dorn 9106 (holotype, RM
not seen; isotype, MO).
I have critically examined an isotype of Boechera
glareosa and compared it with the type collection
386
Novon
and extensive material of B. gunnisoniana. There
is not a single character that reliably separates the
two taxa, especially if the overall variation of the
latter is taken into account. The type collection of
B. glareosa represents the first record of B. gun¬
nisoniana from Utah.
Boechera liastatnla (Greene) Al-Shehbaz, comb,
nov. Basionym: Arabis hastatula Greene, Leaf!.
Hot. Obs. Grit. 2: 79. 1910. TYPE: U.S.A.
Oregon: mountains of Imnaha National Forest,
6000 ft., 25 June 1907, A. IE Sampson & G.
A. Pearson s.n. (holotype, US).
Boechera hirshbergiae (S. Boyd) Al-Shehbaz,
comb. nov. Basionym: Arabis hirshbergiae S.
Boyd, Aliso 17: 203. 1998. TYPE: U.S.A. Cal¬
ifornia: San Diego Co., Cuyamaca Mts., E of
Cuyamaea Reservoir along Sunrise Hwy. (S-l),
0.5 mi. SE of junction with Hwy. 79, about 50
ft. N of the road near the Pedro Pages Trail
marker, near 33°00'N, 116°30'W, ca. 1400 m,
19 Mar. 1995, Hirschberg s.n. (holotype, RSA;
isotype, GH).
Boechera hofTiiianii (Munz) Al-Shehbaz, comb,
nov. Basionym: Arabis maxima var. hoffmanii
Munz, Bull. S. Calif. Acad. Sci. 31: 63. 19.32.
TYPE: U.S.A. California: Santa Cruz Island,
ledges in sea cliffs E of Dick’s Harbor, 28 Feb.
1932, R. Hoffmann 653 (holotype, POM).
Boechera holhoellii (Hornemann) A Love & I).
Love, Rot. Not. 128: 513. 1976. Basionym: Ar¬
abis holhoellii Hornemann, El. Danica 11 (Heft
32): tab. 1879. 1827. TYPE: Greenland. “In¬
sulae Disco ad Jacobshavn detexit Inspector
Gronland,” C. Holboll s.n. (holotype, C not
seen).
fhe plate accompanying the original description
is superb and shows all the fine details of the flow¬
ers, seeds, trichomes, and fruits. Boechera holhoellii
is one of the most complex species in the genus,
and Rollins (1993b) divided it into five varieties.
Love and Love (1976, 1982) treated some of those
as B. collinsii (Eernald) A. Love & D. Love, B. re-
trofracta (Graham) A. Love & I). Love, and B. ten¬
uis (Bocher) A. Love & I). Love, whereas Mulligan
(1996) recognized Arabis boivinii G. A. Mulligan,
A. exilis A. Nelson, and A. pinetorum Tidestrom.
Boechera inyoensis (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis inyoensis Rollins, Rho-
dora 43: 457. 1941. TYPE: U.S.A. California:
Inyo Co., Sierra foothills W of Big Pine, near
rocks in coarse granite sand, 15 May 1906, A.
A. Heller 8259 (holotype, GH; isotypes, CAS,
MO, NY not seen, US).
Boechera johnstonii (Munz) Al-Shehbaz, comb,
nov. Basionym: Arabis johnstonii Munz, Bull.
S. Calif. Acad. Sci. 31: 63. 1932. TYPE:
U.S.A. California: Riverside Co., Kenworthy,
Thomas Valley, San Jacinto Mts., 4500 ft., 19
May 1922, P. A. Munz & I. M. Johnston 5485
(holotype, POM; isotypes, CAS, GH).
Boechera koehleri (T. J. Howell) Al-Shehbaz,
comb. nov. Basionym: Arabis koehleri T. J.
Howell. El. Northw. Amer. 1: 44. 1897. TYPE:
U.S.A. Oregon: Douglas Co., Cost Mts., along
Umpqua River at Roseburg, cliffs, 17 Apr.
1887, T. J. Howell s.n. (holotype, ORE not
seen; isotypes, GH, NY not seen, US, WTU
not seen).
Although no chromosome count is known lor the
species, the presence of falcate fruits and subden-
dritie trichomes supports the assignment of the spe¬
cies to Boechera.
Boechera laevigata (Muhlenberg ex Willdenow)
Al-Shehbaz, comb. nov. Basionym: Turritis lae¬
vigata Muhlenberg ex Willdenow, Sp. Pi. 3(2):
543. 1801. Arabis laevigata (Muhlenberg ex
Willdenow) Poiret, Encycl. Suppl. 1: 411.
1810. TYPE: U.S.A. Pennsylvania: without lo¬
cality, G. H. E. Muhlenberg s.n. (holotype, PH
not seen).
On the basis of having a chromosome number of
2n = 14 (Rollins, 1993b) and falcate fruits, the
species is easily assigned to Boechera.
Boechera lasiocarpa (Rollins) Dorn. Brittonia 55:
3. 2003. Basionym: Arabis lasiocarpa Rollins,
Syst. Bot. 6: 58. 1981. TYPE: U.S.A. Utah:
Rich Co., 6 mi. W of Garden City, off U.S.
Hwy. 89, Wasatch Range, high ridge, sedimen¬
tary rock, 22 June 1979. R. G. & K. IE Rollins
79307 (holotype, GH; isotypes, MO |2], RM.
UC).
Boechera leniiiionii (S. Watson) W. A. Weber,
Phytologia 51: 370. 1982. Basionym: Arabis
lemmonii S. Watson, Proc. Amer. Acad. 22:
467. 1887. TYPE: U.S.A. California: Lassen
Peak, Sep. 1872, J. G. Lemmon 23 (lectotype,
designated by Rollins (1941: 383), GI1).
Watson (1887) cited nine syntypes under the
original description of Arabis lemmonii. Rollins
(1941) listed Lemmon’s collection as the type, and
this was taken as the lectotype. Rollins (1993b)
recognized four varieties, of which two were treated
by Mulligan (1996) as distinct species, A. depau-
perata A. Nelson & Kennedy and A. drepanoloba
Volume 13, Number 4
2003
Al-Shehbaz
Boechera
387
Greene. I have not examined the holotypes of these
two and therefore refrain from recognizing them at
any rank. The accounts in Rollins (1993b) and
Mulligan (1996) of these two species, as well as A.
lemmonii, A. lyallii, anti A. microphylla, hardly
show any agreement. For example. Mulligan listed
A. nubigena J. F. Macbride & Payson as a synonym
of A. depauperata, whereas Rollins treated it as a
v ariety of A. lyallii and recognized A. depauperata
as a variety of A. lemmonii. Furthermore, Mulligan
described the new species A. codyi G. A. Mulligan,
but this is morphologically indistinguishable from
A. nubigena.
Boechera lignil'era (A. Nelson) W. A. Weber.
Phytologia 51: 370. 1982. Basionym: Arabis
lignifera A. Nelson, Bull. Torr. Bot. Club 26:
123. 1899. TYPE: U.S.A. Wyoming: Sweet¬
water Co., Green River, 4 June 1898, A. Nel¬
son 4711 (holotype, RM; isotypes, GH, MO,
US).
Boechera lyallii (S. Watson) Dorn, Vase. PI.
Wyom. ed. 3: 376. 2001. Basionym: Arabis
lyallii S. Watson, Proc. Amer. Acad. 11: 122.
1875. TYPE: U.S.A. Washington: Fort Coville
to the Rocky Mts., 1861, Lyall s.n. (lectotype,
designated by Rollins (1941: 367), GH).
Arabis murrayi G. A. Mulligan, Rhodora 97: 151. 1996.
Syn. nov. TYPE: Canada. Yukon: Kaskawulsh nun-
atak, junction N & Central arms Kaskawulsh Gla¬
cier, W of Kluane Lake, 6000 ft., unstable slopes, 1
July—1 Aug. 1965, I). F. A' I). H. Murray 91b (ho¬
lotype, DAO).
Watson (1871) cited four collections (two of his
and one each by Lyall and Brewer) under Arabis
drummondii var. alpina S. Watson. However, in the
original description of A. lyallii, he (Watson, 1875)
listed A. drummondii var. alpina as a synonym but
did not cite any collections. It is possible that A.
lyallii was intended as a new name at the specific
rank because of the pre-existence of A. alpina L.
Both Hopkins (1937) and Rollins (1941) indicated
that the type of A. lyallii is Lyall’s specimen above,
and this should be an effective lectotypification of
the species. Hopkins also indicated that Waston 75
(GH) is the type of A. drummondii var. alpina, and
this led Mulligan (1996) to take this collection as
the lectotype of both A. lyallii and A. drummondii
var. alpina. In my opinion, the typification of A.
lyallii by both Hopkins (1937) and Rollins (1941)
should be accepted.
Mulligan (1996) reduced Arabis davidsonii
Greene to synonymy of A. lyallii, whereas Rollins
(1941, 1993a, b) treated these as distinct species.
An examination of the types of both taxa shows that
they are not sufficiently distinct, and 1 tend to agree
with Mulligan's opinion.
A critical study of the type collections of Boech¬
era lyallii and Arabis murrayi, along with a very
broad spectrum of material from throughout the
range of the former, clearly shows that the alleged
difference on which A. murrayi is based (cauline
leaves minutely auriculate or not aurieulate vs.
leaves strongly auriculate in B. lyallii) does not
hold at all. The development of auricles in this spe¬
cies, as well as several others (including B. hol-
boellii), can be dramatic and can vary within a sin¬
gle population. If this character is taken at face
value, one would have to recognize A. murrayi with¬
in populations of the entire range of B. lyallii, not
just in Canada and neighboring Washington. The
alleged differences in the trichomes of both species
(Mulligan, 1996: 153), “mostly 0.125 mm wide” in
A. murrayii versus “mostly 0.25 mm wide” in A.
lyallii, cannot be justified as a reliable difference
either. Trichomes, just as other characters, can vary
in size within the same population, f or these rea¬
sons, I do not find adequate grounds that support
the recognition of A. murrayi.
Boechera microphylla (Nuttall) Dorn, Vase. PI.
Wyom. ed. 3: 376. 2001. Basionym: Arabis mi¬
crophylla Nuttall, in Torrey & A. Gray, FI. N.
Amer. 1: 82. 1838. TYPE: U.S.A. Rocky
Mountains, T. Nuttall s.n. (holotype, PH: iso¬
type, BM).
Boechera missouriensis (Greene) Al-Shehbaz,
comb. nov. Basionym: Arabis missouriensis
Greene, Repert. Sp. Nov. Regni Veg. 5: 244.
1908. TYPE: U.S.A. Missouri: Montier, 15
May 1894, B. F. Bush 51 (holotype, NDG
4416; isotype, GH).
Mulligan (1996) reduced this species to synon¬
ymy of Arabis laevigata, but the differences in leaf
shape, margin, and apex, as well as petal length in
relation to sepals (see Rollins, 1993b: 117) justify
the recognition of two closely related species.
Boechera ophira (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis ophira Rollins, Syst.
Bot. 6: 56. 1981. TYPE: U.S.A. Nevada: Nye
Co., Toiyabe Range, near Ophir Pass, T13N,
R41E, open rocky area. 13 June 1979, R. C.
& K. W. Rollins 79231 (holotype, GH; isotype,
GH).
Boechera oxylohula (Greene) W. A. Weber. Phy¬
tologia 51: 370. 1982. Basionym: Arabis oxy-
lobula Greene, Pittonia 4: 195. 1900. TYPE:
U.S.A. Colorado: Garfield Co., Glenwood
Springs, 18 June 1899, G. E. Osterhout (ho¬
lotype, NDG 4427; isotype, RM).
388
Novon
Boechera pallidifolia (Rollins) W. A. Weber. Phy-
tologia 79: 65. 1996. Basionym: Arabis palli-
difolia Rollins, Cruciferae of Continental
North America 181. 1993. TYPE: U.S.A. Col¬
orado: Gunnison Co., high hill above Sapinero,
steep hillside among rocks & low sagebrush,
18 June 1983, R. C. Rollins , K. W. Rollins &
A. G. Roads 8376 (holotype, GH; isotypes, GH.
UC).
Boechera parisliii (S. Watson) Al-Shehbaz, comb,
nov. Basionym: Arabis parisliii S. Watson, Proe.
Amer. Acad. Arts 22: 468. 1887. TYPE:
U.S.A. California: San Bernardino Co., Bear
Valley, San Bernardino Mts., 6500 ft., June
1886, S. II. Parish 1793 (holotype, GH; iso¬
type, UC).
Boechera patens (Sullivant) Al-Shehbaz, comb,
nov. Basionym: Arabis patens Sullivant, Amer.
J. Sci. 42: 49. 1842. TYPE: U.S.A. Ohio: rocky
banks of Scioto River, near Columbus, 1840.
W. S. Sullivant s.n. (holotype, PH; isotype,
GH).
Boechera pendiilina (Greene) W. A. Weber. Phy-
tologia 51: 370. 1982. Basionym: Arabis pen-
dulina Greene, Leafl. Rot. Obs. Grit. 2: 81.
1910. TYPE: U.S.A. Nevada: Clark Co.,
Charleston Mts., 7000—8000 ft.. May—Oct.
1898, C. A. Purpus 6104 (holotype, US; iso-
types, RM, UC |2]).
Boechera perennans (S. Watson) W. A. Weber,
Phytologia 51: 370. 1982. Basionym: Arabis
perennans S. Watson, Proe. Amer. Acad. Arts
22: 467. 1887. TYPE: U.S.A. Arizona: Pima
Co., Santa Catalina Mts., rocky canyon, 15
Apr. 1881, C. G. Pringle s.n. (holotype, GH;
isotype, MO).
Boechera perstellata (E. Braun) Al-Shehbaz,
comb. nov. Basionym: Arabis perstellata E.
Braun, Rhodora 42: 47. 1940. TYPE: U.S.A.
Kentucky: F ranklin Co., Elkhorn Creek, wood¬
ed hillsides, 6 May 1936, E. L. Braun 1226
(holotype, GH).
Boechera pinzlae (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis pinzlae Rollins, Contr.
Gray Herb. 212: 110. 1982. TYPE: U.S.A. Ne¬
vada: Esmeralda Co., White Mts., bowl on NE
side of Boundary Peak, near center of bowl,
ca. 1 1,200 ft., A. Pinzl 4442 (holotype, GH;
isotype, GH).
Boechera platysperina (A. Gray) Al-Shehbaz,
comb. nov. Basionym: Arabis platysperma A.
Gray, Proe. Amer. Acad. Arts 6: 519. 1865.
TYPE: U.S.A. California: Alpine Co.. Ebbetts
Pass, 1863, W. H. Brewer 1989 (lectotype, des¬
ignated by Rollins (1941: 473), GH; isotypes,
UC, US).
Gray (1865) cited two syntypes under the origi¬
nal description of Arabis platysperma. Rollins
(1941) listed Brewer’s collection as the type, and
this was accepted as the lectotype of the species.
Boechera piihcrula (Nuttall) Dorn, Brittonia 55:
3. 2003. Basionym: Arabis puberula Nuttall, in
Torrey & A. Gray, El. N. Amer. 1: 82. 1838.
TYPE: U.S.A. Oregon?: Blue Mts., T. Nuttall
s.n. (holotype, PH).
Boechera pulchra (M. E. Jones ex S. Watson) W.
A. Weber, Phytologia 51: 370. 1982. Basion¬
ym: Arabis pulchra M. E. Jones ex S. Watson,
Proe. Amer. Acad. 22: 468. 1887. TYPE:
U.S.A. Nevada: Ormsby Co., Empire City. 19
June 1882. M. E. Jones 3765 (lectotype, des¬
ignated by Rollins (1941: 459). GH; isotypes,
1)S. MO. NY not seen. POM, UC, US [2|).
Watson (1887) cited seven syntypes under the
original description of Arabis pulchra. Rollins
(1941) listed Jones 3765 collection as the type, and
this was taken here as the lectotype.
Boechera pusilla (Rollins) Dorn, Vase. PI. Wyom.
ed. 3: 376. 2001. Basionym: Arabis pusilla
Rollins, Contr. Gray Herb. 212: 107. 1982.
TYPE: U.S.A. Wyoming: Fremont Co., off Wy¬
oming State Hwv. 28, 39 mi. SW ol Pander,
cracks & crevices ol huge metamorphosed
rocks, 20 June 1981, It C. & K. W. Rollins
81366 (holotype, GH; isotypes, GH, MO. RM.
UC).
Boechera pygmaea (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis pygmaea Rollins, Rho¬
dora 43: 476. 1941. TYPE: U.S.A. California:
Tulare Co., basin of the Upper Kern River,
Volcano Meadows (formerly Whitney Mead¬
ows), 21 July 1904. //. M. Hall & II. I). Bab¬
cock 5465 (holotype, GH; isotypes not seen,
MO, NY, OSC, US).
Boechera rectissima (Greene) Al-Shehbaz, comb,
nov. Basionym: Arabis rectissima Greene, Pit-
tonia 4: 191. 1900. TYPE: U.S.A. California:
Fresno Co., 1890, Peckinpah s.n. (holotype,
NDG 4469; isotype. NY not seen).
Boechera repanda (S. Watson) Al-Shehbaz,
comb. nov. Basionym: Arabis repanda S. Wat¬
son. Proe. Amer. Acad. Arts 11: 122. 1875.
TYPE: U.S.A. California: Mariposa Co., Yo-
semite Valley, 1866, //. /V. Bolander 4881 (ho¬
lotype, G11; isotype, UC).
Volume 13, Number 4
2003
Al-Shehbaz
Boechera
389
Boechera rigidissima (Rollins) Al-Shehbaz,
comb. nov. Basionym: Arabis rigidissima Rol¬
lins, Rhodora 43: 380. 1941. TYPE: U.S.A.
California: Trinity Co., Mary Blaine Mt., grav¬
elly or rocky soil. 6400 ft., 3 Aug. 1935, ,/. R.
Tracy 14469 (holotype, GH; isotype, DS).
Boechera rollei (Rollins) Al-Shehbaz, comb. nov.
Basionym: Arabis rollei Rollins, Harvard Pap.
Bot. 4: 43. 1993. TYPE: U.S.A. California:
Siskiyou Co., divide betw. Applegate & Klam¬
ath Rivers, Red Butte-Kangaroo Mt.—Lilypad
Lake-Towhead Eake area, T47N, RI2W, Sec.
13, 5400-5900 ft., 4 Aug. 1983, W. E. Rolle
831 (holotype, GH; isotypes, GH, JEPS).
Boechera schislacea (Rollins) Dorn. Brittonia 55:
3. 2003. Basionym: Arabis schistacea Rollins,
Contr. Dudley Herb. 3: 370. 1946. TYPE:
U.S.A. Nevada: Nye Co., 15 mi. W of Round
Mt.. Toiyabe Mts., open slope, 9000 ft., I 1 July
1938. R. C. Rollins & T. S. Chambers 2512
(holotype, DS; isotype, GH).
Boechera selhyi (Rvdberg) W. A. Weber, Phyto-
logia 51: 371. 1982. Basionym: Arabis selbyi
Rydberg, Bull. Torr. Bot. Club 31: 557. 1904.
TYPE: U.S.A. Colorado: Ouray Co., W of Our¬
ay, 9 Sep. 1901, L. M. Underwood & A. D.
Selby 207 (holotype. NY not seen).
Boechera shocklevi (Munz) Dorn, Brittonia 55:
3. 2003. Basionym: Arabis shockleyi Munz,
Bull. S. Calif. Acad. Sci. 31. 62. 1932. TYPE:
U.S.A. Nevada: Esmeralda Co., Mellin Mt.,
near Candelaria, May 1884, W. H. Shockley
366 (holotype, GH: isotypes, JEPS. NDG, US).
Boechera shortii (Fernald) Al-Shehbaz, comb,
nov. Basionym: Arabis perstellata var. shortii
Fernald, Rhodora 48: 208. 1946. Sisymbrium
dentatum Torrey, in Short, 3rd Suppl. Cat. PI.
Kentucky 338. 1833, non S. dentatum Allioni,
FI. Pedem. 1: 275. 1785. TYPE: U.S.A. Ken¬
tucky: sandy banks of the Ohio River, C. W.
Short s.n. (lectotype, designated by Mulligan
(1996: 143). DWC; isotvpe, GH).
For full synonymy of the species, see Rollins
(1993b).
Boechera sparsiflora (Nuttall) Dorn, Vase. PI.
Wyom. ed. 3: 376. 2001. Basionym: Arabis
sparsiflora Nuttall, in Torrey & A. Gray, FI. N.
Amer. 1: 81. 1838. TYPE: U.S.A. Rocky Mts.,
toward sources of Oregon. T. Nuttall s.n. (ho¬
lotype, BM; photo, GH; isotype. PH).
Rollins (1941) indicated that Nuttalls sheet at
PH is the isotype, whereas Mulligan (1996) consid¬
ered that sheet a mixture of more than one species
and designated the specimen to the far right as the
lectotype. I have examined the type at BM and
agree with Rollins’s conclusion.
Boechera siricta (Graham) Al-Shehbaz, comb,
nov. Basionym: Turritis stricta Graham, Edin¬
burgh New Philos. J. 7: 350. 1829. Arabis
drummondii A. Gray, Proc. Amer. Acad. Arts
6: 187. 1864. TYPE: U.S.A. Rocky Mts.,
grown from seeds, Drummond s.n. (holotype,
E).
The nomenclature of this species is confusing,
and the species epithet has not been used since the
mid 19th century. The species is far better known
as Arabis drummondii A. Gray, a name proposed
by Gray (1864) to avoid the creation of a later hom¬
onym that would have resulted from the transfer of
Turritis stricta to Arabis because of the existence of
A. stricta Hudson published in 1777 (FI. Angl. I:
292. 1777). The transfer by Love and Love (1976)
of A. drummondii to Boechera is illegitimate be¬
cause these authors did not use T. stricta, the le¬
gitimate, validly published, and earliest basionym
for the species.
Boechera suhpinnatifHla (S. Watson) Al-Sheh-
baz, comb. nov. Basionym: Arabis subpinnati-
fida S. Watson, Proc. Amer. Acad. Arts 20:
353. 1885. TYPE: U.S.A. Oregon: Josephine
Co., Waldo, 3 June 1884, T. Howell s.n. (lec¬
totype, designated by Rollins (1941: 454), GH;
isotypes, NY not seen, US).
Watson (1885) cited two syntypes under the orig¬
inal description of Arabis subpinnatifida. Rollins
(1941) listed Howell’s collection as the type, and
this was taken as the lectotype of the species.
Boechera suffrutescens (S. Watson) Dorn, Brit¬
tonia 55: 3. 2003. Basionym: Arabis suffrutes¬
cens S. Watson, Proc. Amer. Acad. Arts 17:
362. 1882. TYPE: U.S.A. Oregon: Baker Co.,
bluffs of Snake River & vicinity, Apr. 1881,
W. C. Cusick 919 (holotype, GH).
Boechera tiehmii (Rollins) Al-Shehbaz, comb,
nov. Basionym: Arabis tiehmii Rollins, J. Ar¬
nold Arbor. 64: 496. 1983. TYPE: U.S.A. Ne¬
vada: Washoe Co., Sierra Nevada, Carson
Range, ridge N of Mt. Rose, 3/4 air mi. N-NW
of peak, T17N, R IRE, Sec. 11, 10,200 ft., near
rock outcrops on decomposed granite, 19 Aug.
1982. A. Tiehrn 7561 (holotype, (ill; isotypes,
CAS, GH. RSA).
390
Novon
Boechera williamsii (Rollins) Dorn, Vase. PI.
Wyom. ed. 3: 376. 2001. Basionym: Arabis wil-
liamsii Rollins, Syst. Bot. 6: 62. 1981. TYPE:
U.S.A. Wyoming: Fremont Co., off Wyoming
Hwy. 28, E of Continental Divide on SE ap¬
proach to Wind River Mts., coarse granitic
soil, 7800 m, 26 June 1979, R. C. & M. L.
Rollins 79332 (holotype, GH; isotypes, Gil,
MO, RM. UC).
Acknowledgments. I am most grateful to Thom¬
as G. Lammers, Neil A. Harriman, Steve L. O’Kane,
Jr., and Victoria C. Hollowed lor their critical re¬
views of and comments on the manuscript. 1 thank
Nicholas J. Turland for advice on certain nomen-
clatural problems and Anthony R. Brach and Judy
Warnement for help with the literature. 1 also thank
the directors and curators of the herbaria cited for
the loans of material.
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A Synopsis of Tropidocarpum (Brassicaceae)
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. Twisselmannia californica is trans¬
ferred to Tropidocarpum (Brassicaceae). A key to
the four species of Tropidocarpum is presented.
Disjunction between the North and South American
species of the genus is discussed.
Key words: Agallis, Brassicaceae, Iruil diversi¬
ty, Tropidocarpum , Twisselmannia.
As presently delimited, Tropidocarpum Hooker
(Brassicaceae) consists of lour species, of which
Twisselmannia californica Al-Shehbaz and Agallis
lanata (Barneoud) Gilg & Muschler ex (). E. Schulz
were previously placed in monotypic genera. The
narrowly endemic T. californica is restricted to a
limited number of scrub habitats in the Central Val¬
ley of California (Al-Shehbaz, 1999). Although
Twisselmannia Al-Shehbaz was reduced to synon¬
ymy of Tropidocarpum by Appel and Al-Shehbaz
(2002), formal transfer of its species to Tropidocar¬
pum was not proposed. Agallis lanata is very rare
and occurs disjunctly in central Chile in habitats
similar ecologically to those where species of Tro¬
pidocarpum grow. It was transferred to Tropidocar¬
pum by Al-Shehbaz and Price (2001). Of the other
two species originally assigned to Tropidocarpum ,
T. gracile Hooker is a relatively widespread and
abundant species occurring in open habitats and
lower montane slopes from Baja California north to
central California (see below), whereas T. cappari¬
deum Greene is an extremely rare species previ¬
ously known ordy from the vicinity of Mt. Diablo in
the San Francisco Bay area of California (Rollins,
1993a, 1993b). Tropidocarpum capparideum was
not collected for nearly 45 years and was last seen
in 1957 and presumed extinct (Rollins, 1993b).
However, the species has been collected in 2000
by Meredith Osborne (CDA) and in 2001 by Fred
Hrusa (CDA) and Dieter Wilken (SBBG). These
collections, all cited below, were made from Fort
Hunter biggett in Monterey County, and I have con¬
firmed their identity.
All four species are annuals with a remarkably
similar morphology, especially in their indumentum
(with coarse simple trichomes mixed with smaller
forked ones), petiolate and pinnatisect to pinnatifid
cauline leaves, racemes bracteate throughout, yel¬
low flowers occasionally tinged with purple, sta-
minal filaments with dilated bases, angustiseptate
fruits (flattened at a right angle to the septum), mu¬
cilaginous seeds, and incumbent cotyledons. How¬
ever, they have drastically different fruits (Fig. 1).
Twisselmannia has a very distinctive obdeltoid siI-
icle (4 —5 X 3-4 mm) with leathery-thickened and
tuberculate-rugose valves, whereas Tropidocarpum
gracile has narrowly linear siliques up to 7 cm X
2 mm. Both T. capparideum and T. lanatum (Bar¬
neoud) Al-Shehbaz & R. A. Price have oblong
fruits, but the former has four rather than two
valves, a condition rarely seen elsewhere within the
Brassicaceae (Al-Shehbaz, 1984).
Molecular studies of Twisselmannia californica ,
Tropidocarpum gracile, anti T. lanatum (Price, un¬
published) demonstrate that the three species are
almost identical in sequence, differing by only I or
2 base pair substitutions, for the chloroplast gene
n<7/iF and the nuclear ribosomal DNA ITS region.
The three species form a monophyletic clade, with
100% bootstrap support, and this clade is weakly
separated (54% bootstrap support) from a mono¬
phyletic Descurainia Webb & Bert helot including
Hugueninia Reichenbach (Price, unpublished). Be¬
cause of the extremely small amount of sequence
divergence observed among the three species for
the rapidly changing rulhF and ITS regions, it ap¬
pears that both the drastic change in fruit shape
between Twisselmannia and its Tropidocarpum con¬
geners and the North American—South American
disjunction involving T. lanatum are relatively re¬
cent events in the history of the Brassicaceae.
Fruit anil seed morphology have traditionally
been heavily emphasized in generic and tribal clas¬
sification of the Brassicaceae (Hedge, 1976; Al-
Shehbaz, 1984), but recent molecular phylogenetic
studies suggest that fruit and seed morphology have
been subject to frequent homoplasy (Koch et al.,
1999, 2001) and that many of the commonly rec¬
ognized tribes in the family are highly artificial.
Remarkable differences in fruit morphology are
found in several other genera of the Brassicaceae.
For example, Rorippa Scopoli (ca. 75 spp., distrib¬
uted worldwide) has fruits ranging from narrowly
Novon 13: 392-395. 2003.
Volume 13, Number 4
2003
Al-Shehbaz
Synopsis of Tropidocarpum
393
linear to oblong or globose and with two or rarely
three to six valves (Appel & Al-Shehbaz, 2002).
Another striking example of fruit diversity is found
in Vella L., a genus of seven species of shrubs en¬
demic to Algeria, Morocco, and Spain (Appel & Al-
Shehbaz, 2002). All species of Vella have seg¬
mented. 1- to 8-seeded fruits, connate median
staminal filaments, and a base chromosome number
of x = 17 (Warwick & Al-Shehbaz, 1998). One spe¬
cies, V. bourgaeana (Cosson) Warwick & Al-Sheh¬
baz, previously treated as the monotypic genus Eu-
zomodendron Cosson (Lopez Gonzalez, 1993), has
linear-lanceolate fruits and seeds broadly winged
all around, whereas the remaining species have glo¬
bose to ellipsoid fruits and seeds with vestigial
wings. Molecular data (Warwick & Black, 1994;
Crespo et al., 2000) clearly show that Euzomoden-
dron is nested within Vella, and thus they form a
monophyletic genus. Several other genera of Bras-
sicaceae show very wide ranges of fruit morphology.
These include Draba L. (Schulz. 1936), Heliophila
L. (Marais, 1970), Graellsia Boissier (Poulter,
1956), /salts L. (Davis, 1964), Leavenworthia Torrey
(Rollins, 1963), Ornithocarpa Rose (Rollins, 1969).
and Sphaerocardamum Schauer (Bailey, 2001).
The almost identical sequence similarity be¬
tween Twisselmannia and species of Tropidocarpum
contrasts with the apparent rapidity with which ma¬
jor changes in fruit morphology can sometimes oc¬
cur among closely related taxa. A classic example
of a similar situation was elucidated earlier by Syts-
ma and Gottlieb (1986) in the evolution of the strik¬
ingly divergent Heterogaura from within the genus
Clarkia Pursh (Onagraceae). It would be particu¬
larly interesting to conduct comparative genetic and
developmental studies of the fruits of Tropidocar¬
pum, including Twisselmannia.
Another interesting aspect of the genus Tropi¬
docarpum is the large disjunction between the Med¬
iterranean climate zones of California and Chile
(Fig. 1), which was unrecognized until Al-Shehbaz
and Price (2001) transferred Agallis lanata to Tro¬
pidocarpum. The pattern of disjunction among spe¬
cies of many genera between these environmentally
similar areas of North and South America was el¬
egantly covered by Raven and Axelrod (1978).
Key to the Species oe Tropidocarpum
la. Fruits obdeltoid, spreading or antrorsely hirsute,
4- to 8-seeded; valves thick leathery, tubercu-
late-rugose distally. T. californicum
lb. Fruits linear, oblong, or elliptic, retrorsely hir¬
sute (except when glabrescent), 16- to 70-seed-
ed; valves thin-leathery, smooth.
2a. Fruits narrowly linear, (2.5—)3—6(-7) cm X
1.5—2 mm. T. gracile
2b. Fruits oblong to elliptic, 1— 2(—2.5) cm X 3—
5 mm.
3a. Fruit valves 4; anthers oblong, 0.4—0.5
mm; fruit 25- to 40-seeded; California
. T. capparideum
3b. Fruit valves 2; anthers ovate, 0.15—0.25
mm; fruit 16- to 26-seeded; Chile . . .
. T. lanatum
1. Tropidocarpum californicum (Al-Shehbaz)
Al-Shehbaz, comb. nov. Basionym: Twissel¬
mannia californica Al-Shehbaz. Novon 9: 132.
1999. TYPE: U.S.A. California: Kings Co., S
of Kettleman City, E side of Interstate Hwy. 5,
3 mi. N of Kern Co. line, grazed valley salt¬
bush scrub, 30 Mar. 1994. Ed LaRue s.n. (ho-
lotvpe, UCR).
Distribution. U.S.A. California: Kings County.
Specimens examined. U.S.A. California: Kern Co.,
Kern National Wildlife Refuge, Twisselmann 10447
(CDA); Kings Co., ca. 13 mi. S of Kettleman City, along
pipeline access road lying 0.3 mi. E of Interstate 5, 3.1
mi. N of Kern Co. line, Taylor 17098 & Ertter (MO. UC).
To my knowledge, the collections above repre¬
sent all that is known for the species.
2. Tropidocurpum capparideum Greene, Pit-
tonia 1: 217. 1888. TYPE: U.S.A. California:
[Contra Costa Co.], alkaline valley lands skirt¬
ing the San Joaquin River, [Byron Springs], 25
Mar. 1888, E. L. Greene s.n. (holotype, NDG
02806).
Distribution. U.S.A. California: Contra Costa
and Monterrey Counties.
Specimens examined. U.S.A. California: Contra Cos¬
ta Co., Byron Hot Springs, Hoover 806 (MO); Monterry
Co., Fort Hunter Liggett, adjacent to Gabilan Road, ca.
0.8 mi. SSE of intersection with San Migelito Loop, Hrusa
15805, with Wilken & Hazebrook (CDA, MO), Osborne 16,
with Guzman & Witmore (CDA, MO). Hrusa 15804, with
Wilken A' Hazebrook (CDA, MO); \K of Gabilan Creek, ca.
0.4 km S of junction of Gabilan and San Migelito Loop
roads, Wilken 15876, with Morosco (MO, SBBG).
The single specimen in Greene’s herbarium col¬
lected in March 1888 was from Byron Springs, a
locality not given in the original publication of the
species. This specimen is taken herein as the 1 io-
lotype.
3. Tropidocarpum gracile Hooker, Icon. PI. t.
43. 1836. TYPE: U.S.A. California: Monterey
Co., Monterey, David Douglas s.n. (holotype,
K).
Distribution. Mexico. Northern Baja California.
U.S.A. California: Inner Coastal Range, Cascade
394
Novon
Figure 1. Fruits and distributions of Tropidocarpum species. —A, B. T. californicum (dot on map). —C, I). T. cap-
parideum (asterisk). —E, F. T. gracile (stippled area). —G, H. T. lanatum (star). Cross sections in B, D, F, and H are
made in widest portion of fruit. Scale: A, C, E, G = 5 mm; B, D, F, H = 1 mm. Dotted lines represent Tropic of
Cancer. Equator, and Tropic of Capricorn. Drawn by Al-Shehbaz: A, B from LaRue s.n. (holotype, UCR); C, D from
Hoover 806 (MO); E, F from Blankinship s.n. (MO); G. H from Claude Joseph 2791 (US).
Volume 13, Number 4
2003
Al-Shehbaz
Synopsis of Tropidocarpum
395
Range foothills, Tehachapi Mountain, Great Central
Valley, San Francisco Bay Area, Inner South Coast
Range, Southwest California, and western Mojave
Desert (Rollins, 1993b).
Specimen examined. U.S.A. California: Lake Co., Mt.
Konocti, 29 Apr. 1929. Blankinship s.n. (MO).
4. Tropidocarpum laiiatiim (Barneoud) Al-
Shehbaz & R. A. Price, Novon 1 1: 293. 2001.
Basionym: Lepidium lanatum Barneoud, in
Gay, FI. Chile 1: 167. 1846. TYPE: Chile.
Santiago, C. Gay s.n. (holotype, P; photos, F,
US).
Distribution. Chile, Regibn IV and Region Me-
tropolitana.
Specimens examined. CHILE. Region IV: Pray Jorge,
Las Papas, Jiles 720 (S); Ovalle, Bosque Fray Jorge, Spar-
re 3029 (S). Region Metropolilana: Santiago, Claude-
Joseph 2232. 2267 (US); Lo Prado, Claude-Joseph 2791
(US).
The above collections are the only ones known
for the species, and the latest, Jiles 720, was made
on 22 August 1948. No more recent collections are
knowti for the species.
Acknowledgments. I am grateful to Dean Wil¬
liam Taylor and Barbara Ertter (both at UC) for
plant material of Twisselmannia and information
concerning the population status of the species. I
am much indebted to Fred Hrusa, Dieter H. Wilk-
en, and Meredith Osborne for sending the material
ol Tropidocarpum capparideum cited above. I thank
Barbara J. Hellenthal for providing information on
the holotype of T. capparideum, Robert A. Price for
his unpublished molecular data on Tropidocarpum,
and Neil A. Harriman, Victoria C. Hollowell,
Thomas G. Lammers, and one anonymous reviewer
for their critical comments on the manuscript.
Literature Cited
Al-Shehbaz, I. A. 1984. The tribes of Cruciferae (Brassi-
caceae) in the southeastern United States. ,1. Arnold Ar¬
bor. 65: 343-373.
-. 1999. Twisselmannia (Brassicaceae), a remark¬
able new genus from California. Novon 9: 132-135.
- & B. A. Price. 2001. The Chilean Agallis and
Californian Tropidocarpum (Brassicaceae) are congener¬
ic. Novon 11: 292-293.
Appel. 0. & I. A. Al-Shehbaz. 2002. Cruciferae. Pp. 75—
174 in K. Kubitzki (editor), The Families and Genera
of Vascular Plants, 5. Springer-Verlag, Berlin-Heidel¬
berg.
Bailey, C. 1). 2001. Systematics of Sphaerocardamum
(Brassicaceae) and Related Genera. Ph.l). Dissertation,
Cornell University, Ithaca, New York.
Crespo, M. B., M. D. Lledo. M. F. Fay & M. W. Chase.
2000. Subtribe Vellinae (Brassiceae, Brassicaceae): A
combined analysis of ITS nrDNA sequences and mor¬
phological data. Ann. Bot. 86: 53—62.
Davis, P. H. 1964. Materials for the Flora of Turkey: VIII.
Cruciferae. I: Isatis. Notes Roy. Bot. Card. Edinburgh
26: 11-25.
Hedge, I. C. 1976. A systematic and geographical survey
of the Old World Cruciferae. Pp. 1—45 in J. G. Vaughan
et al. (editors). The Biology and Chemistry of the Cru¬
ciferae. Academic Press, London.
Koch. M.. J. Bishop & T. Mitchell-Olds. 1999. Molecular
systematics and evolution of Arabidopsis and Arabis. PI.
Biol. 1: 529-537.
-. B. Haubold & T. Mitchell-Olds. 2001. Molecular
systematics of the Brassicaceae: Evidence from coding
plastidic matV. and nuclear Chs sequences. Amer. J.
Bot. 88: 534-544.
Ldpez Gonzalez, G. 1993. Euzomodendron. In: S. Castro-
viejo et al. (editors), FI. Iberica 4: 344—346. Real Jardfn
Botanico, Madrid.
Marais, W. 1970. Cruciferae. In: L. F. Code! et al. (editors),
FI. Southern Africa 13: 1-118. Government Printer.
Pretoria.
Poulter, B. A. 1956. The genus Graellsia. Notes Roy. Bot.
Gard. Edinburgh 22: 85—93.
Raven, P H. & D. I. Axelrod. 1978. Origin and relation¬
ships of the California flora. Univ. Calif. Publ. Bot. 72:
1-134.
Rollins, R. C. 1963. The evolution and systematics of
Leavenworthia (Cruciferae). Contr. Gray Herb. 192: 3—
98.
-. 1969. A remarkable new crucifer from Mexico.
Contr. Gray Herb. 198: 3—8.
-. 1993a. The Cruciferae of Continental North
America. Stanford llniv. Press, Stanford.
-. 1993b. Tropidocarpum. Pp. 447—448 in J. C.
Hickman (editor). The Jepson Manual. Univ. California
Press, Berkeley.
Schulz, O. F. 1936. Cruciferae. In: A. Engler & H. Harms
(editors). Die natiirlichen Pflanzenfamilien, ed. 2, 17B:
227-658. Verlag von Wilhelm Engelmann, Leipzig.
Sytsma, K. J. & L. D. Gottlieb. 1986. Chloroplast DNA
evidence for the origin of the genus Heterogaura from
a species of Clarkia (Onagraceae). Proc. Natl. Acad.
Sci. U.S.A. 83: 5554-5557.
Warwick, S. 1. & I. A. Al-Shehbaz. 1998. Generic evalu¬
ation of Boleum, Euzomodendron, and Vella (Brassica¬
ceae). Novon 8: 321—325.
- & L. I). Black. 1994. Evaluation of the subtribes
Moricandiinae, Savignyinae, Vellinae and Zillinae
(Brassicaceae, tribe Brassiceae) using chloroplast DNA
restriction site variation. Canad. J. Bot. 72: 1692—1701.
Transberingia, a New Generic Name Replacing the Illegitimate
Beringia (Brassicaceae)
l It sun A. Al-Shehbaz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, II.S.A.
ihsan.al-shehhaz@mobot.org
Steve L. O'Kane, Jr.
Department of Biology, University of Northern Iowa, Cedar Falls, Iowa 50614-0421, U.S.A.
steve.okane@uni.edu
ABSTRACT. Transberingia Al-Shehbaz & O'Kane
(Brassicaceae) replaces Beringia R. A. Price, Al-
Shehhaz & O’Kane, a later homonym to a genus of
the red algae. The new combinations T. bursifolia
and T. bursifolia subsp. virgata are made for the
sole species of this genus.
Key words: Arabidopsis, Beringia , Brassica¬
ceae, North America, Russia, Transberingia.
Beringia R. A. Price, Al-Shehbaz & O'Kane
(Brassicaceae) was named by Price et al. (2001) as
a monotypic genus to accommodate one species of
mustard excluded by Al-Shehbaz et al. (1999) from
Arabidopsis (DC.) Heynhold. The species, hereafter
Transberingia bursifolia, is distributed in western
North America across the Bering Strait, hence the
generic name Transberingia, into the Russian Far
Fast (Berkutenko. 1988; Price et al.. 2001; Rollins,
1993). As indicated by Aiken et al. (2003), Ber¬
ingia is a later homonym of a genus of Rhodophyta
(red algae) named by Perestenko (1975). Therefore,
the following nomenclatural adjustment is proposed
to replace the illegitimate name Beringia of the
Brassicaceae, as well as a new combination at the
species and subspecies ranks recognized by Price
et al. (2001).
Transberingia Al-Shehbaz & O'Kane, nom. nov.
Replaced name: Beringia R. A. Price, Al-
Shehbaz & O’Kane, Novon 11: 333. 2001, non
Beringia Perestenko, Bot. Zhurn. (Moscow &
Leningrad) 60: 1683. 1975. TYPE: Transber¬
ingia bursifolia (DC.) Al-Shehbaz & O'Kane.
Transberingia bursifolia (DC.) Al-Shehbaz &
O’Kane, comb. nov. Basionym: Nasturtium
bursifolium DC., Syst. Nat. 2: 194. 1821.
TYPE: [Russia]. Kamtchatka, Friedrich F. /,.
von Fischer s.n. (holotype. G-DC).
For a detailed synonymy of the species and fol¬
lowing subspecies, see Price et al. (2001).
Transberingia bursifolia subsp. virgata (Nultall)
Al-Shehbaz & O'Kane, comb. nov. Basionym:
Sisymbrium virgatum Nutlall, in Torrey & A.
Gray, FI. N. Amer. 1: 93. 1838. TYPE: U.S.A.
Rocky Mts. [probably Wyoming], “sources of
Sweet Water of the Platte,” Thomas Nuttall s.n.
(holotype. BM; isotypes, Gil. NY. PH).
Acknowledgments. We thank Victoria C. Hol¬
lowed and Charlotte M. Taylor for their critical
comments on the manuscript.
I.iterature Cited
Aiken, S. G.. M. J. Dallwitz, L. L. Consaul, C. I.. Mc-
Jannet, L. J. Gillespie, R. b. Boles, G. W. Argus, J. M.
Gillett. P. J. Scott. R. F.lven, M. C. LeBlanc, A. K.
Brysting & If. Solstad. 2003. Flora of the Canadian
Arctic Archipelago: Descriptions, Illustrations, Identi¬
fication, and Information Retrieval. Version: 29th April
2003. (http://www.mun.ea/biology/delta/arcticf/).
Al-Shehbaz. 1. A., S. I.. O’Kane, Jr. & R. A. Price. 1999.
Generic placement of species excluded from Arabidopsis
(Brassicaceae). Novon 9: 296—307.
Berkutenko, A. N. 1988. Brassicaceae. Pp. 38—115 in S.
S. Charkevicz [Kharkevich] (editor), Plantae Vasculares
Orientis Extremi Sovietiei, Vol. 3. Leningrad.
Perestenko, L. P. 1975. The red algae of the Far-Eastern
Seas of the U.S.S.R. Foliose Cryptonemiacean algae
(Cryptonemiales, Rhodophyta). Bot. Zhurn. (Moscow &
Leningrad) 60: 1676—1697.
Price, R. A., 1. A. Al-Shehbaz A 1 S. L. O’Kane, Jr. 2001.
Beringia (Brassicaceae), a new genus of arabidopsid af¬
finities from Russia and North America. Novon 11:
332-336.
Rollins, R. C. 1993. The Crueiferae of Continental North
America. Stanford Univ. Press, Stanford.
Novon 13: 396. 2003.
An Overlooked New Species of Habenaria (Orchidaceae) from
Central Brazil
Joao A. N. Batista and Luciano de Bern Bianchetti
Embrapa Recursos Geneticos e Biotecnologia, Parque Estagao Biologiea. Final Av. W5
Norte, CP 02372, Brasilia-DF, 70770-901, Brazil, janb@cenargen.embrapa.br;
bianchet@cenargen.embrapa.br
Eduardo G. Gongalves
Universidade de Sao Paulo, Dept, de Botanica, CP 11461, Sao Paulo-SP, 05422-970, Brazil.
edggon@hotmail.com
Abstract. Habenaria pabstii, a new species (Or-
chideae, Orchidaceae) from the core region of the
cerrado vegetation of central Brazil, is described
and illustrated. The species has been collected
since 1961, but was repeatedly misidentified as
other taxa. The species is similar to those in section
Macroceratitae Kraenzlin, but distinct by the short¬
er, forward-arched, free spur and the shorter, non¬
involute stigmas.
RESUMO. Habenaria pabstii, uma nova especie
(Orchideae, Orchidaceae) da regiao nuclear do cer¬
rado do Brasil central 6 descrita e ilustrada. A es¬
pecie tern sido coletada desde 1961, mas foi re-
petidamente identificada erradamente. A especie e
similar aquelas da seyao Macroceratitae , mas dis-
tinta pelo calcar mais curto, livre. projetado para
frente e pelos estigmas mais curtos e nao involutos.
Key words: Brazil, cerrado, Habenaria , Orchi¬
daceae.
Habenaria Willdenow is the second-largest or¬
chid genus in Brazil, with about 165 species ac¬
cording to the last major survey of Brazilian orchids
(Pabst & Dungs, 1975). The genus is composed ex¬
clusively of terrestrial species and is typical of open
grasslands. Most of the species have small, green,
inconspicuous flowers that are of little interest to
cultivators and thus rarely seen in cultivation. As
a consequence, work on the genus has relied almost
completely on botanical collections and on the
study of dried material. The three major treatments
of the genus for the country so far are those of
Cogniaux (1893) in Flora Brasiliensis, Hoehne
(1940) in Flora Brasilica, and Pabst and Dungs
(1975, 1977) in Orchidaceae Brasilienses. The large
number of species, the poor characterization or lit¬
tle information on many taxa, and the existence of
several species complexes has given rise to a sit¬
uation where identification of the species is often
troublesome, with different names being applied to
the same species and some species being over¬
looked.
The main center of diversity of Habenaria in
Brazil is the cerrado. The cerrado is a species-rich
savanna vegetation covering 2 million km 2 of cen¬
tral Brazil (Ratter et al., 1997). Among the Haben¬
aria species-rich areas in central Brazil is the core
region of the cerrado vegetation, including the Dis¬
trito Federal where 77 taxa are known (Batista &
Bianchetti, 2003) and neighboring areas in the
state of Goias. In the course of a survey of the or¬
chid flora of this cerrado region, several taxa of
Habenaria have been collected that we have been
unable to assign to any known New World species
in the genus, one of which, with large anti showy
flowers, is hereby described as new.
Habenaria pabstii J. A. N. Batista & Bianchetti,
sp. nov. TYPE: Brazil. Distrito Federal: Bra¬
silia, Fazenda Agua Limpa (UnB), 15°57'S,
47°55'W, 1070 m, Corrego da Onya, 2 Feb.
1994, L. B. Bianchetti, J. A. /V. Batista & B.
M. T. Walter 1489 (holotype. CEN; isotvpes,
HB, K, MO. SP). Figures 1 and 2.
Habenaria pabstii florum generali aspectu speciebus
sect. Macroceratitae similis sed calcari breviore, libero,
frontaliter arcuato et stigmatibus brevioribus, non involutis
differt. Etiam H. urbaniana similis sed floribus frequenter
longioribus et struetura columnae prominenti lobo medi-
ano rostelli praedito appendicem similitudine cavernae
formans, haec ultra antheras producta differt.
Terrestrial herb. Tuber basal, oblong to subspher-
ic, 1.2—2.8 X 0.8—1.7 cm; roots ca. 1 mm wide,
sparsely hairy, whitened; stem erect, leafy, 20—60
X 0.3—0.4 cm including the inflorescence. Leaves
7 to 11, the 3 or 4 lowermost sheath-like, the larg¬
est around the center of stem, (3)4 to 5(6), spread.
Novon 13: 397^102. 2003.
398
Novon
Figure 1. Habenaria pabstii Batista & Bianchetti. —A. Habit. —B. Flower, 3 A view. —C. Floral diagram. —I). Column,
% view. —E. Pollinaria. Drawn from the type collection, Bianchetti et at. 1489, by Eduardo Gongalves.
linear-lanceolate, 11-21 X 1 — 1.4 cm, acute, the
1(2) upper leaves decreasing in size and similar to
the lower bracts. Inflorescence 5—10 cm long, few-
flowered; bracts ovate-lanceolate, 1.8—3.1 X 0.8-1
cm, caudate, the lowermost almost the same size as
the pedicellate ovary, decreasing in size toward the
apex of the inflorescence, green. Flowers (1)2 to
5(7), spreading, superimposed, large for the genus,
green-white; pedicellate ovary, 2.5—3.5 cm long.
slightly arched, green; sepals mucronate, light
green with the margins whitened, the veins slightly
marked in darker green; dorsal sepal concave,
when flattened somewhat elliptic, 12—16 X 8—13
mm; lateral sepals obliquely oblong, 16—20 X 4.5—
6.5 mm. acute, deflexed in fully open flowers; pet¬
als bipartite, base white or creamy white, green to¬
ward the apex; posterior segment linear-falcate, 13—
15 X 2—3 mm, acute, base free, toward the apex
Volume 13, Number 4
2003
Batista et al.
Habenaria pabstii (Orchidaceae)
399
t igure 2. Habenaria pabstii Batista & Bianchetti. —A. Flower, % view. —B. Column, side view, with the pollinaria removed; An, anther; Ac, anther canal; Au, auricle; Cn,
connective, Ro, rostellum; Ra, rostellum arm; St, stigmas; Sp. spur; Ov, ovary. —C. Rostellum, side view. Scale bars = 1 cm (A) and 2 mm (B, C). Photographed from living specimens
from the same collection as Batista 185.
400
No von
connivent with the dorsal sepal; anterior segment
linear-filiform, 25—13 mm long, 2—2.5(2.8) times
longer than the posterior segment, projected for¬
ward and somewhat parallel to the lateral lip seg¬
ments; lip tripartite, projected forward, base white
or creamy while and light green toward the apex of
the segments; undivided base 2—3 X 2—3 mm; lat¬
eral segments linear-filiform, 32—51 mm long, 1.7—
2.3 times longer than the medium segment; medium
segment shorter and wider, linear-ligulate, 17—24
X 1.5(2) mm, the apex bent upward; spur pendent,
clavate, free from the bracts, slightly to strongly
arched forward, narrower at the base, progressively
thickened toward the apex, 3.3—3.8 cm long, base
1.5 mm wide, apex 2.5—3 mm wide, green; column
6—10 mm high; anthers ca. 4—6 mm tall, white-
yellow; anther canals 7-9 mm long, slender, erect,
somewhat perpendicular to the anthers; connective
emarginate, green; rostellum mid lobe prominent,
elongate, forming a tunnel-like structure projected
beyond the anthers, ea. 10—1 1 mm long including
the lateral arms, 6—7 mm tall at the apex, fleshy,
cuspidate, acute, light green, the base white, slight¬
ly bent, forming a discrete groove that extends
through the lateral arms and which supports the
anther canals; rostellum arms white, 3—4 mm long;
auricles erect, ea. 2.5—3 mm tall, fleshy, flattened,
verrucose toward the apex, white; stigmas 2, closely
parallel, 6—8 mm long including the base, the re¬
ceptive, exposed surface 3-4 mm long and 2-2.5
mm wide, projected forward, non-involute, upper
surface flatly convex, light green; pollinaria 2, yel¬
low, viscidium ea. 1 X 1 mm, spaced 3—5 mm
apart; caudicles elongate, 8—10 mm long, about 2—
3 mm from the apex, ea. 90° bent; pollinia 4, 3-4
X 1—1.3 mm, laterally flattened, perpendicular to
the caudicles main axis. Fruit not examined.
Etymology. The new species is named in honor
of Guido F. J. Pabst (1914—1980), the late well-
known Brazilian orchidologist, author of Orchida-
ceae Brasilienses, who greatly contributed to the
knowledge of the orchid flora of central Brazil and
who described some new species of Habenaria.
Distribution. Habenaria pabstii is an occasional
species, so far only known from the Distrito Federal
and the state of Goias. It should also be expected
in western Minas Gerais, which is floristically re¬
lated to this core cerrado region, which shares
many orchid species in common with the Distrito
Federal and Goias. and probably also in southern
Tocantins state, which is close to the known range
of the new species.
Habitat, ecology, and phenology. Habenaria
pabstii is more commonly found in seasonally wet
grassy fields, in either hydromorphic or sandy soils.
Occasionally, it can also be found growing under
drier conditions or, rarely, in permanently wet
ground with water at the surface. I.ike most Haben¬
aria species in central Brazil, flowering occurs dur¬
ing the rainy season, from December to late March,
extending from the peak to almost the end of the
rainy season. Plants frequently appear in great
numbers, with apparently only the older and more
developed individuals flowering among many still
growing younger plants. The flowers usually do not
last more than a few days in the field (pers. obs.).
Also, like many other Habenaria and terrestrial or¬
chids from open grasslands, the flowering of this
species is greatly enhanced by bushfires (Jones,
1993; Oliveira et ah, 1996). The greatest concen¬
tration of flowering plants is usually found in burnt
areas, though occasionally some plants can also be
found in flower in unburnt places (pers. obs.). The
pollination mechanism and pollinators of the spe¬
cies are unknown.
Habenaria pabstii was apparently first collected
by Ezechias P. Heringer in 1961 in the Distrito
Federal. Analysis of dried specimens collected at
that time and examined by Guido F. J. Pabst has
revealed that Pabst misidentified the species as H.
fastor Warming, //. candolleana Cogniaux, //. ur-
baniana Cogniaux, and //. vaupellii Reichenbach
fil. & Warming. These mistakes probably avoided
the identification of //. pabstii as a new species in
the past.
Among Brazilian Habenaria, II. pabstii is mostly
similar in general flower morphology and size to a
species group within section Macroceratitae, rep¬
resented by H. bractescens Findley. H. fastor Warm¬
ing, H. gourlieana Gillies ex Findley, H. johanenn-
sis Barbosa Rodrigues, //. longicauda Hooker, and
H. macronectar (Vellozo) Hoehne, hence prior Pabst
identifications as //. fastor and H. vaupellii (a syn¬
onym of H. johannensis). However, distinct from all
these species, H. pabstii has a shorter, forward-
arched free spur (Figs. I IF 2A) and shorter, non-
involute stigmas, which are closely parallel forming
a single receptive surface turned up and frontward
(Figs. ID, 2B). All these other species have pen¬
dent spurs, which are completely or partially en¬
closed by the bracts and which are longer than the
ovary and pedicel, ranging from 4 to more than 15
cm, and reaching up to 25 cm in H. longicauda
(Snuverink & Westra, 1983). Also, their stigmas are
always longer (the free part 6—9 mm long in other
species with similar flower size), separated anti in
contact only at the apex, the margins are involute,
that is, rolled inward spirally on each side (Steam,
1983), and the receptive surface at the apex is
Volume 13, Number 4
2003
Batista et al.
Habenaria pabstii (Orchidaceae)
401
mostly turned to the sides. Thus, though H. pabstii
undoubtedly shares some degree of similarity with
these species, it differs in some important details
of the column structure, and its exact infrageneric
classification is unclear. As current infrageneric
classification of Habenaria needs a radical reas¬
sessment (Pridgeon et al., 2001), further studies,
including a molecular analysis, will be necessary
to clarify the relationship between //. pabstii , the
species in section Macroceratitae, and other species
in the genus.
Among other species in section Macroceratitae,
Habenaria pabstii is also similar in general flower
morphology to //. quinqueseta (Michaux) A. Eaton.
However, this species has the leaves more patent,
ovate to lanceolate and broader (2-6 cm wide); pet¬
als and lip white; a longer, pendent spur (5-10 cm
long, reaching up to 18 cm in //. quinqueseta var.
macroceratitis (Willdenow) Luer); and the rostellum
mid lobe smaller, completely placed between the
anther sacs (Luer, 1972; Correll, 1978). Addition¬
ally. //. quinqueseta has a distinct distribution, oc¬
curring in southern North America, Central Amer¬
ica, the Antilles, and northern South America, and
has never been recorded and is not known to occur
in Brazil.
Beyond the species in section Macroceratitae,
the only other Brazilian species to which Habenaria
pabstii is similar and could be confused is II. ur-
baniana (sect. Nudae Cogniaux). Both species are
similar in the large (lowers anti long and filiform
lateral segments of the petals and lip. However //.
urbaniana is vegetatively taller (86-122 cm high),
has lanceolate leaves up to 2 cm wide; a longer
(12-19 cm) and many-flowered inflorescence ((6)12
to 25 flowers) with overall smaller flowers with pro¬
portionally less developed anterior petals (petals
anterior segment 19-33 mm long, 1.6-1.9 times
longer than the posterior segment); a shorter floral
spur (ea. 2—2.5 cm), which at most equals the ovary
plus its pedicel in length, and prominently broad¬
ened at the apex (I mm wide at the base, 3-4 mm
wide in the apex). Additionally, the column struc¬
ture of both species is very' different, with the mid¬
dle lobe of the rostellum of H. urbaniana smaller
(about 1.5 mm long and 2 mm tall), triangular, and
completely enclosed by the anthers. Lastly, plants
of H. pabstii became black when dried or preserved
in ethanol, while those of H. urbaniana and the
species in section Macroceratitae became brown.
I he identification as //. candolleana is unwarranted
since this species has much smaller flowers (dorsal
sepal and lip lateral segments 6-8 mm long), and
a very' distinct column structure.
In column structure, mainly in relation to the
rostellum morphology, Habenaria pabstii is very
similar to H. johannensis. In both species, the ros¬
tellum mid lobe forms a prominent tunnel-like
structure, which is partially erect and projected be¬
yond the anthers (Fig. 2B. 2C). Despite this mor¬
phological similarity, it is not known, and would be
interesting to investigate, whether both species
share any similarities in their mechanisms of pol¬
lination.
Paratypes. BRAZIL. Distrito Federal: Gama, Saia
Velfia. 22 Mar. 1961, Heringer 8123 (AMES, HB, NY);
Sobradinho, Paranoazinho stream, 9 Jan. 1966, Irwin et
al. I ISOS (HB, NY, UB); Nueleo Bandeirante, Vargem
Bonita, 5 Eel). 1966, Irwin et al. 12301 (HB, NY); Nueleo
Bandeirante, Mansoes Setor Bark Way. 14 Jan. 1996, Ba¬
tista & Oliveira 585 (CEN); Brasilia, University of Brasilia
farm, 5 Jan. 1989. Bianchetti & Salles 743 (CEN): Bra¬
silia, Setor de Mansoes do Lago Norte, 14 Jan. 1992, Ba¬
tista 231 (CEN); Brasilia, near Colorado gas station. 10
Eeh. 1991, Batista 185 (CEN); Brasilia, Lago Sul, (JI-17,
16 Eeh. 1995, Oliveira 93 (UB). Goias: Mun. Alto Par-
aiso, Chapada dos Veadeiros, 9 Eeh. 1966, Irwin et al.
12382 (HB, NY. UB); 20 km N of Alto Paraiso, 1250 m,
19 Mar. 1971, Irwin et al. 32181 (HB. NY, UB); GO-118,
ea. 26 km N of Alto Paraiso. 13°58’S, 47°27'W. 8 Feb.
1987, Pirani et al. 1889 (SPE); Alto Paraiso/Colinas do
Sul, ea. 3 km of GO-118, 1200 m, 14°09'22"S,
47°32'43"W, 27 Jan. 1997, Walter et al. 3636 (CEN); Cor¬
umba de Goias, 14 Jan. 1968, Irwin et al. 18598 (HB,
NY, UB); Caldas Novas, 12 Jan. 1981, Menezes in CFCR
642 (SP. SPE); Pirenopolis, Santuario Eoologieo do Morro
do Cabeludo, 22 Jan. 1995, Batista 504 (CEN); Minayu.
29 Jan. 1997, Ass is et al. 404 (CEN, SPE); Mun. Caval-
cante, ea. 17 km SO of Cavaleante, 8 Jan. 2001, Batista
& Pansarin 1137 ( CEN. UEC).
Acknowledgments. We are grateful to Carolyn
E. B. Proenga and Fabio de Barros for helpful com¬
ments and revision of the manuscript and Tarciso
Filgueiras for the Latin diagnosis. We also thank
the curators of AMES, HB, HEPH, MBM. NY. HB.
SP. SPE, UB. and UFG for loans or access to their
collections.
Literature Cited
Batista, J. A. N. & L. B. Bianchetti. 2003. Lista atualizada
das Orchidaceae do Distrito Federal, Brasil. Acta Bot.
Bras. 17(2): 183-201.
Cogniaux, A. 1893. Habenaria. Pp. 18—102 in C. E. P.
Martius, A. G. Eichler & I. Urban (editors). Flora Bras-
iliensis, Vol. 3, Part 4. B. Oldenbourg, Munich.
Correll, I). S. 1978. Native Orchids of Nortli America
North of Mexico. [Reprint of the 1950 ed.) Stanford
Univ. Press, Stanford, California.
Hoehne, E. C. 1940. Habenaria. Pp. 52-254 in E. C.
Hoehne (editor). Flora Brasilica, Vol. 12, Part. 1. Se-
cretaria da Agriculture, Industria e Comercio de Sao
Paulo, Sao Paulo.
Jones, I). L. 1993. Native orchids of Australia. Reed
books, Australia.
Luer, C. A. 1972. The Native Orchids of Florida. New
York Botanical Garden, the Bronx.
402
Novon
Oliveira, B. S., J. A. N. Batista, C. E. B. Proen$a & L. B.
Bianchetti. 1996. InHuencia do fogo na floragao de es-
pecies de Orchidaceae em eerrado. Pp. 61—67 in Anais
do Simpdsio Impaeto das Queimadas sobre os Eoossis-
temas e Mudan$as Globais. 3" Congresso de Eeologia
do Brasil, Brasilia-DF.
Pabst, G. F. J. & F. Dungs. 1975. Orchidaceae Brasilien-
ses, Vol. 1. Briioke-Verlag Kurt Schmersow, Hildes-
heim.
- & -. 1977. Orchidaceae Brasilienses, Vol.
2. Briicke-Verlag Kurt Schmersow, Hildesheim.
Pridgeon, A. M.. P. J. Cribb, M. W. Chase & F. N. Bas-
mussen. 2001. Genera Orchidacearum, Vol. 2, Orchi-
doideae, Part 1. Oxford Univ. Press, New York.
Batter. J. A., J. F. Bibeiro & S. Bridgewater. 1997. The
Brazilian eerrado vegetation and threats to its biodiver¬
sity. Ann. Bot. 80: 223—230.
Snuverink, J. H. & L. Y. Th. Westra. 1983. Studies on the
flora of the Guianas. 3: A survey of Habenaria Willd.
in Suriname (Orchidaceae). Proceedings of the Konink-
lijke Nederlandse Akademie van Wetenchappen, Ser. C,
Bot. 86(4): 567-594.
Stearn, W. T. 1983. Botanical Latin, 3rd rev. ed. David
and Charles Publishers, Newton Abbot.
Felicia fischeri (Asteraceae) Refound in Northwest Tanzania
Henk Beentje
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England, h.heentje@kew.org
Abstract. Felicia fischeri from Tanzania has
been treated as a probable synonym of F. welwit-
schii for the last 50 years; the syntypes had been
lost. Recent collections from northern Tanzania
show that it is a species in its own right.
Key words: Asteraceae, Compositae, Felicia,
Tanzania.
Hoffmann (1895) described Felicia fischeri 0.
Hoffmann (Compositae: Astereae) from two speci¬
mens collected by Fischer in Kagehi. Mwanza Dis¬
trict, northern Tanzania, in the early 1880s—the
dates are not clear. The syntypes were lost when
the Berlin herbarium was bombed in the Second
World War. Grau (1973) decided that F. fischeri was
probably synonymous with F welwitschii (Hiern) J.
Grau, a species otherwise known from the far south
of Tanzania, southern Congo (Kinshasa), and south
to Angola and Zimbabwe. The epithet welwitschii
is of a slightly later date (1898), but as the types
of F fischeri were lost, Grau treated the earlier F
fischeri as a doubtful synonym of F. welwitschii. In
the Flora of Tropical East Africa (Beentje, 2002), I
agreed with Grau.
Over a century after the original collection, the
Tanzanian collectors Festo, Bayona, and Wilbard
collected a small Composite in the Minziro Forest
Reserve in northwest Tanzania:
TANZANIA. Kagera, Bukoba Rural District, Minziro
Forest Reserve, Kakindo Village, SW of forest, altitude
1150 m, rare at forest edge, 01°07'39"S, 31°30'00"E, 1
July 2001, L. Festo, W. Bayona & lb Wilbard 1572 (NUT,
K!, MO); Minziro Forest Reserve, Kalagara-Kayunga Sub¬
village, near top of hill, altitude 1207 in, occasional in
open grassland, 01°03'49"S, 31°32'48"E, 20 July 2001.
/.. Festo. W. Bayona & It. Wilbard 1659 (NUT, K!, MO).
The description of these two plants is as follows:
Herb with annual shoots 40-60 cm high from a
perennial root; stem unbranched except for the in¬
florescence, pale green, pilose to thinly pubescent.
Most leaves in a sub-rosette near ground level, ob¬
long, 5—11 X 0.4—0.7 cm, base slightly narrowed,
margin entire and minutely ciliate to hispidulous,
apex obtuse, with sparse scattered tubercle-based
hairs; cauline leaves sparse, similar to basal ones
but rapidly decreasing in size. Capitula 30 or more
in laxly branched bracteate cymes (the proximal
part of which is probably a leafy branching), radi¬
ate; stalks of individual capitula 6-24 mm long,
bracteolate, pubescent; involucre campanulate,
3.5—4 mm long, ± 5 mm diam.; phyllaries 2-seri-
ate, lanceolate, green with pale scarious margin and
apex, 2.5—3.8 X 0.4—0.6 mm, with a few scattered
hairs. Ray florets 1-seriate, 17-19, whitish, tube
cylindrical, ± 2 mm long, glabrous, limb ± 3.7 X
1.4 mm, style 2.5 mm long. Disc florets many, tube
ca. 2.3 mm long, lobes ca. 0.8 mm long, glabrous.
Achenes compressed, brown with 2 paler margins,
1.2-1.3 mm long, pilose; pappus of ± 15 barbellate
bristles 2—2.3 mm long.
Hoffmann’s original description of Felicia fischeri
is: “herbacea ramis hispidulis; foliis alternis sparsis
linearibus sessilibus obtusis integerrimis hispidis;
capitulis parvulis laxe paniculatis pedicellatis; in-
volucri bracteis pauciseriatis subaequilongis oblan-
ceolatis acutis; ligulis oblongis, achaeniis compres-
sis hirtis; pappi seris 1-seriatis tenuibus ionge
ciliatis.—Untere B. 3 cm long, 2 mm breit.—17
(Kagehi— Fischer n. 371, 336)—Gebirgssteppe.”
Hoffmann did not mention the basal rosette, but
it is possible that this part was missing from Fi¬
schers specimens; many collectors do not collect
the basal parts of Compositae (luckily for us, Festo
and his colleagues did!). Otherwise the description,
while being rather general, agrees neatly with the
recent collections. Kagehi is in Mwanza District,
just south of Lake Victoria and not very far from
the Minziro Reserve (about 220 km as the crow
flies).
The new collections of Felicia fischeri differ from
F. welwitschii sensu stricto in the rosette leaves (F.
welwitschii has leafy stems with proximal leaves op¬
posite), the phyllaries (2-seriate, not 4-seriate), the
number of ray florets (17 to 19, not 25), the smaller
disc florets (tube ca. 2.3 mm, not 3.3-4.5 mm long),
and the fewer and shorter (2—2.3 mm, not 3—4 mm
long) pappus bristles.
I believe these new collections are the taxon that
Fischer collected all those years ago, and the recent
collections resolve a mystery: Felicia fischeri is dis¬
tinct from F. welwitschii, it is a species in its own
right, and a rare endemic restricted to northwest
Novon 13: 403-404. 2003.
404
Novon
Tanzania and probably the adjacent part of Uganda.
I hereby designate tin* neotype:
Felicia fischeri (). Hoffmann, in Engler, die Pflan-
zenwelt Ost-Afrikas und der Nachbargebiete
C: 407. 1895. TYPE: Tanzania. Mvvanza Dis¬
trict: kagchi, Fischer 336, Fischer 371 (syn-
types. I? destroyed); Tanzania. Bukoba District:
Minziro Forest Reserve, Kalagara-Kayunga
Subvillage, 01°03'49"S, 31°32'48''E, 20 July
2001, L. Festo, W. Bayona & W. Wilbard 1659
(neotype, here designated, k). [Duplicates of
the neotype are at NHT and MO but have no
official status, according to the Code (Greuter
et al., 2000); nevertheless, 1 would call them
iso-neotypes. It would have been more appro¬
priate if the neotype had been the top speci¬
men. deposited at the National Herbarium of
Tanzania, rather than the third set deposited
at kew, but the kew sheet was the specimen
studied.)
The conservation status of Felicia fischeri should
probably be Data Deficient (DD); the habitat, open
grassland, is probably still common, but the species
seems to be rare within this habitat. I have no in¬
formation on threats to the vegetation of this area.
Acknowledgments. I thank John Pruski, Victo¬
ria Hollowed, and Roy Gereau for their suggestions
to improve this paper.
Literature Cited
Beentje, II. J. 2002. Compositae Part 2. Flora of Tropical
Fast Africa. Balkema, Rotterdam.
Grau, J. 1973. Revision der Gattung Felicia (Asteraceae).
Mitt. Rot. Staatssamml. Miinchen 9: 195—705.
Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). H. Nicolson, P. C. Silva,
J. E. Skog. 1*. Trehane, N. J. Turland & 1). L. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Hoffmann, (). 1895. Compositae. Pp. 402-A-22 in A. Fn-
gler. Die Pflanzenwelt Ost-Afrikas und der Nachbarge¬
biete C. Geographische Verlagshandlung Dietrich Rei-
mer, Berlin.
Nomenclatural Novelties in the Orchidaceae from
Venezuela and Guyana
German Carnevali Ferndndez-Concha 1 and Ivon M. Ramirez-Morillo
Herbario CICY, Centro de Investigation Cientffica de Yucatan, A.C., A.P. 87, Cordemex,
Merida 97310, Yucatan, Mexico, carneval@cicy.inx, ramirez@cicy.mx
ABSTRACT. The following represent new specific
taxa in the Orchidaceae from Venezuela: Epiden-
drum cesar-fernandezii Carnevali & I. Ramirez, E.
reclinatum Carnevali & I. Ramirez. Pleurothallis lo-
reae Carnevali & 1. Ramirez, P. escalerensis Car¬
nevali & C. A. Luer, and P. stellata I. Ramirez &
Carnevali. A new nothospecies, Epidendrum
Xgransabanensis Carnevali & 1. Ramirez, is pro¬
posed as the product of the introgression between
Epidendrum ibaguense HBK and E. secundum Jac-
quin. A new subspecies, Liparis nervosa suhsp.
granitica Carnevali & I. Ramirez, is proposed to
accommodate populations of Liparis nervosa (Thun-
berg) Lindley restricted to granitic outcrops in the
northwestern Guayana in Venezuela. Descriptions,
illustrations, and discussions of affinities are pro¬
vided for the newly proposed taxa.
Key words: Epidendrum, Guyana, Liparis, Or¬
chidaceae, Pleurothallis, Venezuela.
Field and herbarium studies of Venezuelan Or¬
chidaceae relevant to several floristic projects, pri¬
marily the treatment for the Flora of the Venezuelan
Guayana project (Carnevali et al.. 2003), have un¬
covered several new taxa. These orchid taxa are
herein proposed as new.
SUBTHIHK LaKUINAE
Kpidcndriim cesar-fernandezii Carnevali & I.
Ramirez, sp. nov. TYPE: Venezuela. Tachira:
along road between Rubio and Delicias, 25 km
above Rubio, $V of Rubio, 23 Aug. 1976, T.
Croat & G. Punting ,38489 (holotype, VEN; iso¬
type, MO, isotype fragment, CICY). Figure 1.
Species haec Epidendro goodspeedia.no A. I). Hawkes
similis sed floribus multiminoribus; E. agoyanense et E.
aguaricoense affinis sed ovario sepaloque verruculosis dif-
fert: E. densifloro affinis sed labello latiore, petalis pen-
dulis recedit.
Epiphytic herbs, to 2 m tall; roots unknown;
stems simple, multifoliate, slightly compressed, to
15 mm thick near apex; apical leaves 12—17 cm
long, 15-25 mm wide, elliptic, acute, acuminate,
margin entire; sheaths 5—6 cm long, tubular,
smooth; inflorescence 35—55 cm long, terminal on
the mature stem without a spathe, flowering once,
paniculate, the primary branches pinnate again in
the lower !4 of their length; peduncle 2^4 cm long,
subterete or slightly compressed, surface smooth in
the lower Vi, becoming laxly verruculose above; pri¬
mary branches 25—30 cm long, secondary branches
3—5.5 cm long bearing 5 to 8 flowers; bracts of the
peduncle 8—1 I cm long, ca. 1 cm wide at base,
very narrowly triangular and acuminate, bracts of
the primary branches similar to those of peduncle
hut much smaller, 3—5 cm long; rachis on the
branches subterete and slightly angulate, laxly ver¬
ruculose; floral bracts 3.8—4.1 mm long, ca. I mm
wide, narrowly triangular; flowers resupinate, rela¬
tively small for the group; sepals green, suffused
with honey brown on the apical half, petals trans¬
lucent green, the labellum white, suffused with
pink and some areas with a green hue around the
edges of the callosities, column pale pink on the
apical half, the basal half dull dark green with dull
maroon tinges dorsally and where it joins the lateral
sepals; sepals widely spreading, fleshy, slightly con¬
cave along the midnerve on the apical half, dorsally
conspicuous and coarsely verruculose; dorsal sepal
11-12 X 2.8—3.1 mm, elliptic spathulate, obtusely
acute, 5-nerved; lateral sepals 9— ] 1 X 3.5—4 mm,
elliptic spathulate, obtusely acute, 6-nerved, sub-
apically with fleshy, conical, acute mucro 1 mm
long; petals 9.8-10.2 mm long, 0.9 mm wide at
apex, 0.3 mm wide at the base, thin, flexed down¬
ward and pendent behind the lateral sepals, I-
nerved, filiform-ohlanceolate, rounded to broadly
obtuse at apex, margin entire; labellum 5.9—6.1 X
8.9—9.2 mm across the forcefully spread lateral
lobes, convex in natural position, fused up to the
apex of the column, the margins finely fimbriate, in
general outline transversely oblong elliptic, apically
conspicuously but shallowly emarginate, the disk
' Author to whom reprint requests should be addressed.
Novon 13: 405-418. 2003.
406
Novon
shape. (Based on isotype, T. Croat & G. Hunting .'18489, MO.) Drawing hy Carlos Yam.
Volume 13, Number 4
2003
Carnevali & Ramirez
Novelties in Orchidaceae
407
4 cm
Figure 2. —A. Epidendrum ibaguen.se (based on Carnevali el al. 3241, VEN; 70% EtOH CICY). —B. Epidendrum
Xgransabanensis Carnevali & I. Ramirez (based on Carnevali et al. 3239, VEN; 70 % EtOH CICY). —C. Epidendrum
secundum (based on Carnevali et al. 3240. VEN; 70% EtOH CICY). Drawings by Bruno Manara.
bicallose, with 3 prominent blunt keels that run up
to the apex of the labellum, the one in the middle
thicker and higher; the whole blade 4-lobed, the
basal lobes 4.4-4.6 mm long, reniform, basally cor¬
date, the apical lobes ca. 3.5 mm long. ca. 2 mm
wide, transversely reniform or subquadrate, the api¬
ces diverging, the basal margins separated from the
basal lobes by very narrow sinuses or fused to them;
column 7.5—8 mm long including the clinandrium,
ca. 3.5 mm thick at apex; clinandrium obsolete,
margin entire; anther ca. 2 mm long, ovoid-sub-
spheric, 4-locular, pollinia 4 of the “bird-wing”
type, strongly compressed laterally, the inner ones
somewhat smaller and partially fused on their ven¬
tral surface, caudicles smooth, soft, viscidium
transparent; rostellum subapical, cleft, nectary
shallow.
Etymology. The specific epithet refers to Cesar
Fernandez, a collector of the paratype of the spe¬
cies and an active researcher of the Venezuelan
Andean orchid flora. The hyphen in the epithet is
to be maintained, according to Article 60.9 of the
ICBN (Greuter et al., 2000).
This new species is similar to Epidendrum den-
siflorum Hooker, but the labellum is proportionally
wider, the margins are more erose, anti the ovary'
and outer faces of the sepals are more verruculose.
The petals are pendulous in the new species where¬
as they are spreading in E. densifiorum. The large
plants arul the general structure of inflorescences
and flowers suggest an affinity with the Ecuadorian
Epidendrum agoyanen.se Hagsater & Dodson and E.
aguaricoen.se Hagsater & Dodson, but these spe¬
cies, also from the eastern Amazonian slopes of the
Andes, feature labella with entire margins and
much more divaricate apical lobes. Epidendrum
goodspeedianum, from the western Andes, is also
similar but it is a much larger plant anti has larger
flowers than E. cesar-fernandezii. with the labellum
more than 2.5 cm wide and pedicels of up to 6 cm
long.
Paratype. VENEZUELA. Tachira: Mun. Junfn, El
Playbn, 1115 m, 3 Aug. 2(XH, C. Fernandez 0341 (VEN).
Epidendrum Xgransabanensis Carnevali & I.
Ramirez, nothospecies nov. TYPE: Venezuela.
Bolivar: road El Dorado—Santa Elena tie Uair-
en, km 138 S of El Dorado, Canaima National
Park, NE Gran Sabana general area, approx.
6°19'N, 61°25'W, shrublands over sandy soils
along the road, in a mixed mosaic along with
gallery forests, sandstone outcrops, and boggy
associations, 1500 m, 3 Aug. 1993, G. Car¬
nevali, E. Lorea, /V. Warford , B. Whitlock , K.
Rivero. W. Diaz & C. Fleming 3238 (holotype.
VEN; isotype, CICY [70% EtOH]). Figure 2.
Hybrid formula: Epidendrum ibaguen.se HBK X
Epidendrum secundum Jacquin.
Nothospecies intermedia quam Epidendrum ibaguense
et E. secundum ; haec Epidendro ibaguensi similis sed flo-
ribus non resupinatis, sepalis petalisque proportione lat-
iore brevioribus, callo crassiore, columna breviore recedit.
Terrestrial herbs ; growing over sandy soils, sun-
loving; stems, leaves, and inflorescences indistin¬
guishable from those of Epidendrum ibaguense,
flowers non-resupinate or rarely resupinate, petals
408
Novon
and sepals widely spreading, purple, lal>el 1 urn pur¬
ple with a yellow callus; dorsal sepal I 1.5—12.5 X
3.1—3.5 mm, narrowly elliptic, acute; lateral sepals
12-13 X 3.7-4 mm, elliptic, oblique, acute; petals
10-11 X 2.3—2.8 mm, narrowly elliptic, acute; la-
bellurn cordiform in general outline, with the mar¬
gins irregularly dentate to erose or even laciniate,
flat, 7.5—8 mm long from the apex of the column,
3-lobed, 13.5-14 mm wide across the apices of the
lateral lobes; central lobe 3.7—4 X 8.5—8.7 mm,
broadly obovate, apically emarginate; lateral lobes
reniform, 0.5-0.66 mm long and wide, (labellate
obovate; disk with a pair of laminar, obovate calli
at each side of the apex of the column and that can
be interpreted as being a continuation of it. and a
laminar keel running down the center; column
straight, 0.55—0.65 mm long.
This new entity is almost certainly the result of
the introgression of Epidendrum ibaguense and E.
secundum. Both taxa grow on rocky, exposed places,
in a landscape of a mixed mosaic of savanna with
gallery forests over sandy, acidic soils along the
road from El Dorado to Santa Elena de Dairen, the
former at low elevations (800—1000 m), the latter
at relatively higher elevations (1000—1300 m).
Mixed populations of both parental taxa and the
hybrid taxon are common at intermediate altitudes.
Epidendrum Xgransabanensis was treated as Epi¬
dendrum ibaguense X Epidendrum secundum in
Camevali and Ramirez (in Camevali et al.. 2003;
338).
Epidendrum ibaguense and related taxa, e.g., E.
splendens Schlechter, E. radicans Lindley, E. ma-
crocarpum Richard, E. calanthum Reichenbach f.
& Warscewicz, and others, are characterized by re-
supinate flowers and a simple callus consisting of
two laminar projections at each side of the apex of
the column often joined with two small, globular or
obovate-laminar, keel-like calli at the base of the
disk, and a central laminar keel running down the
center, rarely to the apex of the emargination of the
central lobe, but more often ending midway along
the isthmus of the central lobe. This thin laminar
keel frequently curves or waves one way or the oth¬
er. The column is usually relatively long, as long
as or longer than the blade of the labellum. The
perianth segments are of a relatively soft texture. A
few taxa of this complex, e.g., E. mimeticum Car-
nevali & G. Romero. E. flexuosum G. Meyer, and
clones of E. radicans, consistently feature non-re-
supinate flowers, but are otherwise clearly members
of this grouping when other morphological traits
(e.g., callus, labellum. and perianth segment shape)
are taken into account.
On the other hand, members of the extremely
difficult Epidendrum secundum complex, e.g.. E.
catillus Reichenbach f. & Warscewicz, E. elonga-
tum Jacquin, E. jamesonii Lindley, E. tricallosum
Schlechter, etc., feature non-resupinate flowers and
a complex callus composed of several to many tu¬
bercles and thick, intertwined keels, and a rela¬
tively short column, usually shorter than the label¬
lum blade and much shorter than the petals and
sepals. Further, the lateral lobes of the labellum
appear to originate from the sides of the apex of
the column, not from an independent, free, laminar
blade, as is the case in the labella found in the
members of the E. ibaguense complex. Good illus¬
trations of members of this complex (probably sev¬
eral species on the same plate) are to be found in
Dunsterville and Garay (1979: 224, plate 187) as
E. elongation. Good color photographs of members
of the E. ibaguense and the E. secundum complexes
of the Guayana are found in Couret (1982) and
Romero (1997). In this paper, we follow the ap¬
proach taken by Camevali and Ramirez (in Car-
nevali et al.. 2003: 342-343) in treating most mem¬
bers of this complex as a single entity. Epidendrum
secundum. The group of taxa around E. secundum
requires an urgent revision to ascertain their
boundaries and distributions. In the meantime, it is
safest to treat them as a single, widely ranging, and
variable species.
Within the mixed population in the general Gran
Sabana area in Venezuela mentioned above, colo¬
nies of plants that are intermediate between species
of the two complexes are commonly found. On oc¬
casion. isolated plants featuring intermediate com¬
binations of characters are also encountered in a
landscape of a mixed mosaic of savanna with gal¬
lery forests. These putative hybrid plants are char¬
acterized by non-resupinate flowers (reminiscent of
E. secundum parentage) and a simplified callus
(from the E. ibaguense parentage) consisting of two
thick lateral calli following the lateral, apical edges
of the column, and a laminar keel running down
the center. This laminar keel is thicker and pro¬
portionally shorter than those found in the members
of the Epidendrum ibaguense complex.
The two specimens cited in this protologue rep¬
resent two extremes in labellum shape variation
within the nothospecies, though both display non-
resupinate flowers. The holotype corresponds to an
individual showing an E. ibaguense- type labellum,
which is, however, somewhat broader and shorter.
On the other hand, Figure 2B depicts a second
plant of this grex (G. Camevali et al. 3239), which
features a more E. secundum- like labellum, but
w ith simplified calli. Both were collected along the
Volume 13, Number 4
2003
Carnevali & Ramirez
Novelties in Orchidaceae
409
El Dorado—Santa Elena de Uairen road growing
within a perimeter of a couple of meters and along
with many other plants showing various combina¬
tions of parental characters. Populations of both E.
ibaguense and E. secundum, both in pure stands or
in mixed communities, were found nearby. We de¬
cided to name these apparent hybrids since they
are a common sight along the road and are becom¬
ing increasingly common in both herbarium and
live collections. The introgression between the two
species in the general Gran Sabana area is appar¬
ently associated with disturbed, anthropogenic con¬
ditions. This situation has probably created novel
microhabitats for the survival of the hybrids, or
more likely, has somehow allowed for the break¬
down of former pollination barriers between mem¬
bers of the E. ibaguense and E. secundum popula¬
tions. A more thorough sampling program is
required to ascertain whether the introgression pro¬
cess is restricted to disturbed areas close to roads
or human settlements, or if it occurs elsewhere, un¬
der more pristine conditions.
An additional taxon, recently described as a spe¬
cies, but not mentioned as a nothospeeies, is prob¬
ably also the result of the introgression of members
of the Epidendrum ibaguense and E. secundum
complexes. This is Epidendrum thermophilum Hag-
sater & Dodson (Hagsater & Dodson, 1993) from
northwestern Ecuador. This entity features non-re-
supinate, small red-purple flowers with the overall
morphology of an intermediate between the two
complexes. The labellum is shallowly 3-lobed with
the simpler callus of the E. ibaguense complex. Ep¬
idendrum calanthum (a member of the E. ibaguense
complex with small, pink, soft-textured flowers with
an almost simple labellum) grows nearby, and some
members of the E. secundum complex must occur
in the immediate neighborhood, since populations
of these species (such as E. jamiesonis Reichen-
bach f.) are locally common in open places in the
Andean slopes, e.g., D. Rubio & C. Quelal 692,
MO). It would appear that non-resupination is a
dominant feature in hybrids involving members of
both complexes.
Paratype. VENEZUELA. Iiolivar: locality same as
the holotype, G. Carnevali el al. 3229 (V EN, CICY
[Eton D.
Epidendrum reclinatum Carnevali & I. Ramirez,
sp. nov. TYPE: Guyana. Mazaruni-Potaro:
Guyana—Venezuela border area, Roraima sum¬
mit. la Proa Camp, E of border, near Lake
Gladys, mostly open rocky areas with ravines
and lake, 5°15'36"N, 60°13'00"W, 2800 m, 14
Apr. 1988. R. Liesner 23315 (holotype, VEN;
isotypes, CICY, MO). Figures 3, 4.
Figure 3. Epidendrum reclinatum Carnevali & I. Rami¬
rez. Habit (based on F. Delascio-Chitty 13972, VEN; dig¬
ital scan by German Carnevali).
Species haee Epidendro montigeno similis sod plantis
prostratis, bracteis majoribus, floribus albidis vel flavidis,
petalis proportione latioribus recedit.
Terrestrial herbs, prostrate, sprawling and creep¬
ing, growing on grassy or boggy areas, drying dark
brown to black with an oily sheen; roots carnose,
1—2 mm thick; stems (7—)10—20(—35) cm long, 0.4—
0.6 mm thick, totally clothed by sheaths; rooting at
the base and above as it sprawls over the substrate;
foliose from the base when young, eventually leaves
restricted to the upper half or third of the stem,
simple at first, then branching after flowering;
branches occurring at several levels on the stems,
often about mid-length of the flowering stem, 7—13
cm long; leaves 3.5—5 cm long, 8—1 I mm wide on
flowering stems, smaller on young stems or branch¬
es, narrowly ovate, acute to subacuminate, margins
revolute, minutely erose toward the apex, entire
410
No von
I cm
figure 4. Epidendrum reclinatum Carnevali & i. Rami¬
rez. Floral parts flattened (based on holotvpe. R. Liesner
23315, YEN). Drawing by Rossana Marrufo.
elsewhere; blades articulate with their sheaths, con¬
spicuously striate and minutely cross-veined in dry
material, all in one plane, sheaths 10—20 mm long,
tubular, scarious, rugulose; inflorescences terminal
from the main stems or the branches, racemose, 3—
7 cm long, 6- to 8-flowered, lacking a spathe; pe¬
duncle 2.5—3 cm long; rachis 3—4 cm long, erect
or apparently nutant; floral bract 6-9 X 3—5 mm,
surpassing the pedicellate ovary at anthesis, cu-
cullate, fleshy; flowers white or yellow-tinged,
fleshy, apparently non-resupinate; pedicellate ovary
4—5 mm long; sepals concave, fleshy, dorsal sepal
7—8 X 2.5—2.7 mm, elliptic-oblong, obtuse, dorsal-
ly keeled, 3- to 5-nerved; lateral sepals 7.9— 8 . 1 X
4-4.5 mm, broadly elliptic, oblique, obtusely acute
margins concave, erose particularly toward the
apex, 3- to 5-nerved; petals 6.5—8 X 2.1—2.6 mm,
narrowly spathulate, elliptic, apically broadly ob¬
tuse to subacute, margins erose, I - to 3-nerved;
labellum fleshier than other perianth segments, en¬
tire, deeply concave in natural position, fused to
the column to its apex, the free portion 8—9 mm
long from the insertion of the sepals and labellum,
6.5—8 mm wide when forcefully spread, transverse¬
ly rhombic, the apex very fleshy, obtuse-acuminate,
1.2—1.5 mm long; column 3.5—4 mm long, ca. 2.5
mm thick, the clinandrium and anther smooth.
Epidendrum reclinatum is one of several taxa in
the Epidendrum dendrobioides Thunberg complex,
a group ol taxa that has undergone extensive di¬
versification in the Guayana Highlands, although
several species of the group grow elsewhere. The
complex is characterized by its mainly terrestrial or
lithophytic habit (often at high elevations), usually
on exposed, rocky or boggy conditions, the stems
usually branching about mid-length, the fleshy
flowers that are often campanulate, the relatively
large, fleshy floral bracts, and the simple, deeply
concave, often unspreadably fleshy labellum.
Among other taxa in the complex, E. reclinatum is
readily recognized by the prostrate, sprawling stems
and short unbranched inflorescences. It is appar¬
ently most closely related to the sympatric or par-
apatric Epidendrum montigenum G. S. Ridley ex
Oliver, with which it shares the softly fleshy flowers,
the labellum being spreadable (though somewhat
forcefully) in herbarium material. It is, however,
easy to distinguish from that taxon by its white or
yellowish flowers (vs. pink), the prostrate habit, the
broader petals, and the more transverse labellum.
This species was treated in Carnevali and Ramirez
(in Carnevali et ah. 2003: 345) as Epidendrum
sp. A.
Several taxa in this complex are found sympat-
rically or parapatrically in the Roraima-Tepui mas¬
sif, where E. reclinatum seems apparently restrict¬
ed. Among these, Epidendrum alsum G. S. Ridley,
E. montigenum , and this new r taxon are endemic
there. The several taxa ol the Epidendrum dendro¬
bioides complex occurring in the Venezuelan Gua¬
yana can he distinguished with the use of the fol¬
lowing key.
Key to the Members of the Epidendrum dendrobioides
Complex of the Venezuelan Guayana
1. Inflorescences shorter than the leaves; labellum
longer than wide . E. imthurnii Ridley
IInflorescences longer than the leaves; labellum as
broad as or broader than long. 2
2(1). Flowers pale to deep pink; labellum fleshy co¬
riaceous, flexible and thus easily flattened,
thicker toward base. E. montigenum
2'. Flowers green, yellowish or cream-colored; la¬
bellum very fleshy (rarely fleshy coriaceous in
E. reclinatum ), rigid and impossible to flatten
without breaking it, thicker toward the apex . . 3
3(2). Plants prostrate, sprawling and creeping over
boggy substrate; plants apparently restricted to
Roraima tepui and surrounding massifs (llu Te-
pui, Kukenan Tepui). E. reclinatum
3'. Plants erect, rarely weakly prostrate over
shrubs on tepui shrublands; plants of many
places in the Guayana Highlands .
4(3). Floral bracts 9—13 mm wide (since they are
4
Volume 13, Number 4
2003
Carnevali & Ramirez
Novelties in Orchidaceae
411
conduplicate, the width of one half can be dou¬
bled for easy measurement) . 5
4'. Floral bracts 3—7(—8) mm long . 7
5(4). Floral bracts 3—4 times longer than the pedi¬
cellate ovary . E. urbanianum Cogniaux
5'. Floral bracts shorter, up to 2 times longer than
the pedicellate ovary. 6
6(5). Inflorescences strictly erect, unbranched; ra-
chis straight; leaves acute; floral bracts with
straight apices; dorsal sepal 11—12 mm long;
petals 10—12 mm long; plants from Cerro de La
Neblina.
. . . E. commelinispathum Carnevali & I. Ramirez
6'. Inflorescences arching, decurving or pendent,
often branching; rachis zigzag; leaves obtuse;
dorsal sepal 6—7 mm long; petals 4.5—5.5 mm
long; plants from Roraima and neighboring te-
puis. E. alsum G. S. Ridley
7(5). Dorsal sepal 5—7 mm long, 1.9—2.1 mm wide;
labellum 4—4.5 mm long, 6—7 mm w ide ....
. E. durum Findley
7'. Dorsal sepal 8—9 mm long, 4—4.5 mm wide;
labellum 5—5.5 mm long, 7.5—9 mm wide . . .
. E. ulei Schlechter
Paratypes. VENEZUELA. Bolivar: R oraima, NE
sector, Lago Gladys, 2700 m, 3—5 Apr. 1989. F. Delascio-
Chitty 13972 (VEN); llu-tepui. lower plateau. 2500 m, 16
Apr. 1988. . . sterile, decumbent spreading ... ,” R.
Liesner 23431 (CICY, MO, VEN); Kukenan Tepui summit,
17 Apr. 1988, R. Liesner 23452 (MO, VEN).
Tribe Malaxideae
Liparis nervosa (Thunberg) Lint I ley. Gen. Sp.
Orch. PI. 26: 1830. Ophrys nervosa Thunberg
ex Murray, Syst. Veg. (ed. 14): 814. 1784.
TYPE: Japan. Thunberg s.n. (UPS not seen).
Liparis nervosa is widespread in the Neotropics
and tropical Africa and Asia. For such a wide¬
spread taxon there seems to be little variation in
flower shape. However, there is much vegetative
variation, and some forms might merit subspecific
recognition. One such distinctive series of popula¬
tions is here treated at the subspecific level. An¬
other set of Guianas plants with different floral col¬
oration is discussed here in relation to the other
subspecific taxa considered.
Liparis nervosa subsp. granitica Carnevali & 1.
Ramirez, subsp. nov. TYPE: Venezuela. Boli¬
var: 1 km S of Quebrada La Elore, affluent of
Rio Ore, affluent of Rio Parguaza, 85 m, 9 Sep.
1985, Julian A. Steyermark, Bruce Holst &
Bruno Manara 131667 (holotype, VEN; iso¬
type, MO), figure 5.
Species haec Lipari nervosae subsp. nervosae affinis sed
plantis et floribus minoribus, foliis angustioribus, acutis,
habito graniticola recedit.
Lithophytic, growing on cracks of rocks on ver-
t-1
5 mm
Figure 5. Liparis nervosa subsp. granitica Carnevali &
I. Ramirez. Floral parts flattened (based on Julian A. Stey¬
ermark et al. 131667, VEN). Drawing by Rossana Ma-
rrufo.
tical granite walls. Pseudobulbs ovoid, 1—2 cm long,
entirely concealed in shredded-nerved formless
bracts; leaves 2(to 4) per pseudobulb, narrow to
rarely wide-elliptic to ovate-elliptic, acute, basally
attenuate into a pseudopetiole; laminae with 5 lon¬
gitudinal nerves, prominent, (3-)5(—8) cm long,
(1.6—)2.9(-3.9) cm wide, membranaceous, pale yel¬
low-green, usually erect. Inflorescence terminal,
erect to pendulous, apical, 8—14 cm long, a raceme,
usually 3-flowered to rarely 9-flowered; peduncle
pinkish (fide Steyermark et al. 131667). Flowers
small as compared to other members of the species,
yellow 7 (fide Steyermark et al. 131667 ), arranged
from the mid-point to the apex of the inflorescence.
Sepals and petals membranaceous, labellum curved
backward. Sepals oblong, 7 X 3 mm, with 5 nerves,
apically oblique, rounded, margins entire. Petals
narrowly elliptic, apically rounded, 9X2 mm, mar¬
gins entire. Labellum 2-lobed, in general shape ob-
ovate, strongly decurved, 5.1 mm long, 4.1 mm
wide, apically notched, with a small mucro, 5-
nerved, three central and two lateral from where
reticulate nerves expand to the margins, margins
crenulate, with a 1.5 mm claw 7 at the base of la-
bellum from which two small (0.5 mm long) apical
horn-like calli project. Column strongly incurved
412
Novon
apically, 4.5 mm long, with two small wings, 0.4
mm long.
Paratype. VENEZUELA. Amazonas: Man. Atures,
carretera Fto. Ayacucho—Samariapo, km 12.5, 100—150 m,
20 July 1993, A. Groper 1034 (MO, VEN).
This entity is known from few collections and
grows in seasonally dry areas in environments that
are usually much drier than those where the typical
subspecies for Liparis nervosa grows. It was treated
as Liparis nervosa subsp. A in Carnevali and Ra¬
mirez (in Carnevali el al., 2003: 413). Although
similar to the typical subspecies, the plants of the
new subspecies are smaller (see key below), and
the leaves are narrower ((1.6—)2.9(—3.9) cm vs. usu¬
ally more than 4 cm in the typical subspecies) and
dry a distinctive pale yellow-green or [tale yellow-
brown color, fhe inflorescence is also proportion¬
ally shorter, l’he llowers are somewhat smaller (dor¬
sal sepal up to 7 mm long as opposed to ca. 8 mm
in the typical subspecies) but otherwise identical
to those of the typical subspecies. More material
may indicate that this entity should be recognized
at the species level. I his new subspecies is locally
abundant on granitic outcrops in northern Amazon¬
as State and northwestern Bolivar Stale in Vene¬
zuela (possibly ranging into neighboring Colombia).
There it grows on shady, rocky vertical walls or
otherwise under the shade of shrubs. On the other
hand, the typical subspecies is usually found grow¬
ing on soil, often in boggy places or as subepiphytes
in extremely humid places and not in dry. rocky
places.
Liparis nervosa f. kappleri (Reichenbach f.)
Christenson & Carnevali, Lindleyana 11: 17.
1996. Sturmia kappleri Reichenbach f., Lin-
naea 22: 833. 1849. Leptorchis kappleri (Rei¬
chenbach f.) Kuntze, Revis. Gen. PI. 671.
1891. Liparis kappleri (Reichenbach f.) Rei¬
chenbach I., Ann. Bot. Syst., 6: 218. 1864.
TYPE: Surinam. Feb.—Apr. 1844. Kappler
1491 (MO).
This entity is rare and so far only known from
the type collection (Christenson, 1996). However,
since it is likely to appear elsewhere in the Guianas
and the Venezuelan Guayana, we have included it
in this account. It seems to be nothing more than
an anthocyanin-free form of this species (Christen¬
son, 1996). Liparis nervosa f. kappleri seems to be
a distinctive morph of the typical subspecies, and
should stand as a good taxon. However, more ma¬
terial may indicate closer affinities with subspecies
granitica than with the type of the species.
The following key allows the identification of the
infraspecific taxa of L. nervosa in the Guianas.
Kk\ to the Infrasi’kcific Taxa of Liparis \rrvosa
in the Guianas
la. Leaves (3—)5(—8) cm long, (1.6—)2.9(—3.9) cm
wide, usually narrowly elliptic, membranous,
pale yellow-green upon drying; inflorescences 8-
14 cm long. L nervosa subsp. granitica
lh. Leaves usually longer than 8 cm in mature
plants, often 10—30 cm long, usually broader
than 4 cm, often to 10 cm wide; usually elliptic,
of a firmer texture, drying greenish or brownish;
inflorescences (7—) 1 0— 30( —50) cm long; leaves
drying several shades of brown.
. L. nervosa subsp. nervosa
2a. Flowers tinged with purple or brown ....
. /,. nervosa f. nervosa
2b. Flowers entirely greenish yellow .
. L. nervosa f. kappleri
Sub rm hk Pi.kukothallidinak
Pleiirothullis loreae Carnevali & I. Ramirez, sp.
nov. TYPE: Venezuela. Falcon: Sierra de San
Luis, Zambrano—Curimagua road, cloud forests
over karst, 1300-1500 m, 9 Aug. 1993, G.
Carnevali , F. Lorea, N. Warford, II. Whitlock &
F. Morillo 3263 (holotype, VEIN; CICY [EtOH,
flower)). Figure 6.
Species haec Pleurothalli chamae affinis, sed plantis
floribusque minoribus, sepalis dorsalibus 3-nervis, labello
profunde concavo abhorret; affinis P. cordatae sed sepalo
dorsali proportione angustiore, labello concavo, petalis
earnosioribus glabris reeedit.
Lithophytic, sun-loving, cespitose herbs ; rhizome
abbreviated; stems erect, straight, terete, 1 1-19 cm
long, with two sheaths shorter than internodes, the
shorter one basal, the longer at stem mid-point,
sheaths 1.5—3.2 cm long, margins entire; leaves
6.5—6.7 X 2.9—3.2 cm, coriaceous, ovate, basally
cordate, basal lobes overlapping by 1.5 mm, api¬
cally long attenuate, acute; inflorescences emerging
from two fibrous sheaths, fascicles of 4 or 5 flowers,
I or 2 open at a time; pedicels ca. 2 mm long;
pedicellate ovary 2.5 mm long, slightly verrucose;
flowers fleshy, resupinate, concave, basically deep
wine-purple, tbe sepals marginally somewhat or¬
ange-yellow, the petals and labellum rich purple-
red; sepals concave, thick, the margins incurved
and minutely verruculose; dorsal sepal 5.5 mm
long, 3.5 mm wide when expanded, elliptic-oblong,
obtuse, 3-nerved; lateral sepals totally connate into
a synsepal 4.4—4.6 X 6.4—6.6 mm when expanded,
widely ovate in general outline, fleshy, with 6
nerves, apically notched; petals 3—3.1 X 1.1—1.2
mm when deliberately expanded, thickly fleshy and
concave, narrowly elliptic, acute, falcate, margins
Volume 13, Number 4
2003
Carnevali & Ramirez
Novelties in Orchidaceae
413
3 mm
Figure 6. Pleurothallis loreae Carnevali & I. Ramirez. —A. Habit. —B. Floral segments flattened. —C. Flower in
natural position. — D. Labellum. —E. Column with labellum attached. (Based on the holotype, G. Carnevali el al.
3263, C1CY, VEIN.) Drawing courtesy of C. A. Luer, sketch #17007.
414
Novon
minutely crenate-denticulate, I-nerved; labellum
1.9—2.1 X 1.2— 1.4 min, thick, oblong-ovate in gen¬
eral outline, apieally rounded to broadly obtuse,
with a central depression, with a transversal ridge
near base below which it is shallowly concave; col¬
umn 1.2—1.4 mm long, proportionally short as com¬
pared to labellum.
Etymology: Named after Francisco Lorea Her¬
nandez, curator at the herbarium of the Institute de
Ecologfa, A.C., Xalapa, Mexico (XAL), and a spe¬
cialist in the Lauraeeae, who participated in the
collection of the type specimen.
This species is vegetatively identical to Pleu-
rothallis cordata (Ruiz & Pavon) Findley, a taxon
widespread from Bolivia to Venezuela. In the new
species, however, the petals are thicker, falcate,
glabrous, and the labellum is ventrally concave.
Florally and vegetatively it seems closest to P. cha-
ma Flier, known from Ecuador. The Ecuadorian
species is a much larger plant (stems to 19 cm long
in P. loreae vs. up to 20—35 cm in P. chama ) with
larger bowers (dorsal sepal 5.5 mm long in P. loreae
vs. 12 mm long in P. chama), featuring a 7-nerved
dorsal sepal (vs. 3-nerved in P. loreae ), and a flat
or convex, papillose labellum (vs. deeply concave,
glabrous in P. loreae).
Pleurothallis loreae is one of several distinctive
endemic orchids of the Sierra de San Fuis in north¬
western Venezuela. Other endemics occurring there
include Masdevallia garciae Fuer, Pleurothallis
sanluisii Foldats, Pleurothallis transversilabia Car-
nevali & I. Ramirez. Prosthechea garciana (Garay
& Dunsterville) W. Higgins, Stelis sanluisensis Fol¬
dats, and several other still undescribed orchid
taxa.
Pleurothallis escalerensis Carnevali & C. A.
Fuer, sp. nov. TYPE: Venezuela. Bolivar: car-
retera El Dorado-Santa Elena de Uairen, km
95 al S of El Dorado, light forest over granitic
outcrops, 300—400 m, 20 June 1994. G. Car¬
nevali, IF Fritz !().
letted area in Venezuela. It may have been over¬
looked due to its diminutive size. This species was
included as “Pleurothallis sp. A” in the Pleurothal¬
lis treatment of the Flora of the Venezuelan Gua-
yana (Carnevali & Ramirez in Carnevali et al..
2003: 523). The type locality, i.e., the lower slopes
of the Escalera in northeastern Bolfvar state at
200—500 m, is a place of high orchid diversity.
Here a mixture of tepui species meet their lower
altitudinal ranges and grow along with taxa asso-
Volume 13, Number 4
2003
Carnevali & Ramirez
Novelties in Orchidaceae
417
Table 1. A comparison of Pleu
rothallis stellata and P. index.
Feature
Pleurothallis stellata
Pleurothallis index
Leaf shape
narrowly ovate
ovate
Leaf base
with non-overlapping basal lobes
basal lobes overlapping for 1 cm or more
Leaf dimensions and proportions
9 X 2.5 cm; w/1 = 0.27
5.5/3; w/1 = 0.55 7.5/3.S; w/1 = 0.50
Structure subtending inflorescence
a horizontal sheath 2 cm long
a suberect spathe 12—18 mm long, 5 mm
wide (as in P. cardiostola)
Dorsal sepal
12 X 6 mm; w/1 = 0.5
16 X 4 mm; w/l = 0.25
Synsepal
12 X 5 mm; w/1 = 0.41
16 X 4 mm; w/1 = 0.25
Petals
(N
II
£
CO
X
LO
15 X 2.2 mm; w/1 = 0.14
Labellum
2X2 mm; w/1 = 1
4.5 X 2 mm; w/1 = 0.45
Labellum shape
ovate, sub 3-lobed, obtuse to
rounded
ovate triangular, acute
ciated with sandstone and granitic outcrops as well
as other taxa from lowland Guianan forests. Orchids
include Brassia bidens Lindley, Catasetum barba-
tum (Lindley) Lindley, Catasetum discolor (Lindley)
Lindley, Braemia vittata (Lindley) Jenny, Chaubar-
diella tigrina (Garay & Dunsterville) Garay, Cal-
eottia jorisiana (Rolfe) Schlechter, and many others,
including not fewer than 10 distinct Octomeria spe¬
cies.
Etymology. The name refers to the only locality
where this novelty is known, the ascent to Cerro
Venamo, along the road from El Dorado to Santa
Elena de Uairen, in Bolivar State, Venezuela. The
topography is steep and broken on this ascent and
so is the road, thus being commonly known as La
Escalera, “the stairway.”
I'leurothallis slellata 1. Ramirez & Carnevali, sp.
nov. TYPE: Venezuela. Distrito Federal: tall
cloud forest at ca. 4500 ft., above El Limon,
Mar. 1961, C. C. K. Dunsterville 610 (holotype,
AMES). Figure 8.
Pleurothallis haec P. indice affinis sed foliis proportione
angustioribus auriculis minoribus, spathis non obovatis
nec suberectis, synsepalo quam sepalo postico angustiore,
sepalis petalisque proportione latioribus, labello subtrilo-
bato obtuso discrepat.
Epiphytic, cespitose herbs; rhizome abbreviated;
stems erect, terete, 10—12 cm long, bearing 2 thin,
close-fitting sheaths below the middle; leaves 7—9
cm long, 2.0—2.5 cm wide, narrowly ovate to lan¬
ceolate, acute, thin but fairly rigid; inflorescences of
a single flower borne in succession from a grav-
brown. horizontal, ca. 2 cm long sheath at base of
leaf; pedicels ca. 10 mm long, ovary ca. 3 mm long;
flowers subfleshy, resupinate, widely spreading
(thus flower stellate), relatively showy for the sec¬
tion, sepals and petals deep, rich, very beautiful
“wine-purple” (fide type), with veins of sepals (par¬
ticularly dorsal sepal) showing up in even darker
color, all inner surfaces covered with microscopic
short glandular projections, rather too short to be
called hairs (fide type, ex schedulae); dorsal sepal
12X6 mm, narrowly ovate, acuminate, acute, 5-
nerved; lateral sepals totally connate into a lance¬
olate synsepal 12 X 5 mm, acute, acuminate, 6-
nerved; petals 8.T-8.6 X 1.7-1.9 mm, narrowly
elliptic, acuminate, thick and fleshy at base, mar¬
gins finely ciliate, 1-nerved; labellum 1.9—2 mm
long, 1.9—2 mm wide when spread, fleshy, dark
wine purple, surface finely glandular, ovate-trian¬
gular. somewhat 3-lobed, obtuse to rounded, with a
transverse ridge at base, flexibly attached to col¬
umn. glenion well-developed; column 1.4—1.6 mm
long, stout, oblongoid, dorsoventrally depressed,
puce-colored, stigmatic surfaces two, elliptic,
shiny; rostellum acicular; anther creamy white; pol-
linia 2, hard, yellow.
Etymology. The name refers to the stellate as¬
pect of the flower, which is due to the relatively
similar shapes ol the widely spreading dorsal sepal,
synsepal, and petals.
This showy new species is superficially reminis¬
cent of the Colombian species Pleurothallis itulex
Luer. However, Pleurothallis stellata differs from P.
index in a number of features, especially the lack
of the suberect spathe. In addition, the large, over¬
lapping basal lobes found in the leaves of P. index
are much reduced and non-overlapping in the Ve¬
nezuelan P. stellata. Moreover, the generally small¬
er, differently proportioned flowers of P. stellata are
distinct. In particular, the labella of the two taxa
are very different. See Table 1 for a comparison of
the two species.
Acknowledgments. The authors are grateful to
Carl A. Luer, Bruno Manara, Rossana Marrufo, and
Carlos Yam for the line drawings, to the curators of
418
Novon
the herbaria cited herein for their assistance, and
to Gustavo A. Romero for informative discussions
and lor his editorial help.
Literature Cited
Carnevali, G., I. Ramfrez-Morillo, G. Romero-Gonzalez, C.
Vargas & E. Foldats. 2003. Orchidaceae. Pp. 200—765
in P. Berry, k. Yatskievycli & B. Holst (editors). Flora
of the Venezuelan Guayana, Vol. 7. Missouri Botanical
Garden Press, St. Louis.
Christenson, F. A. 1996. Notes on Neotropical Orchida¬
ceae—II. Lindleyana 11: 12-26.
Couret, P. 1982. Jewels of the Venezuelan Orchids. Li-
brerfa Alemana Oscar Todtmann, Caracas, Venezuela.
Dunsterville, G. C. k. & L. A. Garay. 1979. Orchids of
Venezuela, An Illustrated Field Guide, A—G. Botanical
Museum, Harvard Univ. Printing, Cambridge, Massa¬
chusetts.
Greuter, W., .1. McNeill, F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
J. F. Skog, P. Trehane, N. J. Turland & I). L. Hawks-
worth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Hagsater, E. & C. H. Dodson. 1993. Epidendrum ther-
mophilum Hags. & Dodson, in E. Hagsater & G. Salazar
(editors), Icon. Orchid. 2: 193.
Luer, C. A. 1986. leones Pleurothallidinarum ML System-
atics of Pleurothallis (Orchidaceae). Monogr. Syst. Bot.
Missouri Bot. Card. 20.
Pridgeon, A. M. & M. k. Chase. 2001. A phylogenetic
reclassification of Pleurothallidinae (Orchidaceae). Lin¬
dleyana 10: 235-271.
Romero, G. A. 1997. Venezuela, Paraiso de Orqufdeas.
Armitano eds. Caracas, Venezuela.
A New Species of Cipura (Iridaceae) from
Colombia and Venezuela
Marcela Cells
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495 Bogota,
Colombia, rncelis@ciencias.unal.edu.co
Peter Goldblatt
B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299,
St. Louis, Missouri 63166-0299, U.S.A. peter.goldblatt@mobot.org
Julio Betancur
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogota,
Colombia. jbetane@cieneias.unal.edu.co
ABSTRACT. Cipura gigas is a new species of this
Neotropical genus of nine species that extend from
Paraguav in the south to Cuba and Mexico in the
north. Apparently most closely related to the wide¬
spread C. paludasa, C. gigas is distinguished hy its
height (50-100 cm), the large bulb (ca. 3 cm diam.)
with resinous tunics, basal leaves that are mostly
8—10 mm wide, lilac-purple flowers, and the inner
tepals that form a narrow-mouthed urn-like cup 10-
13 mm long. The species is restricted to the Ori¬
noco basin of eastern Colombia and adjacent Ve¬
nezuela. We provide a revised key to the genus.
Key words: Cipura, Colombia, Iridaceae, Iri-
doideae, Venezuela.
Cipura Aublet (Iridaceae. Iridoideae) is a New
World genus in the tribe Tigridieae that is charac¬
terized by a bulbous rootstock, plicate leaves, and
an ancestral, basic chromosome number of x = 7
(Goldblatt, 1990, 1998). Cipura is distinguished hy
the presence of a subapical cauline leaf inserted at
the base of the subterminal inflorescence(s); inner
tepals that are held erect, often overlapping and
thus forming a wide- to narrow-mouthed cup; and
inner tepal limbs that are short or obscurely distin¬
guished from the claws (Goldblatt & Henrich,
1987, 1999). The genus extends from Paraguay and
southern Brazil to central Mexico and Cuba, and
now includes nine species (Ravenna, 1988).
Two species were unnamed in Goldblatt and
Henrich’s review (1987) of the genus. One of these,
Cipura sp. 1, was subsequently described as C. for-
rnosa by Ravenna (1988). The second species, Cip¬
ura sp. 2, from the frontier region between Colom¬
bia and Venezuela and now known from ample
material, is described here as C. gigas.
In his review of Cipura, Ravenna (1988) also
added another new species, C. paradisiaca Raven¬
na from Brazil, evidently closely related to C. xan-
thomelas Martius ex Klatt. Ravenna also presented
an argument for the recognition of C. inornata Ra¬
venna, a species included by Goldblatt and Hen-
rich (1987) in C. campanulata Ravenna. The latter
was regarded as a synonym of C. cubensis Grise-
bach by Ravenna (1988). Lastly. Ravenna provided
a new name, C. insularis, for the Cuban species C.
cubensis Wright ex Saville, a homonym in Cipura,
dating from 1871. The latter species, evidently
closely related to C. cubensis, was described when
collected as having yellow flowers. It otherwise
closely resembles a slender C. cubensis. Cipura
thus currently has nine species. Cipura cubensis
and C. insularis are difficult to distinguish from C.
inornata from herbarium material, and we question
their status as separate species. According to Ra¬
venna, C. cuberjsis may be separated by the almost
flat leaves, bell-shaped perianth, inner tepals near¬
ly as long as the outer, and minutely funnel-shaped
stigma versus lightly plicate leaves, cup-shaped
perianth, inner tepals shorter than the outer, and
minutely 3-lobed stigma in C. inornata. Illustra¬
tions showing the floral differences are sorely need¬
ed.
Previous studies reported three species of Cipura
from Colombia, C. paludosa Aublet, C. rupicola
Goldblatt & Henrich, and C. gigas. described here,
before reported as Cipura sp. 2 (Goldblatt & Hen¬
rich, 1987) or Cipura sp. A (Goldblatt & Henrich,
Novon 13: 419-422. 2003.
420
Novon
1999). Cipura rupicola, however, only occurs in
lowland Venezuela adjacent to Colombia.
In this paper we describe Cipura gigas as a new
species and provide a key to the genus as a pro¬
visional contribution anticipating the publication of
Ravenna’s promised revision of the genus (Raven¬
na, 1988).
Cipura gigas Celis, Coldblatt Betancur, sp. nov.
TYPE: Colombia. Guaviare: municipio de Mir-
aflores, Serranfa La Lindosa, trocha al Nuevo
Tolima, finca Casa Blanca, cercam'as de la
casa, en sabana potrerizada, 2°29'12"N.
72°41'13"W, 465 m, 1 June 1997, J. Betancur
3.1 mm; corolla tube
funnelform; pollen white; tube 35-45 mm
long; calyx greater than 15 mm long ....
. M. macrantha A. Gray
2b. Corollas pale white or pastel yellow or pastel
pink; corolla tube width al opening <3.1
mm; corolla tube cylindrical; pollen pink to
lavender; tube 20-30 mm long; calyx less
than 15 mm long . M. nana A. Gray
1 h. Caly ces less than 12 mm, lobes ± equal length
of tube; plants generally subshrubs; stems gen¬
erally ascending to erect; plants not or only
weakly rhizomatous; leaves narrowly lanceolate
to ovate {Monardella subg. Monardella).
3a. Leaves distinctly bicolored, green above and
paler beneath. M. hypoleuca A. Gray
3b. Leaves concolorous, similar on both surfac¬
es.
4a. Pubescence very dense, stems and
leaves whitish with epidermis obscured;
foliage very faintly scented; glands in¬
conspicuous; dry interior, montane
slopes.
. M. linoides A. Gray subsp. linoides
4b. Pul tescence sparse, stems and leaves
green with epidermis visible; foliage
strongly scented; glands conspicuous;
ephemeral drainages (moist or season¬
ally wet environments) in coastal foot¬
hills and mesas.
5a. Leaves broadly linear to narrowly
lanceolate with attenuate bases, 2-
4 mm wide, length to width ratio 7-
10:1; leaf glands dense below; stem
glands greater than 100/cm on one
side below the third leaf node from
the apical glomerule; middle bracts
10—15 mm long (longer than caly¬
ces); seeds less than 1.9 mm long;
plants in ephemeral washes (water
only running directly after rain) on
secondary alluvial benches . . .
. M. viminea Greene
5b. Leaves lanceolate to lance-ovate
with acute bases, 4—10 mm wide,
length to width ratio about 3-5:1;
leaf glands sparse below; stem
glands less than 50/cm on one side
below the third leaf node from the
apical glomerule; middle bracts 7—
432
Novon
10 mm long (roughly equal to ca¬
lyces); seeds greater than 1.9 mm
long; plants in streambeds and
banks of intermittent streams (water
running longer than directly after
rain) .
. . M. stoneana Elvin & A. C. Sanders
Acknowledgments. We thank Fred Roberts and
the San Diego Chapter of CNPS for the artwork and
map; Victoria Hollowed for tier tremendous efforts
during the review and publication process; Steve
Weller and Peter Bowler for reviewing the docu¬
ment and making constructive comments prior to
submittal; Gary Wallace for reviewing and making
constructive comments on earlier versions of the
key; Fia DeMarco for immeasurably improving the
literacy of our Latin description; IJC1 Arboretum for
allowing space to grow our material; San Diego Nat¬
ural History Museum (SD), Rancho Santa Ana Bo¬
tanic Garden (RSA), and the U.S. National Her¬
barium (US) for the loan of plant material to UCR;
and the reviewers who made constructive comments
that helped us improve the manuscript. We are par¬
ticularly grateful to our families for their support
and tolerance during the many long hours spent
away from them to complete this project.
Literature Cited
Abrams, L. R. 1912. The Monardellas of southern Cali¬
fornia— II. Muhlenbergia 8: 26—45.
-. 1951. Illustrated Flora of the Pacific States, Vol.
111. Stanford Univ. Press, Stanford, California.
Allen, L. S. 1994. Morphometries of Monardella subg.
Macranthae (Lamiaceae). California Polytechnic State
University, San Luis Obispo. Unpublished.
Beauchamp, R. M. 1986. A Flora of San Diego County,
California. Sweetwater River Press, National City, Cal¬
ifornia.
Botti, S. J. 2001. An Illustrated Flora of Yosemite National
Park. Yosemite Association, El Portal, California.
Briquet, J. 1896. Monardella. P. 809 m H. G. A. bugler
& K. A. Prantl, Die natiirlichen Pflanzenfamilien IV.
3a.
Epling, C. C. 1925. Monograph of the genus Monardella.
Ann. Missouri Bot. Card. 12: 1 — 107.
Gray, A. 1876. Miscellaneous botanical contributions.
Proe. Amer. Acad. Arts Sri. .5: 71-104.
Greene, E. L. 1902. New species of Monardella. Pittonia
5; 80-87.
Hickman, J. C. (editor). 1993. The Jepson Manual, Higher
Plants of California. Univ. California Press, Berkeley.
Jepson, W. L. 1925. Manual of the Flowering Plants of
California. Univ. California Press, Berkeley.
-. 1943. Monardella. Pp. 431—444 in A Flora of
California. Univ. California, Berkeley.
Jokerst, J. I). 1993. Monardella. Pp. 718-727 in J. C.
Hickman (editor), The Jepson Manual, Higher Plants of
California. Univ. California Press, Berkeley.
Munz, P. A. 1935. Monardella. Pp. 449—451 in Manual of
Southern California Botany. Claremont Colleges, Clare¬
mont, California.
-. 1959. A California Flora. Univ. California Press,
Berkeley and Los Angeles.
-. 1974. A Flora of Southern California. Univ. Cal¬
ifornia Press. Berkeley.
Three New Species of Cordia (Boraginaceae) from the Guianas
Christian Feuillet
Department of Systematic Biology-Botany, MKC-166, Smithsonian Institution, P.O. Box
3712, Washington, D.C. 20013-7012, U.S.A. feuillet.christian@nmnh.si.edu
ABSTRACT. Three new species of Cordia (Boragi-
naceae) are described from die Guianas. Cordia
cremersii and C. marioniae belong in section Var-
ronia, and C. fanchoniae in section Myxa. Cordia
marioniae is known from the Rupununi savanna in
the south of Guyana; the two other species have
been collected in French Guiana.
Key words: Boraginaceae, Cordia, French Gui¬
ana, Guyana, sect. Myxa, sect. Varronia.
Only a few species of Boraginaceae have been
added to those known from the flora of the Guianas
since the last treatments by Johnston (1932, 1935),
and addition by Uittien (1937). In contrast, recent
treatments from neighboring countries (Estrada S.,
1995; Gaviria, 1987; Taroda & Gibbs, 1986) de¬
scribed many new taxa, but none of those have
been documented to extend into the Guianas. Dur¬
ing the preparation of the treatment of the Boragi¬
naceae for Flora of the Guianas, some of the col¬
lections studied proved to be new species. Cordia
wurdackiana was published recentlv (Feuillet.
1997) and three new species of Cordia L. are de¬
scribed here, namely C. cremersii and C. fanchon¬
iae, both from French Guiana, and C. marioniae
from Guyana. Cordia is a large panlropical genus
with an estimated 350 species (Miller, 2001); it is
the largest genus in the Boraginaceae and mostly
Neotropical in distribution. Cordia has not been
treated as a whole since de Candolle (1845), the
many modern studies have mostly been of limited
geographic scope, and the genus is badly in need
of monographic study.
Cordia sect. Myxa (Endlicher) I Hi.
Cordia sect. Myxa (Endlicher) DC. is by far the
largest section of Cordia, and the only one of the
six currently recognized sections to be well repre¬
sented in all tropical regions. It is a group of ca.
200 species (Miller, 2001) that are mostly trees
with small white flowers in loose panicles and ca¬
lyces that are explanate under the fruits. It includes
all but two of the Old World species and the ma¬
jority of the New World rain forest species. Sev¬
enteen of the 29 species of Cordia in the Guianas
belong in section Myxa.
Cordia fanchoniae Feuillet. sp. nov. TYPE:
French Guiana. Piste de Saint-filie, 20 km
from RN 1, 5°20'N, 53°00'W, 8 Nov. 1995 (fl
& fr), M.-F. Prevost & A. Cortes 3188 (ho-
lotype, US; isotypes, CAY(2), NY, P). Figure
1 A-C.
Species ad Cordiam sectionem Myxam pertinens; brev-
ibus ramosisque cymosis inflorescentiis, supra glabris et
infra minuti-pilosis foliis, rubris, atromaculatis, obliquis
(ructibus notabilis.
Woody monocaulous plant 0.5—1.5 m tall, bloom¬
ing as small as 0.50 m tall. Internodes 3-20 mm
long. Stems densely short strigose. Leaves: petiole
canaliculate, short, up to 1.5 cm long, densely short
pubescent; blades narrowly oblong to elliptic or ob-
lanceolate, 14—32 X 4—8 cm, acute and slightly
decurrent at base, acuminate at apex, margin entire
and recurved, with 8 to 12 main lateral veins on a
side, the adaxial face drying brown, glabrous to gla-
brescent, veins concolorous, main lateral veins im¬
pressed, smaller veins raised, sometimes somewhat
while, the abaxial face drying tan to olive-green,
seeming glabrescent except on the veins, but ac¬
tually wi th some obscure pubescent, main lateral
veins raised, brown, tertiary veins yellowish, some¬
what obscure. Inflorescence axillary, condensed,
cymose, branched, branches scorpioid, to 2 cm
long, blooming acropetally, densely 15- to 20-flow-
ered; calyx barrel-shaped at anthesis, 4—5 mm long,
5-lobed, with a dense indumentum of thick straight
hairs outside, short appressed pubescent inside,
lobes triangular, up to 1.5 mm long, later torn into
a 5- to 8-toothed, nearly flat plate under the fruit,
whitish, glossy and glabrous inside; corolla white,
lube 5 mm long, lobes oblong, rounded at apex.
1.5—2 X 1.5 mm, reflexed against the calyx, gla¬
brous: in male flowers: stamens inserted on the co¬
rolla tube below middle, exserted, nearly 2 mm out
of the corolla tube, filaments 3.0-3.5 mm long, long
hirsute at base, glabrous above, anthers sagittate or
V-shaped; ovary lacking, replaced by a tuft of long
hairs; in female flowers: no stamen observed, ovary
Novon 13: 433-437. 2003.
434
Novon
Figure 1. A-C. Cordia fanchoniae Feuillet. —A. Leafy stem with inflorescences, the apical leaf showing venation
pattern. —B. Open corolla and stamens. —C. Gynoecium. D—F. Cordia cremersii Feuillet. —D. Leafy stem with
inflorescence, the apical leaf showing venation pattern. —E. Open corolla and stamens. —F. Gynoecium. G—I. Cordia
marioniae Feuillet. —G. Leafy stem with inflorescences, the apical leaf of the lateral branch showing venation pattern.
—II. Open corolla ami stamens. —1. Gynoecium. (A—C based on the US holotype, Provost & Corlds 31HH m . D—F based
on the US holotype. Cromers & de Granville 13934, and a photograph of a live plant by J.-J. de Granville; (i-l based
on the US holotype, Jansen-Jacobs et al. 3931.)
Volume 13, Number 4
2003
Feuillet
New species of Cordia from the Guianas
435
glabrous, broadly conical, 0.5-0.8 mm, 5-lobed at
base, style hirsute below 1st branching, glabrous
above, branched 1.0—1.2 and 1.5-1.7 mm above
the base, 4 terminal segments 1.2-1.5 mm long,
swollen and somewhat 2-lobed at apex. Fruit a
drupe, oblique in the calyx, 10 X 6 mm, base
asymmetric, round, tapering to the slightly acute
(when fresh) apex, mucronate, glabrous, abscission
of the style when fruit becomes bright red. black
around insect exit hole; stone not seen (all fruits
emptied by insect larvae).
Cordia fanchoniae is known only from the basin
of the lower Sinnamary River in the center of north¬
ern French Guiana. The type was collected on an
abandoned timber unloading track. Flowering in
June—July and November; fruiting in June and No¬
vember.
This new species most closely resembles Cordia
nervosa Lamarck of C. sect. Myxa, also from the
Guianas. Cordia fanchoniae differs from C. nervosa
in having longer and narrower leaf blades that are
acute and slightly decurrent at the base, in its calyx
densely hairy outside and clearly 5-lobed, and gla¬
brous fruits. It differs from all other species of Cor¬
dia sect. Myxa of the Guianas in its axillary inflo¬
rescences. All collections, Prevost & Cortes 3188
and Loubry 1194 (0.5 m tall), and Molino et al.
1854 (1.5 m), have a monocaulous habit and axil¬
lary inflorescences. Whether or not the stem later
branches and has terminal inflorescences, this
monocaulous growth may be viewed as a sign of
neoteny equivalent to the sterile first growth unit
typical of Cordia sect. Myxa.
The architectural model corresponding to the
growth pattern of Cordia fanchoniae is Corner's
model as defined by Halle and Oldeman (1970) and
Halle et al. (1978). As shown in Prevost & Cortes
3188 (CAY, sheet 2), after a traumatic reiteration
as defined by Oldeman (1974), the new shoot is
monopodial, unbranched, and the inflorescences
are axillary. Previously known architectural models
in Cordia are Prevosts (sect. Myxa: spp.), Fager-
I ind's (sect. Gerascanthus (Browne) G. Don: C. al-
liodora (Ruiz & Pavon) Oken), and Champagnat’s
(sect. Varronia: spp.).
Cordia fanchoniae is named to honor my friend
and collector of the type, Marie-Fran<;oise '"Fan-
chon" Prevost, who has a keen eye for unusual spe¬
cies and plant architecture.
Paratypes. 1RFNCH GUIANA. Near the lake of Petit
Saut dam (Sinnamary River), 5°03'N, 52°54'W, 3 July
1997 (fl), J.-F. Molino. I). Sabatier & I). Paget 1854 (CAY.
US); Piste de Saint-Flie, 22 km from RN 1, 5°20'N,
53°00'W, 18 June 1991 (fl & fr), D. hmbry 1194 (CAY).
Cordia sect. Varronia (Browne) G. Don
Cordia sect. Varronia (Browne) G. Don is well
represented in open vegetation in Neotropical re¬
gions. It is a group of ea. 100 American species
(Miller. 2001) with a shrubby habit and inflores¬
cences condensed in a spike or a head and calyces
cupular or campanulate, partially or totally enclos¬
ing the fruits. In the Guianas, 8 of the 29 species
of Cordia belong in section Varronia.
Cordia cremersu Feuillet, sp. nov. TYPE: French
Guiana. Bassin de l’Oyapock, Roche Touatou,
150 m, 2°57'N, 52°32'W, 16 May 1995 (fl), G.
Cremers & de Granville 13934 (holotype,
US; isotypes, CAY, NY, P, U). Figure 1D—F.
Species ad Cordiam sectionem Varroniam pertinens;
1.2 X 1 cm globosis capitibus inflorescentiarum, 0.7—1.0
cm longis pedunculis sub anthesi, 1.5-2 cm longis post
anthesin, pubescentibus foliis notabilis.
Shrub 1.5—2 m tall. Twigs with internodes 6—9
mm long, pubescent near the apex. Leaves: petioles
to 5 mm long, densely hairy; blades ovate or less
often elliptic, 3-8 X 1.2-3 cm, cuneate at base,
acute at apex, margin serrulate to doubly serrate,
upper surface drying dark brown, densely pubes¬
cent becoming glabrescent, veins impressed except
a narrow and raised midrib, lower surface with a
white and dense pubescence, venation brown show¬
ing through, veins raised including a wide midrib,
5 to 7 main veins on a side. Inflorescence terminal
and axillary, condensed heads 1.2 X 1 cm, on a
short peduncle 0.7—1.0 cm long at anthesis, becom¬
ing 1.5-2 cm long when fruiting, 15- to 25-flow-
ered; calyx funnel-shaped, 4 mm long, including
five 0.5—1.5 mm long teeth, yellow-green at anthe¬
sis, with scattered white trichomes outside, slightly
denser toward the apex, much denser in bud; co¬
rolla tube 4—5 mm long, 5 lobes broadly elliptic,
rounded at apex, reflexed, 7-10 X 5 mm, white,
pubescent to glabrescent; stamens inserted on the
corolla tube at or slightly below middle, barely ex-
serted, filaments 1.5-2 mm long, with a tuft of hair
at base, anthers subglobose, ca. 0.5 mm long; ovary
conical, obtuse at apex, 0.5—0.6 mm long, glabrous,
style glabrous, branched 0.7-0.8 mm and 1.1—1.3
mm from base, the 4 terminal segments ca. 0.7 mm
long, not swollen at apex. Fruit not seen.
Cordia cremersii is known only from southeast
French Guiana from the type collection discovered
on the southern slope of Roche Touatou at about
150 m elevation. The population was growing at the
limit of the brush-islands on the “savane roche.”
Flowering in May.
436
Novon
This new species most closely resembles Cordia
harleyi Taroda of C. sect. Varronia, known only
from the state of Bahia (Brazil). Cordia cremersii
differs from C. harleyi in its shorter petiole's, longer
and narrower leaf blades that are enneate (vs. trun¬
cate at base) and acute to long acute at apex (vs.
obtuse or rounded), its denser and longer pubes¬
cence on the adaxial surface of the leaves, and its
shorter inflorescence peduncle and somewhat
smaller corolla.
Cordia cremersii is named to honor Georges Gre¬
niers, a discoverer of this new species, as well as
a tireless field botanist with whom I have collected
plants many times in French Guiana.
Cordia marioniae Feuillet, sp. nov. TYPE: Guy¬
ana. Rupununi District: Dadanawa, 120 m
elev., 2°49'N, 59°31'W, 0 June 1995 (fl & fr),
M. J. Jansen-Jacobs, B. J. //. ter Welle, C. Gus-
tafsson & V. James 393J (holotype, US; iso¬
types, BRG, GB, MO, U). Figure 1 (4—1.
Species ad Cordiam subg. Varroniam pertinens; spicatis
inflorescentiis, 3—5 cm longis pedunculis, obovatis foliis,
supra scabridis, infra pubescentibus, late rotundato apice
notabilis.
Shrub. Twigs with long internodes 1—2 cm long.
Leaves: petioles 5—fl mm long, with very short pu¬
bescence; blades obovate, 3.5—8 X 2—4.5 cm, at¬
tenuate at base, broadly rounded at apex, margin
obscurely and loosely serrate above middle, tri-
chomes oppressed, short, stiff, white above, longer,
soft, dense, erect, curved beneath, midrib and sec¬
ondary veins visible above, beneath tertiary vena¬
tion netting visible, drying chocolate brown above,
paler with dark veins beneath, 4 to 6 main veins
on a side. Inflorescence spikes blooming basipetal-
ly, terminal and in a few apical axils, peduncle not
adnate to petiole, 3—5 cm long, fertile part 3—6 cm
long, densely 15- to 40-flowered; calyx pale green,
cup-shaped, about 2 mm long, including 5 trian¬
gular lobes, 0.5 mm long, with a white, oppressed
indumentum outside; corolla white, tube 5 mm
long, salveriform, 5 lobes reflexed, round, emargin-
ate at apex, ca. 0.5—0.6 X 0.7—0.8 mm, glabrous;
stamen inserted on the corolla tube X A of the length
from the base, barely exserted, filaments 2 mm
long, with a tuft of hair on the proximal Vs, anthers
broadly elliptic; ovary broadly conical, ca. 0.7 mm
long, glabrous, style glabrous, divided twice in 2
branches (nearly at the same point) ca. 1.8 mm
from base, 4 terminal segments ca. 1.7 mm long,
narrowly swollen in the apical half. Young fruit
spherical.
Cordia marioniae is known from two collections
discovered 150 years apart in the Rupununi Dis¬
trict, near Dadanawa close to the southwest corner
of the Kanuku Mountains. The type collection is
from a shrub growing in a small, rather dry bush-
island in the savanna. The older paratype, Schom-
burgk 1304, from the bank of the Rupununi does
not have any flower or fruit left on the peduncles,
but shows the typical leaf shape of the species.
Dadanawa is close to the Rupununi River where
the plant probably grew in the clear, narrow gallery
forest. Flowering and fruiting in May-June.
Cordia marioniae is easily set apart from all oth¬
er species of section Varronia by the obovate shape
of the leaf blades. It is also characterized by the
straight, thin inflorescence peduncle.
Cordia marioniae is named after Marion Jansen-
Jacobs, collector of the type, colleague, and long¬
time student of the flora of die Guianas with whom
I had the privilege to collect plants both in Guyana
and French Guiana.
Paratype. GUYANA. “Ufer des I'hisses Rupununi in
Brit. Guiana,” May 1843, Rich. Schomburgk 1404 (B).
Key for Cordia sect. Varronia in the Guianas
la. Inflorescences capitate.
2a. Inflorescences branched; flower heads < 5
mm diam .
.... C. polycephala (Lamarck) I. M. Johnston
2b. Inflorescences unbranched; flower heads >
5 mm diam.
3a. Peduncles > 5 cm long; corollas > 3
cm long C. grand if lor a (Desvaux) kunth
3b. Peduncles < 3 cm long; corollas < 2
cm long . C. cremersii Feuillet
lb. Inflorescences spieate.
4a. Inflorescence peduncles adnate to the peti¬
ole base.
5a. Leaf blades adaxially dull; inflorescenc¬
es densely flowered (calyces touching
each other and without gaps in the fer¬
tile part on the inflorescences); calyx
lobes acuminate, acumen free in closed
buds.
. C. tomentosa (Lamarck) Roemer &
Schultes
5b. Leaf blades adaxially shiny; inflores¬
cences loosely flowered (with gaps in
the fertile part on ihe inflorescences be¬
tween flowers or groups of flowers); ca¬
lyx lobes not acuminate.
. C. schomburgkii DG.
4b. Inflorescence peduncles not adnate to the
petiole base.
6a. Leaf blades adaxially glabrous; inflores¬
cences only terminal.
. C. curassavica (Jacquin) Roemer &
Schultes
6b. Leaf blades adaxially hairy; inflores¬
cences terminal and axillary.
7a. Leaf blades obovate, widely round¬
ed at apex .... C. marioniae Feuillet
Volume 13, Number 4
2003
Feuillet
New species of Cordia from the Guianas
437
71). Leaf blades ovate, acute at apex
.... C. stenostachya Killip ex Gaviria
Acknowledgments. I thank the curators of the
herbaria B, CAY, MO. NY, P. and U for lending the
material in their care. I am grateful to Cathy Pas-
quale-Johnson for the fine illustration. I thank J. S.
Miller for helpful comments and J.-J. de Granville
for making available to me a photograph of the live
plant (not included herein). This paper is published
as Studies on the Flora of the Guianas no. 98. This
is number 06 in the Smithsonian’s Biological Di¬
versity of the Guianas Program publication series.
Literature Cited
Candolle, A. P. de. 1845. Boraginaceae. Prodromus Sys-
tematis Naturalis Regni Vegetabilis 9a: 466—501. Tur¬
pin, Masson et al., Paris.
Estrada Sanchez, J. 1995. Cordia subgenero Varronia
(Boraginaceae). In: Flora de Colombia 14: 1-176.
Feuillet, C. 1997. Cordia wurdackiana, a new species of
Boraginaceae from Guyana. BioLlania, Ed. Esp. 6: 331 —
333.
Gaviria, J. 1987. Die Gattung Cordia in Venezuela. Mitt.
But. Staatssamml. Mtinchen 23: 1-279.
Hall* 4 , F. & R. A. A. Oldeman. 1970. Essai sur
l’Architecture et la Dynamique de Croissance des Ar-
bres Tropicaux. Masson, Paris.
-.-& P. B. Tomlinson. 1978. Tropical frees
and Forests. An Architectural Analysis. Springer, Ber¬
lin.
Johnst on, I. M. 1932. Boraginaceae. In: A. Pulle (editor).
Flora of Suriname 4(1): 306-333.
-. 1935. Studies in the Boraginaceae X. The Bor¬
aginaceae of northeastern South America. J. Arnold Ar¬
bor. 16: 1—64.
Miller, J. S. 2001. Two new species of Cordia L. (Bora¬
ginaceae) from Madagascar. Adansonia, ser. 3. 23: 289-
295.
Oldeman, R. A. A. 1974. L’architecture de la Foret Guy-
anaise. Memoires ORSTOM 73.
Taroda, A. & P. Gibbs. 1986. A revision of the Brazilian
species of Cordia subgenus Varronia (Boraginaceae).
Notes Roy. Bot. Gard. Edinburgh 44: 105—140.
Uittien, H. 1937. Boraginaceae. Additions and correc¬
tions. In: A. Pulle (editor). Flora of Suriname 4(1): 496-
497.
Chamaecrista mwangokae (Fabaceae, Caesalpinioideae), a New
Species from the Southern Highlands of Tanzania
Roy E. Gereau and Gretchen M. Walters
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, L.S.A.
roy.gereau@mohot.org; gretchen.walters@mobot.org
Abstract. We discuss the historical usage of the
genus Chamaecrista (Fabaceae, Caesalpinioideae)
in Africa, describe and illustrate the new species
C. mwangokae from southern Tanzania, reduce C.
hildebrandtii to taxonomic synonymy under C. zam-
besica, and provide a key to the Last African spe¬
cies of Chamaecrista with stipitate petiolar glands.
Key words: Caesalpinioideae, Chamaecrista,
Fabaceae, Tanzania.
Within Fabaceae subfamily Caesalpinioideae, Ir¬
win and Barneby (1982) placed Cassia L., Cha¬
maecrista Moench, and Senna Miller in tribe Cas-
sieae, subtribe Cassiinae. Chamaecrista is a genus
of approximately 330 species (G. P. Lewis, pers.
comm.), with greatest diversity in Central and South
America and a secondary center of diversity in
tropical Africa (Polhill & Thulin, 1989; Ihulin.
1993; Du Puy & Rabevohitra, 2002). Although
Meyer (in Meyer 100 stamens; stamens 0.4-0.7 X ca. 0.4
mm, with a very short filament and two lateral the¬
cae ca. 0.8 length of stamen. Pistillate flowers with
sepals broadly ovate, 6—8 X 6—9 mm, dark red,
chartaceous, appressed puberulous without, gla¬
brous within; petals 4, fleshy, lanceolate to broadly
lanceolate, acute at apex, 15—17 X 5—7 mm, con¬
nate at base for 3—4 mm, tomentose without, velu¬
tinous within; gynoeeium a globose receptacle 7—8
X 7—8 mm with 50 to 60 ovaries; ovary cylindrical
to prismatic, 1.5—3.5 X 0.8—1.5 mm. villous; stigma
sessile, glabrous, globular, ca. 0.5 nun diam.;
ovules 9 to 12 in ranks of 4 to 6. Fruit composed
of 9 to 25 nearly sessile, densely packed mono¬
carps; monocarps pale green ripening dark yellow,
sparsely appressed pubescent, oblong-ovoid, 1.7—
8.0 X 1.3—5.5 cm, apiculate, constricted in dried
specimens around two offset ranks of 3 to 6 seeds;
seeds ovate-ellipsoid, 18—25 X 8—13 mm.
Habitat. Wet submontane forest on well-
drained slopes at 780—1300 m allitiude.
Distribution. Cameroon. South West Province,
Rumpi Hills to Mount Kupe; Littoral Province, near
’t ingui.
Phenology. Flowers collected in January, Feb¬
ruary, and May; fruit collected in June and July.
Vernacular name. “Michile” in Bakossi ( Cheek
7131).
Paratypes. CAMEROON. South West Province:
hunters’ patli from Kupe Village to mountain top, 4°47'N,
9°41'E, 840 m. Cheek 7131 (K, SC A); below Kupe rock
near Esense river, 4°47'N, 9°41 E. 9(H) m. Elad 69 (SCA);
Ml. Kupe. Shrike trail, 12(H) m, Wheatley 398 (SCA); Ml.
Kupe, Shrike trail, 1 1(H) m. Cable 1221 (K): Ml. Kupe,
Shrike trail, 4°49'22"N, 9°41'2r'E, 1000 m, Kenfack
Volume 13, Number 4
2003
Kenfack et al.
Uvariopsis from Tropical Africa
445
f igure 1. Uvariopsis submontana Kenfack, Gosline & Gereau. —A. Apex of branchlet with leaves and detail of
venation. —B. Pistillate flower bud. —C. Staminate flower. —D. Staminate flower with one petal removed, lateral view.
—E. Inner surface of sepal. —F. Outer surface of sepal. —G. Gynoecium. —H. Detail of one carpel, lateral view. —
I. Infructescence. —J. Longitudinal section of mericarp. —K. Transverse section of mericarp. A-11 drawn from the
holotype, Kenfack 1334 (YA); I—K drawn from Kenfack 1373 (YA).
446
Novon
Figure 2. Uvariopsis submontana showing extreme trunciflory. Habitat shot in forest from Mt. kupe, Cameroon, of
Cheek 7131 photographed by Martin Cheek in January 1995.
Volume 13, Number 4
2003
Kenfack et al.
Uvariopsis from Tropical Africa
447
1373 (YA); Mt. Kupe, Shrike trail, 4°49'13"N, 9°4 I' I 7"E,
1110 m, Kenfack 1375 (YA); Mt. Kupe, Shrike trail, 1300
in, Kenfack 1376 (YA); path onto ridge leading to Kupe
rock. 4°47'N, 9°41 , E, 1000 m. Cable 2736 (K); Mt. Kupe,
permanent sample plot at 1200 m on Shrike trail. 4°50'N,
9°40'E, Lane 131 (K); forest trail 2 km S from Etuhe-Tape
Village, 4°51'N, 9°42'E. I 100 m. Cine 460, 534 (K); for¬
est on Mt. Kupe, 2 km E of Nyassosso Government School,
4°49'N, 9°42'E, 1000 m, I). W. Thomas 6039 (MO). Lit¬
toral Province: Nkam Division, Bataha, ca. 10 km W of
Yingui, 780 m. Kenfack 1602 (YA).
Uvariopsis submontana is most similar to and
probably most closely related to U. korupensis Ger-
ean & Kenfack, and can be distinguished from it
by the characters in the key to species (below).
These two species occupy closely adjoining geo¬
graphic ranges in Cameroon’s South West Province,
Uvariopsis submontana in submontane forest at 780
to 1300 m altitude, and U. korupensis in extremely
wet lowland forest at 50 to 160 m. Uvariopsis sub¬
montana is apparently never sympatric with any
other species of Uvariopsis.
Gereau and Kenfack (2000) cited two specimens
from submontane elevations as paratypes of Uva¬
riopsis korupensis. C. Doumenge 446, from forested
slopes in the Bakossi Mountains at 1 100 m. is a
fruiting collection with leaf measurements (32 X 12
cm) at the low' end of the range for U. korupensis ,
and may represent another population of U. submon¬
tana, although flowering material is needed for a
positive determination. I). W. Thomas 4477, from the
side of Mount Cameroon at 600 m. has flowers and
fruit but no leaves. Although floral measurements
(sepals ca. 3 mm long; petals ca. 17 X 10 mm) are
consistent w ith those of U. korupensis, more complete
material is needed, and Thomas 4477 may represent
another, still undescribed species.
Uvariopsis tripetalu (Baker f.) G. E. Schatz,
comb. nov. Basionym; Dennettia tripetala Bak¬
er f.. Cat. pi. Oban 5. t. 2. 1913. TYPE: Ni¬
geria. Benin City, Jan., R. E. Dennett 44 (lec-
totype, designated here, K).
I he monotypic genus Dennettia has traditionally
been distinguished from Uvariopsis on the basis of
its bisexual flowers with 3 sepals and 3 petals, ver¬
sus unisexual flowers with 2 sepals and 4 petals.
However, the description of U. bisexualis Verdcourt
expanded the generic concept of Uvariopsis to en¬
compass bisexual flowers. Dennettia occasionally
exhibits flowers with 2 sepals and 4 petals, and
Uvariopsis congolana (I)e Wildeman) R. E. fries
has 3 petals, making the number of perianth seg¬
ments unreliable as a generic distinction. Pollen of
Dennettia is very similar to that of Uvariopsis, being
borne in tetrads and possessing a rugulate-perforate
tectum, columellar infratectum, and basal foot-layer
of irregular foliations; it differs from the pollen of
Uvariopsis only by the absence of any granules in
the infratectum (Le Thomas, 1981). All these ar¬
guments justify the unification of Dennettia with
Uvariopsis. Baker (1913) described Dennettia with
its one species, D. tripetala, on the basis of two
syntypes, of which Dennett 44 is here chosen as the
lectotype over Talbot 1496 to honor further the col¬
lector R. E. Dennett.
I he key below distinguishes the 16 species of
Uvariopsis, including the new species and the sin¬
gle species here transferred from Dennettia, pri¬
marily by floral characters.
Key to the Species of Uvariopsis in Tkopicai, Akiuca
la.
lb.
Staminate (or hermaphrodite) inflorescences axillary or ramiflorous; pistillate inflorescences axillary, ramiflo-
rous, or trunciflorous.
2a. f lowers bisexual.
■3a. Sepals 2. petals 4; Tanzania . U. bisexualis Verdcourt
3b. Sepals 3; petals 3 (or 4); Cameroon, Nigeria. lb tripetala (Baker f.) G. E. Schatz
2b. f lowers unisexual, either staminate or pistillate.
4a. Petals connate al base; Cameroon. U. zenkeri Engler
4b. Petals separate at base.
5a. Leaf blade 5.5-16.1 cm long; petals ovate, 5-11 mm long; Angola, Cameroon, Democratic
Republic of Congo. Kenya. Sudan, Uganda, Zambia. U. congensis Robyns & Ghesquifere
5b. Leaf blade 12-23 cm long; petals suborbicular, 9-12 mm long; Ghana, Liberia .
. U. globiflora Keay
Staminate (or hermaphrodite) and pistillate inflorescences all trunciflorous.
6a. Flowers sessile or very shortly pedicellate; Cameroon .
.[/. sessiliflora (Mildbraed & Diels) Robyns & Ghesquifere
6b. f lowers distinctly pedicellate.
7a. Petals connate at base.
8a. Flower buds ovoid-conic to pyramidal.
9a. Leaf blade 12-30 cm long; pedicel 10-45 cm long; petals 3, broadly ovate; Cameroon,
Democratic Republic of Congo, Gabon. U. congolana (De Wildeman) R. E. fries
9b. Leaf blade 16—52 cm long; pedicel 1.5-8.0 cm long; petals 4, lanceolate to ovate.
448
Novon
10a. Leaf blade 30—52 X 9—14 cm; inflorescences mostly borne at base of Irunk;
sepals 2-5 mm long; petals 10-35 X 5-12 mm; Cameroon .
. U. korupensis Gereau & Kenfaek
10b. Leaf blade 16—38 X 5—11 cm; inflorescences densely covering trunk from base
to 2 m or more above ground; sepals 5—10 mm long; petals 15—18 X 5-8 mm;
Cameroon . (J. submontana Kenfaek, Gosline & Gereau
8b. Flower buds globose.
1 la. Staminate flower with petals 18-20 mm long; Cote d'Ivoire, Guinea-Konakry, Liberia.
Sierra l,eone . JJ, guineensis Keay
I l b. Staminate flower with petals 10—13 mm long; Cameroon, Democratic Republic of Con¬
go, Gabon. [J. vanderystii Robyns & Ghesquibre
7b. Petals separate at base.
12a. Petals ovate or elliptic-ovate, obtuse or acute.
13a. Leaf blade 12-20 cm long; staminate flower with petals 8-13 X 4.5—7.0 mm; pistiHate
flower with pedicel to 18 cm long and petals to 25 X 15 mm; Cameroon, Nigeria . .
. LI. dioica (Diels) Robyns & Ghesquibre
13b. I>eaf blade 19-29 cm long; staminate flower with petals 7—8 X 3—5 mm; pistillate
flower with pedicel 4.5—11.0 cm long and petals 8—9 X ca. 5 mm; Gabon.
. U. letestui Pellegrin
12b. Petals linear-lanceolate to oblong-lanceolate or ovate-lanceolate, acute to acuminate.
14a. Petals linear-lanceolate; Cameroon, Nigeria.
. U. bakeriana (Hutchinson & Dalziel) Robyns & Ghesquibre
14b. Petals oblong-lanceolate or ovate-lanceolate.
15a. Leaf blade narrowly elliptic or oblanceolate, 11—20 X 4—6 cm; staminate flower
with pedicel ca. 0.4 cm long; pistillate flower with pedicel ca. 7 cm long and
petals 14—15 X ca. 7.8 mm; Angola . II. noldeae Exell & Mendonga
15b. Leaf blade obovate-oblong, 16.6-29.0 X 5.0-9.5 cm; staminate flower with ped¬
icel 1.5—1.7 cm long; pistillate flower with pedicel 8—16 cm long and petals 15—
23 X 4.5-7.0 cm; Cameroon, Central African Republic, Congo Republic, Dem¬
ocratic Republic of Congo, Gabon.
. U. solheidii (De Wildeman) Robyns & Ghesquibre
Acknowledgments. For funding ol fieldwork lead¬
ing to the collection of the type anil paratypes, the
first author gratefully acknowledges the support of
the International Cooperative Biodiversity Group
for work in the Rumpi Hills, the Mount Cameroon
Project for work on Vlt. Kupe, and the WWF Coast¬
al Forest Program for work at Yingui. file second
author thanks the Farthwateh Institute and the IIK
Darwin Initiative lor their generous financial sup¬
port.
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J. Ramsbottom, A. Gepp & A. L. Smith, Catalogue of
the Plants Collected by Mr. & Mrs. P. A. Talbot in the
Oban District of South Nigeria. British Museum, Lon¬
don.
Doyle, J. A. & A. Le Thomas. 1994. Cladistic analysis
and pollen evolution in Annonaceae. Acta Bot. Galliea
14: 149-170.
-. 19%. Phylogenetic analysis and character evo¬
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Engler, A. & L. Diels. 1899. Diagnosen neuer afrikan-
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Exell, A. W. & F. A. Mendonga. 1951. Novidades da flora
de Angola. Rol. Sue. Brot., Sbr. 2, 25: 101-1 12.
Fries, R. E. 1959. Annonaceae. In: H. Melchior (editor).
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Duncker & Humblot, Berlin.
Gereau, R. E. & I). Kenfaek. 2000. Le genre Uvariopsis
(Annonaceae) en Afrique tropicale, avec la description
d'une espbee nouvelle du Cameroun. Adansonia, sbr. 3,
22: 39-43.
Heusden, E. C. H. van. 1992. Flowers of Annonaceae:
Morphology, classification, and evolution. Blumea,
Suppl. 7: 1-218.
Keay, R. Vi. J. 1952. Revision of the “Flora of West Trop¬
ical Africa”—I. Kew Bull. 1952: 149-1%.
-. 1954. Flora of West Tropical Africa, revised 2nd
ed. Vol. 1(1). Crown Agents for Oversea Governments
and Administrations, London.
Le Thomas, A. 1969. Annonacbes. In: A. Aubrbville (ed¬
itor), Flore du Gabon 16: 1-371. Musbum National
d’Histoire Naturelle, Paris.
-. 1981. Ultrastructural characters of the pollen
grains of African Annonaceae and their significance for
the phylogeny of primitive angiosperms. Pollen &
Spores 22: 267-342, 23: 5—36.
Pellegrin, F. 1948. Les Annonacbes du Gabon. III. Pa-
chypodanthium, Letestudoxa, Annona, Uvariopsis. Hex-
alobus , Enantia. Bull. Soc. Bot. France 95: 136—141.
Robson, N. K. B. I960. Annonaceae. In: A. Vi. Exell &
II. Wild (editors). Flora Zambesiaca 1: 104-149. Crown
Agents for Oversea Governments and Administrations,
London.
Robyns, W. & J. Ghesquibre. 1933. Essai de rbvision des
genres Uvariopsis Engl. & Diels et Tetrastemma Diels
(Annonacbes). Ann. Soc-. Sci. Bruxelles, Sbr. B, 53:
312-322.
Setten, A. K. van & J. Koek-Noorman. 1992. Fruits and
seeds of Annonaceae: Morphology and its significance
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Uvariopsis from Tropical Africa
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for classification. Studies in Annonaceae XVIf. Bib-
lioth. Rot. 142: 1-101.
Verdcourt, B. 1971. Annonaceae. In: E. Milne-Redhead
& R. M. Polhill (editors). Flora of Tropical East Africa.
Crown Agents for Oversea Governments and Adminis¬
trations, London.
-. 1986. New laxa of East African Annonaceae.
Kew Bull. 41: 287-297.
Hemipiliopsis, a New Genus of Orchidaceae
Yibo Luo and Singchi Chen (Xinqi Chen)
Laboratory of Systematic* & Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, Nanxincun 20. Xiangshan, Beijing, 100093, People’s Republic of China.
luoyb@ns.ibcas.ac.cn
Austkact. Hemipiliopsis, a monotypic* new genus
of Orchidaceae, is described based on //. purpureo-
punctata (K. Y. bang) Y. B. Luo & S. C. Chen (Hu-
benaria purpureopunctata K. Y. Lang) from south¬
eastern Xizang (Tibet). Its possible relationships to
Brachycorythis, Hemipilia, and Habenaria are dis¬
cussed.
Key words: China, Hemipiliopsis , Orchidaceae.
While the senior author worked on the genus
Hemipilia Lindley, he felt it difficult to treat a spe¬
cies that is very similar in habit to Hemipilia but
was described by Lang (Lang & Tsi, 1978) as Ha¬
benaria purpureopunctata K. V. Lang. Moreover,
Lang mentioned that this generic placement was
based on the presence of a small rostellum, naked
viseidia, and two protruding clavate stigmas (Lang
& Tsi, 1978). Based on these floral features, this
species appeared to be allied with Habenaria
Willdenow, while its purple-spotted stem, leaf, and
other parts, as well as a large-mouthed spur witli a
globular-swelled tip indicate its close relationships
to Hemipilia. Two years later, Lang (1980) stated
that Habenaria purpureopunctata is intermediate
between Habenaria and Hemipilia.
During a trip of the senior author to southeastern
Xizang (Tibet) in 1996, several plants of this spe¬
cies were collected and later transplanted to our
greenhouse. They flowered the next year, so we had
the opportunity to investigate its floral morphology
and ontogeny. As a result, Habenaria purpureo¬
punctata was tentatively proposed as a new genus
of Orchidiinae s. str. (sensu Dress ler, 1993) rather
than a member of Habernariinae (sensu Dressier,
1993; Luo el al., in prep.). Moreover, the nuclear
ribosomal DNA internal transcribed spacers (ITS)
data supported the placement of Habenaria pur¬
pureopunctata in the subtribe Orchidiinae s. str.
(sensu Dressier, 1993; Bateman el al., 2001). It is
treated here as a separate new genus differing from
both Habenaria and Hemipilia.
Hemipiliopsis Y. B. Luo & S. C. Chen, gen. nov.
TYPE: Hemipiliopsis purpureopunctata (K. Y.
Lang) Y. B. Luo & S. C. Chen.
Hoc genus novum Hemipiliae et Habenariae simile, sed
ah ambobus forma ealcaris, a priore viseidiis plus minusve
expositis, stigmatis lobulis duobus elongato-pulvinatis, a
posteriore planta (cum caule, folio, pedunculis, raehidi,
braeteis, pedicellis, ovariis, sepalis et petalis) purpureo¬
punctata, stigmatis lobulis parieti postico cavitatis alfixis,
atque rostello magno differt.
Terrestrial herb; tubers ellipsoid or subellipsoid,
fleshy. Stem erect, usually with one leaf near the
base. Leaf elliptic to ovate-oblong, acuminate or
acute, amplexicaul at base. Inflorescence loosely
several- to many-flowered, spotted with purple on
rachis and peduncle; bracts ovate-lanceolate, with
evident purple spots dorsally; pedicel and ovary
with purple spots. Flowers spotted with purple ex¬
cept the lip; dorsal sepal erect, oblong, concave,
forming a hood together with petals; lateral sepals
obliquely ovate-elliptic, usually ± reflexed; petals
ovate-orbicular; lip subobovate or ± fan-shaped, 3-
lobed at apex, the lobes nearly oblong, the lateral
lobes broader than the central one, spur abruptly
contracted in basal part and then strongly expanded
to form a globose apex; anthers sessile, erect, with
two parallel thecae; lateral gynostemium append¬
ages obvious and positioned next to anther; rostel-
lum three-lobed. with two separate lateral lobes
bearing naked viseidia at apex, the lateral lobes
extending forward, but not along w alls of cavity and
their apices completely separate, the viseidia po¬
sitioned directly above spur mouth; the extended,
two-folded central lobe approximately half as long
as anther; fertile stigma situated in a cavity and
consisting of two elongated pads adnate to its pos¬
terior wall, positioned below rostellum and directly
above spur mouth; pollinia sectile, fragile, each
proximally attenuated into a caudicle attached to
viscidium.
Hemipiliopsis purpureopunctata (K. Y. Lang) Y.
B. Luo & S. C. Chen, comb. nov. Basionym:
Habenaria purpureopunctata K. Y. Lang, Acta
Phytotax. Sin. 16(4): 127, fig. 2. 1978. T\ PE:
China. SE Xizang (Tibet): Bomi, 2500 m, 19
July 1965, Y. 7. Cluing & K. Y. Lang 384 (ho-
lotype, PE). Figure 1.
Novon 13: 450-453. 2003.
Volume 13, Number 4
2003
Luo & Chen
New Genus of Orchidaceae
451
Figure I. Hemipiliopsis purpureopunctata (K. Y. Lung) Y. B. Luo & S. C. Chen. —A. Plant. —B. Plant with two leaves.
—C. Flower, front view. —D. Flower, lateral view. —E. Column, front view. ——F. Column, lateral view. —G. Longitudinal
section of column. —H. Pollinium with viscidium, vertical view. —I. Pollirtium with viscidium, lateral view. Scale har:
A—C = 1 cm; E = 0.5 mm; I), F, G = 1 mm; II. I = 0.25 mm. Drawn from specimen of Y. B. Liio 14 in cultivation.
452
Novon
Plant covered with purple spots throughout, 20—
50 cm tall; tubers 2^1 X 1.5—2.0 cm; stem, usually
with 2 to 5 sterile bracts above the leaf. Leal slant¬
ing upward, 5—15 X 2-5 cm, green with purple
spots above, light purple beneath. Inflorescence 8—
20 cm long, loosely 2- to 20-flowered: bracts usu¬
ally shorter than ovary; pedicel and ovary 13—15
mm long, with purple spots. Flowers light purple or
white purple; dorsal sepal ca. 5X2 mm, rounded
or obtuse, 3-nerved; lateral sepals, ca. 6X4 mm,
obtuse, 3-nerved; petals ca. 5 X I mm, obtuse, 3-
nerved; lip 10—1 1 mm long, 9—10 mm wide in up¬
per part, 3-lobed at apex, the lobes irregularly un¬
dulate along apical margins; spur 9-10 mm long,
2—3 mm thick, contracted at a distance of ca. 1.5
mm from spur mouth, the globose apex ca. 2—3 mm
diam. Capsule ellipsoid, with purple spots, 15 mm
long, 2—3 mm diam. Flowering: June to July; fruit¬
ing: August to September.
This species is distributed in Nyingch, Lhiinze,
Mainling, Bomi, and Zayu, southeastern Xizang,
China, between 2100 and 3400 m, growing in
broad-leaved evergreen forest, alpine oak forest,
grassy places, and sandy soil along rivers. It also
occurs in Assam.
In habit Hemipiliopsis is similar to Hemipilia
(Lang & Tsi, 1978: Lang, 1980: Lang. 1999), but
differs from the latter by being covered with purple
spots throughout (Chen (‘I al., 1999).
As to its floral morphology, Hemipiliopsis has
pad-like stigmas adnate to the walls of the cavity,
a typical feature in the subtribe Orchidinae (Kur-
zweil. 1987: Kurzweil & Weber, 1991; Luo & Chen.
2000). It also lacks some typical features of Ha-
benaria, such as obvious anther canals and strongly
convex or stalked stigmas (Kurzweil & Weber.
1992; Luo et al., in prep.). Therefore, the floral
morphological evidence does not support a close
relationship between Hemipiliopsis and Habenaria.
On the other hand, this species is quite different
from Hemipilia in many floral features. For exam¬
ple, the genus Hemipilia is characterized by (I) a
well-developed three-lobed rostellum with the cen¬
tral lobe conspicuously projecting upward as high
as or higher than the anther; (2) lateral lobes of the
rostellum being completely separate and extending
along the walls of the cavity, forming a shallow
channel together with the prominently extended
basal parts of the lateral gynostemium appendages
in which the anther caudicle is positioned; and (3)
the viscidia being positioned on the inner sides of
the wall of the cavity (Luo & Chen. 1999. 2000).
In Hemipiliopsis, however, the three-lobed rostellum
does not have well-developed lobes, the lateral
lobes extend straight forward but not along the
walls of the cavity, and the viscidia are positioned
directly above the spur mouth (Luo et al., in prep.).
'fhe rostellum structure of Hemipiliopsis is simi¬
lar to that of Brachycorythis Lindley (Kurzweil &
Weber, 1991). especially that of B. tanganyikensis
Summerhayes (Cribb, 2001). The lateral rostellum
lobes of Brachycorythis are adjacent at their apexes
as illustrated by Kurzweil and Weber (1991, figs.
l, 2) except in B. tanganyikensis, where they are
obviously separated at their apexes as shown by
Cribb (2001. fig. 65.1). However, the lateral rostel¬
lum lobes of Hemipiliopsis are completely separated
at their apexes. In addition, Brachycorythis has
many canline leaves, foliaceous bracts, and a bi¬
partite lip. by which it is clearly differentiated from
Hemipiliopsis.
As demonstrated above, this entity has so many
distinct features, both in vegetative and floral char¬
acters, as compared with other members of the tribe
Orchideae (sensu Dressier, 1993) that it is reason¬
able to treat it as a new genus. Based on its Moral
morphology, especially the rostellum structure, it
seems more closely related to Brachycorythis than
to Hemipilia.
Specimens examined. CHINA. Xizang: Ma Gu Dong,
hillside, P. C. Tsoong 6685 (PE); Bomi. grassy slope, Y. T.
Chang & A. I. Ixing 828 (PE); forested valiey, 2050 in.
collector unknown 1522 (KUN, PE), Wu Chen-Yih 5638
(KEN): 2100 m, Xiao Pei-Gen et al. 2256 (KUN); 2300
m, conifer & oak forest, A Ludlow, G. Sherriff II. H.
Ellitot 13194 (BM), A Ludlow, G. Sherriff & //. //. El I i tot
15814 (BM); Nyingch, grassy place in forest, 2300-27(X)
m. Medicinal Plant Collecting Team (Xizang) 3422 (PE);
forested slope, 2700—2800 m. Medicinal Plant Collecting
Team (Xizang) 3539 (PE); under shrubs on sandy banks
of a river, 2550 in, Y. B. Luo 14 (MO. PE); 3000 m, 11.
I\. Qing et al. 533 (PE); Mainling, alpine oak forest,
3300-3400 m. collector unknown 4145 (PE); Additional
Collecting Expedition to Qinghai-Xizang 750836 (KUN,
PE); Lhiinze, 2740 in. Additional Collecting Expedition to
Qinghai-Xizang 750564 (KUN, PE); Lung, Chanyul Chu,
open dry hillside, 3000—3300 m. A Ludlow & G. Sherriff
2324 (K).
Acknowledgments. This study was supported by
the National Natural Science Foundation of China
(NSFC) (30270104). Henrik Pedersen and other re¬
viewers are thanked for their positive suggestions.
Thanks to Yingbao Sun for kindly preparing the
line drawing.
Literature Cited
Bateman, B. M.. P. M. Hollingsworth, .1. Preston, V. B.
Luo, A. M. Pridgeon & M. W. Chase. 2001. Phyloge¬
netics. Pp. 224—232 in A. M. Pridgeon, P. J. Crihb. M.
W. Chase & F. N. Rasmussen (editors). Genera Orchi-
dacearum Vol. 2, part 1: Orehidioideae. Oxford Univ.
Press, Oxford.
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2003
Luo & Chen
New Genus of Orchidaceae
453
Chen. S. C., Z. H. Tsi & Y. IL Luo. 1999. Native Orchids
ol China in Colour. Science Press, Beijing.
Cribb, P. J. 2001. Brachycorythis. Pp. 265—269 in A. M.
Pridgeon, P. J. Cribb. M. W. Chase & F. N. Rasmussen
(editors). Genera Orchidacearum Vol. 2, part I: Orchi-
dioideae. Oxford Univ. Press, Oxford.
Dressier. R. L. 1993. Phylogeny and Classification of the
Orchid Family. Timber Press, Portland, Oregon.
Kurzweil, H. 1987. Developmental studies in orchid flow¬
ers II: Orchidoid species. Nordic J. Bot. 7: 443—451.
- & A. Weber. 1991. Floral morphology of southern
African Orchideae. 1. Orchidinae. Nordic J. Bot. 11:
155-178.
-& -. 1992. Floral morphology of southern
African Orchideae. II. Habenariinae. Nordic J. Bot. 12:
39-61.
Lang, K. Y. 1980. A study on the geographical distribution
and floristic features of the Xizang (Tibet) orchid flora.
Acta Phytotax. Sin. 18: 391—407.
-. 1999. Habenaria. P. 429 in k. Y. Lang (editor).
Flora Reipublicae Popularis Sinicae Vol. 17. Science
Press, Beijing.
-& Z. H. Tsi. 1978. Some new taxa of Orchidaceae
from Tibet, China. Acta Phytotax. Sin. 16(4): 126-129.
Luo, Y. B. X S. C. Chen. 1999. Observations of putative
pollinators of Hemipilia flabellata Bur. et Franch. (Or¬
chidaceae) in north-west Yunnan Province, China. Bot.
J. Linn. Soc. 131: 45—64.
-& -. 20(H). The floral morphology and on¬
togeny of some Chinese representatives of orchid sub-
tribe Orchidinae. Bot. J. Linn. Soc. 134: 529-548.
Passiflora boenderi (Passifloraceae), a New Egg-Mimic
Passionflower from Costa Rica
John M. MacDougal
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
threehrane@sigmaxi.org
Abstract. Passiflora boenderi, a new species of
Passifloraceae endemic to Costa Rica, is described
and illustrated. It is assigned to section Decaloba
DC. in the species group that includes P. gilberti-
ana J. M. MacDougal and P. ornithoura Masters.
This rare small-flowered species is notable for its
intensely colorful and variegated leaves, with rows
of conspicuous golden egg mimics.
Key words: Costa Rica, egg mimic. Passiflora,
Passifloraceae.
For more than a century, a collection of a Costa
Rican passionflower with bilobed purple-tinged
leaves with yellow stripes and dots has hidden in
herbaria under different names. Only after plants
of this taxon were collected alive and then culti¬
vated have we been able to study the fresh flowers
and sec that it is an undescribed species, perfectly
morphologically distinct. While close relatives in
section Decaloba have similar variegated leaves,
this new species is most dramatic, with leaves dark
green above and purple or red-purple beneath, with
light yellow stripes along the two major lateral veins
and conspicuous bright golden spots in two lines.
It is a choice plant for ornamental cultivation for
its foliage, but is a local endemic in primary rain
forest in central Costa Rica and is of conservation
concern.
Passiflora boenderi J. M. MacDougal, sp. nov.
TYPE: U.S.A. Florida: Broward Co., Coconut
Creek, cultivated at Butterfly World, Ltd.,
1992—2002, type specimens collected Nov.
2001, R. Roender 361, grown from plants col¬
lected 19 May 1992 in Costa Rica. Alajuela:
W of main road San Ramon to Bajo Rodriguez,
between Rio Cataratas and Rfo San Lorenzo
on side rd. to Colonia Palmarena, “near dead
volcano,” ca. 1 km N of Finca Santa Cecilia,
10°13'43"N, 84°33'34"W, BOO m (holotype,
CR; isotypes, BM. CAS, INB. GOET, K.
MKXU, MO. TEX). Figures I. 2.
Species scandens ad Passifloram sectionem Decalobam
pertinens. Folia petiolo eglanduloso; lamina supra vittata.
infra purpurata vel atrovinosa, hiloha; lobis lateralibus
aeutis vel acuminatis, interdum rotundatis, lobo centrali
obsolete, angulo inter lobos laterales 11—62°, marginibus
integris, neetariis aureis. Flos parvus, petalis 3—6 mm lon-
gis; coronae filamentis biseriatis, exterioribus luteolis 3.5-
5.5 mm longis; opereulo plicato; androgynophoro 2.9-4
mm longo; semina (7 vel) 8 ad 10-sulcata.
Vine 1.5—5 m, minutely puberulent throughout
at the eernuous shoot lip. glabrescent below; stems
± terete, striate. Stipules 1.7—3.5 X 0.2—0.3 mm.
linear-triangular, subfalcate; petioles 1.2—3.8 cm,
eglandular; leaf blades 4— 12(— I 7) cm long in out¬
line, 4.0—8.0(—9.5) cm wide, 2.4-7(—9.0) cm along
central vein, lateral veins 4.0-8(-9.2) cm, entire,
glabrous or glabrescent except margins minutely
lightly strigillose and sometimes with a few tri-
chomes on the larger veins abaxially, variegated
adaxiallv with light yellow along the three major
veins, especially on lateral lobes, (often deep) pur¬
ple or red-purple abaxially. truncate elliptic to ob-
ovate to widely obovate (to very widely obovate) in
general outline, bilobed %—V 2 (-Zf) their length, the
lateral lobes triangular to lanceolate, acute to acu¬
minate, the very apex sometimes rounded, the cen¬
tral lobe absent or nearly so, sometimes represented
by a mucronate cusp less than 5 mm long, the angle
between the lateral lobes (14—)23—50(—62)° (juve¬
nile plants with leaves less deeply lobed). the ratio
of lateral to central lobe length 1.7—2.3(—2.5); lam¬
inar nectaries (5 to)8 to 13(to 16), borne between
the main veins, appearing bright yellow or yellow-
orange adaxiallv, and yellow-orange with purple
border abaxially, the larger leaves often with a sin¬
gle nectary proximal (exmedial) to each lateral vein
at base of lamina; prophyll of vegetative bud 1,
lanceolate to narrowly triangular. Peduncles (1)2
per node, 1—3 cm, uniflorous; bracts 3, 1—2.8 X
0.1—0.15 mm. linear-triangular, usually early ne¬
crotic and stramineous. Flowers light yellow-green,
the corona yellowish with purple or purple-red ba-
sally and/or distally; flowers with little or no de¬
tectable odor, borne sub-horizontally at anthesis;
floral stipe (3.5—)4.0—7.0 mm (6.5—9 mm in fruit);
hypanthium 5.5—7 mm diam.; sepals 7.0— 11 (— 14)
X 3.0—5.0 mm, triangular-oblong to narrowly ovate-
Novon 13: 454-458. 2003.
Volume 13, Number 4
2003
MacDougal
Passiflora boenderi from Costa Rica
455
Figure 1. Leaves of Possiflora boenderi (clone of type material, Boender 36]). Egg-mimic laminar nectaries can be
seen as round light spots on the leaves.
456
Novon
Figure 2. Flower of Passiflora boenderi with inner coro¬
nal series well expressed (clone of type material, Boender
B6I).
triangular, obtuse to rounded, w ith no apical or sub-
apical horn, light yellow-green outside, slightly pal¬
er light yellow-green or green-yellow inside, at
anthesis reflexed (to wrapped under hypanthium);
petals (2.7—)3.0—6.0 X 2.0—3.0 mm. triangular-ob¬
long to broadly triangular-oblong, broadly obtuse,
strongly reflexed at anthesis, sometimes erase api-
eally, very pale green (green-white) to [tale yellow-
green; coronal filaments 2-seriate, the outer coronal
filaments (31 to)35 to 43 (samples N = 10), 3.5—
5.5 mm long, at base narrow and greenish suffused
with light purple or red-purple, or dull purple
whole proximal half, conspicuously geniculate near
middle, conspicuously thicker and dull to light yel¬
low in middle or distal half, sometimes also purple
or purple-brown distally, but always with yellow or
pale yellow at least in middle, at anthesis the outer
corona forming a shallow bowl or saucer with a
strongly spreading and reflexed edge, the filament
tips pointing downward, the anther-corona clear¬
ance 2.5—3.5 mm; inner coronal series inconspic¬
uous, few to many, ea. 7 to 39 per flower, 1.0—1.8
mm, capillary, elavate to capitellate, dull light pur¬
plish with yellow head; operculum 2.0 mm, plicate,
pale greenish, flushed pale violet or red-purple be¬
low the white apex; androgynophore 2.9-4.0 mm
long, pale green basally, suffused purplish or red-
purple proximally, the free portions ol the staminal
filaments 2.6—3.5 mm long, red-purple or green suf¬
fused with pale purple; anthers 2.0—2.3 mm long,
purple-edged, the pollen yellow; ovary 1.5—2.0 X
1.3—1.5 mm, widely (to very' widely) ellipsoid,
bright green, minutely puberulent in all specimens
seen except one where the ovary is glabrous (En-
dres 70); styles 4—5 mm long including stigmas,
green and unmarked, or with a faint overall flush
of purple, the stigmas 1.5—2 mm diam. Fruit 11 —
21 X 10—20 mm, (ellipsoid) widely ellipsoid to
subglobose, estipitate, purple-black with glaucous
bloom, the mesocarp light green to white; arils 7—
8 mm long, transparent white to very pale trans¬
parent orange, gelatinous, sweet, not or only slightly
fruity, nearly odorless; seeds 3.5—4.2 X (2.4—)2.7—
3.0 X 1.4—1.7 mm, transversely sulcate with (t)8
to 10 sulci, the intervening ridges strongly sculp¬
tured and verrucose, the funiculus conspicuous and
white on fresh seed; seeds per fruit 24 to 67 (N =
10); germination epigeal.
Ecology. Passiflora boenderi is known from
only two sites in the Caribbean drainage of central
Costa Rica at 725—800 m elevation. These are
nearly identical habitats at the elevational transi¬
tion of very wet to pluvial premontane lorest. At
both sites nearly all trees and branches are fes¬
tooned with bryophytes, and the ground is often
saturated and muddy. The vines are found growing
up and into small trees on steep slopes, and on
large shrubs at forest edges. Narrowly endemic spe¬
cies such as this passionflower, restricted to very
small ranges, are typically in danger of extirpation
by habitat conversion, and are therefore of special
conservation concern. At the field site of the col¬
lection of the type clones, P. boenderi was associ¬
ated with or near P. ambigua Hemsley, P. lobata
(Fillip) J. M. MacDougal, P guatemalensis S. Wat¬
son, P costaricensis Fillip, P vitifolia Funth, and
P. oerstedii Masters (R. Boender, pers. comm.). At
the Cariblanco site 1 found P capsularis L and P.
lobata.
Phenology. Flowering plants have been col¬
lected in January, April, and August through No¬
vember.
This new T species was first collected more than
125 years ago by A. R. Endres and was cited by
Hemsley (1880: 481) as Passiflora , sp. (I 1 , capsu¬
laris [L.] ajf.)." The specimens are without exact
locality. One of Endres’s known collecting sites is
Quebrada Verde near San Ramon, very close to the
tvpe locality. We now know it from two localities,
but it is rare at both. Earrv Gilbert’s 1978 collection
was the second discovery of the species, and was
brought into cultivation for a short while but per¬
ished before studies could be made of it. Despite
repeated searches by botanists, the species was not
found again until 1984. In 1991 .Andres Vega found
it near San Ramon, took cuttings, and reared the
butterfly Heliconius cydno from eggs and larvae on
the plant. He guided Ron Boender there the next
Volume 13, Number 4
2003
MacDougal
Passiflora boenderi from Costa Rica
457
year and assisted getting more living material, some
of which ultimately furnished the type specimens.
Clones of the type collection of P. boenderi were
introduced to horticulture in late 1992 and had
spread to Europe by 1995. The collection number
of the type clones has been cited variously as Boen-
der 361, BW361, and BW92-361. Several recent
popular books on passionflowers have included this
species under a nomen nudum, and color photo¬
graphs can be seen in Vanderplank (1996: 61;
2000: 60—61). Kloek (1996: 105—106), and Ulmer
and Ulmer (1997: 110).
Passiflora boenderi is assigned to subgenus De-
caloba (DC.) Reichenbaeh sect. Decaloba DC. on
the basis of its plicate operculum, cernuous shoot
tips, transverse testal sculpturing, and position of
laminar nectaries. It is part of the species group
that includes 1\ ornithoura, P. gilbertiana, P. ape-
tala Killip. P. jorullensis HBK, P. mexicana Jussieu,
and P. affinis Engelmann. Most similar is an un¬
described relative of P. ornithoura from the moun¬
tains of Chiapas and Guatemala represented by
Matilda 3971. That can have remarkably similar
leaves, but P. boenderi differs by its longer outer
corona (3.5—5.5 mm vs. 1.8—3 mm), a slightly short¬
er androgynophore (2.9-4.0 mm vs. 3.4—-4.9 mm),
and a usually pubescent ovary versus an always
glabrous ovary. The bilobed variegated leaves of P
boenderi can be similar also to l 1 . gilbertiana and
P apetala, which both grow in Costa Rica. Passi¬
flora boenderi can be distinguished from P. gilber¬
tiana by its shorter androgynophore (3—4 mm vs.
6.2-9 mm), its shorter outer corona (3.5—5.5 mm
vs. 5—7 mm) that is strongly dilated proximally (vs.
filiform), and its habitat (elevations of 725—800 m
vs. 1600—2300 m). From P. apetala it may be rec¬
ognized by its often longer outer corona (3.5—5.5
mm vs. 2-5 mm) that is strongly dilated proximally
(vs. filiform), its longer petals (2.7-6 mm vs. absent
or to 2.5 mm), a usually pubescent ovary vs. always
glabrous ovary, and its habitat (elevations of 725—
800 m vs. 1280—3270 m). flic angles between the
leaves’ lateral lobes are usually wider in P. apetala
and narrower in P. gilbertiana . but there is overlap.
The inner corona was variably expressed in the
flowers of the six individuals I examined closely,
varying from 7 to almost 40. This is similar to the
variable expression of the inner corona in its very
close relative, P. gilbertiana, where their number
can vary greatly even on one plant (MacDougal,
1989). The conspicuous bright golden laminar nec¬
taries closely resemble Heliconius butterfly eggs
(Gilbert, 1982), and I consider this species to have
one of the most conspicuous eases of egg-mimicry
known (pers. obs.).
Fruit and aril observations were made by the au¬
thor from ten fresh fruits produced in cultivation
by Ron Boender through manual cross-pollination
of several individuals of the type material.
Etymology. This species is named for Ronald
Boender. founder of the Passiflora Society Interna¬
tional, and also president for many years. As prime
mover of that society, since 1989 Ron has brought
together several hundreds of persons interested in
the genus Passiflora, both amateurs and profession¬
als. He established a seed bank, living collections
database, newsletter, and annual national and in¬
ternational meetings. In this way, he has set the
foundation for a forum that has truly benefited sci¬
ence and the diffusion of knowledge of this plant
group. In Coconut Creek, Florida, he maintains the
world’s largest collection of germplasm of Passiflor-
aceae. Additionally, Ron brought this new species
into horticulture for closer study, unselfishly spread
it widely, and hounded me for more than a year
with carefully collected evidence that it was new.
Through his studies and educational presentation
of both butterflies and their host plants, he contin¬
ues to champion and support numerous conserva¬
tion activities in North, Central, and South Amer¬
ica.
Paratypes. COSTA RICA. Locality unknown: 2500 ft..
“Costa Rica,” [anno 1867-1875], Endres 70 (BM, K). Al-
ajuela: along Rio Sarapiquf near Ua Virgen tie Socorro
near Cariblanco, Apr. 1978, L. E. Gilbert s.n. (TEX); cam-
ino Colonia Virgen del Socorro, Cariblanco de Sarapiquf,
4 Jan. 2000, Kay & Vega 196 (MO); cuttings of Boender
361 from Costa Rica cultivated 1992—2002 at MO Cli-
matron® greenhouse, 8 Sep. 1993, MacDougal 4697 (CR,
MO); rd. to l.a Virgen de Socorro Irorn Rte. 9, near small
cascade along the rd. on E side of the Rfo Sarapiquf, ca.
10°15.4'N, 84° 10.3'W, 29 Aug. 1994, I). Smith 1106
(DUKE); Sarapiquf, Cariblanco, carnino a Virgen del So¬
corro, 8 Oct. 1992, A. Vega s.n. (CR, USJ).
Acknowledgments. Great appreciation is due
Andres Vega, the indefatigable field worker and
Costa Rican passionflower expert who helped re¬
discover this species and arranged for the collec¬
tion of the type material. I am grateful to the Pas¬
siflora Society International for support for field
studies and to Mario Posla for field assistance. 1
thank Armando Estrada for checking the herbaria
in CR, INB, and USJ, and Alexander Rodriguez of
INB for his assistance. John Vanderplank of the
National Collection of Passiflora in Britain gener¬
ously offered resources and shared photographs
that assisted in preparing the description of the
species. The curators of BM, DUKE, K, and TEX
generously offered specimens on long-term loan.
458
Novon
Literature Cited
Gilbert, L. E. 1982. The coevolution of a butterfly and a
vine. Sci. Amer. 247: 110—121.
Hemsley, W. B. 1880. Biologia Centrali-Americana, Bot¬
any, Vol. 1(6). 11. II. Porter and Dulau, London,
kloek, P. 1996. Das grosse Buch der Passionsblumen.
Lagerstroemia-Verlag, Hamburg.
MacDougal, J. M. 1989. Two new species of Pass if! ora
section Decaloba (Passifloraceae) from Costa Rica. Ann.
Missouri Bot. Card. 76: 608—614.
Ulmer, T. & B. Ulmer. 1997. Passionsblumen: Eine lasz-
inierende Gattung. Laupenmiihlen Druck, Witten.
Vanderplank, J. 1996. Passion Flowers, 2nd ed. MIT
Press, Cambridge.
-. 2000. Passion Flowers, 3rd ed. MIT Press, Cam¬
bridge.
A New Section of Passiflora, Subgenus Decaloba (Passifloraceae),
from Central America, with Two New Species
John M. MacDougal
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
threebrane@sigmaxi.org
A. Katie Hansen
Section of Integrative Biology, University of Texas, Austin, Texas 78713, U.S.A.
k.hansen@mail.utexas.edu
Abstract. The new section Pterosperma in Pas¬
siflora subg. Decaloba (Passifloraceae) is described
from southern Mexico and Central America on the
basis of leaf, inflorescence, seed, and fruit mor¬
phology. Species in this section are the sole hosts
of the specialized herbivore Eueides lineata (Heli-
conidae). A morphological key to its four species is
given, and two species, Passiflora lancetillensis and
P. pedicellaris, are newly described and illustrated.
Key words: Central America, Eueides, Mexico,
Passiflora, Passifloraceae. Pterosperma.
Exploration of the lowland wet forests of southern
Mexico and Central America has revealed the ex¬
istence of a group of four closely related species of
passionflower vines. When the first species to be
named was grown and studied by L. Gilbert, it was
obvious from the co-occurrence of the numerous
petiolar nectaries with the stalked, relatively un¬
specialized inflorescence structure that the species
did not fit into any known sections of the genus
Passiflora L., and would not key out to any of the
subgenera in Kill ip’s monograph (1938). This
unique group of large woody vines, distinguished
by morphological characters and a shared special¬
ized herbivore, is best recognized as a separate sec¬
tion of the genus and placed in subgenus Decaloba
(DC.) Reichenbach. This subgenus is characterized
by a plicate membranous floral operculum, varie¬
gation on juvenile leaves, and laminar nectaries not
marginal. Two of the four species are newly de¬
scribed here; the other two recently have been de¬
scribed with detailed ecological observations (Gil¬
bert & MacDougal, 2000; Knapp & Mallet, 1984).
Passiflora L. subg. Decaloba (DC.) Reichenbach
sect. Pterosperma L. Gilbert & J. M.
MacDougal, sect. nov. TYPE: Passiflora mi-
crostipula L. Gilbert & J. MacDougal.
Plantae scandentes ramulis juvenis apice cernuis. Folia
stipulis minutis caducis; petiolo neotariis (0 vel) 1 ad 1 I
munito; lamina elobata, integra. Inflorescentia ex floribus
uno ad sex sicut capreolo uno pedunculo portatis constans;
pedicello bracteis tribus 0.8—3.5 mm longis praedito. Flos
corona 2- ad 4-seriata; operculo plicato. Seminibus amplis,
complanatis, reticulatis, marginibus 4-alatis.
Medium-sized woody vines to canopy lianas;
stem and abaxial surface of leaves puberulent as
seedlings, erect trichomes sometimes glandular,
lightly puberulent to glabrous at maturity; stems te¬
rete, pith chambered, posture of shoot tip eemuous;
stipules extremely reduced, caducous; petioles
(0)1- to 11-glandular; laminas narrowly ovate to
(very) widely ovate (conspicuously peltate in one
species) with conduplicate ptyxis, entire, acute to
abruptly acuminate (rarely obtuse), variegated
adaxially or not in juveniles, usually glaucous abax-
ially, laminar nectaries present submarginally, or
absent; prophylls of vegetative bud 2, collateral;
tendrils in seedlings distally swollen and adhesive
in at least one species; peduncle 1 per node, bear¬
ing both a tendril and 1 or 2 pedicels, these often
branched so that the inflorescence is (1)2- to 6-
flowered, the tendril expressed in the inflorescence
sometimes aborting, the common peduncle some¬
times reduced to nearly nothing so that the pedicels
appear to arise from the leaf axils; bracts 3 per first
order pedicel, 0.8—3.5 mm long, narrowly triangular
to linear-lanceolate; floral stipe (pedicel distal to
articulation) conspicuous, 9-42 mm long; flowers
white with a yellow or white outer corona, this with
a purplish red band or not; sepals and petals sub¬
equal, 17—25 mm long, coronal filaments in (2)3—
4 series, the outermost 9-20 mm long, the inner
1.5—8 mm long; operculum plicate; androgynophore
7—14 mm; ovary puberulent to densely pubescent;
fruit 4—6 cm diam., conspicuously stipitate, inflat¬
ed, the pericarp very thin and leathery to parch-
ment-like; seeds 6—12 X 5—1 1 mm, the testa retic-
Novon 13: 459-466. 2003.
460
Novon
ulate in center and surrounded by striate wings
witli erose to laciniate margins, the chalazal beak
antiraphal; pollen 6-colporate; chromosome number
n = 9 (one species known). Germination epigeal.
These four rather similar species are lianas of
primary moist to wet tropical forest that form a co¬
hesive and well-marked group here recognized as
section Pterosperma. They flower in the canopy and
thus are rarely seen or collected, and although con¬
sidered to be uncommon to rare, they are some¬
times locally common as sterile juveniles in light
gaps (Gilbert, pers. obs.; Meerman, pers. comm.).
Three of the four are known to be the sole host/
food plant in their geographical area for the larvae
of the uncommon heliconid butterfly, Eueides linea-
ta (Mallet & Longino, 1982; Knapp & Mallet, 1984;
Meerman, 1999). The stipules of each species are
minute and deciduous, the new growth is cernuous,
there are multiple petiolar nectaries, the minute
laminar nectaries are submarginal when present,
and the inflorescence type is very rare in the genus,
i.e., a pedunculate dichasial cyme with the central
pedicel modified to be a tendril. This is the type of
inflorescence present in most of the other genera in
the tribe Passifloreae. The white flowers have the
characteristics of a bee-pollination syndrome, and
indeed, one is known to be bee-pollinated (Meer¬
man, pers. comm.). Each species has similar un¬
usual marginally winged seeds, hence the name of
the section. The arils and the shape, color, and per¬
sistence of the fruits are known for two of the four
species of this section, and suggest dispersal by
frugivorous bats (Gilbert & MacDougal, 2000).
Leaves of P. eueidipabulum and P. microstipula
were studied by Klucking, who published [(holo¬
graphs of leaf clearings (1992: 239, pi. 88, fig. 4;
245-246, pi. 98, fig. 5).
This distinctive section is placed in Passiflora
subg. Decaloba based on the presence of a plicate
operculum, cernuous stem growth, and variegation
along main veins of the leaf. Preliminary analysis
of two genes also supports this placement (Hansen,
in prep.). The new section is discordant in Decaloba
by its multiple petiolar nectaries, 6-colporate pol¬
len without secondary opereula (L. Escobar, pers.
comm.), and the fundamentally cymose inflores¬
cence. Though the stalked inflorescence type is like
that of most other genera in the tribe Passifloreae,
the placement of the bracts in the inflorescence is
not. Here, as in most passifloras, the displacement
of bracts onto the branches they normally subtend
(the recaulescence of Gusset, 1968) has produced
the evolutionarily derived state of 3-bracteate ped¬
icels. Unpublished studies have found the ehloro-
plast intron rpoCl to be present in P. microstipula,
an evolutionarily basal state in the genus (Hansen,
unpublished). The loss of this intron has occurred
in some other species of subgenus Decaloba. Snow
and MacDougal (1993) reported and illustrated the
one chromosome count for this group, 2 n = 18
(from P. microstipula as “P. sp. nov. A”). The chro¬
mosomes were relatively small, more like those in
the 2 n — 12 group, subgenus Decaloba, than in the
subgenus Passiflora, in which 2 n = 18 is the norm.
Thus, they concluded that the 2 n = 18 count is
not homologous between this section and subgenus
Passiflora.
Key to the Species ok Passiflora Section Ptkrospfrma
la. Leaves peltate 0.9—2.0 cm from the margin; floral
stipe 9—15 mm (Costa Rica, Panama).
.4. P. eueidipabulum
lb. Leaves not peltate, or subpeltate 0.2—0.5 cm
from the margin; floral stipe 16-42 mm (Mexieo
to Honduras).
2a. Leaves puberulent or pubescent abaxially at
maturity; petioles 6—11-glandular; (lowers
with white or whitish outer corona (Belize,
Honduras). 1 . P lancetillensis
2b. Leaves glabrous abaxially, at least at matu¬
rity; petioles 1 —10-glandular or glands ab¬
sent; flowers with yellowish outer corona
(Mexico, Guatemala).
3a. Petioles 1-glandular or glands absent;
lloral stipe 28—40 mm, pubescent; flow¬
er buds and sepals pubescent; flowers
1(2) per node (Guatemala) .
.2. P. pedicellaris
3b. Petioles 4—8(—l())-glandular; floral stipe
1C)—26 mm. glabrous; flower buds and
sepals glabrous; flowers 2 to 4 (to 6) per
node (Mexico) .3. P microstipula
1. Passiflora lancetillensis J. M. MacDougal &
J. Meerman, sp. nov. TYPE: Belize. Cayo: near
San Ignacio, cultivated at the National Pas¬
sionflower Collection of the Belize Tropical
Forest Studies at Green Hills. J. C. Meerman
s.n., 9 June 1996, from several seedlings col¬
lected by Meerman on 2 Dec. 1994 in Belize,
Cayo District, Tapir Mountain Nature Reserve,
17°09'N, 88°51'W, 100 m (holotype, BRH;
isotypes, BM, GOET, MEXU, MO (2), TEX).
Figures 1, 2. 3.
Haec species Passiflorae microstipulae similis, sed ab
ea flore omnino albo filamcntis coronalibus exterioribus
filiformibus tortuosisque ornato, foliis lalius ovalis ad no-
dos floriferos abaxialiter pubescentibus, sepalis pubescen-
tibus atque nectariis petiolaribus plerumque multioribus
distinguitur.
Large vine to low canopy liana, 9 m to at least
20 m, puberulent throughout with erect trichomes
0.3—0.6 mm long, these sometimes glandular, ex-
Volume 13, Number 4
2003
MacDougal & Hansen
New Section of Passiflora
461
Figure 1. Passiflora lancelillensis J. M. MacDougal & ,1. Meerman. Flower and peduncle of clone of type material
(Meerman s.n.). Note nectary glands on pubescent petiole at left. Photograph by Jan C. Meerman.
cept stem and adaxial surface of lamina glabrous
at maturity; stems terete, 0.8—3 cm diam., with
chambered pith, pubescent in younger plants, gla¬
brous at maturity except pubescence persisting
near nodes. Stipules minute to setaceous, 0.4—1.0
X 0.15—0.2 mm, deciduous; petioles 8- to I 1 -glan¬
dular, the nectaries 1.5—2.5 mm diam. in mature
plants, obloid to depressed obovoid, (petioles 6-11-
glandular in juvenile plants, the nectaries narrowly
cylindrical, 0.3—1.0 X 0.1—0.3 mm); laminas
(8)10—16(—22) X 7— 15(—19) cm, not variegated (nor
in seedlings seen, grown, or collected in Belize),
widely ovate to veiy widely ovate, unlobed, not pel¬
tate, entire, abruptly acuminate to obtuse, often
with submarginal laminar nectaries associated with
the end of major secondary veins, sessile or short-
cylindrical, borne 0.5—5 mm from margin; prophylls
of vegetative bud 2. 0.6—1.5 mm long, widely to
narrowly triangular, 3-toothed, collateral. Peduncle
(common peduncle ol tendril and flower) 1 per
node, bearing both a tendril and I or 2 pedicels,
0—2(—3) cm long to the first branch, the pedicels
0.3—1.3 cm long, often 1- or 2-branched so that the
inflorescence at the leaf axil is (1)2- to 4(to 6)-
flowered, the tendril expressed in the inflorescence
sometimes aborting, the common peduncle often re¬
duced to nearly nothing so that the pedicels appear
to arise from the leaf axils; bracts 0.8—2.0 X 0.2—
0.5 mm, triangular to oblong-lanceolate. Flowers
white with a whitish corona and light green center,
borne upward; stipe 25—12 mm; sepals 18—23 X
6.0—9.0 mm, narrowly ovate-oblong, green abaxi-
ally, pale greenish yellow to white adaxially, with
no projection; petals 19-22 X 7.0—8.5 mm, narrow¬
ly ovate-oblong, slightly narrowed at base, white;
coronal filaments in 3 or 4 series, the outer 15—20
mm, filiform, tortuous in distal half, slightly atten¬
uate toward tips, white or very pale cream, not
banded, the inner 2 or 3 series 3.5—6 mm, filiform
to capillary, straight, white; operculum 3.5—1.5 mm,
membranous, plicate, green with white-fringed
apex; limen floor apparently unmarked, androgyn-
ophore base not colored; androgynophore (8—)9—9.5
mm, light to pale green, in the type sparsely pu¬
bescent near the middle; stamina] filaments free 6—
8 mm, anthers 3.2—3.5 mm, not marked; ovary 3.7—
4.5 X 2.7—3.5 mm, on a 1.5 mm gynophore, widely
ellipsoid, densely pubescent; styles 9.5—1 1 mm in-
462
Novon
F igure 2. Passijlora lancetillensis J. MacDougal & J. Meerman. Two nodes along a stem, with associated leaves and
peduncles. Interpretation of the structure of the inflorescence is aided by the letter “T” marking the tendrils. The
articulations of the pedicels marking the floral stipes can be seen as darker lines on the pedicels. Scan by Jan C.
Meerman of living plants of the type material (Meerman s.n.).
eluding stigmas, green and unmarked, usually
lightly pubescent proximally. Fruit 6.5—8 cm long
including 2.5—3.0 cm long stipe, 4.0—4.8 cm diam.,
widely ellipsoid or subglobose, slightly conical at
attachment of stipe, blunt at apex, light green or
yellowish green at maturity, inflated, indehiscent
(but easily splitting into three parts with slight pres¬
sure), without strong odor, short-pubescent inside
on placental walls with erect thick trichomes, exo¬
carp very thin, leathery, drying parchment-like, ca.
1 mm thick; arils translucent whitish, insipid; seeds
ca. 20 to 36 per fruit, 9.5—11.5 X 7.0—8.0 X 2 mm.
dark brown, punetate-reticniate in center, the pits
sometimes weakly organized into a few wavy rows,
with 4 conspicuous marginal wings, (1.5—)2-2.5(—
3.0) mm wide, striate, deeply erose to laciniate (to
deeply laciniate at micropylar end), the chalazal
beak antiraphal; germination epigeal.
Herbivory. The butterfly Eueides lineata (Nym-
phalidae: Heliconiinae) appears to be restricted to
this species of vine in Belize. The females lay eggs
singly on the underside of young but nearly fully
expanded leaves and cause a distinctive feeding pat¬
tern of little windows in the leaf blade. The only
other Heliconiinae butterfly utilizing this species is
Dione juno, which is known to utilize a variety of
passifloras as the larval food plant. Passijlora lan¬
cetillensis is rather toxic to Dione, however, and the
larvae only rarely reach maturity (Meerman, pers.
comm.).
Distribution and habitat. Passijlora lancetillen¬
sis is a low canopy-dwelling species of tropical ev¬
ergreen or seasonal broadleaf lowland hill forest at
elevations of 20-750 m where rainfall varies from
2000 to 4000 mm annually (Meerman & Sabido,
2001). In Belize, it is found in strongly karstie re¬
gions, with forest canopies at 20—35 m, and at least
in some areas, with a marked dry season (Parker
et al., 1993: 18). Because it has been found on the
coast of Honduras and in extreme southwestern Be-
Volume 13, Number 4
2003
MacDougal & Hansen
New Section of Passiflora
463
■ Passiflora lancetillensis collections
□ Passiflora lancetillensis observations
'tt Passiflora pedicellaris type collection
0 50 100 150 200 Km
Figure 3. Map of northern Central America showing dis-
trihutions of Passiflora lancetillensis and P. pedicellaris.
Map courtesy of Jan C. Meerman.
lize, it might lie expected in hilly forest patches in
the intervening and neighboring departments of
Izabal and Peten of Guatemala. Associated species
of Passiflora include P. obovata Killip and P gua-
temalensis S. Watson in southern Belize, and P.
mayarum J. M. MacDougal in northern Belize.
This distinctive new species is presently known
from five collection sites, one the Lancetilla Valley
on the Atlantic coast of Honduras, and four from the
hills of southern and central Belize. Additional lo¬
calities where the species has been observed by
Meerman. but not collected, are noted in Figure 3.
The Honduran specimens, sterile and with large [ire-
reproductive leaves, were cited in the Flora of the
Lancetilla Valley as a probable new species (Stan-
dley. 1931: 293, citing the monographer Killip).
The species is clearly closely related to. but dis¬
tinct from. Passiflora microstipula and P. eueidipa-
bulum. Large leaves at prereproductive nodes are
non-peltate and evenly puberulent to pubescent be¬
neath, and the petioles have significantly more
glands. The stem is lightly puberulent. As in P.
microstipula, there is a pronounced change of mor¬
phology of the petiolar glands from narrowly cylin¬
drical in juvenile leaves to larger depressed ob-
ovoid glands in leaves on large plants or at
reproductive nodes (Gilbert & MacDougal, 2000).
Unlike P. microstipula, however, the leaves remain
pubescent on the undersides at maturity, and the
flower buds and longer floral stipes are pubescent.
The flower is distinctive in the group, with an all-
white filiform corona instead of stockier filaments
that are laterally compressed and yellow 7 or yellow
with reddish purple bands.
Fruits are known from a few 7 produced by open
cross pollination of the type plants in cultivation,
and from one fallen empty fruit of a canopy liana in
the field ( Holst & Davidse 5459). Seeds are known
only from the cultivated type material. The seeds are
similar to those ol Passiflora microstipula and P.
eueidipabulum, with the distinctive erose marginal
wings found in all the species of the section, but the
margins of the wings are cut more deeply, even la-
ciniate to base at the micropylar end.
At the National Passionflower Collection of the
Belize Tropical Forest Studies in Cayo District (240
m), the flowering season appears to be May through
August. Flowers stand erect and open before dawn.
On hot days, the flowers may wilt before noon, but
mostly remain open all day, even into the evening
on cooler days. Usually only one flower per pedun¬
cle opens each day. The flowers emit a strong and
pleasant coumarin-like odor. Medium-small bees
frequent the flowers in cultivation, with most activ¬
ity ceasing in mid-morning around 10 a.m. (Meer¬
man. pers. comm.). One of these visitors, a 1.0 cm
long bee in the Anthophoridae, closely matched the
functional size of the flowers and while foraging for
nectar, appeared to carry 7 pollen on its hairy 7 thorax
from flower to flower. The usual pollinators in wild
habitat are probably bees of similar size. Hum¬
mingbirds have also been seen occasionally to visit
the flowers (Meerman, pers. comm.).
Healthy FI seedlings have been produced in the
greenhouse by crossing between this new species
and Passiflora microstipula (Gilbert & MacDougal,
2000), but these have not flowered yet.
This new passionflower was listed (as a nomen
nudum, P. lancetillensis M[a|cDougal) in Parker et
al. (1993: 10, 38. 51), and its associated vegetation
was discussed. Jan Meerman introduced the spe¬
cies to horticulture by 1996, and briefly described
and figured it in the Passiflora Society Internation¬
als newsletter (Meerman, 1996). A color photo¬
graph and description in German soon appeared in
a popular handbook on passionflowers (Ulmer &
Ulmer. 1997). By 1998 it was listed in plant col¬
lections from at least four countries (Schappert,
1998: 23) and is now available from a few com¬
mercial plant dealers. The name also appeared in
the 1997 version of the IIJCN Red Booh (Walter &
Gillett. 1998: 446) where it was scheduled as R
(Rare) from information provided by MacDougal.
464
No von
The specific epithet refers to the Lancetilla Val¬
ley in northern Honduras where this species was
first collected.
Paralypes. U.S.A. Missouri: cultivated at Missouri
Botanical Garden 1996—2002, from seeds from Meerman’s
cultivated type plants, received Oct. 1996, MacDougal
6035 (MO). BELIZE. Belize: Graey Rock, 17°23'N,
88°26"W, 1996, Meerman JM048 /JMP034 (MO). Cayo:
Tapir Mountain Nature Reserve, 17°09'N, 88°5TW, UTM
16.303 E, 1897 N, 2 Dec. 1994, Meerman s.n./JM017H
(MO). Toledo: southwestern Maya Mountains, Columbia
River Eorest Reserve, Union Camp, dry hills E and N of
camp, I6°23'N, 89°09'W, 6 Apr. 1992, Holst 4144 (MO);
Maya Mountains, Bladen Nature Reserve, Upper Bladen
Branch basin, hill immediately E of “AC Camp,” ca. I
km SW ol Ek Xux ruin, I6°29'40"N, 88°54’30"W, 21 Mav
1996, Holst & Davidse 5459 (BRH). HONDURAS. Atlan-
lida: Lancetilla Valley, near Tela, ca. 15°42'N. 87°28'W.
6 Dec. 1927—20 Mar. 1938, Standley 55285 (F, US), Stan-
dley 56831 (F, photo: DUKE, HUA, MO; US).
2. PassiHora pedieellaris J. M. MacDougal, sp.
nov. TYPE: Guatemala. Baja Verapaz: Nino
Perdido, 15°10'N, 90°06'W, on Arroyo El Car-
acol, 6 km, 300-600 m, 23 June 1977 (bud,
(1, imtii. fr), C. L. Lundell & E. Contreras
21209 (holotype, LL; isotypes, LL, MO). Fig¬
ures 3, 4, 3.
Haec species Passiflorae microstipulae similis, sed ah
ea nectariis petiolaribus deminutis carentibusve, foliis an-
gustius ovatis, androgynophoro breviore atque nodis (lores
unimi duosve tantum gerentibus distinguitur.
Vine, glabrous except minutely puberulent with
straight, possibly glandular trichomes on the pro-
phylls ol the vegetative bud, the hypanthium and
sepals, anti the ovary; stems terete. Stipules 0.6—
1.2 X 0.1—0.2 mm, linear-triangular, late decidu¬
ous; petioles eglandular or I-glandular near or
slightly distal to the middle, the nectary 0.8—1.8 X
0.2—0.4 mm; laminas 9.5—16 X 5.5—10 cm, not pel¬
tate, or subpeltate 2—5 mm from the margin, the
base cordate, entire, not variegated at maturity,
ovate (narrowly ovate), unlobed, acute; laminar nec¬
taries absent; prophylls of vegetative bud 2, 1.1 —
2.5 mm long, narrowly triangular, slightly keeled at
base, one usually slightly shorter, sometimes with
a small marginal tooth. Peduncle (common pedun¬
cle ol tendril and (lower) 1 per node, reduced to
near absence (then the llower appearing axillary at
the base of the tendril) or up to 0.5 cm long, 1(2)-
llowered, the pedicels 0.5—1.5 cm; bracts 1.1 —1.9
X 0.3 mm, linear-triangular. Flowers white, the co¬
rona yellow; (loral stipe 28—36 mm (to 40 mm in
fruit); sepals 17—19 X 9 mm, ovate-oblong, green¬
ish abaxially, whitish adaxially, with no projection;
petals 17—18 X 9 mm, ovate-oblong, white; coronal
filaments in ca. 3 series, the outer 11-13 mm, lin¬
ear, basally yellow-green, distally bright to dark
yellow, no purplish red banding seen, the inner ca.
2 series 2—5 mm, light greenish or yellow-green
marked with purplish red; operculum 3 mm, mem¬
branous, plicate; limen floor pale with purplish ring
or markings, the base of the androgynophore not
colored; androgynophore 7.0—8.5 mm; anthers ca.
4 mm; ovary 3.0 X 1.7 mm, ellipsoid, sparsely to
lightly puberulent with straight, possibly glandular,
erect trichomes, styles ca. 4—6 mm long including
stigmas, glabrous. Fruit known only from immature
but apparently full-sized specimen, ca. 13 X 5.5—
6 cm including a 20—25 mm long stipe, ellipsoid,
apically somewhat conical, exocarp very thin, im¬
mature color green, apparently inflated; arils un¬
known: immature seeds 6 X 5 mm. punctate-retic¬
ulate in the center, with conspicuous I mm wide
wings at the margins.
This species is known only from the type collec¬
tion from tall moist forest in Baja Verapaz, Guate¬
mala. A single unbranched pedicel is borne off a
short to obsolete peduncle at or near the base of
the tendril; rarely a second pedicel is borne more
distally. Out of the dozen inflorescences in the type
material only one tendril/peduncle had a second
branch. With such limited material it is difficult, of
course, to generalize about a complex character
such as the inflorescence structure, especially when
that is known to be variable in the related species.
Nevertheless, it seems that this species normally
sports only one flower per node, unlike its sisters
in the section, which usually have 2 to 5 flower
buds per node. The inflorescence structure, with
the common peduncle much reduced to basically
lacking, is most similar to P. lancetillensis, which
also has very short peduncles. The floral stipe (that
part of the pedicel distal to the articulation) is un¬
usually long in both these species. Before the flow¬
ers of P. lancetillensis had been discovered, I chose
the specific epithet, pedieellaris, to highlight this
feature. The flower has an outer corona that is blunt
and yellow, like most of the rest of the section, but
unlike P. lancetillensis.
Passifiora pedieellaris is notable in this section
in that the petiolar nectaries are very reduced or
absent, the few single narrow' stubs seen on the
petioles of the type perhaps not even functional.
These glands are inconspicuous and probably are
not functioning as egg mimics. The glands are rath¬
er similar to those seen on seedlings and juveniles
of its relative, P. microstipula (Gilbert & Mac¬
Dougal, 2000). The fruit is similar in shape to that
of the Oaxacan clone of P. microstipula, but is
slightly larger. This poorly understood species is
Volume 13, Number 4
2003
MacDougal & Hansen
New Section of Passiflora
465
LUNDELL HERBARIUM
The University ol Texas
at Austin
FLORA OF GUATEMALA
I,undell Herbarium
peUt'oe/tAA* s
Ref '
^ issouri Botanical Garden (MO)
"Granndia" Vine* flowers white* fruit preen.
DEPARTMENT OF BAJA VHRAPAZ: Nino Perdido,
on Arroyo El Caracol* in high forest,
6 kin.
C. I. l.umlell
Eli.. Contrcriu No. 21209 June 23, 1977
Figure 4. Passiflora pedicellaris J. M. MacDougal. Holotype specimen conserved at LL. Scan by Fred Keusenkothen
(MO).
466
Novon
IfiL-
pressed dried flower from the Id, holotype. Scale marked
in mm.
most probably a medium-sized woody vine to can¬
opy liana like its sisters in the section. The ver¬
nacular name, “ granadia appears on the label of
the type collection.
3. Passiflora niicrostipiilu h. Gilbert & J. M.
MaeDougal, Lundellia 3: 1, fig. 1. cover plate.
TYPE: U.S.A. Texas: Travis Co., cultivated at
I’he University of Texas at Austin, 1978—2000,
/,. E. Gilbert 9271 . anno 1979, specimens
grown from seeds collected 13—16 July 1978
in Mexico, Veracruz, Estacidn Biologiea Los
Tuxtlas, (seed Gilbert 7828) (holotype, MEXU;
isotypes, MO. TEX. UPCB, XAL).
Distribution. Mexico: Oaxaca, Veracruz, 20-
350 m.
4. Passiflora eueidipaluilum S. Knapp & Mallet.
Ann. Missouri Bot. Card. 71: 1070, figs. 2, 3b.
1984. TYPE: Panama. Colon: Santa Rita
Ridge Road 7 km from the Transisthmian
Hwy., 9°22'!\, 79°40'W, 200 m. 21 May 1982,
S. Knapp & Schmalzel 5256 (holotype, MO;
isotypes, PM A, TEX).
Distribution. Costa Rica: Puntarenas. Panama:
Code, Colon, 200—1000 m.
Acknowledgments. Lawrence E. Gilbert at the
University of Texas at Austin first recognized the dis¬
tinctiveness of this group and proposed a higher tax¬
onomic placement. He recorded the main character¬
istics and biology ol the section, and he has given
the nama. flic other nomenclatural co-author is Jan
Meerman of Green Hills Butterfly Ranch in Belmo¬
pan, Belize. Jan was able to grow P. lancetillensis to
maturity and make specimens and studies of the
flowers, fruit, and natural history of the species. Jan’s
photographs, map, and observations have greatly
contributed to this work. Together we were able to
fashion a detailed description for this new species,
otherwise known from fragmentary material. Roy
Gereau generously translated the Latin diagnoses.
We thank Tom Wendt of TEX for his help w ith Lun-
dells collecting localities, and Rick Clinebell of MO
for his identification of the bee. We acknowledge
financial support to A. K. Hansen from an NSF Doc¬
toral Dissertation Improvement Grant. The curators
of BRH, DUKE, E, LL, TEX, and US kindly allowed
loans of material from their herbaria.
Literature Cited
Gusset. G. 1968. Les vrilles des Passiflorac6es. Bull. Soc.
Bot. France 115: 45-61.
Gilbert, I,. E. X J. M. MaeDougal. 2000. Lundellia 3: 1-
5, fig. 1, cover plate.
Killip, E. P. 1938. The American species of Passiflora-
eeae. Publ. Field Museum Nat. Hist.. Bot Ser. 19: 1-
613.
Mucking, F. P. 1992. Leaf Venation Patterns, Vol. 6. J.
Cramer, Berlin.
Knapp, S. & J. Mallet. 1984. Two new species of Passi-
flora (Passifloraceae) from Panama, with comments on
their natural history. Ann. Missouri Bot. Card. 71:
1068-1074.
Mallet. J. & J. T. Longino. 1982. Hostplant records and
descriptions of juvenile stages for two rare species of
Eueides (Nymphalidae). J. Lop. Soc. 36: 136-144.
Meerman, J. 1996. Vegetative key to the passionflowers of
Belize. Passiflora 6(3): 25—28, fig. 16. |Journal is the
newsletter of the Passiflora Society International, Co¬
conut Creek, Florida.]
-. 1999. Lepidoptera of Belize. 1. Butterflies, 2.
Emperor Moths and Hawk Moths. Tropical Lepidoptera
10 (supplement 1).
-& W. Sabido. 2001. Central American Ecosystems
Map: Belize. Programme lor Belize, Belize City. 2 vols.
Parker, T. A., Ill, B. K. Holst, L. II. Emmons & J. IL
Meyer. 1993. A Biological Assessment of the Columbia
River Forest Reserve, Toledo District, Belize. Conser¬
vation International, RAP Working Papers 3.
Schappert, P. 1998. Member’s plant list. Passiflora 8(3):
23.
Snow, N. & J. M. MaeDougal. 1993. New chromosome
reports in Passiflora (Passifloraceae). Syst. Bot. 18:
261-273.
Standley, P. C. 1931. Flora of the Lancetilla Valley, Hon¬
duras. Publ. Field Mus. Nat. Hist.. Bot. Ser. 10: 292-
293, pi. 52.
Ulmer, T. & B. Ulmer. 1997. Passionsblumen: Fine fasz-
inierende Gattung. Laupenmuhlen I truck, Witten.
Walter. K. S. & II. J. Gillett (editors). 1998. 1997 IUCN
Redlist of Threatened Plants. Compiled by the World
Conservation Monitoring Centre. IUCN—The W'orld
Conservation Union, Gland, Switzerland, and Cam¬
bridge. UK. Page Bros. (Norwich) Ltd.
Notes on Grasses (Poaceae) for the Flora oj China, II: Paniceae
Sylvia M. Phillips
Herbarium, Royal Botanic Gardens, Kew, Surrey TW9 3AB, United Kingdom.
s.phi 11 ips@kew.org.uk
Chen Shouliang
Herbarium, Jiangsu Institute of Botany, Academia Sinica, 1 Qianghuhuocun, Nanjing,
Jiangsu 210014. nasa@mail.cnbg.net
Abstract. This paper comprises nomenclatural
changes needed for the Flora of China account of
Poaceae tribe Paniceae. The new combinations
Digitaria fujianensis and Pseudoraphis sordida are
proposed for nomenclatural reasons. The varietal
combination Ottochloa nodosa var. micrantha is
validated and lectotvpified. Pseudoraphis balansae
is lectotvpified. and P. longipaleacea is placed in
synonymy. Urochloa cordata is reduced to synony¬
my under U. setigera. the three species of the ge¬
nus Pseudoraphis occurring in China are reviewed
and a key is provided. African specimens previ¬
ously considered nonspecific with Asian Urochloa
setigera are shown to belong to a different species.
Urochloa trichopodioides, which is lectotypified.
Key words: Africa, Asia, China, Digitaria, Ot¬
tochloa, Poaceae, Pseudoraphis, tribe Paniceae,
Urochloa.
During work on the tribe Paniceae for the grass
family account for the Flora of China, the following
nomenclatural changes were found to be necessary.
The opportunity is also taken here to briefly review
the species of Pseudoraphis occurring in China, as
there has been much confusion in the application
of names.
Digitaria Haller
Digitaria fujianensis (L Liou) S. M. Phillips & S.
L. Chen, comb. nov. Basionym: Leptoloma fuji¬
anensis L. Liou, Bot. Res. Academia Sinica 1:
41, f. I. 1983. TYPE: China. Fujian: Lian-cheng,
20 Sep. 1932. Ling Yung 3339 (holotype, PE).
This species belongs to the small group of Digi¬
taria that was formerly separated as Leptoloma be¬
cause of the diffuse paniculate inflorescence, lor ex¬
ample by Hitchcock (1935: 563). However, the
spikelets are typical of Digitaria, and the inflores¬
cence may merely be an extreme expression of a
loosening of the racemes seen elsewhere in this var¬
iable genus. Digitaria fujianensis is similar to D.
cognata (Schidtes) Pilger from the eastern United
States, but this has smaller (2.5—3 mm) spikelets and
shorter leaf blades. Digitaria tomentosa (Koenig)
llenrard from Thailand and southern India also has
an open paniculate inflorescence, but differs by its
broader leaf blades up to 8 mm wide, and especially
by the smaller (2.2—2.5 mm) spikelets with tiny
glumes.
Ottochloa Dandy
Ottochloa nodosa (Kunth) Dandy var. micrantha
(Balansa ex A. Camus) S. M. Phillips & S. L.
Chen, comb. nov. Basionym: Hemigymnia ar-
nottiana Stapf var. micrantha Balansa ex A.
Camus, in Lecomte, FI. Indo-Chine 7: 455.
1922. TYPE: Vietnam. Tonkin. Lankok valley,
11 Oct. 1887, H. Balansa 1609 (lectotype,
designated here. K).
This is a small-spiculate variant of Ottochloa no¬
dosa, a rambling grass widespread in shady places
in the Old World tropics. The paniculate inflores¬
cence can he variable, with the spikelets usually ar¬
ranged in bunches or small dense racemelets along
the primary branches, but sometimes looser with less
obvious spikelet clusters. The spikelets are normally
about 3 mm long, but some specimens from southern
China (Guangdong and Hainan) and Vietnam have
smaller (2—2.5 mm) spikelets that are always borne
in neat appressed racemelets. This variant has long
been recognized as meriting separate status, hut the
name has never been validly published in Ottochloa.
The genus Hemigymnia Stapf (1920) is illegiti¬
mate, as it is a later homonym of Hemigymnia Grif¬
fith (1842), but the epithet micrantha is validly
published and available.
P. C. Keng (1976: 160) was the first to transfer
the taxon to Ottochloa as 0. nodosa var. micrantha
(Balansa ex A. Camus) P. C. Keng. but tlie combi¬
nation is invalid because it lacks the basionym ref-
Novon 13: 467-470. 2003.
468
Novon
erenee (Art. 33.3 of the ICBN; Greuter et al., 2000).
T. D. Zhuang also placed it in Ottochloa (1990: 222).
attributing the combination to I’. C. Keng, but again
the combination is invalid because it lacks a correct
basionym author and reference (Art. 33.3).
This taxon was first recognized by llalansa
(1890: 142) as Panicum nodosum Kunth var. mi-
crantha, a nomen nudum based on three of his col¬
lections ( Balansa 480, 1609, 1610). These collec¬
tions are all represented in the Kew herbarium, and
the best is selected here as lectotype.
PSEUDORA PHIS Gr IKFITH
Pseudoraphis is a small genus of closely related
species occurring from India to China anti Japan and
in southeast Asia anti Australia. There is no overall
account of the genus, and specimens have frequently
been assigned to the most widespread species, P.
spinescens (R. Brown) Vickery, both in the herbarium
and literature, which in fact belong to other species.
One of these species ol Pseudoraphis is known under
an incorrect name. The species of Pseudoraphis oc¬
curring in China are set out below, with notes on
differences from P spinescens. The following key lists
the main distinguishing characters.
Key to the Species of Pseudoraphis in China
la. Inflorescence open, lanceolate to ovate; racemes
spreading, 2—3-spiculate . P. brunoniana
Ih. Inflorescence contracted, linear or oblong; ra¬
cemes erect, usually 1-spiculate.
2a. Spikelets numerous (more than 30); upper
glume longer than lower lemma, narrowly
acuminate; lower lemma 7-veined; 2 sta¬
mens in lower floret . /\ sordida
2b. Spikelets 10 or less; upper glume slightly
shorter than lower lemma, acute; lower lem¬
ma 13-veined; 3 stamens in lower floret . .
. P. balansae
Pscudorufdiis brunoniana (Wallieh & Griffith ex
Griffith) Griffith ex Pilger, Not. But. Cart. Ber¬
lin 10; 210. 1928. Basionym: Panicum bru-
nonianum Wallieh & Griffith ex Griffith, J.
Asiat. Soc. Bengal 5: 574. 1836. Chamaera-
phis spinescens (R. Brown) Poiret var. brunon¬
iana (Wallieh & Griffith ex Griffith) J. I).
Hooker, FI. Brit. bid. 7: 62. 1896. Chamaera-
phis brunoniana (Wallieh & Griffith ex Griffith)
A. Camus, in Lecomte, FI. Gen. Indo-Chine 7:
479. 1922. TYPE: Bangladesh (“Bengal").
Sylhet district, near Goalnuyar, 28 Sep. 1835.
W. Griffith (locality uncertain).
Distribution. Northeastern India, Myanmar,
Thailand, southeastern China, Taiwan. Vietnam, the
Philippines.
This species was originally described as Pani¬
cum brunonianum by Griffith (1836), but in a later
publication (Griffith, 1851a, Notulae: 29) he gave
it a new name in Panicum, P. intermedium, based
on the same specimen. Panicum intermedium Grif¬
fith is therefore an illegitimate superfluous name
according to the ICBN (Art. 52.1: Greuter et ah.
2000). In the 1851 Notulae the combination Pseu¬
doraphis brunoniana is also listed, apparently as a
synonym or alternative name for Panicum inter¬
medium, and is therefore invalid (Art. 34.1). To
complicate matters further, the illustrations pub¬
lished to accompany the Notulae (Griffith, 1851b.
leones, t. 145, fig. 1), revert to the earlier valid
name Panicum brunonianum. Both the Griffith No¬
tulae and leones of 1851 were arranged and pub¬
lished posthumously by John M'Clelland. which
probably accounts for the confusion. The combi¬
nation Pseudoraphis brunoniana was first made val¬
idly by Pilger in 1928.
The name Holcus nutans Roxburgh ex J. D.
Hooker (1896: 62) is invalid, as it was published
as a synonym of Chamaeraphis spinescens var. bru¬
noniana ( Art. 34.1; Greuter et ah. 2000). It is based
on a specimen from “Lower Bengal" in Herb. Rox¬
burgh (K).
There is a Griffith specimen in the Kew herbar¬
ium. collected on 28 September 1835, but with the
locality “Jheel of the Magna." The inflorescence is
young with almost erect branches, and therefore
does not correspond to the ovate inflorescence de¬
scribed in the protologue. The original drawing by
Griffith for his leones (1851b, t. 145) is in the ar¬
chives ol the Kew library. This depicts an ovate
inflorescence, but bears the date “Sept. 17.37." No
lectotypification is made here because of the pos¬
sibility that the holotype from Goalnuyar may still
be extant in Calcutta.
Pseudoraphis brunoniana was mistakenly stated
by Bor (1960: 353) to occur in Ceylon. The grass
described under this name from Ceylon by Lazarides
(1994: 383) is a specimen of P. spinescens. In fact,
specimens from throughout the range of this species
have usually been assigned to P. spinescens. It was
evident, when examining specimens from China, that
they differed in several important respects from ma¬
terial from Australia (including the type of P. spi-
nescens) and also most material from India.
Bor distinguished Pseudoraphis spinescens and P.
brunoniana on spikelet size, and this has been the
cause of confusion. Both species are variable in this
character, and specimens used by Bor from north¬
east India (Assam, Bengal) and adjoining parts of
Myanmar have exceptionally large spikelets. Pseu¬
doraphis spinescens has racemes with 5 to 10 or
Volume 13, Number 4
2003
Phillips & Chen
Notes on Chinese Grasses
469
more spikelets lying more or less end-to-end,
whereas racemes in P. brunoniana bear only 2 or
3 (occasionally 1) distant spikelets, usually with a
longer terminal bristle, resulting in an immediate
difference in facies. The culm nodes in P. spines¬
cens are sericeous, appearing as a shiny white band
of appressed silky hairs, whereas the nodes in P.
brunoniana are merely pubescent. There is also a
subtle difference in spikelet shape, as P. spinescens
has a caudate upper glume much exceeding the
lower lemma, while in P. brunoniana the glume is
narrowly acuminate and extends beyond the lower
lemma to a lesser extent.
The few specimens seen from China are cited be¬
low.
CHINA. Guangdong: Guangzhou [Canton], Apr. 1878,
T. Sampson 367 (K), May 1864, Sampson in Herb. Hance
11077 (K), May 1884, T. Sampson s.n. (K).
Pseudoraphis sordida (Thwaites) S. M. Phillips
& S. L. Chen, comb. nov. Basionym: Panicum
sordidum Thwaites, Enum. PI. Zeyl. 443.
1864. TYPE: Sri Lanka. G. Thwaites C.P. 3857
(isotype, K).
Chamaeraphis spinescens var. depauperata Nees ex J. D.
Hooker, FI. Brit. Ind. 7: 62. 1896. Pseudoraphis de¬
pauperata (Nees ex J. D. Hooker) Keng, Sinensia 11:
43. 1940. Chamaeraphis squarrosa (L. f.) Merrill var.
depauperata (Nees ex J. D. Hooker) Masamune,
Trans. Nat. Hist. Soc. Formosa 30: 18. 1940. Pseu¬
doraphis squarrosa (L. f.) Chase var. depauperata
(Nees ex J. I). Hooker) Hara, J. Jap. Bot. 17: 398.
1941. Pseudoraphis spinescens var. depauperata
(Nees ex J. I). Hooker) Bor, Grasses of Burma, Cey¬
lon, India and Pakistan: 354. 1960. TYPE: India. H.
Wight 1654 (holotype, K).
Pseudoraphis ukishiba Ohwi, Act. Phytotax. Geobot. 10:
273. 1941. TYPE: Japan. Hondo, Shimamura, 25
Aug. 1931, Hashimoto s.n. (holotype, KYO).
Distribution. India, Ceylon, China, Japan, Ko¬
rea.
Bor (1960: 352) aptly described this grass as
having a u Pennisetum- like” inflorescence. The con¬
tracted inflorescence is indeed quite different from
the open inflorescence of P. spinescens, of which it
was made a variety by Bor and others. Both Keng
and Ohwi recognized its separate specific status,
hut unfortunately overlooked the epithet from Cey¬
lon. l’lie habit is more slender than that of P. spi¬
nescens, and the internodes are frequently purple-
tinged. It also differs by its clearly ciliate ligule and
the presence of only two stamens. This species was
omitted from the recent account of the genus in
Ceylon (Lazarides, 1994).
Steudel (1854) made it quite clear that he was
placing this taxon as a variety under Panicum as-
perum Wight, citing the manuscript name Chama¬
eraphis depauperata Nees as basionym, but he did
not definitely make the varietal combination (Art.
33.1; Greuter et ah, 2000). The name C. depauper¬
ata was also not validly published as a species by
Steudel, as it is included under Panicum asperum
(itself an illegitimate homonym). The earliest valid
name at specific level for this grass is Panicum
sordidum Thwaites. The necessary new combination
under Pseudoraphis is made here.
Pseudoraphis halansae Henrard, Blumea, Suppl.
1: 230, t. 17. 1937. TYPE: Vietnam. Annam,
25 Oct. 1886, Balansa in Herb. Lugd. Bat.
908. 85-1414 (lectotype, designated here, L).
Pseudoraphis longipaleacea Chia, El. Hainan. 4: 442, 540,
f. 1232. 1977. Syn. nov. TYPE: China. Hainan I.:
Dongfang, 12 Sep. 1936, Liou 27870 (holotype,
SCBI not seen).
Distribution. Vietnam, Thailand, China (Hai¬
nan).
This species appears to be of restricted distri¬
bution and is known from very few collections. It
is distinctive in the genus because of its merely
acute spikelets lacking the drawn-out narrow apex
on the upper glume found in most species. The
short subacute leaves, white-membranous truncate
ligule, and few-spiculate inflorescence are also
characteristic.
There are three sheets ol this species collected
by Balansa in the Leiden herbarium, which have
been annotated by Henrard as types, and seven oth¬
er Balansa sheets, all of which are probably dupli¬
cates of the same collection. None bear collecting
numbers, only herbarium sheet numbers. Henrard
in his protologue stated there are “specimina multa
legit Balansa.” The sheet selected here as lectotype
is chosen because it corresponds to a sheet number
given by Henrard. The specimen itself has not been
seen, but its image has been checked on the Inter¬
net (http://www.nationaalherbarium.nl).
A photocopy of the type specimen ol Pseudora¬
phis longipaleacea is available at Kew and this, to¬
gether with the illustration in the protologue, clear¬
ly shows that this species is a good match with P.
balansae.
Urochloa P. Beauvois
When writing the account of Urochloa for the Flo¬
ra of Tropical Africa, Stapf (1920: 598) included
some specimens from East Africa under Panicum
setigerum, at the same time transferring the species
to Urochloa. This species was known from India and
Sri Lanka, where it is widespread, and even then
470
Novon
Stapl pointed out that the African specimens differed
hy their glabrous spikelets. He also wrote that the
type was “stated to have been collected in China by
Bladh; but there is no other record of its occurrence
iu that country," perhaps implying that he consid¬
ered the collecting locality incorrect. However, U.
setigera is now known from Myanmar and Thailand,
and U. cordata is a good match, confirming the pres¬
ence of this species in tropical southern China.
Urochloa cordata is therefore considered to be a new
synonym of U. setigera (see below). The type of
Urochloa setigera Retzius, collected by Bladh in the
18th century, is in the herbarium at Lund. Although
there has been no possibility to see the type, there
is ample material of U. setigera from Asia at Kew
for comparative purposes.
Stapfs treatment has been followed in recent
Floras, but study of the species for the Flora of
China has revealed a number of other differences,
enumerated below. It now seems clear that two spe¬
cies should be recognized, as follows:
Urochloa setigera (Retzius) Stapf, in Brain, FI.
Prop. Afr. 9: 598. 1920. Basionym: Panicum
setigerum Retzius, Obs. Bot. 4: 15. 1786. Bra-
chiaria setigera (Retzius) C. E. Hubbard, in
Hooker’s Icon. Bl. 34: t. 3363. 1938. TYPE:
China (Canton). P. J. Bladh s.n. (holotype, I.D
not seen).
Panicum affine Poiret, in Lamarck, Encycl., Suppl. 4:
273. 1816. TYPE: “Indes orientales," Herb. Desvaux
(holotype. P not seen).
Urochloa cordata Y. L. Keng ex S. L. Chen & Y. X. Jin,
Acta Phytotax. Sin. 22: 472. 1984. Syn. nov. TYPE:
China. Hainan: C. Wang 33805 (holotype, SYS).
Leaf blades 8—15 cm long, HI—24 mm wide, mar¬
gins pectinate-setose, at least to the middle; nodes
bearded; inflorescence axis 6-13 cm long bearing
6 to 12 racemes; spikelets pubescent, herbaceous;
fertile floret with a very brief mucro 0.1 mm long.
Distribution. India, Ceylon, Myanmar, Thai¬
land, southern China.
I Jrochloa trichopodinitlrs (Mez & Schumacher)
S. M. Phillips & S. L. Chen. comb. nov. Bas¬
ionym: Panicum trichopodioides Mez & Schu¬
macher, Notizbl. Bot. Cart. Berlin-Dahlem 7:
60. 1917. TYBE: Tanzania. Amboni, C. Holst
2844 (lectotype, designated here, K).
Leaf blades 5.5—10 cm long, 10-18 mm wide,
margins scabrid (rarely a few setae at base); nodes
glabrous or pubescent; inflorescence axis 1—7 cm
long bearing 1 to 7 racemes; spikelets glabrous,
thinly cartilaginous, dully shining; fertile floret with
a pronounced mucro 0.4—0.6 mm long.
Distribution. Kenya, Tanzania, southern Ethio¬
pia, eastern Congo.
Urochloa trichopodioides was described based on
two syntype specimens from Tanzania: Amboni, C.
Holst 2844 (syntype, B; isosyntype, K), and Buko-
ba. Herb. Amani 5333 (syntype, B). The species is
lectotypified here on the Holst specimen at Kew,
which is a good, fully representative collection.
I’lie most important differences between these
two species are summarized in the following cou¬
plet.
Kky to Distinguish Urochloa setigera and
U. TRICHOPODIOIDES
la. Spikelets pubescent; fertile floret with tiny mucro
ca. 0.1 mm; racemes 6 to 12; margins of leaf
blades pectinate-setose . . Urochloa setigera (Asia)
lb. Spikelets glabrous; fertile floret with pronounced
mucro 0.4—0.6 mm; racemes 2 to 6; margins of
leaf blades scabrid .
. Urochloa trichopodioides (Africa)
Acknowledgment. We thank J. F. Veldkamp of
the Leiden Herbarium for providing details of the
type material of Pseudoraphis balansae Henrard.
Literature Cited
Balansa, 8. 1890. Catalogue des Gramin£es de l’lndo-
Chine franyaise. J. Bot. (Morot) 4: 135—145.
Bor, N. L. 1960. The Grasses of Burma, Ceylon, India and
Pakistan. Pergamon, Oxford.
Greuter, W., J. McNeill. F. II. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. C. Silva,
.1. F. Skog, P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Griffith, W. 1836. Description of some grasses which form
part of llie vegetation in the Jheels district of Sylhet. .1.
Asia!. Soc. Bengal 5: 570-575.
-. 1843 [1842], Remarks on a few plants from cen¬
tral India. Calcutta J. Nat. Hist. 3: 361—363.
-. 1851a. Notulae ad plantas asiaticas, Vol. 3.
Charles Serrao, Calcutta.
-. 1851b. leones plantarum asiaticarum, Vol. 3.
Bishop’s College Press, Calcutta.
Hitchcock, A. S. 1935. Manual of the Grasses of the Unit¬
ed Slates. U.S. Government, Washington, D.C.
Hooker, .1. I). 1897 11896]. Flora of British India, Vol. 7.
L. Reeve, Ashford, Kent.
Keng, P C. 1976. Iconographia Cormophytorum Sinico-
rum, Vol. 5. Science Press.
Lazarides, M. 1994. Pseudoraphis. Pp. 383-385 in M. I).
Dassanayake, F. R. Fosberg & W. D. Clayton (editors),
A Revised Handbook to the Flora of Ceylon,Vol. 8. Am¬
erind. New Delhi.
Stapf. (). 1920. Urochloa. Pp. 586—604 in D. Prain (edi¬
tor), Flora of Tropical Africa, Vol. 9. Gramineae. Reeve,
Ashford.
Steudel, F. G. 1855 [18541. Synopsis Plantarum Gluma-
cearum. I. Gramineae. Metzler, Stuttgart.
Zhuang, T. I). 1990. Ottochloa. Pp. 221-224 in Flora Rei-
publicae Popularis Sinicae, Vol. 10(1). Science Press,
Beijing.
Validation of the Name Astragalus yumenensis (Fabaceae), a
Species Endemic to China
Dieter Podlech
lnstitut fur Systematische Botanik der Universitat Miinchen, Menzinger StraBe 67,
D-80638 Munchen, Germany, podlech@botanik.biologie.uni-muenchen.de
Nicholas J. Turland
Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166, U.S.A.
nicholas.turland@mobot.org
Xu Langran
North-western Institute of Botany, Yangling, Xianyang, Shaanxi 712100,
People’s Republic of China
ABSTRACT. While preparing the account of As¬
tragalus L. (Fabaceae) for the Flora of China, Vol¬
ume 10, it was noticed that one species, A. yume¬
nensis S. B. Ho. described from Gansu Province in
1983, was invalidly named because two types (one
flowering and one fruiting) were designated. The
name is validated here, with the fruiting specimen
designated as the holotype.
Key words: Astragalus, China, Fabaceae.
Astragalus yumenensis will be accepted as a spe¬
cies endemic to Gansu Province, China, by Xu and
Podlech in their forthcoming account of Astragalus
in Flora of China, Volume 10 (in prep.). Astragalus
yumenensis was accepted, without comment on its
validity, by Ho (1993: 316) in his account of Chi¬
nese Astragalus subg. Cercidothrix Bunge. It is
most similar to A. scoparius Schrenk (subsp. sco¬
parius; see Ghahremani-nejad, 2001), from north¬
ern Xinjiang Autonomous Region and Kazakhstan,
and A. sogotensis Lipsky, from the same regions as
well as Kyrgyzstan. All three species belong to A.
subg. Cercidothrix, characterized by the presence
of medifixed or forked (vs. simple and basifixed)
trichomes and a non-inflated fruiting calyx that
does not fully envelop the legume. This subgenus
has nearly 800 species distributed in the Old World
(Ghahreman et ah, 2002), from the Mediterranean
region to central Asia, with about 70 species in
China. Astragalus scoparius, A. sogotensis, and A.
yumenensis share the following characters: stems
usually less than 5 cm or plants acaulescent; stip¬
ules free or connate at base; leaves imparipinnate,
raehis not hardening; peduncles equaling or longer
than leaves; racemes lax; calyx not bracteolate; co¬
rolla with standard oblong or narrowly so, wings
emarginate or 2-lobed at apex; legume non-inflated.
They may be distinguished with the following key:
Key to Astragalus scoparius A. sogotensis, and
A. YUMENENSIS
la. Calyx teeth subulate, Vs-Vi as long as tube . . .
. A. yumenensis
lb. Calyx teeth subtriangular, '/ 6 —14 as long as tube.
2a. Legume with both black and white tri¬
chomes; leaflets in 3 to 6 pairs . . A. scoparius
2b. Legume with only white trichomes; leaflets
in 2 or 3 (or 4) pairs. A. sogotensis
A nomenclatural problem exists with Astragalus
yumenensis. Two specimens, representing two gath¬
erings, were simultaneously designated as types.
Both are from Gansu: one flowering, “Jiuquan ad
Yumen,” 19 May 1957, K. J. Kuan 421 (PE), and
one fruiting, “Sunan Yugurzu Zizhi Xian, daheba-
tangeltan, in declivitate sicco, alt. 2540 m." 2 Aug.
1967, Hexi Expedition 126 (PE). The name is there¬
fore invalid under Articles 8.1 and 37.1 of the
ICBN (Greuter et ah, 2000). The name is validated
here by reference to the previously and effectively
published Latin diagnosis and full Latin description
by Ho (1983: 65-66) and by designating the fruit¬
ing specimen, Hexi Expedition 126, as the holotype.
The fruiting specimen is preferable to the flowering
one because it is the more useful for diagnostic
purposes.
Novon 13: 471-472. 2003.
472
Novon
Astragalus yiinienensis S. 15. Ilo ex Podlech & I,.
H. Xu, sp. nov. Astragalus yumenensis S. B.
Ho, Bull. Bot. lies., Harbin 3(1): 65—66, pi.
18. 1983, nom. inval. TYPE: China. Gansu:
Sunan Yugurzu Zizhixian, “Dahebatangeltan
[sic!], in deelivitate sicco,” 2540 rn, 2 Aug.
1967 (fr), Hexi Expedition 126 (holotype, PE,
photograph seen).
Plants 15-30 cm tall, shortly caulescent; caudex
strong, with a many-headed root crown; stems sev¬
eral, short, 2—10 cm, densely covered with medi-
fixed, appressed, white trichomes 0.6—1 mm.
Leaves 5-12 cm; stipules 4—5 mm, triangular,
shortly adnate to petiole, with white, appressed tri¬
chomes; petiole 2-5 cm, together with rac his with
trichomes like stem; leaflets in 2 or 3 pairs, linear,
10—25 X 1—3 mm, on both surfaces sparsely to
rather densely covered with appressed trichomes
1 — 1.5 mm. Peduncle 7—15 cm, with trichomes like
stem but toward raceme with some black trichomes
intermixed; raceme rather dense, 3^4 cm; bracts
narrowly triangular, 2-3 mm, membranous, loosely
with predominantly black trichomes; pedicels 1-2
mm, with black trichomes. Calyx narrowly tubular,
obliquely cut at mouth, 8-12 mm, rather densely
covered with appressed black (predominantly) and
white trichomes (0.3—)0.5—0.8 mm; teeth subulate,
2-4 mm; corolla purple-red; standard 17-20 mm,
limb narrowly obovate, 6—7 mm wide, narrowed into
a short claw at base, apex retuse; wings 14—18 mm,
claw 7—9 mm, auricle rounded, ca. 0.2 mm, limb
narrowly oblong, 7-8 X 2-2.5 mm, apex distinctly
incised; keel 10—15 mm, claw 6—9 mm, auricle mi¬
nute, limb obliquely obovate, distally with broadly
curved lower edge and slightly concave upper edge.
5.5—6 X 2—2.5 mm, apex acute; stamen tube sub¬
truncate at mouth; ovary subsessile, linear, covered
with appressed while and black trichomes; style
glabrous. Legume sessile, 2-valved, straight to
slightly curved, 10-15 mm, ca. 2 mm wide and
high, grooved abaxially, slightly keeled adaxially,
apex shortly acuminate; valves with appressed
white and black or predominantly black trichomes.
Seeds 3 or 4 in each valve.
Phenology. Flowering in May and June, fruit
ripening from June to August.
Habitat. Dry slopes in steppes, gullies of allu¬
vial fans in deserts; 2000-3000 m.
Distribution. Endemic to northwestern Gansu
Province, China.
Vernacular name. Yu men huang qi.
Paratypes. CHINA. Gansu: Aksay Kazakzu Zizhixi¬
an, 1950 m. 14 May 1996, /.. H. Xu 96-188 (MSB); “Jiu-
quan ad Yumen," 19 May 1957 (fl), K. ./. Kuan 421 (PE).
I .iterature Cited
Ghahreman, A., A.-A. Maassoumi & F. Ghahremani-ne-
jad. 2002. Astragalus tuyehensis (Fabaceae), a new spe¬
cies from Iran. Novon 12: 47—19.
Clialir emani-nejad, F. 2001. Notes on Astragalus sect.
Corethrum (Fabaceae), including a new combination.
Ann. Bot. Fenn. 38: 47^49.
Greater, W., J. McNeill. F. K. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, I). H. Nicolson, P. G. Silva,
J. E. Skog, P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Kegnum Veg. 138.
Ho, S. B. 1983. Prascursores [sic!; Praecursores] Horae
Astragalorum sinensium (V). Bull. Bot. lies., Harbin
3(1): 39-91.
-. 1993. Astragalus subgen. Cercidothrix Bunge.
Pp. 252—326 in K. T. Fu (editor). Flora Reipublicae
Popularis Sinicae, 42(1). Science Press, Beijing.
A New Species of Bashania (Poaceae: Bambusoideae) from Ml.
Qinling, Shaanxi, China
Ren Yi
College of Life Science, Shaanxi Normal University, Xi’an 710062, People’s Republic of
China, renyi@snnu.edu.cn
Li Yun
School of Life Science, Northwest University, Xi'an 710069, People’s Republic of China
Dang Gao-di
Foping National Nature Reserve Management Bureau, Foping, Shaanxi 723400, People’s
Republic of China
ABSTRACT. The new species Bashania aristata of
Bambusoideae (Poaceae) from the giant pandas
habitat of Mt. Qinling, Shaanxi, China, is described
and illustrated. The new species differs from B.far-
gesii in its fully developed culm sheath auricles and
aristate lemmas anti from B. qingchenshanensis in
its fully developed culm sheath auricles and in its
lack of auricle leaf sheaths. Its relationship to the
other species of Bashania is discussed along with
the status of this genus.
Key words: Bambusoideae, Bashania, China,
Poaceae.
Bashania Keng f. & T. P. Yi was described (Keng
& Yi, 1982) for Arundinaria fargesii E. G. Carnes
because Arundinaria Michaux is endemic to North
America, and they differ from each other both in
the flowers and the number of primary branches per
culm node. It is readily distinguished from the re¬
lated Chinese Gelidocalamus T. H. Wen by longer
branches growing from the culm, and the presence
of secondary branches, and more than two leaves
on the terminal twigs. Gelidocalamus has short
branches from the culm, without secondary branch¬
es, and with one or two leaves on the twigs.
The genus Bashania consists of seven species:
B. fargesii, B. qingchengshanensis (Keng & Yi,
1982, 1996), B. faberi (Yi, 1993) (= B. fangiana,
Wen, 1985; Keng & Yi, 1996), B. spanostachya (Yi,
1989; Keng & Yi, 1996), B. baoxingensis (Yi,
2000), B. auctiaurita (Yi, 1986), and the new spe¬
cies B. aristata, most of them endemic to Sichuan
and adjacent Shaanxi, Gansu, Hubei. Hunan, and
Yunnan Provinces of China.
The new species, Bashania aristata, is common
in Foping National Nature Reserve and can also be
found in Changqing National Nature Reserve and
other areas of Shaanxi Province. These reserves are
the main distribution of the giant panda on Mt.
Qinling. When we surveyed the plants in the hab¬
itat of the giant panda in Mt. Qinling in 1999, we
found that some individuals of Bashania were flow¬
ering but some were not. The flowering individuals
and non-flowering individuals were distributed in
staggered plots in the area named Sanguanmiao in
Foping National Nature Reserve. This phenological
pattern was abnormal according to the common
know ledge of the flowering behavior of bamboo, and
it was assumed that there was only one species, B.
fargesii, distributed in the reserve. After compari¬
son of the flowering ami non-flowering individuals,
we found that the flowering individuals differed in
having aristate lemmas and auricular culm sheaths.
Further survey indicated that the same phenome¬
non also existed in another giant panda reserve,
Changqing National Nature Reserve located south
of Foping National Nature Reserve, as well as other
areas in Mt. Qinling. The staggered distribution of
B. aristata and B. fargesii correlates with the ab¬
sence of any report of wholesale bamboo death after
flowering over a large area in the forest of Bashania
in the giant panda’s habitat on Mt. Qinling.
Bashania aristata Y. Ren, Y. Li & G. D. Dang, sp.
nov. TYPE: China. Shaanxi: Foping County,
Sanguanmiao, Chaoyangpo, 33°39'16.875N,
107°47' 11.25E, 1600 m, 28 Apr. 1999, Ren Yi
906 (holotype, WNU; isotype, MO). Figure 1.
Species differt a B. fargesii vaginis auriculatis, lemmis
aristatis, a B. qingchenshanensi vaginis auriculatis, auri-
culis foliorum carentibus.
Rhizome leptomorph, monopodial, internodes
Novon 13: 473-476. 2003.
474
Novon
Figure 1. Bashania aristata Y. Ren, Y. Li & G. I). Dang. — A. Rhizome and culm base; the transverse rhizome and
the left branches show the monopodial growth pattern and the right branches show the sympodial growth pattern. —
B. Part of culm. —C. New shoot. —D. Upper part of a culm leaf, showing the sheath auricles and blade. —E. Base
of a leaf. —F. Inflorescence. —G. Spikelet. —H. Lemma (left) and palea (right). —I. Lodicules. —J. Stamens. —K.
Pistil. Seale bars: A—C = 5 cm; D, E, G = 1 cm; F. H, J = 5 mm; I, K = 1 mm. Drawn by Guo Mu-seng from the
holotype, Ren Yi 906 (WNU), at Sanguanmiao.
Volume 13, Number 4
2003
Ren et al.
Bashania aristata (Poaceae)
475
(1)3—5 cm long, 5—20 mm diam., hollow. Culms
erect except the tips nodding, 2—8(—13) m in
height, 2—4(—6.5) cm diam., deep green and with
white powder when young, (lavescenl when old. In-
temodes 25—40 cm long, slightly flattened above
branching complement. Walls 4-8 mm thick; culm
nodes slightly bulging, nodal line salient with rem¬
nant of sheath. Bud I per node, flat; nodal region
6—14 mm long, glabrous. Primary branches 3, sec¬
ondary branches many, but more slender than pri¬
mary branches. Culm sheath leathery, shorter than
the internode, green when fresh and flavescent
when dry, densely covered with black to dark brown
setae abaxially, dents or papillae left when hairs
fall off; inner ligule 2—4 mm tall, with small setae
on it, denticulate at apex; auricle fully developed,
half-moon-shaped, with erect hairs on the edge;
blade lanceolate, erect or curved, not easily cadu¬
cous, green when young with caducous cilia at the
base of upper surface. Leaves 4 to 6 on each ulti¬
mate branch, glabrous adaxially anti sparsely ve-
lutinate abaxially when young, usually 7-17 X 1—
2.3 cm, with 6 pairs of long veins and compact
cross veins; sheath 5—10 cm long; inner ligules
1.5—4 mm tall, tomentulose and with irregular ser-
ra; auricle absent; petiole usually 0.7—0.8 cm long.
Inflorescence a panicle, about 9 cm long, 3—4 cm
wide, eventually exserted; the major axis and
branches of the inflorescence with white microse-
tae. Spikelets slender cylindrical, dark brown when
mature, 2—3 cm long, 3 mm diam., with 4 to 7
florets in each; rachilla internodes 2—3.5 mm long,
flat, with white microsetae. Glumes ovate-lanceo¬
late; outer glume 3—6 mm long, 1—3-veined, the
inner glume 6-8 mm long, 5- to 7-nerved. Lemmas
ovate to lanceolate, with white microsetae at the
base, 12—13 mm long, with 7 long veins and sparse
cross veins, obviously aristate. Paleas 9—10 mm
long, with 2 ridges on the back and bifid at the
apex. Lodicules 3. ciliate at the edge, unequal,
veined at the base. Stamens 3, with anthers 4—5
mm long. Ovary ovoid; stigmas 3, featherv, about 2
mm long. Caryopsis unknown.
Habitat and distribution. The new species lives
on slopes with brown-yellow forest soil under de¬
ciduous broadleaved forest anti in temperate conif¬
erous broadleaved mixed forest ranges, from 1100
to 1600 m. The soil is thick, slightly wet, and acid¬
ic, with thin humus and leaf litter.
The community of the new species is simple. The
arboreous layer consists of species of Quercus or
species of Quercus and Pinus tabulaeforrnis. The
shrub layer consists mainly of the new species and
rarely species of Acer and Viburnum. The herb layer
is weakly developed and consists mainly of species
of Poaceae and Cyperaceae.
The new species is distributed in Foping,
Yangxian, and Zhenba Counties of Shaanxi Prov¬
ince, China. These counties are located on the mid¬
dle part of the south slope of Mt. Qinling, which is
the main distribution area of the giant panda on
Mt. Qinling.
Bashania aristata is readily distinguished from
the six other species of the genus. It is obviously
different from B. fangiana and B. spanostachya by
its longer leaves (more than 10 cm long), abaxially
sparsely velutinous foliage leaves when young, with
brown or dark brown setae on the back of the culm
sheath and large culm sheath auricles. Bashania
aristata is closer to B. fargesii, B. qingchenshanen-
sis . B. baoxingensis , and B. auctiaurita, but is also
readily distinguished from B. fargesii by its large
culm sheath auricles and the aristate lemmas, and
from B. qingchenshanensis and B. baoxingensis by
its large culm sheath auricles and its lack of leaf
sheath auricles. Bashania aristata is distinguished
from B. auctiaurita by its larger stature and less
falcate auricles.
Paratypes. CHINA. Shaanxi: Foping County, Yueba,
Baimagou, 28 Apr. 2000, Li Yun 101 (WNU); Foping
County, around Sanguanmiao, 18 Oct. 2001, Li Yun 111
(MO); Zhengba County, Tang Jian-weng 94 (SIFS).
Key to Chinese Species of Bashania
la. Leaves longer than 10 cm; culm sheath with
brown or dark brown setae on the back.
2a. Leaves sparsely velutinous on lower surface
when young, leaf sheath without auricles;
culm sheath without basal hair ring.
3a. Culm sheaths without auricles; lemmas
not aristate .
. B. fargesii (E. G. Camus) Keng f. &
T. P. Yi
3b. Culm sheaths with auricles; lemmas
aristate.
. . B. aristata Y. Ren. Y. Li & G. 1). Dang
2b. Leaves glabrous, with auricles; culm sheath
with basal hair ring.
4a. Culm sheath without auricles.
5a. Internodes of rhizome and the base
of culm almost solid .
.... B. qingchengshanensis Keng 1. A
T. P. Y i
5b. Internodes of rhizome and the base
of culm hollow.
. B. baoxingensis T. P. Yi
4b. Sheath on upper part of culm with au¬
ricles . B. auctiaurita T. P. Yi
lb. Leaves shorter than 10 cm; culm sheaths gla¬
brous or with yellow-gray setae on the back.
6a. Rhizome with 2 to 5 roots at the node; ra¬
ceme with 3 to 5 spikelets; spikelets 2.5—
4.5 cm long; lemmas 9—14 mm long; paleas
476
Novon
7-12 mm long; anther 5—6 mm long ....
. B. faberi (Rendle) T. P. Yi
6b. Rhizome with 4 to 6 roots at the node; ra¬
ceme with 2 or 3 spikelets; spikelets 1.8—3
cm long; lemmas 7—10 mm long; paleas 5—
6.5 mm long; anthers 3.5—4.5 mm long . . .
. B. spanostachya T. I*. Yi (including
B. fansipanensis)
Species Exceeded in the Key to Chinese
Species
Bashania victorialis (Keng f.) T. P. Yi was first
described as Indocalamus victorialis by Keng f.
(1951), but Yi (1993) removed this species to Bas-
hania according to die specimens collected from
the area where the type was collected. According
to Keng 1. (1951), there are one or rarely three pri¬
mary branches on the culm node. This species
needs further study.
Bashania abietina T. P. Yi & L. Yang was de¬
scribed based on the specimens without flowers (Yi
& Yang, 1998). According to the description, this
species has one branch on the culm node, which is
similar to the species of Indocalamus rather than
Bashania. This species needs further study.
Acknowledgments. This research was supported
by the State Key Basic Research and Development
Plan of China (No. G200004680001), the Founda¬
tion from the Memphis Zoo, U.S.A., and the Foun¬
dation from the Educational Department of Shaanxi
Province.
Literature Cited
Keng, P. C. 1951. A new' species of Indocalamus from
Szechuan. Acta Phytotax. Sin. 1(1): 121 — 122.
- & T. P. Yi. 1982. Bashania, a new bamboo genus
from western China. ,1. Nanjing Univ., Nat. Sci. Ed.
1982(3): 722—732. |ln Chinese with English summary.]
-& -. 1996. Bashania. In: P. C. Keng & Z.
P. Wang (editors), Flora Reipublicae Popularis Sinicae
9(1): 612—620. Science Press, Beijing. |ln Chinese.]
Wen, T. H. 1985. New taxa of Bamboo from China (II). .1.
Bamboo Res. 4(2): 9—19.
Yi, T. P 1986. New material on Chinese Bambusoideae.
J. Bamboo Res. 6(4): 25—30.
-. 1989. Two new species of Bamboo from south¬
western Sichuan. Acta Rot. Yunnan. I 1(1): 35—38.
-. 1993. A new species and five new combinations
of Bambusoideae. J. Bamboo Res. 12(2): 49—54. [In
Chinese with English summary.]
-. 2000. Some new taxa of Bambusoideae in West¬
ern Sichuan, China. J. Bamboo Res. 19(1): 9—26. 11n
Chinese with English summary.]
- & L. Yang. 1998. A new species of the alpine
bamboo from China. J. Bamboo Res. 17(4): 1-3.
Schradera reticulata (Rubiaceae), una Nueva Especie de
Cosla Rica y Panama
Joaquin Sanchez- Gonzdlez
Herbario Nacional, Museo Nacional de Costa Rica, Apdo. 749-1000, San Jose, Costa Rica.
museohn@racsa.co.cr; holcusl6@hotmail.com
RESUMEN. Se describe e ilustra una nueva espe¬
cie, Schradera reticulata (Rubiaceae, Schradereae).
La nueva especie es similar a S. rotundata Standley
ex Steyermark de Colombia. No obstante ambas se
diferencian en que S. reticulata tiene liojas mas
angostas, en su mayorfa oblongas a elfpticas, la ve-
nacion terciaria es ligeramente prominente y fina-
mente reticulada en ambas earas de la lamina fo¬
liar. el apice es corto acuminado, pedunculo mas
largo, involucre) con dientes marginales, hipanto
mas corto y lobulos de la corola mas largos.
Abstract. A new species, Schradera reticulata
(Rubiaceae, Schradereae), is described and illus¬
trated here. This species is similar to S. rotundata
Standley ex Steyermark from Colombia. The two
species differ in that S. reticulata has narrower and
generally oblong to elliptic leaves, the tertiary ve¬
nation is slightly prominent and finely reticulate on
both foliar sides, the apex is short-acuminate, the
peduncle longer, the involucre has marginal teeth,
the hypanthium is shorter, and the corolla lobules
are longer.
key words. Costa Rica, Panama, Rubiaceae,
Schradera, Schradereae.
El genero Schradera Vahl (Isertieae), comprende
40 especies distribuidas desde Rrasil hasta Costa
Rica, las Antillas, el sureste de Asia y Nueva Gui¬
nea. Se distingue por su habito epifito. inflorescen-
cias dispuestas en capftulos pedunculados, subten-
didos por un pequeno involucro, flores
semisuculentas que secan de color negro (Burger &
Taylor, 1993). El genero es poco conocido debido,
entre otras razones, a que el habito epifito dificulta
la recoleccion de buenos espeefmenes. En Mesoa-
merica se considera que el genero Schradera se en-
cuentra representado por cuatro especies, de las
cuales una esta presente en Costa Rica y tres en
Panama (Taylor, en prep.). Ademas hay que mencio-
nar la existeneia de una especie inedita, compartida
por ambos pafses (Taylor, 2003). Por tanto, con esta
nueva especie que se describe aquf, el numero total
aumenta a seis.
Schradera reticulata J. Sanchez-Gonzalez, sp.
nov. TIPO: Costa Rica. Limon: canton de Ta-
lamanca, Sixaola, Gandoca, El Llano entre h-
las Manzanillo y Rfo Mile Creek, 09°37'N,
82°4 1 ' W, 50-100 m, 27 Mar. 1995 (fl), G. He¬
rrera 7585 (holotipo, CR; isotipos, F, K, MO.
US, USJ). Figura 1.
Schraderae rotundatae Standley ex Steyermark affinis
qua imprimis differt folds angustioribus, oblongis usque
elipticis, nervis tertiis utrinque suhtiliter reticulatis pro-
minentibusque; apice breve acuminatis; pedunculis lon-
gioribus, involucri margine minute denticulate (sub lente
10X); hypanthio breviore; lobulis corollae longioribus.
Arbusto epifito, glabro; corteza exfoliante. Hojas
5.5—12.5 X 2—6.5 cm, elfptico-oblongas, elfpticas,
oblanceoladas u obovadas, cartaceas, de color cafe
oscuro en la haz y cafe claro a pardo en el enves
cuando secas, la base cuneada a obtusa, el apice
breviacuminado a obtuso; nervaduras secundarias
6—8 pares, ligeramente broquidodromas, impresas
en la haz, prominentes y de color mas claro en el
enves; venacion terciaria reticulada, prominente en
ambas caras, formando areolas de 0.5—1 mm de
ancho; pecfolos 1-2.5 cm de largo; estfpulas ca. 25
mm de largo, tempranamente caedizas, el apice ob¬
tuso. Inflorescencias terminales, solitarias; pedun-
culos 1—3.5 cm de largo; eabezuelas 1.5—2.0 cm de
diametro (sin incluir las corolas), hemisfericas a
subglobosas, con 6-20 flores; involucro 0.8—1.2 cm
de diametro, margen ondulado con dientes de 0.7—
2.0 mm de largo. Flores protandras, muy fragantes;
hipanto turbinado, de 3^4 mm de largo; limbo ca-
lieino triangular, pulverulento hacia los margenes,
verde claro, secando negro, 3 mm de largo; corola
blanca, negra cuando seca, pulverulenta interna y
externamente; tul >o de 0.8—1.2 cm de largo, lobulos
5, ca. 10 mm de largo, lineares, camosos, con un
apendice de 2 mm de largo en forma de espolon
reflexo e inserto en el apice; anteras 5, exsertas,
dorsifijas, introrsas, de color amarillo tenue, secan¬
do con el conectivo oscuro y las tecas claras, 4 mm
de largo; estilo villoso, estigma exserto, villoso, 25
mm de largo. Frutos jovenes ca. 5 mm de diametro,
subglobosos.
Novon 13: 477-479. 2003.
478
Novon
Figura 1. Schradera reticulata J. Sdnchez-Gonzalez, sp. nov. —A. Kama fertil con ampliackm de la superficie abaxial
de la lamina foliar mostrando el fino reticulado de la venaeion lereiaria (tornado del holdtipo, G. Herrera 7585, CK).
—K. Flor (tornado del pardtipo G. Stiles 85-7, USJ). —C. Infrutescencia (tornado del paralipo J. Kirkbride, jr. & J.
Duke 1800, MO).
Distribution, habitat y fenologta. Habita en
Bosque Htfmedo y muy Humedo Tropical y en
Bosque Pluvial Premontano, segun Hoblridge (Tosi.
1969; FAO, 1971), de 50 a 1200 m. El holdtipo es
el especimen recolectado a la menor elevacion del
ambilo de distribucidn altitudinal de la nueva es-
pecie. Se distribuye por la vertiente atldntiea de
Costa Rica hasta la provincia de Darien en Pana¬
ma. Se ban recolectado especfmenes con flores en
abril, mayo y julio y eon frutos en abril. En el es¬
pecimen J. Kirkbride, Jr. & J. Duke 1300 se men-
ciona que los nativos del Chocd la utilizan para
tenii' la lengua de negro.
Relacidn con otras especies. Schradera reti-
Volume 13, Number 4
2003
Sanchez-Gonzalez
Schradera reticulata (Rubiaceae)
479
culata es similar a S. rotundata Standley ex Ste-
yermark de Colombia. No obstante, ambas especies
difieren en algunas caracterfsticas: S. reticulata tie-
ne hojas de 1-6.5 cm de ancho, en su mayorfa
oblongas a elfpticas, la venacion tereiaria es lige-
ramente prominente y finamente reticulada en am¬
bas earas de la lamina foliar, el apice es corto acu-
minado, peduneulo de 1—3.5 cm de largo, involucro
con dientes marginales, hipanto de 3—4 mm, tur-
binado y lobulos de la corola de 10 mm. En con-
traste, S. rotundata tiene hojas de 4—8 cm, amplia-
tnente obovadas, la venacion tereiaria inconspicua,
apice redondeado a subtruncado, peduneulo de 0.7
a 1.5 cm de largo, involucro sin dientes marginales,
hipanto de 11 mm de largo, urceolado y lobulos de
la corola de 8 mm.
Etimologia. El nombre de Schradera reticulata
liace referencia a la venacion tereiaria finamente
reticulada en ambas superficies de la lamina foliar.
El epftelo fue utilizado como nombre de herbario
por C. M. Taylor.
Pardtipos. COSTA RICA. Liindn: canton de Tala-
manca, cerro entre cerro Chimu y cerro Matama, 30 abr.
1985, L I). Gomez et al. 23581 (MEXU, MO). Heredia:
Parque Naeional Braulio Carrillo, 8 abr. 1983, G. Stiles
83-7 (USJ). PANAMA. Code: vicinity of La Mesa, be¬
yond El Valle, on Eastern ridge, along trail to summit of
Cerro Gaital, 13 July 1987, G. McPherson 11256 (MO).
Comarca de San Bias: El Llano—Cartf Road, 12 mi.
from Pan American Hwy., along deeply shaded large
stream, wet tropical forest, 10 May 1981, h. Sytsma & L.
Andersson 4504 (MO). Darien: Cuasi—Cana Trail between
Cerro Campamiento & La Escalera E of Tres Bocas, 30
Apr. 1968, Kirkbride, Jr. & J. Duke 1300 (MO). Pana¬
ma: Cerro Jefe, 17 May 1968, R. Dressier 3509 (MO).
Agradecimientos. Agradezco a Charlotte Taylor
(MO) por el acceso a los espeefmenes, por cederme
la descripcion preliminar preparada para la Flora
Mesoamericana y el texto de su artfculo sometido a
publicat ion y por la revision critiea del manuscrito.
A Piero Delprete (NY) por la revision critiea del
manuscrito. A Jorge Gomez Laurito (USJ) por la
preparacion de la diagnosis en latfn y revision del
manuscrito. A Jose Alberto Perez (Cope) por la ex-
celente ilustracion. Al Museo Naeional de Costa
Rica por el auspicio y financiamiento de este tra-
bajo.
Literatura Citada
Burger, W. & C. M. Taylor. 1993. Rubiaceae. In: W. Bur¬
ger (editor). Flora Costaricensis. Fieldiana Bot. n.s. 33:
309.
FAO. 1971. Inventariacidn y demostraciones forestales,
Panamd. Zonas de Vida, basado en la labor de J. A.
Tosi. FO:SF/PAN 6. Informe tdcnico 2. Roma. 89 pp. 1
mapa.
Taylor, C. M. 2003. Rubiacearum Americanarum Magna
Hama Pars XIII. A new species of Schradera (Schra-
dereae) from Mesoamerica. Novon 13: 220—222.
Tosi, J. A., Jr. 1969. Republiea de Costa Rica. Mapa Eco-
Idgico segun la clasificacidn de zonas de vida del mun-
do de L. R. Holdridge. Esc. 1: 750.000. Centro Cien-
tffico Tropical. San Josd, Costa Rica.
Aspidistra guangxiensis (Convallariaceae), a New
Species from China
Saichun Tang
South China Institute of Botany, Chinese Academy of Sciences, Wushan, Guangzhou,
Guangdong 510650, People’s Republic of China, tangs0448@sina.com
Yan Liu
Guangxi Institute of Botany, Guangxi Zhuangzu Autonomous Region, Chinese Academy of
Sciences, Yanshan, Guilin, Guangxi 541006. People’s Republic of China
Abstract. In the course of our investigation of
the genus Aspidistra (Convallariaceae), a new spe¬
cies was discovered in the Guangxi Zhuangzu Au¬
tonomous Region of southern China. It is described
below as A. guangxiensis S. C. Tang & Y. Liu, and
an illustration is provided.
Key words: Aspidistra, China, Convallariaceae,
Guangxi.
Aspidistra Ker Gawler (Convallariaceae) consists
of approximately 60 species of perennial, rhizo-
matous, evergreen herbs distributed in eastern Asia
(China, India, Japan, Laos, Thailand, and Vietnam).
Its center of distribution and differentiation is in
southern China, where 56 native species (54 en¬
demic) are known, most of them in the Guangxi
Zhuangzu Autonomous Region (Lang et ah, 1999;
Liang & Tamura, 2000; Fang & Yu, 2002; He,
2002; Li & Tang, 2002). The plants grow in sea¬
sonal rain forests (monsoon iorests) and evergreen
broad-leaved forests. In the course of our investi¬
gation of the genus, a new species was recently
discovered in Long’an County, about 75 km north¬
west of Nanning in southwestern Guangxi.
Aspidistra guangxiensis S. C. Tang & Y. Liu, sp.
nov. TYPE: China. Guangxi: Guilin Botanical
Garden, cultivated, 12 May 2002, Y. Liu 0001 ,
originally collected from Guangxi, Long’an
County, limestone mountains, 350 m, date and
collector not recorded (holotype, 1BK; isotype,
IBK). Figure 1.
Aspidistrae patentilobae affinis, sed staminibus in parte
tertia infera tubi affixis, stigmate conspicue inferioro, an-
theris reniformibus; stigmate facie supremo alutaceo, cen¬
tra 3-carinatis conspicue purpureo, infero purpureo differl.
Herbs perennial, evergreen, rhizomatous. Rhi¬
zome creeping, subterete, 3—5 mm diam., densely
covered with scales. Leaves solitary, spaced; petiole
14-40 cm, partly enveloped by sheaths; leaf blade
ovate to elliptic, 6—20 X 7—8 cm, base decurrent
into petiole, apex acuminate. Flowers solitary; pe¬
duncle decumbent, 4—5 cm; bracts purple, trian¬
gular, terminal 2 or 3 close to flower; perianth abax-
ially purple, adaxially yellow, urceolate, fleshy;
tube subglobose, 1—2 cm diam.; lobes 6 or 8, free,
spreading, linear-lanceolate, 2—3 cm X 2—6 mm,
base with an adaxial, rounded, toothlike appendage
extending part of the way across the mouth of the
perianth tube, apex subobtuse. Stamens 6 or 8, in¬
serted in proximal Vs of perianth tube, positioned
conspicuously lower than stigma; filament incon¬
spicuous; anther reniform, 2—5 mm. dehiscing by a
slit. Pistil ca. 1 cm; ovary much narrower than stig¬
ma, 3-loculed; stigma sessile, dilated and bowl¬
like. 1-1.5 cm diam., thickened, abaxiallv with 12
or 16 longitudinal ridges, adaxially pale yellow and
slightly concave, with 3 purple ridges at center (4
ridges not observed in 4-merous flowers), margin
with 12 or 16 triangular lobes.
Phenology. Flowering in May.
Distribution. Known in the wild only from lime¬
stone mountains at 350 m elevation in Long’an
County, Guangxi Zhuangzu Autonomous Region,
southern China.
This new species is most similar, and probably
related, to Aspidistra patentiloba Y. Wan & X. 11.
Lu (Wan, 1989: 99) in its dilated, thickened stigma,
which is longitudinally ridged abaxially, slightly
concave adaxially, and triangular-lobed at the mar¬
gin. However, the latter species, distributed in Liu-
jiang County in central Guangxi (200 km northeast
of Long’an), differs in having the stamens inserted
at the middle of the perianth tube, positioned level
with the stigma, the anthers ovate, and the stigma
adaxially purple with four ridges at its center.
The floral structure of Aspidistra, especially the
stigma, is the most important taxonomic character
Novon 13: 480-482. 2003.
Volume 13, Number 4
2003
Tang & Liu 481
Aspidistra guangxiensis (Convallariaceae)
A
Figure 1. Aspidistra guangxiensis S. C. Tang & Y. Liu. —A. Flowering plant. —B. Flower. —C. Flower with half of
perianth removed showing stamens and pistil. —D. Perianth bisected. —E. Stigma lateral view. —F. Stigma apical
view. Drawn from the holotype, Y. Liu 0001 (IBK), by He Shunqing.
482
Novon
because of its great variation (Li et al., 2000). Both
A. guangxiensis and A. patentiloba have been cul¬
tivated under the same environmental conditions at
the Guilin Botanical Garden, where their floral dif¬
ferences hold true. Furthermore, one of us (Liu)
collected material of A. guangxiensis from the bo¬
tanical garden in 2003 and observed the floral
structure to be the same as that of the type material
collected in 2002. This demonstrates that the dis¬
tinctness of A. guangxiensis from A. patentiloba is
not an artifact of cultivation and that the diagnostic
floral characters of the new species are constant.
Paratypes. CHINA. Guangxi: Guilin Botanical Gar¬
den, cultivated, 12 May 2002, I. Liu 0002, originally col¬
lected from Guangxi, Long’an County, limestone moun¬
tains, 350 m, date and collector not recorded (IBK).
Acknowledgments. We thank Deng Yunfei,
South China Institute of Botany, Chinese Academy
of Sciences (IBSC). and Nicholas Turland, Missouri
Botanical Garden (MO), for their help in preparing
the paper, and two anonymous reviewers for valu¬
able comments. We also thank He Shunqing, Gu¬
angxi Institute of Botany, Guangxi Zhuangzu Au¬
tonomous Region, Chinese Academy of Sciences
(IBK), for drawing the illustration. This project is
supported by the Special Fund for Taxonomy and
Flora of the Chinese Academy of Sciences.
Literature Cited
l ang, I). & L. Y. Yu. 2(X)2. Three new species of Aspi¬
distra Ker-Gawl. (Liliaceae) from Guangxi, China. Acta
Phytotax. Sin. 40: 159—163.
lie, S. Z. 2002. A new species of Aspidistra Ker-Gawl.
(Liliaceae) from Guizhou, China. Acta Phytotax. Sin.
40: 377-379.
Lang, K. Y„ G. Z. Li. Y. Liu, Y. G. Wei & R. X. Wang.
1999. Taxonomic and phytogeographic studies on the
genus Aspidistra Ker-Gawl. (Liliaceae) in China. Acta
Phytotax. Sin. 37: 468—508.
Li. C. Z. & S. C. Tang. 2002. New taxa of Aspidistra Ker-
Gawl. from Guangxi, China. Guihaia 22: 289-291.
-, K. Y. Lang, R. X. Wang & Y. G. Wei. 2000. On
the trends of morphological differentiation and a new
system ol classification in Chinese Aspidistra Ker-Gwal.
(sic-J (Liliaceae). Guihaia 20: 201—217.
Liang, S. Y. X M. N. Tamura. 2000. Aspidistra. Pp. 240—
250 in: Z. Y. Wu & P. II. Raven (editors), flora of
China, Vol. 24. Flagellariaceae through Marantaceae.
Science Press, Beijing, and Missouri Botanical Garden
Press, St. Louis.
Wan. Y. 1989. Three new species ol the genus Aspidistra
from Guangxi. Bull. Rot. Res.. Harbin 9(2): 97—102.
Nomenclatural Actions in Whytockia (Gesneriaceae)
Yin-Zheng Wang
Laboratory of Systematic & Evolutionary Botany and Herbarium, Institute of Botany, Chinese
Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of
China, wangyz@ns.ibcas.ac.cn
Abstract. On the basis of morphological, ana¬
tomical, and karyotypic evidence, a change in sta¬
tus is proposed for Whytockia wilsonii (A. Weber)
Y. Z. Wang, which is elevated from varietal to spe¬
cies level. Further, the affinity of another variety
currently accepted within W. tsiangiana is recog¬
nized, and it is now transferred from W. tsiangiana
as the new combination W. hekouensis var. minor
(W. W. Smith) Y. Z. Wang. Illustrations of W. wil¬
sonii are included along with morphological de¬
scriptions and diagnostic characters. The relation¬
ships of these and related species are discussed as
well as geographic distribution. A key distinguish¬
ing these species is provided.
Key words: China, Gesneriaceae, Whytockia.
The genus Whytockia W. W. Smith includes eight
species endemic to China. It was proposed by W. W.
Smith in 1919 for the accommodation of Stauranthera
chiritiflora, in which he distinguished a variety minor.
With limited material, Weber (1982) conducted a pre¬
liminary revision of Whytockia in which three species
were recognized. The current author has carried out
systematic and evolutionary studies on Whytockia anti
its allies since 1991. Intensive research in herbaria,
field, and laboratory over the last ten years has led to
a better understanding of this group and to the dis¬
covery of several species previously unknown to sci¬
ence. The recognition of significant characters con¬
tributes to a natural delimitation of the taxa of
Whytockia. The nomenclature of two taxa previously
treated as varieties needs adjustment herein.
Whytockia wilsonii (A. Weber) Y. Z. Wang, stat.
nov. Basionym: Whytockia tsiangiana (H. Han-
del-Mazzetti) A. Weber var. wilsonii A. Weber,
Notes Roy. Hot. Card. Edinburgh 40: 365.
1982. TYPE: China. Sichuan (Szechuan): side
of stream, Sep. 1903, E. II. Wilson 3292 (ho-
lotype, BM; isotypes, E, K). Figure 1.
Leaves opposite, strongly unequal. Larger leaf
blades broadly obovate or widely ovate-oblong, 9—11
X 3^4 cm, nearly sessile or with a short petiole 2—
3 mm long, base oblique, widely cuneate or rounded
on the wide side to cuneate on the narrow side, mar¬
gins serrate-dentate, teeth 1.5—3 mm long. Smaller
leaf blades ovate, 6—13 mm long, sessile. Pair-flow¬
ered cymes emerging from the axils of the larger
leaves. Sepals 5, free or connate at base, ovate or
ovate-triangular, 2—3 X 1-2 mm. Corolla red or red¬
dish, tubular and bilabiate, 1.5-2 cm long with co¬
rolla tul ie 10 mm long, outside glabrous, inside with
2 rows of yellow-green unicellular and clavate hairs
under and between lobes of the lower lip; upper lips
3 mm long, 2-lobed; lower lips about 6.5 mm long,
3-lobed. Stamens 4, didynamous, attached to corolla
base for 2 mm. Staminode long ovate, 1.1 mm long.
Ovaries ovate, about 1.5 mm long; styles about 4.6
mm long; 2 stigmas only connate at base. Seeds mi¬
nute, many, testa obliquely striate.
Whytockia wilsonii resembles W. hijieensis Y. Z.
Wang & Z. Y. Li by its large corolla (1.5—2 cm
long), 5 sepals free or connate at base, and 2 stig¬
mas free or only connate at base. In addition, W.
wilsonii and W. hijieensis have a karyotype (2 n —
18) more similar to each other than to other Why¬
tockia, i.e., similar frequencies of median and sub¬
median chromosomes, and lacking a subterminal
chromosome (Wang et ah, 1998). However, W. wil¬
sonii is easily distinguished from W. hijieensis by
the following morphological characters, i.e., the
large leaf blades of unequal leaf pairs broadly ob¬
ovate or widely ovate-oblong with base widely cu¬
neate or rounded on the wide side; blade margins
serrate-dentate with teeth 1.5—3 mm long; seed tes¬
ta obliquely striate. The X-shaped ventral carpel-
lary bundle of the ovary and the anchor-shaped ax-
ile placentas are remarkably distinctive from those
of other species in Whytockia (Wang & Pan, 1998).
W hytockia hekouensis Y. Z. Wang var. minor
(W. W. Smith) Y. Z. Wang, comb. nov. Basion¬
ym: Whytockia chiritiflora (Oliver) W. W. Smith
var. minor W. W. Smith, Trans. Bot. Soc. Ed¬
inburgh 27: 338. 1919. Whytockia tsiangiana
(H. Handel-Mazzetti) A. Weber var. minor (W.
W. Smith) A. Weber. Notes Roy. Bot. Card. Ed¬
inburgh 40: 365. 1982. TYPE: China. Yunnan:
Yuanyang, Feng Chunling (Feng Chen Lin), in
mountain forest, 7000 ft. alt., A. Henry 11232
(holotype, E; isotype, K).
Novon 13: 483-486. 2003.
484
Novon
F igure 1. Diagrammatic illustrations of the holotype of Whytockia wilsonii (E. //. Wilson 3292, BM). —A. Plant with
flowers, scale har = 10 mm. —B. Opened calyx taking view from abaxial side, scale bar = 4 mm. —C. Corolla opened
to reveal two internal rows of clavate bairs under and between lobes of the lower lip, scale bar = 4 mm. —I). Anthers,
scale bar = 2 mm. —E. Pistil, showing longer styles 2—2.5 times as long as ovary, and two stigmas connate at their
base, scale bar = 2 mm.
Volume 13, Number 4
2003
Wang
Whytockia (Gesneriaceae)
485
Whytockia hekouensis var. minor differs from va¬
riety hekouensis in its calyx lobe narrowly ovate,
apex ascending-spreading; flower pedicel not in-
erassate on upper portion after anthesis.
Whytockia wilsonii and W. hekouensis var. minor
were previously treated as varieties wit Inn W.
tsiangiana (Weber, 1982). The former was de¬
scribed as a new variety for W. tsiangiana, while
the latter was transferred from W. chiritiflora to IT.
tsiangiana (Weber, 1982). The species IT tsian¬
giana is characterized by its small, white corolla
(8—12 mm long), its shorter style equal to the ova¬
ry or less than 1.5 times as long as the ovary, and
its wholly connate stigma. Four other species, IT
bijieensis, W. wilsonii, W. chiritiflora, and IT. hek¬
ouensis, have large, red corollas (1.5—2 cm long),
longer styles 2—2.5 times as long as ovary, and 2
stigmas free or connate only at their base or in
their lower half. Among the four species, the 5-
lobed calyx united in a tube to the mid-point of
the calyx makes IT. chiritiflora and IT hekouensis
cluster together, while W. wilsonii is closely relat¬
ed to IT bijieensis in five sepals that are free or
connate at base, and two stigmas that are free or
only connate at base. In addition to the above mor¬
phological characters, the karyotype of IT. tsian¬
giana, i.e., 2 n = 18 = 4m + 8sm (2sat.) + 6st,
demonstrates a high degree of asymmetry that is
remarkably distinctive from those of other species
above. Whytockia wilsonii and IT bijieensis have
closely similar karyotypes, for the two species
have similar frequencies of median and subme¬
dian chromosomes and lack a subterminal chro¬
mosome, i.e., 2 n = 18 = 2M + 8m (lsat.) + 8sm
(2sat.) in IT wilsonii and 2 n = 18 = 2M + 8m +
8sm (lsat.) in IT bijieensis (Wang et al., 1998).
Moreover, IT tsiangiana is widely distributed from
eastern Guizhou Province to central China, i.e.,
Hunan, western Hubei, and northern Guangxi
Provinces. In contrast, IT wilsonii is limited to
southern Sichuan Province adjacent to the distri¬
butional area of IT bijieensis in western Guizhou
and northeastern Yunnan Provinces (Wang & Li,
1997). Whytockia chiritiflora and IT hekouensis,
two species standing sharply apart from the rest
of Whytockia in their 5-lobed calyx united in a
tube to the mid-point of the calyx, are restricted
to separate localities in southeastern Yunnan
Province (Wang, 1995). The four species, i.e., IT
bijieensis, W. wilsonii, IT. chiritiflora, and IT hek¬
ouensis, are located in the area where the plants
of Gesneriaceae are most diverse in the world. A
diagnostic key for these five species (including va¬
rieties) is provided below.
Rev to the Whytockia Species and Varieties
Discussed Above
la. Corolla white, 0.8—1.2 cm long; style 0.6—0.8
mm long, equal to ovary or less than 1.5 times
as long as ovary; stigma wholly connate, elliptic-
disciform . W. tsiangiana
lb. Corolla red or reddish, 1.5—2 cm long; style 4—
6 mm long, 2-2.5 times as long as ovary; two
stigmas free or connate at base or in lower half.
2a. Five-lobed calyx united in a tube to the mid¬
point of calyx.
3a. Two stigmas free; fruit a capsule, biv-
alvate dehiscent. IT chiritiflora
3b. Two stigmas connate in lower half; fruit
a capsule, irregularly dehiscent . . IT hek¬
ouensis
4a. Calyx lobes widely ovate or ovate-
triangular, apically reflexed; pedi¬
cel incrassate on upper portion al¬
ter anthesis. IT hekouensis var.
hekouensis
4b. Calyx lobes narrowly ovate, apical¬
ly ascendingly spreading; pedicel
not incrassate on upper portion af¬
ter anthesis .
. W. hekouensis var. minor
2b. Calyx with 5 sepals, free or connate at base.
5a. Larger leaf blade of unequal leaf pair
narrowly ovate-oblong, margins irregu¬
larly repand-serrate and glandular-ser¬
rate, these serrations 1—1.5 mm long,
base auriculate or rounded on wide
side; the two stigmas free; seed tesla re¬
ticulate . W. bijieensis
5b. Larger leaf blade of unequal leaf pair
widely obovate or broadly ovate-oblong,
margins serrate-dentate, these teeth
1.5—3 mm long, base widely cuneate or
rounded on wide side; the two stigmas
connate at base; seed testa obliquely
striate. IT wilsonii
Acknowledgments. The author thanks Wang
Wen-tsai and Li Zhen-yu for providing valuable
discussion and comments on this manuscript. I am
grateful for borrowing type specimens from flu 1 fol¬
lowing herbaria: BM, E, and K. This study was sup¬
ported by NSFG Projects 30270093 and the State
Key Basic Research and Development Plan
(G200046803 -4).
Literature Cited
Smith, W. W. 1919. Whytockia, a new genus of Gesneri¬
aceae. Trans. Bot. Soc. Edinburgh 27; 338—339.
Wang, Y. Z. 1995. Two new species of Whytockia (Ges¬
neriaceae) from Yunnan, China. Acta Phytotax. Sin.
33(3): 297-301.
-& Z. Y. Li. 1997. A new species of the genus
Whytockia W. W. Smith (Gesneriaceae) from Guizhou,
China. Acta Phytotax. Sin. 35(1): 67-69.
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No von
-& K. Y. Pan. 1998. Comparative floral anatomy
of Whytockia (Gesneriaeeae) endemic to China. Pp.
352-366 in A. L. Zhang & S. G. Wu (editors), Floristic
Characteristics and Diversity of Cast Asian Plants. Chi¬
na Higher Education Press. Beijing.
-. Z. J. Gu & 1). Y. Hong. 1998. Karyotypes of
Whytockia (Gesneriaeeae). Acta Phytotax. Sin. 36(1):
28-35.
Weber, A. 1982. Contributions to morphology anti system¬
atic^ of Klugieae and Loxonieae (Gesneriaeeae) IX. The
genus Whytockia. Notes Roy. Bot. Gard. Edinburgh 40:
359-367.
Delphinium hillcoatiae var. pilocarpum (Ranunculaceae), a New
Variety from Xizang, China
Yang Qiner and Luo Yan
Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of
Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People's Republic of China.
qeyang@ns.ibcas.ac.cn; luoyan@ns.ibcas.ac.cn
ABSTRACT. Delphinium hillcoatiae var. pilocar¬
pum Q. E. Yang & Y. Luo (Ranunculaceae) is de¬
scribed as a new variety from Xizang. China. It
differs from the typical variety by having densely
puberulent ovaries.
key words: China. Delphinium, Ranuncula¬
ceae.
In the course of examining specimens of Ran¬
unculaceae at Kew in 2002. four collections of the
genus Delphinium L. from the neighborhood of Lha¬
sa, the capital city of Xizang, China, caught our
attention. At first glance all four collections, F.
Spencer Chapman 136, 943, ./. F. Duthie s.n., and
L. A. Waddell s.n., show no essential difference in
appearance from D. hillcoatiae Munz, a species
also described from this region. However, upon
careful examination we found that one of the col¬
lections, Chapman 943, has glabrous ovaries, while
the remaining three all have densely puberulent
ovaries. In the protologue, Munz (1968) described
his species as having glabrous carpels. This has
been confirmed by our examination of the holotype
of Munz’s species, Ludlow & Sherriff 2424, and the
paratype, Kingdon Ward 12370, both deposited at
BM. These two collections have totally glabrous
ovaries indeed. One gathering at PE, Qinghai-Xiz-
ang Fxped. 7938, which was made from Qiixii
County very near Lhasa and named as D. hillcoa¬
tiae, also has densely puberulent ovaries. The de¬
scriptions of D. hillcoatiae in the Flora Reipublicae
Popularis Sinicae (Wang, 1979), Flora Xizangica
(Wang. 1985), and Flora oj China (Wang & War-
nock, 2001) may all have been mainly based on
this collection because the ovaries were described
as puberulent in these works.
Delphinium hillcoatiae is only known from the
Lhasa region, Xizang, China. This species is dis¬
tinguished by its woody root, leaves 3-partite within
1 cm from the base, stem, inflorescence rachis, and
pedicels all densely spreading glandular puberu¬
lent, 4- to 20-flowered terminal raceme, and stam-
inodes and petals black-brown. Based on these
characters it should be a member of D. subg. Del-
phinastrum (DC.) Petermann sect. Elatopsis Huth,
but we cannot yet find with certainty its closest ally
in the Chinese species of Delphinium.
As the specimen at PE cited above as well as
the three specimens at Kew, all with densely pu¬
berulent ovaries, show no other difference from
Delphinium hillcoatiae and occur in nearly the
same region as the species, it seems reasonable to
treat them as a new variety.
Delphinium hillcoatiae Munz var. pilocarpum
Q. E. Yang X Y. Luo, var. nov. TYPE: China.
Xizang: (Qiixii County, shady places, 3600 m,
25 Sep. 1975 (H), Qinghai-Xizang Fxped.
7938 (holotype, PE; isotype, PE).
A typo ovariis dense puberulis differt.
This variety differs from Delphinium hillcoatiae
var. hillcoatiae by its densely puberulent ovaries.
In the typical variety, the ovaries are totally gla¬
brous.
The two varieties are only known from the Lhasa
region and its neighborhood, Xizang, China. Ac¬
cording to the collectors’ note, the typical variety
grows on open dry stony hillside, whereas the va¬
riety pilocarpum grows in shady places.
Pciratypes. CHINA. Xizang: Lhasa, 18 Sep. 1904, A.
A. Waddell s.n. (K), May-July 1947, J. F. Duthie s.n. (K);
Cheesul (= Qiixii ?), 11,500 ft., 21 Aug. 1936, F. Spencer
Chapman 136 (K).
Acknowledgments. We are most grateful to the
Keepers of BM, K, and PE for permission to ex¬
amine specimens, and to Michael G. Gilbert (BM,
now at K) for his kind help in the course of writing
this paper. This work was supported by the National
Natural Science Foundation of China (grant no.
30070057) and a Key Project of the Chinese Acad¬
emy of Sciences (KSCXZ-SW-101A).
Literature Cited
Munz, P. A. 1968. A synopsis of the Asian species of
Delphinium, sensu stricto. J. Arnold Arbor. 49: 73—166.
Novon 13: 487-488. 2003.
488
Novon
Wang, W. T. 1979. Delphinium. Pp. 32(>—14)2 in Flora Hei-
publicae Popularis Sinicae, Vol. 27. Science Press, Beijing.
-. 1985. Delphinium. Pp. 39—59 in Z. Y. Wu (edi¬
tor), Flora Xizangica, Vol. 2. Science Press, Beijing.
- & M. J. Warnock. 2001. Delphinium. Pp. 223—
274 in Z. Y. Wu & P. H. Baven (editors). Flora of China.
Vol. 6. Science Press, Beijing, and Missouri Botanical
Garden Press, St. Louis.
Volume 13, Number 1, pp. 1—160 of NOVON was published on 25 March 2003.
Volume 13. Number 2. pp. 161—280 of NOVON was published on 24 June 2003.
Volume 13, Number 3, pp. 281—376 ol NOVON was published on 10 September 2003.
Volume 13, Number 4, pp. 377—488 of NOVON was published on 5 December 2003.