Flora Anatomy Alliaceae

A revision of Asterogyne (Arecaceae:
Arecoideae: Geonomeae)
FRED W. STAUFFER, CONNY B. ASMUSSEN,
ANDREW HENDERSON, AND PETER K. ENDRESS
Stauffer, F. W. (Institute of Systematic Botany, University of Zurich, Zolliker-

strasse 107, CH-8008, Zurich, Switzerland; email: f.stauffer@access.unizh.ch), C.
B. Asmussen (Department of Ecology, Royal Veterinary and Agricultural Univer-
sity, Rolighedsvej 21, DK-1958, Frederiksberg C, Denmark; email: con@kvl.dk),
A. Henderson (Institute of Systematic Botany, The New York Botanical Garden,
Bronx, NY 10458-5126, U.S.A.; email: ahenderson@nybg.org) & P. K. Endress
(Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008,
Zurich, Switzerland; email: pendress@systbot.unizh.ch). A revision of Asterogyne
(Arecaceae: Arecoideae: Geonomeae). Brittonia 55: 326–356. 2003.—A taxonomic
revision of the neotropical genus Asterogyne Hook. (Arecaceae: Arecoideae: Geon-
omeae) is presented. The genus is characterized by one autoapomorphic character,
the separation of anther thecae by a bifid connective, and the combination of small
to medium-sized stems, bifid simple leaf blades, floral pits in bud covered by a
rounded upper lip that overlaps a lower lip, and pistillate flowers with staminodial
lobes that are displayed in a starlike shape. Five species are recognized in this
treatment: one of them (Asterogyne martiana) is widely distributed from Belize in
Central America to northern Ecuador, three species (A. ramosa, A. spicata, and A.
yaracuyense) are endemic to small areas in the Venezuelan Coastal Range, and one
species (A. guianensis) is endemic to eastern French Guiana. The taxonomic history,
morphology, reproductive biology, distribution and ecology, intrageneric relation-
ships, and conservation status are reviewed.
Key words: Arecaceae, Palmae, Geonomeae, Asterogyne, Neotropics.
Stauffer, F. W. (Institute of Systematic Botany, University of Zurich, Zolliker-

strasse 107, CH-8008, Zurich, Switzerland; email: f.stauffer@access.unizh.ch), C.
B. Asmussen (Department of Ecology, Royal Veterinary and Agricultural Uni-
versity, Rolighedsvej 21, DK-1958, Frederiksberg C, Denmark; email:
con@kvl.dk), A. Henderson (Institute of Systematic Botany, The New York Bo-
tanical Garden, Bronx, NY 10458-5126, U.S.A.; email: ahenderson@nybg.org) &
P. K. Endress (Institute of Systematic Botany, University of Zurich, Zollikerstrasse
107, CH-8008, Zurich, Switzerland; email: pendress@systbot.unizh.ch). A revi-
sion of Asterogyne (Arecaceae: Arecoideae: Geonomeae). Brittonia 55: 326–356.
2003.—Se presenta una revisión taxonómica del género neotropical Asterogyne
Hook. (Arecaceae: Arecoideae: Geonomeae). El género se caracteriza por un car-
ácter autoapomórfico, tecas separadas por un conectivo bı́fido, además de tener
tallos de pequeño a mediano tamaño, láminas foliares enteras y bı́fidas, fóveas
florales cubiertas por un labio superior redondeado que sobrelapa el labio inferior,
flores pistiladas con lóbulos estaminodiales dispuestos de manera estrellada. Para
este tratamiento se reconocen cinco especies: una de ellas (Asterogyne martiana)
ampliamente distribuida desde Belice en América Central hasta el norte de Ec-
uador, tres especies (A. ramosa, A. spicata, and A. yaracuyense) endémicas de
pequeñas áreas en la Cordillera de la Costa de Venezuela y una especie (A. gui-
anensis) endémica del oriente de la Guyana Francesa. Se revisa la historia tax-
onómica, morfologı́a, biologı́a reproductiva, distribución, ecologı́a, relaciones in-
tragenéricas y estado de conservación.
Brittonia, 55(4), 2003, pp. 326–356. ISSUED: 07 October 2003

q 2003, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) 327
FIG. 1. Type specimens of Asterogyne. A, B. Isotype of A. guianensis (de Granville 7124, K); C. Isotype
of A. martiana (Wendland 56, K); D–F. Isotype of A. ramosa (Steyermark et al. 96088, VEN). Scale bars:
8 cm.
The small genus Asterogyne H. Wendl. already known in other genera of Geono-
ex Hook. f. is one of six genera in the neo- meae, e.g., bifid simple leaf blades and sta-
tropical tribe Geonomeae (Dransfield & minodia basally united into a tube. Al-
Uhl, 1998; Henderson et al., 1995) and though modern treatments of the genus
consists of five species in Central America have proposed several characters to differ-
and northern South America (Figs. 1A–F, entiate Asterogyne from other genera of
2A–F, 4). In a letter to J. D. Hooker in Geonomeae, floral pits in bud covered by
1883, Wendland proposed Asterogyne as a a rounded upper lip that overlaps the lower
new genus to be included in the monu- lip (Fig. 9D) and free thecae on a bifid
mental treatment of Genera Plantarum connective (Figs. 8B, 10D, 12C, 14C),
(Bentham & Hooker, 1883). The genus have always been regarded as the two most
was described with several character states relevant features (Burret, 1930; Hen-
328 BRITTONIA [VOL. 55
FIG. 2. Type specimens of Asterogyne. A–C. Isotype of A. spicata (J. Steyermark 90010, VEN); D–F.
Holotype of A. yaracuyense (Steyermark et al. 100311, VEN). Scale bars: 8 cm.
derson et al., 1995; Moore, 1966; Uhl & though there was no original explanation of
Dransfield, 1987; Wessels Boer, 1968). the generic name, it was probably suggested
Asterogyne was published without a type because of the starlike shape of the stami-
species in the original description in 1883. nodial lobes of the pistillate flowers
The typification of the new genus was made (Moore, 1978) (Figs. 7F, 8D). Moore (1966)
two years later in 1885 by Wendland, with described the genus Aristeyera, separating
the transfer of Geonoma martiana to the ge- it from Asterogyne on the basis of differ-
nus Asterogyne. The type specimen selected ences in the number of stamens and stam-
by H. Wendland for this taxon was obtained inodes. Wessels Boer (1968) observed great
from a plant cultivated at the gardens of plasticity of these characters, especially in
Herrenhausen (Fig. 1C). This plant at Her- flowers of Aristeyera spicata, and consid-
renhausen originated from plants cultivated ered that they were inconsistent for a delim-
by L. van Houtte in Gent, apparently from itation at the generic level. He merged both
seeds sent by Warscewicz during his trip to genera and argued that the number of sta-
Central America (Wendland, 1856). Al- mens and staminodes reported by Moore
FIG. 3. Pollen of Asterogyne. A. Pollen grain of A. martiana. B. Detail of exine in A. martiana. C. Pollen
grain of A. ramosa. D. Detail of exine in A. ramosa. E. Pollen grain of A. spicata. F. Detail of exine in A.
spicata. G. Pollen grain of A. yaracuyense. H. Detail of exine in A. yaracuyense. Scale bars: A, C, G 5 5 mm;
B, D, F, H 5 1 mm; E 5 6 mm.
FIG. 4. Distribution of Asterogyne in Central and South America.
FIG. 5. Strict consensus tree of the two most parsimonious trees based on a phylogenetic analysis of mor-
phological data. Bootstrap values are given on the branches.
(1967) for Asterogyne ramosa filled the gap scription of the widespread type species, A.
between the two genera. Due to the vari- martiana (Fig. 4), in 1856, four additional
ability of stamens and staminodes of Aster- taxa were added to the genus within a rel-
ogyne, Wessels Boer (1968, 1988) did not atively short period, from 1967 to 1988.
recognize the two populations in the states These recently discovered species show re-
of Sucre and Yaracuy of the Venezuelan markably limited distributions for the neo-
Coastal Cordillera (Fig. 4) as representing tropical palms, and compared to A. marti-
two different species, but included them in ana, they are known from few herbarium
a broadly defined A. ramosa. After the de- specimens (limited to two for A. guianen-
critical-point dried, and sputter-coated with

gold. Micrographs were obtained with a
Cambridge S4 scanning electron micro-
scope.
For anatomical investigations, flowers in
bud or at anthesis were dehydrated and em-
bedded in Kulzer’s Technovit 7100 (2-hy-
droxyethyl methacrylate, HEMA). The ma-
terial was sectioned at 7 microns using a
rotary microtome, then stained with ruthe-
nium red and toluidine blue and embedded
in Histomount.
A preliminary phylogenetic analysis of
all species of Asterogyne was made for this
revision. Asterogyne was recently shown to
be sister to a clade formed by Geonoma and
Calyptronoma (Asmussen & Chase, 2001),
therefore, we have included Geonoma in-
terrupta and Calyptronoma rivalis as out-
groups. Nineteen morphological characters
(Table I), 10 of them parsimony-uninfor-
mative, were included in the data matrix
(Table II) constructed in McClade 4.0b10
FIG. 6. Habit of Asterogyne guianesis. (Photo: J. J. (Maddison and Maddison, 2000). Clear
de Granville, Camopi River, French Guiana). Scale gaps between the character states of quan-
bar: 80 cm.
titative features were observed. A parsi-
mony analysis was performed using Paup
sis); thus their morphological and anatom- 4.0 (Swofford, 1998). All the characters
ical features are incompletely known. This were treated as ‘‘unordered’’ and were
revision aims to improve the taxonomic equally weighted. Support for the clades
knowledge of the genus and to compile and was calculated by doing a 10,000 replicate
reevaluate the published information on its bootstrap analysis using the branch and
systematics and natural history. bound option.
Morphology and Anatomy

Materials and Methods
STEMS AND ROOTS
Morphology and distribution of the spe-
cies were studied based on 334 specimens Stems of Asterogyne are solitary and
(see Exsiccatae) deposited at AAU, B, BH, mostly erect; only A. martiana and A. yar-
BM, C, CAY, CR, G, K, MER, MO, MYF, acuyense may be slightly decumbent at the
NY, P, U, US, VEN, WIS, and Z. A review base. Basal vegetative branching was de-
of critical literature for the genus (Burret, scribed by Henderson and Steyermark
1930; de Granville & Henderson, 1988; (1986) for A. yaracuyense but the branches
Henderson et al., 1995; Uhl & Dransfield, do not fully develop into mature stems. Lat-
1987; Wessels Boer, 1968) was made. Ad- eral vegetative branches are normally pre-
ditional information on the distribution of sent in A. yaracuyense and are especially
A. martiana was obtained from the curators numerous in A. ramosa, in which 5 to 30
at BRH, PMA, and SCZ, and of A. gui- branches were observed, attached to the
anensis from J. J. de Granville (CAY). stem, 20–40 cm below the crownshaft (Fig.
Specimens were collected in Venezuela by 9A). Each of these lateral branches had 4
the first author in 1996, 1997, 2001, and to 7 small bifid simple leaf blades, and were
2002. For scanning electron microscopy, richly rooted. Our observations showed that
the dissected specimens were dehydrated, these lateral branches do not develop into
FIG. 7. Habit, inflorescence, flowers, and fruits of Asterogyne martiana. A, B. Habit. C. Inflorescence. D.
Detail of rachilla at staminate anthesis. E & F. Detail of rachilla at pistillate anthesis. G. Rachilla with fruits.
(Photos: R. Schmid, La Selva, Costa Rica). Scale bars: A, B 5 1.2 m; C 5 5 cm; D 5 0.4 cm; E, F, G 5
0.3 cm.
new plants. Stem anatomy remains un- The inner cortex is usually homogeneous,
known for the genus. and the aerenchyma, only weakly devel-
Adventitious roots that form a basal cone oped, may occur in the center. In contrast
are present in several species. In Asterogyne to all other genera of the Arecoideae, the
guianensis, the aerial roots are remarkably cells of the inner cortex of the Geonomeae
long, to 90 cm, and contain well-developed are moderately thick-walled, and thin-
white pneumatodes (Fig. 6). Seubert (1998) walled cells are lacking. Narrow fibers are
studied the root anatomy of A. martiana scattered throughout the inner cortex. En-
and found that it is similar to that of other dodermis cells are tanniferous, radially
Geonomeae. The rhizodermis cells are thin- stretched, and u-shaped, with thin outer
walled and disappear in mature roots. In walls and thick inner walls. The pericycle
transverse section, the exodermis and the is one-layered and the vascular cylinder
outer cortex, which is reduced to a few lay- shows the normal pattern observed in the
ers, are not distinct. In longitudinal section, family (Seubert, 1998).
the exodermis and outer cortex are distinct.
The cells of the one-layered exodermis are LEAVES
stretched with blunt ends; those of the outer The leaves, at least the young ones, in
cortex are long stretched with pointed ends. Asterogyne are spirally arranged and rather
FIG. 8. Flowers in Asterogyne martiana. A. Staminate flower. B. Stamen. C. Epidermis of the filament. D.
Pistillate flower. E. Ovary in abaxial view. F. Ovary in adaxial view. G. Protuberances at the base of the styles.
H. Stigmatic branches at anthesis. I. Cross section of the trilocular ovary, arrow points to the septal nectary. J.
Cross section of the secretory epithelium of the nectary. (From Knudsen & Asmussen 604, AAU). Scale bars:
A, D, H 5 1 mm; B 5 300 mm; C 5 25 mm; E, F, I 5 200 mm; G 5 12 mm; J 5 50 mm.
FIG. 9. Habit, leaf, and inflorescence of Asterogyne ramosa. A. Habit. B. Detail of blade. C. Inflorescence
at anthesis. D. Detail of rachillae in bud. E. Detail of rachilla at pistillate anthesis. F. Detail of rachilla at
staminate anthesis. (Photos: F. J. Stauffer, Cerro Humo, Venezuela). Scale bars: A 5 28 cm; B 5 18 cm; C 5
5 cm; D 5 7 mm; E 5 2 mm; F 5 3 mm.
erect and often purplish. The blade is nor- a colourless hypodermal cell layer) only on
mally bifid and simple but it may be lon- the abaxial side. The mesophyll lacks a dis-
gitudinally split with age or damaged by the tinct palisade layer and the cells are some-
wind (Figs. 6, 7A). Longitudinal splits were what transversely extended. The fibers are
especially conspicuous in some collections numerous, often solitary, each with a rather
of A. martiana (Stevens 13505; Quesada wide lumen, scattered throughout the me-
43) from Costa Rica, and A. ramosa (Stey- sophyll but commonly found in the adaxial
ermark & Liesner 120915) and A. yaracu- layer. Some of the vascular bundles are
yense (Stauffer et al. 267) from Venezuela. very small, not producing ribbing of the
According to Wessels Boer (1968) the lamina. Wessels Boer (1968) noted that the
blades of A. martiana and A. spicata show leaf anatomy of Asterogyne shared most of
a hypodermis (sensu Tomlinson 1961, i.e., the characters already observed in other
FIG. 10. Flowers of Asterogyne ramosa. A. Staminate flower in bud. B. Staminate flower at anthesis. C.
Pistillate flower in young bud. D. Stamen. E. Ovary in abaxial view. F. Gynoecium shortly before anthesis. G.
Epidermis of the filament. H. Longitudinal section of the ovule, arrow points to the micropyle. I. Cross section
of the trilocular ovary. J. Cross section of the style and the staminodes, arrow points to the raphide idioblasts
in the mesophyll of the staminodes. K. Cross section of the style showing the ventral slits. (From Stauffer et
al. 824, Z). Scale bars: A, D 5 450 mm; B, C 5 1 mm; E, H–K 5 400 mm; F 5 500 mm; G 5 25 mm.
FIG. 11. Habit, leaf, inflorescence, and fruits of Asterogyne spicata. A. Habit. B. Detail of blade. C. Inflo-
rescence and infructescence. D. Detail of rachilla at staminate anthesis. E. Detail of rachilla with fruits. (Photos:
F. J. Stauffer, Parque Nacional Guatopo, Venezuela). Scale bars: A 5 1.2 m; B 5 20 cm; C 5 8 cm; D, E 5
4 mm.
Geonomeae, but pointed out that the one- 7A, 9A, 11A). Once-branched inflorescenc-
cell-layered abaxial hypodermis of the es are present in A. martiana, A. ramosa,
blades could be interpreted as less devel- and A. yaracuyense (Figs. 7C, 9C, 13B),
oped within the tribe. Vegetative anatomy while A. guianensis and A. spicata show
of A. spicata (as Aristeyera spicata) was well-developed spikes, which are thyrses in
studied by Tomlinson (1966). His obser- a comparative morphological sense (Figs.
vation of two wide vessels per bundle in the 1B, 2C, 11C). A remarkable deviation from
petiole of A. spicata was also found in other the branched inflorescences was observed
species of Geonomeae by Klotz (1978). in some collections of A. martiana restrict-
ed to the Comarca of San Blas in Panama
INFLORESCENCES AND FLORAL TRIADS (e.g., Elias 1769; de Nevers et al. 5209,
Inflorescences of Asterogyne are always 5881, 7914; McDonagh, J. et al. 246) and
interfoliar, solitary, and protandrous (Figs. Colombia (Duke 11338), in which the in-
FIG. 12. Flowers in Asterogyne spicata. A. Staminate flower in late bud. B. Staminate flower at anthesis. C.
Stamen. D. Epidermis of the filament. E. Pistillode. F. Pistillate flower in bud showing staminodes and tip of
the gynoecium. G. Gynoecium. H. Ovary in abaxial view. I. Ovary in adaxial view, arrow points to one of the
openings of the septal nectary. J. Stigmatic branches shortly before anthesis. K. Staminodes, arrows point to
possible secretory slits. L. Cross section of the trilocular ovary. M. Cross section of the style and the staminodes.
N. Cross section of the stigmatic branches before anthesis. (From Stauffer & Stauffer 822, Z). Scale bars: A, E
5 1 mm; B 5 3 mm; C 5 500 mm; D, K 5 50 mm; F, G 5 1.5 mm; H 5 600 mm; I, J, L–N 5 400 mm.
FIG. 13. Habit and inflorescence of Asterogyne yaracuyense. A. Habit. B. Inflorescence. C. Detail of rachilla
at staminate anthesis. (Photos: F. J. Stauffer, Cerro Humo, Venezuela). Scale bars: A 5 30 cm; B 5 8 cm; C 5
3 mm.
florescences are spicate (Fig. 7B). In one of show a reduction to only one. Moore (1967)
these collections (de Nevers et al. 5881) the suggested that the reduction in the number
spike is rather thick. of bracts observed in A. spicata (as Aris-
Morphology and anatomy of the inflores- teyera spicata) represented a derived char-
cence axis and especially the floral vascu- acter within the genus. As first reported by
lature of Asterogyne spicata (as Aristeyera Wessels Boer (1968) for A. martiana, all
spicata) was examined by Uhl (1966). The the species studied in the field showed that
vascular bundles of the flowers of this spe- the small unexpanded inflorescence remains
cies are small because the xylem does not straight and unfolded within closed bracts;
contain large vessels and has a few helical during expansion it elongates rapidly and
elements. The number of vascular traces to emerges through a terminal split of the per-
perianth parts is variable. In the petal tube sistent bracts. The inflorescences are always
of staminate flowers and sepals of pistillate erect at anthesis and become pendent in
flowers, some of the traces appear to dif- fruit (Fig. 7B). In branched inflorescences
ferentiate over a long period of time and as the rachis is normally well developed, but
a result the number of traces increases with in A. martiana (Cuatrecasas 17375; Knapp
the age of the flower. The traces developing 3777; Moore 6530) it may be almost absent
later do not become connected to the vas- and then all the rachillae appear to emerge
cular strands in the floral receptacles. from the apex of the peduncle. The inflo-
The peduncle of Asterogyne is long and rescence raches are normally salmon to
slender. The prophyll is fibrous to membra- pink at anthesis and become brown to red-
nous and normally inserted at the base of dish brown in fruit. Floral pits are spirally
the peduncle or just above it. The inflores- arranged on the rachillae. As described by
cences are enclosed by one or two tubular, Uhl and Dransfield (1987) for Welfia, Ca-
chartaceous to membranous, peduncular lyptrogyne, and Pholidostachys, the floral
bracts. Asterogyne ramosa and A. yaracu- pits in bud of Asterogyne are covered by an
yense always have two peduncular bracts, immersed, rounded, lower lip, overlapped
whereas the species with spicate inflores- by a short upper lip (Fig. 9D). Wessels Boer
cences and A. martiana (with the exception (1968) interpreted the immersed lower lips
of one collection from Panama [Pittier as less specialized in a morphological sense
4226], which has two peduncular bracts), and therefore as phylogenetically basal
FIG. 14. Flowers of Asterogyne yaracuyense. A. Staminate flower in late bud. B. Staminate flower at anthesis.
C. Detail of stamen. D. Epidermis of the filament. E. Pistillate flower in young bud. F. Gynoecium in late bud.
G. Abaxial view of the ovary. H. Adaxial view of the ovary. I. Stigmatic branches before anthesis. J. Cross
section of the trilocular ovary. (From Stauffer & Stauffer 823, Z). Scale bars: A 5 900 mm; B, E 5 1 mm; C
5 400 mm; D 5 50 mm; F 5 450 mm ; G, H, J 5 200 mm; I 5 300 mm.
within the tribe. The lower lip rolls back cinnus. Based on the vasculature, however,
when the flowers become exserted. The un- Uhl (1966) suggested that the triads in As-
common spicate inflorescence observed in terogyne (as Aristeyera) originated from a
one collection of A. martiana (de Nevers et more ramified type of partial inflorescence.
al. 5881) also has the lip margins densely As also reported by Stauffer et al. (2002)
covered by short whitish hairs. The hairs for Geonoma interrupta, the first staminate
are lacking in the normal-branched inflo- flower of the cincinnus, and the cincinnus
rescences on other plants of this species. As as a whole, is subtended by the pit-closing
is common in the Arecoideae, the pits con- bract, which represents the lower lip of the
tain triads composed of one pistillate and pit. According to the interpretation of Wes-
two staminate flowers. Each triad is sunken sels Boer (1968), the largest bract inside the
in the floral pit and represents a short cin- pit subtends the second staminate flower, a
TABLE I
CHARACTERS AND UNORDERED CHARACTER STATES
1. Leaf blade: (0) entire-bifid; (1) divided

2. Inflorescence branching: (0) branched; (1) spicate
3. Upper lip of the pit: (0) absent; (1) present
4. Peduncle (length): (0) #80 cm; (1) $85 cm
5. Peduncular bracts (number): (0) one; (1) two
6. First peduncular bract (length): (0) #60 cm; (1) $64 cm
7. Number of stamens: (0) 6; (1) 8–10; (2) 11–12; (3) 21–27
8. Filament color: (0) white; (1) yellow
9. Filament surface: (0) smooth; (1) papillate
10. Anther shape: (0) divided; (1) with a bifid connective; (2) sagittate
11. Complete fusion of the staminodial tube with the corolla: (0) absent; (1) present
12. Upper part of the sterile androecium: (0) calyptrate; (1) lobed; (2) crenulate
13. Complete fusion of the petals of the pistillate flowers in a tube: (0) absent; (1) present
14. Gynoecium: (0) with 3 more or less equally developed ovaries; (1) pseudomonomerous
15. Central protrusion between the three carpels: (0) absent; (1) present
16. Position of the ovule: (0) at midlength of the locule; (1) basal
17. Lateral vascular bundles in the styles at anthesis: (0) absent; (1) present
18. Upper part of the style curved: (0) absent; (1) present
19. Fruit length: (0) 0.3–7 cm; (1) 1.2 cm; (2) 1.5–1.8 cm; (3) 2.5 cm
medium-sized bract subtends the pistillate 10B, 12B, 14A). Anthers of A. yaracu-
flower, and a small bract represents the pro- yense, and especially A. martiana, are ver-
phyll of the pistillate flower. satile because of the thinner apical regions
of the filaments (Fig. 8B). Wessels Boer
STAMINATE FLOWERS (1968) found that the junction between the
filament and the connective in Asterogyne
Two staminate flowers occur laterally to
is less rigid than in Geonoma. The separa-
the pistillate flower. The perianth is com-
tion of anther thecae on a bifid connective
posed of three basally united and apically
in Asterogyne (Figs. 8B, 10D, 12C, 14C)
free sepals and three basally united and api-
has been considered unique within the
cally valvate petals (Figs. 8A, 10B, 12B,
Geonomeae (Uhl & Dransfield, 1987). The
14B). The number of stamens ranges from
connective is conspicuously swollen in A.
six (Asterogyne martiana, Fig. 8A) to 27
martiana. The anthers are inflexed toward
(A. guianensis); the higher number of sta-
the center of the flower and introrse in bud
mens being present in species with spicate
but spread and extrorse at anthesis. A three-
inflorescences. Taxa of Geonomeae with
lobed pistillode is normally fused with the
unbranched inflorescences and flowers with
staminodial tube to its middle (Fig. 12E).
more than six stamens are rather uncom-
Morphology and vasculature of the stami-
mon. They are present in only two species
nate flowers of A. spicata (as Aristeyera
of Asterogyne (A. guianensis and A. spica-
spicata) were studied by Uhl (1966) and
ta; Fig. 12B) and three species of Geonoma
Moore (1966), and A. martiana by Uhl and
(G. camana Trail, G. chococola Wess. Boer,
Moore (1977).
and G. polyandra Skov). Wessels Boer
(1968) considered the increased number of
POLLEN MORPHOLOGY
stamens in some species of Asterogyne (as
Aristeyera) as a derived character state Pollen in Asterogyne has been described
within the tribe. Based on the increased by Uhl and Dransfield (1987) as elliptic,
number of stamens in A. spicata, Moore monosulcate, with tectate exine, which is
(1966) suggested a closer relationship be- vermiculate or reticulate. In a detailed study
tween Asterogyne (as Aristeyera) and Wel- of the Geonomeae, Punt and Wessels Boer
fia, than with the other Geonomeae. The (1966) stated that Asterogyne species show
stamens are united into a tube up to mid- two different pollen types, the ‘‘Asterogyne
length and free near the top (Figs. 8A, 10A, type,’’ occurring in A. martiana and the
TABLE II
MATRIX OF 19 MORPHOLOGICAL CHARACTERS SCORED FOR THE FIVE SPECIES OF Asterogyne AND TWO OUTGROUP
TAXA (Geonoma interrupta AND Calyptonoma rivalis). THE QUESTION MARK (?) INDICATES DATA NOT AVAILABLE
Character
Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19
A. guianensis 0 1 1 0 0 0 3 0 0 1 1 1 0 0 ? 0 ? 0 3
A. martiana 0 0 1 0 0 0 0 0 0 1 1 1 0 0 1 0 0 0 1
A. ramosa 0 0 1 1 1 0 2 0 0 1 1 1 0 0 1 0 0 0 2
A. spicata 0 1 1 0 0 0 3 1 1 1 1 1 0 0 1 0 1 0 2
A. yaracuy 0 0 1 1 1 0 1 0 0 1 1 1 0 0 1 0 0 0 ?
G. interrupta 1 0 0 0 0 0 0 0 0 0 0 2 0 1 1 1 0 0 0
C. rivalis 1 0 0 0 0 1 0 1 0 2 0 0 1 0 0 0 0 1 0
‘‘Calyptronoma occidentalis type,’’ occur- tegument could be identified. It is five to

ring in A. spicata (as Aristeyera spicata). six cell layers thick at midlength and six to
Pollen of four of the five species of As- ten cell layers thick toward the micropyle.
terogyne was studied for this revision (A. The nucellus protrudes into the micropyle,
martiana, Knudsen & Asmussen 604; A. ra- which faces the ventral side of the locule.
mosa, Stauffer et al. 824; A. spicata, Stauf- A well-developed septal nectary occurs be-
fer et al. 822; A. yaracuyense, Stauffer & tween the carpel flanks (Fig. 8I). It becomes
Stauffer 823). The pollen (Fig. 3A–H) is a triradiate cavity at about midlength of the
asymmetric and monosulcate with a sulcus ovary and extends upward and gradually
length as long as the main axis. The exine outward to the apical region of the ovaries.
is tectate and finely perforate-rugulate (per- The epithelium, especially differentiated in
forate fossulate in A. spicata). This agrees the gynoecium of A. martiana, is formed by
with the study of Harley (unpubl.) of A. large columnar, uninucleate cells (Fig. 8J).
martiana (Schipp 392). Punt and Wessels The three openings of the nectary are to-
Boer (1966) described the pollen of A. mar- ward the top of the ovary and are nonsec-
tiana (Archer 1978, Schipp 392) as easily retory ducts lined by isodiametric epider-
recognized by its fine, but distinct reticu- mal cells. Even though Asterogyne was de-
lum, and the distinctly visible muri (bacu- scribed (Uhl and Dransfield, 1987; Wessels
la), which support the reticulum. The pollen Boer, 1968) as having apical styles, our
of A. spicata (Tamayo 4177) was described study shows they are basal to lateral, which
by Punt and Wessels Boer (1966) as having is probably a common pattern within Geon-
a distinctly vermiculate ornamentation. Re- omeae (Stauffer & Endress, unpubl. data).
sults of our study could not differentiate the Epidermal protuberances at the basal and
two groups proposed by Punt and Wessels midregion of the styles were observed in
Boer (1966) and show that the pollen is anthetic flowers of A. martiana (Schmid,
rather morphologically uniform within the 1970a) and A. yaracuyense (Stauffer & En-
genus. dress, unpubl. data) (Fig. 8G). These pro-
tuberances are absent in young buds and
PISTILLATE FLOWERS seem to develop shortly before anthesis.
The perianth of the pistillate flowers is The three stigmatic branches are papillate
composed of three free, imbricate sepals and reflexed at anthesis (Fig. 8H). The
and three petals, which are united to their staminodes are united to their middle into
middle (Fig. 8D). The carpels alternate with a tube and free above this level (Figs. 8D,
the petals. The ovary is superior and triloc- 10C, 14E). The number of staminodial
ular (Figs. 8E, 8I, 10E, 10I, 12H, 12L, 14G, lobes ranges from five in A. ramosa to 22
14J). Each carpel bears a pendent, anatro- in A. guianensis. A well-developed papil-
pous, crassinucellate ovule (Fig. 10H). One late epidermis in the staminodial lobes is
of these per gynoecium develops fully to present in A. spicata (Fig. 12K). Reduction
maturity. Only one well-differentiated in- of the number of staminodes with respect
to the number of fertile stamens in the same cence of a plant is in flower at a time, while
species is common within the genus; how- the other inflorescences are still in bud or
ever, in Asterogyne martiana the number of in fruit. This pattern was already observed
fertile and sterile stamens is always six in A. martiana by Schmid (1970a) and may
(Fig. 8A, D). The morphology and vascu- be common for the other species in the ge-
lature of the pistillate flowers of A. spicata nus. Staminate flowers emerge first, one af-
were studied by Uhl (1966) and for A. mar- ter another, and when both of them have
tiana by Uhl and Moore (1977). The struc- fallen, the pistillate flower emerges. Two
ture of the gynoecia in four species of As- staminate flowers at anthesis in the same pit
terogyne is part of a current study including were occasionally observed in A. yaracu-
representatives of all Geonomeae (Stauffer yense and A. martiana (Schmid, 1970a). In
& Endress, unpubl. data). The chromosome A. martiana the anthesis of individual sta-
number in the genus is unknown (M. Röser, minate flowers lasts from two to six days
pers. comm.). and the anthesis of the pistillate flowers
then follows and lasts one to two days
FRUITS (Schmid, 1970a; Uhl & Moore, 1977).
Young fruits were observed to occur si-
Fruits of Asterogyne are ellipsoid to el-
multaneously with pistillate flowers on the
lipsoid-ovoid, purplish, black, or garnet red
rachillae of A. martiana (Bonifaz & Cor-
drupes, with a smooth surface and a single
nejo 3758).
seed (Figs. 7G, 11E). They are conspicu-
Several structural features of Asterogyne
ously keeled at the apex. The remains of
flowers suggest insect pollination. The
the stigma are basal. The seed is ellipsoid
well-developed septal nectary (Fig. 8I) ob-
and slightly dorsiventrally compressed. The
served in the pistillate flowers of four of the
endosperm is homogeneous (not ruminate),
five species in the genus (pistillate flowers
and the embryo is basal (Uhl & Dransfield,
of A. guianensis were not available for the
1987). The morphology and anatomy of the
structural study), and the rudimentary septal
fruits in A. martiana and A. spicata were
nectary observed by Schmid (1970a, 1983)
studied by Wessels Boer (1968). Fruits of
in the pistillode of the staminate flowers of
A. martiana resemble those of A. spicata,
A. martiana, may provide nectar for polli-
but the arrangement of the vascular bundles
nators. Schmid (1970b) found that the
in the parenchyma of the mesocarp is less
translucent protuberances at the base of the
regular. In A. spicata, bundles of fibers
styles and the fleshy staminodes in A. mar-
without conductive elements are present in
tiana readily exude fluid when slightly
the outer layer and the endocarp is not ad-
damaged. The production of fragrance is
herent to the seed (Wessels Boer, 1968).
well known in both staminate and pistillate
Germination is adjacent-ligular (Uhl &
flowers of Asterogyne but its histological
Dransfield, 1987). Seeds of A. spicata have
basis is unknown. Floral fragrances were
been reported to germinate within three and
found in the male flowers of all of the Ve-
four months (Braun, 1970; Nehlin, 1994).
nezuelan species. Scent was especially
strong in A. spicata, and has also been re-
Reproductive Biology
ported in A. martiana (Knudsen, 1999;
What we know about the reproductive bi- Schmid, 1970a, 1983; Knudsen & Asmus-
ology of Asterogyne is limited almost en- sen 620; Moore 6535). The presence of os-
tirely to A. martiana (Schmid, 1970a, mophores was preliminarily tested on a vis-
1970b; Uhl & Moore, 1977). Asterogyne is it by the first author (between 2001 and
monoecious and the inflorescences are 2003) to the Venezuelan populations of As-
markedly protandrous. Male and female terogyne. The staminate flowers of A. spi-
flowers on the same inflorescence of A. cata that were submerged for five minutes
martiana are rarely open at the same time, in a water solution containing neutral red
thus maximizing pollen transfer among dif- showed traces of the dye at the base of the
ferent plants (Schmid, 1970a). In the spe- petals and filaments. Floral scent composi-
cies in Venezuela usually just one inflores- tion in A. martiana and its potential impor-
tance in maintaining reproductive isolation ciation with Euterpe precatoria var. longev-
were studied by Knudsen (1999). aginata (Mart.) Henderson, Geonoma inter-
Insect pollination of Asterogyne marti- rupta (Ruiz & Pav.) Mart. var. interrupta,
ana was discussed by Schmid (1970b). In and especially Prestoea pubigera (Griseb.
two populations in Costa Rica, he found in- & H. Wendl.) Hook. f. ex Benth. & Hook.
sects of five different orders visiting the f. The general habit and the bifid simple
flowers; syrphid flies probing for nectar leaf blades of P. pubigera surprisingly re-
were considered effective pollinators for the semble A. ramosa and this might have ob-
species. The flies also visited staminate scured the recognition of the latter in the
flowers to eat pollen. Syrphid flies have relatively nearby cloud forest of Trinidad.
also been reported by de Nevers & Hen- As pointed out by Henderson and Steyer-
derson 6323 to visit flowers of A. martiana mark (1986), all the Venezuelan popula-
in Panama. Ants (Formicidae, Hymenop- tions of Asterogyne are located in three
tera) were reported by Schmid (1970b) to plant refugia of the Coastal Cordillera (Nir-
visit inflorescences of A. martiana (Fig. 7E) gua, Guatopo, Paria) mentioned by Steyer-
and were observed by the first author in sta- mark (1979). Asterogyne spicata is endemic
minate anthesis in A. spicata. Asterogyne to the Guatopo National Park and was re-
spicata has bright orange staminate flowers, cently collected in the forests NW of Gua-
which emit a very strong, sweet scent and tire, in the state of Miranda, where it grows
are visited by bees (Henderson, 1986). Dis- at 600 to 1100 m and is restricted to the
persal of the fruits has never been studied, ecotone between the lowland rain forest and
but the ripe, dark purplish, black, or garnet the low altitude cloud forest. In the locali-
red fruits contrasting with the brown, red- ties that the first author has visited, Aster-
dish brown, reddish, or salmon to pink ogyne spicata is rare and associated with
rachillae suggest bird dispersal. Bactris setulosa H. Karst., Euterpe preca-
toria var. longevaginata, and Geonoma un-
Distribution and Ecology data Klotzsch. Asterogyne yaracuyense was
once thought to be endemic to the cloud
The five species of Asterogyne are dis- forest on the Cerro La Chapa (Yaracuy
tributed in Central America and northern State), normally above 1200 m, but recent
South America (Fig. 4). They are understo- field work of W. Meier (pers. comm.) con-
ry palms; A. martiana and A. ramosa oc- firmed its presence in the nearby cloud for-
casionally are dominant in that stratum of est on Cerro Zapatero. In Cerro La Chapa,
the forest. The only widespread species is A. yaracuyense is rare and usually associ-
A. martiana; the other four species are nar- ated with Dictyocaryum fuscum (H. Karst.)
row ecological endemics, restricted to small H. Wendl., Geonoma interrupta var. inter-
areas in Venezuela and French Guiana. As- rupta, G. spinescens var. braunii F. W.
terogyne martiana grows from Belize to Stauffer, Socratea altissima Balslev & F. W.
northern Ecuador, usually forming large Stauffer (spec. ined.), and Wettinia prae-
stands in the understory of the lowland non- morsa (Willd) Wess. Boer. Appun (1871)
flooded rain forest, but sometimes also provided a short description of a palm in
along streamlets and seasonally flooded the cloud forest of the Pico Hilaria (central
patches of forest. It has also been collected Venezuelan Coastal Cordillera) that match-
in the cloud forests of Costa Rica and Co- es an Asterogyne species, probably A. yar-
lombia, sometimes to 1400 m. acuyense.
Asterogyne ramosa, A. spicata, and A. After 20 years of intensive field work in
yaracuyense are restricted to the Venezue- French Guiana, J. J. de Granville discov-
lan Coastal Cordillera (Fig. 4). Asterogyne ered a fifth species of Asterogyne, A. gui-
ramosa is known only from the cloud for- anensis, from a single locality in the eastern
ests of the Penı́nsula de Paria, state of Su- part of the country, on the Brazilian border
cre, at 730 to 1250 m elevation. There it (Fig. 4). The genus had never been reported
forms large stands in particularly moist ar- for the Guianas; its presence in French Gui-
eas of the Cerro Humo and grows in asso- ana suggests it may occur in other regions
of northeastern South America. The very oma occidentalis. More taxa and more char-
limited range of A. guianensis may be re- acters are needed to obtain a robust hypoth-
lated to low adaptive and reproductive ca- esis of the relationships within Geonomeae.
pacity (de Granville and Henderson, 1988). Two studies, one based on rpl16 and the
Asterogyne guianensis is found in the Oya- other on rps16 plastid DNA sequences, spe-
pock refugium described by Brown (1982), cifically addressed the relationships of the
also known as the East Guiana refugium six genera of Geonomeae (Asmussen,
(Prance, 1982). De Granville and Hender- 1999a, 1999b). However, Asterogyne was
son (1988) suggest that A. guianensis and not included in the study based on rpl16
A. spicata may have once been a single because it was not possible to get a rpl16
population that became isolated as a result sequence for A. martiana at the time (As-
of climate changes. mussen, 1999a). In the study based on
In a study at La Selva Biological Station, rps16, Asterogyne was part of a poorly re-
Costa Rica, Braker and Chazdon (1993) as- solved monophyletic group containing spe-
sessed the levels of damage produced by cies of Geonoma, Calyptrogyne, and Calyp-
the neotropical forest grasshoper Microty- tronoma, with Welfia and Pholidostachys as
lopteryx hebardi (Acrididae: Ommatolam- sisters to this clade (Asmussen, 1999b). A
pinae) that feeds on leaves of A. martiana number of studies of Arecoideae and Geon-
and two species of Geonoma. Asterogyne omeae include Asterogyne, and a robust hy-
martiana was the least preferred species, pothesis of the position of Asterogyne can
probably due to lower water content, lower be expected to be published in the near fu-
fresh/dry mass per unit of leaf area, and ture. A study (Lewis, unpubl. data) of DNA
higher crude lignin content. sequences of the nuclear gene phosphori-
bulokinase from selected genera within sub-
The Placement of Asterogyne Based on family Arecoideae resolves species of As-
Molecular Phylogenetic Analyses terogyne as distinct and Geonoma as a
monophyletic group with Asterogyne as sis-
The taxonomic position of Asterogyne ter to Geonoma. An unpublished study
within the tribe Geonomeae is corroborated (Borchsenius et al., unpubl. data) based on
by phylogenetic analyses of DNA sequenc- morphological characters corroborates the
es. A clade of Geonoma congesta, Calyp- result from the nuclear phosphoribulokinase
tronoma occidentalis, and Asterogyne mar- DNA sequences. In conclusion, most of the
tiana is resolved as monophyletic in the data available, preliminary as they may be,
analysis of three plastid DNA sequences, point to a sister group relationship of Geon-
rbcL, rps16, and trnL-trnF by Asmussen oma and Asterogyne, with the remaining
and Chase (2001). Several other phyloge- genera of Geonomeae outside this mono-
netic analyses of palms have included A. phyletic clade.
martiana. Some of these studies (Hahn,
2002; Lewis & Doyle, 2001; Uhl et al., Relationships within Asterogyne Based
1995) had A. martiana as the only repre- on Molecular Phylogenetic Analyses
sentative for tribe Geonomeae and therefore
did not add information about tribal rela- Asterogyne martiana has a wide natural
tionships of Asterogyne. In other studies distribution and is widely cultivated in bo-
(Asmussen et al., 2000; Baker et al., 1999), tanical and private gardens. It is more dif-
A. martiana and other representatives of ficult to obtain material of the remaining
Geonomeae were part of a poorly resolved four species of Asterogyne, which are re-
clade of Arecoideae; these studies are also stricted in their natural habitats and are not
uninformative with respect to the position cultivated. This is the main explanation for
of Asterogyne. the lack of phylogenies of Asterogyne spe-
In the family level study by Asmussen cies. Attempts made to extract DNA from
and Chase (2001), Asterogyne martiana is herbarium material of A. spicata and A. yar-
resolved as sister to a poorly supported acuyensis have failed, but succeeded only
clade of Geonoma congesta and Calyptron- for A. ramosa (Asmussen & Deichmann,
pers. comm.). Based on preliminary data of Central America and Colombia (Bernal,
plastid DNA, A. ramosa forms a clade with 1989). Most of the populations of A. spi-
A. martiana that is sister to Geonoma (As- cata grow in the Guatopo National Park
mussen et al., unpubl. data). and therefore are protected by the rules of
Fresh leaf material for DNA analyses the national park. Asterogyne spicata is list-
from Asterogyne guianensis, A. ramosa, A. ed as vulnerable in the IUCN red list of
spicata, and A. yaracuyensis is critical for threatened species (Johnson, 1996). The
examining the infraspecific relationships of difficult access to the populations of A. ra-
Asterogyne. The combination of data from mosa in Venezuela and A. guianensis in
morphology, a number of variable DNA se- French Guiana is a major factor contribut-
quences, and AFLP characters for all spe- ing to the conservation of these species. As-
cies of Asterogyne is likely to provide a ro- terogyne yaracuyense was also listed as
bust interpretation of the interspecific rela- vulnerable by Johnson (1996). A recent vis-
tionships for the genus. it to its natural habitat on Cerro La Chapa
has confirmed more extensive alteration and
Relationships within Asterogyne Based destruction of the forest than originally re-
on a Morphological Phylogenetic ported by de Granville and Henderson
Analysis (1988) and Stauffer and Duno de Stefano
(1998). Small agricultural areas for the cul-
The strict consensus tree (Fig. 5, length:
tivation of bananas, coffee, and tubers have
26, CI: 0.962, RI: 0.889) based on mor-
gradually taken possession of the cloud for-
phology resolved Asterogyne as a well-sup-
est atop Cerro La Chapa. Cattle raising and
ported monophyletic group with a bootstrap
construction associated with agricultural ac-
value of 97%. However, it is clear that more
tivities have also altered the species content
outgroup taxa should be included in the
and structure of the cloud forest. Therefore,
analysis to test this hypothesis. Asterogyne
Asterogyne yaracuyense should be listed as
guianensis and A. spicata form a clade with
critically endangered in the IUCN red list
a bootstrap support of 78%; this group is
of threatened species.
characterized by spicate inflorescences and
male flowers with more than 21 stamens.
Systematic Treatment
Asterogyne ramosa and A. yaracuyense
form a clade with a bootstrap support of ASTEROGYNE H. Wendl. ex Hook. f.
90%; the species of this group grow in the
cloud forest of the Venezuelan Coastal Cor- Asterogyne H. Wendl. ex Hook. f. in Ben-
dillera and have a peduncle longer than 85 tham & Hook. f., Gen. Pl. 3: 914. 1883;
cm. The relationships between these two Burret, Bot. Jahrb. Syst. 63: 140. 1930;
clades and A. martiana were unresolved Wessels Boer, Verh. Kon. Ned. Akad.
(Fig. 5). The parsimony search detected two Wetensch., Afd. Nat., 2 Reeks, 58: 79.
most parsimonious trees. One of the two 1968; Uhl & Dransfield, Genera Palma-
most parsimonious trees had the same to- rum, 539. 1987; Dransfield & Uhl, Pal-
pology as the strict consensus tree, whereas mae, in Kubitzki (editor), Fam. gen. vasc.
the other showed A. martiana as sister to pl., fl. pl., 4: 385. 1998. TYPE: Asterogyne
the remaining species of Asterogyne. Our martiana (H. Wendl.) H. Wendl. ex
phylogeny supports the close relationship Hemsl. (Geonoma martiana H. Wendl.),
between A. guianensis and A. spicata, al- typified by H. Wendl. ex Hemsl. in God-
ready suggested by de Granville and Hen- man & Salvin, Biol. Centr. Am. Bot. 3:
derson (1988). A clarification of the exact 409. 1885.
position of the widespread A. martiana will Aristeyera H. E. Moore, J. Arnold Arbor. 47: 3.
have to await additional data. 1966. TYPE: A. spicata H. E. Moore.
Small to medium-sized, solitary, un-

Conservation Status
armed, pleonanthic, monoecious palm. Stem
Asterogyne martiana is a widespread smooth, brown to brownish cream, erect,
species considered to be nonthreatened in sometimes basally decumbent, occasionally
with basal and/or lateral vegetative branch- asymmetric in shape, monosulcate, the sul-
es, unarmed; internodes short; adventitious cus as long as main axis, the exine tectate
roots forming a basal cone. Leaves spirally and finely perforate-rugulate or perforate-
arranged, 6–31, erect to arching, marces- fossulate (Fig. 3A–H); pistillode irregularly
cent; sheath brown or cream colored, short, 3-lobed (Fig. 12E). Pistillate flowers ob-
tubular, splitting opposite petiole, the mar- long to oblong-ovoid; sepals 3, unequal,
gins stiff fibrous, the tomentum brown to imbricate in bud, dorsally keeled; petals 3,
reddish brown; petiole slender, adaxially united up to midlength, distally valvate;
flat or channeled, glabrous, abaxially round- staminodes 5–22, fused to corolla and unit-
ed or angled, brown tomentose; blade bifid ed basally, fleshy, with tanniferous tips; gy-
simple, sometimes becoming irregularly noecium syncarpous, trilocular, triovulate,
split with age or damage caused by wind, the nectary septal; style basal to lateral,
narrowly cuneate at base; veins 25–70, elongate; stigmatic branches 3, papillate, re-
emerging at 20–358 angle from midvein flexed at anthesis; ovule anatropous, insert-
(rachis), prominent at one or both surfaces ed at midlength of locule, pendent. Fruit a
of blade. Inflorescence interfoliar, solitary, drupe, ellipsoid to ellipsoid-ovoid, 1-seed-
branched to one order, rarely spicate or ed, dorsiventrally compressed, slightly to
forked, erect at anthesis, becoming pendent conspicuously keeled at apex, the remains
in fruit, protandrous; peduncle long, slen- of the stigma basal; epicarp smooth, pur-
der, brown to whitish tomentose; prophyll plish black or garnet red at maturity; me-
tubular, fibrous, membranous or charta- socarp fleshy and juicy, with an inner layer
ceous, bicarinate, brown tomentose, insert- of closely appressed, longitudinal fibers;
ed near base of peduncle; peduncular bracts endocarp thin, crustaceous, and shiny (Uhl
1 or 2, tubular, chartaceous or membranous, and Dransfield, 1987). Eophyll with a bifid
inserted 1 to several cm above base of pe- simple blade.
duncle; inflorescence rachis with short, Distribution. Five species distributed in
ovate, pointed bracts, subtending rachillae, Central America (Belize, Guatemala, Hon-
salmon to pink at anthesis, brown to reddish duras, Nicaragua, Costa Rica, and Panama)
brown in fruit; rachillae 1–16, ca. equal in and northern South America (Colombia,
length, ending in a short, sterile apex, same Ecuador, Venezuela, and French Guiana).
color as inflorescence rachis. Flower pits (Fig. 4)
containing triads comprising two staminate
flowers and one pistillate flower, the pits Key to the species of Asterogyne
spirally arranged on rachis, covered by a
rounded upper lip that overlaps the lower, 1. Stamens 21–27; inflorescences spicate.
immersed lip when in bud; floral bracts 3, 2. Inflorescence rachis and rachillae pale
the largest bract subtending the second sta- green at anthesis; filaments white
minate flower, a medium-sized, 2-keeled and smooth; fruits 2.5 cm long, 1.5
cm diam. --------------------------------- A. guianensis
bract subtending the pistillate flower, and a 2. Inflorescence rachis and rachillae
small bract representing the prophyll of the brown at anthesis; filaments conspic-
pistillate flower. Staminate flowers oblong uously yellow orange and papillate;
to oblong-obovoid, borne abaxially to pis- fruits to 1.8 cm long, 0.8 cm diam.
tillate flowers; sepals 3, unequal, united ba- --------------------------------------------------------- A. spicata
sally, narrow, elongate, keeled dorsally; 1. Stamens 6–12; inflorescences branched,

rarely spicate.
petals 3, ca. as long as or longer than sepals, 3. Peduncular bract 1, rarely 2; stamens
united into a tube for ca. ⅔ their length, 6; staminodial lobes 6 ------------ A. martiana
distally valvate; stamens 6–27; filaments 3. Peduncular bracts 2; stamens 8–12;
united into a tube up to midlength, white or staminodial lobes 5–9.
yellow, with or without a papillate epider- 4. Inflorescence rachis 0.5–2.5 cm
long; rachillae 4–6; stamens 11–
mis; connective bifid, tanniferous, bearing 12 ------------------------------------------ A. ramosa
separated thecae, the anthers inflexed to- 4. Inflorescence rachis 5–10 cm long;
ward center of flower and introrse in bud, rachillae 9–16; stamens 8–10 ---
spread and extrorse at anthesis; pollen ----------------------------------------- A. yaracuyense
1. ASTEROGYNE GUIANENSIS Granv. & A. J. in bud, 5 mm long, 2.5–3 mm diam., the

Hend. (Fig. 1A, B) pedicel to 1 mm long; sepals oblong-ob-
ovoid, 4–5 3 1 mm; petals oblong-obovoid,
Asterogyne guianensis Granv. & A. J. 4 3 1.4–1.6 mm; ovary 3 mm long, 2 mm
Hend., Brittonia 40: 76. 1988. TYPE: diam.; style 3 mm long; stigmatic branches
FRENCH GUIANA. Camopi River, ca. 1 mm long; staminodial tube 4 mm long,
1.5 km NE of Mont Belvédère, in swamp the lobes 20–22, 1–1.5 mm long. Fruit nar-
forest, 150 m, 4 Dec 1984, J. J. de Gran- rowly ellipsoid, strongly keeled apically,
ville 7124 [HOLOTYPE: P; ISOTYPES: B, 2.5 cm long, 1.5 cm diam.; epicarp garnet
BR(n.v.), CAY, K, NY, US] red and shiny at maturity; seed narrowly el-
lipsoid, 2 cm long, 0.8 cm diam.
Stem solitary, erect, 1.5–2 m tall, 3–6 cm Local names and uses. None reported.
diam., brown, not branched; internodes Distribution and ecology. Known only
short; aerial roots branched, forming a basal from two collections of eastern French Gui-
cone, each root 38–90 cm long, 0.2–0.3 cm ana, remarkably more than 200 km from
diam., yellowish brown; pneumatodes each other. Henderson and Balick (1987)
white, conical, to 0.2 cm long. Leaves 15– suggested that this species, at that time not
18, the older ones not persistent; sheath 7 yet described, may occur also in northern
3 4–5 cm, brown; petiole 53–60 3 0.8–1 Brazil.
cm, channeled adaxially, rounded abaxially,
Additional specimens examined: FRENCH GUI-
green, the tomentum reddish brown; mid- ANA. Crique Rapatea, Bassin de lApprouague, jonc-
vein 80–90 3 0.5 cm, green, the indumen- tion de la Kourouaı̈e et de la RN2, 48129N, 52879W, 24
tum same as on petiole, channeled adaxi- Oct 1996, Blanc 189 (CAY); plantlets cultivated at
ally, rounded abaxially; blade 1–1.1 3 0.3– Cayenne, 22 Oct 1985, de Granville 7124-A (CAY).
0.4 m, deeply bifid at apex for ca. 35 cm;
primary veins 29 per side, emerging at 20– 2. ASTEROGYNE MARTIANA (H. Wendl.) H.
308 angle at midvein, prominent on abaxial Wendl. ex Hemsl.
side, almost yellow when dry, the lepidote
tomentum slightly reddish brown. Inflores- Asterogyne martiana (H. Wendl.) H. Wendl.
cence spicate; peduncle 40–55 cm long, ex Hemsl. in Godman & Salvin, Biol.
0.4 –0.5 cm diam., dorsiventrally com- Centr. Am. Bot. 3: 409. 1885; Geonoma
pressed, the tomentum brown to whitish; martiana H. Wendl., Linnaea 28: 342.
prophyll 20 cm long, 1.5 cm diam., mem- 1856. TYPE: cult. hort. Herrenhausen, H.
branous, green, inserted at base of pedun- Wendland 56 [HOLOTYPE: GOET; ISOTYPE:
cle; peduncular bract 1, membranous, K(F neg. 20765, BH neg. 20368)] (Fig.
green, 40–42 3 1.3–1.5 cm, inserted 1 cm 1C)
above the base of the peduncle; inflores-
Geonoma trifurcata Oerst., Kjoeb. Vidensk. Med-
cence rachis 20–30 cm long, 0.9–1.5 cm del. 1858: 34. 1859. TYPE: Nicaragua, ad flumen
diam., terete, pale green at anthesis, reddish San Juan, A. S. Oersted 6566 [HOLOTYPE: C(F neg.
brown in fruit, the tomentum whitish to 21069)].
brownish, the sterile apex acute, 1–2 mm Asterogyne minor Burret, Bot. Jahrb. Syst. 63: 143.
long; flower pits spirally arranged, 3–5 mm 1930. TYPE: SW Colombia, above Barbacoas, 500
m, F. C. Lehmann 708 [HOLOTYPE: B(destroyed)].
apart from each other, covered with an in-
flexed lower lip, 3–4 3 2–3 mm. Staminate Stem solitary, erect, sometimes basally de-
flowers obovoid, 9–10 mm long, 3–4 mm cumbent, 0.4–1.6(–3) m tall, 2.5–5 diam.,
diam., white at anthesis, the pedicel to 1–2 dull brown to orange, not branched; inter-
mm long; sepals oblong, 9–9.2 3 2–3 mm; nodes (1–)2.5–3 cm long; aerial roots form-
petals oblong-obovate, 8 3 2.5–3 mm; sta- ing an irregular basal cone, each root 4–8
mens 25–27; filaments united into a tube up cm long, 0.3–0.4 cm diam.; pneumatodes
to 4–5 mm long, the free parts 2.5–3 mm brownish cream-colored, conical, to 1 mm
long, 1 mm diam., white; connective 0.8 long. Leaves (6–)12–20, yellowish red when
mm long; thecae 1.2–1.5 mm long; pistil- young, green when mature, the old ones
lode 1–2 mm long. Pistillate flowers oblong sometimes persistent; sheath (8–)11–16(–38)
3 3 cm, with reddish brown tomentum, lon- diam., remarkably thinner at apex and
gitudinally splitting, the margins slightly fi- therefore the anther versatile, white, with-
brous; petiole (4–)20–68(–75) 3 0.5–1.5 out a papillate epidermis; connective slight-
cm, keeled to channeled adaxially, rounded ly swollen, 0.7–1.1 mm long; thecae 0.7–
to angled abaxially, almost glabrous to 1.6 mm long; pistillode 2–5 mm long, 1
densely brown tomentose, toward the base mm diam., basally fused with staminodial
slightly fibrous at margins; midvein (33–)55– tube. Pistillate flowers oblong in bud, 6–8
79(–108) 3 0.2–0.7 cm, greenish yellow, mm long, 2.7–3.1 mm diam.; sepals oblong,
glabrous or with pinkish brown tomentum, 4–5 3 1–1.5 mm; petals oblong, 4.4–5.5 3
slightly keeled adaxially, rounded abaxial- 1.1–1.8 mm; ovary 1 mm long, 0.8 mm
ly; blade 0.4–1.3 3 0.15–0.3 m, light to diam.; style 6–8 mm long; stigmatic
dark green, deeply bifid at apex for 18–32 branches 1–2 mm long; staminodial tube
cm; primary veins 28–40 per side, emerg- 3.9–5 mm long, the lobes 6, 1.5–2.1 mm
ing at 20–358 angle at midvein, prominent long. Fruit ellipsoid to ellipsoid-ovoid, 0.8–
at both surfaces of blade, yellow when dry. 1.2 cm long, 0.4–0.7 cm diam.; epicarp or-
Inflorescence branched to 1 order or very ange to red when young, purplish black at
rarely spicate; peduncle (20–)34–70(–125) maturity; seed ellipsoid, 0.7–0.8 cm long,
cm long, 0.4–0.7 cm diam., dorsiventrally 0.4–0.5 cm diam.
compressed; prophyll 11–26(–36) cm long, Local names and uses. capoca (Guate-
0.6–1.5(–2) cm diam., inserted at base of mala); pacuquilla (Honduras); pata de gallo,
peduncle; peduncular bract 1, rarely 2, suita (Nicaragua); pacaya, suita (Costa
chartaceous, brownish cream-colored, the Rica), dobo-gaw (Guaymi of Panama); cor-
indumentum same as on prophyll; first pe- tadera, panda, panga, rabihorcao, rabo de
duncular bract 20–31(–40) 3 0.5–1.5 cm, zorro (Colombia); guacamayo, pico (Ecua-
inserted 0.5–2 cm above base of peduncle; dor). The use of the leaves for thatching
second peduncular bract, when present, 15 houses has been reported in Nicaragua
3 0.5–0.9 cm, inserted 25–27 cm above
(Neill 1723, Salick 8106), in Costa Rica
base of peduncle; inflorescence rachis ab-
(Chazdon, 1986), in Panama (Gordon 70),
sent or 0.5–3(–7) cm long, 0.3–0.5(–1.2)
in Colombia (Bernal & Galeano 887), and
cm diam., dorsiventral to irregularly com-
in Ecuador (Aulestia & Aulestia 1306).
pressed, salmon to pink at anthesis and in
Distribution and ecology. Widespread
fruit, glabrous or with indumentum same as
on peduncle; rachillae (2–)3–5(–14), terete, in Central America: Belize (El Cayo, Stann
erect in bud, slightly pendent in fruit, the Creek, Toledo), Guatemala (Izabal), Hon-
color same as inflorescence rachis, the to- duras (Atlántida, Colón, Cortés, Gracias a
mentum white, farinose or short and ap- Dios, Santa Bárbara), Nicaragua (Chonta-
pressed; basal rachillae (9–)14.5–22(–27) les, Jinotega, Matagalpa, Rı́o San Juan, Ze-
cm long, 0.4–0.6(–1.1) cm diam.; middle laya), Costa Rica (Alajuela, Cartago, He-
rachillae (8.5–)14–21(–23.5) cm long, 0.4– redia, Limón, Puntarenas, San José), Pana-
0.6(–1) cm diam.; apical rachillae (9–)15– ma (Bocas del Toro, Chiriquı́, Coclé, Colón,
22 cm long, 0.4–0.6(–1.2) cm diam., the Darién, Panama, San Blas, Veraguas), and
sterile apex acute, 0.2–1.1(–5.5) cm long; in northwestern South America: Colombia
flower pits spirally arranged, ca. 2–3 mm (Antioquia, Chocó, Córdoba, Nariño, Valle
apart from each other, covered with a lower del Cauca), Ecuador (Esmeraldas) (Fig. 4).
semicircular lip, 1–2 3 1–2 mm, the lip be- Common in the understory of the lowland
coming reflexed in fruit. Staminate flowers and mountain rain forests, in flooded and
oblong-obovoid in bud, 6–9 mm long, 2–3 nonflooded areas; from sea level to 1400 m,
mm diam., white at anthesis, the pedicel to but normally below 600 m.
0.7 mm long; sepals oblong-lanceolate,
3.6–4.8 3 1 mm; petals oblong-lanceolate, Selected specimens examined: BELIZE. El Cayo
District: 47 mile section, Humming Bird Hwy., 28 Apr
5–7 3 1–1.3 mm; stamens 6, the filaments 1956, Gentle 9072 (NY), 60 m, 6 Sept 1929, Schipp
united into a tube up to 5.4–7 mm long, the 392 (G, K, MO, NY, Z); W of Isla Rivas, 29 Jan 1903,
free parts 2.4–3 mm long, 0.3–0.5 mm Wilson 224 (NY, US). Toledo: Monkey River, in high
‘‘Monkey Tail,’’ 1 Nov, 1941, Gentle 3743 (MO, NY, Almirante, 30 Nov 1971, Gentry 2803 (MO); Rı́o San
US). Pedrito, 16 Jul 1978, Gordon 70 (MO); without local-
GUATEMALA. Izabal: Wet wooded slopes nr. Km ity, 24 Sept 1940, von Wedel 952 (BH). Chiriquı́: Bur-
824 past Villa Jesús on rd. from Puerto Barrios to Gua- ica Peninsula, 19 Feb 1973, Croat 21997 (K, MO);
temala, 100 m, 8 Nov 1959, Moore & Cetto 8213 Fortuna Dam region, nr. Chiriquı́ Grande, 88459N,
(BH); valley of tributary of Rı́o San Francisco del Mar, 828159W, 200 m, 18 Jan 1986, McPherson 8080 (MO);
2 miles NE of Hopi, 12 miles from Entre Rı́os, 10–20 vic. Chiriquı́ Lagoon, 9 Oct 1940, von Wedel 1108
m, 21 Apr 1940, Steyermark 39785 (US); E portion of (BH, MO). Coclé: Rd. to Coclesito, logging camp, 12
Vera Paz and Chiquimula, 1885, Watson 238 (BH). miles from Llano Grande, 88479N, 808289W, 200 m, 16
HONDURAS. Atlántida: Vic. Lancetilla, 45 m, 8 Dec 1983, Churchill et al. 4122 (K, MO); Boca del
Oct 1934, Yuncker 5002 (MO, NY). Gracias a Dios: Toabré at confluence of Rı́o Toabré and Rı́o Coclé del
Laimos Creek, 15 km SW of Waspan, 7 Mar 1961, Norte, 11 Apr 1969, Lewis et al. 5558 (BH, MO). Co-
15–30 m, Bunting & Licht 401 (NY); Rı́o Plátano, 4 lón: Santa Rita Ridge, 11 May 1999, Henderson &
hours upriver from village of Ras, nr. sea level, 23 May Ferreira 3048 (NY); Santa Rita Ridge, end rd. from
1973, Gentry et al. 7542 (MO). Transisthmian Hwy., ca. 10 miles from Hwy., 29 Mar
NICARAGUA. Chontales: Chontales, 600 m, Jun 1969, Porter et al. 4736 (BH, MO, NY); Santa Rita
1870, Levy 505 (G, P); 1870, Seemann s.n. (BM). Jin- Lumber Rd., ca. 3.1 km E of Agua Clara Rain Gauge,
otega: Salto Kayaska, Rı́o Bocaz, ca. 138519N, 9 Sep 1975, Witherspoon et al. 8322 (MO). Darién:
858229W, 190–340 m, 7 Mar 1980, Beach et al. 16545 Parque Nacional del Darién, ridge between Rı́o Topal-
(US); Comarca de Bocaycito, Cordillera Isabelia, nr. isa and Rı́o Pacuro, ca. 13 km E of Pacuro, Quebrada
Rı́o Bote, 6 May 1976, Neill 7151 (MO, US). Mata- Pobre to Mi Casita, 88039N, 778209W, 450–600 m, 14
galpa: El Comején, 1 km W de la carretera a Waslala, Oct 1987, de Nevers et al. 8320 (MO, US). Panama:
138159N, 858349W, 600 m, Moreno & Robleto 20602 Canal Zone, Apr 1925, Cook s.n. (US); Cerro Jefe
(MO). Rı́o San Juan: Sabalo, 1 km al N de Rı́o San roadside, 2 km N of turnoff to radio tower on rd.
Juan, 118029N, 848279W, 100 m, 9–10 Jul 1985, Mo- [from?] Alto de Pacora, 790 m, 30 Sep 1978, Hammel
reno 26083 (MO); ‘‘Los Filos,’’ nr. Loma Los Filos, 4863 (MO). San Blas: 10 km from Pan-American
Rı́o Santa Cruz, 11889N, 848209W, 6 Sept 1993, Salick Hwy. on El Llano–Cartı́ Rd., 98209N, 798059W, 250 m,
8106 (MO). Zelaya: Ca. 1 km N of El Zapote, 6 km 6 Mar 1987, Barfod & McPherson 13 (AAU); 44 km
S of Colonia Verdún, 118349N, 848249W, 250–300 m, N of Penonome on rd. to Coclecito, 90–150 m, 21 Feb
8 Sept 1983, Nee & Vega 27905 (NY, US); El Hor- 1978, Hammel 1682 (BH, MO); Nusagandi, El Llano–
miguero, caño above camp and on W slope at Loma, Cartı́ Rd., 19 km from Inter-American Hwy. 98199N,
138459N, 848599W, 800–1000 m, 17 Mar 1980, Pipoly 788559W, 300–400 m, 9 Oct 1996, Knudsen & Asmus-
6143 & 6168 (MO, US); ca. 6.3 km S of bridge at sen 604, 620 (AAU); El Llano–Cartı́ Rd., Km 10–15,
Colonia Yolania and ca. 0.8 km S of ridge of Serranias 300–400 m, 3 Sept 1977, Maas et al. 2803 (U). Ver-
de Yolania on rd. to Colonia Manantiales (Colonia So- aguas: Atlantic slope, Rı́o Concepción to Rı́o Barrera,
moza), 11836–379N, 848229W, 200–300 m, 29–31 Oct ca. 60 m, 16 Oct 1978, Hammel 5153 (MO).
1977, Stevens 4810 (BH, MO, NY). COLOMBIA. Antioquia: Mun. Frontino, Corregi-
COSTA RICA. Alajuela: Vic. Guatuso de San Ra- miento de Murrı́, rd. from Nutibara to La Blanquita,
fael (on Rı́o Frı́o), 108439N, 848489W, 80–100 m, 4 950 m, 18 Mar 1982, Bernal & Galeano 256 (NY);
Aug 1949, Holm & Iltis 914 (BH, BM, MO, US); 25 Mun. de San Rafael, 15 km al E del pueblo en la
km NNW of San Ramón by rd. on way to San Lor- carretera a San Carlos, 960 m, 19 Sept 1987, Bernal
enzo, 108179N, 848349W, 500 m, 24 Apr 1983 Liesner & Tobón 1380 (AAU, BH, NY); Villa Arteaga, Mu-
& Judziewicz 14825 (MO, WIS). Cartago: Bois de tata, ca. 1000 m, 5 Apr 1976, Dransfield et al. 4859
Peralta, 300 m, Jun 1903, Pittier 16658 (US). Heredia: (BH, K). Chocó: Area of Baudó, on left bank of river
Finca La Selva, 12 Dec 1984, Henderson 53 (NY); Baudó, ca. 11.5 km upstream of estuary, slightly
along ‘‘Starkey Rd.,’’ 4.5 km SE of bridge at Puerto downstream of estuary of Quebrada Angueradó, 50 m,
Viejo, 108269N, 838589W, 50 m, 7 Aug 1979, Stevens 7 Feb 1967, Fuchs & Zanella 21808 (U); north ridge
13505 (K, MO, US). Limón: Finca Castilla, 30 m, 24 of Alto de Buey, above Dos Bocas del Rı́o Mutatá,
Jul 1936, Dodge & Goerger 9284 (MO); 7 km SW of tributary of Rı́o El Valle, ESE of El Valle, 200–500
Bribrı́, 100–250 m, Gómez et al. 20341 (MO, NY, m, 8 Aug 1976, Gentry & Fallen 17445 (MO). Cor-
WIS); woodlands south of La Lola on the railroad, 120 doba: Junction of Rı́o Tigre and Rı́o Manso, Paramillo
m, 15 Apr 1953, Moore 6717 (BH). Puntarenas: Gol- National Park, transect 1, 78309N, 768059W, 200 m, 25
fito, 15 m, 22 Oct 1950, Hari s.n. (G); hills above Jul 1988, Gentry & Cuadros 63735 (MO). Nariño:
Palmar Norte, 550 m, 6 Mar 1953, Moore 6530 (AAU, Mun. de Tumaco, 3–5 km al E de Guayacana, 100 m,
BH, K); Osa Peninsula, Agabuena, 4 km W of Rincón, 3 Oct 1985, Bernal & Galeano 887 (AAU, NY). Valle
at cleared path dividing lands of Don Fortunato and del Cauca: Buenaventura, 6 May 1926, Cook 84 (US);
Don Contreras, SSW of 4-ha sample plot, 88439N, Bajo Calima, Juanchaco Palmeras, ca. 10 km NW of
838319W, 250 m, 18 Nov 1992, Thomsen 187 (C). San Buenaventura, 38569N, 778089W, 13 Apr 1987, Gentry
José: Rı́o Negro, Cerro La Cangreja, Santa Rosa de et al. 56871 (MO, NY).
Puriscal, 400–700 m, 20 Jun 1986, Chacón & Chacón ECUADOR. Esmeraldas: Reserva Etnica Awá,
1958 (CR); Tarrazú, San Lorenzo, Llanos de Santa Parroquia Alto Tambo, Centro de la Unión, cañón del
Marı́a, 98329300N, 848019200W, 500–600 m, 28 Jan Rı́o Mira, 008529N, 788269W, 250 m, 22 Mar 1993,
1998, Valverde 665 (K). Aulestia & Aulestia 1306 (MO); Playa de Oro on the
PANAMA. Bocas del Toro: Top of ‘‘Fila’’ above Rı́o Santiago, ½ hour walk S of village, 250 m, 25 Feb
1994, Borchsenius 242 (AAU, NY); San Lorenzo Can- regularly compressed, pale green at anthe-
tón, Reserva Etnica Awá, Centro Ricaurte, a orillas del sis, reddish in fruit, glabrous; rachillae 4–
Rı́o Palabı́, 18109N, 788329W, 300 m, 28 Oct 1992,
Tipaz et al. 2250 (AAU, MO). 6, terete, erect, same color as inflorescence
Cult. Hort. Herrenhausen, Wendland 56 (GOET, HO- rachis, each rachilla subtended by a curved
LOTYPE; K, ISOTYPE). Probably originated from seeds bract up to 0.5 cm long, covered with a
sent by Warscewicz from Central America. whitish farinose indumentum; basal, medi-
al, and apical rachillae 17–19 cm long, 0.5
3. ASTEROGYNE RAMOSA (H. E. Moore) cm diam., the sterile apex acute, 0.2–0.4 cm
Wess. Boer. (Fig. 1D–F) long; flower pits spirally arranged, ca. 2–3
mm apart from each other, covered with an
Asterogyne ramosa (H. E. Moore) Wess. inflexed lower lip, 1.5–2 3 1.5–2 mm. Sta-
Boer, Verh. Kon. Ned. Akad. Wetensch., minate flowers oblong-obovoid in bud, 6–8
Afd. Nat., 2 Reeks, 58: 81. 1968. Aris- mm long, 3–4 mm diam., slightly fragrant,
teyera ramosa H. E. Moore, J. Arnold white at anthesis, the pedicel to 1.5 mm
Arbor. 48: 144–146. 1967. TYPE: VE- long; sepals oblong-obovate, 5–6 3 2–3
NEZUELA. Sucre: Penı́nsula de Paria, mm; petals oblong, 4–6 3 2 mm; stamens
Cerro Espejo, en la cumbre entre Man- 11–12, occasionally 1 sterile; filaments
tecal y Paujı́, arriba de Mundo Nuevo y united into a tube to 3 mm long, the free
Rı́o Seco de Irapa, 750–850 m, 6 Aug parts 3 mm long, 0.5 mm diam., white, not
1966, J. Steyermark & M. Rabe 96088 remarkably thinner at apex, the epidermis
(HOLOTYPE: BH; ISOTYPES: NY, US, VEN) not papillate; connective 1 mm long; thecae
1.2 mm long; pistillode 2 mm long. Pistil-
Stem solitary, erect, 3–6 m tall, 6–10 cm late flowers oblong-ovoid in bud, 5–7 mm
diam., creamy brown, occasionally with lat- long, 3–4 mm diam., the pedicel to 1 mm
eral vegetative branches; internodes 5–10 long; sepals oblong to ovoid, 4–5 3 2–2.5
cm long; aerial roots forming a short basal mm; petals ovate, 5–6 3 2 mm; ovary 1.5
cone, each up to 15 cm long, 0.5–1 cm mm long, 1.5 mm diam.; style 1.5–2 mm
diam., brown, conical; pneumatodes very long; stigmatic branches 1 mm long; stam-
short to almost absent. Leaves 15–20, the inodial tube 4 mm long, the lobes (5–)7–9,
older ones briefly persistent; sheath 9–10(– 2.8–3.4 mm long. Fruit ellipsoid, 1.5–1.7
17) 3 7 cm, densely brown tomentose, the cm long, 0.5–0.6 cm diam.; epicarp dark
margins slightly fibrous; petiole 30–35 3 purplish red when young, purplish black at
0.6–1 cm, channeled adaxially, rounded maturity; seed ellipsoid, 1.3 cm long, 0.4
abaxially, brown tomentose; midvein 39 3 cm diam.
0.5 cm, the indumentum brown, slightly Local name and uses. anare. Uses not
channeled adaxially, almost flat abaxially; known.
blade 0.5–0.7 3 0.18–0.27 m, green, deeply Distribution and ecology. Known only
bifid at apex for ca. 33 cm; primary veins from the understory of the montane cloud
34–45 per side, emerging at 25–308 angle forest in the northeastern Coastal Cordille-
at midvein, prominent on both surfaces of ra, state of Sucre, Venezuela (Fig. 4); from
blade, pale green abaxially. Inflorescence 730–1250 m. Although the voucher for col-
branched to 1 order; peduncle 90–120 cm lection Steyermark 94803 mentions the
long, 0.4–0.5 cm diam., dorsiventrally com- palm growing at 1600 m on the Cerro
pressed, green; prophyll 15–20 cm long, 1.8 Humo, this must be an error because the
cm diam., chartaceous, inserted at base of summit of this mountain has been reported
peduncle; peduncular bracts 2, chartaceous, to be 1250 m.
brown, the indumentum same as on pro-
phyll; first peduncular bract 31–49 3 1.4 Additional specimens examined: VENEZUELA.
cm, inserted 0.5 cm above base of pedun- Sucre: Penı́nsula de Paria National Park, above Las
cle; second peduncular bract 41–53 3 1– Melenas, N of Rı́o Grande Arriba, trail to Cerro de
Humo, 108429N, 628379W, 900–1000 m, 3 Jan 2002,
1.2 cm, inserted 1 cm above base of pedun- Stauffer et al. 824 (MY, MYF, VEN, Z); Penı́nsula de
cle; inflorescence rachis 0.5–1.2(–2.5) cm Paria, Cerro Humo, a lo largo de uno de los afluentes
long, 0.4–0.5 cm diam., dorsiventral to ir- del Rı́o Santa Isabel, arriba de Siparo, 1600 m (prob-
able erroneous elevation), 1 Mar 1966, Steyermark duncle; peduncular bract 1, chartaceous,
94803 (BH, NY, U, VEN); Penı́nsula de Paria, above brown tomentose, 22–40 3 1.3–2 cm, in-
Las Melenas, N of Rı́o Grande Arriba, SE of Cerro de
Humo, 108429N, 628379W, 730–1050 m, 1 Dec 1979, serted 1.4 cm above base of peduncle; in-
Steyermark & Liesner 120915 (MO, NY, VEN); Pen- florescence rachis 23–40 cm long, 1–1.5 cm
ı́nsula de Paria, Distrito Marino and Distrito Arismen- diam., terete, pale brown at anthesis, brown
di, trail between crossing of Rı́o Tacarigua to summit in fruit, with same indumentum as on pe-
of slopes E of Cerro Humo, descending to Las Mele-
nas, N of Rı́o Grande Arriba, 10841–429N, 62836– duncle, the sterile apex acute, ca. 0.6–3 cm
379W, 760–1000 m, 24 Feb 1980, Steyermark et al. long; flower pits spirally arranged, ca. 2–4
121713 (MO). mm apart from each other, covered with an
inflexed lower lip, 3 3 2–4 mm. Staminate
flowers obovoid, 0.8–1.3 cm long, 0.4–0.5
4. ASTEROGYNE SPICATA (H. E. Moore)
cm diam., fragrant, the pedicel almost ab-
Wess. Boer. (Fig. 2A–C)
sent; sepals oblong-obovoid, 5–6 3 1.4–2
mm; petals obovoid, 7–8 3 2–3 mm; sta-
Asterogyne spicata (H. E. Moore) Wess.
mens 21–26, occasionally 1 or 2 partially
Boer, Verh. Kon. Ned. Akad. Wetensch.,
or totally sterile; filaments united into a
Afd. Nat., 2 Reeks, 58: 82. 1968; Aris-
tube to 6–7 mm long, conspicuously yellow
teyera spicata H. E. Moore, J. Arnold
orange, not remarkably thinner at apex, the
Arbor. 47: 5–9. 1966. TYPE: VENEZUE-
epidermis papillate, the free parts 3–4 mm
LA. Miranda, Parque Nacional Guatopo,
long, 0.4 mm diam.; connective 1.2–1.5
moist rich mixed evergreen forest on
mm long; thecae 0.5–1.2 mm long; pistil-
steep slopes bordering Rı́o Santa Cruz
lode 4–5 mm long. Pistillate flowers oblong
and Altagracia de Orituco, 14.5 km from
in bud, 8 mm long, 4 mm diam., the pedicel
Los Alpes, 12 km from Rancherı́a Mi
to 1 mm long; sepals oblong, 5–7 3 2–3
Querencia, 520 m, 23 Nov 1961, J. Stey-
mm; petals oblong, 7–8 mm 3 2–3 mm;
ermark 90010 (HOLOTYPE: BH; ISOTYPES:
ovary 0.9 mm long, 1–1.2 mm diam.; style
NY, VEN)
2.5–3 mm long; stigmatic branches 1–1.5
Stem solitary, erect, 2–4(–8) m tall, 4–7 mm long; staminodial tube 2 mm long, the
cm diam., brown, occasionally with basal lobes 15–21, 2–2.2 mm long. Fruit ellip-
vegetative branches; internodes 0.5 cm soid-ovoid, 1.5–1.8 cm long, 0.5–0.8 cm
long; aerial roots forming a short basal diam.; epicarp purplish red in young fruits,
cone; pneumatodes very short. Leaves 15– purplish black at maturity; seed ellipsoid,
19(–25), sometimes reddish brown when 0.8–1 cm long, 0.5 cm diam.
young, green when mature, the older ones Local name and uses. palma, palmito,
not persistent; sheath 10–15 3 9 cm, red- palmiche. Moore (1966) noted that the
dish brown tomentose, the margins slightly fruits are said to be edible and the leaves
fibrous; petiole 35–40 3 0.7–1.5 cm, chan- used for a durable thatch supposed to last
neled adaxially, rounded to angled ab- ten years. Nehlin (1994) reported that the
axially, brown tomentose, slightly fibrous at endocarp has a sweet acidic taste.
margins toward base; midvein 68–84 3 Distribution and ecology. Known only
0.5–0.7 cm, green, with same indumentum from the understory of the lowland and
as on petiole, ridged adaxially, flat abaxi- mountane cloud forest in the northern
ally; rachis with a filiform extension of 1.5– Coastal Cordillera, state of Miranda, Ve-
5 3 0.1 cm; blade 0.7–1 3 0.2–0.3 m, dull nezuela (Fig. 4); normally between 600–
green adaxially, silver green abaxially, 1000 m.
deeply bifid at apex for ca. 45 cm; primary Additional specimens examined: VENEZUELA.
veins 25–27 per side, emerging at 25–308 Miranda: Parque Nacional Guatopo, 20 Dec 1961,
angle at midvein, prominent abaxially, yel- Agostini 10 (VEN); Carretera Santa Teresa–Altagracia
lowish, slightly brown tomentose. Inflores- de Orituco, 600–700 m, Jun 1953, Aristeguieta 1772
cence spicate; peduncle 40–50 cm long, (NY, VEN); selvas de Guatopo, 500 m, Jun 1958, Ar-
isteguieta 3185 (NY, VEN); selvas pluviales de Gua-
0.5–0.6 cm diam., dorsiventrally com- topo, 400–600 m, 13 Nov 1956, Bernardi 5633 (NY);
pressed; prophyll 14–15(–25) cm long, 1.5 Parque Nacional de Guatopo, N of Altagracia de Ori-
cm diam., fibrous, inserted at base of pe- tuco, 400 m, 16 Feb 1979, Gentry & Troth 24823 (BH,
MO); Dist. Paez, Fila La Tigra, Quebrada San Juan, sheath, the margin slightly fibrous near
18 km al SW de Cúpira, 10804–059N, 65845–479W, base; midvein 56–95 cm 3 0.3–0.6 cm,
600 m, 2–7 Sep 1977, González & Ortega 1296 (BH,
MO); NW de Guatire, excursión Fila Juan Torres–Fila loosely brownish gray tomentose, flat to
Las Perdices, por el Rı́o Guayabal hacia el pueblo channeled adaxially, rounded to flat ab-
Guayabal, 108319N, 668209O, 900–1100 m, 19–22 Feb axially; blade 0.70–1.20 3 0.23–0.30 m,
1993, Meier 3386 (NY, VEN); Parque Nacional de dull green adaxially, grayish green and cov-
Guatopo, steep slopes bordering Rı́o Santa Cruz, be-
tween Santa Teresa and Altagracia de Orituco, 14.5 km ered with a whitish farinose indumentum
from Los Alpes, 12 km from Rancherı́a Mi Querencia, abaxially, deeply bifid at apex for ca. 42
520 m, 6 Aug 1970, Moore et al. 9601 (BH); Guatopo cm; primary veins 40–70 per side, emerg-
National Park, along S end of La Macanilla trail, 32 ing at 25–308 angle at midvein, mostly
km (by air) NW of Altagracia de Orituco, 27 Aug prominent abaxially, loosely brown tomen-
1979, Nee 17789 (NY); Guatopo National Park, region
of ‘‘La Macanilla,’’ 108069430N, 668319020W, 570 m, tose. Inflorescence branched to 1 order; pe-
18 Dec 2001, Stauffer et al. 822 (MY, MYF, VEN, Z); duncle 85–95 cm long, 1.5–2 cm diam.,
Parque Nacional Guatopo, along Rı́o Santa Cruz, be- dorsiventrally compressed; prophyll 27–35
tween Santa Teresa and Altagracia de Orituco, 450 m, cm long, 2–3.5(–5) cm diam., chartaceous,
27 Oct 1966, Steyermark 97573 (G, US); Cerros del
Bachiller, nr. E end, above Quebrada Corozal, S of inserted at base of peduncle or 1 cm above
Santa Cruz, 10 km (by air) W of Cúpira, 10899N, it; peduncular bracts 2, with indumentum
658489W, 20–700 m, 20–26 Mar 1978, Steyermark & same as on prophyll, chartaceous, brown;
Davidse 116530 (BH, MO); Cerros del Bachiller, W first peduncular bract 45–58 3 2–2.5(–4)
sector, between base and summit, above Quebrada cm, inserted 3 cm above base of peduncle,
Bachiller, S of Caño, 1086–79N, 658539W, 20–690 m,
Steyermark & Davidse 116565 (BH, MO, NY, VEN); second peduncular bract 58–60 3 2.5–3.5
Parque Nacional Guatopo, entre Santa Teresa y Alta- cm, inserted 4 cm above base of peduncle;
gracia de Orituco, 14.5 km de Los Alpes, 520 m, 29 inflorescence rachis 5–10 cm long, 0.5 cm
Nov 1970, Steyermark & Braun, 104041 (BH, US, diam., dorsiventrally to irregularly com-
VEN); montañas de Guatopo, 22 Mar 1956, Tamayo
4177 (US); Altagracia de Orituco, Santa Teresa, pressed, pale green at anthesis, almost gla-
Parque Nacional Guatopo, 11 May 1969, van Rooden brous; rachillae 9–16, slightly pendent, the
& Akkermans 95 (U). same color as inflorescence rachis, gla-
brous; basal rachillae 28–32 cm long, 0.5
cm diam.; middle rachillae 27–29 cm long,
5. ASTEROGYNE YARACUYENSE A. J. Hend. &
0.5 cm diam.; apical rachillae 24–28 cm
Steyerm. (Fig. 2D–F)
long, 0.5 cm diam., the sterile apex acute,
0.2–0.6 cm; flower pits spirally arranged,
Asterogyne yaracuyense A. J. Hend. &
2–3 mm apart from each other, covered
Steyerm., Brittonia 38: 309–313. 1986.
with an inflexed lower lip, 2.5–3 3 2–5
TYPE: VENEZUELA. Yaracuy: Cerro La
mm. Staminate flowers obovoid, 6–8 mm
Chapa, selva nublada al norte de Nirgua,
long, 2 mm diam., slightly fragrant, white
1200–1400 m, 9–10 Nov 1967, J. A.
at anthesis, the pedicel to 1 mm long; sepals
Steyermark, G. Bunting & J. G. Wessels
oblong-obovoid, 4 3 1–2 mm; petals ob-
Boer 100311. (HOLOTYPE: VEN; ISOTYPE:
long-obovoid, 6–7 3 1.5–2 mm; stamens
BH)
8–10; filaments united into a tube to 4 mm
Stem solitary, erect, sometimes basally long, the free parts 3–3.5 mm long, 0.5 mm
decumbent, 3–6 m tall, 5–8 cm diam., diam., white, conspicuously thinner at apex,
brown, occasionally with basal and lateral lacking a papillate epidermis; connective 1
vegetative branches; internodes 2–3 cm mm long; thecae 1.1–1.2 mm long; pistillode
long; aerial roots forming a short basal 2 mm long. Pistillate flowers oblong-ovoid
cone; pneumatodes very short. Leaves 18– in bud, 4 mm long, 3 mm diam., the pedicel
25(–31), reddish brown when young, green to 1 mm long; sepals oblong-ovate, 3 3 2–
when mature, the older ones briefly persis- 2.5 mm; petals oblong, 3 3 2 mm; ovary 1
tent; sheath 12–30 3 7–9 cm, reddish mm long, 1.2 mm diam.; style 1.5 mm long;
brown tomentose, the margins slightly fi- stigmatic branches 1 mm long; staminodial
brous; petiole (18–)25–35 3 1–1.5 cm, flat tube 2 mm long, the lobes 7–8 or some-
to channeled adaxially, angled to rounded times fewer by congenital union of 2 of
abaxially, the indumentum same as on them, 2 mm long. Fruits not known.
Local names and uses. not reported. Naturales Renovables (MARNR) and the
Distribution and ecology. Known only Instituto Nacional de Parques (Inparques)
from the understory of the Cerro La Chapa kindly provided the permits to collect in
cloud forest, in the northern Coastal Cor- Venezuela.
dillera, state Yaracuy, Venezuela; recently
reported on the Cerro El Zapatero of the Literature Cited
same state (W. Meier, pers. comm.) (Fig. 4);
between 1200–1400 m. Appun, K. F. 1871. Unter den Tropen. Hermann Cos-
tenoble, Jena.
Additional specimens examined: VENEZUELA. Asmussen, C. B. 1999a. Toward a chloroplast DNA
Yaracuy: Dist. Nirgua, selva nublada del Cerro La phylogeny of the tribe Geonomeae (Palmae). Pages
Chapa, N of Nirgua, 12 Nov 1985, Henderson 502 121–129. In: A. Henderson & F. Borchsenius, ed-
(MYF, NY); 5 km N of Nirgua, 10812–139N, 688379W, itors. Evolution, variation, and classification of
28 Nov–1 Dec 1996, Stauffer et al. 265, 267 (VEN- palms. Mem. New York Bot. Gard., vol. 83. Bronx,
Z), 5 km N of Nirgua, 10812–139N, 688379W, 4 Apr New York.
1997, Stauffer et al. 319 (VEN), 5 km N of Nirgua, ———. 1999b. Relationships of the tribe Geonomeae
10812–139N, 688379W, 27 Dec 2001, Stauffer & Stauf- (Arecaceae) based on plastid rps16 DNA sequenc-
fer 823 (MY, MYF, VEN, Z), 9–10 Nov 1967, Stey- es. Acta Bot. Venez. 22: 65–76.
ermark et al. 100307 (BH, VEN), 100311 (VEN, HO- ———, W. J. Baker & J. Dransfield. 2000. Phylog-
LOTYPE; BH, ISOTYPE), 108159N, 688359W, 9–10 Nov eny of the palm family (Arecaceae) based on rps16
1967, Wessels Boer et al. 2032 (BH, MER, NY, U), intron and trnL-trnF plastid DNA sequences. Pages
2036 (BH, MER, NY, U); Cerro Picacho, selva nub- 525–537. In: K. L. Wilson & D. A. Morrison, ed-
lada en la fila, N de Nirgua, 1200–1360 m, 11 Mar itors. Systematics and evolution of monocots. CSI-
1967, Steyermark & Bunting 97661 (BH, US, VEN). RO, East Melbourne, Victoria, Australia.
——— & M. Chase. 2001. Coding and noncoding
plastid DNA in palm systematics. Amer. J. Bot. 88:
Acknowledgments 1103–1117.
Baker, W. J., C. B. Asmussen, S. Barrow, J. Drans-
Special thanks to the curators of the her- field & T. A. Hedderson. 1999. A phylogenetic
baria AAU, B, BH, BM, C, CAY, CR, G, study of the palm family (Palmae) based on chlo-
K, MER, MO, MYF, NY, P, U, US, VEN, roplast DNA sequences from the trnL-trnF region.
WIS, and Z. Hector May kindly provided Pl. Syst. Evol. 219: 111–126.
information on specimens deposited at Bentham, G. & J. D. Hooker. 1883. Genera Plantar-
um. Vol. 3(2). Reeve & Co., London.
BRH and Mireya Correa provided databas- Bernal, R. 1989. Endangerment of Colombian palms.
es containing information on the specimens Principes 33: 113–128.
deposited at PMA and SCZ. Thanks to Ur- Braker, E. & R. Chazdon. 1993. Ecological, behav-
sula Hofmann for providing information on ioral and nutritional factors influencing use of
the specimens at GOET, and to Mary En- palms as host plants by a neotropical forest grass-
hopper. J. Trop. Ecol. 9: 183–197.
dress and Jean-Jacques de Granville for Braun, A. 1970. Palmas cultivadas de Venezuela. Acta
constructive comments on the manuscript. Bot. Venez. 5: 1–94.
Fred J. Stauffer, L. Pappaterra, Ricarda Ri- Brown, K. S. 1982. Paleoecology and regional pat-
ina, Rodrigo Duno, and W. Meier helped terns of evolution in neotropical forest butterflies.
during field work in Venezuela, and Rudolf Pages 255–308. In: G. T. Prance, editor. Biological
diversification in the tropics. Columbia University
Schmid kindly provided photos and infor- Press, New York.
mation on Asterogyne martiana in Central Burret, M. 1930. Geonomeae Americanae. Bot. Jahrb.
America. Mike Thiv is greatly thanked for Syst. 63: 123–270.
helping with the phylogenetic analysis. Ma- Chazdon, R. L. 1986. Physiological and morpholog-
deline Harley provided important com- ical basis of shade tolerance in rain forest under-
story palms. Principes 30: 92–99.
ments on the pollen morphology. Urs Jauch
de Granville, J. J. & A. Henderson. 1988. A new
provided guidance with the SEM work. species of Asterogyne (Palmae) from French Gui-
Visits to AAU, BM, C, K, NY, and US by ana. Brittonia 40: 76–80.
F. W. Stauffer were granted by the Inter- Dransfield, J. & N. Uhl. 1998. Palmae. Pages 306–
national Palm Society (IPS), Program for 389. In: K. Kubitzki, editor. Families and genera of
Short Term Visits of the Smithsonian Insti- vascular plants, flowering plants: monocotyledons.
Vol. 4. Springer, Berlin.
tution (NMNH), and the Kew Latin Amer- Hahn, W. J. 2002. A molecular phylogenetic study of
ica Research Fellowships Program. The the Palmae (Arecaceae) based on atpB, rbcL, and
Ministerio del Ambiente y de los Recursos 18S nrDNA sequences. Syst. Biol. 51: 92–112.
Henderson, A. 1986. A review of pollination studies ———. 1983. Septal nectaries of Asterogyne martiana
in the Palmae. Bot. Rev. 52: 221–259. and other Palmae. Principes 27: 168–174.
——— & M. Balick. 1987. Notes on the palms of the Seubert, E. 1998. Root anatomy of palms. IV. Are-
Amazonia Legal. Principes 31: 116–122. coideae, part 1, general remarks and descriptions of
———, G. Galeano & R. Bernal. 1995. Field guide the roots. Fedde Rep. 109: 89–127.
to the palms of the Americas. Princeton University Stauffer, F. W. & R. Duno de Stefano. 1998. Notes
Press, Princeton. on the conservation status of Asterogyne yaracu-
——— & J. Steyermark. 1986. New palms from Ve- yense in Venezuela. Principes 42: 57–58.
nezuela. Brittonia 38: 309–313. ———, R. Rutishauser & P. K. Endress. 2002. Mor-
Johnson, D., editor. 1996. Palms, their conservation phology and development of the female flowers in
and sustained utilization: status survey and conser- Geonoma interrupta (Arecaceae). Amer. J. Bot. 89:
vation action plan. IUCN/SSC Palm Specialist 220–229.
Group, Gland, Switzerland. Steyermark, J. 1979. Plant refuge and dispersal cen-
Klotz, L. 1978. The number of wide vessels in petiolar ters in Venezuela: their relict and endemic element.
vascular bundles of palms: an anatomical feature of Pages 185–221. In: K. Larsen & L. B. Holm-Niel-
systematic significance. Principes 22: 64–69. sen, editors. Tropical botany. Academic Press, New
Knudsen, J. 1999. Floral scent chemistry in geono- York.
moid palms (Palmae: Geonomeae) and its impor- Swofford, D. L. 1998. PAUP*. Phylogenetic analysis
tance in maintaining reproductive isolation. Pages using parsimony (and other methods). Version 4.
141–168. In: A. Henderson & F. Borchsenius, ed- Sinauer Associates, Sunderland, Massachusetts.
itors. Evolution, variation, and classification of Tomlinson, P. B. 1961. Palmae. Anatomy of the
palms. Mem. New York Bot. Gard., vol. 83. Bronx, monocotyledons. Clarendon Press, Oxford.
New York. ———. 1966. Notes on the vegetative anatomy of Ar-
Lewis, C. E. & J. J. Doyle. 2001. Phylogenetic utility isteyera spicata (Palmae). J. Arnold Arbor. 47: 23–
of the nuclear gene malate synthase in the palm 29.
family (Arecaceae). Molec. Phyl. Evol. 19: 409– Uhl, N. 1966. Morphology and anatomy of the inflo-
420.
rescence axis and flowers of a new palm, Aristeyera
Maddison, W. P. & D. R. Maddison. 2000. Mac-
spicata. J. Arnold Arbor. 47: 9–22.
Clade. Version 4.0. Sinauer Associates, Sunder-
——— & J. Dransfield. 1987. Genera Palmarum: a
land, Massachusetts.
classification of palms based on the work of H. E.
Moore, H. E. 1966. Aristeyera, a new genus of geon-
Moore, Jr., L. H. Bailey Hortorium, and the Inter-
omoid palms. J. Arnold Arbor. 47: 1–8.
———. 1967. Two new American palms. J. Arnold national Palm Society. Allen Press, Lawrence, Kan-
Arb. 48: 143–151. sas.
———. 1978. What’s in a name. Principes 22: 112– ———, ———, J. I. Davis, M. A. Luckow, K. S.
113. Hansen & J. J. Doyle. 1995. Phylogenetic rela-
Nehlin, S. 1994. A gem for tissue culture: Asterogyne tionships among palms: cladistic analyses of mor-
spicata of Venezuela. Principes 38: 211–213. phological and chloroplast DNA restriction site
Prance, G. T. 1982. Forest refuges: evidence from variation. Pages 623–661. In: P. J. Rudall, P. J.
woody angiosperms. Pages 137–158. In: G. T. Cribb, D. F. Cutler & C. J. Humphries, editors.
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tropics. Columbia University Press, New York. Botanic Gardens, Kew.
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15: 266–275. pollination in some palms. Biotropica 9: 170–190.
Schmid, R. 1970a. Notes on the reproductive biology Wendland, H. 1856. Einige neue Palmen Amerikas.
of Asterogyne martiana (Palmae). I. Inflorescence Linnaea 28: 333–352.
and floral morphology; phenology. Principes 14: Wessels Boer, J. G. 1968. The geonomoid palms.
3–9. Verh. Kon. Ned. Akad. Wetensch, Afd. Natuurk.,
———. 1970b. Notes on the reproductive biology of Tweede Sect. ser. 2, 58: 1–202.
Asterogyne martiana (Palmae). II. Pollination by ———. 1988. Palmas indı́genas de Venezuela. Pittier-
syrphid flies. Principes 14: 39–49. ia 17: 1–332.
Numerical List of Species

1. Asterogyne guianensis Granv. & A. J. 3. A. ramosa (H. E. Moore) Wess. Boer
Hend. 4. A. spicata (H. E. Moore) Wess. Boer
2. A. martiana (H. Wendl.) H. Wendl. ex 5. A. yaracuyense A. J. Hend. & Stey-
Hemsl. erm.
Exsiccatae
Only the first collector has been mentioned.

Agostini, G. 10 (4) 40972, 41092, 48546, 56871, 63735 (2);
Aguilar, R. 2140, 2318 (2) 24823 (4).
Alverson, W. 94 (2) Gómez, L. D. 19506, 19655, 20341 (2)
Anderson, R. 126 (2) González, A. 1296 (4)
Archer, W. 1978 (2) Gordon, B. 70 (2)
Aristeguieta, L. 1772, 3185 (4) Grayum, M. 4052 (2)
Aulestia, C. 350, 455, 508, 1306 (2) Grijalva, A. 262, 380, 3481 (2)
Baker, R. 89021 (2) Gudiño, E. 1283 (2)
Barfod, A. 13 (2) Haber, W. 11580 (2)
Beach, J. H. 16545 (2) Hamilton, C. 1087, 2927, 2945 (2)
Bernal, R. 256, 259, 887, 1327, 1380 (2) Hammel, B. 1682, 1797, 4863, 5153,
Bernardi, A. 5633 (4) 12377(2), 12648 (2)
Blanc, M. 189 (1) Hari, P. s.n. (2)]
Bonifaz, C. 3758 (2) Henderson, A. 53, 714, 3048 (2); 502 (5)
Borchsenius, F. 234, 242 (2) Herrera, H. 1188, 1332 (2)
Bunting, G. 401 (2) Holdridge, L. 5122 (2)
Burger, W. 5599, 8076, 10616, 11173 (2) Holm, R. 914, 970 (2)
Callejas, R. 2759, 4220, 8528, 9202 (2) Hoover, W. 1241 (2)
Chacón, A. 973 (2) Janzen, D. 11098 (2)
Chacón, E. 1958 (2) Juncosa, A. 633, 732, 1311, 1604, 1720,
Chacón, I. 1311, 1615 (2) 2532 (2)
Chazdon, R. 16 (2) Killip, E. 35293 (2)
Churchill, H. 4122, 4186 (2) Knapp, S. 1417, 1772, 3777 (2)
Cogollo, A. 4290 (2) Knudsen, J. 604, 620, 623 (2)
Cook, O. F. 61, 84, s.n. (2) Koch, R. 5048 (2)
Correa, M. 624 (2) Kress, W. J. 3778 (2)
Croat, T. 9933, 13202, 13884, 16834, Lehmann, F. C. 708 (2)
21997, 24524, 26052, 26161, 27201, Lems, K. 5199 (2)
33683, 35353, 35740, 36459, 36461, Lent, R. 80 (2)
59889, 67689, 68156, 68405 (2) Levy, P. 505 (2)
Cuatrecasas, J. 17375, 21300, 24195 (2) Lewis, W. 5558 (2)
Danin, A. 77–5–2, 77–14–12 (2) Liesner, R. 2924, 3223, 14825 (2)
Davidse, G. 23727, 31109, 31267 (2) Maas, P. J. M. 2803 (2)
de Granville, J. J. 7124, 7124-A (1) McDonagh, J. 246, 322 (2)
de Nevers, G. 3700, 3795, 3974, 4460, McDowell, T. 886 (2)
5209, 5556, 5881, 5896, 5941, 5942, McPherson, G. 9716, 8080 (2)
6111, 6167, 6282, 6323, 7914, 8320 (2) Meier, W. 3386 (4)
Denslow, J. 2745 (2) Miller, J. 1095 (2)
Dodge, C. 9284 (2) Molina, A. 17559 (2)
Dransfield, J. 4859 (2) Mondragón, M. 119 (2)
Duke, J. 11334, 11336 (2) Monsalve, M. 671, 927 (2)
Elias, T. 1769 (2) Moore, H. E. 6530, 6535, 6717, 8213, 9420
Folsom, J. 5901, 6179 (2) (2); 9601 (4)
Fonnegra, R. 2839, 3024, 4756 (2) Morales, J. 1118 (2)
Forero, E. 8975, 9039 (2) Moreno, P. 12280, 12580, 12771, 12979,
Fuchs, H. P. 21808, 21994 (2) 12985, 13032, 13151, 13287, 14821,
Gentle, P. 3743, 8635, 9072, 9074, 9669 (2) 15013, 19142, 20602, 20640, 20661,
Gentry, A. 2803, 7542, 8818, 8868, 17445, 23621, 23890, 24015, 26083 (2)
Mori, S. 2312, 2319, 4095, 4589, 4905, Stauffer, F. W. 824 (3); 822 (4); 265, 267,
6383 (2) 319, 823 (5)
Navarro, E. 76 (2) Stevens, W. D. 4810, 4943, 6336, 6795,
Nee, M. 17789 (4), 27905 (2) 6820, 8322, 12080, 12246, 12686,
Neill, D. 1723, 4442, 7151 (2) 13065, 13505, 14927, 19343, 20743,
Oersted, A. 6566 (2) 20936, 23489, 23651, 24026, 24331,
Ortiz, F. 297, 733 (2) 24451 (2)
Picado, A. 123 (2) Stevenson, D. 1204 (2)
Pipoly, J. 5111, 5323, 6112, 6143, 6168, Steyermark, J. 94803, 96088, 120915,
121713 (3); 39785, 90010, 97573,
6223, 6290 (2)
104041, 116530, 116565 (4); 97661,
Pittier, H. 4226, 16658, 16717, 16922 (2) 100307, 100311 (5)
Porter, D. 4736 (2) Sudgen, A. 351 (2)
Quesada, F. 43 (2) Sullivan, G. 159 (2)
Ramı́rez, J. 46, 167, 1131, 1364, 1770, Sytsma, K. 1660 (2)
1860 (2) Tamayo, F. 4177 (4)
Ramı́rez, V. 183 (2) Tessene, M. 1329 (2)
Read, R. 640 (2) Thomsen, K. 187 (2)
Rı́os, D. 187 (2) Tipaz, G. 2250, 2693 (2)
Robles, R. 1148, 1192 (2) Tomlinson, P. B. s.n. (2)
Rowlee, W. 1010 (2) Utley, K. 1093, 5975 (2)
Rueda, R. 2670, 3110, 3459 (2) Valverde, O. 665 (2)
Salick, J. 8075, 8106 (2) van Rooden, J. 95 (4)
Sandino, J. 4782 (2) Vincelli, P. 164, 228 (2)
Schipp, W. 392 (2) von Wedel, H. 952, 1108, 2186 (2)
Watson, S. 238 (2)
Schmalzel, R. 1156 (2) Wendland, H. 56, s.n. (2)
Seemann, B. 163, s.n (2) Wessels Boer, J. G. 2032, 2036 (5)
Seymour, F. 5905 (2) Wilbur, R. L. 15071 (2)
Shepherd, J. 863 (2) Wilson, P. 224 (2)
Soza, D. 347 (2) Witherspoon, J. 8322 (2)
Sperry, J. 606 (2) Yuncker, T. 5002 (2)
Standley, P. 25015, 53156 (2) Zarucchi, J. 5056 (2)

Flora Anatomy Alliaceae

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Flora Anatomy Alliaceae

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A revision of Asterogyne (Arecaceae:

Stauffer, F. W. (Institute of Systematic Botany, University of Zurich, Zolliker-

Stauffer, F. W. (Institute of Systematic Botany, University of Zurich, Zolliker-

Brittonia, 55(4), 2003, pp. 326–356. ISSUED: 07 October 2003

FIG. 4. Distribution of Asterogyne in Central and South America.

critical-point dried, and sputter-coated with

Morphology and Anatomy

1. Leaf blade: (0) entire-bifid; (1) divided

‘‘Calyptronoma occidentalis type,’’ occur- tegument could be identified. It is five to

Small to medium-sized, solitary, un-

sally, narrow, elongate, keeled dorsally; 1. Stamens 6–12; inflorescences branched,

1. ASTEROGYNE GUIANENSIS Granv. & A. J. in bud, 5 mm long, 2.5–3 mm diam., the

Numerical List of Species

Only the first collector has been mentioned.

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