Table Of ContentRHODORA, 240-249, 2003
No.
Vol, 105, 923, pp.
AND
ALUM PILOSUM
A REAPPRAISAL OF PASP
PANICEAE)
PANICOIDEAE:
PEREGRINUM (POACEAE:
P,
Denham and Sandra Aliscioni
Silvia S.
S.
B1642HYD
San
de Botanica Darwinion, Casilla de Correo 22, Isidro,
Institute
Buenos Argentina
Aires,
larwin
mc
un
pnosum peic^umwi mcinucia irm
raspalum ana r. die ui
ABSTRACT,
Decumbentes group of Paspalum. and grow from Mexico and Mesoanierica to Brazil
and Bolivia South America. Paspahim pilosiim is characterized by its erect to
in
decumbent culms, glabrous to pilose sheaths and blades, one or two racemes in
on
terminal inflorescences, glabrous spikelets arranged in pairs in 2 irregular series
dimorphic lower glumes in the pair of spikelets, longitudinally sulcate
the rachis,
gynoecium. Paspalum pcregriniim has sheaths and
lower lemmas, and glabrous
racemes occasionally branched, hairy spikelets arranged in to
blades densely pilose,
1
morphology
4 series on the rachis, and a hairy gynoecium. These modifications in the
Another Thrasya
of P. peregrinum are due to a funga! infection. species,
xenezuelana, known only from the type collection, is the same taxon as P.
We
peregrinum. conclude that P. peregrinum and T. venezuelana are in fact
abnormal specimens of pilosum, whose morphology modified by the fungal
P. is
synonymy
Therefore, both species are reduced to the of/', pilosum. This
infection.
abnormal
caused by fungi, and the first in the tribe Paniceae.
Key Words: Paspalum, Paniceae, Poaceae, abnormal morphology, fungal infection
member
Paspalum pilosum Lam. a of the informal group
is
Decumbentes Paspalum (Chase grows from southern
of L. 1929).
It
Mexico and Mesoamerica Bolivia and southem Brazil (Figure
to
1),
wooded
inhabiting open savannas, and slopes. frequent in
fields, It is
m
grazed places, from 60 to 1700 elevation.
Burman
In a revision of the genus Thrasya Kunth, (1987) excluded T.
xillosa Hitchc. from the genus and placed in Paspalum, in the informal
it
&
group Decumbentes under the new name P.peregrimim A. G. Burm.
Filg., since P. xillosum was pre-occupied by P. villosum Thunb. The
Burman Panama
specimens studied by were from and
central Brazil,
peregrmum
and
the specific epithet refers to the unusual pattern of
by The
disjunction exhibited the specimens. type specimen,
Pittier
from Panama.
5363,
is
During our revision of the Decumbentes group of Paspalum, we
found Thrasya venezuelana Chase same
that the as P. peregrinum.
is
new
The analysis of collections from Venezuela and Brazil, addition
in
240
—
Denham
and PaspaJum
2003] Aliscioni pilosuin 241
of Paspaluin pilosum.
Figure Distribution
1
.
made by Dr. O. Morrone (pers. comm.), led us to
observations
to field
A
pilosum.
consider P. peregn'mim as a doubtful species, similar to P.
rm
and
AND METHODS
MATERIALS
w
and
mexu, mo,
Material studied came from: bm, g, k, lpb, p, si, us,
some
Paspalwn peregn'mim included of the
Specimens of
(Appendix).
Hummel new
Burman 69133 and 5506) and
by {Partch
seen
material
2219
45102) and Brazil {Filgiieiras
from Venezuela {Trejos
material
was
& The 5363, not seen.
Zuloaga 2101). type, Pittier
and FUgueiras
Chase was
The of Thrasya venezuelana. 12407,
type collection
examined.
upper anthecium and
microphotographs of the
Scanning electron
Soderstrom and Zuloaga (1989).
gynoecium were prepared following
A
940 scanning electron
The specimens were viewed in a Zeiss
10-20 kV.
th^ rinrwininn oneratine
r«;^r-^c-/^r^t^^ at institute, at
242 Rhodora
[Vol. 105
The
specific histochemical test for chitin was carried out order
in
to detect fungal hyphae walls using an aqueous 1%
solution of
aniline-
85%
blue and
acid (Clark
lactic et 1983).
al.
RESULTS
The
material determined
40-100
The
culms. sheaths and blades are glabrous The
to scarcely pilose.
terminal inflorescences have 1-2 unbranched racemes 4-15
that are
cm
The
(-20) long. last sheath usually subtends a cymose system of
1 -3 axillary inflorescences, each with a single raceme;
other
axillary
The
glabrous adaxially and
hispid abaxially, the margins
are glabrous
to
mm
The
sparcely
pilose. spikelets are glabrous, 2.2-3.0
long, arranged
two
in pairs, in irregular series on the rachis. The lower glumes
are
when
dimorphic
comparing
this bract in the upper and lower
spikelets
of a upper
pair: in the spikelet the lower glume
generally reduced, up
is
to 1/6 the length of the spikelet, while in the lower
spikelet, the lower
glume
well-developed, up
is to 3/4 the length of
the and
spikelet,
The
The
flower is usually absent. The upper anthecium
papillose and
is glabrous.
The
The
cm
their shorter culms (25-35
sheaths and
tall), blades
that are densely
(3-6
branched
The
arranged
in
The
1
The
pilose or rarely glabrous
(Table
Hairs of
1). the ovary
are unicellular,
90-330
urn long, with rounded
a or dome-shaped
apex and
with
a basal
constriction (Figure
Characters shared by
2). two
the
species
are: the
terminal
inflorescences with 1-2
racemes
and
the axillary ones with
a single raceme;
the dimorphic lower glumes
two
in the
spikelets of
a
pair; the 5-nerved,
longitudinally
sulcate and
finely papillose lower
lemma;
the well-developed lower
palea; and
the absence
of lower
the
flower.
The
specimen
Filgueiras 2219
(mo)
presents
with
tillers the
moiphology
Paspalum
of
neresrimmi
and
with th^ mr.r^v,,.i„„„
tillers
Denham
2003] and AYiscioni^Paspalum
piloswn
243
Table Comparison of Paspalum
pilosum and pere^nnum.
1. P.
Plant
part pilosum
P. peregrinum
P.
Culms
40-100 cm
25-35 cm
tall
tall
Sheaths and
blades Glabrous
to scarcely pilose Densely
pilose
Racemes cm
14-15 (-20) 3-6 cm
long,
long,
never branched
occasionally branched
Rachis of the racemes Adaxial surface glabrous, Both surfaces and
abaxial surface hispid, margins
pilose
and margins glabrous
to scarcely pilose
mm
mm
Spikelets Glabrous, 2.2-3 long
Pilose, 2.5-3.5 long
on
Spikelets the rachis In 2 irregular series In to 4 irregular series
1
Upper
anthecium Glabrous
Pilose toward the apex
Gynoecium
Glabrous
Hairy, rarely glabrous
of pilosum same By
P. in the plant. using a histochemical reaction
we
to detect chitin, verified the presence of fungal hyphae on
the
reproductive structures of the with features of P. peregriminu
tillers
Fungal hyphae were on
also detected reproductive structures of the
rest
of the specimens of P. peregrinum.
known
Thrasya venczuelana only from the type. Chase 12407,
is
from Venezuela. This specimen matches the morphology of Paspalum
peregrimmi, with sheaths, blades, and rachis of the racemes densely
mm
pilose; pilose spikelets, 3.5-3.7 (-4) long; the upper anthecium
papillose and pilose toward the apex; and the gynoecium densely
pilose.
DISCUSSION
The gynoecium
presence of a hairy a frequent feature within the
is
subfamily Pooideae; can be hairy throughout surface or only near
it its
the top, sometimes as a fleshy hairy appendage (Clayton and Renvoize
A
1986). hairy gynoecium present in some species of Festuca L.,
is
Megalachne Steud., and Vulpia C. C. Gmel. of the tribe Poeae; Arena
L., Arrhenatherum P. Beauv., and Gaudinia P. Beauv. of the tribe
Aveneae; and Elymus L. and Agropyron Gaertn. of the tribe Triticeae. In
contrast, a hairy gynoecium unusual within the subfamily Panicoi-
is
Watson
deae. Within the tribe Paniceae, and Dallwitz (1992) cited the
presence of a hairy ovary only for the African genus Chaetopoa C. E.
Hubb., although there no reference of this character in the original
is
description of the genus.
Rhodora
244 105
[Vol.
400
)jm
pm
200
pm
40
Figure Scanning
2. electron microphotographs of Paspalum
pilosum. All
abnormal
2101 a. Upper lemma
anthecium, Upper
(si), side. B. anthecium, palea
side. C,
lemma
ovary
—
Denham
and Paspahim
2003] Aliscioni pilosum 245
Structural modifications on vegetative and reproductive of
parts the
plants, such as culm length, branching vegetative axes and
at
inflorescences, and proliferation and size of the spikelets, have been
cited as malformations caused by fungal infections (Fischer and Holton
Examples
1957). of morphological changes caused by other
fiingi in
documented
grass taxa are well in Ichnanthus P. Beauv. (Sticber 1982),
grov^ing America, and
in tropical Asia, Africa, Australia; Strcptostachys
Desv. (Zuloaga and Soderstrom which grows from Venezuela
1985),
to
Brazil and Paraguay; and Holcus L. (Menezes de Sequeira and Almaraz
2001), native from Eurasia, but introduced in another areas. Another
species of the Decumbentes group oi Paspahim, P. unispicatum (Scribn.
&
Men*.) Nash, which grows from Texas to Argentina, has been reported
by Chase (1929) as being frequently affected by a fungus that distorts
the inflorescense. The discovery of specimens such as Filgiieiras 2219,
which normal and abnormal morphology on
presents with the
tillers
same plant, allows us to conclude that the specimens that Burman (1987)
recognized as P. peregrinum represent affected plants, constituted only
by with abnormal morphology.
the
tillers
In contrast, we have no evidence that pilosity on sheaths, blades, axis,
may Uphof
and bracts of the spikelets be caused by fungal pathogens.
mutations can cause glabrous and
(1962) reported that plausible that
is
it
Paspalum
pubescent varieties of a species. Li her treatment of pilosum,
Chase (unpubl. manuscript) mentioned specimens collected in grazed or
trodden places in Brazil, with short culms, blades densely pilose,
racemes 4-8 cm long, spikelets densely hispid with glistening hairs, and
stamens and gynoecium abnormal. She described these plants as being
affected by a fungus, probably belonging to the genus Balansia, Also.
Even
and normal culms. though
she noticed plants with both affected
pereghmim
the abnormal morphology of the specimens of P. is evi-
we
dently caused by a fungal infection, were unable to identify the
However, White
pathogen due lack of Hving material. (1987),
the
to
Clay and Clay and Leuchtmann (1989) have pointed out that the
(1988),
Poaceae commonly infected by fungal endophytes of the family
are
(Ascomycetes).
Clavicipitaceae
Examples based on infected specimens include Holcus
of '*taxa"
which was synonymized under Holcus
Pam.,
mollis L. var. parvifiorus
by Menezes de Sequeira and Almaraz (2001), and Agrostis
mollis L.
pumila U, which Roemer and Schultes (1817) included as an infra-
Paspahim peregrinum and
With.
category of vulgaris
specific
i4.
new
Thrasya venezuelana represent two incidents of taxa that are based
and
on characters resulting from a fungal infection, the first in the
246 Rhodora
[Vol. 105
tribe Paniceae. The remaining species of the Decumbentes group of
Paspalum, approximately 17 species, have been examined for this
phenomenon and no was
infection detected.
abnormal morphology observed
In short, the in the studied speci-
mens of Paspalum peregrinum and Thrasya venezuelana
in represent
pathological deformations carmot be taxonomic
that treated as differ-
we
ences. For this reason, propose the reduction of P. peregrinum and
synonymy
venezuelana under pilosum.
r. to P.
The
following nomenclatural treatment should be adopted:
Paspalum pilosum Lam., Tabl. Encycl. 175. 1791. Dimorphostachys
1:
pilosa (Lam.) E. Foum., Mexic. PI. 2: 14. 1886. Type: America
Richard
Tropical, (holotype:
s.n, p-la!; isotype:
usi).
Thrasya villosa Hitchc, syn. nov., Proc. Biol. Soc. Wash. 40: 84. 1927.
&
Paspalum peregrinum A. G. Bunn. Acta Venez.
Filg., Rot. 14(4): 91.
1987. Type: panama. Cen-o Vaca, 25-28 Dec
eastern Chiriqui, 1911,
5363
Pittier (holotype: us not seen).
Thrasya
venezuelana Chase, syn. now, Wash. Acad.
J. Sci. 42(4): 122. 1952,
non Paspahim
villosum Thunb., 1784. Paspalum venezuelanuni (Chase)
A. G. Burm., Acta Bot. Venez. 14(4): 90. 1987. Type: Venezuela. Federal
Mar
Chase 12407
District, 11 1940, (holotype: us-1762130!;
isotype:
VEN
not seen^.
We
acknowledgments.
thank Drs O. Morrone and Zuloaga who
F.
suggested
to us the idea of the paper and comments
for their on the
manuscript.
literature
cited
Burman,
A. G. 1987. The
7-93.
Chase, A. 1929. The Non
1-310.
28:
EITE
techniques
for the detection of Acremonium
coenophialum
in fescue and
tall
methods of
in vitro culture of the fungus.
Microbiol. Meth. 149-155.
J.
I:
Clay, K. 1988. Fungal endophytes A
of
grasses: defensive mutualism between
plants
and fungi. Ecology 69: 10-16.
AND A. Leuchtmann.
1989. Infection woodland grasses by endophytes.
Mycologia 805-811.
81:
Graminium
Kew
World.
Bull. Additional Series Her
XIII. Majesty's
Stationery
Office,
London,
U.K.
W.
Fischer, G. and Holton.
C.
S. 1957. Biology and
Control of Smut
the Fungi.
1st
The Ronald Co New
ed. Press Ynrlr
—
Denham
and Paspalum
2003] Aliscioni pilosuni 247
Menezes de Sequeira, M. and Almaraz. Hokus
T. 2001.
mollis L. var. parvijfoms
An
Par. (Poaceae): erroneous interpretation of abnomial morphology caused by
fungi. Bot. Linn. Soc. 135: 295-297.
J.
RoEMER, AND
J. J. A. ScHULTES. 1817. Syst. Veg., ed. 16, Vol. 2. Stuttgardtiae.
A
SoDERSTROM, T. R. AND F. O. ZuLOAGA. 1989. revisiou of the genus Olyra and the
new
segregate genus Parodiolyra (Poaceae: Bambosoideae: Olyreae). Smithso-
nian Contr. Bot. 69: 1-79.
M.
SxrEBER, T. 1982. Revision of Ichnanthifs sect Ichnanthus (Gramineae,
Panicoideae). Syst. Bot. 7: 85-1 15.
Uphof, Band W. Zimmemiann
J. C. 1962. Plant hairs. IV. Teil 5. hi: and P. G. Ozenda,
eds.. Encyclopedia of Plant Anatomy, 2nd ed. Gebriider Bomtraegen Berlin.
Watson, and M.
L. J. Dallwitz. 1992. Grass Genera of the World, 1st ed. C.A.B.
International Publishing, Wallingford, Oxon, U.K.
White, J. F. 1987. The widespread distribution of endophytes in the Poaceae. PI. Dis.
340-342.
71:
Zuloaga, F. O. and T. R. Soderstrom. 1985. Classification of the outlying species
new
of world Panicum (Poaceae: Panrceae). Smithsonian Contr. Bot. 59: 1-60.
APPENDIX
SPECIMENS EXAMINED.
&
urman
Fils. are
with an
identified asterisk.
BELIZE. Stann Creek, Possum Point Biological Station, 23 Jun 1993, Vincent et al.
6184
(mo).
km
BOLIVIA. La Paz: Franz Tamayo, Apolo, 52 hacia Charasani, 1550 m, 7 Jun
&
1990, Beck Foster 18538 (k, lpb, Nor-Yungas: camino Yolosa-Caranavi,
si);
km &
desvio a Coroico, a 9 de Yolosa, Jun 1993, RiigoJo ViUaxicencio 1936
1 (si).
Santa Cruz: Velasco, Parque Nacional Noel Kempff M., Calarala Ahlfeld, 300 m, 24
Apr 1993, Killeen et al. 545J (mexu, mo, si).
SW
brazil. Bahia: 24 km of Belmonte on road to Itapebi, 24 Mar 1974, Hat ley
17387 do Rio de Contas, Harley aL 15330 mo, Mocuge,
et al. (k, mo); Serra et (k, p);
&
Mocuge Mar Zuloaga Morrone 6959
de a Barra de Estiva, 15 1999, (si). Distrito
W
Nov
Federal: of Taguatinga, 1200 m, 25 1965, Invin et al. 10652 (mo); D.F. 205,
W
km
Sobradinho Corregom d'Ouro, 15 de Sobradinho, 22 Feb 1992,
entre y
&
Filgueiras Zuloaga 2101"^ Goias: Padre Bernardo, entre PaWdre Bernardo e
(si).
N km
Curralinho. 20 km de Curralinho, 28 Feb 1992, Filgueiras 22 (mo); 15
ca.
N
of Corumba de Goias on road to Niquelandia, ca. 150 m, 15 Jan 1968, Invin etal
1
km
m, Mar
18610 Chapada dos Veadeiros, 10 south of Cavalcante, 1000 10
(mo);
1969, Irwin etal. 24239 (us); Padre Bernardo, entre Padre Bernardo e Curralinho, ca.
N
20 km de Curralinho, 28 Feb 1992, Filgueiras 2219 (mo); Pirinopolis, Pirineus, 17
km
Feb 1956, without collector 4375 Serra do Pirineus, 9 S of Corumba de
(si);
Goias, 1000 m, Dec 1965, Irwin et al. 10892 (us); without locality, 1894-1895,
1
Glaziou 22434 (c, k, p, w). Minas Gerais: 13 Jan 1951, Pires et al. 2907 (us); ca.l 1
&
km E Campanha Hwy. 267 Caxabu, 960 m, Davidse Ramamoorthy
of along to
10609 (mo); ca. 22 km S of Padre Parafso along Hwy. BR-1 16, 700 m, 29 Mar 1976,
248 Rhodora
[Vol. 105
Davidse et al. 11516 (mo). Para: Belem, 10 Nov 1964, Soderstrom & Canalho 1158
(mo). Parana: Itarare, 20 Jan 1915, Dusen 13507 Jaguariaiva, Parque Estadual do
(si);
Dec
Cerrado, 13 1992, Rugolo et al. 1725 (mo). Rio de Janeiro: Macieiras, Serra de
Itatiaia, 1000 m, 18 Jan 1925, Chose 8346 (us). Sao Paulo: Piracununga, 31 Dec
1964, Clayton 4104 (us); Sao Paulo, Parque Estadual das Pontes do
Ipiranga, 13 Jan
1976, da Silva 235 (mo).
COLOMBIA. Buenaventura: 13 Jun 1923, Hitchcock 19909 Cauca: Around
(us). Calf,
western side of Cauca valley, 1000-1200 m, Dec 1905, 659
Pittier (us); carretera
Madomo
Calipopayan, &
entre y Pescador, 1630 m, 26 Jun 1989, Zuloaga Londono
W
4190 (MO, SI); Cauca Valley, Rio Sucio, of Popayan, 1500-1700 m,
3 Jul 1922,
Pennel 8164 La
et al. (us); Esmeralda near Jamundi, Cauca Valley, 1200 m, Jan
1906, Pittier 1539 (us). Choco: Hoya del Rio San Juan, Andagoya, 12 Apr 1979,
SW
Forero 5117 km
et al. (mo). Cundinamarca: 4 of Fusagusuga, 1450 m, 8 Jan
1974, Davidse 5556 Mesa
et al. (mo). Santander: de Los Santos, 1500 m, 1-15 Dec
1
&
1926, Killip Smith 15112 Without
(us). locality, 1846, Moritz 650
(bm,
g).
COSTARICA. Guanacaste: Parque Nacional Rincon de La Vieja, 900-1200 m, 27-28
Jan 1983, Davidse 23441
et al. (mo). Puntarenas: Buenos 380 m,
Aires, 17 Jul 1966,
&
Pohl Calderon 10075 San km km NE
(mo). Jose: S of Rivas, 6 of San
1 Isidro del
General, 725 m, 14 Jul 1966, Pohl & Calderon 10050
(mexu).
EL SALVADOR. Chalatenango! along Hwy. 4 km SSE of La Palma 950 m
4, Jun
II
&
Pohl
1970, Davidse 11896
(mo).
FRENCH
GUIANA. Cayenne:
1835, Leprieur
s.n. (c,
p).
GUATEMALA. km
Chiquimula:
3 S of Quezaltepeque on CA-10, Dec
8 1970,
Harmon
Dwyer
3690
(mo).
iSc
kmNW
HONDURAS. Comayagua: new
along Hwy.,
10 of Siguatepeque, 1080 m,
14
Aug
1971, Pohl 12762 (mo). Cortes Lago de Yojoa: 60 km SE de San Pedro
Sula' 21
Feb
1982, haguirre 148 (mo). El Paraiso: along Hwy. 25 km E of Zamorano
4, El
600 m, 14 Jun 1970, Pohl & Davidse 11921
Quebrada
(mo); Honda,
por
la carretera
Manzaragua
entre San
y Lucas, 1100 m, 11 Oct 1994, Linares & Metsger 1805
(mexu). Morazan: 800 m, Nov SW
1943, Rodriguez 1561 km
(mexu);
ca. 16 of
Talanga
along the road to Tegucigalpa (Hwy. 750 m, 4 Oct &
3), 1986, Davidse
Pilz
SW
31554 km
(mexu,
mo). Olancho: 6 of Dulce Nombre Hwy
de Culmi, along
3
550 m, 23 Jul 1970, Pohl & Davidse 12443
(mo); Vaguada no
del de Culmf, 500 m'
17-22 &
Nelson Romero
Jul 1978, 4703
(mexu),
MEXICO. Tabasco:
Huimanguillo, 10 Oct Cowan
1980, 3249
et al. (mexu).
NICARAGUA. Comarca
Del Cabo:
Puente Pozo
Azul, 10 Jul 1972, Seymour 5802
(mo). Zelaya: between Waspan
and Ulwas, &
12 Jul 1970, Pohl Davidse 12287
(mo)
PANAMA.
Chiriqui: Alto Boquete, 1125 m, 25
Jan 1969, Partch 69133^
(mo)-
Foothills, vicinity of El Boquete, lOOft-1300 m, 28 Sep-7
Oct
1911, Hitchcock 8192
Panama:
(SI). Altos de Campana, a unos 85 km Moel
del
Sulin, Jun 1977 Mendez
1 1
39 (MO); Cerro Campana, 1000 m, 9 Nov Hammel
1978, 5506^
(mo); Chorrera
16
Sep
1911, Hitchcock 8130 Veraguas,
(si); along road
to radio tower on
Cerro San
W
Cnstobla, near Pam. Am. Hwy., km
9 of bridge over Rio
Cobre, 450 m,
25 Feb
Nee
1974, 10163 (mo). Zona del Canal: Ancon 26 Nov-9 Dec
Hill,
1923, Standley
25205
(mo).
PERU. Ayacucho: Aina, between Huanta and Rio Apurimac m
75(>-10(K) 7-17
May &
1929, Killip Smith 22528
(us).
TRINIDAD Y TOBAGO. Poit of Spain, 26 Nov
1912, Hitchcock 9987
Aripo
(us);
Savanna, Cumuto
northeast of
Village, 18 Jun 1963, Soderstrom 980
Aripo
(us)';
Denham
2003] and AUscioni—Paspalum
pilosum 249
Cumutu
Savanna, Dec
Station, 5 1912, Hitchcock 10075 (us); Joseph, 23 Dec
St.
1912, Hitchcock 10189
(us).
VENEZUELA. Amazonas:
Rio Negro, Solano, along Brazo
the Casiquiare, 75 m, 26
Jun
1984, Dav/6?5^c£M///^/-266J5 (mo). Anzoategui: Bolivar, 100 m, 24 Nov
1981,
1
&
Davidse
Gonzalez 19389
(si). Anzoategui: Freites, Burro trail between San Durrial
and Los 1200-1400 m, Dec &
Pajaritos, 1981, Davidse Gonzalez 19794
(mo,
I
si).
E
Atures: near Capuana, bank of the Rio Orinoco, 90 m, 27 Apr Davidse
1979,
et al.
N
km
16795
(mo). Bolivar: a 4 al de Santa Elena de Uairen, 900 m, 14 Aug 1989,
N
Zuloaga al 4441 Hato La km
et (mo); Roscio, Divina Pastora, approx. 8 de
al
&
Santa Elena de Uairen, 830 m, 25 Jul 1983, Huher Alarcon 7831
(mo); Sifontes,
W
km
sector La Hoyada, 7 al de Sta. Elena de Uairen, 850^1 100 m, 23 Oct 1986,
Aymard 4797
(mo). Distrito Federal: El Avila, 1600 m, Oct 1959, Trejos 45102''
km
(mo). Guarico: Estacion Bioldgica Los Llanos, 10 SSE of Calabozo, 60 m, 9 Nov
Monagas
1661
(mo).
