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Manual de Plantas de Costa Rica

Main | Family List (MO) | Family List (INBio) | Cutting Edge
Draft Treatments | Guidelines | Checklist | Citing | Editors

Draft Treatments

ARECACEAE
By M. H. Grayum
English, final draft: placed 1/Apr./2000

Acoelorraphe

1 sp., S Florida, Bahamas, Cuba, S Mex. (Chis., Ver.) to CR, I. San Andrés (Col.).

Bailey, L. H. 1940. Acoelorraphe vs. Paurotis.--Silver-saw palm. Gentes Herb. 4: 361--365.

Acoelorraphe wrightii (Griseb. & H. Wendl.) H. Wendl. ex Becc., Webbia 2: 109. 1907. [Copernicia wrightii Griseb. & H. Wendl., in Griseb., Cat. pl. Cub. 220. 1866; Paurotis wrightii (Griseb. & H. Wendl.) Britton]. PALMA CUBA--Stems erect, cespitose, mostly 1.5--6 m tall and 5--12 cm diam., unarmed (except by persistent petiole bases). Petioles beyond sheath mostly 50--100 cm, fiercely armed marginally with triangular spines, with a conspicuous adaxial hastula, the sheath weathering to persistent mats of coarse, brownish fibers. Lf.-blades palmately deeply divided into bifid segments, to ca. [90 X 60 cm], with very brief midrib, ± silvery abaxially. Plants hermaphroditic, the infls. interfoliar. Prophyll short, ± enclosed in petiole sheath. Peduncle elongate, with 2 bracts much longer than prophyll. Infls. paniculately branched to 4 orders, exceeding the lvs., the rachillae ± densely white-tomentose. Fls. bisexual, ca. 1.8 mm long, yellowish, sweetly fragrant, spirally arranged in minutely bracteate clusters of (1--) 2--3 on low spurs. Sepals 3, ± connate basally. Petals 3, connate basally in a tube. Stamens 6, the filaments connate basally. Pistils 3 (just one developing to frt.), the styles connate, erect. Ripe frts. ca. 7--9 X 6--9 mm, globose, smooth, orange or becoming black, with apical stigmatic residue. Wet forests, 0--50 m; N Atl. slope, Llanura de los Guatusos (RNVS Caño Negro). Fl. 2, 7. [K. Flores et al. 34, CR, INB, MO.]

This is our only fan-lvd. palm with cespitose stems, as well as the only one with spiny petioles. In Costa Rica, it is known only from near the Nicaraguan border in the vicinity of Laguna Caño Negro (Prov. Alajuela)--the southernmost station for the genus. Here, it grows in large stands between the forest and the lake.

Acrocomia

2 spp., Mex. to Ven., S. Braz., Arg., Parag., Bol., Antilles.

Bailey, L. H. 1941. Acrocomia--preliminary paper. Gentes Herb. 4: 421--476.

Acrocomia aculeata (Jacq.) Lodd. ex Mart., Hist. nat. palm. 286. 1845. [Cocos aculeata Jacq., Select. stirp. amer. hist. 278, t. 169. 1763; A. belizensis L. H. Bailey; A. mexicana Karw.; A. panamensis L. H. Bailey; A. vinifera Oerst.]. COYOL--Stems erect, solitary, ca. 2--8 (--10) m tall and 25--40+ cm diam., armed with terete spines. Petioles beyond sheath [to ca. 50 cm long], spiny and ± tomentose, the sheath disintegrating into fibers. Lf.-blades pinnately compound, ca. 2--3.5 m long, the rachis spiny, the lflts. to at least 160 per side, linear, ± regularly spaced and arranged in various planes, abaxially whitish and usually variously pubescent. Plants monoecious, the infls. bisexual, interfoliar. Prophyll usually hidden within petiole sheath and soon splitting. Peduncle ca. 30--50 cm long, spiny, erect to arching or pendulous, with 1 much enlarged (ca. 70--100 cm), densely brown-tomentose and ± spiny, beaked, persistent, basal bract (spathe). Infls. racemosely branched, shorter than lvs. Fls. unisexual, spirally arranged in few, distant, bracteate triads (basally on rachillae) and solitary, densely packed male fls. (distally). Male fls. ca. 4--6 mm long, with 3 distinct sepals and 3 basally connate petals; stamens 6, adnate to base of corolla. Female fls. ca. 7--10 mm long, with 3 distinct sepals and 3 ± distinct petals; staminodia connate in a 6-toothed cup, the teeth usually bearing rounded, sterile anthers; pistil 1, 3-locular, pubescent; stigmas 3, reflexed. Ripe frts. ca. 3.0--3.5 X 3.0--4.5 cm, globose to somewhat oblate, smooth, greenish to yellow, with apical stigmatic residue. Dry and moist forests, 0--500 m (mostly in pastures and other disturbed sites); N Atl. slope (Llanura de los Guatusos), Pac. slope S to Río Grande de Tárcoles, and sporadically southward (Río Grande de Térraba, Pen. Burica). Fl. 2--6. Mex. (Jal.) to N Col., Venez., E Braz., N Arg., Parag., Bol., Gr. Ant. (PR, Rep. Dom.). [Little & Budowski 20309, USJ.]

Acrocomia aculeata is easily recognized by its stout, solitary stems, spinose, persistent lf.-blades with numerous, narrow lflts. arranged in various planes (giving the lvs. a plumose appearance), and dry-forest habitat. Acrocomia is one of the "cocoid" palms (see under Cocos for additional information); the genus is very close to Astrocaryum, from which it may be distinguished by the characters given in the key (couplet 13); Astrocaryum confertum and A. standleyanum are especially similar in having plumose foliage.

This is an abundant and characteristic species in the Guanacaste lowlands, where the frts. are consumed and the stones dispersed by cattle. The endosperm is also edible. A palm wine, vino de coyol, is produced seasonally from the sap of the trunk, and offered for sale locally.

Aiphanes

22 spp., CR to Venez., Trin., W. Braz., Peru, Bol., Gr. Ant. (Haiti, PR), Less. Ant.

Borchsenius, F. & R. Bernal. 1996. Aiphanes (Palmae). Fl. Neotrop. Monogr. 70: 1--95.

Aiphanes hirsuta Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11: 573. 1932. [A. fuscopubens L. H. Bailey].--Stems erect, cespitose, ca. 4--10 m tall and 3--8 cm diam., armed with terete to flattened spines. Petioles beyond sheath [ca. 9--35 (--55?) cm], spiny, the sheath disintegrating into fibers. Lf.-blades pinnately compound, ca. 0.8--1.7 m long, the rachis spiny and densely blackish hispidulous, the lflts. ca. 9--20 [(--26?)] per side, cuneate basally, praemorse distally, ± clustered and arranged in various planes, abaxially glabrescent. Plants monoecious, the infls. bisexual, interfoliar. Prophyll flattened, ± beaked, splitting but persistent. Peduncle ca. 43--145 cm long, spiny, with 1 much enlarged, basal bract. Infls. racemosely branched, the rachillae ca. 9--12 [--28?], blackish hispidulous. Fls. unisexual, spirally arranged in bracteate triads (basally on rachillae) and solitary or paired male fls. (distally). Male fls. 1.5--3.4 mm long, with 3 distinct sepals and 3 ± distinct or basally connate petals; stamens 6, ± distinct. Female fls. ca. 4.5--7 mm long, with 3 distinct sepals and 3 petals connate in basal half; staminodia connate in 6-toothed ring; pistil 1, 3-locular; stigmas 3, reflexed. Ripe frts. ca. 0.9--1.4 X 0.7--1.4 cm, globose to ± ellipsoid, smooth to minutely pustulate, red to purple, rostrate, with apical stigmatic residue. Wet forests, 900--1300 m; S Fila Costeña (C. Anguciana). Fl. 12. CR to NW Ec. [Aguilar et al. 2716, CR, INB, MO.]

Aiphanes belongs to the group of "cocoid" palms (see under Cocos for further information). Aiphanes hirsuta is the only Costa Rican palm with spinose foliage and wedge-shaped lflts. (cuneate at the base and praemorse distally, as in Iriartea or Socratea). This is a very rare species in Costa Rica, where it is known only from the steep, rocky, coastal slopes of Cerro Anguciana, the highest peak in the Fila Costeña. This represents the westernmost station for the genus. Our material belongs to subsp. hirsuta (CR to W Col.).

The South American Aiphanes aculeata Willd. [syn. A. caryotifolia (Kunth) H. Wendl.], which differs from A. hirsuta in having solitary (rather than cespitose) stems, is sometimes cultivated in Costa Rica as an ornamental (Aguilar V. MEAV-106, USJ)

Asterogyne

5 spp., Bel. and Guat. to Pac. Ec., Venez., Fr. Guiana.

Wessels Boer, J. G. 1968. The geonomoid palms. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 58(1): 1--202+.

Asterogyne martiana (H. Wendl.) H. Wendl. ex Hemsl., Biol. cent.-amer., Bot. 3: 409. 1885. [Geonoma martiana H. Wendl., Linnaea 28: 342. 1857]. SUITA--Stems usually decumbent in basal part, solitary, obsolete to 2+ m tall, 0.2--0.4+ cm diam., unarmed. Petioles beyond sheath ca. 30--68 cm, unarmed, the sheath reddish scaly and eventually splitting. Lf.-blades simple and bifid, rarely few-divided or (at least on herbarium sheets) -torn, ca. 47--130 X 15--28 cm, narrowly cuneate at base, unarmed. Plants monoecious, the infls. bisexual, interfoliar. Prophyll thin, papery, reddish tomentose. Peduncle ca. 40--93 cm, with 1 (--2) bracts similar to prophyll, but larger. Infls. spicate (very rarely) to racemosely or subdigitately branched with 2--7 (--8) rachillae; rachis obsolete to 3.5 cm. Fls. unisexual, fragrant, borne in bracteate triads in bilabiate pits. Male fls. ca. 4--5 mm long, with 3 narrow, briefly connate sepals and 3 petals connate for ca. 2/3 their length; stamens 6, the filaments connate basally. Female fls. ca. 6 mm long, with 3 distinct sepals and 3 petals connate for 2/3 their length; staminodia connate basally in 6-lobed tube; pistil 1, 3-locular; style elongate; stigmas 3, triangular. Ripe frts. ca. 9--10 X 3--5 mm, narrowly ovoid to, ellipsoid, smooth, dark purple or black, with basal stigmatic residue. Wet forests, 0--900 m; entire Atl. slope, Pac. slope S from RB Carara. Fl. 1, 4--9, 11--12. Bel. and Guat. to Pac. Ec. [A. Chacón 973, INB, MO.]

Asterogyne martiana is an abundant and familiar species throughout the humid lowlands of Costa Rica. It is easily recognized by its short-stemmed, solitary habit, simple, bifid, elongate lf.-blades, and long-pedunculate, subdigitate infls. Asterogyne is one of the geonomoid palms, characterized by having their floral triads deeply immersed in rachillar pits (see under Geonoma for further information on this group); it is most similar to Geonoma, from which it is technically distinguished by the characters given in the genus key (couplet 20). The combination of simple, bifid lf.-blades and a branched infl. immediately distinguish A. martiana from all other geonomoid palms in Costa Rica, except for the very rare and rather different Geonoma monospatha (which see), of higher elevations. Exceptional individuals of A. martiana with spicate infls. (reduced to a single rachilla) may closely resemble certain species of Geonoma, especially G. cuneata, which usually grows in the same places.

Although Asterogyne martiana is monoecious, plants may appear unisexual, since anthesis of both male and female fls. is synchronized, and the phases are well separated in time; furthermore, male fls. are immersed and not evident during female anthesis, and vice versa.

Astrocaryum

Stems erect, robust and usually arborescent, solitary (ours), densely armed with flattened spines or (A. alatum) only by the persistent, armed petiole-bases. Petioles abundantly spiny and often ± tomentulose or furfuraceous, the sheath splitting. Lf.-blades pinnately compound, the rachis spiny, the lflts. ± numerous, subequal to unequal, clustered or ± regularly spaced, arranged in one or various planes, abaxially whitish tomentulose, marginally spinulose (ours). Plants monoecious, the infls. bisexual, interfoliar. Prophyll thin, unarmed, mostly included within petiole sheath. Peduncle ± elongate, ± spiny, erect to pendulous, with 1 much enlarged, spiny, beaked bract (spathe). Infls. racemosely branched. Fls. unisexual, in bracteate triads basally on rachillae or on infl. rachis, with densely packed paired or solitary male fls. distally. Male fls. with 3 reduced, ± distinct sepals and 3 basally connate petals; stamens 6 (ours), adnate to base of corolla. Female fls. much larger, with sepals connate in truncate to shallowly 3-lobed cup or tube and petals (in ours) connate in a 3-toothed ovoid to cylindrical tube; staminodia (ours) connate in a truncate to sinuate or 6-toothed ring or cup; pistil 1, 3-locular; stigmas 3, ± erect or reflexed, borne on a beak. Ripe frts. smooth or asperous to spinose, ellipsoidal to obovoid, prominently beaked (ours), yellow-brown to orange, with apical stigmatic residue. 18--25 spp., Mex. (Oax., Ver.) to Guianas, Trin., SE Braz., Bol.

Astrocaryum is one of the "cocoid" palms (see under Cocos for further information). Among Costa Rican palm genera, Astrocaryum is distinctive in having stout, solitary stems, a subarborescent to arborescent habit, a fierce armature of flattened spines, and pinnately compound lf.-blades with ± numerous, abaxially whitish lflts. Only Acrocomia, distinguished by the characters given in the genus key (couplet 13), is likely to be confused. Because of their large stature and fierce armature, these are the most imposing of our palm genera for the botanical collector; thus, herbarium records are few and unrepresentative.

Burret, M. 1934. Die Palmengattung Astrocaryum G. F. W. Meyer. Repert. Spec. Nov. Regni Veg. 35: 114--158.

de Nevers, G. C., M. H. Grayum & B. E. Hammel. 1988. Astrocaryum confertum, an enigmatic Costa Rican palm rediscovered. Principes 32: 91--95.

1 Fertile stems usually < 7 m tall, unarmed except by the spiny, persistent petiole bases; lflts. < 70 per side, unequal (1--5+-costate), ± regularly spaced, arranged in a single plane; female fls. (and frts.) borne on main infl. axis at base of rachillae, the rachillae entirely staminate and deciduous in frt.; staminodial tube prominently 6-toothed, the teeth aristate; ripe frts. greenish brown to yellow-brown, prominently spinose in distal half...A. alatum

1' Fertile stems usually > 7 m tall, densely armed with bands of ± deflexed spines; lflts. > 70 per side, ± equal (mostly 1-costate), ± clustered and arranged in various planes (the lf.-blades with a plumose appearance); female fls. borne along basal portion of rachillae, only the distal (staminate) portion of the rachillae deciduous in frt.; staminodial tube truncate to sinuate; ripe frts. orange, ± uniformly asperulous or appressed-spinulose, virtually smooth to the touch

2 Infls. and infrs. erect, the peduncle < 1 m long; anthers ca. 1.8--1.9 mm long; staminodial ring truncate, glabrous; frts. with minute, appressed, blackish spinules; Atl. slope...A. confertum

2' Infls. erect to spreading, the infrs. long-pendent, the peduncle usually becoming > 1 m long; anthers ca. 2.9--3.3 mm long; staminodial ring sinuate, blackish fimbriate; frts. papillate and ± asperulous, but lacking blackish spinules; both slopes...A. standleyanum

Astrocaryum alatum H. F. Loomis, J. Wash. Acad. Sci. 29: 142, figs. 1--2. 1939. COQUILLO, COQUITO--Stems ca. 2--6 m tall and 12--18 cm diam., unarmed except by spiny, persistent petiole bases. Petioles beyond sheath ca. 1.0--3.0 m, spiny and densely and minutely spinulose. Lf.-blades ca. 2.5--4.5 m long, the lflts. ca. 18--37 per side, unequal (1--5+-costate), ± regularly spaced and arranged in a single plane, coarsely serrate or (the narrower ones) unequally bifid distally. Peduncle ca. 70--120 cm, erect to pendulous, with bract ca. 50--55 cm long, densely spiny and ± floccose. Infl. rachis ca. 17--25 cm. Fls. in triads restricted to infl. rachis at base of rachillae, with densely packed, paired or solitary male fls. on rachillae. Male fls. ca. 4--5 mm long, creamy white; anthers ca. 1.8--1.9 mm long. Female fls. ca. 8--10 mm long, with petals connate in an ovoid, distally 3-toothed tube; staminodia connate in a 6-toothed ring, the teeth aristate. Ripe frts. ca. 3.8--5.0 X 3.2--3.8 cm, spiny in distal half, smooth in basal half, ± obovoid, beaked, greenish brown to yellow-brown, borne in compact, spicate clusters (the entirely staminate rachillae deciduous). Wet forests, 0--800 m; entire Atl. slope, Pac. slope S from RB Carara. Fl. 5, 9, 11. E Hond. (Gracias a Dios) to E Pan. [Davidse & Herrera 30866, CR, MO.]

This is the most widespread Costa Rican Astrocaryum species, as well as the smallest one. It differs additionally from our other two species in having fewer, unequal lflts. arranged in a single plane, and larger, prominently spinose frts.

Astrocaryum alatum is particularly abundant in swamp forests, such as at PN Tortuguero. At the last-mentioned locale (and perhaps elsewhere), it may grow intermixed with the superficially very similar Manicaria saccifera (which see).

The endosperm of A. alatum seeds has somewhat the flavor and texture of coconut meat, and is sometimes consumed by local people.

Astrocaryum confertum H. Wendl. ex Burret, Repert. Spec. Nov. Regni Veg. 35: 136. 1934. [A. polystachyum H. Wendl. ex Hemsl., nom. nud.]. PEJIBALLE DE MONTAÑA, SURUBRE--Stems ca. 10--20 m tall and 14--20 cm diam., densely armed with bands of flattened, ± deflexed spines, with dense, basal cone of roots to ca. 0.5 m tall. Petioles beyond sheath ca. 0.7--1.3 m. Lf.-blades ca. 2.9--3.5 m long, the lflts. ca. 114--130 per side, subequal (mostly 1-costate), ± clustered and arranged in various planes, unequally bifid distally. Peduncle ca. 30--60+ cm, erect, with bract ca. 93--98 cm long, densely spiny (especially distally) and minutely brown-furfuraceous. Infl. rachis ca. 50--70 cm. Fls. in distant triads in proximal portion of rachillae, with densely packed, solitary male fls. in distal portion. Male fls. ca. 3--4 mm long, purplish distally, ± immersed in rachillae; anthers ca. 1.5--1.7 mm long. Female fls. ca. 9--11 mm long, with petals connate in a ± cylindrical, distally 3-toothed tube; staminodia connate in a narrow, truncate, marginally glabrous ring. Ripe frts. ca. 2.8--3.2 X 1.4--1.8 cm, ± uniformly appressed-spinulose (the spinules blackish), ellipsoidal to narrowly obovoid, beaked, orange. Wet forests, 0--500 m; N Atl. slope, Cord. Central (Sarapiquí region) to RNVS Barra del Colorado and PN Tortuguero. Fl. 5--6. CR (and ostensibly SE Nic.) to E Pan. [de Nevers & Hammel 7820, CR, MO.]

Astrocaryum confertum is the only spiny, solitary-stemmed, canopy or subcanopy palm in the wet forests of the N Atl. slope. This uncommon species is very similar superficially to A. standleyanum, from which it differs most conspicuously in its relatively short-peduncled infls., erect even to the stage of ripe frts.

Astrocaryum standleyanum L. H. Bailey, Gentes Herb. 3: 88, figs. 67--70. 1933. PEJIBAYE DE MONTAÑA--Stems ca. 8--15+ m tall and 12--25 cm diam., armed with bands of flattened, deflexed, ± whitish spines. Petioles beyond sheath ca. 0.5--1.5 m. Lf.-blades ca. 2.5--4 m long, the lflts. ca. 100--110 per side, subequal (mostly 1-costate), ± clustered and arranged in various planes, glossy adaxially, ± unequally bifid distally. Peduncle ca. 50--95 cm (to ca. 1.2 m in frt.), ± erect or spreading at anthesis but soon becoming pendulous, with bract ca. 84--145 cm long. Infl. rachis ca. 50--95 cm. Fls. in distant triads in proximal portion of rachillae, with densely packed male fls. in distal portion. Male fls. ca. 5--6 mm long, cream-colored, intensely and unpleasantly fragrant, ± immersed in rachillae; anthers ca. 2.9--3.3 mm long. Female fls. ca. 10--14 mm long, with petals connate in a narrowly ovoid, distally 3-toothed tube; staminodia connate in a sinuate, marginally blackish fimbriate ring. Ripe frts. ca. 3.3--4.5 X 2.1--3.2 cm, uniformly asperulous (but lacking blackish spinules), ± broadly ellipsoidal to obovoid, beaked, orange. Wet forests, 0--300+ m; S Atl. slope (RNVS Gandoca-Manzanillo), Pac. slope S from Punta Catedral. Fl. 5--9 (Pan.). CR to extreme NW Col. [Ocampo S. 3109, CR.]

Though vegetatively very similar to Astrocaryum confertum of the N Atl. slope, A. standleyanum is easily distinguished in fertile condition, even from a distance, by its massive, long-pendent infrs. It may also be mistaken for the cultivated pejibaye, Bactris gasipaes, which shares an arborescent habit, spiny vestiture, and plumose foliage. The last-mentioned species differs, however, in having cespitose stems and terete (rather than flattened) spines. Both the palmito and the fruits of A. standleyanum are edible, and the wood has been put to a variety of uses. Guaymí Indians make sombreros from the fibers (Ocampo S. 3109).

Attalea

Ca. 30 spp., Mex. (Nay.) to Venez., Trin., SE Braz., Parag., Bol., Haiti.

Glassman, S. F. 1977a. Preliminary taxonomic studies in the palm genus Orbignya Mart. Phytologia 36: 89--115.

----. 1977b. Preliminary taxonomic studies in the palm genus Scheelea Karsten. Phytologia 37: 219--250.

Attalea butyracea (Mutis ex L. f.) Wess. Boer, Pittieria 17: 312. 1988. [Cocos butyracea Mutis ex L. f., Suppl. pl. 454. 1782; Scheelea costaricensis Burret; S. gomphococca (Mart.) Burret; S. lundellii Bartlett; S. preussii Burret; S. rostrata (Oerst.) Burret; S. zonensis L. H. Bailey]. PALMA REAL--Stems erect, solitary, ca. 3--20 (--30) m tall and 25--50+ cm diam., unarmed. Petioles beyond sheath ca. 30--60 cm, with stout, stiff marginal fibers toward base, the sheath short, open. Lf.-blades pinnately compound, ca. 3.5--9 m long, often held in a ± vertical plane, the rachis unarmed, the lflts. ca. 180--235 per side, subequal, regularly spaced or ± clustered (especially basally), arranged in a single plane (or weakly disposed in various planes), unequally bifid apically, the terminal segment fenestrate (at least on juvenile plants). Plants monoecious, the infls. unisexual (male, and perhaps also female) or bisexual, interfoliar. Prophyll unknown. Peduncle to at least 1.2 m long, becoming pendulous, with 1 much enlarged bract (spathe) ca. 1--3 m long, becoming thick and woody, deeply grooved, splitting and expanding, persistent (much smaller distal bracts also present). Infls. racemosely branched, mostly ca. 70--110 cm long. Fls. unisexual, spirally arranged in bracteate triads (basally on rachillae) and solitary or paired male fls. (distally, or throughout in male infls.). Male fls. ca. 1.3--2.0 cm long, ochroleucous, intensely fragrant, with 3 minute, distinct sepals and 3 ± distinct, linear, fleshy petals; stamens 6, much shorter than petals, the anthers straight. Female fls. ca. 2.0--2.6 cm long, with 3 distinct sepals and 3 distinct petals; staminodia connate in a thick, truncate ring; pistil 1, 3-locular; style narrow; stigmas 3, reflexed. Ripe frts. ca. 4.5--6.2 X 2.1--2.8 cm, oblong-ellipsoid, ± smooth, orange-brown, with apical stigmatic residue. Moist and (occasionally) wet forests, 0--800 m; N Atl slope (Llanura de los Guatusos) and scattered southward, entire Pac. slope. Fl. 5, 9, 12. Mex. (Oax., Ver.) to N Col. and Venez., Trin., Tobago, Amaz. Col. to Bol. [de Nevers et al. 7766, CR, MO.]

This species is recognized by its large size, solitary stems lacking a crownshaft, spinelessness, long, multiply pinnate lf.-blades usually held in a vertical plane with the lflts. ± in the same plane, long-pedunculate clusters of rather large frts., and preference for seasonally dry forests. Attalea belongs to the group of "cocoid" palms (see under Cocos for further information); in fact, A. butyracea bears a rather strong resemblance to the coconut palm, Cocos nucifera (which has fewer but much larger frts.). It is also superficially somewhat similar to Welfia regia (see genus key, couplet 29), of wet forests.

Attalea butyracea, long known in Costa Rica as Scheelea rostrata, is a familiar and conspicuous species, spottily distributed yet locally abundant (as in the lower Coto Brus valley). It has become quite rare in the Guanacaste region. The lvs. have been much utilized for thatching, and the frts. are edible.

A vegetatively extremely similar species, Attalea cohune Mart. [syn. Orbignya cohune (Mart.) Dahlgren ex Standl.] occurs from cent. Mex. (Nay., Jal.) to E Nic., and has been reported (though not reliably) from CR. This species, which should be sought in NE CR, differs from A. butyracea in lacking marginal petiole fibers, and in having very different male fls. (with ovate, flattened petals and to ca. 24 stamens with curled anthers) and larger, relatively broader frts. (to at least 4.5 cm wide).

Bactris

Stems erect, slender or (occasionally) arborescent, cespitose, armed with terete or rarely (B. maraja) flattened spines. Petioles beyond sheath usually spiny, the sheath splitting and always spiny, usually ± ocreate (i.e., prolonged above petiole insertion). Lf.-blades simple and bifid to pinnately compound, the rachis usually spiny, the lflts. equal or unequal, linear, regularly spaced or clustered, arranged in one or various planes, abaxially glabrous to pubescent, usually spiny at least marginally. Plants monoecious, the infls. bisexual, interfoliar or becoming infrafoliar. Prophyll short, usually membranous and concealed in petiole sheath, unarmed. Peduncle usually short, ± curved, usually spiny, with 1 often spiny, usually expanding, persistent, basal bract (spathe). Infls. racemosely branched (ours). Fls. unisexual, spirally arranged in bracteate triads, sometimes also with solitary or paired male flowers especially distally on rachillae. Male fls. with 3 sepals connate in trilobed cup and 3 basally connate petals; stamens (in ours) 6. Female fls. with sepals connate in truncate or shallowly 3-lobed ring and equal to much longer petals similarly connate; staminodia absent, or connate in a membranous ring; pistil 1, 3-locular; stigmas 3, short. Ripe frts. smooth (ours), ovoid to obovoid or oblate, sometimes beaked, yellow-orange to red-orange or purplish black, with apical stigmatic residue. Ca. 65--100 spp., S Mex. (Oax., Ver.) to Guianas, Trin., SE Braz., Parag., Bol., Gr. Ant. (Cuba, Hispan., Jam.).

The combination of erect, cespitose stems and spiny stems and petioles distinguishes Bactris from all other Costa Rican palms except the rare and local Aiphanes (see genus key, couplet 14) and Desmoncus stans (which has cirrhous lflt. apices and spicate infls.). Bactris belongs to the group of "cocoid" palms (see under Cocos for additional information).

Bactris species are ubiquitous components of wet and moist lowland and mid-elevation forests in Costa Rica. Except for the cultivated B. gasipaes, ours are all relatively slender-stemmed, understory to subcanopy plants, rarely exceeding 10 m in height or 8 cm diam. All have subterete spines, except B. maraja, which has flattened spines.

de Nevers, G., A. Henderson & M. H. Grayum. 1996. Mesoamerican Bactris (Palmae). Proc. Calif. Acad. Sci. 49: 171--210.

1 Stems 4--20+ m tall and 8--14 cm diam.; lf.-blades pinnately compound, with > 55 lflts. per side; frts. to 5 X 3 cm, yellow to red; cult. only...B. gasipaes

1' Stems to ca. 10 m tall and 8 cm diam.; lf.-blades simple and bifid or pinnately compound, with < 55 lflts. per side; frts. < 5 X 3 cm or (if nearly as large) brown to purple-black; indigenous

2 Petioles, lf.-rachis, and peduncular bract armed with strongly flattened, mostly yellowish spines; ripe frts. purple-black; 0--450 m, both slopes...B. maraja

2' Petioles, lf.-rachis, and peduncular bract virtually unarmed, or armed with terete or subterete, variously colored spines; ripe frts. red to orange, or brownish to purplish black

3 Lf.-blades ± uniformly pubescent abaxially (and sometimes adaxially), the pubescence obvious to the touch

4 Petioles beyond sheath > 50 cm; lf.-blades pinnately compound, with > 15 lflts. per side; infl. rachillae > 50...B. glandulosa

4' Petioles beyond sheath < 50 cm; lf.-blades simple and bifid, or pinnately compound with < 15 lflts. per side; infl. rachillae < 50

5 Lf.-blades > 0.75 m long, tawny pubescent; infl. rachis 4.5--7.2 cm; rachillae ca. 24--45; Pac. slope...B. herrerana

5' Lf.-blades < 0.75 m long, whitish pubescent; infl. rachis (0.7--) 1.0--3.0 cm; rachillae 3--12; widespread

6 Lf.-blades always pinnately compound, with (4--) 7--10 lflts. per side, conspicuously pubescent on both surfaces; lflts. ca. 4.2--5.1 (--7.0) cm wide, strongly convex adaxially; infl. rachillae 8--12; Pac. slope...B. ana-juliae

6' Lf.-blades simple and bifid, or pinnately compound with 2--9 lflts. per side, usually conspicuously pubescent only abaxially; lflts. (when present) ca. 1.2--4.0 cm wide, plane or weakly convex adaxially; infl. rachillae 3--9; both slopes...B. hondurensis

3' Lf.-blades ± glabrous to minutely squamulose or spinulose abaxially, or inconspicuously pubescent only along veins (the pubescence not apparent to the touch)

7 Lf.-blades simple and bifid (rarely irregularly pinnately divided, then lflts. < 9 per side)

8 Lf.-blades < 2 m long and < 5 X longer than wide, slightly bicolored (± whitened abaxially); peduncle to ca. 10--12 cm, strongly recurved...B. grayumii

8' Lf.-blades > 2 m long and > 5 X longer than wide, ± concolorous; peduncle ca. 15--38.5 cm, straight and suberect (even in frt.)...B. militaris

7' Lf.-blades pinnately compound, with 9+ lflts. per side

9 Petioles beyond sheath obsolete to ca. 5 cm long; lf.-blades ca. 20--50 cm long; petioles and lf.-rachis with spines to at least 9.5 cm long, these yellowish medially and blackish at base and apex; ripe frts. purple-black; dry forest, 0--50 m, Pac. slope...B. guineensis

9' Petioles beyond sheath > 10 cm long; lf.-blades > 40 cm long; petiole and rachis virtually unarmed to variously spiny, the spines uniformly blackish or (B. dianeura) bicolored; ripe frts. orange to red or (B. major) brown to purple-black; widespread

10 Lflts. regularly spaced and arranged in a single plane throughout

11 Petioles and lf.-rachis formidably spiny with ± blackish spines to 11+ cm long; lflts. 25--48 per side, 1--3.5 cm wide (medial ones); peduncle 15--40 cm; infl. rachillae (3--) 5--10 (--17), 14--23 cm long; ripe frts. 3--4.5 X 1.7--3.5 cm, brown to reddish purple or purple-black; N Atl. slope, Pac. slope...B. major

11' Petiole and lf.-rachis unarmed, or sparsely spiny; lflts. 15--31 per side, 3.5--7 cm wide (medial ones); peduncle ca. 10 cm; infl. rachillae 20--30, 4.5--11 cm long; ripe frts. ca. 1.5--1.8 X 0.8--1.6 (--2.0) cm, orange; N Atl. slope...B. polystachya

10' Lflts. clustered and arranged in two or more planes, at least in basal half of lf.-blade

12 Lflts. with prominent marginal spines, the larger ones ca. 2.5--4 cm long; ca. 30--250 m, Atl. slope Cord. Central...B. longiseta

12' Lflts. with inconspicuous marginal spines, always < 1 cm long; widespread

13 Peduncle 9.5--23 cm long; infl. rachis 3.5--11 cm; infl. rachillae ca. 14--29 cm long; 0--150 m, Atl. slope...B. coloradonis

13' Peduncle ca. 5.5--12.5 (--16) cm long; infl. rachis ca. (0.5--) 1--4.5 cm; infl. rachillae ca. (2--) 4.3--11.5 cm long; widespread

14 Infl. rachillae ca. 40--60+, filiform; 0--700 m, Pac. slope...B. baileyana

14' Infl. rachillae ca. (2--) 7--31, stouter; Atl. slope, or above 1100 m on Pac. slope

15 Lflts. usually with prominent cross-veins; petioles and lf.-rachis sparsely but usually formidably armed, the spines (especially on younger lvs.) often reddish (at least medially); infl. rachillae (2--) 7--17; calyx in frt. often 3-lobed; frts. prominently rostrate; (550--) 800--1600 m...B. dianeura

15' Lflts. usually with inconspicuous cross-veins; petioles and lf.-rachis unarmed, or armed with uniformly blackish spines; infl. rachillae 8--31; calyx in frt. truncate; frts. scarcely or weakly rostrate; 0--800 (--1300) m

16 Petioles beyond sheath < 1/2 length of lf.-blade, sparingly spiny; lflts. slightly bicolored (± whitened abaxially); infl. rachis 1--2.5 cm long, the rachillae (13--) 17--31; frts. 0.8--1.8 X 1.1--1.9 cm, ± evidently rostrate...B. caudata

16' Petioles beyond sheath > 1/2 length of lf.-blade, unarmed; lflts. ± concolorous; infl. rachis 3--4.5 cm long, the rachillae 8--23; frts. 1.0--1.3 X 0.8--1.1 cm, scarcely rostrate...B. gracilior

Bactris ana-juliae Casc., in press.--Stems to ca. 1.8--3 m tall and 1.2 cm diam., the internodes spiny. Petioles beyond sheath ca. 11.2--16 (--25.5) cm, covered with spines, the spines ca. (0.5--) 1.6--3.5 cm, yellowish medially and brownish black at either end. Lf.-blades pinnately compound, (0.33--) 0.4--0.55 m long, the rachis whitish pubescent and with spines as petiole, lflts. (4--) 7--10 per side, clustered [and arranged in various planes], 15--22.5 (--28.7) X 4.2--5.1 (--7) cm (medial ones), [lanceolate to oblanceolate], without conspicuous cross-veins, whitish pubescent on both surfaces, marginally spinulose at least distally. Peduncle 5--8 cm, becoming curved, with bract ca. 11--14.6 cm long, covered with brownish black spines. Infl. rachis (0.7--) 1.4--1.7 cm; rachillae 8--12, (2--) 3.2--5 cm long, glandular-furfuraceous. Male fls. unknown. Female fls. ca. 2.1--2.8 mm long. Ripe frts. to ca. 1.2 X 1.0 cm, obovoid, abruptly rostrate, reddish orange, striate. Wet forests, 900--1000 m; S Pac. slope, Fila Costeña (Fila Tinamaste). Fl. 3--4. ENDEMIC. [Valverde 773, CR, MO, USJ.]

This local endemic is distinctive among all Costa Rican congeners in its pinnate lf.-blades that are conspicuously pubescent on both surfaces (rather than just abaxially); only B. hondurensis may exceptionally exhibit this combination of features (see key, couplet 6). Bactris ana-juliae bears a general resemblance to B. dianeura, but the latter sp. has larger lf.-blades with more numerous, relatively narrower lflts. In life, plants of B. ana-juliae are described as striking by virtue of their strongly convex, sigmoid lflts., glossy dark green and becoming paler toward the long-aristate apex, which points vertically toward the ground.

Bactris baileyana H. E. Moore, in L. H. Bailey & H. E. Moore, Gentes Herb. 8: 155, fig. 59. 1949. [B. glandulosa var. baileyana (H. E. Moore) de Nevers].--Stems ca. 2--5 m tall and 3.5--8 cm diam., the internodes spiny. Petioles beyond sheath 40--80 cm, white- to brownish tomentose, formidably (rarely weakly) spiny at least basally, the spines to 3--7 cm, uniformly blackish. Lf.-blades pinnately compound, 1.1--1.8 m long, the rachis unarmed to remotely spiny, lflts. 21--30 per side, clustered and arranged in various planes (except distal 4--5 pairs), 45--76 X 3--8 cm (medial ones), narrowly oblanceolate, sparingly and inconspicuously pubescent along veins to glabrescent abaxially, marginally spinulose distally. Peduncle (5--) 10.5--15 cm, recurved, with bract ca. 22--30 cm long, densely long-hirsute with tawny to reddish spines. Infl. rachis 5.5--10 cm; rachillae ca. 50--70, ca. 4-11 cm long, densely glandular and pubescent. Male fls. pedicellate, 2.5--3.5 mm long. Female fls. ca. 4--5 mm long, the corolla ca. 2--3 times longer than calyx, ± puberulent to glabrescent. Ripe frts. 1.0--1.2 X 0.9--1.2 cm, globose to turbinate, prominently rostrate, bright orange to red, striate. Wet forests, 0--700 m; Pac. slope, Pen. Nicoya (Bejuco, RNVS Curú), vic. Miramar, and S from RB Carara. Fl. 5--7. CR and W Pan. [I. Chacón 1957, CR.]

This and the pubescent-lvd. Bactris glandulosa are the only Bactris species in the wet forests of the Costa Rican Pac. lowlands which have fully pinnate lf.-blades with the lflts. clustered and arranged in various planes. Bactris baileyana and B. glandulosa have very similar infls., unique among our species in their very numerous (50--80+), filiform rachillae (B. herrerana also has filiform rachillae, but fewer). De Nevers et al. (1996) treated B. baileyana as a variety of B. glandulosa, but the former taxon differs not only in its comparatively glabrous foliage, but also in having generally much longer (> 3 cm) petiole spines, lflts. clustered and arranged in various planes over most of the rachis, and shorter (relative to the calyx), non-setose pistillate corollas. At sites where they co-occur, such as RB Carara, these two taxa have different flowering and fruiting phenologies, and appear to behave as separate species.

Bactris caudata H. Wendl. ex Burret, Repert. Spec. Nov. Regni Veg. 34: 230. 1934. [B. dasychaeta Burret].--Stems ca. 1--6 m tall and 2--3.5 cm diam., the internodes spiny or not. Petioles beyond sheath ca. 50--96 cm, sparsely spiny. Lf.-blades pinnately compound, ca. 1--2 m long, the rachis spineless or remotely spiny (with spines to ca. 7 cm), lflts. (3--?) 9--26 per side, clustered and arranged in various planes, (21--) 40--55 (--61.5) X 2.5--6.2 cm (medial ones), linear to lance-elliptic, concave and ± glaucous abaxially, marginally spinulose. Peduncle 7--12.5 cm, strongly recurved, with bract ca. 11--36 cm long, sparsely to densely black-spiny. Infl. rachis ca. 1--2.5 cm; rachillae (13--) 17--31, 5--10 cm long, densely glandular-furfuraceous. Male fls. ca. 3--4 mm long. Female fls. ca. 2.5--5 mm long. Ripe frts. ca. 0.8--1.8 X 1.1--1.9 cm, oblate to obovoid or turbinate, ± evidently rostrate {verify!}, orange to orange-red. Wet forests, ca. 50--850 (--1300) m; entire Atl. slope. Fl. 2, 4, 6, 8. SE Nic. to W Pan. (Bocas del Toro). [Grayum 11070, INB, MO.]

Bactris caudata is a little-known species, one of several Atl. slope Bactris with pinnately compound lf.-blades in which the lflts. are clustered and arranged in various planes. Among these species, B. caudata is best distinguished by its abaxially ± glaucous lflts. and very congested infls. and infrs., with the rachis < 3 cm long. It is perhaps most closely related to the generally simple-lvd. B. grayumii (which see). {Two spp. may be included here: one with rostrate frts. and a lobed perianth, the other with non-rostrate frts. and a truncate perianth.}

Bactris coloradonis L. H. Bailey, Gentes Herb. 3: 104, fig. 81. 1933. [B. porschiana Burret].--Stems ca. 3--12 m tall and 3--8 cm diam., the internodes spiny. Petioles beyond sheath ca. 24 cm, whitish tomentose and spiny (especially toward base). Lf.-blades pinnately compound, ca. 0.9--2.1 m long, the rachis whitish tomentose but usually spineless, lflts. 17--38 (--40) per side, clustered and arranged in various planes, 21--86 X 3--8.5 cm (medial ones), linear-lanceolate, sometimes with spines abaxially, marginally unarmed to spinulose. Peduncle 9.5--23 cm, recurved, with bract ca. 27--37+ cm long, sparsely to densely spinose. Infl. rachis ca. 3.5--11 cm; rachillae 20--51, 14--34 cm long, glandular-furfuraceous. Male fls. 3+ mm long. Female fls. 2.5--6 mm long. Ripe frts. (1--) 1.3--1.9 (--2.5) X (0.9--) 1.2--1.6 (--2) cm, broadly ellipsoidal to obovoid, briefly rostrate, bright orange to red. Wet forests, 0--400 m; Atl. slope Cords. Central (to RNFS Barra del Colorado and Tortuguero region) and Talamanca. Fl. 8--10. SE Nic., polytypically to Pac. Col. and perhaps Ec. [Davidse & Herrera 31101, CR, INB, MO.]

This is the largest Costa Rican Bactris species with pinnately compound lf.-blades in which the lflts. are clustered and arranged in various planes (except for the cult. B. gasipaes). The infls. of B. coloradonis are also notably large in all of their parts (peduncle, rachis, rachillae, etc.). The frts. are distinctive in being only briefly rostrate and not or inconspicuously striate, and in having a truncate corolla (cf. also B. gracilior). Collections from Cord. Talamanca [e.g., Gómez et al. 20453 (MO), Herrera & Chacón 2354 (CR? INB?)] have unusually large frts. (ca. 2.5 X 2.0 cm).

Costa Rican material (for which the name Bactris porschiana is available) differs in some respects from the Panamanian type of B. coloradonis, and may prove to merit taxonomic distinction at some level.

Bactris dianeura Burret, Repert. Spec. Nov. Regni Veg. 34: 217. 1934.--Stems ca. 1--5 (--8) m tall and 1--3 cm diam., the internodes spiny. Petioles beyond sheath ca. (10--) 20--80 cm, sparsely spiny. Lf.-blades pinnately compound, (0.45--) 0.6--0.9(--1.25) m long, the rachis sparsely spiny (with spines to ca. 9 cm), lflts. (10--) 12--20 per side, clustered and arranged in various planes, (14--) 23--50 (--56) X (1.4--) 2.7--4.5 (--6.3) cm (medial ones), lanceolate to oblanceolate, obscurely to conspicuously cross-veined, marginally spinulose at least distally. Peduncle 5.5--12.5 (--16) cm, becoming tightly curved, with bract ca. (6--) 9--26 cm long, densely spinose. Infl. rachis obsolete (rarely) or ca. (0.5--) 1.6--3.4 cm; rachillae (2--) 7--17, (2--) 4.3--11 cm long, glandular-furfuraceous. Male fls. 3--4 mm long. Female fls. ca. 3--5.5 mm long. Ripe frts. 1.1--1.8 X 1.1--1.6 cm, subglobose to (usually) obovoid or turbinate, prominently rostrate, bright orange, finely striate. Wet forests, (550--) 800--1600 m; Atl. slope all major cords., rare (> 1100 m) on Pac. slope, Cords. Guanacaste, Tilarán, and Talamanca, Montes del Aguacate, Fila Costeña. Fl. 2, 4--8. N Nic. to W Pan. [Grayum 7028, CR, MO.]

This species has the highest average elevational range of any Mesoamerican Bactris, and is the only Bactris species found at classic cloud-forest sites such as Monteverde, La Palma, and Tapantí. It is further distinguished by its usually formidably spiny petioles and lf.-rachises (with the spines often reddish medially), clustered and staggered lflts. with conspicuous cross-veins, smallish infls. with relatively few rachillae, and usually obovoid, prominently rostrate, finely striate frts. with a ± trilobate perianth. Bactris dianeura is perhaps most similar to the rare B. ana-juliae, which differs in having smaller, densely pubescent lf.-blades with fewer, shorter, and relatively broader lflts.

Bactris gasipaes Kunth, in Humb., Bonpl. & Kunth, Nov. gen. sp. Quarto ed. 1: 302, Folio ed. 1: 242. 1816. [Guilielma utilis Oerst.]. PEJIBAYE, PEJIVALLE--Stems ca. 4--20+ m tall and 8--14 cm diam., the internodes typically spiny (though spineless varieties exist). Petioles beyond sheath ca. 48--120 cm, with spines in 3 rows abaxially. Lf.-blades pinnately compound, 1.9--3.2 m long, the rachis virtually unarmed to densely spiny, lflts. 60--127+ per side, weakly clustered, arranged in various planes, 52--85 X 2--5.5 cm (medial ones), linear, asymmetrically bifid apically, marginally unarmed to spinulose. Peduncle 20--28 cm, gently recurved, unarmed, with bract ca. 47--71 cm long, moderately to densely spiny. Infl. rachis ca. 15--23 cm; rachillae 46--62, 17--40 cm long, glandular-furfuraceous Male fls. 3--5.5 mm long. Female fls. 5--7 mm long. Ripe frts. to 5 X 3 cm, ovoid, yellow to red. Wet forests, 0--700 (--1200) m; both slopes, always in association with humans. Fl. 1--12, but mainly 1, 5--7 (Atl. slope) or 1--5 (Pac. slope). Widely cult. (Guat. to Guianas, Trin., Braz., Bol.), but unknown in wild state. [Mattos S. & Castro E. 2886, CR.]

Bactris gasipaes, known only in cult., is by far the largest Bactris species in Costa Rica; it also has the longest lf.-blades with the most lflts., and the largest frts. It might be confused with some spp. of Astrocaryum, especially A. standleyanum, but differs in its cespitose habit and subterete, rather than flattened, spines. This species is best known for its edible frts., a staple market item throughout Costa Rica, but has also become the most important commercial source of palmito.

Bactris glandulosa Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1858: 184. 1859. [B. bifida Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1858: 44. 1859, non Mart.; B. alleniana L. H. Bailey; B. fusca Oerst.].--Stems ca.(1.5--) 3.5--5 m tall and 1.5--4 cm diam., the internodes with scattered spines. Petioles beyond sheath (19--) 62--160 cm, virtually unarmed to weakly spiny basally, the spines usually < 3 cm, uniformly blackish. Lf.-blades simple and bifid (very rarely) or pinnately compound, (0.6--) 0.8--1.5 m long, the rachis unarmed to spiny (as petiole), lflts. (2--3--) 23--30 per side, clustered and arranged in various planes in basal 1/2 to 2/3, or regularly spaced and arranged in a single plane virtually throughout, 45--62 X 2--6 cm (medial ones), linear-lanceolate to narrowly elliptical, narrowly oblanceolate, ± densely and uniformly tawny-pilose (at least along veins) abaxially (and sometimes adaxially as well), marginally spinulose. Peduncle 4--8.5 cm, recurved, with bract ca. (13--) 15--24 cm long, densely tawny-hirsute. Infl. rachis ca. 3.5--6 cm; rachillae (24--) 60--80+, 4--8 cm long, densely glandular-furfuraceous and puberulent. Male fls. pedicellate, ca. 2--3 mm long. Female fls. ca. 4--7 mm long, the corolla ca. 4--5 times longer than calyx, covered with long, contorted, blackish setae. Ripe frts. 0.8--1.0 X 0.7--1.2 cm, globose to turbinate, prominently rostrate, orange-red, striate. Wet forests, 0--1150 m; Atl. slope S from vic. Guápiles, Pac. slope S from RB Carara. Fl. 4--7, 9, 12. CR to Pac. Col. [Grayum & Herrera 9235, CR, MO.]

This is the only Costa Rican Bactris species with pinnately compound lf.-blades and numerous (> 20 per side) lflts. that are conspicuously pubescent abaxially. The likewise pubescent-lvd. B. herrerana and B. hondurensis may also have pinnately compound blades, but with fewer lflts. (see key, couplet 4). Bactris glandulosa is further distinctive in its numerous, filiform infl. rachillae, a feature shared only with B. baileyana and B. herrerana (which see), among Costa Rican species.

Bactris gracilior Burret, Repert. Spec. Nov. Regni Veg. 34: 216. 1934. [B. aureodrupa L. H. Bailey; B. longipetiolata H. Wendl. ex Hemsl., nom. nud.].--Stems 1--4 m tall and ca. 2--2.5 (--5?) cm diam., the internodes spiny. Petioles beyond sheath ca. 20--110 cm, unarmed (the sheath densely short-spiny). Lf.-blades pinnately compound, 0.4--1.5 m long, the rachis unarmed or (rarely) spiny, lflts. 10--24 per side, clustered (at least in basal portion of blade) and arranged in various planes, 18--41 (--50) X 2.0--4.4 cm (medial ones), narrowly elliptical to oblanceolate, ± spinulose to spinose along veins abaxially, marginally spinulose distally. Peduncle 6--10 cm, curved, with bract (10--) 15--27 cm long, virtually glabrous to sparsely short-spiny. Infl. rachis 3--4.5 cm; rachillae 8--23, (3--) 7--11.5 cm long, densely glandular-furfuraceous. Male fls. ca. 2--3.5 mm long, mostly pedicellate, the pedicels to ca. 3 mm. Female fls. 2--4 mm long. Ripe frts. ca. 1.0--1.3 X 0.8--1.1 cm, ellipsoidal to obovoid, scarcely rostrate, orange to red. Wet forests, 0--800 (--1050) m; entire Atl. slope. Fl. 5, 7--9. SE Nic. to E Pan. [Robles 1668, CR {sterile}, MO.]

Bactris gracilior is one of several Atl. slope species with pinnately compound lf.-blades in which the lflts. are clustered and arranged in various planes. Though rather nondescript, it is distinctive in the field by its smallish size, relatively long petioles unarmed beyond the sheath, generally unarmed lf.-rachises, and soft foliage. As noted in the original description, the frts. are in every way like smaller versions of B. coloradonis frts. (see under the latter species). Herbarium specimens of B. gracilior are recognizable by the reddish brown coloration of the lflts.

Bactris grayumii de Nevers & Andrew Hend., in de Nevers, Andrew Hend. & Grayum, Proc. Calif. Acad. Sci. 49: 188, fig. 4. 1996.--Stems 1.5--3.5+ m tall and ca. 2.2--3 cm diam., the internodes usually spiny. Petioles beyond sheath ca. 21--65 cm, spiny at least toward base. Lf.-blades simple and deeply bifid (perhaps occasionally irregularly pinnate), to 0.6--1.3 (--1.6) m X 22--45 cm, the rachis unarmed or formidably spiny (with spines to at least 7.5 cm), concave and ± glaucous abaxially, marginally spinulose to (distally) spiny. Peduncle to ca. 10--12 cm, strongly recurved, virtually unarmed to appressed-spiny, with bract ca. 18--27 cm long, ± densely short-black-spiny. Infl. rachis 1.8--4.5 cm; rachillae 13--30, 4--9 cm long, densely glandular-furfuraceous. Male fls. unknown. Female fls. 2--4 mm long. Ripe frts. ca. 1.1--1.4 X 0.9--1.6 cm, obovoid to turbinate, neither striate nor rostrate, orange or red, with calyx truncate. Wet forests, 0--50 (--800) m; N Atl. slope, Cord. Guanacaste (Lago Cote) to RNFS Barra del Colorado, PN Tortuguero. Fl. 5, 9, 12. Cent. Nic. to CR. [Grayum et al. 8995, CR {sterile}, MO.]

This is one of just four Costa Rican Bactris species that have exclusively or potentially simple lf.-blades. It differs from B. herrerana and B. hondurensis in having subglabrous lf.-blades and more numerous infl. rachillae, and from B. militaris in having relatively broader lf.-blades and shorter, recurved peduncles (see key, couplet 8). It may be most closely related to the pinnate-lvd. B. caudata, however, with which it shares abaxially ± glaucous lf.-blades and condensed infls. (with the rachis very short).

Bactris grayumii may be locally abundant at some sites, such as RNFS Barra del Colorado (where it grows together with B. militaris).

Bactris guineensis (L.) H. E. Moore, Gentes Herb. 9: 251. 1963. [Cocos guineensis L., Mant. pl. 137. 1767; B. minor Jacq., nom. illeg.; B. oraria L. H. Bailey]. HUISCOYOL, VIZCOYOL--Stems 0.6--3 m tall and ca. 2.6--3 cm diam., [the internodes spiny or not]. Petioles beyond sheath obsolete to ca. 5 cm, moderately to densely spiny, the spines to at least 9.5 cm, black at base and apex, yellowish medially. Lf.-blades pinnately compound, 0.2--0.5 m long, the rachis spiny (as petiole), lflts. 20--42 per side, regularly spaced or weakly clustered, arranged in ± the same or in various planes, 12--30 X 0.9--2 cm (medial ones), linear, sometimes spinose along veins abaxially, marginally spinulose, ultimately deciduous. Peduncle 10--20 cm, straight to slightly curved, with bract 19.5--35 cm long, moderately yellowish spiny. Infl. rachis 2--5.5 cm; rachillae 9--30, 8--11.5 cm long, densely glandular-furfuraceous. Male fls. ca. 4 mm long. Female fls. 3--4 mm long. Ripe frts. 1.0--1.6 X 1.2--2 cm, oblate, prominently rostrate, reddish purple to purple-black. Dry forests (moist transition), 0--50 m; Pac. slope S to vic. Tivives. Fl. 2. W cent. Nic. to N Col., Venez. [Crow & Rivera 6218, CR {not!}, MO.]

Bactris guineensis is easily recognized by its usually small stature, very short or obsolete petioles, formidable spines yellowish medially, small lf.-blades with ± closely spaced lflts., and purple-black frts. This and B. major are the only Bactris species in the drier, northern portions of the Pac. slope, though B. guineensis occurs in more exposed sites. An alcoholic beverage may be prepared from the frts.

Bactris herrerana Casc., in press.--Stems to 2--3 m tall, 1.5--2.5 cm diam., usually spiny. Petioles beyond sheath (15--) 20--28 (--43) cm, covered with short, black spines. Lf.-blades simple and bifid or (rarely) irregularly pinnately compound, 0.8--1.2 m X [???] cm, the rachis pubescent abaxially and with black spines, lflts. (when present) 2--3 per side, [regularly spaced to weakly clustered, 18--23 X 1.2--4.0 cm (medial ones), narrowly elliptical to linear], short-tawny-pubescent abaxially, marginally spinulose distally. Peduncle 3.5--7 cm, curved, with bract 12--18 cm long, covered with blackish brown and yellowish spines. Infl. rachis 4.5--7.2 cm; rachillae 24--45, 4.5--7.5 cm long, densely glandular-furfuraceous. Male fls. ca. 2--3.5 mm long. Female fls. ca. 3--4.5 mm long. Ripe frts. (0.7--) 1.1--1.3 X (0.9--) 1--1.2 (--1.5) cm, obovoid, abruptly rostrate, red, striate. Wet forests, 200--950 m; Pac. slope S from ZP Cerros de La Cangreja (Puriscal). Fl. 3--5. ENDEMIC. [Henderson et al. 1824, INB, MO.]

Its larger lvs. and larger inflorescences with more numerous, filiform rachillae sharply distinguish Bactris herrerana from B. hondurensis, the only other Costa Rican congener that may have simple, pubescent lf.-blades. The filiform infl. rachillae of B. herrerana are shared only with B. baileyana and B. glandulosa. The latter more closely resembles B. herrerana in its pubescent foliage and identical pistillate corollas (see under B. baileyana); indeed, the entity now known as B. herrerana was previously interpreted as comprising merely aberrant specimens of B. glandulosa. However, the last-mentioned sp. is distinct in having fully pinnate lf.-blades and more numerous (ca. twice as many) infl. rachillae.

Bactris hondurensis Standl., Trop. Woods 21: 25. 1930. [B. paula L. H. Bailey; B. pubescens Burret; B. standleyana Burret; B. wendlandiana Burret].--Stems (0.5--) 1--2.5 (--4) m tall and 0.5--1.5 cm diam., unarmed. Petioles beyond sheath (6--) 16--27 cm, unarmed to sparsely spiny, the spines to 7+ cm, uniformly blackish or yellowish medially and blackish at either end. Lf.-blades simple and bifid or pinnately compound, (0.21--) 0.35--0.7 m X (16.5--) 26--39 cm, the rachis unarmed or spiny as petioles, lflts. (when present) 2--9 per side, regularly spaced to weakly clustered, 18--23 X 1.2--4.0 cm (medial ones), narrowly elliptical to linear, densely and softly white-pubescent abaxially (and sometimes adaxially), marginally spinulose distally. Peduncle 3--12.5 cm, becoming arched, with bract 8--18 cm long, moderately to densely soft-spiny. Infl. rachis 1--3 cm; rachillae 3--9, 2--6 cm long, heavily glandular-furfuraceous. Male fls. ca. 2.5 mm long. Female fls. ca. 2--3.5 mm long. Ripe frts. 0.8--1.3 X 0.7--1.2 cm, broadly ellipsoidal to turbinate, briefly rostrate, orange to red. Wet forests, 0--1200 m; Atl. slope all major cords. (to RNVS Caño Negro and RNFS Barra del Colorado), Pac. slope Cord. Guanacaste, and (infrequently) S from RB Carara. Fl. 4--7, 11. Atl. Hond. to N Pac. Ec. [Ivey & Haber 123, CR {sterile!}, MO.]

This and B. herrerana (see key, couplet 5) are the only Costa Rican Bactris species that may have simple lf.-blades that are conspicuously pubescent abaxially. Less frequent pinnate-lvd. specimens of B. hondurensis may also be confused with the very local B. ana-juliae (see key, couplet 6) or the more widespread B. glandulosa (which has larger lf.-blades with more numerous lflts.). Like many other Bactris species, B. hondurensis is quite variable in spininess; both formidably spiny and unarmed lvs. may occur even on the same stem. If the lf. pubescence is not noted, unarmed or weakly armed individuals may be easily mistaken (in sterile condition) for superficially similar Geonoma species, e.g., G. cuneata.

Bactris hondurensis is one of the smallest Costa Rican Bactris species (but see also B. guineensis). Though sometimes described as having solitary stems, it is (like all our other Bactris species) cespitose by nature; however, the plants tend to have relatively few stems, and may be reduced temporarily to a single stem by mortality.

This is one of the most common, widespread, and frequently collected Bactris species in lowland forests of the Atl. slope; however, it is rare on the Pac. slope.

Bactris longiseta H. Wendl. ex Burret, Repert. Spec. Nov. Regni Veg. 34: 213. 1934. Stems ca. 3--5.5 m tall and ca. 3--3.5 cm diam., ± densely spiny just below the nodes, with spines to ca. 5 cm long. Petioles beyond sheath (34.5--) 62--74 cm, sparsely spiny with spines to at least 3.7 cm long. Lf.-blades pinnately compound, ca. (1.2--) 1.5--1.8+ m long, the rachis "woolly-tomentose," at least potentially with spines to ca. 5 cm long, lflts. (15--) 18--21 per side, clustered and arranged in more than one plane, 21--72 X 3.7--11 cm (medial ones), narrowly elliptical to oblong-lanceolate, with larger marginal spines ca. 2.5--4 cm long. Peduncle at least 5--11 cm, curved and ± flattened, with bract 16--32 cm long, ± densely black-spiny. Infl. rachis ca. 4--5 cm; rachillae ca. (20--) 30--35 (--40?), "to 8--15 cm" long. Male fls. unknown. Female fls. ca. 4.5--5.5 mm long. Ripe frts. ca. 1.5--1.9 X 1.7--2.1 cm, subglobose, obscurely rostrate, "green to yellow to purple." Wet forests, 35--250+ m; Atl. slope Cord. Central (V. Barva). Fl. 5--6. ENDEMIC. [Sears et al. 145, CR.]

This is the only Costa Rican Bactris species that has clustered lflts. with prominent marginal spines. Several disparate sources have characterized B. longiseta as having purple or purplish frts. (all quoted data in the above description are from labels of Sears collections from EB La Selva). Although de Nevers et al. (1996) included Bactris polystachya in their concept of B. longiseta, their report of "purple-brown" frts. derived from a La Selva collection (Holdridge 5118, BH) of B. longiseta s. str. The color of the ripe frts. of this species needs verification; if it does indeed prove to be purple, then frt. color should be added to couplet 12, as an additional character distinguishing B. longiseta from the species that key out below it (all of which have bright orange to red frts.).

Bactris longiseta is apparently a very local endemic, best known from EB La Selva.

Bactris major Jacq., Select. stirp. amer. hist. 134. 1780--1781. [B. augustinea L. H. Bailey; B. balanoidea (Oerst.) H. Wendl.; Pyrenoglyphis balanoidea (Oerst.) H. Karst.; B. superior L. H. Bailey].--Stems 2--5 (--10) m tall and 2--6 cm diam., the internodes spiny. Petioles beyond sheath ca. 22--54 (--150?) cm, formidably spiny, the spines to ca. 11 cm, uniformly blackish. Lf.-blades pinnately compound, 0.8--1.8 m long, the rachis formidably spiny (as petioles), lflts. 25--48 per side, regularly spaced and arranged in a single plane, 25--66 X 1--3.5 cm (medial ones), linear, marginally spinulose or (rarely) spinose. Peduncle 15--40 cm, densely spinulose (at least distally), recurved, with bract 28--60 cm long, moderately to densely spiny. Infl. rachis 1--4 cm; rachillae (3--) 5--10 (--17), 14--23 cm long. Male fls. 3--8 mm long. Female fls. 4--8 mm long. Ripe frts. 3--4.5 X 1.7--3.5 cm, ± ellipsoid to obovoid, bluntly beaked, brown to reddish purple or purple-black, minutely spinulose. Mainly dry and moist forests, often near swamps or along creeks, 0--50 (--100) m; N Atl. slope (Llanura de los Guatusos), Pac. slope on Pen. Nicoya (Sámara, RNA Cabo Blanco, RNVS Curú, Cabuya, I. San Lucas) and S from Río Tempisque basin, RB Isla del Caño. Fl. 8--9, 11. S Mex. (Oax., Ver.) to N Col., Trin., NE Braz., Bol. [Grayum et al. 9074, INB, MO.]

Bactris major is well characterized by a number of distinctive features, including its large size, fierce armature of long, uniformly blackish spines, pinnately compound lf.-blades with the lflts. regularly spaced in a single plane, very large, purplish black frts., and (typically) Pac. coastal habitat. It is one of just three Costa Rican Bactris species definitely known to have purplish (as opposed to orange or red) frts. (see also B. guineensis and B. maraja, both with much smaller frts.). This and B. guineensis are the only Bactris species recorded from the drier, northern portion of the Pac. slope (though B. major also extends southward).

The fruits of Bactris major are edible, and have a pleasant, acidic taste.

All Cent. Amer. material of Bactris major is referrable to var. major (fide Andrew Hend., Palms Amaz. 203. 1995), which extends to Trin. and NE Braz.

Bactris maraja Mart., Hist. nat. palm. 93, t. 71, fig. 1. 1826. [B. divisicupula L. H. Bailey; B. fuscospina L. H. Bailey; B. monticola Barb. Rodr.].--Stems 3--7 (--10) m tall and 2--4 cm diam., the internodes spiny. Petioles beyond sheath 13--125 cm, moderately to densely spiny, the spines flattened, black at base and apex, yellowish medially. Lf.-blades pinnately compound, 0.8--1.4 m long, the rachis sparingly spiny (as petiole), lflts. 15--22 per side, clustered and arranged in at least two planes, 20--48 X 3--9 cm (medial ones), sigmoid, marginally spinulose. Peduncle ca. 14--18 cm, spinulose at least distally, curved, with bract 20--38 cm long, brown-velvety-tomentose and moderately spinose (as petioles). Infl. rachis 2--5 cm; rachillae 6--15, 6--15 cm long. Male fls. to ca. 3.5 mm long. Female fls. ca. 2.5--3 mm long. Ripe frts. 1.2--1.5 X 1.3--2 cm, obturbinate to turbinate, prominently rostrate, purple-black. Wet forests, 0--450 m; rare, Atl. slope S from vic. Siquirres, Pac. slope from Palmar Norte region to Pen. Osa. Fl. 10--11 (Pan.); Fr. 9 (CR). CR to Guianas, Amaz. Venez. to Bol. [Grayum et al. 8003, CR, MO.]

This is the only Costa Rican Bactris species that has strongly flattened (as opposed to terete or subterete) spines. It might therefore be compared with our Astrocaryum species, also characterized by flattened spines; however, those all comprise much larger plants with stouter, solitary stems, lflts. abaxially whitened, and fertile organs much larger in every respect. Bactris maraja is additionally distinctive, among our Bactris species, in its broadly sigmoid lflts. and prominently beaked, purple-black frts. Herbarium specimens have characteristically reddish brown foliage, as B. gracilior.

Though widespread in the neotropics, Bactris maraja is quite rare in Costa Rica, where it is known by just a handful of collections from the southern portion of the country on both slopes. Our material corresponds to var. maraja (fide Andrew Hend., Palms Amaz. 207. 1995).

Bactris militaris H. E. Moore, Gentes Herb. 8: 229, figs. 88, 94--95. 1951. [B. neomilitaris de Nevers & Andrew Hend.].--Stems ca. 0.5--5 m tall and 2.5--4 cm diam., the internodes spiny (at least distally). Petioles beyond sheath ca. 27--70 cm, unarmed to formidably spiny, the spines to at least 6.5 cm, uniformly blackish; sheath dark and varnished within. Lf.-blades simple and bifid, ca. 1.25--3.1 m X 21--30 cm, the rachis unarmed or remotely spiny (as petioles), marginally spiny. Peduncle 15--38.5 cm, straight and erect (in fruit), with bract 20--49 cm long, moderately spiny. Infl. rachis ca. 2--10 cm; rachillae 7--23, 1.5--6 cm long. Male fls. ca. 4 mm long. Female fls. ca. 3 mm long. Ripe frts. 1.1--1.6 X 1.0--1.8 cm, ± turbinate, bluntly rostrate, red or orange. Wet forests, 0--50 (--460) m; Atl. slope Cords. Central (to RNFS Barra del Colorado) and Talamanca (to vic. Puerto Limón), Pac. slope at head of Golfo Dulce. Fl. 4--7. CR (and probably extr. SE Nic.) to E cent. Pan. [Grayum et al. 4476, CR, MO.]

Bactris militaris is easily distinguished by its long and relatively extremely narrow, consistently simple and bifid lf.-blades. As implied by the epithet, the plants may be formidably spiny; however, as in many Bactris spp., spininess is a variable feature in B. militaris, and both armed and unarmed lvs. may occur even on the same plant. The only other Costa Rican Bactris species with which B. militaris could possibly be confused is B. grayumii (see key, couplet 8); these two species are sometimes found growing together, as at RVS Barra del Colorado.

Although it is most characteristically found in swamp forests near sea level (as at the type locality), Bactris militaris has also been collected on ridges at somewhat higher elevations, as in PN Braulio Carrillo.

Bactris militaris is here circumscribed to include the recently segregated B. neomilitaris, described as "very similar" even by its own authors. The new taxon is supposed to differ in having shorter lf.-blades (to just 1.5 m) and frs. with non-pitted endocarps and numerous fibers with attached juice-sacs (as opposed to pitted endocarps and few fibers without juice-sacs). According to this notion, B. militaris is restricted to the Golfo Dulce region. Although B. neomilitaris may yet prove to correspond to a real taxonomic entity (perhaps better ranked as a subspecies), the issue cannot be resolved on the basis of the very few available collections.

Bactris polystachya H. Wendl. ex Grayum, in press. [B. polystachya H. Wendl. ex Hemsl., nom. nud.].--Stems 3--6 m tall and ca. 2--2.5 cm diam., the internodes spiny. Petioles beyond sheath 14--49 cm, unarmed or sparsely spiny. Lf.-blades pinnately compound, to ca. 1.35 m long, the rachis densely brownish scurfy, unarmed or remotely spiny, lflts. 15--31 per side, regularly spaced, arranged in a single plane, 30--83 X 3.5--7 cm (medial ones), linear to narrowly elliptical, usually with larger marginal spines ca. 0.4--2.5 cm long. Peduncle ca. 10 cm, strongly recurved, with bract 23--37 cm long, densely spiny. Infl. rachis 1.5--5 cm; rachillae 20--30, 4.5--11 cm long. Male fls. ca. 4--5 mm long. Female fls. ca. 2.5--3 mm long. Ripe frts. ca. 1.5--1.8 X 1.5--1.8 cm, oblate to obovoid, bluntly rostrate, orange. Wet forests, 0--150 m; Atl. slope Cords. Guanacaste and Central (to Boca Tapada, RNFS Barra del Colorado). Fl. ?; Fr. 11. ENDEMIC. [Grayum et al. 9044, INB, MO.]

Bactris polystachya may be said to combine features of B. caudata (congested infls. and infrs., orange frts.) and B. major (regularly spaced lflts. arranged in the same plane). The ± long (0.4--2.5 cm) marginal spines on the lflts. are unusual among Costa Rican pinnate-lvd. Bactris, shared only with B. longiseta (with clustered lflts. arranged in different planes) and some specimens of B. major (see key, couplet 11).

According to the label of Poveda 1017 (CR, USJ), the palmito of this species is "dulce" and "muy agradable."

Calyptrogyne

Stems obsolete to erect, slender, solitary, unarmed. Petioles beyond sheath ± short, unarmed, the sheath splitting. Lf.-blades simple and bifid to pinnately compound, in which case the lflts. usually unequal, arranged in a single plane. Plants monoecious, the infls. bisexual, interfoliar. Prophyll ± thin, striate, pointed. Peduncle elongate, erect, with 1 bract similar to prophyll, but inserted distally (just below spike), caducous (in ours) and leaving a conspicuous annular scar. Infls. spicate (ours). Fls. unisexual, borne in bracteate triads in bilabiate pits. Male fls. with 3 distinct sepals and 3 petals connate in basal half; stamens 6, the filaments mostly connate and adnate to receptacle. Female fls. with 3 distinct sepals, the petals connate in a shallowly 3-lobed tube which is ultimately circumscissile; staminodia connate in a minutely 6-lobed, fleshy tube; pistil 1, 3-locular; style elongate, triangular in section; stigmas 3, linear, reflexed. Ripe frts. smooth, ellipsoidal to obovoid, black, with basal stigmatic residue. 8 spp., S Mex. (Chis.) to Pac. Col.; Calyptronoma [3 spp., Gr. Ant. (Jam., Cuba, Hisp.)] is probably congeneric.

The Costa Rican species of Calyptrogyne are characterized by their subacaulescent habit, typically pinnately compound lf.-blades with the lflts. of various widths, and long-pedunculate, spicate infls. These features are shared with certain species of Geonoma, e.g., the widespread and variable G. cuneata; however, Calyptrogyne is sharply distinct from these in having the peduncular bract attached high on the peduncle, just below the rachilla, where it leaves a conspicuous, annular scar upon falling.

Calyptrogyne is one of the geonomoid palms, characterized by having their floral triads deeply immersed in rachillar pits (see under Geonoma for further information on this group). It is unique in this group (along with Calytronoma) in the calyptrate corolla and fleshy staminodial tube of the female fls.

de Nevers, G. 1995. Notes on Panama palms. Proc. Calif. Acad. Sci. 48: 329--342.

Wessels Boer, J. G. 1968. The geonomoid palms. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 58(1): 1--202+.

1 Infl. stipe and rachilla glabrous, or virtually so; both slopes...C. ghiesbreghtiana

1' Infl. stipe and rachilla densely reddish-brown-tomentose; Atl. slope only

2 Petioles beyond sheath 25--65 cm; infl. stipe 3.8--14 cm long; male fls. with sepals densely tomentose in distal half; Cords. Guanacaste, Tilarán, and Central...C. trichostachys

2' Petioles beyond sheath (7?--) 11.5--38 cm; infl. stipe 0.9--1.6 (--2.7?) cm long; male fls. with sepals virtually glabrous or marginally ciliate; Cord. Talamanca

3 Lflts. 3--8 (--21?) per side; infl. rachilla 9.3--11.8 (--13.5?) per side; 0--200 (--750+?) m...C. condensata

3' Lflts. 10--14 per side; infl. rachilla ca. 22--24 cm; 900--1400 m...C. herrerae

Calyptrogyne condensata (L. H. Bailey) Wess. Boer, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 58(1): 69. 1968. [Geonoma condensata L. H. Bailey, Gentes Herb. 6: 209, fig. 106. 1943].--Plants acaulescent to subacaulescent [with stems to at least 1 m]. Petioles beyond sheath ca. [7--21] 30--33 cm. Lf.-blades pinnately compound, the rachis ca. [50--58] 68--114 cm, lflts. 3--8 [18--21] per side, ca. [12--22.5] 38--71 cm long (medial ones). Peduncle (to scar) ca. [58--127] 93--142 cm, with bract ca. 14--17 cm long. Infl. rachilla 9.3--11.8 [12.5--13.5] cm; stipe 1.0--1.6 [1.8--2.7] cm long. Male fls. ca. 4.0 mm long, the sepals glabrous or virtually so; anthers ca. 1.0--1.5 mm. Female fls. ca. 3.5--4 mm long. Ripe frts. [1.0--1.4 X 0.5--0.7 cm, oblong-ellipsoidal], color unknown. Wet and moist forests, 0--200 [--750+] m; Atl. slope S from vic. Puerto Viejo de Talamanca. Fl. 1, 4, 9, 11 [1, 12]. CR and W Pan. [Grayum et al. 8001, CR, MO.]

This rare species is known from just a handful of collections, mainly from the Sixaola region of Costa Rica and adjacent Panama. See the key (couplet 3) for differences from the similar, upland C. herrerae. Panamanian material of C. condensata (square-bracketed data above) differs somewhat from Costa Rican material.

Calyptrogyne ghiesbreghtiana (Linden & H. Wendl.) H. Wendl., Bot. Zeitung (Berlin) 17: 72. 1859. [Geonoma ghiesbreghtiana Linden & H. Wendl., in H. Wendl., Linnaea 28: 343. 1857; C. brachystachys H. Wendl. ex Burret; C. donnell-smithii (Dammer) Burret; C. glauca (Oerst.) H. Wendl.; C. sarapiquensis H. Wendl. ex Burret].--Plants acaulescent to caulescent, with stems to 1+ m tall and to at least 3 cm diam. Petioles beyond sheath 27--60 (--116) cm. Lf.-blades simple and bifid or (much more frequently) pinnately compound, the rachis 22--135 cm; compound lvs. with lflts. 2--11 per side, (23--) 29--73 cm long (medial ones). Peduncle (to scar) (57--) 95--182+ cm, with bract ca. (26--) 37--74 cm long. Infl. rachilla (11--) 20--39.2 cm; stipe 0.9--8.4 (--12?) cm long. Male fls. ca. 5.0 mm long, the sepals glabrous or virtually so; anthers 1.2--2.5 mm. Female fls. ca. 3.5--4.0 mm long. Ripe frts. 0.8--2.1 X 0.5--1.2 cm, ovoid or ellipsoid (often narrowly so) to obovoid, purple to black. Wet forests, 0--1550 m; entire Atl. slope, Pac. slope Cords. Guanacaste and Tilarán (> 900 m) and in Golfo Dulce region. Fl. 1--12. S Mex. (Chis.) to extreme W Pan. (La Fortuna region). [A. Chacón 702, CR, MO.]

Calyptrogyne ghiesbreghtiana is the most widespread and variable species in the genus. It is distinguished, in Costa Rica, by a negative character: the absence of pubescence on the infl. rachilla and stipe. Several morphologically and geographically ± discrete entities occur within Costa Rica. Most similar to the Chiapan type is material from ca. 0--200 m elevation from the Atl. slope of the Cord. Central to RNFS Barra del Colorado and PN Tortuguero, represented by the above-cited voucher. Two populations [Grayum 8498 (MO), Vargas 304 (INB); Morales et al. 1850 (INB, MO)] from ca. 400--700 m on the Atl. slope of Volcán Barva differ in having simple, bifid lf.-blades, while material from 0--900+ m elevation in the Golfo Dulce region [e.g., Cordero 151 (INB, MO)] has long-caudate lflt. apices and unusually large (1.7--2.1 X 0.8--1.0 cm, vs. 0.8--1.0 X 0.5--0.7+ cm), obovoid frts. Most distinctive are collections [e.g., Fuentes et al. 219 (INB, MO)] from the Atl. slope of Cords. Guanacaste, Tilarán, and Central at ca. 700--1550 m elevation, for which the name Calyptrogyne brachystachys is available; these differ from the aforementioned races in having much shorter lf.-rachises (22--51 cm, vs. 62--135 cm), infl. rachillae (11--20.5 cm, vs. 21--39.2 cm), and anthers (ca. 1.2 mm, vs. 2--2.5 mm). Three collections [Barringer 3039 (CR), Cowan 4598 (MO), Hammel et al. 17527 (CR, MO)] from the Atl. lowlands of Cord. Talamanca at ca. 200 m are ± inseparable from the otherwise upland "C. brachystachys" (though anthers are not available), and material from above 1000 m in the La Fortuna region of W Pan. seems to bridge the morphological gap between the last-mentioned entity and typical C. ghiesbreghtiana.

The infls. of Calyptrogyne ghiesbreghtiana are markedly protandrous. This, in conjunction with the separation in time of successive infls., guarantees that male and female anthesis never coincide on the same plant. Flowers of both sexes smell strongly of garlic. The fleshy, sweet staminodial tubes, along with the fleshy filament tubes of the male fls., are consumed by pollinating bats. This is the only well-documented case of bat-pollination in the Arecaceae.

Calyptrogyne herrerae Grayum, in press.--Plants subacaulescent. Petioles beyond sheath ca. 11.5--31 cm. Lf.-blades pinnately compound, the rachis 62--90 cm, lflts. 10--14 per side, 16--48 cm long (medial ones). Peduncle (to scar) 123--162 cm, with bract [???] cm long. Infl. rachilla ca. 22--24 cm; stipe ca. 0.9--2 cm long. Male fls. ca. 5.0 mm long, the sepals ± glabrous, or marginally ciliate; anthers ca. 2.0 mm. Female fls. ca. 2.5--3 mm long. Ripe frts. described as "lila, dulces" (Herrera 8836, CR), otherwise unknown. Wet forests, 900--1400 m; Atl. slope Cord. Talamanca. Fl. 8. ENDEMIC. [G. Herrera & Rojas A. 8492, CR.]

Calyptrogyne herrerae is known from just three collections, all from mid-elevations on the Atl. slope of Cord. Talamanca N of Fila Matama. It is most similar to C. condensata, of the adjacent lowlands, but has longer, relatively more slender spikes with less congested fls. (see key, couplet 3).

Calyptrogyne trichostachys Burret, Bot. Jahrb. Syst. 63: 135. 1930.--Plants acaulescent to briefly caulescent, with stems to at least 10 cm tall and 2.5 cm diam. Petioles beyond sheath 25--65 cm. Lf.-blades pinnately compound, the rachis 50--94 cm, lflts. 3--8 (--13) per side, 46--72 cm long (medial ones). Peduncle (to scar) 92--183 cm, with bract 28--44 (--66) cm long. Infl. rachilla 16--31 cm; stipe 3.8--14 cm long. Male fls. ca. 4.0--5.0 mm long, the sepals densely tomentose in distal half; anthers 1.5--2.0 mm. Female fls. ca. 4.0 mm long. Ripe frts. 0.8--1.1 X 0.5--0.7 cm, narrowly obovoid, black. Wet forests, 100--1150 m; both slopes Cord. Guanacaste, Atl. slope Cords. Tilarán, and Central. Fl. 3--4, 6, 9. ENDEMIC. [I. Chacón 2451, CR, MO.]

Calyptrogyne trichostachys is sharply distinct from the sympatric C. ghiesbreghtiana in its reddish tomentose infl. rachilla and stipe; in these respects it more closely resembles the allopatric C. herrerae and C. condensata (see key, couplet 2). All known collections of C. trichostachys are from the Cord. Guanacaste and the Atl. slope of Cord. Tilarán down to the Llanura de San Carlos, with the exception of a single collection (Chazdon 128, CR) from 460 m on the Atl. slope of Volcán Barva. Henderson et al. (1995) reported this species from several provinces of W Pan., but I have seen no bona fide material from outside Costa Rica.

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