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 Terrestrial plants (sometimes paludal), acaulescent or stems rhizomatous to (usually) erect, often massive and arborescent, solitary or cespitose (but the aerial portion unbranched in ours), occasionally (Desmoncus) scandent, smooth to spiny. Petioles unarmed or spiny, with or without a distal hastula (ligule-like structure). Lvs. simple or bifid to pinnately compound or palmately divided, spiraled or (rarely) di- or tristichous, ± clustered at apex of stem, ± strongly plicate, glabrous to pubescent and/or spiny, rarely (Attalea, Reinhardtia) fenestrate, the margins entire to serrate. Plants hermaphroditic, monoecious, or (Chamaedorea) dioecious, the infls. axillary (rarely apparently terminal), interfoliar (borne among the lvs.) or infrafoliar (borne below the lvs.), subtended by a usually 2-keeled prophyll. Infl. spicate to racemosely or paniculately branched. Peduncle obsolete to elongate, bearing 0--many, often conspicuous bracts. Fls. usually sessile and much reduced, bisexual or unisexual (in which case often borne in triads of 2 male fls. and 1 female fl.), ± actinomorphic, usually bracteate. Perianth usually clearly differentiated into calyx and corolla, of usually 3 distinct or ± connate sepals and petals (respectively). Stamens 3 or 6--indefinite, distinct or ± connate; staminodia usually evident in female fls., sometimes connate. Pistils 1 (--4); if 1, monomerous (or pseudomonomerous) to compound; ovary superior, with 1 or 3 (+) locules; styles 1--3 (+), or not evident; stigmas erect or recurved; ovules 1 per locule; placentation axile, basal, or subapical. Frt. a usually 1-seeded drupe, glabrous to spiny or scaly, sometimes (Iriartea, Socratea) ± dehiscent. Seeds with copious endosperm. Ca. 200 genera and perhaps 2500 spp.; pantropical, with very few spp. in temperate regions; many spp. are cultivated for ornament, and some for food. Palms are most abundant and diverse in Costa Rica in humid, lowland forests. They are usually easily recognized by their familiar, plicate lf.-blades. The only other family in Costa Rica with which Arecaceae can easily be confused is Cyclanthaceae, which have many-seeded berries rather than single-seeded drupes. See under the last-mentioned family for further information. Numerous exotic palms are cult. for ornamental purposes in Costa Rica. Probably the most frequent of these is the Madagascan Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. (Chrysalidocarpus lutescens H. Wendl.) [Poveda 680 (CR)], commonly called areca. This is a smallish (to ca. 7 m), cespitose species with yellowish petioles and lf.-rachises and a glaucescent crown-shaft. Henderson, A., G. Galeano & R. Bernal. 1995. Field guide to the palms of the Americas. Princeton Univ. Press, Princeton, NJ. Uhl, N. W. & J. Dransfield. 1987. Genera palmarum. L. H. Bailey Hortorium & International Palm Soc., Lawrence, KS. 1 Lf.-blades palmately divided and veined, ± suborbicular in outline, the midrib obsolete or very short; plants hermaphroditic, the fls. bisexual 2 Stems cespitose; petioles armed marginally with sharp, triangular spines; rare, N Atl. slope...Acoelorraphe 2' Stems solitary; petioles unarmed (sometimes sharp-edged, but lacking spines) 3 Stems ± densely armed with usually branching root-spines, especially toward base; lf.-blades split centrally nearly or quite to the base; ripe frts. white; widespread...Cryosophila 3' Stems unarmed; lf.-blades with central split not or scarcely deeper than lateral splits; ripe frts. blackish; rare and local 4 Stem apex densely covered with matted, reddish brown petiole-sheath fibers; midrib of lf.-blades obsolete (ca. 6--8 cm); pistils 3 at anthesis (the styles connate); ripe frts. ca. 1.3--2.2 X 1.4--2.2 cm, with apical stigmatic residue; ca. 700--1000 m, Atl. slope Cords. Guanacaste and Talamanca, Pac. slope Fila Costeña...Colpothrinax 4' Stem apex mostly bare, the petiole sheaths sparingly fibrous marginally; midrib of lf.-blades elongate (ca. 90--125 cm); pistil 1; ripe frts. ca. 0.9--1.1 X 0.9--1.1 cm, with basal stigmatic residue; ca. 0--200+ m, Talamanca coast, lower Río Tulín (R. Tusubres) basin...Sabal 1' Lf.-blades simple and bifid to pinnately compound, pinnately veined, ± elongate in outline; plants monoecious or dioecious, the fls. unisexual 5 Stems solitary, to 20--30 m tall, supported at base by dense or open cone of spiny stilt-roots to 1--2+ m tall; petiole sheaths closed, forming a distinct, greenish crownshaft; lflts. large, wedge-shaped, cuneate at base, praemorse distally, splitting longitudinally into narrow segments arranged in various planes; stamens 10--65; frts. ± dehiscent, the seeds ultimately falling free 6 Stilt-roots to ca. 100, blackish, forming a very dense cone; terminal lflt. entire; infls. decurved in bud; stamens ca. 10--17; frts. (and seeds) ± globose...Iriartea 6' Stilt-roots to ca. 25, brown, forming an open cone; terminal lflt. bifid; infls. erect in bud; stamens ca. 30--65; frts. (and seeds) ovoid-cylindric...Socratea 5' Stems various, but never supported by spiny stilt-roots; petioles forming a crownshaft (some Chamaedorea spp., Euterpe, Hyospathe, Prestoea, Oenocarpus) or not; lflts. wedge-shaped (Aiphanes, Reinhardtia) or not, arranged in a single or various planes; stamens 3--6 (--28, Reinhardtia; --45, Welfia); frts. indehiscent, the seeds not falling free 7 Plants at least sparingly armed with spines on stem, petioles, lf.-rachis, and/or lf.-blades 8 Plants scandent; lf.-blades pinnately compound, with rachis prolonged distally into a long appendage (cirrus) bearing rigid, retrorse acanthophylls (modified lflts.)...Desmoncus 8' Plants erect; lf.-blades simple or pinnately compound, then with rachis not prolonged distally, or prolonged in a filamentous extension lacking acanthophylls 9 Lvs. pinnately compound, the lflts. cirrhous at apex; infl. spicate; rare, Pen. Osa...Desmoncus 9' Lf.-blades simple or pinnately compound, then lflts. acute to acuminate or caudate at apex; infl. branched 10 Stems and petioles without spines, the lf.-rachis ± inconspicuously spiny along the adaxial keel; lf.-blades with spines restricted to main veins (on adaxial side) and margins of lflts.; lf.-blades ca. 5--10+ m long, pinnately compound, the lflts. very numerous (ca. 95--205 per side) and arranged in various planes; infls. paniculately branched; frts. covered in 9--11 vertical rows of closely overlapping, glossy scales; locally abundant in lowland swamps...Raphia 10' Stems, petioles, or lf.-rachis spiny conspicuously; lf.-blades spiny or not; lf-blades to ca. 5 m long, simple or pinnately compound, the lflts. various; infls. racemosely branched; frt. smooth to spinulose, but not scaly; various habitats 11 Stems stout, solitary; spines short and stout, restricted to petiole margins, the plants otherwise unarmed; infls. unisexual, massive and condensed, the rachillae with spinelike tips...Elaeis 11' Stems various; spines slender, terete or flattened, ± uniformly distributed on petiole (not restricted to petiolar margins) and usually present elsewhere (stem, lf.-rachis, and/or lf.-blades); infls. bisexual, condensed to open, the rachillae [not prominently spinelike at tips] 12 Stems solitary, stout, subarborescent to arborescent; lf.-blades pinnately compound, ± whitish below 13 Dead lvs. persistent, skirting upper half of trunk; spines terete or subterete (flattened adaxially); female fls. with distinct sepals and petals; seasonally dry forests, mostly on Pac. slope...Acrocomia 13' Dead lvs. caducous, not skirting trunk; spines strongly flattened; female fls. with sepals [and sometimes petals] connate in a truncate to shallowly 3-lobed cup or tube; wet forests of both slopes...Astrocaryum 12' Stems cespitose, slender or rarely (Bactris gasipaes) stout and arborescent; lf.-blades simple to pinnately compound, ± concolorous (but see Bactris caudata, B. grayumii) 14 Lf.-blades pinnately compound, the lflts. wedge-shaped, cuneate at base, praemorse distally; peduncles elongate, divergent; very rare, Fila Costeña...Aiphanes 14' Lf.-blades simple to pinnately compound, then with lflts. linear to narrowly elliptical or oblanceolate, variously narrowed at base, acute to caudate distally; peduncles usually relatively short, recurved; widespread...Bactris 7' Plants unarmed, lacking spines 15 Lf.-blades simple, bifid at apex 16 Infls. spicate 17 Stems cespitose 18 Stems ca. 1--2.5 m tall; lf.-blades with primary lateral veins ca. 17--32 per side, the margins subentire; floral triads deeply sunken in bilabiate rachillar pits; ripe frts. ca. 0.5--0.6 X 0.5--0.55 cm, subglobose, with basal stigmatic residue; 1300--1900 m, cent. Pac. slope...Geonoma 18' Stems ca. 0.15--1.2 m tall; lf.-blades with primary lateral veins ca. (7--) 8--10 (--11) per side, the margins very coarsely serrate or doubly serrate; floral triads not sunken in rachillar pits; ripe frts. ca. 0.9--1.3 X 0.6--0.8 cm, ± broadly ellipsoidal to obovoid, with apical stigmatic residue; 50--250 m, N Atl. slope...Reinhardtia 17' Stems solitary 19 Stems green; foliage virtually glabrous; lf.-blades marginally toothed (sometimes ± obscurely), at least distally; plants dioecious; fls. essentially ebracteate, solitary or in irregular groups, superficial or slightly immersed...Chamaedorea 19' Stems brownish; foliage usually ± scurfy or scaly, especially petioles and main lf. veins on abaxial side; lf.-blades marginally entire; plants monoecious; fls. borne in bracteate triads in bilabiate pits [Proceed to couplet 35]. 16' Infls. branched 20 Stems cespitose; lf.-blades with margins coarsely serrate or doubly serrate; male fls. with ca. 14--17+ stamens; frts. with apical stigmatic residue...Reinhardtia 20' Stems solitary or cespitose, if cespitose then lf.-blades with margins subentire; male fls. with 3 or 6 stamens; frts. with basal stigmatic residue 21 Stems green; foliage virtually glabrous; peduncular bracts usually > 1; fls. essentially ebracteate, solitary or in lines, superficial or slightly immersed [Proceed to couplet 38]. 21' Stems brownish; foliage usually ± scurfy or scaly, especially petioles and main lf. veins on abaxial side; peduncular bract usually 1; fls. borne in bracteate triads in bilabiate pits 22 Labia of pits immersed, the margins later recurved; ovary 3-celled at anthesis, the style apical; frts. with fibrous mesocarp...Asterogyne 22' Labia of pits elevated, later splitting medially; ovary 1-celled at anthesis, the style basal; frts. with mesocarp virtually lacking fibers...Geonoma 15' Lf.-blades (at least some) pinnately divided to compound 23 Frts. > 10 cm diam., with thick, dry, fibrous mesocarp...Cocos 23' Frts. < 10 cm diam., with usually thin, fleshy mesocarp 24 Frts. > 3 cm diam., globose to 3-lobed, ± woody, covered with pyramidal, wartlike projections; swamp forests near coast...Manicaria 24' Frts. usually < 3 cm diam., globose to ovoid or ellipsoid (never lobed), usually fleshy, smooth to (rarely) minutely echinulate 25 Petiole sheaths closed and forming a conspicuous crownshaft usually > 0.5 m long; ± slender canopy or subcanopy trees, the fertile stems mostly > 4 m tall; lf.-blades with numerous, ± narrow pinnae; infls. racemosely branched; ovary 1-celled 26 Stems solitary; crownshaft bright green; infl. rachillae densely covered with whitish hairs; widespread, 0--1150 m...Euterpe 26' Stems cespitose; crownshaft usually grayish to purplish; infl. rachillae glabrous or inconspicuously pubescent 27 Lf.-blades whitish or glaucous-looking below; infl. rachillae pendent, closely spaced, all borne on abaxial side of short rachis, the whole resembling a horse's tail; frts. > 1.2 cm diam.; 0--350 m, Pac. slope...Oenocarpus 27' Lf.-blades ± concolorous; infl. rachillae ± divergent to arching, well distributed along and around the ± elongate rachis; frts. < 1.2 cm diam.; widespread, (1000--) 1400--2450 m...Prestoea 25 Petiole sheaths open, or closed and forming a brief crownshaft < 0.5 m long; understory subshrubs to canopy trees; lf.-blades simple to pinnately compound; infls. spicate to racemosely or paniculately branched; ovary 1- (rarely) or 3-celled 28 Stout, subcanopy trees, the stems solitary, 10+ cm diam. and usually > 7 m tall; lf.-blades > 3 m long, regularly pinnate with > 40 lflts. per side; frts. to > 1.5 cm diam. 29 Lflts. blunt and strongly inequilateral at apex; young. lvs. green; infls. interfoliar, the rachillae > 15, shorter than peduncle, < 1.3 cm wide; stamens 6; frts. ± ellipsoidal, > 4.5 cm long; seasonally dry forests, mainly on Pac. slope...Attalea 29' Lflts. acuminate and ± symmetrical at apex; young lvs. reddish brown; infls. quickly becoming infrafoliar, the rachillae < 15, longer than peduncle, > 1.3 cm wide; stamens ca. 39--45; frts. ± dorsiventrally compressed, almond-shaped, < 4.5 cm long; wet forests, Atl. slope and Golfo Dulce region...Welfia 28' Slender understory plants or (rarely) subcanopy trees, the stems solitary or cespitose, to ca. 10 cm diam., to ca 6 (--10) m tall; lf.-blades < 3 m long, regularly or irregularly pinnate with up to 45 lflts. per side; frts. < 1.5 cm diam. 30 Stems cespitose; lflts. ± praemorse distally, sometimes fenestrate at base, very unequal [the terminal lflt(s). somewhat to much wider]; stamens ca. 7--28...Reinhardtia 30' Stems solitary or cespitose; lflts. acute to acuminate distally, never fenestrate, equal or unequal; stamens (3) 6 31 Infls. spicate; stems solitary 32 Stems green; lvs. virtually glabrous; plants dioecious; fls. essentially ebracteate, solitary or in irregular groups (not regularly in triads)...Chamaedorea 32' Stems brownish; lvs. ± scurfy or scaly, especially on the petiole and main lf. veins on abaxial side; plants monoecious; fls. bracteate, mostly in triads 33 Stems > 2.5 m tall; infls. infrafoliar, the peduncle shorter than the rachilla; ripe frts. to ca. 2.5 cm long...Pholidostachys 33' Stems obsolete to (exceptionally) ca. 2.5 m tall; infls. interfoliar, the peduncle usually longer than the rachilla; ripe frts. to 2.0 cm long 34 Fls. superficial or scarcely immersed in rachilla; male fls. with sepals basally connate in a cup; fls. of both sexes with distinct petals; female fls. lacking staminodia; frts. with apical stigmatic residue...Neonicholsonia 34' Fls. deeply sunken in bilabiate pits; male fls. with distinct sepals; fls. of both sexes with petals connate, at least basally; female fls. with staminodia connate in a truncate to 6-toothed or -lobed tube; frts. with basal stigmatic residue 35 Peduncular bract inserted distally on peduncle (just below flowering rachilla), leaving a conspicuous annular scar; labia of pits immersed, the margin later recurved (sometimes also splitting medially); petals of female fls. connate for most of their length in an apically 3-lobed tube; ovary 3-locular, the style apical; frts. with fibrous mesocarp...Calyptrogyne 35' Peduncular bract inserted basally on peduncle, not leaving a distal, annular scar; labia of pits elevated, later splitting medially; petals of female fls. connate basally, free distally; ovary 1-locular, the style basal; frts. with mesocarp virtually lacking fibers...Geonoma 31' Infls. branched; stems solitary or cespitose 36 Stems brownish; infls. racemosely to paniculately branched; fls. deeply sunken in bilabiate pits; ovary 1-celled, the style basal...Geonoma 36' Stems brownish or green; infls. racemosely or (rarely) subpaniculately branched (with some basal rachillae forked); fls. superficial or scarcely immersed; ovary 1- or 3-locular, the style central 37 Stems green; foliage virtually glabrous; peduncular bracts usually > 1; fls. essentially ebracteate, solitary or in lines; ovary 3-locular; frts. with basal stigmatic residue 38 Plants dioecious; fls. usually solitary, occasionally in small fascicles; ripe frts. < 2.0 cm (usually < 1.5 cm) long, purplish to black (or, if yellow to red, infls. unbranched)...Chamaedorea 38' Plants monoecious; fls. borne in lines (acervuli) of 1 (--2) female and 5--15 male fls.; infls. branched; ripe frts. to 1.5--2.0+ cm long, yellow to red...Synechanthus 37' Stems brownish or green; foliage ± scurfy or scaly, especially on the petiole and main lf. veins on abaxial side; ovary 1-locular; frts. with subapical to basal stigmatic residue 39 Stems green, cespitose; male fls. (at least some) pedicellate; female fls. with sepals connate in basal half; frts. narrowly ovoid, with basal stigmatic residue...Hyospathe 39' Stems brownish, solitary or cespitose; male fls. sessile; female fls with distinct sepals; frts. subglobose, with lateral stigmatic residue...Prestoea TOP