www.mobot.org Research Home | Search | Contact | Site Map  
 
Research
W³TROPICOS
QUICK SEARCH

MO PROJECTS:
Africa
Asia/Pacific
Mesoamerica
North America
South America
Floras
General Taxonomy
Photo Essays
Training in Latin
  America

MO RESEARCH:
Wm. L. Brown Center
Bryology
GIS
Graduate Studies
Research Experiences
  for Undergraduates

Imaging Lab
Library
MBG Press
Publications
Climate Change
Catalog Fossil Plants
MO DATABASES:
W³MOST
Image Index
Rare Books
Angiosperm
  Phylogeny

Res Botanica
All Databases
INFORMATION:
The Unseen Garden
What's New?
People at MO
Visitor's Guide
Herbarium
Jobs & Fellowships
Symposium
Research Links
Site Map
Search

Projects

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

Geonoma

Stems obsolete to erect, ± slender, sometimes (G. edulis, G. interrupta) subarborescent, solitary or cespitose, unarmed. Petioles unarmed, the sheath splitting. Lf.-blades simple and bifid to pinnately compound, the rachis unarmed, the lflts. unequal to equal, arranged in a single plane. Plants monoecious, the infls. bisexual, inter- or infrafoliar. Prophyll short to long, beaked. Peduncle obsolete to elongate, with (0--) 1 (--2) bracts similar to prophyll. Infls. spicate to paniculately branched. Fls. unisexual, borne in spirally arranged to verticillate, bracteate triads in bilabiate pits. Male fls. with 3 distinct sepals and 3 petals connate for ca. 2/3 their length; stamens 6 (ours), the filaments connate in a basal tube. Female fls. with 3 basally connate sepals and petals connate for most of their length in a 3-lobed tube, the lobes chaffy, spreading; staminodia connate in a truncate to 6-toothed or -lobed tube; pistils 3 (but 2 ovaries vestigial), the styles connate, basal, elongate; stigmas 3, linear, recurved. Ripe frts. smooth, [globose or ellipsoid], usually deep purplish or black, with basal stigmatic residue. Ca. 60--70 spp., S Mex. (Oax., Ver.) to Guianas, Trin., S Braz., Parag., Bol., Haiti, Less. Ant.

Geonoma is the central genus in the group commonly known as "geonomoid" palms (tribe Geonomeae), characterized by having their floral triads deeply sunken in ± bilabiate rachillar pits. The other geonomoid genera occurring in Costa Rica are Asterogyne, Calyptrogyne, Pholidostachys, and Welfia, all represented by one or (Calyptrogyne) few spp. These genera are distinguished mainly by technical, reproductive characters. Geonoma is unique in the group in having psuedomonomerous ovaries with basal styles (as opposed to trilocular ovaries with apical styles). See under the other genera for additional distinctions.

This is one of the most important palm genera in Costa Rica, ubiquitous in primary forests from sea level to nearly 3000 m elevation. Our Geonoma spp. all comprise understory to subcanopy plants, rarely if ever exceeding 10 m in height. The lowland species are relatively straightforward (at least within Costa Rica), whereas the montane taxa are difficult and poorly understood (see especially G. edulis, G. hoffmanniana, G. talamancana).

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

1 Infls. spicate; lf.-blades simple and bifid to pinnately compound;

2 Stems cespitose; peduncle < 8 cm; infl. rachilla usually < 5 cm long; very rare, 1300--1900 m, cent. Pac. region... G. monospatha

2' Stems solitary; peduncle > 8 cm; infl. rachilla > 5 cm long; widespread

3 Stems to ca. 2 m tall (fertile plants never acaulescent); lf.-blades mottled below with reddish or purplish in life; infls. infrafoliar; peduncle < 15 cm; very rare, 300--900 m, Atl. slope V. Barva... G. epetiolata

3' Stems to ca. 1.5 m tall, usually < 1 m tall (fertile plants often acaulescent); lf.-blades concolorous; infls. interfoliar; peduncle > 15 cm; widespread

4 Peduncular bract absent; rare, 1200--1600 (--2000+) m, Atl. slope Cord. Talamanca...G. talamancana

4' Peduncular bract usually present and conspicuous, very rarely obsolete or absent; widespread

5 Peduncular bract attached near base of peduncle (< 3 cm above prophyll); 0--1200 (--1900) m, widespread

6 Lf.-blades ± membranous to chartaceous, simple and bifid to pinnately compound with (1--) 2--10 lflts. per side, these rarely contracted basally; lf.-rachis ca. 18--76 cm; peduncle ca. 23.5--88.5 cm, with bract subequal to prophyll or exceeding it by up to 50% of prophyll length; rachilla ca. 6--31 X (0.2--) 0.3--0.5 (--0.8) cm; 0--1200 (--1900) m, widespread... G. cuneata

6' Lf.-blades ± soft-coriaceous or subsucculent, pinnately compound, with 7--23 lflts. per side, these ± contracted basally; lf.-rachis ca. (63--) 70--94 cm; peduncle ca. 56--133+ cm, with bract exceeding prophyll by > 50% of prophyll length; rachilla ca. 15--52 X (0.3--) 0.5--1.0 cm; 50--700 m, N Atl. slope...G. procumbens

5' Peduncular bract attached higher on peduncle (> 3 cm above prophyll; very rarely lower); (850?--) 1000+ m, mainly Cord. Tilarán

7 Peduncular bract ca. (1.5--) 4.2--15.5 cm (rarely obsolete), attached ca. (0.8--) 3.3--11 (--16.5+) cm above prophyll and enclosed by it; infl. rachilla at anthesis ± densely pubescent with mostly branching or stellate hairs; female fls. with staminodial tube deeply crenately to subdigitately lobed... G. brenesii

7' Peduncular bract ca. (0.5--) 3--43 cm long, attached 6--18.5 (--34+) cm above prophyll and usually exserted (or even attached) beyond its orifice; infl. rachilla at anthesis subglabrous to moderately or densely pubescent with simple hairs; female fls. with staminodial tube subtruncate to crenulate... G. hoffmanniana

1' Infls. forked (rarely) or racemosely to paniculately branched; lf.-blades pinnately compound, very rarely (G. monospatha) simple and bifid

8 Infl. rachillae 2--3, ca. 2--4 (--8.5) cm long; enlarged peduncular bract lacking (but 2--5 scalelike bracts < 3 cm long present); lf.-rachis ca. 13--22+ cm; lflts. ca. 8--14.5 cm long; very rare, 1300--1900 m, cent. Pac. region... G. monospatha

8' Infl. rachillae > 3 or, if 2--3, > 6 cm long; enlarged peduncular bract present, ca. (0.5--) 3--43 cm long (additional, smaller bracts may also be present); lf.-rachis ca. 17--250 cm; lflts. ca. 14--86 cm long; widespread

9 Infl. rachillae with floral pits verticillate in triads throughout; 0--450 (--750) m, both slopes... G. deversa

9' Infl. rachillae with floral pits decussate to laxly or densely spiraled, not verticillate in triads (except sometimes in distal 1--2 cm); widespread

10 Infl. rachillae filiform, ca. 0.5 mm diam; peduncle ca. 2.1--4.6 cm; lf.-rachis ca. 23--32 cm; 0--300+ m, Golfo Dulce region... G. scoparia

10' Infl. rachillae thicker, ca. 1.5--8 mm diam; peduncle ca. 4--62 cm; lf.-rachis ca. 17--250 cm; widespread

11 Floral pits ± pilose or tomentose internally in upper part (this feature often obscure), with upper lip obsolete (sometimes represented by a line of cilia); stems solitary; 0--1100 m

12 Stems ca. 1.6--6+ m tall; lf.-rachis ca. 110--180 cm; lflts. ca. 12--52 per side; infl. rachis ca. 32--58+ cm; rachillae very numerous, (5--) 13--33 cm long, pubescent with hairs to ca. 0.15 mm, at least some forked to dendritic or stellate...G. interrupta

12' Stems ca. 0.1--2.5 m tall; lf.-rachis ca. 44--80 (--100) cm; lflts. ca. 3--7 (--22) per side; infl. rachis ca. 6.5--31.5 (--56.5) cm; rachillae ca. 10--30 (--37++), 5.5--17 (--24) cm long, pubescent with simple hairs ca. 0.1--0.5 mm... G. oxycarpa

11' Floral pits glabrous internally, with upper lip ± prominently developed; stems cespitose or, if solitary, (700?--) 1000--3000 m

13 Stems solitary; peduncle ca. 7--62 cm, with bract ca. (0.5--) 3--43 cm long, attached ca. 1--18.5 (--34+) cm above prophyll; infl. rachis (0--) 0.9--34.5 (--72) cm; (700?--) 1000--3000 m

14 Stems (1--) 2--10+ m tall; infls. infrafoliar; peduncle ca. 7--28 cm, with bract 7.7--28.5+ cm long, attached ca. 1--4.4 (--7.5) cm above prophyll; infl. rachillae (5--) 9--indefinite; lf.-rachis ca. (31--) 42--250 cm; lflts. (2--) 9--39 per side; (700?--) 1000--2500 m... G. edulis

14' Stems obsolete to ca. 4 (--7?) m tall; infls. interfoliar; peduncle ca. 21.5--62 cm, with bract ca. (0.5--) 3--43 cm long, attached ca. 6--18.5 (--34+) cm above prophyll; infl. rachillae (2--) 3--17; lf.-rachis 22--57+ cm; lflts. (1--) 2--6 (--10) per side; 1400--3000 m... G. hoffmanniana

13' Stems cespitose; peduncle ca. 4--15.5 cm, with bract ca. 4--15.5 cm long, attached < 2 cm above prophyll; infl. rachis ca. (0.7--) 1.5--18 (--20.5) cm; 0--1400 m

15 Lflts. 2--6 per side, 25--86 cm long; peduncular bract ca. 10--15.5 cm long; infl. rachillae ca. 4--8 mm diam, pubescent with at least some branched to stellate hairs; ripe frts. ca. 0.9--1.4 X 0.8--1.3 cm, ± echinulate-roughened...G. congesta

15' Lflts. 3--17 per side, 14--45 cm long; peduncular bract ca. 4--10 cm long; infl. rachillae ca. 1.5--6 mm diam, pubescent with simple hairs; ripe frts. ca. 0.6--0.8 X 0.5--0.8 cm, tessellate-striate

16 Petioles beyond sheath ca. 4.5--21 cm; lf.-rachis ca. 17--52 cm; peduncle 4--9 cm; infl. rachis (0.7--) 1.5--9.5 cm, the rachillae 2.5--16 cm long; (250--) 400--1400 m... G. ferruginea

16' Petioles beyond sheath ca. 38--75 cm; lf.-rachis 56.5--95 cm; peduncle 4--15.5 cm; infl. rachis 5--18 (--20.5) cm, the rachillae (9--) 14--41 cm; 0--500 m

G. longevaginata

Geonoma brenesii Grayum, ined.--Stems solitary, obsolete to ca. 0.5 m tall and to at least 3 cm diam. Petioles beyond sheath ca. 14--53 cm. Lf.-blades pinnately compound, the rachis 11--30.5 cm, lflts. 2--5 (--7) per side, ca. 19--37 cm long (medial ones). Infls. interfoliar, spicate; peduncle 16--39.5 (--56) cm, with narrow bract ca. (1.5--) 4.2--15.5 cm long (rarely obsolete), enclosed by prophyll and inserted ca. (0.8--) 3.3--11 (--16.5+) cm above it; rachilla 6--19 X 0.2--0.5 cm, ± densely pubescent at anthesis with whitish, mostly branched or stellate hairs with ± flattened arms (later glabrescent). Male fls. ca. 3.0 mm long. Female fls. ca. 3.0--4.0 mm long; staminodial tube deeply crenately to subdigitately lobed. Ripe frts. ca. 0.7--0.8 X 0.5--0.6 cm, striate, broadly ellipsoid to obovoid, black. Wet forests, (850?--) 1000--1600 m; Atl. slope and near CD, Cords. Tilarán and Central Fl. 1--7, 10. ENDEMIC. [de Nevers et al. 7789, CR, MO.]

This sp. and the sympatric Geonoma hoffmanniana (see key, couplet 7) are the only Costa Rican Geonoma spp. with spicate infls. found above ca. 1200 m elevation in Cords. Tilarán and Central. Geonoma brenesii closely resembles, and was formerly confused with, the widespread, lowland G. cuneata, from which it differs in its generally smaller lf.-blades, narrow peduncular bracts attached high (usually > 3 cm) above the prophyll and enclosed within it, densely stellate-pubescent infl. rachillae, and female fls. with deeply crenately to subdigitately lobed staminodial tubes.

Most of the collections of Geonoma brenesii are from the Cord. Tilarán, mainly from the Monteverde region.

Geonoma congesta H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 112. 1869. CAÑA DE DANTA--Stems cespitose, 1--5+ m tall and ca. 2--5 cm diam. Petioles beyond sheath ca. 2--35 cm. Lf.-blades pinnately compound, the rachis ca. 34--90 cm, lflts. mostly 2--6 per side, ca. 25--86 cm long (medial ones). Infls. infrafoliar, racemosely to subpaniculately branched; peduncle 4.5--10 (--15) cm, with bract ca. 10--15.5 cm long, enclosed by prophyll and inserted near it; rachis ca. 1.5--11 cm; rachillae 4--14, mostly 7--24.5 X 0.4--0.8 cm, pubescent with simple to forked or stellate hairs. Male fls. ca. 4.0--5.0 mm long. Female fls. ca. 5.0--6.0 mm long; staminodial tube sinuate to crenulate. Ripe frts. ca. 0.9--1.4 X 0.8--1.3 cm, tessellate to (usually) echinulate, broadly ellipsoid or obovoid to subglobose, black. Wet forests, 0--850 (--900?) m; entire Atl. slope, Pac. slope S from ZP Cerros de La Cangreja. Fl. 1, 4--6, 8--11. E Hond. (Gracias a Dios) to NW Col. [I INBio 63, INB, MO.]

Among Costa Rican Geonoma spp. with cespitose stems and branched infls., G. congesta is distinctive in its stout infl. rachillae and comparatively large, usually ± echinulate-roughened frts. The lf.-blades are ± irregularly pinnate, with some very broad segments.

This is a locally abundant, even dominant understory sp. in lowland primary forest (as at EB La Selva). Populations on the Pac. slope differ somewhat in tending to have more extensively branched infls. with longer rachillae.

Geonoma cuneata H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 104. 1869. [G. gracilis H. Wendl. ex Spruce; G. decurrens H. Wendl. ex Burret; G. obovata H. Wendl. ex Spruce].--Plants acaulescent to caulescent, with stems solitary, to ca. 1.5 m tall (often ± decumbent at base) and ca. 3 cm diam. Petioles beyond sheath (5--) 14--59 (--62) cm. Lf.-blades simple and bifid or pinnately compound with rachis ca. 18--76 cm, if simple ca. (26--) 34--92 X 17--36 cm, obovate to narrowly obdeltate or oblanceolate, incised distally ca. 1/5--2/5 the total length, with ca. 20--39 primary lateral veins per side, if pinnate with lflts. (1--) 2--10 per side, ca. 19--55 cm long (medial ones). Infls. interfoliar, spicate; peduncle 23.5--88.5 cm, with bract ca. 11--50.5 cm long, to ca. 50% longer than prophyll and inserted near it; rachilla ca. 6--31 X (0.2--) 0.3--0.5 (--0.8) cm, subglabrous to sparsely pubescent with simple and forked hairs. Male fls. ca. 3.0--3.5 mm long. Female fls. ca. 3.5 mm long; staminodial tube subtruncate. Ripe frts. ca. 0.6--0.9 X 0.5--0.6 cm, finely striate, ellipsoid to subglobose or (rarely) ovoid, purplish-black. Wet forests, 0--1200 (--1900) m; entire Atl. slope, Pac. slope Cords. Guanacaste and Talamanca (> 1100 m) and S from RB Carara. Fl. 1--2, 4--11. SE Nic. to Pac. Ec., W Venez. [G. Rivera 720, CR, MO.]

This is the most widespread and commonly collected Costa Rican Geonoma with spicate infls. It is especially similar to the local (but sympatric) G. procumbens (which see), but might also be confused with G. brenesii, G. hoffmanniana, or G. talamancana, of generally higher elevations. The three last-mentioned spp. differ from G. cuneata principally in having the peduncular bract absent (G. talamancana) or attached high above the prophyll--an important taxonomic character that may be difficult to evaluate on herbarium material. Because of its generally subacaulescent habit, Geonoma cuneata is often confused (especially in sterile conditions) with Asterogyne martiana, Calyptrogyne spp., or even some Chamaedorea spp. (e.g., C. deckeriana).

Some material of Geonoma cuneata has traditionally been segregated, for varying reasons, under the name G. gracilis, recently demoted to varietal status (see Henderson et al., 1995). In my opinion, the type of G. gracilis falls well within the range of variation of G. cuneata, and I find no basis for recognizing G. gracilis at any rank. Geonoma cuneata is indeed a variable sp., especially as regards leaf pinnation, with both simple and pinnate lf.-blades occurring in the same population, or even on the same plant. However, this is not an uncommon situation in Geonoma.

Henderson et al. 1803 (INB, MO), from 1750 m elevation on the S slope of Cerro de la Muerte, resembles Geonoma cuneata in overall aspect, but is described as having the "stem cespitose, with basal offshoots." This is a unique condition among acaulescent Costa Rican Geonoma species with spicate infls. Additionally, this specimen exhibits an unusual rachillar indumentum. It may represent an undescribed species.

Geonoma deversa (Poit.) Kunth, Enum. pl. 3: 231. 1841. [Gynestum deversum Poit., Mém. Mus. Hist. Nat. 9: 390, t. 18. 1822; Geonoma flaccida H. Wendl. ex Spruce; Geonoma longepetiolata Oerst.].--Stems cespitose, ca. 1.7--5 m tall and ca. 0.8--3 cm diam. Petioles beyond sheath 10--46+ cm. Lf.-blades pinnately compound, the rachis ca. 29--90 cm, lflts. 2--30 per side, 13--46 cm long (medial ones). Infls. infrafoliar, subpaniculately to paniculately branched; peduncle ca. 2.3--10 cm, with bract 3--9 cm long, enclosed by prophyll and inserted near it; rachis 5.5--22 cm; rachillae ca. 11--55+, ca. 6--33 X 0.1--0.2 cm, pubescent with simple to stellate hairs. Male fls. ca. 2.0--2.5 mm long. Female fls. ca. 2.5 mm long; staminodial tube crenate. Ripe frts. ca. 0.5--0.75 X 0.4--0.65 cm, tessellate, broadly ellipsoid to subglobose, purplish black. Wet forests, 0--450 (--750) m; Atl. slope Cords. Guanacaste and Central (to vic. Los Chiles and RNFS Barra del Colorado), rare southward (Cocles, Talamanca), Pac. slope V. de El General, Golfo Dulce region, Pen. Osa. Fl. 1, 3--7, 9--11. Bel. and Guat. to Guianas, Braz., Bol. [Grayum & Jacobs 5343, CR, MO.]

Geonoma deversa is the only Costa Rican Geonoma sp. in which the floral pits are regularly arranged in alternating, verticillate triads along the rachillae. It is further distinguished by its cespitose habit and infrafoliar, highly branched infls. with slender rachillae. The lf.-blades dry with a distinctive, rather metallic sheen adaxially, so that even sterile herbarium specimens are identifiable. In terms of leaf pinnation, this sp. is highly variable; plants with numerous (to 25+), narrow lflts. are especially frequent on the Pen. Osa, where they may occur in the same populations as very different-looking conspecifics with trijugate lf.-blades.

Although widespread, Geonoma deversa is spottily distributed in Costa Rica. It seems to be most abundant and most frequently collected on the Pen. Osa and vicinity.

Geonoma edulis H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 106. 1869. [G. polyneura Burret; G. seleri Burret]. SÚRTUBA--Stems solitary, (1--) 2--10+ m tall and 2--15+ cm diam. Petioles beyond sheath ca. (4--) 18--50 cm. Lf.-blades pinnately compound, the rachis (31--) 42--250 cm, lflts. (2--) 9--39 per side, ca. 17.5--80+ cm long (medial ones). Infls. infrafoliar, racemosely (rarely) or subpaniculately to (usually) paniculately branched; peduncle ca. 7--28 cm, with bract ca. 7.7--28.5+ cm long, enclosed by prophyll and inserted ca. 1--4.4 (--7.5) cm above it; rachis ca. (4--) 6--34.5 (--72) cm; rachillae ca. (5--) 9--indefinite, 6--38 X 0.2--0.5 cm, pubescent with mostly simple hairs. Male fls. ca. 3.0--4.0 mm long. Female fls. ca. 3.5--4.5 mm long; staminodial tube subtruncate to crenate. Ripe frts. ca. 0.6--1.1 X 0.5--0.8 cm, tessellate-striate, broadly ellipsoid to subglobose, blackish. Wet forests, ca. (700?--) 1000--2500 m; Atl. slope and near CD all major cords, Pac. slope Cords. Guanacaste and Talamanca. Fl. 1--3, 5--8, 10--11. S Mex. (Chis.) to W cent. Pan., and probably to Peru. [Grayum 10377, CR, INB, MO.]

The name Geonoma edulis is here applied to a highly variable assemblage of montane material characterized by solitary stems, infrafoliar infls. with the peduncular bract attached (usually) < 5 cm above the prophyll, and bilabiate, spirally arranged floral pits. Occasional specimens with the peduncular bract attached rather high [e.g., Grayum 7025, CR, MO] may be virtually indistinguishable from the sympatric G. hoffmanniana (see key, couplet 14), especially if the attachment of the infl. relative to the lvs. is unknown.

Collections from the Cords. Tilarán and (especially) Guanacaste [e.g., III INBio 6 (CR, MO)] here referred to Geonoma edulis agree with the Guatemalan type of G. polyneura in having smallish infls., with a short rachis and relatively few rachillae [the name Geonoma polyneura was synonymized by Wessels Boer (1968) under G. seleri, a name of equal priority but without an extant type]. Specimens with the smallest infls. are easily confused with the sympatric G. ferruginea, especially if the growth habit is unknown. Material from Cord. Talamanca and adjacent Prov. Chiriquí, Panama, looks like a different species, comprising much taller plants (the largest of the genus in Costa Rica) with larger lvs. and very much larger infls. (matching the type of G. edulis), with much more numerous and longer rachillae. I have seen similar collections from S Mex. to Venez. and Peru.

Henderson et al. (1995) included all the material here referred to Geonoma edulis under the much older name G. undata Klotzsch, typified by a Venezuelan collection. Indeed, our Talamancan plants bear a closer resemblance in overall aspect to topotype collections of G. undata than to the material from N Costa Rica described in the previous paragraph. However, as G. undata is characterized by glabrous infl. rachillae and all of our specimens have pubescent rachillae, I have tentatively retained the name G. edulis. Clearly, the entire complex is in need of taxonomic revision.

Despite the specific epithet, the palmito of this species has been described as "muy amargo" (Aguilar 1092, INB).

Geonoma epetiolata H. E. Moore, Gentes Herb. 12: 28, fig. 2. 1980.--Stems solitary, ca. 0.5--2 m tall and 1 cm diam. Petioles beyond sheath ca. 0--3 cm. Lf.-blades simple and bifid, ca. 40--63 X 9.5--17.5 cm, narrowly obdeltate to oblanceolate, incised distally ca. 1/5--1/4 the total length, with ca. 32--40 primary lateral veins per side. Infls. infrafoliar, racemosely to subpaniculately branched; peduncle ca. 9--14 cm, with bract ca. 7.5--8.5 cm long, enclosed by prophyll and inserted near it; rachilla ca. 15--40 X 0.3--0.5 cm, ± densely pubescent with long, curled, brownish hairs (but ultimately glabrescent). Male fls. ca. 2.5--3.0 mm long. Female fls. ca. 3.0--3.5 mm long; staminodial tube crenate. Ripe frts. ca. 0.6--0.8 X 0.5 cm, finely striate, ellipsoid to subglobose, black. Wet forests, 300--900 m; Atl. slope V. Barva. Fl. 2, 4, 8--9. CR, cent. Pan. [Grayum et al. 5021, CR, MO.]

The combination of solitary stems and infrafoliar, spicate infls. immediately distinguishes this rare sp. from all other Costa Rican congeners. As if that were not enough, living plants are conspicuous in having their lf.-blades handsomely mottled abaxially with purplish or reddish. Geonoma epetiolata is additionally distinctive in its obsolete petioles and relatively short peduncles.

Because of its uniquely patterned foliage, Geonoma epetiolata has received some attention as a horticultural subject.

Geonoma ferruginea H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 110. 1869. [G. microspadix H. Wendl. ex Spruce; G. microstachys H. Wendl. ex Burret; G. versiformis H. Wendl. ex Spruce].--Stems cespitose, 1--5 m tall and ca. 0.5--2 cm diam. Petioles beyond sheath ca. 4.5--21 cm. Lf.-blades pinnately compound, the rachis ca. 17--52 cm, lflts. 3--17 per side, ca. 14--33.5 cm long (medial ones). Infls. infrafoliar, racemosely to subpaniculately or (rarely) paniculately branched; peduncle ca. 4--9 cm, with bract ca. 4--10 cm long, enclosed by prophyll and inserted near it at base of peduncle; rachis ca. (0.7--) 1.5--9.5 cm; rachillae ca. 5--20 (--37), ca. 2.5--16 X 0.15--0.35 cm, pubescent with unbranched, whitish hairs. Male fls. ca. 2.5--3.0 mm long. Female fls. ca. 2.0--2.5 mm long; staminodial tube truncate. Ripe frts. ca. 0.6--0.8 X 0.5--0.65 cm, tessellate-striate, broadly ellipsoid to subglobose, black. Wet forests, (250--) 400--1400 m; entire Atl. slope, Pac. slope Cord. Guanacaste. Fl. 1--5, 7--9, 11--12. E Guat. to W Pan. [Grayum et al. 4986, INB, MO.]

Except for the very rare and local Geonoma monospatha (see key, couplet 8), G. ferruginea is the only Costa Rican Geonoma with cespitose stems that occurs above ca. 700 m elevation. It is most similar to the more lowland (but conceivably sympatric) G. longevaginata, larger in all of its parts (see key, couplet 16). These two taxa are well-delimited in Costa Rica, but not (as here circumscribed) in Panama, where two to three other, different spp. in this complex might be distinguished (or, alternatively, the entire assemblage could be lumped into a single, polytypic sp. with various subspp.).

The lf.-blades of Geonoma ferruginea are most commonly trijugate, but specimens with numerous, narrow lflts. also occur, and are especially frequent in the Cord. Guanacaste.

Geonoma hoffmanniana H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 106. 1869. [G. molinae Glassman].--Stems solitary, obsolete to ca. 4 (--7?) m tall and 2.5--5+ cm diam. Petioles beyond sheath ca. 9--66 cm. Lf.-blades pinnately compound, the rachis (17.5--) 22--57+ cm, lflts. (1--) 2--6 (--10) per side, ca. (24--) 28--62 cm long (medial ones). Infls. interfoliar (sometimes becoming infrafoliar), spicate or forked to racemosely, subpaniculately, or (rarely) paniculately branched; peduncle (17--) 21.5--62 cm, subglabrous to densely arachnoid- or floccose-pubescent (later glabrescent), with bract ca. (0.5--) 3--43 cm long, included or exserted from prophyll and inserted 6--18.5 (--34+) cm above it (sometimes with 1--2 smaller, distal bracts); rachis 0--30 cm; rachillae ca. 1--17+, (6.5--) 8.5--31 X 0.2--0.5 cm, subglabrous to moderately or densely pubescent with mainly whitish, simple, ± tortuose hairs (then ultimately glabrescent). Male fls. 3.5--5.5 mm long. Female fls. ca. 4.0--5.0 mm long; staminodial tube subtruncate to crenulate. Ripe frts. ca. 0.5--1.0 X 0.4--0.8 cm, tessellate-striate, broadly ellipsoid to ovoid or subglobose, black. Wet and oak forests, 1400--3000 m; Atl. slope and near CD, Cords. Tilarán, Central (V. Barva), and Talamanca. Fl. 1--12. Hond. (Cortés) to W Pan., and probably to Peru. [Grayum et al. 7124, CR, MO.]

Geonoma hoffmanniana is here defined on the basis of its solitary stems, interfoliar, usually branched infls. with the peduncular bract attached high (6--18.5+ cm) above the prophyll, and montane (1400--3000 m) habitat. Taken in conjunction, these features distinguish G. hoffmanniana (as here circumscribed) from all other Costa Rican congeners, excepting rare individuals of G. edulis (which see) that have the peduncular bract attached unusually far (to a maximum of ca. 7.5 cm) above the prophyll.

As the other common montane Costa Rican Geonoma species, G. edulis, Geonoma hoffmanniana is highly variable and taxonomically difficult. The present treatment must be regarded as tentative. Specimens from V. Barva and the W portion of Cord. Talamanca (PN Tapantí, C. de la Muerte), with a short infl. rachis (ca. 0.9--10.7 cm) and ca. 3--7 (rarely 2) rachillae, best match the type (which is from V. Barva). In the Cord. Tilarán, the plants are somewhat smaller and usually have spicate infls. (rarely with 2--3 rachillae), with the peduncular bract often much reduced and usually attached higher on the peduncle (sometimes well above the orifice of the prophyll). This material [e.g., Grayum & Sleeper 3854, CR, MO] closely approaches Geonoma jussieuana Mart., and is partly the basis for the attribution of that species to Costa Rica by Henderson et al. (1995). I have been unable to cleanly delimit the Costa Rican material from G. hoffmanniana, even though G. jussieuana, as defined by its Bolivian type, may well be a different species.

Many collections of Geonoma hoffmanniana from the E portion of Cord. Talamanca [e.g., G. Herrera 3427 (CR, MO)] and into Prov. Chiriquí, Panama, are more robust than "typical" plants, with a longer infl. rachis (ca. 12--30 cm) and more numerous (ca. 12--17+) rachillae. However, intermediates occur in northern Cent. Amer. (Hond., Nic.) and S Amer. This more robust Talamancan material may approach Geonoma edulis.

I have seen collections matching even the type of G. hoffmanniana from as far south as Peru, begging the question of whether an older, S. Amer. name may exist for this sp. Indeed, Henderson et al. (1995) have subsumed G. hoffmanniana and a long list of other names under Geonoma orbignyana Mart., typified by a Bolivian collection. However, I consider this judgment premature, and prefer to retain a name that applies unequivocally to Costa Rican plants.

Geonoma interrupta (Ruiz & Pav.) Mart., Hist. nat. palm. 8. 1823. [Martinezia interrupta Ruiz & Pav., Syst. veg. fl. peruv. chil. 296. 1798]. SÚRTUBA--Stems solitary, 1.5--6+ m tall and ca. 2--5 cm diam. Petioles beyond sheath 1.5--90 cm. Lf.-blades pinnately compound, the rachis ca. 110--180 cm, lflts. 12--52 per side, ca. (21--) 33--68 cm long (medial ones). Infls. infrafoliar, paniculately branched; peduncle ca. 12--30+ cm, with bract ca. 12--38+ cm long, enclosed by prophyll and inserted ca. 1.5--5 cm above it; rachis ca. 32--58+ cm; rachillae indefinite, (5--) 13--33 X 0.15--0.3 cm, pubescent with simple and forked, dendritic, or stellate hairs. Male fls. ca. 2.5--3.0 mm long. Female fls. ca. 2.0--2.5 mm long; staminodial tube subtruncate to crenulate. Ripe frts. ca. 0.45--0.65 X 0.3--0.55 cm, ± smooth to finely striate, broadly ellipsoid to subglobose or ovoid, black. Wet and moist forests, 0--850+ m; Atl. slope Cords. Tilarán and Central (to Llanuras de los Guatusos and Santa Clara), Pac. slope Cord. Guanacaste, Pen. Nicoya (Las Huacas), and S from Río Grande de Tárcoles. Fl. 1, 4--5, 7--8, 10--12. S Mex. (Chis.) to Peru. [Grayum et al. 7539, CR, MO.]

Geonoma interrupta is characterized by solitary stems, large, pinnately compound lf.-blades with 12 or more lflts. per side, infrafoliar, extensively branched infls., floral pits lacking an upper lip, and a lowland habitat. Only one other Costa Rican Geonoma sp., G. oxycarpa, has floral pits lacking an upper lip. These two spp. have sometimes been lumped (under G. interrupta; e.g., by Henderson et al., 1995), but seem adequately distinguished in our region (see under G. oxycarpa and key, couplet 12).

Plants of this sp. are potentially among the tallest in the genus, and are especially striking by virtue of their long, fully pinnate lf.-blades and large, infrafoliar infls. Among Costa Rican Geonoma spp., only the more upland G. seleri (some forms) could be confused with G. interrupta in terms of size and general aspect.

The somewhat bitter palmito of this sp. is sometimes roasted and eaten.

Geonoma longevaginata H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 109. 1869.--Stems cespitose, 1--6 m tall and 1.5--3 cm diam. Petioles beyond sheath ca. 38--75 cm. Lf.-blades pinnately compound, the rachis ca. 56.5--95 cm, lflts. 3--14 per side, ca. 18--45 cm long (medial ones). Infls. infrafoliar, racemosely to subpaniculately branched; peduncle ca. 4--15.5 cm, with bract ca. 6.5--7 cm long, enclosed by prophyll and inserted near it; rachis ca. 5--18 (--20.5) cm; rachillae ca. 5--18, ca. (9--) 14--41 X 0.15--0.6 cm, pubescent with simple, ± spreading hairs. Male fls. ca. 2.5--4.0 mm long. Female fls. ca. 2.5--3.5 mm long; staminodial tube subtruncate to crenulate. Ripe frts. ca. 0.6--0.8 X 0.5--0.8 cm, tessellate-striate, broadly ovoid or ellipsoid to subglobose, black. Wet forests, 0--500 m; Atl. slope Cords. Central (to RNFS Barra del Colorado, PN Tortuguero) and Talamanca, Pac. slope vic. Golfito and head of Golfo Dulce. Fl. 1--2, 4, 6--7, 9--10, 12. Extr. SE Nic. to E Pan. [Grayum et al. 9185, CR, MO.]

Geonoma longevaginata is distinguished by its cespitose habit and infrafoliar, branched infls. with the floral pits spirally disposed on the rachillae. It is most similar to G. ferruginea, of generally higher elevations (see key, couplet 16, and discussion under G. ferruginea), but is perhaps more likely to be confused with the sympatric G. congesta (which has thicker infl. rachillae and larger, echinulate-roughened frts; see key, couplet 15).

Like many Geonoma sp., G. longevaginata exhibits considerable variability in the pinnation of its lf.-blades. Most plants have many (to 12+ per side), narrow lflts, but plants with trijugate lf.-blades often occur in the same populations. The reverse is true of G. ferruginea (i.e., trijugate lf.-blades are more common, in most areas).

Though ± abundant at some sites (especially EB La Selva), Geonoma longevaginata is in general an uncommon and spottily distributed sp. It is known from the Pac. slope by just two collections.

Geonoma monospatha de Nevers, in de Nevers & Grayum, Principes 42: 98. 1998.--Stems cespitose, 1--2.5 m tall and ca. 0.5--1 cm diam. Petioles beyond sheath ca. 4.5--12.5 cm. Lf.-blades simple and bifid or pinnately compound (even on same plant) with rachis ca. 13--22+ cm, if simple [ca. 19--24 X 5--5.5 cm], "parallel-sided" [oblong?], incised distally ca. 1/6--1/4 the total length, with [ca. 17--32 primary lateral veins per side], if pinnate with lflts. 3--4 (--?) per side, ca. 8--14.5 cm long (medial ones). Infls. inter- or infrafoliar, spicate or (most commonly) forked, rarely racemosely branched; peduncle ca. 4.3--7 cm, without enlarged bract (but with 2--5 scalelike bracts); rachis 0--0.6 cm; rachilla(e) (1--) 2 (--3), ca. 2--4 (--8.5) X 0.2--0.4 cm, subglabrous to sparsely or densely puberulent with stellate hairs. Male fls. ca. 2.5--3.5 mm long. Female fls. ca. 2.8--3.2 mm long; staminodial tube ± crenulate. Ripe frts. ca. 0.5--0.6 X 0.5--0.55 cm, tessellate-striate, subglobose, black. Wet forests, 1300--1900 m; Pac. slope, C. Turrubares to vic. C. Caraigres, Tarrazú. Fl. 3, 9--11 (Pan.). CR to cent. W Pan. (C. Tute). [Cascante et al. 1219, CR.]

The combination of a cespitose growth habit and very few (1--3) infl. rachillae immediately set this rare species apart from all of its Costa Rican congeners. Geonoma monospatha is the only Costa Rican Geonoma sp. that lacks an enlarged peduncular bract, as well as the only one that commonly has exactly two infl. rachillae (but see also G. hoffmanniana). It also has the smallest lf.-blades, on average, of any of our spp., and the smallest infl. rachillae (2--4 cm long). In color and general aspect, herbarium specimens of Geonoma monospatha suggest G. ferruginea, with every part much reduced.

This sp. is known from Costa Rica by just three collections, but is apparently somewhat more common in W Pan.

Geonoma oxycarpa Mart., in A. D. Orb., Voy. Amér. mér. 7(3) Palmiers 30. 1843. [G. mexicana Liebm.; G. binervia Oerst.].--Stems solitary, 0.1--2.5 m tall and 2.5--4 cm diam. Petioles beyond sheath 45--86 cm. Lf.-blades pinnately compound, the rachis ca. ca. 44--80 (--100) cm, lflts. 3--7 (--22) per side, ca. 26--58 cm long (medial ones). Infls. inter- or infrafoliar, subpaniculately to paniculately branched; peduncle ca. 14--42 cm, with bract ca. 10 cm long, ± equal to or barely exserted from prophyll exserted and inserted near it or to ca. 5 cm above; rachis ca. 6.5--31.5 (--56.5) cm; rachillae 10--30 (--37++), 5.5--17 (--24) X 0.15--0.3 cm, pubescent with ± spreading, simple, whitish hairs ca. 0.1--0.5 mm. Male fls. ca. 2.0--2.5 mm long. Female fls. ca. 3.0--4.0 mm long; staminodial tube subtruncate. Ripe frts. ca. 0.5--0.6 X 0.35--0.5 cm, striate, ovoid to ellipsoid, black. Wet forests, ca. 50--1100 m; both slopes Cords. Guanacaste and Tilarán, Atl. slope Cords. Central (to Llanura de San Carlos and RNFS Barra del Colorado) and Talamanca (Fila Matama), Pac. slope Pen. Nicoya (Las Huacas) and S from ZP Cerros de La Cangreja. Fl. 1--2, 4--6, 9--11. S Mex. (Oax., Chis.) to NW Col., Haiti. [Grayum & Herrera 4845, CR, MO.]

Its internally pubescent floral pits lacking an upper lip distinguish Geonoma oxycarpa from all other Costa Rican Geonoma spp. except G. interrupta. Although these two spp. have sometimes been lumped (under G. interrupta; see, e.g., Henderson et al., 1995), they seem ± sharply distinct in our region, as best appreciated at sites where they co-occur (e.g., EB La Selva). Generally speaking, G. oxycarpa comprises smaller plants with less extensively divided lf.-blades and smaller infls. with fewer, shorter, and more coarsely pubescent rachillae (see key, couplet 12).

A few collections from the N Atl. slope here referred to G. oxcarpa [e.g., Hammel et al. 20503 (INB); Liesner et al. 15028 (MO); Zamora et al. 2487 (INB)] approach G. interrupta in some (but not all) of the following characters: lf.-rachis length, number of pinnae, infl.-rachis length, number of rachillae, and rachillar length (parenthetical maxima for these categories). This material matches G. oxycarpa in terms of rachillar pubescence, and corresponds with the type of G. binervia (collected along the Río San Juan), considered a synonym of G. oxycarpa by Wessels Boer (1968).

Geonoma oxycarpa is spottily distributed in Costa Rica, but may be locally abundant, as in the canyon of the Río Reventazón below CATIE, Turrialba.

The application to our material of the name Geonoma oxycarpa, based on a Haitian type, is perhaps questionable.

Geonoma procumbens H. Wendl. ex Spruce, J. Linn. Soc., Bot. 11: 105. 1869.--Stems solitary, obsolete to ca. 1.5 m tall (may be decumbent at base) and to 5--6 cm diam. Petioles beyond sheath ca. 35--68 cm. Lf.-blades pinnately compound, the rachis (63--) 70--94 cm, lflts. 7--23 per side, 18--47 cm long (medial ones). Infls. interfoliar, spicate; peduncle ca. 56--133+ cm, with bract ca. 24--56+ cm long, exserted far beyond prophyll and inserted near it; rachilla ca. 15--52 X (0.3--) 0.5--1.0 cm, erect to arching or drooping, subglabrous or sparsely and inconspicuously pubescent. Male fls. ca. 4.5--5.0 mm long. Female fls. ca. 4.0--4.5 mm long; staminodial tube subtruncate to crenulate. Ripe frts. ca. 0.7--1.0 X 0.4--0.55 cm, finely striate, ellipsoid, black. Wet forests, 50--700 m; N Atl. slope, Cord. Guanacaste (to Llanuras de los Guatusos and San Carlos) and S to Río Sarapiquí. Fl. 5--7. SE Nic. to Pac. Col. [Hammel 20326, INB.]

Geonoma procumbens differs from all other Costa Rican congeners with spicate infls. in its generally larger, thicker lf.-blades with more numerous (usually > 11 per side), basally contracted lflts., and also in having the peduncular bract at least 1.5 X (and usually twice) as long as the prophyll. This sp. has sometimes been lumped in G. cuneata (see, e.g., Henderson et al., 1995), but the two spp. are strikingly distinct in life, as best appreciated at sites where they co-occur (e.g., near Boca Tapada de San Carlos). The leaf texture of G. procumbens, variously described as "soft," "rubbery," "somewhat succulent," or "waxy," has provoked comparisons with cycads.

This is sp. is rare in Costa Rica, where it is restricted to the N Atl. slope, but is widespread and frequently collected in Panama (where some local forms may have simple lf.-blades). I have seen just two collections from the Cord. Central.

Geonoma scoparia Grayum & de Nevers, Principes 32: 111, figs. 9--10. 1988.--Stems solitary, ca. 1.5--3 m tall and 1 cm diam. Petioles beyond sheath ca. 14--52 cm. Lf.-blades pinnately compound, the rachis ca. 23--32 cm, lflts. 3 per side, ca. 21--32 cm long (medial ones). Infls. infrafoliar, paniculately branched; peduncle ca. 2.1--4.6 cm, with bract ca. 5.5--7.5 cm long, enclosed by prophyll and inserted near it; rachis ca. 7.5--13.3 cm, zigzagging; rachillae indefinite, ca. 7--14 X 0.05 cm, glabrous, tuberculate-roughened. Male fls. ca. 2.0 mm long. Female fls. ca. 1.5 mm long; staminodial tube crenate. Ripe frts. ca. 0.4--0.6 X 0.4--0.5 cm, tessellate to tessellate-striate, broadly ellipsoid to subglobose, black. Wet forests, 0--300+ m; S Pac. slope, about head of Golfo Dulce from vic. Golfito to Estero Guerra and Laguna Chocuaco (N Pen. Osa). Fl. 1--3, 6, 10--11. ENDEMIC. [Hammel et al. 19137, INB, MO.]

Geonoma scoparia is highly distinctive, and unique among Costa Rican Geonoma spp., in its abundantly branched infls. with extremely slender (ca. 0.5 mm) rachillae. It is additionally characterized by its solitary stems, consistently trijugate lf.-blades, very short peduncles, and restricted (Golfo Dulce region) distribution. The only sp. with which G. scoparia could possibly be confused is the sympatric G. deversa, which may also have trijugate lf.-blades, and has ± slender infl. rachillae. The latter sp. differs in its cespitose stems and somewhat thicker infl. rachillae with the floral pits disposed in verticillate triads.

Geonoma scoparia is a rare sp. even within its small geographic range, and should be regarded as highly endangered.

Geonoma talamancana Grayum, ined.--Stems solitary, obsolete to ca. 1.5 m tall and ca. 1--2 cm diam. Petioles beyond sheath 9.5--30 (--35) cm. Lf.-blades simple and bifid or pinnately compound with rachis 17--43 (--58) cm, if simple 44.5--81 X 9--23.5 cm, oblanceolate, incised distally ca. 2/5--3/5 the total length, with 19--24 (--40) primary lateral veins per side, if pinnate with lflts. (1--) 2--4 (--7) per side, 26--50 cm long (medial ones). Infls. interfoliar, spicate; peduncle 27.5--78.5 (--81.5) cm, with no enlarged bracts above prophyll; rachilla 9--26 X 0.25--0.5 cm, subglabrous or pubescent with short, appressed, inconspicuous, brownish hairs. Male fls. ca. 3.5--4.0 mm long. Female fls. ca. 3.5--4.0 mm long; staminodial tube subtruncate. Ripe frts. ca. 0.7--0.9 X 0.6--0.8 cm, obscurely tessellate-striate, broadly ellipsoid to ovoid, black. Wet forests, 1200--1600 (--2000+) m; Atl. slope Cord. Talamanca. Fl. 4, 7--8. ENDEMIC? [Grayum 11033, CR, INB, MO.]

Geonoma talamancana comprises subacaulescent plants with solitary stems and spicate, interfoliar infls. with no enlarged peduncular bracts. It is our only Geonoma, other than the very different G. monospatha, that consistently lacks peduncular bracts. In the field, the plants differ in aspect from the similar and widespread G. cuneata by virtue of their comparatively rigid lf.-blades with plicate venation. Geonoma talamancana is the only Geonoma species with spicate infls. generally found above 1200 m in the Costa Rican Cord. Talamanca (but see also G. cuneata and G. monospatha).

Specimens of Geonoma talamancana have sometimes been identified as G. jussieuana Mart. (including G. lehmannii Dammer ex Burret), typified by a Bolivian collection, though perhaps ranging northward to W Pan. or even CR (see discussion under G. hoffmanniana). Although superficially very similar, G. jussieuana differs from G. talamancana in having an elongate peduncular bract, attached high on the peduncle and well exserted from the prophyll.

Hyospathe

2 spp., CR to Guianas, Braz., Bol.

Skov, F. & H. Balslev. 1989. A revision of Hyospathe (Arecaceae). Nordic J. Bot. 9: 189--202.

Hyospathe elegans Mart., Hist. nat. palm. 1, t. 1--2. 1823. [Chamaedorea falcaria L. H. Bailey].--Stems erect, cespitose, ca. 2.5--5 (--7?) m tall and 1.5--5 cm diam., unarmed. Petioles beyond sheath ca. 15--38 cm, unarmed, the sheath closed and forming a brief, green crownshaft ca. 24--50 cm long. Lf.-blades pinnately compound (ours), the rachis unarmed, ca. 44--93 cm, the lflts. ca. 13--25+ per side, subequal to ± unequal, ca. 27--45+ X 0.9--4.5 (--13) cm (medial ones), narrowly to broadly lanceolate, regularly spaced and arranged in a single plane. Plants monoecious, the infls. bisexual, infrafoliar. Prophyll ca. 27--37+ cm long, chartaceous, bluntly pointed and becoming bifid, deciduous. Peduncle ca. 2.2--6.1 cm, with 1 (--2) terete, beaked, deciduous bract(s), the largest ca. 26--34 cm long. Infls. racemosely branched; rachis ca. 3.5--14 cm; rachillae ca. 15--36, ca. 11.5--29.5 cm, subglabrous. Fls. unisexual, bracteolate, spirally arranged in triads (basally on rachillae) and solitary or paired male fls. (distally), the male fls. usually pedicellate. Male fls. ca. 2.5--3.5 mm long, borne on pedicels ca. 0.5--2 mm, with sepals connate for 2/3 their length or more in a 3-lobed tube and 3 distinct petals; stamens 6 in 2 whorls of 3, the inner whorl with much longer filaments adnate to pistillode. Female fls. ca. 1.5--2 mm long, subsessile, with sepals connate in basal 2/3 in 3-lobed tube and 3 distinct petals; staminodia, ligulate, inconspicuous; pistil 1, 1-locular; style short, thick; stigmas 3, recurved. Ripe frts. ca. 1.2--1.6 X 0.4--0.6 cm, narrowly ovoid and ± falcate, smooth, black, with basal stigmatic residue. Wet forests, 400--1300 m; Atl. slope Cord. Talamanca, Pac. slope S Fila Costeña (Tinamaste, Fila Retinto, Las Cruces). Fl. 3, 5, 9. CR to Guianas, Braz., Bol. [Grayum et al. 8784, CR, MO.]

This rare sp. is recognized by its smallish size, cespitose growth habit, brief, greenish crownshaft, pinnately compound lf.-blades with ± numerous and narrow lflts., and infrafoliar, racemosely branched infls. with slender, spreading rachillae. Hyospathe belongs to the subtribe Euterpeinae (see under Euterpe for more details). Among the Costa Rican members of this group, H. elegans is the only sp. combining an understory habit with green crownshafts. It is perhaps more likely to be confused with the more distantly related but vegetatively very similar Chamaedorea costaricana, which has ligulate petioles, longer peduncles and smaller, subglobose frts. Compare also our Synechanthus spp. (much longer peduncles and orange to red ripe frts.).

Hyospathe elegans is locally abundant in primary forests on the grounds of JB Wilson near Las Cruces, but is unlikely to be encountered elsewhere in Costa Rica. It is known from the Atl. slope by just two collections.

The use here of the name Hyospathe elegans is in accord with the suspiciously inclusive concept of this sp. espoused by Skov & Balslev (1989), who cited no Costa Rican collections. Our material differs rather consistently from Panamanian material identified by them as H. elegans in having somewhat larger, more finely pinnate lf.-blades (often simple and bifid in Panama), much longer prophylls ca. as long as the peduncular bract (vs. ca. half as long), and more numerous infl. rachillae. The name Hyospathe lehmannii Burret, though based on a Colombian collection, was applied to Costa Rican material in P. H. Allen's (1956) The Rain Forests of Golfo Dulce; this name, which lacks an extant holotype, was treated as a nomen dubium by Skov & Balslev (1989).

Iriartea

1 sp., extr. SE Nic. to Venez., Braz., Bol.

Henderson, A. 1990. Arecaceae. Part I. Introduction and the Iriarteinae. Fl. Neotrop. Monogr. 53: 1--100.

Iriartea deltoidea Ruiz & Pav., Syst. veg. fl. peruv. chil. 298. 1798. [I. corneto (H. Karst.) H. Wendl.; I. gigantea H. Wendl. ex Burret]. MAQUENQUE, CHONTA NEGRA--Stems erect, solitary, ca. 8--30 m tall and 10--45 cm diam., unarmed, supported by a dense cone of black, spiny stilt-roots to 1+ m tall. Petioles beyond sheath ca. 50--100 cm, unarmed, the sheath closed and forming a distinct, greenish crownshaft ca. 0.5--1.3 m long. Lf.-blades pinnately compound, ca. 2--3.5 m long, the rachis unarmed, the lflts. ca. 12--25 per side, large, cuneate at base, praemorse distally, splitting longitudinally into linear segments arranged in various planes; terminal lflt. entire. Plants monoecious, the infls. bisexual, infrafoliar, decurved in bud. Prophyll ca. 8 cm long, early caducous. Peduncle ca. 15--25 cm, curved, with 8--15 spirally arranged, ± pubescent, deciduous bracts, the distal longest. Infls. subpaniculately branched; rachis ca. 18--20 cm; rachillae ca. 33--40, ca. 49--105+ cm, whitish at anthesis. Fls. unisexual, whitish, spirally arranged in ebracteate, ± sunken triads (most of rachillae) and solitary or paired male fls. (distally). Male fls. ca. 5.5--6 mm long, with 3 distinct, saccate, rounded sepals and 3 much longer, distinct petals; stamens ca. 10--17, with very short filaments. Female fls. ca. 4.5--5 mm long, with 3 distinct sepals and 3 distinct petals; staminodia 10--13, very small and toothlike; pistil 1, 3-locular; stigmas 3, low. Ripe frts. ca. 1.6--2.5 X 1.5--2.4 cm, subglobose, smooth, pale green, ± dehiscent, with apical stigmatic residue. Wet forests, 0--800 m; entire Atl. slope, Pac. slope vic. Palmar Norte (Fila Retinto) to Pen. Osa. Fl. 1--5, 11--12. [Stevens & Montiel 24495, CR?, MO.]

Iriartea deltoidea is a widespread and familiar canopy palm, characterized by its dense cone of blackish, spiny stilt-roots and pinnately compound lf.-blades with "fish-tail" shaped, praemorse lflts. disposed in various planes. These features distinguish it from all other Costa Rican palms except for the related and vegetatively very similar Socratea exorrhiza. Close at hand, adult specimens of Iriartea may be told from S. exorrhiza by their darker, more numerous, and more densely aggregated stilt roots; from a distance, the conspicuously recurved infl. buds of Iriartea contrast markedly with the straight, erect buds of S. exorrhiza. Juvenile plants (and even seedlings) of Iriartea may be readily identified by having lf.-blades with the terminal (or sole) lflt. entire, rather than bifid as in S. exorrhiza (this difference also prevails in adult plants, but the terminal lflt. may be difficult to see in the canopy, or may become torn by the wind). See the genus key (couplet 6) for additional distinctions.

Iriartea deltoidea is highly esteemed in Costa Rica for its sweet cabbage, or palmito, which is palatable even raw. The leaves are used for thatching. However, the wood of this sp. is extremely hard, and, possibly because of this, the plants are often left standing even in pastures.

Manicaria

1 sp., Guat. and Bel. to Guianas, Trin., Braz., Peru.

Burret, M. 1928. Die Palmengattung Manicaria Gaertn. Notizbl. Bot. Gart. Berlin-Dahlem 10: 389--394.

Manicaria saccifera Gaertn., Fruct. sem. pl. 2: 469, t. 176. 1791. PALMA REAL--Stems erect, probably solitary (though often appearing cespitose), sometimes apparently bifurcate, ca. 1.5--8 m tall and 15--30 cm diam., unarmed. Petioles beyond sheath ca. 1--2 m, unarmed, the sheath with fibrous margins, splitting. Lf.-blades variously pinnately divided, ca. 3--4 m long, the rachis unarmed, the lflts. ± unequal (ca. 1--18+ cm wide), with narrow, apically bifid segments and some broader, serrate segments, arranged in a single plane. Plants monoecious, the infls. bisexual, interfoliar. Prophyll to at least 1.1 m long, netlike, tapering to solid, terete apex, completely covering the infl. Peduncle ca. 34--50++ cm, densely red-tomentose, with several bracts, the largest (basal) similar to prophyll and attached near midpoint. Infls. racemosely branched; rachis ca. 14--65 cm; rachillae ca. 16--35, ca. 14--34 cm, reddish lanate. Fls. unisexual, arranged in 1--3 bracteate triads (basally on rachillae) and more densely packed, ± sunken male fls. (distally). Male fls. ca. 5--6.5 mm long, with 3 basally connate sepals and 3 distinct petals; stamens 26--35. Female fls. ca. 8--13 mm long, with 3 distinct sepals and 3 distinct petals; staminodia ca. 10--15, linear, flat, thin; pistil 1, 3-locular; styles 3, deltate, connate; stigmas 3. Ripe frts. ca. 4--5 X 3.8--7.5 cm, subglobose to oblong (2-lobate) or depressed-triangular (3-lobate), densely covered with woody, broad-based, pyramidal tubercles to ca. 1 cm, brown, with subbasal stigmatic residue. Wet (swamp) forests, 0--100 m; N Atl. slope (RNFS Barra del Colorado, PN Tortuguero), very rare on Pac. slope (Río Coto Colorado, near mouth). Fl. 5, 9. [Davidse & Herrera 31447, CR, MO.]

The brownish, woody, densely tuberculate frts. of Manicaria saccifera are unique among Costa Rican palms. The plants may otherwise be recognized by their colonial habit (often appearing cespitose), ± finely divided lf.-blades, with the lflts. unequal, closely spaced, and sharply bifid to serrate apically, and coastal, swamp-forest habitat. From a distance, this sp. bears a striking but superficial resemblance to the sympatric Astrocaryum alatum in terms of size, shape, and pinnation of the lf.-blades. Close at hand, the latter sp. differs in its more obviously solitary stems, fiercely armed, abaxially whitish lf.-blades, and frts. armed with slender, needle-like spines.

According to literature sources, the lf.-blades of Manicaria are said to be ± entire, at least when newly emerged. This does not appear to be the case in Costa Rican populations, as observed in the field. The frts. of M. saccifera contain 2--3 seeds, and it is not uncommon to find closely paired seedlings recently germinated from the same frt. (A. Cascante, pers. comm.). This circumstance may partly explain the pseudo-cespitose growth habit of these palms, as well as occasional individuals with apparently bifurcate stems.

Manicaria saccifera is a common palm in the low-lying forests of PN Tortuguero, but is not likely to be encountered elsewhere in Costa Rica. The collection cited above is the first and (to date) only record from the Pac. coast of Cent. Amer. The lvs. are a locally important source of roofing thatch in the Tortuguero region, as also in Panama, where this sp. is widespread and much more commonly collected. There, the elastic, reticulate prophylls are used by Guaymí Indians to make fish-nets and "stocking caps" (Prov. Bocas del Toro), and the seeds are much sought after by forest mammals [Gordon 15d (MO), 20 (MO)].

Neonicholsonia

1 sp., E Hond. (Gracias a Dios) to W Pan. (Pen. Azuero).

Henderson, A. & G. Galeano. 1996. Euterpe, Prestoea, and Neonicholsonia (Palmae: Euterpeinae). Fl. Neotrop. Monogr. 72: 1--90.

Neonicholsonia watsonii Dammer, Gard. Chron. ser. 3, 30: 179. 1901. [N. georgei Dammer; Woodsonia scheryi L. H. Bailey].--Stems obsolete to decumbent or erect, solitary, ca. 0--0.5 (--1?) m tall and [????] cm diam., unarmed. Petioles beyond sheath ca. 16--108 cm, unarmed, the sheath splitting. Lf.-blades pinnately compound, the rachis ca. 61--155 cm, unarmed, the lflts. ca. 12--27 per side, subequal, ca. 17--52+ X 1.5--6.5 cm, narrowly elliptical to lanceolate or linear, regularly spaced and arranged in a single plane, long-caudate apically, glossy or satiny abaxially. Plants monoecious, the infls. bisexual, interfoliar. Prophyll ca. 5.5--45 cm long, splitting apically. Peduncle ca. (11--) 31--100 cm, slender, with 1 slender, terete, beaked bract ca. (36--) 67--153 cm long, at anthesis puplish externally and whitish within. Infls. spicate; rachilla ca. 20--61 cm, hirsutulous, cream-colored at anthesis, later becoming reddish. Fls. unisexual, spirally arranged in bracteate triads (basally on rachillae) and solitary male fls. (distally). Male fls. ca. 5--6.5 mm long, with 3 sepals basally connate in a cupule and 3 distinct petals; stamens 6, with alternate filaments of different lengths. Female fls. ca. 3--4 mm long, with 3 distinct sepals and 3 distinct petals; staminodia lacking; pistil 1, 1-locular; style not evident; stigmas 3, fleshy, slightly reflexed. Ripe frts. ca. 0.9--1.1 X 0.6--0.8 cm, broadly ellipsoid or obovoid to subglobose, smooth, black, with prominent apical stigmatic residue. Wet forests, 0--800 m; Atl. slope Cords. Guanacaste and Tilarán and very rare southward (Río Aguas Zarcas, Llanura de Santa Clara), Pac. slope Cord. Guanacaste and S from RB Carara. Fl. 1, 3, 6--10. [Henderson et al. 1807, INB.]

The combination of a subacaulescent habit, pinnately compound lf.-blades with 15 or more lflts. per side, and spicate infls. distinguishes Neonicholsonia watsonii from all other Costa Rican palms except the partly sympatric Geonoma procumbens. The latter sp. differs in having the floral triads deeply sunken in bilabiate rachillar pits, and frts. with a basal stigmatic residue. The prominent, apical stigmatic residue of N. watsonii is unusual in its own right, and suffices to distinguish this sp. from all other related or grossly similar Costa Rican palm spp. Because of its abaxially satiny or glossy lflts., N. watsonii bears a strong vegetative resemblance to Chamaedorea macrospadix, with which it may co-occur.

The overall impression of Neonicholsonia is that of an acaulescent Prestoea with unbranched infls. Indeed, the two genera are closely related, both belonging to subtribe Euterpeinae (see under Euterpe for further details).

Though inconspicuous and rather poorly known, Neonicholsonia watsonii is not a rare species; on the contrary, it is usually abundant in primary forests within its ± restricted geographic range.

Oenocarpus

9--10 spp., CR to Guianas, Trin., Braz., Bol.

Balick, M. J. 1986. Systematics and economic botany of the Oenocarpus--Jessenia (Palmae) complex. Advances Econ. Bot. 3: 1--140.

Oenocarpus mapora H. Karst., Linnaea 28: 274. 1857. [O. panamanus L. H. Bailey].--Stems erect, cespitose, ca. 4--18 m tall and 6--15 cm diam., unarmed. Petioles beyond sheath ca. 10--60 (--120?) cm, unarmed, the sheath closed and forming a ± bulging, purplish or blackish crownshaft ca. 25--100 cm long. Lf.-blades pinnately compound, ca. 2.5--4.5 m long, the rachis unarmed, the lflts. ca. 40--78 per side, subequal, ca. 22--130 X (0.6--) 1.2--6.5 cm, narrowly oblanceolate, regularly to ± irregularly spaced and arranged in one to several planes, ± pleated longitudinally, ± glaucous abaxially. Plants monoecious, the infls. bisexual, becoming infrafoliar. Prophyll ca. 28--33 cm long, woody, flattened, the margins coarsely toothed. Peduncle ca. 5--14 cm, with 1 terete, beaked, scaly bract ca. 65--75 cm long. Infls. racemosely branched; rachis ca. 7--22 cm; rachillae very numerous (to at least 130), ca. 36--75 cm, pendulous, ± strongly zigzag, glandular-puberulent to -furfuraceous. Fls. unisexual, arranged in bracteate triads (basally on rachillae) and solitary or paired male fls. (distally). Male fls. ca. 3--5 mm long, with 3 distinct sepals and 3 distinct petals; stamens 6, the filaments inflexed at apex. Female fls. ca. 4--5 mm long, with 3 distinct, suborbicular sepals and 3 distinct petals; staminodia lacking; pistil 1, 1-locular; style short, thick; stigmas 3, fleshy, reflexed. Ripe frts. ca. 1.8--2.5 X 1.3--2.0 cm, broadly ellipsoid or obovoid, smooth, deep purplish black, waxy, with apical to subapical stigmatic residue. Wet forests, 0--350 m; Pac. slope S from ZP Cerros de La Cangreja. Fl. 7, 10, 12. CR to Venez., Braz., Bol. [Grayum & Herrera 9184, CR, MO.]

The distinctive, "horse-tail" or "fly-swat" infls. of Oenocarpus mapora (i.e., with pendent rachillae borne on one side of the rachis) are unique among Costa Rican palms. This sp. is also the only cespitose, arborescent Costa Rican palm with a prominent crownshaft, with the exception of the vegetatively very similar Prestoea acuminata. The latter sp. differs in having concolorous lflts., divergent infl. rachillae borne all around the rachis, and much smaller, subglobose frts., and occurs at much higher elevations. Compare also Chamaedorea costaricana and Hyospathe elegans, both comprising generally smaller plants with green (rather than dark-colored) crownshafts.

Oenocarpus belongs to subtribe Euterpeinae, together with Prestoea; see under Euterpe for additional information.

This sp. was described as "frequent...throughout the [Palmar] area" by Allen (The Rain Forests of Golfo Dulce, 1956), but that seems no longer to be the case. Recent collectors have found this to be a rare or uncommon palm throughout the Costa Rican portion of its range.

Costa Rican material of Oenocarpus mapora belongs to the widespread subsp. mapora fide Balick (1986), but no infraspecific taxa were accepted by Henderson et al. (1995).

Pholidostachys

4 spp., Atl. Nic. [?] to Pac. Ec., Amaz. Col. to Peru.

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

Pholidostachys pulchra H. Wendl. ex Burret, Bot. Jahrb. Syst. 63: 130. 1930. [Calyptrogyne pulchra (H. Wendl. ex Burret) Wess. Boer].--Stems erect, solitary, ca. 2.5--5 m tall and 3--5 cm diam., unarmed. Petioles beyond sheath ca. 53--121 cm, slender, unarmed, the sheath densely reddish tomentose, splitting. Lf.-blades pinnately compound, the rachis ca. 54--65 cm, the lflts. ca. 7--10 per side, subequal, ca. 41--64 X 1.1--9.5 cm, lanceolate, regularly spaced and arranged in a single plane. Plants monoecious, the infls. bisexual, infrafoliar. Prophyll ca. 10--18 cm long, flattened, ± reddish tomentose. Peduncle ca. 8.5--23 cm, becoming pendulous, with several bracts, the largest (basal) ca. 28--37 cm long, short-beaked, thinly reddish tomentose, caducous. Infls. spicate; rachilla ca. 32--53 X 0.5--1.2 cm, densely reddish tomentose Fls. unisexual, borne in bracteate triads in bilabiate pits. Male fls. ca. 5--6 mm long, with 3 distinct sepals and 3 petals connate in basal half; stamens 6, the filaments connate in tube in basal 2/3. Female fls. ca. 5.5--6.5 mm long, with 3 distinct sepals and 3 petals connate for ca. 1/3--1/2 their length; staminodia 6--8, fleshy, connate in tube in basal 2/3; pistil 1, 3-locular; style elongate; stigmas 3, slender, spreading, exserted. Ripe frts. ca. 1.6--2.5 X 0.9--1.3 cm, ellipsoid to narrowly obovoid, substipitate basally, smooth, blackish, with basal stigmatic residue. Wet forests, 0--700+ m; Atl. slope Cord. Central (to RNFS Barra del Colorado and PN Tortuguero), Pac. slope Pen. Osa. Fl. 4, 6--7. CR [and Nic.?] to Pac. Col. [Grayum 6757, CR, MO.]

Pholidostachys is one of the "geonomoid" palms, characterized by having their floral triads deeply immersed in rachillar pits (see under Geonoma for additional information). It is immediately distinguished from virtually all Costa Rican members of this group by the combination of pinnately compound lf.-blades and infrafoliar, spicate infls. (but see the very rare Geonoma monospatha, of much higher elevations, which may occasionally exhibit this combination of characters). The relatively long petioles, short peduncles, and large frts. of P. pulchra are also distinctive.

Because of the thickish infl. rachillae, herbarium specimens of Pholidostachys pulchra are sometimes mistaken for Welfia regia, a much larger palm with many more lflts., branched infls., and larger frts. The frts. of P. pulchra, coarsely corded by thick mesocarp fibers, much resemble those of Calyptrogyne spp. The two genera are indeed narrowly related, and have sometimes been lumped, but Pholidostachys lacks the calyprate pistillate corollas that characterize Calyptrogyne.

Pholidostachys pulchra is an uncommon species in Costa Rica, best known from EB La Selva, where it occurs only in primary forest. It is very rare on the Pac. slope (just one collection has been seen). This species appears to be patchily distributed throughout its geographic range.

Prestoea

Stems decumbent to erect, obsolete to arborescent, solitary or (mostly, in ours) cespitose, unarmed. Petioles beyond sheath short (P. acuminata) to elongate, unarmed, the sheath splitting or (P. acuminata) closed and forming a greenish or purplish crownshaft. Lf.-blades pinnately compound (ours), the rachis unarmed, the lflts. subequal, long, narrow, regularly spaced and arranged in a single plane. Plants monoecious, the infls. bisexual, interfoliar (often becoming infrafoliar) or infrafoliar. Prophyll ± flattened and scaly. Peduncle short to long, with (usually) 1 terete, beaked bract, much longer than prophyll and inserted well above it. Infls. forked (rarely, in ours) or racemosely branched, the rachillae white becoming pinkish to reddish, glabrous to variously pubescent or tomentose. Fls. unisexual, arranged in bracteate triads (basally on rachillae) and solitary or paired male fls. (distally). Male fls. with 3 distinct sepals and 3 distinct petals, stamens 6, the filaments briefly inflexed at apex. Female fls. with 3 distinct sepals and 3 distinct petals; staminodia 6, small, toothlike; pistil 1, 1-locular; style not evident; stigmas 3, erect to reflexed. Ripe frts. ± smooth, subglobose, blackish, with lateral stigmatic residue. 11 spp., N Nic. to Guyana, Trin., Tobago, Braz., Bol., E Cuba, Hispan., PR, Less. Ant.

This is a ± nondescript genus of cespitose, understory to canopy palms, ours with pinnately compound lf.-blades with regularly spaced, narrow lflts. and racemosely branched inflorescences with long rachillae and smallish, subglobose, blackish frts. Prestoea belongs to subtribe Euterpeinae (see under Euterpe for further details); though perhaps closest to Neonicholsonia (with spicate infls.), it is most commonly confused with Euterpe. The last-mentioned genus is distinctive in possessing a conspicuous crownshaft, lacking in all Costa Rican Prestoea spp. except P. acuminata (which see).

The five Costa Rican Prestoea spp. are rather similar, distinguished mainly by overall size and by dimensions and pubescence of the peduncle and infl. axes. Shoddy herbarium specimens (e.g., with poor notes and only selected portions of the infl.) may be identifiable (if at all) only by the locality data.

Henderson, A. & G. Galeano. 1996. Euterpe, Prestoea, and Neonicholsonia (Palmae: Euterpeinae). Fl. Neotrop. Monogr. 72: 1--90.

1 Stems (3.5--) 5--12 m tall; petiole sheaths closed, forming a conspicuous, usually brownish to maroon crownshaft; peduncle ca. 8--15.5 (--18) cm; infl. rachis ca. 48--89+ cm, the rachillae ca. 43--125+; ripe frts. ca. 0.8--1.4 X 0.8--1.3 cm; (1000--) 1400--2450 m... P. acuminata

1' Stems 0.3--8+ m tall; petiole sheaths splitting, not forming a crownshaft; peduncle 18--101 cm; infl. rachis (0--) 0.9--53.5 cm, the rachillae (2--) 3--50; ripe frts. ca. 0.6--1.1 X 0.6--1.1 cm; 0--1800 (--2050) m

2 Lflts. ca. (25--) 30--50 per side; infl. rachis 22.5--53.5 cm, the rachillae ca. 27--50, 13.5--69 cm

3 Petioles beyond sheath ca. 60--150 cm; prophyll ca. 20.5--27.5 cm; peduncle 18--33.5 cm; infl. rachis 22.5--33 cm, the rachillae ca. 39--50, pubescent with whitish, stellate hairs ca. 0.1--0.3 mm; male (and often female) fls. with petals pilose or hirsute abaxially; 40--850 (--1000) m, Atl. slope... P. decurrens

3' Petioles beyond sheath ca. 44--70 cm; prophyll ca. (13--?) 24--46 cm; peduncle ca. (26--) 40.5--72+ cm; infl. rachis ca. 25--53.5 cm, the rachillae ca. 27--46, pubescent with simple hairs to ca. 0.3 mm and (sometimes) stubby, ± stellate hairs to ca. 0.1 mm; male (and female) fls. with petals glabrous abaxially; ca. 1050--1200 m, Atl. slope and Fila Costeña... P. ensiformis

2' Lflts. ca. (11--) 17--43 per side; infl. rachis (0--) 0.9--18.3 cm, the rachillae ca. (2--) 3--13 (--16), 11--47.5 cm

4 Stems ca. 0.3--1.5 (--3?) m tall; petioles beyond sheath ca. 74--200; lflts. (11--) 17--33 per side, 20--68 cm long; prophyll 10.5--32 (--39.5) cm long; peduncle (25+--) 31.5--101 cm; infl. rachillae (2--) 3--12 (--16), 11--36 cm, pubescent with reddish hairs; male fls. with petals glabrous abaxially; 800--1800 (--2050) m, both slopes...P. longepetiolata

4' Stems ca. 2.5--6 m tall; petioles beyond sheath ca. 25--75 cm; lflts. ca. 31--43 per side; 12--54 cm long; prophyll ca. 10.5--15.5 cm long; peduncle (21?--) 29.5--66 cm; infl. rachillae 7--13, 15.5--47.5 cm, pubescent with whitish hairs; male fls. with petals pilose toward apex abaxially; 0--400 (--800+) m, mainly on Pac. slope ...P. schultzeana

Prestoea acuminata (Willd.) H. E. Moore, Gentes Herb. 9: 286. 1963. [Oreodoxa acuminata Willd., Mém. Acad. Roy. Sci. Hist. (Berlin) 1804: 35. 1807; P. allenii H. E. Moore]. PALMITO MORADO--Stems cespitose, (3.5--) 5--12 m tall and 5.2--8.5 cm diam. Petioles beyond sheath 10--60 cm, the sheath closed and forming a green to (most frequently) brownish or maroon crownshaft ca. 40--100 cm long. Lf.-blades ca. 1.9--3+ m long, the lflts. ca. 33--53 per side, 17--95 X 1--8 cm (medial ones). Infls. infrafoliar, racemosely branched; prophyll ca. 28--50+ cm long; peduncle ca. 8--15.5 (--18) cm, with bract ca. 87--150 cm long; rachis ca. 48--89+ cm; rachillae ca. 43--125+, ca. 20--72++ cm, glabrescent to inconspicuously pubescent with stubby, scale-like, ± dendritic or substellate hairs to ca. 0.1 mm. Male fls. ca. 3.5--7.0 mm long, pinkish, the petals subglabrous or very minutely puberulent abaxially. Female fls. ca. 2.5--4.5 mm long, pinkish. Ripe frts. ca. 0.8--1.4 X 0.8--1.3 cm. Wet (oak) forests, (1000--) 1400--2450 m; Atl. slope all major cords., Pac. slope Fila Costeña (C. Anguciana). Fl. 4--6, 11--12. Nic. (I. Ometepe) to N. Venez. and Bol., Gr. Ant. (except Jam.), Less. Ant., Tobago. [Grayum 7338, CR, MO.]

This is the largest of the Costa Rican Prestoea spp., and the only one with closed petiole sheaths forming a crownshaft. It also has the largest infls., borne on the shortest peduncles (both relatively and absolutely) of any of our spp., as well as the largest frts., and has the highest altitudinal range. Prestoea acuminata is most similar to Euterpe precatoria (with solitary stems and green crownshafts) and, especially, Oenocarpus mapora (which see), both of which occur at lower elevations.

Prestoea acuminata, like Euterpe precatoria, is much sought-after for its tasty palmito. As a result, it has become scarce near areas of present or recent human habitation, e.g., PN Tapantí. Unlike Euterpe, however, plants of P. acuminata are cespitose, and thus less liable to be extirpated.

All Central American material of Prestoea acuminata was referred by Henderson & Galeano (1996) to var. acuminata, which accounts for the mainland portion of the species range.

Prestoea decurrens (H. Wendl. ex Burret) H. E. Moore, Gentes Herb. 9: 286. 1963. [Euterpe decurrens H. Wendl. ex Burret, Bot. Jahrb. Syst. 63: 63. 1929].--Stems cespitose, 3--8+ m tall and 6.5--12 cm diam. Petioles beyond sheath 60--150 cm, the sheath splitting. Lf.-blades ca. 1.5--3+ m long, the lflts. ca. (25--) 30--50 per side, 22--83 X 1.3--6 cm (medial ones). Infls. infrafoliar, racemosely branched; prophyll ca. 20.5--27.5 cm long; peduncle ca. 18--33.5 cm, with bract ca. 58+--130 cm long; rachis ca. 22.5--33 cm; rachillae ca. 39--50, ca. 27--61.5 cm, sparsely to densely pubescent with whitish, stellate hairs ca. 0.1--0.3 mm. Male fls. ca. 3.0--4.5 mm long, the petals pilose or hirsute abaxially, especially toward apex (but later glabrescent). Female fls. ca. 1.0--3.0 mm long. Ripe frts. ca. 0.6--0.9 X 0.6--0.9 cm. Wet forests, 40--850 (--1000) m; Atl. slope Cords. Guanacaste, Tilarán, and Central (to PN Tortuguero). Fl. 1--2, 4--7. NE Nic. to Pac. Ec. [Grayum & Jermy 6783, CR, MO.]

Prestoea decurrens is distinguished from the other Costa Rican Prestoea sp. with splitting petioles sheaths (not forming a crownshaft) by its relatively large size, numerous lflts., and large infls. on short (relatively and absolutely) peduncles. It is by far the most common Prestoea sp. below ca. 900 m elevation on the Atl. slope (but see also P. schultzeana). The coarsely stellate rachillar hairs and abaxially pilose or hirsute petals of the male fls. are shared only with P. ravenii (which see), mainly of the Pac. lowlands.

The palmito of Prestoea decurrens has been described as "sweet" and "edible" [Gómez et al. 19185 (MO)].

Prestoea ensiformis (Ruiz & Pav.) H. E. Moore, Gentes Herb. 9: 286. 1963. [Martinezia ensiformis Ruiz & Pav., Syst. veg. fl. peruv. chil. 297. 1798; P. sejuncta L. H. Bailey].--Stems cespitose, ca. 3.5--6 m tall and 4.8--13 cm diam. Petioles beyond sheath ca. 44--70 cm, the sheath splitting. Lf.-blades ca. 2--2.5 m long, the lflts. ca. 38--42 per side, 24.5--88 X 0.8--8 cm (medial ones). Infls infrafoliar (subtended by persistent petiole-sheaths), racemosely branched; prophyll ca. (13?--) 24--46 cm long; peduncle ca. (26--) 40.5--72+ cm, with bract ca. 76--143+ cm long; rachis ca. 25--53.5 cm; rachillae ca. 27--46, ca. 13.5--69 cm, sparsely villous with simple hairs to ca. 0.3 mm, sometimes also with much smaller, ± stellate hairs to ca. 0.1 mm. Male fls. ca. 3.0--5.0 mm long, the petals glabrous abaxially. Female fls. ca. 1.5--3.0 mm long. Ripe frts. ca. 0.7--1.1 X 0.7--1.1 cm. Wet forests, 1050--1200 m; Atl. slope Cord. Tilarán (RB Alberto Manuel Brenes), Pac. slope Fila Costeña (JB Wilson). Fl. 5--6. CR to Peru. [de Nevers et al. 7780, CR, MO.]

Prestoea ensiformis is the only Costa Rican Prestoea sp. to combine long peduncles (usually > ca. 30--35 cm) with long (> 20 cm) infl. rachises and numerous (> 15) rachillae; in our other species, peduncle length is ± inversely correlated with infl.-rachis length and rachilla number (i.e., short peduncles with long rachises and numerous rachillae in P. acuminata and P. decurrens; long peduncles with short rachises and few rachillae in P. longepetiolata and P. schultzeana).

This is apparently a very rare species in Costa Rica, where it is known by just three collections. It is has been collected frequently in Panama, especially on Cerro Jefe and vicinity. The above-cited collection, from San Ramón, is well-prepared and with detailed notes. It is an excellent match for the Panamanian material. Gillis & Plowman 10158 (MO), from Las Cruces, is a poor specimen assigned here tentatively; I have yet to see Moore & Parthasarathy 9443 (BH), a third specimen cited by Henderson & Galeano (1996), also from Las Cruces.

The name Prestoea ensiformis, based on a Peruvian type, is here applied to Costa Rican material following Henderson & Galeano (1996).

Prestoea longepetiolata (Oerst.) H. E. Moore, Gentes Herb. 9: 286. 1963. [Euterpe longepetiolata Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1858: 32. 1859; E. brachyspatha Burret].--Stems apparently solitary to (most frequently) cespitose, 0.3--1.5 (--3?) m tall and ca. 2--5 cm diam. Petioles beyond sheath 74--200 cm, the sheath splitting. Lf.-blades ca. 1.2--2.5 m long, the lflts. ca. (11--) 17--33 per side, 20--68 X 0.9--4.5 cm (medial ones). Infls. interfoliar, forked (rarely) or racemosely branched; prophyll ca. 10.5--32 (--39.5) cm long; peduncle ca. (25+--) 31.5--101 cm, with bract ca. 45+--101 (--148) cm long; rachis (0--) 0.9--18.3 cm; rachillae (2--) 3--12 (--16), ca. 11--36 cm, sparsely to densely reddish pubescent with simple to branched or stellate, ± curled hairs to ca. 0.2 mm. Male fls. ca. 3.0--6.0 mm long, reddish, the petals glabrous abaxially. Female fls. ca. 1.5--3.0 mm long. Ripe frts. ca. 0.7--1.1 X 0.7--1.1 cm. Wet forests, 800--1800 (--2050) m; Atl. slope and near CD, Cords. Tilarán, Central, and Talamanca, Pac. slope Cord. Talamanca, Fila Costeña. Fl. 1--9, 11--12. N Nic. to E cent. Pan., W Ven. [Grayum & Hammel 8456, CR, MO.]

Prestoea longepetiolata is characterized by its short or obsolescent stems, long petioles, comparatively few lflts. (the fewest of any Costa Rican Prestoea), and mid-elevation habitat. The short (< 20 cm) infl. rachises and few (< 15) rachillae are features shared only with P. schultzeana (see key, couplet 4), of lower elevations.

According to Henderson & Galeano (1996), most Costa Rican material of Prestoea longepetiolata belongs to var. longepetiolata, ranging from N Nic. to extr. W Pan. However, one specimen [Hammel 14961 (NY), from 1800 m in the Monteverde area], which I have not seen, was cited under var. roseospadix (L. H. Bailey) Andrew Hend. & Galeano (Fl. Neotrop. Monogr. 72: 67. 1996) [Euterpe roseospadix L. H. Bailey, Gentes Herb. 6: 201. 1943; Prestoea roseospadix (L. H. Bailey) H. E. Moore]. The latter variety, which ranges S to E cent. Pan., is said to differ in having narrower terminal lflts., a shorter peduncle (relative to the infl. rachis) and longer infl. rachis, and more numerous infl. rachillae (with the proximal ones swollen and sterile proximally).

Prestoea schultzeana (Burret) H. E. Moore, Gentes Herb. 12: 34. 1980. [Euterpe schultzeana Burret, Notizbl. Bot. Gart. Berlin-Dahlem 14: 326. 1939].--Stems cespitose, 2.5--6 m tall and ca. 3--3.5 cm diam, often decumbent at base. Petioles beyond sheath 25--75 cm, the sheath splitting. Lf.-blades ca. 1.3--1.6 m long, the lflts. ca. 31--43 per side, 12--54 X 0.8--5 cm (medial ones), ± prominently plicate. Infls. interfoliar (among old lvs.) or infrafoliar, racemosely branched; prophyll ca. 10.5--15.5 cm long; peduncle ca. (21?--) 29.5--66 cm, with bract ca. 56.5--95 cm long; rachis ca. 3.5--18 cm; rachillae ca. 7--13, ca. 15.5--47.5 cm, ± sparsely pubescent with whitish, stellate hairs to ca. 0.15 mm. Male fls. ca. 3.5--4.0 mm long, the petals pilose abaxially toward apex. Female fls. ca. 1.5--2.5 mm long, pinkish. Ripe frts. ca. 0.6--0.9 X 0.7--1.0 cm. Wet forests, 0--400 (--800+) m; N Atl. slope (RNFS Barra del Colorado, PN Tortuguero), Pac. slope S from RB Carara. Fl. 3, 5, 6, 8. CR (and ostensibly extr. SE Nic.) to Amaz. Peru. [Grayum & Herrera 9228, CR, MO.]

The only Prestoea occurring in the Pac. lowlands of Costa Rica, P. schultzeana differs from all of our other spp. except the more upland P. longepetiolata (see key, couplet 4) in having short (< 20 cm) infl. rachises and few (< 15) rachillae. It is perhaps most closely related to the Atl. slope P. decurrens, with which it shares whitish, stellate rachillar hairs and abaxially pilose petals of the male fls. Henderson & Galeano (1996: 61) referred Cent. Amer. material here treated as Prestoea schultzeana to P. decurrens, while noting that it differed in having "thicker, shorter rachillae" than typical P. decurrens, as well as "very closely spaced triads borne in slight depressions." However, they overlooked many other, more significant differences: somewhat smaller habit, shorter petioles, generally fewer and shorter lflts. with ± plicate venation, shorter prophylls, longer peduncles, much shorter infl. rachises, many fewer rachillae, etc. These features are mostly consistent with the concept of P. schultzeana.

Over its geographic range, Prestoea schultzeana exhibits marked variation in certain characters, e.g., peduncle length, rachillar pubescence, and fruit size. Although it is possible that more than one species or subspecies may be involved, I have been unable to delimit discrete taxa.

Raphia

Ca. 20 spp., Atl. Nic. to NW Col., Braz. (Pará), trop. Afr. and Madag.

Otedoh, M. O. 1982. A revision of the genus Raphia Beauv. (Palmae). J. Nigerian Inst. Oil Palm Res. 6: 145--189.

Raphia taedigera (Mart.) Mart., Hist. nat. palm. 216. 1838. [Sagus taedigera Mart., Hist. nat. palm. 54, t. 45, 48. 1824]. YOLILLO--Stems erect, apparently solitary to (most frequently) cespitose, ca. 3--12+ m tall and 25--60 cm diam., unarmed. Petioles beyond sheath ca. 1.5--5+ m, unarmed, deeply concave adaxially, the sheath splitting. Lf.-blades pinnately compound, ca. 5--10+ m long, the rachis armed (except with single row of short spines along adaxial keel), the lflts. ca. 95--205 per side, subequal, narrow (ca. 0.8--6 cm wide), ± regularly spaced, arranged in various planes, with ± prominent cross-veins adaxially, sometimes ± pruinose abaxially, ± inconspicuously spiny along margins (especially distally) and on midvein adaxially. Plants monoecious, the infls. bisexual, interfoliar. Prophyll tubular, 2-keeled. Peduncle "short" [??? cm], pendulous, with several (ca. 6) basally inflated bracts with triangular limbs. Infls. paniculately branched to 3 orders, ± elongate-cylindrical, all axes concealed by overlapping, truncate, sparsely scaly, ciliate-margined bracts; rachis ca. 1--3 (--5) m; rachillae (ultimate axes) ca. 3.5--11 cm. Fls. unisexual, bracteate and bracteolate, mostly solitary, usually distichously to tetrastichously arranged, the female fls. borne basally on the rachillae, the male fls. distally. Male fls. ca. 7--10 mm long, with sepals connate in a shallowly 3-lobed tube and 3 much longer petals connate in a tube in basal 1/4--1/3; stamens 8--11, the filaments distinct or basally connate in a tube. Female fls. ca. 7--9 mm long, with sepals connate in an irregularly 3-lobed tube and 3 petals connate in a tube in basal 1/2; staminodia connate in an epipetalous ring with ca. 6--9 irregular teeth of various lengths bearing short, flattened, sagittate, sterile anthers; pistil 1, 3-locular; style short; stigma 3-lobed, conical. Ripe frts. ca. 4.5--6.8 X 3.2--4 cm, narrowly obovoid to ellipsoid-oblong, hard and glossy, covered in 9--11 vertical rows of overlapping scales, reddish brown, with apical stigmatic residue. Wet (swamp) forests, 0--100 m; entire Atl. slope, Pac. slope at head of Golfo Dulce and on Pen. Osa. Fl. 6--8. Atl. Nic. to NW Col., Braz. (Pará), arguably in trop W Afr. [Henderson et al. 1815, INB.]

Raphia taedigera is characterized by its cespitose habit, extremely long, pinnately compound lf.-blades with the narrow pinnae arranged in various planes, elongate, pendulous infls., and lowland, swamp-forest habitat. The large, glossy, scaly frts., unique among all Central American palms, are diagnostic (when available). The plants are surrounded at the base by ± dense mats of branching pneumatophores, and the stems bear negatively geotropic adventitious roots among the dead petiole bases. The "trunks" consist mainly of petiole bases.

This species is remarkable for usually growing in extensive, virtually pure stands (yolillal) in low-lying, poorly drained sites. Many such stands have been razed for banana or oil-palm plantations.

Raphia taedigera is essentially hapaxanthic, in that each flowering stem produces several infls. over a relatively short period, then dies. It is probable that this species flowers throughout the year.

Recent paleoecological studies by Urquhart (J. Trop. Ecol. 14: 783--791. 1997) establish conclusively the indigenous status of R. taedigera in the New World, long a hotly debated issue.

Reinhardtia

Stems erect, very short to moderate, cespitose (or temporarily solitary), unarmed. Petioles brownish scaly, the portion beyond sheath ± elongate, sometimes ± winged marginally, the sheath tubular, becoming fibrous. Lf.-blades simple and bifid or pinnately compound, when compound the lflts. sometimes fenestrate basally, ± praemorse distally, regularly spaced and arranged in a single plane; terminal lflt. ± larger. Plants monoecious, the infls. bisexual, interfoliar. Prophyll membranous, with 2 triangular distal lobes, ± included in petiole sheath. Peduncle relatively long, with usually 1 elongate, papery, disintegrating bract. Infls. spicate, forked, or racemosely to paniculately branched. Fls. unisexual, white to yellowish, ± distichously arranged in bracteate triads (basally on rachillae) and solitary or paired male fls. (distally). Male fls. with 3 distinct sepals and 3 much longer, basally connate petals; stamens ca. 7--28, the filaments short, basally connate. Female fls. with 3 distinct sepals and 3 basally connate petals, spreading distally; staminodia connate in a ring ± adnate to the petals, with 2--9 usually erect, exserted teeth; pistil 1, 3-locular; style stout; stigmas 3, recurved. Ripe frts. smooth or striate, ovoid to obovoid, purplish black to black, with apical stigmatic residue. 6 spp., S Mex. (Oax., Ver.) to Pac. Col., Gr. Ant. (Rep. Domin.).

Our Reinhardtia species are characterized by their very small to moderate size, cespitose habit, interfoliar, long-pedunculate infls., male fls. with numerous (7--28) stamens, fruits with an apical stigmatic residue, and lowland (< 900 m) habitat. Reinhardtia gracilis and R. latisecta are the only Costa Rican palm species with fenestrate lf.-blades on the adult plants (but see also the very different Attalea butyracea). The combination of a cespitose habit and simple lf.-blades (or pinnately compound blades with just one pair of subterminal lflts.) distinguishes R. koschnyana and R. simplex from all other Costa Rican palms except some forms of the rare Geonoma monospatha, of higher elevations (see also Synechanthus warscewiczianus, which may very rarely have simple lf.-blades).

Palms of the genus Reinhardtia are generally uncommon and inconspicuous in Costa Rican forests, though R. latisecta and R. simplex may be locally ± abundant. All are prized and much sought as ornamental subjects.

Moore, H. E., Jr. 1957. Synopses of various genera of Arecoideae. 23. Reinhardtia. Gentes Herb. 8: 541--576.

1 Lf.-blades paripinnately compound, with (1--) 2--10 lflts. per side; lflts. (at least proximal and distal pairs) fenestrate at base

2 Stems 0.4--2+ m tall; petioles beyond sheath 9--32.5 cm; lf.-rachis < 20 cm; lflts. (1--) 2 (--3) per side; peduncle < 65 cm; infls. racemosely branched; rachis < 8 cm; rachillae < 15; male fls. with ca. 8--11+ stamens; ripe frts ca. 1.3--1.5 X 0.5--0.7 cm; Atl. slope all major cords., very rare on Pac. slope (Tarrazú)... R. gracilis

2' Stems ca. 1.8--6 m tall; petioles beyond sheath ca. 31--92.5 cm; lf.-rachis > 20 cm; lflts. 2--10 per side; peduncle 65+ cm; infls. subpaniculately to paniculately branched; rachis > 8 cm; rachillae > 15; male fls. with ca. 18--28 stamens; ripe frts. ca. 1.6--1.9 X 0.65--1.05 cm; local, N Atl. slope, S. Pac. slope (Pen. Osa)... R. latisecta

1' Lf.-blades simple and bifid, or imparipinnately compound (with the distal pair of lflts. almost completely fused); lflts. never fenestrate

3 Lf.-blades always simple, bifid at apex by ca. (3.3--) 3.9--6.5 (--9.3) cm; infls. spicate; male fls. with ca. 7--10 stamens; 50--250 m, N Atl. slope... R. koschnyana

3' Lf.-blades simple and bifid at apex by ca. 0.6--3.0 (--5.2) cm, or else imparipinnately compound with 0--1 subterminal lflts. per side; infls. forked to (usually) racemosely or (very rarely) subpaniculately branched, with 2--6 (--7) rachillae; male fls. with ca. 14--17+ stamens; 0--700 (--1100) m, widespread... R. simplex

Reinhardtia gracilis (H. Wendl.) Drude ex Dammer, Palmenz. Palmenpfl. 90. 1897. [Malortiea gracilis H. Wendl., Allg. Gartenzeitung 21: 26. 1853].--Stems 0.4--2+ m tall and 0.3--1 cm diam. Petioles beyond sheath 9--32.5 cm. Lf.-blades simple and bifid (very rarely) or pinnately compound with rachis ca. 4--9.8 cm, usually prolonged distally in a slender, free filament to at least 8.5 cm, if simple ca. 21 X 17 cm, broadly elliptical, incised distally ca. 2/3 the total length, with 13--14 primary lateral veins per side, the margins coarsely doubly serrate, if pinnate with lflts. (1--) 2 (--3) per side, (7.3--) 11--20.5 X 0.8--6.3 cm (subterminal ones), narrowly to broadly cuneate, fenestrate proximally, praemorse distally. Infls. racemosely branched; peduncle ca. 21--61 cm, with bract ca. 7--19.5 cm long; rachis ca. 0.2--5 cm; rachillae 3--7, ca. 4--14.5 cm. Male fls. ca. 4.5--5.0 mm long; stamens ca. 8--11+. Female fls. ca. 6.0 mm long. Ripe frts. ca. 1.3--1.5 X 0.5--0.7 cm, narrowly oblong-ovoid or ellipsoidal. Wet forests, 40--850 m; Atl. slope all major cords. (to PN Tortuguero), very rare on Pac. slope (Tarrazú region). Fl. 1--2, 5--12. S Mex. (Oax., Ver.) to extr. NW Col. [Haber et al. 11254, CR, MO.]

Reinhardtia gracilis is immediately recognized by its fenestrate lf.-blades and small size of all of its parts (as compared with R. latisecta, the only other Costa Rica palm with fenestrate blades). This is an exceptionally handsome species, much cultivated as an ornamental. Although widespread in Costa Rica, it is nowhere abundant.

Costa Rican material of Reinhardtia gracilis corresponds to var. rostrata (Burret) H. E. Moore (Principes 1: 145. 1957; R. rostrata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11: 554. 1932), which ranges from SE Nic. to Col.

Reinhardtia koschnyana (H. Wendl. & Dammer) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11: 554. 1932. [Malortiea koschnyana H. Wendl. & Dammer, in Dammer, Gard. Chron. ser. 3, 29: 341. 1901].--Stems 0.15--1.2 m tall and 0.3--0.7 cm diam. Petioles beyond sheath 4--14 cm. Lf.-blades simple and bifid, 14.5--22 X 4.4--8.3 cm, ± narrowly ellliptical or obovoid to oblanceolate, incised distally ca. 1/5--1/3 (--2/5) the total length, with (7--) 8--10 (--11) primary lateral veins per side, the margins very coarsely serrate or doubly serrate. Infls. spicate; peduncle ca. 11.5--35 cm, with bract ca. 10.5--18 cm long; rachilla ca. 3.8--9.8 cm. Male fls. ca. 3.0--4.5 mm long; stamens ca. 7--10. Female fls. ca. 3.0--4.0 mm long. Ripe frts. ca. 0.9--1.3 X 0.6--0.8 cm, ± broadly ellipsoidal to obovoid. Wet forests, 50--250 m; N Atl. slope (Llanuras de los Guatusos and San Carlos). Fl. 1, 3--5, 12 (Pan.). E Hond. (Gracias a Dios) to Pac. Col. [Holm & Iltis 913, CR, MO.]

Reinhardtia koschnyana, one of the most diminutive of palms, is further distinguished by its cespitose habit, simple lf.-blades, and spicate infls. This combination of characters is virtually unique among Costa Rican palm species, seen otherwise only in some individuals of Geonoma monospatha (comprising somewhat larger plants of much higher elevations). The differences emphasized in the key (couplet 3) appear to separate R. koschnyana consistently from the sympatric R. simplex, to which this species bears a striking vegetative resemblance.

A widespread and frequently collected species in Panama, Reinhardtia koschnyana is known from Costa Rica by just a few collections from the now largely deforested N Atl. plains. This is the only region in which all four of our Reinhardtia species occur (or occurred) sympatrically.

Reinhardtia latisecta (H. Wendl.) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11: 554. 1932. [Malortiea latisecta H. Wendl., Allg. Gartenzeitung 21: 146. 1853].--Stems ca. 1.8--6 m tall and 3.5 cm diam. Petioles beyond sheath ca. 31--92.5 cm. Lf.-blades pinnately compound, the rachis ca. 33--80+ cm; lflts. 2--10 per side, ca. 44--69 X 1--11.5 cm (subterminal ones), very narrowly to broadly cuneate, the broader (proximal and distal) pairs fenestrate proximally, praemorse distally. Infls. subpaniculately to paniculately branched; peduncle ca. 65--125+ cm, with bract ca. 43+ cm long; rachis ca. (11--) 16.5--36 cm; rachillae ca. 30--60+, ca. (2.5--) 6--26 cm. Male fls. ca. 4.5--5.5 mm long; stamens ca. 18--28. Female fls. ca. 4.0--4.5 mm long. Ripe frts. ca. 1.6--1.9 X 0.65--1.05 cm, obovoid. Wet forests, 50--700 m; Atl. slope Cord. Guanacaste (to Llanura de San Carlos), Pac. slope on Pen. Osa. Fl. 6. Bel. and Guat. to CR. [Henderson et al. 1814, INB.]

This species and its more familiar congener, Reinhardtia gracilis, are our only palms with fenestrate lf.-blades. Reinhardtia latisecta differs substantially from the last-mentioned species in its taller stems, much longer petioles and lf.-blades, and much larger, paniculately branched infls. (see key, couplet 2). Indeed, R. latisecta comprises by far the largest plants of any Costa Rican Reinhardtia species.

Reinhardtia latisecta is generally a rare species, absent from most of Costa Rica, though it may be locally abundant, e.g., at some sites in the Cord. Guanacaste.

Reinhardtia simplex (H. Wendl.) Drude ex Dammer, Palmenz. Palmenpfl. 91. 1897. [Malortiea simplex H. Wendl., Bot. Zeitung (Berlin) 17: 5. 1859].--Stems 0.3--1.5 m tall and 0.3--0.8 cm diam. Petioles beyond sheath 2.5--16 cm. Lf.-blades simple and bifid or imparipinnately compound with rachis 12--20.5 cm, if simple ca. 12--22.5 X 5--10 cm, narrowly to ± broadly oblong or ellliptical, incised distally ca. 1/20--3/20 (--1/4) the total length, with (8--) 9--12 primary lateral veins per side, the margins coarsely serrate or (usually) doubly serrate, if imparipinnate with subterminal lflts. 0--1 per side, 7.2--15 X 1.3--3.4 cm, narrowly to broadly cuneate, praemorse distally. Infls. forked to (usually) racemosely or (very rarely) subpaniculately branched; peduncle ca. 10.5--42+ cm, with bract ca. 10.5--15 cm long; rachis 0--3.6 cm; rachillae 2--6 (--7), ca. 2.5--8.5 cm. Male fls. ca. 2.5--4.0 mm long; stamens ca. 14--17+. Female fls. ca. 4.5--5.5 mm long. Ripe frts. ca. 0.9--1.3 X 0.6--0.8 cm, ± broadly ellipsoidal to obovoid. Wet forests, 0--700 (--1100) m; Atl. slope all major cords., Pac. slope Cord. Guanacaste and S from RB Carara. Fl. 5--7, 9--11. S Mex. (Chis.), Hond. to extr. NW Col. [Grayum & Herrera 4838, CR, MO.]

Its simple or minimally pinnate, non-fenestrate lf.-blades separate R. simplex from all Costa Rican Reinhardtia species except the rare and local R. koschnyana, which has consistently simple and generally more deeply bifid lf.-blades, spicate infls., and male fls. with only ca. half as many stamens (see key, couplet 3). Its cespitose habit and coarsely serrate or doubly serrate lf. margins combine to distinguish R. simplex from grossly similar, small, understory palms in genera such as Chamaedorea and Geonoma.

This is the most widespread and generally most common Reinhardtia species in Costa Rica, and thus the one most likely to be encountered.

Sabal

16 spp., SE U.S. (S. Okla., Ark., N. Carol.) and N Mex. (Chih., Son.) to Pac. Col., Venez., Trin., Gr. Ant. (Cuba, Hispan., Jam., PR), Anegada (BVI), Bahamas.

Zona, S. 1990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso 12: 583--666.

Sabal mauritiiformis (H. Karst.) Griseb. & H. Wendl., in Griseb., Fl. Brit. W. I. 514. 1864. [Trithrinax mauritiiformis H. Karst., Linnaea 28: 244. 1856; S. allenii L. H. Bailey; S. morrisiana Bartlett ex L. H. Bailey; S. nematoclada Burret]. GUAGARÓN--Stems erect, solitary, ca. 5--12 (--25?) m tall and 15--45 cm diam., unarmed. Petioles beyond sheath ca. 1.3--3 m, the margins sharp but not toothed, with an acuminate to acute adaxial hastula ca. 3.5--11 X 2--3 cm, the sheath splitting, with fibrous margins. Lf.-blades palmately deeply divided into drooping, twice bifid segments, ca. 2--3 X 2--3 m, whitish abaxially, with elongate midrib ca. 90--125 cm. Plants hermaphroditic, the infls. interfoliar. Prophyll short, 2-lobed. Peduncle ca. 1--2 m, with several bracts, these tubular basally and narrowly beaked distally, ± tomentose. Infls. paniculately branched to 4 orders, erect, well exceeding the lvs.; main axis ca. 1--3.5 m long; secondary axes to at least 70 cm. Fls. bisexual, ca. 3 mm long, cream-colored, rankly fragrant, solitary and subsessile, spirally arranged on bracteate, low spur branches. Sepals connate in a shallowly 3-lobed tube. Petals 3, connate basally in a tube, suberect and spreading distally. Stamens 6, the filaments connate in basal 1/2 in a tube adnate to corolla. Pistil 1, 3-locular; style elongate; stigma capitate, 3-lobed. Ripe frts. ca. 0.9--1.1 X 0.9--1.1 cm, smooth, globose to turbinate, purplish black, with basal stigmatic residue. Moist forests, 0--200+ m; very rare and local, S Atl. coast (RNVS Gandoca-Manzanillo), cent. Pac. coast (basin of Río Tusubres). Fl. 8. S Mex. (Oax., Ver.) to E Hond. (Gracias a Dios), CR to Venez., Trin. [Grayum et al. 11228, CR, INB, MO.]

Sabal and Colpothrinax are the only Costa Rican fan-palms that are not armed with thorns along either the trunk (Cryosophila) or the petiole margins (Acoelorraphe). Sabal may by told from Colpothrinax by its conspicuously costate lf.-blades, more openly branched infls., smaller frts., and lower-elevation habitat; see the generic key (couplet 4) for additional distinguishing features.

Sabal mauritiiformis, spottily distributed throughout its range, is known from Costa Rica only by a small and endangered population on coral headlands in RNVS Gandoca-Manzanillo, on the S Atl. coast, and by several small, scattered, unprotected, and much perturbed populations in the central Pac. region. The lvs. are prized for thatching, said to last as long as 20 years.

Plants of an exotic Sabal sp. are occasionally seen in Costa Rican gardens (e.g., J. González et al. 1087, INB; "hojas usadas para fabricar escobas"). The specific identity of these planted specimens has not been established with certainty.

Socratea

5 spp., SE Nic. to Pac. Ec., Guianas, Braz., Bol.

Henderson, A. 1990. Arecaceae. Part I. Introduction and the Iriarteinae. Fl. Neotrop. Monogr. 53: 1--100.

Socratea exorrhiza (Mart.) H. Wendl., Bonplandia 8: 103. 1860. [Iriartea exorrhiza Mart., Hist. nat. palm. 36, t. 33--34. 1824; S. durissima (Oerst.) H. Wendl.]. CHONTA, MAQUENQUE--Stems erect, solitary, ca. 6--25+ m tall and 5--25 cm diam., unarmed, supported by an open cone of brown, spiny stilt-roots ca. 1--2+ m tall. Petioles beyond sheath ca. 10--30 (--46) cm, unarmed, the sheath closed and forming a distinct, greenish crownshaft ca. 0.95--1.5 m long. Lf.-blades pinnately compound, ca. 2--4 m long, the rachis unarmed, the lflts. ca. 8--25 per side, large, cuneate at base, praemorse distally, each splitting longitudinally into 3--7 (usually) narrow segments arranged in various planes; terminal lflt. bifid. Plants monoecious, the infls. bisexual, infrafoliar, erect in bud, later reflexed or pendent. Prophyll to ca. 11 cm long, thinly coriaceous. Peduncle ca. 16.5--28+ cm, ± densely tomentose at anthesis, with ca. 3--5 ± flattened, beaked bracts, the central ones largest (ca. 30--50+ cm long). Infls. racemosely branched; rachis ca. 8--17 cm, tomentose as peduncle; rachillae ca. 9--16, ca. 19.5--64 cm, tomentose as peduncle. Fls. unisexual, spirally arranged in obscurely bracteate triads. Male fls. ca. 6--9 mm long, white, with 3 basally connate sepals and 3 much larger, distinct, asymmetrical petals; stamens ca. 30--65, yellow, with very short filaments. Female fls. ca. 3--5 mm long, with 3 ± distinct, ± rounded, pubescent sepals and 3 distinct, ± rounded, ciliate petals; staminodia very minute or obsolete; pistil 1, 3-locular; stigmas 3, fleshy. Ripe frts. ca. 2--3.3 X 1.3--2.6 cm, subglobose or ovoid to (much more frequently) oblong-ellipsoidal or obovoid, ± smooth, green to yellowish, ± dehiscent, with apical stigmatic residue. Wet forests, 0--750 m; Atl. slope Cords. Central (to RNFS Barra del Colorado and PN Tortuguero) and Talamanca, Pac. slope V. de El General, Golfo Dulce region. Fl. 2--3. Extr. SE Nic. to Bol., Fr. Guiana, Braz. [Stevens & Montiel 24559, CR {check!}, MO.]

Socratea exorrhiza is a canopy or subcanopy palm, recognized by its tall, open cone of brownish, spiny stilt-roots and pinnately compound lf.-blades with broadly cuneate, praemorse, longitudinally splitting. lflts., with the segments disposed in various planes. These features distinguish it sharply from all other Costa Rican palms except for the very similar and sympatric (though perhaps more widespread) Iriartea deltoidea, which differs in having a denser cone of blackish stilt-roots and lf.-blades with an entire, rather than bifid, terminal segment. For additional differences, see under I. deltoidea and the generic key (couplet 6).

The wood of this species, like that of Iriartea deltoidea, is very hard, and has been used in construction. However, unlike I. deltoidea, the palmito of Socratea exorrhiza is unpalatable (though edible).

Socratea exorrhiza, like most large palms, is poorly represented in herbaria. Therefore, it is undoubtedly distributed more widely within Costa Rica than indicated above.

Synechanthus

Stems erect, slender, solitary or cespitose, unarmed. Petioles unarmed, with sheath soon splitting. Lf.-blades simple and bifid to (most commonly) unequally to equally pinnately compound, the rachis unarmed, the lflts. ± straight to evidently sigmoid, regularly spaced or clustered, arranged in one or various planes. Plants monoecious, the infls. bisexual, interfoliar or often becoming infrafoliar. Prophyll short, disintegrating into fibers. Peduncle usually long, with 4--5 bracts, these longer than the prophyll, the distal ones exceeding the peduncle. Infls. racemosely to subpaniculately branched, the rachillae slender, flexuous, angled. Fls. unisexual, very small, ebracteate, borne in distichously arranged files (acervuli) of 1 (--2) proximal female fl(s). and 5--15 distal, biseriate male fls. Male fls. with sepals connate in a low, acutely 3-lobed cup and 3 distinct petals; stamens 3 or 6. Female fls. with sepals connate in a 3-lobed cup and 3 distinct petals; staminodia not evident, or connate in a 6-lobed ring partially adnate to the petals; pistil 1, 3-locular; stigmas 3, short, recurved. Ripe frts. smooth, relatively large, subglobose to narrowly ellipsoid, very fleshy, yellow to red, with basal stigmatic residue. 2 spp., S Mex. (Oax., Ver.) to Pac.

Palms of the small genus Synechanthus are recognized by their understory habit, usually pinnately compound lf.-blades, and long-pedunculate infls. with numerous, slender, angled rachillae, tiny fls., and large, bright yellow to red frts. In Costa Rica, Synechanthus is the only close relative of the much larger genus Chamaedorea. The two genera are similar and easily confused, but Chamaedorea differs in being dioecious. Moreover, Synechanthus is unique among Costa Rican palms in having its fls. disposed in narrow rows called acervuli, each consisting of a single female fl. (rarely 2) and 5--15 male fls. As in most Chamaedorea spp., the female fls. of Synechanthus may be told from the male fls., even in bud, by the imbricate (as opposed to valvate) aestivation of their petals. On fruiting rachillae, scars betray the former position (and linear arrangement) of the abscised male fls.

The rare Hyospathe elegans, which has racemosely branched infls. with slender rachillae, thus also bears a resemblance to our Synechanthus spp. (especially S. warscewiczianus), but differs in its closed petiole sheaths and much shorter peduncles (ca. 2.7--6.1 cm). Synechanthus differs from H. elegans and all our Chamaedorea spp. with branched infrs. in having yellow to orange or red (rather than purplish to black) ripe frts. The only Costa Rican Chamaedorea spp. with orange frts. (C. crucensis, C. deckeriana, C. zamorae) have spicate infrs.

Moore, H. E., Jr. 1971. The genus Synechanthus (Palmae). Principes 15: 10--19.

1 Lf.-blades pinnately compound, the lflts. ca. 13--19+ per side, clustered and ± arranged in various planes (at least in proximal half of blade), ± sigmoid; peduncle ca. 41--103 cm; infls. subpaniculately to paniculately branched; rachillae ca. 7--28.5 cm, minutely papillate-scaberulous; male fls. 5--7 per acervulus; stamens 6, erect in bud, included at anthesis, the anthers ca. as long as or longer than the filaments; Atl. slope... S. fibrosus

1' Lf.-blades simple and bifid (very rarely) to pinnately compound, with the lflts. ca. (2--) 5--34 per side, regularly or ± irregularly spaced (but not in discrete clusters), ± straight; peduncle ca. (24--) 28--73 (--78) cm; infls. racemosely branched; rachillae ca. 8--48 cm, glabrous; male fls. 6--15 per acervulus; stamens 3, inflexed in bud, long-exserted at anthesis, the anthers much shorter than the filaments; both slopes...S. warscewiczianus

Synechanthus fibrosus (H. Wendl.) H. Wendl., Bot. Zeitung (Berlin) 16: 145. 1858. [Chamaedorea fibrosa H. Wendl., Index palm. 57. 1854].--Stems solitary, obsolete to 2.5+ m tall and 2--9 cm diam. Petioles beyond sheath ca. 22--75 cm. Lf.-blades ± equally pinnately compound, the rachis ca. 74--104 cm; lflts. ca. 13--19+ per side, clustered (in groups of 2--6) and ± arranged in various planes (at least in proximal half of blade), ca. 16.5--41+ X 1.2--5.7 (subterminal ones), ± sigmoid. Peduncle ca. 41--103 cm. Infls. subpaniculately to paniculately branched; rachis ca. 8.2--26.5 cm; rachillae ca. 25--75, 7--28.5 cm long, minutely papillate-scaberulous. Male fls. 5--7 per acervulus, ca. 0.8--1.0 mm long, whitish to yellowish; stamens 6, erect in bud, included at anthesis, the anthers ca. as long as or longer than the filaments. Female fls. ca. 1.0 mm long. Ripe frts. 1.2--2.0 X 0.5--1.1 cm, ellipsoidal to oblong-ellipsoidal (often ± falcate), yellow to bright orange. Wet forests, 0--1200 m; Atl. slope Cords. Central (to RNFS Barra del Colorado and PN Tortuguero) and Talamanca. Fl. 1, 3--12. S Mex. (Oax., Ver.) to CR (and ostensibly extr. W Pan.). [Grayum et al. 8919, CR, MO.]

Synechanthus fibrosus is most readily distinguished from the more common S. warscewiczianus by its usually clustered lflts. (reminiscent of some Bactris spp.) and subpaniculately to paniculately branched infls. (i.e., with at least some proximal branches forked or more extensively branched). The infls. of S. warscewiczianus are consistently branched racemosely; rarely, S. fibrosus may have racemosely branched infls. (in which case the other key characters must be used), but I have seen no such collections from Costa Rica.

This is an uncommon, if widely scattered species, though it may be locally abundant (as near Chilamate de Sarapiquí). It has been collected, at both extremes of its altitudinal range, from very near the Panamanian border, but curiously, not yet from Panama itself.

Synechanthus warscewiczianus H. Wendl., Bot. Zeitung (Berlin) 16: 145. 1858. [S. angustifolius H. Wendl.].--Stems solitary (at least apparently) or cespitose, ca. 0.6--5 m tall and 1.5--7 cm diam. Petioles beyond sheath ca. 18.5--65 cm. Lf.-blades simple and bifid or (much more frequently) equally to unequally pinnately compound, the rachis ca. 67--128 cm; compound lvs. with lflts. ca. (2--) 5--34 per side, regularly or ± irregularly spaced, arranged in a single plane, ca. 19--60.5 X 0.9--13.5 (subterminal ones), ± straight . Peduncle ca. (24--) 28--73 (--78) cm. Infls. racemosely branched; rachis ca. (5.2--) 7.4--20 (--23) cm; rachillae ca. (17--) 25--61, 8--48 cm long, glabrous. Male fls. 6--15 per acervulus, ca. 0.5--1.0 mm long, whitish to yellowish or orange; stamens 3, inflexed in bud, long-exserted at anthesis, the anthers much shorter than the filaments. Female fls. ca. 0.8--1.0 mm long. Ripe frts. 1.2--2.7 X 0.6--1.1 cm, subglobose (especially in Pac. lowlands) or ovoid (rarely) to narrowly or broadly oblong-ellipsoidal (often ± falcate), yellow to orange or red. Wet forests, 0--1450 m; entire Atl. slope, Pac. slope Cords. Guanacaste and Tilarán (> 800 m) and S from Tarrazú region and Fila Chonta. Fl. 2--12. N Nic. to Pac. Ec. [I INBio 47, CR, MO.]

Synechanthus warscewiczianus is a common understory palm in wet forests throughout Costa Rica. It is especially conspicuous in fruit, with its relatively large, usually bright orange to red fruits borne on slender, divergent rachillae. Perhaps for this reason it is one of the most frequently collected Costa Rican palms. See under the genus heading for differences from similar species in other genera, and under S. fibrosus and in the species key for differences from the last-mentioned species.

Some of the ostensible distinctions between the two Synechanthus species suggested by Moore (1971) are spurious, or, at best, of limited use. Synechanthus fibrosus was described as having solitary stems and S. warscewiczianus as cespitose; however, stems of the latter species are described as solitary on most labels. The character of "minutely scaberulous" vs. "not minutely scaberulous" infl. rachillae, though employed in the above key, is scarcely appreciable even at high magnification. Moore's contention that female fls. of S. fibrosus lack staminodia or have ("perhaps") only three is false; they have (at least potentially) the same complement of six connate staminodia as female fls. of S. warscewiczianus.

Synechanthus warscewiczianus is an unusually variable species, and some of the variation is correlated with geography. The lf.-blades of this species are typically pinnately compound, with ca. (2--) 5--15 irregularly spaced lflts. of variable width per side. I have seen just two collections (from throughout the range of the species) with simple, bifid lf.-blades; both are from ca. 900--1200 m elevation in the southern Fila Costeña of Costa Rica (Aguilar 2697, INB; I. Chacón 323, USJ). Populations from ca. 200--1150 m elevation on the Atl. slope of Cords. Tilarán, Central and Talamanca (e.g., Grayum et al. 8735, CR, MO), and extending into Panama, are consistently characterized by pinnately compound lf.-blades with relatively numerous (ca. 22--34), ± regularly spaced lflts. of uniformly narrow width (ca. 0.9--3.6 cm); the name Synechanthus angustifolius corresponds to this entity. Because neither of these unusual lf.-forms appears to differ in any other respect from the more typical material, formal taxonomic recognition seems unwarranted.

This species is even more variable in Panama, with organs other than lvs. involved (e.g., several collections provisionally referred here have as few as 3--4 infl. rachillae). The above description applies only to Costa Rican material.

Welfia

1--2 spp., E Hond. to Pac. Ec., Amaz. Peru.

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

Welfia regia H. Wendl. ex André, Ill. Hort. 18: 93, t. 62. 1871. [W. georgii H. Wendl. ex Burret].--Stems erect, solitary, ca. 5--25 m tall and 10--30 cm diam., unarmed. Petioles beyond sheath ca. 30--100 cm, unarmed, densely tomentose, the sheath splitting. Lf.-blades pinnately compound, ca. 2.5--5+ m long, the rachis unarmed, the lflts. ca. 70--90 per side, subequal, ca. 24--135+ X 1--12.5 cm, lanceolate, regularly spaced and arranged in a single plane. Plants monoecious, the infls. bisexual, becoming infrafoliar. Prophyll ca. 60--85+ cm long, flattened, woody, tomentose. Peduncle ca. 6--40+ cm, stout, recurved, with several bracts, the largest (basal) ca. as long as but thinner than prophyll. Infls. racemosely branched; rachis ca. 3--7.5 cm; rachillae 4--10 (--12), ca. 36--80 X 1.5--2.7 cm, minutely stellate-scurfy. Fls. unisexual,white arranged in 7--9 vertical rows of bracteate triads in bilabiate pits. Male fls. ca. 1.1--1.6 cm long, with 3 ± distinct sepals and 3 petals connate in a tube for ca. 1/3--1/2 their length; stamens ca. 39--45. Female fls. ca. 1.0--1.3 cm long, with 3 distinct sepals and 3 petals connate in tube for 2/3+ their length, the lobes triangular; staminodia 12--16, adnate to corolla tube for 2/3 their length; pistil 1, 3-locular, 3-angled; style elongate, cylindrical; stigmas 3, recurved. Ripe frts. ca. 2.5--4.0 X 1.3--1.9 cm, ± narrowly ovoid to elliptical or obovoid, ± dorsiventrally compressed, keeled distally, smooth, reddish brown to dark purple, with basal stigmatic residue. Wet forests, ca. 50--750 m; Atl. slope Cords. Central (to Llanura de los Guatusos) and Talamanca, Pac. slope at head of Golfo Dulce and Pen. Osa. Fl. 4--5 (6, 8, 11; Pan.). E Hond. (Gracias a Dios) to Pac. Ec. (and possibly Amaz. Peru). [Aguilar 2234, CR.]

Welfia is one of the "geonomoid" palms, characterized by having their floral triads deeply sunken in rachillar pits (see under Geonoma for additional information). Welfia regia is by far the largest Costa Rican palm in this group, with the thickest infl. rachillae and the largest frts. It is further distinguished by its very numerous (ca. 39--45) stamens (all our other geonomoids have just 6) and staminodia.

This is a species of wet, lowland, primary forests, recognized vegetatively by its arborescent habit, solitary stems, and very long, evenly pinnate lf.-glades, with relatively narrow lflts. In overall aspect, it is somewhat reminiscent of Attalea butyracea or Cocos nucifera, but those species are absent from wet forests. The new lvs. of W. regia are distinctively reddish brown in coloration, even on juveniles.

Welfia regia is generally an uncommon species in Costa Rica, but may be very abundant locally, as at EB La Selva. Like most large palms, it is poorly represented in herbaria, and for this reason may be more widespread in Costa Rica than indicated above.

The sole Peruvian collection (Gentry et al. 63540, MO) has several morphological peculiarities, and is from a much higher elevation (> 1500 m) than other recent collections (all from < 900 m). It may well represent a different species. The name Welfia regia is here applied in a broad sense following Henderson et al. (1995). In fact, the application of the name (based on an old collection from montane Colombia) is still unclear, and merits further investigation. It is conceivable that the Peruvian entity represents the "real" W. regia, and that all remaining material should be called Welfia georgii (typified by a Costa Rican collection).

The palmito of W. regia has been described as sweet and edible. The large, almond-shaped fruits are much sought after by forest mammals and birds.

TOP

 
 
© 1995-2014 Missouri Botanical Garden, All Rights Reserved
P.O. Box 299, St. Louis, MO 63166-0299
(314) 577-5100

E-mail
Technical Support