KEY TO THE SPECIES OF LACISTEMACEAE
Inflorescences less than 3 cm long; capsules more than 5 mm diam; axils of lower lateral veins not tufted Lacistema aggregatum (Berg) Rusby Inflorescences more than 4 cm long; capsules less than 4 mm diam; axils of lower lateral veins usu-
ally conspicuously tufted (axils glabrous elsewhere).... Lozania pittieri (S. F. Blake) L. B. Smith
their reduced flowers bearing a single stamen and by their small, fleshy, capsular fruits with arillate seeds. The family will be treated by H. Sleumer (pers. comm.) as Flacourtiaceae in his revision of that family.
Flowers are minute and clustered, each with a single anther. Pollination system is unknown.
The fruits display arillate seeds, and though dispersed principally by birds, they are also taken by monkeys (Oppenheimer, 1968).
Two genera and about 27 species; tropical America.
LACISTEMA Sw.
Lacistema aggregatum (Berg) Rusby, Bull. New York Bot. Gard. 4:447. 1907
Tree, usually 6-20 m tall, glabrate or with sparse puber- ulence on young stems and underside of leaf. Petioles 5-10(20) mm long; blades lanceolate, oblong-elliptic or oblanceolate-acuminate, acute to rounded at base, 10-16 cm long, 4.5-7 cm wide, entire or remotely crenate, the veins drying wrinkled, usually barbate in the axils of lateral veins on the lower surface, often pubescent all along midrib at least when young; stipules to 1 cm long, caducous. Flowers greenish, sessile, in narrow, cylindri- cal, bracteated spikes, the spikes 1-3 cm long, 4-12 per axil; bracts cupulate, broader than long, one subtending each flower; perianth segments 4, ca 0.5 mm long, un- equal, erose; ovary and stamen centrally situated on a broad fleshy disk, the disk subtended by free bracteoles (usually 3) exceeding width of disk; stamen solitary, ca 2 mm long, exceeding pistil, the connective bifurcate;
styles 3, short. Capsules ovoid, ca 1 cm long, red, short- stalked, splitting ± irregularly into 2 or 3 valves; seed 1, ca 7 mm long, surrounded by a fleshy, bitter, white aril.
Croat 5691, 8402.
Common in the forest, especially in the young forest.
Flowers in the dry and early rainy seasons, from January to May (rarely to July), usually in the latter half of the dry season. The fruits mature from April to June (some- times August).
The inner wall of one of the valves of the capsule becomes free and folds along the median. After being forced out, the seed is suspended on a slender white fiber from near the apex of the inner wall. Though the seeds are no doubt principally dispersed by birds, Oppen- heimer (1968) reported that white-faced monkeys eat the white aril associated with the seed.
Mexico to Colombia, Venezuela, the Guianas, and Peru; Trinidad, Greater Antilles. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Colon, Veraguas, Los Santos, Herrera, and Darien, from premontane wet forest in Code and Panama, and
from tropical wet forest in Colon (Guasimo). Reported from premontane rain forest in Costa Rica (Holdridge etal., 1971).
See Figs. 191 and 192.
LOZANIA MutisexCald.
Lozania pittieri (S. F. Blake) L. B. Smith, Phytologia 1(3):138. 1935
L. pedicellata (Stand!.) L. B. Smith
Shrub or small tree, to 8 m tall; stems and twigs brittle;
younger stems, petioles, and axes of inflorescences stri- gose. Petioles 3-8 mm long; blades elliptic to obovate- elliptic, acuminate, acute at base, 8-14 cm long, 2.5-4.5 cm wide, entire to obscurely toothed, glabrous above, the veins below weakly strigose and often pilose as well, especially in axils of lateral veins; stipules 1.5-2 mm long, caducous, the scar at most half-encircling stem. Flowers green, ca 1.5 mm diam, in solitary, axillary racemes to 9 cm long; axis and bracts pubescent; bracts minute;
pedicels ca 1.5 mm long; perianth oblique, 4-lobed, the lobes ± rounded, concave, the uppermost at first erect and ± enclosing the pistil, later spreading to expose the pistil, the others ± spreading, subtending the fleshy reniform disk; stamen 1, to 1 mm long, at first held near the upper perianth lobe with the anthers unopened, the filament later elongating, spreading away from the pistil, the anthers opening; anthers longitudinally dehiscent, broader than long, the thecae distinct, directed upward;
ovary minute, pubescent; styles 3, recurved (appearing as 3 hooks). Capsules globose, ca 4 mm diam, sparsely puberulent; seeds 3, minute, with a red aril. Foster 2364.
Apparently rare, but at least locally common. Collected by R. Foster on Balboa Trail 10 {Foster 2364) and by O. Shattuck at Gross Point (Shattuck 972). Flowering on BCI mostly in July and August. The fruits mature in August and September (mostly in August). Rarely flow- ering elsewhere in Panama in March, with the fruits maturing in April. Allen (1956) reported the species to flower in December on the Osa Peninsula in Costa Rica.
Costa Rica to northernmost Venezuela. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, and San Bias, from premontane wet forest in Colon and Panama, and from tropical wet forest in Colon and Veraguas.
38. ULMACEAE
Trees and shrubs, sometimes scandent and armed. Leaves alternate, petiolate; blades simple, generally serrate, pal- mately veined at base; stipules present, caducous. Flowers
Fig. 19\. Lacistema aggregation
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38.
ULMACEAE/TREMA341
KEY TO THE SPECIES OF ULMACEAE
Plants lianas or climbing shrubs, the stems armed Celtis iguanaeus (Jacq.) Sarg.
Plants unarmed trees:
Leaf blades ± glabrous except for midrib, not cordate at base; fruits more than 1 cm long Celtis schippii Standl.
Leaf blades pubescent, asperous especially above, cordate at base; fruits less than 5 mm long ...
Trema micrantha (L.) Blume unisexual or bisexual (monoecious or polygamomonoe-
cious), apetalous, sometimes obscurely zygomorphic, solitary or in axillary cymes; sepals 5, basally connate;
stamens 5, free, opposite the sepals; anthers 2-celled, dehiscing longitudinally; ovary superior, 1-locular, 2-carpellate; placentation apical; ovule 1, anatropous, pendulous; styles 2, basally fused (in Celtis, the stigmas bifurcate). Fruits drupes; seed lacking endosperm.
Members of the family resemble both the Moraceae (39), from which they differ by lacking milky or viscid sap and by having usually inequilateral leaf bases, and the Urticaceae (40), from which they differ by having a two- carpellate ovary and two styles. Some Urticaceae also have unequal leaf bases.
Pollination system unknown. Wind pollination is com- mon in the order Urticales (Faegri & van der Pijl, 1966).
Fruits are probably all animal dispersed. Trema mi- crantha has small red fruits and is probably bird dis- persed. Van der Pijl (1968) reported that the seeds of Celtis iguanaeus are eaten by iguanas.
Sixteen genera and about 300 species; widely dis- tributed.
CELTIS L.
Celtis iguanaeus (Jacq.) Sarg., Silv. North Amer.
7:64.1895 Hackberry, Cagalara
Andromonoecious (with staminate and bisexual flowers on the same plant) liana or climbing shrub, 10-25 m long, the trunk usually less than 10 cm diam, at least the smaller branches armed with recurved spines. Petioles 5-10 mm long; blades ± ovate, acute to acuminate at apex, obtuse to subcordate, 3-veined, and sometimes inequilateral at base, 3-11 cm long, 1.5-4.5 cm wide, sparsely pubescent to glabrous, serrate, the veins on underside prominently raised. Inflorescences cymose, axillary, the smaller branches and often pedicels and calyces minutely puberulent; flowers usually 5-parted, pedicellate, 1.5-2 mm long, green or greenish-yellow;
sepals ± oblong, the margins scarious, ± ciliate; petals lacking; staminate flowers with a rudimentary pistil ca 1.5 mm long, the stamens opposite the sepals, the fila- ments at first curved inward, ca 1.7 mm long, the anthers attached apically near the base of the pistillode, becoming free, the filament then straightening rapidly to fling pol- len from the anthers; pollen powdery; bisexual flowers similar to staminate flowers, the ovary 1-locular. Drupes ovoid, ca 1 cm long, ± 2-edged, yellow, orange, or red.
Croat 14649.
Occasional, in the forest, climbing high into the can- opy. Flowers in the early rainy season. The fruits mature in the middle to late rainy season.
Mexico to Argentina; the Antilles. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien.
Celtis schippii Trel. ex Standl., Field Mus. Nat. Hist., Bot. Ser. 12:409.1936
Tree to 30 m tall, to 60 cm dbh, often buttressed 1.5 m;
bark thin, planar, unfissured or with minute cracks; inner bark thick, with brown streaks; sparsely short-pubescent on young stems and leaves, ± glabrous in age except for midrib above and below. Petioles 5-9 mm long; blades ovate-elliptic to elliptic or ovate, acuminate, obtuse and slightly inequilateral at base, ± decurrent onto petiole, 6-14 cm long and 3.5-7 cm wide (to 24 cm long and 10 cm wide on juveniles), 3-veined at base, the larger lateral veins with inconspicuous cavelike domatia in axils be- neath (especially the apical axils) with 2-4 pairs of major laterals above the basal pair. Flowers axillary, solitary;
pedicels to 7 mm long in fruit; sepals 5, suborbicular, to 1 mm long, persistent, ciliate. Drupes narrowly ovoid, with a pungent odor, green at maturity, to 1.8 cm long, glabrous; pericarp leathery, ca 1 mm thick; seed with a brown testa, ca 1 cm long. Croat 16212.
Rare; known from a few individuals in the old forest.
Flowering unknown. Fruits in July; elsewhere in Central America mature-sized fruits have been seen in March and September.
Belize, Costa Rica, and Panama. In Panama, known only from tropical moist forest on BCI.
TREMA Lour.
Trema micrantha (L.) Blume, Mus. Bot. Lugduno- Batavum 2:58. 1856
T. integerrima (Beurl.) Standl.
Capulin macho, Capuli'n, Jordancillo
Monoecious tree, 2-20 m tall, the trunk to ca 16 cm dbh;
bark light brown, thin, prominently lenticellate; inner bark moderately thick, tan; pubescent all over with erect trichomes, the branchlets becoming glabrate; sap ± pungent, somewhat foul-smelling. Petioles 5-10(18) mm long; blades narrowly lanceolate, long-acuminate, cordate and often inequilateral at base, 5-15 cm long, 1.5-5 cm wide, asperous especially above, serrate to subentire;
stipules lanceolate, minute. Cymes axillary, to 1.5 cm long; flowers minute, 5-parted, green, greenish-yellow, or whitish; sepals to 1.3 mm long, minutely pubescent
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.