Adelobotrys adscendens (Sw.) Tr., J. Bot. 5:210. 1867 Liana or scandent shrub climbing to several meters; stems terete to ribbed, becoming glabrate in age, sometimes with fine roots along their length; younger parts, petioles, and inflorescences densely brown-strigose. Petioles 6-30
mm long; blades ovate to subrotund, acuminate, rounded to cordate at base, 4.5-15 cm long, 3-7 cm wide, thick, entire or denticulate, glabrous or strigose above, strigose below; veins usually 5. Panicles 10-40 cm long, of umbels of 2-6 flowers; pedicels 2-5 mm long; flowers 5-parted;
hypanthium ca 5 mm long, higher than ovary; calyx tube widely spreading, persisting and to 6 mm wide in fruit, the teeth minute or obscure; petals obovate, white to pink- ish, ca 1 cm long, rounded at apex, spreading; stamens
10, about as long as petals; filaments flattened, sometimes blue; anthers linear, deflexed, 6-10 mm long, the connec- tive elaborate, with a short part directed upward and a longer, dorsal, forked appendage directed basally ± parallel to the thecae; ovary superior; style shorter than stamens, becoming recurved near apex. Fruiting hypan- thia conspicuously ribbed at maturity, to 8 mm long, the
664
DICOTYLEDONEAEouter wall of fruit and the calyx eventually weathering away, the thin inner wall held in place by the stout ribs, the ribs then attached only at base; seeds very numerous, obcylindrical, ca 1 mm long, wind dispersed. Foster 2090.
Rare; collected once in the old forest. Probably flowers and fruits throughout the year.
The fruits are not obviously capsular until maturity.
Seeds are slowly dispersed as if by a saltshaker.
Mexico to Peru (Loreto), Bolivia, and Brazil; Jamaica.
In Panama, known from tropical moist forest on BCI and in Code and Darien, from premontane wet forest in Chiriqui, Code, and Panama, and from tropical wet forest in Colon, Code, Panama, and Darien.
ARTHROSTEMA Pav. ex D.Don
Arthrostema alatum Tr., Trans. Linn. Soc. London 28:35. 1871
Herb, to 60 cm tall; stems succulent, square, the edges narrowly winged; trichomes sparse, red, gland-tipped, on stems and inflorescences, including hypanthia and petals. Petioles to 3.5 cm long, winged; blades ovate, acute to acuminate, truncate to subcordate at base, 3-7 cm long, 1.5-4.5 cm wide, setose above, glabrous below, serrulate and ciliate; veins 5. Cymes terminal, divaricately branched, usually with a solitary flower between branches;
flowers 4-parted, ± sessile; hypanthium campanulate, to 4 mm long; calyx tube obscurely lobed, accrescent in fruit; petals pink to lavender, ca 3 mm long; stamens 8, ± equal; anthers with the connective prolonged some- what at base; filaments united to calyx tube; ovary supe- rior; style straight, about as long as stamens. Capsules campanulate, ca 1 cm long, 4-valved, loosening at base and acting as a wind ballast; seeds numerous, minute, cochleate, with grooves and minute tubercles on the rounded side. Croat 6150.
Uncommon, in the Laboratory Clearing. Flowers from August to October. Mature fruits seen in October.
Southern Mexico to Venezuela. In Panama, known from tropical moist forest in the Canal Zone and Panama and from premontane moist forest in Panama.
BELLUCIA Neck, ex Raf.
Bellucia grossularioides (L.) Tr., Trans. Linn. Soc.
London 28:141. 1871
Small tree, to 5 (10) m tall, glabrous except sometimes for puberulence on stems and lower leaf surfaces. Petioles to 6 cm, stout; blades broadly elliptic, obtuse to short- acuminate, obtuse to rounded at base, 15-35 cm long, 10-20 cm wide, entire, pliveined, the major veins 3-5, soon becoming glabrous. Cymes axillary, frequently on leafless branches and on trunk; pedicels 1-2.4 cm long;
flowers with an intense, sweet aroma at anthesis, ca 6 cm wide; hypanthium hemispherical; calyx 1.5-2 cm long, hyaline, dehiscing irregularly at anthesis and falling free from the hypanthium; petals 5 or 6 (to 8), oblong, thick, white or pink, to 3 cm long, broader and often very oblique at apex, spreading at anthesis; stamens 10-16,
to 1.8 mm long; filaments about as long as anthers; an- thers obovate, turned inward, yellowish, with 2 small terminal pores, both the thecae and connective swollen;
ovary inferior; style straight, stout, at first medially posi- tioned, soon pushed to one side of flower; stigma large, to 4 mm wide, capitate, held well above stamens. Berries
± globose, to ca 3 cm diam, pale green at maturity; seeds tan, numerous, ca 0.7 mm long, about twice as long as broad. Shattuck 1119.
Collected by O. Shattuck on Gigante Bay and possibly still occurring there, although the species is undoubtedly more abundant elsewhere. Flowers during July and August; mature fruits have been seen in October.
The species has been confused with B. axinanthera Tr., which is more abundant and apparently more eco- logically variable as well, occurring in Panamanian life zones ranging from premontane moist to premontane rain forests. B. costaricensis Cogn., reported by Gleason (1958) in the Flora of Panama, is a synonym of B. axinan- thera. The calyx of B. axinanthera differs from that of B. grossularioides by being regularly dehiscent, with lobes that are thick, regularly acute, and persistent in fruit.
The calyx lobes of B. grossularioides are hyaline, and their dehiscence is irregular and often circumscissile.
Southern Mexico to Amazonian Brazil and Peru. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, and Darien.
CLIDEMIA D.Don
Clidemia is distinguished by having axillary inflores- cences and petals rounded at the apex. The ovary is infe- rior. Most species are conspicuously pubescent.
Clidemia capitellata (Bonpl.) D. Don ex DC, Prodr.
3:159. 1828 C. neglecta D. Don
Shrub, 1-3 m tall; stems, inflorescences, and lower leaf surfaces densely pubescent with both stalked-stellate and simple trichomes, the simple trichomes often glandular- tipped, usually longer than the stellate ones. Petioles
1-5 cm long; blades ovate to ovate-oblong, acuminate, obtuse to rounded at base, 6-15 (20) cm long, 3-7 (9) cm wide, minutely toothed, pubescent above, the trichomes long, simple, set on bulbous projections of surface ap- pearing as depressions on lower surface; veins 5-7. Flow- ers sessile, 5-parted, clustered and well spaced on a strong central axis and at the ends of short branches; hypanthium and calyx tube together 3-4 mm long; calyx lobes 1-1.5 mm long, the exterior teeth slender, ca 1 mm longer than calyx lobes; petals white, 4.5-5 mm long, spreading at anthesis and later closing; stamens folded in bud, all directed to one side at anthesis, the terminal pore of the anthers held somewhat beneath style; style erect or held to one side; stigma held just above the ± closed petals after stamens shrivel. Berries subglobose, purple, densely pubescent, ca 6 mm diam; seeds minute, numerous. Croat 11770.
Though the species is common in the Canal Zone in disturbed areas, it is uncommon on the island and has
107.
MELASTOMATACEAE/CLIDEMIA665
KEY TO THE SPECIES OF CLIDEMIA
Flowers 4-parted, the flowers or at least the fruits often occurring on a leafless stem below leaves:
Leaf blades usually attenuate at base and decurrent on petiole, not cordate C. septuplinervia Cogn.
Leaf blades cordate or subcordate at base C. purpureo-violacea Cogn.
Flowers 5- to 8-parted:
Pubescences of simple trichomes only:
Plants with stems, blades, and inflorescence branches setose, the trichomes to 1 cm long C. collina Gleason Plants ± hirsute all over, the trichomes ca 1 mm long or less C. dentata DC.
Pubescence (at least on stems) of 3 types: stellate, simple, and simple gland-tipped:
Trichomes of upper blade surface often on conspicuous projections appearing as depressions from the underside; simple trichomes only slightly or a few times longer than the stellate ones; petals 4-5 mm long C. capitellata (Bonpl.) DC.
Trichomes of upper blade surface distributed evenly, the blades lacking projections; simple trichomes usually many times longer than the stellate ones; petals 8-13 mm long C. octona (Bonpl.) L. O. Wms.
been collected only at Gross Point. Flowers and fruits throughout the year.
Recognized by its unusual inflorescences and pustular leaves.
Southern Mexico to Colombia; West Indies. In Pan- ama, known principally from tropical moist forest in the Canal Zone, Bocas del Toro, Chiriqui, Veraguas, Pa- nama, and Darien Provinces; known also from premon- tane moist forest in the Canal Zone, from premontane wet forest in Bocas del Toro, Colon, Chiriqui, Code, and Panama, and from tropical wet forest in Darien and on the Atlantic slope in Veraguas, Colon, and Code.
See Fig. 427.
Clidemia collina Gleason, Phytologia 3:359. 1950 Slender shrub, to 2 m tall; stems, petioles, major veins of lower blade surfaces, and inflorescences conspicuously setose, the trichomes of stems mostly to 1 cm long, often deciduous in age. Petioles 1-3 cm long, the basal part densely setose and usually encrusted with ant detritus, the apical part with swollen domatia inhabited by small brown ants; blades ± elliptic, gradually acuminate, narrowly rounded at base, 15-32 cm long, 5.5-11.5 cm wide, very sparsely setose above, more densely so on major veins below, conspicuously ciliate, pliveined, the veins 7. Flowers 5-parted, in densely setose, branched, axillary clusters; pedicels pink; hypanthium and calyx tube campanulate, ca 3 mm long, the upper part with setae 6-10 mm long; calyx lobes blunt, 1-2 mm long, the exterior teeth slender, setose, about as long as or longer than sepals; petals white, oblong-obovate, rounded at apex, 4-6 mm long; stamens straight, yellow, to 5.5 mm long; anthers to 3 mm long with a single terminal pore; style only slightly exceeding stamens. Fruiting hypanthia unknown. Foster 2355.
Apparently rare on BCI, collected once south of Zetek Trail 800. Seasonal behavior not determined. Flowers at least in April and May, with some fruits maturing in August.
Known only from Panama, from tropical moist forest on BCI and in Bocas del Toro and from premontane wet forest in Code (El Valle).
Clidemia dentata D. Don ex DC, Prodr. 3:158. 1828 Shrub, 1-4 m tall, densely pubescent with long, stiff, erect or recurved trichomes. Leaves paired, one often smaller than the other; petioles to 1.5 cm long; blades narrowly ovate-elliptic, acuminate, acute to rounded and usually unequal at base, 6-20 cm long, 2.5-8 cm wide, entire or minutely serrate, pliveined, the veins 5-7. In- florescences short, paniculate or racemose, bearing few flowers; hypanthium and calyx bell-shaped, 3-4 mm long;
calyx truncate, the exterior teeth subulate, to 4 mm long;
petals 5, white, turning brown on drying, oblong, 5-7 mm long; stamens to 4 mm long; filaments arcuate below anthers; style slightly shorter than petals; stigma simple.
Berries subglobose, to 1 cm diam, green turning blue- black at maturity; seeds minute, numerous. Croat 11940.
Uncommon, within the forest and at the edges of clearings. Flowers and fruits throughout the year.
Southern Mexico to Bolivia and Brazil. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Code, and Darien, from pre- montane moist forest in the Canal Zone, from premon- tane wet forest in Bocas del Toro and Code, and from tropical wet forest in Colon, Code, and Darien.
See Fig. 428.
Clidemia octona (Bonpl.) L. O. Wms., Fieldiana, Bot.
29:558. 1963
Heterotrichum octonum (Bonpl.) DC.
Shrub, (1)2-3(5) m tall; stems, petioles, peduncles, and hypanthia with both stellate and simple trichomes, the simple ones in part very long, in part short and glan- dular. Leaf pairs sometimes unequal; petioles 2-10 cm long; blades ovate, acuminate, cordate at base, 8-20 cm long, 6-12 cm wide, chiefly with long simple trichomes above and short stellate ones below, denticulate; veins 7. Panicles upper-axillary and subterminal; flowers 7- or 8-parted; hypanthium and calyx lobes together to 5 mm long, the exterior teeth slender, much longer than calyx lobes, ca 1.5 mm long; petals obovate, 8-13 mm long, white, broadly spreading; stamens folded in bud, erect and radially symmetrical at anthesis; anthers ca 6 mm long, gradually tapered to apex with a single termi-
Fig. 427.
Clidemia capitellata
Fig. 429.
Clidemia octona
107.
MELASTOMATACEAE/CONOSTEGIA667
nal pore; style stout, erect in bud, elongating after an- thesis, held to one side well above anthers, to ca 1 cm long. Berries purple at maturity, 1—1.5 cm diam, the pubescence of the hypanthium persisting; seeds minute, numerous. Croat 11848.
Occasional, at the edge of the forest. Flowers and fruits throughout the year.
Southern Mexico to Peru and Brazil; Cuba, Jamaica.
In Panama, known from premontane moist forest in the Canal Zone and Veraguas, from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Veraguas, Los Santos, Herrera, Panama, and Darien, from premontane wet forest in the Canal Zone, Chiriqui, and Panama, and from tropical wet forest in Panama and Darien.
See Fig. 429.
Clidemia purpureo-violacea Cogn., Bull. Soc. Roy.
Bot. Belgique ser. 2, 30(1):263. 1891
Slender shrub, 1-5 m tall, often arching; stems glabrous in age. Petioles 1-8 cm long, bearing minute, furfur- aceous, stellate pubescence; blades broadly ovate, acumi- nate, rounded to cordate at base, 12-22 cm long, 3-14 cm wide, minutely stellate-pubescent on veins below and above near base, minutely denticulate and ciliate, ± ma- roon below; veins 5-7. Inflorescences axillary, small, pa- niculate, to ca 1.5 cm long, mostly below leaves; branches of inflorescence inconspicuously stellate-pubescent;
flowers 4-parted, pale red, ca 4 mm diam; hypanthium
± globose, 2-3 mm long, minutely pubescent, the tri- chomes borne on minute papillae, often deciduous; ex- terior teeth ending in deflexed setae ca 0.5 mm long; calyx lobes 0.5-0.7 mm long, markedly recurved with a sub- marginal projection forming the crown; petals oblong, rounded at apex, ca 1.5 mm long; stamens erect, ca 3 mm long; style ca 2 mm long, becoming directed to one side.
Berries to 5 mm long, ± globose, at first red-violet, turn- ing purple-black at maturity; seeds minute, numerous.
Croat 14551.
Rare, known only from deep ravines in the vicinity of Creek #8. Flowers and fruits from April to December.
Most fruits mature during the rainy season.
Costa Rica to Colombia. In Panama, known principally from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, and Darien; known also from premon- tane wet forest in Panama and from tropical wet forest in Darien.
Clidemia septuplinervia Cogn. in Mart., Fl. Brasil.
14(4):506. 1888
Shrub, 1.5-3(7) m tall; stems, petioles, lower leaf sur- faces, axes of inflorescences, and hypanthia bearing very minute, reddish-brown, granular puberulence. Leaf pairs sometimes unequal; petioles to 6 cm long; blades ovate to elliptic, acuminate, attenuate (rarely rounded) and sometimes inequilateral at base, decurrent onto petiole, 11-24 cm long, 5-12 cm wide, glabrous above, sparsely ciliate, subentire, pliveined, the veins 5-7. Cymes axil- lary, minute, ca 1 cm long, usually occurring on leafless stems; hypanthium ovoid or oblong, to 3 mm long; calyx lobes 4, rounded, ca 1 mm long, reddish, the exterior teeth slender, 3-4 times as long as sepals with a few long bristles; petals 4, pink or white. Berries globose, 3-5 mm diam, green then red turning blue at maturity, minutely puberulent; seeds minute, numerous. Foster 1096.
Collected once west of Armour Trail 100-200. Flowers and fruits throughout the year.
Belize to Colombia, Peru, and Brazil; Jamaica. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, and Darien, from trop- ical wet forest in Veraguas (Atlantic slope), Colon, Pa- nama, and Darien, and from premontane rain forest in Darien.
CONOSTEGIA D.Don
Conostegia is distinguished by having terminal inflores- cences, petals obtuse or rounded at apex (acute in C.
cinnamomea), and flower buds with a calyptrate calyx (i.e., the entire calyx deciduous as a unit leaving an ir- regular scar). The ovary is inferior.
Conostegia bracteata Tr., J. Bot. 5:209. 1867 Shrub, to 3.5 m tall, conspicuously hirsute except on petals and inner flower parts, the trichomes long, simple, sometimes slightly branching, especially on main veins and below inflorescences. Petioles to 2 cm long; blades narrowly elliptic to oblanceolate, acuminate, acute to obtuse at base, 9.5-20 cm long, 4-8 cm wide, obscurely serrulate; veins 3. Panicles to ca 10 cm long; flowers in small glomerules subtended by oblong-ovate bracts to 8 mm long; hypanthium 2.5-3 mm long at anthesis;
petals (5)6(7), obovate, rounded to truncate at apex,
KEY TO THE SPECIES OF CONOSTEGIA
Leaf blades glabrous above or nearly so at least at maturity; trichomes of lower surface sessile, stellate (minute in C. cinnamomea):
Lower leaf surface completely concealed by a fine close mat of brownish stellate trichomes C. xalapensis (Bonpl.) DC.
Lower leaf surface not concealed by trichomes, sparsely pubescent at maturity (often superfi- cially glabrous) C. cinnamomea (Beurl.) Wurdack Leaf blades conspicuously pubescent above, the trichomes chiefly or entirely simple:
Trichomes of lower blade surface all simple (rarely a few on principal veins branched but not stellate) C. bracteata Tr.
Trichomes of lower blade surface often both simple and stellate, the stellate trichomes mostly stalked C. speciosa Naud.
107.
MELASTOMATACEAE/CONOSTEGIA669
white, 7-8 mm long; bracts and petals burnt-orange when dry; stamens 12-18, 5-6 mm long; filaments geniculate below anthers; anthers with terminal pores all directed toward style in vicinity of stigma; style straight. Berries round, hirsute, black and shiny, 8-12 mm wide; seeds minute, numerous. Croat 11079.
Occasional, in the forest. Flowers from March to July (rarely later). The fruits mature from August to November.
Nicaragua to Panama. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Colon, Panama, and Darien, from premontane wet forest in Code and Panama, from tropical wet forest in Panama and Darien, and from premontane rain forest in Darien.
See Fig. 430.
Conostegia cinnamomea (Beurl.) Wurdack, Phytologia 38:287. 1978
C. micromeris Standl.
Shrub, usually 1.5-3.5 m tall; younger parts furfuraceous with stellate trichomes, soon glabrate. Petioles 2-5(10) mm long; blades broadest at about middle, gradually tapered to both ends, acuminate, mostly 6-22 cm long, 2.5-8.5 cm wide, bicolorous, glabrate above, sparsely stellate-pubescent below especially on veins, pliveined, the margins entire or minutely wavy (sinuate), the veins 5-7. Terminal compound dichasia 3-6 cm long, divari- cately branched, the branches opposite, each subtended by a pair of narrow bracts; flowers sessile or short- pedunculate, minutely furfuraceous, the buds 3-4.7 cm long, distinctly apiculate, short-pedicellate; hypanfhium 2-3 mm long at anthesis; petals (4)5, white to pink, 4-4.7 mm long, acute at apex, recurved, soon falling;
stamens (8)10, erect at anthesis; filaments flattened, slightly longer than the anthers, markedly curved; an- thers with terminal pores; style ca 5 mm long. Berries depressed-globose, to 6 mm diam, green turning light blue then deep purple at maturity; seeds minute, num- erous. Croat 6250.
Frequent in the forest. Flowers and fruits throughout the year, principally from July to September.
Recognized by its small, inconspicuous inflorescences with beaked buds. The flowers fall off soon, and the plant seldom appears to be conspicuously in flower.
Nicaragua to western Colombia. In Panama, known principally from tropical moist forest in the Canal Zone, Colon, Panama, and Darien; known also from premon- tane wet forest in Code and from tropical wet and pre- montane rain forests in Darien.
See Fig. 431.
Conostegia speciosa Naud., Ann. Sci. Nat. Bot, ser.
3, 16:109. 1851
Shrub, seldom more than 2 m tall, very densely pubescent on all external surfaces, the trichomes of stems and in- florescences stalked-stellate. Petioles 1-4 cm long; blades
± ovate, acuminate, obtuse to rounded at base, 10-27 cm long, 5.5-13 cm wide, rugulose and mostly simple- pubescent above, with both stalked-stellate and simple
trichomes below, minutely serrate, pliveined, the veins 5-7, raised below. Panicles congested, ± maroon, ter- minal; flowers ca 1.5 cm diam, opening few at a time, often on same inflorescence with fruits, the buds violet, ovoid; petals 6 or 7, white to pink, ± rounded; stamens 12 or 14, directed to one side, geniculate below anther;
anthers all with apices clustered around stigma; style short and thick, strongly bent to one side near apex;
stigma green. Berries densely pubescent, dark purple, ca 1.5 cm diam; seeds minute, numerous. Croat 6340.
Common in clearings; occasional along the margin of the lake. Flowers and fruits throughout the year.
Foster 1134 and 1211, collected along the shore on Pena Blanca Point, represent an extreme variation of C.
speciosa or possibly a distinct variety of it. It has been suggested by C. Schnell (pers. comm.) that they are hy- brids between C. speciosa Naud. and C. subcrustulata (Beurl.) Tr. In general appearance they look like C. speci- osa, but they have only sessile stellate trichomes on the lower leaf surface as in C. subcrustulata.
Costa Rica to northern South America; Jamaica. In Panama, known principally from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Veraguas, Panama, and Darien; known also from tropical dry forest in Panama (Taboga Island), from premontane moist forest in the Canal Zone and Panama, from premontane wet forest in Code, and from tropical wet forest in Colon and Code.
Conostegia xalapensis (Bonpl.) D. Don ex DC, Prodr.
3:175. 1828 Canallito
Shrub or small tree, to 10 m tall (often only 3-4 m on BCI); all parts except interior parts of flowers and upper blade surfaces densely pubescent, the trichomes ± ses- sile, brown, stellate. Petioles 1-2.5 cm long; blades ± lanceolate, acuminate, obtuse to cuneate at base, 5-25 cm long, 1.7-8 cm wide, glabrate above in age, usually serrate, pliveined, the veins 3-5. Inflorescences terminal, paniculate; flower buds pyriform; petals 5, glabrous, white (drying yellow), obovate, ca 5 mm long; stamens
10; filaments bent sharply below anthers; anthers with terminal pores, all directed to one side of the flower (as is the stigma), the connective thickened on upper side;
style bent to one side, thickened at apex; stigma papillate.
Berries densely stellate-pubescent, green to purple at maturity, to 8 mm diam; seeds minute, numerous. Croat 12281.
Uncommon, collected only along the shore on the north side of the island. Flowers and fruits throughout the year.
Southern Mexico to northern South America; Cuba. In Panama, ecologically variable; known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, and Darien, from premontane wet forest in Chiriqui, Code, and Panama, from tropical wet forest in Veraguas (At- lantic slope), Colon, Panama, and Darien, and from premontane rain, lower montane moist, and lower mon- tane wet forests in Chiriqui.