A revision of Asterogyne (Arecaceae: Arecoideae: Geonomeae) FRED W . STAUFFER, C O N N Y B . ASMUSSEN, A N D R E W HENDERSON, AND P E T E R K . ENDRESS Stauffer, E W. (Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008, Zurich, Switzerland; email: f.stauffer@access.unizh.ch), C. B. Asmussen (Department of Ecology, Royal Veterinary and Agricultural University, 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: Geonomeae) 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. L~picata, 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 relationships, and conservation status are reviewed. Key words" Arecaceae, Palmae, Geonomeae, Asterogyne, Neotropics. Stauffer, E W. (Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008, Zurich, Switzerland; email: f.stauffer@access.unizh.ch), C. B. Asmussen (Department of Ecology, Royal Veterinary and Agricultural University, 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) & R 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.--Se presenta una revisi6n taxon6mica del gdnero neotropical Asterogyne Hook. (Arecaceae: Arecoideae: Geonomeae). El g6nero se caracteriza por un carficter autoapom6rfico, tecas separadas pot un conectivo bffido, ademfis de tenet tallos de pequefio a mediano tamafio, lfiminas foliares enteras y bffidas, f6veas florales cubiertas por un labio superior redondeado que sobrelapa el labio interior, flores pistiladas con 16bulos estaminodiales dispuestos de manera estrellada. Para este tratamiento se reconocen cinco especies: una de ellas (Asterogyne martiana) ampliamente distribuida desde Belice en Amdrica Central hasta el norte de Ecuador, tres especies (A. ramosa, A. spicata, and A. yaracuyense) enddmicas de pequefias fireas en la Cordillera de la Costa de Venezuela y una especie (A. guianensis) enddmica del oriente de la Guyana Francesa. Se revisa la historia taxondmica, morfolog/a, biologfa reproductiva, distribuci6n, ecologfa, relaciones intragendricas y estado de conservaci6n. Brittonia. 55(4), 2003, pp. 326 356. 92003, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. ISSUED: 07 October 2003 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) A I 327 B 1 1 P 41 i i j D ':.'::7c'=~ 22o~2,2~'::z'=~ ._. c c.': : ; ~ - ' '::x . . . . . . . . . . -- 1. Type specimens of Asterogyne. A, B. Isotype of A. guianensis (de Granville 7124, K); C. Isotype FIG. of A. martiana (Wendland 56, K/; D-F. Isotype of A. ramosa (Steyermark et al. 96088, VEN). Scale bars: 8 cm. The small genus A s t e r o g y n e H. Wendl. ex Hook. f. is one o f six g e n e r a in the neotropical tribe G e o n o m e a e (Dransfield & Uhl, 1998; H e n d e r s o n et al., 1995) and consists of five species in Central A m e r i c a and northern South A m e r i c a (Figs. 1 A - E 2 A - F , 4). In a letter to J. D. H o o k e r in 1883, W e n d l a n d p r o p o s e d A s t e r o g y n e as a new genus to be i n c l u d e d in the m o n u mental treatment of Genera Plantarum (Bentham & Hooker, 1883). The genus was d e s c r i b e d with several character states a l r e a d y k n o w n in other genera of G e o n o meae, e.g., bifid s i m p l e l e a f b l a d e s and stam i n o d i a b a s a l l y united into a tube. A l though m o d e r n t r e a t m e n t s of the genus have p r o p o s e d several characters to differentiate A s t e r o g y n e f r o m other g e n e r a o f G e o n o m e a e , floral pits in bud c o v e r e d by a r o u n d e d u p p e r lip that overlaps the l o w e r lip (Fig. 9D) and free thecae on a bifid c o n n e c t i v e (Figs. 8B, 10D, 12C, 14C), have a l w a y s been r e g a r d e d as the two m o s t r e l e v a n t f e a t u r e s ( B u r r e t , 1930; H e n - 328 BRITTONIA [VOL. 55 ': .<ij i FIG. 2. Type specimens of Asterogyne. A-C. lsotype of A. ~picata (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 & Dransfield, 1987; Wessels Boer, 1968). A s t e r o g y n e was published without a type species in the original description in 1883. The typification of the new genus was made two years later in 1885 by Wendland, with the transfer o f G e o n o m a m a r t i a n a to the genus A s t e r o g y n e . The type specimen selected by H. Wendland for this taxon was obtained from a plant cultivated at the gardens of Herrenhausen (Fig. 1C). This plant at Herrenhausen originated from plants cultivated by L. van Houtte in Gent, apparently from seeds sent by Warscewicz during his trip to Central America (Wendland, 1856). AI- though there was no original explanation of the generic name, it was probably suggested because o f the starlike shape of the staminodial lobes o f the pistillate flowers (Moore, 1978) (Figs. 7E 8D). Moore (1966) described the genus A r i s t e y e r a , separating it from A s t e r o g y n e on the basis of differences in the number of stamens and staminodes. Wessels Boer (1968) observed great plasticity of these characters, especially in flowers of A r i s t e y e r a spicata, and considered that they were inconsistent for a delimitation at the generic level. He merged both genera and argued that the number of stamens and staminodes reported by Moore 2003] S T A U F F E R ET AL.: A S T E R O G Y N E ( A R E C A C E A E ) 329 | ~ ~.,~/. ~ "*,," .~ ~''~ ~ y-- FIG, 3. grain of A. spicata. G. B,D, EH Pollen of A s t e r o g y n e . A. Pollen grain of A. m a r t i a n a . B. Detail of exine in A. m a r t i a n a . C. Pollen r a m o s a . D. Detail of exine in A. r a m o s a . E. Pollen grain of A. ,spicata. F. Detail of exine in A. Pollen grain of A. y a r a c u y e n s e . H. Detail of exine in A. y a r a c u y e n s e . Scale bars: A, C, G = 5 p~m; = 1 b t m ; E = 6btm. 330 BRITTONIA 20~ 90~ J / ~ 80~ ,'-~_ ~ [VOL. 55 70~ 60~ ) 10o - A ~ ~ 5o "~ Asterogyne r162A. guianensis 9 A. martiana FIG. 4. Distribution of Asterogyne in Central and South America. 78 t Aslerogyne guianensis A. spicata A. martiana A. YOOTOSa 97 90 A. .vs Geonoma interrupta C~/vptron~mu~I'ivalis FIG. 5. Strict consensus tree of the two most parsimonious trees based on a phylogenetic analysis of morphological data. Bootstrap values are given on the branches. (1967) for A s t e r o g y n e r a m o s a filled the gap between the two genera. Due to the variability of stamens and staminodes o f Asterogyne, Wessels Boer (1968, 1988) did not recognize the two populations in the states of Sucre and Yaracuy of the Venezuelan Coastal Cordillera (Fig. 4) as representing two different species, but included them in a broadly defined A. ramosa. After the de- scription o f the widespread type species, A. m a r t i a n a (Fig. 4), in 1856, four additional taxa were added to the genus within a relatively short period, from 1967 to 1988. These recently discovered species show rem a r k a b l y limited distributions for the neotropical palms, and compared to A. martiana, they are known from few herbarium specimens (limited to two for A. g u i a n e n - 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) FIG. 6. Habit ofAsterogyne guianesis. (Photo: J. J. de Granville, Camopi River, French Guiana). Scale bar: 80 cm. sis); thus their morphological and anatomical features are incompletely known. This revision aims to improve the taxonomic knowledge of the genus and to compile and reevaluate the published information on its systematics and natural history. Materials and Methods Morphology and distribution of the species were studied based on 334 specimens (see Exsiccatae) deposited at AAU, B, BH, BM, C, CAY, CR, G, K, MER, MO, M Y E NY, R U, US, VEN, WlS, and Z. A review of critical literature for the genus (Burret, 1930; de Granville & Henderson, 1988; Henderson et al., 1995; Uhl & Dransfield, 1987; Wessels Boer, 1968) was made, Additional information on the distribution of A. martiana was obtained from the curators at BRH, PMA, and SCZ, and of A. guianensis from J. J. de Granville (CAY). Specimens were collected in Venezuela by the first author in 1996, 1997, 2001, and 2002. For scanning electron microscopy, the dissected specimens were dehydrated, 331 critical-point dried, and sputter-coated with gold. Micrographs were obtained with a Cambridge $4 scanning electron microscope. For anatomical investigations, flowers in bud or at anthesis were dehydrated and embedded in Kulzer's Technovit 7100 (2-hydroxyethyl methacrylate, HEMA). The material was sectioned at 7 microns using a rotary microtome, then stained with ruthenium red and toluidine blue and e m b e d d e d in Histomount. A preliminary phylogenetic analysis o f 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 irlcluded Geonoma interrupta and Calyptronoma rivalis as outgroups. Nineteen morphological characters (Table I), 10 o f them parsimony-uninformative, were included in the data matrix (Table II) constructed in McClade 4.0b10 (Maddison and Maddison, 2000). Clear gaps between the character states of quantitative features were observed. A parsim o n y analysis was performed using Paup 4.0 (Swofford, 1998). All the characters were treated as " u n o r d e r e d " and were equally weighted. Support for the clades was calculated by doing a 10,000 replicate bootstrap analysis using the branch and bound option. Morphology and Anatomy STEMS AND ROOTS Stems of Asterogyne are solitary and mostly erect; only A. martiana and A. yaracuyense may be slightly decumbent at the base. Basal vegetative branching was des c r i b e d by H e n d e r s o n and S t e y e r m a r k (1986) for A. yaracuyense but the branches do not fully develop into mature stems. Lateral vegetative branches are normally present in A. yaracuyense and are especially numerous in A. ramosa, in which 5 to 30 branches were observed, attached to the stem, 2 0 - 4 0 cm below the crownshaft (Fig. 9A). Each of these lateral branches had 4 to 7 small bifid simple leaf blades, and were richly rooted. Our observations showed that these lateral branches do not develop into 332 BRITTONIA [VOL. 55 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 = 1.2 m: C = 5 cm: D = 0.4 cm: E, E G 0.3 cm. n e w plants. S t e m a n a t o m y r e m a i n s unk n o w n for the genus. A d v e n t i t i o u s roots that form a b a s a l c o n e are p r e s e n t i n several species. In Asterogyne guianensis, the aerial roots are r e m a r k a b l y long, to 90 cm, a n d c o n t a i n w e l l - d e v e l o p e d white p n e u m a t o d e s (Fig. 6). S e u b e r t (1998) studied the root a n a t o m y o f A. martiana and f o u n d that it is similar to that o f other G e o n o m e a e . T h e r h i z o d e r m i s cells are thinw a l l e d a n d d i s a p p e a r in m a t u r e roots. In t r a n s v e r s e section, the e x o d e r m i s a n d the outer cortex, w h i c h is r e d u c e d to a f e w layers, are n o t distinct. In l o n g i t u d i n a l section, the e x o d e r m i s a n d outer cortex are distinct. T h e cells o f the o n e - l a y e r e d e x o d e r m i s are stretched with b l u n t ends; those o f the outer cortex are l o n g stretched with p o i n t e d ends. T h e i n n e r cortex is u s u a l l y h o m o g e n e o u s , and the a e r e n c h y m a , only w e a k l y developed, m a y o c c u r in the center. In contrast to all other g e n e r a of the A r e c o i d e a e , the cells o f the i n n e r cortex of the G e o n o m e a e are m o d e r a t e l y t h i c k - w a l l e d , a n d t h i n w a l l e d cells are lacking. N a r r o w fibers are scattered t h r o u g h o u t the i n n e r cortex. End o d e r m i s c e l l s are t a n n i f e r o u s , r a d i a l l y stretched, a n d u-shaped, with thin outer walls a n d thick i n n e r walls. T h e pericycle is o n e - l a y e r e d and the v a s c u l a r c y l i n d e r shows the n o r m a l pattern o b s e r v e d in the f a m i l y (Seubert, 1998). LEAVES T h e leaves, at least the y o u n g ones, in Asterogyne are spirally arranged and rather 2003] S T A U F F E R ET AL.: A S T E R O G Y N E ( A R E C A C E A E ) 333 i\i:l7 ( FIG. 8. Flowers in Asterogyne martiana. A. Staminate flower. B. Stamen. C. Epidermis of the filament. D. Pistillate flower. E. O v a r y in abaxial view. F. O v a r y 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 = 1 mm; B = 300 ~m; C = 25 ixm: E, E 1 - 200 ttm; G 12 I~m: J 50 ~rn. 334 BRITTONIA [VOL. 55 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: E J. Stauffer, Cerro Humo, Venezuela]. Scale bars: A = 28 cm: B = 18 cm; C = 5 cm; D 7 ram; E 2 ram; F 3 ram. erect a n d often purplish. T h e b l a d e is norm a l l y bifid a n d simple b u t it m a y be long i t u d i n a l l y split with age or d a m a g e d by the w i n d (Figs. 6, 7A). L o n g i t u d i n a l splits were e s p e c i a l l y c o n s p i c u o u s in s o m e c o l l e c t i o n s o f A. martiana (Stevens 13505; Q u e s a d a 43) f r o m C o s t a Rica, a n d A. ramosa (Steyermark & Liesner 120915) a n d A. yaracuyense (Stauffer et al. 267) f r o m V e n e z u e l a . A c c o r d i n g to Wessels B o e r (1968) the blades o f A. martiana a n d A, spicata show a h y p o d e r m i s (sensu T o m l i n s o n 1961, i.e., a c o l o u r l e s s h y p o d e r m a l cell layer) o n l y on the a b a x i a l side. T h e m e s o p h y l l lacks a distinct p a l i s a d e layer and the cells are s o m e what t r a n s v e r s e l y extended. T h e fibers are n u m e r o u s , o f t e n solitary, each with a rather wide l u m e n , scattered t h r o u g h o u t the mesophyll b u t c o m m o n l y f o u n d in the adaxial layer. S o m e o f the v a s c u l a r b u n d l e s are v e r y small, n o t p r o d u c i n g r i b b i n g o f the l a m i n a . W e s s e l s Boer (1968) n o t e d that the leaf a n a t o m y o f Asterogyne shared m o s t o f the characters already o b s e r v e d in other 2003] S T A U F F E R ET AL.: A S T E R O G Y N E ( A R E C A C E A E ) 335 : .~c'-'~ H ~ , ' r ~ ' ~ ~;~j,~ ~,~ ~--'~ 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..I. 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 = 450 ixm; B, C 1 ram: E, H - K = 400 p~m: F 500 txm; G = 25 l,zm. 336 BRITTONIA [VOL. 55 FIG. ll. Habit, leaf, inflorescence, and fruits of Asterogyne spicata. A. Habit. B. Detail of blade. C. Inflorescence and infructescence. D. Detail of rachilla at staminate anthesis. E. Detail of rachilla with fruits. (Photos: E J. Stauffer, Parque Nacional Guatopo, Venezuela). Scale bars: A = 1.2 m: B = 20 cm: C = 8 cm; D, E 4 mm. G e o n o m e a e , but p o i n t e d out that the onec e l l - l a y e r e d a b a x i a l h y p o d e r m i s o f the blades c o u l d be interpreted as less developed w i t h i n the tribe. Vegetative a n a t o m y of A. spicata (as Aristeyera spicata) was studied b y T o m l i n s o n (1966). His observation o f two wide vessels per b u n d l e in the petiole o f A . spicata was also f o u n d in other species of G e o n o m e a e b y K l o t z (1978). INFLORESCENCES AND FLORAL TRIADS Inflorescences of Asterogyne are always interfoliar, solitary, and p r o t a n d r o u s (Figs. 7A, 9A, 11A). O n c e - b r a n c h e d i n f l o r e s c e n c es are p r e s e n t in A. martiana, A. ramosa, and A. yaracuyense (Figs. 7C, 9C, 13B), while A. guianensis a n d A. spicata show w e l l - d e v e l o p e d spikes, w h i c h are thyrses in a c o m p a r a t i v e m o r p h o l o g i c a l sense (Figs. 1B, 2C, 11C). A r e m a r k a b l e d e v i a t i o n f r o m the b r a n c h e d i n f l o r e s c e n c e s was o b s e r v e d in s o m e c o l l e c t i o n s of A. martiana restricted to the C o m a r c a of S a n Blas in P a n a m a (e.g., Elias 1769; de Nevers et al. 5209, 5881, 7914", McDonagh, J. et al. 246) and C o l o m b i a (Duke 11338), in w h i c h the in- 2003] S T A U F F E R ET AL.: A S T E R O G Y N E (ARECACEAE) i l D ~i 84 i 337 .... ~i:" .... , I m N FIG. 12. Flowers in Asterogyne .v~icata. 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 Stm~ler & Stauffer 822, Z). Scale bars: A, E 1 mm: B = 3 ram; C 500 ixm: D, K = 50 p,m: E G = 1.5 l n m : H = 600 p,m; I , J , L - N = 400 p,m. 338 BRITTON|A [VOL. 55 FIG. 13. Habit and inflorescence of Asterogyne yaracto,ense. A. Habit. B. Inflorescence. C. Detail of rachilla at staminate anthesis. (Photos: E J. Stauffer, Cerro Humo, Venezuela). Scale bars: A = 30 cm; B 8 cm; C 3 mm. florescences are spicate (Fig. 7B). In one of these collections (de Nevers et al. 5881) the spike is rather thick. M o r p h o l o g y and anatomy of the inflorescence axis and especially the floral vasculature of Asterogyne spicata (as Aristeyera spicata) was examined by Ubl (1966). The vascular bundles of the flowers o f this species are small because the xylem does not contain large vessels and has a few helical elements. The number of vascular traces to perianth parts is variable. In the petal tube of staminate flowers and sepals of pistillate flowers, some of the traces appear to differentiate over a long period o f time and as a result the number of traces increases with the age of the flower. The traces developing later do not become connected to the vascular strands in the floral receptacles. The peduncle of Asterogyne is long and slender. The prophyll is fibrous to membranous and normally inserted at the base of the peduncle or just above it. The inflorescences are enclosed by one or two tubular, c h a r t a c e o u s to m e m b r a n o u s , p e d u n c u l a r bracts, Asterogyne ramosa and A. yaracuyense always have two peduncular bracts, whereas the species with spicate inflorescences and A. martiana (with the exception of one collection from Panama [Pittier 4226], which has two peduncular bracts), show a reduction to only one. Moore (1967) suggested that the reduction in the number of bracts observed in A. spicam (as Aristeyera spicata) represented a derived character within the genus. As first reported by Wessels Boer (1968) for A. martiana, all the species studied in the field showed that the small unexpanded inflorescence remains straight and unfolded within closed bracts; during expansion it elongates rapidly and emerges through a terminal split o f the persistent bracts. The inflorescences are always erect at anthesis and become pendent in fruit (Fig. 7B). In branched inflorescences the rachis is normally well developed, but in A. martiana (Cuatrecasas 17375; Knapp 3777; Moore 6530) it may be almost absent and then all the rachillae appear to emerge from the apex of the peduncle. The inflorescence raches are normally salmon to pink at anthesis and become brown to reddish brown in fruit. Floral pits are spirally arranged on the rachillae. As described by Uhl and Dransfield (1987) for Welfia, Calyptrogyne, and PholMostachys, the foral pits in bud of Asterogyne are covered by an immersed, rounded, lower lip, overlapped by a short upper lip (Fig. 9D). Wessels Boer (1968) interpreted the immersed lower lips as less specialized in a morphological sense and therefore as phylogenetically basal 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) 339 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 Staz~fJer &Smuffcr 823, Z). Scale bars: A 900 Ixm: B, E l ram: C 400 ~Lm:D = 50 ixm; F 450 gtm : G, H, J = 200 ~xm; I = 300 Ixm. w i t h i n the tribe. T h e l o w e r lip rolls b a c k w h e n the f l o w e r s b e c o m e exserted. T h e unc o m m o n spicate i n f l o r e s c e n c e o b s e r v e d in one c o l l e c t i o n o f A. martiana (ide N e v e r s et al. 58811) also has the lip m a r g i n s d e n s e l y c o v e r e d by short w h i t i s h hairs. T h e hairs are l a c k i n g in the n o r m a l - b r a n c h e d inflor e s c e n c e s on o t h e r plants o f this species. A s is c o m m o n in the A r e c o i d e a e , the pits c o n tain triads c o m p o s e d o f one pistillate and t w o s t a m i n a t e flowers. E a c h triad is s u n k e n in the floral pit and r e p r e s e n t s a short cin- cinnus. B a s e d on the v a s c u l a t u r e , h o w e v e r , Uhl (1966) s u g g e s t e d that the triads in Asterogyne (as Aristeyera) o r i g i n a t e d f r o m a m o r e r a m i f i e d t y p e o f partial i n f l o r e s c e n c e . A s also r e p o r t e d by S t a u f f e r et al. (2002) for G e o n o m a interrupta, the first s t a m i n a t e f l o w e r o f the c i n c i n n u s , and the c i n c i n n u s as a w h o l e , is s u b t e n d e d by the p i t - c l o s i n g bract, w h i c h r e p r e s e n t s the l o w e r lip o f the pit. A c c o r d i n g to the i n t e r p r e t a t i o n o f W e s sels B o e r (1968), the largest bract i n s i d e the pit subtends the s e c o n d s t a m i n a t e flower, a 340 BRITTONIA [VOL. 55 TABLE I CHARACTERS AND UNORDERED CHARACTER STATES 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. Leaf blade: (0) entire-bifid: (1) divided Inflorescence branching: (0) branched; (1) spicate Upper lip of the pit: (0) absent: (1) present Peduncle (length): (0) <80 cm; (1) -->85cm Peduncular bracts (number): (0) one: (1) two First peduncular bract (length): (0) --<60cm; (1) >64 cm Number of stamens: (0) 6: (1) 8-10; (2) 11-12; (3) 21-27 Filament color: (0) white; (1) yellow Filament surface: (0) smooth; (1) papillate Anther shape: (0) divided: (1) with a bifid connective: (2) sagittate Complete fusion of the staminodial tube with the corolla: (0) absent; (1) present Upper part of the sterile androecium: (0) calyptrate: (1) lobed; (2) crenulate Complete fusion of the petals of the pistillate flowers in a tube: (0) absent; (1) present Gynoecium: (0) with 3 more or less equally developed ovaries: (1) pseudomonomerous Central protrusion between the three carpels: (0) absent: I l) present Position of the ovule: (0) at midlength of the locule: (1) basal Lateral vascular bundles in the styles at anthesis: (0) absent: (1) present Upper part of the style curved: (0) absent: (1) present Fruit length: {0) 0.3-7 cm; (1) 1.2 cm: (2) 1.5-1.8 cm: (3) 2.5 cm m e d i u m - s i z e d bract s u b t e n d s the pistillate flower, a n d a s m a l l bract r e p r e s e n t s the prop h y l l o f the pistillate flower. STAMINATE FLOWERS Two s t a m i n a t e flowers o c c u r laterally to the pistillate flower. The p e r i a n t h is c o m posed o f three b a s a l l y u n i t e d a n d a p i c a l l y free sepals a n d three b a s a l l y u n i t e d a n d apically v a l v a t e petals (Figs. 8A, 10B, 12B, 14B). T h e n u m b e r of s t a m e n s r a n g e s from six (Asterogyne martiana, Fig. 8A) to 27 (A. guianensis); the h i g h e r n u m b e r o f stam e n s b e i n g p r e s e n t in species with spicate i n f l o r e s c e n c e s . Taxa of G e o n o m e a e with u n b r a n c h e d i n f l o r e s c e n c e s a n d flowers with m o r e than six s t a m e n s are rather u n c o m m o n . T h e y are p r e s e n t in o n l y t w o species of Asterogyne (A. guianensis a n d A. spicata', Fig. 12B) a n d three species o f Geonoma (G. camana Trail, G. chococola Wess. Boer, a n d G. polyandra Skov). W e s s e l s Boer (1968) c o n s i d e r e d the i n c r e a s e d n u m b e r o f s t a m e n s in s o m e species o f Asterogyne (as Aristeyera) as a d e r i v e d c h a r a c t e r state w i t h i n the tribe. Based o n the i n c r e a s e d n u m b e r o f s t a m e n s in A. spicata, M o o r e (1966) s u g g e s t e d a closer r e l a t i o n s h i p bet w e e n Asterogyne (as Aristeyera) a n d Welfia, than with the other G e o n o m e a e . The s t a m e n s are u n i t e d into a tube up to m i d length a n d free n e a r the top (Figs. 8A, 10A, 10B, 12B, 14A). A n t h e r s o f A. yaracuyense, a n d e s p e c i a l l y A. martiana, are versatile b e c a u s e o f the t h i n n e r apical r e g i o n s o f the f i l a m e n t s (Fig. 8B). W e s s e l s B o e r (1968) f o u n d that the j u n c t i o n b e t w e e n the f i l a m e n t a n d the c o n n e c t i v e in Asterogyne is less rigid than in Geonoma. T h e separation o f a n t h e r thecae on a bifid c o n n e c t i v e in Asterogyne (Figs. 8B, 10D, 12C, 14C) has b e e n c o n s i d e r e d u n i q u e w i t h i n the G e o n o m e a e ( U h l & Dransfield, 1987). The c o n n e c t i v e is c o n s p i c u o u s l y s w o l l e n i n A. martiana. T h e anthers are i n f l e x e d t o w a r d the c e n t e r o f the flower and introrse in b u d but spread a n d extrorse at anthesis. A threel o b e d p i s t i l l o d e is n o r m a l l y f u s e d with the s t a m i n o d i a l t u b e to its m i d d l e (Fig. 12E). M o r p h o l o g y a n d vasculature o f the staminate flowers o f A. spicata (as Aristeyera spicata) were studied by U h l (1966) a n d M o o r e ( 1 9 6 6 L a n d A. martiana b y U h l and M o o r e (1977). POLLEN MORPHOLOGY P o l l e n i n Asterogyne has b e e n d e s c r i b e d b y U h l a n d D r a n s f i e l d (1987) as elliptic, m o n o s u l c a t e , with tectate exine, w h i c h is v e r m i c u l a t e or reticulate. I n a detailed study of the G e o n o m e a e , P u n t a n d W e s s e l s B o e r (1966) stated that Asterogyne species show two d i f f e r e n t p o l l e n types, the "'Asterogyne t y p e , " o c c u r r i n g in A. martiana a n d the 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) TABLE II M A T R I X O F 1 9 M O R P H O L O G I C A L C H A R A C T E R S ,SCORED F O R T H E FIVE SPECIES O F TAXA (Geonomct interrw~ta CLltVtgtotloltlCt rivalis). AND 341 Asterog)'tze AND TWO OUTGROUP T H E Q U E S T I O N M A R K ( ' ? ) I N D I C A T E S DATA N O T A V A I L A B L E Character A. A. A. A. A. G. C. Species 1 2 3 4 5 6 7 8 9 If) 11 12 13 E4 15 16 17 18 19 guianensis martiana ramosa spicata yaracuy interrupta rivalis 0 0 0 0 0 1 1 t 0 0 1 0 0 0 1 1 1 1 1 0 0 0 0 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 3 0 2 3 1 0 0 0 0 0 l 0 0 1 0 0 0 1 0 0 0 1 1 l 1 1 0 2 1 1 1 1 1 0 0 1 1 1 1 1 2 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 "? 1 1 1 1 1 0 0 0 0 0 0 1 0 ? 0 0 1 0 0 0 0 0 0 0 0 0 1 3 1 2 2 9 0 0 "Calyptronoma occidentalis type," occurring in A. spicata (as Aristeyera spicata). Pollen of four of the five species of Asterogyne was studied for this revision (,4. martiana, Knudsen & Asmussen 604; A, ramosa, Smt~'er et al. 824; A. spicata, Stauffer et al. 822; A. yaracuyense, Stauffer & Stauffer 823). The pollen (Fig. 3 A - H ) is asymmetric and monosulcate with a sulcus length as long as the main axis, The exine is tectate and finely perforate-rugulate (perforate fossulate in A. spicam). This agrees with the study of Harley (unpubl.) of A. martiana (Schipp 392). Punt and Wessels Boer (1966) described the pollen of A. martiana (Archer 1978, Schipp 392) as easily recognized by its fine, but distinct reticulum, and the distinctly visible muri (bacula), which support the reticulum. The pollen o f A . spicata (Tamayo 4177) was described by Punt and Wessels Boer (1966) as having a distinctly vermiculate ornamentation. Results of our study could not differentiate the two groups proposed by Punt and Wessels Boer (1966) and show that the pollen is rather morphologically uniform within the genus. PISTILLATE FLOWERS The perianth of the pistillate flowers is c o m p o s e d of three free, imbricate sepals and three petals, which are united to their middle (Fig. 8D). The carpels alternate with the petals. The o v a r y is superior and trilocular (Figs. 8E, 8I, 10E, 10I, 12H, 12L, 14G, 14J). Each carpel bears a pendent, anatropous, crassinucellate ovule (Fig. 10H). One of these per g y n o e c i n m develops fully to maturity. Only one well-differentiated in- tegument could be identified. It is five to six cell layers thick at midlength and six to ten cell layers thick toward the micropyle, The nucellus protrudes into the micropyle, which faces the ventral side of the locule. A well-developed septal nectary occurs between the carpel flanks (Fig. 8I). It b e c o m e s a triradiate cavity at about midlength of the ovary and extends upward and gradually outward to the apical region of the ovaries. The epithelium, especially differentiated in the gynoecium o f A . rnartiana, is formed by large columnar, uninucleate cells (Fig. 8J). The three openings o f the nectary are toward the top o f the o v a r y and are nonsecretory ducts lined by isodiametric epidermal cells. Even though Asterogyne was described (Uhl and Dransfield, 1987; Wessels Boer, 1968) as having apical styles, our study shows they are basal to lateral, which is probably a c o m m o n pattern within Geonomeae (Stauffer & Endress, unpubl, data). Epidermal protuberances at the basal and midregion of the styles were observed in anthetic flowers of A. martiana (Schmid, 1970a) and A. yaracuyense (Stauffer & Endress, unpubl, data) (Fig. 8G). These protuberances are absent in young buds and seem to develop shortly before anthesis. The three stigmatic branches are papillate and reflexed at anthesis (Fig. 8H). The staminodes are united to their middle into a tube and free above this level (Figs. 8D, 10C, 14E). The n u m b e r of staminodial lobes ranges f r o m five in A. ramosa to 22 in A. guianensis. A well-developed papillate epidermis in the staminodial lobes is present in A. spicata (Fig. 12K). Reduction of the n u m b e r o f staminodes with respect 342 BRITTONIA to the n u m b e r o f fertile stamens in the same species is c o m m o n within the genus; however, in Asterogyne martiana the n u m b e r of fertile and sterile stamens is always six (Fig. 8A, D). The m o r p h o l o g y and vasculature of the pistillate flowers of A. spicata were studied by Uhl (1966) and for A. martiana by Uhl and Moore (1977). The structure of the gynoecia in four species of Asterogyne is part of a current study including representatives of all G e o n o m e a e (Stauffer & Endress, unpubl, data). The c h r o m o s o m e number in the genus is unknown (M. R6ser, pets. comm.). FRUITS Fruits o f Asterogyne are ellipsoid to ellipsoid-ovoid, purplish, black, or garnet red drupes, with a smooth surface and a single seed (Figs. 7G, 11E). They are conspicuously keeled at the apex. The remains of the stigma are basal. The seed is ellipsoid and slightly dorsiventrally compressed. The endosperm is homogeneous (not ruminate), and the e m b r y o is basal (Uhl & Dransfield, 1987). The m o r p h o l o g y and a n a t o m y of the fruits in A. martiana and A. spicata were studied by Wessels Boer (1968). Fruits of A. martiana resemble those o f A. spicata, but the arrangement of the vascular bundles in the p a r e n c h y m a of the mesocarp is less regular. In A. spicata, bundles of fibers without conductive elements are present in the outer layer and the endocarp is not adherent to the seed (Wessels Boer, 1968). Germination is adjacent-ligular (Uhl & Dransfield, 1987). Seeds o f A . spicata have been reported to germinate within three and four months (Braun, 1970; Nehlin, 1994). Reproductive Biology What we k n o w about the reproductive biology of Asterogyne is limited almost entirely to A. m a r t i a n a ( S c h m i d , 1970a, 1970b; Uhl & Moore, 1977). Asterogyne is m o n o e c i o u s and the i n f l o r e s c e n c e s are markedly protandrous. Male and female flowers on the same inflorescence of A. martiana are rarely open at the same time, thus m a x i m i z i n g pollen transfer a m o n g different plants (Schmid, 1970a). In the species in Venezuela usually just one inflores- [VOL. 55 cence o f a plant is in flower at a time, while the other inflorescences are still in bud or in fruit. This pattern was already observed in A. martiana by Schmid (1970a) and may be c o m m o n for the other species in the genus. Staminate flowers emerge first, one after another, and when both of them have fallen, the pistillate flower emerges. Two staminate flowers at anthesis in the same pit were occasionally observed in A. yaracuyense and A. martiana (Schmid, 1970a). In A. martiana the anthesis of individual staminate flowers lasts from two to six days and the anthesis of the pistillate flowers then follows and lasts one to two days (Schmid, 1970a; Uhl & Moore, 1977). Young fruits were observed to occur simultaneously with pistillate flowers on the rachillae of A. martiana (Bonifaz & Cornejo 3758). Several structural features o f Asterogyne flowers s u g g e s t insect p o l l i n a t i o n . T h e well-developed septal nectary (Fig. 8I) observed in the pistillate flowers of four of the five species in the genus (pistillate flowers of A. guianensis were not available for the structural study), and the rudimentary septal nectary o b s e r v e d by Schmid (1970a, 1983) in the pistillode of the staminate flowers of A. martiana, m a y provide nectar for pollinators. Schmid (1970b) found that the translucent protuberances at the base of the styles and the fleshy staminodes in A. martiana readily exude fluid when slightly damaged. The production of fragrance is well k n o w n in both staminate and pistillate flowers of Asterogyne but its histological basis is unknown. Floral fragrances were found in the male flowers of all of the Ven e z u e l a n s p e c i e s . Scent w a s e s p e c i a l l y strong in A. spicata, and has also been reported in A. martiana (Knudsen, 1999; Schmid, 1970a, 1983; Knudsen & Asmussen 620; M o o r e 6535). The presence of osmophores was preliminarily tested on a visit by the first author (between 2001 and 2003) to the Venezuelan populations of Asterogyne. The staminate flowers o f A. spicata that were submerged for five minutes in a water solution containing neutral red showed traces of the dye at the base of the petals and filaments. Floral scent composition in A. martiana and its potential impor- 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) tance in maintaining reproductive isolation were studied by Knudsen (1999). Insect pollination o f Asterogyne martiana was discussed by Schmid (1970b). In two populations in Costa Rica, he found insects of five different orders visiting the flowers; syrphid flies probing for nectar were considered effective pollinators for the species. The flies also visited staminate flowers to eat pollen. Syrphid flies have also been reported by de Nevers & Henderson 6323 to visit flowers of A. martiana in Panama. Ants (Formicidae, H y m e n o p tera) were reported by Schmid (1970b) to visit inflorescences ofA. martiana (Fig. 7E) and were observed by the first author in staminate anthesis in A. spicata. Asterogyne spicam has bright orange staminate flowers, which emit a very strong, sweet scent and are visited by bees (Henderson, 1986). Dispersal of the fruits has never been studied, but the ripe, dark purplish, black, or garnet red fruits contrasting with the brown, reddish brown, reddish, or salmon to pink rachillae suggest bird dispersal. Distribution and Ecology The five species of Asterogyne are distributed in Central America and northern South America (Fig. 4). They are understory palms; A. martiana and A. ramosa occasionally are dominant in that stratum of the forest. The only widespread species is A. martiana: the other four species are narrow ecological endemics, restricted to small areas in Venezuela and French Guiana. Asterogyne martiana grows from Belize to northern Ecuador, usually forming large stands in the understory of the lowland nonflooded rain forest, but sometimes also along streamlets and seasonally flooded patches of forest. It has also been collected in the cloud forests of Costa Rica and Colombia, sometimes to 1400 m. Asterogyne ramosa, A. spicata, and A. yaracuyense are restricted to the Venezuelan Coastal Cordillera (Fig. 4). Asterogyne ramosa is known only from the cloud forests of the Peninsula de Paria, state of Sucre, at 730 to 1250 m elevation. There it forms large stands in particularly moist areas of the Cerro H u m o and grows in asso- 343 ciation with Euterpe precatoria var. longevaginata (Mart.) Henderson, Geonoma interrttpta (Ruiz & Pay.) Mart. var. interrupta, and especially Prestoea pubigera (Griseb. & H. Wendl.) Hook. f. ex Benth. & Hook. f. The general habit and the bifid simple leaf blades of P. pubigera surprisingly resemble A. ramosa and this might have obscured the recognition of the latter in the relatively nearby cloud forest of Trinidad. As pointed out by Henderson and Steyermark (1986), all the Venezuelan populations of Asterogyne are located in three plant refugia o f the Coastal Cordillera (Nirgua, Guatopo, Paria) mentioned by Steyermark (1979). Asterogyne spicam is endemic to the Guatopo National Park and was recently collected in the forests N W of Guatire, in the state o f Miranda, where it grows at 600 to 1100 m and is restricted to the ecotone between the lowland rain forest and the low altitude cloud forest. In the localities that the first author has visited, Asterogyne spicata is rare and associated with Bactris setulosa H. Karst., Euterpe precatoria var. Iongevaginata, and Geonoma undata Klotzsch. Asterogyne yaracuyense was once thought to be endemic to the cloud forest on the Cerro La Chapa (Yaracuy State), normally above 1200 m, but recent field work of W. Meier (pers. comm.) confirmed its presence in the nearby cloud forest on Cerro Zapatero. In Cerro La Chapa, A. yaracuyense is rare and usually associated with DicO, ocarvum f u s c u m (H. Karst.) H. Wendl., Geonoma interrupta var. interrupta, G. spinescens var. braunii E W. Stauffer, Socratea altissima Balslev & E W. Stauffer (spec. ined.), and Wettinia praemorsa (Willd) Wess. Boer. Appun (1871) provided a short description of a palm in the cloud forest of the Pico Hilaria (central Venezuelan Coastal Cordillera) that matches an Asterogyne species, probably A. yaracuyense. After 20 years of intensive field work in French Guiana, J. J. de Granville discovered a filth species of Asterogyne, A. guiattensis, from a single locality in the eastern part of the country, on the Brazilian border (Fig. 4). The genus had never been reported for the Guianas: its presence in French Guiana suggests it may occur in other regions 344 BRITTONIA of northeastern South America. The very limited range of A. guianensis may be related to low adaptive and reproductive capacity (de Granville and Henderson, 1988). Asterogyne guianensis is found in the Oyapock refugium described by Brown (1982), also known as the East Guiana refugium (Prance, 1982). De Granville and Henderson (1988) suggest that A. guianensis and A. spicata may have once been a single population that became isolated as a result of climate changes. In a study at La Selva Biological Station, Costa Rica, Braker and Chazdon (1993) assessed the levels of damage produced by the neotropical forest grasshoper Micro~lopte~x hebardi (Acrididae: Ommatolampinae) that feeds on leaves of A. martiana and two species of Geonoma. Asterogyne martiana was the least preferred species, probably due to lower water content, lower fresh/dry mass per unit of leaf area, and higher crude lignin content. The Placement of Asterogyne Based on Molecular Phylogenetic Analyses The taxonomic position o f Asterogyne within the tribe Geonomeae is corroborated by phylogenetic analyses of D N A sequences. A clade o f Geonoma congesta, Calyptronoma occidentalis, and Asterogyne martiana is resolved as monophyletic in the analysis o f three plastid D N A sequences, rbcL, rpsl6, and trnL-trnF by Asmussen and Chase (2001). Several other phylogenetic analyses of palms have included A. martiana. Some of these studies (Hahn, 2002; Lewis & Doyle, 2001; Uhl et al., 1995) had A. martiana as the only representative for tribe Geonomeae and therefore did not add information about tribal relationships o f Asterogyne. In other studies (Asmussen et al., 2000; Baker et al., 1999), A. martiana and other representatives of Geonomeae were part of a poorly resolved clade of Arecoideae: these studies are also uninformative with respect to the position of Asterogyne. In the family level study by Asmussen and Chase (2001), Asterogyne martiana is resolved as sister to a poorly supported clade o f Geonoma congesta and Calyptron- [VOL. 55 oma occidentalis. More taxa and more characters are needed to obtain a robust hypothesis o f the relationships within Geonomeae. Two studies, one based on rpll6 and the other on rpsl6 plastid D N A sequences, specifically addressed the relationships of the six g e n e r a o f G e o n o m e a e ( A s m u s s e n , 1999a, 1999b). However, Asterogyne was not included in the study based on rpll6 because it was not possible to get a rpll6 sequence for A. martiana at the time (Asmussen, 1999a). In the study based on rpsl6, Asterogyne was part of a poorly resolved monophyletic group containing species of Geonoma, Calyptrogyne, and Calyptronoma, with Welfia and Pholidostachvs as sisters to this clade (Asmussen, 1999b). A number of studies of Arecoideae and Geonomeae include Asterogyne, and a robust hypothesis of the position of Asterogyne can be expected to be published in the near future. A study (Lewis, unpubl, data) o f D N A sequences of the nuclear gene phosphoribulokinase from selected genera within subfamily Arecoideae resolves species of Asterogyne as distinct and Geonoma as a monophyletic group with Asterogyne as sister to Geonoma. An unpublished study (Borchsenius et al., unpubl, data) based on morphological characters corroborates the result from the nuclear phosphoribulokinase D N A sequences. In conclusion, most of the data available, preliminary as they may be, point to a sister group relationship o f Geonoma and Asterogyne, with the remaining genera of Geonomeae outside this monophyletic clade. Relationships within Asterogyne Based on Molecular Phylogenetic Analyses Asterogyne martiana has a wide natural distribution and is widely cultivated in botanical and private gardens. It is more difficult to obtain material of the remaining four species of Asterogyne, which are restricted in their natural habitats and are not cultivated. This is the main explanation for the lack of phylogenies of Asterogyne species. Attempts made to extract D N A from herbarium material ofA. spicata and A. yaracuyensis have failed, but succeeded only for A. ramosa (Asmussen & Deichmann, 2003] STAUFFER ET AL.: A S T E R O G Y N E ( A R E C A C E A E ) pers. comm.). Based on preliminary data of plastid DNA, A. ramosa forms a clade with A. martiana that is sister to Geonorna (Asmussen et al,, unpubl, data), Fresh leaf material for D N A analyses from Asterogyne guianensis, A. ramosa, A. spicata, and A. yaracuyensis is critical for examining the infraspecific relationships of Asterogyne. The combination of data from morphology, a number of variable D N A sequences, and A F L P characters for all species of Asterogyne is likely to provide a robust interpretation of the interspecific relationships for the genus. Relationships within Asterogyne Based on a Morphological Phylogenetic Analysis The strict consensus tree (Fig. 5, length: 26, CI: 0.962, RI: 0.889) based on morphology resolved Asterogyne as a well-supported monophyletic group with a bootstrap value of 97%. However, it is clear that more outgroup taxa should be included in the analysis to test this hypothesis. Asterogyne guianensis and A. spicata form a clade with a bootstrap support o f 78%: this group is characterized by spicate inflorescences and male flowers with more than 21 stamens. Asterogyne ramosa and A. yaracuyense form a clade with a bootstrap support of 90%; the species of this group grow in the cloud forest of the Venezuelan Coastal Cordillera and have a peduncle longer than 85 cm. The relationships between these two clades and A. martiana were unresolved (Fig. 5). The parsimony search detected two most parsimonious trees. One of the two most parsimonious trees had the same topology as the strict consensus tree, whereas the other showed A. martiana as sister to the remaining species of Asterogyne. Our phylogeny supports the close relationship between A. guianensis and A. spicata, already suggested by de Granville and Henderson (1988). A clarification of the exact position of the widespread A. martiana will have to await additional data. Conservation Status Asterogyne martiana is a widespread species considered to be nonthreatened in 345 Central America and Colombia (Bernal, 1989). Most of the populations of A, spicam grow in the Guatopo National Park and therefore are protected by the rules o f the national park, Asterogyne spicata is listed as vulnerable in the IUCN red list o f threatened species (Johnson, 1996). The difficult access to the populations of A. ramosa in Venezuela and A. guianensis in French Guiana is a major factor contributing to the conservation of these species. Asterogyne yaracuyense was also listed as vulnerable by Johnson (1996). A recent visit to its natural habitat on Cen'o La Chapa has confirmed more extensive alteration and destruction of the forest than originally reported by de Granville and Henderson (1988) and Stauffer 'and Duno de Stefano (1998). Small agricultural areas for the cultivation of bananas, coffee, and tubers have gradually taken possession of the cloud forest atop Cerro La Chapa. Cattle raising and construction associated with agricultural activities have also altered the species content and structure o f the cloud forest. Therefore, Asterogyne yaracuyense should be listed as critically endangered in the I U C N red list of threatened species. Systematic Treatment ASTEROGYNE H. Wendl. ex Hook. f. Asterogyne H. Wendl. ex Hook. f. in Bentham & Hook. f., Gen. P1. 3: 914. 1883; Burret, Bot. Jahrb. Syst. 63: 140. 1930; Wessels Boer, Verh. Kon. Ned. Akad. Wetensch., Afd. Nat., 2 Reeks, 58: 79. 1968; Uhl & Dransfield, Genera Palmaruin, 539. 1987; Dransfield & Uhl, Palmae, in Kubitzki (editor), Faro. gen. vasc. pl., ft. pl., 4: 385. 1998. TYPE: Asterogyne martiana (H. Wendl.) H. Wendl. ex Hemsl. (Geonoma martiana H. Wendl.), typified by H. Wendl. ex Hemsl. in Godman & Salvin, Biol. Centr. Am. Bot. 3: 409. 1885. Aristeyera H. E. Moore, J. Arnold Arbor. 47: 3. 1966. T'cPE:A. ~v)icata H. E. Moore. Small to m e d i u m - s i z e d , solitary, unarmed, pleonanthic, monoecious palm. Stem smooth, brown to brownish cream, erect, sometimes basally decumbent, occasionally 346 BRITTONIA with basal and/or lateral vegetative branches, unarmed; internodes short; adventitious roots forming a basal cone. Leaves spirally arranged, 6-31, erect to arching, marcescent; sheath brown or cream colored, short, tubular, splitting opposite petiole, the margins stiff fibrous, the tomentum brown to reddish brown; petiole slender, adaxially flat or channeled, glabrous, abaxially rounded or angled, brown tomentose, blade bifid simple, s o m e t i m e s b e c o m i n g irregularly split with age or damage caused by wind, narrowly cuneate at base; veins 25-70, emerging at 20-35 ~ angle from midvein (rachis), prominent at one or both surfaces of blade. Inflorescence interfoliar, solitary, branched to one order, rarely spicate or forked, erect at anthesis, becoming pendent in fruit, protandrous; peduncle long, slender, brown to whitish tomentose; prophyll tubular, fibrous, membranous or chartaceous, bicarinate, brown tomentose, inserted near base o f peduncle; peduncular bracts t or 2, tubular, chartaceous or membranous, inserted 1 to several cm above base of peduncle; i n f l o r e s c e n c e rachis with short, ovate, pointed bracts, subtending rachillae, salmon to pink at anthesis, brown to reddish brown in fruit; rachillae 1-16, ca. equal in length, ending in a short, sterile apex, same color as inflorescence rachis. Flower pits containing triads comprising two staminate flowers and one pistillate flower, the pits spirally arranged on rachis, covered by a rounded upper lip that overlaps the lower, immersed lip when in bud; floral bracts 3, the largest bract subtending the second staminate flower, a medium-sized, 2-keeled bract subtending the pistillate flower, and a small bract representing the prophyll of the pistillate flower. Staminate flowers oblong to oblong-obovoid, borne abaxially to pistillate flowers; sepals 3, unequal, united basally, narrow, elongate, keeled dorsally; petals 3, ca. as long as or longer than sepals, united into a tube for ca. 2,4 their length, distally valvate; stamens 6-27; filaments united into a tube up to midlength, white or yellow, with or without a papillate epidermis; connective bifid, tanniferous, bearing separated thecae, the anthers inflexed toward center of flower and introrse in bud, spread and extrorse at anthesis; pollen [VOL. 55 asymmetric in shape, monosulcate, the sulcus as long as main axis, the exine tectate and finely perforate-rugulate or perforatefossulate (Fig. 3A-H); pistillode irregularly 3-lobed (Fig. 12E). Pistillate flowers oblong to oblong-ovoid; sepals 3, unequal, imbricate in bud, dorsally keeled; petals 3, united up to midlength, distally valvate; staminodes 5-22, fused to corolla and united basally, fleshy, with tanniferous tips; gynoecium syncarpous, trilocular, triovulate, the nectary septal; style basal to lateral, elongate; stigmatic branches 3, papillate, reflexed at anthesis; ovule anatropous, inserted at midlength of locule, pendent. Fruit a drupe, ellipsoid to ellipsoid-ovoid, 1-seeded, dorsiventrally compressed, slightly to conspicuously keeled at apex, the remains of the stigma basal; epicarp smooth, purplish black or garnet red at maturity; mesocarp fleshy and juicy, with an inner layer of closely appressed, longitudinal fibers; endocarp thin, crustaceous, and shiny (Uhl and Dransfield, I987). Eophyll with a bifid simple blade. Distribution.--Five species distributed in Central America (Belize, Guatemala, Honduras, Nicaragua, Costa Rica, and Panama) and northern South America (Colombia, Ecuador, Venezuela, and French Guiana). (Fig. 4) Key to the species of Asterogyne Stamens 21-27; inflorescences spicate. 2. Inflorescence rachis and rachillae pale green at anthesis; filaments white and smooth; fruits 2.5 c m long, 1.5 c m di a m . . . . . . . . . . . . . . A. guianensis 2. I n f l o r e s c e n c e rachis and r a c h i l l a e brown at anthesis; filaments conspicuously yellow orange and papillate; fruits to 1.8 cm long, 0.8 cm diam. .............................. A. spicata Stamens 6-12; inflorescences branched, rarely spicate. 3. Peduncular bract 1, rarely 2: stamens 6; staminodial lobes 6 A. martiana 3. Peduncular bracts 2: stamens 8-12: staminodial lobes 5-9. 4. Inflorescence raehis 0.5-2.5 cm long; rachillae 4 - 6 ; stamens 1112 ...... A. ramosa 4. Inflorescence rachis 5 - 1 0 c m long; rachillae 9 16; stamens 8 - 1 0 --....... A. yaracuyense 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) 1. ASTEROGYNE GUIANENSIS Granv. & A. J. H e n d . (Fig. 1A, B) A s t e r o g y n e g u i a n e n s i s G r a n v . & A . J. Hend., B r i t t o n i a 40: 76. 1988. TYPE: F R E N C H G U I A N A . C a m o p i River, ca. 1.5 k m N E o f M o n t B e l v 6 d b r e , in s w a m p forest, 150 m, 4 D e c 1984, J. J. de Granville 7124 [HOLOTYPE: P; ISOTYPES: B, BR(n.v.), CAY, K, NY, US] Stem solitary, erect, 1 . 5 - 2 m tall, 3 - 6 c m diam., brown, not branched; internodes short: aerial r o o t s b r a n c h e d , f o r m i n g a b a s a l cone, e a c h r o o t 3 8 - 9 0 c m long, 0 . 2 - 0 . 3 c m diam., yellowish brown; pneumatodes white, c o n i c a l , to 0.2 c m long. L e a v e s 1 5 18, the o l d e r o n e s not persistent; sheath 7 • 4 - 5 cm, b r o w n ; p e t i o l e 5 3 - 6 0 X 0 . 8 - 1 cm, channeled adaxially, rounded abaxially, green, the t o m e n t u m r e d d i s h b r o w n ; m i d v e i n 8 0 - 9 0 • 0.5 c m , green, the i n d u m e n turn s a m e as on p e t i o l e , c h a n n e l e d a d a x i ally, r o u n d e d a b a x i a l l y ; blade 1-1.1 x 0 . 3 0.4 m, d e e p l y bifid at a p e x for ca. 35 cm; p r i m a r y v e i n s 29 p e r side, e m e r g i n g at 2 0 30 ~ a n g l e at m i d v e i n , p r o m i n e n t on a b a x i a l side, a l m o s t y e l l o w w h e n dry, the l e p i d o t e t o m e n t u m s l i g h t l y r e d d i s h b r o w n , hlflorescence spicate; p e d u n c l e 4 0 - 5 5 c m long, 0.4-0.5 cm diam., dorsiventrally comp r e s s e d , the t o m e n t u m b r o w n to w h i t i s h ; p r o p h y l l 20 c m long, 1.5 c m d i a m . , m e m b r a n o u s , g r e e n , i n s e r t e d at b a s e o f p e d u n cle; peduncular bract l, membranous, green, 4 0 - 4 2 • 1 . 3 - 1 . 5 cm, i n s e r t e d 1 c m a b o v e the b a s e o f the p e d u n c l e : infloresc e n c e r a c h i s 2 0 - 3 0 c m long, 0 . 9 - 1 . 5 c m d i a m . , terete, p a l e g r e e n at anthesis, r e d d i s h b r o w n in fruit, the t o m e n t u m w h i t i s h to b r o w n i s h , the sterile a p e x acute, 1 - 2 m m l o n g : f l o w e r pits s p i r a l l y a r r a n g e d , 3 - 5 m m a p a r t f r o m e a c h other, c o v e r e d with an inf l e x e d l o w e r lip, 3 - 4 • 2 - 3 m m . S t a m i n a t e f l o w e r s o b o v o i d , 9 - 1 0 m m long, 3 - 4 m m d i a m . , w h i t e at a n t h e s i s , the p e d i c e l to 1 - 2 m m long; s e p a l s o b l o n g , 9 - 9 . 2 • 2 - 3 m m ; p e t a l s o b l o n g - o b o v a t e , 8 X 2 . 5 - 3 ram; stam e n s 2 5 - 2 7 ; f i l a m e n t s u n i t e d into a t u b e up to 4 - 5 m m long, the free parts 2 . 5 - 3 m m long, 1 m m d i a m . , w h i t e ; c o n n e c t i v e 0.8 m m long; t h e c a e 1 . 2 - 1 . 5 m m long; pistill o d e 1 - 2 m m long. Pistillate f l o w e r s o b l o n g 347 in bud, 5 m m l o n g , 2 . 5 - 3 m m d i a m . , t h e p e d i c e l to 1 m m l o n g ; sepals o b l o n g - o b ovoid, 4-5 • 1 ram; petals oblong-obovoid, 4 X 1 . 4 - 1 . 6 m m ; o v a r y 3 m m long, 2 m m d i a m . ; style 3 m m l o n g ; s t i g m a t i c b r a n c h e s 1 m m long; s t a m i n o d i a l tube 4 m m l o n g , the l o b e s 2 0 - 2 2 , 1 - 1 . 5 m m long. Fruit narrowly ellipsoid, strongly keeled apically, 2.5 c m long, 1.5 c m d i a m . ; e p i c a r p g a r n e t r e d and s h i n y at m a t u r i t y : seed n a r r o w l y ell i p s o i d , 2 c m l o n g , 0.8 c m d i a m . Local n a m e s a n d u s e s . - - N o n e r e p o r t e d . Distribution a n d ecology. K n o w n o n l y f r o m t w o c o l l e c t i o n s o f eastern F r e n c h G u i ana, r e m a r k a b l y m o r e than 200 k m f r o m each other. H e n d e r s o n and B a l i c k (1987) s u g g e s t e d that this s p e c i e s , at that t i m e n o t y e t d e s c r i b e d , m a y o c c u r also in n o r t h e r n Brazil. Additional specimens examined: FRENCH GUIANA. Crique Rapatea, Bassin de 1Approuague, jonction de la Kouroua'/e et de la RN2, 4~ 52~ 24 Oct 1996, Blanc 189 (CAY); plantlets cultivated at Cayenne, 22 Oct 1985, de Granville 7124-A (CAY). 2. ASTEROGYNE MARTIANA (H. W e n d l . ) H. W e n d l . ex H e m s l . A s t e r o g y n e m a r t i a n a (H. W e n d l . ) H. W e n d l . ex H e m s l . in G o d m a n & Salvin, Biol. Centr. A m . Bot. 3: 409. 1885; G e o n o m a martiana H. W e n d l . , L i n n a e a 28: 342. 1856. TYPE: cult. hort. H e r r e n h a u s e n , H. W e n d l a n d 56 [HOLOTYPE:G O E T ; ISOTYPE: K ( F neg. 2 0 7 6 5 , B H neg. 20368)] (Fig. 1C) Geonoma trifi~rcata Oerst., Kjoeb. Vidensk. Meddel. 1858: 34. 1859. TYPE: Nicaragua, ad flumen San Juan, A. S. Oersted 6566 [HOLOTYPE:C(F neg. 21 O69)]. Asterogyne minor Burret, Bot. Jahrb. Syst. 63: 143. 1930. TYPE: SW Colombia, above Barbacoas, 500 m, F. C. Lehnlann 708 [HOLOTYPE:B(destroyed)]. Stem solitary, erect, s o m e t i m e s b a s a l l y d e c u m b e n t , 0 . 4 - 1 . 6 ( - 3 ) m tall, 2 . 5 - 5 diam., dull b r o w n to o r a n g e , not b r a n c h e d ; internodes ( 1 - ) 2 . 5 - 3 c m long; aerial roots f o r m ing an irregular b a s a l cone, each root 4 - 8 c m long, 0 . 3 - 0 . 4 c m diam.; p n e u m a t o d e s b r o w n i s h c r e a m - c o l o r e d , conical, to 1 m m long. Leaves ( 6 - ) 1 2 - 2 0 , y e l l o w i s h red w h e n young, green w h e n mature, the o l d ones s o m e t i m e s persistent; sheath ( 8 - ) 1 1 - 1 6 ( - 3 8 ) 348 BRITTONIA X 3 cm, with reddish brown tomentum, longitudinally splitting, the margins slightly fibrous; petiole ( 4 - ) 2 0 - 6 8 ( - 7 5 ) • 0.5-1.5 cm, keeled to channeled adaxially, rounded to a n g l e d abaxially, almost g l a b r o u s to densely brown tomentose, toward the base slightly fibrous at margins; midvein (33-)55791-108) X 0.2-0.7 cm, greenish yellow, glabrous or with pinkish brown tomentum, slightly keeled adaxially, rounded abaxially; blade 0.4-1.3 • 0.15-0.3 m, light to dark green, deeply bifid at apex for 18-32 cm: primary veins 2 8 - 4 0 per side, emerging at 2 0 - 3 5 ~ angle at midvein, prominent at both surfaces of blade, yellow when dry. h ~ o r e s c e n c e branched to 1 order or very rarely spicate: peduncle ( 2 0 - ) 3 4 - 7 0 ( - 1 2 5 ) cm long, 0.4-0.7 cm diam., dorsiventrally compressed: prophyll 11-26(-36) cm long, (/.6-1.5(-2) cm diam., inserted at base of peduncle: peduncular bract 1, rarely 2, chartaceous, brownish cream-colored, the indumentum same as on prophyll; first peduncular bract 2 0 - 3 1 ( - 4 0 ) X 0.5-1.5 cm, inserted 0 . 5 - 2 cm above base of peduncle: second peduncular bract, when present, 15 X 0.5-0.9 cm, inserted 2 5 - 2 7 cm above base of peduncle; inflorescence rachis absent or 0 . 5 - 3 ( - 7 ) cm long, 0.3-0.5(-1.2) cm diam., dorsiventral to irregularly compressed, salmon to pink at anthesis and in fruit, glabrous or with indumentum same as on peduncle; rachillae ( 2 - ) 3 - 5 ( - 1 4 ) , terete, erect in bud, slightly pendent in fruit, the color same as inflorescence rachis, the tomentum white, farinose or short and appressed: basal rachillae ( 9 - ) 1 4 . 5 - 2 2 ( - 2 7 ) cm long, 0.4-0.6(-1.1) cm diam.: middle rachillae (8.5-) 14-21 (-23.5) cm long, 0 . 4 0.6(-1) cm diam.; apical rachillae ( 9 - ) 1 5 22 cm long, 0.4-0.6(-1.2) cm diam., the sterile apex acute, 0.2-1.1(-5.5) cm long; flower pits spirally arranged, ca. 2 - 3 mm apart from each other, covered with a lower semicircular lip, 1-2 • 1-2 ram, the lip becoming reflexed in fruit. Staminate flowetw oblong-obovoid in bud, 6 - 9 m m long, 2-3 mm diam., white at anthesis, the pedicel to 0.7 m m long; sepals oblong-lanceolate, 3.6-4.8 • 1 ram: petals oblong-lanceolate, 5-7 • 1-1.3 ram: stamens 6, the filaments united into a tube up to 5 . 4 - 7 m m long, the free parts 2 . 4 - 3 m m long, 0.3-0.5 mm [VOL. 55 diam., r e m a r k a b l y thinner at apex and therefore the anther versatile, white, without a papillate epidermis; connective slightly swollen, 0.7-1.1 mm long; thecae 0 . 7 1.6 m m long: pistillode 2-5 m m long, 1 m m diam., basally fused with staminodial tube. Pistillate flowers oblong in bud, 6-8 mm long, 2,7-3.1 mm diam.; sepals oblong, 4 - 5 • 1-l,5 mm; petals oblong, 4,4-5.5 • 1,1-1.8 ram; ovary 1 m m long, 0.8 mm diam.; style 6 - 8 m m l o n g : s t i g m a t i c branches 1-2 mm long; staminodial tube 3 . 9 - 5 m m long, the lobes 6, 1.5-2.1 mm long. Fruit ellipsoid to ellipsoid-ovoid, 0 . 8 1.2 cm long, 0.4-0.7 cm diam.; epicarp orange to red when young, purplish black at maturity; seed ellipsoid, 0.7-0.8 cm long, 0.4-0.5 cm diam. Local names and uses.--capoca (Guatemala); pacuquilla (Honduras); pata de gallo, suita ( N i c a r a g u a ) ; p a c a y a , suita ( C o s t a Rica), dobo-gaw (Guaymi of Panama); cortadera, panda~ panga, rabihorcao, rabo de zorro (Colombia): guacamayo, pico (Ecuador). The use of the leaves for thatching houses has been reported in Nicaragua (Neill 1723, Salick 8106), in Costa Rica (Chazdon, 1986), in Panama (Gordon 70), in Colombia (Bernal & Galeano 887), and in Ecuador (Aulestia & Aulestia 1306). Distribution and ecology. Widespread in Central America: Belize (El Cayo, Stann Creek, Toledo), Guatemala (Izabal), Honduras (Atl~intida, Col6n, Cort6s, Gracias a Dios, Santa B~irbara), Nicaragua (Chontales, Jinotega, Matagalpa, Rfo San Juan, Zelaya). Costa Rica (Alajuela, Cartago, Heredia, Lim6n, Puntarenas, San Jos6), Panama (Bocas del Toro, Chiriquf, Coc16, Col6n, Dari6n, Panama, San Blas, Veraguas), and in northwestern South America: Colombia (Antioquia, Choc6, C6rdoba, Narifio, Valle del Cauca), Ecuador (Esmeraldas) (Fig. 4). C o m m o n in the understory of the lowland and mountain rain forests, in flooded and nonflooded areas; from sea level to 1400 m, but normally below 600 m. Selected specimens examined: BELIZE. El Cayo District: 47 mile section, Humming Bird Hwy.. 28 Apr 1956, Gentle 9072 (NY). 60 m, 6 Sept 1929. Schipp 392 (G, K, MO, NY, Z); W of Isla Rivas, 29 Jan 1903, Wilson 224 (NY, US). Toledo: Monkey Rivel; in high 2003] S T A U F F E R ET AL.: A S T E R O G Y N E ( A R E C A C E A E ) "Monkey Tail," 1 Nov, 1941, Gentle 3743 (MO, NY, US1. GUATEMALA. Izabal: Wet wooded slopes nr. Km 824 past Villa Jesfis on rd. from Puerto Barrios to Guatemala, 100 m, 8 Nov 1959, Moore & Cetto 8213 (BH): valley of tributary of Rio San Francisco del Mar, 2 miles NE of Hopi, 12 miles from Entre Rfos, 10-20 m, 21 Apr 1940, Stevermark 39785 (US), E portion of Vera Paz and Chiquimula, 1885, Watson 238 (BH/. HONDURAS. Atl~intida: Vic. Lancetilla, 45 m, 8 Oct 1934, Ynncker 5002 (MO, NY). G r a e i a s a Dios: Laimos Creek, 15 km SW of Waspan, 7 Mar 1961, 15-30 m, Bunting & Licht 401 (NY): Rio Plfitano, 4 hours upriver from village of Ras, nr. sea level, 23 May 1973, Gent O, et al. 7542 (MO). NICARAGUA. Chontales: Chontales, 600 m, Jun 1870, Levv 505 (G, P): 1870, Seemann s.n. (BMI. Jinotega: Salto Kayaska, Rio Bocaz, ca. 13~ 85~ 190-340 m, 7 Mar 1980, Beach et al. 16545 (US); Comarca de Bocaycito, Cordillera Isabelia, nr. Rio Bote, 6 May 1976, Neill 7151 (MO, US/. Matagalpa: E1 Comej6n, 1 km W de la carretera a Waslala, 13~ 85~ 600 m, Moreno & Robleto 20602 (MO). Rio San Juan: Sabalo, 1 k m a l N de Rio San Juan, ll~ 84~ 100 m, 9-10 Jul 1985, Moreno 26083 (MO): "Los Filos,'" nr. Loma Los Filos, Rio Santa Cruz, 1 l~ 84~ 6 Sept 1993, Salick 8106 (MO). Zelaya: Ca. 1 km N of El Zapote, 6 km S of Colonia Verdfin, 11~ 84~ 250-300 m, 8 Sept 1983, Nee & Vega 27905 (NY, US); El Hormiguero, carlo above camp and on W slope at Loma, 13~ 84~ 800-1000 m, 17 Mar 1980, Pipoly 6143 & 6168 (MO, US/: ca. 6.3 km S of bridge at Colonia Yolania and ca. 0.8 km S of ridge of Serranias de Yolania on rd. to Colonia Manantiales (Colonia Somoza/, 11~ 37'N, 84~ 200-300 m, 29-31 Oct 1977, Stevens 4810 (BH, MO, NY). COSTA RICA. Alajuela: Vic. Guatuso de San Rafael (on Rio Frfo), 10~ 84~ 80-100 m, 4 Aug 1949, Hohn & lltis 914 (BH, BM, MO, US): 25 km NNW of San Ram6n by rd. on way to San Lorenzo, 10~ 84~ 500 m, 24 Apr 1983 Liesner & Judziewicz 14825 (MO, WIS). Cartago: Bois de Peralta, 300 m, Jun 1903, Pittier 16658 (US). Heredia: Finca La Selva, 12 Dec 1984, Henderson 53 (NY): along "Starkey Rd.,'" 4.5 km SE of bridge at Puerto Viejo, 10~ 83~ 50 m, 7 Aug 1979, Stevens 13505 (K, MO, US/. Lim6n: Finca Castilla, 30 m, 24 Jul 1936, Donee & Goerger 9284 (MO/: 7 km SW of Bribrf, 100 250 m, G6mez et al. 20341 (MO, NY, W1S): woodlands south of La Lola on the railroad, 120 m, 15 Apr 1953, Moore 6717 (BH). P u n t a r e n a s : Golfito, 15 m, 22 Oct 1950, Hari s.n. /G); hills above Palmar Norte, 550 m, 6 Mar 1953, Moore 6530 (AAU, BH, K): Osa Peninsula, Agabuena, 4 km W of Rinc6n, at cleared path dividing lands of Don Fortunato and Don Contreras, SSW of 4-ha sample plot, 8~ 83~ 'W, 250 m, 18 Nov 1992, Thomsen 187 (C). San Jos& Rio Negro, Cerro La Cangreja, Santa Rosa de Puriscal, 400-700 m, 20 Jun 1986, Chac6n & ChacDn 1958 ICR/: Tarrazfi, San Lorenzo, Llanos de Santa Maria, 9~ 84~ 500 600 m, 28 Jan 1998, Valverde 665 (K). PANAMA. Boeas del Toro: Top of - F i l a " above 349 Almirante, 30 Nov 1971, Gentry 2803 (MO); Rio San Pedrito, 16 Jul 1978, Gordon 70 (MO); without locality, 24 Sept 1940, yon Wedel 952 (BH). Chiriquf: Burica Peninsula, 19 Feb 1973, Croat 21997 (K, MO); Fortuna Darn region, hr. Chiriquf Grande, 8~ 82~ 200 m, 18 Jan 1986, McPherson 8080 (MO): vic, Chiriquf Lagoon, 9 Oct 1940, yon Wedel 1108 (BH, MO/. Coc16: Rd. to Coclesitn, logging camp, 12 miles from Llano Grande, 8~ 80~ 200 m, 16 Dec 1983, Churchill et al. 4122 (K, MO), Boca del Toabrd at confluence of Rio Toabr6 and Rio Coc16 del Norte, 11 Apr 1969, Lewis et al. 5558 (BH, MO/. Co16n: Santa Rita Ridge, 11 May 1999, Henderson & Ferreira 3048 (NY); Santa Rita Ridge, end rd. from Transisthmian Hwy., ca. 10 miles from Hwy., 29 Mar 1969, Porter et al. 4736 (BH, MO, NY); Santa Rita Lumber Rd., ca. 3.1 km E of Agua Clara Rain Gauge, 9 Sep 1975, Witherspoon et al. 8322 (MO), Dari6n: Parque Nacional del Dari6n, ridge between Rio Topalisa and Rio Pacuro, ca. 13 km E of Pacuro, Quebrada Pobre to Mi Casita, 8~ 77~ 450 600 m, 14 Oct 1987, de Nevers et al. 8320 (MO, US). Panama: Canal Zone, Apr 1925, Cook s.n. (US); Cerro Jefe roadside, 2 km N of turnoff to radio tower on rd. [from?] Alto de Pacora, 790 m, 30 Sep 1978, H a m m e l 4863 (MO). San Bias: 10 km from Pan-American Hwy. on El Llano Cartf Rd., 9~ 79~ 250 rn, 6 Mar 1987, BarJ~)d & M e Phe r s on 13 (AAU/: 44 km N of Penonome on rd. to Coclecito, 90 150 m, 21 Feb 1978, Hammel 1682 (BH, MO/: Nusagandi, E1 L l a n o Cart/ Rd., 19 km from Inter-American Hwy. 9~ 78~ 300 400 m, 9 Oct 1996, Knudsen & Asmussen 604, 620 (AAU): E1 Llano-Cart/Rd., Km 10 15, 300-400 m, 3 Sept 1977, M aas et al. 2803 (U). Veraguas: Atlantic slope, Rio Concepci6n to Rio Barrera, ca. 60 m, 16 Oct 1978, H am m e l 5153 (MO). COLOMBIA. Antioquia: Mun. Frontino, Corregimiento de Murrf, rd. from Nutibara to La Blanquita, 950 m, 18 Mar 1982, Bernal & Galeano 256 (NY), Mun. de San Rafael, 15 k m a l E del pueblo en la carretera a San Carlos, 960 m, 19 Sept 1987, Bernal & Tobdn 1380 (AAU, BH, NY); Villa Arteaga, Mutata, ca. 1000 m, 5 Apr 1976, Dran~sfield et al. 4859 (BH, K). Choe6: Area of Baud6, on left bank of river Baud6, ca. 11.5 km upstream of estuary, slightly downstream of estuary of Quebrada Anguerad6, 50 m, 7 Feb 1967, Fuchs & Zanella 21808 (U): north ridge of Alto de Buey, above Dos Bocas del Rio Mutat~i, tributary of Rio El Valle, ESE of E1 Valle, 200 500 m, 8 Aug 1976, Gentry & Fallen 17445 (MO). C o r doba: Junction of Rio Tigre and Rio Manso, Paramillo National Park, transect 1, 7~ 76~ 200 m, 25 Jul 1988, Gentry & Cnadros 63735 (MO). Narifio: Mun. de Tumaco, 3 5 km al E de Guayacana, 100 m, 3 Oct 1985, Bernal & Galeano 887 (AAU, NY). Vaile del Cauca: Buenaventura, 6 May 1926, Cook 84 (US); Bajo Calima, Juanchaco Palmeras, ca. 10 km N W of Buenaventura, 3~ 77~ 13 Apr 1987, Gentry et al. 56871 (MO, NY/. ECUADOR. E s m e r a l d a s : Reserva Etnica Awfi, Parroquia Alto Tambo, Centro de la Uni6n, carl6n del Rio Mira, 00~ 78~ 250 m, 22 Mar 1993, Aulestia & Aulestia 1306 (MO): Playa de Oro on the Rio Santiago, ~ hour walk S of village, 250 m, 25 Feb 350 BRITTONIA 1994. Borchsenius 242 (AAU, NY); San Lorenzo Cant6n, Reserva Etnica Awli, Centro Ricaurle, a orillas del Rio PalabL ~~ 78~ 300 m, 28 Oc~ 1992, Tipaz et aL 2250 (AAU. MO). Cult. Holt. Herrenhausen, Wendland 56 (GOET, HOLOTYPE; K, ISOTYPE). Probably originated from seeds sent by Warscewicz from Central America. 3. ASTEROGYNE RAMOSA (H. Wess. Boer. (Fig. 1 D - F ) E. Moore) A s t e r o g y n e ramosa (H. E. M o o r e ) Wess. Boer, Verb. Kon. Ned. Akad, Wetensch., Aid. Nat,, 2 Reeks, 58: 81, 1968. Aristevera ramosa H. E. Moore~ J, A r n o l d Arbor. 48: 144-146. 1967. TYPE: VEN E Z U E L A . Sucre: Peninsula de Paria, Cerro Espejo, en la c u m b r e entre Mantecal y Paujf, arriba de M u n d o N u e v o y Rio S e c o de Irapa, 7 5 0 - 8 5 0 m, 6 A u g 1966, J. S t e y e r m a r k & M. R a b e 96088 (HOLOTYPE:BH; ISOTYPES: NY, US, V E N ) Stem solitary, erect, 3 - 6 m tall, 6 - 1 0 c m diam., c r e a m y brown, o c c a s i o n a l l y with lateral vegetative branches; internodes 5 - 1 0 cm long; aerial roots f o r m i n g a short basal cone, each up to 15 cm long, 0.5-1 c m diam., brown, conical; p n e u m a t o d e s very short to almost absent. L e a v e s 15-20, the older ones briefly persistent; sheath 9 - 1 0 ( 17) X 7 cm, densely brown tomentose, the margins slightly fibrous; p e t i o l e 3 0 - 3 5 • 0.6-1 cm, channeled adaxially, rounded abaxially, b r o w n tomentose; m i d v e i n 39 • 0.5 cm, the i n d u m e n t u m b r o w n , slightly channeled adaxially, almost flat abaxially; blade 0 . 5 - 0 . 7 • 0.18-0.27 m, green, deeply bifid at apex for ca. 33 cm; p r i m a r y veins 3 4 - 4 5 per side, emerging at 2 5 - 3 0 ~ angle at midvein, prominent on both surfaces o f blade, pale green abaxially. Inflorescence branched to 1 order; peduncle 9 0 - 1 2 0 c m long, 0 . 4 - 0 . 5 c m diam., dorsiventrally c o m pressed, green; prophyll 1 5 - 2 0 c m long, 1.8 c m diam., chartaceous, inserted at base o f peduncle; peduncular bracts 2, chartaceous, brown, the i n d u m e n t u m same as on prophyll; first peduncular bract 3 1 - 4 9 • 1.4 cm, inserted 0.5 cm above base o f peduncle; second peduncular bract 4 1 - 5 3 • 1 1.2 cm, inserted 1 cm above base o f peduncle; inflorescence racbis 0 . 5 - 1 . 2 ( - 2 . 5 ) cm long, 0 . 4 - 0 . 5 c m diam., dorsiventraI to ir- [VOL. 55 regularly compressed, pale green at anthesis, reddish in fruit, glabrous; rachillae 4 6, terete, erect, same c o l o r as inflorescence rachis, each rachilla subtended by a c u r v e d bract up to 0,5 c m long, c o v e r e d with a whitish farinose i n d u m e n t u m ; basal, medial, and apical rachillae 1 7 - 1 9 c m long, 0.5 c m diam., the sterile apex acute, 0 . 2 - 0 . 4 c m long; flower pits spirally arranged, ca. 2 - 3 m m apart f r o m each other, c o v e r e d with an inflexed l o w e r lip, 1.5-2 • 1.5-2 m m . Stam i n a t e f l o w e r s o b l o n g - o b o v o i d in bud, 6 - 8 m m long, 3 - 4 m m diam., slightly fragrant, white at anthesis, the pedicel to 1.5 m m long: sepals o b l o n g - o b o v a t e , 5 - 6 • 2 - 3 m m ; petals oblong, 4 - 6 • 2 m m ; stamens 1 1 - 1 2 , o c c a s i o n a l l y 1 sterile; f i l a m e n t s united into a tube to 3 m m long, the free parts 3 m m long, 0.5 m m diam., white, not r e m a r k a b l y thinner at apex, the epidermis not papillate; connective 1 m m long; thecae 1.2 m m long; pistillode 2 m m long. Pistillate f l o w e r s o b l o n g - o v o i d in bud, 5 - 7 m m long, 3 - 4 m m diam., the pedicel to 1 m m long; sepals o b l o n g to ovoid, 4 - 5 • 2 - 2 . 5 m m ; petals ovate, 5 - 6 x 2 ram; o v a r y 1.5 m m long, 1.5 m m diam.; style 1 . 5 - 2 m m long; stigmatic branches l m m long; staminodial tube 4 m m long, the lobes ( 5 - ) 7 - 9 , 2 . 8 - 3 . 4 m m long. Fruit ellipsoid, 1.5-1.7 cm long, 0 . 5 - 0 . 6 c m diam.; epicarp dark purplish red w h e n y o u n g , purplish black at maturity; seed ellipsoid, 1.3 c m long, 0.4 c m diam. L o c a l n a m e a n d uses.--anare. Uses not known. D i s t r i b u t i o n a n d ecology. K n o w n only f r o m the understory o f the m o n t a n e cloud forest in the northeastern Coastal Cordillera, state o f Sucre, Venezuela (Fig. 4); f r o m 7 3 0 - 1 2 5 0 m. A l t h o u g h the v o u c h e r for coll e c t i o n S t e y e r m a r k 9 4 8 0 3 m e n t i o n s the palm g r o w i n g at 1600 m on the Cerro H u m o , this must be an error b e c a u s e the s u m m i t o f this m o u n t a i n has b e e n reported to be 1 2 5 0 m . Additional specimens examined: VENEZUELA. Sucre: Peninsula de Paria National Park, above Las Melenas, N of Rio Grande Arriba, trail to Cerro de Humo, 10~ 62~ 900-1000 m, 3 Jan 2002, Stauffer et al. 824 (MY, MYE VEN. Z); Peninsula de Paria, Cerro Hnmo, a 2o largo de uno de los affuer~tes del Rio Santa Isabel arriba de Siparo, 1600 m (prob- 2003] S T A U F F E R E T AL.: A S T E R O G Y N E able erroneous elevation), l Mar 1966, Steyermark 94803 (BH, NY, U, VEN); Peninsula de Paria, above Las Melenas, N of Rio Grande Arriba, SE of Cerro de Humo, 10~ 62~ 730 1050 m, 1 Dec 1979, Steyetwlark & Liesner 120915 (MO, NY, VEN); Penfnsula de Paria, Distrito Marino and Distrito A r i s m e n dL trail between crossing of Rio Tacarigua to s u m m i t of slopes E of Cerro Humo, descending to Las Melenas, N of Rio Grande Arriba, 10~ 62o36 37'W, 7 6 0 - 1 0 0 0 m, 24 Feb 1980, Steyermark et al. 121713 (MO). (ARECACEAE) 351 duncle; peduncular bract 1, chartaceous, brown tomentose, 2 2 - 4 0 • 1.3-2 cm, inserted 1.4 cm above base of peduncle; inflorescence rachis 2 3 - 4 0 cm long, 1-1.5 c m diam., terete, pale brown at anthesis, brown in fruit, with same indumentum as on peduncle, the sterile apex acute, ca. 0.6-3 c m long; flower pits spirally arranged, ca. 2 - 4 m m apart from each other, covered with an inflexed lower lip, 3 X 2 - 4 mm. Staminate flowers obovoid, 0.8-1.3 cm long, 0.4-0.5 4. ASTBROaYNE SPICATA (H. E. Moore) c m diam., fragrant, the pedicel almost abWess. Boer. (Fig. 2 A - C ) sent; sepals oblong-obovoid, 5 - 6 • 1.4-2 m m ; petals obovoid, 7 - 8 x 2 - 3 ram; staAsterogyne 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 Aid. Nat., 2 Reeks, 58: 82. 1968; Aristube to 6 - 7 m m long, conspicuously yellow tevera spicata H. E. Moore, J. Arnold orange, not r e m a r k a b l y thinner at apex, the Arbor. 47: 5-9. 1966. TYPE: V E N E Z U E epidermis papillate, the free parts 3 - 4 m m LA. Miranda, Parque Nacional Guatopo, long, 0.4 m m diam.; connective 1.2-1.5 moist rich mixed evergreen forest on m m long; thecae 0.5-1.2 m m long; pistilsteep slopes bordering Rio Santa Cruz lode 4 - 5 m m long. Pistillate flowers oblong and Altagracia de Orituco, 14.5 k m f r o m in bud, 8 m m long, 4 m m diam., the pedicel Los Alpes, 12 k m f r o m Rancherfa Mi to 1 m m long; sepals oblong, 5 - 7 X 2 - 3 Querencia, 520 m, 23 Nov 1961, J. Steyram; petals oblong, 7 - 8 m m • 2 - 3 m m ; ermark 90010 (HOLOTYPE: BH; ISOTYPES: o v a r y 0.9 m m long, I - 1 . 2 m m diam.; style NY, VEN) 2.5-3 m m long; stigmatic branches 1-1.5 Stem solitary, erect, 2 - 4 ( - 8 ) m tall, 4 - 7 m m long; staminodial tube 2 m m long, the cm diam., brown, occasionally with basal lobes 15-21, 2 - 2 . 2 m m long. Fruit ellipv e g e t a t i v e b r a n c h e s ; i n t e r n o d e s 0.5 c m soid-ovoid, 1.5-1.8 c m long, 0.5-0.8 c m long; aerial roots forming a short basal diam.; epicarp purplish red in young fruits, cone; pneumatodes v e r y short. Leaves 1 5 - purplish black at maturity; seed ellipsoid, 19(-25), sometimes reddish brown when 0.8-1 c m long, 0.5 cm diam. young, green when mature, the older ones Local name and uses.~palma, palmito, not persistent; sheath 10-15 X 9 cm, red- palmiche. M o o r e (1966) noted that the dish brown tomentose, the margins slightly fruits are said to be edible and the leaves fibrous; petiole 3 5 - 4 0 X 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 6 8 - 8 4 • 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- m o u n t a n e c l o u d f o r e s t in the n o r t h e r n ally; rachis with a filiform extension of 1.5- Coastal Cordillera, state of Miranda, Ve5 X 0.1 cm; blade 0.7-1 x 0.2-0.3 m, dull nezuela (Fig. 4); normally between 6 0 0 green adaxially, silver green abaxially, 1000 m. deeply bifid at apex for ca. 45 cm; p r i m a r y Additional s p e c i m e n s e x a m i n e d : V E N E Z U E L A . veins 2 5 - 2 7 per side, emerging at 2 5 - 3 0 ~ M i r a n d a : Parque Nacional Guatopo, 20 Dec 1961, angle at midvein, prominent abaxially, yel- Agostini 10 (VEN): Carretera Santa Teresa-Altagracia lowish, slightly b r o w n tomentose. Inflores- de Orituco, 6 0 0 - 7 0 0 m, Jun 1953, Aristeguieta 1772 cence spicate; peduncle 4 0 - 5 0 cm long, (NY, VEN); selvas de Guatopo, 500 m, Jun 1958, Aristeguieta 3185 (NY, VEN); selvas pluviales de Gua0.5-0.6 cm diam., dorsiventrally comtopo, 4 0 0 - 6 0 0 m~ 13 N o v 1956, Bernardi 5633 (NY); pressed; prophyll 1 4 - 1 5 ( - 2 5 ) cm long, 1.5 Parque Nacional de Guatopo, N of Altagracia de Oric m diam., fibrous, inserted at base of pe- tuco, 400 m, 16 Feb 1979, G e n t ~ & Troth 24823 (BH, 352 BRITTONIA MO/; Dist. Paez, Fila La Tigra, Quebrada San Juan, 18 km al SW de C0pira, 10~ 65~ 47'W, 600 m, 2-7 Sep 1977, Gonzdlez & Ortega 1296 (BH, MO); NW de Guatire, excursidn Fila Juan Torres Fila Las Perdices, por el Rfo Guayabal hacia el pueblo Guayabal, 10~ 66~ 900 1100 m, 19-22 Feb 1993, Meier 3386 (NY, VEN); Parque Nacional de Guatopo, steep slopes bordering Rio Santa Cruz, between Santa Teresa and Altagracia de Orituco, 14.5 km from Los Alpes, 12 km from Rancherfa Mi Querencia, 520 m, 6 Aug 1970, Moore et al. 9601 (BH); Guatopo National Park, along S end of La Macanilla trail, 32 km (by air/ NW of Altagracia de Orituco, 27 Aug 1979, Nee 17789 (NY): Guatopo National Park, region of "La Macanilla,'" 10~ 66~ 570 m, 18 Dec 2001, SmuffOr et al. 822 (MY. MYE VEN, Z): Parque Nacional Guatopo. along Rfo Santa Cruz. between Santa Teresa and Altagracia de Orituco, 450 m, 27 Oct 1966, Steyermark 97_573 (G, US); Cerros del Bachiller, nr. E end, above Quebrada Corozal, S of Santa Cruz, 10 km (by air) W of Cfpira, 10~ 65~ 20-700 m, 20-26 Mar 1978, Stevermark & Davidse 116530 (BH, MO); Cerros del Bachiller, W sector, between base and summit, above Quebrada Bachiller, S of Carlo, 10~ 65~ 20-690 m, Steyermark & Davidse 116565 IBH. MO, NY, VEN); Parque Nacional Guatopo, entre Santa Teresa y Altagracia de Orituco, 14.5 km de Los Alpes. 520 m, 29 Nov 1970. Steyermark & Braun. 104041 (BH, US, VEN): montafias de Guatopo, 22 Mar 1956, Tamayo 4177 (US); Altagracia de Orituco. Santa Teresa, Parque Nacional Guatopo, 11 May 1969, van Rooden & Akkermans 9_5 (U). [VOL. 55 sheath, the margin slightly fibrous near base; midvein 56-95 cm • 0.3-0.6 cm, loosely brownish gray tomentose, flat to channeled adaxially, rounded to flat abaxially; blade 0.70-1.20 X 0.23-0.30 m, dull green adaxially, grayish green and covered with a whitish farinose indumentum abaxially, deeply bifid at apex for ca. 42 cm; primary veins 4 0 - 7 0 per side, emerging at 2 5 - 3 0 ~ angle at midvein, mostly prominent abaxially, loosely brown tomentose. b!florescence branched to 1 order; peduncle 8 5 - 9 5 cm long, 1.5-2 cm diam., dorsiventrally compressed; prophyll 27-35 cm long, 2 - 3 . 5 ( - 5 ) cm diam., chartaceous, inserted at base of peduncle or 1 cm above it: peduncular bracts 2, with indumentum same as on prophyll, chartaceous, brown; first peduncular bract 4 5 - 5 8 • 2 - 2 . 5 ( - 4 ) cm, inserted 3 cm above base of peduncle, second peduncular bract 5 8 - 6 0 • 2.5-3.5 cm, inserted 4 cm above base of peduncle: inflorescence rachis 5 - 1 0 cm long, 0.5 cm diam., dorsiventrally to irregularly compressed, pale green at anthesis, almost glabrous; rachillae 9 - 1 6 , slightly pendent, the same color as inflorescence rachis, glabrous; basal rachillae 2 8 - 3 2 cm long, 0.5 cm diam.: middle rachillae 2 7 - 2 9 cm long, 5. ASTEROGYNE YARACUYENSE A . J. H e n d . & 0.5 cm diam.: apical rachillae 2 4 - 2 8 cm Steyerm. (Fig. 2 D - 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 m m apart from each other, covered Steyerm., Brittonia 38: 3 0 9 - 3 1 3 . 1986. with an inflexed lower lip, 2.5-3 • 2-5 TYPE: V E N E Z U E L A . Yaracuy: Cerro La ram. Staminate flowers obovoid, 6 - 8 mm Chapa, selva nublada al norte de Nirgua, long, 2 m m diam., slightly fragrant, white 1200-1400 m, 9 - 1 0 Nov 1967, J. A. at anthesis, the pedicel to 1 m m long: sepals Steyermark, G. Bunting & J. G. Wessels oblong-obovoid, 4 X 1-2 mm: petals obBoer 100311. (HOLOTYPE: VEN: ISOTYPE: long-obovoid, 6 - 7 X 1.5-2 ram; stamens BH) 8-10: filaments united into a tube to 4 mm long, the free parts 3-3.5 m m long, 0.5 mm Stem solitary, erect, sometimes basally 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 m m long, 3 mm diam., the pedicel 25(-31), reddish brown when young, green to 1 m m long; sepals oblong-ovate, 3 X 2 when mature, the older ones briefly persis- 2.5 mm; petals oblong, 3 x 2 mm; ovary 1 tent; sheath 12-30 X 7 - 9 cm, reddish mm long, 1.2 m m diam.: style 1.5 m m long: brown tomentose, the margins slightly fi- stigmatic branches 1 m m long; staminodial brous; petiole ( 1 8 - ) 2 5 - 3 5 X 1-1.5 cm, flat tube 2 m m long, the lobes 7 - 8 or someto channeled adaxially, angled to rounded times fewer by congenital union of 2 of abaxially, the indumentum same as on them, 2 m m long. Fruits not known. 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) L o c a l n a m e s a n d u s e s . - - n o t reported. Distribution and ecology. Known only f r o m the u n d e r s t o r y o f the C e r r o La C h a p a c l o u d forest, in the n o r t h e r n C o a s t a l Cordillera, state Yaracuy, V e n e z u e l a ; r e c e n t l y r e p o r t e d on the C e r r o E1 Z a p a t e r o o f the s a m e state (W. Meier, pets. c o m m . ) (Fig. 4); b e t w e e n 1 2 0 0 - 1 4 0 0 m. Additional specimens examined: VENEZUELA. Yaracuy: Dist. Nirgua, selva nublada del Cerro La Chapa, N of Nirgua, 12 Nov 1985, Henderson 502 (MYE NY): 5 km N of Nirgua. 10~ 68~ 28 Nov-1 Dec 1996, Stauffer et al. 265, 267 (VENZ), 5 km N of Nirgua. 10~ 68~ 4 Apr 1997. Stauffer et al. 319 (VEN/, 5 km N of Nirgua, 10~ 13'N, 68~ 27 Dec 2001, Stauffer & Stauffer 823 (MY, MYE VEN, Z), 9-10 Nov 1967, Steyermark et al. 100307 (BH, VEN), 100311 (VEN, HOLOT'~'PE; BH, ISOTYPE), 10~ 68~ 9-10 Nov 1967, Wessels Boer et al. 2032 (BH, MER, NY, U), 2036 (BH, MER, NY, U); Cerro Picacho, selva nublada en la fila, N de Nirgua, 1200-1360 m, 11 Mar 1967, Steyermark & Bunting 97661 (BH, US, VEN). Acknowledgments S p e c i a l thanks to the curators o f the herbaria A A U , B, B H , B M , C, CAY, C R , (3, K, M E R , M O , M Y E NY, P, U, U S , V E N , W I S , and Z. H e c t o r M a y k i n d l y p r o v i d e d i n f o r m a t i o n o n s p e c i m e n s d e p o s i t e d at B R H and M i r e y a C o r r e a p r o v i d e d databases c o n t a i n i n g i n f o r m a t i o n on the s p e c i m e n s d e p o s i t e d at P M A and S C Z . T h a n k s to U r sula H o f m a n n fo r p r o v i d i n g i n f o r m a t i o n on the s p e c i m e n s at G O E T , and to M a r y E n dress and J e a n - J a c q u e s de G r a n v i l l e f o r c o n s t r u c t i v e c o m m e n t s on the m a n u s c r i p t . F r e d J. Stauffer, L. P a p p a t e r r a , R i c a r d a Riina, R o d r i g o D u n o , and W. M e i e r h e l p e d d u r i n g field w o r k in V e n e z u e l a , and R u d o l f S c h m i d k i n d l y p r o v i d e d p h o t o s and inform a t i o n on A s t e r o g y n e m a r t i a n a in C e n t r a l A m e r i c a . M i k e T h i v is g r e a t l y t h a n k e d f o r h e l p i n g with the p h y l o g e n e t i c analysis. M a deline Harley provided important comm e n t s on the p o l l e n m o r p h o l o g y . U r s J a u c h p r o v i d e d g u i d a n c e w i t h the S E M w o r k . Visits to A A U , B M , C, K, NY, and U S by F. W. S t a u f f e r w e r e g r a n t e d by the Intern a t i o n a l P a l m S o c i e t y (IPS), P r o g r a m f o r S h o r t T e r m Visits o f the S m i t h s o n i a n Institution ( N M N H ) , and the K e w Latin A m e r ica Research Fellowships Program. The M i n i s t e r i o del A m b i e n t e y de los R e c u r s o s 353 Nat u r al es R e n o v a b l e s ( M A R N R ) and the Instituto N a c i o n a l de P a r q u e s ( I n p a r q u e s ) k i n d l y p r o v i d e d the p e r m i t s to c o l l e c t in Venezuela. Literature Cited Appun, K, F. 1871. Unter den Tropen. Hermann Costenoble, Jena. Asmussen, C 9B. 1999a. Toward a chloroplast DNA phylogeny of the tribe Geonomeae (PalmaeI. Pages 121-129. bz: A. Henderson & E Borchsenius, editors. Evolution, variation, and classification of pahns. Mere. New York Bot. Gard., vol. 83. Bronx, New York. 91999b. Relationships of the tribe Geonomeae (Arecaceae) based on plastid rpsl6 DNA sequences. Acta Bot. Venez. 22: 65-76. , W. J. Baker & J. Dransfield. 2000. Phylogeny of the palm family (Arecaceae) based on rpsl6 intron and trnL-trnF plastid DNA sequences. Pages 525-537. In: K. L. Wilson & D. A. Morrison, editors. Systematics and evolution of monocots. CSIRO, East Melbourne, Victoria, Australia. - & M. Chase 92001. Coding and noncoding plastid DNA in palm systematics. Amer. J. Bot. 88: 1103-1117. Baker, W. J., C. B. Asmussen, S. Barrow, J. Dransfield & T. A. Hedderson. 1999. A phylogenetic study of the palm family (Palmae) based on chloroplast DNA sequences from the trnL-trnF region. P1. Syst. Evol. 219: 111-126. Bentham, G. & J. D. Hooker. 1883. Genera Plantarum. Vol. 3(2). Reeve & Co., London. Berna], R. 1989. Endangerment of Colombian palms. Principes 33:113-128. Braker, E. & R. Chazdon. 1993. Ecological, behavioral and nutritional factors influencing use of palms as host plants by a neotropical forest grasshopper. J. Trop. Ecol. 9: 183-197. Braun, A. 1970. Palmas cultivadas de Venezuela. Acta Bot. 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H e n d . & S t e y erm. 2003] STAUFFER ET AL.: ASTEROGYNE (ARECACEAE) 355 Exsiccatae Only the first collector has been mentioned. Agostini, G. l0 (4) Aguilar, R. 2140, 2318 (2) Alverson, W. 94 (2) Anderson, R. 126 (2) Archer, W. 1978 (2) Aristeguieta, L. 1772, 3185 (4) Aulestia, C. 350, 455, 508, 1306 (2) Baker, R. 89021 (2) Barfod, A. 13 (2) Beach, J. H, 16545 (2) Bernal, R. 256, 259, 887, 1327, 1380 (2) Bernardi, A. 5633 (4) Blanc, M. 189 (1) Bonifaz, C. 3758 (2) Borchsenius, E 234, 242 (2) Bunting, G. 401 (2) Burger, W. 5599, 8076, 10616, 11173 (2) Callejas, R. 2759, 4220, 8528, 9202 (2) Chac6n, A. 973 (2) Chac6n, E. 1958 (2) Chac6n, I. 1311, 1615 (2) Chazdon, R. 16 (2) Churchill, H. 4122, 4186 (2) Cogollo, A. 4290 (2) Cook, O. E 61, 84, s.n. (2) Correa, M. 624 (2) Croat, T. 9933, 13202, 13884, 16834, 21997, 24524, 26052, 26161, 27201, 33683, 35353, 35740, 36459, 36461, 59889, 67689, 68156, 68405 (2) Cuatrecasas, J. 17375, 21300, 24195 (2) Danin, A. 77-5-2, 77-14-12 (2) Davidse, G. 23727, 31109, 31267 (2) de Granville, J. J. 7124, 7124-A (1) de Nevers, G. 3700, 3795, 3974, 4460, 5209, 5556, 5881, 5896, 5941, 5942, 6111, 6167, 6282, 6323, 7914, 8320 (2) Denslow, J. 2745 (2) Dodge, C. 9284 (2) Dransfield, J. 4859 (2) Duke, J. 11334, 11336 (2) Elias, T. 1769 (2) Folsom, J. 5901, 6179 (2) Fonnegra, R. 2839, 3024, 4756 (2) Forero, E. 8975, 9039 (2) Fuchs, H. R 21808, 21994 (2) Gentle, R 3743, 8635, 9072, 9074, 9669 (2) Gentry, A. 2803, 7542, 8818, 8868, 17445, 40972, 41092, 48546, 56871, 63735 (2); 24823 (4). G6mez, L. D. 19506, 19655, 20341 (2) Gonzfilez, A. 1296 (4) Gordon, B. 70 (2) Grayum, M. 4052 (2) Grijalva, A. 262, 380, 3481 (2) Gudifio, E. 1283 (2) Haber, W. 11580 (2) Hamilton, C. 1087, 2927, 2945 (2) Hammel, B. 1682, 1797, 4863, 5153, 12377(2), 12648 (2) Hari, R s.n. (2)] Henderson, A. 53, 714, 3048 (2); 502 (5) Herrera, H. 1188, 1332 (2) Holdridge, L. 5122 (2) Holm, R. 914, 970 (2) Hoover, W. 1241 (2) Janzen, D. 11098 (2) Juncosa, A. 633, 732, 1311, 1604, 1720, 2532 (2) Killip, E. 35293 (2) Knapp, S. 1417, 1772, 3777 (2) Knudsen, J. 604, 620, 623 (2) Koch, R. 5048 (2) Kress, W. J. 3778 (2) Lehmann, E C. 708 (2) Lems, K. 5199 (2) Lent, R. 80 (2) Levy, P. 505 (2) Lewis, W. 5558 (2) Liesner, R. 2924, 3223, 14825 (2) Maas, R J. M. 2803 (2) McDonagh, J. 246, 322 (2) McDowell, T. 886 (2) McPherson, G. 9716, 8080 (2) Meier, W. 3386 (4) Miller, J. 1095 (2) Molina, A. 17559 (2) Mondrag6n, M. 119 (2) Monsalve, M. 671, 927 (2) Moore, H. E. 6530, 6535, 6717, 8213, 9420 (2); 9601 (4) Morales, J. 1118 (2) Moreno, R 12280, 12580, 12771, 12979, 12985, 13032, 13151, 13287, 14821, 15013, 19142, 20602, 20640, 20661, 23621, 23890, 24015, 26083 (2) 356 BRITTONIA Mori, S. 2312, 2319, 4095, 4589, 4905, 6383 (2) Navarro, E. 76 (2) Nee, M. 17789 (4), 27905 (2) Neill, D. 1723, 4442, 7151 (2) Oersted, A. 6566 (2) Ortiz, E 297, 733 (2) Picado, A. 123 (2) Pipoly, J. 5111, 5323, 6112, 6143, 6168, 6223, 6290 (2) Pittier, H. 4226, 16658, 16717, 16922 (2) Porter, D. 4736 (2) Quesada, E 43 (2) Ramirez, J. 46, 167, 1131, 1364, 1770, 1860 (2) Ram/rez, V. 183 (2) Read, R. 640 (2) Rfos, D. 187 (2) Robles, R. 1148, 1192 (2) Rowlee, W. 1010 (2) Rueda, R. 2670, 3110, 3459 (2) Salick, J. 8075, 8106 (2) Sandino, J. 4782 (2) Schipp, W. 392 (2) Schmalzel, R. 1156 (2) Seemann, B. 163, s.n (2) Seymour. E 5905 (2) Shepherd, J. 863 (2) Soza, D. 347 (2) Sperry, J. 606 (2) Standley, R 25015, 53156 (2) [VOL. 55 Stauffer, E W. 824 (3); 822 (4): 265, 267, 319, 823 (5) Stevens, W. D. 4810, 4943, 6336, 6795, 6820, 8322, 12080, 12246~ 12686, 13065, 13505, 14927, 19343, 20743, 20936, 23489, 23651, 24026, 24331, 24451 (2) Stevenson, D. 1204 (2) Steyermark, J. 94803, 96088, 120915, 121713 (3); 39785, 90010, 97573, 104041, 116530, 116565 (4); 97661, 100307, 100311 (5) Sudgen, A. 351 (2) Sullivan, G. 159 (2) Sytsma, K. 1660 (2) Tamayo, F. 4177 (4) Tessene, M. 1329 (2) Thomsen, K. 187 (2) Tipaz, G, 2250~ 2693 (2) Tomlinsom R B. s.n. (2) Utley, K. 1093, 5975 (2) Valverde, O. 665 (2) van Rooden, J. 95 (4) Vincelli, R 164, 228 (2) von Wedel, H. 952, 1108, 2186 (2) Watson, S. 238 (2) Wendland, H. 56, s.n. (2) Wessels Boer~ J. G. 2032, 2036 (5) Wilbur, R. L. 15071 (2) Wilson, R 224 (2) Witherspoon, J. 8322 (2) Yuncker, T. 5002 (2) Zarucchi, J. 5056 (2)