Volume 14
Number 1
2004
NOVON
Suaeda neuquenensis (Chenopodiaceae), a New Species
from Argentina
M. A. Alonso
GIB (Institute Universitario de la Biodiversidad), Universidad de
Alicante, Apartado 99, E-03080 Alicante, Spain, ma.alonso@ua.es
L. Conticello and M. B. Cerazo
Departamento de Botanica Aplicada, Universidad Nacional del Comahue,
Rio Negro, Argentina
Abstract. Suaeda neuquenensis , a new species of
Suaeda (Chenopodiaceae), is described from Ane-
lo’s Bowl near Anelo, Neuquen Province, Argenti¬
na. It differs from other perennial Argentinean spe¬
cies in being a decumbent shrub with creeping
stems that root at the lower nodes bearing short,
oblong, often reflexed leaves and a large seed.
Key words: Argentina, Chenopodiaceae, Neu¬
quen, Suaeda.
Two perennial species of Suaeda Forsskal ex J.
F. Gmelin (Chenopodiaceae), S. divaricata Moquin-
Tandon and .S', argentinensis Soriano, are currently
known from Argentina (Soriano, 1942; Giusti,
1997). Suaeda fruticosa (L.) Forsskal was reported
from the Boca district of Buenos Aires by Soriano
(1945), who nonetheless suggested it was a recent
introduction and not likely to persist; there is now
no evidence that it did. Species of the genus are
highly variable as to succulence, habit, and size of
leaves (Morello, 1958; Hopkins & Blackwell, 1977;
Bassett & Crompton, 1977; Pedrol & Castroviejo,
1988). Other features are more constant: herba¬
ceous versus woody habit, type of inflorescence (in¬
cluding the density of the glomerules), relationship
of leaf length and internode length, shape of the
calyx, stigma shape and length, and seed size (Hop¬
kins & Blackwell, 1977; Pedrol & Castroviejo,
1988).
Material and Methods
Dried specimens of Suaeda at ARC, CORD, and
MERI , were studied, as well as fresh material col¬
lected in the Neuquen Province.
Taxonomic and nomenclatural treatments follow
Standley (1916), Morello (1958), Bassett and
Crompton (1977), Hopkins and Blackwell (1977),
Correa (1984), Pedrol and Castroviejo (1988), Teill-
er (1996), Guisti (1997), Greuter el al. (2000), and
Schenk and Ferren (2000, 2001). Herbarium sheets
of Suaeda neuquenensis are conserved at ABH of
the University of Alicante (Holmgren & Holmgren,
1993).
Results and Discussion
Suaeda neuquenensis M. A. Alonso, Conticello &
Cerazo, sp. nov. TYPE: Argentina. Neuquen
Province: Anelo’s Bowl near Anelo,
38°11'56"S, 69°00'46”W, 350 m.s.m. on sa-
line-sodic soil, 24 Mar. 1998, L. Conticello &
M. A. Alonso s.n. (holotype, ABH 41516; iso¬
type, ABH 42972). Figure 1.
Nanophanerophitus in fructicibus pulvinatis usque ad
50 cm altitudinis et 2.50 m diametri se extendentihus.
Caulis atramineus leviler strialus, glaber cum internodiis
1 mm, interdum inaestimabilis in gemmis recentibus, 2-
3 mm longis in ramis adultis. Folia adulta patentia aut
reflexa, iuvenilia perpendicularia, alterna oblonga, sue-
culenta cum apiee obtuso ac leviter mucronato, parum aut
Novon 14: 1-5. 2004.
2
Novon
Figure 1. Suaeda neuquenensis M. A. Alonso, Conticello & Cerazo. — A. Habit (holotype, ABH 41516). — B, C.
Flowering branches. — I). Young sterile shoot. — E. Detail of an axillary floral glomerule from the holotype.
nihil ad basim attenuata, dorsi ventral iter compressa, mar-
gines integri et rotundi, cum nitore virentia ant glauea vel
albescentia propter salina eorpuscula (|uae in superficie
deponuntur, 5—15 mm longa et 3—8 mm lata. Gynodioe-
cius. sessiles llores iuncti in glomerulis axillaribus 2-6,
hermaphroditi cum sterili gynaecio, 5 staminibus et calyx
5 sepalorum. Flores feminei cum gynaecio fecundo unil-
oculari, stylis 3 ad 0.5 mm longis, stigmatibus papillis
elongatis atque pilis acutis vestitis. Flores isti minores
sunt quam hermaphroditi, sed dia parte eius, et cum apice
Volume 14, Number 1
2004
Alonso et al.
Suaeda neuquenensis from Argentina
3
abtuso glabris ac inflexis, 2—3 mm longi. Fructus utricu-
lalus, ovoideus, 3X4 mm, in calycis segmentis inclusus
sed illis non affixus. Semen vertieale, nigrum, lenticulare,
testa reticulata.
Plants woody, cushion-shaped, dwarf shrubs with
creeping stems, forming mats 20—50 cm high and
25 dm across, the decumbent branches rooting at
the nodes. Stems light brown, weakly striated, the
internodes of new branches 1 mm long or less, 2—
3 mm long in mature branches. Leaves alternate
and positioned perpendicular on new branches, be¬
coming reflexed on mature branches, succulent, ob¬
long, 5—15 mm long, 3—8 mm wide, entire, shining
green to glaucous or white due to saline deposited
on the surfaces, the apices obtuse to rounded or
slightly mucronate, the bases attenuated. Flowers
gynodioecious, axillary, sessile, 2 to 6 per glomer-
ule, the perianth composed of 5 sepaloid perianth
segments, 2—3 mm long, these inflated in the mid¬
dle, apically obtuse and indexed, the hermaphro¬
dite flower with 5 stamens and a sessile, sterile
gynoeeium, the female flowers with a solitary, 1-
loeular gynoeeium with 3 stigmas up to 0.5 mm
long, bearing elongate papillae and acute hairs.
Fruit an ovoid utricle, 3 X 4 mm, enclosed by the
perianth segments, which are not closely attached
to it; seeds vertical, black, lens-shaped, reticulate
on the testa.
Ecology. Suaeda neuquenensis is known only
from Anelo’s Bowl near Anelo in the province of
Neuquen, where it occurs on saline-sodic soils in
monospecific communities or in association with
Atriplex undulata (Moquin) Dietrich, Distichlis spi-
cala (L.) Greene. Halophytum ameghinoi Spegaz-
zini, Sesuvium portulacastrum (I,.) L., and Suaeda
divaricata.
Taxonomic Discussion
In a recent paper on the nomenclature of Suae¬
da, Schrenk and Ferren (2001) stressed features
characterizing sections in the genus. According to
their arrangement, Suaeda neuquenensis belongs to
section Limbogermen lljin (sect. Salsina Moquin,
pro parte), which includes American shrubs and
subshrubs, with radial flowers, stigmas from a pit
at the top of the ovary, and leaves with suaedoid C4
anatomy, consisting of a ring of dark green palisade
and Kranz cells surrounding internal aqueous tis¬
sue, among other characters.
Relationships among taxa of section Limboger¬
men are shown in Table 1. Morphologically, S. neu-
quenensis shows a syndrome of characters that is
not found in other South American taxa in the sec¬
tion. Stems are decumbent and root at the basal
nodes, a feature that is absent in both S. argenti-
nensis and S. divaricata ; seeds are vertical and
have a reticulate testa (neither horizontal and
smooth as in S. argentinensis, nor tuberculate as in
S. divaricata). With regard to other American spe¬
cies, S. neuquenensis resembles ,S. conferla (J. K.
Small) 1. M. Johnston by its delicate cushion¬
shaped habit, though it differs by its robust, strong¬
ly woody stems, large fruits up to 4 X 3 mm (the
largest in the section), and its narrower and shorter
leaves, among other features. Connections to other
Central and North American taxa are very weak.
It is unlikely that Suaeda neuquenensis is an ex¬
otic, introduced taxon. Its natural habitat is quite
far from populated or frequented areas, and the soil
in which it grows has a high salt content, making
it unsuitable for agriculture. It is difficult therefore
to support an alien origin for this plant, since no
taxon is currently known to which it could be re¬
ferred.
Key to Related Species ok Suaeda in the Argentine
Republic
The following is part of Giusti's (1997) kev for the genus
Suaeda, to which S. neuquenensis has been added.
la. Shrubs or small shrubs.
2a. Plants with ramification in acute angles.
Leaves appressed to stem. Flowers in axil¬
lary, pedicellate glomerules. Seeds usually
horizontal . Suaeda argentinensis
2b. Plants with ramification in divergent angles.
Leaves divergent, not appressed to stem.
Flowers in axillary, sessile glomerules.
Seeds vertical.
3a. Shrubs erect, 1—3 m high. Leaves sem-
icylindric, perpendicular to the stem.
Young stems downy. Stems brown to
reddish . Suaeda divaricata
3b. Sbrubs decumbent, 0.2— 0.5 m high.
Leaves flat, reflexed on the stem. Young
stems glabrous. Stems light brown . . .
. Suaeda neuquenensis
lb. Annual herbs . . Suaeda patagoniea, Suaeda altisima,
and Suaeda densiflora
Acknowledgments. We thank Fidel Roig and Ma¬
nuel B. Crespo for comments on the manuscript, Ar-
eadio Alonso for his assistance with the Latin descrip¬
tion, and Pedro Marco for the illustration of this
species.
Literature Cited
Bassett, I. J. & C. W. Crompton. 1977. lhe genus Suaeda
(Chenopodiaceae) in Canada. Canad. .1. Bot. 56: 581—591.
Correa, M. N. 1984. Flora Patagoniea. Parte I\ a. Tomo \ 111.
Coleccion Cientffica del INTA. Buenos Aires.
Ferren. W. R. & H. J. Schenk. 2000. Flora of North America.
(http://hua.huh.harvard.edu/FNA/sueda 1 1 .16.00.html)
Gehu, J. M. & S. Rivas-Martfnez. 1982. Notions fondamen-
Table 1 . Comparison of morphological features among the species of Suaeda sect. Limbogt
4
Novon
15
CA)
3
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Southeast Asia (2 spp.)
INovoin 14: 12-16. 2004.
Volume 14, Number 1
2004
Arbelaez
Meliosma from Bolivia
13
Meliosma subg. Meliosma
sect. Meliosma — > Southeast Asia (12 spp.)
sect. Lorenzanea (Liebmann) Beusekom — > Neo¬
tropics (ca. 50 spp.)
Fifteen species were recognized by Van Beuse¬
kom (1971) in the revision of the Southeast Asian
sections of Meliosma. Section Lorenzanea is re¬
stricted to the Neotropics and has never been
monographed. All but one of the New World spe¬
cies, M. alba (Schlechtendal) Walpers of Mexico,
belong to subgenus Kingsboroughia. Meliosma alba
also occurs in south-central China, and it is the
only New World species of the genus with com¬
pound leaves. A few revisions of the genus in some
countries exist, e.g.. Flora lalis, petiolis 4—5 mm
longis et bracteolis pilis 1.5-2 mm longis. 8— It) seplatis
obtectis differt.
Shrub or small tree to 6 m tall; branches and
5 cm
5 mm
Figure 2. Aegiphila hystrieina Aymard & Cuello. — A. Flowering branch with detail of lower leaf surface. — B. Flower
subtended by the three bracteoles covered by the longer triehomes, ovary, pistil, and stigma represented by dotted
lines. — C. Medial section of flower showing the stamens with pistil removed. Drawn from the holotype, Koyama &
Agostini 7254 (VEN).
Volume 14, Number 1
2004
Aymard & Cuello
New Species of Aegiphila
23
Table 2. Comparison of the distinguishing character states of Aegiphila hystricina and A. arcta.
Characters Aegiphila hystricina Aegiphila arcta
Branches, branchlets, and peduncles
I, eaves
Petiole
Bracteole vestiture
Fruit
yellow-gray pubescent
elliptic or ovate-elliptic, 8—13 cm
long, 3—6 cm wide, glandulose vil¬
lose on both surfaces, glandulose
triehomes, ca. 2 mm long, 8— 9-sep-
tatc, margin densely ciliate
4— 5 mm
triehomes 1.5—2 mm long, 8— 10-sep-
tate
5— 6 mm long, sparsely glandular-pu¬
bescent
brown pubescent
oblanceolate or obovate, 10—18 cm
long, 5—8 cm wide, sparsely glandu
lose villose on both surfaces, glan¬
dulose triehomes, ca. 0.4 mm long,
2— 4-septate, margin sparsely ciliate
0.5—1 mm
triehomes ca. 5 mm long, 3— 4-septate
ca. 10 mm long, glabrescent
branchlets obtusely tetragonal, densely appressed-
villous w ith yellow-gray hairs on the younger parts,
glabrescent when mature. Leaves membranous, el¬
liptic or ovate-elliptic, 8—13 X 3—6 cm, apex acu¬
minate, base acute, margin entire, densely ciliate,
glandular villose on adaxial and abaxial surfaces,
denser on the midrib and secondary veins, glan¬
dular triehomes ca. 2 mm long, 8-9-septate; sec¬
ondary veins 8 to 10 per side, elevated on the ab¬
axial surface; petioles 4—5 mm long, densely
appressed-villous with yellow-gray hairs. Inflores¬
cence terminal thyrses, pendent, very congested,
peduncle elongate, 8—1 1 cm long, rachis and ped¬
icels densely appressed-villous with yellow-gray
hairs. Flowers subtended by three bracteoles at the
base of the pedicels, bracteoles linear, 5—10 mm
long, densely covered by glandular triehomes, 1 .5—
2 mm long, 8— 10-septate, pedicel 2—3 mm long;
calyx 1—1.5 mm long, sparsely glandular pilose ex¬
ternally, glabrous internally, the lobes 4, ca. 0.2 mm
long, triangular-ovate, ciliate-glandular, corolla
tube 6—10 mm long, yellow-green, glabrous on both
sides, the lobes 4, 2—4 mm long, imbricate, shortly
glandular-ciliate at the margins; the stamens 4, in¬
serted ca. 5 mm from the base of the tube, exserted,
filaments 8—10 mm long, glabrous, anthers oblong,
ca. 1 mm long; ovary obconical, ca. I mm long,
glabrous, style ca. 4 mm long, inserted, glabrous,
bifid, stigmas 2. Fruit globose, 5—6 mm long, ap-
pressed glandular-pubescent, mature calyx mem¬
branous and persistent, 3—4 mm long, sparsely
glandular-pubescent, subtended by the three per¬
sistent bracteoles.
Distribution and habitat. Aegiphila hystricina is
known from savanna and forest ecotone areas in Ve¬
nezuela anil farther southeast in the Rio Araguari,
Amapa state, in Brazil. In both regions, the species
occurs at elevations between 100 and 450 m.
Local names. “Hueraya dad eo” (Piaroa), from
./. Con t reras 117 ( V E N) .
Because of its four stamens and four corolla
lobes, terminal inflorescence, and calyx not deeply
dentate, Aegiphila hystricina belongs to section Ae-
giphila, group Paniculatae, subgroup Edentatae
(Moldenke, 1934; Ldpez-Palacios, 1977). flic inflo¬
rescence very congested, and the linear bracteoles
subtending the flowers, covered by glandulose tri-
chomes at the base of the pedicels, places this new
species next to A. arcta Moldenke, from Yaracuy
state in the Coastal Cordillera in Venezuela (Mol¬
denke, 1972). Aegiphila hystricina can be distin¬
guished from A. arcta by the characters indicated
in Table 2.
Herbarium specimens of Aegiphila hystricina
were previously referred to A. fendleri Moldenke, a
species from the mountain forests in the Andes of
Colombia and Venezuela. However., the new species
differs from A. fendleri in its leaves elliptic or
ovate-elliptic and the base acute (vs. lanceolate-
oblong to oblong and the base rounded or subcor-
date); petiole 4—5 mm long (vs. 7—12 mm long);
branches, branchlets, petioles, leaves on the lower
surface, and calyces covered by a yellow-gray in-
dument (vs. all parts of the plant covered by a fer¬
ruginous indument); and inflorescences thyrses (vs.
cymes).
Paratypes. VENEZUELA. Amazonas: Atures, alto
no Carinagua, 05°45'N, 67°20’W, ./. Contreras 117 (VEN).
Bolivar: Cedeno, Maniapure, Hato Pisurca, 06°55'N,
66°32'W, A. Fernandez 16938 (PORT). BRAZIL. Amapa:
Rio Araguari, betw. Cachoeiras Travessao & Santa Maria,
01°09'N, 51°52'W, J. M. Pires, W. Rodrigues A 6’. C. Ir¬
vine 50394 (NY).
Acknowledgments. We are grateful to H. van
der Werff (MO) and B. Manara (\ EN) for revising
the Latin descriptions; Victoria C. Hollowell and
two anonymous reviewers for comments on the
24
Novon
manuscript; B. Manara (VEN) for preparing the il¬
lustrations; and the Missouri Botanical Garden and
New York Botanical Garden staff for making their
laeilities available for our research.
Literature Cited
Berry, I’.. (). Huber A B. Holst. 1995. Holistic analysis
and phytogeography. Bp. 161—192 in I’. E. Berry, B.
Holst & K. Yatskievych (volume editors), Flora of the
Venezuelan Guayana, Vol. I (Introduction). Missouri
Botanical Garden I’ress, St. Louis.
Huber. (). 1994. Recent advances in the phytogeography
of the Guyana region. South America. Mem. Soe. Bio-
geogr. 4: 53—63.
- . 1995. Geographical and physical features. Bp. 1 —
62 in B. E. Berry, B. Holst & K. Yatskievych (volume
editors), Flora of the Venezuelan Guayana, Vol. 1 (In¬
troduction). Missouri Botanical Garden Bress, St. Louis.
Jansen-Jaeobs, M. J. 1988. Ease. 4. 148. Verbenaceae. In:
A. R. A. Corts-van Rijn (editor), Flora of the Guianas.
Koeltz, Konigstein.
Lopez- Balacios, S. 1977. Verbenaceae. Flora de Venezue¬
la, U.L.A., Merida.
Michelangeh. F. 2000. Species composition and species-
area relationships in vegetation isolates on the summit
of a sandstone mountain in southern Venezuela. J. Trop.
Ecol. 16: 69-82.
Moldenke, H. 1934. A monograph of the genus Aegiphila.
Brittonia I: 245—477.
- . 1953. Botanical exploration in Venezuela, Ver¬
benaceae. Field iana (Hot.) 28: 512—516.
- . 1972. A new species of Aegiphila from Venezue¬
la. Acta Bot. Venez. 6: 93—94.
France, C. T. & I). M. Johnson. 1992. Blant collections
from the Blateau of Serra do Araed (Amazonas, Brazil)
and their phytogeographic affinities. Kew1 Bull. 47: 1 —
24.
Senecio nemiae (Asteraceae, Senecioneae), a New Species from
Sierra Grande in Patagonia (Argentina)
Adriana Hartal i, 1 Roberta I). Tartosa,'2 and Susana E. Ere ire2'
‘Laboratories de Botanica “Lorenzo R. Parodi,” Facultad de Agronomfa, Universidad de
Buenos Aires, Av. San Martin 4453, 1417 Buenos Aires, Argentina.
cbartoli@mail.agro.uba.ar; tortosa@mail.agro.uba.ar
2Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET)
‘Division Plantas Vaseulares, Museo de La Plata, Paseo del Bosque, 1900 La Plata.
Argentina. freire@museo.fcnym. unlp.edu. ar
Abstract. A new species, Senecio nemiae, from
Sierra Grande, Rio Negro. Argentina, is described
and illustrated. Senecio nemiae most closely resem¬
bles S. adenotrichius from Chile, but is clearly dif¬
ferent by the shorter leaves without auricles and
smaller eapitula, with a calyculus of few bracts and
length shorter than the involucre. A new status, Se¬
necio sect. Senecio ser. Corymbosi subser. Adenotri-
chi, is also proposed to accommodate this taxon in
the previous classification of Senecio for Argentina.
Included is a key to those species of Senecio subser.
Adenotrichi with glabrous achenes.
key words: Argentina, Asteraceae, Compositae,
Patagonia, Senecio, Senecioneae.
Senecio L. has been subject to diverse taxonomic
interpretations varying from a single, broad genus
(Barkley, 1985) to numerous segregate genera (e.g.,
Nordenstam, 1978; Robinson & Bretell, 1973,
1974; Jeffrey, 1992). These new taxa were estab¬
lished based on microfeatures such as stylar ap¬
pendages, stigmatie surfaces, shape of filament col¬
lars, anther apex, anther base, and endothecial
tissue (Wetter, 1983; Vincent, 1996).
In the present paper, Senecio is treated in its
wide concept, containing about 3000 species world¬
wide, with the most diversity in the mountainous
areas or deserts of southern South America, tropical
South Africa, as well as central and eastern Asia.
Senecio s.l. includes monoecious herbs, subshrubs,
shrubs, vines, and small trees, with eapitula radi¬
ate, discoid, or rarely disciform; phyllaries of equal
length in one row, usually subtended by a calycu¬
lus; ray florets, if present, female with ligulate or
short filiform corollas; disc florets hermaphroditic
with tubular 5-dentate or 5-lobed corollas; the an¬
thers ecaudate or slightly tailed, filament collar
slender or dilated; pappus of many slender and sca-
brid bristles, and achenes 5- to 10-ribbed, glabrous
or pubescent.
On a regional basis, a large number of species
of Senecio has been treated by Cabrera in southern
South America, i.e., 208 from Chile (Cabrera,
1949), 90 species from Brazil, Paraguay, and Uru¬
guay (Cabrera, 1957), and 1 14 from Bolivia (Ca¬
brera, 1985). In Argentina Senecio is represented
by 266 native species, arranged in 3 sections and
16 series, with 1 14 species endemic to this country
(Cabrera el ah. 1999). Since the publication of this
treatment for Argentinian species, only one new
species was added (S. calingastensis Tombesi,
2001).
The new species described here belongs to sec¬
tion Senecio, series Corymbosi, which is character¬
ized by the eapitula disposed in terminal corymbs
and leaves sessile, attenuate into a pseudopetiole
or auriculate at the base. We include it within the
subsection Adenotrichi, established by Cabrera
(1949) for Chilean species of the genus, not cited
until now for Argentina. We propose the following
new status to accommodate this taxon in the clas¬
sification sustained by Cabrera et al. (1999).
Senecio sect. Senecio ser. Corymbosi subser.
Adenotrichi (Lindley) A. Bartoli, Tortosa &
S. E. Freire, stat. nov. Basionym: Adenotrichia
Lindley, Bot. Reg. 14: 1 19(1. 1828. Senecio
subsect. Corymbocephalus subsect. Adenotrichi
(Lindley) Cabrera, Lilloa 15: 380. 1949.
TYPE: Senecio adenotrichius DC.
The subseries Adenotrichi within Senecio appears
to be closely related to series Corymbosi subser.
Viscosi. Both subseries share glandular pubescence,
whereas the rest of the subseries of series Corym¬
bosi include species either tomentose or glabrous.
Subseries Adenotrichi differs from subseries Viscosi
in having deeply lobed to pinnatisect leaves (vs.
entire, crenate, or serrate leaves). In addition, sub¬
series Viscosi ranges from Brazil to northeastern Ar-
Novon 14: 25-28. 2004.
26
Novon
Figure 1. Senecio nemiae A. Bartoli, Tortosa & S. E. Freire. —A. Habit. — B, C. Leaves. — I). Phyllary. — E. Trichome.
— F. Capitulum. — G. Bay lloret. — H. Disk floret. — I. Anther. — J. Style branches. Scale bars: A = 3 cm; B-D = I
cm; E = 0.1 mm; F = 5 mm; G, H = 2 mm; I, J = 1 mm. Drawn from holotype, Correa el al. 7090 (BAB).
Volume 14, Number 1
2004
Bartoli et al.
Senecio nemiae from Patagonia
27
gentina, whereas subseries Adenotrichi occurs in
Chile and is now reported from southern Argentina.
Senecio nemiae A. Bartoli, Tortosa & S. K. Freire,
sp. nov. T\ PE: Argentina. Rio Negro: San An¬
tonio, Sierra Grande, 20 Oct. 1979, Correa el
al. 7090 (holotype, BAB). Figure I.
Quoad formam foliorum, fabricam capitulorum et in¬
dumentum ad Senecionem adenotrichium accedit, setl ah
ea differt capitulis, calyculo et foliis minoribus, bracteis
involucri in numero minori, foliis non aurieulatis et pilis
capitatis brevioribus.
Shrublet 20—25 cm, much branched at tin* base,
densely hirsute glandulose on stems, leaves, and
phyllaries, with glandular capitate trichomes, most¬
ly 0.5— 0.8 mm long; stems ascending. Leaves al¬
ternate, sessile, deeply pinnatilobed, 2.5—4 X 0.4—
0.6 cm, linear-elliptic, obtuse at the apex and
slightly dilated at the base, margin often revolute,
with midvein prominent, pubescent on both surfac¬
es; lobes 5 or 6 at each side, irregularly dentate,
acute and mucronate at the apex. Capitula 2 or 3
pedunculate, peduncles 1—2 cm long, arranged in
terminal corymbs; involucre calyculate, cylindric,
8-9 X ea. 8 mm; phyllaries 28 to 30, linear ovate,
8—9 X ea. 1 mm, dorsally pubescent, apex acu¬
minate with a tuft ol short trichomes; ealycular
bracts ca. 9, linear, ea. 6 mm long, dorsally pubes¬
cent. Ray florets ca. 15, female, corollas yellow,
ligulate, 3-dentate, ca. 6 mm long, the basal tube
pilose, 2—4 mm long, the limb 3-dentate, ca. 4 mm
long; disc florets ca. 50, hermaphroditic, corolla
yellow, tubular, 5-lobed, ca. 7 mm long, glabrous,
tube ca. 4 mm long, not basally broadened, limb
narrowly campannlate, only slightly broader than
the tube and not markedly distinct, throat ca. 2.2
mm long, lobes broadly triangular, ca. 0.80 mm
long, at base 0.5— 0.6 mm wide; anthers minutely
tailed, ca. 3 mm long; filament collars balusterform,
ca. 0.5 mm long; apical anther appendage ovate-
lanceolate, ca. 0.4 mm long; the basal tails ca. 0.2
mm long. Style branches penicillate, ca. 1.5 mm
long, apically truncate. Achenes oblong-obovate,
glabrous, ca. 1.8 mm (immature length), slightly
costate. Pappus bristles many, slender and scabrid,
ca. 7 mm long, white.
The specific epithet honors Maevia Noemi Cor¬
rea, Argentinian botanist, in recognition of her work
as director of Flora Patagdnica.
Distribution and habitat. Senecio nemiae is
known only from Sierra Grande, in the political
province Rio Negro of southern Argentina. Sierra
Grande represents a mountain range ol ca. 500 m
altitude, at approximately 42s south latitude, and
together with the Meseta de Somuncura constitutes
a particular district of the vegetation of Patagonia
since both regions contain a number of local en¬
demics, e.g., Adesmia serrana M. N. Correa (Fa-
baceae), Boopis raffaelli Spegazzini (Calyceraceae),
Conyza magnimontana Cabrera and Senecio rne-
gaoreinus Zardini (Asteraeeae), and Silene melan-
opotamica var. agrostophylla Pedersen (Caryophyl-
laceae).
Senecio nemiae is closely related to S. adenotri-
chius DC. from Chile. Both species have deeply
pinnatilobed leaves with dentate-acute lobes and
ligulate corollas with trichomes on the tube. How¬
ever. the new taxon has shorter leaves without au¬
ricles and smaller capitula with fewer phyllaries
and ealycular bracts.
Kk'i to Species w ith Glabrous Achenes ok Senecio
sect. Senecio ser. Corymbosi subser. Adenotrichi
la. Phyllaries few (ca. 13) . . Senecio rivularis J. Bcmy
l b. Phyllaries numerous (ca. 30 to 40).
2a. Leaves auriculate, 4—10 cm long; calyeulus
of many bracts as long as the phyllaries.
3a. Leaves deeply lobed, lobes dentate-
acute . Senecio adenotrichius DC.
3b. Leaves lobed, lobes entire or dentate,
obtuse to subacule .
. Senecio murinus Philippi
2b. Leaves without auricles, 2. ,5-4 cm long; ca-
lyculus of few bracts shorter than the phyl¬
laries . Senecio nemiae A. Bartoli,
Tortosa & S. E. Freire
Acknowledgments. We thank John Pruski and
an anonymous reviewer for reviewing the manu¬
script and for useful comments. Thanks to the cu¬
rator of BAB for the loan of specimens that made
this study possible, and to Victor H. Calvetti for
inking the illustration. We also thank Victoria C.
Hollowell for her accurate editorial work. This
study was supported by the Consejo Nacional de
Investigaciones Cientfficas y Tecnicas (CONICET),
Argentina.
Literature Cited
Barkley, T. M. 1985. Generic boundaries in Senecioneae.
Taxon 34: 17-21.
Cabrera, A. L. 1949. El genero Senecio (Compositae) en
Chile. Lilloa 15: 27—501.
- . 1957. El genero Senecio (Compositae) en Brasil,
Paraguay y Uruguay. Arch. Jard. Bol. Rio de Janeiro
15: 163-269.
- . 1985. El genero Senecio (Compositae) en Bolivia.
Darwiniana 26(1—4): 79—217.
- . S. E. Freire & L. Ariza Espinar. 1999. Tribu VIII.
Senecioneae. Tribu VIII bis. Liabeae, in A. T. Hunziker
(editor), Flora Eanerogamica Argentina 62: 3—180.
Jeffrey, C. 1992. The tribe Senecioneae (Compositae) in
the Mascarene Islands with an annotated world check-
28
Novon
list of the genera of ihe tribe, Notes on Compositae: VI.
Kew Bull. 47: 49-109.
Nordenstam, B. 1978. Taxonomic studies in the tribe Se-
necioneae (Compositae). Opera Bot. 44: 1-88.
Robinson, H. & R. I). Bretell. 1973. Studies in the Se-
necioneae (Asteraeeae), IV. The genera Mesadenia, Sy-
neilesis, Miricacalia, Koyamacacalia and Sinacalia.
Phytologia 27: 265—276.
- & - . 1974. Studies in the Senecioneae (As¬
teraeeae), V. The genera Psacaliopsis, Barkley anthus,
Telanthophora and Roldana. Phytologia 27: 402—439.
Tombesi, T. 2001. Novedades en Senecio (Senecioneae,
Asteraeeae). Hickenia 3(24—31): I I 1 — 1 14.
Vincent, P. L. I). 1996. Progress on clarifying the generic
concept of Senecio based on an extensive worldwide
sample of taxa. Pp. 597—611 in I). J. N. Hind & H. J.
Beentje (editors), Compositae Systematics. Proceedings
of the International Compositae Conference, Kew, Vol.
1. Royal Botanic Gardens, Kew.
Wetter, M. A. 1983. Micromorphological characters and
generic delimitation of some New World Senecioneae
(Asteraeeae). Brittonia 35: 1—27.
Bulbophyllum filifolium (Orchidaceae), a New Species from
Southeastern Brazil
Eduardo L. Borba* and Eric C. Smidt
Laboratorio < le Taxonomia Vegetal, Departamento de Ciencias Biologicas, l) n i vers it lade
Estadual de Feira de Santana, Rod. BR 116, km 03, 44031-460, Feira de Santana-BA,
Brazil. ^Author for correspondence: borba@gmx.net
ABSTRACT. Bulbophyllum filifolium, a new orchid
species from Brazil belonging to section Xiphizusa ,
is described and illustrated. The new species is
rare and endemic to the municipality of Grao Mo-
gol, Minas Gerais state, growing on rocks in campo
rupestre vegetation. Bulbophyllum filifolium is eas¬
ily recognized by its thread-like leaves, one-flow¬
ered inflorescences, and ovate petals with long
club-like appendages on the margin.
hey words: Brazil, Bulbophyllum, Orchidaceae.
Bulbophyllum Thouars is the largest orchid ge¬
nus, comprising over 1000 species throughout the
tropics (Vermeulen, 1991; Dressier, 1993). The
species occur mostly in the Old World, although
nearly 70 species are present in the New World.
Pabst and Dungs (1975, 1977) cited 54 species for
Brazil, and with the addition of some recently de¬
scribed species and new occurrences for this coun¬
try (e.g., Borba et al., 1998; Fraga, 1999; Toscano
de Brito, 2000), this number is increased to about
60.
Twelve of the Brazilian Bulbophyllum species be¬
long to section Xiphizusa (Reichenbach f.) Cog-
niaux, characterized mainly by the long fused se¬
pals. Those species are mainly epiphytic in forest
areas in southeastern Brazil, and occasionally lith-
ophytic in campo rupestre vegetation, as B. plu-
mosum (Barbosa Rodrigues) Cogniaux. The campo
rupestre of the Espinhago Range is the Brazilian
vegetation with the highest number of endemisms
and greatest diversity (Joly, 1970; Giulietti & Pir-
ani, 1988). Due to the discontinuity of these moun¬
tain ranges and outcroppings, many species, mainly
the 1 it hophy tic ones, are distributed in disjunct
populations. Because of that, some sites are espe¬
cially rich in the number of endemic species and/
or present populations differentiated (e.g., geneti¬
cally, morphologically, chemically) from the core
area of the species, as in the region of Grao Mogol
in the north of Minas Gerais (Giulietti & Pirani,
1988; Borba et ah, 2001a, 2001b, 2002). In the
course of a revision of Brazilian Bulbophyllum and
the population genetics and reproductive biology
studies of the orchids of Grao Mogol, we found a
newT species of Bulbophyllum in section Xiphizusa
with remarkable leathery thread-like leaves, de¬
scribed as follows:
Riilhopliyllum filifolium Borba & Smidt, sp. nov.
TYPE: Brazil. Minas Gerais: Grao Mogol. Ser-
ra do Barao, 16°33'S, 42°54'W, 7 Jan. 2002
(fl), E. L. Borba 1999 & C. van den Berg (ho-
lotype, IIUEFS). Figure 1.
Species liaee Bulbophyllo pabstii, B. bidentato , B. plu-
rnoso et aliis speeiebus sectionis Xiphizusa similis; ab om-
nis speeiebus folio filiforme inflorescenlia uniflora et com-
binatione labelli lobo medio carnoso. lobis marginibus
lateralibus dense curto-ciliatis ornatis et petalis ovatis ap-
pendicibus claviformibus ornatis longioribusque ad mar-
ginem differt.
Eithophytie herbs; roots 20--40 mm long,
thread-like, fasciculate; rhizome 2—8 X 1 mm,
creeping; pseudobulbs 4—8 X 6—7 nun, aggregate,
ovoid, slightly 3- to 4-angular, monophyllous.
Leaves 25—30 mm long, 1 nun diam., thread-like,
leathery, folded, erect, apex acute, base canalic¬
ulate and attenuate. Inflorescence basal, one-flow¬
ered; scape 40—50 mm long, curved, with 4
sheath-like bracts, 3—4 nun long, adpressed, not
imbricate; floral bracts 4X3 mm, ovate-lanceo¬
late, glah rous, apex acute; flower pendent; ovary
3 mm long, suleate; sepals membranaceous, sub¬
equal, erect-patent, boat-shaped, glabrous; dorsal
sepal 14 X 3 mm, linear-lanceolate, 3-nerved,
apex acute; lateral sepals fused up to the apex,
synsepal 14X5 mm, lanceolate, 7-nerved, apex
acute, slightly gibbous at base; petals 4X3 mm,
oblique-ovate, erect, 1-nerved, margins eiliate at
the base, with club-like appendages up to 1 mm
long increasing in size from base to apex; 1 i p 9 X
1.7 mm, fleshy, erect, parallel to column, movable;
lateral lobes 4 mm long, erect, auriculate, densely
Novon 14: 29-32. 2004.
30
Novon
XIaah
E
O
id
o
E
o
Figure 1. Bulbophyllum Jilifolium Borba & Smidt. — A. Dorsal sepal. — B. Fused lateral sepals. — C. Petal. — D.
Lip, lower view. — E. Lip, upper view. — F. Lip (below) and column (above), in side view. — G. Habit. Scale bars: 0.5
cm, A-F; 1 cm, G. Drawn from the holotype, Borba 1999 & ran den Berg, by Alarm Duraes.
Volume 14, Number 1
2004
Borba & Smidt
Bulbophyllum filifolium from Brazil
31
short-ciliate; mid-lobe 5 mm long, linear-elliptic,
adaxially with one distinct longitudinally sulcate
ridge, adaxial surface glabrous, apex rounded; col¬
umn 3 X 1.1 mm, with two sinuate stellidia at
apex and two falcate teeth on the ventral face;
anther versatile, 3-globose.
Etymology. This species is named for the ex¬
tremely narrow, linear (thread-like) leaves, the
narrowest among the Brazilian Bulbophyllum spe¬
cies.
Distribution. Bulbophyllum filifolium grows on
rock outcrops of quartzite in the eampo rupestre
vegetation of Serra do Barao, in the municipality
of Grao Mogol, in the state of Minas Gerais, Brazil,
a disjunct area of the main Espinhayo Range. The
eampo rupestre vegetation is characterized by
open, herbaceous vegetation on sandy, stony soils
mixed with herbs and shrubs growing on outcrop¬
pings of quartzite, sandstone, gneiss or “canga”
(iron) rocks (Giulietti & Pirani, 1988; Borba et ah,
2001a). This species appears to be endemic to the
Grao Mogol region and is rare, as only three in¬
dividuals have been found despite several field
trips made to the type locality. A fragment of the
same individual as the holotype is currently under
cultivation at the Universidade Estadual de Feira
Santana (another fragment of the same individual
and the other two individuals remain undisturbed
at the original locality). The Serra do Barao was
recently defined as a State Reserve (Parque Es¬
tadual de Grao Mogol), and B. filifolium possibly
occurs at other sites there. Further studies are
necessary to assess the conservation status of the
species.
Bulbophyllum filifolium belongs to section
Xiphizusa as suggested by its floral and vegetative
morphology, such as the erect rachis not genicu¬
late and not thickened, lateral sepals long and
connate to the apex forming a synsepal, a column
with two long stellidia at the apex and two small
teeth on the ventral face, and pseudobulbs aggre¬
gate and sulcate. The movable lip is a common
feature in the genus and probably plays an im¬
portant role in the pollination mechanism of this
species (wind-assisted fix pollination; Borba & Se-
mir, 1998), as observed in other species of section
Xiphizusa presenting similar flower structure (e.g.,
B. plumosum (Barbosa Rodrigues) Cogniaux and
B. bidentatum (Barbosa Rodrigues) Cogniaux; Ver-
ola, 2002). This species is easily recognizable by
its thread-like leaves and one-flowered inflores¬
cences, unique among the members of B. sect.
Xiphizusa. The other two Brazilian Bulbophyllum
species with narrow leaves are B. insectiferum
Barbosa Rodrigues and B. adiamantinum Brade,
belonging to sections Bulbophyllaria and Micran-
tha, respectively. However, conversely to B. fili¬
folium , these two species have succulent cylindri¬
cal leaves and multi-flowered inflorescences with
very distinct flowers; they also occur in Minas
Gerais, but are not sympatric with B. filifolium.
The long club-like appendages on the petal mar¬
gins are uncommon in other species of the section,
being observed only in B. pabstii Garay. However,
B. filifolium has ovate petals and the mid-lobe of
the lip is fleshy, whereas B. pabstii has ovate-lan¬
ceolate petals and the mid-lobe of the lip is mem¬
branaceous; it also differs by the oblong-lanceo¬
late leaf (ea. 10 mm w ide) and the multi -flowered
inflorescence. The lip mid-lobe of B. filifolium is
similar to that of B. bidentatum, but the latter dif¬
fers by the petals without the club-like append¬
ages, the lateral lobes of the lip not ciliate, the
oblong-lanceolate leaves (ca. 8 mm wide), and the
multi -flowered inflorescence.
Because we found only plants with old (lowers
we cannot describe the flower color accurately, but
they appear to be entirely purple or dark red and
similar to those of B. bidentatum.
Acknowledgments. Thanks are due to Cassio
van den Berg for the Latin diagnosis and other im¬
provements to the manuscript.
Literature Cited
Borba, E. I,. & J. Semir. 1998. Wind-assisted fly polli¬
nation in three Bulbophyllum (Orchidaceae) species oc¬
curring in the Brazilian campus rupestres. Lindleyana
13: 203-218.
- . - & F. Barros. 1998. Bulbophyllum ano¬
int urn Borba, Semir & F. Barros (Orchidaceae), a new
species from the Brazilian campus rupestres. Novon 8:
225-229.
- . J. M. Felix, V. N. Solferini & J. Semir. 2001a.
Fly-pollinated Pleurothallis (Orchidaceae) species have
high genetic variability: Evidence from isozyme mark¬
ers. Amer. J. But. 88: 419-428.
- . ,1. R. Trigo & .1. Semir. 2(H)] b. Variation of dia-
stereoisomeric pyrrolizidine alkaloids in Pleurothallis
(Orchidaceae). Biochem. Syst. Ecol. 29: 45—52.
- , G. .1. Shepherd, C. van den Berg & J. Semir.
2002. Floral and vegetative morphometries of five Pleu¬
rothallis (Orchidaceae) species: Correlation with tax¬
onomy, phylogeny, genetic variability and pollination
systems. Ann. Bot. 90: 219—230.
Dressier, B. I.. 1993. Phylogeny and Classification ol the
Orchid Family. Cambridge Univ. Press, Cambridge.
Fraga, C. N. 1999. Bulbophyllum gomesii Fraga (Orchi¬
daceae), uma nova espdcie da floresta atlantica do Es-
pfrito Santo. Brasil. Bradea 8: 135—138.
Giulietti. A. M. & J. R. Pirani. 1988. Patterns of geo¬
graphic distribution of some plant species from the
Fspinhayo Range, Minas Gerais and Bahia, Brazil. Pp.
39—69 in W. R. Heyer & P. F. Vanzolini (editors). Pro-
32
Novon
ceedings of a Workshop on Neotropical Distribution
Patterns. Academia Brasileira de Cieneias. Rio de Ja¬
neiro.
Joly, A. B. 1970. Conhecja a Vegeta^ao Brasileira.
EDUSP, Sao Paulo.
Pabst, G. K. .). & K Dungs. 1975. Orehidaceae Brasilien-
sis, Vol. I. Briicke-Verlag, Hildesheim.
- & - . 1977. Orehidaceae Brasiliensis, Vol.
2. Briicke-Verlag, Hildesheim.
Toscano de Brito, A. L. V. 2000. Two new species of Or-
chidaeeae from Brazil. Lindleyana 15: 184—188.
Vermeulen, .1. .). 1991. Orchids of Borneo, Vol. 2 — Btil-
bophyllum. Royal Botanic Gardens, kew.
Verola, C. F. 2002. Biologia Floral e Sistemas de Reprod-
m,ao em Fspeeies de Bulbophyllum (Orehidaceae)
Oeorrentes em Mata de (Valeria, Campo Rupestre e Flo-
resla Estaeional. M.Sc. Thesis, Universidade E^stadual
de Campinas. Campinas.
Paronychia revoluta , a New Species of Caryophyl laceae
from Brazil
Claudia Ciena Carneiro
Universidade Estadual de Feira de Santana, Departamento de Cieneias Biologicas,
Laboratorio de Mieromorfologia Vegetal (LAJV1 IV), BR I 16, km 3, Campus U niversitario,
CEP 44031-460, Feira de Santana. Bahia, Brazil, cameiro@uefs.br
Antonio Furl an
Universidade Estadual Paulista, Instituto de Biociencias, Departamento de Botanica, Av.
24A, 1515, Caixa Postal 199, Bela Vista, CEP 13506-900, Rio Claro, Sao Paulo, Brazil.
afurlan@rc.unesp.br
ABSTRACT. A new species of Paronychia, P. re¬
voluta, from the state of Rio Grande do Sul, Brazil,
is described and illustrated. This species is closely
related to P. camphor osmoides, distinguished from
it by linear-subulate leaves with a revolute margin
and prominent midribs. It is also allied to P. Jas-
ciculata, from which it differs in the lanceolate,
acuminate, fimbriate, and whitish stipules, and lin¬
ear-subulate leaves.
Key words: Brazil, Caryophyl laceae. Paronych¬
ia.
Paronychia Miller (Caryophyllaceae) comprises
about 1 10 species distributed among three subgen¬
era (Chaudhri, 1968; Bittrich, 1993). This genus is
characterized by the presence of stipules and
bracts, very small and clustered flowers often con¬
cealed by the bracts, filiform petals, episepalous
stamens, uniovulate ovary, utricle rupturing irreg¬
ularly at base, globose seeds, and a curved embryo.
Although occurring mainly in the Mediterranean
region, the species of Paronychia are widely dis¬
tributed throughout the world, inhabiting warm-
temperate, montane, and occasionally alpine re¬
gions. They may be found in dry stony or rocky
places, calcareous soils, mountain steppes or along
sea coasts and river banks.
In this paper a new Paronychia species, discov¬
ered during a study of Caryophyllaceae from Brazil,
is described and illustrated.
Paronychia revoluta C. E. Carneiro s and S. sipapoanus Maguire. There¬
fore, I have elevated S. duidae subsp. neblinae to
the species level.
Additional specimens examined. VENEZUELA. Ama¬
zonas: Dept. Rio Negro, Cerro Aracamuni, summit, Proa
Camp, 31 Oct. 1987, A'. Liesner A' G. Carnevali 22081
(MO, U); Sierra de la Nehlina, Rfo Yatua, escarpment
above Camp 4, 30 Dee. 1953, B. Maguire et at. .37007
(NY); Sierra de la Nehlina, Rfo Yatua, escarpment edge
betw. Cumbre Camp & N Escarpment, 18 Jan. 1954. B.
Maguire et al. .37240 (NY); Sierra de la Nehlina. Rfo Ya¬
tua, W of Cumbre Camp, 1-2 Dec. 1957. B. Maguire et
id. 4228.3 (NY); Dept. Rfo Negro, Sierra de la Nehlina.
above Rfo Marawinuma, E of Puerto Chimo camp, 26 Apr.
1984, W. Thomas 32.37 (CAS).
Styrax nunezii P. W. Eritscli, sp. nov. TYPE: Peru.
Cuzco: provineia La Convencidn, distrito
Eeharati, E Rfo Apurimac, NE Pueblo Libre,
Anchihuay and Bellavista Mountains, S Cor¬
dillera Vilcabaniba, 2445 m, 12°51'S,
73°30'W, 3 Aug. 1998 ((I), T. V. Nunez, W.
Nan ray, R. de la Colina & S. Udvardy 23320
(holotype, CAS; isotypes, AAU, RR. COL, F,
GH, K. LOJA, LPB, MICH. MO, NY, TEX.
US, W). Figure 3.
Arbor sempervirens. Lamina 7. 0-9. 3 X 3. 5-4.0 cm
crassi-chartacea ovata, apiee aeuminato, subtus pilis den-
sis minutis albi-stellato-tomentosis instrueta, magis dis-
persis sed densis lanatis atro-ferrugineo-stellatis braehiis
usque 0.2 mm longis intermixtis, venis quaternariis leviler
vel baud elevatis, paginis venarum non manifestis subter
indumentum, ubi veteribus pilis ferrugineis absentibus et
margine revoluto. Flos 10 mm longus; calyx 3 X 3—4 mm.
dense pallidus fusco-stellato-tomentosus, ad medium bra-
eliiis pilorum usque 0.2 mm longis; stamina 10; filament a
auriculata pilis albi-stellatis undulatis vel aliquantum rig-
idis braehiis usque 0.8 mm longis instrueta; anthera 3.5
mm longa loeulis lineari-triangularibus ad apicem con-
traetis; connectivum locula leviter superante; stylus filifor-
mis; ovula multa.
Evergreen tree 6 m tall; young twigs densely
brown stellate-tomentose. Petiole 10—1 I mm long;
lamina 7.0— 9.3 X 3. 5-4.0 cm, 1.8— 2.5 times as
long as wide, thiek-chartaeeous. ovate, secondary
veins 7 to 10 on each side of midvein; apex acu¬
minate; base broadly cuneate to subrounded; adax-
ially glabrous except on midrib; abaxially appear¬
ing light brown to the naked eye, with a dense base
tomentum of minute white stellate trichomes, and
more scattered but dense dark ferrugineous woolly
stellate trichomes with arms to 0.2 mm long, sur¬
face ol the veins not visible beneath the pubes¬
cence. ferrugineous trichomes often absent on older
leaves, tertiary veins slightly raised, quaternary
veins faintly or not at all raised; margin entire, rev-
olute on old leaves. Inflorescences axillary or ter¬
minal, racemose (or I - or 2-llowered or rarely pa¬
niculate), I to 3 per node, 2-3 cm long, 3- to
14-flowered, densely light brown stellate-tomen¬
tose; lower pedicels 4—6 mm long. Flowers her¬
maphroditic. 10 mm long; calyx 3 X 3-4 mm, eu-
puliform, densely light brown stellate-tomentose,
trichomes at mid-calyx with arms to 0.2 mm long,
margin truncate between the minute teeth, inner
margin eglandular; corolla 9 mm long; tube 1 mm
long, extending 1—2 mm beyond the calyx margin;
lobes 5, 9 X 1.5 mm, spreading, thickened; sta¬
mens 10; free portion ol stamen tube none; distinct
portion of filaments 1 mm long, ventrally with au¬
ricles bearing white, undulating to somewhat still
stellate trichomes with arms to 0.8 mm long, tri¬
chomes nearest the proximal end of the filament
with arms predominantly pointing downward, those
nearest the distal end with arms predominantly
pointing upward, also stellate-pubescent in sinuses
of filaments and dorsally, trichomes with arms to
1.3 mm long; anthers 3.5 mm long, the glabrous
connectives slightly exceeding the linear-triangular,
apically tapered, stellate-pubescent thecae; ovary
densely gray-green stellate-tomentose; stvle fili¬
form, glabrous; ovules many. Fruit unknown.
Habitat, distribution , and phenology: Known
only from the type collection (I have only seen flow¬
ers, but the specimen label states “young flowres
| sic ) and fruits”) in the cloud forest zone of the
Cordillera Vilcabaniba in southern Peru; flowering
August.
This species resembles Styrax rigidifolius Idrobo
& R. E. Schultes and several species of Styrax from
the G uayana Highland, such as .S’, guanayanus and
S. sipapoanus, in its ahaxial leaf surfaces. These
are abundantly covered with woolly ferrugineous
stellate trichomes in addition to the white stellate
base tomentum, rendering the surface light to dark
brown to the naked eye. Styrax nunezii is easily
distinguished from these species, however, by the
presence (vs. absence) of auricles on the ventral
portion of the stamen filaments. These auricles, as
well as the presence of linear-triangular (vs. linear)
anthers, allies S. nunezii with a group of Andean
species with the same combination of features (e.g..
S. pavonii A. DC., S. pentlandianus .). Remy, S.
peruvianas Zahlbruekner). No other species of this
group is known to have the type of abaxial leaf
surface possessed by S. nunezii. In addition, the
small (10 mm long) flowers and short (2-3 cm) in¬
florescences are unusual in the group.
The species is named for my colleague Percy
Nunez Vargas, curator of Herbario Vargas (CUZ) at
the Universidad Nacional San Antonio Abad del
Cusco, Cuzco, Peru, who collected the type and
Volume 14, Number 1
2004
Fritsch
Sty rax from South America
51
Figure 3. Styrax nunezii P. W. Fritsch. — A. Flowering branch. — B. Leaf, abaxial view. — C. Leaf surface, abaxial
view. — D. Stellate trichome from the base tomentum of the abaxial side of the leaf. — E. Larger stellate triehome from
the abaxial side of the leaf. — F. Flower. — G. Flower, median long-section. — H. Part of corolla + androecium, opened.
— I. Stamen, lateral view. Based on the holotype, Nunez el al. 23320 (CAS).
52
Novon
generously made specimens available to me for ex¬
amination. Additional isotypes are to be expected
at CUZ and USM.
Styrax rotundatus (Perkins) P. W. Fritsch, slat,
nov. Basionym: Styrax martii var. rotundatus
Perkins, Pflanzenr. 4. 241: 40. 1907. TYPE:
Brazil. Bahia: Jaeobina, ./. S. Blanchet 3614
(leetotype, selected here, BB: isotypes, C, F[2],
W; photo of C at F, MICH, MO).
Evergreen shrub to 3(4) m tall: young twigs
densely tawny, gray-brown, dark ferrugineous, or
brown stellate-tomentose. Petiole (2-)4— 8 mm long;
lamina 3.3— 5.8(-6.6) X 2. 1-4.3 cm, 1.3-2 times as
long as wide, thick-chartaceous to subcoriaceous,
elliptic, ovate-elliptic, ovate, or suborbicular, sec¬
ondary veins 4 to 8 on each side of midvein; apex
acute to subrounded; base broadly euneate, round¬
ed, truncate, or cordate; adaxially densely brown
stellate-tomentose when young, glabreseent except
on major veins; abaxially dark ferrugineous or
brown to the naked eye, with a dense base tomen-
tum of green-white minute stellate trichomes, and
abundant but more scattered woolly ferrugineous
trichomes with arms to 0.2— 0.3 mm long that are
sparse to absent in old leaves, tertiary and quater¬
nary veins distinctly differentiated from one anoth¬
er, distinctly raised, their surfaces visible through
the pubescence to completely obscured; margin en¬
tire, sometimes revolute. Inflorescences axillary or
terminal, racemose or paniculate (axillary inflores¬
cences occasionally 1- or 2-flowered), 1 or 2 per
node, 2—6 cm long, 3- to 8-flowered, subtending
leaves occasionally strongly reduced, densely gray,
dark ferrugineous, or brown stellate-tomentose;
lower pedicels 4—7 mm long. Flowers hermaphro¬
ditic, 10—13 mm long; calyx 3.5—5 X 2.5— 3.5 mm.
cupuliform to narrow-cupuliform, densely gray-
green stellate-pubescent mixed with larger evenly
scattered woolly golden or brown stellate trichomes
with arms at mid-calyx to 0.5 mm long, margin
truncate between the minute teeth, inner margin
eglandular; corolla 8-1 I mm long, white; tube 2-3
mm long, extending up to the calyx margin; lobes
5, 5—10 X 0.5— 1.5 mm, recurved to reflexed, thin;
stamens 10; free portion of stamen tube none; dis¬
tinct portion of filaments 2 mm long, ventrally with¬
out auricles, bearing a dense mass of white stellate
trichomes with thin, ± straight arms to 1.5 mm
long, trichomes nearest the distal end of the fila¬
ment with arms predominantly pointing upward,
those nearest the proximal end with arms predom¬
inantly pointing downward, dorsally glabrous, tri¬
chomes along the edges of the filament and at the
sinus between filaments with arms to 2 mm long;
anthers 3—6 mm long, the glabrous connectives
slightly exceeding the linear, apieally tapered, stel¬
late-pubescent thecae; ovary densely golden yellow
stellate-tomentose; style filiform, glabrous; ovules
many. Drupe 5—7 X 4—6 mm, gray-green to dull
orange, subglobose, apex slightly to strongly de¬
pressed, annular-suleate around the remains of the
style base; wall irregularly and coarsely rugose on
herbarium specimens; fruiting calyx 3—5 X 4—6
mm, 0.4— 0.8 times the length ol the drupe proper.
Habitat, distribution , and phenology. Endemic
to Brazil, occurring in the states ol Bahia and Minas
Gerais in campo rupestre, among rocks, rarely on
sandy soil at 900— 1850(— ?2000) m; flowering No-
vember-May, fruiting March-June.
Styrax rotundatus was originally described as a
variety ol S. martii by Perkins (1907; as S. martii
var. rotundatus), who distinguished it from the typ¬
ical variety by its orbicular leaves with rounded or
broad-acuminate apices. In my view, the two taxa
differ consistently not only through the shape of the
leaves (although more subtly than Perkins has sug¬
gested, as indicated in the descriptions herein), but
also through numerous other characters that, taken
together, have prompted me to recognize .S', martii
var. rotundatus at the specific level. The following
characters serve to delimit S. rotundatus from .S'.
martii: petiole (7-) 10— 19 mm long (vs. (2— )4— 8 mm
long), lamina 3.3-5.8(-6.6) cm long (vs. 5.9-11.7
cm long), tertiary and quaternary veins distinctly
differentiated from each other and conspicuously
raised (vs. not well differentiated from each other
and moderately raised), lower pedicels 4—7 mm
long (vs. 1—5 mm long or nearly absent), calyx 3.5—
5 X 2.5— 3.5 mm (vs. 5—8 X 4—5 mm), not striate
(vs. striate) proximally. flowers 10-13 mm long (vs.
13—18 mm long), corolla tube 2—3 mm long (vs. 5—
8 mm long), corolla lobes 0.5— 1.5 mm wide (vs.
1 .5— 2.5 mm wide) and recurved to reflexed (vs.
spreading), and drupes 5—7 X 4-6 mm (vs. 9—16
X 6—7 mm), gray-green to dull orange (vs. olive).
In addition, S. rotundatus is always a shrub, where¬
as S. martii can be either a shrub or a tree; S.
rotundatus occurs exclusively in campo rupestre
vegetation, whereas S. martii can occur in campo
rupestre but also occurs in forests, forest edges, and
campo de altitude. The ranges of the two species
are nearly non-overlapping, with .S’, rotundatus oc¬
curring from extreme northern Bahia to south-cen¬
tral Minas Gerais, and S. martii occurring from
south-central Minas Gerais to northern Rio Grande
do Sul. The two species are apparently sympatric
Volume 14, Number 1
2004
Fritsch
Styrax from South America
53
only in the Serra do Cipd in south-central Minas
Gerais, Brazil.
Perkins (1907) listed two collections in the pro-
tologue of Styrax martii var. rotundatus: Blanchet
234 and Blanchet 3614. 1 have leetotypified this
taxon on the BR specimen of Blanchet 3614 be¬
cause it was annotated by Perkins, and because
duplicates of Blanchet 3614 are apparently more
widely distributed than those of Blanchet 234 (1
have detected only two sheets of Blanchet 234. from
P and W, whereas Blanchet 3614 is housed at BR,
C, F, and W).
Additional specimens examined. BRAZIL. State not
specified: Glocker 234 (BM (the specimen has the same
appearance and is at the same stage of development as
collections of Blanchet 234 ; the numeral “ 234 ” is written
on a label separate from another containing the words
“Glocker. Brazil. Ex Herb. Shuttleworth.,” suggesting that
this is actually a duplicate of Blanchet 234 that has been
incorrectly attributed to Glocker)); C. F. P. von Martins
s.n. (M no. 47). Bahia: Jacobina, ra. 24 km from Jacobi na
to Morro do Chapeu, 28 Oct. 1995, A. M. A. Amorim el
al. IBOI (CAS[3]); Palmeiras, Morro do Pai Inayio, 25 Jan.
1998, A. M. A. Amorim et al. 2137 (CAS); Utinga. Morro
Felix Joao (Morro da Torre da Embratel), 14 km from Utin¬
ga. 16 Mar. 1985, A. P. de Araujo & IE N. da Fonseca
404 (UB); Morro do Chapeu. 27 Aug. 1980, //. P. Bautista
414 (K). 5 Apr. 1984. H. P. Bautista & O. A. Salgado 941
(GLIA. MBM, RB), ./. S. Blanchet 234 (P. W); Bomfim,
Mount Taboa, 8 May 1918. II. M. Curran 174 (GH. NY,
US); 34 km E of Morro do Chapeu along Highway BA-
052, Chapada da Diamantina, 3 Apr. 1976, G. Davidse et
al. 1 1303 (MO); Morro do Chapeu, 2 Mar. 1997, F. Franga
et al. 5924 (k, W); 11-13 km from Mucuge along the new
A ndaraf— Mucuge road. 8 Sep. 1981, A. Furlan et al. 2120
(k); Rio tie Contas, Pico das Almas, 14 Dec. 1984,4. M.
Giulietti et ex of all five anthers. The two novelties bear
apical triehomes only on the ventral pair of anthers.
In point ol fact, there is no a priori reason to restrict
the search for relatives of C. duvalliorum and C. mar¬
itae to species with dissected adult foliage. The avail¬
able evidence suggests that deeply dissected adult fo¬
liage has evolved several times within Cyanea
(Carlquist, 1974; Lammers, 1990a, 1990b; Givnish et
ah, 1994, 1995). Cyanea aspleniifolia and C. ship¬
manii are related to species with toothed leaves, in¬
cluding C. dunhariae Rock and C. profuga C. N.
Forbes (Lammers, 1990b), while the closest relative
ol the C. grimesiana complex appears to be C. loliata
H. Mann, in which the adult leaves are merely lolled
(Lammers, 1990a). For these reasons, species with
non-dissected foliage must also be considered as po¬
tential relatives of the novelties.
Detailed consideration of the entire genus re¬
vealed that there are two informal species groups
in addition to the C. grimesiana complex, which are
characterized by many of the same features found
in C. duvalliorum and C. maritae: frequent produc¬
tion of prickles; relatively well developed bracts;
relatively large white flowers; corolla tubes stout,
arcuate, laterally compressed, and tallest above the
middle; and orange or yellow berries crowned by
the persistent calyx lobes.
The first consists of the ten species formerly seg¬
regated as the genus Rollandia (Lammers et ah,
1995; Lammers. 1998a): C. calycina (Chamisso)
Lammers; C. crispa (Gaudichaud) Lammers, Giv¬
nish & Sytsma; C. humboldtiana (Gaudichaud)
Lammers. Givnish & Sytsma; C. koolauensis Lam¬
mers, Givnish Sytsma; C. lanceolata (Gaudi¬
chaud) Lammers. Givnish N Sytsma; C. longiflora
(Wawra) Lammers, Givnish & Sytsma; C. parvifolia
(C. N. Forbes) Lammers. Givnish & Sytsma; C. pur-
purellifolia (Rock) Lammers, Givnish cK Sytsma; C.
sessilifolia (0. Degener) Lammers; and C. st.-johnii
(Hosaka) Lammers, Givnish & Sytsma. These
plants are unique within the genus in the dorsal
adnation of the filament lube to the corolla tube
and further differ from the C. grimesiana complex
in their lack of dissection of the adult foliage; mar¬
gins are entire or more often merely toothed. How¬
ever, this complex does show some developmental
dimorphism. Four species (C. calycina, C. lanceo¬
lata, C. longiflora, and C. sessilifolia ) produce dis¬
tinctive juveniles, though in these the margins of
the juvenile leaves are merely lohed rather than
parted or divided. Biogeographieally, the species of
the Rollandia complex are all endemic to O'ahu,
except for C. parvifolia, which is endemic to kaua'i;
no representatives occur on or near East Maui.
The second informal group consists of five spe¬
cies found on the two youngest islands of the ar¬
chipelago: C. glabra (E. Wimmer) St. John, C. lon-
gissima (Rock) H. St. John, C. mceldowneyi Rock
(all from East Maui), C. platyphylla (A. Gray) 1 1 i I-
lebrand (windward Hawai'i), and C. scabra Hille-
brand (West Maui). Like the Rollandia complex,
the C. scabra complex differs from the C. grimesi-
ana complex in its lack ol dissection of adult fo¬
liage and the presence of developmental dimor-
Volume 14, Number 1
2004
Lammers
Endemic Hawaiian Genus Cyanea
89
phism. As before, llie species that do produce
distinctive juveniles ( C . glabra, C. mceldowneyi,
and C. platyphylla ) bear leaves that are merely
lobed rather than parted or divided.
The relationships of C. duvalliorum and C. maritae
lie somewhere among these three informal groups. 01
them, the Rollandia complex seems the most distantly
related to C. duvalliorum and C. maritae. None of its
species has leaves, adult or juvenile, that even ap¬
proach the two novelties in degree of dissection, while
the novelties lack the unique adnation of the staminal
column that characterizes the group; furthermore, the
species of the Rollandia complex occur three or four
islands away in a very' different biogeographic setting.
The C. grimesiana and C. scabra complexes lack this
adnation of the staminal column and are represented
on East Maui.
It is more difficult to decide between the C. gri¬
mesiana and C. scabra complexes, because their
distinctness is blurred to some extent by C. duval¬
liorum and C. maritae. Though the C. grimesiana
complex would seem more similar to the novelties
due to its dissected adult loliage, it does not show
any tendency toward developmental dimorphism.
This seems to me to reflect a fundamental differ¬
ence in life history. Furthermore, the novelties dif¬
fer from C. grimesiana and most of its allies in their
lack of apical pubescence on the three dorsal an¬
thers. Eli is feature is unique to these species within
Cyanea (and very rare in the subfamily) and seems
a valuable indicator of affinity. Biogeographically,
this complex is represented on East Maui by just
one of its five species.
Though the C. scabra complex shows no dissec¬
tion of the adult lamina, three of its five species do
produce distinctive juveniles. In all its species, the
anthers bear triehomes only on the ventral pair, as
in the novelties. Furthermore, East Maui is the cen¬
ter of diversity for this complex, with three of its
five species endemic there. Overall, C. duvalliorum
and C. maritae do seem more closely related to the
members of the C. scabra complex.
Within this complex, one would assume that the
two new species are each others closest relative, as
they are the only members of the group with parted
or divided adult leaves. 01 the species described pre¬
viously, C. scabra and C. mceldowneyi seem most
closely related to the two novelties, based on their
common possession of persistent narrowly oblong
bracts and calyx lobes, rounded at apex (see key be¬
low).
It is noteworthy that C. duvalliorum possesses a
feature unique in the genus: the concrescence of
the apical triehomes on the ventral anthers into a
triangular scale. An identical scale, formed in the
same manner, characterizes a Latin American taxon
of Lobelioideae, Centropogon C. Presl subg. Cen¬
tro/ logon (Lammers, 1998b). Like the branched tri-
chomes found in Cyanea calycina and Centropogon
sect. Siphocampyloides Bentham (Lammers, 1998b,
2002; Batterman & Lammers, in press), this is ap¬
parently a case of parallel evolution.
Although the plants discovered at WaLoltiwi
Stream could not be equated with any named spe¬
cies, they do appear to match a number of speci¬
mens collected on East Maui by earlier botanists.
These specimens were sterile or otherwise inade¬
quate for confident identification, and languished
in herbaria under various names for many years.
Of these earlier specimens, Rock 8797, 8797a,
and 8798 served as paratypes of C. scabra var. var-
iabilis Rock. Although these gatherings are referable
to C. duvalliorum (8797) and C. maritae (8797a,
8798). the holotype of C. scabra var. variabilis is a
specimen of C. longissima and that name is properly
treated as a synonym of this species. Previously
(Lammers, 1990a), all these paratypes were regarded
as juveniles of C. glabra. However, I subsequently
realized that the leaf margin of true juvenile C. gla¬
bra (e.g., Forbes I657.M, BISH, K. L. RSA. S) is
only shallowly lobed, not deeply dissected.
The remainder of the early sterile collections ol
C. duvalliorum had been identified tentatively as
C. horrida. Though these two species are very sim¬
ilar in morphology of both juvenile and adult
leaves, sterile specimens may be distinguished on
the basis of pubescence. In C. horrida, the leaves
are sparsely pubescent adaxially and more densely
so abaxially; in C. duvalliorum, pubescence is con¬
fined to the abaxial venation or totally absent.
These earlier collections considerably extend the
distributional range of both C. duvalliorum and C.
maritae on the windward slopes of Haleakala. Some
were collected 7—12 km east of Wai’ohiwi Stream,
from Honomanu Stream to Wailua Iki Stream; oth¬
ers were obtained approximately 27> km east, at Kt-
pahulu Valley. Efforts should be made to ascertain
whether any of these populations is still extant.
Cyanea duvalliorum Lammers & H. Oppenheimer.
sp. nov. Type: U.S.A. Hawaiian Islands: East
Maui, Makawao District, ko'olau Forest Re¬
serve, Wai'ohivvi. 20°50'04”N, 156°15'55"W,
mesic forest, steep shady slope, 2800 ft.. 15
Oct. 2000. //. Oppenheimer & F. Duvall
H100026 (holotype, BISH [sheet plus liquid);
isotypes, B. k, MO, NY, OSH. ITBG, US). Fig¬
ure 2.
Species of) marginem foliorum maturorum pinnatim (is-
90
Novon
Figure 2. Cyanea duvalliorum Lammers & H. Oppenheimer. — A. Adult leaf. — B. Juvenile leaf. — C. Flower. — I).
Fruil. (A, C drawn from the holotype, Oppenheimer & Duvall H 100026: B. I) drawn from a paratype, Oppenheimer el
al. HI 0003.)
sum partition vel divisum el corollae tubum compressum
arcuatum supra medium altissimum affinitatem Cyaneae
grimesianae simulans, sed ob plantas juveniles proprias
et tubum antherarum cum trichomatibus apiealibus solum
in pari ventrali gregem C. seahrae vero pertinet; inter C.
scabram et species affines habitu arboreo 4—6 m alto, rtiar-
gine foliorum maturorum pinnatim partito vel fisso basin
versus pinnato, bracteolis longioribus 4—5 mm longis,
bracteis longioribus 10—13 mm longis, ealyeis lobis lon¬
gioribus 13—19 mm longis, corolla hypanthio triplo vel
quadruplo longiore, staminali columna pubescenti, tubo
filamentorum antheris dorsalibus triplo longiore, antheris
dorsalibus longioribus 1 1-12 mm longis, antheris ven-
tralibus ad apicem squama triangularis formata trichom-
Volume 14, Number 1
2004
Lammers
Endemic Hawaiian Genus Cyanea
91
atihus concrescentibus, et baccis longioribus 17-21 mm
longis distinguenda.
Trees, 4—6 m tall, producing distinctive juveniles
but these not flowering precociously; stems of ju¬
veniles 0.7— 1.2 cm diam., unbranched, armed with
numerous prickles consisting of a sharp trichome
atop a bulbous multicellular base; stems of adults
1.6—2 cm diam. below the leaves and inflorescenc¬
es, 4—21 cm diam. at base, branching about 1 m
above ground, the 4 to II branches diverging from
trunk at a 45°— 60° angle, ascending, sometimes
branching sparingly once again, armed with stout
conic prickles below, becoming smooth toward
apex, sometimes producing short shoots with juve¬
nile foliage; latex white. Juveniles with lamina el¬
liptic or oblanceolate, 13—27 cm long, 7—17 cm
wide across the segments, 0.2— 0.4 cm wide be¬
tween the segments; upper surface dark green and
glabrous; lower surface light green, glabrous, armed
with prickles consisting of a sharp trichome atop a
bulbous multicellular base; margin pinnately divid¬
ed. becoming pinnately compound at base, the seg¬
ments oblong or elliptic, 1—9 X 0.7— 2.8 cm, 8 to
10 per side, separated by sinuses, their margin pin¬
nately lobed, cleft, parted, or divided (these sec¬
ondary segments oblong or narrowly triangular, 2—
18 X 2—10 mm, the apex obtuse); apex obtuse;
petiole 3—4 cm long, 0.2— 0.3 mm diam., X/-—Vi as
long as the lamina, armed with prickles consisting
of a sharp trichome atop a bulbous multicellular
base. Adults with lamina elliptic or oblanceolate,
31—53 cm long, 10—20 cm wide across the seg¬
ments, 0.5— 4.7 cm wide between the segments; up¬
per surface dark green and glabrous; lower surface
light green and pubescent on veins with short stiff
curved hairs; margin pinnately parted or divided,
becoming pinnately compound at base, the seg¬
ments oblong or elliptic, 1.5—12.5 X 0.5—4 cm, 10
to 14 per side, separated by sinuses, their margin
pinnately cleft or parted (these secondary segments
deflate or triangular, 2—22 X 2—18 mm, the apex
acute or acuminate); apex acute or acuminate; pet¬
iole 4.5— 6.5 cm long, 3.5—9 mm diam., Yu—Vs as
long as the lamina, minutely pubescent. Racemes
6- to 10-flowered, arising below the apical rosette
of leaves, the flowers all blooming ± simultaneous¬
ly; peduncle and raehis horizontal, 35—54 mm long,
4—9 mm diam., glabrous; bracts narrowly oblong,
10-13 X 1.5-1. 6 mm, subtending the pedicels,
sometimes a few empty ones on the peduncle, de¬
ciduous, rounded at apex; pedicels ascending, 9—
14 mm long, 0.8—1 mm diam., medially or suba-
pically bibracteolate, glabrous; bracteoles narrowly
oblong, 4—5 X 0.8— 1.2 mm. Hypanthium obconic.
10—12 mm long, 5-9 mm diam., Vg-Vs as long as
the corolla, glabrous, attenuate at base; calyx lobes
narrowly oblong, 13—19 X 2.6—5 mm, abutting at
base but distinct, erect, glabrous, the margin entire,
rounded and apiculate at apex; corolla white, 38—
47 mm long, glabrous; corolla tube arcuate, later¬
ally compressed, 23—28 mm long, 5—6 mm tall at
base, narrowing gradually to 3—4 mm tall at mouth,
cleft dorsal ly for ca. half its length; dorsal corolla
lobes spreading, 16—19 X 1 mm, ca. % as long as
tube; ventral corolla lobes deflexed. 13—15 X 1
mm; filament tube 32—40 mm long, 1.3— 1.5 mm
diam., pubescent with long white hairs, free from
the corolla; anther tube gently decurved, 2.3— 2.7
mm diam., densely pubescent at base and on the
connectives; dorsal anthers 11 — 12 mm long, ca. Vi
as long as the filament tube; ventral anthers 7.5—
8.5 mm long, their apex crowned by a triangular
scale 3—4.5 mm long, formed from loosely eoneres-
eent white trichomes. Berries bright orange, ellip¬
soid, 17-21 mm long, 10—15 mm diam., crowned
by the persistent calyx lobes; seeds brown, broadly
ellipsoid, 0.7 mm long, 0.5 mm diam., the testa
Type C of Murata (1992. 1995), smooth, shiny.
Distribution, habitat, and phenology. Endemic
to windward East Maui in the Hawaiian Islands.
Recent fertile collections are from rnesic forests on
the northwestern slopes of Haleakala in the
Wai‘ohiwi watershed at an elevation of 855 m; more
than 50 mature trees and numerous juveniles were
observed here during 2000—2002. Sterile collec¬
tions were obtained from the same general region
and points to the east at slightly higher elevations
(1070-1522 m) in 1911, 1920, 1927, and 1980.
Sterile specimens were also collected farther east,
at Klpahulu Valley, in 1919. This species flowers
during October, and the fruits ripen during Decem¬
ber and January.
Etymology. Cyanea duvalliorum is named for
two of its discoverers, ornithologists Renate Gass-
mann-Duvall (b. 1946) and her husband Fern Du¬
vall (b. 1953), who are extremely active in the Ha¬
waiian conservation community. They have brought
the new species (and others) into cultivation and
taken the lead in securing its preservation.
Paratypes. U.S.A. Hawaiian Islands. Flast Maui. !Y1ak-
awao: above Kula pipe line, Munro 724A (BISH): Hale¬
akala pipe line, Munro 724H (BISH. NY, FS), 6. C. Munro
725 (BISH); Ha‘iku Uka, Wai'ohiwi Stream, Oppenheimer
et al. HI 1)005 (BISH [2 sheets plus liquid . MO. NY, OSH
[sheet plus liquid], FTBG, US); Ha'iku l ka, tributary of
Wai‘ohiwi Stream, Oppenheimer et al. H10012 (BISH.
OSH, PTBG); Vt of "Opana Gulch across stream, War-
shauer & McEldowney 2754 (BISH). liana: ridge left kl-
pahulu, Forbes I72H.M (BISH), Forbes I729.M (BISH);
92
Novon
trail K ol Ke‘anae & up ridge between W & E Wailua lki.
Forbes 2659. M (I5I5H); Ke'anae Valley, Rock 8797 (BISH).
Cyuncu maritae Lammers & H. Oppenheimer, sp.
nov. TYPE: U .S.A. Hawaiian Islands: East
Maui. Makawao District, Ko‘olau F. R.,
Wai‘ohiwi drainage basin, 2800 ft., growing
above the confluence of two intermittent trib¬
utaries, near the main stream, 20°50'06"N,
156°15'52"W, degraded mesic forest amidst
forestry plantings of Eucalyptus robusta & E.
globulus, 26 Nov. 2000, H. Oppenheimer & F.
Duvall HI 10029 (holotype, BISH; isotypes,
OSH, PTBG). Figure 3.
Species ob marginem foliorum maturorum pi i mat i m lis-
sum partitum vel divisum, calycis lobum et fforum brac-
team oblongum, et corollae tubum arcuatum compressum
af finis Cyaneae duvalliorum , sed ab hac specie habit u fru-
ticoso, ramis basalibus paucioribus 2-4 valde assurgen-
tibus, foliis juvenilibus et maturis cum segmentis trian¬
gularis serrulatis 5—8 utrobique, petiolis foliorum
maturorum proportione longioribus admodum Vs—Yz lami-
nam aequantibus, hypanthio proportione breviore modum
Yf— Zb corollam aequanti, calycis lobis f)revioribus 3. 5-9.5
mm longis, corolla longiore 59—61 mm longa cum lobis
dorsalibus Vi—% tubum aequantibus, filamentorum tubo
antheris dorsalibus quadruplo vel quintuplo longiore, et
baccis parvioribus 12—13 mm longis bene distinguenda.
Shrubs, 2—2.4 m tall, producing distinctive ju¬
veniles but these not flowering precociously; stems
of juveniles unbranched, armed with numerous
[trickles consisting of a sharp triehome atop a bul¬
bous multicellular base; stems of adults 0.7—1 .5 cm
diam. toward apex, 6.3-7. 4 cm diatn. at base, pro¬
ducing 2 to 4 strongly ascending branches from
near the base (sometimes unbranched), armed with
stout conic prickles below, becoming smooth toward
apex; latex white. Juveniles with lamina broadly el¬
liptic, 6.8—15 cm long, 4.6— 8.7 cm wide across the
segments, 2.3— 3.8 cm wide between the segments,
both surfaces armed with prickles consisting of a
sharp triehome atop a bulbous multicellular base;
upper surface dark green; lower surface light green;
margin pinnately parted; segments triangular, 0.5—
2 X 0.3— 1.8 cm, 5 to 8 per side, separated by
sinuses, their margin minutely serrulate, acuminate
at apex; apex acuminate; base truncate; petiole 2—
4.7 cm long, 1.2—2 mm diam., ca. Vs as long as the
lamina, armed with prickles consisting of a sharp
triehome atop a bulbous multicellular base. Adults
with lamina 17—37 cm long, 8.5—10 cm wide across
the segments, 1.3—10 cm wide between the seg¬
ments, elliptic or oblong, both surfaces armed with
[trickles consisting of a sharp triehome atop a bul¬
bous multicellular base; upper surface dark green;
lower surface light green, glabrous or pubescent on
veins with short stiff curved hairs; margin pinnately
cleft, parted, or divided; segments narrowly trian¬
gular, triangular, or deflate, or shallowly triangular,
often falcate, 0.6-10 X 0.8-4. 5 cm wide, 6 to 8
[ter side, separated by sinuses, their margin mi¬
nutely serrulate, acuminate or acute and apiculate
at apex; apex acuminate; base cuneate or obtuse;
petiole 10—13.5 cm long, 3—4.3 mm diam., V3—Y2 as
long as tilt' lamina, sparsely armed with prickles
consisting of a sharp triehome atop a bulbous mul¬
ticellular base. Racemes 9- to 45-flowered over
their lifetime, but only 4 to 8 flowers present at any
given time, arising within the apical rosette of
leaves; peduncle and rachis horizontal, 45—90 mm
long, 2.5—4 mm diam., sparsely armed with prickles
consisting of a sharp triehome atop a bulbous mul¬
ticellular base; bracts oblong. 1.5— 2.2 X 0.8-1 mm,
subtending the pedicels, deciduous, rounded or ob¬
tuse at apex; pedicels spreading, 12—25 mm long,
0.9—1 mm diam., ebracteolate, glabrous. Hypanthi-
um campanulate or obconic, 9—9.5 mm long, 7-9
mm diam., */7— V6 as long as the corolla, glabrous,
cuneate at base; calyx lobes oblong, 3.5— 9.5 X 1.5—
3 mm, abutting at base but distinct, erect, glabrous,
the margin entire, acute or rounded and apiculate
at apex; corolla white tinged with pink, 59-61 mm
long, sparsely pubescent, especially on the lobes;
corolla tube arcuate, laterally compressed, 42—46
mm long, 5—6.3 mm tall at base, increasing to 7—
9.2 mm tall above the middle before decreasing to
3.8— 5.5 mm tall at mouth, cleft dorsal ly for ca. Yi
its length; dorsal corolla lobes spreading. 14-18 X
1.5—2 mm, Yi—Y5 as long as tube; ventral corolla
lobes deflexed, I 1-14 X 0.9-1. 7 mm, connate ba-
sally for 5—6 mm; filament tube 50—51 mm long,
1.8— 2.1 mm diam., glabrous, Iree from the corolla;
anther tube gently decurved, 2.8— 3.4 mm diam. at
base, tapering to 1.5—2 mm diam. at mouth, gla¬
brous; dorsal anthers I 1—12.5 mm long, Vc—Ya as
long as the filament tube; ventral anthers 8-9 mm
long, their apex crowned by a tuft of white tri -
ebonies 2.5—3 mm long. Berries yellowish orange,
broadly ellipsoid or obovoid, 12—13 mm long, 8.5—
10.5 mm diam., crowned by the persistent calyx
lobes; seeds brown, broadly ellipsoid, 0.7 mm long,
0.5 mm diam., the testa Type C 4 >1 Murata (1992,
1995), smooth, shiny.
Distribution, habitat, and phenology. Endemic
to windward East Maui in the Hawaiian Islands.
Recent fertile collections are from mesic forests on
the northwestern slopes of Haleakala in the
Wai‘ohiwi watershed at an elevation of 855 m; less
than 20 adult and juvenile plants were observed
here during 2000—2002. Sterile specimens were
collected from the same general region and points
Volume 14, Number 1
2004
Lammers
Endemic Hawaiian Genus Cyanea
93
Figure 3. Cyanea marline Lammers & II. Oppenheimer. — A. Adult leaf. — B. Juvenile leaf. C. flower. (A. (,
drawn from the holotype, Oppenheimer & Duvall HI 10029; B drawn from a paratype, Oppenheimer A Duvall HI 0005.)
to the east in 191 1, and farther east at Klpahulu
Valley in 1919. This species flowers during Novem¬
ber and December, and the berries ripen by May.
Etymology. This species is named in honor of
one of its discoverers, Marita Oppenheimer nee
Lewis (b. 1957), long-time Maui resident and avid
hiker.
Paratypes. U.S.A. Hawaiian Islands. Last Maui. Mak-
awao: Ha'iku Uka, Wai'ohiw i Stream, Oppenheimer & Du¬
vall HI0005 (BISH. OSH. PTBG); Ha'iku Uka, Wai'ohiwi
94
Novon
Stream, Oppenheimer & Duvall H10005B (BISH, K, NY,
OSH, ITBG). liana: ridge left side Klpahulu Valley,
Forbes I636.M (BISH), Forbes 1 680a. M (BISH); Hono-
manu Valley, Rock 8707a (BISH, (ill); ke'anae Valley,
Rock 8708 (BISH, GH).
I he species of the Cyanea scabra complex as
defined above may be distinguished using the fol¬
lowing key. In the most recent key to the genus
(Gammers, 1990a), this may be substituted for cou¬
plets 48—51 on p. 443.
Key to the Species ok the Hawaiian Cyanea scabra
Complex
la. Bracts oblong, their apex obtuse; calyx lobes ob¬
long, their apex rounded, truncate, or emargin-
ate, often apiculate.
2a. Margin of adult lamina cleft, parted, or di¬
vided.
da. Trees, 4—6 m tall; stems branching at
least 1 m above the base, producing 4
to 1 I spreading branches; juvenile
leaves with 8 to 10 oblong or elliptic
segments per side, the margin of these
segments lobed, cleft, parted, or divid¬
ed; adult leaves with 10 to 14 oblong or
elliptic segments per side, the margin
of these segments cleft or parted; petiole
of adult leaves 4.5— 6.5 cm long, y,o-K
as long as the lamina; bracts 10—13 X
1.5— 1.6 mm; hypanthium 'A—Va as long
as corolla; calyx lobes 13—19 mm long;
corolla 38-47 mm long, the dorsal lobes
ca. Va as long as the tube; filament tube
32—40 mm long, ca. 3 times longer than
the dorsal anthers, pubescent; anther
tube 2.3— 2.7 mm diam., densely pubes¬
cent at base and on connectives; ventral
anthers tipped by a triangular scale of
concrescent trichomes 3—4.5 mm long;
berries 17—21 mm long, 10—15 mm
diam. (bast Maui) . C. duvalliorum
3b. Shrubs, 2—2.4 m tall; stems branching
near base, producing 2 to 4 strongly as¬
cending branches (rarely unbranched);
juvenile leaves with 5 to 8 triangular
segments per side, the margin of these
segments serrulate; adult leaves with 6
to 8 narrowly triangular, triangular, del-
tate, or shallowly triangular segments
per side, the margin of these segments
serrulate; petiole of adult leaves 10—
13.5 cm long, Va-Va as long as the lam¬
ina; bracts 1.5-2. 2 X 0.8-1 mm; hy¬
panthium 14— ‘/() as long as corolla; calyx
lobes 3. 5-9.5 mm long; corolla 59-61
mm long, the dorsal lobes Va— % as long
as the tube; filament tube 50—51 mm
long, 4—5 times longer than the dorsal
anthers, glabrous; anther tube 2.8-3.1
mm diam., glabrous; ventral anthers
tipped by a tuft of free trichomes 2.5-3
mm long; berries 12—13 mm long, 8.5—
10.5 mm diam. (East Maui) . . C. maritae
2b. Margin of adult lamina dentate or denticu¬
late.
4a. Lamina elliptic' or obovate, 2. 5-4. 5
times as long as the petiole, ihe base
obtuse or rounded; corolla 50—60 mm
long, the tube 37—45 mm long, 3—3.8
mm tall at mouth and 4.7— 7.3 mm tall
near middle; filament tube 43—55 mm
long; dorsal anthers 8-8.5 mm long, '/7-
V- as long as filament tube; ventral an¬
thers 5.5— 6.5 mm long, their apical
hairs 3—3.8 mm long (West Maui) . . .
. C. scabra
4b. Lamina narrowly oblong or oblanceo-
late, 5—9 times as long as the petiole,
the base cuneate; corolla 62-65 mm
long, the tube 48—51 mm long, 5.5— 6.5
mm tall at mouth and 8.2-1 1 mm tall
near middle; filament tube 56-60 mm
long; dorsal anthers 5 mm long, '/, 2 — , ,
as long as filament tube; ventral anthers
4 mm long, their apical hairs 1.5—2 mm
long (Kast Maui) . C. mceldowneyi
lb. Bracts linear, linear triangular, or narrowly tri¬
angular, their apex acuminate; calyx lobes tri¬
angular or narrowly triangular, their apex acu¬
minate.
5a. Leal base long attenuate, narrowing imper¬
ceptibly into a winged petiole; corolla tube
45-49 mm long; dorsal anthers 5.3-6 mm
long, about VH) as long as the filament tube;
ventral anthers 4.2— 4.9 mm long, their api¬
cal hairs 1-1.5 mm long (East Maui) ....
. C. longissima
5b. Leaf base cuneate or obtuse, rarely rounded,
the petiole not winged; corolla tube 26—45
mm long; dorsal anthers 7-10 mm long, Vs-
Va as long as the filament tube; ventral an¬
thers 5-8 mm long, their apical hairs 1.7-
3.5 mm long.
6a. Corolla 55—64 mm long, the tube 43—
45 mm long, 5.5—8 mm tall at base, 9—
1 1 mm tall near middle, the lobes Va—
% as long as the tube; filament tube 48-
57 mm long; hairs at apex of ventral
anthers 1.7—2 mm long (East Maui) . .
. 6’. glabra
6b. Corolla 43-56 mm long, the tube 26-
39 mm long, 3—5 mm tall at base, 4—9
mm tall near middle, the lobes 2/--7s as
long as the tube; filament tube 36—50
mm long; hairs at apex of ventral an¬
thers 2—3,5 mm long (windward Ha¬
waii) . platyphylla
Cyanea sect. Deusseoideae
As circumscribed by Rock (1919), Cyanea sect.
Delisseoideae (Hillebrand) Rock comprises those
members of the genus that are shrubs or treelets
0.3—5 m tall (rarely small branched trees to 7 m
tall) with stems 0.5— 2.5 cm in diameter just below
the apex, totally lacking prickles, with entire or
toothed leaves, minute deciduous triangular bracts,
bibracteolate pedicels, minute calyx lobes, hypan¬
thium only I/,,— V- as long as the relatively small,
slender, suberect or gently curved white corolla.
Volume 14, Number 1
2004
Lammers
Endemic Hawaiian Genus Cyanea
95
purple berries, and small minutely papillose seeds
(Type F testa; Buss et al., 2001). Of the species
recognized in the most recent treatment of the ge¬
nus (Lammers, 1990a) and its supplements (Lam¬
mers & Lorence, 1993; Lammers, 1998a), the fol¬
lowing are referable to section Delisseoideae: C.
angustifolia (Chamisso) Hillebrand (lectotype of the
section; St. John, 1969), C. asarifolia H. St. John,
C. coriacea (A. Gray) Hillebrand, C. dolichopoda
Lammers & Lorence, C. elliptica (Rock) hammers,
C. fauriei H. Leveille, C. habenata (H. St. John)
Lammers, C. hardyi Rock, C. kahiliensis (H. St.
John) Lammers, C. linearifolia Rock, C. mannii
Brigham, C. membranacea Rock, C. obtusa (A.
Gray) Hillebrand, and C. spathulata (Hillebrand)
A. Heller.
In my earlier treatment, 1 followed Rock (1917.
1919) in recognizing C. coriacea and C. fauriei as
distinct anti largely allopatric species, distin¬
guished primarily by inflorescence length: plants of
northern Kaua‘i with short spreading or ascending
condensed racemes were called C. coriacea, while
plants of southern Kaua'i with long pendent elon¬
gate racemes were called C. fauriei. However, field
studies on Kaua‘i in 1991 and detailed examination
of a larger body of herbarium specimens convinced
me that many of the long-racemed plants I had
identified as C. fauriei were otherwise indistin¬
guishable from C. coriacea. Variation in raceme
length among these plants is continuous and essen¬
tially clinal. In northern Kaua‘i (e.g., on the Napali
Coast) are found populations with relatively short
often ascending racemes (cf. Rock. 1919, pi. 109).
These lengthen as one proceeds south, reaching
their greatest development with the highly elongate
and pendent racemes found in Olokele Canyon (cf.
Rock, 1919, pi. 1 10). It is worth noting that Rock
himself had earlier (Rock, 1913. 1914) treated the
two names as synonymous, while Wimmer (1943)
treated C. fauriei as a variety of C. coriacea, a clas¬
sification embraced by Degener and Degener
(1956) and St. John (1973. 1980).
However, not all specimens that I had identified
as C. fauriei resemble C. coriacea. Many of the
plants with relatively long pendent racemes differ
from C. coriacea in a number of characteristics, in¬
cluding slimmer twigs and smaller leaves. In these
features, they more closely resemble C. hardyi ; in
fact, some of these specimens were previously iden¬
tified as that species. However, these smaller-
leaved plants differ from C. hardyi in several fea¬
tures, including their broader leaf base, shorter
rachis and pedicels, shorter hypanthium and calyx
lobes, and shorter filament tube. Furthermore, one
specimen previously identified as C. fauriei differs
from all these plants in its extremely small flowers,
e.g., hypanthium only 2.3— 2.7 mm long, corolla just
15—16.5 mm long, filament tube 12—14 mm long,
and anthers (dorsal pair) 4.5—5 mm long. In fact,
these are the smallest known flowers in the entire
genus, by a significant margin. It thus appears that
three distinct species were confounded under the
name C. fauriei: the long-racemed southern popu¬
lations of C. coriacea, plants with slimmer tw igs and
smaller leaves resembling C. hardyi, and a plant
with the smallest flowers in the genus.
In order to determine which of these takes the
name C. fauriei, it was first necessary to clarify the
typification of that name. In the protologue, Leveille
cited two gatherings: “Kauai: Koloa; Waimea, 1000
m, dec. 1909, mart. 1910, (Faurie, 565, 591).”
These are thus syntypes from which a lectotype
must be selected. Rock (1913, 1914, 1919) appar¬
ently only saw Faurie 565, but never referred to it
as the type. Wimmer (1943) saw both gatherings
and specifically excluded Faurie 591 from C. cor¬
iacea var. fauriei by referring it to C. coriacea var.
hardyi (Rock) E. Wimmer, a homotypic synonym of
C. hardyi. Like Rock, he did not specifically state
that Faurie 565 was the type of C. fauriei. As a
result, neither action can be interpreted as an ef¬
fective lectotypification under Article 7.11 of the
ICBN (Greuter et al., 2000).
The specimens upon which Leveille based his
new taxa were deposited in his personal herbarium,
which was sold to E in 1919 (Stafleu & Cowan.
1979). Staff members of that institution have com¬
piled a catalogue of all names ever published by
Leveille, together with information on typification
(cf. Lauener, 1983). In this catalogue, even' one of
the Hawaiian Lobelioideae is listed as “typus —
n[on]. v [idi].,” even though types for nearly all Chi¬
nese Campanulaceae were seen (Chamberlain,
1977). Indeed, when I requested a loan of Levei lie's
Hawaiian lobelioid types from E in 1985. I received
only a set of photographs showing specimens at
BM. and a note stating that no actual specimens
were to be found at E. Lauener (1980) referred to
the specimens at BM as isotypes (or syntypes, if
two or more numbers had been cited in the proto¬
logue), but they appear to be the original specimens
from Leveille’s personal herbarium. I have been un¬
able to discover why these specimens are at BM
and not E with the remainder of Leveille's personal
herbarium.
In any event, both Faurie 565 and Faurie 591
were found at BM. each represented by a photo¬
graph at E. As noted bv Wimmer (1943), the latter
gathering does not conform to the original diagnosis
of C. fauriei and is referable to C. hardyi. For this
96
Novon
reason, C. fauriei is here leetotypified on the basis
of Faurie .565, which does conform to the diagnosis.
Because lliis specimen represents the long-raeemed
southern populations of C. coriacea, this binomial
is here treated as a synonym of C. coriacea (follow¬
ing Rock, 1913, 1914), along with several varieties
of that species that do not merit recognition (cf.
hammers. 1990a). All are listed here for clarity.
Cyanea coriacea (A. Gray) Hillebrand, FI. Ha¬
waiian I si. 254. 1888. Delissea coriacea A.
Gray, Proc. Amer. Acad. Arts 5: 147. 1861.
TYPE: U.S.A. Hawaiian Islands: Kaua‘i,
1851-1855, ./. Remy 302 (holotype, GH; iso¬
types, A, L, P[5], /).
Cyanea fauriei H. Reveille, Repert. Spec. Nov. Regni Veg.
10: 156. 191 1. Cyanea coriacea var. fauriei (H. Rev¬
eille) K. Wimmer, Pflanzenr. IV.276b: 72. 1943. De¬
lissea fauriei (H. Reveille) II. St. John, Phytologia 63:
342. 1987; non H. Reveille, Repert. Spec. Nov.
Regni Veg. 12: 505. 1913. TYPE: U.S.A. Hawaiian
Islands: Kaua‘i, Koloa, Dec. 1909, V. Faurie 565
(lectotype, selected here, BM, photograph, E; isolec-
totypes. A, BISH, G, P, W).
Cyanea coriacea var. degeneriana E. Wimmer, Pflanzenr.
1V.276I): 761. 1953. Delissea coriacea var. degener¬
iana (E. Wimmer) H. St. John, Phytologia 63: 82.
1987. TYPE: U.S.A. Hawaiian Islands: Kaua'i. ridge
behind Papa'a, It) Jan. 1952, O. Degener & A.
Greenwell 21736a (holotype, W; isotypes, B[2|).
Delissea coriacea var. deltoidea II. St. John, Phytologia 63:
340. 1987. Syn. nov. Delissea perlmanii II. St. John,
Phytologia 63: 346. 1987 (as 'perlmannii'). TYPE:
U.S.A. Hawaiian Islands: Kaua’i. Eimahuli Valley,
common, 8 Sep. 1977, S. Perlman I (holotype,
BISH; isotypes, BISII[2]).
Delissea coriacea var. haupuensis H. St. John, Phytologia
63: 340. 1987. Syn. nov. TYPE: U.S.A. Hawaiian
Islands: Kaua'i, Hoary Head Range, near 1286 ft.
peak N of Ra'aukahi, E side of range, dryland forest,
I 100 It.. 3 Nov. 1976, C. Christensen 91 (holotype,
BISH; isotype. BISH).
Delissea coriacea var. lumahaiensis II. St. John, Phytologia
63: 340. 1987. TYPE: U.S.A. Hawaiian Islands:
Kaua'i, road between Hanalei & Lumaha'i, Sep.
1913, C. I\. Forbes & II. ./. Forbes 464. K (holotype,
BISH; isotypes, BISH. GB, K. NY, P, UC, US).
The two sets of populations that were formerly
identified as C. fauriei hut are not referable to C.
coriacea are described here as new species. Though
no names at species rank are available, a name at
the rank of forma is referable to one species as here
circumscribed. However, I explicitly decline to take
up this epithet. Though its type falls within the cir¬
cumscription of the species, its original description
does not address the characters emphasized here.
For the sake of clarity and precision, I prefer to
exercise my prerogative and treat the species as
new, with its own diagnosis.
Cyanea pseudofauriei Rammers, sp. nov. T\ BE:
U.S.A. Hawaiian Islands: Kaua'i. Koloa Dis¬
trict. Rlhu'e-Koloa Forest Reserve, along ridge
ESE of Pu'u Kolo, lower ridge in diverse forest
with rather stunted vegetation on the windward
slopes of the ridge, common on windward
slope, elev. 1200-1700 ft.. 21 Sep. 1988, T.
Flynn. G. Kawakami & R. Nagata 3144 (ho¬
lotype. F; isotype, PTBG). Figure 4A, B.
Cyanea coriacea f. gratiosa E. Wimmer, Pflanzenr.
IV.276R: 760. 1953. Delissea coriacea var. gratiosa
(E. Wimmer) II. Si. John, Phytologia 63: 82. 1987.
TYPE: U.S.A. II awaiian Islands: Kaua'i, ridge be¬
hind Papa'a, Moloa'a Forest Reserve, 16 Jan. 1952.
0. Degener & A. Greenwell 21737 (lectotype, select¬
ed here, B; isolectotype, W).
I'he protologue of Cyanea coriacea I. gratiosa cit¬
ed "Kauai: Ridges behind Papaa in lower forest,
lf>. 1. 1952 (lib. Deg. n. 21736! common form,
21737! rare form intermediate in coloration betw.
n. 21736 and n. 21738).” I have seen sheets of
Degener c, I). Stone, P. Manos & A. L Thomas: DKH 576 (HN,
LE, MO); Moc Chau Distr., Cho Long, Vu Xuan Phuong
79 (1IN). Thanh Iloa Prov.: Ba Thuoc Distr., Co Lung
Mun., N. T. Hiep, L. Averyanov & P. K. Ijoc: HAL 948 (HN,
LE, MO).
Acknowledgments. The study was supported by
the Institute of Ecology and Biological Resources,
National Center for Science and Technology, Hanoi,
Vietnam. I am grateful to Le Xuan Canh and Nguy¬
en l ien Hiep for their encouragement and advice.
I thank Roy Gereau for the Latin diagnosis, Dan
Harder, Ihsan Al-Shehbaz, and Charlotte M. Taylor
for helping with the English, and Ruth Kiew for
providing distribution information. The United
States National Science Foundation is thanked for
a grant (DEB-9870231 to 1). k. Harder) that sup¬
ported the fieldwork.
I .iterature Cited
Doorenbos, J., M. S. M. Sosef & J. J. F. I',, de Wilde. 1998.
The Sections of Begonia including Descriptions, Keys
and Species Lists (Studies in Begoniaceae VI). Wag-
eningen Agricultural University, Netherlands.
Gagnepain, F. 1919. Nouveaux Begonia d’Asie; quelques
synonymes. Bull. Mus. llist. Nat. Baris 25: 194—201,
276-283.
- . 1921. Begun iacties. In: F. Gagnepain (editor),
Flore Generate de L'lndoehine 2(8): 1095—1120. Mas¬
son et Cie, Paris.
Ku. K. 1999. Begoniaceae. FI. Reipublicae Popul. Sin.
52(1): 126-268. Science Press, Beijing.
Pham, H. H. 1999. Begoniaceae. In: H. H. Pham (editor).
Cay co Viet Nam. I: 577-588. Tre Publishing House,
Vietnam.
Slnii Yu Min, Ching I Peng & Cheng Yih Wu. 2002. Syn¬
opsis of the Chinese species of Begonia (Begoniaceae),
with a reappraisal of sectional delimitation. Bot. Bull.
Acad. Sin. 43: 313-327.
Swenson. S. M. & M. C. Tebbitt. 2001. Evolution, bioge¬
ography and systematics of Begoniaceae. Botany 2001:
Plants and People. Albuquerque: 93. | Abstract. |
- . W. L. Clement, L. L. Forrest & M. C. Tebbitt.
2001. Hillebrandia sandwichensis : Evolutionary rela¬
tionships and biogeography. Botany 2001: Plants and
People. Albuquerque: 95. [Abstract. |
Wan Y. & B. N Chang. 1987. A new species of the genus
Begonia from Guangxi. Acta Phytotax. Sin. 25(4): 322—
323.
A New Species of Ceratolacis (Podostemaceae) from
Minas Gerais, Brazil
C. Thomas Philhrick
Department of Biological and Environmental Sciences, Western Connecticut State University,
Danbury, Connecticut 06810, U.S.A. philbrickt@wcsu.edu
Alejandro Novelo R.
Departamento de Botanica, Institute de Biologfa, Universidad Nacional Autonoma de
Mexico, Mexico. D.F., 04510, Mexico, lanovelo@servidor.unam.mx
Bruno E. / r gang
Departamento de Botanica, Universidad Federal do Rio Grande do Sul. Avenida Bento
Goncalves, 9500-Bairro Agronomia, 91570-950, Porto Alegre, Rio Grande do Sul. Brasil
Abstract. A new species ol Ceratolacis ( C . pe-
dunculatum, Podostemaceae) is illustrated and de¬
scribed. I lie new species is known from four lo¬
cations in central Minas Gerais, Brazil. Ceratolacis
pedunculatum is distinguished from the only other
species of the genus based on the presence of a
prominent peduncle and the oblique orientation of
the ovary on the pedicel.
Rksumo. Uma nova especie de Ceratolacis ( C. pe¬
dunculatum, Podostemaceae) e descrita e ilustrada.
Ksta especie nova e conhecida de quatro local ida-
des no centre plo di¬
visa, rare subdichotoma vel pinnata; stipulae asymmetri-
cae, ex folii base amplexicauli cymbiformi constatae. In-
florescentia uniflora, pedunculata. Flores solitarii in axillis
fnliarihus. Spat In 41a claviformis. Pedunculus et pedicellus
post spathellae rupturam elongautes. Tepala 3, uno in an-
dropodii quoque latere, tertio tepalo ad andropodii apicem
portato. Stamina 2. Andropodium et filamenta peranthesin
elongantia. Pollinis grana in dyadibus. Ovarium ex 2 car-
pellis aequalibus compositum, 2-loculare, in pedicello
oblique disposilum. Stigmata 2, libera, stigmatum base in
fructu indurata, persistenti. Fructus capsularis, 2-locular-
is, apice emarginato, duabus valvis aequalibus, persisten-
tibus, 3-costatis, marginibus suturalibus incrassatis. Pe-
dunculus a pedicello in fructu non distinguibilis.
Aquatic herbs, perennial. Roots green, elongate
and cylindrical when young, becoming broad and
crust-like, irregular in outline when older,
branched, prostrate, flattened to elliptical in cross
section, median = 0.9 (0.3-5. 2) mm |N = 50] wide,
tightly attached to rocks via adhesive hairs, growing
intertwined. Stems monomorphic, arising opposite
or subopposite along the Hanks of roots, 4.9 (0.9-
Volume 14, Number 1
2004
Philbrick et al. 109
Ceratolacis pedunculatum from Minas Gerais
12) mm | IN = 50] apart, obscured by leaf bases,
erect, less than 0.1 mm high, unbranched. Leaves
petiolate, distichous, stipulate, double-sheathed,
upright, compound, 4 (1-6) [N = 50] times divided,
rarely subdiehotomous or pinnate, 17 (2.9—35.2)
mm [N = 100] long, ultimate leaf segments linear
to spatulate, 3.1 (0.2-9.5) mm |N = 100] long X
0.3 (0.1-1) mm [N = 100| wide, midvein faint or
lacking, apices acute or obtuse; petiole 3.9 (0.4—
14) mm | IN = 100] long, elliptical in cross section;
stipules asymmetric, composed of an extension of
the amplexicaul leal base, 0.5 (0.1— 5.6) mm [N =
50] long, larger on leaves subtending flowers, apex
entire. Inflorescence with flowers pedunculate and
solitary. Flowers 2 (1—6) [N = 50] per stem, her¬
maphroditic, zygomorphic, borne singly in a leaf
axil, covered by a spathella, pedicellate; peduncle
(below attachment point of spathella) 0.8 (0.1— 3.1)
mm [N = 100] long prior to spathella rupture, elon¬
gating to 2.6 (0.3— 8.5) mm |N = 100] long after
spathella rupture; spathella clavate, 2.4 (1-4.5) nun
[N = 100] long X l.l (0.6— 1.7) mm |N = 100]
wide, rupturing apically, falling away completely in
fruit; pedicels 0.8 (0.1— 3.1) mm [IN = 100] long
prior to spathella rupture, elongating to 5.1 (1.8 —
8.5) mm [N — 50] long after spathella rupture; te-
pals 3, seale-like, linear, acute, two (one on either
side) at the base of the' stalk from which the sta¬
mens arise (andropodium), 0.6 (0.2— 1.8) mm [N =
50] long; a third tepal occurring at the apex of the
andropodium, andropodial tepal 0.4 (0.3— 0.5) mm
|N = 50] long; stamens 2. deciduous; andropodium
I (0.6—2) mm [N — 50] long prior to spathella rup¬
ture, during anthesis elongating to 4.3 (2-7) mm [N
= 50] long; filaments 0.4 (0.2-1. 8) mm [N = 50]
long prior to spathella rupture, during anthesis
elongating to 0.9 (0.4-1. 8) mm [N — 50] long; an¬
thers quadrangular, with parallel sides, 1.1 (0.7—
1.4) mm [N = 50] long, I (0.6-1. 2) mm [N = 50]
wide, dehiscing introrsely anti longitudinally, inner
and outer thecae of equal length; pollen in dyads,
tricolpate, 35 (31-40) /am |N = 100] long. 24 (19—
28) /tun [N = 100] wide; ovary with 2 equal carpels,
2-loeular. oval in shape, oriented obliquely on the
pedicel, 1.5 (0.7— 2.6) mm [N = 50] long, 1 (0.6—
1.7) mm |N = 50] wide, with 6 longitudinal dark
lines on the ovary wall; ovules 20 (12—32) [N =
25]; stigmas 2, free, each stigma triangular in out¬
line, widest at base, upright prior to spathella rup¬
ture, 0.5 (0.3—1) mm [N = 50] long, during anthesis
upright or divergent, elongating to 0.9 (0.4— 1 . 1) nun
[N = 50] long, base of stigmas becoming hardened
and persisting in fruit. Fruit a 2-locular capsule,
1.8 (1.4— 2.1) mm [N = 50] long, 1.3 (0.9— 1.5) mm
|N = 50] wide, capsule apex emarginate due to
persistent stigma bases, with two equal valves, per¬
sistent, valves 3-ribbed, suture margins thickened
and rib-like; peduncle not distinguishable from
pedicel in capsule (due to shedding of spathella
and parenchymatous tissues of peduncle and ped¬
icel); length of peduncle and pedicel combined 6.8
(1.5—17) mm [N = 100] long. Mature seeds un¬
known.
Ceratolacis pedunculatum is known from four lo¬
cations in the central part of the state of Minas
Gerais, Brazil (Fig. 2), ranging from about 500 to
650 m elevation. The related C. erythrolichen is
known only from the type collection (rio Tocantins),
which is approximately 1500 km north of the lo¬
cations where C. pedunculatum was collected. Al¬
though the type locality of C. erythrolichen was in
the state of Goias at the time the species was de¬
scribed in 1849, the location is presently in the
state of Tocantins.
Additional observations. Two reproductive char¬
acteristics distinguish Ceratolacis pedunculatum
from C. erythrolichen. Ceratolacis pedunculatum
possesses a peduncle (Fig. lb, le, lg, lb), and the
axis of the ovary is offset (oblique) from that of the
pedicel by ea. 30—45° (Fig. If— h). In contrast. C.
erythrolichen lacks a peduncle and the ovary axis
is parallel to that of the pedicel.
One characteristic of the root also seems to differ
between C. erythrolichen and C. pedunculatum. Tu-
lasne (1852: 126) and Engler (1930: 52) referred
to the roots of C. erythrolichen as being red while
living and turning blackish or whitish upon drying.
This character is not evident in C. pedunculatum.
in which the roots are dark green in living and
dried specimens.
Justification for recognizing Ceratolacis pedun¬
culatum as distinct from C. erythrolichen warrants
some discussion of the type material of the latter
species. Tulasne (1849) based his description of C.
erythrolichen on a collection of Weddell. As Wed¬
dell worked at Paris and his collections are housed
there, it has been reported (Tulasne, 1852: 127;
Van Royen, 1954: 224) that the type material of C.
erythrolichen is deposited in P. Recently, however,
the type specimen has not been located there.
There is one specimen located in G that seems to
be the holotype. Isotypes are located in K and M.
The holotype and isotypes are comprised of only
vegetative material (roots, stems, leaves). As a con¬
sequence, it has not been possible to study the re¬
productive structures on the type material for C.
erythrolichen. Van Royen (1954) also noted that the
type material was incomplete.
Ceratolacis erythrolichen is only known from the
110
Novon
Figure 1. CeratolacLs pedunculatum C. T. Philbrick, Novelo & Irgang. Drawings based on the holotype. — a. General
habit of plant showing cylindrical root from which leaves arise; steins are obscured by base of leaves. — b. General
habit of plant showing cylindrical root, short stem (obscured by sheathing leaf liases), and two flowers — one at anthesis,
the other enclosed within an intact spathella. Also showing dichotomously and subdiehotomously divided leaves. — c.
Details of a double-sheathed leaf with two basal stipules (left) and two simple leaves on a short stem (right). — d. Detail
of three leaves showing asymmetric stipule on left side of leaf base and dichotomous to subdiehotomous divisions. A
detail of the apical segment of a leaf is included. — e. Flower enclosed within a spathella, which is partially ruptured
at the apex. Apex of the peduncle is indicated by the arrow. — f. Two views of a flower at anthesis showing the
Volume 14, Number 1
2004
Philbrick et al. Ill
Ceratolacis pedunculatum from Minas Gerais
type collection. Thus, the present authors have had
to rely on t lie descriptions and illustrations in Tu-
lasne (1849, 1852. 1855) for their understanding
of the reproductive morphology of the species. In
the protologue for Ceratolacis erythrolichen (as Di-
craea erythrolichen) Tulasne (1849: 102) did not
make reference to a peduncle (a stalk below the
attachment point of the spathella), nor is one in¬
dicated in the illustration for the species (his PL.
X. 1). Rather, Tulasne (1849) referred only to a
pedicel. (In Podostemaceae the pedicel is the stalk
of the ovary that arises above the attachment point
of the spathella.) Subsequently, Tulasne provided
an expanded description of the species in his
monograph of Podostemaceae (Tulasne, 1852: 12b—
128) and again in Flora brasiliensis (Tulasne, 1855:
253—254), but in neither case is any reference to a
peduncle made. Although illustrations of the spe¬
cies are included in these two later works, they are
the same illustrations that accompanied the 1849
description of the species. Given the prominent na¬
ture of the peduncle, the authors contended that
Tulasne would have noted a peduncle if one had
been present in the type material. Thus, the authors
are confident that C. erythrolichen lacks a pedun¬
cle. Ceratolacis pedunculatum possesses a distinct
peduncle. The peduncle occurs below the attach¬
ment of the spathella (arrows in Fig. le, lg). while
the pedicel occurs above. Lastly, Tulasne (1849,
1852, 1855) did not make reference to the ovary of
C. erythrolichen being oblique on the pedicel either
in his species description or in the accompanying
illustrations. Based on the presence of a peduncle,
the oblique orientation of the ovary, and to a lesser
degree on the green roots, the authors conclude that
C. pedunculatum is distinct from C. erythrolichen.
Species in three genera of Podostemaceae in the
New World possess two stamens, an andropodium,
and an asymmetrical sheathing leaf base (Table 1):
two species of Ceratolacis , all species of Crenias,
and one species of Podostemum (P. muelleri Warm¬
ing). It is thus appropriate to focus on these eight
species in a discussion of the characteristics of Cer¬
atolacis pedunculatum.
The nature of the leaf base is partially correlated
with stipule form among species of Ceratolacis,
Crenias, and Podostemum. A single asymmetrical
Distrito Federal; RS, Rio de Janeiro; ES, Espfrito Santo.
stipule (a stipule on one side of the leaf base) and
subdichotomous or pinnate leaves occur in species
of Ceratolacis, Crenias, and one species of Podos¬
temum (P. muelleri). In contrast, a symmetrical
boat-shaped stipule (entire or divided into apical
teeth) occurs in all species of Podostemum (except
P. muelleri and P. irgangii C. T. Philbrick & Nov-
elo); these species all have dichotomously divided
leaves. Podostemum muelleri deviates from this cor¬
relation with its dichotomously divided leaves and
asymmetrical stipule, whereas leaves of P. irgangii
are uniformly verticillate.
Ceratolacis pedunculatum is the only New World
species that has an asymmetrical stipule, an andro¬
podium, two stamens, a pedicel, and a peduncle.
The peduncle (0. 1-8.5 mm long) occurs below the
location where the sac-like spathella attaches to the
pedicel (arrows in Fig. le, lg). In contrast, the spa-
thellas of species ol Podostemum and Crenias, in¬
deed most other species of Podostemoideae in the
New World, attach at the base ol the pedicel — at
the point of attachment to the stem. Thus a pedun¬
cle is lacking. A peduncle is known from only three
species of Podostemaceae in the New World: Cer¬
atolacis pedunculatum, Macarenia clavigera P. Roy-
en, and Vanroyenella plumosa Novelo & C. T. Phil¬
brick.
Characteristics of the capsule distinguish species
andropodium (stalk from which stamens arise), two anthers, two basal tepals, and one tepal arising from the apex of
the andropodium. A detail of one anther is also shown. — g. A flower at anthesis showing its orientation relative to the
root. Arrow indicates the apex of the peduncle. — h. A post-anthesal flower showing the remnants of the spathella
arising from the apex of the peduncle. — i. Mature capsule prior to dehiscence showing the persistent and divergent
stigma bases and the isolobous capsule. — -j. Mature capsule after dehiscence (seeds have been dispersed) showing the
two persistent capsule valves.
112
Novon
of Ceratolacis from Crenias and Podostemum. The
capsule in the latter two genera is cylindrical in
cross section, with a rounded or hlunt apex. In con¬
trast, the capsule of Ceratolacis is somewhat flat¬
tened with an emarginate apex, a character that is
largely a consequence of the persistent stigma lia¬
ses. The stigma liases are deciduous in species of
Crenias and Podostemum. In addition, species in
these two genera possess capsules in which one
valve is deciduous, the other persists. In contrast,
I nit li valves of die capsule of species of Ceratolacis
persist. Thus, with capsules alone it is possible to
distinguish plants of die genus Ceratolacis from
those of other genera of Podostemaceae.
Species of Ceratolacis possess an isolobous ova¬
ry, while die ovaries of species of Crenias and Po¬
dostemum are anisolobous. The presence of an an-
isolobous ovary is usually associated with the ovary
set at an angle (oblique) on the pedicel. However,
Ceratolacis peduncidatum is unusual in that the is¬
olobous ovary is clearly oblique on the pedicel (Fig.
If-h). There is no indication that C. erythrolichen
shares this character.
Variation in stem and root form also helps dis¬
tinguish species of Ceratolacis. Elongate stems
characterize most species of Podostemum and Cren¬
ias, although stems of some species can be ob¬
scured by the sheathing leaf bases. Ceratolacis pe-
dunculatum lacks an obvious upright stem; the
diminutive stem is obscured among the leaf bases.
Roots of species of Ceratolacis, Crenias, and Po¬
dostemum are elongate and cylindrical when young.
Roots of mature plants retain their cylindrical form
in species of Podostemum and Crenias. In contrast,
mature roots of both species of Ceratolacis often
become broadened, irregular in outline, flattened,
and can resemble a crust. These flattened roots are
difficult to remove from the rocks. Thus, to docu¬
ment this character it is necessary to collect spec¬
imens and the rocks to which they are attached.
Species of Crenias and Podostemum possess
dvad pollen. Dyad pollen also characterizes Cera¬
tolacis pedunculatum. ’file authors predict that C.
erythrolichen has dyad pollen as well, although the
absence of adequate type material does not allow
this prediction to be confirmed.
Paratypes. BRAZIL. IVIinas Gerais: on road from l)i-
amantina (Gouveia) to Curvelo, confluence of rio Parauna
and rio CipA, 0 July 2002, C. T. Philbrick, A. Novelo A'.,
It. P. Irgang & C. V. de Senna Gastal 5649 (BHCB. IGN,
MKXU, \1(). WCSU); ca. 10 km W of Sao Sebasliao,
Route BR-420, on road to Pompeu, 0 July 2001, C. T.
Pliilbriek. A. Novell) R., R. E. Irgang & C. V. de Senna
Gasta! 5651 (BHCB, 1CN, MKXU, MO. WCSU); ca. 16
km SW of Pompeu, along BR-262 on road to Martinhos
Campos, rio Para. 16 July 2001, C. T. Philbrick. A. Novell >
Volume 14, Number 1
2004
Philbrick et al. 113
Ceratolacis pedunculatum from Minas Gerais
R., B. E. Irgang & C. V. de Senna Cast a l 5678 (BHCB,
ICN, MKXU, MO, NY, WCSU); 2 Aug. 2002, C. T. Phil-
brick & A. Nor do R. 5781 (BHCB. ICN, MKXU. MO. NY.
WCSU); 30 km N of Corinto on BR-496, ca. 200 m up¬
stream of Bridge, rio Bicudo, 28 July 2002, C. T. Philbrick
& A. Novelo R. 5761 (BHCB, ICN, MKXU, MO, NY.
WCSU).
Acknowledgments. We thank Fernando Chiang
for the Latin description and Albino Luna for draw¬
ing Figure I. Rolf Rutishauser and an anonymous
reviewer are thanked for their helpful comments on
an earlier version of the manuscript. Claudio V. de
Senna Gastal is thanked for his assistance and en¬
couragement while conducting field studies in
southern Brazil. This work was supported by Na¬
tional Science Foundation Grant DEB-9629767 to
C.T.R and Donald H. Les, and research grants from
the Connecticut State University. The follow ing her¬
baria are thanked for providing access to, or loans
of, specimens: B, BHCB, C, ICN, K, L, M, MFXU,
NY, P, S, US, WCSU. We offer special thanks to
the curatorial staff ol C, K. L, and P for their as¬
sistance.
Literature Cited
Engler, A. 1930. Podostemonaceae. In: A. Kngler & K.
Prantl (editors), Nat. Pffanzenfam. ed. 2 18a. 1-68.
483-484.
Tulasne, I.. R. 1849. Podostemacearum synopsis mono-
graphica. Ann. Sci. Nat. But. Ser. 3, It: 87—1 14.
- . 1852. Monographia Podostemacearum. Arch.
Mas. Hist. Nat. 6: 1-208.
- . 1855. Podostemaceae. In: Martius, FI. bras. 4(1):
229-276.
Royen, P. Van. 1954. The Podostemaceae of the New
World. Iff. Acta Bot. Need. 3: 215-263.
Weddell. 11. A. 1873. Podostemaceae. In: DC., Prodr. 17:
39-89, 300-301.
Validation of Ardisia Section Auriculardisia (Myrsinaceae)
Jon M. Ricketson
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
jon.ricketson@mobot.org
John J. Pipoly III
Fairchild Tropical Garden, 1 1935 Old Cutler Hoad, Coral Gables (Miami),
Florida 33156-4242, U.S.A. jpipoly@faircbildgarden.org
ABSTRACT. In the recent revision of Ardisia subg.
Auriculardisia, the new status, combination, and
basionym for Ardisia sect. Auriculardisia were er¬
roneously omitted. The section is validated here.
Key words: Ardisia seel. Auriculardisia, Myrsi-
naceae.
Because the differences used by l.undell (1981)
to separate his segregate genus Auriculardisia from
Ardisia were found to be unreliable, we included
Auriculardisia within the genus Ardisia and revised
its taxonomy. However, in our recent revision of
Ardisia subg. Auriculardisia (Ricketson & Pipoly,
2003: 203), the new status, combination, and its
basionym for Ardisia sect. Auriculardisia were er¬
roneously omitted. That error is here corrected:
Ardisia sect. Auriculardisia (Lundell) Ricketson
& Pipoly, comb, et stat. nov. Basionym: Auri¬
culardisia Lundell, Phytologia 49: 341. 1981.
TYPE: Ardisia glomerata Lundell.
I .iterature Cited
l.undell, C. I,. 1981. Neotropical Myrsinaceae — IV. I’liy-
tologia 49: 341—354.
Ricketson, J. M. & J. J. Pipoly III. 2003. Revision of
Ardisia subgenus Auriculardisia (Myrsinaceae). Ann.
Missouri Bot. Card. 90: 179—317.
Novon 14: 114. 2004.
Reinstatement of Bonellia (Theophrastaceae)
Her til Stahl
Department of Science and Technology, Gotland University, SE 62167 Visby, Sweden.
bertil.stahl@hgo.se
Mari Kallersjo
Molecular Systematics Laboratory, Swedish Museum of Natural History, P.0. Box 50007,
SE 10405 Stockholm, Sweden, mari.kallersjo@nrm.se
Abstract. Following the results of a recent phy¬
logenetic study showing that the genus Jacquinia is
paraphyletic, the following new combinations in the
genus Bonellia are made (all by Stahl & Kallersjo):
B. albiflora (Lundell), B. brevifolia (Urban), B.
flammea (Millspaugh ex Mez), B. frutescens (Miller),
B. lippoldii (Lepper), B. longifolia (Standley), B.
loeflingii (Carrasquel), B. macrocarpa (Cavanilles),
B. macrocarpa (Cavanilles) Stahl & Kallersjo
suhsp. pungens (A. Gray), B. macrocarpa (Cavan¬
illes) Stahl & Kallersjo suhsp. panamensis (Lun¬
dell), B. montana (Stahl), B. mucronata (Roemer &
Schultes), B. nervosa (Presl), B. nitida (Stahl). B.
paludicula (Standley), B. pringlei (Bartlett), B. pau-
ciflora (Stahl & F. Axelrod), B. shaferi (Urban), B.
seleriana (Urban & Loesner ex Mez), B. sprucei
(Mez). B. stenophylla (Urban), and B. umbellata (A.
DC.). The genus Bonellia is distinguished from Jac¬
quinia s. str. by its mostly orange flowers and al¬
ternate leaves, flattened seeds that are incompletely
covered by placental tissue, and two-layered testa.
A key to the genera of the Theophrastaceae is pre¬
sented.
Key words: Bonellia , Theophrastaceae.
The Theophrastaceae form a small Neotropical
family of about 100 species of shrubs and small
trees. Recent phylogenetic research (Kallersjo et
al., 2000) has shown that Samolus L., a genus tra¬
ditionally included in the Prirnulaceae, is sister
group to the Theophrastaceae and therefore should
he included within the latter family or recognized
as a separate family.
The Theophrastaceae s. str. (excluding Samolus)
were monographed by Mez (1903), who recognized
four genera based mainly on floral morphology, viz.
Clavija, Jacquinia, Deherainia, and Theophrasta.
Votsch (1904), in a leaf anatomical work, trans¬
ferred one of the species of Deherainia to a new
genus, Neomezia, and during more recent taxonom¬
ic work (Stahl, 1993), the genus Votschia was es¬
tablished to accommodate a species previously in¬
cluded in Jacquinia.
Following a new analysis based on a combination
of morphological and epDNA data (Kallersjo &
Stahl, 2003), it has become clear that the genus
Jacquinia is not monophyletic and has to be divid¬
ed into two genera. According to that analysis, Jac¬
quinia s. str. should include species with pseudov-
erticillate leaves lacking or with a poorly developed
niucro, lepidote young shoots, whitish flowers, glo¬
bose seeds completely immersed in placental tis¬
sue, and sunken abaxial foliar sclerenchyma. This
circumscription makes Jacquinia a chiefly Carib¬
bean genus of 13 species. Only one of these spe¬
cies. J. armillaris Jacquin, occurs outside of the
Caribbean islands, in northern coastal South Amer¬
ica and eastern coastal Brazil (Stahl. 1992. 1995).
Jacquinia arborea Vahl, with a wide Caribbean dis¬
tribution, has also been reported a few times from
Mesoameriea, but then only from islands off the
eastern coast of Mexico and Honduras.
The reinstated genus Bonellia is defined by al¬
ternate leaves with a well-developed mucro, gla¬
brous or puberulous young shoots, mostly orange
flowers, flattened seeds only partly immersed into
placental tissue, and abaxial foliar sclerenchyma
located adjacent to the lower epidermis. Two spe¬
cies, B. longifolia and B. paludicola, deviate by
having whitish flowers, and it is therefore interest¬
ing to note that both appear at the base of the Bo¬
nellia clade in the phylogeny proposed by Kallersjo
and Stahl (2003). Of these, B. paludicola is the
most problematical species since it, like most mem¬
bers of Jacquinia s. str. (except J. Iceyensis Mez),
has pseudoverticillate leaves. It should also be not¬
ed that one additional species, B. albiflora, has
white flowers, but that species is morphologically
very similar to B. macrocarpa, indicating a close
relationship between these two taxa.
The genus Bonellia consists of 22 species dis¬
tributed in Mesoameriea, northern and western
Novon 14: 115-118. 2004.
116
Novon
South America, and the Greater Antilles. Within
the genus two morphologically and geographically
distinct groups can be discerned. One of these com¬
prises the South American species B. mucronata
(northern Peru), B. sprucei (Ecuador), B. frutescens
(northern Colombia— Venezuela), B. loefiingii (north¬
ern Venezuela), and the Mesoamerican B. nervosa.
All species in this group have rather large, thick-
walled and opaque fruits, comparatively flexible
leaves (deciduous in B. nervosa ), and whitened
young shoots. The second group consists of species
from Cuba, Hispaniola, and Puerto Rico (B. brevi-
folia, B. lippoldii, B. pauciflora, B. stenophylla, B.
umbellata) with small or very small spine-tipped
leaves, small, thin-walled fruits, and tiny flowers.
I'he remaining, Mesoamerican species are more dif¬
ficult to define as a natural group, although some
species appear to be closely related (Stahl. 1989;
Kallersjo & St&hl, 2003).
fhe name Bonellia was published by Colla
(1824) in a catalog of plants from his private gar¬
den. I’he name was evidently proposed by C. G. F.
Bertero, whose material of a plant from Santa Mar¬
tha in northern Colombia was distributed under the
name of Bonellia cavanillesii to herbaria and bo¬
tanical gardens in Europe. However, since Colla
(1824) cited Jacquinia macrocarpa Cavanilles, a
different Mesoamerican species, as a synonym, the
name B. cavanillesii has to be treated as an obligate
synonym of the latter species.
The name Bonellia honors the Italian zoologist
Franco Andrea Bonelli (1784—1830), evidently a
close friend of Bertero and Colla, but whose con¬
tribution to botany is obscure.
Key to the Genera of Theophrastaceae s. str.
la. Unbranched or sparsely branched shrubs and
trees; fully developed leaves 20-120 cm long,
margins often serrate or serrulate.
2a. Corolla rotate, tube smooth on inside; stam-
inodes ovoid-gibbous; (lowers often unisex¬
ual, male or functionally male flowers with
stamen filaments fused into a permanent
tube; anther not or inconspicuously pro¬
duced at apex; leaf margins entire, serrulate,
or serrate . Clavija Ruiz & Pav6n
2b. Corolla campanulate or bowl-shaped, irreg¬
ularly puckered on inside of tube; stami-
nodes not obovoid; flowers bisexual, stamen
filaments fused at base only; anther connec¬
tive distinctly prolonged at apex; leaf mar¬
gins usually spinose-dentate.
3a. Plants erect, the stem spiny; corolla
deeply campanulate, pale buff turning
black upon drying; staminodes trans¬
versely oblong, inserted within the co¬
rolla tube . Theophrasta L.
3b. Plants semi-decumbent, the stem not
spiny; corolla bowl-shaped, orange, not
blackening upon drying; staminodes tri¬
angular, minute, inserted at the mouth
of the corolla tube . Neomezia Votsch
lb. Branched shrubs or small trees; leaves usually
less than 20 cm long, margins entire.
4a. Corolla green, > 2 cm diam. at anthesis;
staminodes obtrullate to linear; fruits ovoid
. Deherainia Decaisne
4b. Corolla orange, whitish, or pale yellow, < 2
cm diam. at anthesis; staminodes flattened,
petaloid.
5a. Corolla bowl-shaped, pale yellow; style
shorter than the ovary; foliar scleren-
chyma of the lower side arranged in a
continuous or subcontinuous layer . . .
. Votschia StShl
5b. Corolla campanulate or ureeolate, or¬
ange or whitish; style of the same length
as the ovary; foliar sclerenchyma ar¬
ranged in distinct bundles.
6a. Young shoots ± lepidote from ir¬
regularly branched, thick-walled
hairs; leaves mostly pseudoverticil-
late (alternate in ./. keyensis), with¬
out or with a poorly developed api¬
cal mucro; corolla white or whitish;
seeds ± globose, covered by pla¬
cental tissue; abaxial foliar scleren¬
chyma often somewhat immersed
into the mesenchyma . . . Jacquinia L.
6b. Young shoots puberulous from short
uniseriate hairs or glabrous; leaves
alternate (pseudoverticillate in B.
paludicola ), usually distinctly mu-
cronate; corolla orange, sometimes
white or whitish; seeds flattened,
incompletely covered by placental
tissue; abaxial foliar sclerenchyma
adjacent to lower epidermis . .
. Bonellia Colla
Taxonomy and New Comrinations
Roiiellia Colla, Hortus Ripul. 21. 1824. TYPE:
Bonellia cavanillesii Bertero ex Colla (= Jac-
quinia macrocarpa Cavanilles).
Bonellia albiflora (Eimdell) Stall I & Kallersjo,
comb. nov. Basionym: Jacquinia albiflora Fun¬
ded, Wrightia 2: 60. I960. TYPE: Guatemala.
Peten: Tikal, 29 July 1959, Contreras 45 (ho-
lotype, FT; isotypes, S. TEX, US).
Bonellia brevifolia (Urban) Stahl & Kallersjo,
comb. nov. Basionym: Jacquinia brevifolia Ur¬
ban. Symb. Antill. 1: 379. 1899. TYPE: Cuba.
Sine loco, Wright 2913 (leetotype, designated
in Stcihl (1995), GH; isotypes, G, GOET, M).
Bonellia (lannnea (Millspaugh ex Me/.) Sullil &
Kallersjo, comb. nov. Basionym: Jacquinia
flanunea Millspaugh ex Mez, Pflanzenr. IV.
236a: 40. 1903. TYPE: Mexico. Yucatan: Silam,
Gaumer 531 (leetotype, designated in St&hl
(1989), G; isotypes. A, BM, BR, C, CAS, I)S,
E, F, FHO, GH, K. LE, MO. NY, UPS, US. W).
Volume 14, Number 1
2004
Stahl & Kallersjo
Reinstatement of Bonellia
117
Bonellia frutescens (Miller) Stahl & Kallersjo,
coml). nov. Basionym: Ruscus frutescens Miller,
Card. Diet. ed. 8: unpaginated (species no. 8
under Ruscus). 1768. TYPE: Colombia. Boli¬
var: vie. of Cartagena, no date, Heriberto 279
(neotype, designated in Stahl (1995), US).
Bonellia lippoldii (Lepper) Staid & Kallersjo.
comb. nov. Basionym: Jacquinia lippoldii Lep¬
per, Wiss. Z. Friedrich-Schiller-Univ. Jena.
Math.-Naturwiss. Keihe 32: 876. 1983. TYPE:
Cuba. Guantanamo: San Antonio del Sur, Abra
de Mariana. E of Abra. 100—100 m alt.. 9 leb.
1979, Bisse et al. s.n. (holo- and isotype, II AJB
no. 39068).
Bonellia loeflingii (Carrasquel) Stahl & Kallersjo,
comb. nov. Basionym: Jacquinia loeflingii Car¬
rasquel, Acta Bot. Venez. 4: 348. 1970. TYPE:
Venezuela. Anzoategui: about 10 km from Pfr-
itu on road to Clarines, Jan. 1968, Aristeguieta
et al. 6524 (holotype, VEN).
Bonellia longifolia (Standley) Staid & Kallersjo,
comb. nov. Basionym: Jacquinia longifolia
Standley, J. Wash. Acad. Sei. 14: 241. 1924.
TYPE: El Salvador. San Vicente: vie. of San
Vicente, alt. 350-500 in, 2-11 Mar. 1922,
Standley 21159 (holotype, US; isotypes, F, G,
GH. K, NY).
Bonellia macrocarpa (Cavanilles) Stahl Kal¬
lersjo, comb. nov. Basionym: Jacquinia macro¬
carpa Cavanilles, Icon. 5: 55, tab. 483. 1799.
TYPE: Mexico. Acapulco, Eel). 1791, Ned s.n.
(lectotype, designated in D’Arcy (1980), B-
WILLD; isotypes, 11BG, MA).
Bonellia macrocarpa (Cavanilles) Stahl & Kal¬
lersjo subsp. pungens (A. Cray) Stahl & Kal¬
lersjo. comb. nov. Basionym: Jacquinia pun-
gens A. Gray, Proc. Amer. Acad. Arts 5: 325.
1854. TYPE: Mexico. Sonora: hills between
Rayon and Ure; 5, 10 Oct. 1851, Thurber 9 03
(lectotype, designated in Stahl (1989), GH;
isotypes, GH, NY).
Bonellia macrocarpa (Cavanilles) Stahl & Kal¬
lersjo subsp. panamensis ( I. undell) Stahl &
Kallersjo, comb. nov. Basionym: Jacquinia
panamensis Lundell, in Woodson & Schery,
Ann. Missouri Bot. Gard. 27: 329. 1940.
TYPE: Panama. Panama: vie. of Bejuco, 18
Oct. 1938, Allen 985 (holotype, MICH; iso¬
types, GH, MO. US).
Bonellia montana (Stahl) Stahl & Kallersjo,
comb. nov. Basionym: Jacquinia montana
Stahl, Nordic J. Bot. 9: 27. 1989. TYPE: Nic¬
aragua. Jinotega: Cerro de la Cruz and vie., W
of Jinotega, alt. 1200—1400 rn, 7 July 1947,
Standley 10968 (holotype, F).
Bonellia inneronata (Roemer & Schultes) Stahl
Si Kallersjo, comb. nov. Basionym: Jacquinia
mucronata Roemer & Schultes, Syst. Veg. 4:
802. 1819. TYPE: Peru. Colasay, Humboldt
8581 (holotype, B-WIEUD, seen on microfiche;
isotypes, HAU. P).
Bonellia nervosa (Presl) Stahl & Kallersjo, comb,
nov. Basionym: Jacquinia nervosa Presl, Reliq.
Haenk. 2: 67. 1835. TYPE: Mexico. Sine loco,
Haenke s.n. (holotype, PR).
Bonellia nitida (Stahl) Stahl & Kallersjo, comb,
nov. Basionym: Jacquinia nitida Stahl, Nordic
J. Bot. 9: 24. 1989. TYPE: Honduras. Comay-
agua: vie. of Comayagua, alt. 600 m, 12—13
Mar. 1947, Standley & Chacon 6025 (holo¬
type, F).
Bonellia paludirnla (Standley) Stahl Kallersjo,
comb. nov. Basionym: Jacquinia paludicola
Standley, Field Mus. Nat. Hist., Bot. Ser. I l:
138. 1932. TYPE: Belize. Punta Gorda, “For¬
est Home,” alt. 60 m, 1 I Sep. 1932, Schipp
1028 (holotype, F; isotypes. A, BM, G, GH, K,
MICH, MO, NY, S).
Bonellia pauciflora (Stahl & F. Axelrod) Stahl &
Kallersjo. comb. nov. Basionym: Jacquinia
pauciflora Stahl & F. Axelrod, Brittonia 50: I.
1998. TYPE: Puerto Rico. Muriicipio Quebra-
< 1 i 1 las. Barrio Charcas, 0.7 km SSE of Charcas,
200-350 m alt., 23 June 1994, Proctor et al.
49406 (holotype, SJ; isotypes, GB, NY).
Bonellia pringlei (Bartlett) Stahl & Kallersjo,
comb. nov. Basionym: Jacquinia pringlei Bart¬
lett, Proc. Amer. Acad. Arts 44: 630. 1909.
TYPE: Mexico. Guerrero: Iguala Canyon, 8
Oct. 1908, Pringle 10887 (holotype, GH; iso¬
types, BH, BM, C, CM, E, F, G, GOET, HBG,
K, L. LE, M. MICH, MO, MSC, NY, P, PH,
PR, US, VT, W).
Bonellia seleriana (Urban & Loesner ex Mez)
Stahl & Kallersjo, comb. nov. Basionym: Jac¬
quinia seleriana Urban & Loesner ex Mez,
Pflanzenr. IV. 236a: 43. 1903. TYPE: Mexico.
Oaxaca: above San Carlos Yauhtepec, 5 Jan.
1896, Seler & Seler 1759 (lectotype, designat¬
ed in Stahl (1989), GH; isotvpes. A, NY).
Bonellia shaferi (Urban) Stahl & Kallersjo, comb,
nov. Basionym: Jacquinia shaferi Urban,
Symb. Antill. 7: 320. 1912. TYPE: Cuba. Hol¬
guin: vie. of Holguin, 7 Apr. 1909, Shafer
1231 (lectotype, designated in Stahl (1995),
NY; isotypes. A, F, (ill, HAC, K, MO, US).
Bonellia sprucei (Mez) Stahl & Kallersjo, comb,
nov. Basionym: Jacquinia sprucei Mez, Pflan¬
zenr. IV. 236a: 43. 1903. TYPE: Ecuador.
Guavas: Chanduy, Spruce 6478 (lectotype, des¬
ignated in StShl (1990), K; isotypes, BM, BP,
CGE, G, LE, OXF, P, W).
118
Novon
lioucllia slenoplivlla (Urban) Stahl & Kallersjo.
comb. nov. Basionym: Jacquinia stenophylla
Urban, Symb. Antill. I: 378. 1899. TYPE:
Cuba. “Cuba Occ.,” 1803, Wright 2912 />./>.
(lectotype, designated in Staid (1995), C; iso¬
types, BM, GH. GOET, HAC, LE, M. NY, S,
US, W). |See Sttlhl (1995) for comments.]
lioucllia iimbellata (A. DC.) Staid & Kallersjo,
comb. nov. Basionym: Jacquinia umbellata A.
DC., Prodr. 8: 150. 1844. TYPE: Puerto Rico.
Sine loco, Bertero s.n. (holotype, G-DC, seen
on microfiche; isotypes, II. MEE, MO. P).
Literature Cited
Colla, L. A. 1824. Hortus ripulensis. Torino.
Kallersjo, M. A1 8. St&hl. 2003. Pliylogeny oi the Theo-
phrastaeeae (Ericales s.l.). hit. J. PI. Sci. 164: 579-591.
- . G. Bergqvist & A. A. Anderberg. 2000. Generic
realignments in primuloid families of the Ericales s.l.:
A phylogenetic analysis based on ON A sequence from
three chloroplast genes and morphology. Amer. J. Bot.
87: 132.5-1341.
Mez. C. 1903. Theophrastaceae. Pp. 1—21 in A. Engler
(editor). Das Pflanzenreich 15 (IV, 236a).
St&hl, B. 1989. A synopsis of Central American Theo-
phrastaceae. Nordic J. Bot. 9: 15—30.
- . 1992. On the identity of Jacquinia armillaris
( I heophrastaeeae) and related species. Brittonia 44:
54-60.
- . 1993. Votschia , a new genus of the Theophras¬
taceae from southeastern Panama. Brittonia 45: 204—
207.
- . 1995. A synopsis of Jacquinia (Theophrastaceae)
in the Antilles and South America. Nordic J. Bot. 15:
493-51 I.
Votsch, W. 1904. Neue systematische-anatomische Unter-
suchungen von Blatt und Aehse der Theophrastaceen.
But. Jahrb. Syst. 33: 502—546.
New Taxa and Nomenclatural Notes on the Flora of the Marojejy
Massif, Madagascar. VT. Rubiaceae: A New Species of Lemyrea
John l\. Stone
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
john.stone@mobot.org
Aaron P. Davis
The Herbarium, Royal Botanic Gardens, kew, Richmond, Surrey, TW9 3AE, U.k.
a.davis@rbgkew.org.uk
Abstract. Lemyrea marojejyensis J. R. Stone &
A. P. Davis, a new species from t tie Marojejy Na¬
tional Park in Madagascar, is described and illus¬
trated. The thick branchlets, large leaves, shortly
pedicellate flowers, and large number of ovules (ca.
32 per locule) distinguish it from other species ol
Lemyrea. Systematic observations on the genus Le-
myrea are provk led.
Key words: Lemyrea , Madagascar, Marojejy,
Octotropideae, Rubiaceae.
Madagascar is a country rich in endemic species
and remains poorly known botanically (Morat &
Lowry, 1997). In particular, the isolated massifs of
northern Madagascar are extremely diverse and
have yielded many new species. The broken chain
of upland areas that traverses northern Madagascar
contains some of the largest areas of remaining pri¬
mary forest and undisturbed natural areas. While
several of the larger mountains are included within
protected areas, much of the area in between is
unprotected and its vegetation remains threatened.
This northern massif region remains poorly known
botanically, but the available collections indicate
that the region is home to a considerable number
of endemic species. The massif at Marojejy has
been one of the better collected of these isolated
uplands (Humbert, 1955), yet new discoveries are
still being published on a regular basis (e.g.. Miller.
1998; Miller & Randrianasolo, 1998; Miller, 2000).
'file central portion of the Marojejy Massif and
some of the surrounding lowlands comprise the
60,050-hectare protected area. Formerly known as
the Reserve I ntegrale No. 12. established 31 De¬
cember 1927, the status of this protected area was
upgraded to a National Park (Marojejy Reserve Na-
lurelle Integrale) on 19 May 1998.
The middle elevations (ca. 600—1200 m) of the
Marojejy Massif seem particularly rich ami diverse
in woody plant families. Araliaceae, Arecaceae,
Clusiaceae, Cunoniaceae, Myrsinaceae, and Rubi¬
aceae are particularly well represented; the her¬
baceous family Balsaminaceae is represented by
more than 30 species (Morat & Lowry, 1997). I hese
slopes have been the site of many newly described
taxa: e.g., Ardisia marojejyensis J. S. Miller & Pi-
poly (Myrsinaceae) (Miller & Pipoly, 1993); Sabicea
marojejyensis Razafimandimbison & J. S. Miller
(Rubiaceae) (Razafimandimbison & Miller. 1999);
Rhodolaena macrocarpa G. K. Schatz, Lowry & A.
E. Wolf (Sarcolaenaceae) (Schatz et al., 2000);
Weinmannia marojejyensis J. S. Miller & J. Bradford
(Cunoniaceae) (Bradford & Miller, 2001). Recent
collections from these middle-elevation [tarts of the
reserve now yield another new species of Rubi¬
aceae belonging to the genus Dnnyrea (A. Cheva¬
lier) A. Chevalier & Beille.
Lemyrea marojejyensis J. R. Stone & A. P. Da¬
vis, sp. nov. TYPE: Madagascar. Prov. Antsi-
ranana: Marojejy Reserve Naturellc Integrale,
path from Mandena to summit of Marojejy,
700-1000 m, I4°26'S, 49°45'E, 2 Oct. 1994
(fl), Lewis et al. 1249 (holotype, MO; isotypes,
BR, G, WAG, K. P, TAN). Figure I.
Arbor usque 9 m alta, rarnis crassis et nodos tumidis.
Folia coriacea, lamina late elliptica usque late obovata,
7.5-10 cm longa, 4-5.5 cm lata, hdlorescentiae axillares,
binae, 1.9-2. 2 cm longae. Pedicelli 1.5— 2.4 mm longi.
Corolla late infundibuliformis, earnosa, 1.3— 1.5 cm longa,
5—8 mm lata. Ovarium 2-loculare, ovulis in quoque loculo
circa 32.
Tree, 5—9 m tall. Branchlets terete to subquad-
rangular, swollen at nodes, 3.5—7 mm diam.,
smooth to somewhat scaly, brown to light gray or
whitish. Leaves: petioles distinctly articulated at
the junction with the branchlet, 0.3—1 cm long, gla¬
brous, base adaxially canaliculate; lamina broadly
elliptic to broadly obovate, 7.5—10 X (3.4— )4— 5.5(—
Novon 14: 1 19-123. 2004.
120
Novon
Figure 1. lemyrea marojejyensis J. R. Stone & A. I*. Davis. — A. Haliit. — B. Flower, cut to reveal internal corolla
surface, anthers, and style. Drawn from the holotype: Irwis et 4 cm wide; floral pedicel
^ 2.4 mm long; ovules > 30 per loeule
. A. marojejyensis
2b. Leaves usually < 3.5 cm wide; floral ped¬
icel > 5 mm long; ovules < 10 per loeule.
3a. Corolla throat hairs present on almost
the entire surface of the corolla tube;
style glabrous, divided to the hase
but adnate for most of its length
. L ciliolata
3b. Corolla throat hairs present only at
the base of the corolla tube; style pu¬
bescent, divided into two lobes but
not to the base . A. krugii
Acknowledgments. Fieldwork at Marojejy was
funded by the National Geographic Society (3647-
87), the World Wildlife Fund-U.S., and the World
Wide Kund lor Nature (project #6502). We thank
the directors and staff of the following herbaria for
placing herbarium material of Irrnyrea at our dis¬
posal: k. MO. P. and TKK. J. Stone thanks Jim Mill¬
er and Armand Randrianasolo for valuable com¬
ments on the manuscript. Bee Gunn provided the
illustration, and Roy Gereau provided advice on the
Latin description, f ieldwork was conducted under
a collaborative agreement between the Missouri Bo¬
tanical Garden and the Parc Botanique et Zoolo-
gique de Tsimbazaza. We are grateful for courtesies
extended by the Government of Madagascar
through the Direction Generale de la Gestion des
Resources Forestieres and the Association Nation-
ale pour le Gestion des Aires Protegees (ANGAP).
A. Davis acknowledges the Leverhulme Trust,
which funded part ol the research for this paper
under the Rubiaceae of Madagascar Project.
Literature Cited
Beille, L. 1939. Structure anatomique des Lemyrea. Rev.
lut. Bot. Appl. Agric. Trop. 19: 250-255.
Bradford. .1. A .1. S. Miller. 2001. New taxa and nomen-
clatural notes on the flora of the Marojejy Massif. Mad¬
agascar. V. Cunoniaceae: Weinmannia. Adansonia. ser.
3, 23: 219-236.
Capuron, R. 1973 [Unpublished manuscript). Revision
des Rubiaof'es de Madagascar et des Comoros. Copies
at BR. k, MO. P. and TEF.
Chevalier, A. 1938. Fssai d’un groupement systematique
des Cafeiers sauvages de Madagascar et des lies Mas-
careignes. Rev. Int. Bot. Appl. Trop. 18: 825—843.
- . 1939. A propos des Lemyrea de Madagascar. Rev.
Int. Bot. Appl. Agric. Trop. 19: 250.
Humbert. 11. 1955. Une merveille de la Nature & Mada¬
gascar. Premiere exploration botanique du massif du
Marojejy et de ses satellites. Mem. Inst. Sei. Madagas¬
car 6(B): 1—27 1 .
Miller, J. S. 1998. New taxa and nomenclatural notes on
the flora of the Marojejy Massif, Madagascar — I. Cap-
paraceae: A new species of Crataeva. Novon 8: 1 67 —
1 69.
- . 2000. Newr taxa and nomenclatural notes on the
flora oi the Marojejy Massif, Madagascar IV. Myrtaceae:
New species of Eugenia. Adansonia, s£r. 3, 22(1): 1 1 1-
1 16.
- & J. J. Pipoly. 1993. A new species of Ardisia
(Myrsinaceae) from Madagascar. Novon 3: 63-65.
- & A. Randrianasolo. 1998. New taxa and nomen¬
clatural notes on the flora of the Marojejy Massif — II.
Volume 14, Number 1
2004
Stone & Davis
Lemyrea marojejyensis from Madagascar
123
Anacardiaceae: A new species of Campnosperma. No-
von 8: 170-1 72.
Morat, P. & P. P. Lowry II. 1997. Floristic richness in llie
Africa— Madagascar region: A brief history and prospec¬
tive. Adansonia, ser. 3, 19: 101—115.
Razafimandimbison, S. & J. S. Miller. 1999. New taxa and
nomenclatural notes on the flora of the Marojejy Massif,
Madagascar — Ill. Rubiaceae — A new' species of Sabi-
cea. Adansonia, ser. 3, 21: 41—45.
Robbrecht, E. 1980. The Hvpobathreae (Rubiaceae— Ixo-
roideae). 1. Delimitation and division of a new tribe.
Bull. Nat. Pantentuin Belg. 50: 69-77.
- . 1988. Tropical Woody Rubiaceae. Opera Hot.
Belg. 1: 1-127.
- . 1994. Supplement to the 1988 outline of the clas¬
sification of the Rubiaceae. In: E. Robbrecht (editor).
Advances in Rubiaceae Macrosysteniatics. Opera But.
Belg. 6: 173-196.
- , I). M. Bridson & D. B. Deb. 1994. The South
Indian genus Octotropis (Rubiaceae). In: E. Robbrecht
(editor), Advances in Rubiaceae Macrosysteniatics. Op¬
era Bot. Belg. 6: 81-91.
Schatz, 0. E. 2001 . Generic Tree Flora of Madagascar.
Royal Botanic Gardens, Kew, and Missouri Botanical
Garden, St. Louis.
- . P. P Lowry II & A. E. Wolf. 2000. Endemic
families of Madagascar. VI. A synoptic revision of Rho¬
dolite tin (Sarcolaenaceae). Adansonia, shr. 3, 22: 239—
252.
Resolving a Nomenclatural and Taxonomic Problem in Mexican
Oenothera sect. Hartmannia (Tribe Onagreae, Onagraceae)
Warren L. Wagner
United Stales National Herbarium, Department of Systematic Biology, Botany, MRC-166,
P.0. Box 37012, Smithsonian Institution, Washington, DC 20013-7012, U.S.A.
wagner.warren@nmnh.si.edu
Abstract. Study of type material from the trans-
volcanie region of Mexico of two species of Oen¬
othera sect. Hartmannia (tribe Onagreae, Onagra¬
ceae), currently known as Oenothera deserticola
(Loesener) Munz and O. purpusii Munz, indicates
that they represent the same species. Thus, the lat¬
er name O. purpusii becomes a new synonym of O.
deserticola. Also, a neotype is designated herein for
0. deserticola. The oidy available name for the sec¬
ond species, previously incorrectly r ailed O. deser¬
ticola , is Hartmannia montana Hose, but since O.
montana is already occupied, a new name, O. ori-
zabae, is here proposed. A previously unreported
chromosome count of n = 7 is given for O. deser¬
ticola. While examining collections to resolve this
problem, a series of specimens from the Sierra Ma-
dre Occidental was determined to represent a third
species here described as 0. luciae-julianiae.
These three species can be distinguished by a suite
of characters involving habit, petal color, and cap¬
sules. Oenothera deserticola is characterized by de¬
cumbent stems, morning-opening flowers, rose pur¬
ple petals, and angled, but unwinged capsules with
an attenuate apex, while O. luciae-julianiae has
erect or ascending stems, evening-opening (lowers,
white petals, and narrowly winged capsules with an
abruptly acuminate apex. Oenothera orizabae has
decumbent stems, evening-opening flowers, white
petals, and large-winged capsules with a rounded
apex.
Resumkn. El estudio de los tipos de Oenothera
deserticola y O. purpusii de la seeeion Oenothera
seccidn Hartmannia procedentes de la region tras-
voleaniea de Mexico, indica que ambos representan
la misma especie. Como resultado, 0. purpusii es
considerado sindnimo de O. deserticola. IJna segun-
da especie, originalmente descrita como Hartman¬
nia montana , pero cuyo epfteto especffico esta ocu-
pado on Oenothera , es substituido por el nombre
O. orizabae , el eual es aqm propuesto. El estudio
de las colecciones procedentes de la Sierra Madre
Occidental determine que existe una tereera es¬
pecie, la eual es aqui descrita como 0. luciae-ju¬
lianiae. Estas tres especies pueden ser distinguidas
por una serie de caracteres relacionados al hahito,
color de petalo y de las capsulas. Oenothera deser¬
ticola es caracterizada por tallos decumbentes, flo-
res de apertura matutina, petalos purpureo-rosados
y capsulas angulosas, dpteras y atenuadas hacia el
apice, mientras que la especie de la Sierra Madre,
aquf descrita como O. luciae-julianiae, tiene tallos
erectos o ascendentes, flores (pie abren al auoehe-
< er, petalos blaneos y capsulas estrechamente ala-
das con un apice abruptamente acuminado. Oenot¬
hera orizabae tiene tallos decumbentes, flores que
abren al anochecer, petalos blaneos y capsulas ala-
das, grandes, con el apice redondeado.
Key words: evening-primrose, Mexico, Oeno¬
thera sect. Hartmannia , Onagraceae.
Nearly one third of the 120 species of the genus
Oenothera L. occur in Mexico. Recent focus on the
phylogeny of the family Onagraceae, especially of
the most diverse tribe Onagreae (Levin et al.,
2003), along with analyses of biogeographic pat¬
terns of tribes Epilobieae, Gongylocarpeae, and
Onagreae (Kalinas et al., in press) have resulted in
the need to resolve a nomenclatural problem and
describe a new species.
Two relatively uncommon species from tlu1 trans-
volcanie region of Mexico, Oenothera deserticola
(Loesener) Munz and O. purpusii Munz, have had
a confused application of names since the latter was
described in a revision of Oenothera sect. Hart¬
mannia (Munz, 1932). A brief history leading up
to the confusion clarifies the problem. The first spe¬
cies was described as Hartmannia montana J. N.
Rose (1905). The second was published as Xylo-
pleurum deserticolum Loesener (1913). both of
these species were included in Munz's revision of
Oenothera sect. Hartmannia (Munz, 1932), but for
some reason he was confused on the correct appli¬
cation of Xylopleurum deserticolum. Munz's descrip¬
tion ol O. deserticola clearly fits the taxon that .1.
N. Rose described as Hartmannia montana (1905).
Novon 14: 124-133. 2004.
Volume 14, Number 1
2004
Wagner
Oenothera Sect. Hartmannia
125
It was characterized by decumbent stems, white
petals, and large-winged capsules with a rounded
apex. Munz cited a number of collections of it. in¬
cluding the type of Hartmannia montana, and
placed Rose’s name in synonymy of 0. deserticola.
Munz also cited all four syntype collections given
in the original publication ol Xylopleurum deserti-
colum (Loesener, 1913). It is not clear if Munz had
actually seen any of the syntype collections or if he
merely cited them from Loesener’s protologue,
while at the same time selecting one of them as the
type. For reasons unknown to me, Munz was con¬
fused because at least one ol the syntypes, Seler
5614 (photo US), and likely all four judging from
Loesener’s description, is clearly the same taxon as
the one he described in the same paper (1932) as
the new species O. purpusii Munz. This species was
characterized by decumbent stems, rose-purple
petals, and smaller, angled, but unwinged capsules
with an attenuate apex.
A photo of one of the four syntypes was taken
before all four were destroyed in the Berlin her¬
barium during World War II. This photo is repro¬
duced here in Figure 1. I had hoped to confirm the
characteristics of original material of O. deserticola
by either locating duplicates of the leetotype, other
syntype collections, or photographs of any of the
other three syntypes, but none were found by the
collections managers who kindly searched for them
at the herbaria known to have received Seler col¬
lections (A, B, CAS, F, GH, K, MEXU, NY. US).
Despite not having any of Loesener’s original ma¬
terial, it is clear from the leaves and dehisced cap¬
sule shown in the photograph that O. deserticola
represents the same taxon that has gone under the
name 0. purpusii since 1932. The name Oenothera
deserticola has priority over 0. purpusii , and there¬
fore is the correct name. This conclusion is further-
supported by Loesener’s original description
(1913), as characterized earlier.
The other species in question was first described
in 1905 as Hartmannia montana Rose. Since the
specific epithet montana is already occupied in Oe¬
nothera, a new7 name, 0. orizabae, is proposed here.
During the course of study of these two species
using material from a number of primarily Ameri¬
can and Mexican herbaria, it became clear that a
significant number of Mexican collections from the
Sierra Madre Occidental, from Chihuahua south
through Durango to Jalisco and Guanajuato that
were identified as O. deserticola by I’. II. Raven,
myself, or others over the past three decades rep¬
resents an undescribed species characterized by
erect or ascending stems, white petals, and narrow¬
ly winged capsules with an abruptly acuminate
apex. It is here described as 0. luciae-julianiae. It
has a distribution allopatric from both O. orizabae
and 0. deserticola. The following key distinguishes
them from one another.
Key to Species of Mexican Oenothera sect. Hartman
N1A
la. Capsules winged, acuminate to rounded at apex;
petals white at anlhesis, fading purple.
2a. Stems decumbent, ascending toward the
apex; cauline leaf blades 1—3 cm wide, nar¬
rowly ovate, occasionally elliptic, subentire to
subdenticulate; capsule body 15—26 mm
long, obovoid, apex rounded, the margins
winged, the wings 2.3-4' mm wide, the sterile
stipe stout, 15—50 mm long; capsule body ap¬
parently dehiscing only at apex ... O. orizabae
2b. Stems erect to ascending; cauline leaf
blades 0.5— 1.1 (—3) cm wide, elliptic to lan¬
ceolate or narrowly elliptic-ovate, denticu¬
late to serrulate; capsule body 8— 18(— 25)
mm long, clavate or narrowly obovoid, apex
obtuse to bluntly acuminate, the wings 1.3—
2 mm wide; the sterile stipe slender, 7—35
mm long; capsule body dehiscing from about
half its length to nearly throughout . . .
. 0. luciae-julianiae
lb. Capsules angled, but not winged, attenuate at
apex; petals rose-purple at anlhesis, fading dark¬
er . O. deserticola
Oenothera deserticola (Loesener) Munz, Amer.
J. But. 19: 758. 1932. Xylopleurum desertico-
lum Loesener, Repert. Sp. Nov. 12: 238. 1913.
TYPE: M exico. Distrito Federal: Contreras,
Lyonnet 532 (neotype, designated here,
MFXU-238843; isoneotypes, Gil, K not seen,
MEXU [3J, MO, NY, US). Figure 2.
Oenothera purpusii Munz, Amer. J. Hot. 19: 759. 1932.
Syn. nov. TYPE: Mexico. Ixtaceihuatl, open woods,
1903, C. A. Purpus 1839 (holotype, POM 32827 not
seen; isotypes, F, G not seen, GH, MO, NY, UC, US).
Perennial herb with usually several or sometimes
more, decumbent or sometimes ascending when
short, branched or occasionally simple stems 5—
20(— 36) cm long, usually reddish purple or occa¬
sionally pale purple, strigillose, rarely with a few
longer erect hairs, often producing new rosettes via
rhizomes up to 6-7 cm long, from a thick somewhat
woody caudex; root fleshy and thickened in age.
Rosette leaf blades 4-6 X 1-1.7 cm, oblanceolate
to elliptic, sparsely to moderately strigillose, ap¬
parently quickly deciduous; petiole 0.4- 1.6 cm;
cauline leaf blades 1-6.5 X 0.5-1. 8 cm, elliptic to
elliptic-oblanceolate or narrowly so, subserrate to
weakly sinuate-toothed, apex obtuse lo acute,
sparsely to moderately strigillose; petiole 0.1— 1.1
cm long. Flowers opening near sunrise (litis et al.
126
Novon
Museum botanicuni Berolinense.
Cucc. et Ed. Selcr: 1’lnnt mexicnnae.
1 ^,1 HO/rf
T, /?*■(» ..„ 'ir’r-, /*. v>»»
, ...
t/1
figure I. Reproduced photograph of Seler 5611 , a Berlin ( It) syntype of Xylopleurum deserticolum l.oesener no longer
extant, taken as part of the Held Museum’s (f) Types of the Berlin Herbarium project. Copies of the photograph are
in MICH, MO. NY, ROM, and US. The photograph was scanned into Photoshop 6.5, and then a dehisced capsule
showing the long apex of the valves allowing for unambiguous identification was cut. enlarged, and pasted into the
upper left-hand corner (obscuring two small branches evident in the original). A white box on the full photograph
indicates the part cut and enlarged, file leaves and part of a stem were removed from the inset for clarity.
202 says evening); floral tube, sepals, and ovary
densely strigillose, and ovary usually also densely
hirtellous with hairs up to 2 nun long; floral tube
4.5—1 1 mm long; sepals 7.5—16 mm long; free tips
in bud absent or rarely present and ca. 0. 1-0.2 mm
long; petals {1.5-17 mm long, rose-purple fading
darker after pollination; filaments 5.5-8 mm long,
rose-purple; anthers 2.5— 4.5 mm long, cream; pol-
Volume 14, Number 1
2004
Wagner
Oenothera Sect. Hartmannia
127
Figure 2. Oenothera deserticola (Loesener) Munz. — A. Habit, showing decumbent stems from a branched rhizomatous
base (Arroyo 182, MO. and Balogh 1005, US). — B. Bud, floral tube, and ovary, showing the lack of free sepal tips
and pubescence pattern (Arroyo 182, MO). — C. Flower, showing petal shape and overlap of petals (Hinton 9104, US).
— 1). Young capsule showing shape, attenuate apex, and sterile stipe (Lyonnet 532, MEXU). — E. Dehisced capsule
showing length of dehiscence and seeds clustered on placentas (Straw & Gregory 1116, OH). — E Dehisced capsule
showing shape of open valves (Lyonnet 532, US).
len cream, > 90% fertile; style 12-21 mm long;
stigma surrounded by the shedding anthers at an-
thesis or slightly elevated above it; stigma lobes
2.5—4 mm long. Capsule body 10—22 mm long.
rose-purple to green with purple tinge, the body
narrowly ellipsoid to clavate, apex attenuate, angled
or rarely with a narrow wing up to 0.5 mm wide;
sterile stipe slender, 7-17 mm long; capsule body
128
Novon
dehiscing nearly throughout; seeds clustered in
each loeule, 1—1.2 nun long, 0.4— 0.6 mm thick,
brown. Gametic chromosome number, n — 7 (Ha¬
ven, unpublished).
Distribution. Oenothera deserticola occurs in
open forests of Pinns, Pinus-Quercus, or occasion¬
ally Abies, Alnus, and Salix, on sandy open sites or
steep banks, in the mountains of the Trans-Mexican
Volcanic region from the northeastern Michoaean
and central Hidalgo south to Mexico and Orizaba,
Veracruz, from (2100)2500 to 3400 m.
The leetotype selected by Munz (1932), but now
destroyed, was collected in Mexico, D.K., Kl De-
sierto de los Leones, May 1909, IT Hopfer s.n. (C.
tV E. Seler 5280) (B destroyed). The leetotype and
three additional syntypes (without further locality
Uhde 1222; El Desierto de los Leones, IT Hopfer
s.n. (labeled as Seler 56 14) [photos: MICH, MO.
NY, POM, US]; and San Halael, Tlalmanealo, Seler
5810) were destroyed during World War II (Chris¬
toph Oberprieler, pers. comm. 2001).
Oenothera deserticola is a member of section
Hartmannia, which can be subdivided into three
groups of species based on petal color and time of
dowering. With its rose-purple petals and morning
flowering, O. deserticola appears to be in a group
with other morning-flowering species with purple
petals, including O. platanorum I*. H. Haven & 1).
H. Parnell, 0. rosea l.’l lerilier de Brutelle ex Aiton,
and O. texensis P. H. Haven & I). H. Parnell. These
species share angled, but not winged capsules and
no free sepal tips in bud or tips free and minute
(0. 1-0.2 mm long).
The geographic and elevational range of Oenot¬
hera deserticola broadly overlaps the range of 0.
orizabae. One collection {Barkley et al. 2488) is a
mixed collection of O. deserticola and O. orizabae
suggesting that they occur sympatrically at some
localities; however, there is no documentation of
hybridization. Hybridization must be strongly lim¬
ited by the different timing of flowering, with Mow¬
ers of O. deserticola open during the day and those
of O. orizabae open al night.
Specimens examined (those marked by * are chromosome
determination vouchers). MEXICO. Distrito Federal:
Desierto de los Leones, * Raven 15424 (also flavonoid
voucher) (TKX); hwy. 190, betw. Puebla & Mexico City,
Ahshapanek 701 (TKX); Contreras Canyon, SW of Mexico
City, Munz 15000 (GH, MO); Chapultepec III. Valle del
Sileneio, Wonderly 217 (MKXU, NMC); Cerro de Esqui-
Huil, dclcgacidn de Mipa Alta. Ventura 2804 (MKXU, MO.
NY); Kfo Krfo, Bussell & Souviron 82 (US); Volcan Pelado,
ladera K, Mondragdn s.n. (MKXU); near Kstacidn La
Cima, Serranfa del Ajuseo, Rzedowski 10840 (MICH. MO,
TKX); Tlalpan, Volcan Ajuseo, Reyes s.n. (MKXU); lava
fields ea. 2 km SSW of La Cima R.B. Station on either
side of old Hwy. 95 on top of Serjana de Ajuseo, litis et
al. 202 (MICH, MSA); Cerro Tepetlahualo, delegacibn de
Milpa Alta, Ventura 1412 (MKXU); Santa Rosa, delega-
cidn de A. Obregon, Ventura 2720 (MKXU, MO); I mi.
SK of Puerto del Aire, Barkley et al. 2488 (MICH, TKX)
| hath sheets mixed collection with 0. orizabae ]. Hidalgo:
Cerro de los Pitos, Matilda 21562 (MKXU |2|); Kl Chico,
Cerro de Las Ventanas, Gentry et al. 82180 (MO); Parque
Nacional Kl Chico, Ortiz 10 (MKXU); Met/.quitillan, Car-
pinteros. 20 km K Metzquititlan, Hernandez 6806 (MO);
Sierra de Pachuca, Rose & Hay 56/ 1 (US). Mexico: on
hwy. Mexico to Puebla, Munz 15064 (POM); Monte do Rio
Krfo, km 49, road from Mexico City to Pueblo, Mexia
2606a (IJC); near Salazar, Rose & Painter 7027 (US); Ne¬
vada de Toluca, Rose A Painter 7878 (US); El Crucereo,
Temascaltepec, Hinton 521 (POM); Cerro de Ledn, Valle
de Mexico, Matuda 10128 (MKXU |2|): Paraje Provincial.
Balls 4178 (UC, US); San Rafael, ladera Vi del Iztaeei-
luiatl, Sanchez 8 (MKXU); Ixtapaluca, Kstacidn Experi¬
mental de Ensenanza y Investigaeidn de Zoqniapan, Koch
& Magana 75242 (MKXU, MO, NY); Llano Grande Gap,
near Rfo Krfo, Sharp 4406 (MKXU. RSA); Nat. Park La¬
gunas de Zempoala, Cole 40 (MKXU), McAdams 57
(MICH), Traylor 55 (MKXU); Comunidad, Temascaltepec,
Hinton 860 (GH, NY, POM); Cumbre, Temascaltepec,
Hinton et al. 7478 ((41. LL, MICH [2|, MO, NY, POM,
TKX, UC, US); Cumbre Trojes, Temascaltepec, Hinton et
al. 8272 (MKXU, MICH, NY, UC, US); Ml. Popoeatapetl,
Nelson 10 (US); 10 km al NE del Entronque a Sultepee,
Flores A- Terpdn 820 (MKXU); Vallee de Bravo, 0.8 mi. S
ol Cajones, 9500 ft., Balogh 1005 (US); 55 km SK of
Mexico City, 10,500 ft.. Weaver 712 (POM. US); near To¬
luca, Schery 64 (MO); Ocuilan. Matuda 82454 (MKXU).
IYlichouran: near base of Cerro de Tecolote near Zaoapu,
Sharp 45581 (RSA); Cerro del Burro, Santa Clara del Co-
bre, Perez 58 (MKXU, MO), Escobedo 1400 (MO); Cerro
Prieto, Nuevo Parangarieutiro, Medina 2684 (MKXU); En
Kl Salto, a 7 km al E de Ocampo, Melchor Ocampo, Mar¬
tinez & Torres 410 (MKXU): Ocampo, I I km al SK de
Ocampo, en el Cerro Kl Chivati, Solo Nunez & Solorzano
12600 (MKXU); Ouerendaro. cerea de San Jose de la
Cumbre, Santos 1801 (MKXU); ladera SW del Cerro San
Andres, Zinapeeuaro, Zamudio 5520 (MKXU, MO); lado
N do La Presa La Gachupina, Jasso 1020 (MKXU); ea. 18
mi. S of Patzcuaro, King & Soderstrom 5180 (TKX, US);
near Parieutin volcano, Eggler 84 (US). Morelos: km 48.
earretera federal, Vazquez 1871 (MKXU [2]); Lagunas de
Zempoala, * St raw & Gregory 1068 ((ill. MKXU, MICH.
RSA, UC); Huitzilac. ladera NE del l.ago llueyapan, Par¬
que Nacional Lagunas de Zempoala, Avonce s.n. (MEXU).
Puebla: 4 km E of Rfo Krfo. Roe et al. 848 (MICH, US);
side ol Popocatepetl, Miranda & Barkley I7M200 (K.
I EX); Kl Salto, Rfo Krfo, Boege 8078 (MKXU). Veracruz:
Lomogrande. Mt. Orizaba, Balls 4877 (UC, US). Without
locality: Gregg 607 (MO).
Oenothera orizabae W. L. Wagner, nom. nov.
Hartmannia montana Rose, Contr. U.S. Natl.
Herb. 8: 329. 1905, non Oenothera montana
Nuttall. in Torrey & Cray, FI. N. Amer. I: 500.
1840. TYPE: Mexico. El Cima, on the railroad
between Mexico, D.F., and Cuernavaca, Mex¬
ico, 19 Sep. 1903, ,/. N. Rose & ./. II. Painter
7170 (holotype, US-00450745; isotypes. GH,
MEXU, NY, US). Figure 3.
Volume 14, Number 1
2004
Wagner
Oenothera Sect. Hartmannia
129
Figure 3. Oenothera orizahae W. U. Wagner. — A. Habit, showing decumbent steins from a multi-stemmed base ( Straw
& Gregory 1120, GH). — B. Bud, floral tube, and ovary, showing the free sepal tips and pubescence pattern ( Straw &
Gregory 1120, GH). — C. Flower, showing petal shape and expected non-overlap of petals (Straw & Gregory 1120,
MICH). — D. Young capsule, showing shape, wings, rounded apex, and sterile stipe (Rose <& Rainier 7170, holotype,
US). — E. Dehisced capsule, showing known length of dehiscence (Rose & Painter 7170, isolype, US).
130
Novon
Perennial herb with several decumbent to as¬
cending, simple or usually branched stems, 6—46
cm long, the plant hirtellous throughout. Rosette
leaf blades 3—5 X 0.9— 1.5 cm, elliptic to oblan-
ceolate, hirtellous; petioles 0.3—1 cm long. Cauline
leal blades 3—9 X 1—3 cm, narrowly ovate, occa¬
sionally elliptic, subentire to subdentieulate; peti¬
ole 0.1— 2.2 cm long. Flowers opening near sunset;
floral tube, sepals, and ovary densely hirtellous
with hairs up to 1.5 mm long; floral tube 9—15 mm
long; sepals 15—20 mm long; free tips in bud 1.7-
2 mm long; petals 18-22 mm long, white, fading
purple; filaments 15—16 mm long, pale maroon; an¬
thers 3—7 mm long, cream; pollen cream-colored,
> 90% fertile; style 20—28 mm long; stigmas held
above the anthers at anthesis, the lobes 4—5 mm
long. Capsule body 15—26 mm long, obovoid, apex
rounded, the margins winged, the wings 2.3—4 mm
wide; the sterile stipe stout, 15—50 mm long; cap¬
sule body apparently dehiscing only at apex; seeds
1 — 1.4 mm long, 0.5— 0.7 mm thick, brown. Gametic
chromosome number, n = 7 (forming 7 pairs at
meiotie metaphase I; Gregory & Klein, 1960; Ha¬
ven, unpublished).
Distribution. Oenothera orizabae is scattered in
rocky grasslands in Pinus-Quercus forest at middle
elevations in the Trans-Mexican Volcanos from Mo¬
relos and Mexico to Kl Chico to the north in Hidalgo,
and Pico Orizaba in Veracruz, at 2980-3000 m.
Oenothera orizabae appears to be allied to a
group of evening-flowering species with white pet¬
als currently assigned to section Hartmannia , in¬
cluding O. kunthiana (Spach) Munz, and 0. tetrap-
tera Cavanilles, but based on recent molecular
studies this white-petaled group may need to be
distinguished as a separate section (Levin et al.,
2004). Another while-flowered species, Oenothera
dissecta A. Gray ex S. Watson, may also belong in
this group, but was most recently assigned to sec¬
tion Gauropsis (Wagner, 1984). A single collection
was examined for pollen fertility using Alexander
stain (Alexander, 1969).
Specimens examined (those marked by * are chromosome
determination vouchers and voucher for pollen fertility in¬
dicated by +). MEXICO. Distrito Federal: I mi. SE
ol Puerto del Aire, Barkley et al. 2438 (MICH, TEX) [both
sheets mixed with 0. desert icola\\ Estacibn I. a Cima. Prin¬
gle 10293 (F, (41, MEXU [2], MICH [2], MO. NY, UC.
US), *Raven s.n. (Stanford greenhouse #67-354', ffavonoid
voucher) (TEX); Colonia del Valle, Lyonnet 198 (US); ca.
2 km SSW of Ua Cima It. 8. Station, old llwr. 95. litis et
al. 200 (MICH. RSA); Volean Pelado. ladera W, Sandoval
230 (MEXU). Ilidal^n: El Chico, Lyonnet 2205 (MEXU).
Mexico: S of Mexico City, hnwenworth & Leavenworth
954 (F); 55 mi. SE of Mexico City, Weaver 713 (POM);
near Gap, near Kfo Frfo, Sharp 4495 (MEXU, KSA); pub¬
lic park area W of divide betw. Mexico & Puebla al the
Estaeionamento, Pennell et al. 125 (MO); 1 km al N de
Llano Grande sobre la ladera S del Telapon, Espinosa 542
(MO); Llano Grande, Telapon. + Lyonnet 689 (MEXU |2|,
US); near Rio Frfo, Puebla hwy., * Straw & Gregory / 120
((41, MEXU, MICH [2], RSA); Rio Frfo, Kenoyer 2485
((ill); just W of Rfo Frfo, 40 mi. E of Mexico City, Man¬
ning A- Manning 53656a (GH). Morelos: Cuernavaca,
Kenoyer A422 (E); 6 mi. W of Yautepec, Walther 133
(MICH). Veracruz,: Pico de Orizaba, Rose & Hay 5672
(US).
Oenothera luciue-juliuniue W. L. Wagner, sp.
nov. TYPE: Mexico. Durango: San Dimas,
Mesa del Roble, San Miguel de Cruces,
24.99°N, 1 05.99° W, 2740 m, 26 May 1990, A.
Garcia & S. Acevedo 345 (holotype, US-
3441077; isotypes, ANSM, CHAPA, CIIDIR,
ENCB, IEB, MEXU, 1) AMI/ (none seen)). Fig¬
ure 4.
Ab Oenothera orizabae caulibus erectis ad adscenden-
tes, laminis foliorum caulinorum 0.5— 1.1 (—3) cm latis
elliptieis ad lanceolatas et denticulatis ad serrulatas, cor-
pore capsulae 8 — 1 H( — 25) mm longo clavato vel anguste
obovoideo ad apicem obtuso vel acuminato alis ejus 1.3—
2 mm latis stipite sterili tenui 7—35 mm longo.
Perennial herb with few to several, ascending to
erect, simple or occasionally branched stems, the
longer ones becoming decumbent and often devel¬
oping lateral branches, 20-50(-90) cm long,
sparsely (very sparsely) to moderately hirtellous,
the hairs 1.5— 2.5 mm long, also strigillose through¬
out or only in the upper portions of the stem, the
hairs mostly 0.3— 0.4 mm long; root a thickened,
often branched taproot. Rosette leaf blades 4—10 X
0.5— 2.5 cm, oblanceolate, denticulate to serrulate,
or weakly sinuate-toothed, sparsely to moderately
strigillose, apex obtuse to rounded; petiole 0—1.5
cm long; cauline leaf blades (1.8— )3-7 X 0. 5-1.1 (—
3) cm, elliptic to lanceolate or narrowly elliptic-
ovate, denticulate to serrulate, apex attenuate,
acute, sparsely to moderately strigillose; petiole 0—
0.5 cm long. Flowers opening near sunset; floral
tube, sepals, and ovary moderately to occasionally
densely strigillose and rarely with scattered hirte¬
llous pubescence with hairs up to 1.5 mm long;
floral tube 12-21 mm long; sepals 14—24 mm long;
free tips in bud 1-2 mm long; petals 16—26 mm
long, white fading deep purplish red; filaments 7—
10 mm long, rose-purple or yellow; anthers 4.8— 6.5
mm long, cream; pollen cream, ca. 50—60% fertile
(range 2U) — 94%); style 20—30 mm long; stigma el¬
evated above to surrounded by the shedding an¬
thers at anthesis, the lobes 3.6—6 mm long. Capsule
body 8— 18(— 25) mm long, gray-green, clavate or
narrowly obovoid, apex obtuse to bluntly acumi¬
nate, winged in the upper part or throughout the
Volume 14, Number 1
2004
Wagner
Oenothera Sect. Hartmannia
131
Figure 4. Oenothera luciae-julianiae W. L. Wagner. — A. Habit, showing ascending to erect stems from a several¬
stemmed base ( Garcui & Acevedo 345, US). — B. Bud, floral tube, and ovary, showing the free sepal tips and pubescence
pattern (Reveal & Harley 4095 , NY). — C. Flower, showing petal shape and apparent slight overlap (Garcia & Acevedo
345, US). — I). Young capsule, showing shape, wings, obtuse to bluntly acuminate apex, and sterile stipe (Reveal <0:
Harley 4095, TEX). — E. Dehisced capsule, showing known length of dehiscence, and valves (although they can be
spreading) (Garcia & Acevedo 345, US).
132
Novon
body, the wings 1.3—2 mm wide; sterile stipe slen¬
der, 7—35 mm long; capsule body dehiscing in up¬
per half to nearly throughout; seeds clustered in
each locule, 1.4— 1.5 mm long, brown.
Distribution. Oenothera luciae-julianiae is scat¬
tered to common in open Pinas or Pinus-Quercus
forests or mixed forests of Abies, Pseudotsuga, Pi¬
nas, Quercus, and sometimes with Arbutus, Juni-
penis, or Cupressus, on rocky or open sites, steep
slopes, or moist flats or along streams in the Mex¬
ican mountains of the Sierra Madre Occidental from
Chihuahua south through Durango, Guanajuato,
Nayarit, Jalisco, Michoacdn, and Q ue re taro, from
21(H) to 2800(— 3250) m. Flowering nearly through¬
out the year, and documented from February
through November.
Etymology. It is a pleasure to name this attrac¬
tive Sierra Madre Occidental species in honor of
my wife, Lucy C. Julian, on the occasion of our 10th
wedding anniversary.
Oenothera luciae-julianiae has gone undetected
for decades of collecting in t he Sierra Madre Oc¬
cidental. The collections cited herein were deter¬
mined most commonly as O. deserticola by P. II.
Haven, myself, or others, but also occasionally as
O. purpusii, O. tetraptera, or O. kunthiana. It was
treated as part of 0. deserticola (now correctly
known as O. orizabae) by Haven and 1). Parnell
(pers. comm.) in their unpublished revision of sec¬
tion Hartmannia. Munz, who revised the section in
1932, never annotated any of the collections, as
very few were made before 1960. .1. N. Rose, how¬
ever, annotated a single collection from Chihuahua
(Nelson 4848) as a new species, but he never pub¬
lished it. Oenothera luciae-julianiae appears to be
allied to a group of evening-flowering species with
white petals currently assigned to section Hartman-
nia, including Oenothera orizabae, O. kunthiana
(Spach) Munz, and O. tetraptera Cavanilles.
Like Oenothera kunthiana of this white-flowered
species group of section Hartmannia, O. luciae-ju¬
lianiae appears to be a permanent translocation
heterozygote (I XFI 1 ) species. In PTH species a ring
of 14 chromosomes is usually formed in meiotie
metaphase. I have not yet been able to obtain cy-
lological material to examine meiosis in this spe¬
cies, and thus current data are only suggestive of
this condition in O. luciae-julianiae. PTH species
in the Onagraceae are nearly always autogamous
(Haven, 1979). The stigma is usually surrounded
by the shedding anthers at anthesis in O. luciae-
julianiae, suggesting that it is predominately autog¬
amous. Another feature of Hi ll species is the bal¬
anced lethals in pollen and ovules, which can be
observed in lowered pollen fertility and fewer ma¬
ture seeds (see Cleland, 1972). I examined pollen
fertility of as many collections as possible of O.
luciae-julianiae. In the 15 collections I examined
with Alexander stain (Alexander, 1969; vouchers
noted with brackets in the exsiccatae) pollen fer¬
tility ranged from 26% to 94%, a very' wide range,
but suggestive nevertheless that O. luciae-julianiae
is probably a PTH species. Typically, PTH species
have about 50% pollen fertility on average, but all
exhibit a range of fertility. For example, Oenothera
nutans G. F. Atkinson & Bartlett exhibited a similar
wide range of fertility (48% to 84%) in a detailed
study (Wasmund, 1990). Haven (1979) pointed out
that most Pi ll species are annuals or biennials,
with about 10% of the known lyfH species in Ona¬
graceae being perennial. If O. luciae-julianiae
proves to be a PrH species, it will represent an¬
other atypical species, as it clearly has a perennial
habit from an enlarged root. Oenothera luciae-julia¬
niae does not appear to be closely related to other
species of the white-flowered species group of sec¬
tion Hartmannia. It may share a common ancestor
with O. tetraptera , an outcrossing pair-forming spe¬
cies that appears to be directly related to the other
PTH species in this group, O. kunthiana.
Paratyjies (those used to study pollen fertility have %
fertile pollen given in square brackets). MEXICO. Chi¬
huahua: ca. 51 ini. S of (Teel, Straw & Forman 1900
(USA) [26%]; Llano Grande, Pennington 181 (TEX)
[37%]; Guachochi, 8 km al W de Cabbrachi, Hernandez
8778 (MEXU); Ocampo, Cascada de Basaseachi, Tenorio
A' Torres 4510 (MEXU); Urique, Rye el al. 15612 (MEXU)
|90%], Tenorio et al. 9 985 (MEXU) [62%]; base of Ml.
Mohinora, 8 mi. from Guadalupe y Calvo, Nelson 4848
(US). Duriinpi: Canolas, Bolanos 1565 (MEXU); 55-60
km SW of Durango City on road to I, a Elor, Breedlove
44184 (MO); from the Sierra Madre, W ol Durango, Forrer
12 (UC); ca. 77 rd. km S of Durango by hwy. to La Elor,
Worthington 8826 (MO); 48 km WNW of Huejuquilla Id
Alto. Jalisco to Canoas, Breedlove & Almeda 59165 (MO);
6 mi. E of Buenos Aires tow. El Salto, Straw & Forman
1795 (BSA) 1 83% |; 31 mi. N of Estaeion Coyotes. Breed¬
love 18741 (MO); 54 mi. N Estaeion Coyotes, just NW ot
Guachichilas, Breedlove 18788 (MO) [59%]; 71 mi. W of
Durango. E of La Campafia, Oliver 715 (MO); I I km SW
of La Ciudad near Buenos Aires. Breedlove 86465 (MO)
|27%|; llwy. 40. 11.5 mi. W of La Ciudad, Wagner &
Solomon 4808 (MO); llwy. 40, at La Campana, Reveal et
al. 2689 (MO) [74%]; llwy. 40. about 81 mi. W of Du¬
rango, Reveal & Atwood 8477 (MO) [39%]; betw. Pinos
Altos & Conohefto, llewitt 118 ((-II) [88%]; El Salto, El
Tapextle, Tenorio A Romero 81 6 (MEXU); 1 km al SW de
El Salto, Tenorio A Romero 927 (MEXU); “Arroyo de El
Salto” al N de El Salto, Tenorio A Romero 759 (MEXU);
17 mi. W of El Salto, Waterfall 12698 ((41. US) [35% |;
25 mi. E of El Salto along Mexican Hwy. 40, Breedlove
15758 (MO); 7 km W of Llano Grande, Hendrickson 1704
(BSA) 1 72%]; ca. 120 road mi. NW of Santiago Papa-
squiaro, Spellenberg A 'Zimmerman 6688 (MO. NMC); 10
Volume 14, Number 1
2004
Wagner
Oenothera Sect. Hartmannia
133
mi. W of El Salto on Mazatlan rd., Straw & (Gregory 1270
(USA); Sierra Madre Occidental, 5.1 rd. mi. by hwy. 40
SW of El Salto at Arroyo de Agua, Worthington H902
(MO); Durango, Ejido Cienega de Eos Caballos, Ortega &
Pacheco HI (CIIDIR, IEB. MEXU, US); Ejido, Encina,
Ortega 20 (CIIDIR, US); El Mezquital, El Zapote, Soil's
902 (CIIDIR, US); Ea Guajolota, Garcia 260 (CIIDIR. US);
Eaguna del Chivo, Acevedo 445 (CIIDIR, IRUG, US); Eas
Minas, Solis II (CIIDIR, US); Nombre de Dios, San Jose
de Ea Parrilla, Sanchez 455a (US), 544 (CIIDIR, US);
Pueblo Nuevo, El Salto, Valenzuela 3—25 (CIIDIR, US);
Predio Eas Rayas de la UJED, Garcia & Acevedo 9H0 (US);
San Rernardino de Milpillas Chico, Sanchez (INI) 9 (US),
19 (CIIDIR, US); San Francisco de Eajas, I.N.I. 31 (CII¬
DIR, US); Santa Barbara, Garcia 1 137a (US); San Dimas,
Eos Aposentos, Garcia et al. 435 (CIIDIR, US). Guana¬
juato: Guanajuato, Rincdn del Toro, Martinez 760
(MEXU 1 2]); Canada de la Virgen, Agua Sabrosa, Martinez
935 (MEXU); 34—35 mi. E of San Euis de la Paz tow.
Xichu, Straw & Forman 1474 (MEXU. RSA). Jalisco:
14— 18 km SW of Tequila on Volcan de Tequila, Breedlove
39226 (MO) [47%]; Volcan Tequila, Reveal & Harley 4095
ll. MEXU, MO. NY. TEX) [94%]. Michoacan: betw. Los
Cabras & Pueblito, Chilchota to Zaeapu, Sharp 45507
(RSA). Nayarit: 105 km WNW of lluejuquilla El Alto
along road to Jesus Maria, Breedlove 6I50H (MO) [94%|.
Queretaro: Colon, parte alta del Cerro Zamorano, Rze-
dowski 44449 (MICH).
Acknowledgments. The holotype and the distri¬
bution of isotypes of Oenothera luciae-julianiae
were kindly provided by S. Gonzalez. Thanks are
given to Alice Tangerini for tin* preparation of the
three excellent plates t licit amply demonstrate the
distinctions among these difficult species of Oenot¬
hera sect. Hartmannia. I thank Denise Mix for
comments on the manuscript, compilation of the
specimens examined, assistance with the loans, and
preparation of Figure 1. I appreciate translations of
the* diagnosis of O. luciae-julianiae to Latin provid¬
ed bv Dan Nicolson. and the translation of the ab¬
stract into Spanish by Pedro Acevedo. I greatly ap¬
preciate the help of Socorro Gonzalez in obtaining
an excellent set of collections of O. luciae-julianiae
from the region surrounding Durango, including the
type. Finally, I am grateful to Peter Hoch for his
excellent comments during the review process,
which improved the clarity and accuracy of this
paper.
Literature Cited
Alexander, M. P. 1969. Differential staining of aborted and
non-aborted pollen. Stain Technol. 44: 117—122.
Gleland, R. E. 1972. Oenothera Cytogenetics and Evolu¬
tion. Academic Press, London.
Gregory, I). P. & W. M. Klein. I960. Investigations of mei-
otic chromosomes of six genera in the Onagraceae. A I i -
so 4: 505-521.
Katinas, E.. J. Crisci, W. E. Wagner & P. C. Hoch. In
press. Geographical diversification of tribes Epilobieae,
Gongylocarpeae, and Onagreae (Onagraceae) in North
America. Ann. Missouri Rot. Card.
Levin, R. A., W. L. Wagner, P. Hoch, M. Nepokroeff, J. C.
Pires, E. A. Zimmer & K. J. Sytsma. 2003. Family-level
relationships of Onagraceae based on chloroplast rhc\.
and ndhV data. Amer. J. Rot. 90: 107—115.
- , - . - . W. J. Hahn, A. Rodriguez, I).
A. Baum, L. Katinas, E. A. Zimmer & K. J. Sytsma.
2004. Evolutionary relationships among and within
Tribes Onagreae and Epilobieae (Onagraceae). Syst.
Rot., in press.
Loesener, L. E. T. 1913. Mexikanische und zentralameri-
kanische Novitaten. IV. Report. Sp. Nov. 12: 217—244.
Munz, P. A. 1932. Studies in Onagraceae VIII. The sub¬
genera Hartmannia and Gauropsis of the genus Oeno¬
thera. The genus Gayophytum. Amer. J. Rot. 19: 755—
778.
- . 1965. Onagraceae. N. Amer. FI. II. 5: 1-278.
Raven, P. H. 1979. A survey of reproductive biology in
Onagraceae. New Zealand J. Rot. 17: 575—593.
Rose, J. N. 1905. Studies of Mexican and Central Amer¬
ican plants — No. 4. Contr. U.S. Natl. Herb. 8: 281—339.
Wagner, W. E. 1984 [ 1 985 1 . Reconsideration of Oenothera
subg. Gauropsis (Onagraceae). Ann. Missouri Rot. Card.
71: 1 1 14-1127.
Wasmund, O. 1990. Cytogenetic investigations on Oeno¬
thera nutans (Onagraceae). PI. Syst. Evol. 169: 69-80.
Four New Species of Nasa ser. Alatae (Loasaceae) in the
Amotape— Huancabamba Zone of Peru
Maxim ilian Weigend
Inslilut fiir Biologie— Systematische Botanik mid Pflanzengeographie, Freie Universitat Berlin,
Altensteinstr. 6, D- 14195 Berlin, Germany, weigend@zedat.fu-berlin.de
Abstract. Four new species of Nasa ser. Alatae
(Urban & Gilg) Weigend are described from the
Amotape— Huancabamba Zone in northern Peru.
The new species Nasa stolonifera has extensive un¬
derground stolons and forms clonal stands, whereas
N. victorii has angular stems with elevated longi¬
tudinal ridges. New species Nasa pongalamesa and
N. lambayequensis are closely allied to N. carnea
from northern Peru and N. loxensis front Ecuador.
Nasa amaluzensis Weigend, previously only known
from tin* type collection in southern Ecuador, is
reported as new to the Peruvian flora. The flowers
of Nasa ser. Alatae are discussed as funnel-revolver
flowers, based on their morphology. A distribution
map of the newly described taxa and a key to all
known species of Nasa ser. Alatae in Peru are pro¬
vided.
Key words: Amotape— Huancabamba Zone, fun¬
nel-revolver flower, Loasaceae, Nasa ser. Alatae,
Peru.
A revision of Nasa ser. Alatae in Peru was re¬
cently provided together with an overview of its
morphology and ecology (Weigend, 2000a). Nasa
ser. Alatae was originally described as Loasa subg.
Loasa ser. Alatae Urban & Gilg (Urban & Gilg,
1000), but has since been removed to the new ge¬
nus Nasa and includes the type of this genus (Nasa
rubrastra (Weigend) Weigend). Nasa ser. Alatae is
closely allied to Nasa ser. Grandiflorae, and its pre¬
cise limits are currently under investigation (Wei¬
gend et al.. in prep.). Both groups share more or
less campanulate corollas usually with yellow, red,
or orange petals and floral scales with distinct api¬
cal wings.
The series as currently defined comprises ca. 27
species, which range throughout the northern and
central Andes (southern limit in south-central Peru
with an outlier in northern Bolivia) and are usually
restricted to the moister eastern slopes in the drier
central Andes. The geographical range has been
recently expanded northward by the description of
a new species from Central America (N. panamen-
sis Weigend, Weigend, 2002b). The species here
described reported from Peru bring the species to¬
tal of Nasa ser. Alatae in the strictest sense (i.e.,
sensu Urban & Gilg. 1900) up to 17 from 13 in the
year 2000, and an original number of only 5 spe¬
cies before the beginning of the current revisionary
effort. I'he species totals known for Colombia and
Ecuador are 7 and 8. respectively, so that Peru now
houses more than twice as many species as each of
these other two countries. This has nothing to do
with the different sizes of the countries, since Nasa
ser. Alatae in Peru is largely restricted to the north¬
ern part and there mostly to the eastern cordilleras,
i.e., an area substantially smaller than their range
in Ecuador and Colombia.
MATERIALS AND METHODS
New collections were obtained during field stud¬
ies in Peru in October 2000 and March— April 2001
in addition to the collections cited in Weigend
(2000a); vouchers are cited under the respective
species below. Pressed specimens, FA A-material.
and photographs are available from these collec¬
tions. Duplicates have been sent by Asuncion Cano
E. from USM, Segundo Leiva G. (Herbario de La
Universidad Antenor Orrega, Trujillo, Peru), and
Victor Quipuscoa S. from HUSA (Universidad Na-
cional de San Augustin, Arequipa, Peru). Also, two
species, N. olmosiana (Killip) Weigend and N. pon¬
galamesa, have been taken into cultivation in Ber¬
lin.
Nasa si.it. Aiatae in the Amotapk-
Huancabamba Zone
The Amotape— Huancabamba Zone has been pos¬
tulated as a particular center of diversity for the
Loasaceae (Weigend, 2002a), and this fact is fur¬
ther underscored by the four new species and the
new record. All five species here discussed are
found in the area called the Amotape— Huancabam¬
ba Zone (Fig. I, Ayers, 1999; Berry, 1982; Young
Beynel, 1997) and close apparent distribution
gaps ol Nasa ser. Alatae (Weigend. 2000a). The
species show the narrow endemieity and morpho-
Novon 14: 134-146. 2004.
Volume 14, Number 1
2004
Weigend
Nasa ser. Alatae from Peru
135
Figure 1. Distribution of the newly described and the newly reported species of Nasa ser. Alulae, bold circles: Nasa
amaluzensis (Weigend) Weigend; black stars: Nasa lambayequensis ; empty star: Nasa stolonifera ; inverted black trian¬
gles: Nasa victorii ; inverted empty triangle: Nasa pongalamesa (map modified from Weigend, 2002a).
logical divergence that is so typical of many plant
species in this geographical region. A total of 14 of
the 17 species of Nasa ser. Alatae known from Peru
are endemic to the Peruvian part of the Amotape—
Huancabamba Zone, another 3 species are found
in both the Ecuadorean and the Peruvian part of
the Amotape— Huancabamba Zone, and 1 species is
endemic to the Ecuadorean part of the Amotape—
Huancabamba Zone. This means that more than
50% of all the known species of Nasa ser. Alatae
are restricted to that relatively small zone, in spite
of the fact that the group is widespread and ranges
from central Peru to Panama.
Nasa ser. Alatae comprises mostly narrowly en¬
demic species, and all the newly described taxa fall
into that category. In some cases apparent narrow
endemicity may reflect a lack of collecting, espe¬
cially in the poorly known cloud forests on the east¬
ern slopes of the Andes. This is borne out by the
fact that some species previously known from oidy
one locality have now been recollected at different
places (e.g., N. lenta (Urban & Gilg) Weigend).
Nasa ser. Alatae are usually plants from cloud and
mistbelt forests, with only very few species ranging
either into paramo habitats (e.g., N. loxensis
(Kunth) Weigend from Ecuador, the new species N.
136
Novon
Figure 2. I\asa pongalamesa Weigend. — A. Young plant showing typical decussate phyllotaxy of series A/otae. — B.
Flower, lateral view, showing dense cover with black stinging setae. — C. Flower, male phase, showing numerous stamens
in the center of the flower. —1). Ovary with dense cover with black stinging setae. — E. Stinging hairs on back of
petal. — F. Stamens and Moral scales in old flower (lemale phase), showing the funnel-shaped scale apices giving access
to the nectar. — O. Floral scale, full dorsal view with basal sacs, scale back, and apical wings. — H. Base of floral
scale showing dorsal sacs with their oblique folds. — I. Apical part of floral scale showing recurved scale neck and
apical wings. — K. Papillose staminodes alter removal of floral scale. — L. Densely setose ovary roof (androeeium,
scales, petals, and two sepals removed) surrounded by the white annular nectary. — M. Receptacle as in L, lateral view,
showing the elevated ovary roof with black stinging setae, the thick, white annular nectary and the base of the style
(Weigend et al. 852C, type material cultivated at Berlin).
Volume 14, Number 1
2004
Weigend
Nasa ser. Alatae from Peru
137
pongalamesa ) and into dry scree-slope habitats (N.
u re nl ivel utina We i gen d) .
Floral Morphology
The basic structure and terminology of the floral
elements of Nasa ser. Alatae have been discussed
previously (Weigend, 2000a) and are largely con¬
gruent with the morphology observed in Nasa ser.
Grandiflorae. The finer detail, especially petal
shape, color, and orientation, size and shape of the
scale back, presence or absence of dorsal threads,
and die orientation and shape of the apical wings
are crucial for taxon delimitation. A detailed over¬
view is here provided on the basis of photographs
of cultivated Nasa pongalamesa (Fig. 2). The pen-
tamerous flowers have a largely inferior ovary, five
calyx lobes, and five membranaceous, shallowly
cymbiform petals (Fig. 2B, 2C). The petals are usu¬
ally setose on the back (f ig. 2C, 2K), but stinging
hairs are usually densest on the ovary (Fig. 2D).
Phe flowers are proterandric and the mature anthers
are folded into the center of the flower during the
male phase; during the female phase the stigma
elongates and is then situated in the center of the
flower, while the anthers have by then wilted (Fig.
2F). Idle floral scales seen from above (i.e., in the
flower visitor’s view) form a typical “revolver” (Fig.
2F), i.e., each scale apex forms a funnel (Fig. 2H.
21) that guides the beak of visiting hummingbirds
to the basal nectar reservoir (Fig. 211).
Pile term revolver flower is generally used for
flowers where there is more than one access point
to the nectariferous part (Endress, 1994: 116). In
Nasa ser. Alatae (and most Loasaceae sublam. Loa-
soideae) there are five nectaries, each covered by
a floral scale and two staminodes, and thus each
floral scale has to be probed to extract all the nectar
from an individual flower. Each floral scale forms a
distally widened funnel providing mechanical guid¬
ance to the nectaries and also ensuring direct con¬
tact of the beak or the head of the visiting hum¬
mingbird with either the anthers or the stigma. This
is a very peculiar case of a revolver flower, and I
would like to introduce the term funnel-revolver
(lower for this particular situation. Phe basal nectar
sacs are not the place where the nectar is secreted;
they only function as nectar storage containers.
Nectar secretion takes place in five basal nectaries
(f ig. 2F. 2M), and nectar is secreted between the
bases of the two free staminodes (Fig. 2K) into the
base of the nectar scale. The ovary roof, i.e., the
receptacle within the nectary, is densely covered
with stinging hairs (Fig. 2M), presumably to deter
nectar theft by small insects, which could otherwise
land on the nectar scales, climb onto the recepta¬
cle, and then remove the nectar from the scales
without touching either anthers or stigmas.
Key to the Taxa of Nasa ser. Alatae in Peru
1. Calyx tube and fruit without stinging hairs
. Nasa anderssonii
1'. Calyx tube and fruit usually densely covered
with stinging hairs, at least in proximal por¬
tion . 2
2(1). Stem with strongly elevated longitudinal
ridges (usually 6); stinging hairs on ovary
concentrated at base, distal portion with very
few, or without stinging hairs .... Nasa victorii
2'. Stem terete, never with elevated ridges;
stinging hairs ± evenly distributed on fruit,
usually very numerous . 3
3(2). Leaves pentagonous with acute leaf lobes;
stem densely covered with uniseriate glan¬
dular hairs, especially in distal portion, setae
4—5 mm long; petals narrowly oblong and se¬
tose on hack . Nasa amaluzensis
3'. Leaves ovate to widely ovate, il widely ovate
then leaf lobes always rounded to acuminate;
stem without or w ith few uniseriate glandular
hairs, setae typically less than 4 mm long;
lamina typically ovate, if subcircular in out¬
line, then petals widely ovate to subcircular
and esetulose on hack . 4
4(3). Corolla yellow, leaves dark green with white
lines along primary and secondary veins
adaxially . N. driesslei
4'. Corolla pink, orange, or red, never yellow,
leaves without white line along veins or, it
with white veins, then lamina bright green
and flowers bright red (only N. tingamarien-
sis , occasionally) . 5
5(4). Inflorescence with large, sessile, semi-am-
plexieaul bracts; petals very deeply cymbi¬
form and bright red; corolla balloon-shaped,
rarely half spreading . N. olmosiana
5'. Inflorescence with petiolate or sessile bracts,
these not semi-amplexicaul; petals variable
in shape and color, often orange or pink; co¬
rolla campanulate or star-shaped, never bal¬
loon-shaped . 6
6(5). Lamina very densely pubescent, velvety to
the touch with numerous stinging hairs be¬
tween the trichomes; vegetative shoots with
numerous, mostly spirally inserted leaves
(rarely opposite); stiffly erect, sparsely
branched shrub; floral scales yellow' with or¬
ange nectar sacs and apically with filaments
. N. urentivelutina
6'. Lamina hairy and sometimes densely so, hut
never velvety to the touch; leaves never spi¬
rally inserted, opposite, internodes > 3 cm;
plant often branched from base; floral scales
red or yellow, nectar sacs not contrastingly
colored . 7
7(6). (.eaves approximately as long as wide, dark
green callus below' insertion of petiole pre¬
sent; petals either very narrow (5—6 X as
long as wide) or widely ovate, carnose and
completely esetulose on back;; capsule cla-
138
Novon
vate, erect on post-anthetieally elongating
pedicel . 8
7'. Dark green callus below insertion of petiole
absent; petals not narrow (< 4 X as long as
w ide); capsule variable in shape, but pedicel
never elongating post-anthetieally . 10
8(7). Petals ovate (< 4 X as long as wide), carnose
. !\as 6 X as long as wide),
membranaceous . 9
9(8). Leaves ovate, up to 90 X 50 mm; found in
Piura . Nasa solata
9'. Leaves widely ovate to subcircular, 90—150
mm long and wide; Cajamarca . . . Nasa dillonii
10(7). Corolla brilliant red. inside of petals densely
glandular distally; soft-stemmed rain forest
herb with widely ovate leaves 100-180 mm
long and w ide, found below (2000— )1000 m
. N. tingomariensis
10'. Characters not as above, leaves smaller,
found only above 2200 m . II
11(10). Petals half-spreading and corolla star¬
shaped; floral scales white or pale yellow 12
11'. Petals erect and corolla campanulate; floral
scales white, pale yellow, orange, or deep
pink . 13
12(1 1). Sparsely branched, stiffly erect herb with few
to many yellow to red setae on stem and leaf
veins; petals pink, half-spreading; Cajamar¬
ca: Bambamarca & Cutervo . Nasa carnea
12'. Much-branched stoloniferous herb or sub¬
shrub with numerous yellow to red setae on
stem and leaf veins; petals orange; La Lib-
ertad: Bolfvar . Nasa stolonifera
13(12). Erect shrubs; capsule turbinate to clavate,
2—2.5 X as long as wide; Cajamarca, Lam-
bayeque, Piura . 14
13'. Erect herbs or scandent shrubs; capsule glo¬
bose or shortly turbinate, 1—1.5 X as long as
wide; Amazonas, Pasco, Huanueo, San Mar¬
tin . 15
14(13). leaves widely ovate, ca. 1.2— 1.5 X as long
as wide with 2 to 3 leaf lobes on each side;
stem and pedicels densely glandular; petals
orange; Lambayeque: Ferrefiafe .
. Nasa lambayeque nsis
14'. Leaves ovate, ca. 2 X as long as wide with
5 to 7 lobes on each side; stem and pedicels
sparsely glandular; petals pink; Cajamarca:
San Miguel . Nasa pongalamesa
15(13). Erect herb with soft stems (collapsing upon
drying); sepals 10—13 X 4.5—7 mm; petals
30-37 mm long, narrowly ovate, apical ly
acuminate, deeply cymbiform, membrana¬
ceous; plant very densely covered with setae;
Pasco, Junin . N. pascoensis
15'. Subscandent shrub with lignescent stems
(not collapsing upon drying); sepals 8—10 X
2-4- mm; petals < 25 mm long, oblong, api¬
cal ly rounded, shallowly cymbiform to ± pla¬
nar; plants sparsely setose to nearly esetu-
lose . 16
16(15). Petals widely ovate, dorsally with numerous
red setae, membranaceous; Amazonas . .
. N. nubicolorum
16'. Petals oblong, apically rounded, without se¬
tae, carnose; Huanueo, San Martin ... A. tenia
1 . Nasa lambayequensis Weigend, sp. nov. TYPE:
Peru. Dept. Lambeyeque: Province Ferrenafe,
Incahuasi, to Laguna Tenibladera, 3000 m, 13
Sep. 1985, 1. Sagdstegui A., I). Skill man,
Mostacero & L Ramirez 12872 (holotype, HUT;
isotypes, F, MO). Figures 1, 3.
Haec species N. pongalamesae affinis, sed ab ea foliis
late ovatis ambobus lateribus lobis 2 vel 3 instructis (nee
anguste ovatis ambobus lateribus lobis 5—7 instructis), co¬
rolla aurantiaca (nec rosea) atque caule apicem versus
dense (nec parce) glanduloso differ!.
Strictly erect shrub up to 1.5 in tall; stein Idled
with white pith, to 0.5 cm diatn., densely covered
with red-brown setae to 3 mm long and scabrid
hairs, young parts of the plant and pedicels very
densely covered with uniseriate, gland-tipped,
white trichomes consisting of 2 to 5 cells each.
Leaves opposite, petioles 30—45 mm long, densely
setose; lamina ovate, 50—75 X 50-65 mm, mem¬
branaceous, base shallowly cordate (sinus 4—6 mm
deep), apex acuminate, margin lobed with 2 to 3
widely triangular lobes on each side, each 10—15
X 17-20 mm, lobe margin serrate; abaxial surface
setose on veins, otherwise covered with scabrid-glo-
chidiate hairs (0.7-1 mm), adaxial surface densely
setose and densely covered with longer scabrid
hairs (ca. 1 mm); venation pinnate with 2 to 3 lat¬
eral veins on each side. Inflorescence a terminal
monochasium, rarely diehasium, 10-20 cm long,
with 5 to 8 pendent flowers, bracts subsessile, up
to 30 X 12 mm; pedicel 20—40 mm long. Calyx
very densely setose, tube conical to ovoidal, 5—6 X
5 nun, calyx lobes 5, ovate-acuminate, 12-14 X 5
mm, densely setose and covered with scabrid tri¬
chomes on the outside; corolla widely campanulate;
petals obovate, cymbiform, 18—22 X 10 mm, apex
rounded to slightly acuminate, abruptly widening
ca. 5 mm from base, densely setose and set with
scabrid and glochidiate hairs on back, with gland-
tipped, 3-celled uniseriate trichomes, orange; nec¬
tar scales 5, with rectangular back, 8X2 mm, base
incurved, basally on back with two depressed glo¬
bose sacs 2 mm diam.. pale red. scale neck slightly
thickened, recurved and deeply red, without lili-
form appendages, laterally protracted into 2 in¬
curved, suberect wings 4 X 1.5 mm; staminodia 2
per scale, 7 mm long, base dilated, 1 mm wide,
filiform above, base curved, papillose, pale yellow;
stamens numerous, in 5 epipetalous fascicles of 12
to 14 each; filaments 12 mm long, white, anthers 1
X 0.5 mm, black. Capsule clavate, crowned with
the persistent calyx lobes, pedicel erect, 25—35 mm
long, capsule 20—25 X 10 nun, very densely red
setose, covered with very short, white, glochidiate
Volume 14, Number 1
2004
Weigend
Nasa ser. Alatae from Peru
139
Figure 3. Nasa lambayequensis Weigend. — A. Leaf. — B. Capsule. — C. Sepal. — D. Petal. — E. Floral scale. — E
Staminode, lateral view. (Drawn from Sagdstegui el al. 12872, HUT.)
and scabrid trichomes and uniseriate, gland-tipped
trichomes, opening with 3 apical valves; seeds nu¬
merous, ovoid, testa reticulate.
Nasa lambayequensis lias been collected in flow¬
er and fruit in August and September, but is likely
to show peak flowering toward the end of the rainy
season in late April and May; no detailed field data
are so far available. Nasa lambayequensis is evi¬
dently closely allied to N. pongalamesa described
below. They share the shrubby, poorly branched
habit and grow in similar situations along rocks and
dry stone walls in jalca (dry paramo of Peru) hab¬
itats. It differs from N. pongalamesa and closely
allied N. carnea in leaf shape (widely ovate with 2
to 3 lateral lobes vs. ovate blades with more than
4 lateral lobes), petal color (orange vs. pink), and
its much denser glandular indument. Both N. pon¬
galamesa and N. lambayequensis are segregates of
N. carnea (Urban & Gilg) Weigend s.l. (sensu Wei¬
gend, 2000a), and the material here referred to the
new species was previously treated under that tax¬
on. The three taxa are allopatric, with both new
species found on the western slope of the Andes,
while N. carnea is restricted to inner Ande< in val¬
leys in the Central Cordillera (centered around
Dept. Cajamarca, Prov. Bambamarca). The northern
specimens of what now remains in N. carnea (e.g..
Dept. Cajamarca, Prov. Cutervo) remain problem¬
atical, and field studies will be required to clarify
where they actually belong, since the available her¬
barium material is unsatisfactory.
Paratypes. PERU. Dept. Lambeyeque: Prov. Eer-
renafe, between Incahuasi and Sinchihual, 3000 m, 25
Aug. 2000. A. Sagdstegui A. & M. Zapata 18279 (HAO,
HUT. M).
140
Novon
2. INasa pongalamesa Weigend, s[). nov. TYPE:
Peru. Dept. Cajamarca: Prov. San Miguel de
Pallaques, Road Agua Blanea to Oyotun, pass
height “Ponga la Mesa," 3500—3600 m, 14
Oct. 2000 (cultivated in Berlin since Sep.
2002), M. Weigend, //. Fiirlher, N. Dostert &
E. Rodriguez R. 2000/752C (holotype, USM;
isotypes. BM. BSB, HUT, M. MO. NY. W). Fig¬
ures 1, 2, 4.
Haec species l\. carneae affinis, sed ah ea hal)itu fru-
ticis rigide erectis (nec herbae arete ramosae), squamis
nectariferis roseis (nec luteolis) et corolla campanulata
(nec stellata) dilfert.
Strictly erect shrub 1-2 m tall; stem filled with
white pith, to 1.5 cm diam., densely covered with
reddish brownish setae to 3 mm long and scabrid
hairs, young parts ol the plant and pedicels with
scattered to many uniseriate, gland-tipped, white
triehomes consisting ol 2 to 5 cells each. Leaves
opposite, petioles 30—55 mm long, densely setose;
lamina ovate, 100-170 X 50—70 mm, membrana¬
ceous, base shallowly cordate (sinus 3-5 mm deep),
apex acuminate, margin lobed with 5 to 7 widely
triangular lobes on each side, each 7—12 X 15—18
mm, lobe margin serrate; abaxial surface very
densely setose on veins and covered with glochid-
iate hairs (0.5 mm), adaxial surface sparsely setose
and densely covered with longer scabrid hairs (ca.
I mm); venation pinnate with 3 to 5 lateral veins
on each side. Inflorescence a terminal monoehasi-
urn, rarely dichasium, 20—50 cm long, with 8 to 12
pendent flowers, bracts petiolate, up to 55 X 18
mm; pedicel ca. 20 mm long, calyx very densely
setose, tube conical to ovoidal, 6—7 X 4 mm, calyx
lobes 5, ovate-acuminate, 13—15 X 5 mm, densely
setose and covered with scabrid triehomes on the
outside; corolla widely campanulate; petals ob-
ovate, cymbiform, 20—25 X 15 mm, apex rounded
to slightly acuminate, abruptly widening ca. 2 mm
from base, setose and set with scabrid anti glochid-
iate hairs on back, with gland-tipped, 3-celled un¬
iseriate triehomes, pale pink; nectar scales 5, with
rectangular back, 6X2 mm, base incurved, basally
on back with two depressedly globose sacs 2 mm
diam., pale red, scale neck slightly thickened, re¬
curved and deeply red, without filiform appendag¬
es, laterally protracted into two incurved, suberect
wings 3 X 2 mm wide; staminodia 2 per scale, 7
mm long, base dilated, I mm wide, filiform above,
base curved, papillose, pale pink; stamens numer¬
ous, in 5 epipetalous fascicles of 12 to 14 each;
filaments 12 mm long, white, anthers 1 X 0.5 mm,
black. Capsule clavale, crowned with the persistent
calyx lobes, pedicel erect, 25—35 mm long, capsule
25—30 X 10 mm, very densely black setose and
covered with very short, white, gloehidiate tri-
chomes, opening with three apical valves. Seeds
numerous, ovoid, testa reticulate.
Nasa pongalamesa was encountered in fruit in
October 2000, i.e., in the middle of the dry season.
Like most species of Nasa in the region, it likely
flowers March to May at the end of the rainy season.
Nasa pongalamesa is closely allied to three other
species, N. carnea (Urban & (dig) Weigend, N.
urentivelutina Weigend (Weigend, 2000), and N.
lamhayequensis (see above). It differs from N. cor¬
nea, e.g., by the stiffly erect habit and perennial
growth form with a strongly lignified stem, a very
dense cover with dark red-brown stinging hairs (vs.
stinging hairs scattered, pale brown), and the pres¬
ence of stinging hairs on the back of the petals.
Nasa urentivelutina differs in apricot-colored petals
(vs. pink), densely spiral phyllotaxis (vs. opposite
leaves), and a much more strongly branched habit
(shrubs may be as large as 2 X 1.5 m). Its floral
scales have a back that is very strongly narrowed
above the nectar sacs and ends in three apical fi¬
liform appendages, whereas the scales of N. pon¬
galamesa are only slightly narrowed above the sacs
and have no filiform appendages.
The new species is also ecologically clearly dif¬
ferentiated: It grows on an exposed ridge with
strong cloud condensation at an elevation of ca.
3500 m between rocks in otherwise grassy puna
vegetation. Nasa carnea, on the other hand, grows
in shrub forest in inner Andean valleys at ca.
2200—3000 m, and N. urentivelutina is restricted to
dry scree slopes in full sun at elevations below
3000 m. Nasa pongalamesa is thus the species of
Nasa ser. Alatae that reaches the highest elevation
so far known from this group and is found together
with a species of the N. cymbopetala group, the
typical puna group in the genus.
Nasa pongalamesa was found in fruit only in Oc¬
tober 2000 and was then brought into cultivation
in the greenhouses of the Institute of Biology in
Berlin. The type specimens were prepared from the
cultivated material when it came into flower. Nasa
pongalamesa is named after the locality of the orig¬
inal collection, “Ponga la Mesa” (lay the table). The
epithet is here used as a noun in apposition, with
its spelling not changed, sensu Article 23.5. of the
ICBN (Greuter et ah, 2000).
Paratypes. PERU. Dept. Cajamarca: Prov. San Mi¬
guel de Pallaques, road Agua Blanca to Oyotun, pass
height “Ponga la Mesa,” 3500— 36(H) in. 14 Oct. 2000. M.
Weigend et ea ovario
liasi trichomatibus urentibus instructo (nec omnino tri-
chomatibus urentibus carente) et habitu fruticis scanden-
lis (nec herbae rigide erectae), ab omnibus speciebus cet¬
eris Loasacearum caule costis longitudinalibus elevatis
instructo differt.
Erect to ascending shrub 0.6— 2 m tall: stem te¬
rete with typically 6 elevated longitudinal ridges,
with white pith, to 1.5 cm diam., with very few
reddish brown setae 1—3 mm long and densely cov¬
ered with minute scabrid hairs < 0.2 mm long,
eglandular. Leaves opposite, petioles 40—65 mm
long, very sparsely setose; lamina widely ovate,
100-150 X 60—90 mm, membranaceous, base sub-
truncate (sinus < 2 mm deep), apex long acumi¬
nate. margin lobed with 5 to 7 indistinct, triangular
lobes on each side, each up to 5 X 15 mm, lobe
margin serrate to serrate-denticulate; abaxial sur¬
face with scattered setae on veins and covered with
numerous scabrid hairs (< 0.2 mm), adaxial sur-
Volume 14, Number 1
2004
Weigend
Nasa ser. Alatae from Peru
143
Figure 5. Nasa stolonifera Weigend. — A. Habit. — B. Sepal. — C. Petal. — I). E. Moral scale, dorsal and lateral view.
— F. Staminode, lateral view. (Drawn from Weigend el al. 2000/822, BSB.)
144
Novon
Figure 6. A hisa victorii Weigend. — A. Habit. — lb Single flower. — C. Sepal. — I). Petal. — E. Floral scale, lateral
view. — F. Staminode, dorsal view. (Drawn from Quipuscoa cl al. 2480, HSR.)
face very sparsely setose on veins and densely cov¬
ered with short seabrid hairs (< 0.3 mm); venation
pinnate with 3 to 5 lateral veins on each side. In¬
florescence a terminal monochasium 1.3—20 cm
long, with up to 6 pendent flowers, bracts petiolate,
lamina ovate, up to 55 X 20 mm; pedicel 20-40
mm long; calyx setose only at the base, tithe sub-
globose, 5 X 5 nun, calyx lobes 5, narrowly trian¬
gular-ovate, ea. 10 X 3^1 mm, densely setose and
densely covered with short seabrid trichomes on the
outside; corolla widely campanulate; petals ob-
ovate, shallowly cymbiform, 15 X 11 mm, apex
Volume 14, Number 1
2004
Weigend
Nasa ser. Alatae from Peru
145
rounded, without triangular teelli, esetulose and
densely set with short scabrid hairs on hack, eglan-
dular, orange-red; nectar scales 5, with rectangular
back, 8X3 mm, base incurved, basally on back
with 2 indistinct, depressedly globose sacs 3 mm
diam., yellow, scale neck not thickened, without ap¬
pendages. laterally protracted into 2 incurved, hor¬
izontal wings 3 X 2 mm; staminodia 2 per scale.
13 mm long, base dilated, 1 mm wide, filiform
above, base and tip curved, papillose, white; sta¬
mens numerous, in 5 epipetalous fascicles of 12 to
14 each; filaments 12—15 mm long, white, anthers
2 X 1.5 mm, black. Capsule subglobose, ca. 20
nun diam.. with setae only in proximal part and
covered with numerous very short, white, scabrid
trichomes, opening with 3 apical valves. Seeds nu¬
merous. ovoidal. testa reticulate.
Nasa victorii probably flowers and fruits sporad¬
ically throughout the year, since its habitat does not
experience a marked dry season. It has been col¬
lected with flower buds in September and with flow¬
ers and fruits in September.
While most species of Nasa are readily recog¬
nized, N. victorii is certainly one of the most readily
identifiable taxa. Its stems with longitudinal ridges
are unique not only in the species group, but in the
entire family Loasaceae. The species is a subscan-
dent to seandent shrub in dense, wet cloud forest on
the eastern slope of the Andes, a notoriously poorly
known region. Like N. nubicolorum from the same
general region (Weigend, 2000a) it grows in deep
leaf litter and humus in extremely wet habitats. The
species is now known from two collections in the
department San Martin, but may be more widespread
in this inaccessible region. Like most species of the
group it is likely to be sensitive to habitat alteration
and therefore vanishes rapidly from areas as they
become accessible to civilization.
Its sparse indument is reminiscent of Nasa an-
derssonii, which is known from the Cordillera Golan
in Peru and also neighboring Ecuador, but it differs
from that taxon, e.g., in stem and (lower morphology,
and habit. Nasa victorii is named in honor of Victor
Quipuseoa Silvestre (LIUS A. Arequipa. Peru).
Paratypes. PERU. Dept. San Martin: Prov. Rioja,
road from Pedro Ruiz to Rioja, between La Ksperanza and
Nuevo Cajamarca, near km 377, 2000—2300 m,
05°42'09"S, 077°48'27"W, 21 Oct. 2000. M. Weigend el
al. 2000/929 (BSR, HUT, M, USM).
Nkw Rkcoko
5. Nasa amaluzensis (Weigend) Weigend, Mon-
ogr. Syst. Bot. Missouri Bot. Card. 75: 954.
1999. Loasa amaluzensis Weigend, Bot. Jahrb.
Syst. 118: 264. 1996. TYPE: Ecuador. Loja:
26.4 km S of Amaluza on road to Zumba, river
valley, 2800 m. S. Clemants 2392 (holotype,
QGA; isotype, QCNE). Figure I (see also Wei¬
gend. 2000b: fig. I6J-R).
Nasa amaluzensis apparently flowers during the
wet season (January to April) and probably fruits
from April onward, like most species of Nasa from
the western part of the Andes.
Nasa amaluzensis was described from a single
collection from the extreme south of Ecuador in
1996. Material exactly matching the type has now
been collected by S. Leiva G. in the extreme north
of Peru, so that this species is now known from both
countries. Nasa amaluzensis shows clear affinity to
Nasa loxensis from Ecuador (it shares horizontally
indexed wings on its nectar scales, and cymbiform,
setose petals), but differs in distally densely glan¬
dular stems and pentagonous leaves with acute leaf
lobes. Its closest geographical neighbors are Nasa
solata (leaves ovate, petals linear), N. sagasteguii
(similar leaf shape but with ovate leaf lobes, car-
nose, esetulose petals), and N. glabra (plant virtu¬
ally esetulose, carnose, esetulose petals).
Material examined. ECUADOR. Prov. Loja: 26.4
km S of Amaluza on road to Zumba, river valley, 2800 m,
,S. Clemants 2392 (QGA, QCNE). PERU. Dept. Piura:
Prov. Ayavaca, Cerro Aypate, 2650 m, 10 Jan. 2002, .S’.
h>iva G. 2632 (BSB, F, HAD. HUT).
Acknowledgments. I express my sincere grati¬
tude to Dominik Kneer (Berlin, Germany) for the
preparation of the drawings and to Katja Weigend
(Berlin) for the preparation of the photographic
plate. I also thank E. Rodriguez R. ( 1 1 LIT). S. Leiva
G. (HAO), Victor Quipuseoa S. (HUSA), and A.
Cano E. (USM) for providing valuable collections,
and N. Dostert (Berlin) and H. Forther (both Mu¬
nich) for their assistance during fieldwork. I thank
the curators and directors of the following herbaria
lor access to and loans of specimens: E, HAO, HUT,
M, MO, USM. Part of the studies was supported by
the Deutscher Akademiseher Austauschdienst, tin*
Deutsche Eorschungsgemeinschaft, and boteon-
sult GmbH (Berlin), which are here gratefully ac¬
knowledged.
I .iterature Cited
Ayers, T. 1999. Biogeography of Lysipomia (Campanula-
ceae), a high elevation endemic: An illustration of spe¬
cies richness at the Huancabamba Depression, Peru.
Arnaldoa 6: 13—28.
Berry, P. E. 1982. The systematics and evolution of Fuch¬
sia sect. Fuchsia (Onagraceae). Ann. Missouri Bot.
Card. 69: 1-198.
Endress, P. k. 1996. Diversity and Evolutionary Biology
146
Novon
of Tropical Flowers, 2nd ed. Cambridge Univ. Press,
Cambridge, U.k.
Greuter, W., J. McNeill. F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). H. Nicolson, P. C. Silva,
J. F. Skog, P. Trehane, N. J. Turland & 1). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Cotie). Regnum Veg. 1 38.
Urban. I. & W. Gilg. 1000. Monographia I .oasacearum.
Nova Acta Caes. Leop. -Carol. German. Nat. Cur. 76(1):
1-368.
Weigend, M. 1990. Loasaeeae. In P. M. Jprgensen & S.
Le6n-Ydnez (editors). Catalogue of the Vascular Plants
of Ecuador. Monogr. Syst. Bot. Missouri Bot. Card. 75:
537-539, 954-955.
- . 2000a. A revision of the Peruvian species of
AV«a scr. Alulae (Loasaeeae). Nordic J. Bot. 20: 15—32.
- . 2000b. 132. Loasaeeae. In G. Hailing cV L. An-
dersson. Flora of Ecuador 64.
- . 2002a. Observations on the biogeography of the
Amotape— Huancabamba Zone in northern Peru. In: k.
Young et al.. Plant Evolution and Endemism in Andean
South America. Bot. Rev. (Lancaster) 68: 38—54.
- . 2002b. Nasa panamemis, a new species of Nasa
(Loasaeeae) from Central America. Bot. Jahrb. Syst.
124: 21 1-216.
Young, K. R. & C. Reynel. 1997. Huancabamba Region,
Peru and Ecuador. Pp. 465—469 in S. I). Davis, V. II.
Heywood, 0. Herrera-MacBryde, J. Villa-Lobos & A. C.
Hamilton (editors). Centers of Plant Diversity, a Guide
and Strategy for their Conservation, Vol. 3. The Amer¬
icas. 1UCN Publications Unit, Cambridge.
Aconitum ouvrardianum var. acutiusculum (Ranunculaceae), a New
Combination from Yunnan, 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. Close examination of the type speci¬
mens of Aconitum acutiusculum and of A. acutius¬
culum var. aureopilosum has shown that their inflo¬
rescence rachis and pedicels are all yellowish
spreading pubescent, and thus the two taxa show
no essential difference from each other. They are
merged herein. A new combination, A. ouvrardian¬
um var. acutiusculum (Fletcher & Lauener) Q. E.
Yang & Y. Luo, is proposed, considering that A.
acutiusculum is most similar to A. ouvrardianum in
having leaves 3-sect with ultimate lobules linear or
lanceolate-linear, inflorescence rachis and pedicels
yellowish spreading puberulent, and the upper se¬
pal navicular-galeate, yet distinguishable by the
acute tip of the lateral sepals; they are sympatric
in geographical distribution, both occurring in De¬
qen County, northwestern Yunnan.
Key words: Aconitum, China, Ranunculaceae,
Yunnan.
Fletcher and Lauener (1950: 198) described ,4c-
onitum acutiusculum based on Forrest 14696 from
Raima Shan, Deqen County, northwestern Yunnan,
and stated that this species is clearly distinguished
from all others in the genus by the acute tip of the
lateral sepals, an otherwise unimportant feature in
the classification of Aconitum L. because the lateral
sepals in this genus are generally subrounded in
outline. In the original Latin description, the upper
part of the stem and the inflorescence are men¬
tioned as densely and extremely minutely white
crispy pubescent (Caulis . . . praesertim superne
cum inflorescentia dense et minutissime albo-cris-
po-pubescens). Wang (1965) described a new va¬
riety, A. acutiusculum var. aureopilosum, from
Yangtsa, Deqen County, and stated that this variety
is distinguished from the typical one by having the
inflorescence yellowish spreading puberulent.
When describing this new variety, he had no op¬
portunity to examine the type specimens of A. acu¬
tiusculum and thus had to base his judgment on
Fletcher and Lauener’s description (Wang, pers.
comm.). We have checked the holotype and isotype
of A. acutiusculum var. aureopilosum at PE and
found that the inflorescence rachis and pedicels are
indeed densely yellowish spreading puberulent.
Pedicel pubescence is one of the most important
diagnostic characters in the classification of the ge¬
nus Aconitum.
Close examination of the type material of Aco¬
nitum acutiusculum, Forrest 14696, at BM, E, and
K has shown that its inflorescence and pedicels,
like those of A. acutiusculum var. aureopilosum , are
densely yellowish spreading puberulent and that
other important characters, such as the division of
leaves and the shape of the upper sepal, are by no
means different from those of A. acutiusculum var.
aureopilosum. Their leaves are 3-sect with ultimate
lobules linear or lanceolate-linear, and the upper
sepal is navicular-galeate. As this variety shows no
essential difference from A. acutiusculum in all the
characters, they should be merged.
Aconitum ouvrardianum Handel-Mazzetti (1931)
is a common species in Deqen, northwestern Yun¬
nan. We have not seen its type. Handel-Mazzetti
9702. but fortunately this species is quite richly
represented in the two major Chinese herbaria, PE
and KUN, and its identity has long been deter¬
mined with certainty. This species shows no essen¬
tial difference from A. acutiusculum in all the char¬
acters except in the shape of the lateral sepals, with
those in A. acutiusculum being acute at the apex
while those in A. ouvrardianum are subrounded. In
our herbarium observations we found this feature is
quite constant in these two entities, but this differ¬
ence is too tenuous to allow treating them as two
independent species. In fact, the type specimen of
A. acutiusculum var. aureopilosum at PE was once
referred to A. ouvrardianum by Chen and Fiu
(1941). When Handel-Mazzetti (1931 ) described A.
ouvrardianum, he had noticed the similarity of For¬
rest 14696 with his new species but also their dif¬
ferences, particularly in the shape of the lateral se¬
pals. Thus, he did not refer this specimen to any
species then known to him. Considering the con¬
sistent difference in the shape of the lateral sepals
Novon 14: 147-148. 2004.
148
No von
yet the great similarity in other characters between
A. ouvrardianum and A. acutiusculum, as well as
their sympatric distribution, it seems best to reduce
A. acutiusculum as a variety under A. ouvrardian¬
um. The following new combination is thus pro¬
posed.
Acoiiitum ouvrardianum Handel-Mazzetti var.
aciitiiiseulum (Fletcher & Lauener) Q. E.
Yang & Y. Luo, comb. nov. Basionym: Aconi¬
tum acutiusculum Fletcher & Lauener, Notes
Roy. Bot. Card. Edinburgh 20: 198. 1950.
TYPE: China. Yunnan: Deqen County, Baima
Shan, heavy pasture on the margins of pine
forests, 12,000 ft., Aug. 1917, G. Forrest
1469 6 (holotype, E; isotypes, K. BM).
Aconitum acutiusculum var. aureopilosum W. T. Wang,
Acta Phytotax. Sin., addit. 1: 85. 1965. Syn. nov.
TV PE: China. Yunnan: Deqen County, Yangtsa,
Snow Range, Rusala, mountain grassy slope, 4100
m, Aug. 1937, T. T. Yu 9671 (holotype, PE; isotype,
PE).
Aconitum ouvrardianum var. acutiusculum is
only known from Deqen County, northwestern Yun¬
nan, and seems quite uncommon. Although we had
made several botanical expeditions to this area, we
tailed to find it. The typical variety ouvrardianum ,
however, is very common in this area, growing in
alpine meadows or bushes at altitudes from 3000
to 4100 m.
Acknowledgments. We are grateful to the Keep¬
ers of BM, E, K, and PE for permission to examine
specimens or for the loan of material. This work
was supported by the National Natural Science
Foundation of China (grant no. 30070057) and a
Key Project of the Chinese Academy of Sciences
(KSCXZ-SW-101A).
Literature Cited
Chen, F. H. & Y. Liu. 1941. An enumeration of Aconitum
collected by T. T. Y u from Yunnan. Bull. Fan Mem. Inst.
Biol., Rot. ser. 11: 43-49.
Fletcher, H. R. & L. A. Lauener. 1950. Notes on some
Chinese species of Aconitum with an enumeration of the
Chinese species represented in the Edinburgh Herbar¬
ium. Notes Roy. Bot. Card. Edinburgh 20: 181-206.
Handel-Mazzetti, H. 1931. Aconitum. Pp. 282—297 in
Symbolae Sinieae, Vol. 7. Julius Springer, Vienna.
Wang, W. T. 1965. Notulae de Ranunculaceis sinensibus
II. Acta Phytotax. Sin., addit. 1: 49-103.
Miscellaneous Notes on Pteridophytes from China and
Neighboring Regions (IV) — Validation of Some Combinations in
Diplopterygium (Pteridophyta: Gleicheniaceae)
Xian-Chun Zhang
Institute of Botany, Chinese Academy of Sciences, Beijing 100093, People’s Republic of
China, xianchunzhang@yahoo.com.cn
Abstract. Some names of Diplopterygium widely
used in Chinese literature of pteridophytes are not
valid; therefore, formal combinations of five names
for Diplopterygium species from China are validat¬
ed. i.e., D. criticum (Ching & P. S. Chiu) Ching ex
X. C. Zhang, D. rufum (Ching) Ching ex X. C.
Zhang, D. rufopilosum (Ching & P. S. Chiu) Ching
ex X. C. Zhang, D. simulans (Ching) Ching ex X.
C. Zhang, and I). yunnanense (Ching) tilling ex X.
C. Zhang. Diplopterygium rufopilosum is lectotypi-
fied.
Key words: China, Diplopterygium, Gleicheni¬
aceae, Pteridophyta.
Diplopterygium (Diels) Nakai was traditionally
treated as a subgenus of Gleichenia Smith (Holttum,
1957, 1959; Tagawa & Iwatsuki. 1979; Tryon &
Tryon, 1982), but is now generally accepted as a
separate genus (Ching, 1978; Kramer, 1990; Wu &
Ching, 1991; 0stergaard Andersen & 011gaard,
2001 ). The Chinese species of Diplopterygium were
recorded under the generic name Hicriopteris C.
Presl (Ching, 1940; Ching et al., 1959). Holttum
(1959) discussed the reason why Hicriopteris was
misused in this sense. The type of Hicriopteris was
first described as a species of Dicranopteris Bern-
hardi, i.e., I). speciosa (C. Presl) Holttum. When
Presl (1851) described the genus he compared it
with Gleichenia glauca (Thunberg) Hooker, a spe¬
cies of Diplopterygium. Tater authors (Ching, 1940;
Copeland, 1947; Ching et ah, 1959) thought he was
describing a species of Gleichenia sect. Diplopter¬
ygium and consequently misapplied the generic
name Hicriopteris. Hicriopteris is properly a syno¬
nym of Dicranopteris (see Holttum, 1959; 0ster-
gaard Andersen & 011gaard, 2001).
Three genera of the fern family Gleicheniaceae
occur in China: Diplopterygium (Diels) Nakai,
Sticherus C. Presl, and Dicranopteris Bernhardi.
Plants of Diplopterygium can be easily distin¬
guished from those of Sticherus by their bipinnate
or pinnate-pinnatifid branches, and from Dicran¬
opteris by their dormant buds and rhizomes with
scales, veinlets of segments once forked, and spo¬
rangia (2)3 to 5(7) per sorus; in Dicranopteris the
dormant buds and rhizomes with hairs, veinlets of
segments 2 to 4 times forked, and sporangia ca. 8
to 25 per sorus (0stergaard Andersen & 011gaard.
2001) separate it from both Diplopterygium and
Sticherus.
Diplopterygium (Diels) Nakai, Bull. Nat. Sci.
Mus. Tokvo 29: 47. 1950. Gleichenia sect. Di¬
plopterygium Diels, in Engler N Plant I, Nat.
Pflanzenfam. 1(4): 353. 1900. Dicranopteris
sect. Diplopterygium (Diels) Underwood, Bull.
Torrey Bot. Club 34: 251. 1907. Gleichenia
subg. Diplopterygium Holttum. Reinwardtia 4:
261. 1957. TYPE: Diplopterygium glaucum
(Thunberg ex Houttuyn) Nakai (Polypodium
glaucum Thunberg ex Houttuyn).
Diplopterygium contains about 25 species, from
Asia to Queensland, Australia, and Polynesia (in¬
cluding Hawaii), mainly in tropical Asia, not in Af¬
rica. A single species, D. hancroftii (Hooker) A. R.
Smith, occurs in tropical America.
Ching et al. (1959) recorded 16 species of Di¬
plopterygium under the generic name Hicriopteris
from China and one from northern Vietnam, the
region bordering China. Six of these already have
been transferred into Diplopterygium, including:
Diplopterygium blotiunum (C. Christensen)
Nakai, Bull. Nat. Sci. Mus. Tokyo 29: 49.
1950.
Diplopterygium cantonense (Ching) Nakai.
Bull. Nat. Sci. Mus. Tokyo 29: 49. 1950.
Diplopterygium chinense (Rosenstock) De
Vol, in H. L. Li et al., El. Taiwan I: 92. 1975,
“ chine ns is.”
Diplopterygium giguiiteum (Aallich ex Hook¬
er) Nakai, Bull. Nat. Sci. Mus. Tokyo 29: 50.
1950.
Diplopterygium glaucum (Thunberg ex Hout-
Novon 14: 149-151. 2004.
150
Novon
luyn) Nakai, Bull. Nat. Sci. Mus. Tokyo 20: 51.
1950.
Diplnplcrygiimi Inrvissimimi (H. Christ) Na¬
kai. Bull. Nat. Sci. Mus. Tokyo 29: 52. 1950.
Chiu^ (in Shing, 1982) published numerous
combinations for Chinese species of Diploptery-
gium, but these were not validated properly ac¬
cording to Article 33.3 of the ICBN (Greuter et al.,
2000). Two of these were validated later, i.e., Di-
plopterygium maximum Ching & II. S. Rung (in
Rung, 1988: 134), and Diplopterygium glaucoides
(Ching) Ching ex P. S. Wang & X. Y. Wang (in Wang
& Wang, 2001: 279). Some of them are presently
regarded as synonyms of various species and there¬
fore need not be transferred into Diplopterygium.
The following five combinations for Diploptery¬
gium are formally proposed here in order to solve
the nomenclatural problems of these Chinese spe¬
cies and for the revision of fern flora of China.
Diplopterygium eriticuni (Ching & P. S. Chiu)
Ching ex X. C. Zhang, comb. nov. Basionym:
HicriopterLs critica Ching & P. S. Chiu, in S.
S. Chien X Chun, FI. Beipubl. Popul. Sin. 2:
348. 1959. Diplopterygium criticum (Ching &
P. S. Chiu) Ching, in R. II. Shing, Gloss. Term.
Nam. Kerns 50. 1982, nom. inval. TYPE: Chi¬
na. Yunnan: Pingbian, Dawei Shan, R. C.
Ching 55 (holotype, PE).
Diplopterygium rufuin (Ching) Ching ex X. C.
Zhang, comb. nov. Basionym: Hicriopteris rufa
Ching, in S. S. Chien & Chun, FI. Reipubl.
Popul. Sin. 2: 347. 1959. Diplopterygium ruf-
um (Ching) Ching, in R. II. Shing, Gloss.
Term. Nam. Ferns 50. 1982. nom. inval.
T\ PE: China. Yunnan: Pingbina, Dawei Shan,
K. M. Feng 4832 (holotype, PE).
The collection number was misprinted as 4823
when the species was published, but on the label
of the type in PE the number is 4832.
Diploplerygiimi riifopilosmn (Ching & P. S.
Chiu) Ching ex X. C. Zhang, comb. nov. Bas¬
ionym: Hicriopteris rufopilosa Ching & P. S.
Chiu, in S. S. Chien & Chun, FI. Reipubl. Po¬
ind. Sin. 2: 348. 1959, “ rufo-pilosa Diplop¬
terygium rufopilosum (Ching X P. S. Chiu)
Ching, in R. II. Shing, Gloss. Term. Nam.
Ferns 50. 1982, " refo-pilosum nom. inval.
TYPE: China. Yunnan: Dali. Cang Shan, C. Y.
Wu 1 1825 (lectotype, designated here, PE).
A second syntype, H. C. Wang 3541 (PE), is of
a young sterile plant, but C. Y. Wu 1 1825 is a fertile
plant; therefore, it is selected as the lectotype.
Diplopterygium siiniilans (Ching) Ching ex X. C.
Zhang, comb. nov. Basionym: Hicriopteris si-
mulans Ching, Acta Phvtotax. Sin. 8: 133,
101. 1959. Diplopterygium simulans (Ching)
Ching, in R. H. Shing, Gloss. Term. Nam.
Ferns 50. 1982, nom. inval. TYPE: China.
Hainan: C. Wang 35496 (holotype, PE).
Diplopterygium yunnanense (Ching) Ching ex
X. C. Zhang, comb. nov. Basionym: Hicriopteris
yunnanensis Ching, in S. S. Chien & Chun, FI.
Reipubl. Popul. Sin. 2: 350. 1959. Diplopter¬
ygium yunnanense (Ching) Ching, in R. II.
Shing, Gloss. Term. Nam. Ferns 50. 1982,
nom. inval. TYPE: China. Yunnan: Jingdong,
Wuliang Shan, W. //. Hsu 6 (holotype, PE).
Acknowledgments. I am indebted to Alan Smith
for discussion of the generic classification of Di¬
plopterygium and Hicriopteris. Thanks also to Prof.
Werner Greuter for nomenclatural comments.
Literature Cited
Ching, 1C C. 1940. On natural classification of the family
“Polypodiaceae.” Sunyatsenia 5: 201—268.
- . 1978. The Chinese fern families and genera: Sys¬
tematic arrangement and historical origin. Acta Phyto-
tax. Sin. 16: 1-37.
- . S. H. I n, Chu 11. Wang & k. H. Shing. 1959.
Gleicheniaceae. In: S. S. Chien & W. Y. Chun, Flora
Reipuhlicae Popularis Sinicae 2: 1 16—132. Science
Press. Beijing.
Copeland, E. B. 1947. Genera Filieum. Chronica Botani-
ca, Waltham, Massachusetts.
Greuter, W., J. McNeill. F. B. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, I). II. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & I). E. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Holttum, R. E. 1957. On the taxonomic subdivision of the
Gleicheniaceae, with description of new Malaysian spe¬
cies and varieties. Reinwardtia 4: 257—280.
- . 1959. Gleicheniaceae. Flora Malesiana ser. 2, 1:
1—36. N.V. Erven P. Noordhoff, Groningen.
Kramer, k. U. 1990. Gleicheniaceae. Pp. 145—152 in k.
Kubitzki (editor). The Families and Genera of Vascular
Plants, Vol. 1. Pteridophytes and Gymnosperms. Vol.
editors, k. U. kramer X P. S. Green. Springer- Verlag,
Berlin.
kung, S. H. 1988. Flora Sichuanica, Tom. 6. Sichuan Sci¬
ence & Technology Publishing House, Chengdu.
Ostergaard Andersen, E. & B. Ollgaard. 2001. Gleichen¬
iaceae. In: G. Hailing & L. Andersson (editors). Flora
of Ecuador 66: 105—170. Botanical Institute, Goteborg
University, Goteborg, Sweden.
Presl, C. 1851. Abh. kiinigl. Boh. ties. Wiss., ser. 5, 6:
386. [Epimel. Bot. 26. 1851.]
Shing, K. 11. 1982. Glossary and Terms of Ferns. Science
Press, Beijing.
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2004
Zhang
Pteridophytes from China
151
Tagawa, M. & K. Iwatsuki. 1979. Gleicheniaceae. In: T.
Smitinand & K. Larsen, Flora of Thailand 3(1): 50—56.
Tryon, 11. M. & A. F. Tryon. 1982. Ferns and Allied Plants
with Special lieferenee to Tropical America. Springer,
Berlin/Heidelberg/New York.
Wang, P. S. & X. Y. Wang. 2(H) I . Pteridophyte Flora of
Guizhou. Guizhou Science & Technology Press, Gui-
yang.
Wu, S. 11. & If. C. Ching. 1991. Fern Families and Genera
of China. Science Press, Beijing.
Volume 14, Number 1, pp. 1-152 of NOVON was published on 23 March 2004.
Volume 14 NOVON
Number 2
2004
Novelties and Notes on Miscellaneous Asian Brassicaceae
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ihsan.al-shehbaz@mobot.org
Abstract. Draba bartholomewii (Brassicaceae), a
new species from Qinghai, China, is described and
illustrated. The monotypic genera Coelonema and
Nesocrambe are reduced to the synonymy of Draba
and Hemicrambe, respectively. The new combina¬
tions Alyssum dahuricum, Draba draboides, Hemi¬
crambe socotrana, Lepidium coronapus, and Pe-
gaeophyton purii are proposed.
Key words: Alyssum , Asia, Brassicaceae, Coe¬
lonema, Dilophia, Draba, Hemicrambe, Lepidium,
Nesocrambe, Pegaeophyton, Ptilotrichum.
Delimitation of genera of Brassicaceae (Crucifer-
ae) is a well-known problem, and 125 of the 337
genera recognized by Appel and Al-Shehbaz (2003)
are monotypic, with about an additional 100 con¬
taining two to four species (oligotypic). Molecular
studies (see review by Koch et al., 2003) have
clearly demonstrated that the vast majority of
monotypic genera are indistinguishable from and
nested within larger genera. Upon a critical re-eval¬
uation of morphology, two monotypic genera, Coe¬
lonema Maximowicz and Nesocrambe A. G. Miller,
are reduced herein to synonymy of other genera
well established on the basis of morphological and
molecular grounds (Koch & Al-Shehbaz, 2002;
Koch et ah, 2003; Warwick & Black, 1997; Gomez-
Campo, 1999).
I'he present paper deals with the description of
a new species of Draba, as well as nomenclatural
adjustments in the genera Alyssum I.., Draba L,
Hemicrambe Webb, Lepidium L., and Pegaeophyton
Hayek & Handel-Mazzetti.
Alyssum versus Ptilotrichum
Although most authors place Ptilotrichum C. A.
Meyer in the synonymy of Alyssum (e.g., Dudley,
1964; Ball & Dudley, 1993; Zhou et ah, 2001; Ap¬
pel & Al-Shehbaz, 2003), a few others (e.g., Re-
benskaya, 1994; Czerepanov, 1995) maintain both
genera. As indicated by Zhou et ah (2001), the
characters used to distinguish Ptilotrichum from
Alyssum (e.g., white vs. yellow flowers, edentate vs.
dentate filaments, and 1 -seeded vs. 2- or 3-seeded
locules) are unreliable and found within Alyssum
excluding Ptilotrichum. Therefore, these alleged
generic boundaries are artificial, and the two gen¬
era should be united. All except one of the species
originally described in Ptilotrichum have names in
Alyssum. The following new combination is needed.
Alyssum dahuricum (Peschkova) Al-Shehbaz,
comb. nov. Basionym: Ptilotrichum dahuricum
Peschkova, Novosti Sist. Vyssh. Bast. 15: 230.
1979. TYPE: Russia. Ghita: Onon River sys¬
tem, mountain at Bukukun River, near Buku-
kunskij, 18 July 1913, V. Smirnov 1986 (ho-
lotype, LE).
Alyssum dahuricum is easily distinguished at an-
thesis from the closely related A. tenuifolium Ste¬
phan ex Willdenow by having stramineous stems
and by lacking the basal and lowermost cauline
leaves. By contrast, A. tenuifolium has purplish or
greenish stems and persistent basal rosette and
cauline leaves.
Draba versus Coelonema
The monotypic Coelonema was recognized in
various Chinese floras (e.g., Zhou et al., 2001) as
endemic and was said to differ from Draba solely
by including stoloniferous perennials with flattened
Novon 14: 153-157. 2004.
154
Novon
bases of staminal filaments. Draba was said to con¬
sist of annuals, biennials, or perennials with slen¬
der or slightly flattened staminal filaments. A crit¬
ical examination at LE of the type of C. draboides
Maximowicz, as well as numerous other species of
Asian Draba, reveals that these alleged differences
are insignificant because several species of Draba
(e.g., D. sibirica (Pallas) Thellung) are typically sto-
loniferous, and the filament bases vary in this com¬
plex genus of ca. 350 species from slender to dis¬
tinctly flattened. Therefore, the conclusion reached
by Appel and Al-Shehbaz (2003) in uniting Coe-
lonema with Draba is justified, and the following
new combination is needed to accommodate C. dra¬
boides in Draba.
Draba draboides (Maximowicz) Al-Shehbaz,
comb. nov. Basionym: Coelonema draboides
Maximowicz, Bull. Acad. Imp. Sci. Saint-Pe-
tersbourg 20: 424. 1880, TYPE: China. [Gan¬
su]: “Terra Tangutorum. Jugum S a fl. Tetung
ad declivitates denudates mintium rarius, 30
A pri 1 — 1 2 May 1873,” N. M. Prezewalski 14
(holotype, LE).
Draba
During a visit to LE in 2002, I examined the
four syntypes of Draba eriopoda Turczaninow, a
species widely distributed in Bhutan, China, India,
Mongolia. Nepal, and Russia. It became immedi¬
ately evident that the treatment of D. eriopoda in
Zhou et al. (2001) represented two distinct species,
of which one is described below as new.
Draba bartbolomewii Al-Shehbaz, sp. nov.
TYPE: China. Qinghai: Dari (Darlag) Xian,
just N of Manzhang, along Manzhang He, belw.
Dari & Banma (Baima), at base of valley sides
in loose soil, 4000 m, 33°17'51"N,
100°25'55"E, 12 Aug. 1993, Ho Tingnung,
Bruce Bartholomew A- Michael G. Gilbert 1 185
(holotype, MO; isotypes, BM. CAS, E, GH,
HNWP, PE). Figure 1.
Herba annua. Caules decumbentes, ad basim ramosi,
pilis stellatis praediti. Folia basalia nulla; folia caulina 2-
6(— 10), sessilia, nonauriculata, ovata vel elliptica, 1—3.5
X 0.4—1. 5 cm, subtus pilis subsessilibus stellatis 4-radia-
tis praedita, supra pilis subsetosis simplicibus et stipitato-
furcatis praedita. Racemi 8— 20-flori, rharhidibus rectis.
Pedicelli fructiferi horizontales, 5—15 mm longi. Petala
Hava, spathulata, 1.2-1. 6 X 0.5— 0.6 mm. Ovarium 14—
20-ovulatum. Fructus oblongi, D — 1 ()(— 1 2) X (2.5— )3— 3.5
mm, compressi, pilis simplicibus et subsessilif)us furcatis
praediti. Semina 1—1.3 X 0.7-0.9 mm.
Herbs annual, 5-25 cm tall; stems decumbent.
branched primarily at or near base, pubescent with
subsessile stellate trichomes rarely mixed with sim¬
ple ones, rarely glabrous distally. Basal leaves w ith¬
ered by anthesis; cauline leaves 2 to 6(to 10), ses¬
sile, not auriculate; leaf blade ovate to elliptic, I —
3.5 X 0.4— 1.5 cm, abaxiallv pubescent with
subsessile or short-stalked, 4-rayed, stellate tri¬
chomes with simple rays, adaxially with subsetose
simple or long-stalked forked trichomes sometimes
mixed with fewer, smaller, 4-rayed stellate ones,
base cuneate, margin 3- to 5-toothed on each side,
apex acute. Racemes 8- to 20-flowered, ebracteate,
elongated in fruit; rachis straight; fruiting pedicels
5—15 mm long, horizontal, straight, pubescent all
around, or rarely glabrescent. Sepals oblong, 1-1.3
X 0.6-0. 8 mm, erect, abaxially pilose with stellate
trichomes, base of lateral pair not saccate, margin
membranous; petals yellow, spatulate, 1.2— 1.6 X
0.5— 0.6 mm, apex retuse, obscurely clawed at base;
filaments 0.8-1 mm; anthers ovate, 0. 1-0.2 mm;
ovules 14 to 20 per ovary. Fruit oblong, 8-10(-12)
X (2.5— )3— 3.5 mm, often horizontal, latiseptate, not
twisted; valves sparsely puberulent with simple and
subsessile forked trichomes, rarely glabrescent, ob¬
scurely veined, base and apex acute; style obsolete,
rarely to 0.1 mm; seeds brown, ovate, 1-1.3 X 0.7—
0.9 mm. Flowering and fruiting in August.
fhe new species is named after Bruce Barthol¬
omew, one of the collectors of the type collection.
Draba bartholomewii is easily distinguished from
D. eriopoda by having decumbent stems, horizontal
fruiting pedicels pubescent all around or rarely gla¬
brous, pilose sepals with stellate trichomes, small
petals 1.2— 1.6 X 0.5— 0.6 mm, oblong, often hori¬
zontal fruits (2.5-)3— 3.5 mm wide, and fruit valves
puberulent with simple and subsessile forked tri¬
chomes. By contrast, D. eriopoda has erect stems,
divaricate-ascending fruiting pedicels often gla¬
brous adaxially and pubescent abaxially, pilose se¬
pals with simple trichomes, larger petals 2— 3(— 3.5)
X (0.5— )0. 8—1 mm, erect or ascending, ovate or
ovate-elliptic fruits 1.5-2. 5 mm wide, and glabrous
fruit valves.
Draba bartholomewii is also related to I), alticola
Komarov (Afghanistan, China, Kyrgyzstan. Tajikis¬
tan), which it resembles in flower size and fruit
shape. However, the latter has rosulate basal
leaves, leafless stems, a (lexuous rachis in fruiting
racemes, pilose sepals with simple and forked tri¬
chomes, glabrous fruits 1-2 mm wide, and seeds
0.4— 0.5 mm long. Draba bartholomewii lacks the
basal leaf rosette and has 2 to 6(to 10) cauline
leaves, a straight rachis in fruiting racemes, pilose
Volume 14, Number 2
2004
Al-Shehbaz
Asian Brassicaceae
155
Figure 1. Draba bartholomewii Al-Shehbaz. —A. Plant. — B. Trichomes of abaxial leaf surface. — C. Trichomes of
adaxial leaf surface. — D. Petal. Scales: A = 1 cm; B. C = 0.2 mm; D = 0.5 mm. Drawn by Al-Shehbaz from the
holotype, Ho , Bartholomew & Gilbert I IBS (MO).
156
Novon
sepals with stellate trichomes, puberulent fruits
(2.5— )3— 3.5 mm wide, and seeds 0.7— 0.9 mm long.
Paratypes. CHINA. <^inghai: Nangqen Xian, Xiao-
long Gou, W of Nangqen on road to Domba, 32°17'N,
96°16'E, T. /V. Ho, H. Bartholomew, M. Watson & M. G.
Gilbert 2822 (CAS, HNWP, MO); Tungjen Xian, T. P.
Wang 6186 (PE). Sieliuan: Haitzeshan, //. Smith 1 1598
(MO, TI, UPS).
Hemicrambe V ERSUS Nesocrambe
Hemicrambe consists of two species, of which
one, //. fruticulosa Webb, is endemic to Morocco
and the other, H. fruticosa (C. C. Townsend) Gdmez-
Campo, is endemic to the island Soqotra (Yemen).
The two species are subshrubs disjunetly separated
by some 0900 km (Gdmez-Campo, 1977, 1978;
Townsend, 1971). The genus is characterized by
having yellow or white flowers, petiolate and den¬
tate or lyrate-pinnatipartite leaves, patent median
nectaries, reflexed fruiting pedieels. and segment¬
ed, flattened fruits with a 2- to 4-seeded, indehis-
cent distal segment and a seedless or 1 -seeded, de¬
hiscent proximal segment (Appel & Al-Shehbaz,
2003).
The recently described Nesocrambe (Miller et al.,
2002) is indistinguishable from Hemicrambe in al¬
most every morphological aspect. It is said to be a
perennial herb with a woody base instead of a sub¬
shrub, but this is an unreliable distinction. Both
genera have well-developed, [latent, median nectar
glands, a feature unique in the entire tribe Bras-
sieeae (Townsend, 1971; Gdmez-Campo, 1977;
Miller et al., 2002). Miller et al. (2002) did not
include Nesocrambe socotrana A. G. Miller in Hem¬
icrambe because when they compared it with the
Soqotran //. fruticosa they found that H. fruticosa
has white instead of yellow flowers, dentate instead
of lyrate-pinnatisect leaves, and cylindric instead
of flattened fruits. The Moroccan //. fruticulosa has
both yellow flowers and lyrate-pinnatipartite leaves.
The only other difference between the two genera
is that in Hemicrambe the distal fruit segment is
compressed and the proximal segment is sometimes
seedless, whereas in Nesocrambe the distal fruit
segment is cylindric and the proximal segment is
1 -seeded. However, such highly artificial separation
is good only at the species rather than the generic
level. Miller et al. (2002: 62) have indicated that
Nesocrambe “is clearly related to Hemicrambe." In
my opinion, the remarkable similarities in nectar
gland and almost every aspect of the plant (see ta¬
ble 1 in Miller et al., 2002: 63) strongly argue for
the maintenance of one genus. Furthermore, there
is no need to create a new monotypic genus when
its nearest relatives, species of Hemicrambe, are
known. Several other genera of the tribe Brassiceae
(e.g., Conicya Porta & Rigor ex Roy, Vella L.) show
fruit diversity far greater than in Hemicrambe in¬
cluding Nesocrambe (Leadlay & Heywood, 1990;
Warwick & Al-Shehbaz, 1998).
Miller et al. (2002: 66) stated that “the perennial
habit is relatively uncommon in the Brassicaceae.”
However, Al-Shehbaz (1984: 349) indicated that
“more than 62 percent [of the family] are peren¬
nials” and only about 5% are typically woody.
Key to the Species of Hemicrambe
la. Leaves crenate-dentate; flowers white; Soqotra
. H. fruticosa
1 b. Leaves lyrate-pinnatisect or lyrate-pinnatipartite;
flowers yellow.
2a. Perennial herbs woody only at base; fruits
cylindric; Soqotra . H. socotrana
2b. Subshrubs; fruits flattened; Morocco . . .
. H. fruticulosa
Hemicrambe socotrana (A. G. Miller) Al-Sheh¬
baz, comb. nov. Basionym: Nesocrambe soco¬
trana A. G. Miller, in Miller et al., Willden-
owia 32: 63. 2002. TYPE: Republic of Yemen.
Western Soqotra, Sharahin, 12°31'N, 53°20'E,
450 m, 6 Feb. 2001, A. G. Miller, R. Atkinson,
A. W. Kulaidi & N. Taleeb M -2002 1 (holotype,
E; isotypes, B not seen, UPS not seen).
Lepidium
Based on extensive molecular studies (Brtigge-
mann, 2000) and critical evaluation ol morphology
(Appel & Al-Shehbaz, 2003; Al-Shehbaz et al.,
2002), Coronopus Zinn was reduced to the synon¬
ymy of Lepidium. All except one species of Coron¬
opus have correct names in Lepidium. The excep¬
tion is C. squamatus (Forsskdl) Ascherson, a
species originally described in Lepidium by Forss-
kal (1775). However, the earliest validly published
name for the species is Cochlearia coronopus L.
(Linnaeus, 1753), and this specific epithet should
be used in Lepidium when this genus is united with
Coronopus. Therefore, the following new combina¬
tion is needed.
Lepidium coronopus (L.) Al-Shehbaz, comb. nov.
Basionym: Cochlearia coronopus L., Sp. PI. 2:
648. 1753. TYPE: (lectotype, designated by
Jonsell & Jarvis, 2002: 68) Herb. Linn. 826.5
(LINN).
Pegaeophyton versus Dilophia
Rawat et al. (1996) described the new species
Dilophia purii from Garhwal in Northwest India.
Volume 14, Number 2
2004
Al-Shehbaz
Asian Brassicaceae
157
The excellent illustration immediately suggested a
possible affinity to Pegaeophyton. However, the de¬
scription and illustration indicated the presence of
glandular trichomes, a feature not known in any
species of Pegaeophyton and Dilophia Thomson.
When I examined the MO isotype of D. purii, I
found none of the trichomes were glandular, and
the generic affinity of this plant to Pegaeophyton
became abundantly clear. Pegaeophyton differs
from Dilophia by having accumbent instead of in¬
cumbent cotyledons, nonfleshy instead of fleshy fu¬
siform roots, obtuse instead of strongly apiculate
anthers, and smooth instead of apically cristate
fruits.
Pegaeophyton purii (1). S. Rawat, L. R. Dangwal
& R. D. Gaur) Al-Shehbaz, comb. nov. Ras-
ionym: Dilophia purii I). S. Rawat, L. R. Dang¬
wal & R. I). Gaur, J. Bombay Nat. Hist. Soc.
93: 262. 1996. TYPE: India. Northwest Him¬
alaya, Roopknnd, 4850 m, 11 Aug. 1993, D.
S. Rawat .s'.//, (holotype, GUH 22,498 not seen;
isotype, MO).
Except for having flattened instead of terete and
8- instead of 2- to 4-seeded fruits. Pegaeophyton
purii is quite similar in all aspects of leaves and
flowers to P. nepalense Al-Shehbaz, Arai & H.
Ohba, a species originally thought to be endemic
to Nepal (Al-Shehbaz et ah, 1998) but later re¬
ported from Bhutan, China, and India (Al-Shehbaz,
2000). The collection of mature fruiting material of
P. purii should help in the determination of whether
or not the two species are conspecific.
Acknowledgments. I am profoundly grateful to
Henk van der Werff for correcting the Latin, John
H. Wiersma for his note on Lepidium coronopus,
Antony G. Miller for allowing me to examine the
type collection of Nesocrambe socotrana, Steve Caf-
ferty for the typification of Cochlearia coronopus, D.
S. Rawat for sending an isotype of Dilophia purii ,
and Nicholas J. Turland for opinions on various no-
menclatural matters. I am deeply indebted to Vic¬
toria C. Hollowell. Neil A. Harriman, and Steve L.
O'Kane, Jr., for their editorial advice and review of
the manuscript.
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Psychotria carrascoana (Rubiaceae), a New Species from the
Carrasco Vegetation of Northeastern Brazil
Piero G. Delprete
Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York 10458-
5126, U.S.A. pdelprete@nybg.org. Current address: National Herbarium of the Netherlands,
Utrecht University Branch, Plant Systematics, Heidelberglaan 2, 3584 CS Utrecht,
The Netherlands, p.delprete@bio.uu.nl
Elnatan Bezerra de Souza
Curso de Biologia, Universidade Estadual Vale do Acarau, Sobral, Ceara, 62040-370, Brazil.
ebsouza@nvanet.br
Abstract. Psychotria carrascoana (Rubiaceae,
Psychotrieae) is here described and illustrated. It
is endemic to carrasco vegetation, a closed, shrub¬
by, xerophilous vegetation on quartzitic sand soils,
on the plateaus of the Serra da Ibiapaba and Serra
do Araripe, at altitudes of 800 to 000 m, state of
Ceara, Brazil. Because of its reduced, triangular
stipules, leaves drying pale green, commonly soli¬
tary flowers, and dorsally tricostate pyrenes, this
species is placed in Psychotria subg. Heteropsycho-
tria.
Key words: carrasco , Ceara, northeastern Bra¬
zil, Psychotria, Psychotrieae, Rubiaceae, shrub-
land.
As part of the project Rubiaceae do Estado do
Ceara, in northeastern Brazil, fieldwork was con¬
ducted during 1998—2000, which has produced
about 400 collections of species of this family. A
preliminary checklist of the Rubiaceae of Ceara
was recently presented (Delprete et ah, 2001), and
an annotated checklist will be published in the
near future (Delprete & Souza, in prep.). Two new
species of Rubiaceae were recently described from
the arid vegetation of this state (Simira gardneriana
Barbosa & Peixoto (2000); Mitracarpus longicalyx
Souza & Sales (2001)). and the identity of some
other taxa collected during this project is currently
under study. Among these, a species of Psychotria
restricted to carrasco vegetation, and readily distin¬
guishable by its small leaves drying pale green, tri¬
angular stipules, flowers commonly solitary, and
dorsally trieostate pyrenes, is described below.
Ecological Obskhvations About Carrasco
Vegetation
The definition of carrasco vegetation lias long
been debated, as it is not readily distinguishable
from that of caatinga, the widespread xerophilous
vegetation of northeastern Brazil. Caatinga is com¬
monly characterized by two vegetational layers:
sparse to moderately dense trees and shrubs fre¬
quently armed with thorns and almost completely
leafless during the dry season, and the lower laver
with the presence of Bromeliaceae and Cactaceae,
and annual herbs (Rizzini, 1997). Andrade-Lima
(1978) stated that carrasco vegetation is physiog-
nomically distinct from caatinga vegetation be¬
cause of the high density of shrubs and treelets,
single-layered vegetation, and the almost complete
absence of Cactaceae and Bromeliaceae. Rizzini
(1997) and Fernandes (1981) recognized carrasco
vegetation as sufficiently distinct from caatinga
vegetation. Fernandes (1990) and Fernandes and
Bezerra (1990) postulated two possible hypotheses
about the identity and origin of carrasco vegetation:
either as a distinct natural vegetation, or as a result
of partial degradation of the cerradao vegetation,
which gives the general aspect of a dense capoeira
(secondary vegetation that can be originated from
the degradation of several possible vegetation
types). Cerradao is usually characterized by three
vegetational layers: a tree layer 8— 1 2( — 1 8) m tall,
with closed canopy and almost completely devoid
of epiphytes; an intermediate arbustive layer about
3—5 m tall and with sparse shrubs and rare lianas;
and a lower herbaceous layer, composed primarily
of Cyperaceae, Poaceae, and Bromeliaceae (ef. Riz¬
zini. 1997).
Araujo et al. (1998a, 1998b), in a project focused
on the plant community and floristic composition of
carrasco, studied three plots in the southern portion
of the Planalto da Ibiapaba, near Novo Oriente,
Ceara. Brazil. As a result of that study, only 31 out
of 102 shrubby and arboreal species were found to
Novon 14: 158-162. 2004.
Volume 14, Number 2
2004
Delprete & Souza
Psychotria carrascoana from Brazil
159
be exclusive to carrasco, while the other species
were shared with the contiguous vegetation types.
Those data alone confirmed the difficulty of decid¬
ing if carrasco vegetation should be treated as a
degraded cerradao or a distinct vegetation type. Af¬
ter a series of additional studies, Araujo and Mar¬
tins (1999) and Araujo et al. (1999) recently con¬
cluded that carrasco vegetation can be defined as
a deciduous, dense, single-layered shrubland, with
vines, irregular canopy, and sparse trees, with an
average annual precipitation of 668—1289 mm, a
dry season ranging from 5 to 7 months a year, grow¬
ing on deep, acidic, distrofic quartzitic sands, at
700—900 m elevation. According to those authors,
carrasco vegetation occurs within the semi-arid do¬
main of northeastern Brazil, at the Chapada do Ar-
aripe, southern Ceara, and Planalto da Ibiapaba, at
the border area of the states of Ceara and Piauf.
The new species of Psychotria described below is
endemic to this vegetation (hence the specific ep¬
ithet), and it grows in the shade of shrubs and tree-
lets.
Psychotria carrascoana Delprete & E. B. Souza,
sp. nov. TYPE: Brazil. Ceara: Planalto da Ibia¬
paba, Mun. Ubajara, Jaburuna Sul. 830 m, 5
Jan. 1995 (fl), F. S. Araujo 1054 (holotype,
EAC; isotype, NY). Figure 1.
Haec species, suffrutex usque ad 1.3 m altus, quoad
inflorescentiam terminalem subsessilem paucifloram P.
subtriflorae similis, sed ab ea stipulis triangularibus (nec
bilobis), inflorescentia ebracteata (nec bracteis naviculi-
formibus subtenta), tubo corollae 1.7— 2.8 mm (nec 6—7
mm) longo atque fructibus rubris (nec cyano-purpureis)
diversa.
Subshrub 0.4— 1.3 m tall, densely branched from
the base; main stems often tortuous; bark thin,
creamy white; raphides present in all plant parts.
Stipules free at base, triangular. 1-2 X 0.8—2 mm,
margin entire or sometimes denticulate, apex acute,
glabrous outside, pubescent inside; leaves opposite,
sessile to subpetiolate; petioles to 1.5 mm long;
blades 9—30 X 2-9 mm, linear, narrowly elliptic to
lanceolate, base attenuate, apex acute to obtuse,
margins narrowly revolute, chartaceous, glabrous
throughout, pale green; midvein prominent below,
discolorous; secondary veins 5 to 9 each side, em¬
bedded within the lamina; domatia absent. Inflo¬
rescence uniflorous (rarely biflorous), terminal, on
lateral short-shoots of 5 to 7 reduced nodes. Flow¬
ers subsessile to short-pedicellate; pedicels to 1.3
mm long; hypanthium broadly turbinate, ea. 2 X
1.2 mm, glabrous; calyx shallowly cupular; tube
0.2— 0.3 mm long; calyx lobes 5(6), 0.3— 0.5 mm
long, deltoid to narrowly triangular, acute; corolla
narrowly infundibuliform, 4.5— 6.5 mm long, gla¬
brous outside, with a ring of hairs from stamen in¬
sertion to mouth inside; corolla tube 1.7— 2.8 mm
long; corolla lobes 5 (rarely 6), 2. 5-3.8 X 1.4-1. 7
mm, oblong-ovate, acute and slightly thickened at
apex, antrorse pubescent at basal portion inside;
stamens subsessile, partially exserted; filaments
0.2— 0.5 mm long, glabrous; anthers oblong, 0.7— 0.9
X 0.3— 0.5 mm; disk entire; style terete, 4—6 mm
long; style branches spatuiate, 0.5— 0.7 X 0.3— 0.4
mm. Fruits drupaceous, ovoid to ellipsoid, 3—5 X
3—4 mm. red; pyrenes plano-convex, dorsally 3-cos¬
tate; seeds suborbicular, ca. 2.5 X 2 mm, pale
brown; exotesta smooth.
Distribution, habitat, and phenology. Endemic
to the carrasco vegetation of the Planalto da Ibia¬
paba and Chapada do Araripe, slate of Ceara, and
probably occurring in the same vegetation type in
the contiguous state of Piauf, always observed and
collected in the shade of taller shrubs, at 800—900
m elevation. Flowering specimens were collected in
January, and fruiting specimens in March and Sep¬
tember.
Paratypes. BRAZIL. Ceara: Planalto da Ibiapaba,
Mun. Ubajara, Jaburuna Sul. 830 m. 27 Apr. 1994 (st). F.
S. Araujo 679 (LAG), 23 May 1994 (st), 768 (EAC); Plan¬
alto da Ibiapaba, Mun. Novo Oriente, Baixa Fria, 16 Feb.
1991 (st). F. .S'. Araujo 278 (EAC); Planalto da Ibiapaba,
Mun. Tiangua. 19 Nov. 1994 (st). A. G. Fernandes & F. ,/.
A. Matos FAC 21465 (EAC); Planalto da Ibiapaba, BR-
222. 3 km from Tiangud, 3°45'S, 41°01 'W, ca. 800 m, 21
Mar. 2000 (st), Delprete et al. 7196 (EAC. NY. uva [Univ-
ersidade Vale do Acarau|), 7197 (EAC, uva), 7198 (EAC,
NY, uva), 7199 (EAC, NY, uva), 7203 (EAC, uva); Cha¬
pada do Araripe, Mun. Crato, Sftio Mirindiva, 7°17'S,
39°33'W, 900 m. 29 Mar. 2000 (voting f r). Delprete et id.
7304 (EAC. NY, uva), s.n. (EAC.‘ k. MO, NY. US); Cha¬
pada do Araripe, Mun. Crato, 19 Sep. 2001 (fr), F. S.
Cavalcanti tV F. Silveira 843 (EAC, NY, uva).
Psychotria L., as traditionally recognized, is one
of the largest genera of flowering plants, with about
2000 species worldwide. Several authors have tried
to organize this taxonomic conundrum, either by
dividing it into several smaller genera (e.g., Bre-
mekamp, 1934), or by treating it as a large genus
with many sections (e.g., Steyermark, 1972, 1974).
Psychotria s.l. has been shown to be paraphyletic
in the molecular phytogenies presented by Anders-
son and Rova (rp.s l6; 1999), and Nepokroeff el al.
(ITS and rbch; 1999). Following ihe molecular data
published by these authors and additional morpho¬
logical evidence, the genera IS'otopleura Breme-
kamp and Carapichea Aublet have been recently
resurrected by Taylor (2001) and Andersson (2002),
respectively. Notopleura is easily distinguished,
among other morphological characters, by its pseu-
160
Novon
Figure 1. Psychotria carrascoana Delprete & E. B. Souza. — A. Branch with immature fruits. — B. Short-shoot with
solitary, terminal (lower hud. — C. Flower in anthesis. — 1). Style. — E. Section of open corolla. — F. Mature fruit. A
and F from photos of living material; B— E, from Araujo 1054 (types, EAC, NY).
Volume 14, Number 2
2004
Delprete & Souza
Psychotria carrascoana from Brazil
161
doaxillary inflorescences, and the sheathing stip¬
ules with an interpetiolar appendage inserted below
the upper margin (Taylor, 2001); and Cnrapichen
by its terminal, condensed inflorescences subtend¬
ed by foliose bracts, pyrenes with germination slits
on abaxial ridges, and persistent stipules, withering
on the stem (Andersson, 2002; Delprete, 2001,
2003). The rest of the Psychotria complex, cur¬
rently divided into subgenus Psychotria (Steyer-
mark, 1972: 444) and subgenus Heteropsychotria
Steyermark (1972: 484), remains taxonomically un¬
resolved, and its close relationship with Palicourea
Aublet has been addressed (e.g., Muller Argovien-
sis, 1881: Nepokroeff et ah, 1999), but not yet re¬
solved (Taylor, 1996. 1997, and work in progress).
About 1 I species of Psychotria are known from
the state of Ceara (Delprete et al., 2001), and
among them Psychotria carrascoana is the species
with the smallest leaves, and the only one with sol¬
itary (or rarely paired) flowers. This species has of¬
ten been collected when sterile, mostly because of
its minute, usually solitary, greenish white flowers,
and therefore difficult to spot during anthesis. Be¬
cause of the reduced, triangular stipules, leaves
drying pale green, terminal flowers commonly sol¬
itary (or rarely in pairs), fruits red, and pyrenes
dorsally tricostate, Psychotria carrascoana is placed
in Psychotria subg. Heteropsychotria. 'I bis species
is most similar to Psychotria subtriflora Muller Ar-
goviensis (Zappi & Stannard, 1995: 571) because
of the pauciflorous, subsessile, terminal inflores¬
cences, but it differs by having stipules triangular
(vs. bilobed, lobes narrowly triangular in P. subtri¬
flora). inflorescences not subtended by bracts (vs.
subtended by navicular bracts in P. subtriflora), co¬
rolla lube 1.7-2. 8 mm long (vs. 6-7 mm long in P.
subtriflora), and red fruits (vs. purplish blue in P.
subtriflora).
Acknowledgments. We are grateful to Francisca
Simdes Cavalcanti, Curator of the Herbario Prisco
Bezerra (EAC; Universidade Federal do Ceara, For¬
taleza) and Coordinator of the project Bubiaceas do
Ceara. for help with field and herbarium work; to
Afranio Fernandes (EAC) for accompanying us dur¬
ing fieldwork in the carrasco vegetation of the Plan-
alto da Ibiapaba and Chapada do Araripe; to Fran¬
cisco Jose de Abreu Matos (Laboratorio de Produtos
Naturais, Universidade Federal do Ceara, Fortale¬
za) for contributing use of the university vehicle for
our expeditions during 1998-2000; to the Conselho
Nacional de Pesquisa (CNPq) for granting support
and permission for collecting Rubiaceae in Ceara;
and to The New York Botanical Garden, the Univ¬
ersidade Federal do Ceara, and the Universidade
Estadual Vale do Aearau (uva; Sobral, Ceara) for
economic and logistic support. The herbarium of
the Universidade Estadual Vale do Aearau (ca.
4000 specimens), the institution of the junior au¬
thor, is not yet included in the Index Herbariorum.
We are thankful to Michael Nee (NY) for proof¬
reading a draft of the manuscript, Bruno Manara
(VEN) and Roy Gereau (MO) for correcting the Lat¬
in diagnosis, and William Burger (F) and an anon¬
ymous reviewer for helpful comments on the man¬
uscript.
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A. Pulle, Flora of Surinam, Vol. 4. J. H. de Bussy, Am¬
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cies of Psychotria subgenus Heteropsychotria (Rubi¬
aceae), with observations on rubiaceous taxonomic
characters. Brittonia 53: 396—404.
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for Psychotria carapichea (Rubiaceae), with a correction
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Una Nueva Especie de Tibouchina (Melastomataceae)
de Guerrero, Mexico
Jesus Ricardo de Santiago Gomez
Laboratorio tie Plantas Vasculares, Facultad de Ciencias, UNAM, Cireuilo Exterior,
Ciudad Universitaria. Mexico, D.F. 04510. rdesantiago@correo.unam.mx
RESUMEN. Se describe e iluslra Tibouchina ara-
neicalyx de la vertiente del Oceano Pacifico de la
Sierra Madre del Sur en Guerrero, Mexico. Se dis-
cuten las posibles relaciones de esta especie con
T. rnexicana (D. Don) Cogniaux y otras especies con
flores pentameras y estambres fuertemente dimor-
ficos, de las que se distingue principalmente por
tener ramas agudamente tetragonas, lobulos del ea-
liz triangulares mas cortos que el hipantio y petalos
con cilios eglandulares.
Abstract. Tibouchina araneicalyx is described
and illustrated from the Pacific slopes of the Sierra
Madre del Sur in Guerrero, Mexico. Its possible
relationships with T. rnexicana (D. Don) Cogniaux
and another Mexican species of Tibouchina with
pentamerous flowers and strongly dimorphic sta¬
mens are discussed; it is principally distinguished
from these species by the sharply tetragonal
branches, the triangular calyx lobes shorter than
the hypanthium, anti the petals with eglandular cil¬
ia.
Key words: Guerrero, Melastomataceae, Mexi¬
co, Tibouchina.
Tibouchina Aublet, con poco mas de 350 espe¬
cies, es el genero de frutos capsulares con mayor
numero de especies de la familia Melastomataceae
(Almeda, 1993; Renner, 1993). Se distribuye destle
el norte tie Argentina hasta el noroeste tie Mexico,
con su maxima diversidad en la euenea del Ama¬
zonas y regiones antlinas adyacentes, con alrededor
de 10 especies en las Antiilas y en America Cen¬
tral. En Mexico esta representado por 25 especies
de las cuales 21 o 22 son endemicas del pais, para
un endemismo tie 84 a 88%, el mas alto dentro de
los distintos generos tie Melastomataceae presentes
en Mexico, esto de acuerdo con los principales tra-
tamientos que han descrito o revisado las distintas
especies del genero presentes en Mexico (Cog¬
niaux, 1891; Rrantlegee, 1913, 1914; Standley,
1924; Wilson, 1958; Williams, 1963; Wurdack.
1967; Almeda, 1993; Totlzia, 1999).
De las 25 especies antes mencionadas, 9 de
ellas, incluyendo la que se describe en este trabajo,
presentan dos series de estambres fuertemente di-
morfieos, los mayores con las tecas y conectivo ro-
sados a rojizo-violaceos mucho mas largos que en
los estambres menores que tienen anteras y conec¬
tivo amarillos, constituyendo un grupo claramente
diferenciado en el que Tibouchina breedlovei Wur¬
dack. 7. chiapensis Wurdack y 7. roseotincta Totlzia
presentan flores estrictamente tetrameras, mientras
que T. araneicalyx descrita aquf, T. durangensis
Standley, T. galeottiana (Triana) Cogniaux, T. me-
xicana (D. Don) Cogniaux, T. rufipilis (Schlechten-
dal) Cogniaux y T. thulia Todzia son basicamente
pentameras, con algunas flores tetrameras, hexa-
meras o muy raramente heptameras en las inflores-
cencias fobs. pers.).
En los herbarios mexicanos mas importantes Ti¬
bouchina se encuentra representado [tor un numero
abundante de colectas. Debido a la compleja ta-
xonomfa del genero, a su abundante variacion mor-
fologica y a la earencia tie claves y descripciones
adecuadas para las especies mexicanas, en nume-
rosas ocasiones el material se encuentra mal de-
terminado o sin asignacidn de esjtecies. Por tal mo-
tivo, aetualmente se realiza una revision de los
ejemplares de este genero con el objetivo de ubicar
claramente las especies presentes en Mexico. Del
material revisado, tanto de los herbarios antes men-
cionados como de la amplia coleccidn del proyecto
“Flora de Guerrero” de la Facultad de Ciencias de
la Universidad Nacional Autonoma de Mexico
(UNAM) aetualmente en revision, se detectaron va¬
ries espeefmenes de un taxon procedente de la Si¬
erra Madre del Sur de Guerrero con estambres mar-
cadamente dimdrficos. Dado que tiene hipantios
con pelos no glandulares, el taxon se encuentra re-
lacionatlo con /.’ rnexicana, T. rufipilis y T. thulia,
especies de las que se distingue principalmente por
tener las ramas agudamente tetragonas. fits lobulos
del caliz triangulares mas cortos que el hipantio y
los petalos con cilios no glandulares, por lo que se
propone en este trabajo como una especie nueva
para la ciencia.
Novon 14: 163-167. 2004.
164
Novon
Tihoiichinn araneicalyx de Santiago, sp. nov.
TIPO: Mexico. Guerrero: Mun. Atoyac . Hipantio con tricomas adpresos no glandu¬
lares . T. roseotincta
1 1). Flores basieamente pentdmeras (ocasionalmente
hexitmeras o heptameras en T. galeottiana y te-
trdmeras, hexameras o heptameras en T. aranei¬
calyx).
4a. Hipantio con pelos glandulares.
5a. Ramas con pelos alargados adpresos o
abiertos, hojas basalmente nervadas . .
. T. galeottiana
5b. Ramas con pelos cortos adpresos, hojas
pi i nervadas . T. durangensis
41). Hipantio con pelos no glandulares.
6a. Lobulos del cdliz mds cortos que el hi¬
pantio, triangulares o lineares, petalos
con cilios eglandulares ... 7! araneicalyx
6b. Ldbulos del ealiz iguales o mas largos
que el hipantio, generalmente oblanceo¬
lados, petalos con cilios glandulares.
7a. Hipantio con pelos adpresos, tallo
con algunos pelos glandulares . . .
. T. thulia
Volume 14, Number 2
2004
de Santiago
Tibouchina araneicalyx de Mexico
167
71). Hipantio con pelos ascendentes o
patentes.
8a. Ramas con pelos largos (0.8—3
mm) cafe-rojizos adpresos a
patentes, hipantio con pelos
rectos subadpresos . . 7. rufipilis
8b. Ramas con pelos blanquecinos
adpresos de menos de 2 mm,
hipantio con pelos patentes
mas o menos flexuosos ....
. T. mexicana
Pardtipos. MEXICO. Guerrero: Mun. Atoyae de Al¬
varez. 2 km al W de El Molote, 5 die. 2000, de Santiago
868 (MEXU), 5 die. 2000. de Santiago 86 9 (CAS. ENCB.
FCMK, MO, UAGC, UAMIZ, XAL); Canada a 0.5 km V
de El Molote, 5 die. 2000, de Santiago 888 (ENCB.
ECME. MEXU), 18 abr. 2001, de Santiago 902 (MEXU),
21 abr. 2001, de Santiago 911 (FCME, MEXU). 10 die.
2001, de Santiago 939 (CAS, ENCB, FCME, MEXU, MO,
UAGC), 10 die. 2001, de Santiago 9 40 (FCME, MEXU),
10 die. 2001, de Santiago 941 (FCME, MEXU); Mun.
Petatlan. El Venado, 1 nov. 1991, de Santiago 81 (ENCB,
ECME, MEXU. MO); El Venado, 17 nov. 1991. de San¬
tiago 486 (ENCB, ECME, MEXU, MO), 17 nov. 1994, de
Santiago 489 (ENCB, FCME, MEXU, MO).
Agradecimientos. Agradezco el apoyo prestado,
tanto para revisar la coleccion de Melastomataceae
del Laboratorio de Plantas Vasculares de la Eacul-
tad de Ciencias de la UNAM como para la reali-
zacion de coleetas botanicas en el estado de Gue¬
rrero, a la coordinadora del Area de Taxonomfa de
dicho laboratorio, la M. en C. Nelly Diego; al Cu-
rador del Herbario Naeional de Mexico (MEXU) M.
en C. Mario Sousa por el apoyo recibido para la
revision de la coleccion de Melastomataceae del
herbario; al M. en C. Lucio Lozada por su continuo
apoyo en el trabajo de campo y en la redaecion del
escrito; al Dr. Jose Luis Villasenor, investigador del
Instituto de Biologfa de la UNAM por la revision
crftica del manuscrito, y a la dibujante Ana E. Vi-
niegra por la elaboracion de la ilustracion que
acompana este trabajo.
Eileratura Citada
Almeda, F. 1993. Melastomataceae. Pp. 1—38 in J. Rze-
dowski y G. Calderon (editores). Flora del Bajfo y de
regiones adyacentes, fascfculo 10. Instituto de Ecologfa,
A. C., Patzeuaro, Miehoacan.
Brandegee, T. S. 1913. Plantae Mexicanae Purpusianae,
V. Univ. Calif. Publ. Bot. 4: 375—388.
- . 1914. Plantae Mexicanae Purpusianae, VI. Univ.
Calif. Publ. Bot. 6: 51-77.
Cogniaux, C. A. 1891. Melastomataceae (Monographiae
Phanerogamarum). Pp. 1-1256 in A. y C. de Candolle
(editores), Monogr. Phan. 7.
Lozada, I,.. E. Leon, ,1. Rojas y R. de Santiago. 2003.
Bosque mesofilo de montana en El Molote. Pp. 1—35 in
N. Diego y R. M. Fonseca (editores), Estudios fiorfsticos
en Guerrero. Num. 13. Facultad de Ciencias, UNAM.
Renner, S. S. 1993. Phylogeny and classification of the
Melastomataceae and Memecylaceae. Nordic J. Bot. 13:
519-540.
Standley, P. C. 1924. Melastomataceae. In: Trees and
shrubs of Mexico. Contr. U.S. Natl. Herb. 23: 1046-
1074.
Todzia, C. A. 1999. Ten new species of Tibouchina (Me¬
lastomataceae) from Mexico. Brittonia 51: 255—279.
Williams, E. (). 1963. Tropical American plants. V. Fiel-
diana, Bot. 29: 543—586.
Vi i Ison, P. G. 1958. Contributions to the flora of tropical
America LXIII: Plantae Hintonianae XII. Kew Bull. 13:
160-161.
Wurdack, J. J. 1967. Certamen Melastomataceis XI.
Phytologia 14: 257—274.
A New Species of Dyckia (Bromeliaceae) from
Rio de Janeiro State, Brazil
Rafaela Campostrini Forzza and Bruno Rezende Silva
Jardim Botanico do Rio de Janeiro (JBRJ), Rua Pacheco Eeao 915, CEP 22460-030,
Rio de Janeiro, Brasil, rafaela@jbrj.gov.br; brsilva@jbrj.gov.br
ABSTRACT. A new seaside endemic species, Dy¬
ckia martinellii B. R. Silva & Forzza, from Rio de
Janeiro State, Brazil, is described and illustrated.
The genus was represented in this state only hy D.
pseudococcinea L. B. Smith, from which D. marti¬
nellii can he distinguished by a rosette with a larger
diameter; longer and wider leaves hearing straight,
patent-antrorse larger spines; basal scape bracts
densely arranged, with the others longer than or
equaling the internodes; floral bracts castaneous,
entire, white-lepidote; and sepals orange with a
castaneous apex.
Rksumo. Uma nova especie endemiea do litoral,
Dyckia martinellii B. R. Silva & Forzza, do estado
do Rio de Janeiro, Brasil e descrita e ilustrada. 0
genero estava representado neste estado apenas por
I), pseudococcinea L. B. Smith da qual D. marti¬
nellii pode ser diferenciada pela roseta de maior
diametro; pelas folhas maiores e mais largas, es-
pinhos patente-antrosos maiores; pelas br&cteas
basais do escapo congestas as demais maiores ou
igualando os entrenos; I trade as florais castanhas,
inteiras, alvo-lepidotas e pelas s£palas laranja com
dpice castanho.
Key words: Brazil, Bromeliaceae, Dyckia, Pit-
cairnioideae, Rio de Janeiro state.
Dyckia Schultes & Schultes f., with ca. 200 spe¬
cies, is among the largest genera in Bromeliaceae
subfamily Pitcairnioideae and the one with greatest
diversity in Brazil, where approximately 80% of the
species occur (Forzza, 2001). The members of this
genus have an exclusively South American distri¬
bution, with diversity centers in southern Brazil
within the cerrado and campos rupestres formations
(Forzza, 2001). According to a phylogenetic anal¬
ysis based on morphological characters, the genus
Dyckia is monophyletic (excluding D. biflora Mez;
Forzza, 2004). This is supported by four synapo-
morphies: an axial inflorescence; the scape bracts
different from leaves; presence of nectaries on the
sepals; and a petal -staminal ring (Forzza, 2001).
Dyckia martinellii B. R. Silva & Forzza, sp. nov.
TYPE: Brazil. Rio de Janeiro: Mun. Paraty,
Paratymirim, costoes rochosos entre Paratym-
irim e Saco do Mamangud, 5 m elev., 27 Sep.
1990 (fl), G. Martinelli 14413 (holotype, RB).
Figure 1.
Species nova Dyckiae pseudococcineae affinis, sed ro-
sula foliorum circa 60 cm diam., foliis longioribus, latior-
ibusque, spinis rectis vel patente-antrorsis et majoribus,
bracteis scapalibus infernis foliis similibus, dense dispos-
itis, supernis longioribus vel rare internodia aequantibus,
bracteis floriferis castaneis, integris et albo-lepidotis, se-
palis aurantiaceis apice brunneis differt; D. encholirioidi
similis, sed propagulis basalibus brevioribus, rosula fo¬
liorum breviore, foliis brevioribus angustioribusque, flori-
bus per anthesim patentibus, bracteis floriferis integris et
albo-lepidotis, sepalis albo-lepidotis, aurantiaceis apice
brunneis, petalis obtrullatis, aurantiaceis differt.
Plant rupicolous, (lowering 70-100 cm high,
propagating by stout basal shoots; rosettes ca. 60
cm diam. Leaves suberect-arcuate; sheaths elliptic,
2.5—3 X 5-6 cm, castaneous abaxially, whitish
adaxially, margin entire except for a few evanescent
spines in the transition to the blade; leaf blades
20—35 X 2. 2-2. 6 cm, lanceolate, apex attenuate,
whitish abaxially, light green adaxially, margins
spinose, spines straight, patent-antrorse, 3—4 mm
long, 1-1.5 cm apart, light green with a castaneous
apex. Scape erect, 55—97 cm long, 0.7—10 mm
diam., green, sparsely white lepidote; scape bracts
the basal ones subfoliaceous, densely arranged,
spinulose, 11—12.5 X 0.9— 1.2 mm, erect, charta-
ceous, sparsely lepidote; the upper ones exceeding
to rarely equaling the internodes, lanceolate, char-
taceous, stramineous, glabrous to sparsely lepidote,
clasping the scape, apex attenuate, margins spi¬
nulose to inconspicuously serrulate; inflorescence
simple, racemose, 30-43 cm long, laxly flowered,
38- to 65-flowered, rachis orange, sparsely lepidote,
straight; floral bracts elliptic to elliptic-attenuate,
acuminate, exceeding to equaling the middle of the
sepals, 4— 12 X 3—6 mm, castaneous, entire, white-
lepidote. Mowers patent at anthesis, becoming sub-
erect afterward, pedicellate, pedicels cylindrical,
1-2 mm long, orange, sparsely lepidote, reaching 4
Novon 14: 168-170. 2004.
Volume 14, Number 2
2004
Forzza & Silva
Dyckia martinelli from Brazil
169
Figure 1. Dyckia martinellii B. R. Silva & Forzza. — A. Habit. — B. Leaf. — C. Spine. — 1). Scape base. — E.
Inflorescence. — F. Floral bract. — G. Flower. — H. Sepal. — I. Petal and stamens. — J. Gynoecium. — K. Ovule. — L.
Capsule. Drawings based on R. C. Forzza el al. 2386 (A— C), G. Martinelli 14413 (D-K), R. R. Silva 961 (L).
mm long in fructification; sepals ovate, apex round¬
ed, 7-9 X 4—6 mm, white-lepidote, orange with a
castaneous apex; petals orange, obtrullate, gla¬
brous. apex emarginate, erect at antbesis, 9—12 X
9 mm; stamens included; filament connate for 3.5
mm, adnate to petal for 2.5 mm; anthers slightly
sagittate at base, attenuate and reflexed at apex, ca.
3 mm long; ovary 7 mm long, ovule alate, ca. 0.7
170
Novon
Table 1 . Comparison
of Dyckia encholirioides. 1).
martinellii, and 1). pseudoococcinea.
1). encholirioides
D. martinellii
1). pseudococcinea
Vegetative propagation
rhizomatous
basal shoots
basal shoots
Rosette diameter (cm)
ca. I(M)
ca. 60
ca. 40
Foliar blades (cm)
30-100 X 2.5^1
20-35 X 2.2-2 .6
20-30 X 1-1.8
Foliar spines (mm)
antrorse, curved, 5-7
patent-antrorse, straight,
3-4
subfoliaceous, densely ar-
retrorse, curved, 1.5—2
Basal scape bracts
subfoliaceous, densely ar-
not foliaceous, sparse, in-
ranged, spinulose
ranged, spinulose
conspicuously serru¬
late
Scape bracts
longer than to rarely
longer than to rarely
much shorter than inter-
equaling the internodes
equaling the internodes
nodes
Anthesal flower position
reflexed
patent
patent
Floral bracts
ferrugineous-lanate, serru¬
late
white-lepidote, entire
reddish, serrulate
Sepals
ferrugineous-lanate
orange with eastaneous
apex, white-lepidote
reddish, glabrous
Petals
yellow, unguiculate
orange, obtrullate
orange, obtrullate
mm long. Fruits ca. 15 mm long; seeds alate, ea. 3
mm long.
Dyckia martinellii presents morphological affin¬
ities both toward I). pseudococcinea and D. encho-
lirioides (Gaudichaud-Beaupr6) Mez (Table 1),
these being t lie only three species known from
coastal southeastern Brazil.
Although the southern states of Rio Grande do
Sul. Santa Catarina, and Parana possess many Dy¬
ckia species, including seaside endemics, only one
species was known from the southeastern State of
Rio de Janeiro, D. pseudococcinea. Sao Paulo, an¬
other southeastern state of Brazil, is also relatively
poor in seaside species: only D. encholirioides is
known at present, with llha do Cardoso being its
northern limit of distribution.
The new species was found growing on granite
outcrops, from 3 to 10 m above the high tide line,
the rocks often being washed by rainwater. By com¬
parison, Dyckia pseudococcinea is a restricted en¬
demic from an area of periodically inundated Eri¬
caceae restinga in the municipality of Marica, Rio
de Janeiro State. Hie specimen cited by Smith and
Downs (1974) as originating in Atibaia, Sao Paulo,
actually belongs to another species, D. tuberosa
(Vellozo) Beer. The less selective D. encholirioides
occurs on rocks or sand along the coast from Rio
Grande do Sul to Sao Paulo, a completely subtrop¬
ical distribution (Smith & Downs, 1974).
This new taxon has only been collected from a
population on a granite outcrop with an inclination
angle of ca. 45°. The outcrop occupies an area of
approximately one hectare and is densely vegetated
with Vellozia Candida Mikan, Alcantarea edmundoi
(Leme) .1. R. Grant, Vriesea neoglutinosa Mez, Cat-
tleya intermedia Graham ex Hooker, Cyrtopodium
polyphyllum (Vellozo) L. C. Menezes, Cereus, Ti-
bouchina, and Pitcairnia. Well represented here,
Dyckia martinellii grows mainly in the shade of the
shrubs, the deepest shade being the place where
the largest specimens were found. This fact may
explain its late discovery since it renders the spec¬
imens almost invisible from passing boats, the only
means of transportation to the site. Many other sea¬
side outcrops with similar characteristics exist in
the region of Paraty and the nearby Ubatuba in Sao
Paulo and should be surveyed to establish the spe¬
cies' biological range.
Paratypes. BRAZIL. Rio de Janeiro: Mun. Paraty,
Paratymirim, costoes rochosos entre Paratymirim e Ma-
mangud, 5 m elev.. 30 May 1990 (fr), G. Martinelli 1208V
(RB): 23°14'06"S, 44°37'62"W, 17 Nov. 2002, fl cult. Dec.
2002, H. R. Silva 961 (RB), 5 Feb. 2003 (fr), R. C. Forzza
2386. R. R. Silva & E. Borges (MO. RB).
Acknowledgments. The authors are grateful to
Elton Leme and Claudio Nieoletti de Fraga for cor¬
rections and critical comments, and to Paulo Or-
mindo for preparing the illustration.
l.iterature Cited
Forzza, If. C. 2001. Filogenia da tribo Puyeae Wittm. e
revisao taxondmica do genero Encholirium Marl, ex
Schult. & Schult. f. (Pitcairnioideae— Bromeliaceae).
Tese de Doutorado, Instituto de Biociencias, Universi-
dade de Sao Paulo, Sao Paulo.
- . 2004 (in press). Revisao taxondmica de Encho¬
lirium Mart, ex Schult. & Schult. f. (Pitcairnioideae—
Bromeliaceae). Bol. Bot. Univ. Sao Paulo.
Smith, L. B. & R. J. Downs. 1974. Bromeliaceae (Pitcair¬
nioideae). FI. Neotrop. Monogr. 14(1): 1—662. Hafner
Press, New York.
A New Species of Polyalthia (Annonaceae) from China
Hou Xueliang
School of Life Sciences, Xiamen University, Xiamen 361005, China, and South China
Agriculture University, Guangzhou 510642, China
*Zi Shijin
South China Institute of Botany, the Chinese Academy of Sciences, Guangzhou 510650,
China, lisj@scib.ac.cn
Abstract. The new species Polyalthia zhui X. L.
flou & S. J. Li, found from Hainan and southern
Guangdong in southern China, is described and il¬
lustrated. It has previously been confused with Po¬
lyalthia nemoralis A. DC. or P. littoralis (Blume)
Boerlage in China, but is distinguished on the basis
of its green fleshy ovate petals, outer petals spread¬
ing ea. 4 X 2.5 mm, inner petals not fully spread¬
ing, somewhat bent to the inside, ea. 5—6 X 4—4.2
mm, 1 mm thick or more, its carpels pilose only in
the lower half, and its non-lobed stigma.
Key words: Annonaceae, China, Polyalthia.
Polyalthia Blume, with 150 to 160 species, is the
largest annonaceous genus in Asia (Van Heusden,
1992). Its species are mainly distributed in tropical
Asia. New Guinea, Australia, the Solomon Islands,
Fiji, Madagascar, and East Africa (Van Heusden.
1992; Verdcourt, 1971) and are especially numerous
in Southeast Asia (Sinclair, 1955). There are 32 spe¬
cies in Singapore (Sinclair, 1955). 27 in Vietnam
(Ban. 2000), 7 in Sri Lanka (Huber. 1985), and 8 in
Burma (Kurz. 1974). The northern distribution edge
of Polyalthia is in southern China; there are 17 spe¬
cies in Guangdong, Guangxi, Yunnan, and Hainan
Provinces (Tsiang & Li, 1979).
The systematics of Annonaceae from China is the
first author’s Ph.D. thesis, and the present paper is
a part of this study. In the last three years, nearly
all Annonaceae herbarium collections in China
(about 10,000 sheets) were checked, and several
field investigations were carried out concurrently.
At the South China Institute of Botany (IBSC),
we found that some specimens cited as Polyalthia
nemoralis A. DC. by Merrill and Chun (1940) were
heterogeneous with the others, i.e.. How 70423
(IBSC). Lau 2843 (IBSC), Liang 66027 (IBSC),
Liang 62116 (IBSC), Liang 62794 (IBSC). Liang
63904 (IBSC), Liang 66500 (IBSC). Tso & Chun
44581 (IBSC), Tso & Chun 44717 (IBSC). We tem¬
porarily regarded them as doubtful specimens for
lacking flowers, and we did not confirm they are
our new species, P zhui, until we had observed the
flowers of the Hainan Western Expedition 264
(IBSC). Its outer petals are ovate, 4 X 2.5 mm, its
inner petals are ovate, 5—6 X 4-4.2 mm, I mm
thick or more, pubescent outside, while P. nemor¬
alis petals are oblong, 8X4 mm, only 0.4 mm
Table 1 .
Morphologic
al comparison of Polyalthia nemoralis
and Polyalthia zhui.
Polyalthia nemoralis
Polylalthia zhui
Petals
White, di
ensely pubescent outer side, eharta-
Green, pubescent outer side, fleshy, 1 mm thick
ceous.
less than 0.4 mm thick, oblong; inner
or more, ovate-rounded; inner petals 5—6 mm
petals
8 mm long
long
Stamens
Connect i
ve apex convex, densely puberulent
Connective apex slightly concave, nearly gla-
brous
Ovaries
Densely
gray pubescent, stigma bifid
Pilose only in the lower half, stigma unlobed
Fruit walls
Coherent
with seeds
Separate from seeds
heaf blade
Secondar
y veins incurved; intersecondary veins
Secondary' veins straight; intersecondary veins 6
veins
absent
; veinlets unbranched; tracheids 0.3—
to 8 pairs; veinlets 1- or 2-branched; tra-
0.5 mm
cheids less than 0.1 mm
Leaf
Length :
width = 2.5—3 : 1
Length : width = 3-3.6 : 1
* Author for correspondence.
Novon 14: 171-175. 2004.
172
Novon
Figure la-c. Polyalthia nemoralis (from H. R. Liang 61573 , IBSC). — a. Flower. — b. Leaf venation. — c. Leaf
venation magnified, d— f. Polyalthia zhui X. L. Hou & S. J. Li (from the holotype, Hainan Western Expedition 264 ,
IBSC). — d. Flower. — e. Leaf venation. — f. Leaf venation magnified. Scale bars: a, c, d & f = 1 mm; b & e = 1 cm.
Volume 14, Number 2
2004
Hou & Li
Polyalthia zhui from China
173
Table 2. Morphological comparison
of Polyalthia littoralis and Polyalthia
zhui.
Polyalthia littoralis
Polyalthia zhui
Ratio of inner petal length to
2-2.6 : 1
1 .4—1.6 : 1
outer petal length
Petal texture
thin-membranous
fleshy, 1 mm thick or more
Ratio of monocarpel length to
2-3.1 : 1
4-5 : 1
the stalk of monocarpel length
Length of monocarpel stalk
3—6 mm
3 mm
Length ol fruiting peduncles
ca. 12 mm
3—1 mm
Leaf blade
lanceolate, usually 2—3 cm wide
oblong-lanceolate or ovate-lan¬
ceolate, usually 3. 5-4. 2 cm
wide
thick, and pilose outside. Further observation
shows that these specimens also differ from P. ne-
moralis in carpels, fruit, stamens, and leaf venation
(Table 1, Fig. 1).
After having compared the specimens with the
known species of Polyalthia recorded in various
floras and systematic treatments (Blume, 1830;
Hooker & Thomson, 1855, 1872; Merrill, 1912,
1923; Sinclair, 1955; Backer & Bakhuizen, 1963;
Huber, 1985; Ban, 2000), we found a few species
with petals less than 10 mm long. Among these.
Figure 2. Holotype of Polyalthia zhui ( Hainan Western
Expedition 264. I BSC).
only P. littoralis (Blume) Boerlage, whose distri¬
bution extends to Indonesia and Vietnam, is similar
to P. zhui. We made sure, however, that P. zhui was
distinct from P. littoralis. Polyalthia littoralis petals
are thin-membranous, whereas P. zhui petals are
thick-fleshy up to 1 mm or more. Furthermore, the
two species can be distinguished by their leaves,
monocarpels, and fruiting peduncles (Table 2). Po¬
lyalthia littoralis was reported as new to China by
Li (1993). According to the specimens cited by Li
(1993), however, his P. littoralis was Merrill and
Chun’s P. nemoralis, that is, it included P. nemor-
alis and P. zhui.
Polyalthia zhui X. L. Hou & S. J. Li. sp. nov.
TYPE: China. Hainan: Darizhou City, Nada,
Lianchangjiaoyuan, 15 May 1954, Hainan
Western Expedition 264 (holotype, 1BSC; iso¬
types. IBK. KUN, PE). Figures ID— F, 2. 3.
Affinis Polyalthiae nemorali, sed floribus ca. 6 mm
diam., petalis exterioribus ovato-rotundis 4 mm longis, 2.5
mm latis, petalis interioribus ovato-rotundis, 5—6 mm lon¬
gis, 4—4.2 mm latis, externis rare pubescentibus, carpellis
meris pubescentibus in dimidio inferiore, stigmate non
lobato, connectivo parum concavo differt.
Shrubs usually 1.5—3 m, rarely up to 5 m tall;
bark dark brown; young twigs dark purple, striate,
pubescent with appressed gray-yellow hairs, but
quickly glabrescent. Leaves coriaceous, oblong-
lanceolate or ovate-lanceolate, 9-16(— 19) X 2.5-
4.5(— 5) cm, apex acuminate to obtuse-acuminate,
base cuneate, glabrous, dark green above, lucid
and pale green beneath, midrib slightly grooved
above and obviously convex beneath, secondary
veins 8 to 10 pairs, at 60° to 80° with the midrib,
interarching 3—5 mm from the margin, reticulations
somewhat prominent on both sides, intersecondary
veins in 6 to 8 pairs; petioles 3—5 mm long, yel¬
lowish, rugose. Inflorescences extra-axillary with 1
(or 2) flowers; peduncles ca. 1 mm long; pedicels
174
No von
F igure .'1. Polyallhia zhui. — a. Abaxial view of stamen. — b. Adaxial view of stamen. — c. Carpel. — d. Longitudinal
section of carpel. Scale bar = 1 mm. (From the holotype, Hainan Western Expedition 264 , IBSC.)
1—2 mm long, pubescent with gray-yellow hairs,
bract 1. broadly ovate, 1-2 X 2 mm. acuminate.
Flower green, ea. 6 mm diam. when mature; sepals
connate at base, lobes 3, ovate, 2 X 1 .8 mm, pu¬
bescent outside, glabrous inside, persistent; petals
6 in 2 whorls, fleshy, acute, pubescent outside, gla¬
brous inside, outer petals ovate, flat, spreading, ca.
4 X 2.5 mm, inner petals ovate, flat, not fully
spreading, somewhat bent to inside, 5—6 X 4-4.2
mm, ca. 1 mm thick or more; stamens ca. 50 to
100, cuneate, 1—1.3 mm long, ca. 1 mm wide, con¬
nective apex slightly concave, dark purple, gla-
brescent; carpels ca. 30. with one basal ovule each.
1.5—2 mm long, pilose with brown-yellow hairs in
the lower half; stigma capitate, sessile, puberulent,
not lobed. Monocarpels 4 to 7 (to 11), ellipsoid, 12—
15 mm long, 8—10 mm diam.. purple-red when dry,
glabrous, densely tiny-dotted, stalks 3 mm long,
pubescent; seed I per loeule, ellipsoid, red, 10—12
mm long, 6—8 mm diam., separated from fruit walls,
endosperm ruminative.
Distribution and habitat. In wet forest at 250—
600 m, Hainan and southern Guangdong. In forest
edges, sometimes along small streams. This was
collected in flower April through July and in fruit
July through February of the following year.
Polyalthia zhui is distinguished from the other
species of Polyalthia by its oblong-lanceolate or
ovate-lanceolate leaf shape, lucid anti pale green
beneath and 6 to 8 pairs of intersecondary veins,
its relatively short pedicels and small green flowers
with fleshy petals, its nearly glabrous and slightly
concave connective apices, its capitate stigma, and
its seeds not adherent to fruit walls.
Eponymy. Polyalthia zhui was named in honor
of our master supervisor. Prof. Zhu Changshan,
working in the Henan Agriculture University, who
shared his rich knowledge of and correct methods
for studying plant taxonomy.
Paratypes. CHINA. Guangdong: Xuwen Co., Mailao-
wo village, 31 Oct. 1951 (fr), Z. S. Zhu I0,'i6 (IBSC);
Xuwen Co., Kengzai village, 26 Apr. 1936 (fr), Hainan
Southern Expedition 291 (IBK, IBSC, KUN, PE); Xuwen
Co., Xuwenshan hill, near Yugonglou, 27 Sep. 1951 (fr),
S. H. Chun 7515 (IBSC): Xuwen Co., Guooheng village,
7 Nov. 1957 (fr), Hainan Southern Expedition 152 (IBSC);
Xuwen Co., Baishui village, 15 Apr. 1985 (II). Li & Xing
1985 (IBSC). Hainan: Baisha Co., a valley near Hong-
rnaodong, 27 Apr. 1936, S. A. Diu 26524 (IBSC. KUN);
Baoting Co., Diaoluosban Mountain. Baishuiling. 24 Nov.
1954, Diaoluoshan Expedition 2815 (IBK, IBSC, PE);
Cbangjiang Co., Jiaqieshan bill. 16 Oct. 1933 (fr), S. A.
Dm 2842 (A. IBSC); Cbangjiang Co., 20 Dec. 1933 (fr),
II. II. Liang 66027 (A, IBK. IBSC); Cbangjiang Co., Dec.
1933 (fr). II. R. Liang 60102 ( \. IBK. PE); Cbangjiang
Co.. 20 Dec. 1933 (fr), II. R. Liang 66500 (IBK. IBSC,
PE); Dongfang Co., Guangba District, Qicha, 17 Nov.
1956, S. II. Chun II 124 (IBSC); Dongfang Co.. Maanling,
6 Nov. 1978. G. A. Eu 1291 (IBSC); Dongfang Co., Jian-
fengling, near Sanmojue village, 23 Mar. 1934, 6. A. Dm
2522 (SYS). .S'. A. Dm 4924 (IBSC); Dongfang Co., //. R.
Liang 62094 (A, IBK. IBSC); Ledong Co., Taohui village.
Taohuiling, S’. A. Dm 27454 (A. IBK. IBSC. KUN, PE);
Eingshui Co., Nanwanling, 21 Jan. 1985, C. A. Eu 5551
(IBSC); Sanya City, Nanshanling, 30 Dec. 1932, Tso S:
Chun 44717 (A, IBK, IBSC, PE); Sanya City. Nansbanl-
ing. 26 Dec. 1932. Tso A- Chun 44581 (A. IBK. IBSC,
Volume 14, Number 2
2004
Hou & Li
Polyalthia zhui from China
175
PE); Sanya Cily, Xiaobaokang, 16 July 1933 (fr), H. R.
Liang 62116 (A. IBK. IBSC. PE); Sanya City, Daposhan
hill, 23 Aug. 1933, H. R. Liang 62794 (A, IBK. IBSC);
Sanya City, Yanglin, 24 Mar. 1933, F. C. How 70423
(IBSC); Sanya Cily, Ganzhaling, upriver Fotian Reservoir,
30 Oct. 1987, Z. X. Li 3058 (IBSC); Sanya City. Dabao-
kang, 16 July 1933, C. Wang 33087 (IBSC, PE); Sanya
City, Nanlin, Guikaishan hill, 16 Oct. 1933 (fr), C. Wang
34633 (IBSC).
Acknowledgments. The authors thank Li Ping-
tao. South China Agriculture University, for criti¬
cally reading the manuscript and giving valuable
suggestions. We are likewise grateful to Zhang
Dianxiang (IBSC) for helpful comments on the man¬
uscript. We are also grateful to Zeng Feiyan (IBSC),
Wang Jinghua (KLIN), Lin Qi and Ban Qin (PE), Li
Mingguang (SYS), and Wei Fanan and Yu Xixi
(IBK) for offering assistance in consulting speci¬
mens. fhe study was supported by a grant from the
Bureau of Environmental Protection of Guangdong
Province (970165).
Literature Cited
Backer, C. A. & K. C. Bakhuizen van den Brink. 1963.
Pp. 100—116 in Flora of Java, Vol. 1. Walters Noordhoff
N.V., Groningen.
Ban. IN. T. 2000. Flora of Vietnam, Vol. 1. Science &
Technics Publishing House, Ha Noi.
Illume, C. L. 1830. P. 99 in Flora Javae, Vol. 1. Fugduxi
Bataviorum.
Hooker, J. D. & T. Thomson. 1855. Annonaceae. Pp. 86-
153 in Flora Indica, Vol. 1. W. Pamplin, London.
- & - . 1872. Annonaceae. Pp. 45—94 in J.
1). Hooker (editor), The Flora of British India I. Reeve,
London.
Huber. H. 1985. Annonaceae. Pp. 1—75 in M. I). Dassan-
ayake (editor), A Revised Handbook to the Flora of
Ceylon, Vol. 5. A. A. Balkema, Rotterdam.
Kurz, S. 1974. Annonaceae. Pp. 25—50 in Forest Flora of
British Burma. Vol. 1. Bishen Singh Mahendra Pal
Singh. Dehra Dun.
Li, P T. 1993. Novelties in Annonaceae from Asia. Gui-
haia 13(4): 311-315.
Merrill E. D. 1912. Annonaceae. Pp. 205—209 in A Flora
of Manila. Bureau of Printing, Manila.
- . 1923. Annonaceae. Pp. 154—178 in An Enumer¬
ation of Philippine Flowering Plants II. Bureau of Print¬
ing, Manila.
- & W. Y. Chun. 1940. FI. Hainan III. Sunyatsenia
5(1-3): 59.
Sinclair, J. 1955. A revision of the Malayan Annonaceae.
Card. Bull. Straits Settl. 14: 149—516.
Tsiang, Y & P. T. Li. 1979. Annonaceae. Pp. 10—175 in
Flora Reipublicae Popularis Sinicae 30(2). Science
Press, Beijing.
Van Heusden, E. C. H. 1992. Flowers of Annonaceae:
Morphology, classification, and evolution. Blumea
Suppl. 7: 1-218.
Verdcourt, B. 1971. Annonaceae. Pp. 1—132 in E. Milne-
Redhead & R. M. Polhill (editors), Flora of Tropical
Fast Africa. Crown Agents for Oversea Governments
and Administrations, London.
A New Species of Didymoplexiella (Orchidaceae) from China
Jin Xiaohua, Chen Singchi, and Qin Haining
Herbarium (PE), Institute of Botany of Chinese Academy of Sciences, Beijing 100093,
China, xiaohuajin@mail.kib.ac.cn; hainingqin@ns.ibcas.ac.cn
Guanghua /hit
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
guanghua.zhu@mobot.org
Gloria Siu Laiping
Kadoorie Farm and Botanic Garden, Hong Kong. siulpcon@kfbg.org.hk
Abstract. Didymoplexiella hainanensis X. H. Jin
& S. C. Chen, a new speeies of Orchidaceae from
Hainan, China, is described and illustrated. The new
species is related to I). ornata (Ridley) Garay, from
which it differs by the dorsal sepal briefly connate
with the petals, the lateral sepals connate Va to Vi at
base, an articulate lip, and larger lateral lobes.
Key worth: China, Didymoplexiella, Orchidaceae.
Didymoplexiella Garay is a leafless orchid genus
of ca. seven species (Seidenfaden & Wood, 1992)
mainly distributed in Indonesia, with a few species,
e.g., D. siamensis (Rolfe ex Downie) Seidenfaden,
extending to northern Thailand, China, and Japan.
It is similar in habit and floral appearance to Di-
dymoplexis Griffith, with which it was considered
by some authors as congeneric (Holttum, 1953).
Garay (1955) found enough distinction in floral col¬
umn to justify a separation between them. The col¬
umn of Didymoplexiella has a pair of long stelidia
on either side of its stigma and lacks a distinct
column foot, while that of Didymoplexis lacks stel¬
idia and the column foot is distinct. The new spe¬
cies was collected in late March from Hainan Is¬
land and is no doubt a new member of
Didymoplexiella. It is somewhat related to D. or¬
nata (Ridley) Garay from Malaysia and Thailand,
from which the new species is distinct by its almost
separate dorsal sepal and petals, articulate lip, and
much larger lateral lobes.
Didymoplexiella hainanensis X. 11. Jin & S. C.
Chen, sp. nov. TYPE: China. Hainan: Ledong,
Jianfenlin Mt., 17 Mar. 2002 (fl). //. K. Ka¬
doorie Program Team 3087 (holotype, PE; iso-
type [spirit], MO). Figure 1.
Habitu Didymoplexiellae ornatae subsimilis, sed sepalo
dorsali petalis breviter vel vix connato, sepalis lateralibus
inter se tantum Va ad Vi parte inferiore connatis, labello
multo longiore quam columna prope medium mobili-arti-
culato bene differt.
Mycoheterotrophic plants; tuber creeping, cylin¬
drical, 5—6 cm long, 3—10 mm diam., wrinkled,
nodal; stem erect, slender, terete, 5—15 cm long,
brown, nodal, sheathing on the lower half, with in¬
ternodes 2-9 mm long; sheaths 1 .5—2.5 mm long.
Inflorescence 7-9 mm long, compactly 2- to 4-flow-
ered; bracts triangular-lanceolate, 1.5—2 mm long;
petlicel and ovary 5—15 mm long, brown. Flowers
ca. 1.5 cm across, white except the lip mid-lobe
yellow, dorsal sepal elliptic, 7-10 X 13 mm, briefly
or scarcely united at base with petals, 3-nerved,
obtuse; petals similar to dorsal sepal. 6—7 X 10
mm, joined with dorsal sepals for Va to Vi of their
length; lateral sepals elliptic, 10—11 X 3.5-4 nun,
3-nerved, obtuse, connate at base to each other for
Va to Vi of the length; lip 11-12 mm long, consisting
of an apical blade 5.5—6 mm long and a basal claw
4.5—5 mm long, with an articulate joint between
them; blade tri-lobed, with a basal claw, its upper
surface a U-shaped callus covered with small warts;
lateral lobes sub-oblong, ca. 5X4 mm with a
slightly undulate margin, much larger than the mid¬
lobe; mid-lobe fleshy, ca. 3 X 2 mm at the apex,
with stelidia on either side of the stigma; stelidia
curved downward, 2.5—3 mm long; pollinia 4,
mealy. Capsule not seen.
Distribution. China, Hainan Province, Ledong
County, Jianfengling Mountains, alt. 700 to 800 m.
in humid evergreen forest.
Paratype. CHINA. Hainan: Ledong, Jianfengling
Mtn., 15 Mar. 2002 (fl), Hong Kong Kadoorie Program
Team 3081 (MO. |spirit| KUN).
Acknowledgments. We are indebted to the Ka-
Novon 14: 176-177. 2004.
Volume 14, Number 2
2004
Jin et al.
Didymoplexiella (Orchidaceae)
177
Figure 1. Didymoplexiella hainanensis X. H. Jin & S. C. Chen. — A. Habit and inflorescence. — B. Front view of
sepals and petals. — C. Front view of lip. — D. Stelids. — E. The articulate claw. ( Based on Hong Kong Kadoorie
Program Team 3087, PE.)
doorie Farm and Botanic Garden, Hong Kong, for
its financial support and to the CITES officials of
China for their kind help. We also thank Yinbao
Sun for his excellent illustration.
Literature Cited
Garay, L. A. 1955. Notatio Orchidologica 3. Arq. Jard.
Got. Kio. 13: 31-54.
Holttum, R. E. 1953. Didymoplexis. P. 107 in R. E. Holt-
tum (editor). Flora of Malaya. Vol. 1. Orchids of Malaya.
Government Printing Office, Singapore.
Seidenfaden, G. & J. J. Wood. 1992. Didymoplexiella. Pp.
127—139 in G. Seidenfaden & J. J. Wood (editors). The
Orchids of Peninsular Malaysia and Singapore. Olsen
& Olsen, Fredensborg.
A New Species of Holcoglossum (Orchidaceae) from China
Jin Xiaohua, Chen Singchi, and (Jin Haining
Herbarium (PH). Institute of Botany of Chinese Academy of Sciences, Beijing 100093,
China, xiaohuajin@mail.kib.ac.cn; hainingqin@ns.ihcas.ac.cn
Guanghua Zhu
Missouri Botanical Garden, P.0. Box 299. St. Louis, Missouri 63166, U.S.A.
guanghua.zhu@mobot.org
Gloria Sin Lai ping
Kadoorie Farm and Botanic Garden, Hong Kong. siulpcon@klBg.org.hk
ABSTRACT. Holcoglossum weixiense X. H. Jin &
S. C. Chen, a new species of Orchidaceae from
Yunnan, China, is described and illustrated. It re¬
sembles morphologically H. sinicum Christenson
and //. flavescens, but it can be distinguished from
the former by having a transverse callus on the lip,
a papillose spur, and a semi-orbicular mid-lobe,
and from the latter by its pending habit, longer and
semiterete leaves (more than 10 cm), and the semi-
orbicular mid-lobe.
Key words: China, Holcoglossum, Orchidaceae.
Holcoglossum Schlechter (Orchidaceae) is dis¬
tributed mainly in southwestern China, with a few
species, e.g., H. subulifolium (Reichenbach f.)
Christenson, extending to Laos, Thailand. Vietnam,
and Malaysia. Because of its pollinia, three-lobed
lip, and the short bifid rostellum, this genus is sim¬
ilar in floral structure to Aerules, Papilionanthe, and
Vanda, from which several species have previously
been transferred, e.g., H. rupestre (Handel-Mazzetti)
Garay (1972), H. flavescens (Schlechter) Tsi (1982).
and H. subulifolium (Reichenbach f.) Christenson
(1987). Although Seidenfaden (1988) stated that it
was difficult to define Holcoglossum, the new spe¬
cies somewhat coincides with Garay’s (1972: 181)
delimitation of the genus: “. . . characterized by the
short, footless column with prominent wings. Cli-
nandrium deeply cleft in front: rostellum short, bi¬
fid. Pollinia 2, notched, on linear, tapering stipes.
Lip sessile with a slender, arcuate spur. Stem rather
short, completely enclosed by distichously arranged
leaf-sheaths. Leaves articulate, triquetrous or te¬
rete."
The new species was collected from the Langcan
River in northwestern Yunnan. It is somewhat re¬
lated to Holcoglossum sinicum Christenson and H.
flavescens (Schlechter) Tsi, but is distinguished from
the former by its semi-orbicular mid-lobe, papillose
spur, notable callus on the labellum, and from the
latter by its pending habit, longer and semiterete
leaves (more than 10 cm), and the semi-orbicular
mid-lobe.
Holcoglossum weixiense X. H. Jin & S. C. Chen,
sp. nov. TYPE: China. Yunnan: Weixi, La¬
ngcan River, 21 May 2002 (fl), //. K. Kadoorie
Program Team 3490 (holotype, PE; isotypes
[spirit], KUN. MO, PE). Figure 1.
Habitu Holcoglosso sinico subsimile, sed calcari intus
papillato, inflorescentia caule longiore, labello medio callo
nolabili instructo bene differ!.
Epiphytic plant, pendulous; root terete, thick,
white; stem short, 1—2 cm long, enclosed by the
leaf-sheathing bases. Leaves 20—30 X 1.5— 2.5 cm,
coriaceous, semiterete, channeled, articulate at
base. Inflorescence lateral, erect, 3-4 cm long, 2-
to 4-flowered, exceeding the stem, bracts ovate, 0.7
cm long. Flowers spreading, opening fully, white,
tinged with pink; pedicel with ovary 2 cm long;
dorsal sepal erect, ovate, 1 X 0.7 cm, 3-veined;
lateral sepals oblique at base, seculate, 1.5 X 0.9
cm wide, 4-veined; petals spathulate, 1.1 X 0.0 cm,
3-veined; lip adnate to the column foot, fixed;
three-lobed; lateral lobes triangular, erect, with
purple streaks on the inner surface; mid-lobe semi-
orbicular, entire, 1 X 1.2 cm. with a transverse
callus at base toward the entrance of the spur; spur
funnel-shaped, papillose, 0.7 cm long; column 0.4
cm long, column foot 0.1 cm long; pollinia 2,
notched; stipes obcuneate, 3 mm long. Capsule un¬
known.
Distribution. China: Yunnan, Weixi, Bilou
Snow Mountain, at 2300 to 2900 m. in evergreen
forest.
Novon 14: 178-179. 2004.
Volume 14, Number 2
2004
Jin et al.
Holcoglossum (Orchidaceae)
179
Figure 1. Holcoglossum weixiense X. II. Jin & S. C. Chen. — A. Habit and inflorescence. — B. Lateral view of flower.
— C. Front view of flower. — L). Lateral view ol lateral lobes. — E. Inner wall of spur. Drawn from the type specimen
(Hong Kong Kadoorie Program Team 3490) by Yingbao Sun.
Paratypes. CHINA. Yunnan: Langcan River, 20 July
2001, Hong Kong Kadoorie Program Team 1166 (KUN,
PE), 21 May 2002 (fl), Hong Kong Kadoorie Program
learn 3436 (KUN), 3497 (PE).
Acknowledgments. We are indebted to Wen-tasi
Wang for his critical reading of the manuscript, to
the Kadoorie Farm and Botanic Garden, Hong
Kong, for its financial support, and to the CITES
officials of China for their kind help. We also thank
Yinbao Sun for his excellent illustration.
Literature Cited
Christenson, E. A. 1987. An intrageneric classification of
Holcoglossum Schltr. (Orchidaceae: Sarcanthinae) with
a key to the genera of the Aerides-Vanda alliance. Notes
Roy. But. Card. Edinburgh 44: 249—256.
Garay, L. A. 1972. On the systematics of the monopodial
orchids I. But. Mus. Leaf!. 23(4): 149-212.
Seidenfaden, G. 1988. Orchid genera in Thailand XIV.
Fifty-nine genera. Opera Bot. 95: 304—308.
Tsi, Z. H. 1982. A study of the genus Holcoglossum of
Orchidaceae. Acta Phytotax. Sin. 20(4): 439—444.
New Taxa of Cimicifuga (Ranunculaceae) from Korea and the
United Stales
Hyun-Woo I^ee
El A Division, Korea Environment Institute, Seoul 122-706, Korea. hwlee@kei.re.kr
Chong-Wook Park
School of Biological Sciences, Seoul National University, Seoul 151-742. Korea.
parkc@plaza.snu.ac.kr
Abstract. A new species and a new variety of
Cimicifuga (Ranunculaceae) are described and il¬
lustrated. Cimicifuga austrokoreana, restricted to
south-central Korea, is distinguished from the other
species of the genus by its strongly arcuate inflo¬
rescence axis, much shorter pedicels, and three
bracteoles, one at the base and the other two at the
middle of the pedicel. Cimicifuga data var. alpes-
tris, found in southern Oregon in the United States,
is distinguished from variety data and the other
members of the genus by its sheathing scales sur¬
rounding the lower nodes of the stem.
Key words: Cimicifuga, Korea, Ranunculaceae,
United States.
The genus Cimicifuga Wernischeck (Ranuncu¬
laceae) comprises as many as 22 species that are
widely distributed in temperate regions of the
Northern Hemisphere, including Europe, Asia, and
North America; the center of diversity is in eastern
Asia (Tamura, 1966, 1990, 1995; Ramsey, 1965,
1997; Compton et al., 1998). The plants of Cimi¬
cifuga are erect rhizomatous herbs with long-peti-
oled, ternately compound leaves, racemose or pa¬
niculate inflorescences bearing many small flowers,
and follicular fruits (Tamura, 1966, 1990, 1995;
Park & Lee, 1995, 1996; Ramsey, 1965, 1997).
Classification of Cimicifuga and its related gen¬
era has been controversial. Compton et al. (1998)
merged Cimicifuga and Souliea Franchet with Ac-
taea L. mainly on the basis of the analyses of ITS
and /rnL-F sequences. However, Wang et al. (1999,
2001) argued that they are independent genera
based on characters from morphology, palynology,
and cytology. Cimicifuga is distinguished from Ac-
taea in having three (rarely one) bracteoles, five
sepals, staminode-like petals, one to eight pistils,
follicular fruits, seeds with membranous scales on
the surface, and more symmetric karyotypes with¬
out telocentric chromosomes. Cimicifuga is also
distinguished from Souliea in having inflorescences
with numerous flowers, caducous sepals, oblong to
ovoid follicles, seeds with membranous scales on
the surface, and mostly tricolpate pollen grains. In
diis study we follow the generic concept and delim¬
itation of Tamura (1995) and Wang et al. (1999,
2001), in which Cimicifuga was recognized as a
distinct genus.
During the course of a comprehensive systematic
study on the genus Cimicifuga, a new species and
a new variety were discovered from Korea and the
United States, respectively. We describe these new
taxa herein to make their names available for the
Flora of Korea and other works in progress.
1. (.imicifiigu austrokoreana H.-W. Lee & C.-
W. Park, sp. nov. TYPE: Korea. Chungbuk:
Youngdong-gun, Mt. Minjuji-san, Samdo-bong,
Minimigol, 700 m, 19 Sep. 2000, H.-W. Ise
1542 (holotype, SNU; isotypes, MO, SNU, TI).
Figure 1.
Differt a C. simplici inflorescentiis axibus arcuatis, ped-
icellis multo brevibus et unibracteolatis basi.
Herbs perennial, hermaphroditic, 40—80 cm tall;
rhizomes thick, 5—8 cm long, knotted, bearing fi¬
brous roots; stem simple, erect. 1.5-4 mm thick,
glabrous to sparsely pubescent with filiform uni¬
cellular trichomes. Leaves cauline, usually 3, al¬
ternate, 1- to 3-ternately compound, long-petioled;
terminal leaflet broadly elliptic, 7.0—10.2 X 5.4—
7. 1 cm, 3-lobed, acuminate to cuspidate at tip, cu-
ueate at base; both surfaces moderately to densely
pubescent along major veins with filiform unicel¬
lular trichomes ca. 0.2 mm long; petiolules 1.5— 2.5
cm long, grooved, sparsely to moderately pubescent
with filiform unicellular trichomes; lateral leaflets
similar to terminal one, but slightly smaller and
inequilateral; petiole 10—20 cm long, grooved,
sparsely to moderately pubescent with filiform uni¬
cellular trichomes. Inflorescence a terminal ra¬
ceme, 10—20 cm long, often bearing 1 or 2 short
Novon 14: 180-184. 2004.
Volume 14, Number 2
2004
Lee & Park
New Taxa of Cimicifuga
181
Figure 1. Cimicifuga austrokoreana H.-W. Lee & C.-W. Park. — A. Inflorescence anil upper leaf. — B. Stem with
lower leaf. — C. Petals. — 1). Pistils and a pedicel with bracteoles. — E. Follicles. — F. Seed. Scale bars 5 cm (A, B),
5 mm (E), 2 mm (D), 1 mm (C, F). A-D, drawn from the holotype (Lee 1542, SNU), and E, F from the paratype (Oh
1355, SNU), by H.-W. Lee.
182
Novon
lateral branches near base, densely pubescent with
saccate unicellular trichomes 0.1— 0.2 mm long; in¬
florescence axis distinctly arcuate; pedicels very
short, 1.0— 2.5 mm long; bracteoles 3, 1 at base and
2 at middle of pedicel, triangular, 0.4— 0.7 X 0.3—
0.5 mm. Flowers bisexual, actinomorphic, small;
sepals 5, petaloid, pale brown, broadly elliptic,
concave, 3.5— 4.8 X 2.8— 3.5 mm; petal I. broadly
elliptic, 2. 0-4.6 X 1.4— 3.3 mm, bearing 2 small
white appendages at tip, short-stipitate, nectarifer¬
ous at base; stamens 18 to 27; filaments filiform,
5—8 mm long, slightly dilated upward; anthers bi¬
locular, broadly oblong, 0.6 X 0.5 mm. basifixed,
latrorsely dehiscent; pistils usually 2. rarely 1 or 3,
elliptic to broadly elliptic, 1-3 X 0.7-2 mm, stip-
itate, sparsely pubescent with saccate unicellular
trichomes ca. 0.1 mm long; style 1. slender, unci¬
nate when mature, 0.8-1. 7 mm long; stigma I. mi¬
nute, appressed to style. Follicles oblong, 5.5—9 X
1—6 mm, chartaceous, glabrous or sparsely pubes¬
cent with saccate unicellular trichomes; stipe 4—6
mm long; seeds 1 to 5 per follicle, elliptic, 2.2— 2.6
X 1 mm, winged; wings ca. 1 mm long, chaffy,
crenate.
Distribution. Restricted to high mountains of
south-central Korea; moist, shady places along
streamsides and mountain slopes of mixed decid¬
uous forests, alt. 500 to 1000 in.
Phenology. Flowering mid-September to Octo¬
ber (pers. obs.).
Cimicifuga austrokoreana is closely related to C.
simplex (DC.) Turczaninow, but it clearly differs
from the latter species by its strongly arcuate inflo¬
rescence axis, much shorter pedicels ca. 1.0— 2.5
mm long, and three bracteoles, one at base and the
other two at middle of pedicel (Fig. 1); in particular,
the inflorescence axis is distinctly and consistently
bent downward from the early stage of its devel¬
opment. Cimicifuga simplex has an erect, straight
inflorescence axis, relatively long pedicels ca. 4-8
mm long, and bracteoles borne at the base of the
pedicel.
In addition, the karyotypic study and allozyme
analysis showed that Cimicifuga austrokoreana is
genetically distinct from C. simplex (Fee & Park,
1998; Fee et al., 2000). Their karyotypes differ in
the position of a secondary constriction on a pair
of submetacentric chromosomes (Fee & Park.
1998), and populations of C. austrokoreana are dis¬
tinguished from those of C. simplex by their allelic
compositions at four loci ( Fe-3 , Gdh , Lap, anil Pgi-
2) and significantly low genetic identity values
(mean = 0.688) (Fee et al., 2000).
Paratypes. KOREA. Chiinhuk: Muju-gun, Mt. l)u-
kyu-san, Baekryundam, 16 Sep. 1996, Lee 771 (SIN LJ ).
Chuiighuk: Youngdong-gun, Mt. Minjuji-san, Samdo-
bong, 29 Sep. 1995, lee 747, 748 (SNU), 8 Oct. 1995,
lee 775, 776 (SNU). Chiinnain: Gurye-gun, Mt. Cliiri,
Piagol, Sunyugyo, under deciduous forest, 24 Sep. 1990,
//vim 5012 (SNU); Jilmaejae, 6 July 1995, lee 566, 567
(SNU), 18 Sep. 2(KM). lee 1541 (SNU). 30 Sep. 2. g, h from Lorence
& Stone 8380 ; e— f, from Lorence & Stone 8385. Scale bar 10 cm in b, c; 7 mm in d, e; 6 mm in f; 7.5 cm in g; 2.5
cm in h.
Volume 14, Number 2
2004
Lorence & Gemmill
Pritchardia flynnii from Kaua'i
187
felt-like, pale brown or gray. Inflorescences 1 or
occasionally 2 per leaf axil, in flower erect or ar¬
cuate, 58—88 cm long, equaling or generally ex¬
ceeding petiole and reaching Vi (rarely %) length of
blade, in fruit 80—100 cm long, erect to arcuate and
slightly shorter to slightly longer than the blade;
prophyll externally brown or tan lepidote, tomen-
tose basally, disintegrating; peduncular bracts 4,
overlapping, eventually disintegrating, externally
uniformly brown or tan tomentose, denser basally;
peduncle 48—60 cm long, 8—1 1 mm diam., terete
or compressed, brown lanate-tomentose; panicle
branched to second degree, the floriferous portion
1 1-20 cm long, the basal branches 10—13 cm long
with 3 or 4 rachillae, the middle branches with a
single bifurcate branch of 2 rachillae or less com¬
monly unbranched, apical portion of rachis with 9
to I I unbranched rachillae, these sinuous in flower,
6—11 cm long, at first sparsely to densely lanate-
tomentose, in fruit 10—14 cm long, tomentose or
glabrescent; floral bracts subulate-filiform, 1 .5—3
mm, brown. Flowers spirally arranged, 8—10 mm
long in bud, glabrous, calyx plus pedicel 3—5 mm
long, calyx cupular, 1.5— 2.5 mm wide at base, 3—4
mm wide at rim of cup, green when fresh, when dry
with prominulous nerves converging in abruptly
acuminate teeth 0.3— 0.8 mm long; petals 5.7-7 X
2.8— 3.5 mm, acute to acuminate at apex, yellow
when fresh, veins prominulous when dry; staminal
cup 3.5—6 mm long, exserted beyond calyx rim.
deep orange-yellow when fresh; filaments 1—2 mm
long, anthers 3—5 mm long; pistil 5—6.5 mm long,
including style 3-4 mm long; style present in fruit,
ca. 2 mm long. Fruit ellipsoid or ovoid-ellipsoid,
25—35 X 18—23 mm when fresh, 22—25 X 15—16
mm when dry, often slightly asymmetrical or acen¬
tric, with persistent calyx and corolla at base and
accrescent style at apex, smooth, glabrous, shiny
dark green, ripening purplish black; seed ovoid-
ellipsoid, 14—18 X 9—12 mm.
Distribution. Known from south-central to
north-central Kaua'i. Populations are known from
the Wahiawa Drainage (Wahiawa Stream and Moun¬
tains and Mt. Kahili) in the south, the north fork of
the Wailua River (“Blue Hole”), Makaleha Moun¬
tains, Power Line Trail, and Wainiha Valley in the
north. Intervening areas likely harbor additional
plants, in effect linking these known populations.
Habitat. This new species occurs from approx¬
imately 488 to 890 m elevation, usually on mod¬
erate to steep slopes in low-stature lowland wet for¬
est. shrubland, or herbland usually dominated by
Metrosideros polymorpha Gaudichaud, M. waiale-
alae (Rock) Rock var. waialealae, and Dicranop-
teris linearis (Burmann) Underwood. Other associ¬
ates include Antidesma platyphyllum II. Mann var.
hillebrandii Pax & K. Hoffmann, Bobea elatior
Gaudichaud, Diospyros sandwicensis (A. DC.) Fos-
berg, Freycinetia arborea Gaudichaud, Ilex anom-
ala Hooker & Arnott, Machaerina angustifolia
(Gaudichaud) T. Koyama, Melicope, Syzygium sand¬
wicensis (A. Gray) Niedenzu, and species of Cibo-
tium, Cyanea, Cyrtandra, Myrsine, Psychotria,
Scaevola, and Tetraplasandra. Weedy alien species
invading this habitat include Psidium cattleianum
Sabine var. cattleianum and Melastoma candidum
D. Don, both of which pose a significant threat to
native plant communities in the vicinity (Lorence
& Flynn, unpublished), and a variety of alien grass¬
es including Paspalum conjugatum Bergius, Setaria
gracilis Kunth, and Schizachyrium condensatum
( Kunth) Nees.
Population size and conservation status. This
species occurs as scattered individuals and small
groves throughout the Wahiawa Drainage area,
where the population size is estimated at 250 to
300 individuals. Additional populations are known
from: the ridge between Mt. La'auhiha'ihai and Mt.
Kahili (6 individuals), the Blue Hole (exact number
unknown). Power Line Trail (8), the Makaleha Mts.
(84). and Wainiha Valley (exact number unknown),
bringing the total to perhaps 350 to 400 plants.
Plants of different size classes including occa¬
sional juveniles are represented in some localities,
suggesting at least limited regeneration is taking
place. However, invasive weedy alien plant species,
notably Psidium cattleianum Sabine and Melastoma
candidum 1). Don and aggressive grasses, threaten
this species (Lorence & Flynn, unpublished). Ad¬
ditional threats are posed by feral animals includ¬
ing feral pigs (Sus scrofa ) and the Polynesian rat
(Rattus exulans), a known seed predator that ad¬
versely impacts regeneration of the Hawaiian Prit¬
chardia species (Cuddihy & Stone, 1990). Feral
pigs are present in many areas inhabited by this
species and may destroy seedlings and increase op¬
portunities for alien plants to invade through deg¬
radation of the habitat. Based on its relatively low
population status, presence of threats, and low re¬
generation, we suggest this new species should be
considered for threatened status.
Etymology. We are pleased to name this new
species for one ol its initial collectors, Timothy W.
Flynn, in recognition of his extensive botanical col¬
lections and contributions to our knowledge of the
Hawaiian flora.
Affinities. Among the Kaua'i species, Pritchar¬
dia flynnii resembles P. hardyi Rock, which is also
characterized by abaxially densely gray tomentose-
188
Novon
lepidote leaves. This latter speeies differs by its
stouter trunk 4-8 m tall and 30-45 cm diam., in¬
florescences equal to or often exceeding the leaves
with minutely villous rachillae, and long-drooping
infructescences with larger fruits 20—35(40) X 16-
25 mm when dry.
Pritchardia flynnii is distinguished from its
Kaua'i congeners by the following combination of
features: short stature with trunk 0.7-7(11) m tall;
relatively small crown with only 10 to 26 leaves;
leaf blades relatively small, 57-107 cm long, nearly
plane to slightly undulate and abaxially densely ap-
pressed tomentose-lepidote; relatively small inflo¬
rescences branched to the second degree, reaching
Vz (rarely %) length of blade in flower and often
reaching the blade apex in fruit; fruits smooth, el¬
lipsoid or ovoid-ellipsoid.
Several collections from the Makaleha Mountains
(Lorence & Flynn 7424, PTBG; Perlman & Wood
16260) diller in having leaf blades that are abaxi¬
ally sparsely scattered lepidote, but these speci¬
mens are otherwise referable to P. flynnii. The com¬
paratively sparse foliar pubescence may be due to
leaf age or weathering, or it may be genetically con¬
trolled. This variant will key out under 8b in the
key below.
Seven endemic Pritchardia species including P.
flynnii occur on Kaua‘i, and two Pacific species are
commonly cultivated there (P. pacifica Seemann &
H. Wendland and P. thurstonii F. Mueller & Drude).
In order to successfully key out these palms com¬
plete material is required including flowers, fruits,
and leaves. In addition, the number of leaves per
crown, relative lengths and positions of leaves, in¬
florescences, and infructescences should be noted
and photographed when making herbarium speci¬
mens. With adequate herbarium material available,
the native Kaua‘i species generally can be sepa¬
rated by the following key, which was adapted in
part from Gemmill (1668) and Read and Hodel
(1660). Measurements are based on dried herbari¬
um specimens.
Kky to tiik Endemic Species oe Pritchardia in Kaua'i,
Hawai‘1
lu. Inflorescences nearly equal lo or exceeding leaf
blades in flower; infructescences usually extend¬
ing well beyond the blade apex and generally
pendulous . P. hardyi
lb. Inflorescences shorter than petioles or reaching
Vz (rarely f4) length of leaf blade; infructescences
shorter than or equaling but not exceeding the
blade apex, erect, arcuate, or pendulous.
2a. Leaf blades abaxially densely silvery to pale
brown or golden, oppressed lepidote-tomen-
tose, the scales and hairs matted and ob¬
scuring the blade surface.
3a. Inflorescence rachillae densely flocco-
se-lanate at least when young, becoming
glabrate with age . P. minor
3b. Inflorescence rachillae glabrous or
sparsely to densely velutinous-tomen-
tose when young and soon glabrescent.
4a. Inflorescence rachillae glabrous,
viscous with varnish-like resinous
coating; flowers and buds shiny,
viscous-resinous; fruit 33—35 mm
long . P. viscosa
4h. Inflorescence rachillae velutinous-
tomentose or glabrescent, not vis¬
cous or resinous; flowers and buds
dull, not shiny and viscous; fruit
22-25 mm long . P. flynnii
2b. Leaf blades abaxially with scattered to
closely arranged scales and hairs, but these
not obscuring the blade surface.
5a. Large palms, up to 20 m tall; trunk
large and stout, 50 cm or more DBH;
crown with more than 40 leaves; leaf
blades waxy or glaucous pale green . .
. P. waialealeana
5b. Moderate- to small-sized palms, less
than 10 m tall; trunk moderate to slen¬
der, much less than 50 cm DBH; crown
with less than 30 leaves; leaf blades
plain green.
6a. Leaves with petioles 00— 170 cm
long; fruit 30—38 X 18 — 21 mm
. P. perlmanii
6b. Leaves with petioles (31)35—90 cm
long; fruit 18-28 X 12-21 mm.
7a. Abaxial surface of blade with
lepidia 0.2— 0.3 mm long; flow¬
ers with calyx 4- pedicel 1.8—
3 mm; petals 3-3.5 mm ....
. P. napaliensis
7b. Abaxial surface of blade with
lepidia 0.4— 1.5 mm long; flow¬
ers with calyx + pedicel 2.5—
5 mm; petals 4.5—7 mm.
8a. Rachillae glabrous; floral
bracts scarious, ovate-tri¬
angular, 0.5— 1.5 mm; ca¬
lyx + pedicel 2.5—3 mm;
petals 4.5—5 mm .
. P. limahuliensis
8b. Rachillae sparsely to
densely velutinous-tomen-
tose when young, often
glabrescent; floral bracts
stiff, subulate-filiform,
1.5—3 mm; calyx + pedi¬
cel 3—5 mm; petals 5.7—7
mm . P. flynnii
(Makaleha Mountains collections)
Paratypes. HAWAIIAN ISLANDS (U.S.A.). Kaua'i:
koloa District, Li hue— Koloa forest reserve, Wahiawa
Stream ud, F. M. 1846. Crueiferae. Pp. 105-185 in C. Gay,
Historia Ffsica y Polftica de Chile, Vol. 1, Botany. Paris,
Santiago.
Volume 14, Number 2
2004
O’Kane & Al-Shehbaz
Physaria in South America
205
Boelcke, 0. 1967. Crueiferae. Pp. 281-371 in A. L. Ca¬
brera (editor). Flora de la Provincia de Buenos Aires,
Vol. 4, pt. 3. Buenos Aires.
- & C. Bomanezuk. 1984. Crueiferae. FI. Patago-
nica, Vol. 4A: 373-544. INTA, Buenos Aires.
Eichler, A. G. 1865. Crueiferae. Pp. 293-338 in C. F.
Martius (editor), FI. Brasil, Vol. 13, pt. 1. Miinchen,
Wien, Leipzig.
Kurtz, F. 1893. Sertum Cordobense — Observaciones Sobre
Plantas Nuevas, Baras 6 Dudosas de la Provincia de
Cdrdoba. Revista Mus. La Plata 5: 283-303.
Manton, I. 1932. Introduction to the general cytology of
the Crueiferae. Ann. Bot. II, 46: 509-556.
Muschler, B. 1908. Crueiferae andinae. Bot. Jahrb. Syst.
40: 267-277.
O’Kane, S. L., Jr. 1999. Lesquerelln navajoensis (Brassi-
caceae), a new species of the L. hitchcockii complex
from New Mexico. Madrono 46: 88—91.
- & 1. A. Al-Shehbaz. 2002. Paysonia, a new genus
segregated from Lesquerella (Brassicaceae). Novon 12:
379-381.
- , - & N. J. Turland. 1999. (1393) Proposal
to conserve the name Lesquerella against Physaria (Cru-
ciferae). Taxon 48: 163—164.
Payson, E. B. 1921 [issued 1922], A monograph of the
genus Lesquerella. Ann. Missouri Bot. Card. 8: 103—
236.
Philippi, B. A. 1864. Plantarum novarum Chilensium:
Centuriae, inclusis quibusdam Mendocinis et Patago-
nicis. Linnaea 33: 1—308.
Rollins, R. C. 1939. The cruciferous genus Physaria.
Bhodora 41: 392^415.
- . 1993. The Crueiferae of Continental North Amer¬
ica. Stanford Univ. Press, Stanford.
- . 1995. Two Lesquerellas (Crueiferae) of south
central and western Montana. Novon 5: 71—75.
- & E. A. Shaw. 1973. The Genus Lesquerella (Cru-
ciferae) in North America. Harvard Univ. Press, Cam¬
bridge.
- . K. A. Beck & F. E. Caplow. 1996. An undescri¬
bed species of Lesquerella (Crueiferae) from the State of
Washington. Bhodora 97: 201—207.
Schulz, O. E. 1934. Neue Cruciferen aus Sudamerika. No-
tizbl. Bot. Gart. Berlin-Dahlem 12: 39^11.
- . 1936. Crueiferae. Pp. 227—658 in A. Engler &
H. Harms (editors), Die Naturlichen Pflanzenfamilien,
Vol. 17B. Verlag von Wilhelm Engelmann, Leipzig.
Watson, S. 1888. Contributions to American botany. XV II.
I. Some new species of plants of the United States, with
revisions of Lesquerella ( Vesicaria ) and of the North
American species of Draba. Proc. Amer. Acad. Arts 23:
249-267.
A New Species of Pariana (Poaceae: Bambusoideae: Olyreae)
Endemic to the Atlantic Moist Forest in the State of Bahia, Brazil
Reyjane Patricia de Oliveira
Departamento de Ciencias Biologicas, Programa de Pos-graduagao em Botanica/Herbario,
Universidade Estadual de Feira de Santana, BR 116N km 3, 44031-460,
Feira de Santana — BA, Brazil, patricia@uefs.br
Hilda Maria Longhi-Wagner
Departamento de Botanica, Universidade Federal do Rio Grande do Sul. Av. Bento
Gongalves, 9500 (43323), 91501-970, Porto Alegre — RS, Brazil, hmlw@vant.com.br
Victoria C. Hollowell
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
victoria.hollowell@mobot.org
ABSTRACT. A new species of Pariana from the
Atlantic forest of the Brazilian state of Bahia is de¬
scribed. Pariana carvalhoi is an herbaceous bam¬
boo (Poaceae: Bambusoideae: Olyreae) that resem¬
bles P. lanceolata hut differs from it by the very
short to absent fimbriae at the leaf sheath apex.
Both species are endemic to Bahia, Brazil. The new
species is only known from the locality of Una, in
southern Bahia. It grows in remnants of humid for¬
est and should be considered as critically endan¬
gered.
Key words: Bahia, Bambusoideae, Brazil, Oly¬
reae, Pariana, Poaceae.
Pariana Aublet includes about 30 species dis¬
tributed in the Amazon Basin from Costa Rica and
Trinidad, to Bolivia and Brazil (Judziewicz et al.,
1999). It has been included in its own tribe Pari-
aneae by several authors (Calderon & Soderstrom,
1980; Clayton & Renvoize, 1986; Hollowed, 1994),
but presently it is considered a genus of the tribe
Olyreae (Clark et ah, 1995; Hollowed, 1987, 1997;
Zhang & Clark. 2000). Pariana , together with Er-
emitis Doll, is classified in the subtribe Parianinae
(Hollowed. 1987. 1997), which is characterized by
the presence of fimbriae at the apex of the leaf
sheaths, termed oral setae (Calderon & Soderstrom,
1980; Hollowed. 1987, 1997), and by the spiciform
inflorescence with each whorl presenting one fe¬
male central spikelet surrounded by five male
spikelets. Kach of these gynecandrous whorls cor¬
responds to a diaspore of the species, shattering as
intact spikelet whorls. According to Soderstrom and
Calderon (1971, 1974). the showy inflorescences of
Pariana species attract insects especially by the
easily visible anthers, and these bamboo grasses
are probably entomophilous. The pollen grains of
some Pariana species differ from the basic pattern
found in grasses (anemophily) by showing an are-
olate exine, sometimes vestigial columellae, and no
distinct annulus (Salgado-Labouriau & Rinaldi,
1990; Salgado-Labouriau et al., 1993); this would
offer more friction in wind transport than the
smooth surface of other grass pollen grains. The
absence of an annulus in Pariana pollen was con¬
firmed, and the developmental detail of its micro¬
spore has been further investigated by Skvarla et
al. (2003).
According to Clark (1990) the state of Bahia rep¬
resents a significant center of diversity for Neotrop¬
ical bamboos, with many new species being de¬
scribed from this area. The two species of Pariana
found in Bahia, P. lanceolata Trinius (cf. Renvoize,
1984) and the new species described here, differ
from other species of the genus by the barbate style,
the smaller number of pale stamens (2 vs. 6 to 36
or more, with pale to bright yellow anthers, within
Pariana), and the much longer pedicels of the male
spikelets. Due to this reduced number of stamens,
the inflorescences of the new and Bahian species
are less conspicuous than those of other Pariana
species.
The new species epithet, Pariana carvalhoi, hon¬
ors the outstanding botanist Andre Maurfcio de
Carvalho (1951—2002), who for many years was the
curator of the herbarium at CEPLAC (Cocoa Re¬
search Center in llheus, Bahia) and was a great
Novon 14: 206-209. 2004.
Volume 14, Number 2
2004
Oliveira et al.
Pariana carvalhoi from Brazil
207
Figure 1. A— C, Pariana lanceolata Trinius. — A. Habit. — B. Inflorescence whorl of male and female spikelets, the
solitary female spikelet cryptic. — C. Apex of leaf sheath with fimbriae evident and numerous. D— N, Pariana carvalhoi
K. P. Oliveira & 11. M. Longhi-Wagner. — D. Habit. — E. Apex of leaf sheath with fimbriae absent or vestigial. — F.
Inflorescence whorl. — G. Central female spikelet. — H. Articulate inflorescence rachis. — I. Female anthecium, ventral
aspect. — J. Female anthecium, dorsal aspect. — K. Male spikelet with adnate laminar pedicel, dorsal aspect. — L.
Male anthecium, ventral aspect. — M. Male anthecium, dorsal aspect. — N. Caryopsis. A— C drawn from IF IF Thomas
et al. 9399 (CEPEC); I ) — N drawn from the holotype, A. M. Carvalho et al. 4382 (CEPEC). Illustration prepared by
Ivan Farias de Castro and Reyjane Patricia de Oliveira.
208
Novon
defender and advocate for the preservation of the
Atlantic forest in eastern Brazil.
Key to the Species of Pariana in Bahia, Brazil
1. Leaf sheaths with numerous fimbriae at the apex,
0.7—20 mm long; leaf blades lanceolate (7.5— )11-
16(— 21.4) X 0.7— 1.7(— 2.5) cm . . . Pariana lanceolata
1'. Leaf sheaths without fimbriae at the apex, rarely
with 1 to 3 vestigial fimbriae, less than 1 mm
long; leaf blades ovate-lanceolate, (6— )1 1.5-20 X
2.1— 3.4 cm . Pariana earvalhoi
Pariana earvalhoi R. P. Oliveira & H. M. Longhi-
Wagner, sp. nov. TYPE. Brazil. Bahia: Una,
along road Sao Josd/Una, 10 Feb. 1994, A. M.
Carvalho, L. G. Clark, T. S. dos Santos, W. W.
Thomas & S. Sant' Ana 4382 (holotype, CE-
PEC; isotype, ISC). Figure I .
Herba perennis cespitosa; vaginae glabrae rare fimbria-
tae; lolia ovato-laneeolata. Spieulae masculae pedicellis
longis, pilosis; stamina 2.
Plants with monomorphic stems 40—75 cm tall,
2-4 mm diam., glabrous or pilose, bearing 3 to 6
leaves; nodes glabrous, with a small conspicuous
scar beneath. Leaf sheaths glabrous or rarely ciliate
at the margins; apex usually without oral fimbriae,
rarely with 1 to 3 vestigial fimbriae, less than 1 mm
long; leaf blades (6-)l 1 .5—20 X 2. 1-3.4 cm, ovate-
lanceolate, base symmetrical, attenuate, apex sym¬
metrical, acuminate, both blade surfaces adaxially
and abaxially glabrous or ciliate along the adaxial
midrib, blade margins scaberulous; pseudopetioles
1- 1.6 mm long; ligules 0.3-0. 5 mm long. Inflores-
cence 6.5—8 X 0.5— 0.8 cm, terminal and spiciform,
with 6 to 8 spikelet whorls 12.5-14 X 0.5— 0.8 mm;
rachis 9-9.5 mm long, scaberulous. Female spike-
lets 10-11.2 X 2.6—3 mm, oblong, glabrous;
glumes 9-10 mm long, both acuminate or shortly
caudate, totally glabrous or with short trichomes at
the apex; anthecium 9.5—1 1 X 2—3 mm, oblong-
lanceolate, yellowish or dark brown. Male spikelets
2— 2.5 X 1.2— 1.5 mm, oblong-ovate, brown; pedi¬
cels 7—8.2 mm long, laminar, usually pilose; glumes
2—3 mm long, oblong-triangular, pilose; lemmas pu-
berulent, 3-nerved; stamens 2. Caryopsis glabrous,
cryptic but free within the persistent anthecium,
apex with erect stylar remnants, hilum linear.
Pariana earvalhoi differs from P. lanceolata es-
peeially by its wider leaf blades that are ovate-lan¬
ceolate (vs. lanceolate in P lanceolata ), and by the
absence of fimbriae at the apex of the leaf sheaths,
that, i( present, are vestigial. This absence of leaf
sheath fimbriae was noted by Tutin (1936) for other
species of Pariana but only from Amazonia.
Both Pariana lanceolata and P. earvalhoi are en¬
demic to the Brazilian state of Bahia, occurring
within moist Atlantic forest. According to IUCN cri¬
teria (IUCN, 1994), P. earvalhoi should be consid¬
ered “critically endangered,” with only a few pop¬
ulations known, all oceurring outside preservation
areas. Although many field trips were made to dif¬
ferent areas in Bahia, additional populations of the
new species have not been found.
Paratypes. BRAZIL. Bahia: Una, estrada Una a Sao
JosG 18 Feb. 1994, A. M. Carvalho et al. 4395 (CEPEC),
10 Feb. 1994, A. M. Carvalho et al. 4384 (CEPEC); Fa¬
zenda Dendaevea, 30 km from Una, 28 Apr. 1976, C. E.
Calderon et al. 2403 (CEPEC).
Acknowledgments. The authors thank the Univ-
ersidade Estadual de Feira de Santana for making
the field trips possible, and the Cocoa Research
Center for specimen loans and logistic support for
fieldwork. The second author thanks the Brazilian
Research Council (CNPq) for a Research Fellow¬
ship. In memoriam, we are grateful to the unfor¬
gettable Dr. Andre Maurfeio de Carvalho for his
enthusiastic support.
Literature Cited
Calderdn, C. E. & T. R. Soderstrom. 1980. The genera of
Bambusoideae (Poaceae) of the American continent:
Keys and comments. Smithsonian Contr. Bot. 44: 1-27.
Clark, L. G. 1990. Diversity and biogeography of Neo¬
tropical bamboos (Poaceae: Bambusoideae). Acta Bol.
Brasil. 4: 125-132.
- , W. Zhang & J. E. Wendel. 1995. A phylogeny of
the grass family (Poaceae) based on ndhV sequence
data. Syst. Bot. 20: 436-460.
Clayton, W. D. & S. A. Renvoize. 1986. Genera Gramin-
um: Grasses of the World. Royal Botanic Gardens, Kew.
Hollowell, V. C. 1987. Systematics of the Subtribe Pari-
aninae (Poaceae: Bambusoideae: Olyreae). Ph.D. The¬
sis, University of South Carolina, Columbia.
- . 1994. Tribus Parianeae. Pp. 216—218 in G. Dav-
idse, M. Sousa & A. 0. Chater (editors). Flora Mesoam-
ericana, Vol. 6. Alismataceae a Cyperaceae. Universi-
dad Nacional Autonoma de Mexico, Mexico, D.F.;
Missouri Botanical Garden, St. Louis; The Natural His¬
tory Museum, London.
- . 1997. Systematic relationships of Pariana and
associated Neotropical taxa. Pp. 45—60 in G. P. Chap¬
man (editor). The Bamboos. Academic Press, London.
IUCN (The World Conservation Union). 1994. The World
List of Threatened Trees. Appendix 4: 629-643. World
Conservation Press, Gland, Switzerland.
Judziewicz, E. M., L. G. Clark, X. Londono & M. ,1. Stern.
1999. American Bamboos. Smithsonian Institution
Press, Washington, D.C.
Renvoize, S. A. 1984. The Grasses of Bahia. Royal Bo¬
tanic Gardens, Kew.
Salgado-Labouriau, M. L. & M. Rinaldi. 1990. Palynology
of Gramineae of the Venezuelan mountains. Grana 29:
1 19-128.
- , S. Nilsson & M. Rinaldi. 1993. Exine sculpture
in Pariana pollen (Gramineae). Grana 32: 243-249.
Skvarla, J. J., J. R. Rowley, V. C. Hollowell X' W. E. Chis-
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soe. 2003. Annulus-pore relationship in Gramineae (Po-
aceae) pollen: The pore margin of Parinrui. Amer. J.
Bot. 90: 924—930.
Soderstrom, T. R. & C. E. Calderon. 1971. Insect polli¬
nation in tropical rain forest grasses. Biotropica 3: 1—
16.
- & - . 1974. Primitive forest grasses and evo¬
lution of the Bambusoideae. Biotropica 6: 141—153.
Tutin, G. 1936. A revision of the genus Pariana (Grami¬
neae). J. Linn. Soc., Bot. 50: 337—362.
Zhang, W. & L. G. Clark. 2000. Phytogeny and classifi¬
cation of the bamboos (Poaceae: Bambusoideae). Pp.
35^12 in S. W. L. Jacobs & J. Everett (editors). Grasses,
Systematics and Evolution. CS1R0, Melbourne. [Pro¬
ceedings of the 2nd International Conference on the
Comparative Biology of the Monocotyledons, Vol. 2.]
New Taxa of Calyptranthes (Myrtaceae) from Colombia
Carlos Parra- 0.
Instituto de Ciencias Naturales, Universidad Nat ional de Colombia, Apartado Aereo 7495,
Bogota, Colombia, caparrao@unal.edu. co
Abstract. Two new species (C. buchenavioides,
C. estoraquensis) and one new subspecies (C. loz-
anoi subsp. tornentosa ) of Calyptranthes (Myrta-
ceae) from Colombia are described and illustrated.
The taxonomic affinities and the distribution ol the
new taxa are discussed.
Key words: Calyptranthes , Colombia, Myrta¬
ceae.
Calyptranthes Swartz is a genus of ca. 100 spe¬
cies in Myrtaceae, distributed from Florida (United
States), the Caribbean, and Mexico to northern Ar¬
gentina and Uruguay (Legrand, 1962; MeV'augh.
1958, 1963; Sanchez-Vindas, 1990). The genus is
distinguished by frequently dibrachiate trichomes,
inflorescence usually a pair of panicles, calyptrate
calyx, 2(-3)-locular ovary, and myrcioid embryo
(Landrum & Kawasaki, 1997; Holst, 2002). In a
preliminary study of the genus for the Colombian
flora, two new species and a new subspecies of Ca¬
lyptranthes lozanoi C. Parra-0. were recognized as
undescribed.
Calyptranthes buchenavioides C. Parra-0., sp.
nov. TYPE: Colombia. Valle del Cauca: “Hoya
del rfo Cali, vertiente derecha, entre Puente
Sofia y Yanaconas,” 1580-1730 in, 6 Nov.
1944 (fr), J. Cuatrecasas 19725 (holotype, F).
Figure 1.
Arbor 3-^4 m alta; ramulis juvenilibus subteretibus, par-
ee pubescentibus, vetustis teretibus, glabratis. Folia pe-
tiolata, petiolo 0.6— 1.6 mm longo; lamina coriacea, supra
glabra, subtus glabrata, obovata, 1.3— 2.6 X 0.7-1. 4 cm,
apice obtusata, basi attenuata, costa supra impressa, gla¬
bra. subtus convexa, tantum per proximalem unam terliam
partem parce pilosa. Inflorescentia et flores non visi. Fruc-
tus globosus, diametro 1.0— 1.3 cm.
Tree, 3-^4 in tall, almost glabrous; trichomes
when present pale gray (except in the embryo);
young branches terete or subterete, sparsely pu¬
bescent; old branches terete, glabrous; vegetative
buds lanceolate or conical, 0.8— 1.1 mm long, pu¬
bescent. Leaf blades obovate, 1.3— 2.6 X 0.7-1. 4
cm, coriaceous, the upper surface glabrous, with
impressed glandular dots, the lower surface gla¬
brous; apex obtuse, sometimes slightly apiculate;
Novon 14: 210-215. 2004.
base attenuate; midvein convex above, glabrous,
convex below, sparsely pilose only in the proximal
VS; lateral veins 7 to 11 pairs (almost impercepti¬
ble), sulcate above, impressed below, joining in a
straight marginal vein at 0.5 mm from the margin;
petiole 0.6— 1.6 mm long, barely rugose, glabrous
or sparsely pubescent, slightly canaliculate. Inflo¬
rescence terminal, apparently a panicle, the axes
terete or subterete, glabrous; bracts ovate, 0.5 X
0.3— 0.4 mm. pubescent, deciduous; bracteoles un¬
known. Buds and flowers unknown: calyptra (per¬
sistent in the fruit) discoid, ca. 1.4 mm diam., with¬
out prominent apiculum. glabrous. Fruits globose,
blackish, 1-1.3 cm diam., exocarp smooth, gla¬
brous; seeds 1, reniform, 1 — 1.1 X 0.9—1 cm, seed
coat papyraceous, pale orange, smooth, without
glands; embryo myrcioid, lower portion ol the hy-
pocotyl and portions of the cotyledons densely pu¬
bescent with long (1.2— 1.5 mm) gray trichomes
mixed with short (ca. 0.5 mm) red trichomes.
The specific epithet refers to the striking vege¬
tative similarity (profuse ramification, opposite, ob¬
ovate, and coriaceous leaves, and similar size of the
leaves) between this new species and some species
ol Buchenavia Eichler (Combretaceae), especially
with Buchenavia tetraphylla (Aublet) R. A. Howard
and Buchenavia parvifolia Ducke.
Calyptranthes buchenavioides is known from
coastal and premontane forest of the western An¬
dean Cordillera in Colombia. Despite the lack of
complete inflorescences of the studied specimens,
other characters such as the calyptra (falling as a
unit), the circular scar at the top of the fruit, and
the myrcioid embryo place it in Calyptranthes.
Calyptranthes buchenavioides is somewhat simi¬
lar to Calyptranthes pulchella DC., a species known
from Colombia to Brazil. The leaves of the species
are similar in shape (obovate blades) and size. Ca¬
lyptranthes buchenavioides differs from Calyptran¬
thes pulchella by the terete or subterete (vs. 2-
winged) branchlets, the base of the leaf blades
attenuate (vs. acute or cuneate), and the diameter
of the calyptra (ca. 1.4 mm vs. 2-3 mm).
Paratypes. COLOMBIA. Valle de Cauca: “Punta
Arenas, north shore of Buenaventura Bay, near sea level,"
Volume 14, Number 2
2004
Parra-O.
New Taxa of Calyptranthes
211
Figure 1. Calyptranthes buchenavioides C. Parra-0. — A. Vegetative branch. — B. Detail of a partial inflorescence. —
C. View of the calyx scar on the Iruit apex, showing the attached calyptra. — D. Fruit. — E. Embryo. (A, C-E, from
the holotype, Cuatrecasas 19725; B, Cuatrecasas 38628 (F paratype).)
2 June 1944. E. R. Killip & J. Cuatrecasas 38628 (COL,
F).
Calyptranthes estoraquensis C. Parra-O., sp.
nov. TYPE: Colombia. Norte de Santander: l.a
Playa de Helen, “vereda Peritama, Area Nat¬
ural Unica Los Estoraques,” 1650 m, 20 Feb.
2002 (Ir), R. Galindo, J. Penuela & E. Rodri¬
guez 689 (holotype, COL: isotype, MO). Fig¬
ure 2.
Arbor 5 m alta; ramulis juvenilibus subteretibus, parce
pubescentibus, vetustis teretibus, glabratis. Folia petiola-
ta, petiolo 3.8— 5. 5(— 7.6) mm longo; lamina coriacea, supra
glabra vel puberula, subtus puhescente, obovata, 5.0— 7.3
X 3.1— 4.7 cm, apice obtusata, basi cuneata, costa supra
impressa, tantum per proximalem unam tertiam partem
puhescente, subtus convexa, dense puhescente. Inflores-
centia paniculata, 4—6 cm longa, dense pubescens; pedi-
cello nullo. Flos alabastro obovoideo, 2.3— 2.6 mm longo;
calyptra 2.0— 2.2 mm diam.; staminibus (1.6)2. 1—3.1 mm
longis; ovario 2-loculari. Fructus globosus, diametro 5.5-
7.9 mm.
Tree, 5 m tall; trichomes when present light red¬
dish brown; young branches subterete, subcom¬
pressed, sparsely pubescent, the light reddish
212
Novon
Figure 2. Calyptranthes estoraquensis C. Parra-O. — A. Fruiting branch. — B. Detail of the leaf blade pubescence
(lower surface). — C. Flower, after anthesis. — D. Fruit. — E. Seed. — F. Longitudinal section of the fruit and seeds
showing the embryo. (A, B, D— F, from the holotype Galindo el al. 689; G, Galindo el al. 1168 (COL paratype).)
brown trichomes mixed with gray trichomes, the
gray ones located mostly in the margins and apex
of the branches; old branches terete, glabrous; veg¬
etative buds semi -elliptic, 1.4— 1.5 mm long, pilose.
Leaf blades obovate, 5—7.3 X 3.1— 4.7 cm, coria¬
ceous. the upper surface glabrous to puberulous,
sometimes with whitish trichomes, with impressed
glandular dots, the lower surface pubescent; apex
obtuse; base cuneate; midvein sulcate above, pu¬
bescent only in the proximal Yi, convex below,
densely pubescent; lateral veins 17 to 35 pairs in
addition to weaker intermediates, convex on both
surfaces, joining in a straight marginal vein at 1.4
mm from the margin; petiole 3.8-5. 5(— 7.6) mm
long, rugose, puberulous with gray or mixed gray
and light reddish brown trichomes (petioles of
Volume 14, Number 2
2004
Parra-O.
New Taxa of Calyptranthes
213
leaves near the inflorescences are densely pubes¬
cent with light reddish brown trichomes), canalic¬
ulate. Inflorescence axillary or terminal, the paired
and opposite panicles 4-6 cm long, with 20 to 30
flowers per panicle, the axes subterete or com¬
pressed. densely pubescent; peduncles 2. 7-3.3 cm
X 1.7— 2.4 mm; bracts lanceolate, ca. 4.5 X 2 mm,
pubescent, truncate in the base, deciduous; brac-
teoles (1)2, the larger one al the base of the hy-
panthium, ovate, ca. 0.7 X 0.3 mm, pubescent, de¬
ciduous or persistent alter anthesis, the smaller one
incompletely surrounding the base of the hypan-
thium, squamiform, ca. 0.5 X 0.9 mm, pubescent,
persistent after anthesis. Flower buds obovoid, 2.3—
2.6 mm long, pubescent, sessile; ealyptra discoid,
2—2.2 mm diam., without prominent apiculum, pu¬
bescent; petals absent; hypanthium 1.8— 2.1 X 2.6-
2.9 mm, prolonged 0.7— 0.8 mm above the ovary,
pubescent outside, glabrous inside; style generally
not developed, when occasionally developed, 1 .2
mm long, glabrous; filaments (1. 4)1.9— 2.9 mm; an¬
thers globose, 0.2 mm long, bearing one apical
gland; ovary 1 mm diam., 2-locular, 1 ovule per
locule. Fruits globose, dark wine-red, 5.5— 7.9 mm
diam., exocarp smooth, sparsely to moderately pu¬
bescent; seeds 2. reniform, 4.8-5. 8 X 2.8— 4.2 mm,
seed coat papyraceous, orange, smooth, without
glands; embryo myrcioid, glabrous.
Calyptranthes estoraquensis is only known from
the Colombian Natural Area named “Estoraques,”
a semidesert landscape (badlands) made of rock
formations caused by erosion. Estoraques is located
at the eastern Andean Cordillera in Norte de San¬
tander Department, between 1450 and 1900 m. At
present, this particular area is included in the Na¬
tional Park System as a protected area, but the pop¬
ulation of the species there seems to be small, “with
only a few individuals” (R. Galindo, pers. comm.
2002).
Calyptranthes estoraquensis seems to be related
to Calyptranthes brasiliensis Sprengel, and espe¬
cially with Calyptranthes brasiliensis var. mutabilis
(Berg) Legrand. A specimen of C. brasiliensis var.
mutabilis from Brazil, Duarte 9152 (MO), exhibits
some degree of similarity w ith the specimens of C.
estoraquesis from Colombia, particularly in the veg¬
etative morphology (trichomes light reddish brown
in young branches and petioles, and similar size of
the petioles). The pubescence of the adaxial leaf
blade and the inflorescence and the size of the in¬
florescence are also common characteristics of
these two taxa. Nevertheless, Calyptranthes estora¬
quensis differs from Calyptranthes brasiliensis var.
mutabilis by its leaf base (cuneate vs. rounded), the
laminae (obovate vs. ovate or ovate-oblong) and the
size of the fruit (5.5— 7.9 mm vs. 4—4.5 mm).
Paratype. COLOMBIA. Norte de Santander: La
Playa de Beldn, “vereda Peritama, Area Natural tlnica Ixjs
Estoraques,” 1550 m, 26 Feb. 2003 (fl). H. Galindo, J.
Pefiuela & E. Rodriguez 1168 (COL).
Calyptranthes lozanoi C. Parra-O. subsp. to-
mentosa C. Parra-O., subsp. nov. TYPE: Co¬
lombia. Antioquia: Guarne, Laguna de Guar-
ne, 20 July 1938 (bud, fr). Bra. Daniel 2258
(holotype, US; isotype, COL). Figure 3.
Haec subspecies a Calyptranthe lozanoi subsp. lozanoi
alabastris ellipsoideis vel late ellipsoideis (nee obovo-
ideis), ramulis, petiolo, foliis (subtus), inflorescentia et al¬
abastris modice vel dense pubescentibus (nee parce pu-
bescentibus trichomatibus albis vel glabratis, interdum
glabris), atque fructu bullato (nec laevi), exocarpio spar-
sim pubescente (nec glabro) differt.
Shrub, 2^1 m tall; trichomes when present yel¬
lowish or blackish; young branches subcompressed,
sparsely to densely pubescent, sometimes almost
blackish; old branches terete, glabrous; vegetative
buds lanceolate or elliptic, 2.2^1. 1 mm long, pu¬
bescent. Leal blades obovate, sometimes broadly
elliptic, 3.2— 5.2(-6) X 1.6— 3.1(— 4) cm, coriaceous,
the upper surface glabrous to sparsely puberulous,
with impressed glandular dots, the lower surface
sparsely to densely pubescent; apex acute or ob¬
tuse; base cuneate; midvein convex above, puber¬
ulous to sparsely pubescent in the proximal Vs, con¬
vex below, moderately to densely pubescent; lateral
veins 15 to 18 pairs, convex on both surfaces; mar¬
ginal veins 2, the inner one 0.9-1. 1 mm from die
margin, the outer one ca. 0.4 mm from the margin;
petiole 2.6-6.1(— 8) mm long, blackish, lightly ru¬
gose, sparsely to densely pubescent (petioles of
leaves near the inflorescences are densely pubes¬
cent with yellowish trichomes), canaliculate. Inflo¬
rescence subterminal, the paired and opposite pan¬
icles 3.5—7 cm long, with 12 to 25 flowers per
panicle, the axes subterete or compressed, densely
pubescent, sometimes almost blackish, rarely gla¬
brous; peduncles 2.2—4 cm X 1.3— 1.7 mm; bracts
dimorphic, either lanceolate, ca. 6X2 mm, pu¬
bescent with yellowish trichomes, deciduous, or ob¬
ovate, 1.2— 1.7 X 0.4— 0.5 cm. persistent; bracteoles
2, opposite at base of hypanthium, pubescent, de¬
ciduous or persistent after anthesis, the larger one
lanceolate, 2.8— 3.4 X 1 — 1.2 mm, the smaller one
elliptic, 1.7— 2.8 X 0.6— 0.8 mm. Flower buds ellip¬
soid or widely ellipsoid, 4.6— 6.4 mm long, pubes¬
cent, sometimes almost blackish, rarely glabrous,
on pedicel 0.8— 1.3 X 0.7— 0.9 mm, densely pubes¬
cent, rarely glabrescent; ealyptra discoid, 3—3.5
214
No von
Figure 3. Calyptranthes lozanoi C. Parra-O. subsp. tomentosa C. Parra-O. — A. Fertile branch. — B. Flower bud. —
C. Flower, after anthesis. — D. Longitudinal section of hypanthium and ovary. — E. Petal. — F. Fruit. — G. Embryo
without seed coat. (A, B, F. G, from the holotype Bro. Daniel 2258 ; G— E, Cabrera 254 (GOl. paratype).)
Volume 14, Number 2
2004
Parra-O.
New Taxa of Calyptranthes
215
mm diam., weakly to prominently apiculate, gen¬
erally pubescent, rarely glabrous; petals present,
attached to the inner portion of the calvptra, 2 or
3 (early deciduous at anthesis), obovate, 2.2 X ].]
mm, dorsally densely pubescent with white tri-
chomes; hypanthium 2.5—3 X 4. 1-4.7 mm, pro¬
longed 1.3— 1.6 mm above the ovary, pubescent out¬
side, glabrous inside; style 4-5.5 mm long,
glabrous; stamens 50-150; filaments 2.2-6 mm; an¬
thers globose, 0.2— 0.3 mm long, without apical
glands; ovary 1.5 mm diam., 2-locular. 1 ovule per
locule. Fruits globose, light brown, 8 mm diam.,
exocarp bullate, sparsely pubescent with whitish
trichomes; seeds 2, reniform, 5.4—6 X 5 mm, seed
coat papyraceous, red, smooth, without glands; em¬
bryo myrcioid, glabrous.
The subspecific epithet refers to the yellowish or
blackish tomentum of the plant that varies from
moderately dense to dense on tw igs, leaves (petiole
and lower surface of the blade), inflorescence, and
flower buds. Calyptranthes lozanoi subsp. tomentosa
is known only from the montane forest of the Cen¬
tral Andes Cordillera in Colombia.
In 2001 I described Calyptranthes lozanoi (Par¬
ra-O.. 2001) as a conspicuous species from the Si¬
erra Nevada de Santa Marta (Colombia), an isolated
coastal mountain separated from the Andes Cordil¬
lera by plains and semiarid regions. The specimens
cited here for the description of the new subspecies
are similar to the specimens collected in Sierra Ne¬
vada de Santa Marta by the obovate leaf blade (and
also in its size), the size of the panicles, the size of
the hypanthium, the pubescence of petals, and the
size of the fruit. Also, the vegetative morphology of
all the specimens strikingly resemble the general
aspect of Myrcianthes O. Berg species that grow in
the montane forests of Colombia (Parra-O., 2001).
Calyptranthes lozanoi subsp. tomentosa differs
from the typical subspecies by the ellipsoid or
w idely ellipsoid (vs. obovoid) flower buds, the mod¬
erate to dense yellowish or blackish pubescence
(vs. sparse whitish pubescence or glabrescent, to
often glabrous) of the twigs, petioles, lower surface
of the leaf blade, inflorescence, and flower buds (vs.
sparse whitish pubescence or glabrescent, to often
glabrous), and the bullate, sparsely pubescent fruits
(vs. smooth and glabrous).
Paratypes. COLOMBIA. Antioquia: Medellin, “Hoya
de la Estacidn Experimental Eorestal ‘Piedras Blancas,’ ”
June— Oct. 1957 (H), /. Cabrera 86 (COL, MEDEL), /. Ca¬
brera 254 (COL, MEDEL); Medellm-Guarne, “Parque
Eeologico Piedras Blancas, laguna de Guarne,” 75°29'W,
6°18'N, 2350 m, 22 July 1995 (II), F Roldan et al. 2354
(HUA, MO).
Acknowledgments. I express my special grati¬
tude to the Missouri Botanical Garden and the Gen-
tty Fellowship Program, which supported me as a
visiting researcher from June to July 2003. In St.
Louis, I’m indebted to Alina Freire Fierro for her
help in all the different aspects related to this pro¬
gram. I am grateful to F and US for sending loans
of Calyptranthes to COL, and to HUA and MEDEL
for providing me access to their collections. Thanks
to Henry Arellano, who prepared the illustrations,
and to Peter Hoeh (MO) for reviewing an earlier
version of the manuscript. Thanks to Roy Gereau
(MO) for reviewing the Latin descriptions and di¬
agnoses, and to B. Holst (SEL) and an anonymous
reviewer for their helpful comments on the manu¬
script. Also, I thank the Herbario Nacional Gol-
ombiano (COL) of the Instituto de Ciencias Natur-
ales, Universidad Nacional de Colombia, for its
support.
Literature Cited
Holst. B. 2002. New species and notes on Myrtaceae from
northern South America. Selbyana 23: 137-180.
Landrum, L. & M. L. Kawasaki. 1997. The genera of Myr¬
taceae in Brazil: An illustrated synoptic treatment and
identification keys. Brittonia 49: 508-536.
Legrand, 1). 1962. El genero Calyptranthes en el Brasil
austral. Lilloa 31: 183—206.
McVaugh, 11. 1958. Myrtaceae. In: ,|. F. Maguire (editor).
Flora of Peru. Field Mus. Nat. Hist.. Bot. Ser. 13, part
4(2): 569-818.
- . 1963. Myrtaceae. Flora of Guatemala. Fieldiana,
Bot. part 24, 7(3): 283-405.
Parra-O., C. 2001. L ria nueva especie de Calyptranthes
Sw. (Myrtaceae) de Colombia. Caldasia 23(2): 435-439.
Sanchez- V indas, P. 1990. Myrtaceae. In: A. G6mez-Pompa
(editor). Flora de Veracruz. Instituto de Ecologfa, Xal-
apa, Veracmz, Mexico 62: 1-146.
New Species and Combinations in Astragalus (Leguminosae) from
China and the Himalayas
Dietrich Podlech
Ludwig-Maximilians-University Munich, Department Biologie 1 — Systematic Botany,
Menzinger StraBe 67, D-80638, Munich, Germany, podlech@lrz.uni-muenchen.de
Lang-Ran Xu
Northwestern Institute of Botany, Academia Sinica, Yangling 712100, Shaanxi,
People’s Republic of China
ABSTRACT. Ten new species of Astragalus are de¬
scribed here. They belong to the following sections:
section Alopecuroidei : A. burqinensis ; section Bra-
chycarpus : A. barclayanus, A. golubojensis, A.
mieherorum, A. nanshanicus, A. pseudojagnobicus,
A. sagastaigolensis ; section Hemiphaca : A. nigro-
dentatus; section Polycladus: A. tibeticola ; section
Skythropos : A. griersonii. Furthermore, six taxa are
raised in rank.
Key words: Astragalus, Bhutan, China, Legu-
minosae, Nepal.
In preparing the treatment of Astragalus L. for
the English version of the Flora of China, several
new species could he detected and furthermore
some taxa, formerly described as varieties, were
found to be good species. In the present paper the
new species are described and the formal combi¬
nations are made. The flora of China is very rich
in Astragalus species. In the Flora, 357 species, of
which 189 are endemic in China, will he treated.
Astragalus L. sect. Alopecuroidei DC., Prodr. 2:
294. 1825. TYPE: Astragalus alopecuroides L.
Astragalus burqinensis Podlech & L. K. Xu, sp.
nov. TYPE: China. Prov. Xinjiang: Burqin, 600
m, 24 July 1998, L. R. Xu 2029 (holotype,
MSB).
Differt ah A. alopecuro planta in partibus vegetativis
glabra vel sub stipulis tantum sparse pilosa, petiolo 2.5—
7 cm (nec 1—3 cm) longo, foliolis 13— 15-jugis (nec 17—
27-jugis), late ovatis, 12-30 X 10-22 mm, apice rotun-
data vel leviter emarginata (nec anguste ovata, (10— )20—
35(_40) x (6—) 1 0— 1 3(— 20) mm, apice obtusa ad
subacuta), bracteolis deficientibus, vexillo 27-28 mm lon¬
go, dorso piloso (nec ca. 20 mm longo, glabro), laminis
carinae ca. 15 X 8-9 mm (nec 7 X 3.5 mm), leguminibus
ca. 14 mm (nec ca. 7 mm) longis, ab A. pseudovulpino
planta in partibus vegetativis glabra vel sub stipulis tan¬
tum sparse pilosa, bracteolis deficientibus, laminis carinae
8—9 mm (nec 4—5 mm) latis.
Plants more than 30 cm tall (lower parts are
missing). Stem erect, finely sulcate, glabrous or just
below the stipules and at the base of stipules
sparsely hairy. Stipules ca. 15 mm long, narrowly
triangular, shortly adnate to the petiole, heaves 20-
30 cm long: petiole 2.5—7 cm long, finely striate-
sulcate. Leaflets in 13 to 15 pairs, widely ovate,
12-30 X 10-22 mm, at the apex rounded to slight¬
ly emarginate. Peduncle very short, up to 1 cm
long, densely covered with spreading hairs 1—2 mm
long. Racemes (in fruit) cylindric, 7-9 cm long,
densely many-flowered; axis loosely covered with
spreading hairs. Bracts linear-acute, 12-18 mm
long, ciliate at the margins with spreading hairs
1.5-2. 5 mm long. Bracteoles absent. Flowers sub-
sessile. Calyx 12-19 mm long, campanulate, dense¬
ly covered with ascending to spreading hairs 2—5
mm long; teeth subulate, 5-7 mm long. Petals yel¬
low. Standard 27-28 mm long; blade elliptic, ca. 9
mm wide, emarginate at the apex, at the base sub-
abruptly narrowed into the long c law. Wings un¬
known. Keel ca. 27 mm long; blades triangular,
with gibbously curved lower edge and ± straight
upper edge, obtuse at the apex, ca. 15 X 8—9 mm;
auricle acute, ca. 1 mm long, claw ca. 13 mm long.
Legumes sessile, oblong, 12-14 mm long, 7-9 mm
high and 4-7 mm wide, obtusely carinate ventrally,
slightly grooved dorsally, abruptly contracted at the
apex into a beak ca. 2 mm long, bilocular; valves
netlike rugulose, loosely covered with subappres-
sed soft hairs up to 3 mm long. Seeds 3 X 2—3 mm,
dark brown, smooth.
Astragalus burqinensis differs from A. alopecurus
Pallas by having vegetative parts glabrous or
sparsely hairy only, by the petiole 2.5— i (vs. 1—3)
cm long, the leaflets in 13 to 15 (vs. 17 to 27) pairs,
widely ovate, 12-30 X 10-22 mm, rounded or
slightly emarginate (vs. narrowly ovate, (10— )20—
35(_40) X (6—) 1 0-13(— 20) mm, obtuse to sub-
Novoin 14: 216-226. 2004.
Volume 14, Number 2
2004
Podlech & Xu
Astragalus from China
217
acute), bracteoles absent, standard 27—28 mm long,
hairy on the back-side (vs. ca. 20 mm long, gla¬
brous), limbs of keel ca. 15 X 8-9 mm (vs. 7 X
3.5 mm), legumes ca. 14 (vs. ca. 7) mm long, from
A. pseudovulpino N. Ulziykhutag by being in veg¬
etative parts glabrous or only below the stipules
sparsely hairy, bracteoles absent and limbs of keel
8—9 (vs. 4—5) mm wide.
Astragalus sect. Ammodendron Bunge, Mem.
Acad. Imp. Sci. Saint Petersbourg 1 1(16): 128.
1868. TYPE: Astragalus ammodendron Bunge.
Astragalus macrostephanus (S. B. Ho) Podlech
& L. R. Xu, stat. nov. Based on Astragalus
iliensis Bunge var. macrostephanus S. B. Ho,
Bull. Bot. Res. North-East. Forest. Inst. 3(1):
68. 1983. TYPE: |China.] Xinjiang: Kazak
Zizhizhou, Huocheng Xian, Sandohezi, 540 m,
14 June 1959, A. R. Li & J. N. Zhu 10619
(holotype, PE; isotvpe, WUK).
Because of the very short original diagnosis of
this taxon a complete description is given here.
Astragalus macrostephanus differs from A. iliensis
by stems of the current year up to 30 (vs. up to 20)
cm long, leaves 8-12 (vs. 3—5) cm long, petiole 4—
5 (vs. 0.5—1) cm long, leaflets in 3 pairs (vs. in 1
or 2 pairs), 12-25 (vs. 20-40(-50)) mm long, pe¬
duncles 6-10 (vs. 2-5) cm long, calyx 5-6 mm
long, rather densely covered with medifixed, ap-
pressed hairs 0. 3-0.4 mm long (vs. 3-4 mm long,
densely covered with partly flexuose, asymmetri¬
cally bifurcate, subappressed to slightly ascending
hairs), petals longer, standard ca. 13 (vs. 8—9) mm,
wings 12 (vs. 7—8) mm, keel 10 (vs. 6—7) mm long,
legumes 10—12 (vs. 4-5) mm long, 4—5 (vs. ca. 2)
mm high and wide.
Plants suffruticose, ca. 35 cm tall, covered with
medifixed, appressed white hairs. Caudex divided,
with long subterranean stolons, covered with gray-
brownish bark. Stems erect, branched, very densely
covered with white hairs 0.3— 0.6 mm long, shoots
of the current year up to 30 cm long. Stipules whit¬
ish to pale brownish hyaline, 2—4 mm long,
obliquely triangular, acute, adnate to the petiole for
ca. 1 mm, the lower ones distinctly vaginate-con-
nate behind the stem, the upper ones mostly free,
sparsely to loosely hairy, at the margins with strong¬
ly asymmetrically bifurcate to basifixed hairs,
sometimes glabrescent with age. Leaves 8-12 cm
long; petiole 4-5 cm long, like the rachis stout, at
the base up to 1.5 mm thick, hairy like the stem.
Leaflets in 3 pairs, narrowly elliptic to narrowly
ovate, 12-25 X 3—5 mm, acute at the apex, on
upper side loosely covered with hairs 0.6— 0.8 mm
long, on underside loosely to rather densely cov¬
ered with somewhat shorter hairs. Peduncle 6—10
cm long, hairy like the stem. Raceme 5—12 cm
long, very remotely 10- to 12-flowered. Bracts whit¬
ish to pale brownish hyaline, 1-1.5 mm long, nar¬
rowly triangular to narrowly ovate, sparsely white
hairy. Pedicels ca. 1 mm long, hairy. Calyx 5-6 mm
long, campanulate, rather densely covered with
white hairs 0.3— 0.4 mm long; teeth subulate, ca. 1
mm long. Standard ca. 13 mm long; blade ca. 5
mm wide, widely elliptic, emarginate at the apex,
below the middle subabruptly contracted into the
widely cuneate claw longer than the blade. Wings
ca. 12 mm long; blades narrowly oblong, round-
tipped, 6 X 1.5 mm; auricle short, claw ca. 6 mm
long. Keel ca. 10 mm long; blades obliquely ellip¬
tic-curved, with widely curved lower edge and con¬
cave upper edge, obtuse at the apex, 4.5 X 2 mm;
auricle short, clawT 5.5 mm long. Ovary sessile,
ovoid, densely hairy; style glabrous. Legumes ses¬
sile, ovoid, 10—12 mm long, 4—5 mm high and
wide, obtusely carinate ventrally, widely grooved
dorsally, acuminate at the apex into a curved beak
ca. 1 mm long, nearly fully bilocular; valves thin
but tough, densely villous with subbasifixed, ±
spreading, tangled, fine, soft white hairs 2-3 mm
long, mostly sitting on depressed tubercles.
Astragalus sect. Brachycarpus Boriss., FI. URSS
12: 877. 1946. TYPE: Astragalus melanos-
tachys Bentham (lectotype, designated by Pod¬
lech (1990: 466)).
Astragalus barclayanus Podlech, sp. nov. T^ PE:
Nepal. Tilicho Pass path, 4820 m. 7 June
1971. C. Barclay & I1. M. Synge 2620 (holo¬
type, K).
Differt ab A. chateri planta pilis albis nigris immixtis
0.2— 0.3(— 0.5) mm longis, in inflorescentiis nigris ad 0.7
mm longis (nec pilis mere albis 0.3— 0.8 mm longis) ob-
tecta, stipulis 3^4 (nec 4—6) mm longis, breviter connatis
vel superioribus liberis (nec ad medium connatis), foliolis
supra glabris (nec sparse ad laxe pilosis), bracteis angus-
tissime triangularibus, 2.5—3 mm longis (nec ovati-acu-
minatis, 3-5 mm longis), calyce 4 (nec 5—6) mm longo,
pilis ascendentibus ad patentibus (nec subappressis) ob-
tecto, vexillo 5-6 (nec 7—8) mm longo, ovario glabro (nec
piloso).
Plants 7-12 cm tall, covered with subappressed
to slightly ascending hairs 0.2-0.3(— 5) mm long, in
the inflorescence up to 0.7 mm long. Caudex with
slightly elongated stolons provided with stipule-like
bracts. Stems short, 3-5 cm long, prostrate to as¬
cending, in lower part (the stolon part) glabrous,
pale yellowish, in upper part (stem of the current
year) rather densely covered with ± straight, sub¬
appressed to slightly ascending hairs, at the lowest
218
Novon
internode with mostly spreading white hairs, with a
few black hairs mixed in. Stipules greenish, 3-4
mm long, the lower ones shortly connate behind tin*
stem, the upper ones free. Leaves 2.5^1 cm long;
petiole 0.5—1 cm long, like the rachis loosely cov¬
ered with tangled, subappressed to ascending white
and also black hairs. Leaflets in 7 to 9 pairs, nar¬
rowly elliptic, 4—7 X ca. 2 nnn, at the apex mostly
rounded, on upper side glabrous, on underside
rather densely to densely covered with subap¬
pressed hairs up to 0.5 mm long, at the midvein
and the tip often with black hairs mixed in. Pe¬
duncle 3—5 cm long, rather densely to densely cov¬
ered with subappressed to ascending hairs, at the
base mostly with white, in upper part mostly with
predominantly black hairs. Raceme globular to
ovoid, densely many-flowered, ca. 1.5 cm long.
Bracts membranous, very narrowly triangular, 2.5—
3 mm long, black hairy. Pedicels 0.5—1 mm long,
black hairy. Calyx 4 mm long, campanulate, dense¬
ly covered with ascending to spreading, tangled
black hairs up to 0.6 mm long; teeth narrowly tri¬
angular, 1 .5—2 mm long, densely black hairy on
inner side. Petals pale pink and white. Standard 5-
6 mm long; blade ca. 3.5 mm wide, widely elliptic,
shallowly emarginate at the apex, very shortly nar¬
rowed at the base without distinct claw. Wings 4.5
mm long; blades obovate-spathulate, widely round¬
ed at the apex, ca. 3X2 mm; auricle ca. 0.5 mm
long, claw 1.5 mm long. Keel 4 mm long; blades
oblique-elliptic, with widely curved lower edge and
slightly curved upper edge, subacute at the apex,
2.5 X 1.5 mm; auricle indistinct, claws 1.5 mm
long, connate for nearly the entire length. Stamen
tube obliquely cut at the mouth. Ovary subsessile,
ovoid, glabrous; style short. Legume unknown.
Astragalus barclayanus differs from A. chateri
Vassilczenko by the indumentum of white and
black hairs 0.2— 0.3(— 0.5) mm long, in the inflores¬
cence of black hairs up to 0.7 mm long (vs. of only
white hairs 0.3— 0.8 mm long), by stipules 3—4 (vs.
4—6) mm long, shortly connate or the upper ones
free from each other (vs. all connate for half of the
length), leaflets glabrous (vs. sparsely to loosely
hairy) on upper side, bracts very narrowly trian¬
gular, 2.5—3 mm long (vs. ovate-acuminate, 3-5
mm long), calyx 4 (vs. 5—6) mm long, covered with
ascending to spreading (vs. subappressed) hairs,
standard 5—6 (vs. 7— 8) mm long, ovaries glabrous
(vs. hairy).
Astragalus goluhnjensis Podlech & L. R. Xu, sp.
nov. TYPE: [China.] Prov. Xizang [Qinghai ?]:
Kam [Qamdo], river Kam, river Goluboj, near
Nry-tschgo, 3590 m, 25 July 1900, Ladygin
358 (holotype, LE).
Differt ab A. densifloro planta pilis 0.2— 0.3 mm (nee
0.4—0.5 mm) longis obtecta, caulibus ad f8 cm (nec 2-
5(— 7) cm) longis, sparsissime pilis albis nigrisque (nec
laxe ad densiuscule pilis albis) obteetis, dentibus calycis
anguste triangularibus 1—1.5 mm longis (nec subulatis,
1.5-2 mm longis), petalis in siceo albidis (nec pallide
violaceis), vexillo ca. 8 mm longo et 5 mm lato (nec 6—7
mm longo et 3—3.5 mm lato), alarum laminis apice oblique
emarginate (nec obtusa), ovario glabro (nec piloso).
Plants ca. 25 cm tall, covered with appressed
hairs 0.2— 0.3 mm long. Caudex up to 15 mm thick,
branched. Stems up to 18 cm long, slender, pros¬
trate to ascending or erect, angular-sulcate, fur¬
nished with scattered appressed white and black
hairs. Stipules narrowly triangular, 3—4 mm long,
very shortly adnate to the petiole, shortly vaginate-
connate behind the stem, with scattered black
hairs, at the margins with minute sesssile glands.
Leaves 2.5—4 cm long, sessile; rachis distinctly
grooved on upper side, loosely covered with ap¬
pressed white hairs. Leaflets in 5 or 6 pairs, nar¬
rowly elliptic, 6—14 X 1—3 mm, obtuse to minutely
emarginate at the apex, on upper side glabrous, on
underside sparsely to loosely hairy, nearly always
with uprolled margins or folded, often curved. Pe¬
duncle 3-9 cm long, sulcate, sparsely but below
the raceme more densely white and predominantly
black hairy. Raceme capitate to somewhat elongat¬
ed. densely many-flowered. Bracts whitish mem¬
branous, narrowly triangular, 2-3 mm long, with
subappressed to ascending predominantly black
hairs. Flowers nearly sessile. Calyx ca. 3 mm long,
campanulate, rather densely to densely covered
with subappressed to slightly ascending predomi¬
nantly black hairs; teeth narrowly triangular, 1 — 1.5
mm long. Petals in dry state whitish. Standard ca.
8 mm long; blade ca. 5 mm wide, widely elliptic,
slightly emarginate at the apex, at the base nar¬
rowed without distinct claw. Wings ca. 7 mm long;
blades narrowly oblong, obliquely laterally incised
at the apex, 4.5 X 1.2 mm; auricle ca. 0.5 mm long,
claw 2 mm long. Keel 5 mm long; blades obliquely
obovate, with lower edge rectangularly curved in
distal part and upper edge straight, subobtuse at
the apex, 3 X 1 .5 mm; auricle ca. 0.3 mm long,
claw 2 mm long. Stamen tube truncate at the
mouth. Ovary shortly stipitate, glabrous. Legume
unknown.
Astragalus golubojensis differs from A. densiflorus
Karelin & Kirilow by hairs 0.2— 0.3 (vs. 0.4— 0.5)
mm long, the stems up to 18 (vs. 2— 5(— 7)) cm long,
covered very sparsely with white and black hairs
(vs. loosely to rather densely with only white hairs),
petals whitish (vs. pale violet) when dry, standard
ca. 8X5 mm (vs. 6-7 X 3—3.5 mm), limbs of
Volume 14, Number 2
2004
Podlech & Xu
Astragalus from China
219
wings obliquely emarginate at the tip (vs. obtuse)
and the ovaries glabrous (vs. hairy).
Astragalus mieheorum Podlech & L. R. Xu. sp.
nov. TYPE: China. Prov. Qinghai: Kunlun
Shan, Kunlun Shankou, SW of Golmud, 4690
m. 35°41'N, 94°03'E, 2 Aug. 1993, G. & S.
Miehe 9392/OH (holotype, MSB).
Differt ah A. densifloro caule, rachidibus pedunculisque
pilis oblique patentibus ad patentibus (nec appressis) 0.5—
0.8 mm longis (nec 0.4— 0.5 mm longis) obtectis, foliolis
subtus pilis ascendentibus (nec appressis) obtectis, calyce
5—6 (nec 3—5) mm longo, vexillo late ovato, 8—9.5 mm
longo et 5-6.5 mm lato (nec elliptica, 6—7 mm longa et
ad 3.5 mm lata).
Plants 7—12 cm tall. Caudex with ± elongated,
slender stolons partly subterranean, partly aerial.
Stems 1—2 cm long, slender, prostrate to ascending,
densely covered with ascending to spreading, white
or black and white hairs 0. 5-0.8 mm long. Stipules
greenish, narrowly triangular, 3—6 mm long, adnate
to the petiole for ca. 1 mm. the lower ones shortly
vaginate-connate behind the stem, the upper ones
nearly free from each other, sparsely predominantly
black hairy, at the base of margins with minute ses¬
sile glands. Leaves 2— 4(— 5.5) cm long; petiole 0.5—
1 cm long, with the rachis loosely to rather densely
hairy like the stem. Leaflets in (3)4 to 6 pairs, nar¬
rowly elliptic, 5—14 X 1.5— 4.5 nun. obtuse to acute
at the apex, glabrous on upper side, on underside
rather densely covered with tangled, ascending
white hairs up to 1 mm long. Peduncle 2^4 cm
long, densely covered with ascending to spreading,
tangled white or black and white, sometimes only
black hairs up to 1 mm long. Raceme ovate, 1.5—
3 cm long, densely many flowered. Bracts white-
membranous, linear-acute, 4-6 mm long, black
hairy. Flowers nearly sessile. Calyx 5—6 mm long,
eampanulate, ± densely covered with ascending to
nearly spreading, tangled black hairs up to 1 mm
long; teeth subulate, ( 1 .5 — )2 — 3.5 mm long. Petals
violet, in dry state often pale yellowish brown. Stan¬
dard 8—9.5 mm long; blade w idely ovate, 5-6.5 mm
wide, widely retuse to slightly emarginate at the
apex, at the base subabruptly narrowed into the
cuneate claw'. Wings 6.5— 7(— 8) mm long; blades
narrowly oblong, gradually widened toward the
truncate-incised apex, 4—4.5 X 2—2.2 mm; auricle
minute, claw 2.5—3 mm long. Keel 5.5—6 mm long;
blades oblique-obovate, with widelv nearly rectan¬
gular-curved lower edge and nearly straight upper
edge, subacute at the apex, ca. 3X2 mm; auricle
minute, claws 2.5—3 mm long, connate for nearly
the entire length. Stamen tube truncate at the
mouth. Ovary sessile to subsessile. widely ovate to
nearly globular, white hairy; style thick, short, gla¬
brous. Legume sessile, globose, ca. 4 mm thick,
rounded at the apex without beak but with the per¬
sistent glabrous style, bilocular; valves densely cov¬
ered with ascending to spreading, tangled, partly
straight partly curly white and black hairs up to 1
mm long. Seeds probably two.
Astragalus mieheorum differs from A. densiflorus
Karelin & KirilowT by the stem, rachis, and pedun¬
cle covered with ascending to spreading hairs 0.5—
0.8 mm long (vs. by appressed hairs 0.4— 0.5 mm
long), by leaflets covered on the underside with as¬
cending (vs. appressed) hairs, by the calyx 5—6 (vs.
3—5) mm long, and by the standard widely ovate,
8-9.5 mm long and 5—6.5 mm wide (vs. elliptic, 6-
7 mm long and up to 3.5 mm wide).
Etymology. Named alter Prof. (1. Miehe and
Mrs. S. Miehe, University of Gottingen, explorers
of the vegetation of Tibet.
Paratypes. CHINA. Qinghai: Kunlun Shan, Kunlun
Shankou, SVi of Golmud, 4870 m, 35°40'N, 94°03'E. 3
Aug. 1993, G. & S. Miehe 9394/06 (MSB); Kunlun Shan,
E of Kunlun Shankou, SVi of Golmud, 4820 in. 35°41'N,
94°14'E, 1 Aug. 1993, G. & S. Miehe 9374/14 (MSB); NE
Tibet, Northeastern Plateau along Golmud-Lhasa highway,
Kunlun Shankou-Qumar Heyan, 4570 m, 35°30'N,
93°45'E, 3 Aug. 1993, G. & S. Miehe 9403/12 (MSB);
between Madoi and Bayanka La. 4450 m. 34°20'N,
97°55'E, 3 Aug. 1998, G. Miehe, ./. Q. Lm & Sonamco
98-35221 (MSB).
Astragalus naiislianieus Podlech & L. R. Xu, sp.
nov. TYPE: China. Prov. Qinghai: Qinghai Nan
Shan, Caka Basin, 36°40'N. 98°45'E. 3520 m,
27 July 1993, G. & S. Miehe 9329/07 (holo¬
type. MSB).
Differt ah A longiracemoso stipulis 2.5—3 (nec 3-7) mm
longis, foliis 2-4.5 (nec 3-8) cm longis, 4—6 (nec 6-9)
jugis, foliolis 4—8 X 1—3 (nec 9—27 X 2—7) mm, pedun-
culo mere albi-piloso (nec in parte superiore saepissime
mere nigri-piloso), axi inflorescentiae albi-piloso (nec
praecipue nigri-piloso), vexillo ca. 7 (nec 8—10) mm longo.
Plants tall, up to 30 cm or more, basal parts
unknown. Stem up to 30 cm long, slender, loosely
to rather densely covered with irregularly spreading
white hairs and sometimes below the nodes with
few black hairs 0.15—0.3 mm long. Stipules green¬
ish, narrowly triangular, 2.5—3 mm long, free from
petiole, loosely to rather densely covered with as¬
cending to nearly spreading white, at the base also
black hairs, at the base of margins with minute,
sessile glands. Leaves 2—4.5 cm long; petiole 0.3—
I cm long, together with the rachis white hairy like
the stem. Leaflets in 4 to 6 pairs, narrowly elliptic
to narrowly ovate, 4—8 X 1—3 mm, rounded to
slightly emarginate at the apex, on upper side gla-
220
Novon
brous, on underside rather densely covered vvilh
subappressed to ascending hairs up to 0.3 mm long,
mostly folded. Peduncles 1-6 cm long, white hairy
like the stem. Racemes at anthesis short, densely
many-flowered, distinctly elongating in fruit; axis
white hairy. Bracts whitish membranous, narrowly
triangular, 2-2.5 mm long, sparsely black hairy.
Pedicels 0.3-0. 5 mm long, black hairy. Calyx 3 mm
long, campanulate, obliquely cut at the mouth,
rather densely covered with subappressed, black
and white hairs up to 0.5 mm long; teeth narrowly
triangular, 1 — 1.5 mm long, densely white hairy on
inner side. Petals violet with darker tip of keel.
Standard ea. 7 mm long; blade ca. 4.5 mm wide,
widely elliptic, emarginate at the apex, gradually
shortly narrowed at the base without a distinct claw.
Wings ca. 5.5 mm long; blades obovate, widened
toward the rounded apex, 3.5 X 2 mm; auricle 0.5
mm long, claw ca. 2 mm long, curved. Keel ca. 4
mm long; blades half-circular, with widely curved
lower edge and straight upper edge, subacute at the
apex, 2.5 X 1.5 mm; auricle minute, acute, claws
ca. 1 .5 mm long, connate for nearly the whole
length. Stamen tube ± truncate at the mouth. Ovary
sessile, nearly globular, glabrous; style thick, short.
Legumes unknown.
Astragalus nanshanicus differs from A. longira-
cemosus IN. Ulziykhutag by stipules 2.5-3 (vs. 3—7)
mm long, leaves 2-4.5 (vs. 3-8) cm long, leaflets
in 4 to 6 (vs. 6 to 9) pairs, 4—8 X 1—3 (vs. 9—27
X 2—7) mm, peduncles only white hairy (vs. in up¬
per part mostly merely black hairs), inflorescence
axis white (vs. predominantly black) hairy, standard
ca. 7 (vs. 8—10) mm long.
Astragalus pseiidujagnobieiis Podlech & L. R.
Xu, sp. nov. TYPE: China. Kaschgaria, Ugu,
Ulug-tuz, valley of river Tscharlym, 28 June
1909, Divanogorskaja 284 (holotype, LE).
Differt ah A. jagnobico stipulis petiolo vix adnatis, bas-
alibus alte vaginanti-connatis (nec petiolo per 0.5-1 mm
adnatis, inter se liberis), foliolis 4—6 (nee 6-9) jugis, vex-
illo ca. 10 mm longo et 5 mm lata (nec 5—6 mm longo et
2. 2-2. 5 mm lata), lamina alarum obovata, 4.5-5 X 2 mm
(nec anguste oblonga, ca. 3X2 mm).
Plants up to 25 cm tall, with appressed to sub¬
appressed hairs 0.1— 0.3 mm long. Caudex up to
15 mm diam., with a pluricipital root-crown. Stems
several, prostrate to ascending, slender, striate-sul-
cate, sparsely to loosely covered with white, below
the nodes also with black hairs. Stipules greenish,
ovate to triangular, white and black hairy, nearly
free from the petiole, the lower ones highly connate
behind the stem, the middle and upper ones free
from each other. Leaves 4-7 cm long; petiole 1-2.5
cm long, like the raehis slender, loosely covered
with white and interspersed black hairs. Leaflets in
4 to 6 pairs, narrowly elliptic to elliptic, 5-12 X
3—6 mm, rounded to distinctly retuse at the apex,
on upper side glabrous, on underside loosely to
rather densely appressed white hairy. Peduncles 6—
10 cm long, slender, ascending-curved, striate-sul-
cate, loosely covered in lower part with merely
white hairs, in upper part also with increasing
black hairs. Raceme at first globose-ovoid, soon
elongating and up to 4 cm long, many-flowered.
Bracts whitish membranous, narrowly triangular,
1.5— 2 mm long, black hair)'. Pedicels ca. 0.5 mm
long, densely black hairy. Calyx 3-4 mm long, cam¬
panulate, obliquely cut at the mouth, loosely to
rather densely covered with black and few white
hairs; teeth narrowly triangular. 1 .2-1.5 mm long.
Petals in dry state whitish to slightly pale-violet-
suffused, standard sometimes with pale violet lon¬
gitudinal nerves, keel in upper pari mostly violet.
Standard ea. 10 mm long; blade slightly obovate,
ca. 5 mm wide, slightly emarginate at the apex,
gradually narrowed at the base. Wings 7 mm long;
blades obovate, widely rounded at the apex, 4.5-5
X 2 mm; auricle short, claw 2.5 mm long. Keel 5
mm long; blades obliquely obovate, lower edge
curved in distal part and upper edge ± straight,
subobtuse at the apex, 3.5 X 2 mm; auricle indis¬
tinct, c law ca. 1.5 mm long. Ovary ovoid. Legumes
unknown.
Astragalus pseudojagnobicus differs from A. jag-
nobicus Lipsky by the stipules scarcely adnate to
the petiole, the lower ones vaginate-connate high
up (vs. adnate to the petiole for 0.5—1 mm, free
from each other), leaflets in 4 to 6 (vs. 6 to 9) pairs,
standard ca. 10 X 5 (vs. 5—6 X 2. 2-2. 5) mm, limbs
of wings narrowly obovate, 4.5—5 X 2 mm (vs. nar¬
rowly oblong, ca. 3 X 2 mm).
Astragalus sagastaigolensis N. Ulziykhutag ex
Podlech & L. R. Xu, sp. nov. TYPE: [China.
Prov. Qinghai:] Dzhungaria, Sagastaigol, 2740
m, 5 Sep. 1879, A. Regel (holotype, LE).
Differt ab A. imetensi stipulis inter se liberis (nec fo-
liorum inferiorum alte vaginati-connatis) foliolis 6-9-juga
(nec 5— 6-juga), bracteis 4—5 (nec 3) mm longis, vexillo
8.5— 9.5 (nec 6—7) mm longo, lamina alarum obtriangulari
apice distincte emarginata, ca. 2.5 mm lata (nee anguste
oblonga apice obtusa, ca. 1.5 mm lata).
Plants ca. 15 cm tall. Caudex strong, up to 15
mm thick, with a pluricipital root-crown, with short
branches. Stems many, up to 10 cm long, prostrate
to ascending or erect, rather densely covered with
appressed to subappressed white hairs 0.4—0. 6 mm
long, with some black hairs mixed in especially at
Volume 14, Number 2
2004
Podlech & Xu
Astragalus from China
221
the nodes, glabrescent with age. Stipules whitish to
slightly greenish, 4—6 mm long, free from the pet¬
iole and from each other, sparsely to loosely cov¬
ered with suhappressed, predominantly black hairs,
at the margins with minute, sessile glands. Leaves
3— 6 cm long, subsessile; rachis hairy like the stem.
Leaflets in (5)6 to 9 pairs, narrowly elliptic, 8—15
X 2-5 mm, rounded to narrowly obtuse at the apex,
on upper side glabrous, on underside rather dense¬
ly covered with appressed white hairs 0.3— 0.5 mm
long. Peduncle 2.5—5 cm long, slightly angular-sul-
cate, white hairy like the stem, toward the raceme
with increasing black hairs. Racemes 2—3 cm long,
densely many-flowered; axis with ascending black
hairs. Bracts whitish membranous, narrowly trian¬
gular, acuminate, 4—5 mm long, with subappressed
to ascending, predominantly black hairs. Pedicels
ca. 1 mm long, recurved, black hairy. Calyx 4^4.5
mm long, campanulate, loosely to rather densely
covered with subappressed to ascending black
hairs ca. 0.5 mm long; teeth subulate, ca. 2 mm
long, black hairy on inner side. Petals whitish when
dry. Standard 8.5— 9.5 mm long; blade elliptic, ca.
4 mm wide, rounded at the apex, at the base sub-
abruptly narrowed into the short claw. Wings ca. 7
mm long; blades obtriangular, distinctly dilated to
the unequally widely emarginate apex, 4.5 X 2.5
mm; auricle ca. 0.3 mm long, claw 2 mm long. Keel
5 mm long; blades oblique-elliptic, with widely
rectangular-curved lower edge and slightly curved
upper edge, subobtuse at the apex, ca. 3X2 mm;
auricle minute, claw 2 mm long. Ovary shortly stip-
itate, densely white hairy; style glabrous. Legumes
sessile, globose, ca. 3 mm diam., rounded ventrally,
deeply grooved dorsally, rounded at the apex, beak¬
less, bilocular; valves pale brownish, rather densely
covered with subappressed to slightly ascending
while and black hairs up to 0.8 mm long. Seeds
one in each locule, 2—2.5 mm long, pale brownish,
smooth.
Astragalus sagastaigolensis differs from A. ime-
tensis Borissova by the stipules all free from each
other (vs. those of lower leaves vaginate-connate
high up), leaflets in 6 to 9 (vs. 5 or 6) pairs, bracts
4— 5 (vs. 3) mm long, standard 8.5— 9.5 (vs. 6—7)
mm long, limbs of wings obtriangular, distinctly
emarginate, ca. 2.5 mm wide (vs. narrowly oblong,
obtuse, ca. 1.5 mm wide).
Astragalus L. sect. Cenantrum Bunge, Mem.
Acad. Imp. Sci. Saint Petersburg 11(16): 23.
1868. TYPE: Astragalus frigidus (L.) A. Gray
(lectotype, designated by Podlech (1990:
468)).
Astragalus purpurinus (Y. C. Ho) Podlech & L.
R. Xu, stat. nov. Based on Astragalus mem-
branaceus Bunge var. purpurinus Y. C. Ho,
Bull. Bot. Lab. North-East. Forest. Inst.
1980(8): 54. 1880. TYPE: [China.] Gansu:
Xiaho Xian [Hsiahohsien], 2600 m, 1 July
1937, T. P. Wang 6925 (holotype, WUK).
Because of the short original diagnosis of this
taxon a complete description is given here.
Astragalus purpurinus differs from A. mongholi-
cus Bunge by the strong stem up to 8 (vs. 2.5—3)
mm diam., shorter (4(-5) vs. 5—9 mm long) and
more densely hairy calyx with shorter teeth (0.3-1
vs. 0.5— 1.5 mm long), purple petals and legumes
with a stipe only 4—5 (vs. 6—15) mm long.
Plants at least 60—70 cm tall, covered with hairs
0.3— 0.5(— 1.5) mm long. Stem erect, at the base up
to 8 mm diam., angular-sulcate, glabrous or with
scattered spreading hairs up to 1.5 mm long. Stip¬
ules green, very narrowly to narrowly triangular,
free, the lower ones 8—10, the upper ones 4—5 mm
long, glabrous or ciliate at the margin. Leaves 5-9
cm long, nearly sessile; rachis grooved on upper
side, sparsely covered with spreading hairs up to 1
mm long. Leaflets in 3 to 9 pairs, narrowly ovate to
narrowly elliptic, rounded to minutely acuminate at
the apex, 10—17 X 3—7 mm, on upper side loosely
covered with appressed to subappressed hairs 0.3—
0.5 mm long, on underside loosely with appressed
to subappressed hairs 0.5—1 mm long. Peduncles
5-13 cm long, sparsely to loosely covered with ap¬
pressed to ± spreading white hairs. Racemes loose¬
ly many-flowered, 4-5 cm long, soon elongating and
at least up to 10 cm long; raceme axis black hairy.
Bracts green, linear-subulate, 5—8 mm long, ca. 0.5
mm wide, loosely mainly white ciliate. Pedicels 1—
1.5 mm long, later on elongating and up to 3 mm
long, with minute black hairs. Bracteoles whitish,
linear, 0.5—1 mm long, in upper part of the pedicel.
Calyx campanulate, 4(— 5) mm long, slightly gib¬
bous at the base, strongly obliquely cut at the
mouth, the tube loosely covered with short black
subappressed hairs up to 0.3 mm long and with few
long white hairs, at the upper margin and the teeth
more densely black hairy; teeth narrowly triangular
to triangular, 0.3—1 mm long. Petals lilac or dark
purplish at the apex and whitish at the base. Stan¬
dard 13 mm long; blade distinctly recurved, ob-
ovate, 7 mm wide, emarginate at the apex, gradu¬
ally and shortly cuneately narrowed at the base.
Wings 13 mm long, distinctly bent at the base of
the blade; blades narrowly oblong, rounded at the
apex, 5 X 1.5 mm; auricle 0.8 mm long, claw 7—8
mm long, straight. Keel 12 mm long; blades strong-
222
No von
ly curved-elliptic, with widely and nearly rectan¬
gularly curved lower edge and distinctly concave
upper edge, obtuse at the apex, 5X2 mm; auricle
ca. 0.3 mm long, claw 7 mm long. Stamen tube
obliquely cut at the mouth. Ovary with a stipe ca.
5 mm long, densely white hairy. Legume with a
slender stipe 4—5 mm long, narrowly elliptic, acu¬
minate at both ends; valves loosely to densely cov¬
ered with subappressed white and black hairs ca.
0.3 mm long, glabrescent with age and sometimes
subglabrous.
Ollier specimen seen. CHINA. Gansu: Yuzhong, 2750
m, 8 Aug. 1959, Y. He 5864 (WUK).
Astragalus xitaibaieus (k. T. Ku) Podlech & L.-
R. Xu. stat. nov. Based on Astragalus mona-
delphus Bunge subsp. xitaibaieus K. T. Fu,
Aeta Hot. Bor.-Occ. Sin. 1(2): 17. 1981. TYPE:
[China.] Shaanxi: Xitaiba Shan, Ding-peng-
shan, 3300 m, Fu Kun-tsun 10321 (holotype,
WUK).
Because of the short original diagnosis of this
taxon a complete description is given here.
Astragalus xitaibaieus differs from A. monadel-
phus by being totally glabrous with the exceptions
of the inner side of the calyx teeth and sometimes
the bracts (vs. at least hairy on underside of leaf¬
lets, calyx, and legumes) and by the presence of
stipel-like entire or fringed, linear to narrowly tri¬
angular emergences 0.5— 2.5 mm long and/or of dis¬
tinctly stalked minute glands at the base of the leaf¬
lets.
Plants 12-22 cm tall, totally glabrous with the
exception of the inner side ol the calyx teeth and
sometimes the bracts. Caudex strong, 15 mm thick,
with a pluricipital root-crown. Stems several, as¬
cending to erect, up to 4 mm thick, angular-striate,
10—17 cm long. Stipules greenish membranous with
narrowly hyaline margins, ovate, obtuse, 12-15 mm
long and up to 10 mm wide, connate at the base,
free from the petiole. Leaves 4—8 cm long; petiole
0.7—2 cm long, at the base of the leaflets with sti¬
pel-like entire or fringed, linear to narrowly trian¬
gular emergences 0.5-2. 5 mm long and/or with dis¬
tinctly stalked minute glands. Leaflets in 5 to 7
pairs, ovate to elliptic, 10-18 X 5-9 mm, at the
apex rounded and sometimes minutely mucronu-
late. Peduncles 4—7.5 cm long. Racemes 8- to 11-
fiowered. Bracts narrowly ovate, acute, ca. 10 mm
long, sometimes sparsely ciliate at the margins.
Pedicels 2—3 mm long. Calyx campanulate, 7.5—9
mm long; teeth subulate, 2.5-3 mm long, hairy on
inner side. Petals yellow. Standard ca. 1 1 mm long;
blade obovate, 5—6 mm wide, emarginate at the
apex, at the base subabruptly contracted into the
rather long claw. Wings ca. 1 1 mm long; blades
narrowly oblong, obtuse at the apex, ca. 4 X 1.5
mm; auricle short, claw 6.5-7 mm long, keel ca.
10 mm long; blades obliquely elliptic, ca. 3.5 X 2
mm; auricle indistinct, claw ca. 6.5 mm long. Sta¬
men tube closed (monadelphous), slightly obliquely
cut at the mouth. Ovary stipitate, hairy. Legumes
with a stipe 5—6 mm long, narrowly elliptic, shortly
attenuate at both ends, ea. 15 mm long, unilocular;
valves hairy.
Paratype. CHINA. Shaanxi: Xitaihai Shan. Hun-ren-
ping, in summo alpino, 3300 m, 4 July 1959, Yang Jin-
xiang 645 (WUK).
Astragalus sect. Corethrum Bunge, Mem. Acad.
Imp. Sci. Saint Petersbourg 11(16): 98. 1868.
T\ PE: Astragalus scoparius C. A. Meyer (lec-
totype, designated by Podlech (1990: 471)).
Astragalus eliaidamueiisis (S. B. Ho) Podlech &
L. R. Xu, stat. nov. Based on Astragalus kro-
nenburgii B. Fedtschenko var. ehaidamuensis
S. B. Ho, Bull. Bot. Res. North-East. Forest.
Inst. 3(1): 42. 1983. TYPE: [China.] Qinghai:
Haixi Mongolzu Zangzu, Kazakzu Zizhizhou,
Delingha, 3290 m, 21 July 1959. Exped. Gan¬
su et Qinghai Acad. Sin. 80S (holotype, WUk).
Because of the very short original diagnosis of
this taxon a complete description is given here.
Astragalus ehaidamuensis differs from A. kronen-
burgii B. Fedtschenko by stipnles triangular, 2 mm
long (vs. narrowly triangular, 3-5 mm long), adnate
to the petiole for 0.5 (vs. 1—2) mm, densely hairy
(vs. subglabrous to loosely hairy), leaflet in up to 4
(vs. in 3 to 6) pairs, peduncles 5-14 (vs. 9-22) cm
long, only white (vs. toward the raceme increasingly
black) hairy, raceme very short, 3- to 7-flowered
(vs. 3-5 cm long, 10- to 16-flowered), bracts 1-2
(vs. 3—4) mm long, calyx 10 (vs. 5-6) mm long,
teeth 2.2-3 (vs. 1-2) mm long.
Plants ca. 18 cm tall, nearly acaulescent, caes-
pitose, covered in vegetative parts with medifixed,
appressed white hairs 0.3—1 mm long. Caudex ca.
8 mm diam., at the apex strongly divided w ith short
branches, covered with brown bark. Stems
branched at the base, parts of the current year, if
developed, up to 0.5 cm long, densely hairy. Stip¬
ules ca. 2 mm long, triangular, adnate to the petiole
for ca. 0.5 mm, densely hairy. Leaves of sterile
shoots 1—2 cm long; petiole 0.5—1 cm long, with 1
or 2 pairs of small leaflets, 2—3 mm long. Leaves
of the flowering shoots 3—7 cm long; petiole 1—2 cm
long, like the rachis slender, densely hairy. Leaflets
in (2)3 to 4 pairs, linear, 10—15 X 1—1.5 mm.
Volume 14, Number 2
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Podlech & Xu
Astragalus from China
223
acute-tipped, on both sides rather densely to dense¬
ly hairy. Peduncles 5-14 cm long, loosely to rather
densely white hairy. Racemes short, subumbellate,
3- to 7-How ered. Bracts membranous, narrowly tri¬
angular, 1—2 mm long, sparsely furnished with
mostly basifixed white and black hairs. Pedicels ea.
0.5 mm long, white and black hairy. Calyx ca. 10
mm long, tubular, obliquely cut at the mouth, loose¬
ly to rather densely covered with medifixed ap-
pressed white hairs 0.5-1 mm long and mostly
slightly shorter black hairs; teeth subulate, 2.2-3
mm long. Petals probably yellow. Standard 15-17
mm long, elliptic, 6-7 mm wide, emarginate at the
apex, with a very short, indistinct claw. Wings 15
mm long; blades narrowly oblong, at the apex deep¬
ly incised, 7X2 mm; auricle 0.5 mm long, claw
8 mm long. Keel ca. 12 mm long; blades obliquely
obovate, with rectangular-curved lower edge, and
nearly straight upper edge, subacute at the apex, 5
X 2.5 mm; auricle acute, 0.3 mm long, claw 7 mm
long. Stamen tube obliquely cut at the mouth. Ova¬
ry with a stipe ca. 2 mm long, linear, white hairy;
style glabrous. Legumes unknown.
Astragalus sect. Ilcmipliaea Bunge, Mem. Acad.
Imp. Sei. Saint Petersbourg 1.1(16): 20. 1868.
TYPE: Astragalus hemiphaca Karelin & Kiri-
low.
Astragalus nigrodentatus N. Ulziykhutag ex Pod¬
lech & L. R. Xu. sp. nov. TYPE: China. Prov.
Xizang: Distr. Kam, Dabo-Shan pass, 18 July
1893, G. N. Potanin (holotype, LE).
Differt ab A. alaschano indumento e pil is 0.6— 0.8(— 1)
mm (nee 0.1— 0.3 mm) longis consistente, stipulis 6—7 (nec
2-3) mm longis, foliolis anguste ovatis ad anguste ellip-
ticis, 5—14 X 2.5—5 mm (nec ellipticis ad obovatis, 3—7
X 2—5 mm), bracteis 4—6 mm (nec 0.8—1 mm) longis,
calyce ca. 5 mm (nec 2.5—3 mm) longis.
Plants 8-10 cm tall, covered in vegetative parts
with flattened, long-acuminate white hairs widest
near the base and elsewhere with slender black
hairs mixed in. Stems ascending, at the base pale,
stolon-like, glabrous, with few7, remote leafless stip¬
ules, in upper part sulcate, loosely to rather densely
covered with stiff, tangled, ascending to spreading
while, toward the nodes mostly black hairs 0.6— 0.8
nun long. Stipules greenish. 6—7 mm long, narrowly
triangular to triangular, free from the petiole, those
without developed leaves connate up to the middle,
the other ones free from each other, the lower ones
glabrous, the upper ones loosely or, especially to¬
ward the margins and tip, densely white and partly
also black hairy. Leaves 3—5 cm long; petiole 0.8—
1.5 cm long, distinctly grooved on the upper side,
similar to the rachis loosely white hairy like the
stem. Leaflets in 6 to 8 pairs, narrowly ovate to
narrowly elliptic, 5-14 X 2.5—5 mm, obtuse at the
apex, on upper side glabrous but with densely white
hairy margins, on underside loosely covered with
appressed hairs 0.2— 0.5 mm long. Peduncles 1.5—
3.5 cm long, sulcate, loosely to densely covered
with subappressed black hairs 0.3-1 mm long, at
the base with some white hairs mixed in. Racemes
short, densely many-flowered. Bracts whitish mem¬
branous, narrowly triangular. 4—6 mm long, with
predominantly black hairs, at the base with sessile,
minute glands. Elowers subsessile. Calyx ca. 5 mm
long, campanulate, densely covered with ascending
to partly nearly spreading black hairs 0.3— 0.6 mm
long; teeth very narrowly triangular, ca. 2 mm long,
hairy on inner side. Petals probably violet in life,
pale yellowish brown when dry. Standard ca. 9 mm
long; blade ca. 4 mm wide, oblong, at the apex
widely and deeply incised, at the base cuneately
narrowed without distinct claw. Wings ca. 7.5 mm
long; blades obtri angular, widest at the widely and
deeply incised apex, 4.5 X 2 mm; auricle 0.3— 0.4
mm long, claw 3 mm long, curved. Keel 6.5 mm
long; blades oblique-elliptic, with widely curved
lower edge and straight upper edge, subacute at the
apex, 3.5 X 2 mm; auricle ca. 0.3 mm long, acute,
claw ca. 3 mm long. Stamen tube truncate at the
mouth. Ovary subsessile, minute, elliptic, glabrous,
2-ovulate. Legumes unknown.
Astragalus nigrodentatus differs from A. alas-
chanus Bunge by the hairs 0.6— 0.8(-l) (vs. 0.1 —
0.3) mm long, stipules 6-7 (vs. 2—3) mm long, leaf¬
lets narrowly ovate to narrowly elliptic, 5—14 X
2.5— 5 mm (vs. elliptic to obovate, 3—7 X 2—5 mm),
bracts 4—6 (vs. 0.8—1) mm long, calyx ca. 5 (vs.
2.5— 3) mm long.
This collection was named Astragalus nigroden¬
tatus by Ulziykhutag in the herbarium at St. Pe¬
tersburg (LE) without description.
Astragalus sect. Laguropsis Bunge, Mem. Acad.
Imp. Sci. Saint Petersbourg 11(16): 137. 1868.
TYPE: Astragalus laguroides Pallas (lectotype,
designated by Podlech (1990: 479)).
Astragalus iiovissiinus Podlech & L. R. Xu. nom.
et stat. nov. Based on Astragalus laguroides
Pallas var. micranthus S. B. Ho, Bull. Bot. Res.
North-East. Forest. Inst. 3(4): 57. 1983 “non
A. micranthus Desv., 181 4.” TYPE: |China.|
Nei Mongol, Baganmaotasum (patria ignota),
29 May 1931, T. N. Liou 2147 (holotype,
WUK). Figure: Ho (1983: 66, fig. 5; as A. la¬
guroides var. micranthus).
224
Novon
Because of the short original diagnosis of this
taxon a complete description is given here.
Plants 8-12 cm tall, acaulescent, with merely
white, in vegetative parts distinctly warty, hairs.
Caudex slender, simple or with a pluricipital root-
crown. Stipules whitish, 3—4 mm long, triangular,
nearly free from the petiole, appressed hairy, at the
margins with basifixed hairs. Leaves 3-6 cm long;
petiole 1-3 cm long, like the rachis slender, loosely
to rather densely covered with medifixed, appressed
hairs 0.5—1 mm long. Leaflets in 2 to 4 pairs, in
the basal leaves often only in 1 or 2 pairs, narrowly
elliptic to elliptic, 7-12 X 3-5 mm, acute at the
apex, on both sides rather densely to densely cov¬
ered with medifixed, appressed hairs 1.5— 2(— 3) mm
long. Peduncles 3.5-6 cm long, slightly angular-
sulcate, loosely to rather densely covered w ith med¬
ifixed, appressed hairs 0.5—1 mm long, partly gla-
brescent with age. Racemes ovoid, 3—5 cm long,
densely many-flowered. Bracts whitish, ca. 2 mm
long, narrowly triangular, covered with mostly bas¬
ifixed hairs. Flowers subsessile. Calyx 11-12 mm
long, tubular at beginning of anthesis, soon ovoid-
inflated, rather densely covered with subbasifixed
to basifixed, rigid, spreading, straight white hairs
2-3 mm long; teeth subulate, 3-4 mm long. Stan¬
dard ca. 14 mm long; blade ca. 5 mm wide, elliptic,
narrowly triangular toward the slightly emarginate
apex, slightly constricted below the middle, at the
base obtusely angularly passing into the claw.
Wings 13 mm long; blades narrowly oblong, round-
tipped, ca. 5 X 1.5 mm; auricle 0.5 mm long, claw
8 mm long. Keel 12 mm long; blades obliquely ob-
ovate, with rectangular-curved lower edge and
nearly straight upper edge, acute-tipped, 3.5 X 1.5
mm; auricle acute, 0.5 mm long, claw ca. 8 mm
long. Stamen tube truncate at the mouth. Ovary ses¬
sile, hairy. Legumes unknown.
Astragalus sect. Polyoladus Y. (7 Ho, Bull. Bot.
Res. North-East. Forest. Inst. 1(3): 107. 1981.
TYPE: Astragalus polycladus Bureau & Fran-
chet.
The following species was proposed as a new
species in Sendtnera 7: 198. 2001, but unfortu¬
nately without a Latin diagnosis.
Astragalus tihetieola Podlech & L. R. Xu, sp.
nov. TYPE: China. Prov. Xizang: S Tibet, Cha-
yul Dzong, Loro Chu, 23 Apr. 1936, F. Ludlow
& G. Sherrifj 1326 (holotype, BM).
Differt ab A. monbeigii caulibus pilis 0.4— 0.7 mm (nec
0.1— 0.3 mm) longis obtectis, calyce 4—5 mm longo tlen-
tibus ejus 1.5—2 mm longis (nec 5—6 mm longo dentibus
ejus 2.5—4 mm longis), vexillo ca. 6 mm longo et 5 mm
lato, lamina transverse elliptica (nec ca. 10.5 mm longo,
lamina late obovate), ovario sessile (nec stipite 1.5—2 mm
longo suffulto).
Plants 15—25 cm tall. Stems 10-17 cm long,
branched at the base, ascending to erect, striate-
sulcate, sparsely to loosely covered with appressed
to subappressed white, at the nodes also black,
hairs 0.4— 0.7 mm long. Stipules green, 6—9 mm
long, ovate-acuminate to narrowly triangular, free
from the petiole, behind the stem high up vaginate-
connate, sparsely to loosely covered with appressed
white and sometimes also with some black hairs.
Leaves 6—12 cm long; petiole 1—3 cm long, like the
rachis loosely covered with appressed white hairs
0.2-0. 5 mm long. Leaflets in 9 to 1 1 pairs, narrowly
elliptic to elliptic, 7—17 X 3-6 mm, rounded to
slightly emarginate at the apex, on upper side gla¬
brous, on underside sparsely and mainly at the
midrib furnished with appressed white hairs 0.5—1
mm long. Peduncles 5-6 cm long, hairy like the
stem but in upper part with predominantly blac k
hairs. Racemes ovoid, rather loosely 7- to 15-flow-
ered. Bracts whitish, linear-acute, 2.5—3 mm long,
predominantly black hairy. Pedicels ca. 1 mm long,
black hairy. Calyx eampanulate, 4—5 mm long,
slightly oblique at the base, slightly obliquely cut
at the mouth, rather densely covered with ap¬
pressed black hairs 0.2— 0.3 mm long and with few¬
er, slightly longer white hairs; teeth subulate, 1.5—
2 mm long. Petals mauve with whitish center, keel
purplish mauve. Standard ca. 6 mm long; blade 5
mm wide, transverse-elliptic, deeply incised at the
apex, at the base subabruptly contracted into the
very short claw. Wings ca. 6 mm long; blades ob¬
long, rounded at the apex, 3.5 X 2 mm; auricle
wide, ca. 0.5 mm long, claw 2.5 mm long. Keel 5
mm long; blades oblique-elliptic, rounded at the
apex, 3X2 mm; claw 2 mm long. Stamen tube
truncate at the mouth. Ovary sessile, white-hairy;
style glabrous. Legumes unknown.
Astragalus tihetieola differs from A. monbeigii N.
D. Simpson by stems covered with hairs 0.4— 0.7
(vs. 0.1— 0.3) mm long, calyx 4—5 mm long with
teeth 1.5—2 mm long (vs. 5—6 mm long with teeth
2.5— 4 mm long), standard ca. 6 X 5 mm, the limb
transversely elliptic (vs. ca. 10.5 mm long, the limb
widely obovate), ovaries sessile (vs. with a stipe
1 .5— 2 mm long).
Astragalus sect. Skythropos N. D. Simpson,
Notes Roy. Bot. Card. Edinburgh 8: 255. 1915.
TYPE: Astragalus skythropos Bunge (lectotype,
designated by Podlech (1990: 488)).
Volume 14, Number 2
2004
Podlech & Xu
Astragalus from China
225
Astragalus griersonii Podlech, sp. nov. TYPE:
Bhutan, Me La, 27°58'N, 91°37'E, 14,000 ft.,
7 Aug. 1933, F. Ludlow & G. Sherrijf 429 fho-
lotype, BM).
Differt ab A. kongrensi plantis 20—25 (nec 6—14) cm
altis, in partibus vegetativis pilis praecipue nigris (nec
mere albis) obtectis, calyce ca. 8 (nec 9—12.5) mm longo,
campanulati-tubuloso (nec tubuloso), dentibus 3—4 (nec
4—7) mm longis, petalis brevioribus.
Plants 20—25 cm tall, with subappressed to ±
spreading, thin, long white and black hairs. Caudex
with rather long stolons up to 10 cm long, furnished
with leafless, pale, scale-like stipules. Stems erect,
short, terete to angular-sulcate, loosely to rather
densely covered with subappressed to mostly ±
spreading, predominantly black hairs 0.3—1 mm
long. Stipules 8—12 mm long, dirty whitish to
slightly greenish, ovate-acuminate, free from the
petiole and from each other, sparsely to loosely cov¬
ered with tangled, subappressed to ascending white
and black hairs, sometimes subglabrous. Leaves 9—
16 cm long; petiole 3—5 cm long, like the rachis
sulcate. black hairy like the stem. Leaflets in 9 to
1 1 pairs, elliptic to nearly orbicular, 5—12 X 4—9
mm, emarginate at the apex, on upper side sparsely,
on underside loosely covered with subappressed
white or white and black hairs 0.5—1 mm long. Pe¬
duncle 10—12 cm long, in lower part distinctly, in
upper part slightly angular-sulcate, rather densely
covered with ascending to spreading blackish hairs
0.5— 1.5 mm long. Racemes short, often globose, ca.
3 cm long, rather densely 9- to 12-flowered; axis
black villous. Bracts whitish, narrowly oblong to
narrowly elliptic, 5-6 mm long, 1—1.5 mm wide, on
outer side rather densely covered with tangled
black hairs up to 1.5 mm long, on inner side (upper
side) sparsely hairy. Pedicels 1 .5—2 mm long, very
densely black hairy. Bracteoles absent. Calyx ca. 8
mm long, campanulate-tubular, truncate at the
mouth, rather densely to densely covered with sub¬
appressed to ascending black hairs 0.3-1 mm long;
teeth narrowly triangular, 3—4 mm long, sparsely
hairy on inner side. Petals purple to bluish white.
Standard ca. 16 mm long; blade ca. 7 mm wide,
widely rectangular, distinctly emarginate at the
wide apex, at the base with a cuneate claw of equal
length. Wings ca. 15 mm long; blades narrowly ob¬
long, rounded at the apex, 8X3 mm; auricle nar¬
row, 2.5—3 mm long, claw ca. 7 mm long. Keel ca.
16 mm long; blades oblique-obovate, with lower
edge rectangularly curved in distal part and upper
edge nearly straight, subobtuse at the apex, ca. 8.5
X 4 mm; auricle ca. 1 mm long, claw ca. 8 mm
long. Staminal tube ± truncate at the mouth. Ovary
with a stipe ca. 4 mm long, densely covered with
subappressed white and predominantly black hairs
ca. 1 mm long; style thickish, hairy in lower half.
Legumes unknown.
Astragalus griersonii differs from A. kongrensis
Baker by being 20—25 (vs. 6—14) cm tall, in veg¬
etative parts with predominantly black (vs. merely
white) hairs, calyx ca. 8 (vs. 9-12.5) mm long, with
teeth 3—4 (vs. 4—7) mm long, standard ca. 16 (vs.
16-21) mm long.
Astragalus sect. Trachycercis Bunge, Mem.
Acad. Imp. Sci. Saint Petersbourg I 1(16); 1 14.
1868. TYPE: Astragalus humilis M. Bieber-
stein (lectotype, designated by Podlech (1990:
490)).
Astragalus lang-ranii Podlech, nom. et slat. nov.
Based on Astragalus salsugineus Karelin &
Kirilow var. multijugus S. B. Ho, Bull. Bot.
Res. North-East. Forest. Inst. 3(1): 52. 1983,
non Astragalus multijugus DC. TY PL: [China.]
Ningxia: Yinchuan, prope viam ferream, 1200
m, 14 May 1959, Y. C. Ho 2135 (holotype,
WUK; isotvpe, PE).
Because of the short original diagnosis of this
taxon a complete description is given here.
Astragalus lang-ranii is near to A. alaschanensis
H. C. Fu, but differs by smaller leaflets (3-4 X 1.3
mm vs. 5—15 X 3-7 mm), shorter bracts (4—6 mm
vs. 10—15 mm long), shorter calyx (12—13 mm vs.
15—18 mm long; teeth 2.5—4 mm vs. 5—9 mm long).
Plants ca. 10 cm tall, shortly caulescent, densely
covered with symmetrically to asymmetrically bi¬
furcate, spreading white hairs, the branches up to
1 mm long. Caudex slender, simple or shortly
branched. Stems up to 5 cm long, densely tomen-
tose. Stipules whitish, adnate to the petiole for 2—
4 mm, the lower triangular, 3—4 mm long, the upper
narrowly triangular-acuminate 5-7 mm long, all
densely hairy. Leaves 8—12 cm long; petiole 3—4
cm long, like the rachis slender, flexuous or curved,
rather densely hairy. Leaflets in 1 1 to 13 pairs, el¬
liptic, 3-6 X 1.5—3 mm, rounded to acute at the
apex, on both sides covered with tangled, ±
spreading hairs. Peduncles 1-1.5 cm long, hairy
like the stem. Raceme loosely 4- to 6-flowered.
Bracts whitish. 4—6 mm long, ovate-acuminate,
loosely white-hairy, partly w ith basifixed hairs. Ped¬
icels 2—3 mm long, hairy. Calyx 12-13 mm long,
tubular, obliquely truncate at the mouth, densely
covered with short to long spreading, tangled hairs;
teeth subulate, 2.5—4 mm long. Petals pale purple.
Standard 20—22 mm long; blade ca. 7 mm wide,
narrowly ovate, ± rounded at the apex, very shortly
226
Novon
narrowed at the base. Wings ca. 19 mm long;
blades narrowly obovate, obliquely slightly incised
at the apex, 10 X 3 mm; auricle 1 mm long, claw
9 mm long. Keel 15 mm long; blades obliquely el¬
liptic-triangular, with widely curved lower edge and
nearly straight upper edge, obtuse at the apex, 6 X
3 mm; auricle short, claw 9 mm long.
Etymology. Named after Prof. Xu Lang-Ran,
co-worker on the Flora of China , who has recog¬
nized the distinctness of this species. The hyphen
in the Latin name lang-ranii must be retained be¬
cause two forenames are combined.
Literature Cited
Ho, S. B. 1983. Trascursores Florae Astragalorum Sinen-
sium. Bull. Bot. lies. North-East. Forest. Inst. 3(4): 51 —
70.
Podlech, I). 1990. Die Typifizierung der altweltlielien Sek-
tionen der Gattung Astragalus L. (Leguminosae). Mitt.
Bot. Staatss. Mtlnehen 29: 461—494.
A New Species of Ledermanniella (Podostemaceae) from Cameroon
John ./. Schenk
Department of Botany and Plant Pathology, Oregon State University, 2082 Cordley Hall,
Corvallis, Oregon 97331-2902, U.S.A. Author for correspondence; present address:
School of Biological Sciences, Washington State University, Pullman,
Washington 99164-4236, U.S.A. john_sch@mail.wsu.edu
Duncan W. Thomas
Department of Forest Science, Oregon State University, Corvallis, Oregon 97331-2902,
U.S.A. DuncanWT@aol.com
Abstract. Ledermanniella prasina J. Schenk &
I). W. Thomas in the family Podostemaceae, sub¬
genus Ledermanniella, is proposed as a new spe¬
cies. It is known only from its type, collected at a
single waterfall in Cameroon. The ecology, mor¬
phology. and variation among individuals are dis¬
cussed. The new species is compared to the mor¬
phologically similar L. bifurcata, L. bowlingii, L.
guineensis, L. jaegeri, L. minutissima, and L. mon-
andra. Illustrations and a key to the monostaminate
species in the subgenus are provided.
Key words: aquatic plants, Cameroon, Leder¬
manniella, narrow endemic, Podostemaceae.
With its center of diversity in Cameroon, Leder¬
manniella Engler is the second largest genus in
Podostemaceae, consisting of about 44 species,
which are all endemic to tropical Africa (Cook,
1996; Gusset. 1983, 1984). As defined by Cusset
(1974), Ledermanniella is distinguished from other
genera in the family through the combination of the
following characters: the presence of alternate or
whorled leaves, solitary or occasionally clustered
flowers inverted within a closed spathellum, and an
ovoid to ellipsoidal capsule with eight ribs. This
broader classification ignored pollen and stamen
number traits previously used and resulted in the
inclusion of Inversodicraeia Engler and Monati-
driella Engler within Ledermanniella (Cusset.
1974). Subsequently, Cusset divided the genus into
two subgenera, Phyllosoma and Ledermanniella,
distinguished by the presence or absence of scale¬
like leaves, respectively (Cusset. 1983, 1984).
The new taxon is clearly a member of Podoste¬
maceae due to its semi-aquatic habitat, thallus-like
base (root), the presence of spathella, flowers that
consist of two tepals, a syncarpous superior gynoe-
cium with two styles, a single anther, and capsular
fruit. The new taxon is placed in Ledermanniella
based on its alternate leaves, solitary to clustered
flowers each inverted within a closed spathellum,
two filamentous tepals, flowers borne along shoots
above a thalloid-root, and ellipsoidal capsules. It is
placed in the subgenus Ledermanniella based on
the absence of scale-like leaves and the presence
of elongated leaves. The new species is known only
from the type collection, which consists of approx¬
imately six pressed individuals along with 25 stems
and shoots (which were separated from each other
and/or other individuals at the time of collection),
and two individuals along with 13 stems and shoots
preserved in alcohol. Both collections have buds,
flowers, and fruits in all stages of development.
Taxonomy
Ledermanniella prasina J. Schenk & 1). W.
Thomas, sp. nov. TYPE: Cameroon. Southwest
Province: 14 km NE of Mundemba, base of a
large waterfall in the Mana River, between the
villages of Miangwe 2 and Meta, 300 m,
5°02.4'N, 9°01.0'E, 01 Dec. 1998. Duncan
Thomas 11550 (holotype, MO; isotypes, K.
LBV, MO (preserved), NY, OSC, P, SC A, WAG,
YA). Figure 1.
Speciebus monostaminiferis subgeneris Ledermanniel-
lae similis sed distinguibilis seminibus smaragdinis et flo-
ribus (tepalis anthera gynoeeioque includentibus) majori-
bus.
Semi-aquatic herbs from the spray-zone of a wa¬
terfall; thalloid-root disk-shaped, holdfast attaching
plant to rock substratum; plants erect, 9—13 cm tall;
main stem arising from thalloid-root, 1.0—32.0 mm
long, 1. 0-2.0 mm wide, erect; vegetative shoots
arising from main stems, filiform, dichotomously
branched, up to 7 cm long; fertile shoots arising
from main stem, filiform, dichotomously branched,
bearing reproductive structures; leaves up to 4 cm
Novon 14: 227-232. 2004.
228
Novon
Figure 1. Ledermanniella prasina J. Schenk & 1). W. Thomas, based on I). W. Thomas 11550. — A. Detailed habit
of a single main stem showing the structure of the main stem (X'), fertile shoots (X"), vegetative shoots (Xv), leaves
(L), and a thalloid-root (R; note that the apical portion and other mains stems are omitted for clarity). — B. A segment
of the primary and fertile shoots showing the details of a flowering cluster, unruptured spathella (Sp) with leaves
subtending spathella; also seen are stipules (St) located on the intrapeliolar scales (Is). — C. Ruptured spathella, with
emerging, inverted flower (left) and fully emerged flower (right), along with tepals (P; only one of two shown).
long, 0.1— 0.5 mm wide, dichotomously branched,
solitary to verticillate below spathella, basal portion
scale (intrapeliolar scale) with stipules, apical por¬
tion elongating from center of scale, filiform, gla¬
brous; spathella erect, 3.0— 8.0 mm long before de¬
hiscence, 1.0— 2.2 mm wide, ovate at beginning of
development, then elongating to clavate, glabrous,
solitary or in clusters (1—6) in the axils or along
fertile shoots, apex globose to acute, dehiscence
mostly irregular, occasionally bilobed to trilobed;
(lowers solitary within spathella, zygomorphic, in¬
verted prior to the opening of the spathella, upright
after elongation, pedicel up to 13 mm long, tepals
2, acicular, 0.4 — 1 0.7 mm long, gynoecia 1.5( — 1 .7)
mm long, 0.7— 0.9 mm wide, attached to a gyno-
phore, unilocular, styles 2, linear. 0.6— 0.8 mm long,
stigmas bilobed; stamens 1 (very rarely 2); fila¬
ments ( = andropodium) 1.1— 1.3 mm long at spa-
Volume 14, Number 2
2004
Schenk & Thomas
Ledermanniella prasina from Cameroon
229
thellum dehiscence, 5.0— 5.3 mm long after anthe-
sis, alternate to the tepals; anthers basifixed, 2.0
mm long, 0.5— 0.8 mm wide, longitudinally dehis¬
cent; pollen white, in monads; capsules elliptic,
2.2— 2.7(— 3) mm long, 1.0— 1.2 mm wide, bivalved,
one valve persistent after dehiscence, the other ca¬
ducous, ribs 8 per capsule, with 3 ribs per valve
and 2 as the marginal sutures; placentas green;
seeds bright emerald green, 0.3-0. 4 mm long,
0.125—0.15 mm wide, ca. 25 to 37 per capsule.
Ecology
Like many Podostemaceae, species of the genus
Ledermanniella are sometimes narrowly endemic to
a single geographical area (Cook. 1996). From our
current knowledge, L. prasina occurs only at a sin¬
gle site, and is therefore narrowly endemic to the
Mana River valley in the Rumpi Hills of Cameroon.
On the basis of this information, it should be red-
data listed as an endangered species according to
the criteria established by the International Union
for the Conservation of Nature (IUCN). The new
species occupies a very unusual and specific hab¬
itat. The waterfall in the Mana River consists of two
stages with a total height of approximately 100 m.
At the bottom, there is a large, deep pool and a
shelf of exposed gneiss bedrock, about 200 m2 in
extent; L. prasina occurs only on this shelf. The
shelf is above the observed high-water level in the
river, but its largely unvegetated state suggests pe¬
riodic scouring, probably during occasional flash
floods. The area has a very wet but strongly two-
seasonal climate with a total annual rainfall ex¬
ceeding 5000 mm. There is a short dry season De¬
cember to March, with three months averaging less
than 100 mm of rain. The first part of the wet sea¬
son is typified by thunderstorms of varying fre¬
quency and duration, sometimes with dry periods
of several weeks’ duration. The wet season peaks
in August and September, when monthly rainfall
may exceed 1000 mm (Chuyong et a!., 2004). This
wet, seasonal climate, combined with the rugged,
rocky topography of the Rumpi Hills, generates the
type of river that is ideal for Podostemaceae, with
a rocky bed and large seasonal variation in flow.
With the exception of L. prasina, all Podostemaceae
reported from Cameroon follow similar phenological
patterns in response to this climate. The plants are
obligate hydrophytes and grow submerged in fast¬
flowing water during the wet season, attached to the
rocks by thalloid-roots. When the water level in the
rivers subsides at the beginning of the dry season,
the exposed plants flower and fruit rapidly, after
which at least the aerial parts die (Cusset, 1987;
Thomas, pers. obs.). Several species of Podoste¬
maceae exhibiting this phenology have been col¬
lected from the Mana River, including Lederman¬
niella ledermannii (Engler) C. Cusset,
Macropodiella pellucida (Engler) C. Cusset, and
Saxicolella flabellata (G. Taylor) C. Cusset (Cusset,
1987).
Ledermanniella prasina exhibits a very different
habit and phenology, which is unique for the family
in Cameroon. It does not grow submerged as most
podostemads do, but instead has erect aerial stems
and shoots that are kept moist throughout the grow¬
ing period by spray generated from the waterfall.
The spray is continuous during the wetter months
with high water flow in the river, and it is this spray
that supports the growth of the new species. The
aerial shoots are intolerant of desiccation and with¬
er rapidly when removed from the moist environ¬
ment. In the dry season, the waterfall is present but
much reduced in volume, and little spray reaches
the rock shelf that supports the L. prasina popu¬
lation. During this period, the dry rocks appear to
be largely devoid of vegetation and we do not yet
know if L. prasina is annual, or if it passes the dry
season as a dormant thalloid-root. Instead of having
the short flowering period typical of Podostemaceae
when the water level drops, the flowering and fruit¬
ing of L. prasina are prolonged and take place dur¬
ing the latter part of the growing season, when its
fertile branches are produced. During this same pe¬
riod, the other Podostemaceae in the river are sub¬
merged and growing vegetatively, but they do not
flower.
For several reasons, we do not believe that Led-
ermanniella prasina is a terrestrial form of one ol
the many other Podostemaceae that grow in the
area. No other species of subgenus Ledermanniella
have been collected from the Mana River. In ad¬
dition, the L. prasina population is limited to a rock
shelf, at least a meter above the high-water level,
and does not extend down to the water, suggesting
that it is intolerant of extended periods of submer¬
sion. Finally, some of the traits that separate the
new species from similar species are structural or
reproductive, and likely to be independent of hab¬
itat-related morphological plasticity.
Morphology
Podostemads are difficult to describe in terms of
standard angiosperm morphology due to their great¬
ly reduced vegetative anatomy and unusual mor¬
phology. Distinguishing vegetative organs using cell
and tissue orientation is an extremely difficult if not
an impossible task (Rutishauser, 1995; Ameka et
230
Novon
al., 2003). The literature contains descriptive in¬
consistencies among the interpreted structures,
demonstrating a general discordance of terminology
and homology assessment among biologists study¬
ing Podostemaceae (Rutishauser, 1995; Sehgal et
al., 2002). We use the terms “root,” “main stems,”
“fertile shoots,” “vegetative shoots,” and “leaves”
(Fig. I) simply to describe this species; however,
homology of such organs remains uncertain (Rutis¬
hauser, i995, 1997).
The root structure is represented by an undefined
thalloid tissue with a holdfast (= hapteron) located
underneath. The function of the holdfast is to sta¬
bilize the plant by attaching it to the substrate. Bot¬
anists working with Podostemaceae use the term
holdfast or hapteron, borrowed from algae termi¬
nology, to describe the superficially similar struc¬
ture (Taylor, 1954; Hammond, 1937; Philbrick,
1984). The shape of the thalloid-root is described
as being disk-like. However, the variability in thal¬
loid-root shape is not fully understood in this spe¬
cies. The term “thalloid-root” is used to describe
the undifferentiated tissue of which the main stem
arises and that binds the plant to the substrate.
There is no implication of homology to the familiar
idea of a root since the structure has been sug¬
gested to be derived from a stem, a coalesced stem
and root, or simply a modified root (Sehgal et al.,
2002; Rutishauser, 1995). This term conforms to
current literature and the root-shoot (CRS) model
as the structure located underneath the main stem.
However, much anatomical work needs to done on
this plant as well as other podostemads before more
concise determinations of homology can be made.
Approximately four to five main stems develop
directly from the thalloid-root for each individual
(Fig. 1A). The vegetative shoots and fertile shoots
are borne on the main stem. The majority of the
flowers are borne on the fertile shoots; however,
some parts of the thalloid-root contain a shorter
main stem of which there are no apparent vegeta¬
tive or fertile shoots, but do have spathella sub¬
tended by leaves. It is not known if these stems
elongate later in development and give rise to the
vegetative and fertile shoots or remain as shorter
dimorphic stems. This trait is not found in all in¬
dividuals.
Two types of shoots are borne from the main
stem, fertile shoots and vegetative shoots. The veg¬
etative shoots branch dichotomously and appear
leaf-like. There are two to three vegetative shoots
borne on the main stem, before the fertile shoots
arise (Fig. 1 A). Located where the vegetative shoots
diverge from the main stem, ligule-like structures
are found that encircle it (Fig. 1A, B). The fertile
shoots tend to be thicker than the vegetative shoots
where they are borne on the main stem (Fig. 1A,
B) . They also differ by having reproductive struc¬
tures along the branches and in the axils of di¬
chotomous branches (F ig. I A, B).
Leaves are located below the spathella (F ig. 1 A—
C) . The basal portions of the leaves are scale-like,
with an elongating thread-like leaf that dichoto¬
mously branches, similar in appearance to the veg¬
etative stems. Stipules are located along the side of
the scale. Ameka et al. (2003) referred to the basal
scale as an intrapetiolar scale, which can also be
rudimentary lateral stipules. We apply the term
“leaves” to remain consistent with literature de¬
scribing similar species (Ameka et al., 2003; Ru¬
tishauser, 1997; Hall, 1971; Taylor, 1954).
The spathella are quite variable in this species.
The shape can vary from ovate to elliptic to clavate
as they mature. It is also found that the spathella
on the distal parts of the shoots are more ovate than
those located toward the base. There is further var¬
iation in the spathellum margin following dehis¬
cence. Most spathella have irregular margins fol¬
lowing dehiscence; some, however, develop bilobed
or trilobed margins.
The number of stamens is a variable character
in this taxon as well as in other select species of
Podostemaceae (Cusset, 1974). Two out of approx¬
imately 75 shoots have a single flower (each indi¬
vidual has approximately four to five shoots with an
average of ca. 20 (lowers per shoot) that contains
two anthers attached to an andropodium (specimen
housed at OSC). However, variability in anther
number is rare enough that we continue to describe
this species as monostaminate. Furthermore, a sin¬
gle individual was found containing two ovaries at¬
tached to a single gynophore (specimen housed at
OSC). From our understanding of the group, this is
a novel attribute for Ledermanniella. We could not
find any literature documenting this mutation, sug¬
gesting its anomaly, and we therefore did not in¬
clude it in the description.
The specific epithet “prasina” refers to the em¬
erald green seeds that were removed from dehisced
capsules. Less mature seeds seen inside undehis¬
ced capsules are also green.
Detailed comparisons were made between the
new species and similar described species. The
species selected for comparison included all mon¬
ostaminate species from the subgenus, plus one
species with two stamens. Published descriptions
were used for the comparisons in all but two cases,
where we needed to examine herbarium material.
Comparisons were made using herbarium mate¬
rial with the two monostaminate species, Leder-
Volume 14, Number 2
2004
Schenk & Thomas
Ledermanniella prasina from Cameroon
231
manniella bowlingii (J. B. Hall) C. Cusset from
Ghana and L guineensis C. Cussel from Guinea.
These two speeies were examined because they re¬
semble the general habit of L. prasina: a single sta¬
men, elongated shoots, intrapetiolar scale (L. bow¬
lingii only), and linear leaf or leaf-like structures.
The herbarium material along with written de¬
scriptions demonstrate that Ledermanniella bowlin¬
gii differs qualitatively by having papillate spa-
thella, pollen in dyads, inflorescence a cyme, and
brown to reddish brown seeds, along with quanti¬
tative traits such as longer shoots, shorter tepals,
smaller anthers, and smaller gynoecium (Hall,
1971; Cusset, 1984; Ameka et al., 2003). This was
determined after examining the isotype (P) and a
photograph of the holotype (K) along with descrip¬
tions and illustrations. Ledermanniella guineensis is
distinguished from L. prasina by having a total
length of one meter, inflorescence a cyme, a bilobed
spathellum, the lack of leaves directly subtending
the spathellum, smaller overall flower size, and
brown to reddish brown seeds (Hall, 1971; Cusset,
1984). The holotype (P), along with four additional
collections of Ledermanniella guineensis, was used
for this comparison along with written descriptions
and illustrations.
We did not use herbarium material to compare
the three remaining monostaminate species lecler-
manniella jaegeri G. Cusset, L. minutissima C. Cus¬
set, both from Sierra Leone, and L. monandra C.
Cusset from Cameroon, since the published de¬
scriptions of these species show they are very dif¬
ferent. Both L. minutissima and L. jaegeri differ
from L. prasina by containing leaves that subtend
the spathella which lack intrapetiolar scales and
dichotomous branching. Ledermanniella minutissi¬
ma is further separated from L. prasina by pos¬
sessing spathella that always arise from the thalloid
or from a small shoot that is up to 3 mm long, and
are always solitary (Cusset, 1984). ledermanniella
jaegeri has the additional character traits of leaves
borne along the entire shoot instead of solely under
the spathellum, as well as a more elliptic ovary
(Cusset, 1984). Ledermanniella monandra appears
to have an intrapetiolar scale; however, the leaves
are borne on the entire length of the fertile shoot
and are triangular in shape in contrast to the elon¬
gating leaves of L. prasina (Cusset, 1974, 1984).
The only bistaminate species investigated was
Ledermanniella bifurcata (Engler) C. Cusset from
Cameroon, Gabon, and Congo. Although it does
have two stamens, it shares similar morphological
traits, such as irregular spathella dehiscence, pol¬
len in monads, and intrapetiolar scales, which are
characters we felt needed further comparisons. Be¬
sides the anther number, L. bifurcata differs by
having an ovate spathellum (L. prasina also has an
ovate spathellum, though it elongates to become
clavate as it matures), a fertile shoot that widens at
nodes (vs. no widening at nodes in L. prasina ),
leaves subtending the spathellum for the entire
length of the shoot, shorter tepal length, smaller
closed spathella. longer spathella pedicels, smaller
anther size, and smaller gynoecium (Cusset, 1984,
1987; Hall. 1971).
Key to the Monostaminate Species of Ledermanniella
subg. Ledermanniella
la. Leaves present under spathella, intrapetiolar
scale absent, leaves do not branch dichotomous-
ly.
2a. Plants without elongated shoots (0—3 mm);
leaves located only under spathella . . .
. L. minutissima C. Cusset
2b. Shoots greater than 3 mm, leaves located
along entire shoot . L jaegeri C. Cusset
lb. Leaves either absent under spathella, or if pre¬
sent, with dichotomous branching and intrapetio¬
lar scale.
3a. Leaves triangular, 1-2 mm long .
. L. monandra C. Cusset
3b. Leaves linear, greater than 2 mm long.
4a. Spathella papillate, pollen in dyads . .
. /,. bowlingii (J. B. Hall) C. Cusset
4b. Spathella glabrous, pollen in monads.
5a. Stems up to I m long, trailing in
moving water; leaves absent below
spathella; spathella arranged in a
cyme; spathella bilobed at dehis¬
cence; anthers 1 mm long; tepals
0.3— 0.4 mm long; seeds brown/red
. L. guineensis C. Cusset
5b. Plants 9—13 cm, erect on wet rock;
leaves located below spathella; spa¬
thella solitary or in cluster; spa¬
thella irregular to bilobed at dehis¬
cence; anthers 2 mm long; tepals
0.4— 0.7 mm long; seeds emerald
green . L. prasina
J. Schenk & 1). W. Thomas
Acknowledgments. The authors are indebted to
Ffena Schlachter for illustrations, and to Kenton L.
Chambers for the Latin description. We thank Aar¬
on Liston for reviewing drafts of this manuscript,
and two anonymous reviewers who provided useful
comments. Thanks are also due to the curators of
K and P for loaned specimens, as well as to Rich¬
ard R. Halse (OSC) for obtaining and housing the
loans. Funds for the fieldwork were provided by the
International Cooperative Biodiversity Groups
(ICBC).
Literature Cited
Ameka, G. K., G. C. Clerk, E. Pfeifer & R. Rutishauser.
2003. Developmental morphology of ledermanniella
232
Novon
bowlingii (Podostemaceae) from Ghana. PI. Syst. Evol.
237: 165-183.
Chuyong, G. B., R. Condit, 1). Kenfack, E. Losos, VI. Sa-
inge, N. C. Songwe & D. W. Thomas. 2004. Korup Eor-
est Dynamics Plot, Cameroon. Chapter 29 in E. Losos
& E. G. Leigh, Jr. (editors). Tropical Forest Diversity
and Dynamism: Findings from a Large-Scale Plot Net¬
work. Univ. Chicago Press, Chicago.
Cook, C. I). K. 1996. The Aquatic Plant Book, 2nd Rev.
Ed. SPB Academic Publishing, The Hague.
Cusset, C. 1974. Contribution it lYtude des Podostema¬
ceae: 4. Les genres Ledermanniella, Monandriella et
Inversodicraeia. Adansonia, ser. 2, 14: 271-275.
- . 1983. Contribution it lY'tude des Podostemaceae.
7. Ledermanniella Engl., sous-genre Phyilosoma, C.
Cusset. Bull. Mus. Natl. Hist. Nat.. B. Adansonia 5:
361-390.
- . 1984. Contribution i lYtude des Podostemaceae:
8. Ledermanniella Engl, sous-genre Ledermanniella.
Bull. Mus. Natl. Hist. Nat., B, Adansonia 3: 249—278.
- . 1987. Podostemaceae and Tristichaceae. Pp. 51—
99 in B. Satabif4 & P. H. Moral (editors), Flore du Came¬
roon Vol. 30. Ministfere de I'Enseignement Suptirieur et
de la Recherche Scientifique, Yaounde.
Hall, J. B. 1971. New Podostemaceae from Ghana with
notes on related species. Kew Bull. 26: 125-136.
Hammond, B. L. 1937. Development of Podostemum cer-
atophyllum. Bull. Torr. Bot. Club 64: 17—36.
Philbrick, C. T. 1984. Aspects of floral biology, breeding
system, and seed and seedling biology in Podostemum
ceratophyllum (Podostemaceae). Syst. Bot. 9: 166—174.
Rntishauser, R. 1995. Developmental patterns of leaves
in Podostemaceae compared with more typical flowering
plants: Saltational evolution and fuzzy morphology. Can-
ad. J. Bot. 73: 1305-1317.
- . 1997. Structural and developmental diversity in
Podostemaceae (river-weeds). Aquat. Bot. 57: 29—70.
Sehgal, A., M. Sethi & H. Y. Mohan Ram. 2002. Origin,
structure, and interpretation of the thallus in Hydrob-
ryopsis sessilis (Podostemaceae). Ini. J. PL Sei. 163:
891-905.
Taylor, G. 1954. Podostemaceae. Pp. 122—127 in J.
Hutchinson & J. M. Dalziel (editors), Flora of West
Tropical Africa, vol. 1, 2nd ed., revised by R. W. J.
Keay. Crown Agents, London.
A New Species of Minasia (Asteraceae, Vernonieae) from the
Planalto de Diamantina, Minas Gerais, Brazil
Joao Semir
Departamento de Botanica, Universidade Estadual de Campinas, C. P. 6109, 13083-970,
Campinas SP. Brazil, semir@unicamp.br
FI avia F. Jesus
Departamento de Genetica, Universidade Estadual de Campinas, C. P. 6109, 13083-970,
Campinas SP, Brazil, f931719@dac.un iearnp.br
ABSTRACT. A new species of Minasia (Astera-
eeae, Vernonieae) was found in the “campus rupes-
tres" of the Planalto de Diamantina, Minas Gerais,
Brazil. Minasia lewinsohnii is characterized by its
overall small size, linear to elliptical subfleshy
leaves, sessile to pedunculate capitula, and
aehenes with indumentum throughout the surface.
So far, it is known from only one population.
Key words: Asteraceae, Brazil, Minasia, Ver¬
nonieae.
Minasia H. Robinson (Asteraceae, Vernonieae)
was recently established by Robinson (1992). who
created the genus based on two species previously
described as belonging to Vernonia Schreber and
established another new species. According to Rob¬
inson (1992, 1995, 1996, 1999), the genus cur¬
rently includes five species: M. alpestris (Gardner)
H. Robinson, M. scapigera H. Robinson. M. perei-
rae H. Rol >inson, M. splettiae H. Robinson, and M.
cabralensis H. Robinson.
Minasia species are endemic to the campos ru-
pestres of the Kspinhago Range in Minas Gerais,
Brazil. More specifically, they occur in the high-
altitude areas ranging from the Serra do Cipd to the
Planalto de Diamantina, as well as in the isolated
western extension of the campos rupestres in Minas
Gerais, the Serra do Gabral. The campos rupestres
have an immense diversity of species, many of
which are endemic (Giulietti & Pirani, 1988). They
are considered to present the greatest degree of en¬
demism among Brazilian vegetation types (July,
1970). Therefore, finding a new species in these
environments is not surprising.
Recent collecting trips were conducted by fac¬
ulty of the Universidade Estadual de Campinas for
taxonomic, genetic, and ecological studies on the
interactions between Asteraceae and phytophagous
insects. During those trips, a new, seemingly micro¬
endemic species of Minasia was found in the Plan¬
alto de Diamantina. This region seems to be the
one with the greatest diversity of Minasia species,
with four of the five previously described species
occurring there (M. alpestris, M. scapigera, M. per-
eirae, and M. splettiae), in addition to the new spe¬
cies proposed here.
Minasia lewdnsoliiiii .). Semir & F. F. Jesus, sp.
nov. TYPE: Brazil. Minas Gerais: Diamantina,
Estrada Guinda-Sao Joao da Chapada, campo
arenoso com cascalho quartzftico, 18°09.59'S,
43°43.00'W. 7 Sep. 1996 (fl, fr), T. M. Lewin-
sohn, P. I. Prado, A. J. Santos 6696 (EF.C).
of the adaxial face, margin barbellate, purple on
the middle, pale on the margins. Flowers 21 to 43
per capitulum; corolla 7.1-10.6 mm long, lavender
to magenta, densely glandular-verrucous on the
lobes, sparsely so on the tube, tomentose on the
external apex of the lobes, glabrous below; lobes
2.5— 3.2 X 0.6-0. 8 mm, lanceolate; anthers 2. 9-4. 2
mm long, tails ea. 0.7 mm long, apical appendages
Volume 14, Number 2
2004
Semir & Jesus
Minasia lewinsohnii from Brazil
235
0.6— 1.0 X 0.2— 0.3 mm, triangular to sub-lanceo¬
late, apex acute; style ca. 12 mm long. Achene 2.2—
3.5 mm long, 0.8— 1.5 mm diam., ribbed, ca. 8 ribs,
sericeous to setuliferous indumentum, dense to
sub-glabrescent throughout the surface, sometimes
denser at the base, glandular surface; pappus bris¬
tles 0.4— 7.5 mm tall, in indistinguishable series,
the most external ones gradually shorter, lavender
to magenta, barbellate.
The species is comparable in general size — both
of rosette and inflorescence — to Minasia pereirae
and M. splettiae. In inflorescence height it is also
comparable to M. cabralensis ; however, the rosette
leaves are shorter in M. lewinsohnii (up to 9 cm
long) than in M. cabralensis (10-16 cm long), giving
M. lewinsohnii the appearance of an overall smaller
plant. Minasia lewinsohnii is much smaller than M.
alpestris and M. scapigera, both of which can be up
to 1 m tall. In addition to the difference in general
size, the leaves in M. lewinsohnii are linear to el¬
liptical, not oblaneeolate as in M. alpestris , and the
inflorescence is not congested like the one in M.
scapigera.
The capitula in this new species may be pedun¬
culate, as they generally are in Minasia cabralensis,
but very rarely in the other species of the genus.
However, the peduncles are much shorter in M.
lewinsohnii than in M. cabralensis (up to 1.5 cm vs.
up to 35 cm long). Also, the achene in M. lewin¬
sohnii has indumentum throughout the surface,
whereas in M. cabralensis the achene is distally gla¬
brous (Robinson, 1996). Furthermore, the leaves
are shorter and more fleshy in M. lewinsohnii than
in M. cabralensis. To date, M. cabralensis seems to
be restricted to the Serra do Cabral while M. lew¬
insohnii has been found only in the Planalto de
Diamantina. Minasia lewinsohnii differs from M.
splettiae in having wider and thicker leaves and
from M. pereirae in having linear to elliptic rather
than oblaneeolate leaves.
To date, Minasia lewinsohnii is known only from
one population, in the Planalto de Diamantina, oc¬
curring on sand. The species is named in honor of
Thomas Lewinsohn (Departamento de Zoologia,
Universidade Estadual de Campinas), one of the
first collectors of this population.
Paratypes. BRAZIL. Minas Gerais: Diamantina, Es¬
trada Guinda-S. Joao da Chapada, I8°09.58'S,
43°42.96'W, 23 June 2000 (fl, fr). K. S. Yotoko et al. KSY-
00/112 (BllCB, UEC), 29 Apr. 1997, T. M. Lewinsohn et
al. in PIC97032 (MBM, SP, UEC).
Acknowledgments. We thank Thomas M. Lew¬
insohn, Paulo I. Prado, and Vera N. Solferini for
collections and support in the study of this genus;
Angela B. Martins for the Latin diagnosis; Angela
M. F. Pacheco for the drawings; Christina Muirhead
for help with the English text; Fabio A. Vitta and
Mariana E. Mansanares for support in the Depar¬
tamento de Botanica (UEC); Karla S. Yotoko and
Aluana G. Abreu for the collections in 2000; Har¬
old Robinson (US), John Pruski (MO), and Victoria
C. Hollowell (MO) for critical reviews of the man¬
uscript; CNPq and FAPESP (98/05085-2) for finan¬
cial support. This study is part of the program BI-
OTA/FAPESP, The Virtual Institute of Biodiversity
(www.biota.org.br).
Literature Cited
Giulietti, A. M. & J. R. Pirani. 1988. Patterns of geograph¬
ic distribution of some plant species from the Espinhago
Range, Minas Gerais and Bahia, Brazil. Pp. 39-69 in
W. R. Heyer & P. E. Vanzolini (editors), Proceedings of
a Workshop on Neotropical Distribution Patterns. Aca¬
demia Brasileira de Ciencias, Rio de Janeiro.
Joly, A. B. 1970. Conhega a Vegetagao Brasileira. EDUSP
& Polfgono, Sao Paulo.
Robinson, H. 1992. Notes on Lychnophorinae from Minas
Gerais, Brazil, a synopsis of Lychnophoriopsis Schultz-
Bip., and the new genera Anteremanthus and Minasia
(Vemonieae: Asteraceae). Proc. Biol. Soc. Wash. 105:
640-652.
- . 1995. New combinations and new species in
American Vernonieae (Asteraceae). Phytologia 78: 384—
399.
- . 1996. A new species of Minasia from the Serra
do Cabral, Minas Gerais, Brazil (Vernonieae; Astera¬
ceae). Phytologia 80: 350—35 1 .
- . 1999. Generic and subtribal classification of
American Vernonieae. Smithsonian Contr. Bot. 89.
Eugenia myrciariifolia (Myrtaceae), a New Species from
Parana, Brazil
Lucia Helena Soares-Silva
Herbario, Departamento de Botanica UnB, C.P. 04457 — 70919-970 — Brasilia, I)F, Brazil.
lsoares@unb.br
Marcos Sobral
Departamento de Botanica UFMG, Av. Antonio Carlos 6627, — 31270-110 — Belo Horizonte,
MG, Brazil, sobral@mono.icb.ufmg.br
ABSTRACT. Eugenia myrciariifolia, a new species
from the southern Brazilian state of Parana, is de¬
scribed and illustrated. This species is related to
Eugenia oeidocarpa, from which it is set apart by
the presence of cataphylls on the twigs, elliptic to
lanceolate leaves, smaller or absent pedicels, and
axillary, pilose flowers.
RESUMO. Eugenia myrciariifolia, uma nova es-
pdeie do estado do Parana, Brasil, e descrita e il-
ustrada. A nova cspccie e proxima a Eugenia oei¬
docarpa, da qual se distingue pela presen^a de
catafilos nos ramos, folhas elfpticas a lanceoladas,
pedicelos menores ou ausentes e flores axi lares e
pilosas.
Key words: Brazil, Eugenia, Myrtaceae.
Eugenia L. is one of the largest genera of the
Myrtaceae, with about 350 American species (Lan¬
drum & Kawasaki, 1997). The Brazilian species
have remained largely unstudied since the para¬
mount treatment of Berg (1857—1859), except for
local floristic studies (e.g., Kiaerskou, 1893; Le-
grand & Klein, 1969; Kawasaki, 1989; Mattos,
1984; Barroso & Peron, 1994; Marchiori & Sobral,
1997; Sobral, 2003). While studying the family
Myrtaceae in the state of Parana, a small-leaved
tree of the forests from the northern part of the state
came to our attention and is here proposed as new.
Eugenia my reiariifoliu Soares-Silva & Sobral, sp.
nov. TYPE: Brazil. Parana: Sapopema, Salto
das Orqufdeas, margem direita do rio Lageado,
8 Dec. 1995, /.. Soares-Silva 563 (holotype,
UB; isotype, FUEL). Figure I.
Species E. oeidocarpae proxima, a qua ramulis cata-
phyllorum instructa, foliis ellipticis vel lanceolatis (non
obovatis), pedicellis usque 2 mm longis (non 2—5 mm lon-
gis) et floribus axillaribus ovariis pubescentibus reeedit.
Novon 14: 236-238. 2004.
Small tree up to 5 m high; twigs red-brown or
grayish, exfoliating, the younger ones somewhat an¬
gled, brown and puberulent with white trichomes
0. 1—0.3 nun long, bearing at their bases spathulate
to linear cataphylls 2.5— 5.5 X 1 mm, these invo¬
lute, with cilia to 0.1 nun. Leaves chartaceous, with
scattered translucid punctations visible to the eye,
these more evident abaxially, reticulate, slightly
discolored, glabrous except for occasional scattered
white trichomes to 0.1 mm long along the adaxial
central nerve and 0. 1-0.3 mm long at the base of
leaves; blades elliptic to elliptic-lanceolate, 1—3 X
0.5—2 cm, the length/width ratio 2. 2-2. 8:1; apex
obtuse to acute; base acute to attenuate; central
nerve plane to slightly prominent adaxially and ev¬
idently raised abaxially; secondary nerves 6 to 10
pairs, evident on both faces and diverging at angles
of 50°— 75°; marginal vein 0.6— 0.8 mm from margin,
the margin itself with a yellowish ridge to 0.1 mm
wide; petioles plane adaxially, 1-3 X 0.4-0. 6 mm,
with trichomes like those on the branches. Flowers
axillary and solitary; pedicels 1—2 X 0.2-0. 5 mm
or absent; bracteoles persisting after anthesis, ovate
to rounded ovate, 0.5—1 X 0.7— 1.2 mm, ciliate;
flower buds globose, 3—4 mm diam.; calyx lobes
hemispherical, unequal in size, the external ones
1-1.3 X 1.5-2 mm, the internal ones 1.6—2 X 2-
2.5 mm, sparsely puberulent without and sericeous
within, with trichomes about 0.1 mm, the margins
sometimes with red cilia to 0.4 mm; petals orbicular
and concave, 2.5—3 mm diam., glabrous and ciliate;
staminal ring sericeous, 1.5 mm diam.; stamens ca.
100. 4—5 mm, the anthers globose, 0.4— 0.5 mm,
with one apical gland; style glabrous, 4—6 mm; ova¬
ry externally pubescent or sericeous, about 1 mm
long, bilocular, with 7 to 8 ovules per locule on a
central placenta. Fruits not seen.
Habitat and distribution. Eugenia myrciariifol¬
ia is a small tree from seasonal forests of central
Volume 14, Number 2
2004
Soares-Silva & Sobral
Eugenia myrciariifolia from Brazil
237
Figure 1. Eugenia myrciariifolia Soares-Silva & Sobral. — A. Habit. — B. Cataphylls at base of twigs. — C. Detail of
adaxial side of leaf. — I). Detail of inflorescence. — E. Flower bud, with bracteoles detached. — F. Flower after polli¬
nation, with bracteoles detached. — G. Stamen with apical gland. From Soares-Silva 570.
238
Novon
and western Parana; until now it has been collected
in only four municipalities.
Phenology. Blooming specimens were collected
from October to December; fruiting specimens are
still unknown.
Etymology. The specific epithet is allusive to
the resemblance of this species with some of the
genus Myrciaria, especially Myrciaria tenella (DC.)
0. Berg, with which it can be easily confused when
sterile due to its small leaves and dense foliation;
curiously, one of the vernacular names that we have
registered for this species, eambuf, is also largely
applied in southern Brazil lor species of Myrciaria.
Vernacular names. Cambuf, pitangao (from Lin¬
de man & Haas 2892).
Affinities. This species is related to Eugenia
oeidocarpa 0. Berg, another southern Brazilian spe¬
cies growing from Parana to Rio Grande do Sul
(Sobral, 2003). The following key presents the char¬
acters that set them apart.
Key kok Distinguishing Eugenia myrciariifolia and Eu¬
genia OEIDOCARPA IN SOUTHERN BRAZIL
la. Twigs without linear cataphylls at base; leaves
obovate with revolute margins; petioles to 4 X
0.8 mm; the inflorescences ramiflorous, 2- to 6-
florous, the axis to 2 X 0.5 mm; pedicels to 5 X
0.5 mm; flowers glabrous; larger calyx lobes to 3
X 3 mm . Eugenia oeidocarpa
lb. Twigs with linear cataphylls at base; leaves el¬
liptic-lanceolate; petioles to 2 X 0.6 mm; pedi¬
cels to 2 X 0.5 mm; flowers pilose, solitary and
axillary; larger calyx lobes to 2 X 2 mm .
. Eugenia myrciariifolia
Additionally, this species shows close resem¬
blance to the protologue of Eugenia coaetanea ().
Berg (Berg. 1857-1859), another ramiflorous spe¬
cies distinguished from E. myrciariifolia and E. oei-
docarpa by pilose young leaves, linear bracteoles
to 4 mm, and flower buds to 8 mm; nevertheless,
comparison with this species is problematic since
the type was destroyed at B and no isotypes are
known. Moreover, although said to be collected in
the state of Rio Grande do Sul, no further collec¬
tions of E. coaetanea are known, notwithstanding
extensive surveys in the myrtaceous flora of this
state (Mattos, 1984; Marchiori & Sobral, 1997; So¬
bral, 2003).
Paratypes. BRAZIL. Parana: Aranha, Lindeman &
Hass .3432 (MBM, U); Campo Novo, ca. 50 km de Lar-
anjeiras do Sul. Lindeman <£ Hass 2892 (MBM. U, LIB);
Parque Nacional do Iguayu [municipality of C(iu Azul, not
registered in label), 100 m W of rio Floriano, Lindeman
& Hass 3544 (MBM. 0); Sapopema, Salto das Orqufdeas,
margem direita do rio Lageado, Soares-Silva 441 (FUEL,
UB), 9 Dec. 1995, Soares-Silva 570 (FUEL, UB), Chagas
e Silva & Soares-Silva 2056 (FUEL, UB), Silva 2127
(FUEL, UB), Medri 744 (BHCB, FUEL), Pavdo et al. s.n.
(FUEL 27498, MO).
Acknowledgments. We acknowledge Claudenir
S. Cares (UB) for his fine drawing of Eugenia myr¬
ciariifolia, the curators of FUEL and MBM for their
kind collaboration, and Victoria Hollowed (MO)
and an anonymous reviewer for kind and valuable
suggestions for the improvement of this paper.
Literature Cited
Barroso, G. M. & M. V. Peron. 1994. Myrtaceae. In: M.
P. M. Lima & R. Guedes-Bruni (org.), Reserva Ecold-
gica de Maea6 de Cima, Nova Friburgo, Rio de Janeiro.
Aspectos Florfsticos das Esp^cies Vasculares, Vol. 1.
Jardim Botiinico do Rio de Janeiro, Rio de Janeiro.
Berg, 0. 1857-1859. Myrtaceae. In: K. Martius (editor).
Flora Brasil iensis 14(1): 1—656.
Kawasaki, M. L. 1989. Flora da serra do Cip6, Minas
Gerais; Myrtaceae. Bol. Hot. Univ. Sao Paulo II: 121 —
1 70.
Kiaerskou, H. 1893. Enumeratio myrtacearum brasilien-
sium. In: E. Warming (editor), Symbolarum ad floram
Brasiliae Centralis cognoscendam 39: 1-199.
Landrum. L. R. & M. L. Kawasaki. 1997. The genera of
Myrtaceae in Brazil: An illustrated synoptic treatment
and identification keys. Brittonia 49: 508-536.
Legrand, C. I). & R. M. Klein. 1969. Miildceas: Eugenia.
Pp. 45—216 in R. Reitz (org.), Flora llustrada Catari-
nense, Itajaf.
Marchiori, ,1. N. C. — 0.8 mm.
Plants subshrubs, woody at base; trichomes den¬
dritic throughout, 0.1— 0.2 mm long; stems 10—17
cm long, erect to ascending, few from base, many-
branched above, sparsely pubescent below, gla¬
brous and glaucous above. Leaves all cauline, nar¬
rowly lanceolate, 4-13 X 1-2 mm, fleshy,
becoming leathery upon drying, moderately pubes¬
cent, gradually becoming sparsely pubescent up¬
ward, base minutely auriculate, margin entire, apex
acute, auricles to I X 0.5 mm. Raceme ebracteate,
6- to 10-flowered; fruiting pedicels 3—5 mm long.
Volume 14, Number 3
2004
Al-Shehbaz
Synopsis of Neuontobotrys
255
256
Novon
glabrous, strongly reflexed or recurved and forming
half a loop. Sepals narrowly oblong, 3—3.5 X 0.5—
0.7 mm, moderately pubescent; petals yellow, soon
turning reddish with age, linear-oblanceolate, 4.5-
5 X 0.5— 0.7 mm, sparsely pubescent outside; fila¬
ments 2.5—3 mm long; anthers oblong-linear, 0.9—
I mm long; ovules 12 to 14 per locule. Fruits
oblong-linear, 6-8 X ca. 1 mm, not torulose; valves
thick, leathery, with a prominent midvein; gyno-
phore 0. 1—0.2 mm long; style slender 0.8—1 mm
long; seeds ovoid, brown, uniseriate, 0.9-1 X 0.6—
0.8 mm.
Neuontobotrys elloanensis, which is known only
from the type gathering, is easily distinguished from
its nearest relative, N. deserticola, by having entire,
narrowly lanceolate leaves to 2 mm wide and small¬
er petals (4.5—5 mm long) sparsely pubescent out¬
side. Hy contrast, N. deserticola has dentate, broad¬
ly ovate to ovate-lanceolate leaves 4 — 1 0( — 20) mm
wide and larger petals (6-8 mm long) glabrous out¬
side.
5. Neuontobotrys deserticola (Philippi) Al-
Shehbaz, comb. nov. Basionym: Cardamine de¬
serticola Philippi, Anal. Univ. Chile 81: 86.
1892. TYPE: Chile. “Deserto Atacama ad fon-
tem Acerillos,” Oct. 1877, 0. Villanueva 1606
(holotype, SGO-63889).
Sisymbrium philippianum I. M. Johnston, Rev. Chil. Hist.
Nat. 33: 26. 1929. Syn. nov. Based on Sisymbrium
amplexicaule Philippi, FI. Atac. 8: 10. I860, non A.
Cray, U.S. Explor. Exped. Wilkes, Bot. XV. 61.
1854. TYPE: Chile. Atacama: Pajonal, |23°46'S,
3170 m], R. A. Philippi s.n. (lectotype, designated
here, SCO-49243).
Distribution. Argentina (Prov. Catamarca, Ju-
juy, and Salta), Bolivia (Depto. Potosf), and Chile
(Regidn II and III).
Representative specimens. ARGENTINA. Catamarca:
Antofagasta de la Sierra, bos Nacimientos, Cabrera et al.
31862 (MO). Jujuy: Susquesientre Cauchori y Catua, Ca¬
brera el al. 22344 (SI). Salta: Salar de Gallego, Camino
a buea-huasi, Cabrera 8882 (F, GH). B01.1VIA. Potosi:
Prov. Nor bipes, Chiguana, Asplund 3031 (S, UPS).
CHILE. Region II (Antofagasta): San Pedro, near river
Loa, Zollner 8842 (MO, US). Region III (Atacama):
Depto. Copiapd, Cord. Rio Figueroa, Cl. Paredones, Wer-
dermann 96 8 (B, CAS, F, GH, MO, UC).
Although the type of Neuontobotrys deserticola
consists of three racemes and lacks the lower parts
of the plant, it is indistinguishable in every aspect
of (lower, fruit, and indumentum from plants of Sis¬
ymbrium amplexicaule Philippi. In fart, Schulz
(1924) was the first to point out this similarity but
failed to unite the two species.
Munoz-Pizarro (1960) cited two sheets, SGO-
49242 and SGO-49243, as the type of Sisymbrium
amplexicaule. The latter specimen, which is the
most complete and consists of four plants all col¬
lected from one locality, is designated herein as the
lectotype. The other sheet, SCO-49242, has five
fragments each collected from a different place, and
it is impossible to assign any of these fragments to
a given locality.
As delimited hy Knntze (1898), Hesperis intri¬
catissima (Philippi) Knntze included two species
that differ substantially in every morphological as¬
pect except for fruit size and orientation. In order
to simplify the complicated nomenclature of these
two species, they are designated herein as “sp. A”
and “sp. B.” One species (sp. A), which is the bas¬
ionym of //. intricatissima, is a glabrous annual
originally described as Draba intricatissima Philip¬
pi and now recognized as Eremodraba intricatissi¬
ma (Philippi) O. E. Schulz (Schulz, 1924; Al-Sheh-
baz, 1990). I he second of Kuntze’s (1898) species
(sp. B) is a pubescent perennial or subshrub cor¬
responding to Sisymbrium amplexicaule Philippi.
Schulz (1924) mishandled the nomenclature of
Kuntzes (1898) //. intricatissima in four ways.
First, he excluded the material based on sp. A to
Eremodraba, but maintained the epithet “'intricatis¬
sima" in Sisymbrium for sp. B. Second, he called
the species Hesperis intricatissima Knntze, instead
of //. intricatissima (Philippi) Knntze, even though
Knntze (1898) cited the basionym I). intricatissima
and did not treat the plant as a new species. Third,
Schulz (1924, 1928, 1932) overlooked Beiche’s
(1896) earlier transfer of I). intricatissima (sp. A)
to Sisymbrium. Finally, although Johnston (1929)
correctly interpreted the nomenclature of sp. B and
proposed the name S. philippianum I. M. Johnston
to replace the later homonym S. amplexicaule Phi¬
lippi, Schulz (1932) continued to use S. intricatis-
sirnum for sp. B and placed S. philippianum in its
synonymy.
Acknowledgments. I am most grateful to Melica
Munoz-Schick for her help with the Philippi types
at SGO, Carmen Ulloa for translating the abstract
into Spanish. Robert Vogt lor sending the images
of Neuontobotrys berningeri , Anthony R. Brach for
providing needed literature, and the directors and
curators of the herbaria cited. I am most thankful
to Steve L. O’Kane Jr.. Victoria C. Hollowed, Amy
McPherson, Diana Gunter, and an anonymous re¬
viewer for their valuable comments on the manu¬
script.
Literature Cited
Al-Shehbaz, I. A. 1990. The South American Eremodraba
(Brassicaoeae). Ann. Missouri Bot. Gard. 77: 602—604.
Volume 14, Number 3
2004
Al-Shehbaz
Synopsis of Neuontobotrys
257
Appel, 0. & I. A. Al-Shehbaz. 2003. Crueiferae. Pp. 75—
174 in k. Kubitzki (editor). Families and Genera of
Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg.
Baehni, C. & J. F. Macbride. 1937. Remarques sur les
Gruciferae-Sisymbrieae. Candollea 7: 291-296.
Boelcke, 0. 1961. Nota sobre Sisymbrium robustum Chod.
et Wilcz. Bol. Soc. Argent. But. 9: 387—392.
Chodat, R. & F. Wilczek. 1902. Contributions & la (lore
de la Republique Argentine. Bull. Herb. Boissier, Ser.
2. 2: 281-296.
Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Johnston, I. M. 1929. A new Chilean plant and some no-
menclatorial changes. Rev. Chilena Hist. Nat. 33: 25—
27.
Kuntze, 0. 1898. Revisio Genera Plantarum. Vol. 3, part
2. Arthur Felix, Leipzig.
Macbride, J. F. 1938. Crueiferae. Field Mus. Nat. Hist.,
Bot. Ser. 13: 937-983.
Munoz-Pizarro, C. 1960. Las Especies de Plantas Descri-
tas por R. A. Philippi en el Siglo XIX. Ediciones de la
Universidad de Chile, Santiago.
Reiche, C. 1896. Flora de Chile, Vol. 1. Imprenta Cer¬
vantes, Santiago.
Romanczuk, M. C. 1981. Novedades en el genero Sisym¬
brium (Crueiferae) y sinopsis de las especies Patagdni-
cas. Parodiana 1(1): 1—29.
- . 1982. El genero Sisymbrium (Crueiferae) en la
Argentina. Darwiniana 24: 75—156.
Schulz, 0. E. 1924. Crueiferae— Sisymbrieae. Pp. 1—388
in A. Engler (editor), Pflanzenreich IV 105(Heft 86).
Verlag von Wilhelm Engelmann, Leipzig.
- . 1928. Beilrage zur Kenntnis der Flora von Chile.
Notizbl. Bot. Gart. Berlin-Dahlem 10: 44)0-472.
- . 1932. Crueiferae variae. Notizbl. Bot. Gart. Ber¬
lin-Dahlem 11: 389-392.
- . 1936. Crueiferae. Pp. 227—658 in A. Engler &
H. Harms (editors). Die Natiirlichen Pflanzenfamilien,
Ed. 2. 17B. Verlag von Wilhelm Engelmann, Leipzig.
Stafleu, F. A. & R. S. Cowan. 1976. Taxonomic Literature,
Vol. 1: A-G. Bohn, Scheltema & Holkema, Utrecht.
Torrey, J. & A. Gray. 1838. A Flora of North America,
Vol. 1, part 1. Wiley & Putnam, New York.
Warwick, S. I. & I. A. Al-Shehbaz. 2003. Nomenclatural
notes on Sisymbrium (Brassicaceae). Novon 13: 265—
267.
- . - , R. A. Price & C. Sauder. 2002. Phylog-
eny of Sisymbrium (Brassicaceae) based on FI’S se¬
quences of nuclear ribosomal DNA. Canad. J. Bot. 80:
1002-1017.
A Synopsis of t lie South American Weberbauera (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. Twenty-two species are recognized in
Weberbauera (Brassicaceae), and a key to distin¬
guish them is given. Weberbauera dillonii, W. sca-
brifolia, and W. violacea arc described and illus¬
trated. and their distinguishing characters from
nearest relatives are discussed. The new combina¬
tions W. peruviana and W. lechleri are proposed.
Sisymbrium petraeum and .S’, fastigiatum are re¬
duced to synonymy of W. lechleri.
Resumen. Se reconocen veintidos especies en
Weberbauera (Brassicaceae), y se presenta una
clave para distinguirlas. Se describen e ilustran We¬
berbauera dillonii, W. scabrifolia y W. violacea y se
discuten los caracteres que las distinguen de otras
especies cercanas. Se proponen nuevas combina-
ciones de W. peruviana y II' lechleri. Los nombres
Sisymbrium petraeum y S. fastigiatum se reducen a
sindnimos de W. lechleri.
Key words: Brassicaceae, Sisymbrium, South
America, Stenodraba, Weberbauera.
Weberbauera 0. E. Schulz (Brassicaceae), a ge¬
nus widely distributed in Peru, Bolivia, Argentina,
and Chile (Appel & Al-Shehbaz, 2003), was re¬
cently revised and its boundaries critically delim¬
ited (Al-Shehbaz, 1990a, 1990b). The present pa¬
per continues this revision, recognizing 22 species
in Weberbauera, including three new species and
two new combinations. The two names transferred
herein to Weberbauera, lb peruviana (DC.) Al-Sheh¬
baz and W. lechleri (Fournier) Al-Shehbaz, were
originally described in Sisymbrium L.
Sisymbrium was believed to have one of its major
centers of diversity in South America (Schulz,
1924; Al-Shehbaz, 1988), but that opinion has been
shown to be incorrect (Warwick et al., 2002), and
the genus is considered to be restricted primarily
to the Old World, with only one species, S. linifol-
ium Nuttall, native to the New World (western
North America). Ravenna (1981) and Boelcke and
Romanczuk (1984) placed in Stenodraba 0. E.
Schulz some species recognized here as Weber¬
bauera. Schulz (1924, 1936) separated Stenodraba
from Weberbauera solely on the absence instead of
the presence of bracts. This is an unreliable feature
for the delimitation of genera in Brassicaceae, and
numerous other genera (e.g.. Cardamine I... Draba
L., Rorippa Scopoli, Sisymbrium ) have species with
or without bracts (Appel & Al-Shehbaz, 2003). As
indicated by Al-Shehbaz (1990a), Stenodraba docs
not merit recognition.
In the enumeration below, the information pro¬
vided includes the bibliographic citation of each
species and its type collections. If the species is
known from collections other than the type, one
representative collection for every major locale is
cited. For the full synonymy and typification of spe¬
cies 2, 3, 7, 8, 10, 12, 18—21, see Al-Shehbaz
(1990a). For a detailed description of Weberbauera
and full account of its generic synonyms and their
types, see Al-Shehbaz (1990a).
Weberbauera Gilg & Muschler, Bot. Jahrb. Syst.
42: 481. 1909. TYPE: Weberbauera densiflora
(Muschler) Gilg & Muschler = Weberbauera
sputhulifolia (A. Gray) (). E. Schulz.
Novon 14: 258-268. 2004.
Volume 14, Number 3
2004
Al-Shehbaz
South American Weberbauera
259
Key to the Species of Weberbavera
la. Plant, including fruit, densely covered with malpighiaceous trichomes; petals often pubescenl .
. 21. IP. trichocarpa
lb. Plants glabrous or with simple and/or stalked forked trichomes, rarely with malpighiaceous trichomes and
then only on basal leaves; fruit glabrous or rarely sparsely pubescent with simple trichomes; petals glabrous.
2a. Inflorescence bracteate throughout, rarely uppermost flowers ebracteate.
3a. Petals violet. 6.5— 7.5 X 3-3.5 mm . 22. W. violacea
3b. Petals white, 2-3.5(-5) X 0.7-2(-2.5) mm.
4a. Plants densely scabrous with appressed antrorse trichomes; basal leaves entire, densely
scabrous adaxially, glabrous abaxially . 16. IP scabrifolia
4b. Plants glabrous or pilose with spreading trichomes; at least some leaves dentate, repand,
lyrate, or pinnatifid, all glabrous or pubescent but never scabrous.
5a. Basal leaves not ciliale at base, sometimes glabrous or with forked trichomes .
. IB. IP spathulifolia
5b. Basal leaves conspicuously ciliate (at least basally) or densely covered with simple
trichomes.
6a. Fruit apex attenuate; style 1-3 mm long; petals 3.5-5 mm long ... 7. IP herzogii
6b. Fruit apex obtuse; style to 0.2 mm long; petals 2.5-3.5 mm long.
7a. Basal and lowermost cauline leaves lyrate-pinnatifid, ciliate; stems and
leaves glabrous; cauline leaves and bracts entire or repand; seeds 3 to 5
per locule . 1 - 4P bracteata
7b. Basal and lowermost cauline leaves denticulate or dentate, not ciliate;
stems and leaves densely pilose or hirsute; cauline leaves and bracts
coarsely dentate; seeds 6 to 15 per locule.
8a. Cauline leaves and bracts sessile; stems retrorsely pilose; fruits gla¬
brous; pedicels pilose; petals 2—2.5 mm wide . 6. IP dillonii
8b. Cauline leaves and bracts petiolate; stems hirsute; fruits hirsute; ped¬
icels glabrous; petals 0.7—1 mm wide . 14. W. peruviana
2b. Inflorescences ebracteate, rarely lowermost few flowers bracteate.
9a. Sepals persistent into fruit; fruil sparsely pubescent with retrorse simple trichomes .
. 15. IP retropilosa
9b. Sepals soon caducous; fruit glabrous.
10a. Petals yellow; fruit septum perforate; leaves adaxially densely sericeous .... 13. IP perforata
10b. Petals white; fruit septum complete; leaves glabrous or differently pubescent.
11a. Basal leaves numerous; petioles persistent, straw-colored, 3—3.5 cm long, ca. as long
as blade, overlapping and forming a dense crown . 5. IP densifolia
1 lb. Basal leaves few to several; petioles soon caducous or if persistent, then not straw-
colored, less than 3 cm long, shorter than blades, not forming a dense crown.
12a. Stems several-branched above; plants with some dendritic trichomes at least on
stems and sepals.
13a. Basal leaves ciliate with simple trichomes, pubescent with dendritic tri¬
chomes . 12. IP! parvifolia
13b. Basal leaves not ciliate, exclusively with malpighiaceous trichomes .
. 10. IP lechleri
12h. Stems simple above; plants glabrous or with simple and/or forked trichomes.
14a. Basal and cauline leaves not ciliate, sometimes glabrous or with some
branched trichomes.
15a. Leaves fleshy, glabrous; petals 6.5-8 mm long; caudex thick, 1.5-2
cm diam . 1 < . IP smithii
15b. Leaves not fleshy, often pubescent or rarely glabrous; petals 2—3.5
(^4) mm long; caudex slender, almost always less than 1 cm diam.
16a. Basal leaves filiform to narrowly linear; cauline leaves coarsely
dentate-serrate; trichomes minute, 0.03— 0.1 (-0.1 5) mm long. . . .
. 11. IP! minutipila
16b. Basal leaves oblong, ovate, lanceolate, or spatulate, very rarely
linear; cauline leaves entire to repand or dentate; trichomes
coarser, (0.2-)0.4-0.7(-l.l) mm long . 18. IP spathulifolia
14b. Basal and cauline leaves ciliate with simple trichomes.
17a. Basal leaves pinnatifid; inflorescence 1- to 3-flowered; petals less
than 2 mm long; stems less than 2 cm tall . 4. IP cymosa
17b. Basal leaves entire, rarely dentate or sinuate; inflorescence (3- or)4- to
10(to 15)-flowered; petals 2-5.5 mm long; stems more than 2 cm tall.
18a. Basal leaves semiterete, thick, linear . 20. IP suffruticosa
18b. Basal leaves flat, thin, oblong to oblaneeolate or spatulate, rare¬
ly linear-lanceolate.
19a. Style obsolete or rarely to 0.6 mm long in fruit.
260
Novon
20a. Fruil torulose; infructescence lax racemes; fruiting
pedicel slender, divaricate, 4— 8(-l2) mm long; basal
leaves entire, to 1.5 mm wide . 9. IF lagunae
20b. Fruit smooth; infructescence usually dense, sub-
umbellate; iruiting pedicels stout, subappressed,
1.5-4.5(— 7) mm long; basal leaves usually dentate,
2 — 4.5( — 6) mm wide . 3. IF colchaguensis
19b. Style 1-3 mm long in fruit, if shorter then fruits conspic¬
uously flattened.
21a. Leaves abaxially with trichomes shorter than those
adaxially or on margin; fruit torulose; stems usually
leafless; petioles of basal leaves stout, swollen .
. 8. IF imbricatifolia
21b. Leaves abaxially glabrous; fruit smooth; stems I - to
4-leaved; petioles of basal leaves slender, not swol¬
len.
22a. Basal leaves entire; petals (3.5— )4— 5 mm long;
style (0.8-)1.5-2 mm long in fruit .
. 19. IF stenophylla
22b. Basal leaves dentate; petals 2.5— 3.5 mm long;
style 0.5— 0.9(— 1.1) mm long in fruit .
. 2. IF chillanensis
1. Weberbauera bracteata ((). E. Schulz) ,). F.
Macbride, Candollea 5: 356. 1934. Pelagatia
bracteata 0. E. Schulz, Pflanzenreich IV.
l()5(Heft 86): 192. 1924. TYPE: IVru. An-
cash: Pallasca, Cordillera of Pelagatos, 4600
m, 23 J an. 1920, A. Weberbauer 7234 (holo-
type, B; isotypes, F, G).
Distribution. Known oidy from the type collection.
2. Weberbauera chillauensis (Philippi) Al-
Shehbaz, J. Arnold Arbor. 71: 244. 1990. Dra-
ba chillanensis Philippi, Anal. Univ. Chile 2:
377. 1862. TYPE: Chile. [Regibn 1 X ] Termas
de Chilian, s.d., Philippi s.n. (holotype, SCO).
Distribution. Argentina (Prov. Mendoza) and
Chile (Region VIII and IX).
Representative specimens. ARGENTINA. Mendoza:
Malargiie, Banos del Azufre, 19 Jan. 1941, Castellanos
s.n. (BA). CHILE. Region VIII: Volean Peteroa, Werder-
mann 6 04 (B, BM, E, F, G, GH, K. M, MO, NY, S, U,
UC, /). Region IX: Nuble, Cordillera de Chilian, Jaffuel
3722 (GH).
3. Weberbauera colrhugurusis (Barneoud) Al-
Shehbaz, J. Arnold Arbor. 71: 241. 1990. Car-
damine colchaguensis Barneoud, in Gay, FI.
Chile 1: 1 15. 1846. TYPE: Chile. [Region VII]
Colchagua, Cordillera del Cajon del Azufre,
cerca de volean de Talcaregue, 8000-9000 ft.
[2438—2743 rn|. s.d., C. Gay 171 (holotype, P;
isotype, G).
Distribution. Argentina (Prov. Mendoza, Neu-
qubn, and Rfo Negro) and Chile (Region IV, Me-
tropolitana, VII, VIII, X, XI, and XII).
Representative specimens. ARGENTINA. Mendoza:
Laguna Atuel, Bticher, Hjerting & Rahn 1976 (C. MO).
Neuquen: Ixjs Lagos, Filo Machete al Co. Rothleugal,
Dierre 929 (LIL). Rfo Negro: Cordon del Rfo Colorado,
Cerro Gorra. 19 Feb. 1940, Moreau s.n. (BA). CHILE.
Region IV: Choapa, La Vega Redonda, E of La Vega Es-
condida, Morrison 16995 (1)S, G, K, MO, S, UC). Region
Metropolitana: Rio Blancos, Frodin 580 (UPS). Region
VII: Colehaqua, Las Damas, Philippi 9 ll> (SCO). Region
VIII : Curicb, El Valle de los Ciegos, near volcano of Pe¬
teroa, Bridges 1120 (BM, E). Region X: Cautfn, Villa r-
riea, Neger s.n. (M). Region XI: Osorno, Paso Puyehue,
Sparre & Constance 10812 (UC). Region XII: Cerro Agu-
do, 50°49'S, 72°57'W, Arroyo X Squeo 870017 (C0NC).
4. Weberbauera cymosa Al-Shehbaz, J. Arnold
Arbor. 71: 239. 1990. TYPE: Bolivia. La Paz:
Murillo, near Palca, base of Illimani, 4800—
5000 m. 25 Feb. 1979,4. Ceballos, A. Chapin ,
./. Fernandez- Casas & E. Valddz-bermeijo 543
(holotype, C).
Distribution. Endemic to Bolivia (Depto. La
Paz).
Representative specimens. BOLIVIA. La Paz: Murillo,
La Paz-EI Alto-Valle de Zongo, 16°17'S, 68°7'W, Reek-
21752 (LPB, MO).
l he species is known thus far only from the two
collections cited above.
5. Weberbauera densifolia Al-Shehbaz, ,). Ar¬
nold Arbor. 71: 229. 1990. TYPE: Argentina.
Prov. Catamarca: Depto. Santa Marfa, Sierra
del Aconquija, 4600 in. 20 Feb. 1925. 5. Ven¬
turi 6610 (holotype, US).
Distribution. Argentina (Prov. Catamarca) and
Bolivia.
Volume 14, Number 3
2004
Al-Shehbaz
South American Weberbauera
261
Representative specimens. ARGENTINA. Catamarca:
Catamarca, Rodriguez 1360 (MO). BOLIVIA. Without lo¬
cality, Navarro Sanchez 1147 (MO).
The species is known thus far only from the three
collections cited above.
6. Weberbauera dillonii Al-Shehbaz, sp. nov.
TYPE: Peru. La Libertad: Bolivar, Ascenso
Nevado de Cajamarquilla, 3000 m, 7°08'S,
77°42'W, 10 Nov. 2001, /. Sanchez V, M. Dil¬
lon & G. Iberico 1 1 172 (holotype, MO; isotypes
not seen, CPUN, F). Figure 1.
Herba perennis retrorse pilosa, 15—25 cm alta. Folia
basalia rosulata, petiolata, oblonga vel oblanceolata, 2—6
X 0.7— 1.5 cm, denticulata, retrorse pilosa; folia caulina
sessilia, oblonga vel ovata. grosse dentata. Racemi 20-50-
flori, omnino bracteati; pedicelli fructiferi 4.5— 6.5 mm
longi, sigmoidei vel recurvati, retrorse pilosi. Sepala ob¬
longa, 2.5—3 X 0.7—1 mm; petala alba, late obovata, 3—
3.5 X 2—2.5 mm. Fructus lineares vel oblongo-lineares,
8—12 X 1.5—2 mm, glabri, curvati; stylus 0.1— 0.2 mm
longus; semina oblonga, 1—1.3 X 0.6— 0.7 mm; cotyle-
dones incumbentes.
Perennial herbs, retrorsely pilose throughout ex¬
cept for petals and fruits; caudex with petiolar re¬
mains of previous years; trichomes simple, retrorse;
stems 15—25 cm long, erect or ascending, densely
pilose with appressed retrorse trichomes, usually a
few from the base, simple or branched above,
densely pilose with appressed retrorse trichomes.
Basal leaves rosulate; petiole 1—3 cm long, pubes¬
cent as stems; leaf blade oblong to oblanceolate, 2—
6 X 0.7— 1.5 cm, densely and retrorsely pilose on
both sides, base cuneate to attenuate, margin den¬
ticulate, apex subacute; middle cauline leaves ses¬
sile, oblong to ovate, 1—2 cm X 4—8 mm, base cu¬
neate, margin coarsely dentate, apex acute,
pubescent as basal leaves. Raceme 20- to 50-flow-
ered, bracteate throughout, dense, slightly elongat¬
ed in fruit; bracts sessile, cuneate to subauriculate
at base, similar to cauline leaves but progressively
smaller upward; fruiting pedicels 4.5— 6.5 mm long,
sigmoid and slightly recurved, retrorsely pubescent.
Sepals oblong, 2.5—3 X 0.7—1 mm, not saccate at
base, sparsely pubescent below apex, ascending;
petals white, broadly obovate, 3—3.5 X 2-2.5 mm,
not clawed, apex rounded; filaments 2-2.5 mm
long; anthers oblong, ca. 0.5 mm long. Fruits linear
to oblong-linear, 8-12 X 1.5—2 mm, glabrous,
curved, slightly flattened at a right angle to the sep¬
tum; valves smooth, not veined; septum complete;
style 0.1— 0.2 mm long; stigma entire; seeds 8 to 10
per locule, oblong, brown, 1—1.3 X 0.6— 0.7 mm;
funieles slender along entire length; cotyledons in¬
cumbent.
Weberbauera dillonii , which is known only from
the type collections, is named in honor of Michael
Dillon, one of the collectors of the type material
who has done extensive fieldwork in Peru. It be¬
longs to a group of four other species with fully
bracteate racemes. It is most closely related to W.
peruviana , from which it is easily distinguished by
having sessile cauline leaves and bracts, retrorsely
pilose stems and leaves, glabrous fruits, pilose
fruiting pedicels, and broadly obovate petals 2-2.5
mm wide. Weberbauera peruviana has petiolate cau¬
line leaves and bracts, hirsute stems and leaves,
sparsely to densely hirsute fruits, glabrous fruiting
pedicels, and narrowly spatulate petals 0.7—1 mm
wide.
The new species resembles (in its small petals
< 4 mm long, minute styles to 0.2 mm long, and
bracteate racemes) Weberbauera bracteata (0. F.
Schulz) J. F. Macbride, a species known only from
the type collection made by August Weberbauer on
January 1920 from Depto. Ancash and which has
not yet been re-collected. From this, W. dillonii is
distinguished by having densely and retrorsely pi¬
lose stems and leaves, denticulate and non-ciliate
basal leaves, coarsely dentate cauline leaves and
bracts, broadly obovate petals 2-2.5 mm wide, sig¬
moid and slightly recurved fruiting pedicels, linear
to oblong-linear and curved fruits, veinless valves,
and eight to ten seeds per locule. By contrast, W.
bracteata has glabrous stems and leaves, lyrate-pin-
natifid and ciliate basal and lowermost cauline
leaves, entire or repand cauline leaves and bracts,
spatulate petals ca. I mm wide, ascending fruiting
pedicels subappressed at base to rachis, oblong and
straight fruits, prominently veined valves, and three
to five seeds per locule.
From the other two Peruvian species of Weber¬
bauera with fully bracteate racemes, W. spathuli-
folia (A. Gray) 0. E. Schulz and W. herzogii (O. E.
Schulz) Al-Shehbaz, W. dillonii is separated by hav¬
ing stems and leaves densely and retrorsely pilose
(instead of glabrous or sparsely pubescent with
spreading trichomes) and funieles slender along the
entire length (instead of distinctly flattened at
base). From W. spathulifolia it is also separated by
having exclusively simple instead of forked and
submalpighiaceous trichomes mixed with simple
ones, and from W. herzogii it differs by the minute
style 0. 1-0.2 mm long instead of (0.8— )1.2— 2.5 mm
long.
7. Weberbauera herzogii (0. E. Schulz) Al-
Shehbaz, J. Arnold Arbor. 71: 236. 1990. Sar-
codraba herzogii O. E. Schulz, Notizbl. Bot.
Gart. Berlin-Dahlem 10: 563. 1929. TYPE:
Bolivia Felsplatten der Hiigel iiber dem Titi-
262
Novon
Figure 1. Weberbauera dillonii Al-Shehbaz. — A. Plant with remains of infructescence (left) of previous season. — I?.
Sepal. — C. Petal. — I). Bract, fruiting pedicel, and fruit. Drawn from the MO holotype (/. Sdnchez V., M. Dillon & G.
Iberico 1 1 172) by Al-Shehbaz. Seale: A = 1 cm, B— I) = 1 mm.
Volume 14, Number 3
2004
Al-Shehbaz
South American Weberbauera
263
cacasee bei Guagqui, ca. 3900 m, s.d., T. Her¬
zog 2510 (holotype, B; isotypes, G. S, Z).
Distribution. Argentina (Prov. Jnjny), Bolivia
(Depto. La Paz and Potosf), and Peru (Depto. Puno).
Representative specimens. ARGENTINA. Jujuy: Hu-
mahuaca, Esquinas l>lancas, between Tres Cruces and Hu-
mahuaca, Ruthsatz 13/18 (GH). BOLIVIA. I,a Paz: I11-
gavei, Guaqui, Asplund 2226 (S, UPS). Potosf: Frias, ca.
25 kin N of Yocalla towards Ventanilla, Wood 11743 (K,
MO). PERU. Puno: Puno, Shepard 38 (GH, US).
8. Weberbauera imbricatifolia (Barneoud) Al-
Shehbaz. J. Arnold Arbor. 71: 247. 1990. Dra-
ba imbricatifolia Barneoud, in Gay, FI. Chile
I: 158. 1846. TYPE: Chile. [Region IV] Cor¬
dillera de Coquimbo, 12.000 ft. [3658 m], C.
Gay s.n. (holotype, P; isotype. B).
Distribution. Argentina (Prov. San Juan) and
Chile (Region IV, V, and VI).
Representative specimens. ARGENTINA. San Juan:
Dept. Iglesia, 31 km W of Arrequintin, 15 Feb. 1985,
Hunziberg & Gamenro s.n. (OS). CHILE. Region IV: Co¬
quimbo, Cordillera de Combarbala, Laguna Tibia,
31°17'S, 70°45'W, Jib's 4864 (CONC, M). Region V: Pe-
torca, 5 km S of Junta de Piuquenes, Ri'o Sobrante, Mor¬
rison 17290 (l)S, k. UC). Region VI: Santiago, Andes of
San Jos6, Gay 1028 (SGO).
9. Weberbauera laguuae (0. E. Schulz) Al-
Shehbaz. J. Arnold Arbor. 71: 246. 1990. Sten-
odraba suffruticosa (Barneoud) 0. E. Schulz
var. lagunae 0. E. Schulz, Notizbl. Bot. Gart.
Berlin-Dahlem 10: 469. 1928. TYPE. Chile.
Vallenar, Cordillera Laguna Chica, ca. 4000
in, Jan. 1924, E. Werdermann 262 (holotype,
B; isotypes, BM, CONC, E, E, G, GH, k, MO,
UC).
Distribution. Endemic to Chile (Region III).
Representative specimens. CHILE. Region III: vicin¬
ity of Laguna Valeriano, Johnston 6 065 (CONC, GH, US).
10. Weberbauera leehleri (Fournier) Al-Sheh¬
baz, comb. nov. Basionvm: Sisymbrium leehleri
Fournier, Rech. Crucif. 129. 1865. TYPE:
Chile. Terra Pehuenchorum, Dec. 1854, W.
Lechler 3080 (holotype, P; isotypes, G[2], K,
P[ 2 1).
Sisymbrium petraeum Philippi, Linnaea 28: 668. 1856,
non S. petraeum (L.) Delarbre, FI. Auv. ed. 2: 349.
1800. Syn. nov. Stenodraba glareosa Ravenna, Nor¬
dic J. Bot. 1: 141. 1981. TYPE: Chile. [Prov. Nuble]
“in Andibus prope oppidum Chilian,” R. A. Philippi
s.n. (holotype, SGO 49254).
Sisymbrium fastigiatum Philippi, Anal. Univ. Chile 41:
670. 1872. Syn. nov. TYPE: Chile. [Prov. Santiago|:
Mina Cristo, valley of Maipo, B. Ddvilla s.n. (lecto-
type, designated by Munoz-Schick (1973: 30), SGO
45138).
Distribution. Argentina (Prov. Neuqufm and
Rio Negro) and Chile (Region IX).
Representative specimens. ARGENTINA. Neuquen:
Cerro Colohuincu 1, Comber 876 E (K). Rfo Negro: Bar-
iloche, lago Moreno Este, Hosseus 107 (RAF). CHILE. Re¬
gion IX: Cordillera of Chilian, 1856—1857, P. Germain
s.n. (G, K, W).
Munoz-Schick (1973) designated the above lec-
totype of Sisymbrium fastigiatum, but Ravenna
(1981) overlooked that lectotypification and erro¬
neously designated another one of Davilla’s collec¬
tions (SGO 47168) as the lectotype. Ravenna
(1981) treated S. petraeum Philippi, S. leehleri, S.
caespitosum Philippi, and S. fastigiatum as four in¬
dependent species that he placed in the genus
Stenodraba 0. E. Schulz, but the alleged differenc¬
es given by him are trivial and quantitative in
nature. All are indistinguishable in every aspect,
especially in their trichomes, which are mal-
pighiaceous on the basal leaves and dendritic on
the stems, cauline leaves, and sepals. The density
of trichomes is quite variable in Weberbauera lech-
leri, and the holotype has malpighiaceous trichomes
restricted to the margins of basal leaves, whereas
the isotypes have such trichomes on both leal sur¬
faces. As indicated above, Stenodraba does not
merit recognition independent of Weberbauera.
Schulz (1924) first placed Weberbauera leehleri
in Heterothrix (B. L. Robinson) Rydberg and later
(Schulz, 1936) in Pennellia Nieuwland, but he used
the illegitimate later homonym Sisymbrium pe¬
traeum Philippi as the basionym for his combina¬
tions. As indicated by Al-Shehbaz (1990c) and Rol¬
lins (1980), the generic name Heterothrix is an
illegitimate later homonym for a genus in the Apo-
cynaceae, and the correct name for the mustard
genus is Pennellia. Pennellia differs from Weber¬
bauera by having a cup-shaped calyx with purple
sepals about as long as the petals. Weberbauera has
an oblong calyx with green sepals distinctly shorter
than the petals.
1 1. Weberbauera niinutipila Al-Shehbaz, J. Ar¬
nold Arbor. 71: 231. 1990. TYPE: Peru. Puno:
San Roman, Puno— Arequipa road at km 1 12.8,
ca. 4 km (air) E of Tinocopalca (km 119.5),
ca. 10 km W of road turnoff to Sta. Lucia (at
km 101), ca. 4000 m, 12 Jan. 1963, //. //. &
C. M. litis with D. & V. Ugent 1455 (holotype,
GH: isotype, WIS).
264
Novon
Distribution. Bolivia (Depto. La Paz) anti Peru
(l)epto. Arequipa. Puno).
Representative specimens. BOLIVIA. La Paz: Valle
Chuquiaguillo, Asplund 1888 (S, UPS). PERU. Arequipa:
Arequipa, S of Sta. Lucfa on roarl from Puno to Arequipa,
ca. 4—5 km E of Sta. Lucfa, litis & llgent 1415 (Vi IS).
12. Weberbauera parvifolia (Philippi) Al-Sheh-
baz, J. Arnold Arbor. 71: 248. 1990. Sisym¬
brium parvifolium Philippi, Linnaea 28: 667.
1856. TYPE: Chile. Cordillera de Linares,
s.d., Germain s.n. (holotype, SCO).
Distribution. Argentina (Prov. Neuquen) and
Chile (Region Metropolitana, VII, and VIII).
Representative specimens. ARGENTINA. Neuquen:
Cordillera del Viento, cruzada de Trieao, Malal al Cajon
de Butald, Boelcke et al. 11565 (BAA, SI). CHILE. Re¬
gion Metropolitana: Monumento Natural el Morado,
Price 1409 (MO). Region VII: Colchagua, San Fernando,
Trrrnas El Falco, Montero 6043 (SI). Region VIII: Cu¬
bed, Termas del Flaco (Andes de Colchagua), Aravena
33349 (G, GH, MO).
13. Weberbauera perforata Al-Shehbaz, Ann.
Missouri Rot. Card. 77: 841. 1990. TYPE:
Peru. Cuzco: Nevado Auzangate |as Ausanga-
te], 71°24'W, 13°38'S, 4800 m. 1 1 May 1957,
R. Hirsch PI 255 (holotype, GH).
Distribution. Known only from the type collec¬
tion.
14. Weberbauera peruviana (DC.) Al-Shehbaz,
comb. nov. Basionym: Sisymbrium peruvianum
DC., Syst. Nat. 2: 477. 1821. TYPE: Peru. H.
Ruiz & ./. A. Pavdn s.n. (holotype, BM; iso¬
types, B, G-DC).
Distribution. Argentina (Prov. Catamarca, Ju-
juy, and Tucuman). Bolivia (Depto. Chuquisaca,
Cochabamba, La Paz. Oruro, and Potosf), and Peru
(Depto. Ayacucho, Cuzco, Huancavelica, Lima,
Puno, and Tacna).
Representative specimens. ARGENTINA. Catamarca:
Cerro Overo, Rriicher 95017 (M). Jiijuy: Humahuaca,
Mina Aguilar, 14 km N of Molina de Minera Aguilar. Hun-
ziker et al. 10568 (SI). Tucuman: Tafi, cumbres Calcha-
(|ufes. Co. Bayo. 20‘1.1'S. 65°42'W, Gomez-Sosa & Miil-
gura 204 (M(), SI). BOLIVIA. Clniqnisaca: Zudanez,
Cordillera de los Sombreros, along Ida and Azurduy road.
Wood A- Carretero 16149 (k. MO). Cochabamba: (,)uilla-
collo, camino Sipe Sipe a Kami, cuenca del valle de Co¬
chabamba. Beck et id. 18064 (MO). La Paz: 47.2 km NF
of Penas on road to Valle Hiehucota, Solomon 4957 (GH.
MO). Oruro: Sajama, Curahura de Carangas, 10 km SW
on backload, 68°25'W, 17°50'S, Johns 83-39 (F. I PH.
MO). Potosf: 10 km S of Potosf, ILc.sV 6359 (GH, MO).
PERU. Ayacucho: Luncanas, Pampa Galeras, Tovar 6704
(MO). Cuzco: Urubamba, Chichero, Cuper, Qoriwayra-
china, 72°0'W, 13°25'S, C. & E. Franquemonl 314 (F).
Huancavelica: Morococha, Grant 7573 (A, F). Lima:
Huarochiri, laguna de Tuctucocha, Cerrate 1866 (GH).
Puno: Macusani, road from Ollachea to Macusani,
70°30'W, 13°53'S, M. & K. Weigend 2000/120 (MO). Tac-
na: Tarata, Corillera del Barroso, Torre et al. 2135 (MO).
As a result of its exclusion from Sisymbrium
(Warwick et al., 2002) and the discovery herein of
its nearest relative, Weberbauera dillonii , S. peru¬
vianum is appropriately transferred to Weberbauera
after being recognized in Sisymbrium since its dis¬
covery more than 180 years ago. In every aspect of
morphology (see discussion under W. dillonii ), the
species is at home in Weberbauera.
15. Weberbauera relropilosa Al-Shehbaz, J.
Arnold Arbor. 71: 234. 1990. TYPE: Bolivia.
Depto. La Paz: Prov. Pacajes, Charana, 4000
m, 2 Mar. 1921, Asplund s.n. (holotype, US).
Distribution. Endemic to Bolivia (Depto. La
Paz).
Representative specimens. BOLIVIA. La Paz: Pacajes,
Charana, Asplund 2700 (S, UPS).
The species is known thus far only from the two
collections above.
16, Weberbauera seabrifolia Al-Shehbaz, sp.
nov. TYPE: Peru. Iluanuco: Dos de Mayo, road
from Pachas to Plata, 4030 m, 9°42'S,
76°47'W. flat puna, 18 Mar. 2001. M. Weigend,
K. Weigend. M. Hinder & E. Rodriguez 5220
(holotype, M; isotype, MO). Figure 2.
Herba perennis, 3-6 cm alta. Folia basalia petiolata,
oblonga, 7-20 X 3—9 mm, integra, superne antrorse sca-
bra, inferne glabra; folia eaulina sessilia, oblaneeolata vel
linearia, integra. Racemi 10— 27-flori, omnino bracteati;
pedicelli fructiferi 4-7 mm longi, recti, pubescentes. Se-
pala oblonga, 2. 5- .3 X ca. I mm; petala alba, spathulata,
3.5—4 X ca. 1.5 mm. Fructus lineares vel oblongo-line-
ares, 5—9 X ca. 1 .5 mm, glabri, curvati: stylus ca. 1 mm
longus; semina oblonga, ca. 1 X 0.5 mm; cotyledones in-
cumbentes.
Perennial herbs; caudex slender, with leaf re¬
mains of previous years; trichomes simple, op¬
pressed, antrorse; stems 2 to 5 from base, un¬
branched, 3—6 cm long, decumbent or ascending,
densely pubescent with oppressed, antrorse tri¬
chomes to 0.6 mm long and to 0.04 mm wide at
base. Basal leaves rosulate; petiole 4—10 mm long,
ciliate; leal blade oblong, 7—20 X 3—9 mm, abax-
ially glabrous, adaxially scabrous with antrorse, ap-
pressed trichomes to 0.4 mm long and to 0.12 mm
wide at base, base cuneate, margin entire, apex ob¬
tuse; cauline leaves sessile or nearly so, oblanceo-
Volume 14, Number 3
2004
Al-Shehbaz
South American Weberbauera
265
figure 2. Weberbauera scabrifolia Al-Shehbaz. — A. Plant. — B. Trichomes of adaxial surface of basal leaves. — C.
Trichomes of stems and pedicels. — D. Petal. — E. Fruit and fruiting pedicel. — F. Seed and funicle. Drawn from the
M holotype (M. Weigend, K. Weigend, M. Hinder & E. Rodriguez 5220) by Al-Shehbaz. Scale: A = 1 cm. B, C = 0.4
mm, D— F = 1 mm.
266
No von
late to linear, 5—10 X 1—4 mm, base attenuate,
margin entire, upper ones and bracts ciliate and
glabrous except midvein, trichomes longer and nar¬
rower than those of basal leaves. Raceme 10- to
27-flowered, bracteate throughout, dense, slightly
elongated in fruit; bracts sessile, similar to cauline
leaves but progressively smaller upward; fruiting
pedicels 4—7 mm long, straight, suberect to as¬
cending, antrorsely pubescent. Sepals oblong, 2.5—
3 X ca. 1 mm, not saccate at base, sparsely [al¬
bescent, ascending; petals white, spatulate, 3. 5-^4
X ca. 1 .5 mm, apex rounded, claw' ca. 1.5 mm long;
filaments ca. 2.5 mm long; anthers oblong, ca. 0.5
mm long. Fruits linear to oblong-linear, 5-9 X ca.
1 .5 mm, glabrous, curved, terete; valves smooth,
with a distinct midvein; septum complete; style ca.
1 mm long; stigma entire; seeds 5 to 12 per locule,
oblong, brown, ca. I X 0.5 mm; funicles thickened
at base; cotyledons incumbent.
Weberbauera scabrifolia, which is known only
from the type gathering, is easily separated from all
species of the genus by its appressed, antrorse, sim¬
ple trichomes much thicker and shorter on the bas¬
al leaves than on the rest of the plant and by having
basal leaves glabrous abaxially and densely sca¬
brous adaxiallv.
17. Weberbauera smilliii Al-Shehbaz, J. Arnold
Arbor. 71: 233. 1990. TYPE: Peru. Ancash:
Yungay, Huascaran National Park, Llanganuco
Sector, Quebrada Ancosh at Portachuelo,
77°35'W, 9°03'S, 31 Dec. 1984, D. /V. Smith
& h. Godwin BB94 (holotype, MO).
Distribution. Endemic to Peru (Depto. Ancash).
Representative specimen. PERU. Ancash: Yungay,
Huascaran National Park, Llanganuco Sector, Quebrada
Ancosh at Portachuelo, Smith I I298A (MO).
The species is known thus far only from the two
collections above.
18. Weberbauera spatbulifolia (A. Gray) 0. E.
Schulz, Pflanzenreich IV. 105(1 left 86): 193.
1924. Basionym: Sisymbrium spathulifolium
A. Gray, U.S. Expl. Exped. Phan. 15(1): 60.
1854. TYPE: Peru. [Junfn]: Orbajillo, W ilkes
Expedition, Anonymous s.n. (holotype, US; iso¬
type, NY).
Distribution. Argentina (Prov. Catamarca, Jujuy,
and La Rioja), Bolivia (Depto. La Paz, Oruro, and
Potosf), and Peru (Depto. Ancash. Huancavelica. Ju¬
nto, Moquegua, Pasco, Puno, and Santa Rosa).
Representative specimens. ARGENTINA. Catamarca:
Rfo Potrero, Sleumer 1905 (8). jujuy: Hurnahuaca, Mina
Aguilar, Sleumer 5402 (LIE). La Rioja: Sierra Famatina,
Cueva de Pdrez, Hieronymus A ; Niederlein 376 (B). BO¬
LIVIA. La Paz: Larecaja, vicinity of Combaya, Mandon
914 (BM, G, P, k). Oruro: Abaroa, Challepata, Asplund
3293 (UPS). Potosf: Frias, Cerro Potosf, Petersen & Hjert-
ing 1030 (C, LIL). PERU. Ancash: Carhuds, lluascardn
National Park, Quebrada Los Cedros, Smith , Valencia &
Minaya 9924 (F, GH. MO). Huancavelica: Viseo, Mac-
bride & Featherstone 590 (F, G, NY). Moquegua: Mo¬
quegua, above Torata, Weberbauer 7471 (BM, F, G, US).
Pasco: Cerro, Cerro de Pasco, Macbride 3065 (CAS, F,
US). Puno: Carabaya, Antapampa, Vargas 6837 (F). San¬
ta Rosa: Cuzco, Stafford 512 (BM. k).
19. Weberbauera stenopbylla (Leybold) Al-
Shehbaz, J. Arnold Arbor. 71: 245. 1990. Dra-
ba stenophylla Leybold, Anal. Univ. Chile 16:
679. 1859. T\ PE: Chile. Cord. Santiago, Cerro
Colorado, Mapocho Valley, 6000—7000 ft.
[1829—2134 m], s.d., Leybold s.n. (lectotype,
designated here, the plate accompanying the
original publication).
Distribution. Endemic to Chile (Region VIM).
Representative specimens. CHILE: Region VIII: El
Valle de los Ciegos, near volcano of Petreroa, Bridges
1121 (BM, E, GH, k).
As indicated by Al-Shehbaz (1990a), no type
material of this species has been found, and the
well-illustrated plate accompanying the detailed
original description of Draba stenophylla is desig¬
nated herein as the lectotype.
20. Weberbauera suffruticosa (Barneoud) Al-
Shehbaz, J. Arnold Arbor. 71: 247. 1990. Dra¬
ba suffruticosa Barneoud, in Cay, FI. Chile 1:
157. 1846. TYPE: Chile. Cordillera Ovalle,
12,000 ft. [3658 m], C. Gay s.n. (holotype, P;
isotype, B).
Distribution. Endemic to Chile (Region IV).
Representative specimen. CHILE. Region IV: Quebra¬
da Larga [30°44'S, 70°23'W], Jiles 3408 (CONC).
21. Weberbauera tricliocarpa (Muschler) J. F.
Macbride, Candollea 5: 356. 1934. Basionym:
Eudema trichocarpum Muschler, Bot. Jarhb.
Syst. 40: 276. 1908. TYPE: Peru. Above Lima,
near Alpamia [Alpamina], 4500 m, 2 Mar.
1904, A. Weberbauer 5119 (holotype, B).
Distribution. Endemic to Peru (Depto. Ancash,
Lima, and Pasco).
Representative specimens. PERU. Ancash: Pallasca,
Conchucds, Weberbauer 7229 (B. F, GH). Lima: Rfo Blan¬
co, Macbride 2990 (F, GH. MO, NY. US). Pasco: Cerro
de Pasco, Macbride 3073 (CAS, F, GH, MO, NY, US).
22. Weberbauera violacea Al-Shehbaz, sp. nov.
Volume 14, Number 3
2004
Al-Shehbaz
South American Weberbauera
267
Figure 3. Weberbauera violacea Al-Shehbaz. — A. Plant. — B. Bract. — C. Sepal. — D. Petal. — E. Young fruit. Drawn
from the MO holotvpe (A. Sagdstegui A. et al. 11175) by Al-Shehbaz. Scale: A = 1 cm. B-E = 1 mm.
TYPE: Peru. Cajamarca: Cajabamba, Caja-
bamba-Luchubamba, jalca vegetation, 3800
m, 1 7 Nov. 1983, A. Sagdstegui A. et al. 11175
(holotype, MO; isolype, HUT). Figure 3.
Herba perennis, 4—15 cm alta. Eolia basalia rosulata,
petiolata, spathulata, 1. 5-3.5 X 1-2.2 cm, integra vel re¬
panda; folia caulina sessilia, oblonga vel oblanceolata, in¬
tegra vel repanda, basi ciliata. Racemi 20— 35-flori, om-
nino bracteati; pedicelli Horiferi 7-10 mm longi, recti.
268
Novon
subappressi, glabri. Sepala oblonga, 4—5 X 1.5—2 mm;
petala violacea, late obovata, 6.5— 7.5 X 3—3.5 mm; stylus
1 .5—2 mm longus.
Perennial herbs, subglabrous or sparsely pubes¬
cent along petiole ami at base of cauline leaves;
trichomes simple, spreading, to 1.5 mm long; eau-
dex thick, simple; stems 4—15 cm long, decumbent
or ascending, unbranched, glabrous. Basal leaves
rosulate; petiole 0.5-3. 5 cm long, ciliate; leaf blade
spatulate, 1.5— 3.5 X 1-2.2 cm, glabrous, base eu-
neate to attenuate, margin entire or repand, apex
rounded; cauline leaves sessile, oblong, 6—20 X I -
5 mm, base cuneate, margin entire or repand, apex
obtuse, sparsely ciliate at base. Raceme 20- to 35-
flowered, bracteate throughout, dense; racliis
straight, glabrous; bracts sessile, similar to cauline
leaves but progressively smaller upward; flowering
pedicels 7—10 mm long, erect to ascending,
straight, subappressed, glabrous. Sepals oblong, 4—
5 X 1 .5—2 mm, not saccate at base, glabrous or
with a few trichomes below apex, ascending; petals
violet, broadly obovate, 6.5— 7.5 X 3—3.5 mm, not
clawed, apex rounded; filaments 2—3 mm long; an¬
thers oblong, 0.6— 0.8 mm long; style 1.5-2 mm
long. Immature fruits linear, ca. 8X1 mm; valves
glabrous, with a conspicuous midvein; seeds not
seen.
Weberbauera violacea, which is known only from
the type collection, is readily distinguished from all
other species of Weberbauera by having violet, large
petals 6.5— 7.5 mm long and racemes bracteate
throughout. All other species of the genus have
white or rarely yellow (If; perforata) flowers almost
always less than 5 mm long. The only exception is
W. smithii, a species with ebracteate racemes and
white flowers 6.5—8 mm long. Petal size and color
easily separate the new species from those with ful¬
ly bracteate racemes (see lead 2a in the key above).
Weberbauera violacea is most closely related to
W. herzogii and W. scabrifolia. All three differ from
the remaining species of Weberbauera by a combi¬
nation of fully bracteate racemes, attenuate, slender
styles 1-3 mm long, and receptacle considerably
wider than the pedicel.
Acknowledgments. I am much indebted to Olga
Martha Montiel for translating the summary to
Spanish, to Henk van der Werff for correcting the
Latin, to Michael Dillon for sending his Peruvian
collections of mustards for my study, and to Susan-
ne Renner for sending a loan of South American
mustards and for allowing a plant of the type col¬
lection of Weberbauera scabrifolia he deposited at
MO. 1 am grateful to Steve L. O' Kane Jr., Neil A.
Harriman, and Victoria C. Hollowell for valuable
comments on the manuscript. I thank the directors
and curators of the herbaria cited above.
I .iterature Cited
Al-Shehbaz, I. A. 1988. The genera of Sisymbrieae (Cru-
eiferae; Brassicaceae) in the southeastern United States.
.1. Arnold Arbor. 69: 213—237.
- . 1990a. A revision of Weberbauera (Brassicaceae).
J. Arnold Arbor. 71: 221-250.
- . 1990b. Weberbauera perforata (Brassicaceae), a
new species from Peru. Ann. Missouri Bot. Card. 77:
481-482.
- . 1990c. New or noteworthy species in the South
American genera Mancoa, Pennellia, and Sisymbrium.
Harvard Pap. Bot. 2: 1 1—16.
Appel, 0. & I. A. Al-Shehbaz. 2003. Cruciferae. Pp. 75—
174 in: K. Kubitzki (editor). Families and Genera of
Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg.
Boelcke, O. & C. Uomanczuk. 1984. Cmciferae. FI. Pa-
tagonica, Vol. 4A: 373—544. INTA, Buenos Aires.
Munoz-Schick. M. 1973. Complemento de “Las especies
de plantas deseritas por H. A. Philippi durante del siglo
XIX.” Anal. Univ. Chile 128: 5-69.
Ravenna, P. 1981. Taxonomic notes on certain Chilean
Cruciferae. Nordic J. Bot. 1: 140—142.
Rollins, R. C. 1980. The genus Pennellia in North Amer¬
ica. Contr. Gray Herb. 210: 5-21.
Schulz, (). F. 1924. Cruciferae— Sisymbrieae. Pp. 1-388
in: A. Engler (editor). Das Pflanzenreich IV. 105(Heft
86). Verlag von Wilhelm Engelmann, Leipzig.
- . 1936. Cruciferae. Pp. 227—658 in: A. Engler &
H. Harms (editors). Die natiirlichen Pflanzenfamilien,
ed. 2. 17B. Verlag von Wilhelm Engelmann, Leipzig.
Warwick, S. I., 1. A. Al-Shehbaz, R. A. Price & C. Sander.
2002. Phylogeny of Sisymbrium (Brassicaceae) based on
ITS sequences of nuclear ribosomal DNA. Canad. J.
Bot. 80: 1002-1017.
Transfer of Arabidopsis gamosepala and Torularia brachycarpa to
Braya (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
Suzanne I. Warwick
Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central
Experimental Farm, Ottawa, Ontario K1A 0C6, Canada. warwicks@agr.gc.ca
Abstract. The new combinations Braya gamo¬
sepala and B. brachycarpa (Brassicaceae) are pro¬
posed. The distinguishing characters, especially of
habit and fruit and trichome morphology, separating
Arabidopsis, Braya, anti Neotorularia (formerly To¬
rularia) are discussed.
Key words: Arabidopsis, Brassicaceae, Braya,
Neotorularia.
Arabidopsis gamosepala Hedge (Brassicaceae),
which is endemic to Afghanistan (Hedge, 1968),
was transferred to Neotorularia Hedge & J. Leonard
by Al-Shehbaz and O’Kane (1997) and has been
maintained in that genus to the present. It was ex¬
cluded from Arabidopsis Heynhold by these authors
because it has pubescent, torulose fruits, and den-
dritieally branched trichomes. As delimited by
O’Kane and Al-Shehbaz (1997), Al-Shehbaz et al.
(1999), and Al-Shehbaz and O’Kane (2002), Ara¬
bidopsis lias glabrous, non-torulose fruits and a
mixture of simple, forked, or rarely stellate stalked
trichomes.
Torularia brachycarpa Vassilczenko, originally
collected only from Tajikistan (Vassilczenko, 1939),
is now documented as widespread in four provinces
in China (Zhou el ah, 2001). It was transferred to
Neotorularia (Leonard, 1986) because Torularia
(Cosson) 0. E. Schulz is an illegitimate later hom¬
onym of the red algae Torularia Bonnemaison.
Recent phylogenetic studies, based on ITS se¬
quences of nuclear ribosomal DNA and frnL intron
of chloroplast DNA (Warwick et ah, 2004), on
Braya Sterenberg & Hoppe and Neotorularia (dear¬
ly showed that Arabidopsis gamosepala, Neotoru¬
laria brachycarpa (Vassilczenko) Hedge & Leonard,
and B. humilis (C. A. Meyer) B. L. Robinson (N.
humilis (C. A. Meyer) Hedge & Leonard) are well
nested within Braya and are most closely related
to species of this genus with linear fruits, including
B. alpina Sterenberg & Hoppe, the type species of
Braya. These four species differ from all members
of Neotorularia, including the generic type N. to-
rulosa (Desfontaines) Hedge & Leonard, by being
perennials with a well-defined basal rosette (instead
of annuals with no basal rosette) and by having at
least basally bracteate instead of ebracteate ra¬
cemes. On the basis of molecular and morphologi¬
cal data, Arabidopsis gamosepala and Neotorularia
brachycarpa are best accommodated in Braya, and
their transfer to this genus is herein effected.
Braya brachycarpa (Vassilczenko) Al-Shehbaz &
Warwick, comb. nov. Basionym: Torularia bra¬
chycarpa Vassilczenko, in Komarov, Fh URSS
8: 635. 1939. TYPE: Tajikistan. Pamir: near
Ak-baital, confluence of Murgab, ca. 3900 m.
7 July 1901. M. 1. Alexeenko 2391 (holotype,
LE).
Braya brachycarpa is easily distinguished from
the remaining species of Braya by having racemes
bracteate throughout or rarely only along the low¬
ermost part, oblong to linear fruits widest basally
and 3— 10(— 15) mm long, and seeds sub-biseriate
only at the basal portion of the fruit.
Braya gamosepala (Hedge) Al-Shehbaz & War¬
wick, comb. nov. Basionym: Arabidopsis ga¬
mosepala Hedge, in K. II. Rechinger, Flora
Iranica 57: 334. 1968. TYPE: Afghanistan.
Munjan: above Anjuman valley, near Anju-
man, 3100 m, 14 Aug. 1965, I). Podlech
12379 (holotype, M: isotypes, E. M).
The occurrence of gamosepaly, although rather
rare in the Brassicaceae (e.g., Brayopsis gamose¬
pala Al-Shehbaz, Desideria mirabilis Pampanini,
Sisymbrium gamosepalum Hedge), is not consid¬
ered to be a useful character at the generic rank
(Al-Shehbaz, 2001), but readily distinguishes B.
Novon 14: 269-270. 2004.
270
Novon
gamosepala from t lie remaining species of the ge¬
nus.
Acknowledgments. We are grateful to Thomas
G. hammers and Victoria C. Hollowed for their crit¬
ical comments on the manuscript.
I .iterature Cited
Al-Shehbaz, I. A. 2001. A review of gamosepaly in the
Brassicaceae and a revision of Desideria, with a critical
evaluation of related genera. Ann. Missouri Bot. Card.
87: 549-563.
- & S. L. O’Kane Jr. 1997. Arabidopsis gamosepala
and A. tuemurnica belong to Neotorularia (Brassica-
ceae). Novon 7: 93—94.
- & - . 2002. Taxonomy and phylogeny of 4r-
abidopsis (Brassicaceae). (22 August, 2002). In: C. R.
Somerville & E. M. Meyerow it/, (editors). The Arabidop¬
sis Book, American Society of Plant Biologists, Rock¬
ville, Maryland, doi/10. 1 199/tab. 0001 (http://
www.aspb.org/publications/arabidopsis/).
- . - & R. A. Price. 1999. Generic placement
of species excluded from Arabidopsis. Novon 9: 296—
307.
Hedge. I. C. 1968. Sisymbrieae. Pp. 309—342 in: K. 11.
Rechinger (editor), Flora Iranica 57. Akademisehe
Druck-u.Verlagsanstalt, Graz, Austria.
Leonard, J. 1986. Neotorularia Hedge and J. Leonard nom
g^n^rique nouveau de Cruciferae. Bull. Jard. Bot. Nat.
Belg. 56: 389-395.
O’Kane, S. I... Jr. & I. A. Al-Shchbaz. 1997. A synopsis
of Arabidopsis. Novon 7: 323—327.
Vassilczenko, I. T. 1939. Sisymbrium. Pp. 59—69, 635—
636 in: V. E. Komarov (editor), Flora URSS, Vol. 8.
Akademii Nauk SSSR, Moscow and Leningrad.
Warwick, S. I., I. A. Al-Shehbaz, C. Sander. J. G. Harris
& M. Koch. 2004. Phylogeny of Braya and Neotorularia
based on ITS sequences of nuclear ribosomal l)NA and
trn I. intron of chloroplast I)NA. Canad. J. Bot. 82: 376—
392.
Zhou, T. Y., I,. 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.
Shangrilaia (Brassicaceae), a New Genus from China
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ihsan.al-shehbaz@mohot.org
Jipei Yue and Hang Sun
Herbarium, Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan,
Kunming, Yunnan 650204, People’s Republic of China, yuejipei@mail.kib.ac.cn;
hsun@mail.kib.ac.cn
ABSTRACT. The new genus and species Shangri¬
laia nana (Brassicaceae) are described and illus¬
trated, and their relationships to distinguishing
characters from Braya and Baimashania are dis¬
cussed.
Key words: Baimashania, Brassicaceae, Braya,
China, Shangrilaia, Yunnan.
Although a comprehensive account of die Bras¬
sicaceae for the Flora of China has recently been
published (Zhou et al., 2001), many new taxa and
new nomenclatural adjustments have since been
added for China, especially those that are based on
plants from Sichuan, Tibet (Xizang), Yunnan, and
Xinjiang provinces or autonomous regions (Al-
Shehbaz, 2002a, 2002b, 2002c, 2002d, 2003,
2004a, 2004b; Al-Shehbaz & Koch, 2003; Al-
Shehbaz & Warwick, 2004; Al-Shehbaz et ah,
2002a, 2002b). Such additions clearly indicate that
our knowledge of the Chinese mustards, especially
from the provinces and regions above, is far from
being complete, anil further fieldwork is likely to
bring additional novelties. A case in point is the
present paper, which describes a new genus and
species, Shangrilaia nana, from Yunnan Province.
Shangrilaia Al-Shehbaz, J. P. Yue & H. Sun, gen.
nov. TYPE: Shangrilaia nana Al-Shehbaz, J.
P. Yue & H. Sun.
Herba nana, perennis, scaposa, pulvinata. Folia acicu-
lari-linearia, ciliata, integra, persistentia, dense imbricata,
basi complanata, pilis simplicibus instructa. Flores soli-
tarii. Sepala ovata, nonsaccata. Petala alba, spathulata.
Glandulae nectariferae confluentes. Ovula 6—12. Fructus
ovoidei, teretes, breviter stipitati, valvis pubescentibus
coriaceis; stylus tenuis, ad 1 mm longus; stigma integrum.
Semina uniseriata, ovoidea; cotyledones incumbentes.
Dwarf, perennial, scapose, pulvinate herbs, with
simple or few-branched caudex covered with leaves
of previous years. Trichomes simple. Stems densely
leafy, internodes obsolete. Leaves acicular-linear,
thick, densely overlapping, ciliate, persistent, with
flattened, triangular base. Flowers solitary, termi¬
nating stem. Sepals ovate, deciduous, erect, not
saccate at base. Petals white, spatulate, obtuse,
clawed. Stamens 6, tetradynamous; filaments slen¬
der at base; anthers ovate, not apiculate at apex.
Nectar glands confluent, subtending bases of all
stamens; median glands present. Ovules 6 to 12 per
ovary. Fruit dehiscent silicle, ovoid, terete, short
stipitate; valves leathery, with a distinct midvein at
least basally, pubescent; replum flattened at least
basally; septum perforate, veinless; style slender, to
1 mm long; stigma capitate, entire. Seeds uniser-
iate, wingless, ovoid, not mucilaginous when wet¬
ted; cotyledons incumbent.
Initial attempts to identify the plants on which
Shangrilaia is based, using Zhou et al. (2001), Ap¬
pel and Al-Shehbaz (2003), or the interactive keys
((http://flora.huh. harvard. edu:8080/actkey/index.jsp))
to the Brassicaceae genera of the world or that of
the flora of China, yielded unsatisfactory results.
Those keys inadequately led to either Baimashania
Al-Shehbaz or Braya Sternberg & Hoppe, and the
characters of the new genus did not match those of
these two genera. It became abundantly clear that
a new genus is needed to accommodate them.
Shangrilaia, which is named after the city Shangri-
La (formerly Zhong Dian), differs from both Bai¬
mashania and Braya in having exclusively simple
instead of a mixture of simple and branched tri¬
chomes, perforate instead of complete septa, and
acicular-linear instead of distinctly flattened leaves.
It al so differs from Baimashania in having incum¬
bent instead of aceumbent cotyledons, ovoid and
terete instead of linear and latiseptate (flattened
parallel to the septum) fruits, thick leathery instead
of papery valves, veinless instead of veined septum,
and flattened instead of rounded replum. From
Novon 14: 271-274. 2004.
272
Novon
l*'igure I. Shangrilaia nana Al-Shehbaz, J. P. Yue & H. Sun. — A. Plant. — B. Leaf. — C. Portion of leaf showing
abaxial surface. — I). Sepal. — -E. Petal. — F. Fruit and pedicel with valve removed to show seed and perforated septum.
— G. Fruit valve. — H. Section of seed showing incumbent cotyledons. Scale: A = 1 cm; B, I)— G = I mm; C = 0.2
mm. II = 0.5 mm. Drawn by Al-Shehbaz from the holotype (Yue 0366).
Volume 14, Number 3
2004
Al-Shehbaz et al.
Shangrilaia from China
273
Braya, it is also readily distinguished by having
solitary flowers at the end of the stem instead of
flowers in racemes or corymbs on a well-developed
peduncle, as well as in having confluent instead of
four separate nectar glands.
One species assigned to Braya by Zhou et al.
(2001), B. forrestii W. W. Smith, was described as
having exclusively simple trichomes. However, mo¬
lecular studies by Warwick et al. (2004) clearly
demonstrated that this species is quite distinct from
the rest of Braya and perhaps should be excluded
from that genus.
Shangrilaia nana Al-Shehbaz, .). P. Vue & H. Sun,
sp. nov. TYPE: China. Yunnan: Shi-ka Shan,
near Shangri-La, scree slope, sandy area,
27°47'N, 99°35'E, 4200 m, 5 Aug. 2003, Ji-
pei Yue 0366 (holotype, MO; isotype, KUN).
Figure 1.
Herba pulvinata 1—2.5 cm alta, caudice simplici vel
pauciramoso. Pili 0.2-0. 5 mm longi. Folia aciculari-li-
nearia, 2—6 X 0.2— 0.4 mm, dense imbricata, sparse cil-
iata, basi triangulari, 1-1.5 X 1-1.5 mm. Flores solitarii.
Pedicelli fructiferi 0.5—2 mm longi, crassi. Sepala ovata,
glabra, 2.5—3 X 1.5—2 mm. Petala alba, spathulata, 4—5
X 1.5— 2.5 mm, unguibus 2—3 mm longis. Ovula 6—12.
Fructus ovoidei, teretes, breviter stipitati, valvis 3—3.5 X
2—2.5 mm, pubescentibus coriaeeis, septis ad maturitatem
perforatis; stylus 0.4—1 mm longus. Semina ovoidea, 1 —
1 .2 X 0. 6-0.8 mm.
Pulvinate herbs 1-2.5 cm tail, caudex simple or
few-branched. Trichomes simple, 0.2— 0.5 mm long.
Leaves acicular-l inear, 2—6 X 0.2— 0.4 mm, densely
overlapping, with a prominent midvein abaxially;
base flattened, subtriangular, 1—1.5 X 1-1.5 mm,
margin entire, sparsely ciliate to subsetose at least
basally or up to the middle, apex obtuse, often with
1 to 3 trichomes. Flowers solitary, in leaf axils, 1
to 4 terminating the stem. Fruiting pedicels stout,
somewhat angled, 0.5—2 mm long, glabrous or
sparsely pubescent with straight, horizontal tri¬
chomes, distinctly expanded at receptacle. Sepals
ovate, glabrous, 2.5—3 X 1 .5—2 mm, with prominent
veins. Petals white, spatulate, 4—5 X 1.5— 2.5 mm,
obtuse; claw 2—3 mm long, slender at base. F ila¬
ments of median stamens 2—2.5 mm long, those of
lateral stamens 1 — 1.2 mm long; anthers ovate, 0.5—
0.6 mm long. Ovules 6 to 12 per ovary. Fruit ovoid,
terete; valves leathery, 3—3.5 X 2—2.5 mm, with Hat
margin and prominent midvein at least basally, pu-
berulent with simple trichomes; septum initially
complete, becoming perforate at seed's maturity;
gynophore stout, slightly wider than pedicel, 0.2—
0.7 mm long; style slender. 0.4—1 mm long. Seeds
ovoid, 1-1.2 X 0.6— 0.8 mm. Fruiting August— Sep-
tember.
Habitat. Scree slopes on sandy soil; 4180—
4200 m. Yunnan.
Both the type collection and the paratype have
ample fruits, and the floral description is based on
two flowers found on two plants. Collection of flow¬
ering material should be done in late June or early
July. There is some variation in the pubescence of
leaves, and most plants examined have trichomes
at the leaf base and usually one or a few at the leaf
apex. Although there are up to six ovules in each
locule, usually one or two develop into mature
seeds. The septum is complete during early fruit
development, and fruits with mature seeds always
have perforate septa.
Paratypes. CHINA. Yunnan: Shika Shan, Zhong
Dian, 27°47'N, 99°35'E, 4180 m, 27 Sep. 2001, J. P Yue
054 (KUN, MO).
Acknowledgments. We are most grateful to
Henk van der Werff for correcting the Latin and
Victoria C. Hollowell for the editorial advice. We
thank Fernand Jacquemoud and an anonymous re¬
viewer for their valuable comments. We are much
indebted to the Yunnan Natural Science Founda¬
tion (grants-in-aid 2002C0060M), the National Nat¬
ural Science Foundation of China (grants 30270121
and 40332021), and the Innovation Program of the
Chinese Academy of Science (grant KSCX2-1-09),
all of which supported fieldwork and research of
one of us (J. P. Yue).
Literature Cited
Al-Shehbaz, 1. A. 2002a. New species of Alyssum, Aphrag-
mus, Arabis and Sinosophiopsis (Brassieaceae) from Chi¬
na and India. Novon 12: 309—313.
- . 2002b. Six new species of Draba (Brassieaceae)
from the Himalayas. Novon 12: 314—318.
- . 2002c. New combinations in Brassieaceae (Cru-
ciferae): Draba serpens is a Hemilophia and D. william-
sii is a Lepidostemon (Brassieaceae). Edinburgh J. Bot.
59: 443-450.
- . 2002d. Noccaea nepalensis, a new species from
Nepal, and four new combinations in Noccaea (Brassi-
caceae). Adansonia 24: 89—91.
- . 2003. Aphragmus bouffordii, a new species from
Tibet and a synopsis of Aphragmus (Brassieaceae). Har¬
vard Pap. Bot. 8: 25—27.
- . 2004a. Novelties and notes on miscellaneous
Asian Brassieaceae. Novon 14: 153—157.
- . 2004b. Two new species of Draba (Brassieaceae):
I). mieheorum from Tibet and D. sagasteguii from Peru.
Novon 14: 249-252.
- & M. Koch. 2003. Drabopsis is united with Draba
(Brassieaceae). Novon 13: 173—174.
- & S. 1. Warwick. 2004. Transfer of Arabidopsis
gamosepala and Torularia brachycarpa to Braya (Bras-
sicaceae). Novon 14: 269—270.
- . K. Mummenhoff & 0. Appel. 2002a. Cardaria ,
Coronopus, and Stroganowia are united with Lepidium
(Brassieaceae). Novon 12: 5—11.
274
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- , B. Bartholomew, A. Abbas & A. Tumur. 2(H)2b.
New or noteworthy species of Brassicaceae for the Flora
of China. Harvard Bap. Bot. 7(1): 23—24.
Appel, 0. & I. A. Al-Shelibaz. 2003. Cruciferae. Bp. 75-
174 in: K. Kubitzki (editor). Families and Genera of
Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg.
Warwick, S. I., I. A. Al-Shehbaz, C. Sauder, J. G. Harris
& M. Koch. 2004. Bhylogeny of Bray a and A 'eotorularia
(Brassicaceae) based on nuclear ribosomal ITS and
chloroplast trn I . intron sequences. Canad. J. Bot. 82:
376-392.
Zhou, T. Y., L. L. Lu, G. Yang & 1. A. Al-Shehbaz. 2001.
Brassicaceae. Bp. 1-193 in: Z. Y. Wu & B. H. Raven
(editors), F lora of China, Vol. 8. Science Cress, Beijing,
and Missouri Botanical Garden Cress, St. Louis.
New Combinations in Silphium (Asteraceae: Heliantheae)
Jennifer A. Clevinger
Department of Biology, James Madison University, Harrisonburg, Virginia 22807, U.S.A.
clevinja@jmu.edu
Abstract. As part of a revision of Silphium (As¬
teraceae: Heliantheae) for the Flora of North Amer¬
ica (FNA), four new combinations are proposed: S.
asteriscus var. simpsonii (Greene) J. A. Clevinger,
S. asteriscus var. trifoliatum (L.) J. A. Clevinger, S.
asteriscus var. latifolium (A. Gray) J. A. Clevinger,
and .S', radula var. gracile (A. Gray) J. A. Clevinger.
In addition, Silphium confertifolium Small is re¬
duced to synonymy of Silphium asteriscus var. la¬
tifolium, and Silphium simpsonii var. wrighl ii L M.
Perry is reduced to synonymy of Silphium radula
Nuttall. Silphium radula var. gracile is lectotypi-
fied.
Key words: Asteraceae, Compositae, Helian¬
theae, North America, Silphium.
As part of a revision of Silphium L. (Asteraceae:
Heliantheae) in Clevinger’s (1999) dissertation,
7000 herbarium specimens from 16 herbaria were
examined and populations in 22 states were visited.
A molecular phylogenetic study of Silphium ex¬
plored relationships in Silphium resulting in the
recognition of two monophyletic sections: sect. Sil¬
phium anil sect. Composita (Clevinger & Panero,
2000). As a result of these studies and subsequent
eolleeting, four nomenclatural changes in Silphium
sect. Silphium are proposed. Three species names
are reduced to varietal rank, and one varietal name
is transferred to a different species. These nomen¬
clatural changes pave the way for the taxonomic
treatment of Silphium that is currently being [ire-
pared for the Flora of North America.
Silphium asteriscus 1 ..
The most current circumscription of Silphium as¬
teriscus can be found in Cronquist (1980). His con¬
cept of Silphium asteriscus includes three varieties:
var. asteriscus, var. angustatum A. Gray, and var.
scabrum Nuttall. Phis taxonomic concept should be
expanded to include S. dentatum Elliott, S. trifol¬
iatum L., and S. confertifolium Small, all of which
Cronquist recognized as species. It should also in¬
clude S. simpsonii Greene, which Cronquist placed
in synonymy of S. gracile A. Gray. These taxa all
share the characteristics of having a variable num¬
ber of ray flowers ranging from 8 to 21 and lance¬
olate to ovate leaves. Cronquist used phyllotaxy and
presence or absence of pubescence on stems,
leaves, and paleae to distinguish between these
species. However, my field observations reveal var¬
iability in these characters within populations
found where the geographical ranges of these taxa
overlap. This variability is symptomatic of intro-
gression between these taxa. Therefore, because of
the introgression, they should be recognized at the
varietal level rather than as separate species.
Silphium dentatum already has varietal status as
S. asteriscus var. dentatum (Elliott) Chapman and
will be recognized as such (Chapman, 1860). Sil¬
phium trifoliatum var. trifoliatum, S. trifoliatum var.
latifolium A. Gray, and S. simpsonii should he
transferred herein to S. asteriscus. Silphium confer¬
tifolium will be placed in synonymy with Silphium
asteriscus var. latifolium.
Silphium asteriscus L., Sp. El. 2: 920. 1753.
TYPE: “Asteriscus coronae solis flore et facie”
in Dillenius, Hurt. Eltham., 42, t. 37, f. 42.
1732 (lectotype, designated by Perry (1937:
292)).
Silphium asteriscus var. simpsonii (Greene) J. A.
Clevinger, comb. nov. Basionym: Silphium
simpsonii Greene, Pittonia 4: 45. 1899. TYPE:
U.S.A. Florida: Palma Sola, in damp ground
in pine barrens, 8 July 1890, J. //. Simpson
81 (holotype, US).
Silphium asteriscus var. simpsonii is most closely
allied to S. asteriscus var. dentatum and S. asteriscus
var. angustatum. They share the characteristic of
stipitate glands on the paleae. Stipitate glands are
absent in other varieties of Silphium asteriscus but
are also found in Silphium laciniatum L., Silphium
perplexum J. Allison, and Silphium glutinosum J.
Allison. Additionally, Silphium asteriscus var. simp¬
sonii, variety dentatum, and variety angustatum,
along with variety asteriscus and variety scabrum,
share the characteristics of predominantly alternate
phyllotaxy and scabrous to hispid leaves. This is in
contrast to the predominantly opposite or whorled
phyllotaxy and glabrous leaves of variety trifoliatum
Novon 14: 275-277. 2004.
276
Novon
and variety latifolium. Silphium asteriscus var.
simpsonii can be distinguished from its closest al¬
lies, variety dentatum anil variety angustatum, by
its height (0.2— 1.1 in vs. 0.5— 1.5 m) and persistent
basal rosette.
Silphium asteriscus var. simpsonii is restricted to
the southeastern U.S. and is primarily found in
Florida. Perry (1937) divided Silphium simpsonii
into two varieties. Silphium simpsonii var. wrightii
Perry, found in the southwestern U.S., should not
be included here. Rather, because of its large num¬
ber of ray flowers (20 to 30+), I reduce Silphium
simpsonii var. wrightii to synonymy of ,S. radula var.
radula. During the summer of 2002, Silphium as¬
teriscus var. simpsonii was collected in Florida and
added to the Clevinger and Panero (2000) molec¬
ular data set for phylogenetic analysis. In the re¬
sulting phylogenetic tree, Silphium asteriscus var.
simpsonii was allied with Silphium asteriscus rather
than Silphium radula.
Silpliimn asteriscus var. trifoliatiiin (L.) J. A.
Clevinger, comb. nov. Basionym: Silphium tri-
foliatum L., Sp. PI. 2: 920. 1753. Silphium
ternifolium Michaux, FI. Bor. Amer. (Mi-
chaux) 2: 146. 1803. TYPE: “Habitat in Vir¬
ginia” (lectotype, designated by Beveal in
Jarvis & Turland (1998: 367), LINN micro¬
fiche # 1032.8, photos, F. GH).
Fieldwork shows that Silphium asteriscus var. tri-
foliatum is most closely allied to Silphium asteris¬
cus var. latifolium. They share the characteristics
of glabrous leaves, peduncles, and stems in contrast
to all other varieties of Silphium asteriscus. Sil¬
phium asteriscus var. trifoliatum is easily distin¬
guished in the northeastern U.S. because of its tri¬
foliate leaf arrangement. However, these same
populations also contain individuals with alternate,
opposite, or 4-foliate leaf arrangements. Along the
southern edge of the varietal range, where it over¬
laps with S. asteriscus var. asteriscus and S. aster¬
iscus var. latifolium. these alternate, opposite, and
4-foliate leaf arrangements become more predomi¬
nant.
Silpliiuiu asteriscus var. lutil'oliuiu (A. (fray) J.
A. Clevinger, comb. nov. Basionym: Silphium
trifoliatum L. var. latifolium A. Gray, Syn. FI.
N. Amer. 1: 241. 1884. TYPE: U.S. A. Ala¬
bama: July 1840, S. B. Buckley s.n. (holotype,
GH).
Silphium confertifolium Small. FI. S.E. U.S. 1243, 1340.
1903. Syn. nov. TYPE: U.S. A. Alabama: Choctoaw
Co., Cocoa, 13—15 Oct. 1896, C. Schuchert s.n. (ho¬
lotype. NY; isotypes, NY. US).
Like the other varieties of Silphium asteriscus,
variety latifolium has ray flowers that number 8 to
21. This variety is distinguished by its predomi¬
nantly opposite phyllotaxy and glabrous leaves and
stems. In my fieldwork, I found that when Silphium
asteriscus var. latifolium populations come in con¬
tact with variety asteriscus there is a mix of glabrous
and pubescent plants. When its populations come
in contact with variety trifoliatum, there is a mix of
opposite and trifoliate leaf arrangements on the
plants. Because of this introgression that I observed
in the held, variety latifolium is placed in S. aster¬
iscus.
I conclude that Silphium confertifolium repre¬
sents an environmental variant of S. trifoliatum var.
latifolium and should not retain its status as a dis¬
tinct species. This is based upon examination of
herbarium specimens and visits to sites in Dallas
Co., Alabama, which is the locality of historic S.
confertifolium collections {Godfrey 6 9684, Jones
752, NCU). Cronquist (1980) distinguished these
two species on the basis of height and leaf distri¬
bution. Silphium confertifolium ranges in height
from 0.4 to I m with basally disposed leaves, while
Silphium trifoliatum var. latifolium ranges in height
from 1 to 2 m with evenly distributed leaves. In the
field I observed a continuum of plants within pop¬
ulations that varied in stature and leaf distribution
according to proximity to chalk outcroppings. The
plants closest to the chalk outcroppings were al¬
ways the shortest in height with reduced leaves,
and the ones furthest were tallest in height with
evenly distributed leaves.
The recognition of Silphium asteriscus var. trifol¬
iatum and variety latifolium is in agreement with
the dissertation work of W. B. Weber (1968). Weber
conducted extensive hybridization studies and con¬
cluded that S. asteriscus and S. trifoliatum are not
reproductively isolated. Weber also concluded that
phyllotaxy was variable between these two species.
Weber and his adviser, T. B. Fisher, proposed in
Weber’s dissertation that S. trifoliatum var. trifol¬
iatum and S. trifoliatum var. latifolium should be
reduced to varietal rank within S. asteriscus. How¬
ever, these changes were never formally published.
Silphium radula Nuttai.l
Silpliiuiu radula Nuttall, Trans. Amer. Philos.
Soc., n.s. 7: 354. 1841. TYPE: U.S. A. Plains
of Arkansas, T. Nuttall s.n. (holotype, BM).
Silphium simpsonii Greene var. wrightii L. M. Perry, Hho-
dora 39: 288. 1037. Syn. nov. TYPE: U.S. A. Texas:
1848. C. Wright s.n. (holotype, (41).
Silphium simpsonii var. wrightii is reduced here-
Volume 14, Number 3
2004
Clevinger
New Combinations in Silphium
277
in to synonymy of Silphium radula based on the
examination of the type specimen at GH. Gandhi
and Thomas (1989) recognized both S. radula and
S. simpsonii var. wrightii and stated that they differ
only in the number of capitula and leaf texture. I
have observed that these differences are quite var¬
iable, and these two entities should not be sepa¬
rated.
Silpliiiiin radula var. graeile (A. Gray) J. A. Clev¬
inger, comb, et slat. nov. Basionym: Silphium
graeile A. Gray, Proc. Amer. Acad. Arts 8:
653. 1873. TYPE: U.S.A. Texas: Harris Co.,
Houston, 1842, F. ./. Lindheimer s.n. (lecto-
type, selected here, GH; isotype, (ill).
Silphium radula var. graeile is recognized by a
persistent basal rosette and ray flowers numbering
12 to 18. In contrast, Silphium radula var. radula
has a caducous basal rosette and ray flowers num¬
bering 20 to 30. Perry (1937) and others have sug¬
gested that the southwestern U.S. Silphium radula
var. graeile is allied to the southeastern U.S. Sil¬
phium asteriscus var. simpsonii. These two varieties
resemble one another in their persistent basal ro¬
settes and reduced stature. However, Silphium rad¬
ula var. graeile and variety radula have pubescent
phyllaries and are found in Arkansas, Louisiana,
Oklahoma, and Texas, while Silphium asteriscus
var. simpsonii has glabrous phyllaries and is found
in Alabama, Florida, Georgia, Mississippi, and
South Carolina. Silphium radula var. graeile (as Sil¬
phium graeile) was included in the molecular study
of Clevinger and Panero (2000). On the basis of
ITS and ETS sequence data, this variety was dem¬
onstrated to be sister to S. radula var. radula.
Asa Gray cited two collections, those of Hall and
of Lindheimer, in his description of Silphium grae¬
ile. He did not designate either as the holotype;
therefore, they are syntypes. According to Gray, the
Lindheimer specimens have larger leaves and more
pubescence than the Hall specimens. Both descrip¬
tions fit Gray’s concept of S. graeile and my concept
of S. radula var. graeile. I have seen the two Lind¬
heimer collections at GH. The status of the Hall
collections is uncertain; I have seen two 1872 col¬
lections at F and NY labeled //«// 321 (Gray does
not cite a number in his description) but none at
GH. Given this uncertainty, I have chosen the Lind¬
heimer collection at GH to serve as the lectotype.
Acknowledgments. I am grateful to Jose L. Pa¬
nero, Beryl Simpson, Billie Turner, Doug Goldman,
Curtis Clevinger, James Allison, Conley K. Mc¬
Mullen, Lowell and Mary Amiek, and the curators
of the following herbaria for the loan of specimens:
F, FLAS, GH, IA, LL, LSI), MICH, MO, MSC,
NCU, NY, OS, TENN, TEX, US, and USCH. This
paper represents a portion of a doctoral dissertation
by the author submitted to the Department of Bot¬
any at the University of Texas at Austin. Support
for some aspects of this study was provided by
grants from the American Society of Plant Taxon¬
omists and the Virginia Academy of Science.
Literature Cited
Chapman, A. W. I860. Flora of the Southern United
States: Containing an Abridged Description of the Flow¬
ering Plants and Ferns of Tennessee, North and South
Carolina, Georgia, Alabama, Mississippi and Florida:
Arranged According to the Natural System. Ivison,
Phinney, New York.
Clevinger, J. A. 1999. Systematies of Silphium and Its
Subtribe Engelmaniinae (Asteraceae: Heliantheae).
Ph.D. Dissertation, The University of Texas at Austin.
- & J. L. Panero. 2000. Phylogenetic analysis of
Silphium and subtribe Engelmanniinae (Asteraceae:
Heliantheae) based in ITS and ETS sequence data.
Amer. J. Bot. 87: 565-572.
Cronquist, A. 1980. Vascular Flora of the Southeastern
United States, Vol. I: Asteraceae. Univ. North Carolina
Press, Chapel Mill.
Gandhi, K. N. & R. I). Thomas. 1989. Asteraceae of Lou¬
isiana. Sida Bot. Misc. 4.
Jarvis, C. E. & N. Turland. 1998. Typification ol Linnaean
specific and varietal names in the Compositae (Astera¬
ceae). Taxon 47: 347—370.
Perry, L. M. 1937. Notes on Silphium. Rhodora 39: 281 —
297.
Weber, W. R. 1968. Biosystematic Studies in the Genus
Silphium L. (Compositae): Investigations in the Selected
Intraspecific Taxa of Silphium asteriscus L. Ph.D. Dis¬
sertation, The Ohio State University.
Phylloscirpus (Cyperaceae) Revisited
Sandra Dhooge and Paid Goetghebeur
Ghent University, Department of Biology, Research Group Spermatophytes, K.I
Ledeganckstraat 35, B-9000 Gent, Belgium. Sandra.Dhooge@UGent.be;
Paul.Goetghebeur@UGent.be
Abstract. Two species of Scirpus are transferred
to Phylloscirpus: Scirpus deserticola Philippi and
Scirpus boliviensis Barros. Phylloscirpus deserticola
has all the typical characteristics of the genus Phyl¬
loscirpus, such as eligulate leaves, a terminal head
of spikelets. and scabrid hypogynous bristles. The
allocation of Scirpus boliviensis to Phylloscirpus is
morphologically less obvious due to its reduced
structures, but is well supported by a phylogenetic
analysis based on plastid rbcl . anil OtjL-F sequence
data. Phylloscirpus deserticola is lectotypified be¬
cause the holotype consists of a mixed collection.
Key words: Andes, Cyperaceae, Phylloscirpus.
Phylloscirpus C. B. Clarke (1908) is a genus with
a single species, Phylloscirpus acaulis (Philippi)
Goetghebeur & I). A. Simpson (1990), distributed
in Andean regions of South America. Clarke (1908)
described the genus based on the presence of a
dense head of spikelets subtended by bracts, each
spikelet having spirally arranged glumes and bi¬
sexual (lowers comprising five or six scabrid, hy¬
pogynous bristles.
During our ongoing taxonomic study of Scirpus
s.l. in the Andes, it became clear that one of the
species, Scirpus deserticola Philippi, clearly has the
diagnostic features of Phylloscirpus , justifying a
new combination within the genus. Another and
very small species, Scirpus boliviensis Barros, also
has characters in common with the species of Phyl¬
loscirpus, such as eligulate leaves and numerous
red tannin idioblasts in the glumes. Although
Clarke did not mention these characters in his orig¬
inal description, our observations and a molecular
analysis discussed below strongly indicate that
Scirpus boliviensis Barros should be included in the
genus Phylloscirpus. However, as Scirpus boliviensis
Barros has one terminal spikelet (opposed to a head
of spikelets) and no bristle hairs, it is necessary to
redefine Phylloscirpus, to give the genus a broader
circumscription.
This proposal to make two new combinations in
Phylloscirpus is well supported by our phylogenetic
analysis based on rbc L and traL-F sequencing data
of several Andean Scirpus species.
Material and Methods
MORPHOLOGICAL DATA
Morphological data were obtained from the study
of herbarium material with a stereomicroscope.
Herbarium material from the following herbaria was
studied: AAU, B, BM, C, GENT. GH, GOET, ISC,
K, NY, P, OCA. SGO, SI, U. and US.
MOLECULAR DATA
The plant material used for DNA extraction and
its provenance are listed in Table 1. Total DNA was
isolated using the modified CTAB method of Doyle
and Doyle (1987). Amplification was done in over¬
lapping pieces using the IF, 1024F. 724R. 1024R,
and I460R primers for the rbc\ . gene (Muasya et
ah, 1998) and the c, d, e, and f primers for /raL-E
(Taberlet et ah. 1991). The amplified products were
cleaned using the QIAquiek kit according to the
manufacturer’s protocols (Qiagen, Ltd.), and se¬
quenced using standard dideoxy methods and run
on an ABI 377 automated sequencer (all according
to the manufacturer's protocols. Applied Biosys¬
tems, Inc.). Phe sequences were edited in Sequence
Navigator, assembled in Autoassembler (Applied
Biosystems, Inc.), and aligned by eye in a sequence
matrix of Scirpus s.l. (Muasya et ah, 2000).
PH'1 LOGENETIC AN At A SIS
The molecular data set was analyzed using par¬
simony algorithms of PAUP* 4.0b8 (Swofford,
2002). A heuristic search of 1000 replicates with
the TBR (tree-bisection-reconnection) swapping al¬
gorithm and random addition was conducted, re¬
taining five trees at each step. Internal support was
estimated with 1000 bootstrap replicates. For each
bootstrap replicate, 100 heuristic searches with
random sequence addition were conducted.
Taxonomy
Phylloscirpus deserticola (Philippi) Dhooge &
Goetghebeur, comb. nov. Basionym: Scirpus de¬
serticola Philippi, FI. Atacam.: 53. 1860. TYPE:
Chile. Agua de profetas in deserto Atacamensi,
Jan. 1854, R. A. Philippi s.n. (lectotype, select¬
ed here. SGO 046286 A). Figure 1A, I).
Nov ON 14: 278-284. 2004.
Volume 14, Number 3
2004
Dhooge & Goetghebeur
Phylloscirpus (Cyperaceae)
279
Scirpus semisubterraneus Boeckeler, Eirinaea 36: 495.
1870. Scirpus deserticola Philippi var. semisubterra¬
neus (Boeckeler) Barros, Anal. Mus. Argent. Cienc.
Nat. 38: 156. 1935. TYPE: Peru. W. Lechler n° 1977
(holotype, not seen; isotypes, k, P).
Caespitose, small herbs, 1.8— 12 cm tall; rhi¬
zomes creeping, branching; culms 0.6-10 cm X
0.5— 1.3 mm, terete, glabrous, sometimes with little
red spots (tannin idioblasts), covered with old
brown leaf sheaths. Leaves shorter than to as long
as the culm; leaf sheaths 2.5— 6.5 mm long, often
with red spots; leaf blade 0.3— 1.9 cm X 0.4—0.85
mm, scabrous toward the leaf tip; leaf tip muticous,
often orange; ligule absent. Inflorescence a single
spikelet or a terminal, dense head composed of 2
to 6 spikelets (Fig. I A), 4—7.2 X 1.5—8 mm, red-
brown to dark brown; when 2 or more spikelets are
present, each lateral spikelet subtended by a bract
and a small, membranous prophyll; bracts 3.4— 6.5
X 2-4 mm, red-brown with many tannin idioblasts
and a conspicuous, swollen, pale midrib, apiculate,
as long as to slightly shorter than the inflorescence.
All glumes fertile, 2.7-4.7 X 1.5-1. 9 mm, ovate,
red-brown with many red spots (tannin idioblasts)
and a conspicuous, swollen, pale midrib; flowers
bisexual; stamens 3; ovary 3-carpellate; style
Figure 1. A and D. Phylloscirpus deserticola (Philippi) Dhooge & Goetghebeur ( Philippi s.n., lectotype, SGO). - — A.
Inflorescence. — D. Fruit with bristle hairs. — B. Inflorescence of Phylloscirpus acaulis (Philippi) Goetghebeur & D. A.
Simpson ( Ruthsatz 8765, GENT). C and E. Phylloscirpus boliviensis (Barros) Dhooge & Goetghebeur ( Petersen & Hjerting
1048b, holotype, G). — C. Inflorescence. — E. Eruil.
280
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Volume 14, Number 3
2004
Dhooge & Goetghebeur
Phylloscirpus (Cyperaceae)
281
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°y John Manning. Scale bar 10 mm for entire plant, capsules, floral bracts, and side view of flower; detail of the
perianth base, entire gynoecium and stamens of dissected flower, 3X scale bar, seed approximately 6X scale bar.
Volume 14, Number 3
2004
Goldblatt & Manning
African Iridaceae
291
breve quam exteriore, floribus hypocrateriformibus acti-
nomorphis pallide ad intense carneis tepalis purpureis ad
basem nigris leviter dulce odoris, tubo perianthii pauce
curvato cylindrico infime 2—3 mm abrupte expanse in eu-
pulam ca. 3 mm profundam, tepalis obovatis patentibus
subaequalibus (14— )18— 22 X (f> — ) 10 — 12 mm, filamentis
conniventis 5—6 mm longis ereetis nigris, antheris diver-
gentibus 6-7 mm longis nigris, polline flavo, ovario ovo-
ideo ca. 2 mm longo, stylo erecto furcato ad basem an-
therarum ramis 2—3 mm longis nigris, capsulis ovoideis
9—11 X 5 mm, seminibus subglobosis vel angulalis 1.2—
1.5 mm diam. leviter rugosis.
Plants 20—60 cm high; conns globose, 15—20
mm diam., the tunics of medium to coarsely tex¬
tured fibers; cataphylls membranous, the upper one
reaching just above ground level and flushed red¬
dish brown. leaves 3, all basal, the lower two nar¬
rowly to broadly lanceolate, (3— )5— 10 mm wide,
usually about half as long as the stem, the margins
and midribs moderately thickened, the uppermost
leal entirely sheathing, reaching at least to the mid¬
dle or upper third of the stem. Stem erect, simple
or 1- to 3-branched, 0.5—1 mm diam. at the base
of the spike, branches held 45°— 90° to the main
axis; spike almost straight, the main spike usually
3- to 6-flowered, the lateral spikes I - to 4-flowered;
bracts membranous and translucent, the outer 8—10
mm long, with three light purple veins, lightly
three-dentate, the inner as long as to slightly longer
than the outer, forked at the apex, with two purple
veins. Flowers salver-shaped, actinomorphie, pale
to deep pink, the tepals each purple to blackish
toward the base, lightly sweet scented; perianth
tube slightly curved and cylindric for 2-3 mm, wid¬
ening abruptly into a cup ca. 3 mm deep; tepals
obovate, spreading at nearly right angles to the
tube, subequal, (14— ) 18— 22 X (6—) 10— 12 mm, the
inner slightly wider than the outer; filaments in¬
serted at the mouth of the narrow part of the tube,
blocking the mouth, connivent, 5—6 mm long,
straight and erect, black; anthers divergent, 6—7
mm long (before anthesis), black, the pollen yellow;
ovary ovoid, ca. 2 mm long; style straight and erect,
dividing opposite the base of the anthers, the
branches arching outward, 2—3 mm long, black.
Capsules ovoid, 9—1 1 X 5 mm; seeds subglobose or
angled by pressure, 1 .2-1.5 mm diam., yellow-
brown, lightly wrinkled with surface cells collicu-
late.
Phenology. Flowering early to mid September,
probably also in late August.
Etymology. From the Latin superbus , “superb,
excellent,” referring to the large, attractive, deep
pink and purple-black flowers.
Distribution and biology. Ixia superba has a re¬
corded range of just a few acres at the foot of the
Langeberg in the Little Karoo east of Montagu.
Plants grow on loamy sandstone ground that grades
to a light clay at the interface of Table Mountain
Sandstone and Bokkeveld Systems on the farm Kli-
pheuwel. We collected plants in bloom there in ear¬
ly September 2002 close to the end of their flow¬
ering season. Associated plants were Babiana
patula N. E. Brown, Moraea gawleri Sprengel, Wat¬
son ia laccata (Jacquin) Ker-Gawler (Iridaceae), and
the shrub Elytropappus rhinocerotis (L. f.) Lessing.
The flowers evidently have a bimodal pollination
strategy that uses both hopbine scarab beetles
(Scarabaeidae: Hopliini) and worker honey bees.
Apis mellifiera (and possibly female bees of other
genera). The flowers have the typical attributes of
the hopbine beetle pollination system, viz. a bright¬
ly colored perianth with dark markings, darkly pig¬
mented stamens, relatively long anthers, and short
style branches. The closed perianth tube and ab¬
sence of nectar are also features associated with
this pollination system in Ixia (Goldblatt et ah,
2000a). Presence of a sweet floral odor is, however,
an attribute of bee pollination and bees do visit the
flowers, which evidently last three days, but close
at night. Both hopbine beetles and honey bees were
found visiting the flowers, the latter actively col¬
lecting pollen. Early in the morning hopbine bee¬
tles were found in half-closed flowers in which they
had evidently spent the night. In one other collec¬
tion of /. superba we have found ( Steiner 3239,
NBG), field notes indicate that the flowers were vis¬
ited by the hopbine beetle Peritrichia sp.
Relationships. Ixia superba is readily confused
with a second Little Karoo species, Ixia gloriosa G.
J. Lewis (1962). Allhough both have deep pink flow¬
ers with a large blackish glossy central eye and
blackish filaments and anthers, there are several im¬
portant floral differences between them. The darker
red-purple flowers of /. gloriosa have relatively short,
slightly spreading filaments 3—5 mm long, and an¬
thers, said by de Vos (1999b) to be (>-1 mm long
and connivent at the tips. Plants studied by us alive
( Goldblatt , Manning & Porter 121 79, MO, NBG),
however, had anthers ca. 4.5 mm long diverging
throughout, thus not apically connivent. The style
normally divides at the mouth of the tube, thus op¬
posite the base of the filaments, and the style
branches are ca. 4 mm long. The perianth tube is
4—7 mm long and widens abruptly into a small cup
little more than 1 mm deep. Fresh flowers of I. glo¬
riosa have a characteristic umbonate appearance, the
result of the tepals being raised above the rim of the
perianth cup before spreading. This is not apparent
in pressed specimens or closed flowers. In contrast,
/. superba has the perianth tube slender below for 3
292
Novon
mm, widening abruptly into a cup ea. 3 mm deep
and the tepals spread uniformly from the rim of the
tube. The filaments are 5—6 mm long, and the di¬
vergent anthers are ea. 6 mm long. The style is en¬
closed above by the closely connivent filaments and
divides at or shortly above the top of the filaments
into shorter style branches 2—3 mm long. In addition,
/. gloriosa typically has linear leaves 15—55 X 1—3
mm and a long slender spike of 8 to 10 flowers on
the main axis, whereas /. superba has shorter, lan¬
ceolate leaves 10—30 X (3— )5— 10 mm and a shorter,
crowded spike of 3 to 6 flowers. While the bracts of
the two species are similar, those of /. gloriosa have
more pronounced veins t lint when dry remain dark
purple, while the veins in the bracts of /. superba
fade and almost disappear.
There are several other locally endemic species
of Ixia in the Breede River valley that resemble
one another closely in their deep pink flowers with
a dark center and blackish filaments and anthers
(Lewis, 1962). This flower type is adapted for pol¬
lination by hopbine beetles, a pollination system
that seems to favor local differentiation among plant
species, e.g., Aristea subg. Pseudaristea (Goldblatt
& Manning, 1997) and Sparaxis (Goldblatt et al.,
2000b). The species of Ixia in the Breede River
valley with flowers adapted for this pollination sys¬
tem form a geographic al series, replacing one an¬
other across relatively short distances. Moving
down the river valley from its head in the northwest,
the series comprises /. vinacea G. J. Lewis (Tul-
bagh), /. rouxii G. J. Lewis (Wolseley), /. mostertii
M. P. de Vos (Romans River to Worcester), /. van-
zyliae L. Bolus (Worcester to Roberston), /. atran-
dra Goldblatt & J. C. Manning (south of Worcester
to Stettyn), /. superba (Montagu), and /. gloriosa
(Barrydale). Among this alliance, /. superba is read¬
ily distinguished by the perianth tube that forms a
wide cup ca. 3 mm long in die upper half and the
style that divides at or slightly above the base of
the filaments, thus close to the base of the anthers,
rather than at the* mouth of the perianth tube and
we'll below the base of the anthers.
Paratypes. SOUTH AFRICA. Western Cape: 3320
(Montagu), Farm Rietvlei No. 1, ca. 10.5 km SF of Mon¬
tagu, scattered on S-facing rocky slope (CC), ca. 370 m,
5 Sep. 1997, Steiner 32.39 (NBC); Scheepers Rust, 20 mi.
K of Montagu, 22 Aug. 1936, Martley s.n. (BOL 26930).
Tritoniu delpierrei M. P. de Vos, J. S. African But.
49; 403. 1983. Tritoniu marlothii subsp. del¬
pierrei (M. P. de Vos) M. P. de Vos, FI. S. Africa
7(2,1): 121. 1999. TYPE: South Africa. North¬
ern Cape: Richtersveld, top of Helskloof, M.
P. de Vos 2462 (type, STL).
We do not accept de Vos’s (1999a) reduction of
the South African Tritonia delpierrei to subspecific
rank in T. marlothii M. P de Vos. We have seen
and examined both plants in the wild, and while
they are evidently sister laxa, isolated geographi¬
cally and phylogenetieally within Tritonia (de Vos,
1983), there is little overlap in their critical taxo¬
nomic features. We prefer to maintain them as sep¬
arate species differing in floral coloration and di¬
mensions that apparently indicate different primary
pollinators. Tritonia delpierrei has lightly sweet-
scented, pale yellow flowers, the lower tepals with
cream markings, a perianth tube 12—22 mm long,
and the dorsal tepal Fi — 1 2 mm long. In contrast, T.
marlothii has odorless flowers, a pale to deep pur¬
ple perianth (not lilac as indicated by de Vos) with
the lower tepals yellow with purple tips and a me¬
dian dark violet spot, and new collections (e.g.,
Goldblatt & Porter 11757, MO, NBG) show the
perianth tube is 28— 42 mm long and the dorsal
tepal (8-) 10- 1 2 mm long. The flowers of T. mar¬
lothii show the stereotyped adaptations for polli¬
nation by the long-proboscid fly Prosoeca peringue-
yi (Manning & Goldblatt, 1996), and we have
captured this fly while visiting and evidently pol¬
linating the flowers (unpublished obs.). The shorter
perianth tube of T. delpierrei suggests pollination
by a long-tongued anthophorine bee. The ranges of
the two taxa largely overlap. Tritonia marlothii oc¬
curs in the southern Richtersveld of Namaqualand
at Brakfontein, southwest of Eksteenfontein, and
along the eastern foothills of the Stinkfontein
Mountains north of Eksteenfontein, whereas T. del¬
pierrei extends from near Eksteenfontein in the
south to Helskloof in the north, a distance of some
60 km that includes a large part of the range of T.
marlothii. While T. delpierrei seems to favor sandy
slopes, we have only seen T. marlothii in rocky
ground, an apparent habitat difference also consis¬
tent with separate species status.
Thus despite their evident immediate relation¬
ship, we see no reason to consider Tritonia delpier¬
rei as a subspecies of 7. marlothii. Their geographic
ranges partly overlap while their habitats differ, and
their critical morphological features show no evi¬
dence of convergence.
ImnoiDKAK
Ferraria kamiesbergensis M. P. de Vos, J. S. Af¬
rican Bot. 45: 362. 1979. TYPE: South Africa.
Northern Cape: Rondefontein, I Sep. 1976, E.
G. II. Oliver 5970 (lectotype, here designated,
lower left plant on sheet, STE).
Described by M. P. de Vos in 1979, Ferraria ka-
Volume 14, Number 3
2004
Goldblatt & Manning
African Iridaceae
293
miesbergensis is represented by four elements: t he
illustration accompanying the protologue (de Vos,
1979: 363), and three plants mounted on a single
sheet, Oliver 5970. These were evidently all col¬
lected on the same date at Rondefontein in the Ka-
miesberg of central Namaqualand. We have visited
the type locality (or as close to it as is possible)
and found a small, uniform population of plants
with small pale yellow flowers, stems 12—15 cm
high, branching if at all. only from near the base,
with distinctive dark green leaves, and subequal
inflorescence spathes ( Goldblatt cfe Porter 12223,
MO. NBG). Most significantly, the lower leaves all
curve inward in the same direction.
Only one specimen on the type sheet matches
these plants, and it also has the paired inflores¬
cence spathes almost equal in length with the inner
sheathing only below. The other two plants are tall¬
er (one is ca. 30 cm tall), have narrower, evidently
flat leaves, produce branches well above ground
level, and the outer inflorescence spathes are about
half as long as the inner and are largely sheathing.
The plant illustrated in the protologue corresponds
to the larger of these two taller specimens, and both
appear to be a second species, possibly what is
currently called Ferraria divaricata Sweet, either
its subspecies aiirea M. R de Vos or subspecies
arenosa M. R. de Vos. The flower illustrated is prob¬
ably F. kamiesmontana (and is certainly not F. di¬
varicate z), while the smaller plant on the lower left
of the type sheet alone matches plants in the wild
and is the lectotype.
The plants found at Rondefontein are matched
at other sites in the southern Kamiesberg on the
farm Karas, to the north, immediately adjacent to
Rondefontein. Elsewhere on the Kamiesberg, the
only other pale yellow-flowered species of the genus
is the florally nearly identical Ferraria macrochla-
rnys (treated by de Vos as F. uncinata subsp. ma-
crochlamys, see below), which has quite different
leaves.
The range of Ferraria kamiesbergensis according
to de Vos is from Springbok (Thompson 1301 , NBG)
in the north to Wallekraal in the southwest ( Comp¬
ton 5405 , BOL), and extending east as far as the
Calvinia district (e.g., de Eos 2347, NBG). We have
examined all the specimens on which de Vos based
her assessment of the range of the species and con¬
sider them to correspond with what she called F.
divaricata. None of tin* specimens she cited have
the distinctive incurved leaves and subequal
spathes of the southern Kamiesberg populations
(flower color can no longer be determined). As cir¬
cumscribed here, F. kamiesbergensis is restricted to
the southern Kamiesberg at elevations above 1000
m where plants matching the lectotype occur.
Ferraria macroehlamys (Baker) Goldblatt & J.
C. Manning, comb. nov. Basionym: Lapeirousia
macroehlamys Baker, J. Bot. new ser. 5: 338.
1876. Ferraria uncinata subsp. macroehlamys
(Baker) M. P. de Vos. J. S. African Bot. 45:
369. 1979. TYPE: South Africa. Namaqua¬
land, without precise locality or date. Herb.
Forsyth s.n. [probably collected by J. Niven in
1899] (type. K).
In her revision of Ferraria (Iridoideae) de Vos
(1979) treated a series of central and northern Na¬
maqualand populations (Northern Gape, South Af¬
rica) with pale yellow flowers as F. uncinata subsp.
macroehlamys, while she regarded Western Cape
populations with deep blue to violet flowers as F.
uncinata Sweet subsp. uncinata. De Vos believed
that subspecies macroehlamys merited subspecific
status because it differed only in having flowers
with a differently colored perianth. The two taxa
share with F. brevifolia G. J. Lewis and F. kamies¬
bergensis M. P. de Vos a distinctive flower type in
the genus in which the tepal claws are suberect and
collectively form a narrow floral cup that contains
relatively dilute nectar. Both these latter species
also have predominantly pale yellow tepals virtually
the same shade and size as those of subspecies
macroehlamys. De Vos regarded F. macroehlamys
and F. uncinata as sharing similar leaf blades with
the margins thickened and the edges crisped and
often undulate. In her key (1979: 329) to the spe¬
cies, F. uncinata (including subsp. macroehlamys)
is distinguished by “margins of at least some of the
foliage leaves crisped.” De Vos commented of sub¬
species macroehlamys and subspecies uncinata
that they “differ only in flower coloring and geo¬
graphic distribution.” Closer examination of spec¬
imens she included in subspecies macroehlamys,
however, shows that the thickened, hyaline part of
the leaf margins is crenate to lightly serrate and
sometimes shortly velvety, but the leaf edges them¬
selves are only occasionally lightly crisped. In Fer¬
raria uncinata the thickened part of the leaf margin
may be entire, or sometimes lightly serrated or cris-
pulate, and not velvety but rather evidently smooth
(but under 10X magnification papillate), while the
edges of at least some of the leaves are crisped and
usually undulate. Thus, examination of the leaf
margins alone can usually distinguish the two taxa.
In addition, the flowers of F. uncinata have outer
tepals (18— )30— 35 mm long, with claws 9—1 1 mm
long, thus usually slightly less than one-third the
294
Novon
length of the entire tepal. In F. uncinata subsp.
macrochlamys the outer tepals are 24—28 mm long
and the claws 12.5— 11 mm long, thus half as long
as the entire tepal.
We believe the differences are sufficient that
Ferraria uncinata subsp. macrochlamys should be
treated as a separate species. In fact, we are not
convinced that it is most closely related to F. un¬
cinata. The central Namaqualand F kamiesmon-
tana has flowers virtually identical to those of F
uncinata subsp. macrochlamys in size, proportion,
and orientation, as well as color, and can be distin¬
guished from the subspecies solely by a few dark
spots at the bases of the outer tepal limbs. Leaf
margins of F. kamiesmontana are, however, entire,
smooth, and barely thickened. Growth form of the
two is similar: plants of both taxa are low growing
and have a few subequal branches near the stem
apex, whereas F. uncinata subsp. uncinata often
branches above the base, is frequently taller, and
the stems emerge from the leaf sheaths. Relation¬
ships of F uncinata subsp. macrochlamys are by
no means obvious, and it is most useful to treat it
as a separate species, pending phylogenetic anal¬
ysis of Ferraria using molecular techniques.
Regarding the type collection of Ferraria ma¬
crochlamys, the specimen bears the annotation
“Forsyth 1835” suggesting a gathering by a Mr.
Forsyth. No collector of this name in known in
southern Africa, and the name Forsyth, listed by
Gunn and Codd (1981) in their compilation of plant
collecting in southern Africa, is without initials,
dates of birth and death, and lacking additional
information. We now believe the annotation refers
to William Forsyth, son of W. F. Forsyth of the Chel¬
sea Physic Garden, who maintained a herbarium
collection. William Forsyth died in July 1835 and
his library, presumably containing herbarium spec¬
imens, was auctioned in November of that year (C.
Nelson, pers. comm.). Indications in the accounts
of Bahiana spiralis Baker, based on another Forsyth
specimen (Baker, 1892, 1896), have led us to con¬
clude that the Forsyth herbarium was purchased by
Wiliam Banks, then Keeper of the Kew Herbarium,
for the collection there. The most likely collector
ol the type of B. spiralis and Ferraria macrochlamys
is the British botanist James Niven, who collected
in Namaqualand in 1799 and later, and whose
specimens are now widely dispersed (Gunn &
Codd, 1981). The type citation above reflects this
new information.
Moraea simplex Goldblatt & J. C. Manning, sp.
nov. TYPE: South Africa. Western Cape: 21
km N of Piketberg, 18 Sep. 2002, F. Goldblatt
& /,. J. Porter 12231 (holotype, NBG; isotypes,
MO, PRE). Figure 2.
Plantae 28— M) cm altae, cormo ovoideo ca. 10 mm dia-
metro tunicis reticulatis fibrosis brunneis obtecto, foliis
productis 1 canaliculatis 2-3 mm latis, caule perramoso,
spatha interne inflorescentiis 18—22 mm longis, floribus
flavo-cremeis, tepalorum limbis leviter reflexis unguibus
ascendentibus oa. 2 mm longis, limbis externis 18—20 X
7.5— 8.5 mm intends ca. 18X7 mm, filamentis liberis ca.
5 mm longis inferne in colunmam gracilem conniventibus
superne per 1 mm divergentibus, antheris albis ca. 3 mm
longis, stylo ramis filiformibus indivisis ca. 4 mm longis
inter filamenta arcuatis, ovario exserto anguste ovato ca.
3 mm longo.
Plants 28^40 cm high; corm ovoid, ca. 10 mm
diani., the tunics of coarse, light brown fibers, with
a neck of fairly fine fibers surrounding the base of
the stem; cataphylls papery, the uppermost reaching
shortly above the ground, becoming dry and light
brown by flowering time, then often irregularly bro¬
ken. Foliage leaf solitary, inserted on the first aerial
node, the blade channeled, exceeding the stem,
arching outward or trailing distally, 2-3 mm wide.
Stem erect, lightly flexuose, sticky below the nodes
for half the length of an internode, bearing a
sheathing bract-like leaf at each of the upper
nodes, these 24—35 mm long, green, becoming dry
and light brown above, the apices acute, bearing
two or three short branches of one internode each
at the upper three or four nodes, branches at the
lower nodes sometimes two internodes long and
also branched, the branches erect and parallel to
the stem below, flexed outward above the sheathing
leaf. Inflorescence a rhipidium, terminal on the
branches, several-flowered; spathes subequal,
green, with dry, brown tips, 18-22 mm long, the
outer slightly shorter than the inner. Flowers uni¬
formly pale creamy yellow, the outer tepals each
with a pale yellow mark at the base of the limb,
unscented, the tepal claws ascending, forming a
cup enclosing the base of the filaments; tepals
clawed, the outer slightly larger than the inner, the
claws ascending, ca. 2 mm long, the limbs ovate,
obtuse, initially spreading horizontally, later dip¬
ping up to 30° below the horizontal, the outer 18—
20 X 7.5— 8.5 mm, the inner ca. 18 X 7 mm; fil¬
aments ca. 5 mm long, free but conuivent below
and forming a slender column, diverging in the up¬
per I mm; anthers ca. 3 mm long, oblong, ap-
pressed to one another, white, the pollen white; ova¬
ry exserted, narrowly ovoid, ca. 3 mm long; style
dividing shortly below the base of the anthers into
3 filiform branches, each stigmatic apieally and ex¬
tended between the filaments, ca. 4 mm long. Cap¬
sules and seeds unknown.
Volume 14, Number 3
2004
Goldblatt & Manning
African Iridaceae
295
Figure 2. Moraea simplex Goldblatt & J. C. Manning. Drawn from living plants from the type population, Goldblatt
& Porter 12231, by John Manning. Scale bar 10 mm for entire plant, side view of stamens and style branches approx¬
imately 3X scale bar.
296
Novon
Phenology. Flowering in mid to late September,
probably into mid October.
Etymology. From the Latin simplex, “un¬
adorned, plain.” referring both to the pale-colored
(lowers without markings, the spreading subequal
tepals, and the filiform style branches lacking style
crests.
Distribution anil biology. Moraea simplex is
known oidy from the relatively moist clay flats at
the eastern foot of the Piketberg Mountains. Nu¬
merous plants were found at the beginning of tbeir
(lowering or still in bud on September 18 at tbe
single known site for the species, which had been
burned the previous summer. We infer that (low¬
ering continues until at least early October. The
flowers each last a single afternoon, opening at
about 14:00H and wilting shortly after nightfall.
Relationships. The seventh member of the tax-
onomically isolated subgenus Visciramosa (Gold-
blatt, 1986; Goldhlatt Manning, 2000b), Moraea
simplex has the typical attributes of the subgenus,
including sticky internodes, multiple leaves,
branched stems, relatively short inflorescence
spathes, and free, connivent filaments. Plants are
readily mistaken (or the common and widespread
M. inconspicua Goldhlatt in the vegetative state as
both species share narrow leaves, short, subequal
rhipidial spathes, and relatively small flowers.
However, M. inconspicua usually has two foliage
leaves, unequal tepals, the outer larger than the
inner, the tepal limbs strongly reflexed, and rela¬
tively broad, flattened style branches with paired
erect crests, and the anthers appressed to the op¬
posed style branches, the latter condition typical of
the genus. In contrast, M. simplex has solitary fo¬
liage leaves, the inner and outer tepals nearly equal
in size and spreading to slightly dipping below the
horizontal, and simple, filiform stvle branches that
extend between the filaments.
That the species belongs in Moraea cannot be
doubted despite the unusual filiform style branches
that extend between the filaments, for the resem¬
blance to other species of subgenus Visciramosa is
striking. It has firm, channeled leaves, stieky nodes
and internodes, and the distinctive lateral branch¬
ing pattern in which the branch axis is parallel to
the main stem below, before arching outward almost
horizontally above the sheathing bract, but with the
rhipidia held erect, and the subequal, obtuse rhip¬
idial spathes of the subgenus. Only the flowers are
discordant, not only with the subgenus, but with
Moraea in general. Reduced style branches, not or
hardly wider than the anthers they subtend, are rel¬
atively common in Moraea anil were the hallmark
of the genus Homeria (Goldhlatt, 1981), now in¬
cluded in Moraea (Goldhlatt. 1998; Goldhlatt et ah,
2002). Even in Homeria, however, the style branch¬
es have forked apices and lie opposite the stamens
and appressed to the anthers. Completely undivid¬
ed style branches, not opposed to the stamens, were
the feature that distinguished Roggeveldia (Gold¬
hlatt, 1979). The two species of Roggeveldia are
now also included in Moraea (Goldhlatt, 1998) and
are most probably related to Moraea crispa Thun-
berg and its immediate allies (Goldhlatt. 1986).
Thus the filiform, undivided style branches held
alternate to the stamens are a parallel development
in a second lineage of Moraea. Discovery and de¬
scription of Moraea simplex brings the number of
species in the genus to 197 (Goldhlatt & Manning,
2002).
In Moraea tbe following combinations at subspe¬
cies rank were inadvertently not made when spe¬
cies of Hexaglottis and Homeria were transferred to
that genus (Goldhlatt, 1998).
Moraea bulbillifera subsp. anomala (Goldhlatt)
Goldhlatt, comb. nov. Basionym: Homeria bul¬
billifera subsp. anomala Goldhlatt, Ann. Mis¬
souri Hot. Gard. 68: 458. 1981. TYPE: South
Africa. Western Cape: Witsands near Ossekop,
P. Goldhlatt 4885 (holotype, MO; isotypes, BR,
K. NBG. PRE, WAG).
Representative specimen. SOUTH AFRICA. Western
Cape: 3321 (Ladismith) Cloete's Pass to Wagenbooms
(DDK Sep. 1976, Goldhlatt 4157 ( K. MO. PRE, S. WAG).
Moraea lewisiae subsp. secunda (Goldhlatt)
Goldhlatt, comb. nov. Basionym: Hexaglottis
lewisiae subsp. secunda Goldhlatt, Ann. Mis¬
souri Bot. Gard. 74: 558. 1987. TYPE: South
Africa. Northern Cape: near the top of Spek-
takel Pass, P. Goldhlatt 6678 (holotype, PRE;
isotypes, K, MO, NBG, S, US, WAG).
Representative specimen. SOUTH AFRICA. Northern
Cape: 3119 (Calvinia) Nieuwoudtville, karroid hills NE
of Klipkoppies (AC). Sep. 1981, Goldhlatt 7078 (MO).
Moraea virgata subsp. karooica (Goldhlatt)
Goldhlatt, comb. nov. Basionym: Hexaglottis
virgata subsp. karooica Goldblatt, Ann. Mis¬
souri Bot. Gard. 74: 568. 1987. TYPE: South
Africa. Northern Cape: 22 km W of Middelpos,
D. Snijman 765 (holotype, NBG; isotypes, K,
MO, PRE).
Representative specimen. SOUTH AFRICA. Northern
Cape: 3120 (Williston) Roggeveld Escarpment, 71 mi. SE
of Calvinia on Middelpos road via Blomfontein (CC), Sep.
1981, Griddhlatt 4621 (MO. NBG).
Volume 14, Number 3
2004
Goldblatt & Manning
African Iridaceae
297
Nivenioideae
Aristea torulosa Klatt, Abh. Naturf. Ges. Halle
15: 382. 1882. TYPE: South Africa. Eastern
Cape: Transkei, betw. Gekau & Bashee, Dec.
1832. J. F. I)rege 4558 (lectotype, effectively
designated by Weimarck (1949: 27). B not
seen, photo, K; isotype, P).
Aristea torulosa var. monostachya Baker, FI. Cap. 6: 49.
1896. TYPK: South Africa. Natal, without precise
locality or date, Buchanan s.n. (type, K).
Aristea woodii N. E. Brown, Kew Bull. 1931: 192. 1931.
Syn. nov. TYPE: South Africa. KwaZulu-Natal, In-
anda, J. M. Wood 757 (type, K).
Aristea torulosa has been overlooked in recent
times, and Vincent (1985) excluded the species
from his account of Aristea in eastern southern Af¬
rica because he considered that no specimens
could be assigned to it. J. G. Baker (1896) and
Weimarck (1940), however, had no misgivings
about its identity and status, and Baker (1896) ac¬
tually regarded what was later described as A. woo¬
dii as A. torulosa var. monostachya Baker. Aristea
woodii and A. torulosa have identical derived, rel¬
atively large rhipidial spathes and bracts that are
dark green below, aging reddish brown, that grade
unevenly into broad translucent margins, darker
along the veins thus giving them a feathered ap¬
pearance. The pale edges of the spathes and bracts
become torn and irregularly fringed with age.
The only significant difference between plants
assigned to Aristea woodii and the type of A. toru¬
losa is the crowded flowering axis of the latter,
which has the lower nodes bearing short lateral
branches. Typical A. woodii has a more slender
flowering stem usually bearing only sessile flower
clusters. Plants readily referable to ,4. woodii may,
however, have one or even two of the lowermost
nodes bearing stalked flower clusters; thus, in ef¬
fect, the flowering stem is branched. The type of a
second species, regarded as a synonym of A. toru¬
losa by Weimarck (1940) but excluded by Vincent
(1985), A. congesta, described by N. E. Brown in
1931 and based on plants from the Transkei. South
Africa, is just such a plant, and we see no reason
to distinguish it from A. woodii. The more congested
floral axis of A. congesta in turn, is little different
from that of A. torulosa. Plants similar to the types
of A. congesta and A. torulosa are matched by
Krauss s.n. (K) from Port Natal and Rehmann s.n.
(K) from Woodbush in Limpopo Province. The
Krauss collection was referred by Weimarck to A.
torulosa, while the Rehmann collection was not cit¬
ed by either Weimarck (1940) or Vincent (1985).
More extensive collections now available, at NBG,
MO, and PRE, suggest that all three named species
represent single taxon. The earliest name for this
widespread species that extends from Tanzania and
eastern Angola (Goldblatt, 1993, 1996) in the north
to eastern South Africa in the south thus becomes
A. torulosa.
Acknowledgments. We thank Ingrid Nanni and
Lendon Porter for their assistance and companion¬
ship in the field. Support for this study by grants
7103-01 and 7316-02 from the National Geograph¬
ic Society is gratefully acknowledged. Collecting
permits were provided by the Nature Conservation
authorities of Northern Cape and Western Gape
Provinces, South Africa.
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& Sons, London.
- . 1896. Irideae. Pp. 7-71 in W. T. Thiselton-Dyer
(editor). Flora Capensis 6. Ashford, Kent.
Brown, N. E. 1931. XXIX — Notes upon some South Af¬
rican plants. Kew Bull. Misc. Inform. 1931: 191 — 197.
tie Vos, M. P. 1979. The African genus Ferraria. ,1. S.
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- . 1983. The African genus Tritonia Ker-Gawler:
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- . 1988. Three new species of Ixia I,. (Iridaceae)
from the Cape Province. S. African J. Bot. 54: 596—
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- . 1999a. Tritonia (Iridaceae). In G. Germishuizen
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tional Botanical Institute, Pretoria.
- . 1999b. Ixia (Iridaceae). In G. Germishuizen (ed¬
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centrale. Taxonomania 4: 2—3.
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839-844.
- . 1981. Systematics and biology of Homeria (Iri¬
daceae). Ann. Missouri Bot. Gard. 68: 413—503.
- . 1986. fhe Moraeas of Southern Africa. Ann. Kir-
stenboseh Bot. Gard. 14: 1—224.
- . 1990. Phylogeny and classification of Iridaceae.
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- . 1996. Gladiolus in Tropical Africa. Timber Press,
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- . 1998. Reduction of Barnardiella, Galaxia, Gyn-
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tive radiation of pollination mechanisms in Sparaxis (Ir¬
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Tradescantia boliviana (Commelinaceae), a New Combination for
an Overlooked South American Species
Jason R. Grant
Laboratoire tie botanique evolutive, Institut de botanique, Faeulte des Sciences, Universite
de Neuchatel, rue £mile-Argand 1 1, Case Postale 2, 2007 Neuchatel, Switzerland.
jason.grant@unine.ch
Abstract. the new combination Tradescantia
boliviana (Hasskarl) J. R. Grant is made in recog¬
nition that it is a distinct species widespread in
Peru, Bolivia, Paraguay, and Argentina, here sep¬
arated by morphological characters and distribution
from its previous inclusion within T. ambigua Mar¬
tins ex J. H. Schultes, which is restricted to north¬
eastern Brazil.
Key words: Argentina, Bolivia, Brazil, Mandon-
ia, Paraguay, Peru, Tradescantia.
While researching the Tradescantia sect. Man-
donia complex, problems in species limits and no¬
menclature were discovered in the non-Mexican
species. A new combination is proposed as Trad¬
escantia boliviana (Hasskarl) J. R. Grant in recog¬
nition that the material from Peru, Bolivia, Para¬
guay, and Argentina represents a distinct species.
This species is here separated from synonymy un¬
der T. ambigua Martius ex J. H. Schultes, which is
restricted to northeastern Brazil.
Ten species comprise Tradescantia sect. Man-
donia D. H unt. The group has a center of diversity
in Mexico with six species: T. burchii I). Hunt
[found in Jalisco], T. crassifolia Cavanilles (includ¬
ing T. acaulis Martens & Galeotti and T. iridescens
Bindley) [northern Mexico to Guatemala], T. exal-
tata D. Hunt |()axaca], T. llamasii Matuda [Est.
Mexico], T. peninsularis Brandegee [Baja Califor¬
nia], and T. tepoxtlana Matuda [Morelos] (Hunt.
1980, 1986). Four non-Mexican species range from
Guatemala to South America: T. ambigua Martius
[distributed in northeastern Brazil], T. boliviana
(Hasskarl) J. R. Grant [Argentina, Bolivia. Para¬
guay, Peru], T. petricola J. R. Grant [Costa Rica,
Venezuela] (Grant, 2000), and T. velutina Kunth &
C. 1). Bouehe [Guatemala, Honduras, and Nicara-
gua].
While researching Tradescantia petricola , addi¬
tional specimens were located from Venezuela that
matched this previously envisioned Costa Rican
endemic (Grant, 2000). While further searching
among South American indets and among T. am¬
bigua in many herbaria (COL, G, Gil. IAN. INPA,
L, MG, MO, NY, P, R, RB, SP, SPF, and US), it
became evident based on plant size, leaf disposi¬
tion, sepals, and distribution that the specimens
from Peru, Bolivia, Paraguay, and Argentina were
clearly different from those from Brazil. While it
was initially thought that these represented a new
species, the name Mandonia boliviana was found
to apply. Type material of Mandonia boliviana was
located at Leiden (L, 1 sheet), Paris (P, 3 sheets),
and Geneva (G, 2 sheets), and its lectotypification
is proposed below. Examination of this type mate¬
rial, and comparison to recently collected speci¬
mens, clarifies its identity as distinct from T. am¬
bigua. Since the combination into Tradescantia has
not been made, the new combination is proposed
here.
Tradescantia ambigua is restricted to the caatin-
ga of northeastern Brazil, while Tradescantia boli¬
viana is restricted to the Gran Chaco of Argentina,
Bolivia, Paraguay, and Peru (Fig. 1).
Ti •adeseantia boliviana (Hasskarl) J. R. Grant,
comb. nov. Basionym: Mandonia boliviana
Hasskarl, Flora 54: 260. 1871. Skofitzia boli¬
viana (Hasskarl) Hasskarl & Kanitz, Oesterr.
Bot. Z. 22: 147. 1872. TYPE: Bolivia. Lare-
caja: “Viciniis Sorata montis colles Ullontigi
ad scopulorum radicem in regione temperata,”
2700 m, Feb.— Apr. 1858, G. Mandon 1239
(lectotype, designated here, L [Herb. J. K.
Hasskarl]; isotypes, G [2], k not seen. P. P
[originally from ‘Herb. E. Cossen'], P [origi¬
nally from ‘Herbier E. Drake’)).
Specimens examined. ARGENTINA. Cordoba: [entre
Panolma y la Villa del Transito (P)], [zwischen Panolma
unter der Villa del Transito und der Westseite der Tierra
Achala de Cordoba (US)], 23 Feb. 1876, Hieronymus s.n.
(P, US); Rodado, Stuckert 10938 (G); Sierra Cleica de Cor¬
doba, Stuckert 13530 (G), lessen 450 (GH); Punilla, Cap-
ilia del Monte, Cuezzo & Balegno 2144 (G. US). Salta:
M. Sala, Venturi 8180 (GH, US); Cerro San Bernardo, Sal¬
ta, W [no collector?] 6151 (GH); Qrdn, Pintascayo, Pierotti
6558 (G); Cachi, Quebrada de Escoipe, Ruta 30, 2 km
Novon 14: 299-301. 2004.
300
Novon
Figure 1. Distribution ol Tradescantia boliviano in open circles, and T. ambigna in solid squares.
antes de FI Maray, Novara 6067 (G); Rosario de Lerma,
Quebrada del Rio Blanco, 2—3 km al SW (aguas arriba)
del rio Toro, Novara 7603 (G [3]). Tueunian: Capital,
Dunaquito, Venturi 1660 (GH, US), Venturi 1660a ((ill.
US); Tufi del Valle, Venturi 2021 ((ill. US). BOLIVIA.
Chuquisaea: Tomina, Weddell 3330 (P). Cochabamba:
near Taquina, above Cochabamba, Everdam 25231 (G);
Cochabamba, Julio 11.174 (NY, US), Cardenas 5560 (US);
Camino a Saeabe, cerca a Cochabamba, Ochoa 670 (NY);
Cercado de Cochabamba, la Colina de San Pedro, Stein-
bach s.ri. (US); Campero, Mique, Steinbach 755 (GH, NY.
US); Cerro San Pedro, behind Cochabamba, Cutler & Car¬
denas 3830 (elow 500 m, and is possibly amphibious. (See
S. latifolius for further discussion of habitat.)
The holotype of Syngonanthus dichroanthus at
NY is a mixed sheet including an unidentified ro-
sulate species of Eriocaulon of about the same stat¬
ure. The isotype sheet at P is a mixture with the
same Eriocaulon as well as with Eleocharis minima
Kunth (Cyperaceae). both sheets were distributed
as S. simplex (Miquel) Ruhland.
This inconspicuous mud-dwelling species is
known so far only from mixed collections with other
diminutive species. Its relatives include several
other small rosulale annuals with pale glabrous ca-
pitula, including Syngonanthus gracilis (Bongard)
Ruhland, S. biformis (N. E. Brown) Gleason, 5. sim¬
plex (Miquel) Ruhland, and S. latifolius (Moldenke)
Hensold. Syngonanthus dichroanthus is distin¬
guished from these in [tart by its delicate habit,
membranous threadlike leaves, and involucral
bracts exceeding the capitulum. From the closely
sympatric and common taxon S. gracilis var. aureus
Ruhland, it can usually be distinguished by the
lack of glandular hairs on the peduncle sheaths.
However, its most striking feature is the strong con¬
trast between the copper-colored staminate flowers
and the white pistillate flowers, an unusual char¬
acteristic shared only with S. latifolius.
Strong differentiation in the color of the stami¬
nate and pistillate (lowers occurs to my knowledge
in only a few divergent species of Syngonanthus.
These include the dimorphic-stemmed aquatic S.
spongiosus Hensold, with staminate flowers deep
yellow-brown, and the rosette species S. angolensis
Hess of Africa and S. goyazensis (Kornicke) Ruh¬
land of Brazil, in both of which the staminate ca¬
lyces are deeply pigmented only below the lobes.
The specimen from Manaus ( Liitzelburg 21962
p.p.) lacks staminate flowers, which makes positive
identification difficult, but is provisionally placed
here due to the glabrous flowers, involucral bracts
glabrous and surpassing the capitulum (which dis¬
tinguish it from Syngonanthus biformis and S. sim¬
plex), the membranous leaves, and lack of glandular
hairs.
Paratype. BOLIVIA. Santa Cruz: Prov. Velasco, Es-
taci6n Flor de Oro, margen del Rfo It6nez (Guapor6), fron-
tera con Rondonia, ea. 20 km N del Serranfa de Huan-
chaca, ea. 85 km E del Rfo Paragua, 15 May 1991 (fl),
M. Pena et al. 62 p.p. (F, mixed collection with Syngon¬
anthus gracilis var. aureus Ruhland).
Additional specimen examined. BRAZIL. Amazonas:
Manaus, Villa Municipal, 28 Aug. 1916 (9), Liitzelburg
21962 p.p. (LL = fragment ex NY, mixed collection with
Utricularia and sterile graminoid, probably Cyperaceae).
Syngonanthus latifolius (Moldenke) Hensold,
comb. nov. Basionym: Syngonanthus gracilis
var. latifolius Moldenke, Phytologia 21: 418.
1971. TYPE: Brazil. Mato Grosso: Serra do
Roncador, gallery margin, ca. 86 km N of Xav-
antina, 550 m, 31 May 1966 (fl, fr), H. S. Ir¬
win, J. W. Crear Jr., R. Souza, l\. Reis dos San¬
tos 16349 (holotype, NY). Figure 2G— J.
Plants rosulate annuals; roots white, spongy,
branched, the larger 0.5— 0.7 mm diam. Leaves re¬
curved, ± prostrate, ca 1.0-1. 2 cm long, 1.0-1. 5
mm wide, ligulate, subacute, the tip minutely re¬
curved and keeled; 3- to 5-veined, the midvein vis¬
ible adaxially as a white line, at least in the basal
half; pubescent abaxially with short appressed mal-
pighian hairs, and adaxially with long filamentous
arachnoid hairs, both surfaces glabrate. Inflores¬
cences numerous, ca. 25 to 50, of mixed ages, some¬
times with new inflorescences initiating outside the
older ones; peduncle sheaths 10—17 X 0.6— 0.8 mm
diam., the lamina ca. 2 mm long, acute, callous-
tipped, slightly recurved; arachnoid pubescent as
the leaves, also with scattered upright gland-tipped
hairs; white-costate only near the base; peduncles
6—15 cm long, (0.25— )0.3— 0.4 mm wide, 3-costate;
the intercostae green and arachnoid pubescent,
broad near the apex, narrowing to thin furrows be¬
low; the costae stramineous, glabrous, shiny, with a
few gland-tipped hairs when young; capitula 4—5.5
mm diam., ovoid in bud, depressed-hemispheric to
globose at maturity; involucral bracts hyaline, gla¬
brous, lustrous, the outer obovate, rounded-apieu-
late at apex, pale, shredding with age, the inner
oblanceolate-obtuse to linear-acute and gold with a
brown tint; shorter than the pistillate flowers; re¬
ceptacle densely white-villous, trichomes about
equaling the staminate flowers; receptacular bracts
absent. Flowers 3-merous, the staminate and pistil¬
late about evenly mixed and strongly dimorphic,
the staminate flowers pigmented and much shorter
than the pistillate, the pistillate surpassing the in¬
volucral bracts by 0.6— 0.7 mm, abscising after seed
set; staminate flowers : pedicels ca. 0.15—0.25 mm;
calyx coppery brown, zygomorphic, the segments
ca. 1.0 X 1.0 mm, broadly obovate-spalulate,
rounded, slightly cucullate, fused ca. Vi — % of their
length, sparingly pubescent to glabrate on the ab-
axial face; corolla borne on stipe ca. 0.5 X 0.1 mm;
the corolla itself ca. 0.35 mm long, membranous,
the lobes involute; filaments ca. 0.35 mm, adnate
to corolla; anthers while, 0.2 mm long; pistillodes
ca. 0.25 mm long, infundibular, the rim papillate;
pistillate flowers: pedicels ca. 0.5 mm long; sepals
oblong-elliptic, acute to broadly acuminate, navic¬
ular and slightly sclerified along midvein at base.
Volume 14, Number 3
2004
Hensold
South American Eriocaulaceae
307
Figure 2. A— F. Syngonanthus dichroanthus Hensold. — A. Habit. — B. Capitulum. — C. Starninate flower. — I). Corolla
of starninate flower. — E. Pistillate flower. — F. Gynoecium. G— J. Syngonanthus latifolius (Mold.) Hensold. — G. Sta-
minate flower. — H. Pistillate flower. — I. Gynoecium. — J. Habit. Vouchers: A— F Pires & Black 2,165', ( i — J Irwin et al.
16349.
308
Novon
1. 7-2.0 X 1.0 mm. white-hyaline, tinged brown at
apex, glabrous; corolla short-stipitate, the petals
fused at middle margins, free at base and apex,
linear-oblanceolate, acute to acuminate, involute
after anthesis, ca. 1.1— 1.3 X 0.25 mm, membra¬
nous, glabrous at maturity; style ca. 0.6 mm long,
style branches ca. 0.4 mm long, appendages small
and filamentous. Seeds red-brown, ca. 0.4 X 0.2
mm, with raised white longitudinal ribs after wet¬
ting. the epidermis waxy (under SEM).
Ecology and distribution. Known from a single
collection in marshy campo at 550 m, at the margin
of gallery forest in cerrado of northeastern Mato
Grosso, Brazil, ca. 52°W, 14°S.
Syngonanthus latifolius was originally described
as a variety of .S', gracilis (Bongard) Ruhland, dis¬
tinguished from other varieties by the leaves wider
and more closely appressed to the ground. The
marked size and color dimorphism of the flowers
were not noted in the original description, however,
and together with the leaf characters, support rec¬
ognition of the taxon as a distinct species.
The closest affinity of Syngonanthus latifolius is
probably with S. dichroanthus Hensold, which has
a nearly identical floral structure and color dimor¬
phism, but flowers subequal in size, and membra¬
nous filiform leaves. In S. latifolius , the sepals and
pedicels of the pistillate flowers are nearly twice as
long as those of the staminate flowers, such that the
small rounded golden-brown staminate flowers are
dwarfed by the white acute pistillate flowers with
which they are intermixed. Similar differentiation
in flower size and a similar aspect to the capitulum
are matched only in the species .S’, bijorrnis (N. E.
Brown) Gleason, but the latter is distinguished by
its uniformly white flowers, pubescent involucres,
densely pubescent petals in the pistillate flowers,
and prominent style appendages.
Affinity is also suggested with Syngonanthus
gracilis var. aureus, with which S. latifolius shares
the annual habit, nearly glabrous sepals and petals,
reduced filiform style appendages, and the waxy
coating observed on the seeds under SEM (Hen-
sold, unpublished data).
Both Syngonanthus latifolius and S. dichroanthus
have been collected together with S. gracilis var.
aureus in wetlands (veredas) associated with cer¬
rado. Eiten (1975) described in detail the abruptly
delineated “seasonally marshy grassy campos" or
“hillside campos,” which separate gallery forests
from uplantl cerrado in the Brazilian Serra do Ron-
eador and elsewhere. Created by rainy season
drainage, these campos have gleyed soils, espe¬
cially in their lower, wetter portions, where Erio-
caulaceae are reported as a typical element. Syn¬
gonanthus dichroanthus may occur on wetter sites
than S. latifolius. since both the Rfo Itenez site and
the veredao in Maranhao are inundated for longer
periods than the valleys of the Serra do Roneador
(Eiten, 1975), but more collections are needed to
confirm tbis.
Additional specimen examined. BRAZIL. Mato 6 ros¬
so: Serra do Roneador, gallery margin, ca. 86 km N of
Xavantina, 550 m, 31 May 1966, II. S. Irwin et al. 16346
p.p. (till) [mixed sheet with X. gracilis var. aureus Ruh¬
land, probably from the same gathering as Irwin et al.
16349. the type].
Acknowledgments. Many thanks to Creg Muell¬
er and the Botany Department at the Field Museum
for providing research space; to Peter Jprgensen,
editor of the Catdlogo de las Plantas Vasculares de
Bolivia, for help and encouragement; to Luzmila
Arroyo and Mike Net* for crawling specimens at
USZ and NY; and to the curators of GH, LL, MO,
NY. and P for loans of material.
I .iterature Cited
Eiten, G. 1975. The vegetation of the Serra do Roneador.
Biotropica 7: 112—135.
Giulietti, A. M. 1978. Modificayoes taxonomicas no genero
Eriocaulon I.. Bol. But. Univ. Sao Paulo 6: 39-47.
- . 1984. Estudos Taxonomicos no Genero Leiothrix
Buhl. (Eriocaulaceae). Tese de Livre-Docencia, Univ-
ersidade de Sao Paulo, Sao Paulo, Brazil.
Killeen. T. J., E. Garefa & S. G. Beck (editors). 1993.
Guta de Arboles de Bolivia. Missouri Botanical Garden,
St. Louis, and Herbario Nacional de Bolivia, La Paz.
Ruhland, W. 1903. Eriocaulaceae. In: A. Engler (editor),
Das Pflanzenreieh IV. 30: 1—294. Wilhelm Engelmann,
Leipzig.
Stiitzel. T. 1988. Untersuchungen zur Wurzelanatomie der
Eriocaulaceen. Flora 180: 223—239.
- . 1998. Eriocaulaceae. Pp. 197-207 in K. kubitz-
ki (editor). The Families and Genera of Vascular Plants,
Vol. IV. Flowering Plants. Monocotyledons. Alismatanae
and Commelinanae (except Gramineae). Springer- Ver-
lag. Berlin. Heidelberg.
Tomlinson, P. B. 1969. Anatomy of the Monocotyledons
III. t iommel inales— Zingiberales. Clarendon Press, Ox¬
ford.
Zhang, Z. 1999. Monographic der Gattung Eriocaulon in
Ostasien. l)iss. Bot. 313: i— vii, 1—289.
Subtribes of Principally North American Genera of Cichorieae
(Compositae)
Joongku Lee 1,2 and Bruce G. Baldwin'-'
'Jepson Herbarium and Department of Integrative Biology, 1001 Valley Life Sciences
Building #2465, University of California, Berkeley, California 94720-2465, U.S.A.
“’Current address: Korea Research Institute of Bioscience and Biotechnology, 52 Oun-dong,
Yuseong-ku, Daejeon 305-333, Korea
’Author for correspondence: bbaldwin@uclink4.berkeley.edu
ABSTRACT. A new subtribal classification (or
principally North American genera of Cichorieae
allows for a practical taxonomy that reflects recent
phylogenetic hypotheses. We propose the newT sub-
tribes Lygodesmiinae (for Chaeladelpha, Lygodes -
mia sensu Tomb, and Shinnersoseris), Pinaropap-
pinae (for Marshal Ijohnstonia and Pinaropappns),
Pyrrhopappinae (for Picrosia and Pyrrhopappus),
and three monogeneric subtribes (Glyptopleurinae
( Glyptopleura ), Krigiinae ( Krigia ), and Phalacroser-
idinae ( Phalacroseris )) for taxa previously treated in
Malacothrieinae, Microseridinae, and Stephanom-
eriinae. We recognize novel circumscriptions for
Malacothrieinae ( Anisocoma , Atrichoseris, Calyco-
seris, and Malacothrix ), Microseridinae (Agoseris,
Microseris, Nothocalais, Stebbinsoseris, and Uropap-
pus ), and Stephanomeriinae ( Munzothamnus , Pleia-
canthus, Prenanthella, Rafinesquia, and Stephan-
omeria). The revised classification provides a more
precise taxonomic representation of relationships
among genera, insofar as robust resolution of mono-
phyletic groups currently allows.
Key words: Cichorieae, Lactueeae, Malacothri¬
einae, Microseridinae, Stephanomeriinae, sub¬
tribes, taxonomy.
Phylogenetic analyses of 18S— 26S nuclear rl)NA
external and internal transcribed spacer sequences
(Lee et al., 2002, 2003) and chloroplast DNA re¬
striction sites (Jansen et al.. 1991; Whitton et al.,
1995) yielded evidence for new hypotheses of re¬
lationship among all genera of Cichorieae (Lactu-
ceae) that have natural centers of diversity in North
America (plus Picrosia, endemic to South America).
On the basis of molecular findings, Lee et al. (2003)
concluded that taxa previously treated in subtribes
Malacothrieinae, Microseridinae, and Stephano¬
meriinae (e.g., Stebbins, 1953; Bremer, 1993,
1994) or in Jeffrey’s (1966) “Microseris subgroup”
and “Stephanomeria subgroup” represent a single
major radiation, with a large western North Amer¬
ican component. As previously treated, each of
those subtribes and subgroups is evidently para-
phyletie or polyphyletic. To better reflect phyloge¬
netic relationships, we propose revised circum¬
scriptions for Malacothrieinae, Microseridinae, and
Stephanomeriinae and recognize six new subtribes.
Malacothrieinae K. Bremer, Novon 3: 329. 1993.
TYPE: Malacothrix DC.
Annual or perennial herbs. Leaf blades linear,
elliptic, lanceolate, oblanceolate, obovate, or spat-
ulate; margins entire, denticulate, lobed, or pinnat-
ifid. Capitulescences corymbiform or capitula borne
singly (peduncles scapiform). Receptacles not pa-
leate, bristly or not bristly. Involucres cylindrical to
obconic or campanulate. Phyllaries in 3 to 6 series,
unequal (1 or 2 subequal series in Atrichoseris).
f lorets 15 to 250 per capitulum. Corollas yellow or
white (often with abaxial reddish stripes). Cypselae
columnar to fusiform, 5- to 15-ribbed or -veined (5
usually prominent), beaked (Calycoseris) or not
beaked. Pappi 0 ( Atrichoseris ) or o( 15 to 40 sub¬
equal, barbellate or, rarely, plumose (Anisocoma),
persistent or often (all or mostly) caducous or frag¬
ile bristles and sometimes an outer crown of minute
teeth. Pollen echinate (Tomb et al., 1974). Chro¬
mosome number 2 n — 14, 18, 20.
Included genera: Anisocoma Torrey & A. Cray,
Atrichoseris A. Cray, Calycoseris A. Cray, Malaco¬
thrix DC.
Microseridinae Stebbins ex Solbrig, Taxon 12:
234. 1963. TYPE: Microseris I). Don.
Annual, biennial, or perennial herbs. Leaf blades
linear, oblong, lanceolate, oblanceolate, or spatu-
late; margins entire, denticulate, sinuate, lobed, or
pinnatifid. Capitulescences branched or, often, ca¬
pitula borne singly (peduncles scapiform). Recep¬
tacles not paleate, not bristly. Involucres subey-
lindrical to obconic or campanulate. Phyllaries in
Novon 14: 309-313. 2004.
310
No von
2+ series, subequal or unequal, the outer some¬
times forming ealyculi. Florets 5 to 300+ . Corollas
yellow, orange, or white (sometimes with abaxial
reddish stripes). Cypselae columnar or fusiform, ±
10-ribbed, beaked (Agoseris) or not. Pappi of (0)2
to 90 equal or unequal, smooth to barbellate (rarely
plumose) bristles or bristle-tipped scales, persistent
or caducous. Pollen echinolophate (Feuer & Tomb,
1977). Chromosome number 2n = 18, 27 (triploid),
or 36.
Included genera: Agoseris Kafinesque, Microseris
1). Don, Nothocalais (A. Gray) Greene, Stebbinso-
seris K. L Chambers, Uropappus Nuttall.
Steplianomeriinae Stebbins ex Solbrig, Taxon 12:
235. 1963. TYPE: Stephanomeria Nuttall.
Annual or perennial herbs, or shrubs. Leal
blades linear, oblong, lanceolate, oblanceolate, ob-
ovate, or scale-like; margins entire, sinuate-den¬
tate, lobed, or pinnatifid. Capitulescences corym-
biform to thyrsoid-paniculiform. Receptacles not
paleate, not bristly. Involucres narrowly cylindrical
or obconic. Phyllaries usually in 2 series, unequal
(outer forming ealyculi), sometimes graduated (Ste-
pharwmeria cichoriacea). Florets 3 to 15 (to 30 in
Rafinesquia) per capitulum. Corollas pink, purple,
or white. Cypselae clavate, columnar, or fusiform,
smooth, sometimes grooved or ^ 5-ribbed, beaked
(Rafinesquia) or not. Pappi ol 5 to 20+ equal or
unequal ( Prenanthella ), plumose {Rafinesquia, Ste¬
phanomeria), or barbellate, persistent, or deciduous
{Munzothamnus) bristles. Pollen eehinate (Tomb ct
al., 1974). Chromosome number 2 n = 14 {Pre¬
nanthella), 16, 24 (triploid), or 32.
Included genera: Munzothamnus P. II. Raven,
Pleiacanthus Rydberg, Prenanthella Rydberg, Raf¬
inesquia Nuttall, Stephanomeria Nuttall.
Glyptoplcurinue Joongku Lee & B. G. Baldwin,
subtribus nov. TYPE: Glyptopleura I). C. Ea¬
ton.
Herbae annuae. Laminae foliorum spatulatae vel oblon-
gae, profunde lobatae vel pinnatifidae; margines albi, sca-
riosi, dentati vel setoso-ciliati. Capitula portata singulatim
vel capitulescentiae parcc ramosae. Receptacula epaleata,
non setosa. Involucra cylindracea vel urceolata, ad bases
bracteis spatulatis marginihus albis, scariosis, dentatis.
Phyllaria 2-seriata, ± aequalia. Flosculi 7—16. Corollae
cremeae vel luteolae (in sicco subroseae). Cypselae cla-
vatae, tuberculatae, valde costatae, rostratae. Pappi con-
slati ex setis 100+, aequalibus, barbellatis, extimis de-
ciduis. Pollina echinata. 2 n = 18.
Annual herbs. Leaf blades spatulate to oblong,
deeply lobed to pinnatifid; margins white, searious,
dentate or setose-ciliate. Capitula borne singly or
capitulescences sparingly branched. Receptacles
not paleate, not bristly. Involucres cylindrical to ur-
ceolate, al base with spatulate bracts with margins
white, searious, toothed. Phyllaries 2-seriate, ±
equal. Florets 7 to 16 per capitulum. Corollas
cream-colored to light yellow (when dry, pink).
Cypselae clavate, tuberculate, strongly ribbed,
beaked. Pappi ol 100+ equal, barbellate bristles,
the outermost deciduous. Pollen eehinate (Tomb et
al., 1974). Chromosome number 2n = 18.
Included genus: Glyptopleura I). C. Eaton.
ki -igiiuae Joongku Lee & B. G. Baldwin, subtribus
nov. TYPE: Krigia Schreber.
Ilerbae annuae vel perennes. Laminae foliorum line-
ares, oblongae, lanceolatae, vel oblanceolatae; margines
integri, lobati, vel pinnatifidi. Capitula portata singulatim
vel capitulescentiae cymosae vel interdum superne sub-
umbellatae. Receptacula epaleata. Involucra turbinata,
campanulata, vel urceolata. Phyllaria 1— 2 + -seriata, ae¬
qualia. Flosculi 4—30. Corollae llavae vel aurantiacae.
Cypselae columnares vel ovoideae, 10-20-nervatae vel -
costatae, non rostratae. Pappi null i vel ex squamis 5 +
constati (interdum fascientibus coronis minutis), pler-
umque cingentibus setis 5 + , inaequalibus, spiculatis.
Pollina echinolophata. 2n = 8. 10. 12, 18. 20. 30, vel 60.
Plants annual or perennial herbs. Leaf blades
linear, oblong, lanceolate, or oblanceolate; margins
entire, lobed, or pinnatifid. Capitula borne singly
or capitulescences eymiform or subumbelliform.
Receptacles not paleate. Involucres turbinate, cam-
panulate, or urceolate. Phyllaries I - or 2 + -seriate,
equal. Florets 4 to 30 per capitulum. Corollas yel¬
low or orange. Cypselae columnar or ovoid, 10- to
20-nerved or -ribbed, not beaked. Pappi none or of
5+ scales (sometimes forming minute crowns), usu¬
ally surrounding 5 + . unequal, spiculate bristles.
Pollen echinolophate (Feuer & Tomb, 1977). Chro¬
mosome number 2 n = 8, 10, 12. 18, 20, 30, or 60.
Included genus: Krigia Schreber.
Lygodesiniiiiac Joongku Lee & B. G. Baldwin,
subtribus nov. TYPE: Lygodesmia I). Don.
Herbae annuae ( Shinnersoseride ) vel perennes. Laminae
foliorum lineares vel lanceo-lineares (vel squamiformes
distaliter); margines plerumque integri, interdum lobati.
Capitula raro portata singulatim, capitulescentiae pler¬
umque ramosissimae. Receptacula epaleata. Involucra cy¬
lindracea vel subcylindracea. Phyllaria 2-seriata, inae-
qualia (extimis fascientibus calyculis). Flosculi 5—12.
Corollae subrosaceae, purpureae, vel albae. Cypselae col¬
umnares vel fusiformes, laeves, strialae, rugosae, vel cos¬
tatae, non rostratae. Pappi ex setis 24—250, aequalis, bar¬
bellatis (el squamis 5, subulatis in Chaetadelpha), constati
omnibus persistentibus. Pollina echinata vel echinolopha¬
ta {Lygodesmia). 2 n — 12 (Shinnersoseride) vel HI, vel 27
(triploideis).
Volume 14, Number 3
2004
Lee & Baldwin
North America Genera of Cichorieae
311
Annual (, Shinnersoseris ) or perennial herbs. Leaf
blades linear or lance-linear (or scale-like distally);
margins usually entire, sometimes lobed. Capitula
rarely borne singly, capitulescences usually much-
branched. Receptacles not paleate. Involucres cy¬
lindrical or subcylindrical. Phyllaries 2-seriate. un¬
equal (the outer forming calyculi). Florets 5 to 12
per capitulum. Corollas pink, purple, or white. Cyp-
selae columnar or fusiform, smooth, striate, rugose,
or ribbed, not beaked. Pappi of 24 to 250 equal,
barbellate bristles (and 5, subulate scales in Chae-
tadelpha), all persistent. Pollen echinate or eehin-
olophate ( Lygodesmia ) (Tomb et al., 1974). Chro¬
mosome number 2n — 12 ( Shinnersoseris ) or 18, or
27 (triploids).
Included genera: Chaetadelpha A. Gray, Lygo¬
desmia I). Don (sensu Tomb), Shinnersoseris Tomb.
Phalacroseridinae Joongku Lee & B. G. Baldwin,
subtribus nov. TYPE: Phalacroseris A. Gray.
Herbae perennes. Laminae foliorum lineares, lanceo-
lineares, vel oblanceolatae; margines integri. Capitula por-
tata singulatim (pedunculis scapiformibus); receptacula
epaleata; involucra campanulata; phyllaria 2+-seriata, ae-
qualia. Flosculi 13 + , corollae Havae. Cypselae fusiformes,
laeves, non rostratae; pappi nulli. Pollina ecliinolophata.
2 n = 18.
Perennial herbs. Leaf blades linear, lance-linear,
or oblanceolate; margins entire. Capitula borne sin¬
gly (peduncles scapiform). Receptacles not paleate.
Involucres campanulate. Phyllaries 2 + -seriate,
equal. Florets 13+ per capitulum. Corollas yellow.
Cypselae fusiform, smooth, not beaked. Pappi none.
Pollen eehinolophate (Feuer & Tomb, 1977). Chro¬
mosome number 2 n = 18.
Included genus: Phalacroseris A. Gray.
I'inaropappinae Joongku Lee & B. G. Baldwin,
subtribus nov. TYPE: Pinaropappus Lessing.
Herbae perennes vel frutices rosulatae ( Marshalljohn -
stonia). Laminae foliorum lineares (vel squamiformes dis-
taliter), ellipticae, vel rhombeae; margines integri, dentati,
vel lobati. Capitula portata singulatim. Receptacula pa-
leata (. Pinaropappo ) vel epaleata. Involucra obconica vel
campanulata. Phyllaria 3 + -seriata, inaequalia. Flosculi
10—18. Corollae subrosaceae, purpureae, vel albae. Cyp¬
selae suleis 5, angustis, aequaliter dispositis, columnares
vel fusiformes, non rostratae (distaliter attenuatae in Pi¬
naropappo p.p.). Pappi ex setis 40-60, aequalis, barbel-
latis, persistentibus constati. Pollina echinata. 2 n = 18
vel 36.
Perennial herbs or rosette shrubs ( Marshalljohn -
stonia). Leaf blades linear (or scale-like distally),
elliptic, or rhombic; margins entire, dentate, or
lobed. Capitula borne singly. Receptacles paleate
( Pinaropappus ) or not paleate. Involucres obconic
or campanulate. Phyllaries 3 + -seriate, unequal.
Florets 10 to 18 per capitulum. Corollas pink, pur¬
ple, or white. Cypselae columnar or fusiform, with
5 narrow, equally spaced furrows, not beaked (dis¬
tally attenuate in Pinaropappus pro parte). Pappi of
40 to 60 equal, barbellate bristles, persistent. Pol¬
len echinate (Tomb et al., 1974). Chromosome
number 2 n — 18 or 36.
Included genera: Marshalljohnstonia Henrick-
son, Pinaropappus Lessing.
Pyrrhopappinae Joongku Lee & B. G. Baldwin,
subtribus uov. TYPE: Pyrrhopappus DC.
Herbae annuae vel perennes. Laminae foliorum ellip¬
ticae, lanceolatae, vel oblanceolatae; margines integri vel
pinnatifidi. Capitula portata singulatim vel capitulescen-
tiae laxe corymbiformes. Involucra cylindracea vel anguste
campanulata. Phyllaria 2-seriata, inaequalia (extimis fas-
cientibus calyculis). Flosculi 30—150+. Corollae flavae,
purpureae, vel albae. Cypselae fusiformes, 5— 12-coslatae,
rostris fragilibus. Pappi ex setis 119—155, aequalis, bar-
bellatis (cum pilis minutis, reflexis subtentis in Pyrrho-
pappo) constati, persistentibus. Pollina echinata ( Picrosia )
vel ecliinolophata (Pyrrhopappo). 2 n - 12, 14, vel 24.
Annual or perennial herbs. Leaf blades elliptic,
lanceolate, or oblanceolate; margins entire or pin-
natifid. Capitula borne singly or capitulescences
loosely corymbiform. Receptacles not paleate. In¬
volucres cylindrical to narrowly campanulate. Phyl¬
laries 2-seriate, unequal (the outer forming calyc¬
uli). Florets 30 to 150+ per capitulum. Corollas
yellow, purple, or white. Cypselae fusiform, 5- to
12-ribbed, beaks fragile. Pappi of 119 to 155 equal,
barbellate bristles (subtended by minute, reflexed
hairs in Pyrrhopappus), persistent. Pollen echinate
(Picrosia) or eehinolophate ( Pyrrhopappus ) (Feuer
6 Tomb, 1977). Chromosome number 2n =12, 14,
or 24.
Included genera: Picrosia I). Don, Pyrrhopappus
DC.
Kky to Principally North American Subtribes of Cichorieae
la. Capitulescences monocephalic (capitula borne singly); cypselae not ribbed or furrowed; pappi absent ....
. Phalacroseridinae
lb. Capitulescences corymbiform to thyrsoid-paniculiform or subumbelliform, or if monocephalic, then evpselae
ribbed or furrowed and pappi present (rarely absent in Microseris douglasii).
2a. Cypselae beaked.
3a. Some or all pappus bristles caducous or fragile.
312
Novon
4a. Capitulescences stipitate-glandular (gland heads ± disk-shaped); pappus hrislles falling
together . Malacothricinae p.p. ( Calycoseris )
4b. Capitulescences not stipitate-glandular; outer pappus bristles caducous or fragile, the inner
persistent . Glyptopleurinae
3b. Pappus bristles persistent.
3a. Pappus bristles plumose . Stephanomeriinae p.p. (Rafinesquia)
5b. Pappus bristles barbellate or smooth.
6a. Capitulescences monocephalic; corollas yellow or orange; beaks shorter than to much
longer than bodies of cypselae; apices of beaks glabrous . . Microseridinae p.p. (Agoseris)
6b. Capitulescences usually loosely corymbiform, sometimes monocephalic; corollas yel¬
low, purple, or white; beaks much longer than bodies of cypselae; apices of beaks
glabrous or villous (if capitulescences monocephalic and corollas yellow, then apices
of beaks villous) . Pyrrhopappinae
2b. Cypselae not beaked (apieally attenuate in Pinaropappus and Uropappus).
7a. Corollas usually yellow or orange (if white, then pappi of scales, scales and bristles, or bristle-
tipped scales); pollen echinolophate.
8a. Phyllaries in 1 or 2 series, subequal, the outer not forming calyculi; pappi of scales or
scales and bristles . Krigiinae
8b. Phyllaries in 2+ series, subequal or unequal, the outer sometimes forming calyculi; pappi
of bristle-tipped scales or bristles . Microseridinae
7b. Corollas often pink or purple (if yellow or white, then pappi absent (Atrichoseris) or of only
bristles); pollen usually eehinate (echinolophate in Lygodesmia).
9a. Receptacles not paleate, often bristly; florets 15 to 250 per capitulum; corollas usually
yellow, sometimes white; pappus bristles often all or mostly caducous or readily falling . .
. Malacothricinae
9b. Receptacles usually not paleate, rarely paleate (. Pinaropappus ), not bristly; florets 3 to 18
per capitulum; corollas usually pink or purple, sometimes white; pappus bristles usually
persistent.
I()a. Capitulescences monocephalic; phyllaries in 3+ series, unequal . Pinaropappinae
l()b. Capitulescences usually corymbiform to thyrsoid-paniculiform, rarely monocephalic;
phyllaries usually in 2 unequal series, the outer forming a calyculus.
I la. Plants non-thorny herbaceous perennials or opposite-leaved annuals ( Skinner -
soseris); pappus bristles barbellate; pollen usually echinolophate, rarely eehinate
(Chuetadelpha, Shinnersoseris) . Lygodesmiinae
lib. Plants thorny and non-thorny herbaceous perennials, alternate-leaved annuals,
or shrubs; pappus bristles usually plumose (if barbellate and plants herbaceous
perennials, then stems thorny (. Pleiacanthiis )); pollen eehinate Stephanomeriinae
An expanded phylogenetic study of Malcothri-
einae is under way to resolve the memberships of
lineages within the non-monophyletic Malacothrix,
prior to generic redel imitation. We arc also studying
relationships of Thamnoseris (endemic to Islas Des-
venturadas, Chile), which was included by Stebbins
(195.3) in Stephanomeriinae. Data from additional
gene regions are needed to gain refined under¬
standing of the relationships among the subtribes
of New World Cichorieae treated here.
Acknowledgments. We thank Theodore M. Bar¬
kley, Kenton L. Chambers, Victoria C. Hollowell,
David J. Keil, John F. Pruski, and John L Strother
for helpful comments on the manuscript and JLS
and Alan R. Smith for assistance with the Latin
diagnoses; Molly Forbes (University ol California
Botanical Carden), Jon Rebman (San Diego Natural
History Museum), and JLS (University Herbarium)
for providing plant material, loans, or access to
specimens. We especially thank L. I). Gottlieb for
collaborating on the molecular systematic studies
that underlie the taxonomic changes proposed here¬
in. Phis work was supported by a postdoctoral fel¬
lowship (to JL) from the Korea Science and Engi¬
neering Foundation and a grant (to JL) from the
Lawrence R. Heckard Endowment Fund of the Jep-
son Herbarium, University of California, Berkeley.
Literature Cited
Bremer, K. 1993. New subtribes of the I .acluceae (Aster-
aceae). Novon 3: 328—330.
- . 1994. Asteraeeae: Cladistics and Classification.
Timber Press, Portland.
Feuer, S. & A. S. Tomb. 1977. Pollen morphology and
detailed structure of Compositae, tribe Cichorieae, 2;
Subtribe Microseridinae. Amer. J. But. 64: 230—245.
Jansen, It. k., R. S. Wallace, k.-J. kim & k. L. Cham¬
bers. 1991. Systematic implications of ehloroplast I)NA
variation in the subtribe Microseridinae (Asteraeeae:
Lactuceae). Amer. J. Rot. 78: 1015—1027.
Jeffrey, ( 1. 1966. Notes on Compositae: I. The Cichorieae
in east tropical Africa, kew Bull. 18: 427—486.
Lee, J., B. C. Baldwin & L. I). Gottlieb. 2002. Phylogeny
of Stephanameria and related genera (Compositae— Lac¬
tuceae) based on analysis of 18S— 26S nuclear rl)NA
ITS and FI'S sequences. Amer. J. Bot. 159: I0(t-I0R.
- , - & - . 2003. Phylogenetic relation¬
ships among the primarily North American genera of
Volume 14, Number 3
2004
Lee & Baldwin
North America Genera of Cichorieae
313
Cichorieae (Compositae) based on analysis of 18S-26S
nuclear rl)NA ITS and ETS sequences. Syst. Mot. 28:
6 1 6-626.
Stebbins, G. L. 1953. A new classification of the tribe
Cichorieae, family Compositae. Madrono 12: 65—81.
Tomb. A. S.. D. A. Larson & .1. J. Skvarla. 1974. Pollen
morphology and detailed structure of family Composi¬
tae, tribe Cichorieae: I. Subtribe Stephanomeriinae.
Amor. J. Bot. 61: 486-498.
Whitton, J.. R. S. Wallace & R. k. Jansen. 1995. Phylo¬
genetic relationships and patterns of character change
in the tribe Lactuceae (Asteraceae) based on chloroplast
DNA restriction site variation. Canad. J. Bot. 73: 1058-
1073.
A New Combination in Matelea (Apocynaceae— Asclepiadoideae)
Sigrid Liede and IJlrich Meve
Department of Plant Systematics, University of Bayreuth, Universitatsstr. 30, D-95440
Bayreuth, Germany, sigrid.liede@uni-bayreuth.de; ulrich.meve@uni-bayreuth.de
ABSTRACT. Chthamalia cuyabensis is transferred
to Matelea cuyabensis and is lectotypified.
Key words: Apocynaceae, Asclepiadoideae,
Chthamalia, Matelea.
In the course of compiling the Asclepiadoideae
for the Prodromus Florae Matogrossensis (ed. B.
Dubs, Switzerland), a species attributed to the ge¬
nus Chthamalia Decaisne (Apocynaceae— Asclepia¬
doideae; tribe Aselepiadeae, subtribe Gonolobinae)
turned up. The genus Chthamalia, comprising eight
species, was sunk by Woodson (1941) under Ma¬
telea Aublet, and Woodson (1941) combined the
four North and Central American species, including
the type species, C. nummularia Decaisne, under
Matelea. The four Brazilian species were not treat¬
ed by Woodson (1941).
The genus Matelea, with some 280 species, is
one of the least known genera of Asclepiadoideae,
and opinions diverge widely concerning the ques¬
tion whether Matelea is one large, variable genus,
or whether several monophyletic genera are hidden
within Matelea, or whether Matelea even has to be
expanded to comprise the ca. 100 Gonolobus Mi-
chaux species as well. No molecular phylogeny is
yet available, and only some, mostly geographically
restricted, revisions have been carried out in the
last decades (e.g., Stevens, 1985, 1988, 2001). In
Brazil, from where ca. 70 species ol Matelea s.l.
are recorded, none of the four Chthamalia species
has been mentioned in the literature lately. Nev¬
ertheless, Rapini (2001) cited Chthamalia as a syn¬
onym under Matelea. and there is no apparent mor¬
phological reason to reach a different conclusion.
As the name Chthamalia has ceased to exist with
the transfer of the type species to Matelea by Wood-
son (1941), it is necessary to combine Chthamalia
cuyabensis, the only species recorded from Mato
Grosso, to Matelea cuyabensis in order to have a
valid name available for the Prodromus Florae Ma¬
togrossensis, which is about to be published. As
there is, to our knowledge, no ongoing research on
Matelea s.l. in Brazil, and as a new name would
need to be found ( Chthamalia not being available
anymore) if new results were to require the split of
the species from Matelea, we consider it best prac¬
tice to transfer the species in question to Matelea,
while leaving the remaining three Brazilian species,
lor which no immediate need for a name is known,
untouched for luture research. Chthamalia cuya¬
bensis was first described by Malme (1927), who
considered it closely related to C. majori E. Four¬
nier from Sao Paulo, from which it is said to differ
by smaller flowers and the form of the leaves. Both
characters may prove to be unreliable if more ma¬
terial comes to light, and it cannot be excluded that
C. cuyabensis might constitute a subspecies, a va¬
riety, or even only a synonym of C. majori. How¬
ever, currently available data do not permit such
conclusions.
Matelea cuyabensis (Malme) Liede & Meve,
comb. nov. Basionym: Chthamalia cuyabensis
Malme, Ark. Bot. 21A(12): 22. t. 1, f. 2. 1927.
TYPE: Brazil. Mato Grosso: Cuyabd, 12 Nov.
1902, G. (). A. N. Malme / / 2562 (lectotype,
designated here, G not seen).
The protologue is accompanied by an illustration
of the habit, flower, and pollinarium.
Literature Cited
Malme, G. 0. A. N. 1927. Asclepiadaceae mattogrossen-
sis. Ark. Bot. 21 A: 1-27.
Rapini, A. 2001. Asclepiadoideae (Apocynaceae) da cad-
eia do Espinhago de Minas Gerais, Brasil. Bob Bot.
Univ. Sao Paulo 19: 55-169.
Stevens, W. I). 1985. Asclepiadaceae. Pp. 228—241 in J.
Rzedowski & G. C. de Rzedowski (editors), Flora Fa-
nerogdmica del Valle de Mexico, Vol. 2. Instituto de
Fcologfa, Mexico, 1). F.
- . 1988. A synopsis of Matelea subg. Dictyanthus
(Apocynaceae: Asclepiadoideae). Ann. Missouri Bot.
Gard. 75: 1533-1564.
- . 2001. Asclepiadaceae. Pp. 234—270 in W. 1).
Stevens, C. Ulloa Ulloa, A. Pool & 0. M. Montiel (ed¬
itors), Flora de Nicaragua, Tomo I. Monogr. Syst. Bot.
Missouri Bot. Gard. 85.
Woodson, R. E. 1941. The North American Asclepiada¬
ceae. Ann. Missouri Bot. Gard. 28: 193-244.
Novon 14: 314. 2004.
Three New Speeies of Celastraceae (Hippocrateoideae) from
Southeastern Brazil, and a New Combination in Peritassa
Julio Antonio Lombardi
Departamento de Botanica, Instituto de Ciencias Biologieas, Universidade Federal de Minas
Gerais, Av. Antonio Carlos 6627, 31270-110, Belo Horizonte, Minas Gerais, Brazil.
cissus@mono.icb.ufmg.br
Abstract. The new Peritassa sadleri and Peri¬
tassa longifolia are distinguished by their habit,
leaf characteristics, color of flowers, shape of pet¬
als, number of ovules, size of inflorescences, and
size of fruits. The new species Salacia nemorosa is
distinguished from other southeastern Brazilian
species by inflorescences, disc details, and size anti
shape of flowers and fruits. The new combination
in Peritassa is also proposed: Peritassa glabra (A.
C. Smith) Lombardi.
KKSUMO. As novas especies Peritassa sadleri e
Peritassa longifolia sao distintas pelos habitos,
morfologia das folhas, cor das flores, forma das pe-
talas, numero de ovulos e tamanho das infloreseen-
cias e tlos frutos. Salacia nemorosa difere das out-
ras congeneres do sudeste do Brasil pelas
inflorescencias, detalhes do disco e tamanho e for¬
ma das flores e dos frutos. Tambem uma nova com-
binayao em Peritassa e proposta: Peritassa glabra
(A. C. Smith) Lombardi.
Key words: Brazil, Celastraceae, H'PI >ocrateo-
ideae, Peritassa , Salacia.
I he former family Hippocrateaceae is today in¬
cluded in the Celastraceae as a subfamily, which
includes 5 tribes, 24 genera, and ca. 357 described
species (Hal 1^, 1990; Mennega, 1997) distributed
worldwide in tropical and subtropical areas.
The delimitations of the Neotropical genera are
notoriously controversial, ranging from 2, Salacia
and Hippocratea (Peyritsch, 1878), to 17 by Miers’s
(1872) account. In this work I followed Smith
(1940), who recognized three genera segregated
from Salacia L., Cheiloclinium Miers, Peritassa
Miers, and Tontelea Aublet, included in the tribe
Salacieae with the African Salacighia Loesener and
Thyrsosalacia Loesener (Mennega, 1997).
The genus Peritassa is restricted to the Neotrop¬
ics with 14 species (Lombardi & Temponi, 1999;
Mennega, 1997). Peritassa was described by Miers
(1872) and segregated from Salacia by its anthers
with longitudinal dehiscence with a prominent con¬
nective and a tubular nectariferous disc.
Salacia is a genus of ca. 200 species (Mennega,
1997), distributed worldwide in the tropics and
subtropics. Smith (1940) recognized 29 species
from the Neotropics. The last comprehensive taxo¬
nomic study of the genus, although unpublished, is
Hedin (1999), who recognized 38 species in the
Neotropics, including 3 new laxa from Costa Rica,
Ecuador, and Peru.
The probable paraphyly of Salacia s. str. has
been pointed out in recent molecular studies (Sim¬
mons et ah, 2001a, b), but before a more extensive
analysis of worldwide representative species on
both morphological and molecular aspects, it is
premature to propose any generic realignments.
While working on a taxonomic study of Brazilian
species of Hippocrateoideae, as part of a study of
the Neotropical species, I discovered three new
species from the Espirito Santo Atlantic ridge (Ser-
ra do Mar): two in Peritassa and one in Salacia.
These newly described species occur in an area
with few forest fragments, remnants from a vege¬
tation continuous in the past hut almost totally de¬
pleted now in Espirito Santo.
Peritassa sadleri Lombardi, sp. nov. TYPE: Bra¬
zil. Espfrito Santo: Itarana, Alto Jatiboca, Faz.
Frederico Sadler, 20°00'48.6"S, 40°54'39.9"W,
786 m, 26 Feb. 2003, ,/. A. Ixtmbardi 5205. A.
Salino, R. C. Mota & L. G. Temponi (holotype,
BHCB; isotypes, MBM, MBML, MO, NY,
SPE). Figure 1.
Peritassae mexiae disco brevi-tubulari et antherarum
dehiscentia transversali-obliqua affinis, sod habitu, flomm
colore, petalis orbicularibus, ovulorum numero et fruc-
tuum forma et dimensione differt. Etiam Peritassae lon-
gifoliae hie descriptae affinis, sed habitu, pedicelli di¬
mensione et fructuum forma et dimensione differt.
Small tree, 4—5 m, glabrous, branches terete,
lenticellate, sulcate, smooth, with seasonal growth
marked by scale-like cataphylls. Leaves opposite;
stipules 0.4— 0.8 mm long, caducous, triangular.
No VON 14: 315-321. 2004.
316
No von
Figure 1. Peritassa sadleri Lombardi. — A. Lateral view of flower. — If. Branch with leaves and inflorescences. — C.
Lateral and adaxial views of stamens. — D. Lateral view of middle section of open flower showing disc, two stamens,
and pistil. Drawn from the holotype, Lombardi el at. 5205 (BHCB).
apiculate, minutely erase; petioles 0.4(— 0.8) cm,
canaliculate; Blades 5.9—7.2(14.9) X 1.4— 2. 8(4.1)
cm, elliptic, chartaceous, drying brawn, the base
euneate, the apex acute or acuminate, tip cleft or
apiculate, the margin entire and thickened, cream-
colored when dry, venation brochidodromous with
veins slightly prominulous on both sides. Inflores¬
cence thyrsoid-paniculate, 0.9— 1.4 X 1—1.5 cm,
with 28 to 49 (lowers, axillary, sometimes at leafless
nodes; bracts 0.7-1 .2 mm long, triangular, the mar¬
gin minutely erase, scarious; peduncle 0.1 — 0.3( — 0.7)
cm long, terete, in old leafy nodes covered by tri¬
angular scales, on new shoots sometimes at leafless
nodes and naked, then minutely lenticellate-punc-
tulate; inflorescence branches opposite or sub¬
opposite, terete; pedicels (1.2— ) 1.7 mm long, terete,
minutely punetulate, articulate at base; bracteoles
2, opposite at base of pedicel, triangular, 0.4 mm
Volume 14, Number 3
2004
Lombardi
Celastraceae from Southeastern Brazil
317
long. Flowers green, 3.7—4 mm wide at anthesis;
sepals 5, 1.2— 1.3 X 1—1.1 mm. elliptic, unequal,
± carnose, the margin minutely erose; petals 5,
1.5— 1.8 X 1.6— 1.7 mm, subcircular, ± carnose,
spreading at anthesis, the margin minutely erose;
disc annular, short-tubular, ea. 0.4 mm high, 0.2
mm thick, carnose, free from reproductive parts,
the margin entire; stamens 3, ca. 0.8 mm long, in
open flower exserted and reflexed, the filaments
flattened, the connective adaxially thickened, not
surpassing the thecae, the anthers ca. 0.17 X 0.42
mm, oblong, slightly emarginate at base, basifixed.
dehiscence extrorse by almost transverse apical
slits; pistil 0.6 X 0.63 mm. conic, ovary slightly 3-
lobed, 3-locular, with 2 subapical ovules in each
locule, style ca. 0.16 mm long after anthesis, cen¬
tral, terete, stigmas obscure. Mature drupes 4—6.8
X 2.6 X 2. 1 cm, ellipsoid, the epicarp crustaceous,
smooth, drying light brown-pruinose, minutely
cream-colored punctuate; seeds not seen.
Phenology. Collected with flowers and very
young fruits in February; collected with mature
fruits in October.
Distribution and habitat. Peritassa sadleri oc¬
curs at altitudes of 700—800 m in the rain forests
of Espfrito Santo, on the southeastern Atlantic coast
of Brazil. This area has many small properties with
old coffee plantations and pastures; each of these
maintains small portions of forest in varied degrees
ol preservation in order to fulfill federal government
legislation. However protected by environmental
laws, the conservation of this species is not sure
while it is known only from outside of official con¬
servation sites.
Etymology. The specific epithet of this species
is an homage to the Brazilian-Pomeranian Sadler
family who has preserved an apparently almost in¬
tact tract of forest at the intersection of many family
members’ properties.
This new species resembles Peritassa mexiae A.
C. Smith by the disc, stamens, and pistil, but can
be distinguished by the habit (small tree vs. liana),
color of dried leaves (brown vs. cinereous), smaller
inflorescences (0.9-1. 4 X 1-1.5 cm vs. (0.8— )2. 3—
5.0 X (0.8— )2.8— 7. 1 cm), color of flowers (green vs.
yellow), shape of petals (± circular vs. elliptic),
number of ovules in each locule (2 vs. 1), and big¬
ger fruits (4—6.8 X 2.6 X 2.1 cm vs. 2.4— 2.7 X
1.6— 2.2 X f.l cm). Peritassa sadleri is also very
similar to Peritassa longifolia here described: the
flowers are almost identical, but the two species
differ in habit (small tree vs. liana), size of the ped¬
icels (1.2— 1.7 mm vs. 2.9 mm), and by different
fruits (mature 4—6.8 X 2.6 X 2.1 cm, ellipsoid vs.
immature 3.4— 4.8 X 2. 8-4. 2 cm, subpyriform).
Paratypes. BRAZIL. Espfrito Santo: Brejetuba [in
sched. ‘Brejatuba’], Rancho Dantas, 22 Oct. 2000, G.
Hatschbach, M. Hatschbach & J. M. Silva 71523 (BHCB
1 2 J, MBM); Itarana, Jatiboca, Alto de Jatiboca, 3 km da
Vila Jatiboca, Fazenda da familia Seidler [sic], 19°51'S,
40°52'W, 14 Feb. 1999. E. M. NicLughadha 204 , A. Mel-
lo-Silva. B. L Stannard & M. C. Ass is (BHCB, K not seen,
MB ML, SPF).
Peritassa longifolia Lombardi, sp. nov. TYPE:
Brazil. Espfrito Santo: Santa Teresa, Nova
Lombardia, Bes. Biol. Augusto Ruschi, 750 m,
28 Nov. 2001, L. Kollmann 5055 , E. Hansen
& W. Pizziolo (holotype, MBML; isotype,
BHCB). Figure 2.
Peritassae mexiae disco brevi-tubulari et antherarum
dehiscentia transversali-obliqua affinis, sed foliomm for¬
ma, fforum colore, petalis orbicularibus, ovulorum numero
et fructuum dimensione differt. Ftiam Peritassae sadleri
hie descriptae affinis, sed habitu, pedicelli dimensione et
fructuum forma et dimensione differt.
Lianas, glabrous, branches terete, lenticellate,
sulcate, smooth. Leaves opposite; stipules not seen,
caducous; petioles 0.7— 0.9 cm, canaliculate; blades
7.5—17.7 X 1. 9-4.3 cm, elliptic, ehartaceous, dry¬
ing cinereous, opaque, the base cuneate, the apex
acuminate, tip apiculate, the margin entire and
thickened, venation brochidodromous with veins
slightly prominulous on both sides. Inflorescence
thyrsoid-paniculate, 1.1— 1.8 X 1. 1-1.8 cm, with
ca. 15 flowers, axillary; bracts 0.4— 0.6 mm long,
triangular, the margin entire; peduncle obsolete; in¬
florescence branches arising front an axillary tuft of
six triangular sc ales; branches opposite or alter¬
nate, terete; pedicels 2.9 mm long, terete, minutely
punctulate; bracteoles 2, opposite at base or middle
of pedicel, triangular, 0.3 mm long. Flowers green,
4.6 mm wide at anthesis; sepals 5, 0.8-0.9 X 1.2-
1.3 mm, triangular or elliptic, unequal, ± carnose,
the margin minutely erose; petals 5, 1.9— 2.1 X 2
mm, ± circular, ± carnose, spreading at anthesis,
the margin entire to minutely erose; disc annular,
short-tubular, ca. 0.3 mm high, 0.2 mm thick, ear-
nose, free from reproductive parts, the margin en¬
tire; stamens 3, ca. 0.7 mm long, in open flower
exserted and reflexed, the filaments flattened, the
connective adaxially thickened, darkened when
dry, not surpassing the thecae, the anthers ca. 0.22
X 0.48 mm, oblong, slightly emarginate at base,
basifixed, dehiscence extrorse by almost transverse
apical slits; pistil 0.6 X 0.7 mm, conical, ovary
slightly 3-lobed, 3-locular, with 2 subapical ovules
in each locule, style ca. 0.3 mm long after anthesis.
318
Novon
of stamens. Drawn from the isotype Kollmann 5055 et al. (BHCB).
central, terete; stigmas obscure. Mature drupes not
seen, immature drupes 3.4— 4.8 X 2.8— 4.2 cm, ±
pyriform, the epicarp crustaceous, smooth, drying
brown-pruinose, minutely alveolate; seeds ea. 5,
2.3 X 1 cm, reniform.
Phenology. Collected with dowers in Novem¬
ber; collected with immature fruits in April anil
May.
Distribution and habitat. Peritassa longifolia
occurs at altitudes of ca. 700 m in the rain forests
of Espfrito Santo, on the southeastern Atlantic coast
of Brazil, and is known only within the limits of the
Augusto Ruschi Biological Reserve, an area with
4000 ha of primary rain lorest. This area is one of
the lew intact and relatively large forest fragments
in Espfrito Santo. The Biological Reserve provides
protection for this species, although the population
status is unknown.
This new species resembles Peritassa mexiae by
the disc, stamens, and pistil, and the dried leaf
color and texture, but may be distinguished by the
blade apex (acuminate vs. acute), smaller inflores¬
cences (1.1— 1.8 X 1.1— 1.8 cm vs. (0.8— )2. 3— 5.0 X
(0.8-)2.8— 7.1 cm), color of flowers (green vs. yel¬
low), shape of petals (± circular vs. elliptic), num¬
ber of ovules in each locule (2 vs. 1), and bigger
fruits (immature 3.' 4—4.8 X 2.8— 4.2 cm vs. mature
2.4— 2.7 X 1.6— 2.2 X 1.1 cm). Peritassa longifolia
is also similar to Peritassa sadleri described herein,
with the almost identical flowers. However, the two
species are distinguished by their habit (liana vs.
small tree), by the bigger pedicels (2.9 mm vs. 1.2—
Volume 14, Number 3
2004
Lombardi
Celastraceae from Southeastern Brazil
319
1.7 mm), and by different fruits (immature 3.4— 4. 8
X 2.8— 4.2 cm, subpyriform vs. mature 4—6.8 X 2.6
X 2.1 cm, ellipsoid).
Paratypes. BRAZIL. Espfrito Santo: Sarila Teresa.
Nova Lombardia, Res. Biol. Augusto Rusclii, 8 May 2002.
R. R. Vervloet 230 & E. Hausen (BHCB, MBML); trilha
Roda D'Agua, 2 Apr. 2002, R. R. Vervloet 53, E. Hausen
& W. Pizziolo (BHCB, MBML).
Comments on the two Peritassa species described
herein. The transversally dehiscent anther locule
and inconspicuous connective group Peritassa lon-
gifolia and P. sadleri with P. calypsoides (Cambes-
sedes) A. C. Smith, P. rnexiae, and the former Ton-
telea glabra, whose combination in Peritassa is
here proposed; these five species stand apart from
remaining Peritassa species by their almost trans¬
verse anther dehiscence slits and connective not
exceeding thecae. However, an intermediary mor¬
phology is found in P hatschbachii Lombardi, with
its oblique anther dehiscence slits and a connective
which slightly or not at all surpasses the thecae.
Salaeia nemorosa Lombardi, sp. nov. TYPE: Bra¬
zil. Espfrito Santo: Santa Teresa, Res. Biol.
Santa Lucia, trilha do palmiteiro, 27 Jan.
2000, V. Demuner 621 & E. Hausen (holotype,
MBML; isotype, BHCB). Figure 3.
filter species austro-orientales Brasilienses inflorescen-
tia forma singulari distincta. Salaciae mosenii disci forma
similis, sed disci externa margine suleo praedito, floribus
maioribus et fructibus minoribus pyriformibus distincta.
frees, ca. 10—12 m, DBH 35 cm, glabrous,
branches terete, lenticellate, sulcate, smooth, with
seasonal growth marked by scale-like cataphylls.
Leaves opposite; stipules 0.6 mm, persistent, tri¬
angular, margin denticulate; petioles 0.6— 0.7(— 0.8)
cm, canaliculate; blades (8— )8.9— 10.6 X 3-4(— 4.9)
cm. elliptic, chartaceous, drying brown, the base
cuneate, the apex acuminate or acute, tip slightly
emarginate, the margin entire and thickened,
cream-colored when dry, venation broehidodromous
with veins slightly prominulous on both sides,
cream-colored when dry. Inflorescence thyrsoid-pa-
niculate, 1.8— 2.8 X 1—2.5 cm, with ca. 18 flowers,
axillary at leafless nodes; bracts 0.7-1. 3 mm long,
triangular, the margin erose; peduncle 0.8— 1.3 cm,
terete, all branches opposite, terete; pedicels (1.9— )2.9
mm long, terete, minutely punctuate; bracteoles 2,
opposite at base of pedicel, triangular, 0.7 mm long.
Flowers yellow, 4.7 mm w ide at anthesis; calyx syn-
sepalous, sepals 5, lobes 0.5 X 1 mm, triangular
or elliptic, coriaceous, the margin entire, involute;
petals 5, 2.2 X 1—1.2 mm, elliptic, ± chartaceous,
spreading at anthesis, the margin undulate; disc ca.
0.6 X 0.4 mm, conical, carnose, free from repro¬
ductive parts, the margin grooved, entire; stamens
3, ca. 1 mm long, at first appressed together around
pistil, after exserted and reflexed, the filaments flat¬
tened, the connective adaxially thickened, the an¬
thers ca. 0.17 X 0.4 mm, oblong, slightly emargin¬
ate at apex, basifixed, dehiscence extrorse by
oblique almost transverse apical slits; pistil 0.65 X
1 mm, conical, ovary 3-lobed, 3-locular, with 2 ax¬
illary ovules in each locule, style ca. 0.5 mm long
after anthesis, central, conical, 3-angled, stigmas
obscure. Immature drupes 5.4 X 3.5 cm, pyriform,
sometimes ribbed at base, the epicarp crustaceous,
drying brown-pruinose, minutely rugose; seeds I or
2. 2.8 X 2.3 X 1.5 cm, reniform.
Phenology. Collected with flowers in January;
collected with immature fruits in October.
Distribution and habitat. Salaeia nemorosa oc¬
curs at altitudes of ca. 650—850 m in the rain for¬
ests of Espfrito Santo, on the southeastern Atlantic
coast of Brazil. This taxon is known only from Santa
Teresa municipality, which includes the Santa Lu¬
cia and Augusto Rusclii Biological Reserves, areas
with 2500 and 4000 ha, respectively, of primary
and secondary rain forests inside the city limits.
The biological reserves provide adequate protection
for this species, which have reproductive popula¬
tions in different places in Santa Teresa.
This new' species is distinguished from all other
southeastern Brazilian species by its pedunculate
inflorescence; Salaeia elliptica (Martins ex Schul¬
tes) G. Don, Salaeia crassifolia (Martius ex Schul¬
tes) G. Don, and Salaeia grandifolia (Martius ex
Schultes) G. Don have fasciculate inflorescences,
while Salaeia mosenii A. C. Smith has chiefly ses¬
sile thyrsoid-paniculate inflorescences and Salaeia
arborea (Schrank) Peyritseh has simple or com¬
pound dichasia. It resembles Salaeia mosenii in its
conic disc, but it differs because the disc has a
flattened outer margin instead of a groove and larg¬
er flowers (4.7 mm vs. 2.4— 4.3 mm) and different
fruit shape (pyriform vs. spherical).
Paratypes. BRAZIL. Espfrito Santo: Santa Teresa,
Alto Sao Lourengo, Sftio da Cachoeira, 25 Oct. 2000, V.
Demuner 1490, E. Hausen & W. Pizziolo (BHCB, MBML);
Santa Teresa, Est. Biol, de Santa Lucia, 22 Sep. 1993, L.
I). Thomaz 1722 (BHCB, MBML); Santa Teresa, Res. Biol.
Augusto Ruschi, estr. Alto Goiapaba— Agu, 10 Jan. 2002,
L Kollrnann 5294 & E. Hausen (BHCB, MBML), 10 Jan.
2002, L. Kollrnann 5299 & E. Hausen (BHCB, MBML);
trilha da cachoeira, 29 Jan. 2002, /.. Kollrnann 5361 &
E. Hausen (BHCB, MBML); Santa Teresa, Nova Lombar¬
dia, Res. Biol. Augusto Rusclii, estr. Goiapaba— Agu, 24
Oct. 2002, R. R. Verrloet 1272, E. Hausen & W. Pizziolo
(MBML).
320
Novon
Figure 3. Solaria nemorosa Lombardi. — A. Lateral view of flower. — lb branch with leaves and inflorescences. — C.
Lateral and adaxial views of stamens. — D. Lateral view of middle section of open flower showing disc, two stamens,
and pistil. Drawn from the isotype Demuner 621 & Hausen (BHCB).
Iii addition to the aforementioned novelties, a
new combination is proposed.
IVritussa glabra (A. C. Smith) Lombardi, comb,
nov. Basionym: Tontelea glabra A. C. Smith,
Brittonia 3: 500. 1940. TYPE: Bolivia. La Paz:
San Carlos, Mapiri region, 850 m, 5 Feb.
1927, O. Buchtien 1728 (holotype, NY; iso¬
type, F).
Smith (1940) revised the Hippocrateoideae (as
Hippocrateaceae) from the Neotropics, recognizing
I 15 species, including Peritassa mexiae A. C.
Smith and Peritassa calypsoides (Cambessedes) A.
C. Smith. These species have an inconspicuous an¬
ther connective and oblique dehiscence clefts, al¬
most transverse, and are distinguished from other
fleshy-fruited genera (tribe Salacieae, Mennega,
1997) because the anther locules, although dehis¬
cent by transversal clefts, are not confluent at the
apex. The type of Peritassa calypsoides was not
studied hv Smith, who confused this species with a
previously undescribed Peritassa species, P.
hatschbachii Lombardi (Lombardi & Temponi,
Volume 14, Number 3
2004
Lombardi
Celastraceae from Southeastern Brazil
321
1999). Peritassa mexiae was described based on a
single specimen with detached “imperfect” flowers
found between the inflorescence branches (Smith,
1940: 518). I examined the isotype specimens (K
Mexia 5206, A, BM, F, GH, NY, VIC), and the
flowers referred to by Smith appear to be older
dried flowers that had fallen between the inflores¬
cence branches and remained on the sheet. At the
same time. Smith described Tontelea glabra A. C.
Smith (1940: 500), but did not perceive the simi¬
larity of this species with Peritassa mexiae. I went
through the types of Tontelea glabra and a number
of Amazonian collections from several herbarium
collections (CTES, F, IAN. I PA. MG, NY. R, RB,
S, U, UB, US) and did not encounter the anther
character of “confluent horizontal apical clefts"
(Smith. 1940: 501) used by the author for inclusion
of this taxon in Tontelea.
Tontelea in Smith’s (1940: 355) key is distin¬
guished from Peritassa by the dehiscence of anthers
by confluent slits, while Peritassa dehiscence was
by non-confluent slits. Almost all species in Ton¬
telea have oblong, relatively large anthers (0.2— 0.4
X 0.3— 0.6(0.8) mm) with clearly transverse dehis¬
cence slits and 3-lobed stigmas, alternate or op¬
posite with stamens, except for the aberrant species
Tontelea cylindrocarpa, T. myrsinoides, T. nectan-
drifolia, and T. glabra, which have relatively small¬
er anthers (0.1— 0.3 X 0.2— 0.3(0. 5) mm) and unlo-
bed stigmas. The transfer of T. glabra to the genus
Peritassa diminishes the morphological discrepan¬
cy within Tontelea ; the generic placement of the
other three divergent species remains to be studied.
Acknowledgments. The author thanks T. S. Fil-
gueiras for the Latin diagnoses, Victoria Hollowell,
Jon Ricketson, and Mark Simmons for critically re¬
viewing the manuscript, GNPq for the research fel¬
lowship grant (523026/96-0), and FAPEMIG for a
financial research grant (CBS-2080/96) as well as
the support of tin* author’s participation in the Bra¬
zilian Botanical Congress in 2003. I thank the cu¬
rators of the following herbaria for the loans of
specimens: A, BM, CTES, F, GH, IAN, IPA, MBM,
MBML, MG, MO, NY, R, RB, S, SPF, U, UB. US,
VIC. Drawings are by the author.
Literature Cited
Hall6, N. 1990. Cdlastracdes (Hippoerateoiddes). Pp. 1—
247 in 15. Satabie & P. Morat, Flore du Cameroun 32.
Ministdre de l’Enseignement Supdrieur de
l’lnformatique et de la Recherche Scientifique, Yaoun¬
de.
Hedin, J. P. T. 1999. Systematic Studies of the Neotropical
Species of Salacia L. (Hippocrateaeeae) and Its Rela¬
tives. Ph.D. Thesis, Washington University, St. Ixmis.
[Unpublished.]
Lombardi, J. A. & L. G. Temponi. 1999. A new species
of Peritassa Miers (Hippocrateaeeae) from southern Bra¬
zil, and notes on two confused species. Novon 9: 221—
226.
Mennega, A. M. W. 1997. Wood anatomy ol the llippo-
crateoideae (Celastraceae). IAWA .1. IB: 331—368.
Miers, J. 1B72. On the Hippocrateaeeae of South America.
Trans. Linn. Soc. London 2B: 319-432.
Peyritisch, J. 1878. Hippocrateaeeae. Pp. 125-164 in C.
F. P Martius & A. G. Eiehler, Flora Brasiliensis I 1(1).
Frid. Fleischer, Lipsiae.
Simmons, M. P., C. C. Clevinger, V. Savolainen, R. H.
Archer, S. Mathews & J. F. Doyle. 2-
05-014-87 from COLCIENCI AS to the Universidad
del Valle (Jorge E. Ramos-Perez. principal inves¬
tigator). Some of the collections were made during
class field trips financed bv the Departamento de
Biologfa of the Universidad del Valle.
I .iterature Cited
Calderon. E., (7 Galeano & N. Garcfa (editors). 2002.
Libro rojo de plantas fanerdgamas de Colombia, Vol. I.
Chrysobalanaceae, Dichapetalaceae y Lecythidaeeae.
Instituto Alexander von Humboldt, Institutode Ciencias
Naturales-Universidad Nat ional de Colombia, y Minis-
terio del Medio Ambiente, Bogota.
Greuter, W., J. McNeill, F. R. Barrie, II. M. Burdet, V.
Demoulin, T. S. Filgueiras, D. II. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & I). L. Hawk-
sworth (editors). 2000. International Code of Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Hailing. (7 1958. Monograph of the Cyelanthaceae. Acta
Horti Berg. 18: 1—428.
- . 1973. Cyelanthaceae. In: G. Harling & B. Sparre
(editors). Flora of Ecuador I: 1-48.
- & R. Eriksson. 1998. Cyelanthaceae. Pp. 471-
486 in J. A. Steyermark, P. E. Berry & B. K. Holst
(editors), Flora of the Venezuelan Guayana, Vol. 4. Mis¬
souri Botanical Garden Press, St. Louis.
Festuca dinirica and F. guaramacalana (Poaceae, Loliinae), Two
New Species from the Venezuelan Andes
Daniel Stancik
Department of Botany, Faculty of Science, Charles University, Benatska 2, 128 01 Prague,
Czech Republic, dan_stancik@yahoo.com
Abstract. Two new species, Festuca dinirica
Standfk and F. guaramacalana Stancfk (Poaceae,
Loliinae), from the Venezuelan Andes are de¬
scribed. Festuca dinirica is found in the paramo
zone and seems best placed in subgenus Festuca
sect. Aulaxyper ; Festuca guaramacalana is found
in humid mountain forests and belongs in Festuca
subg. Subulatae sect. Subulatae. Both species are
endemic to Venezuela, known from small areas of
Dinira National Park and Guaramacal National
Park, respectively.
Resumen. Se presentan dos especies nuevas,
Festuca dinirica Stancik and F. guaramacalana
StanCfk (Poaceae, Loliinae), endemicas de los An¬
des de Venezuela. La especie F. dinirica pertenece
al subg. Festuca sect. Aulaxyper y se conoce solo
de la zona paramuna del Parque Nacional Dinira.
La especie F. guaramacalana es un representante
del subg. Subulatae sect. Subulatae , eudemica de
la zona del bosque humedo altoandino del Parque
Nacional Guaramacal.
Key words: Andes, endemic. Festuca, Loliinae,
Poaceae, Venezuela.
The genus Festuca L. is a highly diversified cos¬
mopolitan grass genus, represented by over 140
species (Darbyshire et ah. 2003) in the Andes of
South America and one extra-Andean enclave in
southeastern Brazil and northeastern Argentina.
Revision of Festuca in Venezuela has resulted in
recognition of two new species. Festuca dinirica
and F guaramacalana. Type collections of these
new species come from recently organized field
studies in little known regions of Dinira National
Park (Stale Lara) and Guaramacal National Park
(State Trujillo), and represent first collections of
Festuca from these areas (for comparison, see Al¬
exeev. 1986; Briceno & Morillo, 1994; Dorr et ah,
2000; Vareschi, 1970). Both species appear to be
endemic to the Venezuelan Andes.
Festuca dinirica belongs in subgenus Festuca L.
sect. Aulaxyper Dumort. This section is represented
by only a few (probably 2 to 4) species in South
America and remains insufficiently known. The
geographically nearest species with clear affinity to
section Aulaxyper is the markedly distinct species
Festuca andicola Kunth from swampy habitats in
Colombia and Ecuador.
Festuca guaramacalana belongs in subgenus
Subulatae (Tzvelev) E. A I exeev sect. Subulatae
Tzvelev represented in Venezuela by two other spe¬
cies (F. ulochaeta Nees ex Steudel and F. coromo-
tensis Briceno). An artificial key for distinguishing
these species is provided. The survey of known
South American members of Festuca sect. Subula¬
tae was published recently (Stancfk & Peterson,
2002) and included eight species.
Festuca dinirica Stanfnk, sp. nov. TYPE: Vene¬
zuela. Lara: Munic. Humocaro Alto. NP Di¬
nira, 09°35'38.7"N, 70°07'12.2"W, 3170 m, 30
Nov. 2000, Standik 4287 (holotype, PRC; iso¬
types, CAR, COL). Figure If-j.
Haec species a Festuca tolucensi innovationibus mixtis
non intravaginalibus, ligula foliari 0.5 (non 1 .8—3.5) mm
longa, foliis tenuioribus 0.3— 0.5 (non 0.5— 0.7) mm latis,
glabris (non scabris), sclerenchymate abaxiali cum fasci-
culis vascularibus non junctis differt.
Tussocks, 50—60 cm tall; innovations mixed;
culms erect, glabrous, with 1 to 3 nodes in basal
half. Sheaths coriaceous-membranous, brownish
gray, glabrous, fibrous; ligule 0.5 mm long, shortly
ciliate; leaf blades linear, 20—30 cm long, 0.3— 0.5
mm wide, involute, pungent, glabrous, green. Pan¬
icle contracted, lanceolate, 6—10 cm long and 0.7-
1 cm wide; branches scabrous. Spikelets 8-10 mm
long, lanceolate, with 3 to 4 perfect florets; rachilla
scabrous or sparsely covered by bairs; glumes
membranous, upper Vs scabrid; lower glume 3.5—
4.2 mm long, lanceolate, acute, I -nerved; upper
glume 6-7 mm long, lanceolate, acute, 3-nerved;
lemmas 6—7 mm long, 5-nerved, membranous, lan¬
ceolate, scabrous, awned; awn 1-3 mm long; callus
glabrous; palea membranous, 2-carinate, scabrous,
markedly 2-dentate, as long as lemma; lodicules
lanceolate, 2-dentate; stamens 3, anthers 2—2.5 mm
Novon 14: 341-344. 2004.
342
Novon
Figure ]. a— e. Festuca guaramacalana Stanci'k (StanFik 4286, PRC). — a. Habit (bar: 9 cm). — b. Fragment of panicle
(bar: 32 mm). — c. Spikelet (bar: 14 mm). — d. Glumes (bar: 7 mm). — e. Cross section of leaf (bar: 0.4 mm), f— j,
Festuca dinirica Stani'fk (Stanfi'k 4287. PRC). — f. Habit (bar: 5 cm). — g. Panicle (bar: 2.5 cm). — h. Spikelet (bar:
10 mm). — i. Glumes (bar: 9 mm). — j. Cross section of leaf (bar: 0.3 mm).
Volume 14, Number 3
2004
Stancik
Species from Venezuelan Andes
343
long; ovary apex glabrous. Caryopsis lanceolate; hi-
lum linear, %—% of total. Leaf blade section with 5
vascular bundles and 3 to 5 ribs above; scleren-
chyma under abaxial and adaxial epidermis dis¬
continuous, vascular bundles free; adaxial epider¬
mis with microhairs about 0.09 mm long.
Observations. Spikelet structure in the new
species resembles Festuca tolucensis, another fre¬
quent species of the Venezuelan paramos. However,
F. dinirica differs clearly in organization of the tus¬
socks (small, with mixed innovations in contrast to
the big tussocks with intravaginal innovations of F.
tolucensis), in structure of sheaths (brown, striate,
and fibrous sheaths in contrast to the stramineous
and integrated sheaths), length of the ligule (0.5
mm vs. 1 .8— 3.5 mm), and in the anatomic structure
of the leaves (abaxial sclerenchyma isolated and all
vascular bundles free in contrast to the continuous
abaxial sclerenchyma and some vascular bundles
united with sclerenchyma).
The formation of the mixed innovations and
brown, striate sheaths (partly disintegrated into fi¬
bers) make Festuca dinirica close to section Aulax-
yper. In South America, this section is not very fre¬
quent and only poorly studied. The Colombian and
Ecuadorian species Festuca andicola is (geograph¬
ically) the nearest species with clear affinity to sec¬
tion Aulaxyper. Festuca andicola differs from F. di¬
nirica by growing in swampy areas, forming only
solitary culms with long rhizomes, and in having
distinctive spikelets (lower glume 1.4— 1.8 mm long
vs. 3.5— 4.2 mm; upper glume 2—2.5 mm long vs.
6—7 mm; lemma 5—5.5 mm long vs. 6—7 mm; awn
0.5— 0.7 mm long vs. 1—3 mm; anthers 0.8— 1.1 mm
long vs. 2-2.5 mm).
Distribution and habitat. Festuca dinirica is
endemic to Venezuela and known only from the
type locality — Dinira National Park in Lara. This
species forms small tussocks in shrubby-grassy pa¬
ramos, associated with Blechnum L., Espeletia Mu-
tis ex Bonpland, Hypericum L., Calamagrostis
Adanson, Chusquea Kunth, Rhynchospora Vahl,
etc., at an altitude of ca. 3200 m. No more infor¬
mation about the distribution of this species is
available. It is probably steno-endemic and vulner¬
able to extinction.
The specific epithet refers to the type locality,
Dinira National Park.
Paratypes. VENEZUELA. Lara: II umocaro
Alto. NP Dinira, 09°35'38.7"N, 70°07' 1 2.2"W,
grass paramo with shrubby patches, 3170 m, 30
Nov. 2000, Stancik 4289 (CAR, COL, PRC), Stan¬
cik 4288 (CAR, COL).
Key to the Venezuelan Species Festuca sect.
SUBULATAE
la. Lemma with awn markedly flexuous, 7—15 mm
long . 1 . F. ulochaeta
lb. Lemma awnless or with straight awn 0.5—1 mm
long . 2
2a. Spikelets 9—11 mm long, with 3 to 4 (lorets,
awn 0—0.5 mm long . 2. F. coromotensis
2b. Spikelets 15—17 mm long, with 5 to 7 florets,
awn 0.5-0. 1 mm long . . 3 . F. guaramacalana
Festuca guaramacalana Stancik. sp. nov. TYPE:
Venezuela. Trujillo: Bocono, PN Guaratnacal,
09°14'15. 1"N, 70°1 I '14.3"W. 2880 m, 29 Nov.
2000, Stancik 4286 (holotype, PRC; isotypes,
CAR, COL). Figure la-e.
Haec species a Festuca coromotensi ligula folii acuta
non truncata, 1 — 1.5 (non 0.5— 0.7) mm longa, spiculis 15—
17 (non 9—11) mm longis, lemmate 9.5—10.5 (non 7.5—
8.5) mm longo distinguitur.
Rhizomatous grass, forming small tussocks, 100-
130 cm tall; innovations extravaginal; culms erect,
glabrous, with 3 or 4 nodes in distal half. Sheatli
membranous-coriaceous, purplish brown, striate, fi¬
brous at base, margins lree; auricles absent; ligule
1 — 1.5 mm long, (coriaceous)membranous, acute;
leaf blades linear, involute, 30—40 cm long and 0.6—
0.7 cm wide, (olive-)green, glabrous. Panicle
branched, flexuous, pendent, elongate, 15—20 cm
long and 2—5 cm wide; branches finely scabrous.
Spikelets narrowly lanceolate, 15—17 mm long, with
5 to 7 perfect florets; rachilla pilose; glumes pur¬
plish, narrowly lanceolate, acute, coriaceous-mem¬
branous, sparsely scabrous; lower glume 3.7— 4.7
mm long, 1 -nerved; upper glume 5—7 mm long, 3-
nerved; lemmas coriaceous-membranous, lanceo¬
late, 9.5—10.5 mm long, 5-nerved, purplish, apex
entire, papillose, mucronate or shortly awned, the
aivn 0.5—1 mm long; callus sparsely pilose; palea
two-carinate, finely pilose, almost as long as lemma,
apex shortly two-dentate; lodicules ovate, two-den¬
tate; stamens 3, anthers 2.5— 2.8 mm long; ovary
apex glabrous. Caryopsis lanceolate; hilum linear, K-,
of total length. Leaf blade cross section with ca. 1 I
vascular bundles and 5 ribs above; sclerenchyma
under both abaxial and adaxial epidermis discon¬
tinuous; unit with some vascular bundles forming
girders, bulliform cells absent; adaxial epidermis
with scattered microhairs, 0.09 mm long.
Observations. This new species belongs to the
section Subulatae, and among the South American
members of this section Festuca guaramacalana
has the largest spikelets and longest glumes (Stan-
Cfk & Peterson, 2002). Similarly, as in the case of
F. sodiroana Hackel ex E. B. Alexeev and versus
344
Novon
typically long awns elsewhere in section Subulatae,
the lemma is only shortly awned (0.5—1 mm). Mor¬
phologically, Festuca coromotensis B. Briceno ap¬
pears to he the most similar with truncate (vs.
acute), shorter ligules (0.5— 0.7 mm long vs. 1-1.5
mm), shorter spikelets (9—11 mm long vs. 15—17
mm) with 3 to 4 (vs. 5 to 7) florets, and a shorter
lemma (7.5— 8.5 mm long vs. 9.5—10.5 mm).
Distribution and habitat. This species is en¬
demic to Venezuela. It is known only from the type
collection from the National Park Guaramaeal (Tru¬
jillo), where it occurs at the margin of a brook with
Neurolepis Meisner and Cortaderia Stapf in Andean
mountain forest, at about 3000 m. This species
probably occurs rarely, as no additional exemplars
were found in the field or in herbarium collections.
It is possibly a steno-endemic species vulnerable
to extinction.
The specific epithet refers to the type locality
region, Guaramaeal National Park.
Acknowledgments. I thank the staff of all the
herbaria visited (AAU, B. C, CAR, COL, F, HFRZ,
MER, MERC, MERF, MY. PORT, PRC, U. US.
YEN, W). Thanks are due to Silvia Llamozas
(YEN), B. Briceno (Universidad de los Andes, Me¬
rida), and Giuseppe Colonnello (CAR) for their spe¬
cial help and assistance. The investigation was fi¬
nancially supported by a grant from the Czeeh
Grant Agency GACR (No. 206/03/0197).
Literature Cited
Alexeev, E. B. 1986. Festuca I.. (Poaceae) in Venezuela,
Colombia A Ecuador. Novosti Sist. Vyssli. Hast. 23: 5—
23. [In Russian.]
Briceno, B. & G. Morillo. 1994. El genero Festuca I.. (Po¬
aceae) en los paramos de Merida. Ernst ia 4(3-4): 73—
88.
Darbyshire S. J.. B. J. Soreng, I). Stanffk & S. I). Koch.
2003. Festuca. In: R. J. Soreng, G. Davidse, P. M. Pe¬
terson, F. 0. Zuloaga, E. J. Judziewicz, T. S. Filgueiras
& (). Morrone (editors), Catalogue of New World Grass¬
es (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl.
Herb. 48: 1-730.
Dorr, L. J., B. Stergios, A. R. Smith & N. L. A. Cuello.
2000. Catalogue of the Vascular Plants of Guaramaeal
National Park, Portuguesa and Trujillo States, Venezue¬
la. Contr. U.S. Natl. Herb. 40: 1-155.
Stanffk, I). & P. M. Peterson. 2002. Two new species in
Festuca from South America (Poaceae: Loliinae: Sect.
Subulatae). Sida 20(1): 21-29.
Vareschi, V. 1970. f lora de los pdramos de Venezuela, pp.
404—407. Universidad de Los Andes, Merida. Venezue-
Sarocalamus , a New Sino-H imalayan Bamboo Genus
(Poaeeae: Bambusoideae)
Chris M. A. Stapleton
l he Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 BAR,
United Kingdom, c.stapleton@rbgkew.org.uk
Grainne Nf Chonghaile and Trevor R. Hodkinson
Department of Botany, University of Dublin, Trinity College, Dublin 2, Ireland.
hodkinst@tcd.ie
Abstract. Sarocalamus Stapleton, a new genus
distributed from the eastern Himalayas to the Sich¬
uan Province of China, is described. Among all
Asian bamboo genera it shares the greatest number
of morphological characters with the North Ameri¬
can genus Arundinaria Michaux. However. DNA
sequencing has suggested that other genera arc
probably more closely related to Arundinaria, while
Sarocalamus appears instead to be a possible sister
group to PhyUostachys Siebold & Zuccarini. a ge¬
nus with some similarities in branching but very
different flowers. Biogeograph ical implications arc
discussed, and the new combinations Sarocalamus
racemosus (Munro) Stapleton, S. faberi (Rendle)
Stapleton, and .S'. spanostachyus (T. P. Yi) Stapleton
are made.
key words: Arundinaria , Bhutan, biogeography,
China. India. Nepal, North A meriea, Sarocalamus.
Characterization of the Asian temperate woody
bamboos for the Flora oj Bhutan (Stapleton, 1994,
2000) and the forthcoming English-language ver¬
sion of the Flora of China has established that
three species of Sino-Himalayan bamboo are mor¬
phologically most similar to the geographically dis¬
tant type species of Arundinaria Michaux, A. gi-
gantea , from the southeastern United States (Texas
to Maryland). One species , Arundinaria racemosa
Munro, is from eastern Nepal, the Indian States of
West Bengal and Sikkim, and Bhutan. The others,
Arundinaria jaheri Rendle and Bashania spanos-
tachya E P. \i, are from isolated mountains in
southern Sichuan Province, and northeastern Yun¬
nan Province, Southwest China. Bashania fansi-
panensis T. (,). Nguyen from northern Vietnam may
also belong in this genus, but is not well known.
Apart from Sarocalamus, all other Asian bam¬
boos can be separated clearly from the type species
of Arundinaria on the basis of floral or vegetative
morphological characters. This includes species in
the genus Pleioblastus Nakai, which is still fre¬
quently synonymized within Arundinaria but differs
in its fused prophyll margins (Stapleton, 1997). Al¬
though more than 500 species names have been
pu blished in Arundinaria , only four, poorly known
Asian species do not already have combinations in
other genera and a different, modern placement
(Ohrnberger, 1999). More than 30 Asian genera
have been described by Japanese, Chinese, and
European botanists to accommodate species origi¬
nally described in Arundinaria, and the number of
genera currently recognized in this group ranges
from 19 (Li, 1997) to 28 (Wang, 1997). In a mod¬
erate treatment Ohrnberger (1999) included 23
genera. Although the circumscription of Arundi¬
naria has remained controversial, with the number
of Asian species included in the genus by different
authors ranging from 16 (Chao & Chu, 1981) to
none (Wang, 1997; Keng & Wang, 1996; Ohrnber¬
ger, 1999), it is clear that a large number of tem¬
perate Asian genera an; now recognized as being
(list inct from that genus.
Asian bamboos morphologically closest to the
North American type species, Arundinaria gigan¬
tea, such as species of Sarocalamus, Pleioblastus,
and Bashania P. C. Keng & T. P. Yi, arc often still
included in Arundinaria (Clayton & Renvoize,
1986; Chao & Renvoize, 1989; Ei, 1997). However,
molecular evidence from ITS, trn\ . intron, trn\,~ F
intergenic spacer, and rpl 16 intron DNA sequence
data (Zhang, 1996; Nf Chonghaile, 2002) has sug¬
gested that such species are not necessarily its
closest relatives. Instead, Pseudosasa japonica
(Steudel) Nakai has been suggested as the closest
Asian relative of Arundinaria gigantea from both
ndhV and rplAG sequence data (Zhang, 1996; Nf
Chonghaile, 2002), a clade comprising A. gigantea
and P. japonica receiving 86% BS (Nf Chonghaile,
2002), even though P. japonica has never been eon-
Novon 14: 345-349. 2004.
346
Novon
sidered a particularly close relative of A. gigantea
on morphological grounds. Pseudosasa Nakai lias
received almost universal recognition (McClure,
1973; Clayton Si Renvoize, 1986; Chao, 1989; De-
moly, 1996; Keng & Wang, 1996).
Although Sarocalamus shares the principal dis¬
creet macro-morphological character states with
Arundinaria, it still differs in several, more quan¬
titative characters. These include shorter, more cy¬
lindrical. more separated and less imbricate florets
of more consistent size, less than 10 X 2 mm; a
more sinuous rhachilla with much longer inter¬
nodes; completely glabrous pedicels; and more del¬
icate, smoother, rugose rather than ribbed, less
closely spaced leaf blades. The three species of
Sarocalamus have been placed in four other Asian
genera when not included in Arundinaria : Fargesia
Franchet, Yushania P. C. Keng, Bashania, and Gel-
idocalamus T. H. Wen. Sarocalamus differs quite
profoundly from Fargesia and Yushania in having
pachymorph rather than leptomorph rhizomes. It is
closer to Bashania and Gelidocalamus. However,
while those genera do have leptomorph rhizomes,
they also have decurrent, strongly pulvinate synllo-
reseence paraelades, and a much greater degree of
compression of the basal internodes on their central
branches, giving a higher number of branches at
each culm node. Bashania also has thickened,
prominent, long-ciliate prophyll keels, greater
branchlet ramification and tougher leaves, while
Gelidocalamus has much smaller florets, com¬
pressed rhachillas, swollen supra-nodal ridges, sol¬
itary, more lanceolate leaves, and its 7 to 12 sub¬
equal branches do not re-branch at all.
Molecular evidence is generally lacking at the
generic level in Asian woody bamboos, with low or
absent support for many well-recognized taxonomic
groupings. Combined analyses of data from differ¬
ent genes is starting to provide slightly better res¬
olution, however. What support has been shown at
the generic level is generally congruent with mor¬
phological distinctions and current taxonomic treat¬
ments of Asian bamboos. In the most comprehen¬
sive study of woody bamboos (Nf Chonghaile,
2002), one species of Sarocalamus was included
along with Arundinaria gigantea, species of Ba-
shariia, Fargesia, Yushania, and 33 other genera.
While the two species of Bashania, B. fargesii (E.
C. Camus) P. C. Keng Si T. P. Yi and B. qing-
chengshanensis P. (7 Keng Si T. P. Yi, formed a
fairly well-supported elade (70% RS in parsimony
analysis, 82% RS in N.l analysis), Sarocalamus fa-
heri (included there as Bashania fangiana (E. (7
Camus) P. C. Keng & T. H. Wen) was not resolved
within the Bashania elade and did not associate
with any other genera in parsimony analysis. It re¬
mained a largely unresolved member of the well-
supported monophyletic group of mainly Asian
temperate bamboos first reported by Watanabe et
al. (1994). It is now known to include 25 or more
genera from Africa, Asia, and North America
adapted to subtropical or temperate habitats in both
hemispheres (Nf Chonghaile, 2002). hut is widely
referred to as the “north temperate elade.”
The generic name Sarocalamus means broom
bamboo and is derived from saron (trapov: broom,
sweepings) and calamus (Ka\ap.o£: reed, bamboo).
It alludes to the very erect branching, reminiscent
of a traditional broom or besom, and the frequent
use of these bamboos for sweeping.
Sarocalamus Stapleton, gen. nov. I \ PE: Saroca¬
lamus racemosus (Munro) Stapleton.
Bashaniae et Arundinariae rhizomatibus tenuibus,
staminibus 3, inflorescentia ebracteata, gemmis apertis
similis, ab ilia ramis inflorescentiae erectis, intemodiis
ramorum longioribus, foliis tenuibus, ab hac flosculis an-
gustis, ab uterque pedicellis glabris, differt.
Similar to Bashania and Arundinaria in its lep¬
tomorph rhizomes, 3 stamens, ebracteate inflores¬
cences and open buds. Differing from the former in
its erect inflorescence branches, longer branch in¬
ternodes and thinner leaves, from the latter in its
narrower florets, and from both in its glabrous ped¬
icels.
Temperate self-supporting bamboo; rhizomes
leptomorph; culms tillering (plurieaespitose), erect
to drooping; internodes smooth, terete; nodes not
swollen, unarmed; supra-nodal ridge well devel¬
oped; branch buds lanceolate, on promontory, en¬
closed within single, 2-keeled prophylls, always
open at the front; branches initially I to 3, very
erect, appressed, sulcate, basal internodes progres¬
sively increasing in length, basal internode often
long, lateral branching often distant from culm;
complement proliferating to become broom-like, at¬
tenuate; lateral branch axes always subtended by
sheaths, no replication of lateral branches. Leaf
blades linear-lanceolate, thin, less than 12 cm long,
with prominent cross-veins (tessellate). Synfloresc-
enee initially terminal or lateral to leafy branches,
later in leafless complements; synflorescenee
branches glabrous, without pulvini, remaining very
erect and appressed, branching racemose or rarely
partially paniculate, not fasciculate, subtended by
very small remnants of sheaths or lings of hairs,
rarely with very small pulvini. Spikelets 3 to 6 per
synflorescenee, on glabrous promontory (pedicel);
prophyll represented by lower glume; glumes 1 to
2, both small, lower glume usually very small, usu-
Volume 14, Number 3
2004
Stapleton et al.
New Sino-Himalayan Bamboo
347
ally distant from lower lemma, without subtended
buds (flowering semelauctant); rhachilla often sin¬
uous, strongly flattened, more than half length of
palea; palea keels curving, appressed to rhachilla;
stamens 3; stigmas 3. Flowering gregarious.
Distribution. East Nepal, India (West Bengal &
Sikkim), Bhutan, Western China (Tibet, Sichuan
and Yunnan Provinces), North Vietnam, 2800-
3900 m.
1. Sarocalamus racemosus (Illunro) Stapleton,
comb. nov. Basionym: Arundinaria racemosa
Munro, Trans. Linn. Soc. London 26: 17. 1868,
emend. Gamble in Kew Bull. Misc. Inform.
1912: 138. 1912. Fargesia racemosa ( Munro)
T. P. Yi, J. Bamboo Res. 2(1): 39. 1983. Yu-
shania racemosa (Munro) R. B. Majumdar, in
S. Karthikeyan et al., FI. Ind. Enumerat.—
Monocot.: 283. 1989. TYPE: Nepal/Sikkim
(India). Islumbo Pass |Chiya Banjyang], 13
Oct. 1857, Thomson s.n., Kew distribution
number (K 6738) (lectotype, selected by Gam¬
ble (1912: 137). K).
Arundinaria racemosa Munro was first described
from collections of two species from the eastern
Himalayas (Munro, 1868). It was emended by Gam¬
ble (1912), who separated the lower altitude col¬
lections from 6000 to 9000 ft. in the vicinity of
Darjeeling into a new species, Arundinaria moling ,
now known as Yushania moling (Gamble) R. B. Ma-
jumdar, leaving only the “high-level form,” found
above 10.000 ft., in Arundinaria racemosa. The
Munro syntypes were discussed in detail by Gam¬
ble (1912: 137) when describing his new species.
He stated “it seems quite evident that Thomson’s
Islumbo Pass flowering specimen is the real type
of A. racemosa.'' This can be taken as an effective
lectotypification under Article 7.1 1 of the St. I /mis
Code (Greuter et al., 2000), as a type* element was
indicated by direct citation including the term
“type.” Chao and Renvoize (1989) later designated
Munro’s first syntype, a Thomson collection made
in 1857 from 6000 ft. on Birch Hill in Darjeeling,
as lectotype of A. racemosa. They apparently did
not realize that the species had already been lec-
totypified by Gamble, even though they concur¬
rently lectotypified ,4. moling , in the protologue of
w hich Gamble had made his lectotypification of A.
racemosa. According to the label of Thomson’s
Birch H ill collection, the lectotype designated by
Chao and Renvoize was a “pale blue” bamboo. Two
bamboos from Darjeeling, A. moling and tin* rarer
/ / i m alayacal amus hookerianus (Munro) Stapleton,
both have pale blue culms, w hile culms of A. ra¬
cemosa are green or yellow. Gamble (1912: 137)
had even stated that “Birch Hill is the end of the
Darjeeling Ridge . . . and is a very unlikely locality
for the high-level A. racemosa." However, as Gam¬
ble (1912: 137) had also pointed out, Thomson’s
collection of some other, pale blue bamboo truly
from Birch Hill would seem to have become lost,
and the Birch Hill label is currently attached to a
sheet bearing only a flowering specimen from Is¬
lumbo Pass, which bears the correct label as well.
It would appear, therefore, that Chao and Renvoize
1 1989), by simply designating Munro’s first syntype
as lectotype, may have inadvertently designated a
long-lost collection of A. raiding or Himalayacala-
mus hookerianus from Darjeeling as lectotype of A.
racemosa, but fortunately there can be no doubt
that the earlier lectotypification of Gamble (1912)
takes priority. The Islumbo Pass material shows the
distinctive branching and synflorescence of Saro¬
calamus and is accepted here as lectotype of A.
racemosa Munro and type of the genus Sarocala¬
mus. It was collected by Thomson in 1857 in woods
above the Islumbo Pass, now known in Nepali as
Chiya Banjyang, on the Singalila Range, which
separates East Nepal and Sikkim. The name Arun¬
dinaria racemosa is still often misapplied to Arun¬
dinaria moling Gamble in India (Tewari, 1993;
Seethalakshmi & Muktesh Kumar, 1998), even
though that species has been transferred by the Bo¬
tanical Survey of India as Yushania moling (Gam¬
ble) R. B. Majumdar.
Habitat and distribution. Sarocalamus racemo¬
sus is found at an altitude of 2900-3800 m in the
understory of Abies densa and mixed coniferous and
rhododendron forests, in degraded forest, and in
cleared areas of pastureland, in which il can be the
principal species. It is often found on more freely
drained sites than Yushania microphylla, which has
hollow rhizomes and extends to a higher altitude.
The type collection was made on a pass that con¬
stitutes the border between Nepal and Sikkim. Fur¬
ther collections have been made in Sikkim, where
it was reported to be common in forests around Sen-
clial (Gamble, 1896), and across western, central,
and eastern Bhutan. It is to be expected that it will
also be found in further localities in the eastern¬
most districts of Panchtar and Taplejung in Nepal.
It has also been reported from Tibet (Yi, 1983) and
is to be expected in the Indian State of Arunachal
Pradesh.
Additional specimens examined. INDIA. Sikkim: Sin¬
galila Range, Sukumbum, May 1889, l)r. dummies Col¬
lectors s.n. (K). BHUTAN. Puro: Chile La, 1 June 1971,
Bedi 32 (K); Thimphu: Dorchula, 22 Dec. 1989 Stapleton
872 (TH1M). Tongsa: Yotong La, 15 Aug. 1989, Stapleton
348
Novon
832 (TIUM). Hiiiuthniig: Lame Gompa, 23 June 1990,
lliirgi s.n. (K, THIM). Mongar: Thrumsingla, 11 Oct.
1990, Wood 7348 (E). Trashigung: Taahiyangtse, E side
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in K. Kubitzki (editor), The Families and Genera of
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Hutchinson, J. 1959. Peliosanthes. P. 604 in The Families
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Jessop. J. P. 1976. A revision of Peliosanthes (Liliaceae).
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Krause, K. 1930. Mondoideae. Pp. 376-377 in L. Diels
(editor), A. Engler & K. Prantl, Die natilrlichen Pflan-
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Larsen, K. 1966. Two new Liliaceae from the Khao Yai
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Mabberley, D. J. 1987. Neolourya. P. 394 in The Plant-
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Rodriguez, L. 1934a. Homodrae^es nouvelles d’Indochine.
Bull. Mus. Hist. Nat. (Paris), Ser. II. 6: 95-97.
- . 1934b. HOnodrac^es. Pp. 654—673 in F. Gag-
nepain (main editor), Flore G£n£rule de l’Indo-Chine,
Vol. 6. Masson, Paris.
Tanaka, N. 1999. Taxonomic notes on Peliosanthes (Con-
vallariaceae) I. Acta Phytotax. Geobot. 50: 147—155.
Yamashita, J. & M. N. Tamura. 2000. Molecular phylog-
eny of the Convallariaceae (Asparagales). Pp. 387-400
in K. L. Wilson & D. A. Morrison (editors), Monocots:
Systematics and Evolution. CSIRO. Melbourne.
INew Combinations in the Florida Flora II
Daniel B. Ward
Department ot Botany, University of Florida, Gainesville, Florida 32611. U.S.A
ABSTRACT. New combinations are made for the
following species and varieties within the flora of
Florida: Andropogon elliottii var. stenophyllus, Ar-
noglossum ovatum var. lanceolatum, Dalea albida,
Dalea gracilis, Eupatorium album var. petaloideum,
Harrisia eriophora var. fra grans, Harrisia gracilis
var. aboriginum, Harrisia gracilis var. simpsonii,
Houstonia procumbens var. hirsuta. Imperata cylin-
drica var. mexicana, Liatris pauciflora var. secunda,
Pisonia discolor var. floridana, Pityopsis nervosa
var. tracyi, Tephrosia virginiana var. mohrii, Yucca
flaccida var. smalliana.
Key words: Agavaceae, Andropogon , Arnoglos-
sum, Asteraceae, Cactaceae, Compositae, Dalea,
Eupatorium, Fabaceae, Gramineae, Harrisia, Hous¬
tonia, Imperata, Leguminosae, Liatris, Nyctagina-
ceae, Pisonia, Pityopsis, Poaceae, Rubiaceae, Te¬
phrosia, Yucca.
As compilation of Florida’s vascular plant inven¬
tory creeps toward completion, taxa continue to be
encountered for which no currently accepted name
is available. These taxa — all of them familiar to an
earlier generation of botanists, and all of them rec¬
ognizable in the field — have been orphaned and left
nameless by transfer of their generic or specific
name to synonymy elsewhere. Other taxa, though
not wholly abandoned, have been placed by forced
adoption with genera or species to which they do
not comfortably belong.
This listing is a continuation of an effort to re¬
store species and infraspecific taxa to ranks appro¬
priate to their morphological differences (cf. Long,
1970; Ward, 2001; Wunderlin & Hansen, 2001).
Such an effort is, of course, a necessary component
of taxonomic revision, where judgment has changed
as to proper generic/specific alignments and names,
and where subordinate taxa must either be renamed
or abandoned. The number of these unnamed or
misnamed subordinate taxa is surprisingly large. It
is common practice for persons who undertake tax¬
onomic revisions to reduce to synonymy the less
well-defined subordinate taxa that had been rec¬
ognized by previous workers. Such practice is per¬
haps commendable, for mindless, wholesale trans¬
fer of these lesser taxa would burden the literature
with innumerable unneeded names and obscure the
distinctions that the revisionist more fully under¬
stands.
But when a student undertakes reconsideration
of a genus of appreciable size and cannot be as
familiar as he might wish with the plants in the
field, he may omit those lesser taxa of which he is
unsure but that local botanists may recognize. Or
he acknowledges the lesser taxon but assigns it a
rank that is inappropriate to its characteristics.
The Florida flora, rich both in number of species
and in entities that merit infraspecific rank, is un¬
dergoing rapid change, not only in the arrival of
species new to the state and disappearance of its
natives, but change in the names of long-familiar
taxa. At times these name changes are unavoidable,
for agreed-upon new understanding of specific/ge¬
neric alignments requires new names to reflect the
new status.
As previously (Ward, 2001), in each of the fol¬
lowing transfers earlier authors have treated the
transferred taxon as worthy of recognition; no new
taxa are proposed. And where generic realignments
have been proposed and are here accepted, ac¬
knowledgment is thereby given to the merit of those
changes. As before, though the International Code
of Botanical Nomenclature (Greuter et al., 2000)
permits both subspecies and variety to be employed
at infraspecific levels, preference is expressed here
for “varietas” as the sole infraspecific rank. Types
of the taxa discussed here have not been seen ex¬
cept where so indicated.
At bottom, the determination of rank does not
depend solely on explicit morphological criteria.
More critical, if unquantifiable, is the experience
of a writer to equate new data sets with those with
which he is more familiar, the better to develop and
maintain a stable, internally consistent nomencla¬
ture ranking. The present effort is intended as a
step in that direction.
Always, one is reminded of the well-loved phrase
of J. M. Greenman (Anderson, 1969), surely true
also of lesser rankings, that “Species are but judg¬
ments.” It is hoped that the judgments expressed
here may find support.
Novon 14: 365-371. 2004.
366
No von
Acavaceae
Yucca flaccida Haworth var. smalliana (Fernald)
I). B. Ward, comb, et stat. nov. Basionym: Yuc¬
ca smalliana Fernald, Rhodora 46: 8. 1944.
TYPE: U.S.A. Florida: Duval Co., sandy soil,
near Jacksonville, May [1893?], Curtiss 2950
(holotype, GH; isotype, FLAS).
As prescient ly noted by Fernald (1944: 5), Yucca
is “always a baffling genus to work with from her¬
barium material.” He was confronted with just that
challenge when he addressed the nomenclature of
the Yucca familiar to him in eastern Virginia. His
decision appears unchallenged, that the plant
known to Small (1933) as Y. concava Haworth rep¬
resents the true Y. Jilamentosa L.
Two taxa were involved, set apart from other east¬
ern species of the genus by fibers that peel back
from the leaf margins (well described by Godfrey,
1988). Though at no point does Fernald refer to
these distinctive marginal fibers, they were appar¬
ent to early authors; Linnaeus (1753) cited the
Gronovius phrase-name: Yucca foliis lanceolatis ac-
uminatis integerrimis rnargine filamentosis. With
Virginia the source of the Gronovius material, and
with no other Yucca native in that area, there is no
reason to doubt the Virginia plant known to Fernald
is the same as that known to Linnaeus.
Small (1933), however, had applied Yucca jila¬
mentosa to a similar but more southern plant, one
only sparingly extending north to the Carolinas. By
Femald’s transfer of Y. Jilamentosa to his Virginia
plant, he left the more southern plant without a
name. He then chose a Curtiss collection from
Jacksonville, Florida, as representative, and named
it Yucca smalliana in acknowledgment of J. K.
Small, who he believed had done much to elucidate
these southeastern yuccas.
Though Fernald (1944) discussed in detail the
differences between his Virginia Yucca Jilamentosa
and the Florida type of his E smalliana, his unfa¬
miliarity in the field w ith the second, more southern
plant caused him to overlook an aspect difference
between these plants that readily permits them
when in flower to be distinguished even at a dis¬
tance. The robust rosette of basal leaves and stocky,
(loriferous inflorescence of Y. Jilamentosa has made
it a popular ornamental of old gardens and ceme¬
teries. It was carried as a grave marker by the
Scotch-Irish and German pioneers who moved out
of Virginia into early Kentucky, then a generation
later into the newly available Indiana territory and
throughout the Midwest (ef. Deam, 1940: 316). The
southern plant, in contrast, with its tall, mostly na¬
ked scape and relatively few flowers, is little cul¬
tivated. Fernald also understated the minimal (here,
varietal) differences between his new E smalliana
and a third entity recognized by Small, Y. flaccida
Haworth.
Both of Fernald’s taxa were recorded for the Car¬
olinas by Aides (1964. 1968). He accepted the as¬
signment of Yucca Jilamentosa. But he chose to treat
the second entity at varietal level, as Y. Jilamentosa
var. smalliana (Fernald) Ahles. Since Ahles later
(1968) placed Y. flaccida in synonymy, it may be
argued he erred in using smalliana, for there are
prior varieties of E flaccida, which automatically
establish the prior varietal autonym Jlaccida (cf.
Art. 26.3, Greuter et al., 2000). But Ahles’s new
combination was valid when published (1964) since
no synonymy was then listed.
Yet Femald’s selected type of Yucca smalliana, by
description, by examination of a duplicate of Cur¬
tiss’s collection (F’LAS 46959), and by its location
far to the south of known natural E Jilamentosa,
readily falls within E flaccida s.L, a judgment sup¬
ported by Duncan and Kartesz (1981) and Godfrey
(1988). With priority established al varietal level by
variety smalliana (Fernald) Ahles, Femald’s new en¬
tity is properly transferred to E Jlaccida.
A two-part caveat is in order, however. Because
Haworth’s specimens of Yucca Jlaccida are not ex¬
tant, the validity of the new combination rests on
the presumption (and probability) that Haworth’s
material would now be treated as representative of
E Jlaccida s. str., as presently recognized. Validity
requires also that the several varieties described
under E Jilamentosa anti E flaccida by Trelease
(1902) and others either fall within Y. Jilamentosa
or, if E smalliana, outside the parameters of variety
smalliana.
Cactackae
Harrisin eriophora (Pfeiffer) Britton var. Iragrans
(Small) I). B. Ward, comb, et stat. nov. Basio¬
nym: Harrisia fragrans Small, in Britton &
Rose, The Cactaceae 2: 149. 1920. TYPE:
U.S.A. Florida: St. Lucie Co., hammock on sand
dune 6 mi. S of Ft. Pierce, 20 Dec. 1917, ./. K.
Small 8457 (holotype, NY; isotypes, GIL US).
Harrisia gracilis (P. Miller) Britton var. aborigin-
um (Small) D. B. Ward, comb, et stat. nov.
Basionym: Harrisia aboriginum Small, in Brit¬
ton & Rose, The Cactaceae 2: 154. 1920.
TYPE: U.S.A. Florida: Manatee Co., ham¬
mock, Terra Ceia Island, 29 Apr. 1919, ./. K.
Small, Cuthbert & DeWinkler s.n. (lectotype,
designated by Benson (1982: 934), NY;
isolectotype, US).
Volume 14, Number 3
2004
Ward
Combinations in Florida Flora II
367
Harrisia gracilis (Miller) Britton var. simpsonii
(Small) I). B. Ward, comb, et stat. nov. Bas-
ionym: Harrisia simpsonii Small, in Britton &
Rose, The Cactaceae 2: 152. 1920. TYPE:
U.S.A. Florida: Monroe Co., hammock near
Flamingo, May 1919, ./. K. Small s.n. (lecto-
type, designated by Benson (1982: 934), NY).
After Britton (1908) set Harrisia apart from the
all-inclusive genus Cereus Miller, his protege, J. K.
Small (in Britton & Rose, 1920), placed names on
the forms he encountered in southern Florida. The
influence of Benson (1982), however, who preferred
retention of a comprehensive Cereus, has reduced
attention given to these endemic Florida segregates.
The three taxa addressed here are consistently giv¬
en recognition (Benson, 1982; Austin, 1984; Wun-
derlin, 1998), but at different levels.
In the judgment of Austin (1984), whose field
knowledge of these plants is unequaled, the small
differences between the Florida taxa of Harrisia
gracilis (his Cereus gracilis) and the differences that
separate them from their putative parent (the non-
Florida, typical H. gracilis) justify varietal status.
Benson (1982) also employed varietal rank in Ce¬
reus. Both authors retained Small’s H. fragrarts, but
at varietal level in Cereus. The increasing accep¬
tance of a moderate dissection of the larger Cereus,
by recognition of the segregate genus Harrisia,
makes the new combinations necessary.
These plants have been classified by the Florida
Department of Agriculture as “endangered” and as
such have protection of law. Because the Florida
statute (F.S. 581.185) does not provide for classi¬
fication of infraspecific taxa, they are listed as Har¬
risia eriophora and //. gracilis.
CoMI’t >SITAK
Arnoglossum ovatum (Walter) H. Robinson var.
lanceolatuni (Nuttall) I). B. Ward, comb, et
stat. nov. Basionym: Cacalia lanceolata Nut-
tall, Genera N. Amer. Plants 2: 138. 1818.
TYPE: U.S.A. “Florida” [on label], [date?],
Baldwin s.n. (lectotype, designated here, PH).
Though Nuttall stated his new species to have
been provided to him by (William) Baldwin, he im¬
precisely noted its source as “Georgia and Florida”
and did not designate a type. The only appropriate
specimen (PH) is labeled simply “(Nutt.)” ami
“Florida.” The present selection formalizes Nut-
tail's apparent intent.
The conventional treatment of Cacalia L. (cf.
Pippen. 1978) is now abandoned as irretrievably
ambiguous (Wagenitz, 1995; Brummitt, 1998; Nie-
olson, 1999), with all species reassigned to other
genera. Though Krai and Godfrey (1958) under¬
stood (and well illustrated) two distinct varieties in
what is now Arnoglossum ovatum, subsequent au¬
thors have failed to give status to Nuttall’s narrow¬
leaved variant.
Eupatorium album F. var. petaloideuin (Britton)
R. K. Godfrey ex I). B. Ward, comb, et stat.
nov. Basionym: Eupatorium petaloideum Brit¬
ton. in Small, Bull. Torrey Bot. Club 24: 492.
1897 [“ petalodium corr. Small, 1903: 1168].
TYPE: U.S.A. Florida: Duval Co. [1892?],
Curtiss 1190 (lectotype, designated here, NY).
Though the accomplishments of Robert K. God¬
frey in clarifying the southeastern flora were many
(cf. Krai, 2001), one of his major efforts failed to
reach publication and is thus little recognized. This
was Godfrey’s prolonged and detailed field study of
the genus Eupatorium. Through the 1960s and ear¬
ly 1970s he devoted many hundreds of hours to
tracing down and visiting the type localities of the
numerous names assigned to southeastern members
of this genus. Repeatedly he concluded (pers.
comm.) that certain of these names did not deserve
the oblivion to which later authors had assigned
them.
Eupatorium petaloideum Britton in Small (1897)
was one such name. In the field Godfrey found
plants that matched Britton’s description, common¬
ly not in association with E. album and distinguish¬
able from that species, lie maintained that Britton’s
taxon “has to be recognized at some level apart
from E. album ” (pers. comm., Feb. 1987). Though
this judgment — never placed in print — is in con¬
trast with that of Cronquist (1980), Clewell (1985),
and Wunderlin (1998). Godfrey’s superior field ex¬
perience justifies retention of the taxon at a sub¬
ordinate level.
Liatris pauciflora Pursh var. secunda (Elliott) I).
B. Ward, comb, et stat. nov. Basionym: Liatris
secunda Elliott, Sketch 2: 278. 1822? TYPE:
U.S.A. Georgia: Savannah, S. Elliott s.n. (ho-
lotype, CHARE presumed lost).
Following the monograph of Gaiser (1946), two
previously confused Florida taxa of Liatris have
seemed clearly defined. Liatris pauciflora Pursh
was described as having the stem and leaves gla¬
brous, with an erect, racemose, or paniculate inflo¬
rescence; L secunda Elliott was distinguished by
the stem short-pubescent, and the inflorescence ra¬
cemose with heads frequently secund. [Other au¬
thors (Small, 1933; Clewell. 1985; Wunderlin,
368
Novon
1998) supplemented these small differences with
the observation of eglandular phyllaries in L. pau-
ciflora and glandular-dotted phyllaries in L. secun¬
da, a characteristic seen neither by Gaiser nor ap¬
parent on recent collections (FLAS, FSU).] But the
weak morphology was not critical; the two taxa were
allopatric, with L. pauciflora in the central and
northern peninsula and sparingly into southeastern
Georgia, and L. secunda disjunct, in the western
panhandle and extending northward through Geor¬
gia anil into the Carolinas.
The pattern shown by these two taxa is quite
familiar, with a widely distributed continental pop¬
ulation and a near-endemic peninsular population,
a presumed consequence of Pleistocene glacial
flooding and isolation. But, barring knowledge of
origin, the two are separable on little more than the
difference in pubescence, with perhaps a greater
tendency in Liatris secunda toward secund-
branehed inflorescences (itself a consequence of an
arching stem). If they were sympatric, specific rank
would be unlikely to have been proposed. In view
of tin' real but minor observable differences, vari¬
etal rank is sufficient.
Pityopsis nervosa (Willdenow) Dress var. tracyi
(Small) I). B. Ward, comb, et stat. nov. Bas-
ionym: Chrysopsis tracyi Small, Southeast U.S.
1182, 1339. 1903. TYPE: U.S. A. Florida:
Manatee Go., in sand, Palma Sola, 3 Dec.
1901, Tracy 77 Id (holotype, NY).
The judgment as to whether a comprehensive
Chrysopsis Elliott should also encompass Pityopsis
N ul tall (Dress, 1953; Cronquist, 1980) is progres¬
sively moving toward the negative (Small, 1933;
Dress, 1975; Glewell, 1985; Semple iK Bowers,
1985; Wunderlin, 1998). But Semple and Bowers
(1985) misplaced the large-headed Pityopsis tracyi
Small by assigning it varietal status within the
small-headed P. graminifolia (Michaux) Nuttall. As
clearly recognized by Dress (1953, 1975), the
large-headed complex is best distinguished as P.
nervosa. Within P. nervosa s.l., variety nervosa (with
narrow, ascending leaves) occurs throughout the
stati', and variety tracyi (with broad, spreading
leaves) is endemic in the peninsula (north to Mar¬
ion County).
Gramineak
Andropogon elliottii Chapman var. stcnopliyllus
(llackel) I). B. Ward, comb, et slat. nov. Bas-
ionym: Andropogon virginicus E. var. viridis
Curtiss ex Hackel in DC. subvar. stenophyllus
llackel, in DC., Monogr. Phanerog. 6: 411.
1889. TYPE: U.S. A. Florida: Chapman s.n.
(lectotype, designated by Campbell (1983:
217), W; isolectotype, W).
By its very quality, the excellent monograph by
Campbell (1983) of the Andropogon virginicus com¬
plex has deflected later workers from inclination to
challenge one name change imposed on a wide¬
spread member of the group. Within this difficult
genus the species known since Chapman’s first flora
(I860) as A. elliottii Chapman stands out with its
distinctive overlapping and inflated sheaths. Camp¬
bell rejected this name, replacing it with the pre¬
viously questionable and unused A. gyrans Ashe.
Campbell (1983) made the novel argument that
Andropogon elliottii . even though Chapman’s ma¬
terials and description were unmistakably this spe¬
cies, is nevertheless based on A. argenteus Elliott
(1816) since Chapman parenthetically included
that name (a synonym of A. ternarius Michaux) at
the bottom of his original description.
As a replacement name Campbell selected An¬
dropogon gyrans Ashe (1898). This name, however,
had no holotype, and Campbell acknowledged “the
equivalence ... is questionable” of the Ashe spec¬
imen he selected as a neotype. Hitchcock and
Chase (1951: 812) had referred Ashe’s name to A.
elliottii. but with a query.
Campbell is incorrect in claiming A. argenteus
Elliott as Chapman’s basis. Chapman had clearly
indicated he did not accept Elliott’s name (since it
was a homonym of A. argenteus DC.) and in his
later revision (Chapman, 1897) omitted the refer¬
ence entirely. With restoration of Andropogon el¬
liottii Chapman, the infraspecific taxa defined by
Campbell become: Andropogon elliottii var. elliottii
[= A. gyrans var. gyrans] and variety stenophyllus
(Hackel) I). B. Ward [= A. gyrans var. stenophyllus
(Hackel) Campbell],
linperulu eylindrica (L.) P. Beauvois var. mexi-
cana (Ruprecht) I). B. Ward, comb. nov. Bas-
ionym: Imperata brasiliensis Trinius var. mex-
icana Ruprecht, Bull. Acad. Sci. Bruxelles 9:
245. 1842. TYPE: Mexico. Vera Cruz: [date?],
Galeotti 5678 (holotype, K).
Two forms of Imperata have been introduced into
Florida. The first to appear [earliest record: Miami,
Mar. 1905, Britton s.n. (FLAS, NY)] was identified
as /. brasiliensis Trinius; it has remained restricted
to south Florida. The second | earliest record:
Gainesville, 29 Mar. 1937, Ritchey s.n. (FLAS)] was
introduced as /. eylindrica (L.) P. Beauvois; it has
spread aggressively throughout most of the state
(still rare in south Florida), largely by vegetative
Volume 14, Number 3
2004
Ward
Combinations in Florida Flora II
369
means. This second taxon, known as Cogon-grass,
is presently classified in Florida as a Category I
invasive weed; on a global scale il has long been
ranked as one of the world's top ten worst weeds
(Holm, 1969).
Though these taxa are presumed to come from
different continents (South America and Asia), and
are consistently treated as specifically distinct (e.g..
Gabel, 1982), in Florida they carr scarcely be dis¬
tinguished. The most recent field study (Tippincott,
1997), as well as a guide to the state’s flora (Wun-
derlin, 1998), rely on the number of anthers per
floret as the sole discriminator, a cryptic anil in¬
consequential basis for species delimitation (one
anther in I. brasiliensis; two in /. cylirulrica). In¬
deed, this character fails in cultivated materials (C.
Lippincott, pers. comm., Sep. 2001). There are ad¬
ditional aspect differences [inflorescence narrowly
spicate, plants low (to 0.5 m) in /. brasiliensis ; in¬
florescences somewhat diffuse, plants taller (to 1.5
m) in /. cylindrical. But separation, both in the field
and in the herbarium, remains difficult and uncer¬
tain.
Imperata is a poorly understood genus of 39
names (Chase & Niles. 1962) and eight or nine
species (Clayton & Renvoize, 1986; Gabel, 1982).
One is perhaps presumptuous to make a new com¬
bination here, in the absence of a perspective be¬
yond Florida, by uniting taxa that are so widely
assumed to be specifically distinct. But the prac¬
tical difficulty of determining which taxon is [ire-
sent, and the importance of bringing full force to
bear in restricting the spread of this pernicious
weed, suggests the wiser strategy lies in treating the
two taxa as a specific unit.
Lk<;i minosae
Dalen albiila (Torrey & A. Gray) I). B. Ward,
comb, et stat. nov. Basionym: Petalostemon
carneum Michaux var. albidum Torrey & A.
Gray, FI. N. Amer. 1:311. 1838. TYPE: U.S.A.
Georgia; Milledgeville, [ilateY | llaykin s.n. (ho-
lotype, NY).
Dalea gracilis (Nuttall) I). B. Ward, comb. nov.
Basionym: Petalostemon gracile Nuttall, J.
Acad. Nat. Sci. Phil. 7: 92. 1834. TYPE:
U.S.A. “W. Flor.” [on label], [date?], N. A.
Ware s.n. (lectotype, designated here, PH).
Nuttall (1834: 92) reported his new species as
“Hab. In the lower part of Alabama and Florida,”
without designating a type or collector. As indicated
by the slips affixed to his specimens (PH), he had
received materials from Nathaniel A. Ware (from
Florida), from Hezekiah Gates (from Alabama), and
from William Baldwin (source not indicated). The
first of these was marked by I). k. Wemple in 1965
as “typus,” though his published report (1970: 33)
stated only “Tyj le in PH." The present selection
formally confirms his choice.
The excellent treatment of Petalostemon by
Wemple (1970) underlies the revision by Barneby
(1977), who included all species of Petalostemon
within the more inclusive Dalea. But Barneby,
lacking Wemple’s Florida field experience, under¬
stated the magnitude of differences separating Pe-
talostemon albidum (Torrey & A. Gray) Small from
the related P. carneum (cf. Small, 1933; Wemple,
1970); he assigneil the taxon to Dalea carnea var.
albida (Torrey & A. Gray) Barneby. Similarly, he
minimized the differences separating Petalostemon
gracile Nuttall from P. carneum ; he reduced Nut-
tail's species to Dalea carnea var. gracilis (Nuttall)
Barneby.
Barneby’s viewT of the generic limits of Dalea is
accepted. But the taxa so clearly recognized by
Small and by Nuttall need restoration to the level
of species.
Tephrosia virginiana (F.) Persoon var. mohrii
(Rydberg) 1). B. Ward, comb, el stat. nov. Bas¬
ionym: Cracca mohrii Rydberg, N. Amer. FI.
24: 163. 1923. TYPE: U.S.A. Florida: Walton
Co., near Eucheeana, June 1880, Mohr s.n.
(holotype, US).
Tephrosia mohrii (Rydberg) Godfrey N krai is an
endemic taxon of the Florida panhandle, known in
only two counties (Clewell, 1985). Though similar
to the widespread T. virginiana, it has been rec¬
ognized as distinct by Small (1933), Godfrey and
krai (1958), and Clewell (1985). But Wood (1949),
Isely (1990), and Wunderlin (1998) have combined
it without distinction.
The distinction between Tephrosia mohrii and T.
virginiana is not inconsiderable (cl. Godfrey &
krai. 1958; Clewell. 1985); T. mohrii is a much
smaller plant, with smaller, less-pubescent leaflets.
Even so, T. virginiana varies enough within itself
(Isely, 1990) to justify the doubt of authors who are
not familiar with the endemic variant in the field.
An intermediate, varietal status is appropriate. This
taxon, as Tephrosia mohrii, has been classified by
the Florida Department of Agriculture as “threat¬
ened,” under the criteria of Florida Statute
581.185.
370
Novon
Nyctauimackak
I'isonia discolor Sprengel var. floridana (Britton)
I). B. Ward, comb, et stat. nov. Basionym: Pi-
sonia floridana Britton, in Small, FI. Southeast
U.S. 411, 1330. 1903. TYPE: U.S.A. Florida:
Monroe Co., among lime rocks, Bock Key,
| date?], Blodgett s.n. (holotype, NY).
The forms of Pisonia discolor encountered in
Florida may reflect "founder effect” selections from
variable Caribbean populations. Bogle (1974) rec¬
ognized two Florida varieties: variety discolor
Sprengel (Syst. ii: 168, 1825), and variety longifolia
Heimerl (Bot. Jahrb. Syst. 21: 627. 1896), both with
glabrous to sparsely pubescent leaves. A third var¬
iant, Pisonia floridana, a plant with “copiously pu¬
bescent" leaves, was collected in the mid 1800s on
Rock Key, 12 miles west of Key West, and not again
seen (Bogle, 1974: 33). Plants with moderately pu¬
bescent leaves, perhaps representing this taxon,
were found in 1994 on Long Key (R. Hammer, pers.
comm., Dec. 1994).
Though the Florida variants perhaps intergrade
and likely do not represent major segregates of the
circum-Caribbean parental stock of Pisonia discol¬
or , no harm is done by making available this third
varietal combination P. discolor var. floridana, that
may aid in documenting variability of the Florida
entities.
Rubiackak
llniistniiin procunibens (Walter ex .1. F. Gmelin)
Standley var. hirsuta (W. H. Lewis) I). B.
Ward, comb. nov. Basionym: Hedyotis procum-
bens (Walter ex J. F. Gmelin) Standley var. hir¬
suta W. H. l^wis, Ann. Missouri Bot. Gard.
53: 378. 1966. TYPE: U.S.A. Florida: Walton
Co., Villa lasso, 1 mi. W of Choctawfhatchee]
Bay, 28 May 1964, McDaniel 4707 (holotype,
FSU).
The coastal-plain wildflower felicitously known
to schoolchildren as Fairy-footprints varies greatly
in pubescence. This trait was addressed by Lewis
(1966) by recognition of the pubescent extreme as
Hedyotis procunibens var. hirsuta. Neither Wilbur
(1968) nor Terrell (1996) accepted the variety as a
“biologically significant taxon,” though Wilbur pre¬
sented two maps (1968: 310) that recorded distri¬
bution of the glabrous and pubescent entities.
Long observation of the species in the field sup¬
ports the evidence deduced from Wilbur’s maps,
that nearly all individuals may be readily assigned
to one entity or the other (FLAS, D.B.W., annotat¬
ed). It is true, as originally stated by Lewis and
fully confirmed by both Wilbur and Terrell, that the
ranges are largely sympatric (though as noted by
Wilbur, the pubescent form does not extend as far
into South Carolina; it also is essentially absent
from southern Florida (D.B.W., pers. obs.; Wilbur,
1968)). Though no argument can be made that
these two entities are of high taxonomic signifi¬
cance. the lack of ambiguity in sorting specimens
both in the herbarium and in the field makes re¬
tention of Lewis’s taxa worthwhile.
Lewis (1966). however, was then in an inclusive
mode that favored merger of closely related genera;
he made the new combination under Hedyotis. In
more recent years there has been an apparent con¬
sensus that Houstonia (and Oldenlandia) are best
held separate (Terrell & Lewis, 1990; Terrell. 1990,
1991, 1996). With restoration of Houston ia (Terrell,
1996), Lewis’s variety requires a new combination.
Acknowledgments. I thank the curators of the
New York Botanical Garden for the loan of speci¬
mens critical to this study, James A. Macklin for
photocopies of Nuttall types. Carol L. Lippineott for
sharing her intimate knowledge of Florida Impera-
ta, Wilbur H. Duncan for his experience and in¬
sight regarding Yucca, Daniel F. Austin. Kerry A.
Barringer, Mark L. Gabel, Gerald F. Guala, James
L. Reveal, and Edward E. Terrell for their sugges¬
tions in review of this paper, and Victoria C. Hol¬
lowed for her patient, meticulous editorial services.
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in the Florida flora. Novon 11: 366—369.
Arisaema menghaiense (Araceae), a New Epiphytic Species from
South Yunnan, China
Yin Jian-Tao\ Li Heng '-2, and Xu Zai-Fu'
'Xishuangbanna tropical Botanical Garden, Chinese Academy of Sciences, Menglun,
Mengla 666303, Xishuangbanna, Yunnan, People’s Republic of China
‘Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming 650204,
Yunnan, People’s Republic of China. Iiheng@public.km.yn.cn
ABSTRACT. Arisaema menghaiense J. T. Yin, H. Li
& Z. F. Xu. (sect. Pistillata Engler) is described
and illustrated as a new species of Araceae from
South Yunnan, China. It is closely related to Ari¬
saema lobatum Engler in having a globose tuber,
trifoliolate leal blades, a unisexual spadix, and an
erect cylindrical appendix. It differs in its epiphytic
habit, the inflorescence that appears before the leaf,
a green (female) or [tale purple (male) spathe with
a filiform acumen ea. 60 cm (male) or 25 cm (fe¬
male) long, a pale green, erect, sessile cylindrical
spadix appendix with a few acute neuter flowers
basally and anther thecae opening by an apical slit.
Key words: Araceae, Arisaema, China, Yunnan.
During our study of tropical plant resources in
South Yunnan, China, a new species of Arisaema
was discovered; it is described here to make the
name available for the Flora of China.
The genus Arisaema Martius was established in
1831 when it included only Arisaema costatum
(Wallich) Martius (Martius, 1831), which was trans¬
ferred from Arum. The next year. Martius moved
two other species, Arisaema nepenthoules (Wallich)
Martius and A. speciosum (Wallich) Martius, into
Arisaema from Arum (Schott, 1832). Today, Arisae¬
ma comprises more than 170 species (Mayo et ah,
1997) and is one of the larger genera ol Araceae.
Arisaema menghaiense J. T. Yin, 11. Li tK Z. E.
Xu, sp. nov. TY EE: China. Yunnan: Menghai,
Bada, 100°4'E, 22°52'l\, epiphyte on tree,
1800 m. 29 Apr. 2002 (fl), Yin Jian-Tao AH
(holotype, HITBC). F igure 1.
Haec species Arisaemali lobato affinis, sed ah eo habitu
epiphytico, inflorescentia ante folium, spathae limbo vir-
ido (femineus) vel purpurascenti (mas) cum acumine fili-
formi 25 (femineus) vel 60 cm (mas) longo, appendice
viridula, cylindriea, erecta, sessili, infeme florum steri-
lium rudimentis subulatis sparsim obsila, antherarum the-
cis rima apieali dehiscentibus differt.
Perennial, dioecious, herbaceous epiphyte; tuber
depressed-globose, ca. 5 X 10 cm, renewing sea¬
sonally, purple outside with brown spots, 1—1.5 mm
diam.; roots whitish, 2—7 cm long, 1—2 mm diam.
Leaf (from fruiting plant) 1; petiole cylindrical,
green and smooth without spots, 20—30 cm long,
ca. I cm diam.; leaf blade greenish, membrana¬
ceous. trifoliolate; leaflets petiolulate; terminal leaf¬
let ovate, ca. 20 X 10 cm; apex acute; base cu-
neate, lateral veins numerous, connective vein 2—3
mm remote from the margin, petiolule 15 mm long;
lateral leaflets obliquely ovate-lanceolate, (‘a. 21 X
8 cm; apex acuminate with 5—8 mm filiform tail;
base obliquely cuneate; petiolules 8—1 I mm long.
Inflorescences unisexual, appearing before the leaf;
cataphylls 3, membranaceous, pale green, oblong;
apex obtuse, 18—23 cm long; peduncle terete,
white, ca. 26 X 0.3 cm (male), 25 X 1 cm (female);
male spathe tube cylindrical, green, ca. 6 X 1.5
cm, throat margin broadly aurieulate, strongly out¬
ward recurved; limb greenish purple, ovate-lance¬
olate, arched, ca. 6 X 5 cm, apex acuminate with
a purple tail up to 60 cm long; female spathe tube
whitish green, cylindrical, slightly constricted near
the throat, ca. 5 X 1 cm; throat obliquely truncate;
limb erect, whitish green, ovate-lanceolate, ca. 4 X
3.5 cm; apex acuminate, with a greenish tail up to
25 cm long. Male spadix with basal fertile portion
cylindrical, ca. 3.8 X 0.5— 0.6 cm, densely dow¬
ered; synandria subsessile, purple; anthers 4 to 6,
subglobose, dehiscing by an apical slit; appendix
sessile, cylindrical, greenish, ca. 3.8 X 0.3 cm;
apex obtuse, with a few subulate neuter flowers in
lower part; female spadix with basal portion cylin¬
drical, ca. 2.2 X 0.8 cm, densely flowered; ovaries
green. I -loculed, ovoid; stigmas subsessile, discoid;
ovules 5 to 6, basal, erect; appendix as in male
spadix. Fruiting spadix cylindrical, ca. 5.2 X 1-
1.5 cm; appendix ca. 4.5 cm long.
Distribution. China; Menghai County of Yun¬
nan Province.
Phenology. Flowering, April; fruiting. May.
Novon 14: 372-374. 2004.
Volume 14, Number 3
2004
Yin et al.
Epiphyte from Yunnan
373
Figure 1. Arisaema menghaiense .). T. Yin, H. Li & Z. F. Xu. — A. Male flowering plant with tuber. — B. Female
flowering plant. — C. Male spadix. — D. Synandrium. — E. Fruiting spadix. — F. Pistil. — G. Longitudinal section of
pistil showing ovules. — H. Leaves of female plant. A. B, C. I) from photos of Yin Jian-Tao 38 (holotype), and E. F.
G, H from Yin Jian-Tao 33 (female, paratype).
374
Novon
Arisaema menghaiense is an epiphytic species in
the genus Arisaema. This species belongs to section
Pistillata Engler (Engler, 1920; Hara. 1971; Li
Heng, 1979, 1980), which is characterized by a
tuber, 3(to 5)-foliolate leaf blade, unisexual spadix,
anther cells dehiscing by an apical elliptic or ob¬
long slit, and a cylindrical, erect spadix appendix
with nearly truncate base, which exceeds the
spathe tube. The following characters distinguish
A. menghaiense from .4. lobatum and other species
in section Pistillata: petiole and peduncle without
colored marks appearing before leaves, greenish
purple (male) or whitish green (female) spathe limb
with a filiform tail up to 60 cm (male) or 25 cm
(female) long, sessile cylindrical appendix with a
few subulate neuter flowers in the lower part, and
the anther thecae dehiscing by an apical slit.
I he living tubers of the type and paratvpe spec¬
imens were introduced in Xishungbanna Tropical
Botanical Garden in 2002.
I he epithet of the new species is derived from
the type locality.
Paratype. CHINA. Yunnan: Menghai, Mengsong,
I00°34'E, 22°.VN, epiphyte on trunk of tree, 1800 m, 29
May 2002 (young fr), Yin Jian-Tao 33 (HITBC).
Acknowledgments. The field expedition to Men¬
ghai County was supported by the Chinese Acad¬
emy ol Sciences (the Conservation and Study of
Tropical Plant Resources Project and the grant
number 2000WK-7). We are grateful to Zou Shou-
Qing and Huang Jia-Yuan of Xishungbanna Tropi¬
cal Botanical Garden for their comments and help
in preparing the manuscript. We also thank Lei Li-
Gong for checking the Latin and Wu Xi-Lin of the
Kunming Institute of Botany for preparing the il¬
lustration.
Literature Cited
Engler, A. 1920. Araceae— Aroideae und Araceae— Pistici-
deae. In: A. Engler (editor), Das Pflanzenreieh IY-23E
(Heft 73): 1-274 (Arisaema 149-220).
Hara, H. 1971. A revision of the eastern Himalayan spe¬
cies of the genus Arisaema (Araceae). Pp. 321—354 in
H. Hara, The Flora of Eastern Himalaya. Univ. Tokyo
Press, Tokyo.
Li. H. 1979. Araceae, Lemnaceae. Pp. 116—194 in (7 Y.
Wu & H. Li (editors), Flora Reipublicae Popularis Sin-
ieae 13(2). Science Press, Beijing.
- . 1980. Himalayas-Hengduan Mountains — The
centre of distribution and differentiation of the genus
Arisaema. Acta Bot. Yunnan. 2(4): 402-416 |in Chi¬
nese]; also in Geological and Ecological Studies of
Qinghai— Xizang Plateau, Vol. 2: 1321-1327. Science
Press, Beijing | in English].
Martius, C. F. P. von. 1831. Cher die Art der Befruchtung
bei einigen Aroideen und liber die Charakteristik meh-
rerer Gattungen dieser Eamilie. Flora 14: 449—460 (Ar¬
isaema — 159).
Mavo, S. J., J. Bogner & P. C. Boyce. 1997. The Genera
of Araceae: 270—275. Royal Botanic Gardens, Kew.
Schott, II. W. 1832. Araceae. In: II. W. Schott X S. En-
dlicher (editors), Meletemeta Botanica 16—22. C. Ger-
old, Vienna.
Volume 14, Number 3, pp. 245-376 of NOVON was published on 29 September 2004.
Volume 14 NO VON nI|[^
Number 4
2004
Leptodictyum wallacei (Musci: Amblystegiaceae), a New Species
from Texas, U.S.A.
Bruce Allen and Robert E. Magill
Missouri Botanical Garden, P.O. Box 299. St. Louis, Missouri 63166-0299, U.S.A.
bruce.allen@mobot.org; bob.magill@mobot.org
Abstract. A new aquatic moss from Hext Spring.
Texas, is described as Leptodictyum wallacei. The
new species is distinct in the genus by its small
plant size, stiff stem aspect, long leaf cells, firm-
walled basal cells, and undifferentiated alar cells.
Key words: Amblystegiaceae, aquatic moss,
Lept odictyum, Texas .
The southern interior region of the United States
(southern Missouri, Arkansas, eastern Oklahoma,
and central Texas) is an area of numerous limestone
springs. These springs are often fast-running and
have cold, clear water. Associated with these
springs are a number of interesting mosses such as
Hygroamblyslegium noterophilum (Sullivant & Les-
quereux) Warnstorf, Brachythecium rivulare Schim-
per, Donrichardsia macroneuron (Grout) Crum & L.
E. Anderson, Fiss ideas grandifrons Bridel, Fontin-
(dis duriaei Sehimper, and the curious laxirete- form
of Leptodictyum riparium (Hedwig) Warnstorf. Re¬
cently Brad Wallace collected an interesting moss
he found in large masses submerged in Hext
Spring, a limestone spring in San Saba County, Tex¬
as. He sent it to us for identification, and aftercare¬
ful observation we recognized it as a new species
of Leptodictyum (Sehimper) Warnstorf.
Leptodictyum wallacei B. H. Allen & Magill. sp.
nov. TYPE: U.S.A. Texas: San Saba Co.. Hext
Spring, 4 mi. W of Cherokee, off Ranch Road
501, USCS Magill Mountain quad map,
30°58'49"N, 098°46'22"W, 1555 ft. elev., 13
Dec. 2003, Brad Wallace s.n. (holotype, MO;
isotypes. BM. DUKE, EII. H. MICH.’ NY, PC.
S, US). Eigure I .
Species liaec a Leptodictyo ripario plantis rigidis, mar-
ginihus foliorum serrulatis, costis latioribus, foliis valde
tortilibus ubi siccis pilisque axillarihus rubris differt.
Plants slender in green to yellow-green, dense
masses of long, stiff, sparingly branched strands;
stems stiff, irregularly branched, in cross section
w ith sclerodermis of 4 to 5 rows of small, yellow or
reddish, thick-walled cells, cortical cells enlarged,
yellowish to hyaline, firm-walled, small central
stand present; paraphyllia absent; pseudoparaphyl-
lia foliose; rhizoids not seen; axillary hairs 3- to 5-
celled. lower I to 2 cells irregularly quadrate, up¬
per cells rectangular to cylindrical, all cells red.
Leaves erect-spreading and individually strongly
twisted when dry, erect when wet. evenly spaced,
2.0— 2.5 mm long, lanceolate, smooth, long-acumi¬
nate to a broad, blunt tip, concave, not decurrent;
margins plane to erect, obscurely serrulate to sub-
entire; costae single, broad, to 70 pun wide at base,
extending % the leaf length, in cross section cells
± homogeneous, thick-walled; laminal cells linear-
rhomboidal to vermicular, firm-walled, upper cells
to 70 X 6 pni, median leaf cells to 120 X 6 pun,
cells near leaf insertions long-rectangular, firm-
walled, at margins cells often subquadrate, alar
cells not differentiated. Perichaetial and perigonia
not seen. Sporophytes not seen.
Leptodictyum wallacei is a slender moss that
grows in large, permanently submerged masses in
Hext Spring, Texas. Since the plants grew sub¬
merged, it is not possible to determine the orien¬
tation of the stems, i.e., whether the primary stems
typically grow horizontally or erect, and there is
Novon 14: 377-379. 2004.
378
Novon
Figure I. Leptodictyum wallacei B. H. Allen & Magill. — A. Leaf apex and upper leaf cells. — B. Habit. — C. Median
leaf margin and median leaf cells. — 1). E & G. Leaves. — E Axillary hairs. — H. Costa in cross section, lower part of
leaf. 1. Stem in cross section. — J. Basal leal margin and basal leaf cells. Seale in mm: bar = 0.04 (F. H); bar =
0.06 (A. C. J); bar = 0.0B (1); bar = 0.5 (I). E. G); bar = 1.56 (B).
Volume 14, Number 4
2004
Allen & Magill 379
Leptodictyum wallacei (Musci: Amblystegiaceae)
little stem/branch leaf differentiation. When wet,
the leaves of L. wallacei are concave and evenly
spaced on the stems. The dry plants have a stiff
aspect, ami the leaves are individually, strongly
spirally twisted. The leaves of L. wallacei are
rounded at the apex, obscurely serrulate through¬
out. have remarkably long, narrow leaf cells (length:
width ratio up to 20:1), and a thick costa that can
be 70 gm broad at base.
William Buck (NY) examined this collection, and
on the basis of its very broad costa and aquatic
habit suggested it could be placed in the genus
Hygroamblystegium Loeske. This is where we ini¬
tially concluded it should be placed. However, the
exceptionally long, narrow leaf cells and complete
absence of alar cell development in this specimen
appears to undermine this view, bars Hedenas (S)
also examined the collection, and in his opinion
this moss represents an aberrant form of Leptodic¬
tyum riparium (Hedwig) Warnstorf. Bernard Goffi-
net and Alain Vanderpoorten (CONN), on the basis
of preliminary molecular analysis, are also of the
view that this species is best placed in leptodic¬
tyum (Schimper) Warnstorf.
There is in the Interior Highlands of North
America a very odd expression of Leptodictyum ri¬
parium which has been described several times.
This odd expression (the /a.ttre/e-expression of
Crum & Anderson, 1981) often grows submerged
in slow-moving streams or in springs. In the field,
plants of the /a.itre/e-expression have the size and
aspect of Fontinalis Hedwig. Its leaves are often
concave, evenly distributed around the stem, and
commonly rounded at the apex, which can be twist¬
ed when dry. The leaf margins can be obscurely
serrulate, the costa is relatively thick, and the long,
narrow leaf cells are up to 15:1 (lengtb:widtb).
Plants of L. riparium laxirete-ex pression are lax and
fairly large in size, and they differ from typical L.
riparium in having entirely red axillary hairs (see
Allen, Fontinalaceae Exsiccatae S. MO). The axil¬
lary hairs in L. riparium have one or two red basal
cells, but otherwise the cells are hyaline (see Mig-
ula, Cryptogamae Germaniae, Austriae et Helvetiae
401, MO). The plants from Hext Spring, Texas,
share many features with the laxirete-exprcss'um of
L. riparium , including having entirely red axillary
hairs. Although the two taxa are morphologically
similar, the Hext Spring plants are here recognized
as a new species on the basis of their smaller size,
stiff stems and leaves that have longer cells, firm-
walled basal cells, and undifferentiated alar cells.
Many of the features noted above are prone to plas¬
ticity when they occur in aquatic mosses. Never¬
theless, our extensive field experience with the lax-
ire te -expression of L. riparium in Missouri and
Arkansas indicates it never exhibits the distinctive
suite of features that characterizes L. wallacei.
Literature Cited
Crum, H. A. & L. E. Anderson. 1981. Mosses of Eastern
North America. Columbia Univ. Press, New York.
Synopsis of Plinia (Myrtaceae) in Mesoamerica
Fred R. Barrie
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Address
for correspondence: Botany Department, The Field Museum, 1400 S. Lake Shore Drive,
Chic •ago, Illinois 60605, U.S.A. fred. barrie@mobot.org
Abstract. Fhe 14 species of Plinia known from
Mesoamerica are reviewed. Light new species are
described and illustrated: one from Nicaragua, P.
nicaraguensis ; two from Costa Rica, P. guanacas-
tensis and P. moralesii; and five from Panama, P.
cerrocampanensis, P. coclensis, P darienensis, P
gentryi, and P. panamensis. One species, P. sala-
mancana, is transferred from Eugenia. A key to the
Mesoamerican species is provided.
RKSUMKN. Se examinan las 14 espeeies de Plinia
eonocidas de Mesoamerica. Se describen y ilustran
oc ho nuevas espeeies, una de Nicaragua, P. nicar¬
aguensis. dos de Costa Rica, P. guanacastensis y P.
moralesii y cinco de Panama. P cerrocampanensis.
I‘. coclensis, P. darienensis, I1, gentryi y P pana¬
mensis. Sc* transfiere una especie, P. salamancana.
de Eugenia. Se provee una clave para las espeeies
de Mesoamerica.
Key words: Costa Rica, Mesoamerica, Myrta¬
ceae, Nicaragua, Panama, Plinia.
I his report is the product of a revision of Plinia
L. undertaken for the* flora Mesoamerieana Project.
Plinia is a Neotropical genus of shrubs and small
trees, occurring from Brazil and Peru north to the*
West Indies and Cuba, and westward to Panama,
Costa Rica, Nicaragua, and Belize. The genus is
characterized by flowers with the hypanthium pro¬
longed beyond the summit of the ovary, a calyx
fused and nearly closed in bud that splits at anthe-
sis into four irregular lobes, and a bilocular ovary
with two locules per ovule. The fruit is a single- or
several-seeded berry; the seeds have a papery coat
enclosing an embryo with two large, plano-convex
cotyledons and a reduced, indistinct hypocotyl. The
fleshy fruits place Plinia in Myrtaceae tribe Myr-
teae. along with nearly all New World Myrtaceae;
the embryo characters place it in the subtribe Eu-
geniinae (McVaugh, 1968; Landrum Kawasaki.
1997).
Recent estimates ol the number of species in Pli¬
nia have ranged from as few as 6 (Sanchez Vindas.
2001) to approximately 30 (McVaugh, 1989). The
newly described taxa presented here raise the es-
Novoim 14: 380-400. 2004.
timate to ca. 40 species. The nearly seven-fold dif¬
ference is indicative of the confusion as to what,
exactly, constitutes a Plinia. The core group of spe¬
cies in Plinia, including all known Mesoamerican
taxa, shares with the type species, P. pinnata L.,
the character of sessile or subsessile flowers
grouped in cauliflorous, glomerate inflorescences
that are subtended by up to 10 pairs of prominent,
decussate bracts superficially resembling an invo¬
lucre (McVaugh. 1968). However, several authors
have increased the number of included species by
weighting certain floral characters and discounting
inflorescence structure. Kausel (1956) transferred
three species from Myrciaria (47. cauliflora. M. ja-
boticaba, and ,47. trunciflora ) solely on a single
character of the embryo, the presence of separate,
plano-convex cotyledons. But in at least one of
these species, Myrciaria cauliflora (Martins) ().
Berg, the flowers have a tubular hypanthium, cir-
eumcissile at the base and falling as a unit with
the perianth and androecium after anthesis (M. L.
Kawasaki, pers. comm.). This character is common
to all Myrciaria. but is otherwise unknown in Pli¬
nia. McVaugh (1958, 1963) referred to Plinia any
eugenioid species with floral characters as de-
sc ribed al >ove. Included were species with axillary,
racemose inflorescences and pedicellate flowers
(e.g., Plinia clausa McVaugh).
Kausel and/or McVaugh have been followed by
some subsequent authors (e.g., Rotman, 1982; So-
bral, 1993; Kawasaki (V Holst, 2002). though with
varying degrees of enthusiasm. Landrum and Ka¬
wasaki (1997: 532) described Plinia as “an arbi¬
trarily defined group." McVaugh himself considered
it a “heterogeneous assemblage" (1963: 507); his
inclusion of the racemose taxa was provisional,
pending reevaluation when better data on the ge¬
neric limits became available. The confusion is not
unique to Plinia. but indicative of the current un¬
settled state of generic concepts in t lit* Eugeniinae.
The issue will not be satisfactorily resolved until a
comprehensive analysis of the subtribe is per¬
formed.
I he Mesoamerican species of Plinia are shrubs
Volume 14, Number 4
2004
Barrie
Synopsis of Plinia
381
or small trees, 10 m or shorter, of evergreen lowland
and montane forests and cloud forests. All have the
floral anti inflorescence characters of the Plinia
“core group." The bark is smooth, white, gray, or
brown, and exfoliates in plates. Vestiture is com¬
posed of straight or recurved, simple trichomes,
0.5-3 mm long, and is quite dense on young
growth.
The branchlets are weakly to strongly com¬
pressed near the outermost nodes. They are bilat¬
erally channeled between the two outermost nodes
in Plinia guanacastensis, P. nicaraguensis, and P.
panamensis. Young growth is sericeous or velutin-
ous, the trichomes are simple, 0.5—3 mm long, erect
or recurved, and typically white or golden-brown.
The vestiture thins but persists with age.
Unlike Eugenia , where terminal, vegetative buds
tend to be rather uniform and unremarkable, in Pli¬
nia they often have species-specific characters. The
buds are ovate to broadly ovate, up to 1 cm long,
and protected by 8 to 14 ovate to broadly ovate,
imbricate scales, which vary among species in size,
shape, and vestiture. When a bud breaks, the inner
scales tend to erdarge and change shape. However,
the outer 2 to 4 scales retain their conformation
and, when present, are often useful for identifica¬
tion. The most significant characters of the scales
are general shape, whether or not the back is car¬
inate, the shape of the apex, whether rounded,
acute, or acuminate, and whether or not the mar¬
gins are scarious or ciliate. The form of the inflo¬
rescence bracts often approximates that of the veg¬
etative bud scales.
The leaves in Plinia are petiolate. The blades
are narrowly elliptic or lanceolate to elliptic or
ovate. Emergent leaves are commonly sericeous or
arachnoid, densely covered with simple, white tri¬
chomes up to 3 mm long. This covering is soon lost;
in mature leaves the vestiture is thinner, usually
composed of trichomes shorter than 1 mm and often
restricted to the midvein or to one side of the leaf.
Venation is brochidodromous; the submarginal vein
formed by the lateral veins as they curve to meet
the next succeeding lateral vein may be promi¬
nently looped if the lateral veins are few and rather
remote from the margin, as in P. salticola McVaugh,
or it may appear straight if it is close to the margin
or if the lateral veins are numerous, as in P. pan¬
amensis. The veins may be weakly convex or flush
with the surface of the blade (e.g.. P. darienensis)
or moderately to deeply impressed in the upper sur¬
face (e.g., P. panamensis). Leaf bases, typically cu-
neate. are cordate in I1, panamensis and subcordate
to abruptly truncate in P. cerrocampanensis. Ex¬
cepting P. gentryi, which has acute or acuminate,
bluntly tipped leaves, leaf apices are acuminate to
caudate, with a sharply pointed tip. Petioles are
terete or flattened dorsal Iv, with pubescence that
tends to match the branch to which it is attached
more than it does the blade.
fhe glomerate inflorescences are borne on the
trunks and larger branches. The bracts subtending
the1 inflorescence are persistent and conspicuous in
flower, though at times obscured by mature fruit.
The identifying characters are similar to those of
the vegetative bud scales: shape, apices, margins,
and vestiture. The 2 to 10 flowers are sessile, borne
in pairs, and subtended by a pair of bracteoles that
are similar to the bracts.
fhe flowers are 4-merous, with the hypanthium
prolonged into a tube above the summit of the ova¬
ry. The size and shape of the hypanthium, the de¬
gree to which it is ribbed, and vestiture type pro¬
vide useful, species-level characters. I lie calyx is
fused and nearly closed in bud, the tips of the calyx
lobes outlining a terminal pore. At anthesis, the
calyx tears irregularly into 4 lobes and reflexes. I he
petals are caducous and rarely found in herbarium
collections. They are white with irregular margins
and, in the few specimens seen, pubescent on both
surfaces. The stamens are numerous, borne in a
ring at the base of the calyx lobes that is presented
as the calyx reflexes. Like the petals, the stamens
are apparently quickly lost and infrequently seen
in specimens. The ovary is 2-loculed, with 2 ovules
per locule. The single style is glabrous.
The fruit is a weakly to strongly compressed ber¬
ry, usually single-seeded, 3—12 cm in diameter. I lie
pericarp is succulent when ripe, yellow or orange
with a sweet odor. Ribbing may be present or ab¬
sent. The membranous seed coat surrounds the em¬
bryo of a reduced hypocotyl and two massive, sep¬
arate, plano-convex cotyledons. In some species the
embryo is purple or reddish purple.
Plinia L., Sp. PI. 516. 1753. TYPE: Plinia pinnata
E.
Key to the Mesoamkrican Species oe I’ijma
la. Leaves 5.5—10 cm long.
2a. Leaves elliptic or oblong, coriaceous.
3a. Lateral veins 10 to 14 per side, the
leaf apex acute or acuminate, the tip
1-5 mm, blunt or rounded .
. 5. P. gentryi
3b. Lateral veins 16 to 22 per side, the
leaf apex acuminate or caudate, the
tip 10—25 mm, sharply pointed . . .
. 13. P. salamancana
2b. Leaves lanceolate or narrowly elliptic,
chartaceous.
382
Novon
Hi.
4a. Lateral veins 14 to 15 per side;
fruits lenticular . 2. ft coclensis
4b. Lateral veins 15 to 25 per side;
fruits oblate or subglobose .
. 12. ft puriscalensis
Leaves 9—40 cm long.
5a. Lateral veins 20 or more per side.
6a. Leaf base cordate ... 9. ft panamensis
6b. I eat base cuneate.
7a. Lateral veins inscribed or im¬
pressed in the upper leaf sur-
lace . 11. ft povedae
7b. Lateral veins flat or slightly
convex . 4. /I darienensis
5b. Lateral veins fewer than 20 per side.
8a. Lateral veins flat or slightly convex
on both surfaces; not prominent.
9a. Lateral veins 6 to 10 per side
. 14. ft salticola
9b. Lateral veins 12 to 18 per
side.
10a. Leaf blades 9—12 cm,
the petioles 5—10 mm;
Belize .... 10. ft peroblata
10b. Leaf blades 9—21 cm,
the petioles 15— 20 mm;
Costa Rica . . 7 .ft moralesii
8b. Lateral veins inscribed in the upper
leaf surface, prominent on the lower
leaf surface.
11a. Leaf tips 15—20 mm.
12a. Glands on the lower leaf
surface numerous, red or
brown, distinct .
... 1. ft cerrocampanensis
12b. Glands on the lower leaf
surface numerous, con-
colorous and indistinct
. 6. ft guanacastensis
I lb. Leaf tips 20—40 mm.
Ida. Leaf lips 20—30 mm;
petioles 12—20 mm ....
. 8. ft nicaraguensis
13b. Leaf tips 30—40 mm;
petioles 8—10 mm .
. 3. ft cuspidata
l . Plinia cerrocampanensis Barrie, sp. nov.
TYPE: Panama. Panama: Gerro Campana,
800-1000 m, 3 Apr. 1972, A. Gentry 4944 (ho-
lotype, MO; isotype, F). Figure I.
times as long as wide; chartaceous; drying green
above, paler green or yellow below, the veins com¬
monly red or brown on the lower surface; midvein
flat or convex on both surfaces; lateral veins 10 to
14 per side, inscribed in the upper surface, prom¬
inent on the lower surface; submarginal veins sim¬
ilar to the laterals and arched between them, 2—3
mm from the margins; surfaces of young leaves se¬
riceous or arachnoid, the trichomes 2-3 mm, slen¬
der, white, the upper surface ultimately glabrate or
with a few persistent trichomes, often near the base,
the midvein puberulent, the lateral veins glabrate,
the lower surface persistently pubescent, the Iri-
chomes ea. 2 mm. contorted, the midvein and lat¬
eral veins puberulent; glands numerous and small,
red or brown on the lower surface; base subcordate
or abruptly truncate; margins entire; apex abruptly
acuminate, the tip 15—20 mm, sharply pointed; pet¬
ioles 10—15 mm. terete or flattened dorsally, coarse¬
ly puberulent, the vestiture similar to the branch-
lets. Inflorescence cauliflorous, sessile, flowers ea.
8; bracts 2—5 X 1.5—3 mm, ovate, the back cari¬
nate, the base rounded, tomentose, the margins
searious-ciliate, the apex acuminate, paired in ea.
4 decussate ranks; buds 8—10 nun, pyriform, the
ovary tomentose, the trichomes ea. 2 mm. white,
the calyx glabrous, glandular, the pore 1-2 mm
diam.. the tips of the calyx lobes ciliate. Flowers
sessile; braeteoles ea. 6) X 2 mm, lanceolate, the
outer surface tomentose, the base rounded, the mar¬
gins searious-ciliate, the apex acuminate; hypan-
thium campanulate, the tube extending 2-3 nun be¬
yond the summit ol the ovary, the throat glabrous,
ea. 4 mm diam. at the base ol the stamina! ring;
calyx lobes 2—3 X 2—3 mm, oblong or obovate. both
surfaces glabrous, the apex broadly acute, ciliate;
petals 3—4 X 2.5—3 mm, obovate, white, both sur¬
faces puberulent, the trichomes ea. 0. 1 mm, the
margins and apex irregular, ciliate; stamens ea.
200, 5—7 mm; style ca. 10 mm. Fruits 30—50 X
40—60 mm. oblate or turbinate; pericarp fleshy with
numerous, indistinct, vertical ridges, sparsely pu¬
bescent; mature color yellow or orange.
Species haec a Plinia cuspidata foliis laminis basibus
cordatis vel trunoatis et apicibus brevioribus differ!.
Shrubs or trees to ca. 6 m; young growth velu-
tinous, the trichomes 2—3 mm. coarse, coppery at
first, turning white, branehlets compressed below
the terminal node, the bark red, velutinous, ulti¬
mately white, glabrate; outer scales of the vegeta¬
tive buds 3—6 X 2—3 mm, ovate, carinate dorsally,
pubescent, the margins searious-ciliate, the apex
acuminate or apieulate. Leaves narrowly elliptic or
elliptic, the blades 10-20 X 3. 5-6.5 cm. 2.5-3.7
Habitat, distribution, and phenology. Ridge-top
cloud forests and adjacent Atlantic slope montane
wet forests in the Parque Naeional Altos de Cam¬
pana, Panama, 700-1000 m. Collected in flower
May to July; collected in fruit March to May.
Plinia cerrocampanensis is distinguished bv the
carinate bud scales and inflorescence bracts with
acuminate to apieulate apices and the narrowly el¬
liptic leaves, 10—20 cm long, with subcordate or
abruptly truncate bases and numerous, small red
glands on the lower surface. The other Panamanian
Volume 14, Number 4
2004
Barrie
Synopsis of Plinia
383
MISSOURI
BOTANICAL GARDEN
HERBARIUM
N? 5166960
M y£T*c
plants OF PANAMA fiyrtaceae
prov. Panama
Gerro Campana, alt. 800-1000 meters.
Fallen tree 8 meters j fruits green, seed
purple within.
Al Gentry no.
A. Gentry 4944
HOLOTYPE: Plinia cerrocampanensis Barrie, sp nov
F.R, Barrie July 2003
MISSOURI BOTANICAL GARDEN |MOj
OO-A/i o
Figure 1. Plinia cerrocampanensis Barrie (Gentry 4944: holotype, MO).
species with cordate leaf Bases, P. panamensis , lias
broader and typically much longer leaves (14—30
cm) with numerous, deeply inscribed lateral veins.
The ovate, dorsally carinate bracts with acumi¬
nate apices are similar in shape, though about twice
the size, to those of P. cuspidata Gomez-Laurito &
Valverde. The two may be distinguished by several
leaf characters, among them the apices, which are
caudate-aristate in P cuspidata , with the acumen
30-40 nun long. The leaf bases are cuneate, and
384
Novon
the glands on the undersurface are green or clear,
not red. This is one of three Panamanian species,
the others being P. cor I crisis and P. gentryi, that
occur in isolated patches of low-elevation cloud for¬
ests along the Continental Divide. These habitats
are known to be rich in endemics (Myers, 1969;
Lewis, 1971).
Paratypes. PANAMA. Panama: Cerro Campana,
Busey 848 (MO); P. N. Altos de Campana, sendero del
Tigre, Correa A. & Montenegro 9543 (F), Espinosa et al.
709 (F), Galdames 745 (F), Galdames et al. 4318 (MO);
Sendero de Interpretacidn. 1 km al K del Campamento de
los guardabosques de INRENARK, Correa A. & Monte¬
negro 9023 (f ), Galdames 1106 (F); Reeorrido por el sen¬
dero a un eostado del Camino Zamora, Correa A. & Mon¬
tenegro 1 1 150 ( f ); Cordillera del Llornn. Galdames et al.
3790 (f, MO); upper slopes of Cerro Campana within the
boundary of the national park administered by RE-NA¬
RK. LeDoux 2578 (MO).
2. I’linia coclensis Harrie, sp. nov. TYPE: Pan¬
ama. Code: summit at Alto Calvario, low mon¬
tane cloud forest, ridge-type vegetation, 4 Apr.
1977, ,/. I*. Folsom R. Robinson 2399 (ho-
lotype, MO; isotypes, F, MO). Figure 2.
Phniam puriscalensem aemulans, sed Ibliis nervis pau-
eioribus et fructibus lenticularibus differ!.
Shrubs or trees, 1.5-5 m; young growth seri¬
ceous, the trie homes white, 1-2 mm. Branchlets
slender, weakly compressed at the upper nodes;
bark red. smooth, ultimately glabrate or with a few
persistent trichomes; the outer scales of the vege¬
tative buds broadly ovate. 1-2 X 1-2 mm, tin- back
carinate, glabrous, the margins ciliate, the apex
acute. Leaves narrowly elliptic or lanceolate, the
blades 5.5—8 X 1.5— 2.2 cm, 3—4.4 times as long
as wide; ehartaceous; drying a concolorous green;
midvein flat or convex on the upper surface, convex
on the lower surface; lateral veins 14 to 15 per side,
obscure on one or both surfaces of mature leaves;
submarginal veins similar to the laterals and weak¬
ly arched between them, ea. 2 mm from the mar¬
gins; tin- upper and lower surfaces glabrate or per¬
sistently puberulent near the base of the midvein;
glands minute, obscure on mature leaves; base cu-
neate; margins cartilaginous; apex caudate, the tip
15—25 mm; petioles 8— H) mm, puberulent, ulti¬
mately glabrate; terete or weaklv compressed dor-
sally. Inflorescences, (lowers, and attached fruits
not seen. Fruits 10-20 X 20-40 mm, lenticular;
pericarp fleshy, obscurely ribbed, glabrous; mature
color unknown.
Habitat, distribution, and phenology. Wet for¬
ests, Pacific Slope, Code Province, Panama. 400—
900 rn. Collected in fruit January to April.
Plinia coclensis is distinguished by the carinate,
broadly ovate bud scales with acute apices, the nar¬
rowly elliptic leaves with cuneate bases, and
strongly lenticular fruits that are obscurely ribbed
and glabrous. The leaves are similar in size and
shape to those of the Costa Rican species P. pur-
iscalensis P. E. Sanchez & O. Jimenez, which has
oblate or subglobose, rather than lenticular, fruits.
Paratypes. PANAMA. Code: Codecito Road, tran¬
section from I mi. beyond divide to ridge top, de Fevers
et al. 0740 (F, MO); 7 km N of Llano Grande on road to
Coclesito, Hammel 1944 (MO).
3. Plinia enspidala Gdmez-Laurito & Valverde,
Lankesteriana 3: I I. 2002. TYPE: Costa Rica.
Limon: Talamanca, Bratsi, Buena Vista. Finca
ACODEFO, 25 Nov. 2000. 0. Valverde & S.
Mora 1339 (holotype, USJ).
Illustration. Gdmez-Laurito & Valverde, Lan¬
kesteriana 3: 12; f. 1. 2002.
Shrubs or trees, to 15 m; young growth sericeous,
the trichomes 1—3 mm, white. Branchlets com¬
pressed and bilaterally grooved immediately below
the terminal nodes; bark reddish brown, villosu-
lous, the trichomes 0.3— 0.5 mm, straight and re¬
curved. brown, ultimately sordid; outer scales of the
vegetative bracts 1—3 X 1—1.5 mm. ovate, carinate,
appressed-pubescent on the outer surface, the mar¬
gins scarious, the apex acuminate. Leaves elliptic,
the blades 13—23 X 5—10 cm, 2—3 times as long
as wide, ehartaceous, drying a concolorous green;
midvein convex between two parallel grooves on the
upper surface, prominent on the lower surface; lat¬
eral veins 14 to Iff per side, shallowly to deeply
inscribed in the upper surface, prominent on the
lower surface; submarginal veins similar to the lat¬
erals and weakly arched between them, 4—5 mm
from the margins; the upper surface ultimately gla¬
brate. the midvein glabrate or persistently pubes¬
cent. the trichomes erect or recurved, ca. 2 mm.
the lower surface persistently but sparsely pubes¬
cent, the trichomes 0.3— 0.5 mm, the midvein and
lateral veins pubescent to tomentose; glands nu¬
merous on both surfaces, obscure above on mature
leaves; base cuneate to obtuse or rounded; margins
planate; apex abruptly caudate acuminate, the tip
30-40 mm, sharply pointed; petioles 8-10 mm, vil-
losulous, similar to the branchlets, terete. Inflores¬
cences cauliflorous. sessile, flowers 4 to 10; bracts
1.5— 3.5 X 1.5—3 mm. lanceolate to ovate, sparsely
pubescent to tomentose on the outer surface, the
margins searious-ciliate, the apex acute, paired in
4 decussate ranks; buds 9—10 mm. pyriform, to¬
mentose, the pore 1-2 mm across. Flowers sessile;
Volume 14, Number 4
2004
Barrie
Synopsis of Plinia
385
MISSOURI
BOTANICAL GARDEN
HERBARIUM
NS 2601119
New Vork BoUmral G.rden
Cf ?(.*,<-
BET; L. R. Landrum, 1981
1 *7-2*’*'
itnmit at Alto Calvarlo, low montane
oud forest, ridge-type vegetation; 900 m.
sc -shaped fruit, one seeded, with 2
xge purple cotylendons; fruit turning
id; shrub of 2 meters; young leaves silver.
P. Fols
April 1977
Figure 2. Plinia coclensis Barrie {Folsom & Robinson 2399 ; holotype, MO).
bracteoles ca. 2 X 1 mm, lanceolate or ovate, to-
mentose, the margins scarious-ciliate, t he apex
acute or acuminate; hypanthium campanulate, ea.
7 mm, the tube extending 3-4 mm beyond the sum¬
mit of the ovary, the ovary and tube tomentose, the
trichomes ca. 1 mm, white; calyx lobes ca. 4X4
mm, oblong, tomentose without, villosulous within,
the margins irregular, the apex rounded; petals
white, both surfaces puberulent, the trichomes c a.
0.1 mm; stamens ca. 130; style 3^f mm. Fruits 30
386
Novon
X 35—40 mm, oblate or lenticular; pericarp fleshy,
smooth, yellow at maturity; calyx persistent, re¬
duced. die lobes deciduous.
Habitat, distribution, and phenology . Wet for¬
ests, Atlantic slope, from central Panama to central
Costa Rica, 0—900 m. Collected in flower in July;
collected in fruit February to April.
Plinia cuspidata is distinguished by the carinate
vegetative bud scales with acuminate apices, leaf
blades with 14 to 18 lateral veins per side and
abruptly caudate-acuminate apices 30-40 mm
long, and oblate or lenticular, smooth fruits. Many
ol the specimens examined were originally identi¬
fied as P. povedae, the leaves of which tend to be
narrower, with 20 to 40 lateral veins per side and
the leaf apex less abruptly acuminate. The fruits of
P. povadae are costate and typically have 6 to 8
prominent ribs.
Additional specimens examined. COSTA RICA. Car¬
tage: Moravia tie Turrialba, Poveda 9(d) (CR. F). Linidn:
Gudpiles, bos Angeles, San Miguel. Herrera & Schik 3798
(K MO); Kst. Carillo, Sendero a las tomas de agua, Cerro
Coronel, F, ol Laguna Danto, Stevens 2.1805 (MO); Cerro
Coronel, h of Rfo Zapote, F. of new road to Raphia swamp,
within I km of Rfo Colorado, Stevens et al. 24675 (F).
Puntarenas: Osa, Sierpe, Boca Ganado, Fila Ganado,
Herrera et al. 665 t (CR, f); Osa. Rajando Cerro Brujo
hacia el oeste, Sierpe, Marin & Trejos Urena 329 (I NR,
MO). San Jose: P. N. Braulio Carillo, fist. Carillo, San¬
chez V. 566 (CR). PANAMA. Boras del Toro: Siguiendo
el camino que va a Quebrada Bonyic, Santamaria & Lara
1091 (f ). Code: vie. La Mesa, bey. FI Valle, on N slope
ol Cerro Caital, McPherson 11222 (F). Panama: Capita,
NE side of Cerro Trinidad, Foster 2111 (MO).
4. Pliuiu darienensis Barrie, sp. nov. TYPE: Pan¬
ama. Darien: Rfo Pirre, old forest 10 mi. S of
El Real near “Dos Bocas,” 2 Apr. 1971, R.
Foster & T. Lowenbach 2271 (holotype, MO;
isotypes, F, MICH). Figure 3.
Pliniae panamensi similis, sed foliis laminis basibus
cuneatis et venis lateralibus et submarginalibus convexis
vel planis differt.
Shrubs or trees to ca. 7 m; young growth seri¬
ceous, the triehomes 2-4 mm, coarse, coppery
branchlets weakly compressed; bark brown, flaking,
densely to sparsely pubescent, the triehomes ca.
0.2 mm, straight or recurved; outer scales on veg¬
etative buds 3—5 X 5—8 mm, depressed ovate,
densely appressed pubescent, the margins undu¬
late, scarious-ciliate, the apex rounded. Leaves nar¬
rowly elliptic, the blades 14-40 X 6-8 cm, 3-5.5
times as long as wide; coriaceous; midvein flat or
concave on the upper surface, prominent on the
lower surface; lateral veins 20 to 35 per side, flat
or slightly convex; submarginal veins similar to the
laterals and arched between them, 1-2 mm from
the margins; the upper surface ultimately glabrate,
the midvein persistently sericeous or hispidulous,
the triehomes ca. 0.1 mm. erect, the lower surface
glabrate or with a few persistent triehomes near the
base, th<“ midvein hispidulous, the triehomes ca.
0.2 mm; glands numerous and minute on both sur¬
faces or obscure above; base cuneate; margins rev¬
olute near base; apex acuminate or caudate, the tip
15 mm, sharply pointed; petioles 10-15 mm,
densely puberulent, similar to the branchlets, te¬
rete. Inflorescence cauliflorous, sessile; flowers 2 to
4; bracts ca. 2X2 mm, ovate, the outer surface
tomentose, the inner surface with scattered Iri-
chomes, the margins undulate, irregular, scarious,
the apex rounded, paired in 2 to 4 decussate ranks;
buds 12-15 mm, pyriform, the base tomentose, the
tube and calyx sparsely pubescent. Flowers sessile;
bracteoles ca. 2 X 2 mm, similar to the bracts,
ovate, the outer surface tomentose, the inner sur¬
face with scattered triehomes, the margins undu¬
late, irregular, scarious, the apex rounded; hypan-
thium campanulate, the tube elongate 3-5 mm
beyond the summit of the ovary, 6-8 mm diam. at
the base ol the staminal ring, the base tomentose,
the tube and the outer surface of the calyx lobes
glabrate; calyx lobes ca. 9 X 7.5 mm, oblong, pu¬
bescent on the inner surface, the margins irregular,
the apex rounded; petals not seen; stamens ca. 400;
style ca. 18 mm. Fruits 65-80 mm diam., oblate,
the calyx lobes and tube persistent; pericarp fleshy,
4 to 8 prominent ribs plus numerous indistinct in¬
termediates, sparsely pubescent; mature color or¬
ange or yellow.
Habitat, distribution, and phenology. Wet for¬
ests, Darien Province, Panama, 300 m. Collected
in flower and fruit October to April.
Plinia darienensis is characterized by narrowly
elliptic leaves with blades 14— 40 cm long, with lat¬
eral and submarginal veins that are flat or nearly
so on both surfaces. In the other large-leaved Pan¬
amanian species, P. panamensis and P. povedae, the
lateral veins are inscribed. Plinia panamensis dif¬
fers also in having cordate leaf bases. The outer
bud scales are broader than long with undulate,
scarious-ciliate margins. The fruits are similar in
appearance to those of P. panamensis, though pos¬
sibly somewhat smaller. Fruits of the latter with di¬
ameters of 120 mm have been seen, but, as the
measurements for P. darienensis are based on only
two collections and five fruits, it may be that the
apparent size difference is an artifact of inadequate
sampling.
Paratypes. PANAMA. Darien: La Boca de Pirre,
Volume 14, Number 4
2004
Barrie
Synopsis of Plinia
387
It l-awr ,f T. hminhach 22V
Hoi-OTVI't: /’/min ilarieiKnm Bamc. sp iil
FR Barrie
MISSOl'Kl BOTANICAL CAKDKN |MO|
Provincia :J,
Figure 3. Plinia darienensis Barrie ( Poster & Lowenbach 2271 ; holotype, MO).
Bristan 1247 (MO); Camino del Pine, Duke & Bristan 2 SB
(MO); Cerro Pirre, valley betw. Pirre & next most southerly
peak, Folsom 4391 (MO); ca. 3 mi. SE of Pijibasal, ca. 9
mi. S of El Real, vie. of confluence of Dos Bocas & Rfo
Pirre, Hartman 1201 1 (F. MO).
5. Plinia gentryi Barrie, sp. nov. I \ PE; Panama.
Panamd: near top of Cerro Jefe to 1 mi. be¬
yond, 1 Jan. 1972. A. Gentry 3448 (holotype,
MO). Figure 4.
388
Novon
MISSOURI
BOTANICAL GARDEN
HERBARIUM
N? 04968854
Small tree 2" dbh, 5 m tall. Fruits turning
orange.
Near top of Cerro Jefe to 1 ml. beyond.
Figure 4. Plinia gentryi Rarrie (Gentry 344H; holotype, MO).
J44B Jan. 1,
MISSOURI BOTANICAL OAR DEN HERBARIUM (MO)
A Plinia salamancana foliis nervis paucioribus, apici-
bus brevioribus et peliolis longioribus differt.
I rees ca. 5 m; young growth coarsely tomentose,
the trichomes 1-3 mm; branchlets compressed at
tlx 1 outer nodes; hark reddish, puberulent, the tri-
chomes ca. 0.2 mm. contorted, ultimately glabrate;
outer scales of the vegetative buds ca. I X 1 mm,
broadly ovate, the margins rounded, eiliate, the
scales otherwise glabrous. Leaves elliptic, the
blades 3.5— 7.5 X 1.5—4 cm, 1.7— 2.5 times as long
Volume 14, Number 4
2004
Barrie
Synopsis of Plinia
389
as wide; coriaceous; midvein convex or flat on the
upper surface, convex on the lower surface; lateral
veins 10 to 14 per side, often obscure above on
mature leaves, convex below; submarginal veins
similar to the laterals and arched between them. 1-
2 mm from the margins; both surfaces glabrate or
with a few persistent trichomes near the base, the
midvein glabrate or persistently puberulent above,
persistently puberulent below, the trichomes ca. 0.2
mm; glands numerous, minute and punctate on the
upper surface, scattered on the lower surface; base
obtuse; margins revolute; apex acute or abruptly
acuminate, the tip 1-5 mm, blunt or rounded; pet¬
ioles 8—15 mm, puberulent, the vestiture similar to
the branchlets. ultimately glabrate, terete or some¬
what flattened dorsally. Inflorescence cauline, flow¬
ers not seen, apparently 4; bracts not seen; attached
fruits not seen. Fruits 15-25 X 25-35 mm, oblate;
pericarp fleshv. rugulose, pubescent, more densely
near the apex; calyx deciduous in ripe fruit, leaving
a terminal, circular scar; mature color not known.
Habitat , distribution , and phenolo¬
gy. Apparently endemic to the cloud lorest area
on Cerro Jefe, Panama, 900 m. Collected in fruit
in January.
Apparently restricted to the narrow band of cloud
forest along the ridge of the Continental Divide on
Cerro Jefe, Plinia gentryi is most similai to P. sal-
amancana (Standley) Barrie, which gro vs in the
wet montane forests at lower elevations. I he tw o
may be readily distinguished by leaves alone. The
leaf blades of P. gentryi tend to be shorter (3. 5-7. 5
cm long vs. 6-10 cm in P. salamancana ) and
broader (1.7— 2.5 times as long as wide vs. 2.5— 2.8
in P. salamancana :), with shorter apices (1—5 mm
vs. 15—25 mm). The ultimate leaf tip is blunt or
rounded, rather than sharply pointed, as is the case
in P. salamancana. In both species, in mature
leaves the lateral veins are obscure on the upper
surface, which is dotted with numerous, minute,
punctate glands.
Etymology. Named in honor of the late, noted
tropical botanist A1 Gentry, who collected the type
of this species, as well as that of P. cerrocampanen-
sis.
Paratypes. P/ NAMA. Panama; Cerro Jefe, Duke
946H (MO); near >p of Cerro Jefe to I mi. beyond, Gentry
et al. 3471 (MICil, MO).
6. Plinia guanacastensis Barrie, sp. nov. TYPE;
Costa Rica. Guanacaste: P. N. Guanacaste,
Cord, de Guanacaste, Cerro Cacao. Est. Cacao,
10 Mar. 1991, C. Chavez 512 (holotype, INB;
isotypes CR, F). Figure 5.
Species haec a Plinia salticola plus venis lateralibus
foliis et fructibus immaturis puberulis differt.
Trees to ca. 12 m; emergent growth sericeous,
the trichomes 1—2 mm, white; branchlets com¬
pressed, bilaterally channeled below the terminal
nodes, the bark reddish, puberulent, the trichomes
ca. 0.1 mm; outer scales of vegetative bracts ca. 1
X 1 mm, widely ovate, the margins scarious, the
apex rounded or mueronate. Leaves narrowly ellip¬
tic or elliptic, the blades 12—16 X 4-6 cm, 2.7—
3.5 times as long as wide; chartaceous; drying a
concolorous green or paler on the lower surface;
midvein flat or convex on the upper surface, prom¬
inent on the lower surface; lateral veins 10 to 18
per side, inscribed in the upper surface, prominent
on the lower surface; submarginal veins similar to
the laterals and arched between them, 4-5 mm
from the margins; the upper surface persistently
pubescent along the midvein anti the lateral veins,
otherwise glabrate, the lower surface with a few
persistent trichomes, the midvein sericeous, the tri¬
chomes 1-2 mm, the lateral veins puberulent;
glands numerous, obscure above on mature leaves;
base cuneate; margins revolute near the base; apex
caudate-acuminate, the t if) 15—20 mm. sharply
pointed; petioles 10—15 mm, pubendent, similar to
the branchlets, terete. Inflorescence cauliflorous,
sessile; flowers 2 to 8; bracts 1—2 X 1—2 mm, semi¬
circular. convex, a few trichomes on the outer sur¬
face, the margins ciliate, the apex rounded, paired
in 2 to 4 deci ssate ranks of decreasing size; buds
not seen. Flowers and mature fruits not seen. Im¬
mature fruits ca. 2 cm, obconic, sessile, constricted
below the remnants ol the calyx, puberulent, the
trichomes white or golden, ca. 0.1 mm; calyx lobes
deciduous, the tube persistent, forming a crown ca.
3X2 mm on the developing fruit.
Habitat, distribution , and phenology. Montane
wet forests in the Cordillera de Guanacaste, Costa
Rica, 800-1000 m. Collected in flower in August;
collected in fruit February to March.
The developing fruits of Plinia guanacastensis
are similar to those of P. salticola MeVaugh, but are
puberulent rather than glabrous or nearly so. While
the leaf blades of both species are approximately
the same length, 12-16 cm here, 9-17 cm in P.
salticola, the leaves of P. guanacastensis tend to be
narrower, 2.7— 3.5 times as long as wide versus a
ratio of 2-3:1 in P. salticola, with about twice as
many lateral veins, 10 to 18 versus 6 to 10 per
side, which are inscribed, rather than flat or con¬
vex. as in P. salticola.
Paratypes. COSTA RICA. Guanacaste: Liberia, P. N.
Guanacaste, Cord, de Guanacaste, Kst. Cacao, Sendero a
390
Novon
Figure
INSTITUTO NACTONAL DE BIODIVERSID/
11 ^olaboracioii COK IL MSSOURr BOTANICAL CARDEN (HO)
^ ft*
C Chdvez 512
GUANACASTE: Cantdn de Liberia.
P. N. Guanacaste
Cord, de Guanacaste; Cerro Cacao.
Kstaoidn Cacao.
10-55'
//yf £ & f ^ O € dfetJC ^ho£^
FLORA DE NICARAGUA
< irtajnerto rt* 2KJLAYA, AUar.t«» C«fto
M "ifrillo ( la Klcada)
Arbol de ? ata. hojaa » ratings
S ' do da olor m a tacit a
'
corteja lisa id /.
riinia satumancana (Standi ) Barrie, comb nov
Woodson et al. 1570
Holotype: Eugenia salamancana Standi
F R Barrie July 2003
MISSOURI BOTANICAL GARDEN |MOJ
96724?
H0L0 TYPE OF:
Eugenia salamancana Standi.
Ann. Missouri Bot. Gard. 26:295. 1939.
ver. S. Long 1995.
FIELD MUSEUM OF
NATURAL HISTORY
NEGATIVE NO
68884
Figure 9. Plinia salamancana (Standley) Barrie ( Woodson el al. 1570 ; holotype, F).
Additional specimens examined. PANAMA. Colon:
Santa Rita Arriba, 2-4 km ’. Sparre 191 14 (S), B. Sparre 19252 (S). Za-
inora-Chinchi|M‘: Rio Nangaritza, Paehicutza, ./. Jar-
amillo 18943 (QCA), J. Jaramillo 14092 (QCA). ./. Jar-
amillo & E. Grijalva 13413 (QCA), W. Palacios & I). Neill
6474 (MO. QCNE), W. Palacios et al. 8330 (MO, QCNE);
Miazi, detras del Gampamento Militar, IE Palacios et al.
8506 (MO. QCNE). PERU. Amazonas: N\\ del PV l also
Paquisha, Cordillera del Condor, 5. Baldeon 541 (USM);
Cordillera del Condor, PV Alfonso Ugarte, Rio Comainas,
II. Beltran & It. Foster 795 (MO).
Antliiiriiiin pinkleyi Croat & Carlsen, sp. nov.
TYPE: Ecuador. Napo: Limoncocha, 240 m,
20 May 1991, S. Ingram & I). Atwood 977,
originally collected by M. Madison et al. s.n.,
Selby accession #78-1 156 (holotype, SEE; iso¬
type. MO 4224348). Figure 2C. D.
Planta terrestris. Elos lamina profunde triloba usque ad
4—7 cm e basi fissa, lobis omnibus latitudine aequalibus,
lobo medio oblongo vel oblanceolato, lobis lateralibus pa-
tentibus apicibus anguste rotundatis, eostis posticis nudis.
Inflorescentia spatha viridi, purpurascente, fibrosa; spad-
ice sessili, purpureo usque atrobrunneo, cylindrico aut an¬
guste acuminato.
Terrestrial; internodes short, 2—2.5 cm diam.;
cataphylls drying reddish brown, 3—6 cm long, de¬
ciduous. Leaves erect-spreading; petiole (20-)33-55
cm X 3—5 mm, terete (slightly D-shaped); genicu-
lum yellow, 1.5-3 cm X 2-3 mm, drying somewhat
darker than petiole; blades deeply trilobed to 4—7
cm from the base, subcoriaceous, (18— )22-37 X
(20-)26-44 cm, medium green above, pale yellow¬
ish green below, lower surface drying conspicuously
paler than above and silvery, both surfaces matte
to semiglossy, raphides usually absent, base acute
to truncate; central lobe oblong to oblanceolate,
(15— )20-33 X 5-1 l cm, apex acuminate, apiculum
2—3 cm long; midrib raised above, somewhat sunk¬
en toward the apex, raised below; primary lateral
veins 4 to 6 per side, arcuate-ascending, weakly
raised above and below, departing midrib at 30°—
40°; tertiary veins weakly raised below; collective
veins arising from the third (second) pair of primary
lateral veins, 4-6 mm from the margin, sunken
above, raised below, angled; lateral lobes spread¬
ing. 90° from central lobe, 14—22 X 4.5—10 cm,
apex narrowly rounded, inner margin convex to
straight, outer margin concave; basal veins 5 to 6
pairs, the first one joined for 1—3 cm, the rest up
to 8 cm; posterior ribs straight to weakly arcuate,
80°— 90° from midrib, naked for 2—3.5 cm. Inflores¬
cence erect; peduncle 16— 40(— 54) cm X 2—4 mm,
shorter than petiole (equal); spathe green becoming
purplish, lanceolate, fibrous, 5.5— 9.5 X 0.9— 1.5
cm, apex long-acuminate, base meeting peduncle
at 90°, somewhat auriculate; spadix sessile, purple
to dark brown, cylindric to narrowly tapered, stout,
4—9 cm long, 4-8 mm diam. at base, 3-5 mm diam.
at apex; ca. 6 flowers [ter principal spiral, rhombic,
2.8-3.2 X 2.8 mm, sides straight to weakly sig¬
moid; lateral tepals ca. 2.3 mm wide, inner margin
straight to weakly rounded, turned upward when
the pistil emerges, outer margin angled; stamens
weakly exserted; anthers ca. 0.5 X 0.65 mm, thec ae
elliptic, slightly divaricate; pistils emergent; stigma
elliptic, ca. 0.4 mm across. Infructescence dark pur¬
ple, 8-9 X 1-1.4 cm; fruits green becoming dark
purple, roundly ovate, ca. 3 X 2.5 mm.
Anthurium pinkleyi is known only from north¬
eastern Ecuador in Napo and Orellana Provinces
(Fig. 4A). It grows in tropical moist forests at less
than 500 m in elevation. It is named after its ear¬
liest collector, Homer V. Pinkley, who made a spec¬
imen in 1966 ( H . Pinkley 278, ECON) from Dur-
eno, Rfo Aguarico, just east of Lago Agrio, while
conducting ethnobotanical studies in the area.
Anthurium pinkleyi can be recognized by its na¬
ked posterior ribs departing from the midrib at 80°—
90°. its slender lobes, the central one being oblong
and the lateral ones spreading, and its sessile, dark
purple, cylindric spadix subtended by a fibrous
green to purplish green spathe. Also characteristic
408
No von
are the tepals that twist upward after the pistil
emerges.
Right of t fie collections examined, including the
type specimen, were made from cultivated plants
growing at Marie Selby Botanical Gardens (Selby
accessions: <8-1126, <8-1 149, and 78-1156), that
were originally collected by Madison and col¬
leagues in Ecuador, Napo Province, near Limon-
cocha in the vicinity of the Rio Napo. Apparently
no herbarium specimen was made at the time of
collection of the living material for cultivation.
When Madison (1978) redt ‘scribed Anthurium
rimbachii Sodiro, he included as a representative
specimen one of the collections taken from the type
plant of A. pinkleyi (M. Madison 4135). He did not
have the type specimen of A. rimbachii on hand
and was clearly confusing that species with this
new material, because they both have deep purple,
sessile spadices. According to the original descrip¬
tion ol Sodiro (1903), A. rimbachii was collected in
western Ecuador (Guayas Province) and it differs
in having winged posterior ribs and bigger leaf
blades (26—58 X 28—46 cm). It also differs in hav¬
ing very long exserted stamens and an obovate cen¬
tral lobe, constricted at the base. Unfortunately, the
location ol the type specimen ol A. rimbachii is
unknown. However, A. rimbachii clearly differs
from the material included here in A. pinkleyi by
all the above-mentioned characters.
In Ecuador, Anthurium pinkleyi might also be
confused with A. moronense, both having purple,
sessile spadices. However, the latter spec ies has
broader central and lateral lobes (10-17 cm and 8—
I.) cm wide, respectively) and occurs at higher el¬
evations in southeastern Ecuador. Anthurium pink¬
leyi may be confused with A. angustilohum Croat
from Panama, both sharing slender, oblong central
lobes, fhe latter differs in having collective veins
arising from the first pair of basal veins and running
3-4 mm from the margin, and a stipitate green spa¬
dix.
Paratypes. ECUADOR. INapo: bimoncocha, original¬
ly collected by M. Madison et al., cultivated at Marie Sel¬
by Botanical Gardens, A. Christenson 1 108 |SEb 78- 1 149|
(MO, SEE), T. Croat 79395 (MO), S. Ingram 1159 [SEE
l
inflorescence and leaf. — B. Leaves showing spreading and rounded lateral lobes, abaxiul surface. C, I). Anthurium
ternifolium Croat & Carlsen (Croat 72829). — C. Trilobed leaves showing falcate lateral lobes and sunken primary
lateral veins. — I). Close-up of immature infructescence.
52 X 1.3-2 cm; fruits dark purple, white at base,
obovoid, apiculate, 5—7 X 3—4 mm.
Anthurium rotundutum ranges from the Atlantic
slopes of southern Costa Rica to Panama (Atlantic
and Pacific slopes) into the very northern tip of
Colombia (Fig. 4B). It grows from sea level to 1000
m in elevation, in tropical wet or premontane forest.
'Phe specific epithet refers to the almost rounded
apex of the lateral lobes of the leaves. Anthurium
rotundutum is recognized by its naked posterior
ribs, spreading lateral lobes with broadly rounded
apices, and its stipitate yellow spadix.
Most specimens now attributed to this new spe¬
cies were formerly identified as Anthurium guru-
garunum Standley. However. Croat (1986) pointed
out that the Panamanian material he included in A.
garaguranum differed from the other material in
having stout, naked posterior ribs, and shorter,
more rounded lateral lobes. Anthurium rotundutum
and A. trilobum share a yellow, stipitate spadix. I he
410
Novon
I' igure 4. A (top). Distribution map of six new South American species of Anthurium sect. Semaeophyllium. — B
(bottom). Distribution map of the new Central American species, Anthurium rotundatum.
Volume 14, Number 4
2004
Carlsen & Croat
New Species of Anthurium
411
latter differs in having winged posterior ribs, lateral
lobes narrowly rounded at apex, and 8 to 14 pairs
of primary lateral veins.
Anthurium rotundatum can also be confused with
A. chimhorazen.se from Ecuador, both sharing na¬
ked, straight posterior ribs and spreading lateral
lobes, rounded at apex. The latter differs in having
the collective veins (2-)3-5 mm from the margin
and a sessile deep green spadix.
Paratypes. COSTA RICA. Union: Manzanillo de Tal-
amanca, M. Grayum & 1C Burton 4328 (MO); Qda. Mata
de himrin, Finca Anai, M. Grayum et al. 4446 (MO); union
de Rfo Uren y Rfo Sukut. 1.5 km aguas arriba, G. Herrera
3175 (MO). PANAMA. Bocas del Toro: entre la presa
y Valle La Esperanza, rfo Changuinola, I,. Carrasquilla &
A'. Mendoza 1384 (PMA); rd. betw. Fortuna Dam and Chi-
ri(|Lif Grande, 7.3 mi. N of bridge, T. Croat & !M. Grayum
60260 (MO). Chiriqin: rd. betw. Chiriquf Grande and
Fortuna, 7.7 mi. W of Chiriquf Grande, T. Groat & M.
Grayum 60093 (CM. MO. PMA, RSA). Code: trail to Las
Minas. P //. Allen 2462 (US); La Mesa. 7.’ Groat 14397
(MO); Cerro Pilon. T. Groat 37340 (MO). T. Groat 37567
(MO); El Cope, near Continental Divide, T. Croat 44723
(MO), B. Hummel 955 (MO), B. Hammel 1049 (MO). .S.
Knapp & R. Dressier 3488 (MO); Llano Grande-Coelecito,
N of La Pintada, T. Groat 49222 (MO): rd. betw. finca
Mandarinas and Finca Furlong, T. Croat 67194 (MO); Cer¬
ro Gaital, T. Croat 74811 (MO), G. de Nevers et al. 3509
(MO, PMA), S. Knapp 5321 (MO), .S'. Knapp 5800 (MO);
Alto Calvario. J. P. Folsom 2480 (MO); La Junta- Alto Cal-
vario, J. P. Folsom 5908 (MO); from La Mesa towards Los
Llanos, J. Luteyn 4104 (DUKE); Llano Grande-Cascajal,
NW of Penonome, A. Sytsma 3844 (MO); Cerro Caracoral,
A. Sytsma 4083 (MO); Cerro Turega, R. F. Woodson & R.
W. Schery 167 (MO). Colon: along Rfo Guanehe, T. An¬
tonio 4810 (MO), 7.’ Croat 37021 (MO), G. McPherson
8516 (MO); Cerro Jefe region, Santa Rita Ridge Rd.. 10-
21 km from Transistbmanian Hwy., M. Correa 587
(DUKE, MO), T. Croat 34304 (MO), S. Knapp & R.
Schmalzel 5209 (MO), .S'. Knapp & R. Schmalzel 5228
(MO), G. McPherson 10232 (CM. MO), .S'. Mori & ./. Kal-
lunki 5058 (MO); Portobelo-Nombre de Dios, Nuevo To-
nosi-Rfo Indio, T. Groat 33529 (MO, PMA); Portobelo-Rfo
Cascajal, vie. Nuevo Tonosi, 7. Croat 33658 (MO); Por¬
tobelo-Nombre de Dios, 10 km W of Nombre de Dios, .S’.
Knapp & J. Mallet 5686 (MO), S. Knapp & J. Mallet 5705
(MO); SW of Portobelo. R. Liesner 1032 (MO); originally
collected at Cerro Jefe, cultivated at Univ. of Hawaii, R.
Sheffer 176 (MO). R. Sheffer 275 (MO); 5-8 km SW of
Cerro Bruja, A. Sytsma et al. 4296 (MO. I’M A); Donoso,
hacia Valle Grande, A. Zapata et al. 969 (PMA). Pana¬
ma: Cerro Azul, T. Croat 11594 (MO); Cerro Campana,
7.’ Croat 12161 (MO), T. Croat 35942 (MO), P. Hutchinson
& R. Dressier 2955 (DAV), H. van der Werjf & J. Herrera
6209 (MO); rd. beyond Su Lin Hotel, T. Croat 14229
(MO); El Llano-Cartf, 7-20 km from Inter-American Hwy.,
T. Croat 25147 (MO), T. Croat 75141 (CAS. MO), J. P.
Folsom 2564 (MO), ./. P. Folsom & L. Collins 1519 (MO),
S. Hoover 1311 (MO), R. Liesner 1192 (MO, NY), S.
Thompson 4623 (MO); 3-3.5 mi. NE of Altos de Pacora,
T. Croat 68677 (MO); P. N. Altos de Campana, Buena
Vista, Espinosa et al. 3165 (PMA); Torti Arriba, Rancho
Chorro, ./. P. Folsom et al. 6678 (MO); Peluca meteoro¬
logical station. II. Kennedy & R. Dressier 2983 (MO); Rfo
Terable valley, S. Knapp & R. Schlmalzel 5489 (MO); Ser-
ranfa de Maji, S of lpeli, .S'. Knapp et al. 4485 (B, K, MO).
San Bias: El Llano— Cartf Rd., Km 19—28, G. de Nevers
6 II. Herrera 7248 (CM, K. MEXU, MO. NY). G. de Ne¬
vers et al. 5079 (MO); Rfo Cangandi, Pueblo Cangandi, H.
Herrera 208 (MO); Nusigandf, G. McPherson 10788 (MO);
Rfo Ada, Aila Tiwar, A. Sugden 455 (K. MO). Veraguas:
Alto Piedra-Rfo Dos Bocas, 5-8 km from Escuela Agrfcola
Alto Piedra, T. Croat 25966 (MO); Santa Fe-Calovebora,
Escuela Agrfcola Alto Piedra, T. Croat 49010 (MO). T.
Croat & J. P Folsom 33969 (MO). J. P. Folsom & I,
Collins 1614 (MO); Santa Fe to Rfo San Luis. Rfo Segundo
Brazo. T. Croat 66894 (MO); Rfo Primero Brazo, 5 mi.
NW of Santa Fe, R. Liesner 819 (MO). COLOMBIA. Cho-
co: E side of Serranfa del Darien, approached from Acan-
df, A. Juncosa 666 (MO).
Antluiriiiin ternifolium Croat & Carlsen, sp. nov.
TYPE: Ecuador. Pastaza: along road betw.
Puyo & Banos, vie. ol Shell, less than 1 km N
of village, 1°29'39"S, 78°3'52"W, 1096 m, 15
Aug. 2002, T. Croat, L. Hannon & P. Schmidt
86586 (holotype, QCNE; isotypes, B, COL,
IIUA, K, MO 5735452-5735453 [2], NY,
QCA, US, USM, VEN). Figure 3C. I).
Planta terrestris aut epiphytica scandensque, interno-
diis usque ad 7 em longis. Folium petiolo semitereti vel
D-formi; lamina profunde triloba usque ad 1 2— 14 .0 — 6.5 cm
e basi fissa, lobo medio oblongo-lanceolato graeili, lobis
lateralibus falcatis apicibus anguste rotundatis, costis pos-
ticis nudis, costa media utrinque acute elevata. Infiores-
centia spatha viridi; spadice sessili, viridi, cylindrico aut
anguste aeuminato.
Terrestrial or climbing epiphyte; internodes up
to 7 cm long, 1-1.5 cm diam.; cataphylls drying
reddish brown, 4-8 cm long, apex acuminate, tie-
composing into fibers, persisting at upper nodes.
leaves erect; petiole (14— )20— 34 cm X 3—5 mm,
semiterete or D-shaped; geniculum 1—1.5 cm X 3—
5 mm; blades deeply trilobed to [2— ]4— 6.5 cm from
the base, subcoriaceous, 13-28 X | 1 2- 1 16-26 cm,
dark green, semiglossy above, slightly paler, matte
below, raphides visible on upper surface, base cu-
neate; central lobe oblong-lanceolate, (8.5-] 14—26
X 1 3.5 — (5 — 1 0 cm. apex acuminate; midrib raised
above and below, angled; primary lateral veins 5 to
7 per side, arcuate-ascending, sunken above, raised
below, departing midrib at 30°— 45°; tertiary veins
moderately conspicuous; collective veins arising
from one of the lowermost pairs of primary lateral
veins, 4—7 mm from the margin, sunken above,
raised below; lateral lobes falcate [spreading], 35°-
45° [ 70°-90°] from central lobe, 14-22 X 4-8.5
cm, apex narrowly rounded, inner margin straight
to weakly convex, outer margin concave; basal
veins 3 to 5 pairs, the first free, the rest joined for
2.5—4 cm; posterior ribs straight to weakly arcuate,
55°— 65° from midrib, naked for 1—3 cm. In/lores-
412
Novon
cence erect; peduncle 20—35 cm X 2—5 mm, equal
or longer than petiole; spathe green, lanceolate, re-
flexed. 4-8 X 0.6-1 cm, apex acuminate, acumen
3—5 mm long, base obtuse, clasping; spadix sessile,
green, cylindric to narrowly tapered, (3— )5 — !()(-! 4)
cm long, 4—5 mm diam. at base, ca. 3 mm diam.
at apex; 3 to 4 (lowers per principal spiral, rhombic,
3—3.5 X 2—2.5 mm, sides sigmoid; tepals semi¬
glossy, lateral tepals ca. 1.5 mm wide, inner margin
straight to weakly rounded, then curved upward,
outer margin angled; stamens not exserted; anthers
ca. 3.5 X 0.5 mm, thecae elliptic, slightly divari¬
cate; pistils emergent just at tepal level; stigma slit-
like, ca. 0.5 mm across. I hfructescence green, 7—14
X 0.7— 1.5 cm; fruits green, ovate, ca. 5 X 3—4 mm.
Anthurium ternifolium is endemic to the north¬
eastern slopes of the Andes in Ecuador (Morona-
Santiago. Napo, and Pastaza Provinces), growing
between 800 and 1700 m in premontane and mon¬
tane wet forests (Fig. 4 A). The species epithet re¬
fers to the deeply trilobed leaf blades with the cen¬
tral lobe being almost equal to the lateral ones in
width and length.
Anthurium ternifolium is distinguished by its
slender, oblong-lanceolate central lobe, semi-terete
petiole, midrib acutely raised on both sides, falcate
lateral lobes, collective veins 4—7 mm from the
margin, and its short, green, sessile spadix.
Anthurium ternifolium can be confused with A.
grex-avium Madison, which occurs at high eleva¬
tions in southeastern Kcuador, both having falcate
slender lateral lobes, climbing habit, and elongated
internodes. However, A. grex-avium differs in hav¬
ing just one very conspic uous pair of basal veins,
winged posterior ribs, and a stipitate spadix. The
leaves of A. ternifolium may also resemble juvenile
leaves of A. furcatum Sodiro, both having naked
posterior ribs, falcate lateral lobes, and an oblong
central lobe. However, A. furcatum is distinct in its
primary lateral veins departing tlm midrib at (>()°—
70°. in h aving a stipitate, red-purple spadix, and in
occurring on the western slopes of the Andes in
Kcuador.
Paratypes. KCUADOR. Moronu— Santiago: P. N.
Sangay, entre Rfo Sardina y Volran. Montalvo & C. Cerdn
21 (QAP). Napo: carretera Hollfn— Loreto, km 25—31. A.
Alvarado 149 (MO). F. Hurtado & X. Ruiz 1796 (MO). IK
Palacios 406X (MO); rd. betw. Baeza and Tena, 31—59 km
N of Archidona, T. Croat 49606 (MO), 7. Croat 49626
(MO). T. Croat 5X7X2 (MO. OCA); 6.7 km W of Rfo Pay-
amino. 20 km W of Loreto, T. Croat 7262X (MO); Rfo
Granadillo, Canipamento de INECEL, “Codo Alto,” IE
Palacios 5512 (MO): Proyecto Hidroel^ctrico Coca, Rfo
Quijos, 10 km 5 de Reventador, IK Palacios 5X92 (MO.
QCNE). Pasta/.a: vicinity of Mera, E. Asplund IX54X (S),
h. Asplund 1X574 (S), E. Asplund 19596 (S), T. Croat
1 2X29 (MO). M. Lugo 64 (MO, S). /.' Plowman A' E. Davis
1549 (S); vicinity of Shell. X. Cornejo & C. Honifaz 1460
(GUAY), T. Croat 7X547 (MO. QCNE); Colonia Isidro
Ayora, C. Hurling <£• /.. Andersson 16956 (GB); Puyo—
Puerto Napo rd., San Jose ca. 17 km E of Puyo, G. Hurling
& L. Andersson 171 15 (GB); 4 km N of Mera, along Rfo
San Jorge and Rfo Tigre, It. 0llgaurd & II. Balslev 9141
(AAU); 3.5 km \\ of Mera on road to Banos. II. Stein 29X6
(MO).
Acknowledgments. The authors thank Peter
Stevens (MO) for numerous and very helpful com¬
ments on the original manuscript. This publication
is part of a yet unpublished revision of Anthurium
sect. Semaeophyllium that the first author carried
out as a master's thesis project at the Biology De¬
partment of die University of Missouri— St. Louis,
under the guidance of P. Stevens and T. Croat. All
photographs used to illustrate the species were tak¬
en by the second author. We are thankful to P. Ste¬
vens and Roy Gereau for correcting Latin descrip¬
tions, and Victoria Hollowed, Dorothy Bay, Jon
Ricketson, and an anonymous reviewer for helpful
comments on the manuscript. The following her¬
baria are gratefully acknowledged for their speci¬
mens on loan: AAU, B. CAS, CM, DAV, DUKE,
KCON, K. GB, GUAY, K. MKXU, MICH, NY, PM A,
GAP. OCA, QCNE, S, SKI,, US. USM. W IS, WU.
Literature Cited
Croat. T. IK 1986. A Revision of the Genus Anthurium
(Araeeae) of Mexico and Central America. Part II: Pan¬
ama. Monogr. Syst. But. Missouri Rot. Card. 14: 1-204.
- . 1994. Taxonomic status of Neotropical Araeeae.
Aroideana 17: 33—60.
- & R. 1). Sheffer. 1983. The sectional groupings of
Anthurium (Araeeae). Aroideana 6: 85-123.
Madison, M. 1978. The species ol Anthurium with pal-
inately divided leaves. Selbyana 2: 239— 281.
Schott, II. C. I860. Prodronius Systematis Aroidearum.
Typis congregationis mechitharistieae, Vienna.
Sodiro, L. 1903. Anturios Ecuatorianos. Monograffa II.
Contribuciones al eonoeimiento de la flora Ecuatoriana.
l ip. de la Escuela de Arles y Olicios, Quito.
Two New Species of Encyclia (Orchidaceae: Laeliinae)
from Venezuela
German Carnevali Ferndndez-Concha* and Ivon M. Ramirez
Herbarium CICY, Centro de Investigaci6n Cientifica de Yucatan, Calle 43, no. 130, Col.
Chuburna de Hidalgo, 97200 Merida, Yucatan, Mexico. carneval@cicy.mx; ramirez@cicy.rnx
^Research associate of the Orchid Herbarium of Oakes Ames, Harvard University Herbaria,
22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A.
ABSTRACT. Two new species of Encyclia Hooker
(Orchidaceae: Laeliinae) are proposed from Vene¬
zuela. E. betancourtiana Carnevali & I. Ramirez
and E. garciae-esquivelii Carnevali & I. Ramirez.
The first, from Andean Venezuela, is related to E.
aspera, hut has a subquadrate labellum and grows
at higher elevations. The second, from the northern
slopes of the Coastal Range, is related to E. diurna
but has smaller, differently colored flowers, a more
elliptic central lobe, and grows in coastal thorn for¬
ests.
Resum K IN . Dos nuevas especies de Encyclia
Hooker (Orchidaceae: Laeliinae) de Venezuela se
proponen, Encyclia betancourtiana Carnevali & I.
Ramirez y E. garciae-esquivelii Carnevali & I. Ra¬
mirez. La primera de estas, de la region andina,
esta relacionada con E. aspera, pero el lobulo cen¬
tral es subcuadrado y crece a mayores elevaciones.
La segunda. de la vertiente nolle de la Cordillera
de la Costa, esta relacionada con E. diurna pero
tiene flores menores y de colores diferentes, un lob¬
ulo central mas eliptieo y crece en selvas espinosas
costaneras.
Key words : Encyclia, Laeliinae, Orchidaceae,
systematics, Venezuela.
The genus Encyclia Hooker was first treated for
Venezuela by Foldats in 1970 (embedded within
the genus Epidendrum Jacquin). The genus was
then reviewed by Carnevali and Ramirez (1988, as
Encyclia) and later by Carnevali et al. (1994).
These last two treatments included within Encyclia
the species that were recently segregated as the
genus Prosthechea Knowles & Westcott by Higgins
(1998). An overall, encyclopedic view of the South
American species of the genus was presented by
Withner (2000), who essentially treated the Vene¬
zuelan species following Carnevali et al. (1994).
This author for the first time explicitly excluded the
Prosthechea taxa from Encyclia in a generic treat¬
ment. Romero and Carnevali (2000) added two En¬
cyclia species for Venezuela in their new edition of
Dunsterville and Garay’s Orchid of Venezuela: A
Field Guide. The Encyclia species from the Vene¬
zuelan Guayana were recently treated by Carnevali
and Ramirez (2003). Among all these treatments,
we can account to date for 16 Encyclia species in
Venezuela. Since there have been several range ex¬
tensions, circumscription changes, and additions in
the genus for the Venezuelan flora, we have started
a new study that will deal with these problems to
eventually result in a new revision of the genus for
the country.
In this paper, in preparation for the larger over¬
view of Venezuelan Encyclia, we propose two new
species, raising the number of known Venezuelan
members of the genus to 18. 1 lie first new species,
Encyclia betancourtiana, is the result of die recent
collection of a hitherto unrecorded member of the
E. aspera (Lindley) Schlechter group, a complex of
Andean members of the genus. I he second new
species, Encyclia garciae-esquivelii, is the result of
a study of the variation within the broad circum¬
scription of E. diurna (Jacquin) Schlechter taken
by previous authors (e.g., foldats, 1970; Carnevali
& Ramirez, 1988; Carnevali et al., 1994).
Encyclia betancourtiana Carnevali & I. Rannrez,
sp. nov. TYPE: Venezuela. Merida: Mun. Lib-
ertador, Parque Naeional Sierra Nevada, valle
intramontano en los alrededores de LI Morro,
a lo largo tie los taludes del rfo Nuestra Se-
nora, ca. 8°27'N, 71°10'W, 1800-2000 m, 20
feb. 2002, G. Carnevali 6056 (holotype, VEN;
isotype, CICY). Figure 1.
Species haec Encycliae asperae affinis seel planta parv-
iore, labello lobo centrali subquadrato, lobis lateralibus
proportione breviore latiore, petalis latiore paulo basaliter
attenuatis, disco altilamellato recedit.
Epiphytic herbs 15-20 cm tall; rhizome abbre¬
viated, thus plants ± caespitose with the pseudo-
ulbs clustered but eventually shortly creeping;
pseudobulbs 2.5-5 X 1.2-2. 5 cm, ovoid to (more
Novon 14; 413-419. 2004.
414
No von
Figure 1. Eneyclia betancourtiana Carnevali & I. Ramfrez. — A. Plant. — B. Flower. — C. Perianth segments, flattened
(based on G. Carnevali 60S 6 (isotype, CICY), drawn by Rossana Marrufo, CICY).
rarely) ovoid-suborbicular, clothed in 1 or 2 scari-
ous sheaths, apieally (I - or)2-leaved. Leaves 10-20
X 0.9— 1.9 cm, rigidly coriaceous, straight, erect,
oblong, obtuse, apex minutely unequal, blade al¬
most flat or slightly concave. Inflorescence racemose
or with a single, few-flowered branch at base, sur¬
passing the leaves, with 12 to 16 laxly arranged
flowers, the flowers spaced 15—40 mm; peduncle
and rachis purple-brown tinged, coarsely verrucu-
los e\ pedicel and ovary 17.5-19 mm long, ca. 3 mm
thick, verruculose, purple-brown tinged. Flowers
with widely spreading perianth segments, resupi-
uate. petals and sepals dark yellow-green with dark
brown suffusion toward apices, labellum with white
midlobe and with 2 or 3 purple longitudinal, raised
nerves, the apices of the lateral lobes pale yellow-
Volume 14, Number 4
2004
Carnevali & Ramirez
Encyclia from Venezuela
415
Figure 2. Encydia garciae-esquivelii Carnevali & I. Ramirez. Flowers (photograph hy G. Carnevali, based on G.
Carnevali & I. Ramirez 6811).
green, the callus white with a few purple nerves,
the column apically white, at base pale yellow-
green with 3 faint lines of line purple dots, the
anther dull [link with thick, deep dark purple mar¬
gins; sepals 7-nerved, elliptic or oblong-elliptic, ob¬
tuse or obtusely acute, the laterals very slightly
oblique, more attenuated basally than the dorsal
sepal, the apex on the external face with dense ver-
rueae and bluntly keeled; dorsal sepal 15.5—16 X
5.5—6 mm; petals 14.8—15.2 X 5—6 mm, 1.5 mm
wide at base, 5-nerved, spathulate, obtuse to ob¬
tusely acute at apex, apically with a thickened nui-
cro; labellum 3-lobed, the lateral lobes parallel and
laxly embracing the column, upon spreading form¬
ing an angle of less than 45° in relation to the main
labelar axis, 14—15 mm long along its main axis.
13—14 mm across the spread lateral lobes, midlobe
8-9 mm long from the point where lateral lobes
arise, broadly obovate in general outline including
the basalmost, callus-including section; the blade
of the midlobe 6-7 X 9.5 mm, transversally sub¬
quadrate-rhomboid, broadly truncate and shallowly
emarginated or 2-lobed at apex, the upper surface
with the nerves keeled and coarsely verruculose
with the verrueae arranged in 8 longitudinal lines;
lateral lobes 9-9.5 mm long along the external mar¬
gin (ontogenetically the most basal), 6-6.5 mm long
along the internal margin, ca. 3.5 mm wide at base.
oblong-elliptic in general outline, 6- or 8-nerved,
the lobes decurrent on the disk for 4.7—5 mm;
obliquely obtuse to round at apex; disk formed by
4 longitudinal, smooth [dates or broad keels, the 2
central ones fused in an elevated almost flat, oblong
plate, 6—6.5 mm long, the plate tridentate at apex
and with a shallow, narrowly elliptic longitudinal
depression in the central portion; the 2 outermost
keels much lower and divergent centrally; the cal¬
lus barely extending into the lower half of the cen¬
tral lobe; column 7-7.9 mm long, ca. 3.5 mm wide
at apex where it expands and gradually and slightly
attenuates toward base where it is 3 mm wide,
somewhat pandurate in outline; just below apex
provided with 2 small wings, 0.9-1. 1 mm long,
these narrowly triangular, acute, in natural position
bent toward the labellum and very laxly holding it
in position; stigmatic surface ca. 2 X ca. 2.5 mm
wide, obcordate; the clinandrium with 3 shallow1
teeth, the central ca. 0.2 mm long; anther subquad¬
rate in general position, ca. 1.5 mm long. Capsules
unknown.
Encyclia betancourtiana belongs to an Andean
complex of Encyclia characterized by coarsely ver¬
ruculose peduncles, rachis, and ovaries. 1 hese En¬
cyclia species also feature thick, elevated, more or
less flat call i with the central depression less de-
416
Novon
veloped than in other groups of llx* genus. Fur¬
thermore, the keel or nerves that emerge from the
callus and extend into the midlobe of the labellum
are thick and usually coarsely verruculose. The au¬
ricles or wings of the column are also relatively
small and usually acute. The capsules are verru¬
culose to warty in other species of the complex and
so, probably, are those of the newly proposed spe¬
cies. This Andean complex includes such taxa as
Eneyclia aspera (sensu Withner, 2000, including as
synonyms E. trachypiis Sehleehter and E. asperir-
achis Garay), E. angustiloba Sehleehter, E. microtos
(Reichenbach f.) Hoehne, and probably E. naran-
japatensis Dodson, which range from Peru to Co¬
lombia. mostly along the western declivities of the
Andes, although E. microtos is also known from the
eastern drainages in Peru. Some species of this An¬
dean complex also feature reddish peduncles and
rachises similar to the purple-brown ones seen in
E. betancourtiana. Eneyclia betancourtiana is the
first member of this complex to be recorded from
llu' Cordillera de Merida and for Venezuela.
Among the species of this Andean complex, En-
cyclia betancourtiana can be diagnosed by its trans¬
versely subquadrate-rhomboid central lobe of the
labellum (vs. orbicular or orbicular-elliptic in E.
aspera ), and the lateral lobes are proportionally
shorter (barely projecting beyond the apex of the
column in natural position vs. conspicuously longer
than the column in E. aspera), broader, and apical ly
blunter (vs. narrow and acute in E. aspera). It also
grows at much higher elevations than E. aspera.
It is interesting to note that Eneyclia betancour¬
tiana has essentially the same labellum outline as
h. andrichii Menezes, a species from Kspirito San¬
to, Brazil. In E. betancourtiana, however, the leaves
are proportionally broader (to 10.5-11 times longer
than wide vs. ea. 12.5 times as long as wide in E.
andrichii) and the labellum has fewer raised, col¬
ored nerves. We have not seen authentic material
of E. andrichii, but assume the two taxa not to be
closely related on account of the biogeograph ical
disjunction.
Eneyclia betancourtiana is apparently rare and
geographically localized. At present, it is only
known from a few epiphytic plants collected on
trees over chlfs along the Nuestra Senora River, one
of the most important rivers of Barque Nacional Si¬
erra Nevada. I liese plants were brought into cul¬
tivation in the year 2001, and a few of them flow¬
ered in 2002. The type material was prepared from
these inflorescences. The plants seem to thrive un¬
der cultivation but are difficult to bring into flower.
Eponymy. We take great pleasure in naming
this species after Armando and Armando Dimas
Betancourt, amateur orchid growers from Venezue¬
la. who first collected and flowered this species.
I hey also provided the type material.
Eneyclia garciae-esquivelii Carnevali & I. Ra¬
mirez, sp. nov. I 't PE: Venezuela. Aragua: unos
1—2 km al oeste de la cumbre de la earretera
entre Ocumare de la Costa y Cata. unos 4—5
km al F de la Ocumare de la Costa, 10°29'N,
68°42'W, Pari] ue Nacional Ht ‘iiry Pittier, en la
vertiente norte de la Cordillera de la Costa,
300-400 m; flowering under cultivation by G.
Carnevali & I. M. Ramirez in Merida. Yucatan,
Mexico, 10 June 2003, specimens prepared
from plants collected by G. Carnevali in 1986,
C. Carnevali & I. Ramirez bill I (holotype,
VEN; isotypes. AMES, CICY. MFXU, MO.
NY). Figures 2, 3.
Species liaec Encycliae diurtuie a f fin is sell planta et
floribus parvioribus, pseudobulbis 2— 3-foliatis. pelala se-
palaque bruimeis vel viridibrunneis proportione angust-
iore, labello lobo apical i proportione august iore (1.8 long-
iore quam latiore vs. 1.1 — 1., 3 latiore quam longiore) apiee
truncato vel paulo einarginato vel truneato (vs. rotundato)
reeedit.
Epiphytic or lithophytic herbs 25—40 cm tall not
counting the inflorescences; rhizome abbreviated,
thus plants caespitose with the pseudobulbs clus¬
tered. clothed in searious sheaths; pseudobulbs sub-
orbicular to suborbicLilar-pyriform, 3-5 X 3—3.5
cm, slightly less thick, apical ly 2- to 3-leaved,
clothed by 2 to 3 searious sheaths that soon dis¬
integrate. Leaves 20—29 X 1.8— 2.2 cm. linear ob¬
long. acute, apex minutely unequal, margins revo-
lute, blades often twisted 90°-180°. Infl orescences
27-70 cm long, erect, paniculate with I to 5 lateral
branches, lateral branches 6—8 cm long, 2- to 6-
(lowered, peduncle terete, relatively thin and wiry,
to 2 mm thick al base, bearing 2 or 3 tubular, close¬
ly fitting sheaths al lowermost internodes, the
sheaths 14—16 mm long, peduncle and rachis dull
brown-purple, ± densely verruculose, the verrucae
creamy white to green, 0.2-0.3 mm diam floral
bracts ea. I mm long and wide, broadly ovate, apic-
ulate, sub-fleshy when fresh, dark maroon along
and around midnerve, otherwise green. Flowers re-
supinate, with widely spreading perianth segments,
sweetly fragrant, sepals and petals dull pale green,
dull brown-maroon suffused, suffusion lighter to¬
ward the margins, particularly in the petals, the
labellum with white midlobe with 7 entire or par¬
tially broken longitudinal purple nerves at each
side of the midnerve, the lateral lobes cream white
with purple-brown longitudinal nerves on the basal
half, clear white on the apical half, the disk basallv
Volume 14, Number 4
2004
Carnevali & Ramirez
Encyclia from Venezuela
417
t _ i
5 mm
Kigure 3. Encyclia garciae-esquivelii Carnevali & I. Ramirez. Perianth segments, flattened (based on G. Carnevali
2011 (VEN), drawn by Marfa Pfa Bevilacqua, YEN). Both scales = 5 mm.
green, the callus white with some pink suffusion at
base and five faint red-purple nerves on the lower
half, column cream white at apex, green at base
with some faint purplish or maroon suffusion on the
adaxial surface on the lower half; sepals flat or
slightly concave, the dorsal ones 14.5—16 X 4.8—5
mm, elliptic-lanceolate, obtuse. 9-nerved, the lat¬
erals 15—15.5 X 6 nun. elliptic-lanceolate, obtuse,
slightly oblique, 1 1 -nerved; petals 15—16 X 5—
5.2( — 6.5) mm at their widest, attenuated below the
middle and 1—2 nun wide at base, apically concave,
spathulate-elliptic, apically obtuse, minutely apic-
ulate, thick, 5-nerved at base, i -nerved apically;
labellum 3-lobed, the lateral lobes tightly embrac¬
ing the column, upon spreading forming an angle
of less than 45° relative to the main labellar axis,
the labellum 15.5—16.5 mm total length along its
main axis, 14.5—15.3 nun wide across spread lat¬
eral lobes; midlobe 8-9 turn long from the point
where lateral lobes arise, broadly elliptic in general
outline including the basalmost, callus-including
section, the blade 7. 6-7.8 mm long, suborbicular
to elliptic-suborbieular, truncate to shallowly emar-
ginate at apex; lateral lobes 6.5-7. 5 mm long along
the external margin (ontogenetically the most bas¬
al), 5. 5-6.5 nun long along the internal margin, ca.
3-4 mm wide at base, sub-triangular to triangular
oblong in general outline, 8-nerved, the lobes de¬
current on the disk for 4. 5-4. 8 mm, obliquely
round to truncate at apex where they are ca. 2 mm
long; disk formed by four longitudinal, smooth
plates or broad keels that run parallel at base, the
two outermost lower while the higher, central ones
diverge at above the point of emergence of the lat¬
eral lobes to converge again at the base of the mid¬
lobe to form an elliptic depression ca. 2 mm long,
the callus extending into the lower half of the cen¬
tral lobe; column 8.7-9. 2 X ca. 3 mm wide at apex
where it expands and from where it gradually at¬
tenuates toward base, just below apex provided
with two small wings, 0.9— 1.1 X ca. 0.5 mm wide,
the wings oblong, apically rounded, in natural po¬
sition bent toward the labellum and holding it in
position (tightly appressed to the column); stigmatic
surface ca. 2 mm long, obovate-obcordate; anther
sub-rounded in general position, ca. 2 mm long and
wide; the clinandrium with 5 shallow teeth. Pedicel
and ovary 15-17 mm long, 3 mm diam.. subterete,
thickened on the upper half, pale greenish yellow
on the lower half, above darker, maroon-suffused
green, verruculose, the verrucae similar to those of
the peduncle and rachis but somewhat smaller and
more laxly arranged; capsules ellipsoid, verrucu¬
lose.
418
Novon
Encyclia garciae-esquivelii is closely related to
E. diurna (Jacquin) Schlechter and lias long been
thought to be only a variant of it. It was first noted
as potentially distinctive (under E. diurna) by Car-
nevali and Ramfrez (1988). It is distinct enough,
both morphologically and ecologically.
The plants of Encyclia garciae-esquivelii are
smaller with proportionally shorter leaves (mature
leaves 25—30 cm long in E. garciae-esquivelii vs.
exceeding 50 cm in E. diurna). Pseudobulbs are
often 3-leaved in E. garciae-esquivelii, while they
are almost always 2-leaved in E. diurna. The inflo¬
rescences are usually much shorter in E. garciae-
esquivelii than in E. diurna (usually longer than 1
m in fully mature plants) and the flowers are con¬
spicuously smaller (dorsal sepal 14.5-16 mm long
in E. garciae-esquivelii vs. 21-24 mm long in E.
diurna). fhe central lobe of the labellum in E. gar-
ciae-esquivelii is broadly elliptic and apically trun¬
cate, while the same structure is orbicular and api¬
cally rounded in E. diurna, thus proportionally
narrower (1.8 longer than wide in the new species
as opposed to 1.1— 1.3 times longer than wide in E.
diurna). A most striking difference lies in the col¬
oration of the (lowers, since the petals and sepals
of E. garciae-esquivelii are dull pale green, with
dull brown-maroon suffusion, while they are in¬
variably green in tint' E. diurna. The raised nerves
in the labellum of E. diurna are a faint dull purple,
while they are a deeper purple hue in the new spe¬
cies.
Besides these morphological differences, Ency¬
clia garciae-esquivelii is biogeographically distinct
from E. diurna. While E. diurna grows at elevations
of (600-)800- 1500 m mostly on the southern slopes
of the Coastal Range in areas of cloud forests or
premontaue rain forests, E. garciae-esquivelii is an
inhabitant of low-lying, xeromorphic vegetation
along the coast at elevations of less than 100 m,
often at sea level.
Encyclia garciae-esquivelii is a possibly rare,
poorly collected species, known from two sets of
populations, one in central coastal Venezuela,
where they occur as low epiphytes or lithophytes in
tropical thorn forests of north-(sea)facing slopes,
along the roads from Oeumare de la Costa and
Cata, and also along the road from Cata to Cuyagua.
The second known population is from northeast-
oriented, sea-facing c liffs near Chichirivichi in Fal¬
con, probably 80—100 km northwest of the first set
of populations, but in similar vegetation. The spe¬
cies, although possibly rare, is probably composed
ol several dispersed populations widespread along
the xerophytic belt associated with the coast of
north-central Venezuela.
V here known, Encyclia garciae-esquivelii is
commonly associated with orchid taxa common to
coastal thorn forests such as Brassavola nodosa (I,.)
Findley, Encyclia cordigera (HBK) Dressier, Myr-
mecophila humboldtii (Reichenbach f.) Rolfe, and
Oncidium baueri Findley. These forests commonly
feature columnar cacti in several genera (Subpilo-
cereus, Pilosocereus, Praecereus), and herbaceous or
succulent species such as Pitcairnia, Peperomia,
Melocactus, Anthurium (especially members of sect.
Pachyneurium), and Bromelia chrysantha Jacquin.
The low canopy is dominated by spiny legumes.
An additional population of related eneyclias,
represented by the collection Carnevali 3203
(CICY, VEN), remains a mystery. This population
comes from Sierra Imataea, in Bolfvar State in
southeastern Venezuela. The cultivated plants of
this population grown at the Caracas Botanical Gar¬
den came without more precise data. This popula¬
tion was treated by Carnevali and Ramfrez (2003)
as part of a broadly conceived Encyclia diurna. The
authors then expressed their misgivings as to this
hypothesis of relationships. This population is sim¬
ilar to E. garciae-esquivelii, but has a broader, more
rounded central lobe to the labellum, and the api¬
ces of the lateral lobes are more rounded and re¬
flexed in natural position. Otherwise, it conforms
with Encyclia garciae-esquivelii. It may represent
an additional undescribed species in the E. diurna
complex or a distinctive population of Encyclia gar-
ciae-esquivelii. The Guayanan locality, however,
suggests this population may belong to a different
species than E. garciae-esquivelii. Thus, until more
material is collected (to assess ranges of variation)
and more precise locality data are available, we
refrain from proposing an additional novelty.
Paratypes. VENEZUELA. Aragua: unos 1—2 km al
oeste del la cumbre de la carrelera entre Oeumare de la
Costa y Cata, unos 4— 5 km al E de la Oeumare de la
Costa, I0°29'N, 68°42'W, Parque Naeional Henry Pittier,
en la vertiente norte de la Cordillera de la Costa, 300-
400 m, flowering under cultivation in Maracay, Edo. Ar-
agua, 19 June 1994 ( G . Carnevali 3668, VEN); (lowering
in cultivation, 1 July 1986, G. Carnevali 2011 (CICY,
pickled; VEN). Falcon; Dist. Silva, a lo largo del horde
sur del Oolfete de Guare, al pie de las esearpas caleareas
que miran al norte, SW de Chichiriviche, W de la Cueva
de los Indios, 1-4 m, 10°54'— 55'N, 68°17'-19'W, 31 Aug.
1974, ./. A. Steyermark & II. Manara 110489 (NY, VEN);
(lowering in cultivation al the Caracas Botanical Garden,
21 July 1993, G. Carnevali 3204 (CICY, pickled; VEN).
Etymology. We are pleased lo honor Carlos
Garcfa-Esquivel of Caracas, Venezuela, with this
species. Dr. Garcfa-Esquivel, a psychiatrist, is a
world-renowned orchid species grower and one of
the most knowledgeable Venezuelans in orchid
Volume 14, Number 4
2004
Carnevali & Ramirez
Encyclia from Venezuela
419
matters. He lias collected widely in the country and
lias already been honored by several orchid names.
These include such orchids as Masdevallia garci¬
ana Luer, Epidendrum garcia-esquivelii L. Sanchez
& Hagsater. Pleurothallis garciana Luer, and oth¬
ers. The name Encyclia garciana is preoccupied by
E. garciana (Garay & Dunsterville) Carnevali & I.
Ramirez [basionym: Epidendrum garcianum Garay
& Dunsterville = Proslhechea garciana (Garay &
Dunsterville) W. Higgins) that preempted us from
using only the first of Dr. Garcias last names.
The epithet choice " garciae-esquivelii' is al¬
lowed by article 23.5 of the International Code oj
Botanical Nomenclature (Greuter et al., 2000) as a
genitive noun. The hyphen is deliberate and sup¬
ported by Article 60.9., ex. 1 1 .
Acknowledgments. The curators of AMES, MO,
NY, and VEN loaned specimens that were impor¬
tant to this research. The authors thank Gustavo
Romero (AMES) for his comments and assistance
with literature and herbarium specimens. Jose Luis
Tapia-Munoz (CICY) and Rodrigo Duno (CIGY)
provided comments on an earlier manuscript draft.
Lilia Can Itza, Silvia Hernandez Aguilar, and Kil-
ogonio May Pat (CICY) were important in specimen
processing and databasing. The authors thank The
New York Botanical Garden for access to restricted
distributional information of Encyclia taxa con¬
tained in the NYBG specimen database. Rossana
Marrufo at CICY helped the authors with the ico¬
nography of the article.
Literature Cited
Carnevali, C. & 1. M. Ramfrez. 1988. Revisidn tie h.ncy-
clia Hook. (Orchidaceae) para Venezuela. Bol. Comite
Orquideol. Soe. Venez. Ci. Nat. 23: 13—87.
- & - . 2003. Encyclic Hook. Pp. 317—324 in
G. Carnevali, 1. Ramfrez, G. Romero, C. Vargas & h.
Foldats: Orchidaceae. Pp. 200—619 in J. A. Steyermark,
P. L. Berry, K. Yatskievich & B. k. Holst (editors), flora
of the Venezuelan Guayana, Vol. 7. Missouri Botanical
Garden Press, St. Louis.
- , - & G. Romero, 1994. Orchidaceae Dun-
stervillorum VIII: New Encyclia species and combina¬
tions from Venezuelan Guayana. I.indleyana 9: 59—70.
Foldats, E. 1970. Epidendrum. In Orchidaceae, flora de
Venezuela 15(3): 140-447.
Greuter, W„ J. McNeill. F. B. Barrie, H. M. Burdet, V.
Demoulin, T. S. Filgueiras, 1). 11. Nicolson, P. C. Silva,
J. K. Skog, P. Trehane, N. J. Turland & 1). L. Hawk-
sworth (editors). 2000. International Code ol Botanical
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Higgins, W. F. 1997. A reconsideration of the genus Pros-
thechea (Orchidaceae). Phytologia 82: 370—383.
Romero, G. A. & G. Carnevali. 20(H). Orchids of Vene¬
zuela: An Illustrated Field Guide, 2nd ed. Armitano
Editores, Caracas, Venezuela.
Withner, C. L. 20(H). The Cattleyas and their Relatives.
VI: The South American Encyclia Species. Timber
Press, Portland.
A New Vining Species of Swartzia (Fabaceae, Swartzieae) from t lie
Venezuelan Amazon
Nidi a L. Cuello A.
UNELLE/-Guanare, Progrania de Rent rsos Naturalcs Renovables, Herbario Universitario
(PORT), M( *sa de C ^avacas, estado Portuguesa 3323, Venezuela, neuello@canlv.net
Abstract. Swartzia barnebyana, a new legume
species with a viuing habit from the state of Ama¬
zonas in Venezuela, is described and illustrated.
I he new species belongs to section Possira , series
Unifoliolatae, in which it is closely related to both
S. floribunda Spruce ex Benlham and S. iniridensis
B. S. Cowan, by its intramarginally interconnected
primary veins and its densely strigulose-velutinous
pubescence on branchlcts, inflorescence axes, and
external calyx segments. Swartzia barnebyana may
lie distinguished from both related species mainly
by the type and color of the ovary pubescence,
shape and size of leaflet blades, number of calyx
segments, and absence of pubescence in the inter¬
nal surface of the calyx.
RKSUMKN. Se describe e ilustra Swartzia barne¬
byana , una especie nueva de leguminosa con ha-
bito trepador del estado Amazonas, Venezuela. La
especie nueva perteneee a la seceion Possira, serie
Unifoliolatae dentro de la eual se relaciona estre-
chamente con S. floribunda Spruce ex Bentham y
S. iniridensis R. S. Cowan por las venas primarias
interconeetadas intramarginalmente y la pubescen-
eia densamente estrigulosa-velutina sobre ramitas,
eje de las infloresceneias y segmentos extemos del
caliz. Swartzia barnebyana se distingue de las es-
pecies relacionadas, principalmente por el tipo y
color de la pubesceneia en el ovario, la forma y
tamano de la lamina de los folfolos, el mimero tie
segmentos del cdliz y la auseneia de pubesceneia
en la superficie interna del ealiz.
Key words: Fabaceae, section Possira , series
Unifoliolatae . Swartzia, Venezuelan Amazon.
During preparation of the treatment of Swartzia
Schreber for the Horn of the Venezuelan Guayana
(Cuello & Cowan. 1999), a number of undeter¬
mined taxa were set aside for further study. One of
these, represented in the mentioned treatment as
“Swartzia sp. E” (1999: 405), turns out to be an
undescribed species for which a name and descrip¬
tion are provided in this article.
Novon 14: 420-423. 2004.
Swartzia barnebyana Cuello. sp. nov. TYPE: Ve¬
nezuela. Amazonas: Atures, Serram'a Batata.
55 km al SE Puerto Ayaeueho, 2 km al NE de
Salto Colorado, Cano Colorado sobre eorazas
latent icas, 5°33'N, 67°08'W, 550 m, Oct. 1989
(fl), Elio Sanoja, Angel Fernandez & Manuel
Ydnez S37I (holotype, PORT; isotypes, MO,
VEN). Figure I.
Ilaec species venis priniariis prope marginem conju-
galis atque ramulis infforeseentiae axihus cl calycis scg-
mentis extus dense velutinis ad Swartzia floribundarn cl
■S. iniridensem comparabilis, sed a prima foliolis marginc
planis (nee revolutis) et ovario venlralitcr pubescente (nee
glabro), a seeunda ovario in uno marginc barbato trichom-
antihus infuseatis (nee scriceo), al) ambabus calycis seg-
mentis intus glabris et foliolis multo majoribus diversa.
Vine or sarmentose tree; branchlcts terete,
densely strigulose-velutinous; stipules caducous,
3-4 mm long, narrowly lanceolate, acute, densely
pilose-velutinous; leaves unifoliolate. petioles 5—6
mm long, densely strigulose, stipellate apically, sti-
pels caducous, 1 — 1.5 mm long, subulate to trian¬
gular, petiolules 3—5 mm long, 3—5 mm wide,
densely pilose-velutinous, leaflet blades broadly
oblong (18— )24-40 X 9—14 cm, the margin plane,
the base rounded, the apex obtuse to acuminate,
the acumen 12—15 mm long, glabrous on upper sur-
laee. densely strigulose to glabreseent beneath with
the base of the costa densely strigulose beneath,
the costa impressed, the primary veins plane on the
upper surface, costa and primary veins salient be¬
neath, the primary veins interconnected intramar¬
ginally. Inflorescences axillary racemes, sometimes
secondarily becoming branchlcts with young
leaves, also bearing axillary racemes, axes 5-10 X
0.2-0. 4 cm, densely strigulose-velutinous, brown,
bracts persistent. 3—6 X 1.5—2 mm, triangular to
lanceolate, sometimes developing into leaves with
blades 2—3 X 1 — 1.2 cm; pedicels at anthesis 15—
20 mm long, in flower buds 7-15 X 1.5-2 mm,
densely strigulose-velutinous. Flower buds ovate-
oblong in outline. 9-10(13) mm long, 7-8(10) mm
wide, densely strigulose-velutinous; calyx segments
5. persistent, densely strigulose-velutinous exter-
Volume 14, Number 4
2004
Cuello
Swartzia barnebyana from Venezuela
421
Figure 1. A per ion of a branch of Swartzia barnebyana Cuello with inflorescences. (Drawn from the type Sanoja,
Ferndndez & Ydnez 3371.)
nully. glabrous within; petal white, the claw 4—5
mm long, the blade orbicular, 15—1 7 mm diam.,
glabrous; larger stamens 3, the filaments white, 15-
18 mm long, the anthers yellow, 4-5 mm long, ca.
2 mm wide, oblong-lanceolate, medium-sized sta¬
mens 2. opposite the larger stamens, the filaments
12—14 mm long, the anthers 2—3 mm long, ca. 1
mm wide, the smaller stamens ca. 220. the f i la-
422
Novon
Table 1. Characters
that distinguish Swartzia barnebyana from its closest relatives.
.S’, barnebyana
.S’, iniridensis
.S’, floribunda
Habit
sarmentose tree or vine
tree 3—4 m
shrub 1 .5—3 m
Inflorescence axes
2—4 mm wide, densely stri-
1-2 mm w ide, densely
2—3 mm wide, densely
Bracts
gulose-vel ut i nous, dark
brown
velutinous, sericeous
strigulose-velutinous,
light brown
triangular to lanceolate, 3-
6 mm long
lanceolate, ca. 1.5 mm
long
linear. 1.5-2 mm long
Pedicels
15-20 mm long
9- 1 5 mm long
(6)10-15 mm long
Anthers of small stamens oblong, apex emarginate to
slightly mucronate, 1-1.2
mm long
oblong to lanceolate
oblong-ovate, apiculate,
1-2 mm long
Ovary stipe
11-17 mm long
ca. 9.5 mm long
9-15 mm long
merits 8— 10 mm long, the anthers oblong, with the
apex emarginate to slightly mucronate, 1-1.2 mm
long, ea. 0.5 mm wide; gynoecium with t lie ventral
suture barbate, glabrous elsewhere, stigma puncti-
lorm, style 4-6 mm long, ovary elliptic-obovate, lat¬
erally flattened, 0— 8 mm long, 3.5—5 mm w ide, ova¬
ry stipe white, I 1-17 mm long. Fruits not seen.
Distribution and ecology. Swartzia barnebyana
is known only from the type and paratype collec¬
tions. According to information recorded on speci¬
men labels, the new species is a small vining tree
or vine occurring in nonflooded forests at 500-600
m elevation in Cano Colorado in the Serranfa Ha-
tala. which belongs to the Cuao-Sipapo massif in
the Venezuelan Amazon. It has been collected in
flower during September and October.
The genus Swartzia comprises about 150 species
distributed in two sections, Swartzia Cowan and
Possira (Aublet) DC., according to Cowan (1967).
Swartzia barnebyana belongs in Cowan’s (1967)
section Possira , series Unifoliolatae, based on its
axillary inflorescences, simple racemes, the pres¬
ence of bracts that are obviously reduced leaves,
sometimes with leaflets evident, and unifoliolate
leaves.
Due to its intramarginally interconnected pri¬
mary veins and its densely strigulose-velutinous
pubescence on branchlets, inflorescence axes, and
external calyx segments, the new species Swartzia
barnebyana is clearly related to both S. floribunda,
from the Venezuelan Amazon, and S. iniridensis,
from Colombia (Cowan, 1967). In contrast to the
glabrous gynoecium of S. floribunda, the new spe¬
cies. like S. iniridensis, has the ventral part of the
ovary pubescent. I lie ovary pubescence in die new'
species consists of sparse barbate brown hairs,
while in S. iniridensis it is densely appressed se¬
riceous. The new species also differs from the two
mentioned relatives by the glabrous inner surface
of the 5-segmented calyx. In contrast, S. floribunda
has a calyx with 4 to 6 segments, villose but gla-
brescent within, and S. iniridensis has 4 segments,
sericeous within. Additionally, .S’, barnebyana has
the blades of the unifoliolate leaves broadly oblong
and much larger |( 18)24-40 X 9-14 cm] than the
oblong to elliptic-oblong leaflets of both S. floribun¬
da [(8)10.5—19(24) X 3. 5-6(9) cm] and .S’, iniriden¬
sis [ lf>— 22 X 5.5—8 cm]. Also, .S’, barnebyana has
plane leaflet margins and S. floribunda has revolute
margins. Other features considered relevant in dis¬
tinguishing S. barnebyana from the related species
are shown in Table 1.
The combination of vining or sarmentose habit
and unifoliolate leaves suggests a superficial rela¬
tionship between the new species and Swartzia pal-
ustris Barneby of the Venezuelan Amazon (Barneby,
1991); however, 5. barnebyana differs greatly from
.S’, palustris, which has glahrate branches and
densely golden sericeous inflorescences. The latter
species also has smaller, differently shaped leaflets,
smaller, ovate bracts (scarcely 1 mm long), smaller
globose flower buds (4-5 mm diam.), and a shorter
ovary stipe, among other differences.
1 am honored to name this species after Rupert
Barneby, a great gentleman who dedicated much of
his life to the study of legumes and contributed
greatly to our knowledge of the genus Swartzia in
the Venezuelan Guayana.
Paratype. VENEZUFLA. Amazonas: Atures, Serran¬
fa Batata, 55 km al SE de Pto. Ayacucho, 2 km al NE de
Salto Colorado, Cano Colorado sobre eorazas laterfticas,
5°33'N, 67°08'W, 550 m, Sep. 1989 (fl), Elio Sanoja, An¬
gel Ferndndez A1 Manuel Ydnez 3200 (PORT, MO, YEN).
Volume 14, Number 4
2004
Cuello
Swartzia barnebyana from Venezuela
423
Acknowledgments. I am grateful to Bruno Man-
ara (VEN) for preparing the illustration. 1 thank
Benjamin Torke (MO) and Gerardo Aymard (PORT)
for their comments on an early version of the man¬
uscript and Roy Gereau (MO) for help with the Lat¬
in diagnosis. Special thanks are given to \ ictoria
Hollowed (MO), Ronald Liesner (MO), Benjamin
Torke, and Michael Vincent for critical comment
improving the manuscript.
Literature Cited
Barneby, R. C. 1991. Notes on Swartzia (Leguminosae:
Swartziae) preliminary to the Flora of the Venezuelan
Guayana. Ann. Missouri Bot. Card. 78: 177—183.
Cowan, K. S. 1967. Flora Neotropica Monograph No. I.
Swartzia (Leguminosae, Caesalpinioideae, Swartzieae).
Hafner, New York.
Cuello, N. L. & R. S. Cowan. 1999. Swartzia (Fabaceae).
Pp. 394—415 in P. E. Berry, B. K. Holst & K. Yatskiev-
ych (editors). Flora of the Venezuelan Guayana, Vol. 5.
Missouri Botanical Garden Press, St. Louis.
New Combinations in South American Phaseolinae: Dolichopsis
and Macroptilium (Fabaceae: Phaseoleae)
Alfonso Delgado- Salinas and Leticia Torres-Colin
Departamento de Botdniea, Institute) de Biologfa, UNAM, Apartado Postal 70-233, 04510
Mexico, D.F. Mexico, adelgado@servidor.unam. nix; lety@ihiologia.unani.nix
Alls lit ACT. I he new combination Dolichopsis lig-
ulata (Piper) A. I)elgad< ) is made from Phaseolus
ligulatus Piper. Iliis species is readily distin¬
guished from Dolichopsis paraguariensis by ils pre¬
senting the beak ol the keel witli a sigmoid cur¬
vature and by its western distribution in Bolivia and
adjacent Paraguay. The new combination Macrop-
tilium ecuadoriense (Hassler) L Torres-Colin & A.
Delgado is made from Phaseolus atropurpureus Ses-
se & Mocifio ex DC. var. ecuadoriense Hassler, with
Phaseolus affinis newly synonymized. Bed petals
borne from a tubular calyx and presence of hairs
in its wing petals readily distinguish Macroptilium
ecuadoriense from the common and widespread Ma-
croptilium atropurpureum with its deep maroon pet¬
als borne from a campanulate calyx and glabrous
wing petals.
Key words: Chaco, Dolichopsis, Ecuador, Fa-
baeeae, Macroptilium, Pantanal, Phaseoleae, Phas¬
eolinae, Phaseolus.
I he native New World subtribe Phaseolinae
comprises nine genera: Dolichopsis Hassler, Ma¬
croptilium (Bentham) Urban, Mysanthus G. P. Lewis
& A. Delgado, Oryxis A. Delgado & G. P. Lewis,
Oxyrhynchus Brandegee, Phaseolus L„ Ramirezella
Bose, Strophostyles Elliott, and Vigna Savi. Of
these, the most diversified in terms of species is
Phaseolus, and in the past, several species of dif¬
ferent extant genera were originally described un¬
der it. During a phylogenetic study of Pliaseo/us
(Delgado-Salinas el al., 1999), it became evident
that the taxonomy of most genera needed to be
worked out, and thus, the nomenclature of some
requires changes, as follows:
Dolichopsis 1 1 assi .kk
I he genus Dolichopsis as currently known com¬
prises one species. Dolichopsis paraguariensis. I bis
species was described by Hassler (1907), and sub¬
sequent collecting suggested that it was somewhat
broadly distributed in northern Argentina, Para¬
guay, Bolivia, and southwestern Brazil. Dolichopsis
Novon 14: 424-427. 2004.
paraguariensis is ecologically and geographically
unique among species of Phaseolinae by being re¬
stricted to inundated areas of the Chaco and adja¬
cent wetlands, such as the Pantanal, Mato Grosso,
and the I ueuman-Bolivian forest, and Ixiamas (Ba-
mella : 187.
1944. Syn. nov. 1 'i PE: Paraguay. Valenzuela (entre
Villa Rica y Asuncion), eampos-esteros (pantanosos),
mar. 1942, T. Rojas 0537 (holotype, SI not seen).
Although Piper (1926) stated that the type col¬
lection of Phaseolus ligulatus was made by Orland
E. W bite, the labels on both the holotype and iso¬
type indicate that the collector was Martin Carde¬
nas (his number 2000). According to Busby (1927),
both botanists collected together at Ixiamas. This is
further indicated by Busby (1927: 272). who de¬
scribed Mascagnia macrophylla from a collection
at Ixiamas by M. Cardenas (his no. 1943) on 18
December 1921. and Dianthera graminifolia
(1927: 367) from a collection at the same locality
by O. E. White (his no. 2310). collected on 19 De¬
cember 1921.
The habit, foliage, and fruit of Dolichopsis ligu¬
lala, and to a lesser extent the inflorescence, are
Volume 14, Number 4
2004
Delgado-Salinas & Torres-Coh'n
South American Phaseolinae
425
similar to those of I), paraguariensis. The flowers
of the two species, however, differ in a number of
features. Most noticeably, the keel petals of I). lig-
ulata have a beak that is loosely and somewhat
sigmoidally curved, and the style enclosed within
has an identical curvature, is distally thickened,
and bears a highly compact pollen brush 2 mm
long. In contrast, the incurved and upwardly as¬
cending keel in I). paraguariensis encloses a style
that is distally tapered and bears a loose pollen
brush for well over 2 mm long. The stigma is ter¬
minal to oblique in both species, but positioned
toward the front of the flower in D. paraguariensis
due to style rotation. In D. ligulata, the style rotates
laterally toward the right side of the flower. The
fruits of both Dolichopsis species are broad, about
1 cm wide, and laterally compressed. The seeds are
transversely oriented to the fruit length at maturity,
rendering a flattened and wider pod that is distinct¬
ly unlike any other pod in the subtribe Phaseolinae.
Distribution and habitat. Dolichopsis ligulata
occurs in Bolivia and adjacent western Paraguay,
and inhabits savannas or shrublands, usually in
mud or standing water around the margins of sea¬
sonal or semi-permanent ponds often associated
with grasses, sedges, and such species as Tabebuia
insignis. Flowering takes place from September to
March, during the rainy season. Whereas plants of
I). paraguariensis can be found in the same eco¬
logical setting as I). ligulata, there have been no
reports of svmpatry.
Specimens examined. BOLIVIA. Beni: Prov. Yaouma,
Estacion Biologica del Beni, cerea de la Laguna Norman-
dia, 19 Aug. 1991, Beck 18853 (NY); alrededor del Porv-
enir, 21 Sep. 1997, Beck 24296 (MEXU); near Lake Ro-
gagua. 1 Nov. 1921. Rushy 1639 (NY). PARAGUAY. San
Pedro: Alto Paraguay. Primavera, 14 June 1959, Wool-
ston 1097 (NY ).
Macroptiijum (Bentham) Urban
The genus Macroptilium comprises 18 species
distributed from the southwestern United States to
northern Argentina. The species are mainly climb¬
ers or creeping herbs, often confined to tropical and
subtropical habitats. Fifty-five percent of all taxa
are endemic to South America (Torres-Colfn, un¬
published).
Macroptilium is characterized in part by an in¬
dumentum of simple hairs, inflorescences w ith nec¬
tariferous nodes, fascicles of bracts at the base of
the inflorescence, caducous floral bracts and brac-
teoles, corollas with conspicuous large wing petals,
the left-handed one occupying the position of the
standard, and keel petals with an incurved, distally
hooked beak. In contrast, the closely related genus
Phaseolus differs in all of these regards, from un¬
cinate hairs and non-nectariferous inflorescence
nodes, to persistent floral bracts and laterally coiled
keel beaks. Since Urban (1928) raised Phaseolus
sect. Macroptilium to the generic level, species of
Phaseolus continue to be reassigned to Macroptil¬
ium in both hemispheres. Recently, two new com¬
binations have been proposed, one in South Amer¬
ica (I)rewes & Palacios, 1994), and another in
Mexico (Delgado-Salinas . Capitulum. c.
Central floret. — d. Style branches. — e. Apical portion of the pappus bristles. — f. Peripheral floret with the lobes
recurved. — g, h, i. Details of the anthers showing the caudate bases (i) and the apical appendage (h). Drawn from the
holotype at MEXU (Gonzalez- Medrano et al. 17920).
436
No von
Helietta parvifolia (A. Gray ex Hemsley) Bentham
(Rutaceae), Leucophyllum ambiguum Bonpland
(Scrophulariaceae), Machaonia coulteri (Hooker f.)
Standley (Rubiaceae), and Turnera diffusa Will-
denow ex Schultes (Turneraeeae).
The new species is outstanding among the other
Mexican species of the genus because of its larger
capitula (involucre 2 cm or more wide), solitary or
in a cluster of 2 or 3 by branch, sessile and covered
at the I >ase by the leaves. All the other known Mex¬
ican species have smaller capitula (involucre 1.5
cm or less wide), generally arranged in glomerules
or congested cymes. Another feature of this new
species is the number of florets per capitulum (2(H)
or more); no other Mexican species has capitula
with this many florets, generally counting 60 or
less.
By its number of florets and the arrangement of
the heads, Gochnatia hiriarliana belongs to Goth-
natia sect. Glomerata S. E. Freire, L Katinas & G.
Saneho (Freire et ah, 2002). Among the features
that characterize the members of this section are
their heads solitary or 2 to 3 by branch, the number
of phyllary series (mostly 5 or 6), and the pappus
with several short outer bristles. It differs from the
other species of the section (all Mexican) bv its
large number of florets per capitulum: this new spe¬
cies has more than 200 florets, while the other spe¬
cies rarely reach 60 per capitulum.
Etymology. The epithet honors Patricia Hiriart-
Valencia (1947—1991), botanist and student of the
flora of the dry regions, especially from central
Mexico where this new species was found.
Paratype. MEXICO. Hidalgo: Arriba de Chalmita, 2
Nov. 1946, F. Miranda 1027 (MEXU).
Kky to the Mexican Species of Gochnatia
la. Florets 200 or more per capitulum; involucre 2
cm or more wide; slate of Hidalgo . . G. hiriarliana
Medrano, Villasenor & Medina
Ih. florets 60 or fewer per capitulum; involucre 1.5
cm or less wide.
2a. florets 10 or more per capitulum; involucre
5 mm or more wide.
3a. Leaf blade 3.0 cm or less long; state of
Puebla . G. purpusii Brandegee
3b. Leaf blade 3.5 cm or more long.
4a. florets 25 or less per capitulum;
state of Baja California Sur . . .
. G. arborescens Brandegee
4b. Florets 35 to 60 per capitulum;
slates of Guanajuato, Hidalgo, Que-
rdtaro. San Luis Potosf, Tamaulipas
. G. magna M. C. Johnston
21 >. florets 8 or fewer per capitulum; involucre
1 mm or less wide.
5a. Capitula iu terminal rounded glomer¬
ules; leaves gray pubescent abaxially;
state of Oaxaca .
... 6. smithii B. L. Robinson & Greenman
5b. Capitula not in rounded glomerules,
mostly in axillary or terminal glomerate
cymes; leaves mostly yellow-pubescent
abaxially.
6a. Leaf blade 2.0— 5.0 cm long, gen¬
erally lanceolate or elliptic-lanceo¬
late; involucre 5-6(-7) mm long;
Chihuahuan Desert, from Texas to
the states of Hidalgo and Querdtaro
. G. hypoleuca (DC.) A. Gray
subsp. hypoleuca
6b. Leaf blade 0.4— 2.0 cm long, ovate
or elliptic ovate; involucre 6—8 mm
long; states of Oaxaca, Puebla, Ve¬
racruz . . . G. hypoleuca (DC.) A. Gray
subsp. obtusata (S. E. Blake)
Cabrera
Acknowledgments. We thank Albino Lima for
the drawing, and Fernando Gliiang and two anon¬
ymous reviewers for the critical review of the man¬
uscript.
I .iterate re Cited
Bremer, k. 1994. Asteraceae: Cladistics and Classifica¬
tion. Timber Press, Portland, Oregon.
Cabrera, A. L. 1971. Revisidn del gdnero Gochnatia
(Compositae). Bevista Mas. La Plata, Nueva Serie 12:
1-160.
freire, S. E., L. Katinas & G. Saneho. 2002. Gochnatia
(Asteraceae, Mutisieae) and the Gochnatia complex:
Taxonomic implications from morphology. Ann. Missou¬
ri Bot. Gard. 89: 524-550.
Salvia fairuziana (Lamiaceae), a New Species from Lebanon
Ricardus M. Haber
Institute for Conservation of Biodiversity and Ecotourism Development (USER University),
P.O. Box 967, Jounieh, Lebanon, fon@sodetel.net. lb
Myrna T. Semaan
Center for Environmental Development, Awareness & Research (C.E.D.A.R.), P.O. Box 967,
Jounieh, Lebanon
Abstract. A new species of t he genus Salvia L.
(Lamiaceae) is described from die western slopes
of Mount Lebanon characterized by a Mediterra¬
nean climate. Salvia fairuziana R. M. Haber & Se¬
maan is distinguished by the following characters:
perennial herb; hispid with dense tuberculate mul-
liseptate trichomes emerging from wart-like turgid
purple epidermal cells; leaves ovate-elliptic, cor¬
date, shallowly lobed, verrucose; floral bracts dis-
coidal. green with a white center and purple tips;
verticillasters crowded; calyx 1.0— 1.2 cm long,
green, infundibular; corolla 2.4-2. 7 cm long, lilac,
squamulate, ventricose; staminal connectives do-
labriform.
Key words: biodiversity, endemic, flora. Lami¬
aceae, Lebanon, Mediterranean.
According to Mouterde (1983), Lebanon pos¬
sesses 18 species within the genus Salvia (Lami¬
aceae). Most of these have a wide distribution range
in die Mediterranean region or occur eastward into
Iran and Iraq (Hedge, 1982). Four species, how¬
ever, are restricted to the eastern Mediterranean
l Post, 1932; Mouterde. 1983). Salvia rubifolia Bois-
sier, S. viscosa von Jacquin, and .S. pinardii Boissier
are limited to Lebanon and Syria; S. judaica Bois¬
sier occurs in Lebanon, Syria, and Palestine. Salvia
hierosolymitana Boissier has a similar distribution
lo the latter, but also occurs in Jordan (Al-Eisawi,
1998) and northern Cyprus (Greuter & Kaus,
1984).
Salvia fairuziana R. M. Haber & Semaan, sp. nov.
TYPE. Lebanon. Kesrouan Province: Kfour
village, 770 m, 34°02'09.6"N, 35°41'10.1"E,
17 May 2002, R. Haber A' M. Semaan 2002
(holotype, MO; isotypes, BEL K). Figure I.
Herba perennis; indumentum densum hispidum, un-
usquisque trichoma tuberculata multiseptata in tumulo ex
epidermide turgido purpurato; folia verrucosa, ovato-ellip-
lica, basi cordata, lobis lateral i bus non profundis; brarteae
florales virides, centris albidis; verticillastri eongesti; ca¬
lyces 1.0— 1.2 cm longi, infundibulares, lobis non effusis
in fructibus; corollae 2.4-2. 7 cm longae cum squamulis,
ventricosae, labiis lilacinis; connectiva dolabriformia.
Perennial herb 50-80 cm high, with a distinctive
and pleasant fragrance, mostly single-stemmed;
stem 0. 0-0.7 cm wide at base, simple, hispid with
dense tuberculate and multiseptate trichomes, each
emerging from a wart-like mound of turgid, purple
epidermal cells, and also pilose with trichomes
mostly arising from purple bases on the lower stem
to short glandular on the upper. Basal leaves 15.0-
18.5 X 6.5-8.0 cm, petiolate; petiole 5.5— 6.5 cm
long, pilose, with tuberculate multiseptate tri¬
chomes arising from purple mounds, with the in-
dument extending to the midrib below; blade ovate,
obtuse, cordate, shallowly lobed, erose, much ver¬
rucose above with multiseptate, tuberculate lanate-
arachnoid trichomes mainly at the tips of verrucae,
reticulate below with multiseptate, variably tuber¬
culate lanate-arachnoid trichomes; cauline and flo¬
ral leaves 3.0-12.5 X 2. 3-6.0 cm, with indumen¬
tum similar to the basal leaves; the lower leaves
shortly petiolate to sessile, ovate-elliptic, obtuse,
cordate, unequal at base, shallowly lobed, sinuate,
erose; upper leaves amplexicaul, ovate-lanceolate,
acute-acuminate, cordate, unlobed, very sinuate,
slightly undulate, erose; floral bracts 1.0— 2.6 X
0.5— 2.3 cm, the lower orbicular-reniform, acumi¬
nate, cordate, sinuate, erose, glabrous above,
smooth at center, lanate-arachnoid and verrucose at
the periphery, lanate below with tuberculate mul¬
tiseptate trichomes, the upper bracts ovate, acu¬
minate, entire, ciliate, glabrous to pubescent with
tuberculate short, stout septate trichomes, diseoidal
with a white center and a green margin sometimes
tinted purple especially at the tips of the flowering
branches, the margin with some glandular tri¬
chomes above, but with arachnoid and short tuber¬
culate trichomes below. Panicle commencing from
the second or third stem node, 15—30 cm long,
Novon 14: 437-439. 2004.
438
Novon
figure I. Salvia Jairuziana It. M. Haber & Semaan, photos from the typified collection taken by the authors in Kfour
village. — A (left). Whole Plant. — B (right). Verticillasters.
erect, densely covered with purple-based tubereu-
late multiseptate and short glandular triehomes;
panicle branches up to 15, each 7-10 cm long with
5 to 7 verticillasters, sometimes with secondary
branches, the lowermost usually aborted; verticil¬
lasters crowded, each 2- to 6-flowered, with (lowers
longer than the internodes. Flowers 2.6-2. 8 cm
long; pedicels 3 mm long with dense glandular tri¬
ehomes and short stout tuberculate-septate eglan-
dular triehomes; calyx 1. 0—1.2 cm long, infundib¬
ular. 13-veined, green, with a purple tint that
increases gradually in density from the lower part
to the upper part of the panicle; tube 6 mm long,
with dense glandular triehomes and short rigid tu-
bereulate-septate eglandular triehomes especially
on veins, pi lost- inside; lower li/) with two teeth 5—
6 mm long, triangular-subulate, with mucro 1.5 mm
long; upper lip 5.5 X 3.0 mm. truncate, tridentate
with the middle tooth shorter, flat, concave; teeth
muoronate, ciliate with short rigid tuberculate-sep¬
tate triehomes; corolla 2.4-2. 7 cm long, white to
faintly lilac in bud. lilac-violet in flower; tube 8 mm
long, white, ventricose, squamulate, with an inter¬
nal fimbriated scale on squamula; lower lip 8—10
mm long, with short triehomes and sessile glands;
upper lip 17—10 mm long, very falcate, emarginate,
with short tuberculate-septate eglandular tri¬
ehomes, glandular triehomes, and sessile glands;
pistil 36 mm long, exserted by 5-7 mm, lilac to
violet at tin* lip; stigma bifid, unequal; stamina!
connectives clearly longer than, and articulating
with, the filaments, anthers and connectives 22 mm
long; lower theca 2.9 mm long, sterile, reduced to
a dolabriform plate, with a tooth running down the
plate. Seed 2.2— 2.3 X 1.7— 1.8 mm, brown, finely
tuberculate, with darker venation, ovoid, rounded
on the outside, angled on the inside and with a
rounded apex.
Distribution and habitat. Salvia Jairuziana is
local and relatively rare. 1 he authors observed a
population comprising a few hundred specimens
extending between 700 and 900 m altitude near
Kfour village. There it grows among calcareous
Volume 14, Number 4
2004
Haber & Semaan
Salvia fairuziana from Lebanon
439
rocks in Mediterranean red soil in an open Quercus
coccifera L. grove (l)e Vaninas, 1934).
Derelict terraces where Salvia fairuziana grows
indicate that the region was previously managed for
agriculture. This may have allowed the plant to sur¬
vive unnoticed and to regain a foothold as the ter¬
races were abandoned several decades ago. How¬
ever, urban sprawl and encroaching development
on the mountainside have destroyed large expanses
of the original habitat and remain a serious threat
to the extant population. 1 he flowering period of
Salvia fairuziana has been observed by the authors
to extend from May to June.
Salvia fairuziana most closely resembles S. scla-
rea L., S. indica L, S. chrysophylla Stapf, and 5.
argentea L. based on its dolabriform articulate sta¬
mens. squamulate and ventricose corolla tube, and
its lilac-colored corolla upper lip. However, S. fai¬
ruziana differs from S. sclarea in having green, not
pink-mauve, bracts. While the hood and lower lip
of the corolla are similarly colored lilac in S. fai¬
ruziana, the lower lip is variably colored in S.
indica , 3’. chrysophylla, and S. argentea , being dark
violet, yellow, and white, respectively. In addition,
the latter three have lax verticillasters, whereas
those of S. fairuziana are congested.
Salvia fairuziana also differs markedly from oth¬
er eastern Mediterranean Salvia species existing in
Lebanon. Salvia ruhifolia anil S. pinardii occur in
the continental semi-arid habitat while S. viscosa,
S. hierosolymitana, and S. judaica may occur in
habitats and climatic conditions similar to those of
S. fairuziana. Salvia hierosolymitana has a red co¬
rolla and calyx, S. judaica a violet corolla (10 mm
long) and red calyx, and S. viscosa a pink to pale
violet corolla (15 mm long). In contrast, 5. fairu¬
ziana has a green calyx and lilac corolla (24—27
mm long).
Salvia fairuziana is named, on behalf of the mil¬
lions of admirers, as homage, gratitude, and respect
to the Lebanese operatic diva. Lady Fairuz. who
has, for the past 50 years, enchanted the country,
inspiring humaneness, patriotism, and respect for
nature.
Paratype. LEBANON. Kesrouan Province: Kfour
village, 755 m. 34°01'48.3"N, 35°40'39.2"E, 8 June 2002.
R. Haber & M. Semaan 2002 (MO).
Acknowledgments. The authors thank the Khal¬
il F’attal and Sons Establishment for financially
supporting research projects of the Friends of Na¬
ture NGO. The “Wild Flora Project” has already
resulted in the discovery of several probable new
species and many other taxa previously unaccount¬
ed for in the flora of Lebanon.
Literature Cited
Al-Eisawi, I). M. H. 1998. Salvia hierosolymitana Boiss.
P. 173 in Field Guide to Wild Flowers of Jordan and
Neighbouring Countries. Jordan Press Foundation *‘A1
Rai,” Amman.
De Vaumas, E. 1954. La structure et le relief du sol. Pp.
17-83 in Le Liban. Firmin-Didot, Paris.
Greuter, W. & T. Raus (editors). 1984. Med-Checklist No-
tulae, 8. Willdenowia 13.
Hedge, I. C. 1982. Salvia L. Pp. 400-461 in P. II. Davis,
Flora of Turkey, Vol. 7. Edinburgh Univ. Press, Edin¬
burgh.
Mouterde, P. 1983. Salvia I- Pp. 155-171 in Nouvelle
F lore du Liban et de la Syrie, Vol. 3. Dar El-Machreq,
Beirut.
Post, G. 1932. Salvia L. Pp. 347—370 in Flora of Syria,
Palestine and Sinai, Vol. 2. American Press, Beirut.
A New Species of Phegopteris (Thelypteridaceae) from Korea
Chul Hwan Kim and Byung-Yun Sun *
Faculty ol Biological Sciences, Chonbuk National University, Chonju, Chonbuk 561-756,
Korea, plantaxa@hanmail.nel; *author for correspondence: sunby@chonbuk.ac.kr
Soo Hyun Park
I he National Arboretum, Jikdong-ri 51-*, Sohul-myon, Pochun-gun, Gyunggi 487-821.
Korea, natpark@hanmail.net
Austkact. A new species of fern, Phegopteris k<>-
reana ( I helypteridaceae), was collected from south¬
ern Korea and is described here. Phegopteris ko-
reana is most similar to P. decursive-pinnata
morphologically; however, the new species consis¬
tently shows discontinuous morphological gaps
from the latter in the shape of the fronds, pinnae,
and ultimate segments, venation pattern in ultimate
segments, tufted trichomes in the sori, and spore
wall sculpturing.
Key words: Korea, Phegopteris, Thelypterida¬
ceae.
Thelypteridaceae Ching ex Piehi Sermolli are
one of the largest families of ferns, comprising ea.
900 species mostly distributed in tropical regions
((.rimes A Parris, 1986). The family is highly prob¬
lematic, and there has been much confusion in the
taxonomy of the family (Iwatsuki. 1965; Smith.
1990). I he circumscription of genera varies greatly
according to different authors, ranging from the rec¬
ognition of only one genus, Thelypteris Schmidel, in
th(‘ broadest sense to segregating ea. 30 genera by
Holttum (1969, 1971, 1982). Most authors, how¬
ever. agree that Phegopteris (C. Presl) Pee is one of
the most distinctive elements in the family by the
presence of wings on the rachis anti recognize it as
a distinct genus (Grimes & Parris, 1986; Smith,
1990. 1993; Shing et al.. 1999).
Phegopteris is a small genus of only live species
(including the new one described here) in temper¬
ate regions of the Northern Hemisphere in eastern
Asia and North America. It is well distinguished
from other genera of Thelypteridaceae by wings on
the rachis, the wings uniting adjacent pinnae, the
costae lacking an adaxial groove, the naked sori,
and fronds lacking septate trichomes (Grimes &
Parris, 1986; Smith, 1990. 1993; Shing et ah,
1999). Among the species, P. connectilis (Michaux)
Watt usually occurs in cool to cold climate regions
and on mountains or in alpine regions in China,
Japan, Korea, Russia (Siberia), Taiwan, North
America, and northern Europe, whereas P. decur¬
sive-pinnata (H. C. Hall) Fee occurs mainly at low
elevations in eastern Asia, including China, India,
Indonesia. Japan. Korea. Indochina, and Taiwan. In
contrast to the w ide distribution ranges of .nose two
species, P. hexagonoptera (Michaux) Fee is con¬
fined to southeastern North America, and P. tibetica
Ching is restricted to high elevations in Xizang,
China (Iwatsuki, 1965; Park, 1975; Smith, 1993;
Shing et ah. 1999).
During compilation of the fern flora of Korea, we
found ferns that clearly belong to Phegopteris, but
which cannot be placed within any of the known
species. It is here described.
Phegopteris koreana Ih-Y. Sun & C. H. Kim. sp.
nov. TYPE: Korea. Jeollanam-do Province: Ml.
Haekyang. under the edge of deciduous forest,
partially shaded, 13 Oct. 2001, Sun H.-Y. &
C. II. Kan 434 (holotype, JNU; isotypes. J Nil).
Figure 1.
Korean name: Keun-seol-seol-go-sa-ri.
Ilaec species quoad rhizoma breve erectum vel adscen-
dens etiam frondes eaespitosas lamina ambitu lanceolata
latitudinem maximum ad medium attingente ad P. decur¬
sive-pinnata maxime accedit, serl ab ea frondis lamina
uni- vel hipinnata pinnis prof unde pinnatiseetis vel pi n-
natis, pinnulis vel segmentis erenato-lobalis latitudenem
max imam ad medium attingentibus, pinnularum (vel seg-
mentorum) venulis I- ad 3-furoatis, capsula sporangiali
setis usque ad 0.5 mm longis ornata atque sporis tuber-
culatis distinguitur.
Plants terrestrial, green in summer; rhizomes
short, erect or ascending. Fronds monomorphic,
clustered, 50—80 cm long; stipes 10—20 cm long,
densely scaly at base, sparsely scaly elsewhere;
scales 4—8 X 0.4— 0.6 mm, linear-lanceolate, mem¬
branaceous, dark brown, margins setiferous; blades
herbaceous, elliptic, broadest at middle portion,
gradually narrowed toward caudate apex and blade
Novon 14: 440-443. 2004.
Volume 14, Number 4
2004
Kim et al.
Phegopteris koreana (Thelypteridaceae)
441
Figure I. Phegopteris koreana B.-Y. Sun & C. H. Kim. — A. Habit. — B. Pinnae-pinnules or lobes with veinlets and
sori. — C. Seale from stipe base. — I). Seale from raehis. — E. Sorus with hairs. — F. SFM microphotograph of spore.
All are drawn from holotype. Sun, B.-Y. A C. II. Kim 434 (JNU).
base, 40—60 cm long, 20—30 cm witle at middle, 1-
pinnate with deeply pinnatisect pinnae or 2-pin-
nate; rachises winged by continuation of dilated
(auricled) bases of pinnae, glabrous or sparsely pu¬
bescent, sparsely scaly; pinnae 5—13 X 1.5— 2.5 cm
at middle, gradually reduced in size toward base
and apex, ca. 20 pairs, alternate, broadest at mid¬
dle, narrowly elliptic, deeply pinnatisect or pinnate.
442
No von
connected to adjacent pinnae by rachis wing except
near basal ones, apex acuminate or caudate,
sparsely pubescent and scaly; pinnules or pinnati-
sect segments in 15 to 20 pairs, nearly opposite,
acute or obtuse, lobed-crenate; veins free, not
reaching margins, generally forked 2 or 3 times on
pinnules or pinnatisect segments. Sori round, near¬
ly medial in 1 row, generally subterminal on the
veins, with trichome tufts on the receptacles, which
are less than 0.5 mm long; sporangial capsules
sparsely setiferous, to 0.5 nun long, with sparse,
stalked glandular trichomes; indusia lacking.
Spores monolete, bilateral, tuberculate. Chromo¬
some number 2 n = 1 20.
The principal attributes distinguishing Phegop-
teris koreana from P. decursive-pinnata are in char¬
acteristics of the fronds and in such micromorpho-
logical characters as die length of trichome tufts on
the sori and in the spore wall sculpturing. Phegop-
teris koreana usually has fronds 50-80 cm long anil
20—30 cm wide at the middle, while P. decursive-
pinnata usually has smaller fronds, 20—60 cm long
and 5—15 cm wide. Although both species have
once-pinnate blades, the pinnae are basically dif¬
ferent. The pinnae of P. koreana are deeply pin¬
natisect, and hence the fronds are bipinnatisect, or
appear to be twiee-pinnate. In contrast, the pinnae
of P. decursive-pinnata are pinnatifid in the middle
(and hence the fronds are bipinnatifid), and shal¬
lowly pinnatifid to lobed or crenate toward the base
and apex. The shape of pinnae also differs. In P.
koreana, the middle part is the broadest and hence
the pinnae are narrowly elliptic. In P. decursive-
pinnata, the margins are parallel and hence the
pinnae are oblong except for the caudate apex,
hach pinnule or pinnatisect segment (i.e., ultimate
segment) of P. koreana is crenate, whereas the ul¬
timate segments of P. decursive-pinnata are usually
entire to subentire. The two species have trichome
tufts on the receptacle of the sori, which are shorter,
less than 0.5 mm long in P. koreana, and more than
0.5 mm long in P. decursive-pinnata. The venation
pattern in ultimate segments also readily distin¬
guishes the two species. In P. koreana, each pin¬
nule or pinnatisect segment has veins that fork 2
or 3 times; in P. decursive-pinnata the ultimate seg¬
ments have veins that are usually simple or rarely
fork twice.
The spore wall sculpturing is different in the two
species: the tubercles are well developed and rel¬
atively large in P. koreana (Fig. I F), whereas the
tubercles are nearly lacking or rarely are small and
poorly developed in / \ decursive-pinnata, as de¬
scribed by Tryon and Lugardon (1991).
Iwatsuki (1965) commented that P. decursive-pin¬
nata is variable in the size and form of the fronds
and cited a herbarium specimen (Chou 382 ) col¬
lected in Korea that showed bipinnatisect fronds
more than 65 cm long. He treated this as a form of
P. decursive-pinnata. However, in our comparative
study of herbarium specimens collected from Chi¬
na, Japan, and Korea, this large, bipinnatisect or
twiee-pinnate form consistently differs from P. de¬
cursive-pinnata in the shape of the fronds, pinnae,
and ultimate segments, lulled trichomes in the sori,
and spore wall sculpturing. We therefore treat this
specimen and similar plants as comprising a new
species.
Paratypes. KOREA. Jeollabuk-do: Sunchang-gun,
Mt. Hoemun, I Oct. 1997, Hwang. Y. M. & K ./. Ko s.n.
(JNU), Rhyu, II. ./. A- H. R. Kang s.n. (JNU); Temple Bog-
wangsa, 2 Sep. 1973, 7.' H. Lee s.n. (SNUA [4]); Mt. Nae-
jang, 7 Aug. 1974, T B. Lee s.n. (SNUA [2J). Jeollanam-
do: Mt. Suin. near Castle Suin, 16 May 2003, Sun, B. Y.
s.n. (JNU); Gwangyang-gun, Jinsang-myeon. Eochi-ri, 27
Aug. 2001, Park. S. H. et al. s.n. (JNU).
Key m the Species oe Phegopteris in the World
la. Rhizomes long creeping; fronds distantly spaced;
blades deltate or nearly so, broadest at or near
base.
2a. Basal pair of pinnae connected to those next
above by wing along rachis . . P. hexagonoptera
2b. Basal pair of pinnae not connected to those
next above by wing along rachis.
3a. Blades 8—23 X 6— 20 cm, deltate; basal
pair of pinnae slightly longer than broad
. P. connectilis
3b. Blades 18—20 X 10—14 cm, narrowly
deltate; basal pair of pinnae 2 times
longer than broad . P. tibetica
lb. Rhizomes short, erect or ascending; fronds clus¬
tered; blades lanceolate, broadest at middle.
4a. Blades I -pinnate, pinnae lobed or pinnati¬
fid, nearly parallel from base to middle, pin¬
nules nearly entire; veinlets of pinnules sim¬
ple, I- or rarely 2-forked; setae on
sporangial capsules 0.5-0.8 mm long; spore
walls smooth, rarely with a few tubercles
. P. decursive-pinnata
4b. Blades 1 -pinnate; pinnae deeply pinnatisect
or blades 2-pinnate, broadest at middle, pin¬
nules lobed-crenate; veinlets of pinnules I-
to 3-forked; setae on sporangial capsules to
0.5 mm long; spore walls tuberculate ....
. P. koreana
Acknowledgments. This research was supported
in part by a grant from the Eco-teehnopia21 Project
of Environmental Technology Development funded
by the Ministry of Environment of the Korean Gov¬
ernment in 2003. Many thanks to A. R. Smith for
commenting on and improving the manuscript. The
authors also thank D. E. Boufford for improving the
Volume 14, Number 4
2004
Kim et al.
Phegopteris koreana (Thelypteridaceae)
443
manuscript, and the directors of the following her¬
baria for the loan of specimens: PE, SNU, SNIUA.
Literature Cited
Grimes, J. W. & B. S. Parris. 1986. Index of Thelypteri¬
daceae. Royal Botanic Gardens, Kew.
Holttum, R. E. 1969. Studies in the family Thelypterida¬
ceae. The genera Phegopteris, Pseudophegopteris, and
Macrothelypteris. Blumea 17: 5—32.
- . 1971. Studies in the family Thelypteridaceae III.
A new system of genera in the Old World. Blumea 19:
1 7-52.
- . 1982. Thelypteridaceae. Pp. 331-599 in C. G.
G. J. van Steenis & R. E. Holttum (editors). Flora Ma-
lesiana Series 11 — Pteridophyta, Ferns and Fern Allies,
Vol. 1, Part 5. Martinus Nijhoff, The Netherlands.
Iwatsuki, K. 1965. Taxonomy of the thelypteroid terns,
with special reference to the species of Japan and ad¬
jacent regions. IV. Enumeration of the species of Japan
and adjacent regions. Mem. Coll. Sci., Kyoto Imp.
Univ., Ser. B, Biol. 31: 125—197.
Park, M. K. 1975. Phegopteris & Thelypteris. Pp. 258—275
in Illustrated Encyclopedia of Fauna & Flora of Korea,
Vol. 16. Pteridophyta. Ministry of Education, Seoul. [In
Korean.]
Shing, K. H„ Y. X. Fin. P. S. Chiu & G. H. Yao. 1999.
Thelypteridaceae. Pp. 15—353 in K. H. Shing (editor).
Flora Reipublicae Popularis Sinicae, Vol. 4(1). Science
Press, Beijing. [In Chinese.]
Smith, A. R. 1990. Thelypteridaceae. Pp. 263—272 in K.
Kuhitzki (editor), The Families and Genera of Vascular
Plants, Vol. I. Pteridophytes and Gymnosperms. Spring¬
er- Verlag, Berlin.
- . 1993. Thelypteridaceae. Pp. 206-222 in Flora of
North America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 2. Oxford Fniv.
Press, New York.
Tryon, A. F. & B. Lugardon. 1991. Phegopteris. Pp. 394—
395 in Spores of the Pteridophyta. Springer- Verlag, New
York.
Six New Taxa of Passiflora (Passifloraceae), with Nomenclatural
Notes on the Genus in Mesoamerica
John XL MacDougal
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
threebrane@sigmaxi.org
Alts m act. Nomenclatural notes, leetotypifica-
tions, a neotypification, and changes in synonymy
affecting Passiflora (Passifloraceae) in Mesoamerica
are presented, as well as descriptions of newr vari¬
eties and species. Included are six new taxa: P.
cissampeloides, P. ornithoura var. chiapasensis, P.
quetzal, P. quinquangularis, P. subfertilis, and P.
veraguasensis. Two taxa change rank: P. suhlanceo-
lata is elevated from P. palmeri var. sublanceolata
Killip and P. jorullensis var. salvadorensis is low¬
ered from I1, salvadorensis J. I). Smith. The new
combination I1, platyloba var. pubescens (Grisebach)
J. MacDougal must be substituted for P. platyloba
var. williamsii (Killip) A. Gentry, and P. serrulata
var. pubescens Grisebach is a synonym of P. wil¬
liamsii killip. not P. vitifolia IIBK. The well-known
name P. pulchella IIBK is a new synonym of P.
bicornis Miller; P. brevipes killip is a new synonym
of P. cobanensis killip: P guatemalensis S. Watson
is not a synonym of P. hahnii (Fournier) Masters;
P. cookii killip is a new synonym of P. halinii ; and
P. obscura Findley is a new synonym of P. sexflora
Jussieu. Leetotypes are designated for P. divaricata
Grisebach ( Duehassaing s.n., anno 1850, GOET),
l‘. serrulata /3. pubescens Grisebach {Duehassaing,
s.n. 1 1850— 1851 1, GOET). The colored plate op¬
posite page 430 in Revue Horticole 40 (1869) is
designated as the neotype of I*, hahnii.
Key words: Mesoamerica, Passiflora, Passiflor¬
aceae, S. Calderon.
Approximately 20% of the genus Passiflora F.
grows in Mesoamerica, that area from southeastern
Mexico through Panama. These 106 species are
predominately members of the small-flowered and
small-fruited passionflowers, subgenus Decaloba
(DC.) Reichenbaeh. The last monograph of Passi¬
flora (killip, 1938) recognized about 70 species of
the genus in Mesoamerica, and I and others have
added numerous others in the last 20 years (e.g.,
knapp & Mallet, 1984; MacDougal \ killip (1938:
226). B destroyed, photo at E neg. 16554, MO; iso-
tvpe. B-W not seen, P).
Passiflora divaricata Grisebach, Bonplandia 6(1): 7. 1858.
TYPE: Panama. Anno 1850. Duehassaing s.n. (leo-
totype, designated here, GOET).
Miller’s older name supplants the widely known
Passiflora pulchella of floras from Mexico to Vene¬
zuela. Killip (1938: 226) suspected this: “This may
well be . . . Passiflora bicornis ... but Miller’s de¬
scription is indefinite ... in the absence of a type
specimen, it seems best to maintain the name that
is in general use. " Examination of the type found
at BM, with its well-preserved flower, large bracts,
and bilobed leaves, now shows Passiflora bicornis
to be an older name for the same species.
killip (1938) also suggested that the type of the
synonym Passiflora divaricata was a Duehassaing
specimen at P, but in Grisebach’s herbarium at
GOET there are three elements from Duehassaing
that appear to have been part of Grisebach’s pro-
tologue. One is a watercolor painting with violet
flowers, one has a divaricate branch described in
the protologue. and one is marked in Grisebach’s
hand “[Passiflora] divaricata m.” I have chosen the
last element as leetotype.
As a synonym of Passiflora pulchella, Killip
Novon 14: 444^462. 2004.
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
445
(1938) cited the ineffectively published name Pas¬
siflora subtriangularis [no rank] beta S. Calderon,
Datos Bot. Medica El Salvador (Passifloras Dilo-
batas de El Salvador) 11, figures 9. 19, plate opp.
p. 12. pre-1936. For a discussion of the validity
and availability of Calderon’s publication, see be¬
low under P. quinquangularis S. Calderon ex .1.
MacDougal.
Passiflora cissampeloules J. MacDougal, sp. nov.
TYPE: Honduras. Cortes: trail to Cerro Can-
tilles. Cusuco National Park, 15°30'N,
88°14'W, 1840 m, 18 Mar. 1993 (fr), T. Haw¬
kins 634 (holotype, EAP; isotypes, HEH not
seen. MO, TEFH not seen). Figure I.
Haec species Passiflorae guatemalensi si m i I is, sed ah
ea stipnl is glandularibus integris vel leviter crenatis, hrac-
teis minoribus, fructu ellipsoideo manifeste stipitato atque
seminibus insigniter minoribus distinguitur.
Vine, to 10 m, glabrous throughout; stems terete,
glaucous; prophylls of the vegetative bud 2. un¬
equal. 0.9-2. 2 X 0.4— 0.8 mm, both narrowly ovate,
acuminate, entire or rarely with a marginal tooth.
Stipules 4—8 X 1 1-16 mm, depressed-ovate, aurie-
ulate, slightly clasping, truncate at apiculate apex,
the margin entire to obscurely erenulate and 3- to
6-glandular; petioles eglandular; blades 5—6.5 X
5-6.7 cm, peltate 8-10 mm from margin, entire,
not variegated at maturity, widely ovate, at base
subtruncate with convex curve in peltate region,
obscurely 3-lobed, angle between lateral veins 50°-
60°, th<- lateral lobes ± obsolete, the central lobe
acute or bluntly acute; laminar nectaries absent or
marginal when present, as I or 2 glands borne at
edge of peltate region. Peduncles I per node, (1.4—
)2.0-3.2 cm, uniflorous; bracts 2 at apex of pedun¬
cle, variable in size, 6—15 X 9—17 mm, ovate-tri¬
angular. free to base, entire or with a very few mi¬
nute setae proximally. not glandular, acute at apex,
green. Flowers “creme color, centre yellow (Haw¬
kins 198), the corona apparently yellow; stipe ca. 3
mm long (to 4.5 mm in fruit); hypanthium 17-18
mm diam.; sepals 1 1-15 X 12-17 mm, widely ob¬
long triangular, rounded at apex, with no projection;
petals 16-17 X 12-13 mm, widely ovate, abruptly
narrowed at base; coronal filaments in 3 to 4 series,
the outermost 5-8 mm long, linear, the inner 2 to
3 series 3—5 mm long, capitellate; operculum mem¬
branous, plicate; androgynophore 8—9 mm, anthers
ca. 6 mm; ovary 3—4 X 2—2.5 mm, ellipsoid, gla¬
brous; styles 5 mm long including stigmas. Fruit
ca. 8 X 3.5—4 cm including a 1.5— 2.0 cm long
stipe, ellipsoid, lustrous; seeds 4.5— 4.9 X 2.7— 2.8
X 1.3— 1.5 mm thick, narrowly obovate in outline.
campylotropous, the testa reticulate with ca. 60
pits, the chalazal beak inclined toward the raphe.
With a peltate leaf similar to Passiflora guate-
malensis S. Watson or some lowland forms of P.
hahnii (Fournier) Masters, and a stipitate fruit like
P. membranacea Bentham, this new species com¬
bines character states from each of the other spe¬
cies in supersection Hahniopathanthus (Harms) J.
MacDougal & Feuillet, to which this species clearly
belongs. From P. membranacea , which has leaves
peltate only 1 — 3.5( — 5) mm from the margin, pedun¬
cles (6.3— )8—14(— 17.5) cm long, bracts 2.7— 5.0 X
2.5— 5(-6.3) cm. and androgynophore (2.7— )3.0-4.0
cm long, it is easily distinguished by the peltate
leaf, shorter peduncles, smaller bracts, and much
smaller flower. From P. guatemalensis, which has
dentate-fimbriate stipules with filiform teeth, an¬
drogynophore 3.5— 5.0(— 6.0) mm long, and a non-
stipitate fruit or the stipe 1 mm long or less, it
differs by the entire or crenate stipules with glan¬
dular margins, the longer androgynophore, and the
stipitate fruit. It differs from P. Iiahnii, which has
bracts 1 .9— 6.2 X 1 .5 X 6.4 cm, peduncles 2.5— 7.0
cm, and a fruit stipe of 1-4 mm, in the smaller
bracts, usually shorter peduncles, and the stipitate
fruit. Passiflora cissampeloules has generally small¬
er bracts than the others in the supersection, and
the seeds are markedly smaller than the other
known species, which range from (4.6— )5— 6.5 X
(3. l-).3.3-4.2(-4.5) mm.
Distribution anil habitat. Known from Copan
and Cortes, Honduras, at 1460—1840 m. I he type
was collected in moist cloud forest of pine, Liquid-
ambar, Podocarpus, and Quercus. I he paratype is
from dense mixed hardwood forest.
Etymology. The peltate leaves of this new spe¬
cies very much resemble species of Cissampelos E.
in the Menispermaceae, hence the specific epithet.
Paratype. HONDURAS. Copan: S slope of Cerro
Azul directly N and above Quebrada Grande, 12 km NE
of Florida, Cerro Azul National Park, 15°06'N, 88°55'W,
9 Feb. 1992, T. Hawkins 198 (EAP, MO).
Passiflora cobanensis Killip. J. Wash. Acad. Sci.
14: 111. 1924. TYPE: Guatemala. Alta Vera-
paz: between Chama and Coban, 3000 ft.
[“950 m” in protologue], 26 July 1920, H.
Johnson 411 (holotype, US-1083984, photo at
F).
Passiflora brevipes Killip, Publ. Carnegie Inst. Wash. 461:
312, plate 2. 1936. Syn. nov. TYPE: Belize. Toledo:
Jacinto Hills [ca. 16°09'N, 88°55'W], 400 ft. [“120
m” in protologue], I I Mar. 1934, W. A. Schipp 1404
(holotype, F-733685, neg. 52921; isotypes, A not
446
No von
a
>3
i
4
MISSOURI
BOTANICAL GARDEN
HERBARIUM
NV 5618545
Type of
dsn Au(X> in Chiapas) m, SE Oaxaca, W
Chiapas, Mexico, and El Salvador .
. var. salvadorensis
There is no basis lor separation of these laxa by
(lie nearly identical flowers. Moth are the same size,
w ith strictly one row of coronal filaments, a 8—4 mm
long suberect operculum that is sparsely puberu-
lenl basally, and a 9—1 I mm long androgynophore.
I he flowers of both exhibit tin1 same unusual be¬
havior of remaining open more than one day (pcrs.
obs.; Calderon, s.d.). In both, the corona is initially
red or a fiery red-orange, which persists for the first
day, but changes or fades the second day to purple
or purple-pink before closing later that evening el¬
even the morning of the third day. The odor of Pas¬
siflora jorullensis var. salvadorensis is unknown, but
the typical variety is famously stinky with a strong
odor described as between “stereoraire el [nitride”
by Planchon (1849). A two-day flower is unique
among tlu* native species of passifloras in Meso-
ameriea. and is shared in Mexico and North Amer¬
ica only by the red variant of the close relative P.
mexicana Jussieu. The pollination system associ¬
ated with this unusual floral behavior is not known.
Dennis Breedlove has seen hummingbirds at the
flowers of variety salvadorensis in Cintalapa, Chia¬
pas, at 1600 m (pers. comm.), wasps have been
seen at the flowers of the red /( mexicana, and both
wasps and hummingbirds at the flower of the relat¬
ed (and also malodorous) I*, gilbertiana J. Mac-
Dougal.
As noted by killip (1938), the leaf sh ape is often
distinctive, with the leaves of Passiflora jorullensis
var. salvadorensis more deeply lobed and the lobes
generally obtuse or rounded, but there is overlap
between the varieties. Both occasionally retain
some costal variegation into maturity. The leaves of
variety salvadorensis closely resemble several Bau-
hinia species with which the variety often grows.
I he differences in vestiture are generally ones of
quantity rather than quality: although variety sal¬
vadorensis often appears glabrous at first glance,
close examination reveals a slight puberulence at
least on the young growth or around the nodes. The
type and other material from El Salvador is partic¬
ularly glabreseent, with ovaries that are glabrous or
very nearly so. In variety jorullensis, glabrous ova¬
ries are seen only in the westernmost populations
in Mexico.
I hese two disjunct taxa are usually found in eco¬
logically different forest types. Passiflora jorullensis
var. jorullensis mainly inhabits moist montane oak
forest, while variety salvadorensis is usually found
in subdeciduous or tropical deciduous forest, yet
sometimes ranges up to montane moist forest. Nei-
the r variety of the species is known from Guatemala
or Honduras.
As a synonym of Passiflora salvadorensis , killip
(1938) cited the ineffectively published name, Pas¬
siflora subtriangularis (no rank) alfa S. Calderon.
Dates Bot. Medica El Salvador (Passifloras Dilo-
batas de El Salvador) 15. figures II. 12. plate opp.
p. 20, pre-1936. That name is accompanied by a
detailed protologue with valuable morphological
and ecological information about this variety. Fora
discussion of the validity and availability of Cald¬
erons publication, see below under P. quinquan-
g id oris S. Calderon ex .1. MaeDougal.
Representative specimens examined. EL SALVADOR.
Ahuachapan: San Benito, al S del [tie del cafetal El Nar¬
anjo. Co. Davila. E. Sandoval & Chinchilla 574 (MO). San
Salvador: San Salvador, .S'. Calderon RIO (US); eamino al
eerro de San Jacinto, 21 July 1959, E. Montalvo 3662
(ENCB); vicinity of San Salvador, I] Standley & Padilla
2051 (LAP). MEXICO. Chiapas: Mex. Ilwy. 190. 13 mi.
SE of Tapanatepec, I I Sep. 1964. 1). Breedlove 7167 (US,
ENCB, K LL); Arriaga. 1.3 km N ol Arriaga along Mex.
Ilwy. 195, 25 Sep. 1971. I). Breedlove 10848 (DS. ENCB,
MICH); Cintala[ 1a, Sf, of Cerro Haul on the border with
the state of Oaxaca, 16 km NVf of Rizo de Oro along
logging road to Colonia Eigaroa, 6 Sep. 1972. I). Breedlove
27627 (DS); Cintalapa, along 111. 190, 14.7 mi. E of in¬
tersection of Rt. 200 to Tapaehula, 8 Aug. 1978. vicinity
of Tonala, 18 Aug. 1949, E. Matilda 18783 (E. MEXU);
Inxtla Gutierrez— Jalisco, Sep. 1923, Purpus 0257 (DS).
Oaxaca: San Pedro Tapanatepec, along lit. 190, 5.9 mi.
E ol intersection of lit. 200 to Tapaehula, 8 Aug. 1978,
./. MaeDougal 326 (DUKE).
Passiflora ornitlioura Masters var. chiapasensis
J. MaeDougal. var. nov. TYPE: Mexico. Chia¬
pas: Mt. Ovando (15°25'N, 92°36'W], 2000 m,
14-18 Nov. 1939, E. Matilda 3971 (holotype,
MEXU: isotypes, (ill. MICH — 2 sheets, MO.
US). Eigure 2.
Haec varietas a varietate typica floribus seminibusque
majoribus, foliorum plerumque minus profunde lobatorum
lobis lateralibus angulum minorem inter sc formantibus
atque filamentis coronalibus in seriem unicam dispositis
distinguitur.
Herbaceous vine, size unknown but probably 2—
5 m, minutely puberulent throughout at the shoot
tip (except blades), becoming sparsely puberulent
(to glabreseent) below. Stems ± terete, striate, pu¬
berulent or sparsely puberulent (rarely glabreseent)
below; prophyll of vegetative bud 1. narrowly lan-
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
449
FLORA MEXICANA
Mituda Herbarium
*/^a ss
-t. "T e“^i
Z, -20 mm,
filiform, the inner I to 2 series 7—10 mm (1 inner
series at petals. 2 inner series at sepals); operculum
4.5— 5.0 mm, membranous, plicate; limen edge at
least 1.5 mm high; androgynophoro 8.5—9 mm. the
free staminal filaments ca. 6 mm. the anthers ca. 5
mm long; ovary 3.8— 4.0 X 2.3— 2.5 mm, ovoid-el¬
lipsoid, glabrous; styles at least 9 mm long includ¬
ing stigmas. Fruit ca. 6 X 3.5 cm, ellipsoid, stipe
absent or less than 4 mm; seeds 5.3— 6.0 X 3.5—
3.9 X 2.0 mm, obovate in outline, eampylotropous,
the testa reticulate with ca. 50 to 60 pits, the cha-
lazal beak inclined toward the raphe.
Passiflora quetzal is assigned to supersection
Hahniopat hunt bus in subgenus Decaloba, where it
exhibits a number of character states primitive for
that supersection. I his new species vegetatively
closely resembles certain forms of P. membranacea
from Chiapas, Mexico, with subpeltate leaves, a
pair of small stalked petiolar nectaries, and abun¬
dant marginal laminar nectary glands borne not just
at the base of the lamina, but also marginally be¬
tween the primary veins. These characteristics are
primitive in the group, by outgroup comparison,
killip (1938: 23) did not observe glands in this
group, stating “in these . . . species there are no
nectar-secreting glands in any form . . . ." In reality,
extrafloral nectaries are common in this group. The
species are polymorphic for the presence and lo¬
cations of various types of nectaries, and this var¬
iation gives valuable evidence about the relation¬
ships within each species and within the
supersection. In the description above, the leaves
lacking the marginal nectaries in some positions
were smaller ones; the large leaves on all the spec¬
imens bear abundant glands. The flowers are sim¬
ilar to those of P. guatemalensis and to forms of P.
halinii, but the bracts art* smaller. Passiflora quetzal
does have only two floral bracts, an advanced state
in the supersection and in subgenus Decaloba. The
floral buds are obtusely and shallowly conical, like
P. guatemalensis. 1 originally annotated the isotype
(KNUR) as "all. cookii Killip." a name now consid¬
ered a synonym of P. halinii (see discussion above
under P. halinii). Several searches at the F herbar¬
ium for an expected isotype have been fruitless.
Distribution and habitat. Southwestern Chia-
pas, Mexico, and adjacent western Sail Marcos,
Guatemala, on western slopes of the Serrania Tran-
sistmica, or Cordillera Madre, at 1800-2100 m in
wet mountain forest or “bosque mesofdo de mon¬
tafia (moist montane forest). The type locality is
also the type locality for Passiflora eglandulosa J.
MacDougal.
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
453
ISOTYPUS
PL^n-fioro. J. MacD
7 J if rtov in.
I MAC none At
rieil .
frtOTc Dutc£,M£X(J
GUATEMALA
Chicago Natural Huron- Museum
(Wurla Agricob P»ivu»rnca»u
lore nr* mb ran ace a Bo
speuie* A-ff P <+(xk* f r lUp
iVtA \}XA4UJ!A Bt+xlK
Wrt mountain fortat nw Alda* Fratemulad. hetwwn San Uftfarl Pi*
de l« C!u rata and PiUo Gordo, v**t facing "l"t" ,,f ,t"'
Mountain-*, department of San Matron, alt. IMG -100 m Prerm-
tier 10 18, 1<*S.
l)u»e Uni¥t>i%ity Hvibat’um
Utu O. Willi H. Amihko Malm* I. «*vJ T,rw IV WiHw»
Figure 3. Photo -aph of the isotype of Passiflora quetzal at ENCB.
Etymology. This species is named for that
splendor of the forest, the quetzal, and the native
trogons that daily perch among the vines at the type
locality. Following 1CBN Article 23.2 (Greuter et
al., 2000), 1 choose the epithet from the undeclined
American English and Central American Spanish
word for Pharomachrus mocino, the resplendant
quetzal, taken from the Nahuatl word quetzalli.
Paratypes. MEXICO. Chiapas: Union Juarez, en el
454
Novon
F,g.Ure 4' . Fh°t°graP1' of a lr'‘sl' flower of Cassiflora quinquangularis in longitudinal section showing details of
and curved floral stipe {MacDougal 626, DUKE).
Volcdn lacami por el cam i no de Talquidn a la eima del
volcdn, por la lfnea divisoria de Guatemala, 6 Feb. 1987
(bud, fr), Martinez .S', cl al. 19613 (MKXU) 19678
(MKXU, MO).
Passiflora qiiinqiiangulai-is S. Calderon ex J.
MacDougal, sp. nov. I YPF: El Salvador. San
Salvador: S of San Salvador, July 1922 (11), .S’.
Calderon 851 (holotype. US I 151809, photo at
MO neg. 05129; isotypes, (ill not seen, US).
Figure 4.
Haer species P. rubrae et P. capsulari similis, sed ab
eis caule quinquangulari atque alabastris modo araneae
ex pedunculis pergracilis (minus quam 0.5 mm in diam.)
pendentibus distinguitur.
Vine 2 4 tn, herbaceous, or slightly woody al
base, pubescent throughout, often densely so al
shoot lip. on stem, and abaxial surface of leaves,
sometimes the stems glabrescenl with age; stems 5-
angular, ± cemuous at growing lip. pubescent with
trichomes 0.3(0. 4-0.6)-0.8 mm long; prophyll of
the vegetative bud 1, linear-triangular to very nar¬
rowly lanceolate, 2.5— 4(-7) mm long. Stipules 4—7
X 0.4-0.9 mm. linear-triangular, falcate; petioles
eglandular; blades 4~10(-17) X 4— 9(— 1 2) cm, en¬
tire. not variegated, widely oblong-obovate (to wide¬
ly oblong). 2(3)-lobed, the base slightly l<» strongly
cordate, the lateral lobes acute to rounded, the an¬
gles between the lateral lobes (30°-)35o-45o(-60°),
the central lobe (obtuse) rounded to truncate to ob¬
solete; laminar nectaries absent. Inflorescences
sometimes present in older plants as nearly leafless
condensed lateral branches; peduncles I (very' rare¬
ly 2) per node, (1 .5— )3— 5.5(— 9.3) cm, very slender.
less than 0.5 mm diam., uniflorous, not braeteate;
buds abruptly acuminate or short-beaked. Flowers
green-white or cream, with a light purple center,
the corona white with purple, with fruity sweet odor
reminiscent of raspberries; stipe 3-5 mm (to 7 mm
in fruit); hypanthium 8-10.5 mm diam.; sepals 15-
21 (—29) X 5.4— 7.0 mm, narrowly triangular, long-
acuminate but with no projection, pale green-yellow
to yellow-green, abaxially sparsely pilosulous es¬
pecially on the 3 conspicuous nerves, light yellow-
green and often flushed with minute red spots; pet¬
als 10—15 X 3.0— 4.3 mm, lanceolate, acuminate (or
rounded at very apex), white or pale green-yellow;
coronal filaments in 2 series, the outer I 1-18 mm,
filiform, slightly attenuated and white in distal half,
light red-purple to purple-red proximally, purple-
red basal I y, the inner 3— 4.5 mm. capillary, red-pur¬
ple, elavate at the paler tip; operculum ea. 2 mm
high, membranous, plicate, light purple to medium
red-purple, white apieally; limen floor very pale
red-purple; nectar ring (annulus) present in nectary
chamber, ea. 1 mm high, dull purple, not bright
yellow; androgynophore (7— )8— 10 mm, strictly un-
colored except for trace of chlorophyll, the free sta-
minal filaments 1.5-5.5 mm, not colored, anthers
3-7— 4.2 mm long, not colored; ovary 2.3— 4.0 X
l.i— 2.3 mm, ellipsoid, slightly 6-ridged, minutely
densely puberulent; styles 5-6.5 mm long includ¬
ing stigmas. Fruit 3-4.5(-6) X 1 .5— 2(— 3) cm, in¬
cluding a ca. 3 mm stipe, broadly fusiform to ellip¬
soid, conical at both ends or apieally with a beak,
acutely 6-carmate with keels ea. 1—2 mm high, yel¬
low-green. slightly short-puberulent with trichomes
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
455
0.1 mm or less, dehiscent; seeds 3.4— 4.0 X 1.7—
2.4 mm, transversely sulcate with 6 or 7 (or 8) sul¬
ci, the intervening ridges not rugulose.
This species is notable for its long (to 5.5 cm or
longer) and diagnostically very slender peduncles
(less than 0.5 diam.), upon which the acuminate or
short-beaked buds and the flowers are pendent.
Due to the curvature of the floral stipe (ef. fig. 4),
however, the flowers lace horizontally to open. In
Passiflora, it belongs to section Xerogona (Rafin-
esque) Killip in subgenus Decaloba based on the
plicate operculum, absence of laminar nectaries or
floral bracts, the carinate apically dehiscent fruit,
and the seeds' smooth transverse ridges. Most sim¬
ilar to forms of P. rubra L„ P. quirujuangularis dif¬
fers significantly by its slender floral peduncle and
resulting floral orientation, the 5-angled stem, and
the only minutely puberulent, not conspicuously
pubescent, ovary. Although P. rubra does not occur
in Mexico or Central America, another close rela¬
tive does, P. capsularis L. This species prefers wet¬
ter habitats than P. quinquangularis, but in a few
places in Guatemala they grow within a km of each
other (pers. obs.). Passiflora capsularis is easily dis¬
tinguished from the new species by its pure white
or cream, unmarked flower that is borne upward on
a stout peduncle, less pubescence overall, and the
3- to 4-angled stem.
As in many species ol Passiflora subg. Decaloba,
the vestiture is composed of two different classes
of trichomes: very short, curved, cylindrical to cla-
vate antrorsely appressed trichomes 0.08—0.12 mm
long (Harms, 1893; MacDougal, 1994), and longer,
straighter, pointed ones, in this case 0.3— 0.8 mm
long. The ovary, for example, has only the short
curved type, while most other surfaces of the plant
have a mix of the two.
Specimens of this new species have been cited
by a number of authors under the name Passiflora
capsularis, and the descriptions of that species in
florulas of Central America, as well as in Ki 1 lip’s
(1938) monograph, have included measurements
and character states from the new species. The il¬
lustration of P. capsularis in Standley and \\ illiams
(1961), except for the separate fruit, is actually
wholly of this species.
A cleared leaf of Passiflora quinquangularis from
./. MacDougal 626 (MO) was studied and illustrated
by Klucking (1992: 243—244, pi. 96. fig. 3), under
the abbreviated names, “P. quing.” and “P. quing-
- -” [sic]. Katie Hansen (pers. comm.) reports that
the chloroplast intron rpoCl is lacking in samples
of ./. MacDougal 626 (MO). This clone, imported
live from Guatemala in 1980, was grown and widely
distributed. It was self-incompatible in the green¬
house. It appears to be the only clone in cultivation
even now, and has been listed in many nursery cat¬
alogs and illustrated in color in a few popular books
(e.g., Ulmer & MacDougal, 2004; Vanderplank,
1996, 2000).
Named for its 5-angled stem, Passiflora quin¬
quangularis has its origin in a short obscure pub¬
lication by Salvador Calderon, a publication that
survives by only a single known partial copy. It
would have been lost to science except that it was
seen and cited by the monographer Killip (1936,
1938) while he worked at the US herbarium. Killip
cited both text and figures when, as a synonym of
P. capsularis, he invalidly published “ Passiflora
quinquangularis Calderon, Passifloras Dilobatas del
Salvador 6.” A more complete citation would be.
Datos Hot. Medica El Salvador (Passifloras Dilo¬
batas de El Salvador) 6, figures 1-8.
Salvador Calderon (1884—1940), a Salvadorian
botanist and chief of Agricultural Botany of his
country, collected plants in collaboration with Paul
C. Standley of the Field Museum for a flora of El
Salvador. He was the Salvadorian contact for bot¬
anists from the United States in the 1920s and early
1930s, and published articles and booklets between
1925 and 1929 on botany, agriculture, and geology.
The seven-page protologue of his P. quinquangu¬
laris is one of the more detailed ever published for
a passiflora, with astute observations on anatomy,
phyllotaxy, plant movement, ptyxis, floral morphol¬
ogy, chirality, seed morphology, and more.
The publication that heralded this passionflower
was never widely distributed and appears to be
generally lost. After much correspondence, 1 can
locate only one partial copy in the reprint collection
of the United States National Herbarium at US. It
is a booklet with the title Datos Para Iai Botdnica
Medica de El Salvador and the author’s name on
the cover. Also on the cover is “Escuela de Medi-
cina, Qufmica y Farmacia/San Salvador, Republica
de El Salvador, America Central.’ Inside, the work
is entitled Passifloras Dilobatas de El Salvador and
runs from page [1—4 missing] 5 to 20, where the
article clearly continues but is not present in this
bound copy. There are line drawings and two
tipped-in color plates. There is no date. It might
have been published during his productive period
in the 1920s, an idea also supported by the fact
that herbarium specimens (F, US) of all taxa col¬
lected by Calderon are dated between 1921 and
1929, with a very few from 1931. Killip cited the
publication in both 1936 and 1938; allowing for
time in manuscript, Killip may well have seen it
before 1935. This date is critical bec ause P. quin-
456
Novon
quangularis was published in Spanish, without the
Latin diagnosis required after 1934 for valid pub¬
lication. It probably beats this cutoff date, but it
hardly matters, lor there is little question that due
1° the extreme rarity of the publication, the name
is not effectively published (Greuter et al„ 2000,
I CUN Art. 29.1).
The de< ■ision that Passifloras Dilobatas de El . Sal¬
vador was not effectively published and distributed
invalidates two infrageneric botanical names that
were otherwise newly and probably validly pub¬
lished there: "Passiflora subsection Pubescens" S.
Calderon and "Passiflora subsection Raripilus" S.
Calderon. Two other species names are invalidated:
the already doubtfully valid “ Passiflora subtrian¬
gularis (no rank) beta ” S. Calderon |- P bicornis ],
and th«* definitely invalid "Passiflora subtriangu¬
laris (no rank) alfa" S. Calderon. This last was pub¬
lished with P. salvadorensis J. I). Smith | = P. jo-
rullensis var. salvadorensis | in synonymy. Calderon
made no reference to any of these new botanical
names in his two other more widely known botan¬
ical publications (Standley & Calderon, 1925,
1941), probably bee •ause Killip was a collaborator
for the passionflowers there.
Regarding Passiflora quinquangularis , Calderon
did not cite any specific specimens in his booklet,
except to say that his description was based on wild
plants found in the highlands around the capital
(San Salvador). Only two preserved collections of
this species by Calderon are known, and only one,
Calderon 851 (GH, US |2|), is from the area of San
Salvador. Killip (1936), treating Passiflora quin-
quangularis as a synonym of P. capsularis, right¬
fully chose this collection as the “type,” and in
1938 noted that these specimens were held at (ill
and US. At one time I thought the name to be valid
and effective, and thus in 1989 had annotated one
sheet at US as lectotype; to avoid future confusion,
I now designate the same sheet to be the holotype.
Distribution and habitat. Passiflora quiru/uan-
gularis is found in seasonally dry forest, brushy
hillsides, hedges and secondary vegetation, oak
with pine forest, and “ecotone de selva baja cad-
ucifolia y bosque mesofilo” (ecotone of low decid¬
uous forest and moist forest) at elevations of 580-
1600 m sporadically from Oaxaca, Mexico, to El
Salvador, with a disjunct population known from
the Nicoya Peninsula of Costa Rica. It is not re¬
corded from Honduras or Nicaragua. Only in El
Salvador and southwestern Guatemala does it seem
to be common.
Phenology. Elowering and fruiting throughout
the year.
Common names. Ala de inurcielago (R. Villa -
corta et al. RV854 ), bejuco calzoncillo {J. Gonzalez
d il), calzoncillo ( Calderon 851). bejuco calzdn de
mujer (F. Chinchilla s.n.), bejuco calzdn de liernbra
(,/. Gonzalez 409). 1 hese all refer to the curious
shape of the bilobed leaf.
Paratypes. COSTA RICA. Guanacaste: La Cruz,
Santa Elena. Parque Nacional Guanacaste, 2001, A. Rs-
trada s.n. (CR. MO photo). EL SALVADOR. Almaclia-
P«n: 1928, S. Calderon 2428 (F); San Benito, al N del
Eseobal, Parque Nacional El Imposible, I3°49'N,
89°56'W, 27 Apr. 1994, F. Chinchilla s.n. (B, EACH,
MO), R. Villacorta et al. R 1 854 (LAGU, MO); near Ataco,
19 Jan. 1947, P. Standley A- Padilla 2644 (F); vicinity of
Ahuarliapan. 9-27 Jan. 1922. P. Standley 19788 (MO.
NY. US). La Lihertad: Teotepque, 13°25'N, 89°31'W, 4
May 1996, ,/. Gonzdlez 331 (MO); Jayaque, 13°40'N,
89°26'W, ./. Gonzalez 409 (B, EAP, LAGU, MO. UES);
finea El Para iso, Jayage. 13 July 1976, Montalvo 4711
(MO). GUATEMALA. Raja Verapaz: Salama Puente
Barranca, al W de Salama, 15°06'N. 90°18'W, 24 July
1988, P. Tenorio et al. 14740 (MO). Santa Rosa: Cuaji-
niquilapa, Sep. 1893, Heyde & Lux s.n./J. I). Smith 6 142
(US); cultivated at Duke University 1980-1984, from live
stems collected 27 Jan. 1980, 6 mi. E of rd. to Cuilapa
on CA-1, ca. 3 mi. W of intersection of CA-1 with CA-8
to Valle Nuevo, pressed July 1983, ./. MacDougal 626
(BH, RM, COL, CR. DUKE, E, G, GOET, MEXU, MO,
MO-spirit. UPGR, US, UVAL); along road SE of Barber-
ena, 21 Nov. 1940, P. Standley 77729 (E, US); near Cui-
lapilla. 23 Nov. 1940, P Standley 78118 (E). MEXICO.
Oaxaca: Lachiguiri, eerca a Santiago Lachiquiri, 9 June
1985, M. Chazaro A- M. Dutch 3371 ( W IS); Juchitun, I
km al S de Guevea de Humboldt, 16 Mar. 1983. R Torres
el al. 2521 (MEXU).
Passiflora sexflora Jussieu, Ann. Mus. Natl. Hist.
Nat. 6: 1 10. pi. 37, fig. I. 1805. Decaloba sex-
flora (Jussieu) M. Roemer, Earn. Nat. Syn.
Monogr. 2: 164. 1846. TYPE: Dominican Re¬
public. Santo Domingo, Poiteau s.n. (holotype,
P).
Passiflora obscura Lindley, Trans. Roy. Hurt. Soe. London
7: 48. 1830. Syn. nov. Decaloba obscura (Lindley)
M. Roemer, Earn. Nat. Syn. Monogr. 2: 157. 1846.
TYPE: Cultivated in England from seeds brought
from "some part of the North Eastern Coast of South
America” by G. Don in 1823, llerb. ./. Lindley (ho¬
lotype, CGE, photo at MO neg. 051 19).
Examination of type material of Passiflora obscu¬
ra shows this name to be a synonym of the earlier
P sexflora. Killip (1938), not having seen this type,
aligned the name doubtfully under P. rubra. These
two species are often vegetatively very similar, with
pubescent more or less bilobed leaves with no lam¬
inar nectaries, but are unmistakable with fertile
material. Passiflora sexflora has bracts on the usu¬
ally branched peduncles, and purple berries, while
P. rubra has naked, non-bracteate, unbranched pe¬
duncles, and a red, pink, or white and red dehis¬
cent berry-like capsule.
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
457
%
pA-tsi-florg. iu.b-ftrHh*. M/iTbuiui
/ lf> uirtr.
Sf> “'Mr/.
i/nmutt
-
50i PLANTS OF GUATEMALA
L
" FfSiUtUL/iMl/.
J’ +rr4xt-i\tMr-A.1'arM. alt. l»H> 1400 m
Figure 5. Photograph of the holotype of Passiflora subfertilis at F.
Passiflora subfertilis J. MacDougal, sp. nov.
TYPE: Guatemala. Quezaltenango: betw. linca
Pirineos & Finca Soledad, along stream, lower
S-facing slopes of Volcan Santa Marfa, betw.
Santa Marfa tie Jesus & Calahuache, 1300—
1400 m, 5 Jan. 1940 (fl. imm. fr), ./. A. Sley-
ermark 33503 (holotype, F). Figures 5, 6.
Haec species Passiflorae biflorae similis, sed ah ea
gemma vegetativa prophyllum unicum gerente, folii lobis
458
Novon
Figure 6. Flowers of Passijlora biflora compared to the flower of P. subfertilis. From left to right, flowers from three
clones of P. biflora (Costa Mica, MacDougal 463 (DUKE); Costa Mica, MacDougal 464 (DUKE); Guatemala, MacDougal
598GR (DUKE)), and flower of P. subfertilis {MacDougal 597GR). Scale is in cm.
acuminatis acutisve atque petalis sepalisque angustioribus
distinguitur.
Vine 2-4 m. minutely puberulent throughout at
the shoot tip, or the blades sparsely so, sparsely
puberulent to glabrescent below; stem subterete or
slightly angular, striate; prophyll of the vegetative
bud 1 . 1 .2—2.0 mm long, narrowly lanceolate, en¬
tire, sometimes curved over the bud. Stipules 1.6-
2.1 X 0.2-0. 3 mm, linear-triangular, falcate, stra-
mineous-neereseent; petioles eglandular; blades
2.8-6 X 5—9.5 cm, in outline obovate to widely
obovate, shallowly 2-lobed (rarely sub 3-lobed), the
has*1 subtruncate to obscurely widely cordate, the
lateral lobes acuminate to acute, the central lobe
absent or when present obsolete to obtuse, the an¬
gle between the lateral veins 50°-72°, the ratio of
laminar width to central vein length 1. 1-2.1, not
variegated, the margin entire; laminar nectaries nu¬
merous but inconspicuous, borne between the main
veins; juvenile leaves in outline deeply depressed
obovate, deeply 2-lobed, the base broadly rounded
to subtruncate, the lobes very widely spreading,
lanceolate to narrowly triangular, acuminate, the
angle between the lateral veins 7()°-115°. Pedun¬
cles (1)2 per node, 0.8—1 .0 cm, uniflorous; bracts
0.8-2. 0 X 0. 1-0.2 mm, linear-triangular, early
stramineous-necrescent; buds slightly constricted
just distal to hypanthium. flowers white, the corona
light yellow; stipe 5-7 mm (to 9 mm in fruit); sepals
13-15 X 6—i mm, ovate-oblong, pale green abax-
ially, ± white adaxially, with no projection; petals
10-12 X 4 mm, narrowly oblong, white; coronal
filaments in 2 series, the outer 5—6 mm long, slight¬
ly laterally compressed, basally yellow-green, dis-
tally light yellow, the inner ca. 2.5 mm, capillary,
capitellate, near-white with white to green tips;
operculum membranous, plicate; androgynophore
ca. 9 mm, slightly purple; ovary glabrous, styles
and stigmas green. Mature fruit unknown; immature
fruit 2-2.5 X 1.5-1. 8 cm, widely ellipsoid to wide¬
ly obovoid, not stipitate; arils unknown; seeds ap¬
parently transversely sulcate.
This rare species was known only from two col¬
lections made at one place by Steyermark until I
went to, or close to, that locality in 1980 and col¬
lected it alive. I was then able to cultivate, study,
and photograph the species in the greenhouse. As¬
signed to section Decaloba of subgenus Decaloba,
it is generally very similar to the sympatric Passi-
flora biflora Lamarck, fhe historic monographer
Killip had annotated the two older collections as P.
biflora , but it may be distinguished easily from that
species by having only one prophyll at the vege¬
tative bud and leaves with sharply pointed lateral
lobes, 'file flowers are very similar, but the buds of
P subfertilis are constricted distal to the hypanthi¬
um, unlike the buds of P. biflora. In Figure 6, three
clones of P. biflora from Guatemala and Costa Rica
showing a range of floral variability in that species
are compared to a flower of P. subfertilis. The flower
adjacent to that of P. subfertilis is from a P. biflora
collected at the exact same locality, fhe petals of
the new species are proportionally narrower than
those ol P. biflora. The loliage very much resembles
that of the rare P. sanctae-mariae .1. MacDougal,
described from a locality only I km distant, except
P. subfertilis has numerous inconspicuous laminar
nectaries. These nectaries can be seen in the
cleared leaf from MacDougal 597GR (MO) that was
studied and illustrated by Klucking (1992: 253-
254, pi. 109, fig. 4), as “Passiflora subfertilis
[MacDougal]/' 1 1 also vegetatively resembles P. hel-
lert Peyritseh, with which it grows, and the prophyll
of tht‘ vegetative bud is often curved over in the
same way. I his species has a floral corona of only
one row; however, its leaves have a narrower angle
of lobing. from 24°— 40°(— 50°), and the leaves are
Volume 14, Number 4
2004
MacDougal
New Taxa of Pass i flora
459
nearly always longer than wide, with a width to
length ratio of 0.55-0.85(-l .0). Because the new
species resembles the three sympatric species men¬
tioned and is only known from a very small area, I
wondered if it might he a hybrid. Hybrids are rare
in the genus, but are suspected in the P. sexflora
Jussieu— P. rugosissima killip complex in Veracruz
and Oaxaca, Mexico, and in the widespread Neo¬
tropical P. foetida L. and P. ciliata Aiton complex¬
es. Morphologically, it is not clearly intermediate
between any of the three potential parents, though
one of the parents would certainly be P. biflora (cf.
Fig. 6). If it is a hybrid, it is either reproducing or
is being regenerated around the type locality, for
the plant I found was only a few years old. and
probably not a vegetative sprout of the same plant
Steyermark found 40 years earlier. In the green¬
house, fresh pollen showed only 40%— 50% viabil¬
ity as indicated by lactophenol-eotton blue staining
(slide preserved on sheet at MO). Based on this, I
give the name subfertilis, and although there is
some suspicion that it is a hybrid, it is not here
published as one.
The type and paratype localities in Guatemala
are no more than a kilometer from each other on
I lie Pacific slopes of Volean Santa Maria, in dis¬
turbed primary and secondary regrowth ol premon¬
lane moist forest at 1250—1400 m. Associated
there, besides the species discussed above, are the
rare Passiflora prolata Masters, and within 1 km,
P. hahnii. Local vernacular names recorded on
Steyermark s labels are murcielago de montana and
granadilla.
Paratypes. GUATEMALA. Quezaltenango: ca. 6 km
below Santa Maria de Jesus on lit. 9S, near km post 196,
19 Jan. 1980 (juv.), ./. MacDougal A Miley 597 (DUKE,
F. MEXU, MO. UVAL), [thereafter cultivated at Duke Uni¬
versity 1980—1983 as J. MacDougal 597GR (DUKE,
MO)]; betw. Finca Pirineos & Finca Soledad, along
stream, lower S-facing slopes of Volean Santa Maria, be¬
tween Santa Marfa de Jesus & Calahuache, 5 Jan. 1940
(11). J. A. Steyermark 33591 (F).
Passiflora sublanceolata (Killip) J. MacDougal,
comb, et slat. nov. Basionym: Passiflora pal-
rneri var. sublanceolata killip. Publ. Carnegie
Inst. Wash. 461: 322. 1936. TYPE: Guate¬
mala. Petcn: Uaxactun to San Clemente, 30
Apr. 1931, //. H. Bartlett 12788 (holotype, US
1492638; isotype, MICH).
This species has vividly colored flowers variously
described as intense rose, bright purple-pink, or
hot [link. The androgynophore is long (17—21 mm)
and the white or white with pink or purple corona
is reduced to 10 mm long or less. It is humming¬
bird-pollinated, as was documented by Janzen
(1968. as undetermined Passiflora sp.). From the
three specimens known to him, killip named this
as a variety of Passiflora palmeri Rose, a white and
violet-flowered subshrub endemic to Baja Califor¬
nia. We now have abundant material, as well as a
better understanding of the habitat differences be¬
tween the desert in Baja California and the wetter
though seasonally dry Yucatan Peninsula, that
shows these to be distinct. Passiflora sublanceolata
is now known from Tabasco, Mexico, to northeast¬
ern Belize, and is in cultivation.
Passiflora veraguasensis J. MacDougal, sp. nov.
TYPE: Panama. Veraguas: along road betw.
Escuela Agricola & Alto Piedra (above Santa
Ee) & Rio Dos Bocas, ca. 5-8 km from Es¬
cuela, 730-770 m. 26 July 1974 (bud, II. imm
fr), T. B. Croat 25908 (holotype, MO
2292671). Figure 7.
Haec species Passiflorae nelsonii similis, sed ah ea fo¬
lds bracteisque minoribus, neetariis petiolaribus paucior-
ibus minoribusque atque seriebus coronalibus duabus ex-
timis inter se longitudine accedentibus distinguitur.
Vine, glabrous throughout except the bracts, se¬
pals, and styles; stems terete; prophyll of the veg¬
etative bud I. widely triangular, obscurely glan¬
dular-dentate. Stipules 11 — 17 X 3.5—6 mm.
oblong-lanceolate to oblong-oblanceolate, acumi¬
nate to acute, deciduous, both margins glandular-
serrulate; petioles not glandular or 1- to 3(to 4)-
glandular near or just below the apex, the nectaries
0.8-1. 2 X 0.8-1. 7 mm; blades 8.5-13 X 6.5-11
cm. entire to obscurely glandular-denticulate es¬
pecially near the base, not variegated, sometimes
glaucous abaxially, (ovate to) widely ovate, unlobed,
abruptly acuminate; laminar nectaries marginal,
obsolescent except at the laminar base. Peduncle 1
per node, 2.1— 4.5 cm, uniflorous; bracts 4.7— 5.6 X
2.8— 3.2 cm. connate 1.0— 1.6 cm at the base, ovate,
acute to abruptly short-acuminate, glabrous except
for a narrow band of tomentum along the adaxial
margin, the margin minutely glandular-serrulate to
subentire. Flowers green-white, the corona purple
(“dark lavender," T. Croat 25908); stipe 10-13 mm,
deeply recessed into the hypanthium base; sepals
25—28 mm, the margins and apex sparsely pubes¬
cent. the subapical projection 1.5— 2.0 mm long;
petals ± equal to sepals in length, green-white;
coronal filaments in 10 to 1 1 series, the outermost
17—21 mm, the next 11—13 mm and thicker than
the outermost, the inner 8 to 9 series 0.7—3 mm
long; operculum 4.7-5 mm long, basally connate
and membranous, filamentous in distal %; audio-
460
No von
Holotype of;
p&Ujfl&l* V&t A4UA > {>£r.U
FANAMA
Province of Ve
Family
I H»efoH.i l
MISSOURI
BOTANICAL GARDEN
HERBARIUM
-K? 2292G71
del. A. Gentry !«»
Thomas B. Croat July 26, 147-
MISSOUKI BOTANICAl O.AKOFN Ml HHAHfl'V
Figure i. Photograph of the holotype of Passiflora veraguasensis at MO.
gynophore 8.5-9 mm, anthers 8-9 mm long; ovary
4.5—5 X 3—4 mm, ellipsoid, glabrous; styles short
pubescent adaxially. Fruit ca. 6 X 5 cm, widely
ellipsoid, exocarp thin, color unknown, not stipi-
tate; arils unknown; seeds 10.0—10.5 X 6.0— 7.0
mm. shallowly alveolate with 18 to 25 pits, sur¬
rounded by 1-2 mm wide almost winged, flattened
margins that are striate-dentate, the prominent cha-
lazal beak antiraphal, the micropylar end notched
or retuse.
Passiflora veraguasensis is most like the more
Volume 14, Number 4
2004
MacDougal
New Taxa of Passiflora
461
northern P. nelsonii Masters & Rose, and like it. is
assigned to subgenus Passiflora, supersection Laur-
ifolia (Cervi) Feuillet & J. MacDougal, series Tili-
ifoliae Feuillet & J. MacDougal. Passiflora nelsonii
ranges from Veracruz. Mexico, to northeastern Cos¬
ta Rica. Passiflora veraguasensis is apparently re¬
stricted to Panama. It is generally smaller vegeta-
lively, and differences include the smaller bracts
(4. 7-5.6 X 2. 8-3. 2 cm vs. 5.0-7. 5 X 3.0-5.0 cm)
and petiolar nectaries reduced in number and size
(none or 1 to 3, rarely 4 glands vs. 4 or 5, rarely
() glands that are 1-1.5 nun wide vs. 2-4 mm wide).
In the new species, the two outer series of coronal
filaments are more similar in length, with the outer
twice or less as long (outermost 17-21 mm. the next
11-13 mm). Passiflora nelsonii has its outermost
coronal row 3 to 10 times longer than the next row
(outermost 18-22. the next 1-7 mm).
This new species was discussed by Gentry
( 1976) under the name Passiflora tiliaefolia L., and
die present type collection was cited as a range
extension of that species from South America. Gen¬
try said the operculum appeared to be entire, but
it is actually filamentous for more than half its
length. Gentry’s misapplied name was repeated in
D’Arcy (1987). The butterfly Heliconius cydno
I Nymphalidae) is recorded on the label of Knapp
5765 (MO) as an herbivore.
Distribution and habitat. Passiflora veragu¬
asensis is found in western Panama near the Con¬
tinental Divide at 730-1000 m in premontane moist
to pluvial forest, at forest edges, and in forest gaps.
Phenology. Flowering April through Septem¬
ber. fruiting June through December.
Etymology. The species is named for the Pan¬
amanian province of Veraguas, where it was first
collected.
Paratypes. PANAMA. Chiriqm: al E del sitio de pre-
sa en Fortuna, 6 May 1976, Mendoza 512 (MO). Code:
lulls N of' El Valle, E slope & ridges leading to Cerro
Gaital, 27 June 1982, S. Knapp 5765 (MO). Veraguas:
2/10 mi. beyond fork in road at Escuela Agrfeola Alto
Piedra on road to Rio Calovebora, 3 Apr. 1976, T. Croat
A ./. Folsom 55857 (MO): forest & forest remnants to 12
km NW of Santa Ee de Veraguas, 6 Dec. 1975, W. D'Arcy
10546 (MO).
Acknowledgments. Much ol this work was start¬
ed while a postdoctoral researcher for the Flora
Mesoamericana project at the Missouri Botanical
Garden with support from the Jessie Smith Noyes
Foundation. 1 thank the curators of BH, BM, C,
CAS, CR, DUKE, DS. EAP. ENCB. F, GH. GOET,
K. LAGU, LL. MEXU, MICH. NY. TEFH. US. and
W IS for permitting the long-term loans necessary
for this study. Roy Gereau most generously trans¬
lated the diagnoses of the new taxa into Latin, and
Fred Keusenkothen of the MO web group scanned
the illustrated herbarium specimens. George Pilz
provided invaluable measurements from the EAP
herbarium. Our understanding of some of the spe¬
cies here was greatly enabled through their culti¬
vation. and I am indebted to the staff of the Duke
University greenhouses many years ago for their as¬
sistance in caring for those living collections. Don¬
ald E. Stone and Melvin I). Turner provided critical
support and analysis for this study. Ruth E Schal-
lert, librarian at the Smithsonian Institution for
many years, discovered the only known copy ol
Calderon’s passiflora publication. 1 am also indebt¬
ed to Peter M. Jprgensen. Gerrit Davidse, Jan Meer-
man, Connie Wolf. Kristen Porter-Utley. B. Hansen,
and Katie Hansen for assistance. Charlotte laylor
and an anonymous reviewer improved the manu¬
script. Finally, 1 thank Robert Magill for unfailing
support and access to the special research facilities
at the Missouri Botanical Garden.
Eiterature Cited
Calderon, S. [s.d., pre-1936.] Passifloras dilobatas de El
Salvador. Datos para la Rotanica Medica de El Salvador.
Escuela de Medicina, Qufmica, y Farmaeia, San Sal¬
vador.
D'Arcy, W. 1987. Flora of Panama: Checklist and index.
Monogr. Syst. Rot. Missouri Rot. Card. I 7, 18.
Feuillet, C. & J. M. MacDougal. 2003 [June 2004], A new
infrageneric classification of Passiflora E. (Passiflora-
ceae). Passiflora 13(2): 34—35, 37—38.
Gentry, A. II. 1976. Additional Panamanian Passiflora-
ceae. Ann. Missouri Rot. Card. 63: 341—345.
Greuter, W., .1. McNeill, F. R. Rarrie, H. M. Runlet. V
Demoulin, T. S. Eilgueiras, I). 11. Nicolson, P. C. Silva,
J. E. Skog, P. Trehane, N. J. Turland & D. E. Hawks-
worth (editors). 2000. International Code of Rotanieal
Nomenclature (Saint Louis Code). Regnum Veg. 138.
Grisebach, A. 1858. Novitiae florae panamensis, Passiflo-
raeae. Ronplandia 6(1): 7.
Harms, 11. 1893. Cher die Verwertung des anatomischen
Raues fUr die Umgrenzung und Einteilung del Passi-
floraceae. Rot. Jahrb. Syst. 15: 548—633.
Janzen, D. H. 1968. Reproductive behavior in the Passi-
(loraceae and some of its pollinators in Central America,
behaviour 32: 33-48.
Killip, E. P. 1936. Passifloraceae of the Mayan region.
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- . 1938. The American species of Passifloraceae.
Publ. Field Mus. Nat. Hist., Rot. Ser. 19: 1-613.
Klucking, E. P. 1992. Leaf Venation Patterns, Vol. 6. J.
Cramer, Rerlin.
Knapp, S. & J. Mallet. 1984. Two new species of Passi¬
flora (Passifloraceae) from Panama, with comments on
their natural history. Ann. Missouri Rot. Card. 71:
1068-1074.
MacDougal, J. M. 1989. Two new species of Passiflora
section Decaloba (Passifloraceae) from Costa Rica. Ann.
Missouri Rot. Card. 76: 608—614.
- . 1994. Revision of Passiflora subgenus Decaloba
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section Pseudodysosmia (Passifloraceae). Syst. Hut.
Monogr. 41: 1-146.
& A. K. Hansen. 2003. A new section of Passi-
flora. subgenus Decaloba (Passifloraceae), from Central
America, with two new species. Novon 13: 459-466.
Masters, M. T. 1871. Contributions to the natural history
of the Passifloraceae. I rails. Linn. Soc. London 27-
593-645.
- . 1872. Passifloraceae. Pp. 529-628 in Flora Bras-
iliensis 13(1), ed. A. W. Fiehler, Munich.
Matilda. I’.. 1950. A contribution to our knowledge of wild
flora of Ml. Ovando. Amer. Midi. Naturalist 43- 195-
223.
Planchon. J. F. 1849. FI. Serres .lard. Fur. 5: 528.
Standley, P. C. & S. Calderdn. 1925. Passifloraceae. Pp.
154—155 in l.ista preliminar de las plantas de FI Sal¬
vador. Typografia La Unidn-Dutriz Hermanos, San Sal¬
vador.
- & - - • 1941 [ 1 9447 1 . Passifloraeae. Pp. 197-
199 in l.ista Preliminar de las Plantas de FI Salvador,
2nd ed. Imprenta nacional, San Salvador.
& L. (). Williams. 1961. Flora of Guatemala. Pas¬
sifloraceae. Fieldiana Hot. 24(7): 115-146.
I liner. I. A J. M. MacDougal. 2004. Passijlora : Passion¬
flowers ol the World. Timber Press, Portland.
Vanderplank, J. 1996. Passion Flowers, 2nd ed. MIT
Press, Cambridge.
- . 2000. Passion Flowers, 3rd ed. MIT Press, Cam¬
bridge.
New Taxa and Names in North American Ancistrocarphus ,
Diaperia , and Logfia (Asteraceae: Gnaphalieae: P dagininae) and
Related Taxa
James I). Morefield
Nevada Natural Heritage Program, 1550 East College Parkway, Suite 137, Carson City,
Nevada 89706-7921, U.S.A. jdmore@heritage.nv.gov
ABSTRACT. Ancistrocarphus keilii, a distinctive,
rarely collected, previously undescribed, and re¬
cently rediscovered species endemic to Santa Bar¬
bara County, California, is formally named. Diaper¬
ia is resurrected for three North American species
usually placed in, but clearly distinct from, Evax.
The name Evax verna is neotypified to be consistent
with current usage that includes Evax multicaidis
las a new' synonym), and the new combinations Dia¬
peria verna and Diaperia verna var. drummondii are
provided. Diaperia prolifera var. barnebyi, a long-
collected but previously undescribed variety from
southeastern New' Mexico, western Iexas, and
southwestern Oklahoma, is also formally named.
Among Cassini's anagrammatic segregates from Ei-
lago, the priority of Logfia over Oglifa is at best
ambiguous, but is adopted to preserve current us¬
age in the Old World, pending a proposal to con¬
serve. A priorable epithet being available, the new
combination Logfia filaginoides is made for the spe¬
cies long known as Eilago californica (newly syn-
onymized), including Eilago californica var. tomen-
,'osa (also a new synonym). The correct basionym
of Oglifa is established as Gnaphalium subg. Og¬
lifa, contrary to earlier interpretations.
Key words: Ancistrocarphus, Asteraceae, Com-
positae, Cassini, Diaperia, Evax, Kilagininae, Ei¬
lago, Gnaphalieae, Gnaphalium. Ixigfia, Micropsis,
North America, Oglifa.
During field and herbarium studies of several
thousand specimens and populations of Asteraceae
subtribe Kilagininae Bentham & Hooker f. for a
systematic analysis of the subtribe (Morefield,
1992a), for the Jepson Manual (Hickman, 1993),
and soon for the Flora of North America (Flora of
North America Editorial Committee, 1 993+), sev¬
eral previously undescribed taxa have been named,
and the circumscriptions and distributions of cer¬
tain genera and smaller taxa have been clarified
(Morefield, 1992b, 1992c, 1992d). In order to con¬
tinue this process for the Flora of North America
project, several new taxa, names, and circumscrip¬
tions are proposed or reported herein.
The capitula of Filagininae, though small, are
unusually complex for Asteraceae, particularly in
the organization of florets and receptacular bracts
(paleae; see diagram in Felger, 2000: 101: More¬
field & Felger, 2000) and in the minute but system¬
atically important variation of the paleae. for this
reason, certain conventions of terminology have
been adopted for the upcoming Flora of North
America treatments and are used here also for ef¬
ficiency and comparability. The disciform capitula
of Filagininae possess two or more outer series of
pistillate llorets surrounding relatively few, central,
functionally staminate or sometimes bisexual flo¬
rets. The paleae and other structures are referred
to according to the sexual condition of their asso¬
ciated florets as, for example, pistillate paleae or
staminate corollas or bisexual cypselae. All florets,
or at least the outermost pistillate florets, are sub¬
tended (and often enclosed to various degrees) by
a palea. Each palea often is further differentiated
into a central and proximal body and a scarious,
reflexed, marginal and distal wing. 1 he outer pis¬
tillate paleae have assumed the involucral function
in most species, the true involucre and its phy Ma¬
ries being present outside the paleae, but usually
strongly reduced and/or differentiated, and some¬
times vestigial.
The leaves that immediately subtend capitula
and/or glomerules are often differentiated, and are'
termed capitular leaves. Branches may also imme¬
diately subtend capitula or glomerules; il so, ca¬
pitular leaves collectively subtend such branches
and their capitula, and capitula appear to be sessile
in forks of pseudo-dichotomies or -polytomies.
Sometimes capitular leaves subtend only glomer¬
ules and not individual capitula, and individual ca¬
pitula may be difficult to distinguish within glom¬
erules.
The descriptions of new taxa herein include all
differential traits of systematic significance within
Novon 14; 463-475. 2004.
464
Novon
Filagininae, so as to be comparable with previously
published descriptions (Morefield, 1992b, 1992c).
Ancistrocarphi s Expands
I wo peculiar specimens of diminutive, stemless
annuals collected in Santa Barbara County, Cali¬
fornia. by Ralph Hoffmann in the spring of 1929
and labeled as Hesperevax (A. (bay) A. Gray were
found among the collections of the Santa Barbara
Botanical Garden in the late 1980s. I'heir generic
affinities were not immediately evident, and there
was considerable doubt that the species still sur¬
vived amidst the relatively dense human population
of its locality after six decades. Further phyloge¬
netic study based on morphology placed the un¬
named taxon as the sister species of Ancistrocar-
phus filagineus A. Gray, heretofore the sole species
in the genus, with a high degree of confidence
(Morefield, 1992a). A recent collection by David J.
Keil greatly clarified the distribution and habitat of
the species and demonstrated that it still survives
in the wild.
Ancistroearphus keilii Morefield, sp. nov. TYRE:
U.S.A. California: Santa Barbara Co., Vanden-
berg Air Force Base, North Base, Pine Canyon
Road betw. Utah Ave. & Lompoc Gate, Burton
Mesa Chaparral & coastal live oak woodland,
local in minute herb carpet beneath shrubs, 4
Mar. 1995, D. Keil 24561 & L. 1). Oyler (ho-
lotype, OBI 054629; isotype, MO). Figure 1.
Anris! rocarphu filagineo similis, scd caulibus null is; lo¬
ins oninino basalilms valde petiolatis, busibus folioruni
valde expansis eircum capitula arete involucratis; paleis
staminatis paleis pistillatis plerumque brevioribus nee
longioribus, 1 .8— 2.8 (ad invicem 2.7— 4.1) mm longis late
spathulatis non lanceolatis; apieibus palearuin stamina-
tarum obtusis rotundatisve apiculatis non aeurninatis el
apiculis scariosis non spiniformibus; cypselis 1-1.4 X
0.5— 0.6 (ad invicem 1.4—2 X 0.6— 0.9) mm non atrofas-
eiatis; phyllari is vestigialilnis; reeeptaeulis obpandurifor-
m i bus non fungiformibus; et apieibus palearuin pistilla-
tarum al is scariosis indistinetis lanceolatis nee distinetis
nee ovatis; Hesperevaci acauli var. acauli similis, sed re¬
eeptaeulis inter paleas glabris; paleis pistillatis trinervatis
saeeatis tarde deeiduis nee apertis nee permanentibus,
unaquaeque eypselain perfeete includenti; basilius pa-
learum slaminatarum ad maturitates patulescentibus; et
apieibus paleamm staminatarum ad maturitates valde in-
eurvatis apiculatis non rotundatis et apiculis scariosis non
herbaeeis.
Diminutive aeaulous annuals 0.5-1 (-2) cm high
from taproots (sometimes densely clustered, ap¬
pearing as I fibrous-rooted plant), plants grayish
green, arachnoid-sericeous throughout. Leaves all
basal, rosulate. simple, strongly petiolate, up to
10(-18) X I (-2) mm, longest 3-6X capitula lengths.
abruptly differentiated from paleae, or 1 or 2 ±
grading into paleae by reduction of blades and ex¬
pansion of petiole bases, sometimes losing blades
altogether and then resembling long-aristate phyl-
laries; petioles mostly 1-3X blade lengths; petiole
bases strongly expanded, ± parchment-like in tex¬
ture, yellowish to tan. prominently 3-veined, closely
involuerate; blades oblaneeolate to obovate. ± pla¬
nar to folded, pliant, entire, uniformly arachnoid-
sericeous, primary vein I. subordinate venation ob¬
scure. apices acute to rounded, mucronate.
Capitulescences 1, unbranched, each consisting of
I sessile capitulum among leaves, sometimes ap¬
pearing to form dense leafy glomerules of 2 to 10,
but these consisting instead of densely packed I-
headed plants with separate taproots (apparently
having germinated in-place from intact capitula of
previous years plants); capitula disciform, fully pa-
leate, erect, ± spheric, 2.5— 3.5 mm diam.; invo¬
lucres vestigial, functionally simulated by petiole
bases and paleae; receptacles broadly hourglass¬
shaped or obpanduriform, 0.9-1. 3 mm long, I-2X
as long as wide, glabrous; pideae about 10 to 17 in
2 or 8 spirally ranked series, Iree throughout, veins
parallel, not reaching apices, apices shortly gla¬
brous; pistillate paleae falling separately (at least
upon dissection, tardily so or sometimes not at all
in nature), in 1 or 2 series, imbricate, erect to as¬
cending, saccate, 3-veined (central vein thinning,
obscure with age. lateral veins remaining promi¬
nent), each embracing a floret, ultimately complete¬
ly enclosing a cypsela, gently incurved, ± ovate,
longest 2. 3-3. 3 mm long, wings indistinct, contin¬
uous across apices ol paleae, incurved (with styles)
over staminate florets at anthesis, at maturity ± in¬
volute, incurved, shiny, translucent, obscurely stri¬
ate, lanceolate, entire, glabrous, bodies brownish,
± navicular, not gibbous, obcompressed, thinly
cartilaginous between veins, abaxial surfaces ±
rounded transversely, arachnoid distally near mar¬
gins or sometimes wholly glabrate, adaxial surfaces
arachnoid to thinly lanuginose; staminate paleae
persistent, 5(to 7), whorled, erect at anthesis, at
maturity somewhat expanded, open, concave, ob¬
scurely veined, broadly spatulate, 1 .8-2.8 mm long,
slightly surpassed by pistillate paleae, ± uniformly
thickened, cartilaginous, greenish to brownish on
both surfaces, abaxially glabrous or distally arach¬
noid, adaxially arachnoid to thinly lanuginose ex¬
cept for apicula, bases spreading at maturity, ±
cuneate to rounded, apices ± strongly incurved,
somewhat involute, entire, shortly scarious-apieu-
late, otherwise obtuse to rounded, neither spinose
nor uncinate. Pistillate florets 5 to 11 in 1 or 2
spirally ranked series from proximal portions of re-
Volume 14, Number 4
2004
Morefield
North American Asteraceae
465
Figure 1. Ancistrocarphus keilii Morefield. — A. Plant in late maturity. — B-E. Leaves showing variation in size and
shape. — F. Receptacle with staminate florets attached, showing outlines of staminate paleae (two in front removed) and
pistillate paleae (bases only, those in front and hack removed) in attached positions. G— K, Views of pistillate palea.
G. Adaxial view. — II. Diagram of longitudinal cross section in lateral view. — J. Abaxial view. K. Diagram of
transverse cross section in top view. — L. Mature cypsela, showing position within pistillate palea. — M. Pistillate
corolla. — N. Staminate floret. — P. Staminate palea in adaxial view. Drawn from Keil 24561 & Oyler, OBI 054629 (B-
I), G— P) and Hoffmann s.n., SBBG 063224 (A. E. and 1).
eeptacles, each hidden and nearly enclosed by a
subtending palea, pistillate corollas whitish, fili¬
form. limb absent, styles exserted but surpassed by
wings of subtending paleae; staminate florets 3 to 6
from receptacle apices, collectively and loosely
subtended by whorl of staminate paleae. ovaries
vestigial, up to 0.1 mm long, staminate corollas ±
concealed by pistillate paleae at anthesis, later vis¬
ible, somewhat bilaterally symmetric. 0.9— 1.3 mm
long, ± fnnnelform, glabrous, lobes 4, reddish pur¬
ple to yellowish brown, debate, blunt, unequal with
1 or 2 enlarged, stamens included, styles strongly
exserted. Cypselae each remaining enclosed by a
palea. brown, without black band, obovoid, apicu-
late, not angular, slightly incurved, not gibbous, ob-
compressed, 1.0— 1.4 X 0.5— 0.6 mm. glabrous,
smooth, dull, corolla scars apical; pappi none.
Etymology. The tiew species is named in honor
of David J. Keil (b. 1946) of California Polytechnic
State University in San Luis Obispo, consummate
student of and expert on the flora of California and
the southwestern United States in general (and of
Asteraceae in particular) and collector of the ho-
lotype. Because Ancistrocarphus keilii is likely to
become a focus of conservation concern, and ap¬
propriate common names are very helpful in such
circumstances, the common name Santa Ynez
groundstar is here suggested, based on its geo¬
graphic range and its distinctive habit.
466
Novon
I lie three gatherings of Ancistrocarphus keilii
thus far known, including the two historical para-
types cited below, comprise about 180 individual
plants. Based on these collections, the species ap¬
peal's confined to the Santa Ynez Kiver drainage of
southwest Santa Barbara County, California, where
it is known from sandy soils in chaparral vegetation
bordering oak woodlands between 40 and 130 m
elevation. Plants in reproductive condition have
been collected between 4 March and 12 April, and
flowering and fruiting therefore likely occur some¬
time between late February and the end of April or
early May in any given year.
In contrast with the five stiffly and inwardly
hooked staminate paleae of Ancistrocarphus filagi-
neus , the new species has no obvious dispersal
mechanism. Ancistrocarphus keilii appears to grow
annually in dense turf-like associations comprised
of dozens to perhaps hundreds of individuals, sug¬
gesting that the cypselae germinate more or less in-
place from previous years' populations. Lack of dis¬
persal mechanisms is also consistent with the very
limited geographic range of the species. It is allo-
patric with A. filagineus , which in Santa Barbara
County is found farther inland to the east and north.
Unlike many members of the subtribe, no other
species of l ilagininae have been found mixed with
A. keilii, suggesting that its habitat is fairly distinc¬
tive.
Because of its very' limited geographic range and
dispersal ability, and its location in a region of rel¬
atively dense human population with a strong po¬
tential for future impacts, Ancistrocarphus keilii
should be of immediate conservation concern. Its
conservation status should be reviewed and care¬
fully monitored by the State of California, the U.S.
Fish and Wildlife Service, Vandenberg Air Force
Base, and other appropriate agencies. Because of
the small and inconspicuous nature of the plants,
surveys resulting from such reviews may well show
the species to be somewhat more widespread than
current documentation indicates.
In phylogenetic reconstructions based on exten¬
sive morphologic data sets, including the new spe¬
cies and all other known laxa of Filagininae, An¬
cistrocarphus consistently resolved as the sister
genus to Hesperevax (Morefield, 1992a). The two
genera share strongly expanded staminate paleae
among other features. Ancistrocarphus is apparently
allied more with the genus Evax Gaertner and its
segregates than with Stylocline Nuttall, where it has
frequently been placed by v irtue of sharing saccate
pistillate paleae. The pistillate paleae of Ancistro¬
carphus differ from those of Stylocline and all other
filagininae in having two prominent lateral veins
in addition to the central one.
Addition of a second species to Ancistrocarphus
renders the genus considerably more heterogeneous
and partially bridges the gap between it and Hes¬
perevax (near //. acaulis (Kellogg) F. Greene var.
acaulis in particular). With the latter genus the new
species shares the (often) stemless habit, strongly
petiolate leaves, expanded petiole bases (though
thinner and 3-veined in the case of Ancistrocarphus
keilii ), vestigial involucre, spatulate and obtuse sta¬
minate paleae, proximally expanded receptacle,
and somewhat bilaterally symmetric 4-lobed sta¬
minate corollas. It is possible that Ancistrocarphus
keilii is derived from a common ancestor of, or a
hybrid between, the two genera. It currently shares
the majority of its character states with A. filagi¬
neus, however, and the two consistently resolved as
sister species in all phylogenetic analyses (More-
field, 1992a).
Key to Hesperevax and the Species of Ancistrocarphus
la. Receptacles setose among paleae; pistillate pa¬
leae open, obscurely veined, persistent, cypselae
falling free; staminate paleae at maturity erect
proximally, erect to strongly spreading distally,
apices herbaceous throughout, ohtuse to rounded
. Hesperevax
lb. Receptacles glabrous; pistillate paleae saccate,
3-veined (central vein sometimes thinning with
age, becoming obscure), tardily falling, each
completely enclosing a eypsela; staminate paleae
at maturity spreading proximally, strongly in¬
curved distally, apices at least shortly non-her-
baceous, apiculate to acuminate . . Ancistrocarphus
2a. Caulescent (usually strongly so); leaves bas¬
al and eauline, sessile or broadly and indis¬
tinctly petiolate, bases attenuate, not or
scarcely involucrate; mature staminate pa¬
leae 2.7-4.1 mm long, surpassing the pistil¬
late paleae, broadly lanceolate, apices long-
spinose, uncinate, acuminate; cypselae 1.4—
2 X 0.6-0.9 mm, with proximal black band;
involucres of mostly 3 to 6, minute, scarious,
persistent phyllaries; receptacles ± fungi¬
form; apices of pistillate paleae with dis¬
tinct, ovate wings . . Ancistrocarphus filagineus
2b. Acaulous; leaves all basal, strongly petio¬
late, petiole bases strongly expanded, close¬
ly involucrate; mature staminate paleae 1.8—
2.8 mm long, slightly surpassed by pistillate
paleae, broadly spatulate, apices shortly
scarious-apiculate, otherwise obtuse to
rounded, neither spinose nor uncinate; cyp¬
selae 1 — 1.4 X 0.5-0. 6 mm, without black
band; involucres vestigial (simulated by pa¬
leae and petiole bases); receptacles obpan-
duriform; apices of pistillate paleae with in¬
distinct, lanceolate wings .
. Ancistrocarphus keilii
Paratypes. U.S. A. California: Santa Barbara Co.,
Volume 14, Number 4
2004
Morefield
North American Asteraceae
467
Buellton to Lompoc, sandy soil, 30 Mar. 1929, R. Hoff¬
mann s.n. (SBBG 063223); Buellton to Lompoc, sandy
border of woods, 12 Apr. 1929, R. Hoffmann s.n. (SBBG
063224, NY, UC).
Diaperia Dkparts Evax
Three closely similar species of the central Unit¬
ed States and adjacent Mexico have usually been
placed in Evax (or in Filago L. when the two genera
have been merged). As with Hesperevax of Califor¬
nia and Oregon (Morefield, 1992c), however, these
three species form a monophyletic group only dis¬
tantly similar to typical Evax species (Morefield,
1992a), and they therefore warrant treatment as a
separate genus, the earliest available name for
which is:
Diaperia Nuttall, Trans. Amer. Philos. Soc. n.s., 7:
337. 1840. Evax Gaertner sect. Diaperia (Nut-
tall) A. Gray, Syn. FI. N. Amer. 1(2): 229.
1884. TYPE: Evax prolifera Nuttall ex DC.
Calymmandra Torrey & A. Gray, FI. N. Amer. 2(2): 262.
1842. Evax sect. Calymmandra (Torrey & A. Gray)
A. Gray, Syn. FI. N. Amer. 1(2): 229. 1884. TYPE:
Calymmandra Candida Torrey & A. Gray.
Filaginopsis Torrey & A. Gray, FI. N. Amer. 2(2): 263.
1842. TYPE: Evax multicaulis DC. (lectotype, des¬
ignated by Pfeiffer, 1874: 1347).
Based on phylogenetic analysis of 120 morpho¬
logic characters for all known species and varieties
of Filagininae, Diaperia appears most closely re¬
lated to Evax sect. Filaginoides Smoljaninova of the
Mediterranean basin and central Asia, and partic¬
ularly to Evax eriosphaera Boissier & Heldreich of
the eastern Mediterranean region (Morefield,
1992a). The two groups of species are similar in
many ways, but also exhibit several differences that
likely justify placing Evax sect. Filaginoides in yet
another genus separate from Evax (no name at that
rank is yet available).
Key to Diaperia and Related Filagininae
la. Pappi present on some florets; or pistillate paleae
predominantly saccate or spreading at maturity;
or staminate paleae absent or spreading at ma¬
turity or adaxially hairy . other Filagininae
lb. Pappi absent on all florets; pistillate paleae pre¬
dominantly open, flat to loosely folded, remain¬
ing erect; staminate paleae present, remaining
erect, glabrous or abaxially hairy.
2a. Paleae persistent, free throughout ....
. Evax (excluding sect. Filaginoides)
2b. Paleae falling, staminate and at least inner
pistillate ones as a unit, cohering distally by
dense tangled hairs.
3a. Staminate paleae about equaling pistil¬
late paleae, open throughout, apices
acuminate or cuspidate, very shortly
glabrous distal to pubescence, ± hya¬
line, erect, not forming cavities in the
mass of hairs; plants always compact,
less than 3 cm high, pulvinate to round¬
ed . Evax sect. Filaginoides
3b. Staminate paleae slightly enlarged, be¬
coming ± closed proximally, apices ob¬
tuse, sericeous-lanuginose to tips, ±
herbaceous, somewhat spreading at ma¬
turity, thereby forming obconic cavities
in the mass of hairs; plants usually with
well-developed stems 3-25 cm high,
simple or loosely branched .... Diaperia
Diaperia consists of the following three species
and two additional varieties of the central United
States and northern Mexico, for which typification
and current nomenclature are here established, and
of which one variety is described as new.
Diaperia Candida (Torrey & A. Gray) Bentham &
Hooker f., Gen. PL 2(1): 298. 1873. Calym¬
mandra Candida Torrey & A. Gray, FI. N.
Amer. 2(2): 262. 1842. Evax Candida (Torrey
& A. Gray) A. Gray, Syn. FI. N. Amer. 1(2);
230. 1884. Filago Candida (Torrey & A. (fiav)
Shinners, Sida 1: 252. 1964. TYPE: U.S.A.
Texas: s.d., T. Drummond 177 (holotype, GH;
isotypes, BM[3], CGE, GH, K[2], OXF).
Some of the type sheets are dated 1835 or 1836,
but these apparently reflect the years various sets
were received, since Thomas Drummond died in
Cuba in Mareh 1835. The plants on all the sheets
appear to be of the same gathering. The notation of
“San Filipe” on one sheet at K could refer to lo¬
cations in present-day Austin or Hidalgo Counties,
Texas, both areas having produced more modem
specimens of Diaperia Candida. The species occu¬
pies a limited range covering eastern Texas and the
adjacent corners of southwestern Arkansas, north¬
western Louisiana, and southeastern Oklahoma.
Within Diaperia, D. Candida is aberrant in its
bisexual disk florets and its reported chromosome
complement of 2 n = 711 (Keil & Pinkava, 1976)
and might eventually justify resurrection of the
monotypic genus Calymmandra (see above), though
I hesitate to propose this without further study. The
chromosome count is of particular interest, as all
of the other 25 counted species of Filagininae have
2 n = 1411 (representatives of Evax , Filago, Digfia
Cassini, Micropus L., Psilocarphus Nuttall, and
Stylocline) or 2 n = 1311 (a few Diaperia and Evax).
This might imply that the vast majority of Kilagi-
ninae are tetraploid with Diaperia Candida being
an ancestral diploid, a hypothesis with no phylo¬
genetic support (Morefield, 1992a). It is also pos¬
sible that Keil and Pinkava (1976) accidentally
468
Novon
mis-translated a rneiotic count of n = 14 to a mi¬
totic equivalent of 711 instead of 1411.
Diaperia verna (Rafinesque) Morefield, comb,
nov. Rasionym: Evax verna Rafinesque, Atlan¬
tic J. 6: 1 78. 1833. Filago verna (Rafinesque)
Simmers. Sida I: 253. 1964. TYPE: U.S.A.
Louisiana: Claiborne Parish, Salem Cemetery
E of La. 9 in Athens, Sec. 6, T19N, R6W, 5
May 1979. R. I). Tlwma.s et al. 64261 (neo¬
type, designated here, NLU 157605; isotypes,
COLO 388762, NY. RSA 301867, UT 107742.
UTEP 20825, VDB 185027).
Evax multicaulis DC., Prodr. 5: 459. 1836. Syn. nov. Dia-
peria multicaulis (DC.) Nuttall, Trans. Amor. Philos.
Soc. n. s., 7: 338. 1840. Filaginopsis multicaulis
(DC.) Toney & A. Gray, FI. IN. Amer. 2(2): 263.
1842. Filago nivea Small, Dull. Toney Bot. Club 24:
333. 1897, non Filago multicaulis Lamarck, FI.
Franc. 2: 59. 1779. TYPE: U.S.A. Texas: “in campis
Sn Fernando de Bexar," June 1829, ./. L. Berlandier
210V (leetotype, designated here, ((-DC, II)C micro¬
fiche 800. 906.111.4 at BSA).
The name Evax verna Rafinesque (1833a) was
heretofore of uncertain application. I)e Candolle
(1836: 459) mentioned the name (misspelled as
“ Evax vera ”) under “species dubia” at the end of
bis Evax treatment, noting “an huie generi adnu-
meranda?' (“perhaps to be counted with this ge¬
nus?”). Toney and Cray (1842: 263) cited Evax
verna in synonymy with their (thus incorrect) com¬
bination Filaginopsis multicaulis, citing only uRaf.!
herh.' It is unclear from the context whether this
citation referred to a specimen seen or just to the
reprinted description in Herbarium Rafinesquianum
(Rafinesque, 1833b). In any case, neither I nor ap¬
parently any other subsequent workers have been
able to locate original type or leetotype material
either at CH or at numerous other herbaria
checked, thus necessitating designation of a neo¬
type. Gray's (1884) Synoptical Flora of North Amer¬
ica recognized Evax multicaulis and all the other
species now included in Diaperia, but did not men¬
tion Evax verna in any context. Shinners (1964)
took up Rafinesque's epithet again without expla¬
nation for the species including Evax multicaulis,
and subsequent workers have used Evax verna or
Evax multicaulis in about equal proportions.
In the absence of any original type material, only
Rafinesque's (1833a: 178) protologue is available
to guide selection of a neotype: “20. Evax verna
Raf. Caneseens sericea, Caule graeile subramoso
Fol. laxis semiamplex. obi. obtusiusc. infimis eu-
neatis, El. solit term, bract, ineq. fol. similis, per-
iantho semiglob, squatnis paucis subrot. — Texas &
Louisiana, triuncial, II. white, floscules greenish.”
In some respects, Rafinesque's description above
does not match the plants de Candolle (1836: 459)
named Evax multicaulis. In particular |my inter¬
pretations in brackets], the traits “fol. . . . semiam¬
plex.” (with the leaves . . . somewhat clasping), “fl.
solit. . . .” (with the "floscules” [glomerules? capit-
ula? flowers?] solitary), and “periantho . . . squamis
paucis subrot.” (with the perianth [involucre] . . .
with scales [phyllaries] few, subrotund) are not con¬
sistent with the type material of Evax multicaulis.
It is clear from Rafinesque's “triuncial” (3 inches
high), however, that he was probably describing
young and incompletely developed material and
that, allowing for some misinterpretation of struc¬
tures, such material could easily have belonged to
the same species de Candolle later named Evax
multicaulis. The largest plants of the young neotype
material selected above are about 3 inches high,
and their incompletely developed capitular leaves
do very much resemble “subrotund scales” of in¬
volucres (though the true involucres are vestigial in
this species). Based on the description “fl. white,
floscules greenish.” the term floscules would most
likely refer to the glomerules of capitula, and these
can indeed be solitary on young plants before they
have branched, as they are on several plants of the
neotype material. The cauline leaves of the neotype
material are slightly expanded near the bases, giv¬
ing the impression that they could be “somewhat
clasping.” although these bases are usually planar
and not folded about the stem. Given the size of
the material that Rafinesque described, then, tlx*
neotype selected above is not in serious conflict
with the protologue (((renter et al.. 2000. Art. 9.17).
Furthermore, no better lit to the protologue can
be found among 3-inch-high plants of any other
annual Gnaphalieae (as reasonably implied by Raf-
inesque’s placement in Evax) known from Texas
and Louisiana then or now, including species of
Diaperia, Gnaphalium L., and Facelis Cassini.
These species have plants that are not yet in (lower
when 3 inches high, numerous non-terminal glom¬
erules, involucres that are far from hemispheric,
numerous obvious phyllaries that are not subro¬
tund. and/or no structures that could be confused
with a few subrotund phyllaries. The only other
candidate species that comes close to matching the
protologue is Diaperia Candida, and no workers
have ever suggested that Evax verna belonged with
that species. The neotype above was therefore se¬
lected to preserve current and previous application
of Evax verna to the species de Candolle named
Evax multicaulis.
Collection numbers appeared on the neotype la¬
bels as “64261 & 2122,” the second number clear-
Volume 14, Number 4
2004
Morefield
North American Asteraceae
469
lv belonging to I.. Lewis or one of the other sec¬
ondary collectors listed.
The additional type materials of Evax multicaulis
that I examined, but rejected as lectotype, were:
U.S.A. Texas: “in campis area S[an] Antonio de
Bexar," variously June 1829 or s.d., J. L. Berlandier
1958 (syntype, G-DC, I DC microfiche 800.
906. III. 5 at RSA; isosyntypes, GH. K. LE, MO,
0 X F. PH); “Lac de St. Nicolas [ires la . . . | illegible]
. . . St. Espirit,” June 1829, ,/. L. Berlandier I95B
(syntype, G-DC. IDC microfiche 800. 906.111.4 at
USA); “in campis S[an| Fernando de Bexar, vari¬
ously June 1828 or June 1829 or s.d.. Berlandier
2109 (isosyntypes, BM, CH, k, LE). Simmers
(1951) took both locations cited in de Candolle’s
(1886: 459) protologue to be in present-day Texas.
All of the type material seen definitely represents
the same taxon. Among the three G-DC syntvpes
found, the location and dale information on the lec¬
totype chosen above most (dearly matches the pro¬
tologue. (This is contrary to my earlier annotations
of Berlandier 1958 specimens as isolectotypes,
which should now be corrected.)
A specimen of Diaperia verna (var. verna) at CAS
(U.S.A. Texas: “Camp bei la Grange," s.d.. A.
Seheele [or F. Roemerl] s.n., CAS 7254 [Hb. A. Pia¬
get)) may represent type material of Filaga texana
Seheele. The description of Filago texana in the
protologue (Linnaea 22|2|: 164. 1849), however, re¬
fers to a different plant with bisexual and pappose
central florets. The plants named Micropus minimus
DC.. Prodr. 5: 461. 1836. likely also represent this
species, but the type specimens examined (Mexico.
Monterrey: “pres Monterey, Jan. 1828, Berlandier
1861, BM. E [Hb. Delessert photograph |. LE) are
all too young to lie determined with certainty,
though no other similar species are known from that
state. Evax nivea (Small) Cory, Rhodora 38: 407.
1936, is a superfluous combination based on Filago
nivea. The range of Diaperia verna is centered in
Texas, and extends to Arizona. Arkansas. Georgia,
Oklahoma, Louisiana, South Carolina, and northern
Mexico.
I agree with previous workers that the material
named Filaginopsis drummondii Torrey & A. Gray
is best treated as varietally distinct from Diaperia
verna , necessitating the following combination:
Diaperia verna (Rafinesque) Morefield var.
drummondii (Torrey & A. Gray) Morefield,
comb. nov. Basionym: Filaginopsis drummon¬
dii Torrey & A. Gray, Id. N. Amer. 2(2): 263.
1842. Diaperia drummondii (Torrey & A.
Gray) Bentham & Hooker f., Gen. PI. 2(1):
298. 1873. Evax multicaulis DC. var. drum¬
mondii (Torrey & A. Gray) A. Gray, Syn. 1 1.
N. Amer. 1(2): 229. 1884. Filago verna (Raf-
inesque) Shinners var. drummondii (Torrey &
A. Gray) Shinners, Sida I: 253. 1964. Evax
verna Rafinesque var. drummondii (Torrey &
A. Gray) Kartesz & Ghandi, Phytologia 71:
272. 1991. TYPE: U.S.A. Texas: s.d.. T. Drum¬
mond 176 (lectotype, designated by Shinners
(1951: 126), GH; isotypes, BM[2], CGE[3],
GH[2], K[2], NY[2], OXF, PH).
Among all the type material cited above, the
sheet at NY from the Torrey Herbarium and the two
of three sheets at GH bearing “ Filaginopsis drum¬
mondii" in what appears to be Asa Gray’s hand¬
writing are appropriate candidates for the lectotype
specimen. All of the plants on all specimens cited
above are uniform in appearance and seem to be
from the same gathering. All but the lectotype sheet
bear Drummond’s collection number 176. likely re¬
flecting a simple omission. The different years
(1835, 1836, 1839) appearing on a few sheets
doubtless indicate when the specimens were re¬
ceived. since Thomas Drummond died in Cuba in
March 1835. Nothing in the protologue supports
rejection of any of the three lectotype candidates
over the others, so I here follow Shinners’s (1951)
designation and annotation (as “type ) of the un¬
numbered specimen at GH as lectotype.
Some of the isotypes variously bear “Texas 111”
or “Third Collection” or “Coll. Ill,' apparently re¬
ferring to a particular set of Drummond specimens.
One of the sheets at k also indicates the location
“San Felipe.” This could refer to locations in pre¬
sent-day Hidalgo County, Texas, where variety
drummondii is also documented by two gatherings
in 1941, or in Austin County, Texas, whence I have
seen no specimens although occurrence there is
plausible. The variety occurs along and up to 300
km inland from the Gull of Mexico in Texas, with
a disjunct record in coastal Alabama. It likely also
occurs in adjacent Tamaulipas, Mexico.
Diaperia prolifera (Nuttall ex DC.) Nuttall, leans.
Amer. Philos. Soc. n.s., 7: 338. 1840. Evax
prolifera Nuttall ex DC., Prodr. 5: 459. 1836.
Filago nuttallii Shinners, Sida 1: 253. 1964,
non Filago prolifera Pomel, Nouv. Mat. H. All.
I: 47. 1874. TYPE: U.S.A. Oklahoma: Choc-
law Co., Red River at Kiamichi River, prairies.
June 1819. T. Nuttall s.n. (holotype, G-DC,
IDC microfiche 800. 906. 1 1 1.3 at RSA; iso¬
types, BM, CGE, GH[2], K[2], NY, PH[2]).
Diaperia prolifera occupies a broad crescent from
ill*' intermediate and shortgrass prairies of western
470
No von
South Dakota and southeastern Montana, south and
east to eastern New Mexico, Texas, and southwest¬
ern Arkansas, and then to the chalk prairies of
southern Mississippi and southern Alabama. The
following is a long-collected but previously un¬
named variant of Diaperia prolifera from southeast¬
ern New Mexico, western Texas, and southwestern
Oklahoma:
Diaperia prolifera (Nuttall ex DC.) Nuttall var.
barnebyi Morefield, var. nov. TYPE: U.S.A.
New Mexico: Lincoln Co., 12 road mi. E of
Picacho Post Office, along draws m dry lime¬
stone hills, with Fouquieria & Dasylirion,
1440 m, 22 May 1987, II C. Barneby 18233
(holotype, NY; isotypes, GH, MO, NMC, RSA,
TEX, UTEP).
Diaperiae proliferae var. proliferae si nulls, sed plant is
ubique argenteis arete sericeis non luxe lanuginosis; foliis
juxta eapitula valde indureseentibus basaliter carinatis el
semper erectis involucrum rigidum secondarium fomian-
tifius; ramis eapitulescentiarum valde ascendentibus er-
ectisve saepe inaequalibus aliquot valde reductis; glome-
rulis amplissimis I (ad 3) (ad invicem 4 ad 40 + ) eapitula
continentibus; reeeptaeulis anguste eonoideis 0.9— 1. 1 mm
longis 2-2.4 plo longioribus quam latioribus; et paleis pis-
lillalis longissimis 2. 5-3.2 (ad invicem 3.3-4.0) mm lon¬
gis.
Caulescent annuals 3—9 cm high from slender
taproots, plants densely and tightly silvery-seri¬
ceous throughout; stems single, erect, or 2 to 5 from
base and strongly ascending, usually branched
throughout, rarely simple, slender; branches sym-
podial or sometimes appearing monopodial by re¬
duction or failure of development. Leaves basal and
eauline, gradually enlarged and congested up
stems, alternate, simple, sessile or broadly and in¬
distinctly petiolate, up to I I X 3 mm; bases ± at¬
tenuate, herbaceous or becoming thickened and in¬
durate among eapitula, 1 -veined; blades oblanceolate
to spatulate or obovate, ± planar to undulate, pli¬
ant, entire, uniformly sericeous, primary vein 1,
subordinate venation obscure, apices usually ob¬
tuse to narrow ly rounded; capitular leaves generally
broadest, often appearing whorled. usually mixed
with, subtending, and clearly exserted from be¬
tween individual eapitula in glomerules, abruptly
differentiated from phyllaries and paleae, longest
2— I OX eapitula lengths, bases erect, carinate,
thickened, indurate, forming rigid secondary invo¬
lucres. ( iapitulescences I to 5, cymiform, sympodial.
strictly pseudo-polytomous or dichasiform, or some¬
times appearing monopodial by reduction or failure
of branch development, each consisting of 3 to 20+
eapitula; branches all strictly subtending glomer¬
ules, arising among capitular leaves, strongly as¬
cending to erect, equal or often unequal with some
scarcely or not at all elongating; eapitula disciform,
fully paleate, sessile, borne singly or rarely in
dense glomerules of 2 or 3, all terminal at branch
forks and apices, ± erect, cylindric to ± ellipsoid,
3. 5- 4.5 X 1. 5-2.0 mm; involucres of (2 to)4 to 0
reduced, empty phyllaries similar to and seemingly
derived from paleae, otherwise functionally simu¬
lated by paleae and capitular leaves; receptacles
conic, 0.9-1. 1 mm long, ± 2X as long as wide,
glabrous; paleae about 18 to 30 in 4 to 6 spirally
ranked series, erect to ascending, free proximally,
distal ly all or at least the inner coherent by tangled
hairs, falling as a unit (sometimes tardily so or out¬
ermost series persistent), veins parallel, obscure,
midveins not reaching apices, apices densely lan-
uginose to sericeous; pistillate paleae in 3 to 5 se¬
ries, imbricate, mostly open, ± planar to concave,
gently incurved, oblanceolate to oblong, longest
2.5- 3.2 mm long, margins of pistillate paleae con¬
tinuous with and not abruptly differentiated from
bodies, not winged, gradually thinning, ± scarious,
shiny, translucent, not conspicuously striate, entire,
abaxially lanuginose to glabrous, bodies yellowish,
not gibbous, somewhat thickened, papery to carti¬
laginous, abaxial surfaces ± rounded transversely,
glabrous proximally, apices incurved (with styles)
over staminate florets at anthesis, at maturity erect,
— planar to concave; staminate paleae about 3,
slightly enlarged, open distal ly, becoming ± closed
proximally, concave medially, ± spatulate, mostly
1.5- 2.5 mm long, slightly surpassing pistillate pa¬
leae at maturity, papery to somewhat cartilaginous,
greenish to brownish on both surfaces, abaxially ±
lanuginose to tips, adaxially ± glabrous, bases
erect, acute to acuminate, apices somewhat spread¬
ing at maturity, forming obconic cavities in the
mass of hairs, entire, ± herbaceous, obtuse to
rounded, neither spinose nor uncinate. Pistillate
florets 13 to 27+ in 3 to 5 spirally ranked series,
± evenly distributed on receptacle, pistillate corol¬
la* whitish, filiform, limb absent, styles strongly ex¬
serted but surpassed by subtending paleae; stami¬
nate florets about 3 from receptacle centers, each
subtended by a staminate palea, ovaries partly de¬
veloped, abortive, 0.4— 0.6 mm long, staminate co¬
rollas ± concealed by pistillate paleae at anthesis,
later exposed bv spreading staminate paleae, radi¬
ally symmetric, 1.4— 2.0 mm long, funnelform, gla¬
brous, lobes usually 4, yellowish brown, debate,
blunt, equal, stamens included, styles not or
scarcely exserted. Cypselae falling free, light to
dark brown, without black band, ± obovoid, ob¬
tuse, somewhat angular, ± straight, not gibbous,
obcompressed, mostly 0.9-1. 2 X 0.5-0.7 mm, gla-
Volume 14, Number 4
2004
Morefield
North American Asteraceae
471
l>rous, minutely papillate, dull to ± shiny, corolla
scars apical; pappi none.
Etymology. The new variety is named with
gratitude for the late Rupert C. Barneby (1911-
2000), who immediately recognized the unusual
characteristics of this plant as few before him had,
who supplied the type gathering (and the specimens
of Micropsis dasycarpa discussed elsewhere in this
paper), who extended characteristic courtesy and
assistance during and beyond my years as a stu¬
dent. and whose eloquent and voluminous lifetime
contributions to the systematics and taxonomy of
Fabaceae remain a priceless resource and inspi¬
ration to students of botany the world over. Barneby
rabbit tobacco is here recommended as a common
name for this variety, to be consistent with other
vernacular names used in the genus.
As is evident from the paratypes cited below,
specimens of Diaperia prolifera var. barnebyi have
been known since 1852, but the geographic and
ecologic significance of the variation was not pre¬
viously recognized. Based on all the type material,
Diaperia prolifera var. barnebyi occurs from west
Texas to southeast New Mexico and extreme south¬
west Oklahoma, a range from which variety proli-
fera appears largely absent and which comprises
the southwest corner of the species’ range. The va¬
riety barnebyi clearly is the xeromorphic phase of
the species and does not appear to be particularly
weedy. Both varieties approach the Rio Grande just
north of Coahuila, Mexico, and may eventually be
documented from that country.
Diaperia prolifera var. barnebyi is found on open,
dry, shallow, rocky or gravelly soils often derived
from limestone or gypsum, sometimes in drainages,
disturbed areas, or near other sources of extra mois¬
ture, at elevations between about 500 and 1500 m.
Based on phenology apparent in the type material,
plants of the new variety undergo their reproductive
cycles between about mid April and mid June.
The relatively large geographic range of variety
barnebyi ought to render it fairly invulnerable from
a conservation standpoint. There are. however, few
recent collections among the type and paratype
specimens cited herein, which comprise all speci¬
mens currently known to me. Attempts should be
made to relocate as many of the historical sites as
possible, and to find additional, previously undoc¬
umented sites, before the variety is assumed to be
secure from extinction.
Where the ranges of variety barnebyi and variety pro¬
lifera overlap in central Texas anti Oklahoma, specimens
have been collected that are intermediate in characteris¬
tics between the two types. Specimens I have examined
that are of uncertain varietal placement, and that are ex¬
plicitly not considered paratypes (which are listed farther
below), include: U.S.A. Oklahoma: Woods Co., near
Alva, prairie hillside in old road, 10 May 1914, G. W.
Stevens 3053 (US); 16 mi. W & 3 mi. S of Alva, top of
hill, short sparse grass, 17 May 1972, S. Bellah 71 (BRY).
Texas: Bandera Co., Mystic Canyon, Dry' Canyon Creek
0.5 mi. E of Real Co., limestone plateau, 2200 ft., 4 June
1975, ./. Smith 576 (1,1,); Bell Co., 4 mi. NE of Salado,
calcareous gravel, oak-juniper woodland, 12 May 1953,
//. D. Ripley & R. C. Barneby 11059 (NY); McLennan
Co., 3 mi. S of McGregor, 26 May 1946. C. /.. York 46105
(TEX); Menard Co.. Menard, 26 Apr. 1933, H. Mahony
s.n. (TEX); Randall Co., I’alo Duro Canyon, 15 June 1929,
B. C. Tharp 6495 (LL, TEX); Terrell Co., Blackstone
Ranch, 13 mi. S of Sheffield, cedar-sotol mesa top, 2750
ft., 12 June 1949, G. L Webster 244 (MICH, TEX).
Typical material of the two varieties may be separated
as follows:
Key to the Varieties ok Diaperia prolifera
la. Plants loosely lanuginose, grayish to greenish;
capitula 4 to 40+ in largest glomerules, recep¬
tacles wider than long; capitular leaves proxi-
mally pliant to ± rigid, not or scarcely carinate,
often ± spreading, loosely involucrate; capitu-
lescence branches mostly spreading to ascend¬
ing; longest paleae 3. 3-4.0 mm long .
. Diaperia prolifera var. prolifera
lb. Plants tightly sericeous, silvery; capitula borne
singly or rarely in glomerules of 2 or 3, recep¬
tacles ± 2X as long as wide; capitular leaves
proximally thickened, indurate, carinate, erect,
forming rigid secondary involucres; capitulesc-
ence branches strongly ascending to erect; lon¬
gest paleae 2. 5-3. 2 mm long .
. Diaperia prolifera var. barnebyi
Paratypes. U.S.A. New Mexico: Eddy Co., Guada¬
lupe Mountains, near mouth of Dark Canyon, rocky N
slope, 23 Apr. 1932, H. Wilkens 1591 (I’ll); Lincoln Co.,
3 mi. E of Picacho, draw in limestone hills, caliche gravel,
4800 ft., 2 May 1986. R. C. Barneby 18068 (NMC. NY).
Oklahoma: Harmon Co., 5.5 mi. S of Hollis, shallow
land, overgrazed pasture with Prosopis, 13 May 1950, II.
T. Waterfall 9414 (ARIZ). Texas: stony hills of the Pecos,
May 1852, C. Wright 1406 (BM, GH, LE, NY. PH, US);
Hardeman Co., 2 mi. W of Acme, gypsum outcrop, 1 June
1957, I). S. Correll 16528 (LL); Lubbock Co., Lubbock
sewer farm, 3 June 1930, I). Demaree 7752 (GH, LE. MO,
MONTU, WIS); Terrell Co., Langtry to Dryden, 14 June
1931, B. C. Tharp 8814 (TEX).
Logfia Survives Oglifa : A Tale oe Two
Anagrams
Since conservation of the genus Filago L. with
F. pyramidata L. as the type (Buchheim & Wagenitz,
1964; Greuler et al., 2000, Appendix IIIA.E.3)
there have been, among European workers, two di¬
vergent approaches to the circumscription of filago
and its relatives. The first approach, exemplified by
Wagenitz (1969), has been to recognize a large and
heterogeneous Filago with several subgenera and
472
Novon
sections, generally including Evax (and bv impli¬
cation Diaperia and Hesperevax of North America,
also included in Evax until more recently). Justifi¬
cation of this approach has been based on the ap¬
parent recombination, in “intermediate” species
groups, of various character states distinguishing
the type species of Filago and Evax , and also by
the existence of occasional natural hybrids among
species of the various subgenera and sections (Wag-
enitz, 1969).
II applied to North American taxa. this “Wag-
enitz approach” would immediately require inclu¬
sion of Stylocline, a close relative of Filago subg.
Oglifa (Cassini) Grenier (Morefield, 1992a), in a yet
more heterogeneous Filago. By logical extension.
Micropus would fall next, and thereafter all the re¬
maining genera of Asteraceae subtribe Lilagininae
would have to be lumped.
I prefer the other approach, first championed by
Cassini (1819, 1822) and more recently by Holub
(1975, 1976, 1998), that recognizes the various re¬
combinant species groups as smaller, more homo¬
geneous genera. The various combinations of typi¬
cal Filago and Evax character states in these
groups appear to identify natural and easily rec¬
ognizable genera (as l ilagininae go) that are either
monophyletic or minimally paraphyletic and that
often exhibit unique synapomorphies (Morefield,
1992a). This approach to classifying lilagininae
has been applied to the North American taxa bv
resurrection and segregation of the genera Ancistro-
carphus (Morefield, in Hickman, 1993), Diaperia
(this paper, above), and Hesperevax (Morefield,
1992c). Holub (1998: 107) completed the process
by transferring to htgfia the three North American
species traditionally placed in Filago (F. arizonica
A. (nay, F californica Nuttall, and F. depressa A.
(hay).
I iigree with llolub's (1975, 1976, 1998) circum¬
scription of htgfia to include Oglifa (Cassini) Cas¬
sini. When the International Code of Botanical No¬
menclature (ICBN; Greuter el al.. 2000) is applied,
however, priority of Logfia over Oglifa is ambiguous
and may need to be established via conservation in
order to preserve current usage. When Cassiid
(1819) first split Fdago into smaller and more nat¬
ural units, he named three of them using the ana¬
grams Gifola , Ixtgfia, and Oglifa. Kaeh of these taxa
was clearly accepted by Cassini at the time (ICBN,
Art. 34. 1 ) and was provided with a description and
a clear indication of the type species. Each except
htgfia was also provided with a clear indication of
its rank, Gifola as a genus, and Oglifa as a sub¬
genus. The type of Gifola Cassini belongs to the
same genus as the type now conserved for Filago.
All earlier workers have interpreted Cassini’s
(1819: 143-144) subgenus Oglifa to have been as¬
sociated with the genus Filago, by the title and
context of his article and by his designation of Fi¬
lago arvensis E. as type. A careful reading of Cas¬
sini's article, however, shows that he explicitly dif¬
ferentiated subgenus Oglifa from a portion of
Gnaphalium (as translated from the French, from
“many true Gnaphalium, such as G. luteo-album,
sylvaticum and uliginosum ”), not from Filago, and
that he explicitly limited membership in Filago to
a single species, F. pygmaea L. Cassini (1819:
143). therefore, definitely associated his new name
with Gnaphalium (ICBN Art. 33.1) and validated
Gnaphalium L. subg. Oglifa Cassini, a name not
heretofore recognized. Cassini (1822: 564) later
validated Oglifa at the generic rank.
Validation of htgfia in 1819 was questionable,
and may not have occurred until 1822. Cassini
(1819: 143) began his protologue of htgfia by list¬
ing the two Fdago species he included therein,
stating that those species belonged in a genus other
than Gifola Cassini (the latter having been de¬
scribed earlier in the same publication), and listing
the differential traits. He then stated (as translated
from the French), "I thus join together these two
species in a particular genus or subgenus, of which
F. gallica must be regarded as the tvpe . . .,” after
which he went on to name and formally describe
the taxon htgfia.
Even though Cassini (1819: 143) said that the
two species of his new taxon “differ genetically
from Gifola," it does not necessarily follow that he
was ranking htgfia itself as a genus, only that htg¬
fia was part (or all) of a genus other than Gifola.
By designating htgfia as a “genus or subgenus,” it
can be argued that Cassini either (I) published al¬
ternate names in 1819 (as allowed bv ICBN, Art.
34.2). or (2) expressed taxonomic doubt about the
placement of htgfia (as allowed bv ICBN. Art.
34. 1 ). resulting in no “clear indication of its rank"
and leaving it valid but unranked in 1819 (as al¬
lowed by ICBN, Art. 35.3). In the first case, htgfia
was valid at generic rank in 1819 and has priority
over Oglifa. ( htgfia was not valid at subgeneric
rank in 1819, however, because it was not definitely
associated with the name of a genus, both Gifola
and Filago having been explicitly excluded.) In the
second case, htgfia was validated at generic rank
by Cassini (1822: 564) simultaneously with Oglifa,
and then Lessing (1832: 333) established priority
of Oglifa over htgfia in accordance w ith ICBN, Art.
11.5.
\\ Idle I believe that the second case above (ap¬
plication of Arts. 34.1 and 35.3) represents the
Volume 14, Number 4
2004
Morefield
North American Asteraceae
473
more logically rigorous interpretation of Cassini’s
words, Holul) (1975, 1976, 1998; but not Clirtek &
Holub, 1963) and the various nomenelatural indi¬
ces have followed the first interpretation instead,
and it would now be nomenelaturally disadvanta¬
geous to use Oglifa in place of Logfia. Therefore,
pending a proposal to conserve Logfia over Oglifa
(in prep.), I agree that the consequences of the
rules are doubtful in the case of logfia and that
established custom should be followed (1CBN, Pre¬
amble 10). The nomenclature of logfia is summa¬
rized as follows:
Logfia Cassini, Bull. Sci. Soc. Pliilom. Paris 1819:
143. 1819 | as “genre ou sous-genre”], Cassini,
in F. Cuvier, Diet. Sci. Nat., ed. 2, 23: 564.
1822. Filago I.. subg. logfia (Cassini) Grenier,
FI. Jurass. (2): 431. 1869. Filago L. sect, log¬
fia (Cassini) Boissier, FI. Orient. 3: 248. 1875.
TYPE: Filago gallica F.
Oglifa (Cassini) Cassini, in F. Cuvier, Diet. Sci. Nat., ed.
2, 23: 564. 1822. Gnaphalium L. subg. Oglifa Cas¬
sini. Bull. Sci. Soc. Philom. Paris 1819: 143. 1819.
Filago I.. sect. Oglifa (Cassini) DC., Prodr. 6: 248.
1838. Filago L. subg. Oglifa (Cassini) Grenier, FI.
Jurass. (2): 430. 1869. TYPE: Filago arvensis F.
Xerotium Bluff & Fingerhuth, Comp. FI. German, (sect. 1)
2: 343. 1825. TYPE: Gnaphalium minimum J. E.
Smith.
Species of logfia are indigenous to Europe, cen¬
tral Asia, northern Africa, and southwestern North
America, anil have been widely introduced outside
those areas. Combinations under Logfia were vali¬
dated by Holub (1998) for the three North Ameri¬
can species listed earlier. When removed from Fi¬
lago, however, an earlier specific epithet becomes
available for the species long known as Filago cal-
ifornica, and must be used:
Logfia filaginoides (Hooker & Arnott) Morefield,
comb. nov. Basionym: Gnaphalium filaginoides
Hooker & Arnott, Bot. Beechey. Voy. (8): 359.
1839. TYPE: U.S.A. “California”: s.d., D.
Douglas s.n. (lectotype, designated here, K
[HI). Hook]).
Filago californica Nuttall. Frans. Amer. Philos. Soc. ser.
2, 7: 405. 1841. Syn. nov. Non Filago filaginoides
(Karelin & Kirilow) Wagenitz, Willdenowia 5: 417.
1969. Oglifa californica (Nuttall) Bydberg, FI.
Rocky Mts. 911. 1917. logfia californica (Nuttall)
Holub, Preslia 70: 107. 1998. TYPE: U.S.A. Cali¬
fornia: “near Santa Barbara,” Apr. 1836, T. Nuttall
s.n. (holotype, BM |Hb. Nuttall |; isotype, PH 1383).
Filago californica Nuttall var. tomentosa Nuttall. Trans.
Amer. Philos. Soc. ser. 2, 7: 406. 1811. Syn. nov.
TYPE: U.S.A. California: Santa Luis Obispo Co.,
USFS trailhead area on Hwy. 166 NE of Tepusquet
Canyon Rd., 9 Apr. 1998. I). Keil 26673-A (neotype,
designated here, OBI 055308).
Hooker and Arnott (1839: 359) validated Gna¬
phalium filaginoides as “2. G.? filaginoides" (ICBN,
Art. 34.1) but designated no type. Specimens were
found on sheets at BM, GH, and K. each labeled
with some abbreviation or permutation of Gnaphal¬
ium filaginoides. A Douglas specimen on a sheet
stamped “Herbarium Hookerianum 1867” at K is
selected here as the lectotype because it is the only
candidate definitely seen by one of the authors of
the taxon, and was gathered by a frequent collector
of Hooker’s and Arnott’s new taxa. There is also a
pencil drawing of the species on the sheet. Two
other specimens on the same sheet (Nuttall s.n. and
Coulter 352 ) also represent the same species. The
specimens labeled Gnaphalium filaginoides on the
sheets at BM [Hb. Nuttall] and G1I were not pro¬
vided with collector names, but are very similar to
the lectotype specimen, and may represent isolec-
totypes.
Filago parvula Torrey & A. Gray, FI. N. Amer.
2 (3): 432. 1843, was a superfluous name for Gna¬
phalium filaginoides. The indexed name “ Gnaphal¬
ium filaginoides Bojer ex DC.” (de Candolle, 1837
[1838]: 219) was not accepted by its author, who
validated Stenocline filaginoides DC. instead. It is
therefore invalid and should be removed from all
plant name indices. Logfia filaginoides is most
abundant in California and adjacent northwestern
Mexico, extending eastward across a narrowing tri¬
angular area to extreme western Texas.
Other specimens located at GH and K |Hb.
Hook.] likely represent additional isotypes of Fi¬
lago californica, but too lew label data accompany
them to permit certainty. No original material, type
or otherwise, could be located for Filago californica
var. tomentosa, nor was any designated in the pro-
tologue, necessitating selection of a neotype. The
protologue diagnosed the variety as having “leaves
crowded and tomentose, flower clusters approxi¬
mating in spikes,” and indicated that both it and
variety californica were found “near St. Barbara,
Upper California." The neotype specimen selected
above matches this diagnosis exactly, and is the
“nearest” to Santa Barbara of any similar speci¬
mens I have seen. It was collected just across the
Santa Barbara/San Luis Obispo County line near
the Cuyama River (D. J. Keil, pers. comm.. 19 Aug.
2004). As neotypified, Filago californica var. to¬
mentosa appears to represent an occasional ecotype
of coastal and chaparral areas, and does not merit
taxonomic recognition.
474
No von
Micropsis Arrives
Micropsis DC. consists of five species native to
temperate South America and found variously in
moist or dry, often sandy or alkaline soils from cen¬
tral Chile to Paraguay and southern Brazil, south
to the Juan Fernandez archipelago (Bustamante,
1979; Cabrera, 1963). Its species are unique among
Filag ininae for their densely long-hairy eypselae
and for the saccate, nearly closed, and often lac¬
erate bisexual paleae, and may be only conver-
gently related to the rest of the subtribe. Phyloge¬
netic analysis of morphologic characters (Morefield,
1992a) placed the genus closest to Stuartina muel-
leri Sonder of Australia.
In 1988 the late Rupert C. Barneby sent to me
for identification a specimen of Asteraceae from the
central gulf coast area of Texas. This turned out to
be Micropsis dasycarpa (Grisebach) Beauverd, and
the name and typification are here newly reported
for the Flora of North America project:
Micropsis dasycarpa (Grisebach) Beauverd, Bull.
Soc. Bot. Genfcve ser. 2, 5: 224. 1913. Filago
dasycarpa Grisebach, Abh. Konigl. Ges. Wiss.
Gottingen 24: 185. 1879. TYPE: Uruguay.
Concepcidn del Uruguay, Oct. 1877, P. Lorentz
IIBI (holotype, GOET not seen; isotypes, G-
BOIS not seen. NY).
Of his North American collection (cited below),
Barneby (in lilt. 9 Nov. 1988) wrote:
“The habitat in Texas is in low swampy ground, and
one of the associated herbs is Mimosa strigillosa T.
& G. The latter has a remarkable bicentric dis¬
persal , around the Gulf of Mexico in US, and in
warm-temperate South America chiefly in the Pa-
rand basin. There is no obvious reason to think that
the mimosa is not native in both hemispheres —
there are a number of xerophytic species with about
the same range — and the possibility arises that Mi¬
cropsis is not just a casual introduction but an over¬
looked native in Texas. The microhabitat where I
found it was certainly not pristine Gulf lowland
prairie, but that scarcely exists nowadays.”
I agree with Barneby that Micropsis dasycarpa
could be native to Texas. Although its recent dis¬
covery in a part of the world relatively well-ex¬
plored by botanists, and the apparent lack of fur¬
ther collections from the area, would argue against
that interpretation for now. the species will be given
benefit of the doubt and treated as introduced for
the Flora of North America project.
North American .specimen examined. U.S.A. Texas:
Victoria Co., N of Inez, colonial in drying mud at edge of
pool, in Gulf Coast Prairie, 18 May 1987, R. C. Barneby
18201 (NY 622020, KSA 475037).
South American specimens examined. PARAGUAY.
Villa Florida, 2 Oct. 1892, 0. Kuntze s.n. (NY); Cerros de
Tobaty, Sep. 1900, E. Bossier 6 102 (NY|2|). URUGUAY.
Departamento de Colonia, Riachuelo, 1 Nov. 1960, A. L
Cabrera 18201 (RSA).
Acknowledgments. I am grateful to David J.
Keil (OBI), Patricia K. Holmgren (NY), Steve Boyd
(RSA), and the late Rupert C. Barneby (NY) for
generously making specimens and other informa¬
tion available to me. Linda DeVito and Susan Fras¬
er of the New' York Botanical Gardens LuEsther T.
Mertz Library, Linda Oestry of the Missouri Botan¬
ical Garden Library, and Irene Holiman and Gary
Wallace of the Rancho Santa Ana Botanic Garden
were most accommodating in cheerfully and rapidly
supplying photocopied pages of old and/or rare
works ant) other information critical to the comple¬
tion of this study. The comments and suggestions
of Eric B. Peterson, Dieter Wilken (SBBG), Arnold
Tiehm (RENO), John Pruski and Victoria Hollowed
(MO), and Vicki Funk substantially improved ear¬
lier versions of this paper. The advice and opinions
of Fred Barrie, Kanchi Gandhi, Dan Nicolson, and
John Pruski on the nomenclatural status of Logfia
and Oglifa, anti of John Strother on matters of ter¬
minology and description, were also greatly appre¬
ciated. Portions of the work leading to this paper
were supported by the Rancho Santa Ana Botanic
Garden, by the National Science Foundation (Grad¬
uate Fellowship Program, and Dissertation Im¬
provement Grant BSR-9000893), and by the State
of Nevada. I thank the curators of the herbaria cited
above, particularly Steve Junak (SBBG), for loans
(often extended) of materials in their care, and for
their patience. And finally I am grateful for my fam¬
ily, Leslie, Cameron, and Shaelin, who tolerated the
extra hours required to complete this work.
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Buchheirn, G. & G. Wagenitz. 1964. Proposal to conserve
the generic name Filago L. (8969, Compositae). Reg-
num Veg. 34: 61-62.
Bustamante, L. E. N. 1979. Flora de la Cuenca de San¬
tiago de Chile, Vol. 3. Univ. de Chile, Santiago.
Cabrera, A. L. 1963. Flora de la Provincia de Buenos
Aires, Parte VI. Compuestas. Colecci6n Cientffica del
I.N.T.A., Buenos Aires.
Candolle, A. P. de. 1836. Prodromus Systematis Naturalis
Regni Vegetabilis, Vol. 5. Treuttel and Wiirtz, Paris.
- . 1837 [1838]. Prodromus Systematis Naturalis
Regni Vegetabilis, Vol. 6. Treuttel and Wiirtz. Paris.
Cassini, H. 1819. Examen analytique du genre Filago de
Linne. Bull. Sci. Soc. Philom. Paris 1819: 141-144.
- . 1822. lnul^es, Inuleae. Pp. 559—573 in F. Cuvier
(editor), Dictionnaire des Sciences Naturelles, 2nd ed.,
Vol. 23. Le Normant, Paris, and F. G. Levrault, Stras¬
bourg.
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2004
Morefield
North American Asteraceae
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Chrtek, J. & J. Holub. 1963. Poznamky k taxonomii a
nomenklatura rodu Evax Gaertn. a Eilago L. Preslia 35:
1-17.
Felger, R. S. 20(H). Flora of the (Iran Desierto and Rfo
Colorado of Northwestern Mexico. Univ. Arizona Press,
Tucson.
Flora of North America Editorial Committee (editors).
1993 + . Flora of North America North of Mexico. 8 +
vols. Oxford Univ. Press, New York and Oxford.
Gray, A. 1884. Synoptical Flora of North America, Vol. 1,
Part 2. New York.
Greuter, W., J. McNeill, F. R. Rarrie, 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.
Hickman, J. C. (editor). 1993. The Jepson Manual: Higher
Plants of California. Univ. California Press, Berkeley.
Holub, J. 1975. Ifloga, Eilago, Logfia, Evax, Bombycilae-
na, Cymbolaena, Micropus. Pp. 100-1 16 in P. H. Davis
(editor). Flora of Turkey and the East Aegean Islands,
Vol. 5. Univ. Edinburgh Press, Scotland.
- . 1976. Eilago, Ifloga, Logfia, Evax, Bombycilae-
na, Micropus. Pp. 121-126 in T. G. Tutin et al. (editors),
Flora Europaea, Vol. 4. Cambridge Univ. Press, Cam¬
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- . 1998. Reclassifications and new names in vas¬
cular plants 1. Preslia 70: 97—122.
Hooker, W. J. & G. A. W. Arnott. 1839. The Botany of
Captain Beechey’s Voyage, Part 8. London.
Keil, D. J. & D. J. Pinkava. 1976. Chromosome counts
and taxonomic notes for Compositae from the United
States and Mexico. Airier. J. But. 63: 1393-1403.
Lessing, C. E. 1832. Synopsis Generum Compositarum
Earumque Dispositionis Novae Tentamen Monographiis
Multarum Capensium lnterjeetis. Duneker and Hum-
blot, Berlin.
Morefield, J. D. 1992a. Evolution and Systematics of Styl-
ocline (Asteraceae: Inuleae). Ph.D. Dissertation, Clare¬
mont Graduate University, Claremont, California. Pub¬
lished by UMI Dissertation Information Service, order
number 9238862, document number 31 1 188, Ann Ar¬
bor, Michigan.
- . 1992b. Three new species of Stylocline (Astera¬
ceae: Inuleae) from California and the Mojave Desert.
Madrono 39: 114—130.
- . 1992c. Resurrection and revision of Hesperevax
(Asteraceae: Inuleae). Syst. But. 17: 293—310.
- . 1992d. Notes on the status of Psilocarphus berteri
(Asteraceae: Inuleae). Madrono 39: 155—157.
- & R. Eelger. 20re el nervio central y los secundarios, apice acu-
minado, base cuneada a ligeramente asimetriea,
con 5—1 1 pares de nervios secundarios, 2—1 nervios
saliendo desde la base de la lamina, enves usual-
478
Novon
mente con domacios en las axilas del nervio central
y en secondaries, glandulas pelueidas ausentes; es-
tipulas 0.75—1.5 mm, pronto deciduas. Inflorescen-
cias son cimas terminales o axilares, 2—5 cm de
largo, con 2—5 llores, ejes estrellado-puberulentos;
pedunculos 0.3— 0.8 cm, 2-bracteados cerca de la
base, braeteas 0.75—1.25 mm. Flores con pedicelos
7—14 mm, articulados, con bract^olas 0.5—0.75
mm, pronto deciduas; sepalos 4, 9—1 1 mm, lanceo-
lados, denso y estrellado-puberulentos en la parte
externa, esparcido y estrellado-puberulentos inter-
namente; petalos 4, i— 8 mm, lanceolados, blancos,
glabros; falanges 4, opuestos a los petalos, cada uno
con 10 estambres, parte libre de los filamentos
0.25—0.75 nun, anteras 2.25—3.5 mm incluyendo el
mucro, oblongo-triangular, desiguales, mucro 0.25-
0.6 mm; ovario con 5 fuertes costillas, estilo ca. 6.5
mm, estrellado-puberulento en la mitad basal, gla-
bro en la mitad distal. Frutos 2. 5-4. 5 X 2.5-4 cm,
globosos a ovoides, 5-locular, con 5 fuertes costi¬
llas, costillas formadas a lo largo del punto de
union de las valvas, valvas lenosas. pared 4—6 mm
de grosor; semillas ca. 5 por Ideulo, fertiles o es-
feriles, fertiles ca. 5 mm de diametro, subglobosas,
cubiertas ca. lA de su tamano por el arilo.
Fenologia. Flores en julio; frutos entre enero a
marzo y en junio.
Distribut ion. End^mica de Costa Rica, cono-
cida lie los bosques nubosos de la Cordillera de
Talamanca en la vertienle Pacifica, entre 1600-
1850 m.
Mortoniodendron abelianum se reeonoce por sus
grandes frutos prominentemente acostillados, en
oeasiones costillas pareciendo alas, con paredes
valvares gruesas y endurecidas, estambres con la
parte libre de los filamentos muy eorta (igual o me-
nor a 0.75 mm) y por presentar loculos de los frutos
con ea. 5 semillas, entre fertiles y esferiles. Muy
similar a M. moralesii sin embargo, este ultimo tie—
ne tall itos y peciolos estrellailo-tomentosos (vs. es-
trellado puberulentos) y los frutos son nada a leve
acostillados (vs. prominente acostillados). Tambien
es semejante a M. anisophyllum (Standley) Standley
Steyermark, pero se les puede distinguir debido
a (pie este ultimo presenta frutos mas pequenos
(entre 1—2 X 1—2 cm), la pared del exocarpo es
mas delgada (luista 0.5 mm) y su rango de distri-
bueidn es distinto (1—850 m). Mortoniodendron abe¬
lianum ha sido erroneamente identificado bajo el
nombre de M. pentagonum (Donnell Smith) Miran¬
da. una especie restringida a Guatemala (Dorr,
2001), pero esta especie se separa debido a que
presenta hojas plinervadas, en la cual el primer par
de venas sale hasta 2 mm sobre la base, sus hojas
carecen de domacios, sus laminas foliares tienen
glandulas pelueidas, los pedicelos no son articu¬
lados, los estambres son fibres, biseriados, no en
falanges, los pedicelos (2-3 cm), sepalos (15 mm)
y anteras (4.5— 6.5 mm) son mds largos, finalmente
debido a que los loculos de los frutos tienen uni-
camente una semilla, la cual esta cubierta total-
mente por el arilo.
FI epiteto esta dedicado a Abel Rodriguez, mi
padre, a quien agradezco su apoyo incondicional
en todo momento de mi vida.
Pardtipos. COSTA RICA. Puntarenas: Coto Ifius,
Parque Intern. I, a Amistad, cordillera de Talamanca, Est.
Altamira, Sendero Fos Gigantes, Angulo 165 (CR, INB,
MO); Est. Biol. I.as Alturas, El Tajo, Aguilar 97 (CR, INB,
MO); Est. Pittier, NE de la Estacion, Acosta 2330 (CB,
INB, MO); Sendero a Eila Pittier, Gamboa 2 (INB); Zona
Protectora I.as Tablas, Sendero a Quebrada Gemela, Chin¬
chilla 22 (CR, INB, MO); Sitio Las Tablas, Alfaro 2015
(INB, CR).
2. Mortionodcndron apetalum Al. Rodriguez,
sp. nov. I IPO; Costa Rica. Alajuela: Laguna
Pocosol, Rio Penas Blancas, bosque Kterno de
los Ninos, 10°2rN. 84°40'0, 840 m. 27 abr.
1992, B. Haber 1 1 161 (holotipo, INR; isdtipos,
CR. MO). Figura 2.
Haec species inter congeneros foliis glandulis pelluci-
dis praeditis, flore apetaln atque Iructus grandis valvarurn
lignosarum parietibus crassis recognoscitur.
Arbol 25—50 rn; ramitas teretes, estrellado-pu-
berulentas. Hojas con peciolos 9-18 mm, estrella¬
do-puberulentos; lamina 9-19 X 4.5-9 cm, lan-
ceolada a elfptico-lanceolada, coriacea,
estrellado-puberulenta, principalmente en ramitas
jovenes, glabrescente con el tiempo, apice acumi-
nado, base redondeada a leve oblicua, con 5—12
pares de nervios secundarios, 2—4 nervios saliendo
desde la base de la lamina, enves usualmente con
domacios en las axilas del nervio central y en se¬
cundarios, con glandulas pelueidas; estipulas no
observadas, pronto deciduas. Inflorescencias son
panfculas cimosas y de ramificacidn dieotbmiea.
terminales o axilares, 2.5-9 cm, con 4—20 Mores,
pedunculos 1—3.5 cm, ejes estrellado-puberulentos,
a menudo bracteados, braeteas no observadas,
pronto deciduas. Flores con pedicelos 5—12 mm.
articulados, en oeasiones bracteolados, bracteolas
no observadas, pronto deciduas; sepalos 5, 7-9
mm, lanceolados, estrellado-puberulentos, crema;
petalos ausentes; falanges 5, opuestos a los sepalos,
cada uno con 10 estambres, parte fibre de los fi¬
lamentos 2—3 mm, anteras 0.75—2.5 nun. incluyen¬
do mucro, oblongas, subiguales, mucro obsoleto o
no sobrepasando a 0.12 mm; ovario con 5 leves
Volume 14, Number 4
2004
Rodriguez
Mortoniodendron (Tiliaceae)
479
I'igura 2. Mortionodendron apetalum Al. Rodriguez ( Haber 1 1 161). —A. Ramita con hojas e inflorescencia. — R.
Domacios. — C. Flor. — D. Pistilo. — E. S/palo y falange. — F. Fruto. — G. Fruto en seccirtn transversal. — H. Semilla
subtendida por el arilo.
(‘ostillas, estilo ca. 4 mm, estrellado-puberulento en
la mitad basal, glabro en la mitad distal. Frutos
1.5— 2.5 X 2—3 cm, globosos a ovoides, 5-locular.
eon 5 leves costillas, costillas formadas a lo largo
del punto de union de las valvas, valvas lenosas,
pared 3—5 mm de grosor; semillas una por loeulo,
fertiles o esteriles, fertiles 8-10 X 7-8 mm. ovoi¬
des, envueltas completamente [tor el arilo.
480
Novon
Fenologia. Flores entre marzo y altril; frutos
entre julio y agosto.
Distribution. Endemiea de Costa Rica, cono-
cida en bosques humedos de la Cordillera de Gua-
naeaste. Cordillera de Tilaran y Cordillera Central
en las vertientes Pacffica y Atlantiea. entre 600—
1000 rn.
Mortoniodendron apetalum se reeonoce por la
comhinaeibn de los siguientes carac teres: hojas con
glandnlas pelucidas, flores apetaladas y frutos con
valvas lenosas y de pared gruesa; tambien, es im-
portante mencionar como caracterfsticas sobresa-
lientes que los individuos alcanzan gran tamafio,
sus semillas son grandes y ovoides, cubiertas en su
totalidad por el arilo, I < >s peciolos son alargados,
las hojas lanceoladas y las anteras muestran el niu-
cro apical obsolete a pobremente desarrollado.
Mortoniodendron apetalum es muy relacionado a M.
ruizii Miranda y a M. palaciosii Miranda, ambas
especies de Mexico, debido a que todas correspon-
den a grandes arboles. la lamina de sus hojas posee
glandnlas pelucidas, sus flores son apetaladas, sus
frutos son grandes, lenosos y de gruesas valvas. Sin
embargo, se les distingue ya que estas ultimas po-
seen flores con 4 (ocasionalmente 3) sepalos y fru¬
tos con igual numero de valvas (vs. 5 sepalos y 5
valvas); ademas, M. ruizii es mas pubescente en
partes vegetativas, presenta peciolos mas pequenos,
entre 6-10 mm (vs. 9-18 mm) y pedunculos igual-
mente imis pequenos, entre 0.3— 0.7 cm (vs. 0.1-
3.5 cm). Mortoniodendron palaciosii tiene hojas
mds pequenas, entre 7.5-14.5 X 3. 5-7.2 cm (vs.
9-19 X 4.5—9 cm), base a menudo fuerte oblicua
(vs. redondeada a leve oblicua) y frutos hasta 1.8
cm de ancho (vs. 3 cm).
Pardtipos. COSTA RICA. Alajuela: Res. Biol. Mon¬
teverdi'. Rfo IVnas Blancas, Haber 8383 (CR, INB. MO);
San Ram6n, cordillera de Tilardn, El Castillo, Harmon
338 (CR. INB. MO); Upala, cordillera de Guanaeaste, Bar¬
que Nac. Rincbn de La Vieja, Colonia Blanca, Rivera
1498 (CR, INB. MO). Guanaeaste: Liberia, cordillera de
Guanaeaste, Kst. Cacao, Sendero a Est. Marilza. Gonzalez
llll (CR, INB, MO); Parque Nae. Rincdn de La Vieja.
Sendero a la Ionia de agua, Rivera 599 (CR, INB. MO);
Tilardn, cordillera de Tilanin. Palmital, Rfo Carlo Negro,
Ouehrada Malanga, Hello 4897 (CR. INB, MO).
3. Mortoniodendron caulifloruni Al. Rodri¬
guez, sp. nov. TIPO: Costa Rica. Puntarenas:
Peninsula de Osa, Rancho Quemado, cerca de
15 km al Oeste de Rincdn. Reserva Forestal
Golfo Dulce, bosques aledanos a la carretera
a lo largo del Rfo Riyito y cerca riel puente,
8°42 ' IN , 83°33'0, 250-350 m. 31 mayo 1988.
B. Hummel 16924 (holdtipo, I NR; isdtipos,
CR. MO). Figura 3.
Ilaec species a congeneris inflorescentiis eaulifloris ra-
miflorisve atque etiarn terminalibus axillaribusve distin-
guitur.
Arbusto o arbol pequeno de 1-8(15) m; ramitas
teretes, glabras a puberulentas, con diminutos tri-
comas estrellados o simples. Hojas con peciolos 3—
7 mm, glabros a esparcido y estrellado-puberulen-
tos; lamina 11.5—32.5 X 3.5—10 cm, oblonga a
oblongo-elfptica, coridcea. glabra sobre ambas su¬
perficies, aunque a veces con esparcidos y dimi¬
nutos tricomas estrellados o simples sobre el nervio
central o los secundarios, apice largo-acuminado,
base redondeada, cuneada o leve oblicua, con 5—9
pares de nervios secundarios, 3 nervios saliendo
desde la base de la ldmina, enves a menudo con
domacios en las axilas del nervio central y en .se¬
cundarios. glandnlas pelucidas ausentes; estfpulas
0.2—1 mm, pronto deciduas. Inflorescencias son
cortas cimas o racimos terminates, axilares, cauli-
floras o ramifloras, 0.8-5 cm, con 1-5 flores, ejes
denso y estrellado-puberulentos, pedunculos desde
casi obsoletos hasta 1.5 cm, en la mayorfa de los
casos bracteados, bracteas saliendo cerca dr* la
base y ca. 1 mm. Flores con pedicelos 5-10 mm,
artieulados, la mayorfa bracteolados, bracteolas
0.75—1 mm; sepalos 4, 6—8 mm, elfptico-lanceola-
dos, denso y estrellado-puberulento en el lado ex-
terno. disperse y estrellado-puberulento en la parte
interna; petalos 4, ca. de 5 mm, lanceolados, blan-
cos, glabros; falanges 4, opuestos a los petalos,
cada uno con ca. 1 I estambres, |)arte libre de los
filamentos 1-2 mm. anteras 2. 5-3.2 mm incluyen-
do el mucro, oblongas, desiguales, mucro ca. 0.25
mm; ovario con 5 leves costillas, estilo 4-5 mm.
glabro. Frutos 1. 5-2.5 X 0.8-2.5 cm, obloides, 5-
locular. con 5 leves costillas, costillas formadas a
lo largo del punto de union de las valvas, valvas
erustaceas, pared ca. 0.5 mm de grosor; semillas
1—2 por loculo, fertdes a pocas veces esteriles, fbr-
tiles 4—5 mm de diametro. orbiculares, arilo sub-
tendiendo su parte basal.
Fenologia. Flores entre junio y octubre; frutos
la mayor parte del afio.
Distribucidn. End^mica de Costa Rica, se rest-
ringe a los bosques muy humedos del Pacffico Sur,
en la Peninsula de Osa y Golfito, desde el nivel del
mar hasta 600 m.
En el gcnero. Mortoniodendron caulifloruni es la-
cil de distinguir por ser la linica especie con inflo¬
rescencias caulifloras o ramifloras. aunque tambien
terminales o axilares. Ademds, se reeonoce por su
habito frecuentemente arbustivo, inflorescencias
cortas, con pocas flores y partes vegetativas gla-
brcscentes a diminuto puberulentas. Colecciones
Volume 14, Number 4
2004
Rodriguez
Mortoniodendron (Tiliaceae)
481
482
Novon
de esta especie han sido incluidas errdneamente
bajo M. anisophyllum, especie con la cual muestra
gran afinidad y cs simpdtrica, pcro esta ultima sc
distingue debido a que tiene inllorcsccncias con
frecuencia multiHoras, hasta 20 (lores (vs. hasta 5
(lores), nunea son caulifloras o ratnidoras, cl mucro
apical de las anteras es mas largo, entre 0.4— 0.6
mm (vs. ca. 0.25 mm), las semillas fertiles se en-
cuentran subtendidas por cl arilo hasta % partes
tamano o mas (vs. subtendidas basalmente) y final-
mente debido a que es una especie que puede al-
canzar mayor altura, hasta 20 m de altura (vs. hasta
8 m).
En los bosques muy humedos de la Peninsula de
Osa se ha observado que Mortoniodendron cauliflo-
rum es bastante comun. Esta zona se ha caracte-
rizado por cl gran numero de especies endemicas
descritas, lo (]ue sustenta cl valor bioldgico de esta
region c invita a la prescrvacidn de sus habitats
naturalcs. Apartc de GoKito y Peninsula de Osa,
probablemente esta especie se encuentra en otras
areas de la vertiente Pacifica de Costa Rica, e in¬
clusive de Panama.
Pardtipos. COSTA RICA. Punturenas: GoKito, Re-
fugio Natural de Vida Silvestre Golfito, hi la Gamba, Ldpiz
525 (CR, INK); Osa, Parque Nae. Coreovado, Cerro Brujo,
Herrera .1976 (CR, INK, \10); Kst. Sirena , Aguilar 2404
(CR. INK, MO); Cos Planes, Gonzalez 51 (INK. MO); Sen-
dero Mira dor, Angulo 484 (CK. INK. MO); Sendero Rio
Claro, Kernan 877 (MO); Res. For. Golfo Dulce, Agua
Kuena, Aguilar 517 (MO); Kst. Agujas, Azofeifa 214 (INK):
Kst. Kiol. Marenco, Burger et al. 12355 (NY); Cos Mogos.
Aguilar 2946 (CK. INK, MO); Puerto Piro, Gonzalez 436
(CR, INK, MO); Quebrada Huaca, Dodge & Goerger
10309 (MO); Rancho Ouemado. Ilammel 18299 (CR, INK.
MO); Kincdn, Rodriguez 2444 (CR. INK. MO); Sierpe, Mo¬
rales 31 (INK).
4. Mortionodcndron Iniigipcdimculat uni Al.
Rodriguez, sp. nov. T1P0: Costa Rica. Limdn:
Talamanca, Amubri, margen izquierdo del rio
Uren, Coma Sheuab, 9°32.5'N, 82°54'0, 150
m, 25 jun. 1989, 6. Herrera 2992 (holdtipo.
CR; isdtipos, MO, USJ). Figura 4.
Haec species inter eongeneros indorescentia pedunculo
notabiliter elongato insidente atque foliorum nervis basa-
libus vix rnanifestis perfacile reeognoscitur.
Arbusto o arbol pequeno de 3-4 m; ramitas te-
retes, glabras. Hojas eon peciolos 1.5—5 mm, estre-
llado-puberulentos; lamina 9.5—13 X 3—3.8 cm,
eliptiea, eoriacea, haz y envds glabros o eon ditni-
nutos y esparcidos tricomas estrellados, apice acu-
minado, base conspicuo oblicua, eon 6—8 pares de
nervios secundarios, 3—5 nervios ligeramente visi¬
bles saliendo desde la base de la lamina, enves a
menudo con domaeios en las axilas del nervio cen¬
tral y en secundarios, glandulas pelucidas ausentes;
estipulas no observadas, pronto deciduas. Indores-
ceneias son paniculas cimosas, terminales, 4-10.5
cm, 5— 10-flores, ejes glabros a esparcido puberu-
lentos, pedunculos 3.5— 8.5 cm, bracteas no obser¬
vadas, pronto deciduas. Flores con pedicelos 5—10
mm, articulados, bracteolas no observadas, pronto
deciduas; sepalos 5, ea. 7 mm. eliptieos, denso es-
trellado-puberulentos en el lado externo, eorlo to-
mentoso a puberulentos en la parte interna; petalos
5, 4-5 mm, lanceolados, con la base blanca y el
apice amarillo, glabros; falanges 5, opuestos a los
petalos, cada uno con ca. 7 estambres, parte fibre
de los filamentos 1-1.5 mm. anteras 1.5-1.75 mm,
incluyendo el mucro, oblongas, subiguales, mucro
ca. 0.3 mm; ovario con 5 leves cost i I las, estilo ca.
2 mm. glabro. Frutos ca. 1.5 X 1.5 cm, obloides,
5-locular, con 5 leves costillas, costillas formadas
a lo largo del punto de union de las valvas, valvas
cnistaceas, pared ca. 0.25 mm de grosor; semillas
2 por ldeulo, fertiles o esteriles. fertiles 5—6 mm de
diametro, orbiculares, eubiertas totalmente por el
arilo.
Fenologia. Flores en junio; frutos en julio.
DLstrilmeidn. Endemica de Costa Rica, restrin-
gida a los bosques muy humedos de la vertiente
Atlantica, region de Suretka y Amubri en Talaman¬
ca, entre 150—250 m.
Mortoniodendron longipedunculaturn se reconoee
de las otras especies del genero por presentar in-
florescencias eon pedunculos muy alargados. Ade-
tnas, se distingue por su pequeno habito, ramitas y
hojas glabrescente, lamina angosta, con la base
conspicuo oblicua, nervios basales poco visibles y
por sus semillas eubiertas en su totalidad por el
arilo. Esta especie presenta similitud con M. cau-
lijlorum y M. anisophyllum debido a sus frutos pe-
quenos, con valvas crustaceas y delgadas. Sin em¬
bargo, ambas especies se separan debido a que
podrfan mostrar infloresceneias con pedunculos
mas cortos (nunea mds de 1.5 cm), hojas con la
base redondeada, euneada a leve oblicua, nervios
basales conspicuos y flores con mayor numero de
estambres por falange (10-1 I estambres).
Pardtipo. COSTA RICA. Linion: Talamanca, Suret-
ka, bosques cercanos al sitio de explorat ion petrolera,
Ixiurito 9531 (CR, USJ).
5. Mortoniodendron nioralesii Al. Rodriguez,
sp. nov. TIPO: Costa Rica. San Jose: Acosta,
Fila Bustamante, Hacienda Tiquires, bosque
primario y secundario en el camino a Aguas
Buenas, 9°43'10"N. 84°10'50"C), 1600-1800
m, 3 jun. 1995, F. Morales 42BI) (holdtipo,
INB; isdtipos. BM, CR, F. K, MEXU, MO, NY,
IIC, US, W). Figura 5.
Volume 14, Number 4
2004
Rodriguez
Mortoniodendron (Tiliaceae)
483
F'igura 4. Mortii 'odendron longipedunculatum Al. Rodriguez (Herrera 2992). — A. Kamila con hojas e infloresoencia.
— B. Domacio. - C. Flor. — D. Pistilo y s^palos (petalos caedizos). — E. IVtalo y falange. — F. Fruto en seccidn
transversal. — G. r ruto. — H. Semilla subtendida por el arilo.
Haec species inter congeneros fructus ellipsoidei gran-
d is ecostati vel leviter costati loeulis .'5- ad 6-seminalibus
et valvarum parietibus percrassis induratis recognoseitur.
Arbol de 6-15 m; ramitas teretes. estrellado-to-
mentosas en partes terminales. Hojas con peciolos
entre 8—14 mm, estrellado-tomentosos; lamina 7—
14 X 3.5— 6.5 cm, elfptica a obovada-eliptica, co-
riacea, haz glabrado a estrellado-puberulento sobre
los nervios, enves densamente estrellatlo-pul)ern-
lento, apice corto acuminado, base leve asimetrica.
484
Novon
cuneada o rara vez redondeada, con 7—10 pares de
nervios seeundarios, 3 nemos saliendo desde la
base de la lamina, enves a menudo con domacios
en las axilas del nervio central y en seeundarios,
gldndulas pelueidas ausentes; estfpulas 1.5—2.25
mm, deciduas. lnllorescencias son cimas terminales
o axilares, 3—10 cm, ejes es about as long
as th*1 throat, and strigose, generally conspicuously
apically glandular eypselae (see Fig. 1). These
characters are congruent with those of an expanded
Ixiennecia. Moreover, in the protologue of Erigeron
araneosus. Urban (1902-1903) related it to E. ni-
vens Schultz Bipontinus (a synonym of L. gnaphal-
ioides, the generitype), which independently sup¬
ports our relocation of C. araneosa to Ixiennecia. In
Volume 14, Number 4
2004
Sancho & Pruski 487
Laennecia araneosa from the West Indies
1 mm
B
Figure 1. Laennecia araneosa (Urban) G. Sancho & Pruski. — A. Disk corolla. — B. Cypsela with pappus removed.
— C. Twin hair of cypsela. — I). Glandular trichome of cypsela. (Drawn by Gisela Sancho from Ekman 13566 , US.)
addition to making the following needed combina¬
tion, we provide a description of L. araneosa in¬
cluding the diagnostic generic features of Laennecia
lacking in the protologue of E. araneosa Urban, yet
stressed by Nesom (1990),
No West Indian species of Conyza now referable
to Laennecia (other than L. araneosa) were listed
by Adams (1972), Correll and Correll (1982), Gri-
sebach (1864), Howard (1989), Liogier (1962,
1996. 1997), Nesom (2000), Proctor (1984), or Ur¬
ban (1921), nor are any Virgin Islands species re¬
ferable to Laennecia. Thus, the transfer to Laen¬
necia of this species from the Dominican Republic
marks the first report of the genus Laennecia in the
West Indies. The 18 other species of Laennecia oc¬
cur on Continental America between the south¬
western United States and northern Argentina (Ne¬
som. 1990; Sancho & Ariza Espinar, 2003), and 5
of these mainland species are treated in the Laen¬
necia treatment in Flora Mesoamericana (Sancho &
Pruski, in prep.).
Laennecia araneosa (Urban) G. Sancho & Pruski,
comb. nov. Basionym: Erigeron araneosus Ur¬
ban, Symb. Antill. 3: 404. 1903. Conyza ara¬
neosa (Urban) Cronquist, Bull. Torrey Bot.
Club 70: 632. 1943. TYPE: Dominican Re¬
public. Ua Vega: Valle Nuevo, 2270 m. May
sin. annum, Eggers 2200 (holotype, B, de¬
stroyed, photograph (taken by Harvey M. Hall
in 1925) at UC not seen, US). Figure I. Al¬
though we were unable to locate isotype ma¬
terial (in C. MO, NY, 0. UC, US), we are op¬
timistic that isotype material exists in other
herbaria, from which a future lectotype sheet
may be designated.
Low annual or short-lived perennial, tap-rooted
herbs, 8—49 cm tall, few- to several-branched from
near the base, herbage glandular, arachnoid-lanate,
trichomes glandular and non-glandular; stems as¬
cending, subterete, internodes commonly slightly
shorter than leaves. leaves alternate, sessile, mod¬
erately pinnately lobed, (5)7—15 mm long, 1—4 mm
wide, oblanceolate, chartaceous, base slightly
clasping, margins flat, marginal lobes 2 to 4, ellip¬
tic-ovate, ca. 1-1.5 X ca. 0.5 mm, apex of blade
and lobes acute to rounded, sometimes slightly mu-
cron ulate. Capitulescence axillary from the upper¬
most nodes, racemose; peduncles 0.3— 1.2 cm long.
488
Novon
Capitula heterogamous, subradiate, 50- to 70-flow-
ered; involucre campanulate, 3-4(4. 5) mm high. 3—
4.5 mm wide; phyllaries 2(3)-seriate, linear-lance¬
olate, 2-4(-4.5) mm long, ca. 0.7 nun wide, glan¬
dular. basally tan, midrib region greenish, margins
hyaline, apex acuminate, often lacerate or ciliate.
Marginal florets pistillate, 45 to 61, pluriseriate,
corollas subradiate, filiform, 2.5—3 mm long,
cream-colored, tube weakly glandular, limb dis¬
tinct, short, 0.2— 0.5 mm long, apex hi- or triden-
tate, style often exserted to apex of limb; central
disk florets bisexual, 5 to 9, corolla funnelform, 2.5—
3 mm long, cream-colored, glandular at the apex of
the tube and lobes, tube ca. 1.5 mm long, throat
04)— 0.8 mm long, lobes 5, lanceolate, 0.4— 0.7 mm
long, anther thecae ca. 0.5 mm long, style branches
ca. 0.5 mm long, apex triangular-lanceolate. Cyp-
selae compressed, elliptic-obovate, 1.2— 1.4 mm
long, apical ly constricted, moderately 2-nerved
marginally, strigose, generally resinous glandular
distal ly. glands often deciduous rather than persis¬
tent; pappus uniseriate, of 15 to 25 scabrid stra¬
mineous bristles, usually 2.5—3 mm long.
Distribution and ecology. Laennecia araneosa is
endemic to the Dominican Republic. This species
is locally common in grassy pinelands of the Cor¬
dillera Central in the south-central part of the is¬
land at elevations between 2000 and 2300 m. Most
collections are from near Valle Nuevo, at approxi¬
mately 18°48'N, 70°42'W. Plants seemingly flower
year round, having been collected in January, April,
May, August, and September. The genus should be
looked for in mountainous regions elsewhere in the
West Indies, especially on the biogeographically
similar Cuba.
We place Laennecia araneosa in section Laen¬
necia because of its arachnoid-lanate vestiture, pin-
nately lobed leaves, marginal corollas with short
ligules, and cypselae generally with large resinous
glands. Within section Laennecia, L. araneosa
seems, by clasping leaves, glandular leaves and
phyllaries, ligulate marginal corollas, and few (15
to 25) pappus bristles, to be most closely related to
/,. microglossa and L. schiediarui.
Additional material examined. DOMINICAN REPUB¬
LIC. Azua: Cordillera Central, San .luan, Lomas de la
Mediam'a, near Sabana Nueva. Ekman 13566 (NY, US).
La Vega: Valle Nuevo, Bueno 1753 (US). Jimenez A Lith-
gow 1335 (US). Liogier 12207 (NY, US); prope Constanza
infra Valle Nuevo, von Ttirckheim 3160 (MO. NY, US).
Acknowledgments. We thank Guy Nesom, Tom
Xanoni, and Elsa Zardini for helpful discussion and
for reviewing the manuscript. We thank Olof Ryd-
ing (C) and Per Sunding (0) for searching for iso-
tvpe material of Eggers 2200 in their respective
herbaria. The first author was supported during
preparation of this manuscript by Consejo Nacional
de Investigaciones Cientfficas y Ternicas (CONI-
CET), Argentina, and the Elizabeth Bascom Eel-
lows hi, >s of the Missouri Botanical Garden.
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Adams, C. I). 1972. Flowering Plants of Jamaica. Univer¬
sity of die West Indies, Mona. Jamaica.
Correll. I). S. & H. B. Correll. 1982. Flora of the Bahama
Archipelago. Cramer, Vaduz.
Cronquist, A. 1943. The separation of Erigeron from Co-
nyza. Bull. Torrey Bot. Club 70: 629—632.
Cuatrecasas, J. 1963. Notes on Neotropical Compositae,
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- . 1969. Prima Flora Columbiana. 3. Compositae—
Astereae. Webbia 24: 1-335.
Crisebaeh, A. H. B. 1864 1 1861 1. Synanthereae. Pp. 352-
385 in Flora of the British West Indian Islands. Lovell,
Reeve, London.
Howard, R. A. 1989. Flora ol the Lesser Antilles, Leeward
and Windward Islands, Vol. 6. Arnold Arboretum and
Harvard University, Jamaica Plain.
Liogier. II. A. 1962 ( 1963]. Flora de Cuba. Vol. 5, Ru-
biales— Valeriana les— Cucurbi tales— Cam pan id ales— A s-
terales. Universidad de Puerto Rico, Rio Piedras. |As
“Hermano Alain (l)r. F. F. Liogier).v|
- . 1996. La Flora de la Fspanola. VIII. Universidad
Central del Fste, Vol. 72, Ser. Ci. 29: 1 — 5H11. San Pedro
de Macon's, Bepuhlica Dominicana. [As "Alain Henri
Liogier. "|
- . 1997. Descriptive Flora of Puerto Rico and Ad¬
jacent Islands, Vol. 5. Universidad de Puerto Rico, San
Juan.
Nesom, C. L. 1990. Taxonomy of the genus Laennecia
(Asteraeeae: Astereae). Phytologia 68: 205—228.
- . 2000. Generic Conspectus of the Tribe Astereae
(Asteraeeae) in North America, Central America, the
Antilles, and Hawaii. Sida. Bot. Miso., vol. 20.
- . 2001. hienneeia turnerorum (Asteraeeae: Aster¬
eae). a new species from trans-Pecos Texas. Sida 19:
789-793.
Proctor. B. 1984. Flora of die Cayman Islands. Kew
Bull. Addit. Ser.. XI.
Sancho, G. A L. Ari/.a Espinar. 2003. Asteraeeae, parte
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Asterinae (excepto Grindelia > Haplopappus). Flora Fa-
nerogdmiea Argentina Fascfeulo 81 : 1 — 102. PROFLO¬
RA, CONICFT, Crtrdoha.
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Anlillanae, Vol. 3(2—3): 280-420. Fratres Borntraeger,
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- . 1921. Compositae. In: Flora Domingensis. Sym-
bolae Anlillanae, Vol. 8(2): 703—751. Fratres Borntrae¬
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del gtmero Conyza Less. II. Rehabilitaeidn del genero
Laennecia Cass. Darwiniana 23: 159—169.
Ballota byblensis (Lamiaceae), a New Species from Lebanon
Myrna T. Semaan
Center for Environmental Development, Awareness & Research (C.E.D.A.R.), P.0. Box 967,
Jounieh, Lebanon, fon@sodetel.net. lb
Ricardus M. Haber
Institute for Conservation of Biodiversity and Ecotourism Development (USEK University),
P.O. Box 967, Jounieh, Lebanon
Abstract. A new species of the genus Ballota L.
(Lamiaceae) is described from the western slopes
of Mount Lebanon in an eastern Mediterranean cli¬
mate. Ballota byblensis Semaan & K. M. Haber is
distinguished by die following characters: perennial
herb, suffrutescent, brittle; stem densely matted
with patent liny simple to long septate glandular
trichomes, with few sessile glands; eauline leaves
cordate, crenate, rugose, tomentose with glandular
simple-stellate trichomes on both sides; inflores¬
cence lax; verticillaster 6- to 19(22)-Howered; brac-
teoles many, filiform, glandular, shorter than calyx;
calyx 9-10 mm long, infundibular, with glandular
simple-stellate trichomes outside, appressed pilose-
pubescent in the upper half inside; limb erecto-
patent, oblique; teeth 10(11), unequal; corolla 16-
18 mm long, pink-purple; seeds black.
Key words.' Btdlotti , endemic, Lamiaceae, Leb¬
anon. Mediterranean.
The genus Ballota (Lamiaceae) comprises five
species in Lebanon (Mouterde, 1983; Greuter et al.,
1986; Heller & lleyn, 1986). Btdlota antdibanotica
Post, B. undulata (Sieber ex Iresenius) Bentham,
and B. damascena Boissier are basically restricted
to the Levant and Sinai occurring in sub-arid to
arid regions. Ballota nigra L. subsp. uncinata
(Fieri & Beguinot) Patzak and B. saxatilis Sieber
ex J. & C. Presl attain wider geographical distri¬
bution surviving under more temperate conditions
(Davis & Doroszenko, 1975).
Ballota byblensis Semaan & B. M. Haber, sp. nov.
TYPE. Lebanon. Jbeil Province: Jaj Mountain,
1600 m, 34°09'22.8"N, 35°50'10.2"E, 10 Aug.
2002, R. Haber & M. Semaan 2002 (holotype,
MO; isotypes. BE1. K). Figure 1.
Herba perennis, suffrutescens, fragihs; caulis dense im-
plicitus cum patentibus minutis longis septatis glandiferis
pilis et paucis sessilibus glandibus; folia eaulina cordata,
crenata, rugosa, pilis stellatis glandiferis in superficiebus
ambabus folii; inflorescentiae laxae; verticillastri 6— 19(22)
floribus; bracteolae plurimae, 2.0— 6.0 X 0.2-0. 5(0.7) mm,
liliformes, glandiferae; calyx 9-10 mm, infundibularis, ex¬
tra glandifer, stellatis pilis, intra pubescens in dimidio
superiore; limbus erecto-patens, obliquus; dentes 10(11)
inaequales; corolla 16-18 mm, rosea-purpurata.
Perennial herb, suffrutescent, rhizomatous, fra¬
grant, brittle woody and green parts; flowering
stems up to 35 cm long, less than 2 mm wide, qua¬
drangular, branched or simple, brown at base,
green above, densely matted with patent liny sim¬
ple to long septate glandular trichomes, and with
few sessile glands, vegetative stems 5—7 cm long.
Cauline leaves petiolate, petiole 2.0— 2. 5(3.0) cm
long, blade (2.0)2.5-3.3 X 2. 5-3.3 cm, orbicular
to broadly oval, cordate, deeply crenate, rugose, to¬
mentose with stellate glandular trichomes above,
tiny simple to long septate glandular trichomes and
sessile glands below, green above, grayish green
below with denser indumentum; floral leaves peti¬
olate, petiole 3-20 mm long, blade 1.0-2. 5 X 1.0-
2.5 cm, orbicular-oval, truncate-cordate, indumen¬
tum similar to cauline leaves. Inflorescence lax;
verticillaster 6- to l9(22)-flowered, pedicellate;
pedicels 2-3 mm long, with dense patent tiny sim¬
ple to long septate glandular trichomes; bracteoles
many, 2.0-6.0 X 0.2-0.5(0.7) mm. filiform, some¬
times with 1 or 2 teeth, mucronate, with patent tiny
simple to long septate glandular trichomes, denser
on outside and occasionally with stellate trichomes.
Flower 19—20 mm long, sessile; calyx 9—10 mm
long, infundibular; tube 7.5 mm long and termi¬
nating in a circular vein, with dense patent tiny
simple to long septate glandular trichomes outside,
and stellate glandular trichomes in the upper third,
glabrous in the lower half inside and appressed pi¬
lose-pubescent in the upper half; limb 6-7 mm
diam., erecto-patent, oblique (fig. 2); teeth 10(1 1),
unequal, l .0-2.5 X 0.7-1. 4 mm including mucro
of 0.2— 0.5 mm, triangular-oval, pubescent inside,
with glandular stellate, liny simple and long septate
trichomes outside; corolla 16—18 mm long; tube 9—
Novon 14: 489-492. 2004.
490
Novon
Figure 1. Ilal lota byblrnsis Seaman & R. M. Haber, photos of holotype
— A (left). Whole plant. — B (right). Inflorescence.
collection taken In authors in Jaj Mountain.
igure 2. Illustration of calyx limb drawn by tbe authors from the holotype specimen.
Volume 14, Number 4
2004
Semaan & Haber
Ballota byblensis from Lebanon
491
11 mm long, white, well exserted, with sessile
glands and downward appressed long trichomes
outside, and annulus of trichomes inside; upper lip
6.0— 7.5 X 3.5 mm, white outside, pink inside, bifid
with lobes divided and/or gnawed, appressed vil¬
lous outside at base and margins, with dense stel¬
late glandular trichomes at center especially on
hood, bearded inside at hood and lobes, glabrous
below; lower lip 6.0— 6.5 mm long, purple striated
with white, glabrescent below; filaments variegated
white and pink-purple, with papillae below, lanate
above; style 14 mm long, white bifid. Seeds 2.6-
2.8 X 1.2— 1.3 mm, oblong-obovate, black, tip trun¬
cate, with sessile white glands gradually disap¬
pearing with maturity.
Distribution and habitat. Ballota byblensis is
localized and confined to the rugged high Jaj Moun¬
tain of Jbeil Province on the western slopes of the
Mount Lebanon Range at 1600—1800 m in altitude.
The habitat is marked with a harsh winter of low
temperature, stormy winds, frost, and a consider¬
able snow cover. The summer is moderate and hu¬
mid (Service Meterologique, 1966, 1967). I he re¬
gion lost its conifer forests over time, reducing the
terrain to strongly eroded masses of rocks with
pockets of soil (Beals, 1965). The occurrence of B.
byblensis is relatively common. It hangs out of crev¬
ices in sandy limestone rocks with woody branches
extending over rock faces. Grazing is intensive in
the region anti strongly impacts the green cover.
Though B. byblensis seems not consumed by ru¬
minants, its brittle nature exacerbates the impact
of trampling and similar abrasive activity. The flow¬
ering period of B. byblensis extends from August to
early September, as observed by the authors.
Ballota byblensis is closest morphologically to B.
saxatilis and B. antilibanotica of the Lebanese flo¬
ra. The differentiation of B. byblensis from other
Ballota species in Lebanon is elucidated in the fol¬
lowing key (Post, 1932; Davis & Doroszenko, 1975;
Feinbrun-Dothan, 1978; Mouterde, 1983).
la. Calyx limb short with shallow crenation or den¬
tation.
2a. Calyx limb multicrenate to multidentate,
broadly expanded; stem hirsute . . B. undulata
2b. Calyx limb w ith 10 to 12(20) spinulate eren-
ae; stem white floccose-lanate .
. B. damascena
lb. Calyx limb with triangular-ovate teeth.
3a. Calyx teeth 5; bracteoles subulate . . . B. nigra
subsp. uncinata
3b. Calyx teeth at least 10; bracteoles linear-fi¬
liform.
4a. Stem with short stellate appressed pu¬
bescence and longer simple to
branched-stellate glandular trichomes;
leaves round-reniform; bracteoles as
long as calyx . B. antilibanotica
4b. Stem not as above; leaves orbicular-
ovate; bracteoles shorter than calyx.
5a. Stem matted with simple and long
septate glandular trichomes; verti-
cillasters lax; bracteoles mucro-
nate; corolla 16—18 mm .
. B. byblensis
5b. Stem eglandular-villous of shortly
fasciculate or stellate trichomes,
with or without simple trichomes;
verticillasters dense; bracteoles
emucronate; corolla 10—16 mm . .
. B. saxatilis
In addition, Ballota byblensis differs from B. sax¬
atilis by its longer bracteoles with glandular and
only occasionally stellate trichomes, and glandular
trichomes on the calyx. Ballota byblensis differs fur¬
ther from B. antilibanotica by having a well-ex-
serted corolla tube and a darker pink-purple corolla.
I'lie tw7o species are allopatric with considerable
disparity in habitat characteristics: B. antilibano¬
tica is an inland species of semi-arid regions,
whereas B. byblensis is a montane species. Among
the flora of Turkey, B. byblensis shares morpholog¬
ical characters with the two endemic species B. la-
tibracteolata P. H. Davis & Doroszenko and B. lar-
endana Boissier & Heldreich, which are close to
B. saxatilis. Ballota byblensis differs from B. lati-
bracteolata by having both sides of leaves glandu¬
lar. non-ciliate filiform bracteoles, infundibular ca¬
lyx, erecto-patent limb, pink-purple corolla with
white stripes, not white corolla with purple stripes.
Ballota byblensis differs from B. larendana by its
brittle nature, by glandular leaves, bracteoles short¬
er than calyx, glandular infundibular calyx, erecto-
patent limb, and longer corolla. Ballota byblensis is
also distinguished morphologically from the central
Mediterranean B. rupestris (Bivona-Bernardi) \ isi-
ani and the southwest European and northwest Af¬
rican B. hirsuta Bentham (Tutin et ab, 1972). Bal¬
lota byblensis differs from B. rupestris by smaller
and ovate-orbicular cauline leaves, non-membra-
nous bracteoles, larger diameter of limbs, and lon¬
ger corolla. It also differs from B. hirsuta by its
smaller calyx and limb diameter, and longer corol¬
la.
Ballota byblensis is named after the Neolithic
Canaanite city, Jbeil (Byblos), as il was discovered
on the overlooking Jaj Mountain slopes from which
the Phoenicians felled cedars to trade wood with
the ancient world (Meiggs, 1982).
Paratype. LEBANON. Jbeil Province: Jaj Mountain.
1697 m. 24 Aug. 2002. R. Haber & \1. Semaan 2002
(BEI, MO, K).
492
Novon
Acknowledgments. The authors thank the Khal¬
il Fallal and Sons Establishment for financially
supporting research projects of the Friends of Na¬
ture NGO. The “Wild Flora Project” already re¬
sulted in the discovery of several probable new spe¬
cies and many described taxa previously
unaccounted for among tin- flora of Lebanon.
Literature Cited
Beals, E. W. 1965. The remnant cedar forest of Lebanon.
J. Ecol. 53: 679—694.
Davis, 1’. II. A A. Doroszenko. 1975. Hallota I,. Pp. 156—
165 in P. H. Davis, Flora of Turkey, Vol. 7. Edinburgh
Univ. Press, Edinburgh.
Eeinbrun-Dothan, IN. 1978. Hallota L. Pp. 125-128 in
Flora Palaestina, Part III. The Israeli Academy of Sci¬
ences and Humanities, Jerusalem.
Greuter, W., II. M. Burdet & C. Long. 1986. Med-Cheek-
I ist. Vol. 3. Conservatoire el Jardin Botaniques, Villede
GenFve.
Heller, I). & C. C. Ileyn. 1986. Conspectus Florae Orien-
talis, Vol. 3. The Israel Academy of Sciences and Hu¬
manities, Jerusalem.
Meiggs, If. 1982. The cedars of Lebanon, pp. 49—87. De¬
forestation, pp. 371-103. In Trees and Timber in the
Ancient Mediterranean World. Oxford Univ. Press, Ox¬
ford.
Mouterde, P. 1983. Hallota L. Pp. 141-144 in Nouvelle
Flore du Liban et de la Svrie, Vol. 3. Dar El-Machreq,
Beirut.
Post, G. 1932. Hallota L. Pp. 389-392 in Flora of Syria,
Palestine and Sinai, Vol. 2. American Press, Beirut.
Service Meterologique. 1966. Atlas Climatique du Liban:
Pluie, Temperature. Pression. Nebulosity, Tome I. Di¬
rection de 1’ Aviation Civil. Service Meterologique, Bey¬
routh.
- . 1967. Atlas Climatique du Liban: Humiditd At-
mospherique, Statistique Diverses de Frequence, Tome
II. Direction de l’Aviation Civil. Service Metdrologique,
Beyrouth.
rutin, T. G., V. H. Heywood, N. A. Burges, I). M. Moore,
I). H. Valentine, S. M. Walters & 1). A. Webb (editors).
1972. Hallota L. Pp. 149—151 in Flora Europaea, Vol.
3. Cambridge Univ. Press, Cambridge.
Rubiacearum Americanarum Magna Hama Pars XVI. New Species,
a New Subspecies, and an Overlooked Species of Psychotria
subg. Heteropsychotria from Mexico, Central America,
and Western South America
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, LJ.S.A.
charlotte.taylor@mobot.org
Abstract. Described and illustrated are the new
species: Psychotria calidicola C. M. Taylor, found
in Costa Rica and Panama and previously confused
with P. eurycarpa Standley; P. chocoana C. M. Tay¬
lor. found in northwestern Colombia; P. gaitalensis
C. M. Taylor of central Panama, previously con¬
fused with P. simiarum Standley; P. osaensis C. M.
Taylor of southern Costa Rica, previously confused
with P. deflexa DC.; P. paradichroa C. M. Taylor of
Panama, previously confused with P. dichroa
(Standley) C. M. I aylor; Psychotria sanblasensis C.
M. Taylor of eastern Panama, previously confused
with P. croceovenosa Dwyer; P. sinuata C. M. Taylor
of northwestern South America, similar It) P. an-
gustiflora k. Krause and distinguished by its usu¬
ally sinuate to scalloped leaf margins; and P. soe-
jartoi C. M. Taylor of northwestern South America,
previously confused with P. stenostachya Standley.
Also described and illustrated is a new subspecies,
P simiarum subsp. cluapensis of central and south¬
ern Mexico, which is allopatric with subspecies
simiarum of coastal Veracruz, Mexico, to Panama.
The name P. buchtienii (11. W inkler) Standley ap¬
plies to a commonly collected species found from
southern Mexico through Bolivia, for which a de¬
tailed description is presented; P. huclitienii has
previously been confused with P. officinalis (Au-
blet) Sandwith. All of these species belong to Psy¬
chotria subg. Heteropsychotria (Rubiaceae, tribe
Psyehotrieae).
Key words: Neotropics, Psychotria, Psycho-
trieae, Rubiaceae.
The pantropical genus Psychotria L. (Rubiaceae:
Psyehotrieae) includes perhaps 2000 species of
shrubs and small trees. This genus is characterized
by the presences of raphides, relatively small, in¬
sect-pollinated. white flowers, ovules that are basal
and solitary in each of the two ovary locules, four
or usually five valvate corolla lobes, and drupa¬
ceous fruits with hard-walled, single-seeded py¬
renes. In the Neotropics, two principal lineages
have been identified within Psychotria in its broad
sense, as well as several smaller ones. 1 hese large
groups are Psychotria subg. Psychotria and Psycho¬
tria subg. Heteropsychotria Steyermark (Taylor,
1996; Nepokroeff et al., 1999). Psychotria subg.
Psychotria is distinguished by its usually caducous
stipules, of various forms; the generally gray or
brown color of its dried specimens; and its fruits
that are usually red or orange at maturity. I his
group is pantropical, though the plants of each re¬
gion (e.g., Africa, the Pacific Islands, the Neotrop¬
ics) appear to comprise separate lineages (Nepok¬
roeff et al., 1999). Psychotria subg. Psychotria
includes about 600 species worldwide with about
200 species in the Neotropics (Hamilton. 1989).
Psychotria subg. Heteropsychotria is a wholly
Neotropical group that is distinguished by its per¬
sistent stipules, which are usually united around
the stem into a continuous sheath with two lobes
on each interpetiolar side; the generally green color
of its dried specimens; and its fruits that are usually
white, blue, or black at maturity. Psychotria subg.
Heteropsychotria is similar to the Neotropical genus
Palicourea Aublet. which comprises an additional
200 species (Taylor, 1996, 1997) and is apparently
derived from within Psychotria subg. Heteropsycho¬
tria. Palicourea has been separated from Psychotria
subg. Heteropsychotria based on its flowers that are
apparently adapted for hummingbird pollination
(Taylor, 1996) and its typically somewhat larger
fruits. Plants of Psychotria subg. Heteropsychotria
have generally fragrant, diurnal, white to yellow
flowers with corollas generally 2-10 mm long
(though sometimes nocturnal and/or up to 50 mm
long, e.g., Psychotria chiapensis Standley). Their
corollas have a wide range of internal pubescence
patterns, including fully glabrous, barbate in the
throat, or pubescent in part or all of the tube. I hese
corollas are straight-sided to funnelform, or in some
species they may be bent near the base and some-
Novon 14: 493-508. 2004.
494
Novon
times also gibbous (i.e., asymmetrically swollen)
here. I his last corolla form resembles that of Pal-
icourea, in which the corollas have a well-devel¬
oped tube and a swollen, usually gibbous base that
is closed off by an internal ring of pubescence. Ap¬
parently nectar accumulates in this swollen corolla
base and is protected from many insect visitors by
this pubescent ring (Taylor, 1996). In addition to
this swollen base and internal pubescence, flowers
of Palicourea generally lack odor, are larger, reg¬
ularly 5—50 nun long, and are brightly colored.
However, the distinctions between Palicourea and
Psychotria subg. Heteropsychotria are not complete¬
ly clear, and recent molecular information indicates
that pollination mode and the correlated floral mor¬
phology may have shifted repeatedly within this
group. Thus, the hummingbird-pollinated plants
classified in Palicourea apparently have been de¬
rived several times from different insect-pollinated
lineages of subgenus Heteropsychotria. Such shifts
in pollination mode with subsequent diversification
of species have now been documented for several
Rubiaceae genera (e.g., kirkbride, 1997; Malcom-
ber, 2002; Taylor, 1994), and probably are the
source of much of the species diversity in this fam¬
ily.
Most if not all the species of Psychotria subg.
Heteropsychotria are distylous, as are nearly all
those of Palicourea (Taylor, 1997) and Psychotria
subg. Psychotria (Hamilton, 1989). The flowers typ¬
ically last for one day, but usually only one or a
few flowers of the multiflowered inflorescence open
each day, so an individual plant typically produces
flowers for weeks or even months. Similarly, the
fruits generally ripen a few at a time over a period
of weeks or months. The fruits of Psychotria subg.
Heteropsychotria are succulent, and usually watery
and sometimes a little sweet -tasting. They are dis¬
persed by birds (Blake & Loiselle, 1992; Loiselle
& Blake, 1990), and typically are removed very
soon after they are ripe (pers. obs.). In some species
the fruits pass through several color stages before
maturity, usually becoming yellow, then orange,
then red, then finally black (e.g., Psychotria race-
mosa I,. C. Richard); in these species, the black
fruits are removed quickly, and frequently not ob¬
served by collectors.
During preparation of the Rubiaceae for the Flo¬
ra Mesoamericana, the following undescribed spe¬
cies and subspecies of Psychotria subg. Hetero¬
psychotria were discovered, and the commonly
collected species previously treated as P. officinalis
(Aublet) Sandwith was found to differ from the type
ol that name and to be correctly called by a differ¬
ent, little-used name.
Psychotria huchtienii (H. Winkler) Standley,
Publ. Field Columbian Mus., Bot. Ser., 7: 303.
1931. Uragoga huchtienii H. Winkler, Repert.
Sp. Nov. Regni Veg. 8: 4. 1910. TYPE: Bolivia.
| Fa Paz] San Antonio bei Mapiri, 850 m, Dec.
1907, 0. Ruchtien 1608 (lectotype, designated
by Standley (1931: 303), B destroyed, photo at
F; isotypes, F, US not seen microfiche). Figure
1C-F.
Shrubs or small trees flowering at I m tall, to 3
m tall, branched; stems glabrous, Leaves elliptic to
ovate or lanceolate, 7.5-23.5 X 2. 5-8.5 cm, at
apex acute to usually acuminate and often some¬
what falcate, at base obtuse to acute, drying pap¬
yraceous, adaxially glabrous and often rather shiny,
abaxially puberulous to glabrous; secondary veins
7 to 10 pairs, not looping to interconnect or some¬
times doing so weakly in distal part of blade, w ith¬
out domatia, adaxially venation plane, abaxially
costa prominent, secondary veins prominulous, and
remaining venation plane; margins plane; petioles
3-17 mm long; stipules persistent, glabrous to pu¬
berulous, united around stem, sheath 0.8— 1.5 mm
long, truncate or sometimes in those stipules sub¬
tending peduncles concave with lobes very closely
grouped, lobes 2 on each side, deltoid to narrowly
triangular, 1-2 mm long, acute. Inflorescences ter¬
minal, paniculate, pilosulous to puberulous, pale
green to perhaps whitened; peduncles 1.5-4 cm
long; branched portion pyramidal to cylindrical,
1.5-4 X 2-3 cm; secondary axes I to 4 pairs, the
lowermost 1 to 2 pairs frequently reflexed, second¬
ary axes all terminating in a single congested-di¬
chotomous cymule, secondary axes not subtended
by bracts at junction with primary axis; bracts en¬
closing flowers several, ligulate to narrowly so, I —
9 mm long with longer bracts external to shorter
bracts, obtuse to acute, entire to ciliolate; flowers
distylous, sessile in glomerules of 2 to 5; hypan-
thium cylindrical, ca. 0.5 mm, puberulous to pilo¬
sulous; calyx limb 0.3— 0.8 mm long, puberulous,
shallowly lobed, ciliolate; corolla funnelform, white
to cream, externally puberulous or hirtellous some¬
times becoming glabrescent, internally glabrous in
basal half of tube and hirtellous in upper half of
tube, tube 3-5 mm long, lobes triangular, 1-2 mm
long, acute, abaxially smooth; anthers in short-
styled form 1-1.8 mm long, exserted, in long-styled
form ca. 1 mm long, included, situated in upper
part of corolla tube (i.e., opposite hirtellous region);
stigmas in short-styled form ca. 2 mm long, includ¬
ed, situated in upper part of corolla tube, in long-
styled form ca. 0.5 mm, exserted. Infructescences to
8X7 cm, becoming purple; fruits subglobose, 4.5-
Volume 14, Number 4
2004
Taylor
Psychotria subg. Heteropsychotria
495
at anthesis; based on Dnvidse et at. 20351. C-E. Psychotria buchtienii (H. Winkler) Standley; C based on Hammel
4/95; I), E based on Hammel 4200. — C. Stem with leaves and inflorescence. — D. Corolla at anlhesis (short-styled
form). - — E. Detail of inflorescence with one young flower bud and two old flowers (from which corollas have fallen). K
G. Psychotria sinuata C. M. Taylor; based on Forero 6782. — F. Leaf. — G. Detail of inflorescence with two young
flower buds and one flower (apparent long-styled form) at anthesis. H— J. Psychotria sanblasensis C. M. I ay lor; based
on t le Nevers 4123. — H. Stem with leaves and inflorescence. — I. Detail of inflorescence, with three flower buds and
one old flower (from which corolla has fallen). — J. Mature flower bud (short-stvled form), shortly before anthesis,
partially dissected. A, C, F, H to 5-cm scale; B, G to same 5-mm scale; D, F to Same 5- mm scale.
496
Novon
5 mm diam., puberulous or pilosulous to glabrous,
black or purple-black; pyrenes 2, planoconvex, dor-
sally with 4 to 5 rather sharp longitudinal ridges,
adaxially (ventrally) with a central “Y”-shaped de¬
pression.
Habitat, distribution, and phenology. In wet
forest at 0-1300 m, southern Mexico to Bolivia;
collected in flower and fruit generally throughout
the year.
This species is distinguished by its branched in-
llorescences with congested cymules and well-de¬
veloped floral bracts, corolla tubes 3—5 mm long,
subglobose fruits, and pyrenes with a “Y”-shaped
indentation near the center of the adaxial (ventral)
face. It is commonly collected, in particular in Cen¬
tral America and northwestern South America. Psy-
chotria buchtienii has been confused with P. offi¬
cinalis for a long time (e.g., Standley, 1938;
Standley & Williams, 1975; Dwyer, 1980; Burger
6 Taylor, 1993; Taylor, 2001), but in fact that latter
name correctly applies to a species that is restricted
to northeastern South America and apparently al-
lopatric to P. buchtienii. Psychotria officinalis dif¬
fers from P. buchtienii in its slender, usually very
low habit, its slender corymbiform inflorescences
with umbelliform secondary axes and the primary
axis usually not well developed, its very narrow to
linear bracts, its corolla tubes 2.5-3 mm long, and
its pyrenes with a linear longitudinal sulcus on the
adaxial face. Psychotria buchtienii has also been
confused with P. gracilenta Mueller Argoviensis
and P. hoffmannseggiana (Roemer & Schultes)
Mueller Argoviensis, both of which are sympatric
and have inflorescences or at least infructescences
of similar arrangement with well-developed bracts.
However, both of these last species also differ from
P. buchtienii in their pyrenes, which have a linear
longitudinal sulcus on the adaxial face. Details of
pyrene morphology have been considered taxonom-
ically informative in the Psychotrieae for recogniz¬
ing genera and species groups by several authors
(review in Piesschaert, 2001), and pyrene mor¬
phology has generally been found consistent at the
species level (Piesschaert et ah. 2001). The “Y"-
shaped indentation on the adaxial face of the py¬
renes of P. buchtienii does not closely match pat¬
terns documented lor other species of Psychotria
subg. Heteropsychotria. Pyrene morphology shows
considerable variation in many lineages of Psycho¬
trieae (Piesschaert. 2001; Piesschaert et ah. 2001),
and relatively very few species of Psychotria subg.
Heteropsychotria have even been studied.
Because of the large number of specimens of this
overlooked and confused species, selected speci¬
mens are cited as vouchers for geographic distri¬
bution below (with countries arranged from north to
south but their political units alphabetical), and the
remainder of the specimens examined are listed in
Appendix I.
Selected specimens examined (see also Appendix
I). MKXICO. Chiapas: 10 km ENE of Dos hagos above
Santa Elena, l>. Breedlove A' Almeda 57617 (MO). CEN¬
TRAL AMERICA. BELIZE. Cayo: Maya Mtns.. Smokey
Branch, A. Monro A' Helgason 187 (MO). Stann Creek:
Cockscomb Mtns.. tributary of Cocoa Branch of Sitte Biv-
er. 2 km due N of Victoria Peak, A. Gentry 7925 (MO).
Toledo: Camp 3, 6—8 km SW of Union Camp, trail from
Camp 3 toward Cabro on Jimmy Cut trail. T. Ilawkins
1454 (MO). ( ill A If, MALA. Alta Verapaz: bordering Bio
Sebol ea. 12 km below Sebol, E. Contreras 4204 (MO).
Izahal: Cadenas/Puerto Mendez, on Toquela River Road,
E. Contreras 9075 (MO). Peten: San Diego, Bfo Pasibn,
M. Aguilar II. 500 (MO). HONDURAS. Cortes: near
Agua Azul, Lake Yojoa, /,. 0. Williams A- Molina 17917
(MO). NICARACUA. Jinotega: Salto Kayaska, Rfo Bo-
cay, IL D. Stevens et al. 16555 (MO). Zclava: Cerro La
Pimenta Number I. summit and area adjacent to summit,
,/. Pipoly 5091 (MO). COSTA RICA. Cartago: Turrialba,
Tayutic, Eila Vereb, alto entre Cuenca del Rfo Vereh y el
Jieotea, A. Cascanle 625 (MO). Guanacaste: Parque Nac.
Guanacasle, Estacirtn Pitilla. I INBio 59 (MO). Heredia:
Einca La Selva, the OTS field station on the Rfo Puerto
Viejo just E of junction with Rfo Sarapiquf, Damon Smith
510 (MO). I.imdn: 7 km SW of Brihri, E I ). Gdmez et al.
20525 (MO). Piiiitarcnas: top of Ridge Road, near Bin-
con de Osa, II. Kennedy 1 969 (MO). Sail Jose: vicinity
of El General, A. Sketch 21 65 (MO). PANAMA. Boeas
del Toro: along road from Eortuna Dam to Cbiriquf
Grande, C. McPherson 8428 (MO). Chiriipu: along road
in vicinity of branch in road to Cerro Colorado and Es-
eopeta, above Bfo San Eelix near town of San Felix, T.
Croat 55420 (MO). Code: top of El Petroso, W. D'Arcy
11565 (MO). Darien: S of El Real, region called Alturas
de Nique, near Cana mine, G. McPherson 11502 (MO).
Herrera: 18 km W of Las Minas, Hummel 4195 (MO),
4200 (MO). Panama: Cerro Jefe area, near Altos de Pa-
cora, W. D'Arcy A' Sytsma 14746 (MO). Veraguas: above
primero brazo del Bfo Santa Marfa, N of Eseuela Agrfcola
Alto de Piedra, just W' of Santa Ee, S. Knapp A' Dressier
5282A (MO). SOUTH AMERICA. COLOMBIA. Amazon¬
as: mpio. Leticia, Parque Nac. Nat. Amacayacu, Centro
Administrative Mala-mata, A. Hildas et al. 1408 (MO),
1580 (MO). Antioquia: Parque Nac. Nat. has Orqufdeas.
sector Calles, A. Cogollo A Ramtrez 5148 (MO). A. Co-
gollo et id. 55 96 (MO). Chord: mpio. Riosueio, zona de
Uraba, Cerros del Cuchillo, I). Cardenas 269 (MO), 386
(MO), 982 (MO). 1743 (MO), 2106 (MO). Guaviare:
mpio. San Josb del Guaviare, sector de laguna Pavbn II.
lerritorio Nukak, I). Cardenas et al. 7127 (MO). Magda¬
lena: Alto Bfo Buritaca, Alto de Mira, .S'. Madrinon A
Barbosa 320 (MO). Meta: mpio. Mesetas, Inspeceidn de
Policfa Uribe, vereda Brisas del Rfo Duda, ./. Betancur
1278 (MO). Putumayo: mpio. Mocoa, correg. San Anto¬
nio, vereda Alto Campucana, Einca ha Mariposa, J. Be¬
tancur et al. 5205 (MO). Risaralda: mpio. Mistrato, cor¬
reg. Geguadas, zonas de Geguadas y Carrema, ./. /..
Ferndndez A. el al. 9497 (MO). Valle: mpio. Buenaven¬
tura, correg. Cordoba, San Cipriano, Reserva Nat. Bfo Es-
calerete, W. Devin et at. 3967 (MO). 4469 (MO), 5746
Volume 14, Number 4
2004
Taylor
Psychotria subg. Heteropsychotria
497
(MO), 5755 (MO). 5594 (MO). ECUADOR. Carehi: 5 km
from Eita, A. Barfod et al. 41353 (AAU. MO). Esmeral-
ilas: canton Eloy Alfaro, Reserva Ecol6gica Cotacachi-
Caya|>as, Charco Vicente, M. Tirado et al. 364 (MO). Suo-
iuiiImos: canton El Chaco, Rio Oranadillo, campamenlo
de Inecel “Codo Alto," W. Palacios, 5529 (MO). Napo:
Reserva Florfstica El Chuncho, Payamino, Estacion Exp.
INI AP-Napo, C. Cerda 2486 (MO). W. Palacios et al. 1 160
(MO), C. Persson 134 (MO), ./. Zaruma 433 (MO); canton
Arehidona, Parque Nac. Sumaco Napo-Galera, Cordillera
de Galeras, E. Freire & Cerda 545 (MO); Reserva Biol.
Jaturi Sacha, Rfo Napo, B km al E de Misahuallf, C. Cerda
965 (MO), C. Cerda et al. 4622 (MO), I). Neill & Manning
8016 (MO). Pastaza: Rfo Curaray, alrededores de Laguna
Garzayacu, I). Neill & Palacios 6729 (MO). PERU. Ama¬
zonas: Quebrada chigku shinuk, R. Kayap 624 (MO).
Cusco: Acuartambo, Conipata Valley, Rfo Iono Road V
of Patria, R. Foster et al. 10596 (MO). Iluanuco: Leoncio
Prado, Rupa Rupa, al E de Lingo Marfa, cerca al cerro
Queinado, J. Sclwake V. 9935 (MO). Loreto: Maynas.
Yanamono, Explorama Lounst Camp on Rfo Amazonas be¬
tween Indiana and mouth of Rfo Napo, A. Gentry et al.
29038 (MO). Pasco: Oxapampa, V side of Cordillera de
San Matias between Iscozacin and summit, David Smith
2062 (MO). Puno: Sandia, between Rfo Azata-Colorado,
j\ Nunez & Munoz 5315 (MO). San Martin: Lamas, cas-
erfo Bonilla, trail to E of road. Km 75 of Tarapoto-Yuri-
maguas road, S. Knapp A* Mallett 1 132 (MO). I cavali.
Coronel Portillo. Cordillera Azul, Km 64 on Lingo Marfa-
Pucallpa road. I km E of Puente Cholon. K. Young &
Sullivan 686 (MO). BOLIVIA. La Paz: Nor Yungas, 13.7
km NW of San Pedro on road through Incahuara-Mejil-
lones and along trail to 12 de Oetubre, ./. Solomon 9272
(MO).
Psychotria calidicola (.. M. I ay lor. sp. nov.
’ TYPE: C osta Rica. Limdn: Parque Nacional
Tortuguero, sobre el Rfo lortuguero, ca. 3 km
al SO del pueblo, 10°31'N, 83°30'W, 12 Oct.
1988, R. Robles 2100 (holotype, CR; isotype,
MO-03644674). Figure 2A-I).
Haec species a Psychotria eurycarpa inflorescentiae axe
glabro, tubo corollino 13-14 mm longo atque fructu 15-
20 X 12-20 mm distinguitur; quam ea altitudines humil-
iores habitat.
Shrubs or usually small trees In 8 in tall,
branched; stems glabrous. Leaves elliptic to broadly
so, oblanceolate or obovate, 8—20 X (3.0, Osa Pen¬
insula^. 5-12 cm, at base acute to cuneate or
somewhat obtuse, at apex acute to somewhat acu¬
minate, drying papyraceous, on both surfaces gla¬
brous and often shiny; secondary veins 5 to 7 pairs,
looping to inter onnect, without domatia, adaxially
costa and seec alary- veins a little thickened and
remaining venation plane, ahaxially costa promi¬
nent, secondary veins prominulous, anti remaining
venation plane to a little thickened; margins plane;
petioles 1-3.2 cm long; stipules persistent, glabrous,
shortly united around stem, with interpetiolai por¬
tion 0.5— 3.5 mm long, broadly rounded to subtrun¬
cate, entire to denticulate with teeth 2 per side, to
0.2 mm long. Inflorescences terminal, paniculate,
green, glabrous; peduncles 1.5—4 cm long;
branched portion corymbiform. 3-4 X 6-10 cm,
with secondary axes 3 pairs, dichasial, with basal
2 pairs usually closely grouped to subverticillate;
bracts acute, those subtending secondary axes lack¬
ing or usually triangular to linear, 1—2 mm long,
those subtending flowers linear to narrowly ligulate
or triangular, 0.2-1 mm long; flowers distylous, ap¬
parently diurnal, sessile at ends of inflorescence
axes in cymules of 5 to 9; hypanthium turbinate to
ellipsoid, ea. 1 mm long, glabrescent; calyx limb
0. 5-0.8 nun long, glabrous, sinuate to denticulate;
corolla salverform, white, externally puberulous to
glabrescent, internally glabous except puberulous
below middle of tube, tube 13-14 nun long, lobes
5 to 7. narrowly ligulate, 6-8 nun long, ahaxially a
little thickened and rounded; anthers in short-
styled form ca. 3.5 mm long, partially exserted. in
long-styled form ca. 3 mm long, included, situated
near middle of corolla tube; stigmas in short-styled
form ca. 3 mm long, included, situated just above
middle of corolla tube, in long-styled form ca. 2
nun long, exserted. Infructescences similar to inflo¬
rescences; fruits obovoid to ellipsoid, 15—20 X 12—
20 mm, laterally somewhat flattened, glabrescent,
mature color not noted; pyrenes 2, planoconvex,
dorsally with 4 to 5 low obtuse ridges.
Habitat, distribution , and phenology. In wet
forests of Alla itic slopes at 0-800 m, southeastern
Nicaragua to western Panama; collected in flower
February through April, in Iruit January, hehiuary,
and September through December.
This new species has previously been confused
with Psychotria eurycarpa Standley (Taylor, 1991.
2001; Bawa & Beach, 1983; Dwyer, 1980). How¬
ever, Psychotria eurycarpa differs from this new
species in its shorter corolla tubes (11—12 mm
long), generally smaller leaves (5.5—15 X 2.5—7
cm), puberulous inflorescence axes, and smaller
fruits (8-12 mm diam.). Additionally, P eurycarpa
has been collected in flower April through July, and
generally inhabits premontane forests at higher el¬
evations, 700-1500 m. Lhe specific epithet refers
to the habitat of this new species, in warmer forests
at lower elevations, l he floral biology ol P. calidi¬
cola was studied by Bawa and Beach (1983), as
“ Coussarea sp. voucher JHB 1424.
The fruits are described as turning red at ma¬
turity (Robles 2100), which would be highly unusu¬
al in Psychotria subg. Heteropsychotria as a mature
color but not as an intermediate color before the
fruits finally become blue or black, lhe flowers are
reported to be fragrant by several collectors, and
498
Novon
2' 1 iD‘ Psychotria calidicola V1 Taylor; A based on Will 12/08/99/98 ; B, C based on G. Herrera 2597 —
A Stem with leaves and mfructescence. — B. Flower bud. — C. Detail of inflorescence with three old flowers (from
winch corolla has (alien). — -D. Corolla a. anthesis (short-styled form); based on C. Herrera 2597. E, F 'PsyJuZZ
fa " CnSlS ' dy °r‘ ''ltl1 leaves and inflorescence; based on Stein & Hamilton 1011. — F Detail of
u stence showing several flower buds and one bud (before anthesis. with corolla lobes spreading probably as an
a, fact of drying); based on McPherson 11967. -G. Psychotria paradichroa C. M. Taylor stem with leaves and
inflorescence; based on Dressier 3317. A. F. G to 5-cm s. ale; B. C. I) to l-cm scale
Gray um (Grayum 6986) found them particularly
striking: “. . . Flowers emitting an intoxicatingly
sweet, frangipani-like fragrance (1 I’M).”
Paratypes. COSTA RICA. Alajuela: 15 km WNW of
Quesada by air l km W of Jabillos, R. Liesner et al. 15178
(MO). Heredia: N of Puerto Viejo. 12 km to ferry, over
ferry. 6 km along road, N. Garwood et al. 946 (MO); area
between Rio Peje and Rfo Sardinalito, Atlantic slope of
Volcdn Barva, M. Grayum 6986 (MO); Finca Fa Selva, the
Volume 14, Number 4
2004
Taylor
Psychotria subg. Heteropsychotria
499
OTS Biological Station 3 km S of Puerto Viejo, G. Haris-
horn 1135 (MO). 1736 (MO, sterile). B. Loiselle 167 (MO).
Limon: Parque Nac. Tortuguero, K. Barringer et al. 1929
(MO); Estaci6n Cuatro Esquinas, B. Delgado 102 (MO);
Pococf, Estacidn Agua Fria, 3 km al NE siguiendo el cur-
so del Bfo Agua Fria, R. Robles & Flores 1634 (MO);
sendero Agua Fria hasla entrada Sendero Aguacate, ./.
Solano 280 (MO); Hacienda Tape/.co- Hacienda Fa Suerte,
29 km by air W of Tortuguero, C. Davidson 6879 (MO),
6978 (MO); Cordillera de Talamanca, along Quebrada
Cariabral, from Rfo liarbil la to ca. 1.5 km upstream, M.
Grayum et al. 8860 (MO); canton Pococf, cuenca Tortu-
guero-Sierpe, entrada a Finca Bananera Geest. Estacion
Casa Verde, ca. 16 km NE de Cariari, B. Hummel & Gra¬
yum 20745 (MO); Cordillera de Talamanca, Matina, al NE
de la confluencia de Quebrada Canabral con Rfo Barbilla,
Fila Suita, G. Herrera 2172 (MO); canton Limon. El Pro¬
greso, entre Cerro Muchilla y Cerro Avioneta, siguiendo
la Fila Matama, Valle de La Estrella, G. Herrera 2597
(MO); Siquirres, Las Brisas, margen izquierda de quebra¬
da Jesus, eamino a Cerro Tigre, G. Herrera & Valverde
8851 (MO); along river near Experiment Station Los Dia-
montes, GuApiles, R. l^ent 321 (MO); near Rfo Parismina,
8 km W of Dos Bocas, R. Lent 2451 (MO); Reserva Biol.
Hitoy-Cerere, alrededores de la Estacion. Valle de La Es¬
trella, .4. Moreno 23 (MO); Cerro Tortuguero, ladera N\X.
1 km al N de Tortuguero, rumbo a la Barra del Colorado,
R. Robles 2116 (MO). Puntarenas: canton Golfito, Re¬
serva Forestal Golfo Dulce, Aguabuena 3.5 km W of Rin¬
con, K. Thomsen 786 (MO); Aguabuena, sector N, R. Agui¬
lar 684 (MO); Serranfas de Golfito detras de Esquinas
Rainforest Lodge, entre Villa Bricena y Golfito, B. Hum¬
mel et al. 20490 (MO); canton Osa. eamino a la toma de
agua. Rancho Quemado, Rincdn, F. Quesada 305 (MO);
canton Osa, tierras del Grupo de Conservaeion do Rancho
Quemado, Sierpe, J. Mann 226 (MO); La Gamba, Parque
Nacional Esquinas, S. Will 13/08/99/98 (MO). San Jose:
Zona Protectora La Cangreja, Mastatal de Puriscal, J. Mo¬
rales & Jimenez 909 (MO). NICARAGUA. Zelaya: mpio.
Waspan, Reserva Bosawas, sobre el Rfo Waspuk, 300 m
despues del rapido wavla kumbas, luego 0 hasta el cerro
Suyakno Tuna Azan, R. Rueda et al. 4971 (MO); mpio. El
Castillo. Reserva Indio-Mafz, Cerro El Diablo. R. Rueda
et al. 5431 (MO). PANAMA. Bocas del Toro: al 0 del
campamento Corriente Grande (1RHE), M. Correa et al.
3712 (MO, PMA); just above Quebrada Monaco on Chi-
riquf trail. ./. //. Kirkbride & Duke 661 (MO).
Psychotria ehoeoana C. M. Taylor, sp. nov. T\ PE:
Colombia. Choco: Jequedd, 42 km Vi of Las
Animas E of Rfo Palo on Pan American High¬
way (under construction), ca. 250 m, 1 1 Jan.
1979, A. Gentry & E. Renteria A. 23998 (ho-
lotype, HUA- 16395; isotype. MO-2719115).
Figure 3B-D.
Haec species a congeneris Psychotria subgeneris Het-
eropsychotriae stipularum lobulis ligulatis sat grandibus,
infiorescentiae axibus secundariis subverticil latis ffores
sessiles in glomerulos parvos numerosos gerentibus, limbo
calycino extus puberulo glabrescenteve intus dense hir-
tello atque fructuum sat grandium pyrenis dorsaliter lae-
vibus distinguitur.
Shrubs or perhaps small trees flowering at 8 m
tall; stems glabrous, quadrate often becoming
somewhat channeled longitudinally. Leaves elliptic
to broadly so or somewhat oblong, 13-25 X 5-13
cm, at apex rounded or somewhat acuminate, at
base cuneate to obtuse, drying papyraceous to char-
taceous, adaxially and abaxially glabrous; second¬
ary veins 9 to 13 pairs, not or weakly looping to
interconnect, without domatia, adaxially costa and
secondary veins prominulous and remaining vena¬
tion plane, abaxially costa prominent, secondary
veins prominulous, and reticulated higher order ve¬
nation plane to thickened; margins plane; petioles
2.5—4 cm long; stipules persistent, glabrous, united
around stem, sheath subtruncate, ca. 5 mm long,
lobes 2 per side, ligulate to narrowly elliptic, 10-
16 mm long, obtuse to rounded. Inflorescences ter¬
minal. paniculate, puberulous to hirtellous, appar¬
ently green; peduncles 2—7.5 cm long; branched
portion 8-13.5 X 7-13 cm, eorymbiform to broadly
pyramidal; secondary axes 7 to 8 pairs, branched,
pairs often closely grouped to subverticillate; bracts
triangular to ligulate, acute to rounded, those sub¬
tending secondary axes 1 .5—4 mm long, those sub¬
tending flowers 1-2 mm long; flowers sessile in
glomerules of 5 to 15; hypanthium turbinate to cu-
puliform, ca. 0.5 mm long, glabreseent; calyx limb
1-1.5 mm long, externally puberulous to glabres-
cent, internally densely hirtellous, deeply lobed,
lobes broadly rounded to obtuse; corolla tubular-
funnelform, white, externally glabrous, internally
glabrous except densely hirtellous at and just above
stamen insertion, tube ca. 3.5 mm long, ca. 2.5 mm
diam. near middle, lobes 5, narrowly triangular, ca.
3 mm long, acute, abaxially thickened with ap¬
pendage rounded to broadly triangular; anthers ca.
1.5 mm long, partially exserted; stigmas not seen.
Infructescences similar to inflorescences ;fruits ellip¬
soid. ca. 10 X 8 mm, glabrous, mature color not
noted; pyrenes 2, planoconvex, dorsally smooth.
Habitat, distribution, and phenology. In wet
forests at ca. 250 m, northwestern Colombia; col¬
lected in flower in January, in fruit in August.
This new species is distinguished by its stipules
united around the stem into a continuous sheath
with two relatively well-developed lobes (10-16
mm long) on each side, inflorescences with the sec¬
ondary axes often closely grouped to subverticillate,
flowers sessile in numerous small glomerules, calyx
limbs densely hirtellous on the internal (i.e., ad-
axial) surface, and relatively large fruits (ca. 10 X
8 mm) with the pyrenes smooth dorsally. It is so far
known only from Choco Department in northwest¬
ern Colombia, and the specific epithet refers to this
distribution. Many of the plant species with this
500
Novon
M^ure ,5. A. Psychotria soejartoi C. M. Taylor, stem with leaves and infructescence; based on Cogollo 864. B-D.
Psychotria chocoana C. M. Taylor; B based on Arias 65; C, I) based on Gentry & Renteria 23998. — B. Stem with
leaves, stipules, and infructescence. — C. Detail of inlloreseenee with flower bud and three old (lowers (from which
corolla has fallen). —I). Flower at anthesis, partially dissected. E, I. Psychotria osaensis C. M. Taylor; based on Kernan
& Phillips 66 1. — E. Stem with leaves and inflorescence. — F. Detail of inflorescence with three flower buds and one
flower (apparent long-styled form) at anthesis. A, B, E to 5-cm scale; C. I) to same 5-mm scale.
distribution are found from eastern Panama through
coastal Paeific Colombia to northwestern Ecuador,
and / ysychotria chocoana may be more widely dis¬
tributed in tins poorly collected South American
region.
I his new species is similar in general aspect as
well as fruit size and morphology to Psychotria al-
Icnti Standley; P. allenii differs from P. chocoana in
its stipules with deltoid lobes 2—4 mm long, its sec¬
ondary leaf veins 5 to 10 pairs, and its shorter calyx
limbs, ca. 0.3 mm long.
Paratypes. COLOMBIA. Choeo: inpio. Ouibdo, cor-
Volume 14, Number 4
2004
Taylor
Psychotria subg. Heteropsychotria
501
regimiento Tutunendo, carretera a Medellin, /.. Arias 65
(CHOCO, MO); mpio. Bahfa Solano, en la trocha entre
Bahia Solano y el Rio Jua, ,/. Espina et al. 3071 (HUA).
Psychotria gaitalensis C. M. Taylor, sp. nov.
TYPE: 1 ’anama. Code: above El Valle, on trail
to top of Cerro Gaital, 8°40,N, 80°10,W, 1050—
1180 m, 22 Jan. 1988, G. McPherson 1 1967
(holotype, PMA; isotype, MO). Figure 2E, F.
Haec species a congeneris stipulis circum caulem con-
natis vaginam truncatam continuam formantibus, bracteis
(loralibus bene evolutis, limbo calycino profnnde lobato
sed bene evoluto atque tubo corollino 3—4 mm longo dis-
tinguitur.
Shrubs and small trees flowering at 2 m tall, to
3 in tall; sterns glabrous. Leaves narrowly elliptic,
4.5-10 X 1-3 cm, at apex acute to a little acu¬
minate, at base acute to cuneate, drying subcoria-
ceous, glabrous on both surfaces; secondary veins
5 to 7 pairs, not looping to interconnect or doing
so weakly near apex, without domatia, adaxially ve¬
nation plane or costa thickened, abaxially costa
prominent, secondary veins plane to prominulous,
and remaining venation plane and mostly not visi¬
ble; margins plane: petioles 4—9 mm long; stipules
persistent, united around stem, sheatb 1-2 mm
long, truncate, externally glabrous, internally gla¬
brous except at base enclosing a dense ring of col-
leters to 1 mm long. Inflorescences terminal, panic¬
ulate, glabrous, apparently green; peduncles 2.2-4
cm long; branched portion pyramidal, 1 .5—4 X 2—
4 t m; secondary axes 2 to 5 pairs, branched to
several orders; bracts narrowly elliptic to narrowly
ligulate, obtuse to rounded, glabrous, those sub¬
tending secondary axes 3—5 mm long, those sub¬
tending (lowers 1—2 mm long; flowers sessile in di-
chasial cymules of 5 to 7. each usually subtended
by 2 to 4 bracts; hypanthium turbinate, 0.5— 0.8 mm
long, glabrous; calyx limb 0.8—1 mm long, glabrous,
lobed nearly completely, lobes deltoid to ligulate,
obtuse, usually a little unequal on an individual
(lower: corolla tubular, yellow to white sometimes
tinged with purple, externally glabrous, internally
glabrous except densely hirtellous in a ring just
above middle of tube and sparsely pubescent in
tube above this, tube 3-4 mm long, lobes 5, ligu¬
late, 1-1.5 mm long, abaxially a little thickened;
anthers ea. 1 mm long, included, situated just be¬
low corolla throat; stigmas ca. 0.3 mm long, ex-
serted. Infructescences apparently similar to inflo¬
rescences; fruits ellipsoid, ca. 3 X 2.5 mm,
laterally somewhat flattened, glabrous, mature color
not noted; pyrenes 2, planoconvex, dorsally with 4
to 5 narrow longitudinal ridges.
Habitat , distribution , and phenology. In wet
premontane and cloud forests at 900—1400 m in
western Panama; collected in (lower in January and
December, in fruit in January and July.
This new species is distinguished by its combi¬
nation of relatively small leaves, stipules that are
united around the stem into a continuous, unlobed,
truncate sheath, relatively small inflorescences with
the floral bracts well developed, deeply lobed rel¬
atively well developed calyx limbs, and corollas
with tubes 3-4 mm long. The specific epithet refers
to the type locality. In its general aspect Psychotria
gaitalensis is similar to P simiarum Standley; P.
simiarum differs from P. gaitalensis by its smaller
floral bracts, 0.2-1. 2 mm long, that are narrowly
triangular to linear, and its shorter calyx limbs,
0.3-1 nun long, that are only shallowly lobed. The
leaves of P. gaitalensis appear to be rather succu¬
lent in life. The flowers seen of P. gaitalensis are
similar to the long-styled form of distylous Psycho¬
tria species.
Paratypes. PANAMA. Code: slopes and summit of
Cerro Oaital, N of El Valle, S. Knapp et al. 5976 (MO):
divide SW of La Mesa at end of logging road, B. Stein &
Hamilton 1011 (MO).
Psychotria osaensis C. M. Taylor, sp. nov. TYPE:
Costa Rica. Puntarenas: Parque Nacional Cor-
covado, Sirena, Los Patos forest, 8°28'IN,
83°35'W, 1-50 m, 8 July 1989, C. Kernan
1211 (holotype, CR: isotype, MO-4590639).
Figure 3E, F.
Haec species a Psychotria deflexa petiolo 1-3 mm lon¬
go, inflorescentia semper ebracteata atque tubo corollino
2.5— 3.5 mm longo distinguitur.
Shrubs or small trees to 3.5 m tall, branched,
erect; stems puberulous to glabrous. Leaves lance¬
olate, 5-12 X 1-2.8 cm, at apex slenderly acute to
acuminate, at base acute to obtuse, drying papy¬
raceous, adaxially glabrous, abaxially puberulous to
glabrous; secondary veins 5 to 7 pairs, not looping
to interconnect or sometimes doing so near apex,
without domatia, adaxially venation plane or costa
prominulous, abaxially costa and secondary veins
prominulous and tertiary veins plane or sometimes
thickened; petioles 1—3 mm long; stipules persis¬
tent, puberulous, united around stem, sheath 0.3-
0.5 mm long, truncate to concave, lobes 2 per side,
linear to narrowly triangular, 1 .5—3 mm long, acute.
Inflorescences terminal, paniculate, puberulous; pe¬
duncle 2.5—4 cm long, flexuous, slender; branched
portion cylindrical to narrowly pyramidal, 2-4 X
2—3 cm, with secondary axes produced at 2 to 4
nodes, 2 per node, terminating in a 3- to 5(7)-flow-
ered dichasial cyme; bracts none; flowers sessile;
502
Novon
hypanthium cylindrical to cupulifomi, 0.3-0. 5 nun
long, |uil)crulous; calyx limb 0.2-0.3 mm long, pu¬
berulous, sinuate to shallowly lobed; corolla fun-
nelform, white, externally glabrous to puberulous,
internally glabrous except with a hirtellous ring
near middle ol tube, tube ca. 2 mm long, lobes 5,
triangular, ca. I mm long, abaxially smooth; anthers
in apparent long-styled form ca. 0.5 mm long, in¬
cluded, situated in corolla throat; stigmas 2, in ap¬
parent long-styled form linear, ca. 0.3 mm long,
exserted. Infructescences similar to inflorescences,
becoming purple; fruits ellipsoid, ca. 2X3 mm,
laterally a little flattened, puberulous to glabres-
cent, purple; pyrenes 2, planoconvex, dorsally with
3 to 4 obtuse angles and surface irregularly pitted.
Habitat, distribution, and phenology. In wet
forest at 1—500 m on the Osa Peninsula of southern
Costa Rica; collected in flower in July, in fruit in
October.
This new species is similar to Psychotria deflexa
DC., and has been previously combined with it
(e.g.. Burger & Taylor, 1993). However, P deflexa,
which is sympatric (e.g., R. Liesner 3198, MO; I).
Janzen, 10406, MO), differs consistently in its gen¬
erally more robust habit and inflorescences, longer
petioles (3-12 mm long), inflorescences usually
with at least a few bracts, longer corolla tubes (2.5—
3.5 mm long), and larger fruits (2. 5-3. 5 mm long).
Psychotria osaensis is only known from a small re¬
gion on the Osa Peninsula, a phytogeographically
distinct part of Costa Rica; the specific epithet re¬
fers to this distribution. Only a few flowering col¬
lections have been seen, and it is not clear if this
species is distylous. The flowers seen resemble the
long-styled flowers of distylous species.
Paratypes. COSTA RICA. Puntarcnas: Parque Nae.
Corcovado, Los I’atos Forest, C. Kernan <£; Phillips 6 61
(CR, MO); Los Palos, Itfo Sirena Trail, C. Kernan 1331
(OR. MO); trail from Los Chiles to base of hills, R. I.iesner
3199 (MO).
Psychotria paradichroa C. M. Taylor, sp. nov.
TYPE: Panama. I ’anama: Cerro Jefe in Clusia
forest, 2700-3000 ft., 27 Jan. 1966, E. L Ty¬
son, J. Dwyer A: K. Blum 3285 (holotype,
PM A; isotype, MO- 1321251). Figure 2G.
Haeo species a Psychotria dichroa infforescentia fasci-
culata alba cremea vel pallide lutea pedunculo 1—4 cm
longo insidente atque bracteis (loralibus acutis usque ac-
uminatis distinguitur.
Shrubs and small trees flowering at 2 m tall, to
5 m tall; stems glabrous, /waves narrowly elliptic-
oblong to narrowly oblanceolate, 6—25 X 2-6 cm.
at apex acute to usually acuminate, at base acute
to cuneate, drying chartaceous to subcoriaceous,
adaxially glabrous and shiny, abaxially glabrous
and dull: secondary veins 10 to 24 pairs, ascending
to often spreading at 90° or more from costa, usu¬
ally looping to interconnect, without domatia, adax¬
ially costa and secondary veins prominulous and
remaining venation plane, abaxially costa promi¬
nent and remaining venation plane; petioles 3—25
mm long; stipules persistent, glabrous, united
around stem into a continuous sheath or with intra-
petiolar portion reduced, sheath 2-3 mm long, trun¬
cate, lobes 2 per side, triangular to ligulate, 2.5-4
nnn long, obtuse to rounded. Inflorescences termi¬
nal, capitate with heads several and fasciculate,
erect, glabrous, white to cream or pale yellow some¬
times becoming pink to purplish with age; pedun¬
cles 3 or rarely 5, 1-4 cm long, each terminating
in 1 head or sometimes a pair of bracts and 3 axes,
these axes to 2 cm long and each terminating in 1
head, the basalmost bracts ovate, 1-3 cm long,
acute to acuminate; heads 1-2 cm diam., ovoid to
cupuliform, enclosed by 4 involucral bracts in 2
decussate pairs, these involucral bracts lanceolate
to ovate, 1-2.5 cm long, acute to acuminate; floral
bracts lanceolate to triangular, ligulate, or oblan¬
ceolate, 6—10 mm long, acute. Flowers sessile, dis¬
tylous; hypanthium cylindrical, ca. 1 mm long, gla¬
brous; calyx limb ca. 1 mm long, glabrous, sinuate
to denticulate; corolla tubular to tubular-infundib-
uliform. white to usually pink or lavender, exter¬
nally glabrous, internally glabrous except with a
densely hirtellous ring near middle of tube, tube
ca. 12 mm long, lobes 5, narrowly triangular, ca.
4.5 mm long, acute, abaxially smooth; anthers ca.
3 mm long, in long-styled form included, situated
near middle of corolla tube, in short-styled form
exserted; stigmas in long-styled form ca. 0.8 mm
long and exserted, in short-styled form ca. 1.5 mm
long, included, situated near middle of corolla tube,
above its hirtellous ring. Infructescences purple;
fruits obovoid, ca. 7X4 mm, glabrous, deep blue
to purple-black; pyrenes 2, in cross section hemi¬
spherical, dorsally with I to 5 low rounded longi¬
tudinal ridges.
Habitat, distribution , and phenology. In pre-
montane and cloud forests of central and western
Panama at 600—1300 m; collected in flower January
through June, September, and December, in fruit
May through August and October.
I his species has previously been confused with
Psychotria dichroa (Standley) C. M. Taylor by
Dwyer (1980). However, these two species are ap¬
parently allopatric and differ in several morpholog¬
ical features. Notably, in P. paradichroa the inflo-
Volume 14, Number 4
2004
Taylor
Psychotria subg. Heteropsychotria
503
rescences are white to cream or pale yellow
(sometimes becoming pink to purple with age) and
sessile or fasciculate with peduncles 1-4 cm long
and the bracts acute to acuminate, versus in P. di-
chroa respectively always purple to rose, violet, or
magenta, with solitary, entire or tripartite heads
borne on peduncles 3—9 cm long, with the bracts
shortly acuminate to usually obtuse. The type spec¬
imen of P. dichroa is so far known only from the
Chiriquf Highlands of western Panama, while P.
parcidichroa comprises the specimens that have
been included under this name from east of that
region. Specimens of P paradichroa are far more
numerous than those ol P. dichroa.
Paratypes. PANAMA. Code: sawmill 7 km NE of El
Cope, T. Antonio 1139 (MO); vicinity of I, a Mesa, N of el
Valle de Anton, along E edge of Cerro Gaital, /. Croat
67268 (MO); Alto Calvario above El Cope, ca. 6 km N of
El Cope. T. Croat 68751 (MO); Cerro Caracoral, J. Duke
A Dwyer 15103 (MO), ./. H. Kirkbride 1105 (MO); around
Rivera sawmill, Alto Calvario, 7 km N of El Cope, ,/.
Folsom 3231 (MO), ./. Folsom et X 2.5-6 cm, smaller inflorescences with the
branched portion 2^1 X 2-5 cm. and smaller fruits
ca. 6 mm in diameter. Otherwise these plants can¬
not be distinguished and thus the northern group
is here recognized as a subspecies. There does ap¬
pear on herbarium specimens to be some difference
in lengths of anthers and stigmas between the sub¬
species, but few well preserved flowers are avail¬
able of Mexican / * simiarum.
datatypes. MEXICO. Chiapas: mpio. Hcrriozabal. 13
kin N of Berriozahal. I). Hreedlove & Smith 21646 (MO,
NY); mpio. Palenque, along ridges 6-12 km S of Palenque
on road to Ocosingo, / ). Hreedlove 28793 (MO); mpio. Pel-
taleingo, slope of Alik ullial Nab above Petalcingo, D.
Hreedlove 49894 (MO); a 8 km al O de Palenque, cerca
de la cascada Motiepa, E. Cabrera & II. de Cabrera 3949
(MO); en los alrededores de la zona arqueologica de Pal¬
enque, E. Cabrera AL' //. de Cabrera 7017 (MO); mpio.
Ocosingo, Centro Arqueologico Bonampak, ./. Meave et al.
H-507 (MO). Veracruz: mpio. Minatitlan, 13.7 km al E
de La Laguna sobre terracerfa a Uxpanapa, luego 7.1 km
al N sobre camino nuevo a Belisario Dominguez, 7. Wendt
et al. 3484 (MO); 6.5 km al N. T. Wendt et al. 5474 (MO);
mpio. Ilidalgotitlan. Penaseo al NE de Ea Laguna, al E
del Rfo Cuevas y la Estacibn de Luz, T. Wendt et al. 6077
(MO).
Psychotria sinuata C. M. Taylor, sp. nov. TYPE:
Colombia. Valle del Cauca; mpio. Buenaven¬
tura, campamento El Chanco, Proyecto Calima
111 CVC. 4°0()'N, 76°()0'N, 450 m, 15 Mar.
1986, W. Devia I PH) (holotype, TULV; isotype,
MO). Figure IF. (7.
I lace species a congeneris Psychotria subgeneris Het-
eropsvehotriae foliorum nervis secundariis in nervum sin-
uatum submarginalem inanifestum conjunctis marginibus
a I cum plerumque parallelis sinuatisque atque inflores-
cencia laxa sat ampla distinguitur.
Shrubs flowering at 2 m tall, to 4 m tall; stems
puberulous to glabrescent. Leaves elliptic, 14—28 X
4.5—13 cm, at apex acute to usually acuminate with
tips to 12 mm long, at base obtuse to cuneate, dry¬
ing papyraceous, adaxially glabrous, abaxially pu¬
berulous to glabrescent; secondary veins 13 to 18
pairs, uniting in a well-developed undulating sub¬
marginal vein, without domatia, adaxially and abax¬
ially costa prominulous to prominent, secondary
veins prominulous, and remaining venation thick¬
ened to prominulous, subparallel and perpendicu¬
lar to secondary veins; margins thickened, plane,
often sinuate or scalloped in outline parallel to sub¬
marginal veins; petioles 12-20 mm long: stipules
persistent, puberulous to glabrous, united around
stem, sheath 3-6 mm long, truncate, lobes 2 per
side, narrowly triangular to deltoid, 2-4 mm long,
acute. Inflorescences terminal, paniculate, puberu¬
lous to glabrous or sometimes hirtellous in lines or
patches, green, perhaps strongly deflexed; pedun¬
cles 9—14 cm long; branched portion 12-21 X 16-
28 cm, pyramidal; primary axis often flexuous; sec¬
ondary axes 6 to 7 pairs, branched to several or¬
ders; bracts narrowly triangular, acute, those sub¬
tending secondary axes 4—1 I mm long, those
subtending flowers 0.8— 1.2 mm long; flowers ap¬
parently distylous, sessile in dichasial cymules of
5 to 15; hypanthium turbinate, ca. 0.5 mm long,
glabrous; calyx limb ca. 0.3 mm long, glabrous,
deeply lobed, lobes triangular to deltoid, obtuse;
corolla tubular-funnelform, white, externally puber¬
ulous, internally glabrous except hirtellous at and
above stamen insertion, tube 3.5—4 mm long. 0.8-
1 mm diam. near middle, lobes 5, triangular, 1.2—
1.5 mm long, acute, adaxially a little thickened;
anthers in apparent short-styled form ca. 1.2 mm
long, partially exserted; stigmas in apparent long-
styled form ca. 0.2 mm long, exserted. Infructesc-
ences similar to inflorescences or with axes more
spreading; fruits ellipsoid to ovoid, ca. 4 X 4.5 mm.
glabrous, blue-black; pyrenes 2. planoconvex, dor-
sally with 3 to 5 weak acute ridges.
Distribution, habitat, and phenology. In wet
forests at 130—700 m, western Colombia to north¬
western Ecuador; collected in flower in February,
March, and October, in fruit in October.
This new species is distinguished by its leaves
with the secondary veins united in a well-developed
submarginal vein and the margins usually sinuate
to scalloped in parallel to this submarginal vein:
the epithet refers to this margin shape, which is not
known in other species of Psychotria subg. Heter¬
opsychotria. The inflorescences are usually rela¬
tively large and weak to deflexed or even pendu¬
lous. Similarly well-developed, lax inflorescences
506
Novon
are found in P. angustijlora k. Krause of southern
Central America; I1, angustijlora differs from this
new species in its straight leaf margins and inter-
petiolar, shallowly lobed stipules. This new species
is also similar to P. ramiflora Rusby of Amazonian
South America; P. ramiflora differs in its straight
leaf margins and pedicellate flowers. The higher
order leaf venation of this new species often has a
characteristic, very regular organization, which is
similar in general appearance to that seen in some
species of Compsoneura Warburg (Myristicaceae).
Paralypes. COLOMBIA. Chocu: along road between
Pueblo Rico (Risaralda) and Istmina (Chord), along Que-
hrada Antrtn, 15 km W of Santa Cecilia, 6 km W of Cho-
c6-Risaralda border, ca. 20 km K of Playa del Oro. T.
(.root 70897 (MO); mpio. San Josf^ del Palmar, hoya del
Rfo lorito (afluente del Rfo llabita), declive occidental,
finca "los Guaduales," quebrada Santa Fe, afluente del
Rfo Tori to. E. Torero et al. 6782 (MO). Valle del Cauca:
mpio. Buenaventura, correg. Cordoba, vereda San Cipri-
ano, Reserva Natural Rfo Escalerete, in permanent plot,
3°50'N, 76°52'W, W. Devin 56 44 (MO. 'l’ULV), C. M. Tay¬
lor et al. 11975 (COL, JAUM. MO. TULV), 11983 (COL,
J A l M, MO, IT I V). ECUADOR. Carchi: trail through
forest near San Marcos 45 km W of Chical, IL Hoover
1064 (MO).
Psyeliotria soejartoi C. M. Taylor, sp. nov. TYPE:
Colombia. Antioquia: mpio. de Anon, correg.
de I ’rovideneia. Valle del Rfo Anon, entre Dos
Roc as y Anon, zona transitional entre bosque
luimedo y bosque montanoso, Buenos Aires,
primary forest trail, above bridge between
I’rovidencia and Anon, ca. 4 km upriver, 400—
000 m. 6—12 Sep. 1973, I). I). Soejarto, R.
Fonnegra, P. Havens & J. White 1318 (holo-
type, MO -40304 19; isotypes, F- 1809850,
HUA). Figure 3A.
Haec species a Psyeliotria stenostaehya pyrenis duabus.
tlorihus in glomerulos plerumque inllorescentiae ad axem
primarium sessiles dispositis atque lohulis corollinis abax-
ialiter exappendiculatis distinguitur.
Shrubs or small trees to 4 m tall, branched,
erect; stems glabrous, halves elliptic, elliptic-ob¬
long, or elliptic-oblanceolate, 10—23 X 3.8— 10 cm,
at apex acute to usually rather abruptly acuminate
with the acumen 8-15 mm long, at base acute to
obtuse, drying ehartaceous, on both surfaces gla¬
brous; secondary veins 8 to 13 pairs, usually ex¬
tending al least weakly to margins, without domatia,
adaxially venation plane or costa a little sulcate,
abaxially costa prominent, secondary veins promi-
nulous, and remaining venation plane to a little
thickened; petioles 0.8— 3.2 cm long; stipules per¬
sistent, glabrous, united around stem, externally
glabrous, internally glabrous except with a ring of
colletors 1.5-3 mm long al junction of sheath and
stem, sheath I — 4 nun long, lobes 2 per side, 5—18
nun long, deeply 2- to 4-fid (so the stipules appar¬
ently 4- to 8-lobed on each side) with segments
narrowly triangular, unequal in length, acute to
acuminate, often caducous by fragmentation. Inflo¬
rescences terminal, spiciform, congested-cymose,
puberulous to glabrous; peduncles 2-12 mm long;
(lowering portion cylindrical, 3—3.5 X 2 cm, with
5 to 14 glomerules of flowers, these sessile or sub-
sessile on primary axis, subopposite to alternate;
bracts triangular to ligulate, 2-6 mm long, obtuse
to acute; flowers sessile in glomerules of 3 to 5;
hypanthium turbinate, ca. I mm long, glabrous; ca¬
lyx limb 0.8—1 mm long, glabrous, truncate to sin¬
uate; corolla funnelforin, white, externally glabrous,
internally glabrous except hirtellous in upper half
of tube, tube ca. 4 mm long, lobes 5, ligulate, ca.
4 mm long, obtuse, abaxially smooth; anthers ca.
1.5 mm long, exserted on filaments ca. 3.5 mm
long, these inserted just above middle of corolla
tube; stigmas ca. I mm long, included, situated
near middle of corolla tube. Infructescences similar
to inflorescences or to 6.5 X 2.5 cm, usually be¬
coming more elongated and occasionally with sec¬
ondary axes also developing; fruits ellipsoid to ob-
ovoid, 5—5.5 X 6—7 mm, laterally somewhat
flattened, glabrous, mature color unknown; pyrenes
2. planoconvex, dorsally with 4 to 5 obtuse ridges.
Habitat, distribution , and phenology. In wet
forests at 330—900 m in northwestern Colombia and
perhaps eastern Panama (see discussion below);
collected in flower in April and May, in fruit August
through October.
I his new species has previously been confused
in herb, with Psyeliotria stenostaehya Standley,
which it resembles generally in aspect, stipule mor¬
phology, and inflorescence arrangement. However,
P. stenostaehya differs in having usually clearly de¬
veloped secondary inflorescence axes, undivided or
only very shortly fimbriate stipule lobes (the stip¬
ules thus appearing 2-lobed on each side), calyx
limbs that are deeply lobed, and corolla lobes ca.
2 mm long with a linear abaxial appendage 0.5—
1.2 mm long. Ibis new species has been collected
by several botanists but was first documented by
the specimens of 1). I). Soejarto, who established
the herbarium at the University of Antioquia, Co¬
lombia (HUA), and the specific epithet commemo¬
rates him. I lie few flowers seen resemble the short-
styled form of distylous species. The fruits are said
by some collectors to be orange, but the same is
said of several other Psyeliotria species in which
Taylor
Psychotria subg. Heteropsychotria
507
Volume 14, Number 4
2004
the fruits are actually black at maturity, but rarely
seen.
The collection S. Knapp & Mallet 4660 (Panama,
San Bias, trail from Puerto Obaldfa to La Bonga,
tributary of Bfo Armila, ca. 2 hrs. walk from Puerto
Obaldfa, 0-50 m, MO) is similar vegetatively, in
inflorescence size and details, and in calyx mor¬
phology, and this specimen is provisionally includ¬
ed in this new species as the only Panamanian col¬
lection. However, this collection differs from the
Colombian specimens in its corollas in bud with
tubes already ca. 9 mm long, and it may represent
a related but distinct species; the material available
at present is inadequate to evaluate its status.
Paratypes. COLOMBIA. Antioquia: mpio. San Luis,
Parque Fcologico Canon del Bio Claro, sector central,
margen izquierda, A. Cogollo A Borja 749 (J ALM, MO),
A. Cogollo 864 (JAUM, MO); mpio. Anon, correg. Provi-
dencia, valle del Rio Anon, entre Dos Bocas y Anon,
Buenos Aires, 4 km from Providencia, I). Soejarto et al.
4947 (F, MO); vicinity of Planta Providencia, ca. 35 km
SW of Zaragoza, in valley ol Rfo Anon in areas surround¬
ing confluence of Quebrada La lirana and Rio Anon, II.
Waide 62414 (US). Choco: earretera San Jos6 del Palmar-
Novita, alrededores del campamento de Curundo, ca. de
Rfo Ingara, E. Forero et al. 2447 (MO).
Acknowledgments. I thank R. E. Gereau for
preparation of the Latin diagnoses; the curators of
CHOCO, CR, F, HUA, PM A, TULV, and LS for
access to specimens; the National Geographic
Foundation for support for fieldwork in Colombia;
II. Foster, T. Wachter, and the Andrew Mellon
Foundation through a grant to the Field Museum of
Natural History for support for travel to that insti¬
tution; M. Correa and F. Piesschaert for providing
helpful information; and my good colleague W. De-
via for providing information and invaluable collab¬
oration.
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Blake, J. G. & B. A. Loiselle. 1992. Fruits in die diets of
Neotropical migrant birds in Costa Rica. Biotropica
24(2a): 200-210.
Burger, W. C. & C. M. Taylor. 1993. Flora Costaricensis:
Family #202. Rubiaceae. Fieldiana, Bot. n.s. 33: 1 —
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Dwyer, J. I). 1980. Rubiaceae. In: R. F. Woodson (editor).
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Hamilton, C. W. 1989. A revision of Mesoamerican Psy¬
chotria subg. Psychotria (Rubiaceae). Ann. Missouri
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Kirkbride, J. H., Jr. 1997. Manipulus rubiacearum — VI.
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Loiselle, B. A. & J. G. Blake. 1990. Diets of understory
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Malcomber, S. T. 2002. Phylogeny of Gaertnera Lam. (Ru¬
biaceae) based on multiple DNA markers: Evidence of
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verse genus. Evolution 56: 42—57.
Nepokroeff, M., B. Bremer & K. J. Sytsma. 1999. Reor¬
ganization of the genus Psychotria and tribe Psycho-
trieae (Rubiaceae) inferred from ITS and rbc\. sequence
data. Syst. Bot. 24: 5-17.
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Appendix 1
Additional specimens studied of Psychotria buchlienii
(H. Winkler) Standley, arranged by country, then alpha¬
betically. Countries are arranged north to south.
MEXICO: I). Breedlove 56641 (MO); I). Breedlove A Al-
meda 57604 (MO); E. Martinez S. 14500 (MO), 14509
(MO), I5BB7 (MO), 18284 (MO), 18284 (MO).
GUATEMALA: E. Contreras 9946 (MO), 9949 (MO),
10214 (MO), 10308 (MO), 10417 (MO), 11400 (MO); IF
Hannon A Puentes 2064 (MO); IF Kellerman 7470 (US);
C. Landed A Contreras 19488 (MO); E. Martinez .S’. A
Stevens 24424 (MO); E. Martinez S. et al. 14672 (MO); //.
von Tuerckheim 4140 (US).
HONDURAS: D. Aguilar 165 (MO); C. Carets 189 (MO);
/. Montoya 146 (MO); C. Morton 7789 (US); C. Kelson A
Escobar '8581 (MO); V. I, Ochoa 118 (MO); O. Ttllez V. A
Martinez 8443 (MO).
NICARAGUA: A. Grijalva 447 (MO); J. Pipoly 6128
(MO).
COSTA RICA: I). Acevedo 67 (MO); B. Aguilar 2164
(MO); IF Alverson A Flores 3112 (MO); ./. Beach 1420
(MO), 1355 (MO), 1460 (MO); G. Carballo 187 (MO); E.
Castro 294 (MO); /. A. Chacon C. 529 (MO); C. Cordero
17 (MO), 57 (MO); F. Cowan 4587 (MO); T. Croat 46641
(MO), 36740 (MO), 43423 (MO); T. Croat A Grayum
59700 (MO), 59704 (MO), 59885 (MO); T. Croat 67674
(MO); C. Dodge A Thomas 4401 (MO); J. Folsom 8742
(MO). 9252 (MO), 9333 (MO), 9510 (MO), 9521 (MO); G.
508
Novon
Frankie 194 (MO); N. Garwood el al. 1146 (MO); R. God¬
frey 66581 (MO); /.. I). Gomez P. 19435 (MO); R. Gonzalez
II (MO); /. Gutierrez el al. 9 (MO); B. Hummel 7748
(MO), II 160 (MO), 1 1261 (MO). 11736 (MO), 11837
(MO), 12087 (MO), 12352 (MO); li. Ilammel el al. 19391
(MO); G. Herrera 3204 (MO); W. Huber & Weissenlwfer 10
(MO), 9918-4 (MO); G. Induni 153 (MO), 207 (MO, (ml
probably wrong label); A. Jimdnez M. 3691 (MO); G. Ker-
nan 173 (MO); R. Uni 454 (MO); R. Liesner 1797 (MO),
3265 (MO), 14313 (MO); S. Martdn 876 (MO); T. McDow¬
ell & Santana 297 (MO), 299 (MO); T. McDowell 643
(MO); J. Morales 3187 (MO); R. Rueda el al. 1392 (MO);
./■ Solomon 19253 (MO): J. Sperry 605 (MO); C. M. Taylor
& Gereati 9804 (MO); 0. Valverde & Estrada 167 (MO);
S. Will 12/02/99/13 (MO), 25/07/99/69 (MO); /.. (). Wil¬
liams el (d. 24242 (MO).
PANAMA: I Antonio 3104 (MO); ./. Aranda 2862 (MO);
T. Bel iz el al. 3770 (MO), 3845 (MO); 7.’ Croat 13328
(MO), 22650 (MO), 22921 (MO), 23013 (MO), 25946
(MO), 25675 (MO). 27292A (MO), 27457 (MO). 27638
(MO), 27640 (MO); T. Croat & Folsom 34044 (MO); /.’
Croat 37686 (MO), 37703 (MO), 37725 (MO); H. Cuadros
el al. 3851 (MO); G. Davidse & Hamilton 23704 (MO); G.
de Severs et al. 8422 (MO); ./. Duke 4056 (MO). 8748
(MO). 13155 (MO), 13523 (MO); ./. Duke & Lallathin
15001 (MO); J. Dwyer 2223 (MO), 2251 (MO); J. Dwyer
et al. 3461 (MO), 3497 (MO). 4534 (MO); ./. Dwyer 8311
(MO), 8346 (MO); ./. Dwyer & hillathin 8606 (MO). 8620
(MO); ./. Dwyer 8895 (MO); J. Dwyer & Correa 8904 (MO);
8943 (MO); ./. Dwyer & Gentry 9431 (MO); ./. Dwyer
10520 (MO). / 1907 (MO); J. Folsom & Lantz 1882 (MO);
./. holsom et id. 1983 (MO); Folsom & Robinson 2348
(MO); ./. Folsom 4252 (MO); ./. Folsom et