KEY TO THE TAXA OF POLYGONACEAE
Herbs; leaves long and narrow, less than 3.5 cm wide; flowers perfect Polygonum Trees, shrubs, or woody vines; leaves more than 3.5 cm wide (if less, then ovate); plants usually
dioecious or the flowers functionally unisexual:
Tree more than 10 m tall and more than 12 cm dbh, the stems hollow, usually inhabited by fiercely stinging ants; achenes contained within enlarged, papery, reddish, 3-winged calyx more than 4 cm long Triplaris cumingiana Fisch. & C. Meyer Woody vines, shrubs, or small trees usually less than 10 m tall or less than 12 cm dbh, usually
lacking hollow stems (as adults) and not inhabited by stinging ants; achenes drupelike, en- veloped in fleshy perianth lobes Coccoloba
complex system of fruit dispersal. A host-dependent phytophagous insect might be suspected.
Range is difficult to determine because of the incon- spicuous habit of the plant. Probably ranging from Belize to Brazil. In Panama, known from tropical moist forest in the Canal Zone and from premontane wet forest in Code (El Valle).
See Fig. 218.
46. POLYGONACEAE
Trees, shrubs, lianas, or succulent-stemmed herbs (stems hollow in Triplaris). Leaves alternate, petiolate; blades simple, entire; venation pinnate; stipules present as sheathing ocreae. Flowers bisexual or unisexual (func- tionally monoecious in Coccoloba, dioecious in Triplaris), actinomorphic, on terminal spikes, racemes, or panicles in ocreolate fascicles of 1 to few flowers; tepals 4-6, free, in 1 series (in Triplaris, 6, connate, in 2 series), accrescent in fruit, becoming winged or bladdery; stamens 6-9, in basically 2 series, free or adnate; anthers 2-celled, at least the outer introrse, dehiscing longitudinally; ovary superior, 1-locular, 2-4-carpellate, subtended by an annular, nectar-secreting disk; placentation seemingly basal; ovule 1, orthotropous; styles and stigmas 2 or 3.
Fruits achenes (enveloped by the fleshy perianth lobes and drupelike in Coccoloba); seed with mealy endosperm.
Polygonaceae are distinguished by their ocreate stipules.
Pollination systems are unknown. Some members of the family have dimorphic heterostyly (Faegri & van der Pijl, 1966).
The seeds of Triplaris cumingiana are wind dispersed, but those of probably most Polygonaceae are principally bird dispersed. Coccoloba generally has a thin fleshy mesocarp and a well-protected seed, but in the case of C. acuminata the shiny dark seed is displayed against the greatiy accrescent, fleshy, white sepals. The achenes of Polygonum are probably similarly displayed, but since
most occur in aquatic habitats they are perhaps in part water dispersed as well. Van der Pijl (1968) suggested that seeds of Polygonum may be dispersed on the feet of shore birds. Despite their close association with Triplaris, ants of the genus Pseudomyrmex do not function in polli- nation or fruit dispersal (Wheeler, 1910). They do benefit the plant, however, by warding off predators such as leaf-cutter ants (Atta). The sting of the genus Pseudo- myrmex is severe and no doubt provides Triplaris with considerable protection.
About 40 genera with some 800 species; mostly of North Temperate distribution.
COCCOLOBA P.Browne
Coccoloba acapulcensis Standl., Proc. Biol. Soc. Wash.
33:66. 1920
Monoecious tree, 4-8 m tall, glabrous but with the lower midrib of leaf minutely hirtellous; trunk slender, some- times forming adventitious roots. Leaves borne individ- ually at apex of short shoots to ca 1 cm long; blades ovate, narrowly very long-acuminate at apex, rounded at base, 4-10 cm long, 2-6 cm wide, the midrib raised above;
major lateral veins in 3 or 4 pairs. Flowers brownish- purple, borne in racemes to 2 cm long from near apex of short shoot before leaves appear; rachis hirtellous; bracts
± lanceolate, ca 0.5 mm long; pedicels ca 1 mm long;
staminate flowers borne individually along rachis (usually less than 12 per inflorescence); tepals usually 5, oblong- obovate, rounded at apex, ca 1.5 mm long, ± spreading;
stamens usually 9, to 4.5 mm long; pistillate flowers not seen. Fruits narrowly ovoid, red, to 9 mm long and 6 mm wide (dried), often moderately well developed before leaves appear. Croat 8339, 14991.
Apparently rare on the island; collected on Standley Trail 1340 and north of Zetek Trail on the escarpment in the old forest. Flowers in early April before the leaves appear. The fruits usually develop by late April, and the trees have leaves by the time the fruit fully matures (see Gentry 5163, Madden Lake).
KEY TO THE SPECIES OF COCCOLOBA
Midrib sunken above (at least with respect to blade surface) and usually pubescent with conspicu- uous, brown, hispid or puberulent pubescence (the trichomes sometimes deciduous in age) C. manzanillensis Beurl.
376 DICOTYLEDONEAE
Midrib raised above, usually glabrous or inconspicuously puberulent (never with conspicuous long trichomes):
Leaves less than 10 cm long, ovate, narrowly very long-acuminate at apex, rounded at base, borne on a short shoot about 1 cm long; inflorescences ca 1-2 cm long C. acapulcensis Standl.
Leaves usually more than 10 cm long, not shaped as above; inflorescences much longer:
Leaves lanceolate to ovate-lanceolate, gradually acuminate, often less than 5 cm wide, with prominent axillary tufts on lateral veins of lower surface; inflorescences spicate; fruiting perianths red or white; plant usually a shrub C. acuminata H.B.K.
Leaves not as above, often more than 5 cm wide, generally lacking axillary tufts below; inflo- rescences racemose; fruiting perianths never red or white; plants usually a tree or vine:
Plant usually a vine; blades often subrounded, broadly rounded at base, usually blunt to rounded at apex, the midrib often with some short puberulence above; lepidote scales on blades reddish-brown C. parimensis Benth.
Plant a tree; blades acuminate at apex, not subrounded; midribs usually glabrous above; lepi- dote scales usually pale, easily seen under magnification C. coronata Jacq.
Panamanian material, though identified as C. acapul- censis by R. A. Howard, differs radically from specimens elsewhere in Central America. However, the species is quite variable.
Mexico to Panama. In Panama, known from tropical moist forest in the Canal Zone (BCI and near Madden Lake) and from tropical wet forest in adjacent Colon (Rio Pina-Rio Media divide). The species tends to grow only in regions where there are extensive outcrops of limestone.
See Fig. 219.
Coccoloba acuminata H.B.K., Nov. Gen. & Sp.
2:176. 1817
Slender, functionally monoecious shrub or tree, 2-8 m tall; stems glabrous to puberulent, with sparse round lenticels; ocreae to 1 cm long. Petioles from base of ocrea, 4-15 mm long; blades mostly oblong-lanceolate or oblong-elliptic, acuminate, cuneate to nearly rounded at base, mostly 6-18 (22) cm long, 2-5 (8) cm wide, glabrous above, glabrous to weakly pubescent on midrib and veins below, the vein axils tufted, the midrib arched. Inflores- cences terminal, spicate, slender, mostly 15-25 cm long;
rachis ridged, swollen at each flower cluster; staminate flowers red, ca 2 mm long, appearing successively from the apex of a very short, stout, bracteated stalk; perianth lobes 5, ovate, imbricate, 6-8 mm long, fleshy, accrescent, reddish turning white at maturity; stamens 8 (9), included, the filaments fused into a tube ca half their length; anthers blocking entrance to nonfunctional ovary. Pistillate flow- ers usually borne singly at each nodule of rachis; ovary 3-sided; styles 3, short, held somewhat below anthers;
nectar stored within staminal tube. Fruits subglobose, 6-8 mm wide, the achene strongly 3-sided, shiny black or brown, enveloped by 3 of the 5 perianth lobes and contrasting sharply with them in color, protruding at maturity. Croat 15101.
Common along some areas of the shore. Flowers and fruits throughout the year.
Panama to Ecuador, Peru, and Brazil. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien and from tropical wet forest in Darien.
Coccoloba coronata Jacq., Enum. Syst. PL Ins. Carib.
19. 1760 C. obovata H.B.K.
Pena blanca, Papaturra blanca
Apparently dioecious tree, 4-13 m tall, the trunks mostly less than 15 cm diam and clustered, usually producing sucker shoots from near base, the internodes often hollow;
branches glabrous, the younger parts with lepidote scales (striate when dry); bark smooth. Petioles ± terete, 1-2.5 cm long, inserted above base of stipule (about midway on stipule); blades ± ovate to obovate, abruptly to gradually acuminate at apex, acute to obtuse or subcordate and sometimes inequilateral at base, 8-24 cm long, 3-9.5 cm wide, glabrous but with both surfaces bearing ± promi- nent lepidote scales; midrib raised above; juvenile leaves red to wine-colored, to 28 cm long and 14 cm wide or larger. Inflorescences slender, solitary, erect, on short lateral branches; flowers numerous, white, ca 2 mm long, subtended by brown ocreolae; staminate inflorescences to 25 cm long; stamens usually 8, exserted. Pistillate inflo- rescences 16-20 cm long, the flowers sessile or short- pedicellate; staminodia usually 8, short; styles 3; stigmas spatulate, flattened. Fruits sessile or short-stalked, ovate,
± terete in cross section when fresh (prominently striate when dry), to 12 mm long and 8 mm wide. Croat 11962.
Occasional, in the forest. Usually flowering from late June to August, less frequently in September. The fruits mature from September to December.
The hollow stems may be inhabited by small ants.
Guatemala to Colombia and Venezuela; Trinidad, southern Windward Islands. In Panama, known princi- pally from tropical moist forest in the Canal Zone, San Bias, Chiriqui, Veraguas, Panama, and Darien; known also from tropical dry forest in Herrera (Pese) and from premontane wet forest in Chiriqui, Code, and Panama.
See Fig. 220.
Coccoloba manzanillensis Beurl., Prim. Fl. Portobello in K. Vetensk. Acad. Handl. 142. 1854(1856)
C. nematostachya (Griseb.) Lindau Hueso
Monoecious tree, usually less than 13 m tall; branchlets densely appressed-pubescent; stems at first grayish-
'151
pubescent, soon becoming glabrous, gray, conspicuously lenticellate (striate when dried); ocreae divided to middle or below, to 4.5(9) cm long. Petioles 1-2 cm long (to 4.5 cm on juveniles), inserted at base of ocrea, puberulent (also with longer trichomes near base); blades oblong- elliptic to obovate, short-acuminate (often downturned) to rounded at apex, narrowing below middle, narrowly cordate or rounded at base, mostly 9-28 cm long, 4.5-J.9 cm wide, young, thin, and often reddish when in flower, mature, thick, and bullate in fruit, ± glabrous above but with the veins densely brown-hirsute (trichomes some- times deciduous in age), the veins sunken above, conspic- ously raised and puberulent to hirsute below. Inflores- cences spikelike racemes to 40 cm long, pendent from the ends of the main stem or from short lateral branches near apex; flowers greenish, ovate, ca 1.5 mm long, function- ally unisexual; staminate flowers in clusters of 3 or 4;
tepals 4 or 5, green; stamens 6-8, long-exserted, spread- ing; pollen white, moderately tacky; pistillode with 2 or 3 styles. Pistillate flowers densely congested on racemes, borne singly at each nodule. Fruits globose (beaked and striate on drying), ca 6 mm diam, reddish to violet-purple at first, fleshy, sweet, and purple when mature; achenes ovoid, beaked, dark brown, ca 4 mm long, drying trig- onous. Croat 9558, 15071.
Adult plants common along the shore but uncommon within the forest; juveniles are abundant within the forest, especially in the younger forest. Juveniles are typically unbranched with hollow, densely pubescent stems and leaves that are very large (usually 35-50 cm long and
18-27 cm wide), short-petiolate, elliptic to obovate or oblanceolate, with pilose, deeply divided stipules to 11 cm long. Plants lose their leaves shortly before flowering and generally flower as new leaves are emerging. Flowers from late rainy season into the dry season. The fruits are mature in the rainy season, mostly from June to Sep- tember.
Known only from Panama, from tropical moist forest in the Canal Zone, Panama, and Darien and from tropical wet forest in Colon and Panama.
See Figs. 221 and 222.
Coccoloba parimensis Benth., London J. Bot.
4:626. 1845 C. leptostachya Standl.
Monoecious liana, extending into top of canopy (juvenile sometimes appearing to be suberect shrub; plant report- edly a tree elsewhere); stems minutely puberulent, be-
46.
POLYGONACEAE/POLYGONUM379
coming glabrous (striate when dry), often ± flattened.
Petioles mostly 2-3 cm long, (to 5 cm on juveniles) weakly canaliculate above; stipules ocreate, 2.5-5 cm long, subpersistent; blades broadly ovate to elliptic or obovate, rounded to acuminate at apex, rounded to sub- cordate and sometimes inequilateral at base, mostly (5) 10-25 cm long and (3)5-16 cm wide (juveniles to 33 cm long and 22 cm wide), both surfaces with minute, reddish-brown, lepidote scales, the midrib raised, ± glabrous to minutely puberulent above, all veins conspic- uously raised below, usually loop-connected. Inflo- rescences axillary or terminal on short, lateral, leafy branches, usually solitary; all parts puberulent; flowers greenish, ca 2 mm long, each surrounded by a thin brown ocreola; staminate inflorescences to 16 cm long, the flow- ers clustered in distinct, somewhat spiral whorls; stamens slightly exserted; pistil rudimentary. Pistillate inflores- cences usually less than 10 cm long, the flowers solitary at each nodule; stamens rudimentary; pistil 2 mm long;
styles 3. Fruiting pedicels usually 3-4 mm long; fruits ovoid to globose, to 1 cm long, black, with a thin, fleshy mesocarp; seed 1, ovoid, ca 7 mm long, brown. Croat 4777, 15161.
Frequent in the forest. Flowers at the beginning of the rainy season in May and June. The fruits usually mature by September and November, sometimes as early as August.
Less common than C. manzanillensis.
Known from Panama, Colombia, Peru, and Brazil. In Panama, known only from tropical moist forest in the Canal Zone and Panama.
POLYGONUM L.
Polygonum acuminatum H.B.K., Nov. Gen. & Sp.
2:178. 1817
Robust herb, to 3 m tall (usually less), often growing in water; moderately to densely strigose on most parts.
Leaves sessile or subsessile; blades linear-lanceolate, long-acuminate at apex, abruptly decurrent at base, 10-30 cm long, 1-3.5 cm wide, strigose-ciliate, inconspicuously pellucid-punctate; ocreae 2-3 cm long, apically ciliate with long strigose pubescence 1-1.5 cm long. Inflores- cences terminal, spikelike racemes or panicles of few branches; flowers continuous on rachis, fasciculate, whit- ish, subtended by ciliate, imbricate ocreolae 2-3.5 mm long; pedicels exceeding ocreolae ca 1 mm, conspicu-
KEY TO THE SPECIES OF POLYGONUM
Leaves conspicuously strigose all over; ocreae with long apical cilia to 1.5 cm long P. acuminatum H.B.K.
Leaves glabrous or pubescent on veins only; ocreae with apical cilia lacking or less than 1 cm long:
Leaves, ocreae, and tepals conspicuously dark-punctate; midribs of leaves subglabrous P. punctatum S. Elliott Leaves, ocreae, and tepals inconspicuously pellucid-punctate; midribs of leaves strigose
P. hydropiperoides Michx.
380
DICOTYLEDONEAEously articulate at apex; tepals 4(5), ovate, 2.5-3.5 mm long; stamens usually 6, exserted, ca 3.5 mm long; ovary lenticular; styles 2, 2-3 mm long. Achenes lenticular, 2-2.5 mm long, 1.5-2 mm wide, brown to black, beaked.
White 142.
Rare, in marshy habitats. Seasonal behavior uncertain.
Probably flowers and fruits throughout the year.
Mexico to Argentina; West Indies. In Panama, known only from tropical moist forest in the Canal Zone.
Polygonum hydropiperoides Michx., Fl. Bor. Amer.
1:239.1803
Slender herb, to ca 1 m tall, often ± reclining, subgla- brous. Petioles usually less than 5 mm long; blades linear- lanceolate, long-acuminate at apex, acute and decurrent on petiole at base, 4-15 cm long, 0.5-1.5 cm wide, incon- spicuously pellucid-punctate, usually strigose on midrib and veins below, minutely ciliate; ocreae 1-3 cm long, with strigose apical cilia to 8 mm long. Inflorescences terminal, spikelike racemes or panicles of few branches;
flowers interrupted along rachis, fasciculate, light pur- plish to greenish, subtended by ciliate ocreolae 2-3 mm long; pedicels exceeding ocreolae ca 1 mm, articulate at apex; tepals usually 5, ovate, 2-3 mm long; stamens usu- ally 9, ca 1.5 mm long; ovary trigonous; styles 3, ca 1 mm long. Achenes trigonous, 2-3 mm long, brown to black, inconspicuously beaked. Shattuck 840.
Collected once by Shattuck at Gigante Bay. Apparently flowering and fruiting throughout the year.
Canada to South America. In Panama, known only from BCI.
Polygonum punctatum S. Elliott, Bot. S. Carolina &
Georgia 1:455. 1817 Chilillo, Chili de perro
Slender herb, to 70 cm tall, nearly glabrous. Leaves, ocreae, and tepals conspicuously dark-punctate; petioles less than 5 mm long; blades linear-lanceolate, acuminate at apex, narrowly decurrent on petiole at base, 2-10 cm long, 0.5-2 cm wide, glabrous but minutely ciliate and sometimes with the midrib near base sparsely strigose;
ocreae 5-15 mm long, with strigose cilia at apex usually 7-10 mm long. Inflorescences terminal, spikelike racemes or panicles of few branches; flowers interrupted along rachis, fasciculate, white or greenish, subtended by ciliate ocreolae 2-3 mm long; pedicels exceeding ocreolae 1-2 mm, articulate at apex; tepals 5, ovate, 3-4 mm long;
stamens usually 8, 1.5-2 mm long; ovary trigonous;
styles 3, ca 1 mm long. Achenes trigonous, 2.5-4 mm long, brown to black, somewhat beaked. Croat 5247.
Rare, in marshy places, especially sandbars. Flowers throughout the year, possibly with a peak in the late rainy and early dry seasons.
Canada to Argentina; West Indies. In Panama, known from tropical moist forest in the Canal Zone and Panama, from tropical dry forest in Code, and from premontane wet forest in Chiriqui and Code.
TRIPLARIS Loefl.
Triplaris cumingiana Fisch. & C. Meyer, Mem. Acad.
Imp. Sci. Saint-Petersbourg, Ser. 6, 6:149. 1840 Palo santo, Guayabo hormiguero, Vara santa Dioecious tree, usually 10-20 m tall; trunk smooth,
12-30 cm dbh; bark light brown, thin, peeling off; stems hollow. Petioles very short or to 2 cm long, canaliculate;
blades mostly oblong-elliptic, acuminate, obtuse at base, 15-30 cm long, 4-12 cm wide, glabrous but with the veins below strigose (especially midrib). Inflorescences from upper axils; staminate inflorescences of spikes to 35 cm long and 1.5 cm wide; flowers subsessile, usually in pairs, emerging one at a time from densely pubescent, spathaceous ocreolae; perianth greenish, in one series of 3 linear and 3 narrowly triangular tepals 3-4 mm long, connate for about half their length; stamens 9, exserted;
anthers introrse, versatile. Pistillate inflorescences of racemes to ca 20 cm long (to 30 cm in fruit); pedicels 2-9 mm long; calyx sericeous, ca 1 cm long at anthesis (greatly accrescent and becoming red in age), the lobes narrowly triangular to linear, 2-3 times the length of the tube and spreading at anthesis; petals ± linear, exceeding tube, fused to base of tube; styles 3, the inner surface stigmatic in upper two-thirds. Achenes sharply trigonous (the surfaces ± flat), 8-12 mm long, shiny, brown, persistent within and dispersed by the enlarged calyx, the calyx to 6 cm long, pubescent, 3-winged, the wings pinkish, spreading, 3.5-4.5 cm long, 6-7 mm wide. Croat 4633, 8165.
Usually locally common; otherwise only occasional in both the young and old forests. Plants may begin to flower when as litde as 11 m tall and 12 cm dbh. Flowers from February to April, chiefly in March. The fruits begin to mature by February and are dispersed chiefly in March and April but also in May. Plants lose their leaves in July and August.
Stems are inhabited by very aggressive ants (Pseudo- myrmex triplaridis ForeL), whose sting is quite severe.
Costa Rica to Ecuador; cultivated in the West Indies and elsewhere. In Panama, known from tropical moist forest in the Canal Zone, San Bias (Perme), Los Santos, Panama, and Darien and from tropical wet forest in Panama (Cerro Campana).
See Fig. 223.