Selbyana 9: 243-259 THE VASCULAR FLORA OF LA SELVA BIOLOGICAL STATION, COSTA RICA* MORACEAE

BARRY E. lIAMMELt Department of Botany, Duke University, Durham, North Carolina 27706

BERG, C. C. 1972. Olmedieae and Brosimeae (Mo­ often 4-parted or sometimes apparently without raceae). Fl. Neotropica Monogr. no. 7: 1-229. perianth and unorganized; perianth uniseriate --. 1977. U rticales, their differentiation and sys­ with lobes imbricate or valvate; stamens of the tematic position. Plant Syst. Evol., Suppl. 1: 349- same number as or fewer than the perianth lobes 374. and opposite them, the filaments straight or in­ --. 1977. The Castilleae, a tribe ofthe Moraceae, renamed and redefined due to the exclusion of the curved in bud, the anthers usually 2-thecous and type genus Olmedia from the "Olmedieae." Acta longitudinally dehiscent; ovary usually 1(2)-loc­ Bot. Need. 16: 73-82. ular, free and superior or variously adnate to BURGER, W. 1977. Family #52, Moraceae. In W. fused to the perianth and inferior, often imbed­ BURGER, ed., Flora Costaricensis. Fieldiana, Bot. ded in the fleshy receptacle, stigmas 2 and usually 40: 94-215. linear, the single ovule pendulous. Fruits often Dioecious or monoecious trees, shrubs, or drupaceous and succulent due to the accrescent herbs, sometimes epiphytic, and rarely climbing, perianth, the individual drupes often coalescent usually with milky sap. Leaves alternate, simple, to form a compound infructescence, sometimes entire, toothed or rarely lobed, often 2-ranked; the whole infructescence enclosed and then ber­ venation pinnate, often with the basal veins pal­ ry-like. mate; petioles much shorter than the blade; stip­ Moraceae are world-wide in distribution, oc­ ules paired, free or fused, their scars completely curring in both temperate and tropical regions or partially encircling the stem. Inflorescences but are most diverse and conspicuous in the low­ axillary, often paired, highly variable as to form land tropics. The family contains over 1,000 (few-flowered racemes or spikes, condensed and species with approximately half ofthem in Ficus. involucrate discoid heads, globose heads with All four tribes of the family are represented at peltate scales to many-flowered fruitlike syconia La Selva. As treated here Moraceae does not with the flowers enclosed and the syconia usually include Cecropia, Coussapoa, or Pourouma, unisexual but sometimes with male and female which are instead dealt with in the family Ce­ flowers intermixed). Flowers unisexual, small, cropiaceae.

KEy TO THE GENERA

1. Stems herbaceous or suffiuticose at most, less than 50 cm tall; inflorescences bisexual (with male and female flowers intermixed), flat discoid structures about 2-3 cm in diameter...... 5. DORSTENIA. 1. Stems woody, 1-40 m tall; inflorescences unisexual or, if bisexual, then globose. 2. Leaves deeply pinnately lobed, often over 50 cm long by 30 cm wide; cultivated trees with green globose infructescences 20-30 cm in diameter...... 1. ARTOCARPUS. 2. Leaves unlobed or occasionally sagittate, smaller; not cultivated, fruits smaller. 3. Flowers and fruits enclosed in a more or less globose fruitlike, bisexual inflorescence; stipule scars completely encircling the stem; plants often epiphytic...... 6. FICUS. 3. Flowers and fruits exposed, the inflorescences unisexual (rarely bisexual); stipule scars completely encircling the stem or not; plants terrestrial. 4. Inflorescences involucrate (subtended by numerous basally attached, imbricate bracts), somewhat globose, discoid, or funnel-shaped and sometimes laterally flattened. 5. Sap resinous, clear yellow; infructescences globose, beset with large spinelike bracts...... 8. NAUCLEOPSIS. 5. Sap milky; infructescences discoid or funnel-shaped, lacking spinelike bracts. 6. Stipules fused and thus solitary at the node; leaf margin appearing denticulate from tufts of trichomes ...... 3. CASTILLA.

* ROBERT L. WILBUR, Flora Editor, Department of Botany, Duke University, Durham, North Carolina 27706. t Present address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299.

243 244 SELBYANA [Volume 9

6. Stipules free, paired at the node; leaf margin not denticulate from trichomes. 7. Leaf margin coarsely dentate apically, lower surface sparsely pubescent on the veins ...... 9. PEREBEA. 7. Leaf margins entire, the blades glabrous. 8. Anthers apiculate with a slender appendage 0.3-0.4 mm long; leaves small, the longest 6-12 cm long, 2-3 cm wide; pistillate flowers solitary; rare canopy tree 30 m tall...... 10. PSEUDOLMEDlA. 8. Anthers not apiculate; leaves large, mostly 17-35 cm long, 6.5-13 cm wide; pistillate flowers numerous, clustered on the receptacle; common treelets mostly less than 5 m tall ...... 7. MACQUIRA. 4. Inflorescences lacking an involucre (but see Trophis involucrata), globose, spicate or racemose. 9. Leaves, especially the juvenile, with erect uncinate trichomes; inflorescences (whether staminate, pistillate or hermaphroditic) globose to globose-flattened, covered (when young) with peltate bracts...... 2. BROSlMUM. 9. Leaves lacking uncinate trichomes; inflorescences otherwise (the staminate never globose), with or without peltate bracts. 10. Flowers (staminate) in spikes or racemes. 11. Peduncles with uncinate trichomes; perianth lacking, stamens arranged more or less in rows along the rachis and subtended by ligulate and sometimes peltate bracts ...... 4. CLARlSlA. 11. Peduncles with straight trichomes; perianth 4-parted; peltate bracts lacking. 12. Filaments incurved in bud, anther valves indehiscent in their lower third and thus remaining pouchlike when dry; tepals valvate in bud; stigmas linear, densely puberulent ...... 12. TROPHlS. 12. Filaments straight in bud, anther valves dehiscent to the base and laid out flat when dry; tepals decussate in bud; stigmas broadly triangular, glabrous...... 11. SOROCEA. 10. Flowers (pistillate) often solitary or paired. 13. Inflorescence with scattered peltate bracts on the peduncle; fruit surface smooth and glabrous...... 4. CLARISIA. 13. Inflorescence with deltoid bracts on the rachis; fruit surface covered with elongate tubercules, minutely pubescent ...... 12. TROPHIS.

1. ARTOCARPUS J. R. & J. G. A. Forst. 4 cm in diameter on a peduncle 3-6 cm long; pistillate inflorescences globose to ellipsoid cy­ Monoecious trees with milky sap. Leaves spi­ lindrical. Fruit syncarpic, green to yellow with ral and sometimes 2-ranked, simple, entire to acute pyramidal projections (10)15-30 cm in di­ deeply pinnatifid; stipules paired, leaving encir­ ameter; seeds present and then about 2.5 cm long cling, lateral or interpetiolar scars. Inflorescences and 1. 5 cm wide or lacking. unisexual, the flowers numerous and congested The breadfruit or "palo de pan" is an intro­ into large globose to cylindrical clusters, axillary duction from the South Pacific or Melanesia but or borne on old wood, paired or solitary. Sta­ now widespread in the neotropics. Several ma­ minate flowers with a single stamen with a straight ture trees grow along the disturbed woods along filament, the perianth tubular; pistillate flowers the Rio Puerto Viejo and about clearings. with the tubular perianth fused to those of neigh­ boring flowers. Fruit a syncarp ofachenes imbed­ ded in the flesh of the syncarp. 2. BROSIMUM Sw. Artocarpus is an Old World genus of about 48 species. Two species, the breadfruit and the jack­ Dioecious or monoecious trees; twigs of ju­ fruit, are widely cultivated in the neotropics. Only venile plants with uncinate trichomanes; sap the former is planted at La Selva. milky (at La Selva). Leaves distichous, entire, elliptic to oblong, petiolate; stipules free and 1. Artocarpns altms (Parkinson) Fosberg paired, the scars not completely encircling the (=A. communis J. R. & J. G. Forst.) stem (La Selva) or united, solitary, and com­ pletely encircling the stem. Inflorescences axil­ Tree to 29(35) m. Leaves usually deeply pin­ lary, unisexual or bisexual, globose to globose­ natifid with (l )3-7(9) lobes on each side, the blade depressed, covered in peltate bracts at least in 30-100 cm long and 25-65 cm wide, puberulent early stages, paired or solitary; staminate flowers to short pubescent both above and below; petiole with or without a usually 4-merous perianth, the 2-4(6) cm long; stipules 1.0-2.5 cm long, sparse­ filaments straight in bud; pistillate flowers usu­ ly to densely puberulent. Staminate inflores­ ally 1 per inflorescence, fused with the receptacle, cences a cylindrical head 7-30 cm long and 1.5- stigmas 2. Fruit drupaceous or baccate, SUf- 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 245 rounded by the accrescent and usually succulent cording to distributional evidence, La Selva plants receptacle, orange, red or green at maturity. probably belong to subspecies bolivarense (Pit­ Brosimum is a genus of 13 species distributed tier) Berg, which in Costa Rica is restricted to from Mexico to southern Brazil and also in the the Caribbean slope. However, these plants are West Indies. poorly collected so that little is known about in­ traspecific variation. KEY TO THE SPECIES 2. Brosimum guianensis (AubI.) Huber 1. Inflorescences more or less discoid, bisexual; fruits reddish at maturity; leaves silvery beneath...... Trees to 20 m tall, 60 cm in diameter; trunk ...... 2. B. guianensis. dark brown, large horizontal stipule scars reach­ 1. Inflorescences strictly globose, unisexual; fruits green ing nearly half-way around the trunk, branchlets or orange at maturity; leaves green beneath. 2. Stipules narrowly triangular, not prominently reddish. Leaf blades oblong-elliptic, 9-12 cm long, 3-4 cm wide, apically abruptly acuminate nerved, their scars encircling more than 1;' the stem; veins of the leaf scarcely elevated beneath; (acumen 0.5-1.5 cm long), the base acute; lateral mature fruits green...... 1. B. alicastrum. and reticulate veins plane on both surfaces, lat­ 2. Stipules broad at the base, prominently pal­ eral veins arising at an angle of65-75°, marginal mately nerved; their scars encircling less than 112 veins lying within 1 mm of margin; surface of the stem; tertiary veins of the leaf prominently blade dark green, glossy above, much paler, near­ elevated beneath; mature fruits orange...... ly white beneath; petioles 0.7-0.8 cm long, bear­ ...... 3. B. lactescens. ing small uncinate trichomes; stipules 3-4 mm long. Inflorescence bisexual, globose-flattened or 1. Brosimum alicastrum Sw. somewhat peltate 0.5-1.0 cm wide, mostly sta­ minate with I-several pistillate flowers sur­ Trees to 35(45) m tall, 1.5 m in diameter; trunk rounded by numerous staminate flowers, the pe­ with large thick buttresses; bark brown; sap milky duncle 0.7-1.5 cm long, the whole inflorescence but thin. Leaf blades elliptic, 19-20 cm long, covered with peltate bracts. Infructescence usu­ 7-8.5 cm wide, apically acuminate (acumen 1.5- ally with 1 fruit, 1 seed, reddish at maturity. 2 cm long), basally acute to rounded and slightly Flowering in late July. Southern Mexico south cordulate; lateral and reticulate veins paler than into Peru, Bolivia and Brazil. the lamina but only slightly raised; 14-16 pairs Brosimum guranense so far is known from only of major lateral veins, arising at an angle of 70- one mature individual at La Selva although nu­ 80°; petiole 0.7-1.0 cm long; stipules 8-11 mm merous saplings have been found throughout the long; mature plants nearly glabrous, sapling leaves property. This species is distinctive vegetatively and branchlets with small uncinate trichomes. because of the strikingly discolorous leaves: dark Inflorescence unisexual; the staminate (not seen green glossy above while nearly white beneath. at La Selva) globose, 3-8 mm in diameter; pe­ The small uncinate trichomes present on the pet­ duncles 7-10 mm long. Staminate flowers un­ ioles of mature leaves further characterize the organized, individual anthers intermixed with species. Its inflorescences bear both male and peltate scales over the surface of the receptacle; female flowers in contrast to the unisexual inflo­ functional pistillate flowers 1-2 per inflores­ rescences of the other two species at La Selva. cence. Fruit green and glaucous at maturity, suc­ culent, usually 1-seeded; seed embedded in a 3. Brosimum lactescens (T. Moore) C. C. Berg white somewhat pithy and succulent pericarp. Flowering in the dry season, February to March; Dioecious trees 30-40 m tall, 80 cm in di­ dropping mature fruits by April. Mexico south ameter; trunk light gray with small buttresses. into Brazil and Peru; West Indies. Leaf blades elliptic, 12-18 cm long, 5-7 cm wide, At La Selva Brosimum alicastrum is known apically acute to acuminate (rarely rounded), ba­ fromjust two individuals. Both produce fruit and sally acute to rounded or truncate; midrib and both are found in alluvial forest along the West lateral veins impressed above, prominent be­ River Road. The species is known locally as neath, tertiary veins also prominent beneath; lat­ "ojoche" and attracts many "agoutis," "pacas" eral veins 18-24, arising at an angle of 60-75°; and "sahinos" during the fruiting season. leaf surface somewhat scabrous beneath, nearly Brosimum alicastrum can be distinguished glaucous; petioles about 1 cm long; stipules 3- from the other two sympatric species by the large 10 mm long, puberulent and prominently veined. leaves with nonelevated venation. The plants at Inflorescences globose, the staminate about 1.0 La Selva differ from other collections and de­ cm in diameter, the pistillate (not seen at La scriptions of the species in that the fruits are Selva) about 2-6 mm in diameter, peduncles green rather than red or orange at maturity. Ac- bearing small uncinate trichomes; staminate 246 SELBYANA [Volume 9 flowers numerous and intermixed with peltate necting veins reaching to within 1 mm of margin; bracts; pistillate flowers l-several per head. Fruits margin appearing serrate with tufts oftrichomes, 1-3 per infructescence, I-seeded, bright orange upper surface shiny, distantly short pubescent and and succulent at maturity, astringent but sweet. scabrous, lower surface dull hirsute; petioles 1.0- Flowering in April to November; fruiting col­ 1.5 cm long; stipules 2-13 cm long, the margin lections from May. Central Mexico to Costa Rica white tomentellous, the scars diagonal. Inflores­ and down the east side of the Andes to southern cences discoid, the male about 1.8 cm in diam­ Brazil. eter, the female 1.5-2 cm in diameter. Fruits Widespread at La Selva but most common orange, the individual units ellipsoid, about 1 along the Rio Puerto Viejo. The fallen globose cm tall, united in their lower two-thirds, the ve­ staminate inflorescences are conspicuous in places lutinous accrescent perianth becoming orange at along the East River Road and Far Loop trails maturity; seeds surrounded in orange succulent during flowering periods. Plants appear to come flesh. Flowering and fruiting March to July. Hon­ into flower two times a year and mature fruits duras south into Colombia. about six months later. Leaves of saplings of this Known at La Selva only from riverside forest species are nearly three times the size of leaves and secondary woods along the Rio Puerto Viejo, on adult plants and are covered with small un­ where it is quite common. The long arching cinate trichomes. branches with large, more or less 2-ranked leaves are distinctive. The self-pruning branches result 3. CASTILLA Sesse in a straight, unbranched bole topped by a wide, relatively short crown. This feature (cladoptosis) Usually dioecious trees with self-pruning has been considered important in circumscribing branches and buttresses; sap milky and copious. the tribe Castilleae. Although staminate inflo­ Leaves 2-ranked, often large, somewhat scabrous rescences may sometimes accompany pistillate above, pubescent; tertiary venation more or less ones in this species, all of our collections are parallel; petioles short; stipules connate, com­ unisexual. The plants at La Selva belong to sub­ pletelyencircling the stem. Inflorescences axil­ species costaricana (Liebm.) C. C. Berg, restrict­ lary, discoid to funnel-shaped, sometimes lat­ ed to the Caribbean slope in Costa Rica and erally folded, subtended by numerous imbricate apparently isolated from the typical variety, which bracts; staminate inflorescences often paired, in Costa Rica has. been reported only from the conduplicate, reniform, pedunculate, those found Nicoya Peninsula. in pistillate plants smaller and funnel-shaped, the During the late 1800's and early 1900's this flowers with no definite organization, stamens species was economically important in the rub­ paired or solitary on the receptacle, interspersed ber industry and was at that time widely intro­ with bracts; pistillate inflorescences usually sol­ duced throughout the tropics. It has since be­ itary but often accompanied by small staminate come naturalized in many areas outside its natural inflorescences, discoid to globose-flattened, ses­ range. Latex was collected commercially from sile, the flowers with a 4- or 5-10bed perianth, plants in the La Selva area by local workers as the ovary either partly united with the perianth recently as the 1940's. (La Selva species), free, or immersed in the re­ ceptacle. Fruit drupaceous, united basally by the 4. CLARISIA Ruiz & Pavon accrescent perianths, the infructescence globose­ flattened. Dioecious trees, trunk narrowly buttressed, Castilla is a genus of three species, one re­ bark dark gray, sap milky. Leaves entire; stipules stricted to the Amazon basin and the other two paired, usually deciduous, encircling less than occurring in both Central and South America. half the stem. Inflorescences axillary or cauliflo­ This is the only genus of Moraceae at La Selva rous, usually paired; staminate inflorescences with completely fused stipuies. spicate, the numerous stamens densely crowded and irregularly interspersed with bracts and peri­ 1. Castilla elastica Sesse anth parts; pistillate inflorescences axillary, usu­ ally 2-flowered, the pedicel bearing several pel­ Trees 25-35 m tall but often fertile when tate bracts; ovary unilocular and uniovular. smaller; trunk gray; twigs densely appressed pu­ Fruit a drupe, the accrescent perianth becoming bescent with golden trichomes. Leafblades ob­ red and succulent. long, widest above the middle, 28-40 cm long, As is true for many tall trees of the wet Carib­ 13-20 cm wide, apically acute, basally shallowly bean lowlands, the genus Clarisia is poorly rep­ cordate to truncate; midrib and lateral veins plane resented in herbaria. Collections from La Selva above, prominent beneath, lateral veins 20-24 apparently represent the first staminate material pairs arising at an angle of 50-70°, loop con- from southern Central America. Clarisia is a ge- 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 247 nus of two or three species ranging from Mexico 1. Leaves broadly deltoid and deeply lobed; inflores­ to the Amazon Basin. One species with two sub­ cence margin lobed; peduncles 15-20 cm long. .. species occurs in Central America...... 2. D. contrajerva.

1. Clarisia biftora Ruiz & Pavon 1. Dorstenia choconiana S. Wats. Trees to 30 m tall; the largest roots reddish. Herbs to 40 cm tall. Leaf blades elliptic 9- Leaf blades oblong, often widest towards the tip, 17.5 cm long, 3.5-7.5 cm wide, occasionally very 13-22 cm long, 4.5-7 cm wide, the apex abruptly slightly lobed, often toothed in the distal half, acuminate with an acumen 1-2 cm long, basally apically broadly acuminate, basally acute to acute; midrib and lateral veins plane to somewhat truncate, rarely somewhat cordate, the blade impressed above, prominent and drying lighter shortly and narrowly decurrent; midrib and lat­ below; lateral veins 12-14 pairs arising at an eral veins plane above, prominent below; main angle of 60-65°; leaf surface dark green above, lateral veins 7-8 pairs, arising at an angle of 50- paler below, glabrous; petioles about 1 cm long. 70°; surface of blade glabrous above, scaberulous Stipules 2-3 mm long. Staminate inflorescences and minutely pubescent on the veins below; pet­ spicate, 3-4.5 cm long, paired and axillary to iole 1.5-5 cm long; stipules paired, acuminate, somewhat scattered along the internode; pedun­ bilobed with one lobe longer, 2-4 mm long. In­ cle 2-4 mm long, covered with minute uncinate florescence solitary, flat-topped to somewhat cup­ trichomes. Pistillate inflorescences of two axil­ shaped, elliptical, 1.5-2.5 cm in diameter; pe­ lary flowers having a narrowly globose or ovoid duncle 3-6 cm long. Seeds white, about 2 mm perianth 2.5-5 mm long. Fruits ovoid, 1.5-2 cm in diameter, trigonate, papillate. Probably flow­ long, becoming red and succulent. Flowering in ering and fruiting throughout the year. late August, fruit mature by June. Central Mex­ Dorstenia choconiana is uncommon at La Sel­ ico south to Bolivia and Brazil. va and restricted to somewhat disturbed but shady At La Selva this species is known only from alluvial sites. A small population grows in the the two individuals in the arboretum (both pis­ trail on the far side of the bridge over the Q. EI tillate) and a tagged staminate individual at the Salto on the East River Road. base of the steps behind the washroom at the At La Selva the leaf margins ofthis species are original dormitory area. Arboretum tree no. 819 sometimes coarsely toothed in the distal half. produced fruit in June of 1981 while it was less The form with pinnately lobed or incised leaves than 4 m tall. The nearby 30 m tall mature in­ is not knoW!} from La Selva. dividual was also producing fruit at that time. Uncinate trichomes, present on the staminate 2. Dorstenia contrajerva L. peduncles, were not recorded previously for the genus Clarisia. Herbs, acaulescent or from a short tuberous stem; leaves clustered at the top of the tuber. Leaf blades broadly deltoid (7) 15-20(26) cm 5. DORSTENIA L. long, (9)15-20(34) cm wide, basally 3-10bed, the Perennial, rhizomatose herbs, with milky sap lobes themselves often deeply lobed and dis­ scant. Leaves alternate, simple, entire to deeply tantly toothed, apically acuminate, basally trun­ lobed, stipules paired and free. Inflorescence on cate, cordate to hastate, and the blade usually a long peduncle, the male and female flowers decurrent; midrib and lateral veins plane to sessile, intermixed, and exposed on a flat recep­ somewhat impressed above, prominent below; tacle. Fruit achene-like, sometimes explosively main lateral veins 3-5 pairs, often palmate with released from the receptacle. 3-5 main veins, arising at an angle of 40-60°; Dorstenia is pantropical and contains a total petioles (8) 15-20(26) cm long; stipules triangu­ of about 50 species most of which are found in lar, about 5 mm long, usually persistent and Africa and South America. Five species are sheathing the top of the rhizome. Inflorescence known from Central America. Although African with a much lobed and often purplish margin, members of this genus are woody, the genus is 1.5-3 cm wide; peduncle (10)15-20(34) cm long. otherwise unusual in the Moraceae because of its Seeds about 1.5 mm in diameter. Probably fertile herbaceous habit. throughout the year. Mexico south to northern South America and from the West Indies. Although Dorstenia contrajerva is often very KEy TO THE SPECIES common throughout its range in areas similar to 1. Leaves elliptic, entire or slightly lobed or toothed; the abandoned cacao plantation at La Selva, it inflorescence margin entire; peduncles 3-6 cm long. is known here from only one small clump ofthree ...... 1. D. choconiana. to four plants. These plants grow at the base of 248 SELBYANA [Volume 9 the large Leuhea tree at the boat landing below first but becoming succulent, juicy, and in some the original station. This area is frequently flood­ species red or purple at maturity. ed during the rainy season and the population is Ficus is a large pantropical genus of 500 or very likely ephemeral. more species with its largest center of diversity in southeast Asia. In Flora Costaricensis 39 na­ 6. FICUS L. tive species were distinguished by Burger. Only eight of these are apparently restricted to Costa DEWOLF, G. P., JR. 1960. Ficus. In R. E. WOODSON Rica or Costa Rica and Panama, whereas the AND R. F. SCHERY, eds., Flora of Panama. Am. remainder are widespread species found Missouri Bot. Gard. 47: 146-165. throughout Central America with some ranging STANDLEY, P. C. 1917. The Central American species into the West Indies and South America. of Ficus. Contr. U.S. Nat!. Herb. 20: 1-35. At La Selva only one species, Ficus insipida, Trees or shrubs or rarely (at La Selva) lianas, is conspicuous. It is the dominant tree in several often beginning as epiphytes and later rooting in areas along the Rio Puerto Viejo, and belongs to the ground and eventually "strangling" the sup­ the subgenus Pharmacosycea. The members of porting tree; sap usually milky. Leaves alternate; subgenus Pharmacosycea typically do not grow stipules paired, covering the shoot apex, usually as epiphytes, have only one fig per node, and deciduous, their scar completely encircling the only two bracts subtending each fig. Twelve of stem. Inflorescence a syconium, the flowers en­ the 16 species at La Selva belong to subgenus closed in a sessile or pedicellate, globose fruitlike Urostigma characterized by its often epiphytic receptacle (the fig), subtended by 2 or 3 bracts habit, two figs per node and three bracts sub­ and opening only at a small apical pore (ostiole). tending each fig. Several species not treated here, Flowers small, numerous. and unisexual, both including F. brevibracteata, F. macbridei, F. tri­ pistillate and staminate borne in the same fig, gonata, and F. turrialbae, known tentatively only interspersed with bracts. Fruits small achenes held from sterile material at La Selva or from their in the berry-like fig, the figs hard and green at distributional ranges, are to be expected.

KEy TO THE SPECIES 1. Figs solitary at the nodes, subtended by 3 bracts; plants usually free standing trees, seldom beginning as epiphytes. 2. Figs sessile; leaves broadly elliptic, the widest 12-17 cm wide ...... 15. F. tonduzii. 2. Figs pedicellate; leaves narrower, the widest 6-9 cm wide. 3. Epidermis of petioles and branchlets cracked and flaking; figs and leaves scabrous. 4. Leaves and figs glabrous; leaves flat; basal bracts all basal; branchlets reddish. . .. 10. F. maxima. 4. Leaves and figs pubescent; leaves bullate; 1 of the 3 basal bracts placed on the surface of the fig above its middle; branchlets grayish tan...... 8. F. glaucescens. 3. Epidermis of petioles and branchlets not markedly exfoliating; figs and leaves smooth. 5. Leafblades obovate, apically rounded to broadly acute, the largest 12-15 cm long; loop connections as prominent above as the lateral veins; very rare epiphyte, in the forest along streams...... 5. F. crassivenosa. 5. Leaf blades narrowly elliptic, apically acuminate, the largest 20-25 cm long; loop connections obscure; common tree along the river...... 9. F. insipida. 1. Figs usually paired at the nodes, subtended by 2 bracts; plants usually epiphytic, sometimes strangling and becoming free standing, rarely entirely terrestrial. 6. Figs sessile or the pedicels less than 0.2 mm long. 7. Stipules and basal bracts pubescent (trichomes visible without magnification). 8. Figs longer than wide, conspicuously pubescent; leaves scabrous above...... 13. F. popenoei. 8. Figs globose to globose-flattened, glabrous; leaves not scabrous above. 9. Leafblades small, seldom over 8 cm long, with only 2, or more often, 3-4 pairs oflateral veins, the lowest pair extending nearly to the middle of the blade ...... 3. F. colubrinae. 9. Leaf blades larger, usually over 12 cm long, with 7-12 pairs oflateral veins, the lowest pair not extending above the lower quarter of the blade (see also Ficus velutina with short peduncles 2-6 mm long and minutely pubescent figs). 10. Basal bracts conspicuous, often longer than the figs; petioles hirsute .... 4. F. costaricana. 10. Basal bracts small and hidden beneath the figs; petioles glabrous but their epidermis red and flaking...... 7. F. dugandii. 7. Stipules and basal bracts glabrous or only minutely pubescent. 11. Mature figs 0.3-1.5 cm in diameter; leaves elliptic. 12. Plants slender branched epiphytic shrubs or lianas; figs 0.3-0.5 cm in diameter; often more than 2 per node ...... 14. F. schippii. 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 249

12. Plants robust free standing trees; figs 0.8-1.3 cm in diameter, only 2 per node ...... 7. F. dugandii. 11. Mature figs 1.5-2.5 cm in diameter; leaves oblong or cordate. 13. Leaves oblong, not glaucous beneath; petioles less than 3 cm long; stipules persistent and conspicuous; figs glabrous...... 1. F. cahuitensis. 13. Leaves broadly cordate, glaucous beneath; petioles more than 5 em long; stipules deciduous; figs minutely velvety pubescent...... II. F. nymphaeifolia. 6. Figs with more or less conspicuous peduncles 0.2-1.5 cm long. 14. Leaves ovoid, apically rounded to broadly acute, the margins revolute; reticulate venation very prominent beneath; petioles and stipules brownish tomentose...... 16. F. velutina. 14. Leaves elliptic to oblong, apically acuminate, the margins flat; reticulate venation obscure; petioles and stipules glabrous or minutely pubescent. 15. Largest leaves 15-20 cm long, oblong and abruptly narrowed to an acumen; major lateral veins of the blade very distinct from the intermediate veins; petioles 2.5-6 em long...... 2. F. citrifolia. 15. Largest leaves 10-13 em long, elliptic and gradually acuminate to the apex; major lateral veins usually difficult to distinguish; petioles 0.5-2.5 em long. 16. Stipules and leaves minutely pubescent; leaves narrowly lanceolate, often 3-5 times longer than wide; figs 0.5-0.9 cm in diameter...... 6. F. donnell-smithii. 16. Stipules and leaves glabrous; leaves elliptic, less than 3 times longer than wide; figs 1-1.4 em in diameter...... 12. F. pertusa.

1. Ficus cahuitensis C. C. Berg in the canopy. Its long spreading branches and long oblanceolate leaves clustered near the tips Epiphytic shrub, sometimes becoming quite of branches make it distinctive even from a dis­ large, the branches long and spreading, leafy tance. branches about 1 cm diameter with the leaves This species was described since Burger's treat­ usually clustered distally. Leafblades oblanceo­ ment for Flora Costaricensis. It had been con­ late or oblong-obovoid, 20-30 em long, 7-11 cm fused with Ficus obtusifolia from which it differs wide, apically broadly acute or rounded, basally in its persistent stipules and nearly glabrous figs. acute to acuminate; midrib and lateral veins Although F. obtusifolia is not known from La prominent on both surfaces but more so beneath; Selva, it is to be expected. Vegetatively F. ca­ principal lateral veins 8-12 pairs, faintly loop­ huitensis is quite different from F. nymphaeifo­ connected; tertiary veins indistinct above except lia, their figs are very similar suggesting that they slightly raised when dry, distinct beneath, con­ are more closely related to each other than to trasting in color with the darker lamina; surface any other species at La Selva. of blade glabrous, glossy dark green above, pale and dull beneath, glabrous except for minute ap­ pressed scalelike trichomes along the principal 2. Ficus citrifolia P. Mill. veins; petioles 2-2.5 em long; stipules narrowly triangular, 3-5.5 cm long, 1.5-2.5 cm wide, dark Epiphytic shrub or elsewhere small trees; sap brown, persistent. Figs 2 per node, globose and clear; branchlets with reddish exfoliating epi­ somewhat flattened, 1-1.7 cm tall, 1.7-2.3 cm dermis. Leaf blades oblong-elliptic, 11-20 cm in diameter, minutely pubescent (elsewhere) to long, 6-9.5 cm wide, the apex rounded with an glabrous (La Selva) except for minute, appressed abrupt short acumen 0.5-1.5 cm long, basally scalelike trichomes, sessile; basal bracts semicir­ rounded and minutely cordulate; midrib and lat­ cularor broadly triangular, 0.7-1.3 cm long, 0.7- eral veins prominulous above, slightly more 1.4 cm wide, minutely pubescent (trichomes 0.1 prominent beneath, drying whitish above, red­ mm long) on the outer surface; ostiole, 4-5 mm dish beneath; major lateral veins 12-14 pairs, wide, slightly raised when dry. Achenes ovoid arising at an angle of 60-65°, distinctly loop­ and more or less flattened laterally and with a connected; plants glabrous throughout; petioles sagittal furrow, 1.8-2 mm long. Collected with 2.5-6 cm long, deeply grooved above; stipules figs August to December. Caribbean lowlands of about 1 cm long. Figs usually 2 per node, ob­ Costa Rica. ovoid, 0.8-1.1 em long, peduncle 0.6-0.7 cm Common and conspicuous in the La Selva area long; basal bracts 3, about 0.2 cm long; ostiole and elsewhere in the Caribbean lowlands, espe­ flat. Achenes ovoid and slightly triquetrous, about cially in open areas where often found on the 1.3 mm long. Nearly mature figs collected in trunks of large trees below the lowest branches. April. Southern Florida through Central Amer­ In the forest at La Selva the species is found most ica south into Paraguay; West Indies. often near the river and there often grows high Wide ranging but very rare at La Selva where 250 SELBYANA [Volume 9 known from just one individual. It should be trichomes. Figs usually 2 per node, globose but looked for, particularly in areas of the new prop­ slightly wider than long, 1-1.2 em in diameter erty along the Rio Sarapiqui and Rio Peje. This at maturity, sessile; basal bracts broadly trian­ species might be mistaken for a large-leaved form gular, 0.5-0.8 em long with golden tan, ap­ of Ficus pertusa but the clear sap, long petioles pressed pubescence covering the figs in early and peduncles are distinctive. stages; ostiole conical, about 1 mm high. Achenes globose, 0.9 mm in diameter. Specimen with im­ 3. Ficus colubrinae StandI. mature figs collected in June. Guatemala south into Ecuador. Epiphytic shrub or treelet, rarely becoming free Known from only one fertile collection at La standing; branchlets hirsute and the tan epider­ Selva made from a tree along the Rio Sarapiqui mis exfoliating. Leafblades obovate-elliptic, 8- near the river crossing on the new property. This 1O.S em long, 4-S.5 em wide, apically acute, ba­ species has been rarely collected below 300 m sally acute to rounded; midrib and lateral veins elevation. obscure above, prominent beneath; principallat­ eral veins 3-4 pairs, distinctly loop-connected, 5. Ficus crassivenosa W. Burger the upper ones arising at an angle of 50-S5°, the lower pair at a much narrower angle (30°); leaf Epiphytic trees or shrubs; epidermis of surface glabrous above, sparsely appressed pu­ branchlets reddish and flaking. Leafblades ob­ bescent beneath especially along the veins, tri­ long-obovate, 11.5-15 em long, 6-7.5 em wide, chomes 0.5-1 mm long; petioles 1-3.S em long, apically broadly acute or occasionally rounded, pubescent with tan, woolly and matted tri­ the base acute; midrib and lateral veins promin­ chomes; stipules 0.8-2.2 em long, pubescent with ulous on both surfaces but more conspicuous brownish appressed trichomes about 1 mm long. beneath; principal lateral veins 13-16 pairs, aris­ Figs usually 2 per node, 0.8-1 em in diameter, ing at an angle of 80-90°, loop-connected and sessile; basal bracts about 1.3 mm long. Achenes forming a definite submarginal vein; tertiary veins ovoid, 0.9-1 mm long. Mature figs collected from obscure; surface of blade glabrous throughout; February through August. Guatemala south to petioles 2-2.S em long; stipules 1.5-2.5 em long, Central Panama. glabrous. Figs usually 1 per node, globose, 2-2.5 Very common at La Selva where often found em in diameter at maturity; peduncle 1.S-2 em a few meters above the ground on large trees long; basal bracts broadly triangular, about 1.5 along the river and in trees left standing in pas­ em long; ostiole slightly raised, 1 mm in diam­ tures. Ficus colubrinae is easily recognized by its eter. Achenes ovoid to reniform about 2 mm small leaves with relatively few lateral veins and long. Fertile specimens collected August to No­ tawny pubescence. The only other sympatric Fi­ vember. Caribbean lowlands of Costa Rica. cus with a similar pubescence and sessile figs is Reported in Flora Costaricensis from only three F. costaricana, which, however, has much larger, collections. Intensive collecting at La Selva has many-veined leaves. disclosed only two individuals in primary forest along the Sabalo-Esquina trail. Ficus crassive­ 4. Ficus costaricana (Liebm.) Miq. nasa is recognized by its closely parallel lateral veins, which are often perpendicular to the mid­ Tree (8)25(30) m tall, beginning as epiphyte rib. and strangling the host plant; trunk reddish brown. Leafblades oblong-elliptic, 13-17.S em 6. Ficus donnell-smithii StandI. long, S.S-7 em wide, apically acute to rounded, the base narrowly rounded and cordulate; midrib Epiphytic shrubs or rarely trees; branchlets mi­ and lateral veins prominent on both surfaces; nutely puberulent. Leaf blades narrowly lanceo­ principal lateral veins 10-12 pairs, arising at an late to oblong-elliptic, 9-11 em long, 2.5-3.S em angle of 55-60°, distinctly loop-connected near wide, apically acuminate, the base narrowly the margin; tertiary venation conspicuous, es­ rounded to acute, rarely truncate; midrib and pecially beneath; surface of blade glabrous above, lateral veins prominulous on both surfaces; prin­ flecked with minute rusty scales beneath, pubes­ cipal lateral veins 6-8( I 0) pairs, arising at an cence (at La Selva) restricted to a few scattered angle of 65-75°, faintly loop-connected; tertiary trichomes (ca. 0.8 mm long) on the midrib and veins obscure; both surfaces of blade glabrate; along either side, or elsewhere minutely pubes­ petioles 1.2-2 em long, usually minutely pubes­ cent throughout; petioles 2-4.S em long, dis­ cent; stipules small, O.S-1 em long. Figs usually tantly hirsute, trichomes about 1 mm long; stip­ 2 per node, 0.8-1 em in diameter, minutely pu­ ules more or less persistent, 1.5-2 em long, berulent; peduncle 0.5-0.6 em long; bracts densely appressed pubescent with golden brown broadly triangular, about 1 mm long; ostiole 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 251

plane. Achenes about 0.9 mm long, greenish in apically acute to rounded but abruptly acuminate color. Fertile collections from April to Septem­ with the acumen 0.5-1.5 cm long, basally acute; ber. Guatemala south into Costa Rica. midrib and lateral veins prominent on both sur­ Apparently restricted to the Caribbean low­ faces but more so beneath and the lamina bullate; lands in Costa Rica. In the Flora Costaricensis principal lateral veins (7)8-9(10) pairs, arising at Burger knew it from just two collections includ­ an angle of 50-80°, the lowest pair arising at a ing one from La Selva. It is actually quite com­ much narrower angle than those immediately mon in the area, where it is found both in open, above and paralleling the leaf margin, loop-con­ more or less secondary habitats as well as in the nected near the margin; reticulate venation ob­ forest canopy. The long narrow juvenile leaves scure above, very prominent beneath; surface of are especially distinctive. The only other fig at blade smooth, shiny and glabrous above, sca­ La Selva that resembles Ficus donnel/-smithii is brous, dull and pubescent with distant, stifferect F. pertusa, which has broader leaves, larger figs, trichomes 0.5 mm long beneath; petioles 2.5-4 and is glabrous throughout. cm long, gray tan and flaking; stipules 1-1.5 cm long, glabrous. Figs 1 per node, globose, 2.5-3 7. Ficus dugandii StandI. cm in diameter at maturity, pubescent with short, stiff, distant trichomes 0.3-0.4 mm long, sca­ Tree 25-30 m tall; trunk dark gray; branchlets brous; ostiole slightly conical; basal bracts 3, small reddish and the epidermis exfoliating, deeply fur­ triangular, about 1 mm long, 1 of the bracts al­ rowed on drying. Leaf blades oblong-elliptic, ways fused to the surface of the fig about half 15-18 cm long, 6.5-8.5 cm wide, apically broad­ way to the ostiole; peduncle about 1 cm long, ly acute but abruptly acuminate with the acumen puberulent. Achenes reniform, about 2 mm long. 0.5-1.5 cm long, basally rounded to acute; mid­ Bearing figs June and July. Mexico south into rib and lateral veins raised on both surfaces in northern South America. fresh material, plane or slightly impressed above Known from low areas in the primary forest in dried material, lighter colored than the lamina, understory at La Selva. The bullate, pubescent principal lateral veins 9-13 pairs, arising at an and scabrous leaves with very prominent retic­ angle of 60-70°, loop-connected (sometimes ulate venation on the lower surface, and espe­ faintly so) near the margin; tertiary veins indis­ cially the unusual position of one of the basal tinct above, slightly more conspicuous beneath; bracts of the figs, are distinctive. surface ofblade glabrous above, occasionally with Although Ficus glaucescens (as well as many few scattered appressed trichomes along the mid­ other names) was considered synonymous with rib beneath, veins of the lower surface traced by F. maxima in the Flora of Panama, plants here abundant minute (0.1 mm) translucent scalelike described differ even from that broad concept of trichomes; petioles 4-5.5 cm long, reddish and F. maxima in their consistently "misplaced" flaking especially towards the summit; stipules basal bract, the very prominent tertiary venation, 1-1.5 cm long, densely grayish to tan appressed the bullate leaves, and the gray tan rather than pubescent, with the trichomes about 2 mm long. reddish brown flaking periderm. This species was Figs paired at the nodes, globose to oblate, 1- not included in Flora Costaricensis. 1.5 cm wide, 0.8-1 cm wide; basal bracts broadly triangular, 2-3 mm long; ostiole flat but with 9. Ficus insipida Willd. slightly raised ring oftissue. Achenes ovoid, about 1.2 mm long. Collected with nearly mature figs Tall trees, 15-30 m tall with long drooping in June. Costa Rica south into Ecuador. branches; the bark gray. Leafblades oblong-el­ Known from only two individuals in primary liptic, 19-22 cm long, 5-7 cm wide, apically nar­ forest at La Selva and from very few collections rowly acute to acuminate, basally acute; midrib throughout its range. Ficus dugandii is the only and lateral veins prominulous on both surfaces; member of the subgenus Urostigma at La Selva principal lateral veins 19-24 pairs, arising at an that apparently does not begin life as an epiphyte. angle of 70-80°, faintly loop-connected near the This species resembles F. citrifolia in leaf shape margin; tertiary veins obscure on both surfaces; and venation but the latter has pedunculate figs surface of blade glabrous; petioles 4-5 cm long; and gray rather than reddish branchlets. Both stipules 6-10 cm long. Figs globose, 3-4.5 cm species are very rare at La Selva. in diameter, raised on a short stipe 0.5-1 cm long above the basal bracts; pedicel 1-2 cm long; 8. Ficus glaucescens (Liebm.) Miq. basal bracts small, broadly triangular, about 1 mm long; ostiole flat. Achenes reniform, about Buttressed trees 10-25 m tall, 50 cm dbh; the 2.7 mm long. Producing figs at least from June branchlets tan gray with flaking periderm. Leaf through October. Mexico south into Paraguay. blades elliptic, 16-20 cm long, 6-10 cm wide, Ficus insipida is the most conspicuous fig at 252 SELBYANA [Volume 9

La Selva. It grows only along the river and is 14-24 cm long, 13-20 cm wide, apically broadly particularly abundant in certain low sandy areas acute to rounded with an acumen 1-1.5 cm long, such as just upriver from Rafael'S former cabin. the basal lobes 2.5-6 cm long, occasionally over­ Burger reported that this species probably hy­ lapping; midrib and lateral veins prominulous bridizes with F. maxima. A number of possible on both surfaces, strongly contrasting in color hybrids between these species are known from with the lamina, principal lateral veins 7-10 pairs La Selva (see account of Ficus maxima). (2-3 pairs palmate at the base), arising at an angle of 55-75°, faintly loop-connected about 0.5 cm 10. Ficus maxima P. Mill. from the margin; tertiary veins obscure; surface of blade glabrous, glaucous beneath; petioles 8- Trees or spreading shrubs, 5-15 m tall; branch­ 11 cm long; stipules 1-2 cm long, glabrous. Figs lets dark reddish brown with exfoliating peri­ 2 per node, sessile, globose, 1.5-2.5 cm in di­ derm. Leaf blades oblong-elliptic, 16-32 cm ameter, velvety pubescent; ostiole large, raised long, 7-12 cm wide, apically broadly acute to on a ring 5-7 mm in diameter, the flaps broadly rounded and abruptly acuminate with a short triangular, 2-3 mm long; basal bracts ovoid, wid­ acumen 0.5-1.5 cm long, basally acute; midrib er than long, about 3-5 mm long, 7-8 mm wide. and lateral veins prominent on both surfaces but Achenes ovoid, 1.3-l.7 mm long. Fertile speci­ especially so beneath; principal lateral veins 7- mens collected in April and June. Costa Rica 11 pairs, arising at an angle of 60-70°, the lowest south into northern South America. pair much more sharply ascending than the oth­ Known from very few individuals at La Selva ers, loop-connected 1-3 mm from the margin; where found only as an epiphyte. Even with only tertiary veins obscure above, prominulous be­ vegetative material Ficus nymphaeifolia cannot neath; upper surface glabrous, lower surface sca­ be confused with any other in the area. The large brous; petioles 2.5-5 cm long, dark reddish brown cordate leaves with long petioles are distinctive. with flaking periderm; stipules 1.5-2.5 cm long, The figs alone are also sufficient to identify this glabrous. Figs 1 per node, globose, about 2-2.5 species; it is the only fig at La Selva with densely cm in diameter, scabrous, sparsely pubescent with velvety short pubescent figs. trichomes about 0.2 mm long; ostiole flat; basal bracts 3, broadly triangular, about 1 mm long, 12. Ficus pertusa L.f. very rarely 1 of them attached to the surface of the fig a short distance from the base; peduncle Epiphytic shrub or sometimes elsewhere a tree; 1-2 cm long, minutely pubescent apically. branchlets gray. Leafblades elliptic, 9.5-13 cm Achenesreniform, about 2.5 mm long. Producing long, 3.5-4.8 cm wide, apically acuminate, ba­ figs at least from March through August. South­ sally acute; midrib and lateral veins prominulous ern Mexico south into the Amazon Basin; West on both surfaces when dry, principal lateral veins Indies. (7)8-11 pairs, very faint and difficult to distin­ At La Selva known only as a spreading shrub guish from the tertiary venation, arising at an or low tree growing either in open swamps or angle of (50)70-80°, loop-connected near the near the river. Burger reported a number ofpos­ margin; surface of blade glabrous, usually drying sible hybrids between Ficus maxima and F. in­ brownish; petioles 1.5-2 cm long; stipules 0.5- sip ida. Two collections from La Selva (Grayum 1.3 cm long, glabrous. Figs 2 or sometimes only 2200, Hammel & Trainer 10892) also appear to 1 per node, globose, about 1.2 cm in diameter, be of hybrid origin. These specimens have the red to dark purple (rarely green) at maturity; 05- reddish flaking periderm and slightly prominu­ tiole about 2 mm in diameter, sunken within a lous, reticulate venation of F. maxima but the ring of tissue; basal bracts 2, broadly ovate-tri­ leaves are only slightly scabrous, the stipules are angUlar, about 2 mm long, glabrous or minutely much larger than those described for F. maxima pubescent. Achenes nearly globose to slightly re­ and shorter than those of F. insipida. The fig size niform, 1 mm long. Producing figs mostly in the and number of lateral leaf veins are also inter­ dry season. February to May. Mexico south into mediate. This possible hybrid individual is a southern Brazil. large-crowned but low tree growing at the edge Common at La Selva and apparently restricted of the Rio Puerto Viejo on the point of land at to more or less open areas along the Rio Puerto the junction with the Rio Sarapiqui. Viejo or to the edges of open swamps. Ficus pertusa is characterized by its usual gla­ 11. Ficus nympbaeifolia P. Mill. brosity, small leaves, small, usually paired figs with a sunken ostiole, and indistinct lateral veins Epiphytic shrubs or trees, sometimes becom­ that are nearly perpendicular to the midrib. Some ing free standing elsewhere. Leaf blades cordate, individuals of this species (e.g., Hammel 9685) 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 253 have leaves that dry green with fewer and more usually arising at a much narrower angle, the distinct lateral veins arising at a narrower angle, veins distinctly loop-connected 2-5 mm from and figs that are green at maturity and occur only the margin; tertiary veins obscure above, lighter one per node. This form also occurs on Barro colored than the lamina beneath; both surfaces Colorado Island in Panama and as far north as of blade glabrous but often slightly scabrous in Guatemala. Further study of F. pertusa from fresh material; petioles 2.5-4.5 cm long, often throughout its range is indicated. with flaking epidermis; stipules 1.5-2.5 cm long. Figs globose, paired or clustered at the nodes, 13. Ficus popenoei StandI. glabrous, often sessile, 4-8 mm in diameter; os­ tiole flat, 1 mm in diameter; basal bracts broadly Epiphytic trees, the branchlets with rusty tan triangular-rounded, about 1 mm long, glabrous; pubescence. Leaf blades oblong-obovoid, wid­ peduncle 0-1 mm long. Achenes ovoid to tur­ est above the middle, 10-18 cm long, 6-9 cm binate, 0.6-0.8 mm long. Producing figs mostly wide, apically obtuse to rounded, basally trun­ during the dry season, February to May, but also cate, rounded or cordate; midrib and lateral veins some collections from June to August. Caribbean prominent on both surfaces, more so beneath, slopes of Central America from Belize south into rusty tan pubescent with erect trichomes 0.5-1 Panama and northwestern South America. mm long; principal lateral veins 9-11 pairs, aris­ Occasional in trees along the river and near ing at an angle of 65-75°, distinctly loop-con­ swamps. Ficus schippii is the only species at La nected 3-8 mm from the margin; tertiary veins Selva, and perhaps the only one in Central Amer­ prominulous but indistinct above, prominent and ica, that begins life as a root-climbing liana. Al­ contrasting sharply with the lamina beneath; sur­ though it reportedly becomes a free standing tree face ofblade scabrous above, pubescent with erect elsewhere, the species is known only as a climber trichomes 1-2 mm long beneath; petioles 0.5- at La Selva. Its small, clustered, more or less 2.5 cm long; stipules 0.5-0.7 cm long, with ap­ sessile figs, its habit, and its slightly scabrous pressed rusty tan pubescence. Figs oblong-ovoid, leaves are all distinctive. 2-2.5 cm long, 1.3-1.5 cm wide, densely rusty brown pubescent with trichomesO.5-1 mmlong; 15. Ficus tonduzii StandI. ostiole slightly raised, about 1 mm in diameter; basal bracts semicircular, 2 mm long, 3-4 mm Understory trees to 20 m tall, the trunk light wide; peduncle 0.2-0.3 cm long, pubescent. gray; branchlets with light grayish tan flaking Achenes ovoid and somewhat flattened, mostly periderm. Leaf blades oblong-elliptic, 23-33 cm 1 mm long. Apparently producing figs during the long, 11-17 cm wide, apically acute to rounded, dry season, those ofthe one fertile collection found basally acute; midrib and lateral veins plane and on the ground by May. Belize south into Colom­ indistinct above, prominent and contrasting with bia. the lamina beneath; principal lateral veins 9-13 Known from just two individuals at La Selva, pairs, arising at an angle of75-85°, very distinctly one from the gravel bar along the Rio Sarapiqui loop-connected into a submarginal vein 2-4 mm at the end of A venida Marafi6n and the other from the margin; tertiary veins obscure above, along the original west boundary on the hill near prominulous and clearly visible beneath; both the junction with the Sura Trail. At La Selva the surfaces of blade glabrous but slightly scabrous oblong-obovoid, scabrous and pubescent leaves beneath at least when dry; petioles 6-10 cm long, are diagnostic. The oblong-ovoid figs of Ficus light tan with flaking periderm; stipules 3-5 cm popenoei are unique among Central American long, glabrous. Figs 1 per node, globose, 2-3 cm species. Leaves of the sympatric F. velutina are in diameter, slightly scabrous, nearly sessile, the similar to the present species, but are not sca­ peduncles 1-3 mm long; ostiole small, about 1 brous above. mm in diameter, minutely apiculate; basal bracts 3, broadly triangular, 3-4 mm long. Achenes 14. Ficus schippii StandI. ovoid, 1.5-3 mm long. Seen with figs in March and September. Honduras south into Ecuador. Liana or epiphytic shrub with long slender Uncommon but widely scattered at La Selva branches. Leafblades elliptic to oblong, 10-18 where it is found almost always along forest cm long, 5.5-8 cm wide, apically acute but usu­ streams. One individual grows on the point of ally with an abruptly acuminate tip 0.5-1.5 cm land along Quebrada EI Sura just behind the sta­ long, basally acute to rounded, sometimes cor­ tion. Ficus tonduzii is distinctive for its broad dulate; midrib and lateral veins prominent on leaves and distant, very conspicuous lateral and both surfaces; principal lateral veins 9-13 pairs submarginal veins. It shares features of the twigs, arising at an angle of (60)70°, the lowest pair figs, and venation with two other sympatric 254 SELBYANA [Volume 9

species of subgenus Pharmacosycea, F. glauces­ stipules small, paired and free, not encircling the cens and F. maxima and, among the species at stem, deciduous. Staminate inflorescences soli­ La Selva, is probably most closely related to these. tary or paired on short shoots, discoid to globose, shortly pedunculate with imbricate bracts cov­ 16. Ficus velutina Humb. & Bonpl. ex Willd. ering the receptacle; staminate flowers free or connate towards the base with a 4-parted peri­ Small to large trees 7 to 30 m tall, beginning anth, the filaments straight or slightly incurved as epiphytes and becoming stranglers; branchlets in bud; pistillate inflorescences globose, usually thick and sulcate with brownish pubescence of solitary, nearly sessile to pedunculate with, 1- spreading trichomes 0.5-1 mm long. Leafblades many-flowered, the perianth mostly 4-lobed, stiff and leathery, ovoid, 13-21 cm long, 9-12 fused to the ovary. Fruits usually succulent, cm wide, apically acute to rounded, basally trun­ I-seeded drupes, free or connate. cate to rounded and often cordulate, the margin The largely South American Macquira is a ge­ revolute; midrib and lateral veins plane or slight­ nus of five species. Only M. costaricana ranges ly impressed above when dry, very prominent into Central America. In floral and fruit char­ beneath; principal lateral veins 8-12 pairs, aris­ acters Macquira is scarcely distinct from Pere­ ing at an angle of 50-60°, distinctly loop-con­ bea. The two genera are separated largely on the nected within 3-5 mm of the margin; tertiary basis of the entire leaves in Macquira in contrast venation obscure above (slightly impressed when to the toothed leaves of Perebea and stipules that dry), very prominent and conspicuous beneath; completely encircle the stem of Perebea in con­ surface of blade glabrous above, distantly hirsute trast to the partially encircling stipules of Mac­ with brownish trichomes 0.5-1 mm long along quira. the midrib and lateral veins beneath, also mi­ nutely pubescent with short straight trichomes 1. Macquira costaricana (StandI.) C. C. Berg about 0.1 mm long along the tertiary veins; pet­ Shrubs or treelets 2-5(20) m tall; branchlets ioles 1.5-2 cm long, densely pubescent with ap­ drying somewhat reddish. Leaf blades oblong, pressed brown trichomes 0.5-1.5 mm long. Figs 17-29 cm long, (3.5)6.5-14 cm wide, apically 2 per node, globose 1.5-2 cm in diameter, mi­ rounded or broadly acute but abruptly acuminate nutely pubescent with erect trichomes, green with the acumen 1-1.5 cm long, basally acute to mottled with burgundy spots at maturity; ostiole rounded and often inequilateral; midrib and lat­ with a raised collar 6 mm in diameter; basal eral veins prominent on both surfaces; principal bracts broadly triangular, 2-3 mm long, pubes­ lateral veins 8-10 pairs, arising at an angle of cent; peduncle 4-6 mm long, pubescent with 65-90°, distinctly loop-connected and forming a brownish trichomes. Achenes 1.4-1.8 mm long. submarginal vein 3-5 mm from the margin; re­ Specimen with mature figs collected in August. ticulate venation prominent above when dry, Southern Mexico south into northern South prominent beneath and conspicuously lighter America. than the lamina; surface of blade glabrous and Uncommon at La Selva where known from glossy above, obscurely pubescent beneath with just three mature individuals. Ficus velutina usu­ short scattered trichomes; petioles 1-1.5 cm long; ally occurs at elevations above 1,000 m. How­ stipules about 0.5 cm long. Staminate inflores­ ever, it is distinctive throughout its range for the cences usually solitary or occasionally paired; pe­ very prominent reticulate venation, brown se­ duncle to 5 mm long; receptacles discoid, 0.5-1 riceous stipules, thick, leathery and very broadly cm in diameter, many-flowered, the flowers more ovate leaves, and its short pedunculate figs with or less connate at the base. Pistillate inflores­ a raised collar around the ostiole. At La Selva cences solitary, nearly sessile, discoid, 15-35- this species is likely to be confused only with F. flowered; stigmas ovoid, recurved, brownish pu­ popenoei whose leaves, however, are distinctly bescent; ovary completely enclosed in the ac­ scabrous above. crescent and succulent perianth. Fruits bright reddish orange at maturity, the seeds about 1 cm 7. MACQUIRA Aubl. long. Probably fertile throughout the year; sta­ minate material collected February to Novem­ BERG, C. C. 1972. 0lmedieae. Fl. Neotrop. Monogr. 7: 62-75. ber; pistillate material from March to July. Nic­ aragua south into Peru. Trees or shrubs, usually dioecious. Leaves en­ Common and widespread in the primary forest tire, 2-ranked, oblong and abruptly acuminate, at La Selva. Fruiting material is easily mistaken somewhat leathery, glabrous to slightly pubes­ for Annonaceae but the milky sap and paired cent, drying green, glossy above; petioles short; stipules are diagnostic. 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 255

8. NAUCLEOPSIS Miq. about 4 mm long and 0.5 mm wide; anthers nar­ rowly ovate, about 1.3 mm long and 0.5 mm BERG, C. c. 1972. Olmedieae. Fl. Neotrop. Monogr. wide, the connective wider at one end and at­ 7: 104-144. tached at the middle. Pistillate inflorescences dis­ Usually dioecious trees. Leaves 2-ranked, often coid to capitate when young, 2-3 cm wide at leathery, oblong and often widest above the mid­ anthesis, subtended by a series ofimbricate bracts, dle, entire, usually with more than 15 pairs of occasionally accompanied by a staminate inflo­ lateral veins; stipules free, completely encircling rescence; tepals indistinguishable from and in­ the stem. Staminate inflorescences discoid to terspersed with "pseudobracts" accrescent and more or less globose or cupulate, 1-6 per axil, spinelike in fruit, varying in length in different usually borne along leafless portions of the individuals from 0.5-2.5 cm long; stigmas 4-5 branches, usually pedunculate, the bracts of the mm long, black when dry. Infructescences glo­ receptacle numerous and somewhat petaloid; bose, 5-10 cm in diameter, green and more or flowers numerous, free or fused basally; stamens less woody at maturity, but the fruits imbedded straight in bud; pistillate inflorescences discoid in yellowish pulp of the receptacle; seeds ovoid, to globose, solitary, sessile or pedunculate; flow­ about 1.5 cm long. Staminate plants collected in ers few to many; tepals free or connate, similar early April; fruiting in May and June. Honduras to the numerous receptacular protuberances or south into Costa Rica. pseudobracts interspersed among the flowers; Frequent and widespread at La Selva where ovaries completely immersed in the receptacle. found most often on slopes and ridges in primary Infructescence usually globose and hard with forest in the southern half of the property. The spiny, angular protuberances; seeds imbedded in clear rather than milky sap is unusual in the Mo­ the pulpy receptacle. raceae, at least among those at La Selva, and Naucleopsis is a genus of 18 species restricted more like that of the Cecropiaceae. to the neotropics, with most species concentrated Naucleopsis naga was previously recorded in in the Amazon Basin. Only the following species Costa Rica and from just one fertile collection. occurs in Central America and none in the West The scarcity of collections of such locally com­ Indies. mon species is not unusual for the flora of the wet lowland tropics. Staminate material (Folsom 9686) of this species is here described for the first time. In pistillate plants, certain individuals or 1. Naucleopsis naga Pittier perhaps certain stages of all individuals, produce Usuallydioecious trees 10-25 m tall; bark light one to two staminate inflorescences at nodes to­ tan, the sap yellow and resinous; branchlets drying gether with the pistillate inflorescence. furrowed and with a somewhat flaky periderm. The length of the spinelike pseudobracts on Leaf blades oblong, widest at or above the mature infructescences of most trees are about middle, 18-34 em long, 5-12 em wide, apically 2.5 cm long, but some individuals bear infruc­ acute to rounded and sometimes abruptly acu­ tescences with mature spines less than 1 cm long. minate, basally acute to rounded; midrib and lateral veins prominent on both surfaces; prin­ 9. PEREBEA Aubi. cipallateral veins 18-20 pairs, faintly loop-con­ nected near the margin; tertiary veins obscure BERG, C. C. 1972. Olmedieae. Fl. Neotrop. Monogr. above; surface of blade glabrous above, glabrous 7: 38-61. beneath or distantly appressed pubescent along Dioecious or monoecious shrubs or trees. the midrib; petioles 1-2 cm long; stipules 1-1.5 Leaves oblong-elliptic, usually with an acumi­ cm long, minutely appressed pubescent, the scars nate tip, marginally dentate; stipules free, ca­ obscure. Staminate inflorescences clustered, 1-6 ducous, completely encircling the stem. Stami­ per axil, usually found at leafless nodes, cupulate, nate inflorescences cylindrical to discoid, usually 0.5-1.3 cm in diameter; bracts of the receptacle clustered in the axils of leary nodes, sessile to broadly ovate, grading in a series of 8-15 im­ shortly pedunculate or borne on well developed bricate bracts from smallest outside to largest short shoots, the receptacle covered with imbri­ inside, the largest about 5 mm long and 8 mm cate bracts; flowers 1-many; tepals and stamens wide, minutely pubescent on the surface and cil­ usually 4, the filaments straight or slightly in­ iate on the hyaline margin; flowers numerous, curved in bud; pistillate inflorescences discoid, 20-50 per inflorescence, free to the base, 5 mm solitary, occasionally accompanied by 1 to sev­ long, the perianth of 4 more or less free, ligulate eral staminate inflorescences; flowers I-many, lobes about 4 mm long; stamens 2, filaments the perianth mostly 4-parted, the stigmas discoid 256 SELBYANA [Volume 9 to linear. Infructescencereddish orange, the fruits of flowering and fruiting, peaking in February drupaceous in a fleshy perianth. and July. Perebea is a genus of eight species restricted to the neotropics and centered in the Amazon 10. PSEUDOLMEDIA Trecul Basin. Three species range into Central America with two of them reaching as far north as Costa BERG, C. C. 1972. Olmedieae. A. Neotrop. Monogr. Rica. 7: 17-38. Dioecious trees, medium to large. Leaves usu­ 1. Perebea augustifolia (Poepp. & Endl.) C. C. ally entire, usually with microscopic pluricellular Berg trichomes, 2-ranked; stipules free, fully encir­ cling the stem, caducous. Staminate inflores­ Usually dioecious shrubs or trees 2.5-8(20) m cencesusually 1-4 per node, discoid, sessile; peri­ tall; branchlets sparsely appressed pubescent with anth lacking, numerous free stamens interspersed trichomes about 0.5 mm long. Leafblades ob­ with 1igulate bracts, the anther often provided long-elliptic, (5)14-24.5 cm long, (1)4.5-10 cm with an apical awn or tuft oftrichomes. Pistillate wide, apically acuminate with an abruptly nar­ inflorescences solitary or paired, sessile and with rowed tip 1-2.5 cm long, basally acute to round­ only 1 flower, the receptacle bearing imbricate ed; midrib and lateral veins obscurely raised bracts; perianth tubular and fused to the ovary; above when dry, prominent and conspicuous be­ stigmas filiform. Fruits reddish and succulent neath; principal lateral veins 12-16 pairs, arising from the accrescent perianth. at an angle of 60-80°, distinctly loop-connected Pseudolmedia, a genus of nine species, is rep­ 1-6 mm from the margin; tertiary veins obscure resented by two species in southern Mexico above, prominent and conspicuous beneath; sur­ through Central America with one of them, P. face of blade glabrous, sparsely appressed pu­ spuria, also found in the West Indies. The other bescent on the veins beneath with trichomes seven species occur only in South America, rang­ about 0.5 mm long; petioles 0.5-1.5 cm long, ing into southern Brazil and Bolivia. distantly appressed pubescent; stipules 7-12 mm long, not united, sericeous especially towards the 1. Pseudolmedia spuria (Sw.) Griseb. base and along the margin. Staminate inflores­ cences clustered on short shoots in leafaxils, cu­ Trees (5)35 m tall, about 1.5 m in diameter, pulate or cylindrical, 3-5 mm long and 2-3 mm the branchlets smooth and brown, glabrous; bark wide, 1-6 per cluster; receptacle with 20-25 im­ dark brown; sap white. Leaf blades narrowly bricate bracts in several series, the bracts mi­ ovate to elliptic or oblong, (4)6-12(14) cm long, nutely appressed pubescent; peduncles 1-3 mm (1.5)2-3(6) cm wide, apically acute to acuminate long, minutely appressed pubescent; flowers 5- with a narrow blunt tip, basally acute; midrib 20 per inflorescence, the perianth 4-parted. Pis­ and lateral veins prominent on both surfaces; tillate inflorescences solitary (rarely accompan­ principal lateral veins 10-18 pairs, arising at an ied by a staminate inflorescence), cupulate or dis­ angle of 60-75°, distinctly loop-connected 1-3 coid, 5-8 mm wide at anthesis; receptacle covered mm from the margin; tertiary veins prominulous with about 10-25 imbricate bracts in several se­ on both surfaces, very finely reticulate; surface ries, the bracts with lighter colored margins and ofthe blade glossy and essentially glabrous above minutely appressed pubescent; peduncles 2-4 mm and beneath; petioles 3-8 mm long, glabrous; long, minutely appressed pubescent; flowers stipules 2-12 mm long, minutely puberulent to about 5-15 per inflorescence, the perianth tu­ glabrous. Staminate inflorescences subtended by bular, densely pubescent, accrescent, succulent an involucre of lanceolate bracts; bracts among and covering the fruit at maturity, the stigmas the stamens often widest above the middle; an­ narrowly lance01ate. Infructescences 1.5-3 cm in thers apiculate; pistillate inflorescences (none diameter with clusters ofl-15, bright red or red­ found at La Selva) also subtended by an invo­ dish orange, ovoid fruits about 1 cm long. Per­ lucre of deltiod bracts. Fruit an ovoid drupe about haps fertile throughout the year; both flowering 5-9 mm in diameter. Flowering in May. Mexico and fruiting material collected from February south into Panama; West Indies. through November. Costa Rica south into Am­ Known from only one individual at La Selva. azonian Brazil and Peru. In Flora Costaricensis Burger reported only three Widespread and common at La Selva where other collections from Costa Rica. The La Selva found throughout the primary forest. Perebea material was collected from a very large tree on angustijolia occasionally bears staminate inflo­ a forested ridge along the west boundary of the rescences with the pistillate. At La Selva this new property just north of the Quebrada Paralela species appears to go through two major seasons crossing. 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 257

11. SOROCEA St. Hil. above and puberulous on the veins, distantly hir­ sute on the blade beneath but more densely so Dioecious shrubs or trees (elsewhere). Leaves on the veins; petioles 1-4 cm long, hirsute; stip­ usually oblong-elliptic, unlobed but often den­ ules 4-6 mm long, minutely pubescent and mar­ tate in the distal third, glabrous or pubescent, ginally ciliate. Inflorescences spicate, 2-12 cm tertiary venation often conspicuous beneath; long, solitary or clustered, the staminate ones stipules paired, usually deciduous, leaving scars occasionally branched, floriferous to the base or that do not encircle the stem. Inflorescences sol­ nearly so. Staminate flowers rather densely clus­ itary, paired, or clustered spikes or racemes in tered but distinct on the rachis; filaments 0.7-1 the a~:ils of leaves, the rachis usually covered mm long; anthers ovoid or orbicular at dehis­ with peltate bracts; staminate flowers 4-parted, cence; pistillate flowers more distant on the ra­ sessile or pedicellate, the tepals decussate-im­ chis, sessile or shortly pedicellate; perianth tu­ bricate in bud, the filaments straight in bud; pis­ bular, minutely pubescent, the style stout, usually tillate flowers tubular, apically minutely 4-parted less than 3 mm long, hollow; stigmas broadly or irregular, perianth accrescent and covering the triangular, acuminate. Infructescence racemose, mature fruit, often minutely pubescent, style 2- 7-13 cm long, often irregularly twisted or curved; lobed. Fruits drupaceous, single-seeded, en­ fruits ovoid, about 1 cm long at maturity, sur­ closed in the fleshy, succulent perianth, bright rounded by the succulent and minutely pubes­ red or orange at maturity. cent perianth; pedicels 2-5 mm long. Apparently Sorocea is a genus of 22 species ranging from flowering and fruiting throughout the year. Costa Guatemala to Paraguay and northern Argentina. Rica and Panama. Four species occur in Central America and three Common and widespread at La Selva although of them are endemic. All four occur in Costa restricted to somewhat open areas such as Rica. Inflorescences of Sorocea are very similar streambanks and swamp edges. At La Selva the to those of Trophis but the two genera can be species is usually a slender shrub 2 to 2.5 m tall distinguished at La Selva on the basis of vege­ but elsewhere it becomes a tree to 20 m tall. The tative characters; for the most part the leaves of bright reddish orange fruits in contorted racemes Trophis species from La Selva are scabrous and are distinctive and are eaten by tanagers and oth­ have a raised midrib whereas they are never sca­ er birds of forest edges. brous in Sorocea and always have a sunken mid­ rib. 2. Sorocea trophoides W. Burger

KEy TO THE SPECIES Slender shrubs 1-2.5(15) m tall; branchlets glabrous, bright yellowish green. Leaf blades 1. Leaves pubescent beneath especially along the veins, oblong-elliptic, (7)24-40 cm long, (2)6.5-13 cm with the trichomes commonly to 0.4 mm long; sta­ mens erect in bud, 1-1.3 mm long at anthesis... wide, apically abruptly acuminate to an acumen ...... 1. S. pubivena. 1.5-3 cm long, basally acute and decurrent to 1. Leaves essentially glabrous throughout; stamens in­ rounded, the margin shallowly undulate in the curved in bud, 1.8-2 mm long at anthesis...... upper 213; midrib impressed above, the lateral ...... 2. S. trophoides. veins prominulous or slightly impressed, veins prominent and conspicuous beneath; main lat­ 1. Sorocea pubivena Hemsl. eral veins 11-14 pairs, arising at an angle of 50- 70°, distinctly loop-connected 1-4 mm from the Small trees or shrubs 2-10(20) m tall; branch­ margin; tertiary veins obscure above, conspicu­ lets hirtellous, dull grayish green or tan. Leaf ous beneath; surface of blade glabrous through­ blades oblong-elliptic, often widest above the out except very minutely pubescent along the middle, 18-35 cm long by 6-13 cm wide, apically midrib above; petioles 1.5-2 cm long, glabrous; abruptly acuminate with a narrow acumen 1.5- stipules about 4 mm long, glabrous. Inflores­ 4 cm long, basally acute and usually somewhat cences spicate, 1-3 cm long, usually solitary, the decurrent but occasionally truncate, the margin staminate nearly sessile, the pistillate with a short usually shallowly and distantly serrate especially peduncle 3-5 mm long. Staminateflowers sessile in the upper 213; midrib and lateral veins im­ and rather congested on the rachis; filaments of pressed above, prominent and conspicuous be­ the stamens about 1.5 mm long, the anthers neath; main lateral veins 9-12 pairs, arising at ovoid, about 0.6 mm long at dehiscence. Pistil­ an angle of 50-70", distinctly loop-connected (at late flowers globose-depressed or urceolate, 1-2 least in the distal halO 3-5 mm from the margin; mm long, sessile or on a short pedicel about 1 tertiary veins obscure above, prominent and mm long; stigma slender and apparently solid, conspicuous beneath; surface of blade glabrous 2-4 mm long; styles narrowly lanceolate-acu- 258 SELBYANA [Volume 9

minate. Infructescence about 4 cm long; fruits 1. Trophis involucrata W. Burger globose or ellipsoid, 4-8 mm in diameter on pedicels about 3 mm long, minutely puberulent. Shrubs 1.5-3 m tall; new branchlets minutely Flowering specimens from February and August, and densely hirsute but soon becoming glabrate, one fruiting collection (immature) from August. reddish and the periderm somewhat exfoliating. Costa Rica south into Peru. Leaf blades elliptic, oblong to oblanceolate, often Restricted to a single population at La Selva widest above the middle, 9-20 cm long, 4-8 cm along the Quebrada Taconazo. Sorocea tro­ wide, apically abruptly long-acuminate with an phoides is usually found between 750 and 2 000 acumen 2-3 cm long, basally acute to rarelyal­ m elevation. The plants at La Selva have s~me­ most rounded; midrib and lateral veins plane and what larger leaves than usual for the species. obscure above, prominent and conspicuous be­ neath; principal lateral veins mostly about eight pairs not counting the small but distinct veins of 12. TROPHIS P. Br. t~e acumen, arising at an angle of 60-80°, dis­ BURGER, W. 1962. Studies in New World Moraceae: tInc.tly loop-connected 2-6 mm from the margin; Trophis, Ciarisia, Acanthinophyllum. Ann. Mis­ tertIary veins obscure above, prominent and souri Bot. Gard. 49: 1-34. conspicuous beneath; surface of blade not visibly pubescent but scabrous on both surfaces; petioles Dioecious trees or shrubs. Leaves 2-ranked, 0.3-0.8 cm long, minutely hirsute; stipules tri­ often scabrous, apically acuminate, marginally angular, 1.5-2 mm long, leaving no visible scar, shallowly undulate to serrate; stipules leaving a often somewhat persistent. Staminate inflores­ scar encircling less than half the stem. Inflores­ cences racemose, axillary, paired or solitary, 1- cences spicate or racemose in the axils of leaves 2.5 cm long; rachis with scattered, small, nar­ paired or solitary; rachis with peltate subor~ rowly triangular bracts; flowers few to many, bicular or basally attached, triangul~ bracts. rather distant on the rachis, the young buds tur­ Staminate flowers 4-parted, the tepals valvate in binate and pointed, later becoming globose the bud; stamens incurved in bud. Pistillate flowers pedicels about 1-1.5 mm long; tepals bro'adly with a tubular perianth sometimes adnate to the triangular, about 1.5 mm long at anthesis, mi­ fruit; stigmas slender and deeply bifid; perianth nutely and sparsely hirsute with a tuft of longer tUbe.accrescent and succulent in fruit, sometimes trichomes at the tip; filaments 2 mm long, anther beanng small fleshy spinelike protuberances. sacs broadly reniform, medially attached. Pistil­ Fruits globose, drupaceous, becoming black or late inflorescences axillary, racemose, solitary or red at maturity. paired, 1-2 cm long; rachis covered (in early Trophis is a small genus offour to five species stages) with broader, denser, imbricate bracts re­ concentrated in southern Mexico but extending sembling an involucre (especially in uniflorous also to the West Indies and South America. The inflorescences); flowers (1)2-5, distant on the three taxa at La Selva can be distinguished even rachis, the pedicels to 0.5 cm long; perianth of without fertile material, but in general, fertile 4 bractlike tepals, about 5 mm long, much ex­ samples of both male and female plants are nec­ essary. ceeded by the cylindrical and tuberculate, 2-3 mm long ovary; style lacking; stigmas filiform, 3-4 mm long, densely hirsute. Fruits globose to broadly fusiform, 1.3-1.6 cm long, adorned with numerous fleshy tubercles about 1.5 mm long KEy TO THE SPECIES dis~tly and minutely hirsute throughout, be~ comIng succulent and tasty at maturity, dark pur­ 1. Flowers pedicellate, inflorescences racemose (1-2- flowered in female plants); petioles about 0.5 cm ?le to reddish black. Flowering and fruiting mostly long; mature fruits with numerous slender fleshy In the dry season, December through April. Ca­ protuberances; small understory shrubs or treelets ribbean lowlands of Costa Rica in Alajuela He- widely scattered in the primary forest...... redia and Limon provinces. ' ...... 1. T. involucrata. Widespread in the primary forest at La Selva 1. Flowers sessile, inflorescences spicate; petioles but perhaps most common on ridges in the mostly 1 cm or longer; mature fruits plain; subcan­ northern part of the property. It is conspicuous opy trees of old secondary woods along the river. only during the dry season when flowering and ...... 2. T. racemosa. 2. Leaves smooth; mature staminate inflores­ ~~ing. The slightly scabrous leaves are very cences 5-7 cm long; largest leaves about 23 cm slmdar to those of the sympatric Trophis race­ long...... ssp. racemosa. mosa spp. ramon but have much shorter peti­ 2. Leaves scabrous; mature staminate inflores­ oles. Trophis racemosa is also a much larger tree cences 3-4.5 cm long; largest leaves about 15 and is restricted to alluvial soils at La Selva. cm long...... ssp. ramon. Pollen is explosively released in this species, and 1986] HAMMEL: FLORA OF LA SELVA-MORACEAE 259 other morphological and experimental evidence bescent. Fruits, not seen mature at La Selva, glo­ confirm that it is wind-pollinated. bose, turbinate when dry, about 1 cm in diam­ eter, becoming red or blackish at maturity. Flowering during the rainy season, July to Oc­ 2. Trophis racemosa (L.) Urb. tober, fruits probably maturing by December. Mexico south into Venezuela and Peru; West Small to medium-sized trees, 6-25 m tall, the Indies. bark usually reddish brown and flaking. Leaf Known only from secondary or gallery forest blades elliptic to oblong, 14-15 cm long, 5-6.5 along the Rio Puerto Viejo. In fact, all the in­ cm wide, apically acuminate with an acumen dividuals known so far lie along the West River 1.5-2 cm long, basally acute to rounded; midrib Road in the strip of land once planted to cacao and lateral veins obscure above, prominent be­ and pejibaye, or in the pastured property op­ neath; principal lateral veins (6)8-9(10) pairs, posite the station. arising at an angle of 50-70°, distinctly loop­ Most of the La Selva plants of this species have connected, especially above the middle, 2-8 mm scabrous leaves and short, densely flowered pis­ from the margin; tertiary veins obscure but tillate inflorescences, and fit well within the con­ prominulous above when dry, prominent and cept of ssp. ramon. However, the leaves of two conspicuous beneath; surface of blade scabrous individuals are not at all scabrous, they are with­ above and beneath, otherwise glabrous; petioles out teeth on the margin, and the staminate in­ about I cm long, grooved above, minutely pu­ florescences are commonly as much as 6 to 7 cm bescent; stipules broadly triangular, 2.2-3 mm long. These features, taken together, clearlyiden­ long, distantly and minutely pubescent. Stami­ tify these problematic individuals as ssp. race­ nate inflorescences usually paired in the axils of mosa formerly thought restricted to the West leaves, 1.3-4.5 cm long, densely and minutely Indies. In addition to the characters of the key, pubescent throughout, peduncle about 5 mm long; the collections of ssp. racemosa differ from those flowers densely clustered, the perianth sessile, of ssp. ramon at La Selva by the following: bark tepals 1.5-2 mm long, densely and minutely pu­ light gray, not flaking; leaves 22-23 cm long, 7- bescent, filaments about 2.5 mm long. Pistillate 9 cm wide; tertiary veins obscure beneath; pet­ inflorescences usually paired, about 1.3 cm long ioles 1.2-1.5 cm long, glabrous; stipules to 5.5 including a short peduncle about 5 mm long, mm long with a long acuminate tip. densely pubescent throughout, usually 6-8-flow­ The pollen of Trophis racemosa is probably ered; flowers more or less paired, sessile, and dispersed by air currents. Apparently triggered densely clustered along one side of the rachis, the by desiccation in the sunlight, the pollen was perianth urceolate and obscurely lobed at the observed being released like that of T. involu­ apex, densely pubescent, stigmas linear and pu- crata.