342
DICOTYLEDONEAEoutside; petals lacking; anthers of the staminate flowers contained within the boat-shaped tepal in bud, the tension of the elongating filament released when the anther slips from the sepal and springs upward, the thecae directed upward; pollen white, powdery; pistillode columnar;
pistillate flowers similar to staminate flowers, the ovary globose, with 2 bifid styles, the stylar bases persisting in fruit. Fruits ovoid or subglobose, 3-4 mm long, red.
Croat 6242.
Occasional at the edges of clearings, rare in the forest.
Flowers and fruits principally in the rainy season.
Superficially similar to Pouzolzia obliqua (40. Urti- caceae), which has four-parted flowers and an achene with a single long style.
In Ecuador and some other areas, Trema integerrima (Beurl.) Standl. is recognized as a distinct species, being a larger tree occurring in forests rather than in disturbed areas. The differences on BCI seem insignificant, though two collections, Aviles 58a and Hay den 4, have the short erect pubescence and the entire thin leaves attributed to T. integerrima. Both are considered here to be only forms of T. micraniha.
Widely distributed in the tropics and subtropics of the Western Hemisphere. In Panama, ecologically variable and widespread in cutover areas; known from tropical moist forest throughout Panama, from tropical dry forest in Code, from premontane moist forest in the Canal Zone, from premontane wet forest in Chiriqui, Code, and Panama, from tropical wet forest in Colon and Ver- aguas, and from premontane rain and lower montane wet forests in Chiriqui.
See Fig. 193.
39- MORACEAE 343
KEY TO THE TAXA OF MORACEAE
Plants herbs; inflorescences cup-shaped Dorstenia contrajerva L.
Plants trees or shrubs; inflorescences various:
Flowers borne on the inner surface of a ± globose, hollow receptacle, its apex a small opening closed by scales; stems solid; stipules fully amplexicaul, usually conspicuous Ficus Flowers not borne on inner surface of a ± closed receptacle; stems often hollow; stipules fully
amplexicaul or lateral:
Leaves conspicuously lobed:
Leaves pinnately lobed; sap white Artocarpus altilis (Park.) Fosb.
Leaves palmately lobed; sap brown or black:
Flowers in spikes; leaves peltate Cecropia Flowers in cymes; leaves basifixed Pourouma guianensis Aubl.
Leaves not lobed:
Plants armed; staminate flowers in dense globose heads 1-2.3 cm diam; pistillate flowers more than 5 mm long, 5-9 in ovoid heads Poukenia armata (Miq.) Standl.
Plants unarmed; flowers various:
Flowers in spikes or racemes:
Staminate spikes densely flowered; pistillate flowers and fruits sessile; leaves asperous Trophis racemosa (L.) Urban Staminate spikes loosely flowered; pistillate flowers and fruits pedicellate; leaves
smooth Sorocea affinis Hemsl.
Flowers solitary or in globose or discoid heads:
Leaves ovate, spirally arranged; petioles more than 3 cm long; staminate inflorescences dichotomously branched 2-4 times with several ± globose heads ca 5 mm diam:
Leaves usually cordate at base, the major lateral veins in 6-8 pairs above base; pis- tillate inflorescences (heads) 2-4-lobed Coussapoa magnifolia Tree.
Leaves usually obtuse to truncate at base, the major lateral veins in 10-20 pairs above base; pistillate inflorescences (heads) not lobed Coussapoa panamensis Pitt.
Leaves not ovate, distichous; petioles less than 2 cm long; staminate inflorescences not compound:
Inflorescences bisexual, globular, ca 5-10 mm diam with many staminate flowers and 1 or 2 pistillate flowers at center of cluster Brosimum alicastrum Sw. subsp. bolivarense (Pitt.) C. C. Berg Inflorescences unisexual:
Staminate inflorescences fan-shaped; pistillate flowers in discoid heads; perianth lacking; fruits syncarps (all carpels united); blades strongly cordate, dentate Castilla elastica Sesse Staminate inflorescences not fan-shaped; pistillate flowers solitary or in discoid
heads; perianth distinct; fruits solitary or the carpels ± free; blades cuneate to narrowly rounded at base, mostly entire (undulate-serrate in Perebea):
Pistillate flowers dense in discoid heads:
Leaves and stems glabrous or the stems inconspicuously puberulent; blades entire; fruiting heads to 5 cm diam; fruits to 2 cm long Maquira costaricana (Standl.) C. C. Berg Leaves (at least on midrib) and stems conspicuously hispidulous; blades
undulate-serrate toward apex; fruiting heads ca 2 cm diam; fruits less than 1 cm long Perebea xanthochyma Karst.
Pistillate flowers solitary, small, bracteate at base:
Leaves asperous; staminate flowers with a distinct perianth and 4 stamens;
ovary superior; leaves remotely undulate-serrate at apex Olmedia aspera R. & P.
Leaves smooth; staminate flowers with vestigial perianth, the heads unorgan- ized; ovary inferior to semi-inferior; leaves entire Pseudolmedia spuria (Sw.) Griseb.
the genus Azteca do not function in pollination of flowers Reportedly 53 (Willis, 1966) to 73 (Lawrence, 1964) or dispersal of seeds (Wheeler, 1910). These ants are genera and 1,000-1,500 species. The family is worldwide beneficial to the plant, however, in warding off predators in distribution, principally in tropical or subtropical areas such as leaf-cutter ants (Atta) (Janzen, 1969). with comparatively few species in temperate areas.
344 DICOTYLEDONEAE ARTOCARPUS Forst. & Forst.f.
Artocarpus altilis (Park.) Fosb., J. Wash. Acad. Sci.
31:95.1941.
A. communis Forst. & Forst.f.
Breadfruit
Monoecious tree, usually less than 12(20) m tall; trunk lenticellate; outer bark thin; inner bark thick, granular, with copious milky sap; wood yellowish. Petioles stout, (2)4-7 cm long; blades ovate in outline, deeply pinnately lobed, acuminate at apices of lobes, the blades acute at base, 30-80(100) cm long, 15-40(65) cm wide, thick and leathery, scabrous, usually pubescent below and on veins above; stipules fully amplexicaul, to 15 cm long, conspic- uously appressed-pubescent, deciduous. Peduncles stout, 5-8 cm long; staminate spikes club-shaped, 20-40 cm long, ca 3 cm diam at apex, narrower at base, the stami- nate flowers dense; perianth 2- or 4-lobed, ca 3 mm long;
stamen solitary; anther exserted, oblong, bilobed. Pistil- late flowers in globular or oblong spikes; perianth tubular, embedded in the fleshy rachis; style exserted; stigmas entire, 2- or 3-lobed; fruiting pedicels to 7.5 cm long, 1.5-2 cm wide. Syncarp ellipsoid or rounded, 10-30 cm long, green and soft at maturity, the surface asperous with numerous round or isodiametrical segments or echinate;
seeds many. Croat 10121.
Cultivated in the Laboratory Clearing. Flowers in the late dry season. The fruits mature in the middle to late rainy season.
Native to the South Pacific islands; cultivated through- out the tropics of the world and in various places in Panama.
BROSIMUM Sw.
Brosimum alicastrum Sw. subsp. bolivarense (Pitt.) C. C. Berg, Acta Bot. Neerl. 19:326. 1970.
B. bernadetteae Woods.; Helicostylis latifolia Pitt.
Berba
Monoecious tree, 3-35 m tall; bark thin, with prominent leaf scars and irregular horizontal raised lines; inner bark smooth, thick, tan; sap forming milky droplets. Petioles stout, 4-14 mm long; blades elliptic-obovate to elliptic, acuminate to mucronate at apex, broadly cuneate to rounded at base, 6-15 (20) cm long, 3-6.5 (8) cm wide, glabrous, coriaceous, the major lateral veins raised below, with a conspicuous submarginal collecting vein and prominulous reticulate veins; stipules nearly encircling stem, 5-9(15) mm long, deciduous. Flowers dense, in globular clusters 4-9 mm diam, completely concealed before anthesis by short-stipitate, round, peltate bracts;
peduncles obsolete or to 5 mm long; perianth obsolete;
staminate flowers many, the stamen solitary, the anther circular, eccentrically peltate, ca 1 mm diam, dehiscing by 2 basal valves; pistillate flowers 1 or 2 at center of clusters, the stigmas deeply 2-lobed, the lobes exserted 4-7 mm before staminate flowers open, spreading, subu-
late. Fruit a false drupe, ± globose, 1-1.5(2) cm diam, with minute round protuberances, with an apical depres- sion; seed 1. Croat 10306, 11647.
Common in the forest. Seasonal behavior uncertain.
Apparently flowers from November to May, mostly during the dry season. The fruits mature from May to October and are eaten when ripe by monkeys (Hladik &
Hladik, 1969).
Carpenter (1934) reported that fruits of this species are second only to Ficus as food for most animals of the forest. Bats play a principal role in their dispersal (R.
Foster, pers. comm.). The outer shell of the fruit is often thrown to the ground.
The subspecies bolivarense is distinguished from the subspecies alicastrum by having anthers with free thecae;
anthers of the subspecies alicastrum are peltate with the thecae fused.
The typical subspecies ranges from Mexico to Costa Rica and the West Indies. The subspecies bolivarense ranges from Costa Rica through the Andes to Guyana and Brazil (Acre Territory). In Panama, known from tropical moist forest in the Canal Zone, Chiriqui, Pa- nama, and Darien. Reported from tropical wet forest in Costa Rica (Holdridge et al., 1971).
CASTILLA Sesse
Castilla elastica Sesse in Cerv., Gaz. Lit. Mexico, Suppl. 7. 1794
C. panamensis Cook
Rubber tree, Mastate bianco, Caucho, Hule, Ule Monoecious or dioecious tree, 10-30 m tall, with low buttresses; young parts densely yellowish-pubescent;
stems hollow; sap white. Leaves pendulous, deciduous;
petioles to 1 cm long; stipules fully amplexicaul, 3-9 cm long, densely golden, appressed-pubescent, deciduous;
blades oblong to oblong-obovate, acuminate at apex, cordate at base, 20-45 (55) cm long, 8-18 (25) cm wide, minutely ciliate-denticulate, asperous especially below with short, spreading, often golden trichomes. Flowers inserted on large, flattened, unisexual receptacles covered with imbricate bracts, the receptacles axillary or at de- foliated nodes, involucrate; primary staminate inflores- cences (2)4(6) per axil, fan-shaped, conduplicate, to 15 mm long and 25 mm wide; peduncles 3-10 mm long;
bracts in 10-12 series; perianth absent; stamens numerous, scattered among the bractlets; filaments to 3.5 mm long;
anthers oval, ca 1 mm long; complemental staminate inflorescences (accompanying pistillate inflorescences) usually 2, similar to primary ones or funnel-shaped to cup-shaped. Pistillate receptacles usually solitary, thickly discoid, 1-2 cm diam, 1-2 cm thick, nearly sessile; in- volucral bracts in 5-10 series; flowers mostly 15-30;
perianth tubular, (1.5)2-3 mm long, shallowly 4(5)-lobed, fleshy, short-velutinous, accrescent; ovary subinferior;
styles dimorphic, to 1.5 mm long with stigmas 3-6 mm long or 2-3 mm long with stigmas less than 3 mm long;
stigmas 2, rarely 3. Fruits thick, discoid syncarps 2.5-4.5
39-
MORACEAE/CECROPIA345
cm wide; seeds many, ca 1 cm long and enclosed in the orange, fleshy, accrescent perianth. Croat 5335.
Occasional in the forest on the western side of the island. Probably flowers in the late dry and early rainy seasons (April and May). Berg (1972) reported that the species flowers all year. The fruits are mature in May and June. The leaves are lost in the dry season.
Widespread on both coasts from Mexico to Panama and along the western coasts of Colombia and Ecuador.
In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Panama, and Darien, from pre- montane moist forest in the Canal Zone and Panama, from premontane wet forest in Code (El Valle), and from tropical wet forest in Colon (Guasimo).
CECROPIA L. (Guarumo, Trompy, Trumpet tree) Cecropia is distinguished by its slender, pendent, uni- sexual catkins; staminate catkins fall soon after anthesis.
Flowers have a minute tubular perianth, the staminate with two stamens, the pistillate with a barely exserted, pencil-shaped style. Each flower produces a tiny achene.
Leaves are peltate and spirally arranged on stout, hollow branches, which frequently house myrmecophilous ants.
The leaves are a favorite food of sloths. The genus is
characteristic of secondary areas and usually has soft wood and black sap.
Cecropia insignis Liebm., K. Danske Vidensk. Selsk.
Skr. Naturvidensk. Math. Afd., ser. 5, 318. 1851 C. eximia Cuatr.
Guarumo bianco
Dioecious tree, to about 40 m high and 70 cm dbh, broadly branched; trunk with prominent lenticels and often stilt-rooted; outer bark thin; inner bark reddish.
Petioles stout, arachnoid-villous and longitudinally striate, the basal pulvinus brown; stipules commonly red when falling, mostly 17-26(40) cm long and 5-8 cm broad, densely hirtellous on outer surface, with 2 promi- nent ridges; mature leaves prominently clustered at ends of branches, up to 90 cm wide, divided more than three- fourths of the way to center, the lobes 7-9, acute to obtuse at apex, conspicuously narrowed at base, to 22 cm broad, the margins held conspicuously upward, the upper sur- face smooth and shiny, glabrous except for sparse arach- noid trichomes deciduous in age, the lower surface paler and minutely canescent beneath (at least between veins).
Staminate spathes oblong, 10-12(16) cm long, 5-6.5 cm wide, rounded at apex with long apiculate tip, the spa- dices held erect in clusters of usually 6 or 7, 12-15 cm long, 8-10 mm wide, whitish or green, the basal stipes
KEY TO THE SPECIES OF CECROPIA
Leaves smooth above, remaining intact after falling to ground; basal stipe of spadices glabrous, purplish, pruinose (covered with thin, waxy layer); pistillate spadices yellow (except when very young) C. insignis Liebm.
Leaves scabrous or rough on upper surface, usually rolling up after falling to ground; basal stipe of spadices not pruinose, variously pubescent; pistillate spadices white or greenish:
Leaves often divided more than three-fourths of the way to base, usually flat or only slightly folded; staminate spadices usually in clusters of less than 10; pistillate spadices 25-30 cm long when fully expanded C. obtusifolia Bertol.
Leaves usually divided about halfway to base or less, the surface of the blade much folded, not able to be flattened; staminate spadices in clusters of 12-60; pistillate spadices less than 12 cm long when fully expanded:
Pistillate spadices in clusters of 4-6, the common peduncle not more than twice as long as spadix; basal pulvinus of petiole with uniform trichomes, the velvetlike layer of tri- chomes not interspersed with longer white trichomes C. peltata L.
Pistillate spadices in clusters of 6-12, the common peduncle 5-8 times as long as spadix; basal pulvinus of petiole with the brown velvetlike layer of trichomes interspersed with dense, longer, white trichomes C. longipes Pitt.
KEY TO THE SPECIES OF CECROPIA (ON THE BASIS OF STAMINATE SPADICES)
The following key is provided because staminate inflorescences fall soon after anthesis and are often found on the ground.
Staminate spadices less than 10 per peduncle:
Spadices 4 mm thick; basal stipes ± green, puberulent, to 10 mm long; leaves scabridulous above, flat to somewhat pleated C. obtusifolia Bertol.
Spadices ca 1 cm thick; basal stipes glabrous, purple, pruinose, 12-16 mm long; leaves smooth above, much pleated C. insignis Liebm.
Staminate spadices 12 or more per peduncle:
Spadices usually less than 6 cm long C. peltata L.
Spadices usually more than 6 cm long C. longipes Pitt.
purplish, broad, 12-16 mm long, pruinose, the common peduncles 5-10 cm long; pistillate spathes as those of staminate, the spadices in clusters of 4-7, pale yellow, 8-10 cm long and ca 8 mm thick at anthesis, becoming bright yellow in fruit and to 15 cm long and 1.3 cm thick, the basal stipes thick, pruinose, ca 1 cm long, the common peduncles 8-13 cm long, gray-hirtellous. Achenes ovate- elliptic, somewhat flattened, 1.7-2.3 mm long, ca 1 mm wide, conspicuously muricate, reddish-brown. Croat 7023 $.
Common in the forest, even in the older forest. Many individuals grow in the vicinity of the Laboratory Clear- ing. Flowers in the dry season (December to April). The fruits mature from April to August with a peak in July.
Easily distinguished by the large, smooth, very deeply divided blades and the spadices with pruinose stipes.
Nicaragua to Colombia. In Panama, known from tropi- cal moist forest in the Canal Zone, Bocas del Toro, and Los Santos and from tropical wet forest in Colon (Salud).
Reported from premontane wet and rain forests in Costa Rica (Holdridge et al., 1971).
See Fig. 194.
Cecropia longipes Pitt., Contr. U.S. Natl. Herb.
18:227.1917 Guarumo poludo
Dioecious tree, to 15 m tall; branches stout, 4-5 cm diam at apex. Petioles gray-hirtellous, 30-75 cm long, the brown velutinous trichomes interspersed with longer white trichomes; stipules ca 7-9 cm long; mature leaves to ca 90 cm wide, usually divided less than halfway to center, dark green and scabridulous above, paler and subarachnoid-puberulent below; lobes 9-13, broadly rounded at apex, not narrowed at base. Staminate spathes carrot-shaped before anthesis, pendent, reddish-brown, densely hirsute, 9-12 cm long, the spadices in clusters of 16-50, to 13 cm long and 2-3 mm diam, the stipes
1-2 cm long, gray-hirtellous, the common peduncles 7-12 cm long, densely hirsute; perianth tubular; stamens 2. Pistillate spathes reddish, 2-ribbed, the spadices in clusters of 6-12, 8-9 cm long and 5 mm diam at anthesis, 10-12 cm long and ca 1 cm diam in fruit, the stipes hir- tellous, to 1 cm long, the common peduncles greatly accrescent in fruit, 50-80 cm long, moderately covered with sharp and stiff, somewhat urticating trichomes.
Achenes tan, ellipsoid to narrowly ovate, smooth when fresh, ca 2.3 mm long, to 1.3 mm wide. Croat 15248a.
Apparently rare; known only from the vicinity of the Laboratory Clearing. In the Canal Zone, the plant is uncommon, growing as isolated plants in open areas. Both of the pistillate plants growing in the Laboratory Clear- ing set an abundance of fruit. Flowers at the beginning of the rainy season (April to June). The fruits mature from July to September (sometimes November).
Known only from Panama, from tropical moist forest on both slopes of the Canal Zone and in Darien and from premontane moist forest on the Pacific slope of the Canal Zone (Fort Kobbe).
See Fig. 195.
39.
MORACEAE/CECROPIA347
Cecropia obtusifolia BertoL, Fl. Guat. 39. 1840 C. mexicana Hemsl.; C. panamensis Hemsl.
Guarumo, Trumpet tree
Dioecious tree, mostly 5-10 m tall; trunk moderately slender, the young branches stout, ferruginous-hirtellous to glabrate. Petioles densely short-pubescent; stipules 7-11 cm long; mature leaves usually divided more than halfway to center, scabridulous and minutely arachnoid above, paler and minutely cinereous below; lobes usually 9-15, rounded to shortly acuminate at apex, somewhat to moderately narrowed at base, the lower free margin held
± flat to moderately erect, the leaves thus ± pleated, the area surrounding the petiole flat to 3-4 cm in all direc- tions from center; veins, petioles, and spathes often ± maroon. Staminate spathes (11)12-20 cm long, ca 2 cm wide at anthesis, gradually acuminate, the spadices usu- ally in clusters of 3-9, (10)12-18 cm long, 3-4 mm thick, subsessile or with puberulent stipes to 1 cm long, the common peduncle 8-12 cm long; pistillate spathes 16-20 cm long at anthesis, usually arachnoid on outside, villous inside, the spadices in clusters of 2-4, 17-30 cm long, ca 5 mm diam at anthesis, 6-9 mm diam and fleshy in fruit, the common peduncle 8-20 (24) cm long, usually rough with short erect trichomes. Achenes ovate to broadly oblong, somewhat flattened, the edges acute, 3.3-3.7 mm long, to 2.7 mm broad, tan to white. Croat 11716, 11800.
Very common along the edge of the lake, particularly the northern and the eroded eastern shorelines; occasional in the forest, especially in tree-fall areas, some trees being found in the older forest. Flowers and fruits throughout the year.
Most easily confused with C. peltata, which has leaves conspicuously pleated to the center with shorter, broader lobes and shorter spadices.
Fruits are taken by spider monkeys from April to August (Hladik & Hladik, 1969).
Southern Mexico to Panama, Ecuador, and probably Colombia. In Panama, a wide-ranging and ecologically variable species and a characteristic tree of tropical moist forest (Holdridge & Budowski, 1956); known from tropi- cal moist forest in the Canal Zone, Bocas del Toro, Ve- raguas, Panama, and Darien (no doubt elsewhere also), from premontane wet forest in Panama, from tropical wet forest in Colon and Darien, and from premontane rain forest in Panama (summit of Cerro Jefe).
See Fig. 196.
Cecropia peltata L., Syst. Nat. ed. 10,1286. 1759 C. arachnoidea Pitt.
Guarumo, Trumpet tree
Dioecious tree, 6-20 m tall. Petioles densely hirtellous and arachnoid, the basal pulvinus a uniform mass of dense trichomes; stipules 6-9 cm long; leaves lobed less than three-fourths of the way to middle, usually about halfway; lobes 9-11, obtuse to rounded at apex, scarcely or not at all contracted at base, the lateral margins held prominently upward and prominently pleated between